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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(summer/fall) as one volume

by the Botanical Research Institute of Texas.

VOLUME 9 NUMBER 1 24 JULY 2015

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Table of Contents

SYSTEMATICS

Typification of Sedum rubroglaucum (Crassulaceae)

Peter F. Zika 1

Micranthes rufopilosa (Saxifragaceae) comb, nov.: an alpine species from Alaska and Yukon

David F. Murray and Reidar Elven 7

nrDNA sequence (ITS and ETS) from herbarium specimens reveals phylogenetic affinities of

Erigeron geiseri (Asteraceae)

Richard D. Noyes, William Caraway, and Dulcinea V. Groff 11

New combinations in Coryphantha and Escobaria (Cactaceae)

Root Gorelick 25

Scleria bellii (Cyperaceae), a distinctive and uncommon nutsedge from the

southern U.S., Cuba, and Mexico

Richard J. LeBlond, Samantha M. Tessel, and Derick B. Poindexter 31

Styrax peltatus (Styracaceae), a new species from Oaxaca, Mexico

Peter W. Fritsch 43

Nuevas especies de Macherium (Leguminosae: Papilionoideae: Dalbergiae) en

Mexico y Centroamerica

Jose L. Linares 49

Gnaphaliothamnus nesomii (Asteraceae: Gnaphalieae), a new species from Guatemala and

nomenclatorial changes

Michael O. Dillon and Federico Luebert 63

New combinations in Eumachia (Rubiaceae) for species occurring on the Guiana Shield

Piero G. Delprete and Joseph H. Kirkbride, Jr. 75

New species of Acanthaceae from Ecuador

Dieter C. Wasshausen 81

Cuatro nuevas especies y un nuevo registro de Freziera (Pentaphylacaceae) de Ecuador y Peru

Daniel Santamari'a-Aguilar y Laura P. Lagomarsino 89

New combinations in Hexasepalum (Rubiaceae: Spermacoceae)

Joseph H. Kirkbride, Jr. and Piero G. Delprete 103

Observations about Phaseolus lignosus (Leguminosae: Papilionoideae: Phaseoleae),

a bean species from the Bermuda Islands

D.G. Debouck 107

PALEOBOTANY

Dasylarynx anomalus gen. et sp. nov., a tubular monocotyledon-like flower in Mid-Tertiary

Dominican amber

George O. Poinar, Jr. and Kenton L. Chambers 121

Prioria dominicana sp. nov. (Fabaceae: Caesalpinioideae), a fossil flower in Mid-Tertiary

Dominican amber

George O. Poinar, Jr. and Kenton L. Chambers 129

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

Cretaceous Myanmar amber

George Poinar, Jr., Donis da Silva Alfredo, and Iuri Goulart Baseia 135

FLORISTICS, ECOLOGY, AND CONSERVATION

Furrowed blister pods stranded on northern Atlantic Ocean coasts represent an undescribed

Sacoglottis (Humiriaceae) endocarp most similar to the fossil Sacoglottis costata

Raymond van der Ham, Tinde van Andel, Linda Butcher, Izumi Hanno, Theo Lambrechts,

Kim Lincicome, Paul Mikkelsen, Jenifer Mina, Bob Patterson, Ed Perry, and Mike Romance 137

A survey of the woody and climbing plants of the Refugio de Aves Dr. Alexander Skutch,

“Los Cusingos,” Perez Zeledon Canton, Costa Rica

Ronald L. Jones, Humberto Jimenez-Saa, and Allen C. Risk 149

Cuscatlania vulcanicola (Nyctaginaceae) nuevo registro del genero y especie para la

flora de Honduras

Jose Ledis Linares, Frank Sullyvan Cardoza Ruiz y Patricia HernAndez-Ledesma 167

Structural analysis of shrublands adjacent to the Monterrey Metropolitan Area, Mexico

Pamela Anabel Canizales-VelAzquez, Oscar Alberto Aguirre-Calderon,

Eduardo Alani's-Rodri'guez, Eduardo Javier Trevino-Garza, and Jose Manuel Mata-Balderas 173

Matelea chihuahuensis (Apocynaceae): an addition to the flora of the United States and

a synopsis of the species

Angela McDonnell, Mark Fishbein, Meg Quinn, Trevor Hare, and Kevin Keith 187

The first naturalized occurrence of the genus Forsythia (01eaceae)in Arkansas (U.S.A.),

with additional noteworthy angiosperm records for the state

Brett E. Serviss, Tyler L. Childs, Sydney S. Grant, Tiffany A. Graves, Ethan Holicer,

Seth A. McBroom, Logan Thomas, James H. Peck, and Allen Leible 195

Nephrolepis cordifolia (Nephrolepidaceae) naturalized in southern California (U.S.A.):

with notes on unintended consequences of escaped garden plants

Richard E. Riefner, Jr. and Alan R. Smith 201

Buddleja davidii (Scrophulariaceae) naturalized populations in Tennessee (U.S.A.) and

its woody associates

Ralph L. Thompson and Katrina Rivers Thompson 213

Addendum to the vascular flora of the Hancock Biological Station, Murray State University,

Calloway County, Kentucky, U.S.A.

J. Richard Abbott and Ralph L. Thompson 229

Floristics and community ecology of aquatic vegetation occurring in seven large springs at

Ozark National Scenic Riverways, Missouri (U.S.A.), 2007-2012

David E. Bowles and Hope R. Dodd 235

Cyperus granitophilus (Cyperaceae), a granite outcrop endemic, new for Texas and

Oklahoma (U.S.A.)

Robert J. O’Kennon and Kimberly Norton Taylor 251

Phytogeographical relationships and analysis of the flora of South-Central Texas, U.S.A.

A.A. Saghatelyan 259

Book Reviews, Notices, and Announcements 6,42,48,62, 74,80,120,136,148,166,186,200,

228, 234, 250, 258

INDEX to new names and new combinations in J. Bot. Res. Inst Texas 9(1), 2015

Coryphantha sneedii var. orcuttii (Boed.) Gorelick, comb, et stat. nov.—28

Dasylarynx Poinar & K.L. Chambers, gen. nov.—122

Dasylarynx anomalus Poinar & K.L. Chambers, sp. nov.—122

Dyschoriste ecuadoriana Wassh., sp. nov.—83

Escobaria sneedii var. orcuttii (Boed.) Gorelick, comb, et stat. nov.—28

Eumachia abrupta (Hiern) Delprete & J.H. Kirkbr., comb. nov.—76

Eumachia acuifolia (C. Wright) Delprete & J.H. Kirkbr., comb. nov.—76

Eumachia albert-smithii (Standi.) Delprete & J.H. Kirkbr., comb. nov.—76

Eumachia astrellantha (Wernham) Delprete &J.H. Kirkbr., comb. nov.—76

Eumachia boliviana (Standi.) Delprete & J.H. Kirkbr., comb. nov.—76

Eumachia cephalantha (Mull. Arg.) Delprete & J.H. Kirkbr., comb. nov.—77

Eumachia deinocalyx (Sandwith) Delprete & J.H. Kirkbr., comb. nov.—77

Eumachia guianensis (Bremek.) Delprete & J.H. Kirkbr., comb. nov.—77

Eumachia kappleri (Miq.) Delprete & J.H. Kirkbr., comb. nov.—77

Eumachia membranacea (Gillespie) Delprete & J.H. Kirkbr., comb. nov.—76

Eumachia microdon (DC.) Delprete & J.H. Kirkbr., comb. nov.—77

Eumachia nana (K. Krause) Delprete & J.H. Kirkbr., comb. nov.—78

Eumachia pallidinervia (Steyerm.) Delprete & J.H. Kirkbr., comb. nov.—78

Eumachia paupertina (Standi. & Steyerm.) Delprete & J.H. Kirkbr., comb. nov.—78

Eumachia podocephala (Mull. Arg.) Delprete & J.H. Kirkbr., comb. nov.—78

Eumachia wilhelminensis (Steyerm.) Delprete & J.H. Kirkbr., comb. nov.—79

Freziera humiriifolia D. Santam., sp. nov.—90

Freziera neillii D. Santam., sp. nov.—92

Freziera tundaymensis D. Santam., sp. nov.—95

Freziera yanachagensis D. Santam., sp. nov.—96

Gnaphaliothamnus nesomii M.O. Dillon & Luebert, sp. nov.—64

Hexasepalum angustatum (Steyerm.) J.H. Kirkbr. & Delprete, comb. nov.—104

Hexasepalum apiculatum (Willd.) Delprete & J.H. Kirkbr., comb. nov.—104

Hexasepalum gardneri (K. Schum.) J.H. Kirkbr. & Delprete, comb. nov.—104

Hexasepalum lippioides (Griseb.) J.H. Kirkbr. & Delprete, comb. nov.—104

Hexasepalum mello-barretoi (Standi.) J.H. Kirkbr. & Delprete, comb. nov.—104

Hexasepalum radulum (Willd.) Delprete & J.H. Kirkbr., comb. nov.—105

Hexasepalum rosmarinifolium (Pohl ex DC.) Delprete & J.H. Kirkbr., comb. nov.—105

Hexasepalum sarmentosum (Sw.) Delprete & J.H. Kirkbr., comb. nov.—105

Hexasepalum scandens (Sw.) J.H. Kirkbr. & Delprete, comb. nov.—105

Hexasepalum serrulatum (P. Beauv.) J.H. Kirkbr. & Delprete, comb. nov.—105

Machaerium excavatumJ. Linares, sp. nov.—51

MachaeriumfranksullyvaniiJ. Linares, sp. nov.—58

Machaerium paucifoliolatumJ. Linares, sp. nov.—54

Machaerium ramosiaej. Linares, sp. nov.—49

Machaerium rubrinervumj. Linares, sp. nov.—56

Mendoncia hollenbergiae Wassh., sp. nov.—81

Mendoncia sericea Wassh., sp. nov.—83

Micranthes rufopilosa (Hulten) D.F. Murray & Elven, comb. nov.—7

Prioria dominicana Poinar & K.L. Chambers, sp. nov.—130

Pseudognaphalium paramorum (S.F. Blake) M.O. Dillon, comb. nov.—72

Pseudognaphalium stolonatum (S.F. Blake) M.O. Dillon, comb. nov.—69

Scleria bellii LeBlond, sp. nov.—31

Stenostephanus holm-nielsenii Wassh., sp. nov.—86

Styrax peltatus P.W. Fritsch, sp. nov.—43

Typifications inJ. Bot. Res . Inst Texas 9(1), 2015

Eumachia guianensis (Bremek.) Delprete & J.H. Kirkbr.—77

Eumachia nana (K. Krause) Delprete & J.H. Kirkbr.—78

Eumachia podocephala (Mull.Arg.) Delprete & J.H. Kirkbr.—78

Hexasepalum gardneri (K. Schum.)J.H. Kirkbr. & Delprete —104

Hexasepalum lippioides (Griseb.) J.H. Kirkbr. & Delprete —104

Mapouria capituliflora Mull. Arg.—76

Phaseolus lignosus Britton —115

Psychotria pinularis Sesse & Moc.—77

Sedum rubroglaucum Praeger —4

TYPIFICATION OF SEDUM RUBROGLAUCUM (CRASSULACEAE)

Peter F. Zika

WTU Herbarium, Box 355325

University of Washington

Seattle, Washington 98195-5325, USA.

zikap@comcast.net

ABSTRACT

Sedum rubroglaucum was validly published in 1919, but the type specimen at DBN was not found. Sedum rubroglaucum as described, and

later illustrated, can not be readily distinguished from a number of closely related taxa in Sedum sect. Gormania in the absence of a type

specimen. A neotype is designated, based on a 1940 collection at YM from the type locality, in Yosemite National Park, California. Sedum

rubroglaucum becomes a synonym of the typical expression of the widespread California species S. obtusatum, described in 1868.

RESUMEN

Sedum rubroglaucum se publico validamente en 1919, pero el especimen tipo de DBN no se encontro. Sedum rubroglaucum tal como se de¬

scribe, y posteriormente se ilustro, no puede distinguirse facilmente de un numero de taxa relacionados de Sedum sect. Gormania en ausen-

cia de un specimen tipo. Se designa un neotipo, basado en una coleccion de 1940 en YM de la localidad tipo, en Yosemite National Park,

California. Sedum rubroglaucum se hace sinonimo la especie tipica y extendida por California S. obtusatum, descrita en 1868.

INTRODUCTION

Sedum L. sect. Gormania (Britton) R.T. Clausen is a small group of perennial species in California and Oregon,

with persistent rosettes on the flowering shoots, and corollas that are fused a short distance above the base.

One member of sect. Gormania is S. rubroglaucum Praeger. The name was validly published (Praeger 1919) but

lacks a type, as Praeger’s original material is missing. According to Stafleu and Cowan (1983), Praeger’s types

and the majority of his herbarium specimens were deposited in Dublin, Ireland, at DBN, with additional mate¬

rial at A, BEL, BM, CGE, E, GH, ILL, K, LIV, and NOT (herbarium acronyms in Thiers, continuously updated).

A search of those institutions failed to yield any collections labeled Sedum rubroglaucum.

Praeger (1919) cited a single gathering, which must be interpreted as the type. Sedum rubroglaucum was

described from living material sent to Praeger from California, on the Short Trail [now known as the Four Mile

Trail] in Yosemite National Park, California. Praeger (1919) said: “The plant was collected and forwarded alive

by Prof. H.M. Hall in June, 1915, and flowered in the following year.” There is no surviving specimen to serve

as holotype, as no original material from Praeger or Hall is extant. Jepson (1936:110) cited a specimen by Hall

as the type of S. rubroglaucum, but there is no indication that he saw a herbarium sheet. It is more likely Jepson

was citing the solitary collection mentioned in Praeger’s 1919 protologue. Hall was based in California. A

physical search in California collections was unsuccessful; there were no 1915 Sedum sect. Gormania vouchers

prepared by H.M. Hall at CAS, DS, GH, JEPS, POM, RSA, or UC. A wider search in the database of the Consor¬

tium of California Herbaria (2015) yielded 331 Hall specimens gathered in 1915 in California, none were in the

genus Sedum. Clausen (1975) was also unable to locate specimens that Praeger might have had at hand when

describing S. rubroglaucum. Hall’s 1915 shipment to Praeger did not include flowering material, diminishing

the chances he would have retained a pressed duplicate in a California herbarium. Praeger (1919) noted the

living specimen bloomed in September 1916 in a garden, and a flowering shoot was eventually illustrated for

Praeger’s (1921:219, fig. 125) catalogue of cultivated Sedum. Is the 1921 illustration (see Fig. 1) appropriate as a

lectotype for the name S. rubroglaucum, in the absence of a holotype specimen? Apparently not, for the artist

may have worked after 1919. If the graphic was published in 1919, it would qualify as part of Praeger’s original

materials at the time of publication. But Praeger used the artwork in 1921, weakening the argument it was

J.Bot. Res. Inst. Texas 9(1): 1-5.2015

Journal of the Botanical Research Institute of Texas 9(1)

Fig. 125.— S. rubroglaucum Praeger.

Fig. '[.Sedum rubroglaucum, illustrated by Praeger (1921:219, fig. 125) two years after publication of the binomial. Reproduced with permission of The

Garden (formerly The Journal of the Royal Horticultural Society).

original material, suitable for lectotype designation, in the sense of Articles 9.2 and 9.3 of the Melbourne Code

(McNeill etal. 2012).

The protologue of Sedum rubroglaucum mentions the petals are fused at the base, and Praeger (1919,1921)

placed it appropriately in Sedum sect. Gormania. The only illustration (Fig. 1) also shows a typical member of

the section with rosettes on the flowering and sterile shoots. However, neither the 1921 illustration nor the

1919 description are sufficient to distinguish S. rubroglaucum from several other closely related California taxa

more recently described in sect. Gormania, such S. albomarginatum R.T. Clausen, S. oblanceolatum R.T. Clau¬

sen, and the four subspecies of S. obtusatum A. Gray treated in Boyd and Denton (2012).

Clausen (1975:371) deduced that Sedum rubroglaucum must refer to nomenclaturally typical material of S.

obtusatum, based on geography, and that was a reasonable assumption. According to Taylor (2010), only two

perennial rosette-forming Sedum species are present in Yosemite National Park, S. obtusatum and S. spathulifo-

lium Hook. The latter, including its synonym S. yosemitense Britton, has petals separated to the base, not fused,

and so differs from the 1919 description of S. rubroglaucum. Therefore, based on the locality and the proto¬

logue, there seems little doubt that the material sent by Hall to Praeger was the typical subspecies of S. obtusa¬

tum, following current classifications of the genus (Clausen 1975; Denton 1982, 1993; Ohba 2009; Boyd &

Denton 2012; Wilson et al. 2014; Zika 2014).

The 1919 description of Sedum rubroglaucum is ambiguous, as is the 1921 illustration, and neither are sup¬

ported by an extant specimen. Thus a neotype is designated below, in accordance with Articles 9.7 and 9.13 of

the Code. The neotype specimen (Fig. 2) is housed at the herbarium of Yosemite National Park, and was col¬

lected by Reid Moran in 1940 from the same trail where Hall made his collection in 1915. Moran stated on his

specimen label: “... ‘topotype’ of Sedum rubroglaucum .” This choice of neotype is consistent with both Praeger’s

concept and locality, and fixes S. rubroglaucum as a synonym of the typical expression of S. obtusatum, a name

with priority, published by Asa Gray (1868).

Zika, Typification of Sedum rubroglaucum

YOSEMITE NAT'L PARE

HERBA-R1UM

1415

YOSEMITE NATIONAL PARK

HERBARIUM

No. /9W

HERBARIUM OF THE YOSEMITE MUSEUM

PLANTS OF YOSEMITE NATIONAL PARK

Scientific Name SedUOi ( Gomania) obtuaatuffl Gray-

Common Name

Locality 4 mile A to Glacier Point elc.7000'

Habitat Cracke in granite

Topotype of Sedum rubroglaucum Praeger 2

yellow tlui0h glaucous, becoming reddish; flowers

Collector Reid Moran No. 539 Date T/U/'hO

Fig. 2. Sedum rubroglaucum neotype, Moran 539 (YM). Scale bar 10 cm. Reproduced with permission of the Yosemite Museum Herbarium, U.S. National

Park Service.

Journal of the Botanical Research Institute of Texas 9(1)

TAXONOMY

Sedum rubroglaucum Praeger, J. Bot. 57:51. 1919. (Fig. 2). Type: U.S.A. CAUFORNIA [Mariposa Co.]: Yosemite National

Park, Four Mile Trail [formerly known as Short Trail] to Glacier Point, cracks in granite, elev. ca. 7000 ft [2134 m], 4 Jul 1940, R.V.

Moran 539 (neotype, designated here: YM!, catalogue number YOSE65304, ex YM1415, ex YM1955; isoneotype: UCSB71676!).

ACKNOWTEDGMENTS

I am grateful to Howard Fox, Elizabeth Brennan and Aidan Diskin for their assistance in searching for Robert

Lloyd Praeger and Harvey Monroe Hall materials at DBN, including the living collections of the associated

garden. I extend my appreciation to the keepers and staff of all the institutions cited, for their time and effort

searching for Praeger’s original material. Alison Colwell examined the holdings of the herbarium of Yosemite

National Park, and set aside relevant gatherings, while the Curator, Miriam Watson, granted permission to re¬

produce an image of the neotype. Dean Taylor provided valuable collecting notes from Hall’s visit to Yosemite

in May 1915. Chris Young, editor of The Garden (formerly The Journal of the Royal Horticultural Society),

kindly allowed the reproduction of Praeger’s 1921 illustration. I thank both John Wiersema and Kanchi

Gandhi for their insights and suggested revisions. Julie Nelson of the Shasta-Trinity National Forest granted

partial support for Sedum sect. Gormania research, done in conjunction with Barbara Wilson, Richard Brain-

erd, and Nick Otting of the Carex Working Group, and I am indebted to all of them for their lively interest in

the project.

REFERENCES

Boyd, S. & M.F. Denton. 2012. Sedum. In: B.G. Baldwin, D.H. Goldman, D.J. Keil, R. Patterson, TJ. Rosatti, & D.H. Wilkin, eds.

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

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

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

U.S.A.

Consortium of California Herbaria. 2015. [Data provided by the participants of the Consortium of California Herbaria.]

http://ucjeps.berkeley.edu/consortium/

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

ble/2806401.

Denton, M.F. 1993. Sedum. In: J.C. Hickman, ed. The Jepson manual: Higher plants of California. University of California

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

Gray, A. 1868. Characters of new plants of California and elsewhere, principally of those collected by H. N. Bolander in

the State Geological Survey. Proc. Amer. Acad. Arts 7:327-401.

Jepson, W.L. 1936. A flora of California, volume 6. Capparidaceae to Cornaceae. California School Book Depository, San

Francisco, California, U.S.A.

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

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

and plants (Melbourne code). RegnumVeg. 154:1-240. www.iapt-taxon.org/nomen/main.php

Ohba, H. 2009. Sedum. In: Flora of North America Editorial Committee, eds. Flora of North America north of Mexico,

Magnoliophyta: Paeoniaceae to Ericaceae. Oxford University Press, New York, U.S.A. 8:199-222. http://www.efloras.

org/florataxon.aspx?flora _id=1 &taxon _id=129989

Praeger, R.L. 1919. Notes on Sedum —III. J. Bot. 57:49-58.

Praeger, R.L. 1921. An account of the genus Sedum as found in cultivation. J. Roy. Hort. Soc. 46:1-314.

Stafleu, F.A. & R.S. Cowan. 1983. Taxonomic literature. A selective guide to botanical publications and collections with

dates, commentaries and types. Volume IV: P-Sak, 2nd ed. Bohn, Scheltema & Holkema, Utrecht, Netherlands, http://

www.sil.si.edu/digitalcollections/tl-2/index.cfm

Taylor, D.W. 2010. Flora of the Yosemite sierra, being a transect flora of the central Sierra Nevada, including all of Tu¬

olumne, Mariposa and Madera counties, the Mono Basin, and adjacent areas of Mono County. Published by the

author, Aptos, California, U.S.A.

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

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

Zika, Typification of Sedum rubroglaucum

Wilson, B.L., R.E. Brainerd, & N. Otting. 2014. Sedum kiersteadiae (Crassulaceae), a newly described species from the Klam¬

ath region of California, U.S.A. J. Bot. Res. Inst. Texas 8:9-15.

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

159:111-121.

Journal of the Botanical Research Institute of Texas 9(1)

BOOK REVIEW

Johnnie L. Gentry, George P. Johnson, Brent T. Baker, C. Theo Witsell, & Jennifer D. Ogle, eds. 2013. Atlas of

the Vascular Plants of Arkansas. (ISBN-13: 978-0-615-67980-8, pbk). University of Arkansas, De¬

partment of Printing Services, University Services Building, 2801 S. University Ave., Little Rock, Ar¬

kansas 72204, U.S.A. (Orders: cavern.uark.edu/~arkflora/, herb@uark.edu, University of Arkansas

Herbarium, Attn: Jennifer Ogle, Biomass Research Center 141, University of Arkansas, Fayetteville,

Arkansas 72701, U.S.A., 1-479-575-4372). $43.90, 709 pp., 2892 maps, some color introduction maps,

8Vi" x 11".

The Atlas of the Vascular Plants of Arkansas is the complimentary work of the Checklist of the Vascular Plants of

Arkansas, two critical accomplishments of the Arkansas Vascular Flora Committee (AVFC) seeking definitive

documentation of the known native and naturalized vascular plant species in the state of Arkansas. The Atlas

goes one step further than the previously-published Checklist by providing known county-distribution maps

for every species presently known in the state. Each species reported in the atlas has one county distribution

map under which the family, genus, species, and common name are listed. The maps are well-sized and are

conveniently organized alphabetically by scientific name of the family, genus, species, subspecies, variety, and

a few putatively-accepted hybrids under four major sections: Pteridophytes (ferns and fern allies), Gymno-

sperms (conifers), Angiosperm Dicots, and Angiosperm Monocots. AVFC adopts many changes in taxonomic

alignments based on updated phylogenetic research and maintains that the project will update taxonomy as

that research continues. The atlas portion also assigns a numerical code for species that are introduced, en¬

demic to Arkansas, non-native invasive, and/or of special conservation concern in Arkansas.

The enormity of biogeographical information presented by this book is put into context by an excellent in¬

troduction, including information on history of documentation of the flora in the state, geology relevant to the

flora, and natural vegetation. While the atlas portion itself functions well as an updated geographic record of

the flora, the additional information in the introduction, atlas, and appendices converts the Atlas into a rele¬

vant, modern documentation of the state’s natural history that provides an important baseline upon which

future research will build.— Devin Rodgers, Research Associate, Botanical Research Institute of Texas, Fort Worth,

Texas, U.S.A.

J. Bot. Res. Inst. Texas 9(1): 6.2015

MICRANTHES RUFOPILOSA (SAXIFRAGACEAE) COMB. NOV.:

AN ALPINE SPECIES FROM ALASKA AND YUKON

David F. Murray

University of Alaska Museum of the North

907 Yukon Drive

Fairbanks, Alaska 99775-6960, USA.

dfmurray@alaska.edu

Reidar Elven

Natural History Museum, University of Oslo

P.O. Box 1172

Blindern, NO-0318, Oslo, NORWAY

reidar.elven@nhm.uio.no

ABSTRACT

Micranthes rufopilosa comb.nov. is published with comparisons to related taxa: M. gaspensis, M. nivalis, and M. tenuis.

RESUMEN

Se publica Micranthes rufopilosa comb. nov. con comparaciones con los taxa relacionados: M. gaspensis, M. nivalis, y M. tenuis.

INTRODUCTION

Micranthes rufopilosa (Hulten) D.F. Murray & Elven, comb.nov. Basionym: Saxifraga nivalis var. rufopilosa Hulten, Ark.

Bot. ser. 2, 7:69. 1967 (1968), validly published in Madrono 19:223.1968. Type: U.S.A. Alaska: Steese Highway, mountains W of

Mastodon Dome, Eric Hulten s.n., 25-26 Jul 1964 (holotype: S!).

Saxifraga rufopilosa (Hulten) A.E. Porsild, Natl. Mus. Canad., Publ. Bot. 4:41.1974 (1975).

Saxifraga leriophora sensu A.E. Porsild non S. Watson, Canad. Field-Naturalist 79:79-90.1965.

Saxifraga tenuis sensu A.E. Porsild & Cody pro parte, Vase. Pi. Continental NWT Canada, map 710, fide Cody, Fl. Yukon. 1996.

Micranthes rufopilosa must be discussed in the context of its largely sympatric close morphological relatives: M.

nivalis, M. tenuis, and also the geographically distant M. gaspensis. The assertion thatM. tenuis andM. nivalis are

distinguished only with great difficulty (Brouillet & Elvander 2009) is not our experience. The two species are

distinguished by several morphological discontinuities (e.g., Healy and Gillespie 2004).

Hulten was reluctant to recognize M. tenuis at a rank higher than variety. In his manual (1968a) Hulten

treated the entities tenuis and rufopilosa as two varieties of M. nivalis. He later cited (1971) the differences be¬

tween M. tenuis and M. nivalis but treated those differences as inconsistent, characterizing their occurrence as

“here and there”. As of 1973, he still felt that those treating the taxon tenuis at the rank of species did so on cy-

tological grounds and remarked that he had not seen good material of M. tenuis from Alaska, where many

specimens now document the species. Hulten was here not in agreement with other European authors who

had recognized the two taxa as distinct species for nearly a century. However, we note, as Healy and Gillespie

(2004) remarked, “typical” or “good” M. tenuis of Europe is not what is found in North America, at least in the

western parts where Hultsen had his held experience.

After reviewing M. tenuis and M. nivalis for the Canadian Arctic, Savile (1961) concluded that in the High

Arctic those two species are easily separated, but to the south those distinctions become less clear as plants

somewhat intermediate can be found. Thus Savile recognized a third taxon, this one from the “Alaska-Yukon

refugium” meaning the unglaciated lands there, implying that this taxon has a different distribution because it

had a different history from the two species in the glaciated Canadian Arctic. His third taxon is M. rufopilosa.

Healy and Gillespie (2004) demonstrated in excellent detail that M. nivalis and M. tenuis from arctic Can¬

ada differ in both morphology and cpDNA. They also pointed out that the few alpine plants from Alaska and

Yukon they had seen, ones they had treated as M. tenuis, departed from the arctic specimens of that species by

quantitative characters, and these plants appeared to be intermediate between M. nivalis and M. tenuis. They

too had come upon M. rufopilosa.

J. Bot. Res. Inst. Texas 9(1): 7 -10.2015

Journal of the Botanical Research Institute of Texas 9(1)

Porsild’s description of M. tenuis (1951) from along the Canol Road referred, we believe, to M. rufopilosa,

which at that time neither he nor anyone else in North America recognized. In that account he emphasized

the reddish pubescence on lower leaf surfaces and pink to purplish petals, two features that, in part, define M.

rufopilosa.

Then Porsild (1965), when determining material from central Alaska, Steese Highway near Eagle Summit,

referred those plants to Saxifraga leriophora S. Watson.

Hulten (1967, 1968b) published Saxifraga nivalis var. rufopilosa, based on his collections from near Mast¬

odon Dome, not far from Eagle Summit, noting the more or less “ dense rufo-pilosa ” of the under surface of the

leaves and also its purple petals. He also wrote that his var. rufopilosa was the same as Porsild’s ?eriophora.

In 1974 Porsild compared var. rufopilosa, M. tenuis, and M. nivalis, referring to the specimens at CAN and

those recently collected by his brother Robert along the Dempster Highway in the Yukon. He concluded that

var. rufopilosa is more similar to M tenuis than to M. nivalis, but different from both at the rank of species, thus

his combination Saxifraga rufopilosa (Hulten) A.E. Porsild.

Obviously, the inclusive taxonomy in which Scoggan (1978) subsumes M. tenuis and M. gaspensis within

M. nivalis, is not followed by us.

HYBRID ORIGIN?

Morphological intermediacy is a typical feature of hybrids, and M. rufopilosa has been reported to be in some

respects intermediate to M. tenuis and M. nivalis. The assumption of morphological intermediacy needs to be

critically evaluated. After examining a number of specimens of all three species, we do not share it. Are there

other reasons to assume hybrid origin? Micranthes nivalis and M. tenuis are well known to have chromosome

numbers of 2n = 60 and 2n = 20 respectively, based on more than 20 independent countings.

Citing Krause and Beamish (1973), Gervais et al. (1995), and Devyatov et al. (1997), Healy and Gillespie

(2004) pointed to the occurrences of 2n = 40 plants on Wrangel Island in arctic Russia and in southwestern

subarctic Yukon, where both M. tenuis (2n = 20) and M. nivalis (2n = 60) are found, and on the Gaspe as M.

gaspensis (2n = 40) where neither putative parent is extant.

Brouillet and Elvander (2009) in their discussion of Micranthes gaspensis (Fernald) Small agreed with

Gervais et al. (1995) that this species represents a stabilized FI hybrid between M. nivalis and M. tenuis. They

went on to infer that a count of 2n = 40 from southwest Yukon published by Krause and Beamish (1973) was

also the product of hybridization between M. tenuis and M. nivalis. The count was given as “S. nivalis-tenuis

complex” by Krause and Beamish (1973), which we cannot check as the voucher specimen, despite a careful

search by Bruce Bennett (pers. comm.) and by us, has not been found. As for M. rufopilosa there is no evidence

of meiotic abnormalities, often a consequence of hybridization, inasmuch as plants readily form normal an¬

thers (pollen stainability not determined), fruits, and seeds.

Whereas hybridization between M. nivalis and M. tenuis may be the correct explanation for the occur¬

rence of M. gaspensis, further work is required to provide evidence. It is pure speculation to apply this possible

origin also to the Alaska-Yukon M. rufopilosa. That there exists a chromosome count of 2n = 40 in southwestern

Yukon does not mean that M. rufopilosa has that number and, by extension, an origin through a M. nivalis x M.

tenuis cross. Moreover, M. rufopilosa differs from M. gaspensis in important ways (see below). If the stabilized

hybrid explanation is correct, then crosses by the same parents in different areas have produced very different

results.

One must ask why, where both M. tenuis and M. nivalis are sympatric, sometimes growing in mixed

stands, such as throughout the Scandinavian mountains and in the Svalbard Archipelago of arctic Norway,

throughout northern Greenland and arctic Canada, and in most of arctic Russia, intermediates, fertile or ster¬

ile, have not been found and no counts of 2n = 40 have been made except for the one from Wrangel Island.

Hybridization, if that is the explanation for the existence of 2n = 40 cytotypes, must be a very rare event.

Murray and Elven, New combination in Micranthes

TAXONOMY AND DESCRIPTIONS

Micranthes rufopilosa (Hulten) D.F. Murray & Elven

Plants 2-12 cm tall; stems 1-3(5) from short rootstock, stems and inflorescence with mixed long, multicellular

white hairs and brown hairs; leaves with reddish-brown hairs along margins and petioles, typically densely

matted on lower leaf surface; inflorescence initially a compact terminal cluster of short-pedicellate flowers,

racemose to paniculate in fruit; flowers with pink to purple petals.

Micranthes nivalis (L.) Small, N. Amer. Fl. 22:136. 1905. Saxifraga nivalis L. Sp. Pl.:401.1753. Type: Europe Herb. Linn 517.19

far right hand specimen (lectotype, designated byjonsell & Jarvis 2002:72: LINN).

Plants 4-20 cm tall; stems 1-8 from a stout, simple or branched rootstock; stems and inflorescence with long,

multicellular, white to clear hairs, some purple-tipped; leaves glabrous or with reddish-brown hairs on mar¬

gins of blades and petioles, sometimes densely covering lower leaf surface; inflorescence initially one or more

compact terminal clusters of flowers,, racemose, in fruit; flowers with white petals turning pink with age.

Micranthes tenuis (Wahlenb.) Small. N. Am. Fl. 22:136. 1905.

Saxifraga tenuis (Wahlenb.) Harry Sm. ex Lindm.

Saxifraga nivalis var. tenuis Wahlenb. Fl. Lapp.:114. 1812 Type: Sweden. Kajsats, 30 Jul 1807, leg. G. Wahlenberg (lectotype, designated

by Moberg & Nilsson 1991:294: UPS!)

Plants 1-10 cm tall; stems 1-6(10) from a stout rootstock, stems glabrate below with short, multicelular, crisp

white hairs above, sometimes densely so, mixed with short glandular hairs; cell walls violet or purple; leaves

glabrous or with reddish-brown hairs along margins of blades and petioles, sometimes densely covering lower

leaf surface; inflorescence a compact terminal cluster in flower becoming corymbose in fruit; flowers with

pink to purple petals, exceptionally white.

Micranthes gaspensis (Fernald) Small. N. Amer. Fl. 22:552. 1918.

Saxifragagaspensis Fernald. Rhodora 19:141-142.1917. Type: CANADA. Quebec: Gaspe, Tabletopped Mt., Alt. 1000-1100 m, 8 & 12 Aug

1906, Fernald & Collins 600 (holotype: GH digital image!; isotype: CAN!).

Saxifraga nivalis var. gaspensis (Fernald) B. Boivin

Plants 3.5-10 cm tall; stems 1-2(5) from a stout rootstock, stems and inflorescence pubescent with short white

hairs, glabrate below gradually more densely pubescent above, some hairs glandular-tipped; leaves glabrous or

margins ciliate, with tangled reddish-brown hairs on lower leaf surface; inflorescence a compact terminal

cluster of flowers becoming racemose in fruit; flowers with white petals.

Porsild (1974) provided photos (plates 8 and 9) and a brief, useful review of diagnostic features that dis¬

tinguish M. rufopilosa from M. tenuis and M. nivalis. Micranthes rufopilosa takes its name from the dense layer of

reddish-brown hairs on the lower surface of some leaves; however, the leaves are not consistently so. Moreover,

Micranthes tenuis, M. nivalis, and M. gaspensis also have reddish-brown hairs on the petiole and leaf margins

and in some instances also on the lower surface of the leaves, although only occasionally as dense as in M. ru¬

fopilosa.

On stems and branches of the inflorescence M. rufopilosa has mixed long, white and reddish-brown hairs,

similar in length to the long white, crisp hairs of M. nivalis. Micranthes nivalis has only long white hairs, M.

gaspensis has short white hairs, and M. tenuis is typically sparsely short-hairy, more dense distally, with a mix¬

ture of short, glandular hairs.

Specimens of Micranthes rufopilosa we have seen are all from the Yukon-Tanana Upland of central Alaska

and from central and southern Yukon. Cody (1996) has mapped arctic localities well disjunct of its main range

in Yukon; these arctic occurrences have been determined by us as M. tenuis.

Journal of the Botanical Research Institute of Texas 9(1)

KEY TO SPECIES

1. Petals white or margins pink with age; stems glabrous or with only white hairs, hypanthium triangular to hemispheric.

2. Stems and inflorescence with short crisp and long tangled white hairs_M. nivalis

2. Stems and inflorescence glabrate or with short white hairs_M. gaspensis

1. Petals reddish to purple; stems pubescent not as above, hypanthium turbinate.

3. Stems with long tangled white and brown hairs; inflorescence racemose or paniclate in fruit_M. rufopilosa

3. Stems glabrate or with short white and short, glandular hairs; inflorescence corymbose in fruit_M. tenuis

Representative specimens (M icranthes rufopilosa ).—U.S.A. ALASKA. Charley River Quad.: Mt. Sorenson, 65°00'N, 142°45'W, 19 Jul 2002,

Cook & Roland 02-443 (ALA). Circle Quad.: Steese Hwy. Eagle Summit, 26 Jul 1964, Viereck 7397 (ALA) [collected at the same time and place

as the type was gathered by Eric Hulten]. Nabesna Quad.: Chisana R., 62°16'N, 141°55'W, 20 Jun 2003, Roland & Batten 5817B (ALA). Liv-

engood Quad.: White Mts., Vic. VABM Fossil, 65°37'N, 147°22'W, 13 Jun 1995, Parker & Herriges 5639 (ALA). CANADA. Yukon: Klaza Mt.

62°27'N, 137°51'W, 19 Jul 2012, Bennett & Cannings 12-0243 (ALA). Syenite Range, 63°95'N, 137°27'W, 26 Jun 2012, Bennett, Cannings, &

Schroeder 12-0064 (ALA); Mt. Klotz, 65°37'N, 140°17'W, 24 Jun 2007, Bennett, Line, Kennedy, & Mennell 07-118 (ALA); Ptarmigan Heart,

61°49'N, 138°35'W, 16 Jul 1948, Raup, Drury, & Raup, 13749 (CAN); Ogilvie Mts., Dempster Hwy. Mile 51, 7 Jul 1968, Porsild 1574 (CAN)

[basis for photo in A.E. Porsild 1974.]; Mt. Nansen, 62°06'N, 137°17'W, 16-17Jun 1968, Porsild 1372 (CAN); Mt. Archibald, Mile 1026 Alaska

Hwy., 23 Jul 1967, Pearson 67-297A (CAN); Dawson Range, 62°35'N, 138°20'W, 27 Jun 1941, Bostock256 (CAN).

ACKNOWLEDGMENTS

We thank the curators and collections managers at ALA, CAN, and DAO for loans and assistance during visits

to their herbaria. Thanks also to R.J. Gornall and Peter Lesica for their careful readings of the manuscript and

their comments and suggestions, advice we have considered in every instance.

REFERENCES

Brouillet, L. & P.E. Elvander. 2009. Micranthes. In: Flora of North America Editorial Committee, eds. Flora of North America

north of Mexico. Oxford Univ. Press, New York, U.S.A. 8:49-70.

Cody, WJ. 1996 Flora of the Yukon. NRC Press, Ottawa, Canada.

Devyatov, A.G., P.Y. Zhmylev, & A.D. Kozhevnikova. 1997. Chromosome numbers of some arctic species of the genus Saxi-

fraga (Saxifragaceae). Bot. Zhur. 82:122.

Gervais, C., N. Dignard, & R. Trahan. 1995. The chromosome number of Saxifraga gaspensis Fernald. Rhodora 97:171-175.

Healy, C. & LJ. Gillespie 2004. A systematic analysis of the alpine saxifrage complex (Saxifragaceae) in the Canadian Arctic

Islands using morphology and chloroplast DNA data. Canad. Field Naturalist 118:326-340.

Hulten, E. 1945. Flora of Alaska and Yukon. Vol. 5. Acta Univ. Lund. Avd. 2, 56(1):799-978.

Hulten, E. 1967. Comments on the flora of Alaska and Yukon. Ark. Bot. ser.2, 7(1 ):1—147.

Hulten, E. 1968a. Flora of Alaska and neighboring territories. Stanford University Press, Stanford, California, U.S.A.

Hulten, E. 1968b Validity of nomenclature changes undertaken in the flora of Alaska and Yukon. Madrono 19(6):223.

Hulten, E. 1971. Circumpolar plants II. Dicotyledons. Kongl. Svenska Vetenskapsakad. Handl. 13:1-463.

Hulten, E. 1973. Supplement to flora of Alaska and neighboring territories. Bot. Not. 126:459-512.

Jonsell, B. & C.E. Jarvis. 2002. Lectotypification of Linnaean names for Flora Nordica Vol. 1 (Lycopodiaceae-Papavera-

ceae). Nord. J. Bot. 14:145-164.

Krause, D.L. & K.I. Beamish. 1973. Notes on Saxifraga occidentalis and closely related species in British Columbia. Syesis

6:105-113.

Moberg, R. & 0. Nilsson. 1991. Typification of Nordic vascular plants. 1. Names published by G. Wahlenberg. Nord. J. Bot.

11:287-299.

Porsild, A.E. 1951. Botany of southeastern Yukon adjacent to the Canol Road. Natl. Mus. Canada Bull. No 121. Ottawa.

Porsild, A.E. 1965. Some new or critical vascular plants of Alaska and Yukon. Canad. Field-Naturalist 79:79-90.

Porsild, A.E. 1974 (1975). Materials for a flora of central Yukon Territory. Natl. Mus. Canad. Publ. Bot. No. 4. Ottawa.

Savile, D.B.0.1961. The botany of the northwestern Queen Elizabeth Islands. Canad. J. Bot. 39:909-942.

Scoggan, H. 1978. The flora of Canada, Part 3. Dicotyledonae (Saururaceae to Violaceae). Natl. Mus. Nat. Sci., Publ. Bot.

7(3):547-1115.

nrDNA SEQUENCE (ITS AND ETS) FROM HERBARIUM SPECIMENS REVEALS

PHYLOGENETIC AFFINITIES OF ERIGERON GEISERI (ASTERACEAE)

Richard D. Noyes, William Caraway, and Dulcinea V. Groff

Department of Biology

University of Central Arkansas

Conway, Arkansas 72035, US.A.

rnoyes@uca.edu

ABSTRACT

Erigeron geiseri Shinners is a tap-rooted herbaceous daisy occurring from south-eastern Texas to southern Oklahoma. Our study of pollen

quality from 67 herbarium specimens supports the hypothesis that the species comprises sexual diploid populations, restricted to southern

locations, and polyploid, putatively apomictic, populations that are widespread. We successfully amplified and sequenced ITS and ETS

spacer regions of nrDNA for five specimens (two putative sexual diploids, three putative apomictic polyploids). Phylogenetic analysis con¬

ducted using maximum likelihood and Bayesian methods support a topology showing that 1) sexual and apomictic populations of E. geiseri

form a well-supported monophyletic group, and, 2) E. geiseri is sister to E. strigosus of eastern North America. The results support the clas¬

sification of E. geiseri with E. strigosus in E. sect. Phalacroloma.

RESUMEN

Erigeron geiseri Shinners es una margarita herbacea que se da desde el sureste de Texas al sur de Oklahoma. Nuestro estudio de la calidad del

polen en 67 espechnenes de herbario soporta la hipotesis de que especie comprende poblaciones sexuales diploides, restringidas a locali-

dades surenas, y poblaciones poliploides, putativamente apomicticas, que estan esparcidas. Amplificamos y secuenciamos con exito las re-

giones de los espaciadores ITS y ETS del nrDNA de cinco espechnenes (dos diploides sexuales putativos, tres poliploides apomicticos puta-

tivos). Los analisis filogeneticos realizados usando maxima verosimilitud y metodos Bayesianos soportan una topologia que muestra 1) las

poblaciones sexuales y apomicticas de E. geiseri forman un grupo monofiletico bien soportado, y, 2) E. geiseri is es hermano de E. strigosus

del este de Norte America. Los resultados soportan la clasificacion de E. geiseri con E. strigosus en E. sect. Phalacroloma.

INTRODUCTION

Erigeron geiseri Shinners (Asteraceae), Geiser’s fleabane or Basin fleabane, was originally described (Shinners

1947) from collections within a few-county radius around Austin in central Texas (Burnet, Bastrop, Falls, Gua¬

dalupe, Llano, and Robertson counties). Subsequent study documented the range of the species from the

south-eastern coast of Texas north to southern Oklahoma (Nesom 1979). Although the global conservation

status of G4 for E. geiseri indicates it is apparently secure (NatureServe 2014), most of its relevant biological or

ecological attributes have not been assessed. Further, the taxon is ranked S3 for Texas (vulnerable), and either

not ranked or under review for Oklahoma. The species (Fig. 1) is a tap-rooted annual to 40 cm with narrow

oblanceolate leaves, the basal ones often exhibiting 2-4 pair of conspicuous lobes. The inflorescence is a diffuse

corymbiform array with 1-20 capitula, with peduncles often slightly dilated distally. The capitula feature ob-

ovate phyllaries with scarious margins, and low-conic receptacles. Ray florets in the species are white, the rela¬

tively few disc flowers are yellow, and cypselae bear a biseriate pappus consisting of about 10 scabrous bristles

and an equal number of low scales (Nesom 2006). The species flowers from late March-May (-June) and is

distributed with broad ecological tolerance, occurring on sandy to clay soils associated with prairie, roadside,

and fencerow habitats. Upon publishing E. geiseri, Shinners also described the intraspecific taxon E. geiseri

var. calcicola Shinners for plants with longer stem pubescence and a pappus reduced to a low crown. These

plants have subsequently been synonymized with E. versicolor (Greenm.) Nesom (Nesom 2006).

While a chromosome number for E. geiseri has not been published, specimen annotations by Dr. B.L.

Turner, University of Texas, indicate 2n=18 (Turner #4526, TEX), the typical number for diploid Erigeron, as

well as triploids with 2n=27 (Turner #4527, TEX). The unpublished numbers intimate that E. geiseri, similar to

J. Bot. Res. Inst. Texas 9(1): 11 -24.2015

Journal of the Botanical Research Institute of Texas 9(1)

ASTERACEAE

Erigeron geiseri Shinners

TX: BurrojlCo ; Carr, W R. 11752 (TEX). l3Apr m2

pm VP1QID APOWICT; Pollen large (>l7ym dtam > and of poor quaWy

Estonaled UViong 30.62160‘N ■ -98 30222^

R D Noyes and 0 V Groff, December 2013

UNIVERSITY OF CENTRAL ARKANSAS (UCAC)

Fig. 1. Herbarium specimen of Erigeron geiseri. Carr 11782 (TEX) exhibits the tap-root, small stature, erect habit, diffuse inflorescence, and occasional leaf

lobing that characterize the species. The largest specimen in the center, including the root, is 20.3 cm tall. Inset shows the entire herbarium specimen.

congeners such as E. strigosus and E. tenuis, likely comprises diploid, presumably sexual populations, and trip-

loid populations that likely reproduce via apomixis (Noyes 2006; Groff 2010; Noyes & Givens 2013).

Differing hypotheses regarding the taxonomic affinities of Erigeron geiseri have been proposed. Shinners

(1947) allied E. geiseri with the Mexican taxon E. coronarius Greene, presently classified within E. sect. Ge-

niculactis (Nesom 2008). Nesom (1989) placed E. geiseri within a broadly defined E. section Olygotrichium,

calling attention to morphological similarities among E. geiseri, E. tenuis Torr. & A. Gray E. tenellus DC., and

E. turnerorum Nesom. Nesom (2008) later included E. geiseri within the segregate E. sect. Quercifolium, along

with 18 other species including E. tenellus and E. turnerorum. Morphological similarity between E. geiseri and

E. tenuis (Shinners 1947; Nesom 1979; Nesom 1989; Nesom 2008) adds an additional twist as E. tenuis has been

Noyes et al., Phylogenetic affinities of Erigeron geiseri

shown to be genetically similar to E. strigosus Muhl. ex Willd. and E. annuus (L.) Pers. of eastern North Amer¬

ica (Noyes 2000; Groff 2010) all presently classified as E. sect. Phalacroloma (Nesom 2008).

The objective of this study was to utilize herbarium specimens 1) to estimate, from study of pollen quality,

the distribution of putative sexual diploid and apomictic polyploid populations of Erigeron geiseri, and, 2) to

evaluate the phylogenetic affinities of E. geiseri using nuclear ribosomal DNA spacer sequence data (ITS and

ETS). Results provide data relevant to the conservation of E. geiseri and help to clarify our understanding of the

relationships among the Erigeron species of eastern North America.

METHODS

Mode of reproduction and geographic mapping

A total of 67 herbarium specimens of Erigeron geiseri were studied. The majority were obtained on loan from

the University of Texas (TEX & LL; 44 specimens) and from the Botanical Research Institute of Texas (BRIT;

18 specimens). Five specimens were also studied that were originally misidentihed and included among loans

for the study of other Erigeron species (Missouri Botanical Garden, MO, 4 specimens; Mississippi State Univer¬

sity, MISSA, 1 specimen).

Pollen from the 67 herbarium specimens was evaluated to estimate mode of reproduction (Noyes & Alli¬

son 2005; Noyes & Groff 2011). Samples from mature but unopened florets were stained in Cotton Blue in

lactophenol (Stanley & Linskens 1974) for a minimum of four days and evaluated at 400X on an Olympus BX51

compound microscope using bright held optics. A pollen sample was interpreted to represent the sexual dip¬

loid condition if the grains were uniformly small (12-15 pm diam.) with a high percentage darkly stained, pu¬

tatively viable, grains (Noyes & Allison 2005). The polyploid apomictic condition, by contrast, was indicated

by samples that included large grains (>17 pm diam.), high proportion of non-staining aborted grains, and

frequent non-staining pollen micrograins. Sample measurements of grains were performed using AnalySIS (v.

3.1; soft Imaging System, GmbH) on images captured with an Olympus FV12 monochrome CCD camera.

For distribution mapping, the geographic coordinates for each specimen were estimated from label local¬

ity information using TOPO USA (version 5.0, DeForme, Yarmouth, ME). The data were then plotted using

layer (tl_2009_US_county.shp) was obtained from the US Census Bureau, Geographic Products Branch

(http://www2.census.gov/cgi-bin/shapehles2009/national-hles). Records from duplicate specimens were

culled so that only unique collections were mapped. As a rough check of the accuracy of our plotted data, we

verihed that county names extracted from plotted data points matched input county names. Annotation labels

were affixed to all specimens indicating putative mode of reproduction and estimated geographic coordinates.

DNA isolation, amplification, and sequencing

To investigate the genetic relationships of F. geiseri, we selected eight relatively recently collected specimens

(four putative sexual diploid, four putative polyploid apomicts) with ample leaf material (Table 1). Total DNA

was isolated for each specimen using the DNeasy Plant Mini kit (#69106, QIAGEN Inc., Valencia, CA, USA)

using about 0.5 cm 2 dried leaf tissue. The prescribed protocol was followed except that to 500 pi warmed API

within a 1.6 ml Eppendorf tube we added a small quantity of sterilized sand to facilitate grinding, plus lOmM

sodium metabisulhte (#S244, Fisher Scientific, Pittsburgh, PA, USA) as a supplemental antioxidant (Horne et

al. 2004). In addition, we heated the grindate in API for 0.5 hour, and employed a 13K centrifugation for 5 min.

after incubation with AP2 prior to application to the shredder column to facilitate removal of protein precipi¬

tate and undigested leaf tissue. Samples were eluted into a total of 60 pi AE (rather than the recommended 200

pi) to foster higher yield concentrations. The resulting genomic DNA was quantified via fluorometry (Hoefer

DyNA Quant 200, Amersham Biosciences, Piscataway, NJ, USA) and evaluated for quality by agarose electro¬

phoresis. Samples exhibiting at least a portion of genomic DNA greater than lkb in size were selected for se¬

quence analysis.

We sequenced the 18S-26S nrDNA spacer regions ITS (ITS1,5.8S, ITS2) and ETS to facilitate comparisons

Journal of the Botanical Research Institute of Texas 9(1)

Table 1 . Herbarium specimens ofErigeron geiseri selected for DNA study.

Specimen

Designation

Isolated DNA Suitable

for Sequence Analysis?

GenBank Accession

Numbers

1. Fleetwood, R. J. 9049 (TEX);

Texas: Brazoria Co., 13 April 1967.

Putative sexual diploids

2. Fleetwood, R.J. 10811 (LL);

Texas: Galveston Co., 04 March 1974.

3. Lieb, C. 5. 337 (TEX);

Texas: Matagorda Co., 09 April 1983.

YES 1

ITS: KP318874

4. Nesom, G.L 5389 (TEX);

Texas: Gonzales Co., 27 March 1986.

YES

ETS: KP318879

ITS: KP318875

1. Carr, W. R. 11782 (TEX);

Texas: Burnet Co., 13 April 1992.

Putative apomictic polyploids

YES

ETS: KP318880

ITS: KP318876

2. Corr, W. R. 23324 (TEX);

Texas: Goliad Co., 27 May 2004.

YES

ETS: KP318881

ITS: KP318877

3. Smith, B. 77 (BRIT);

Texas: McCollough Co., 06 April 1966.

ETS: KP318882

4. Turner, B. L. 4527 (TEX);

Texas: Gonzales Co., 27 March 1959.

YES 1

ITS: KP318878

ETS: KP318883

Required internal amplification for ITS and ETS from initial weak PCR products.

with other Erigeron species (Noyes 2000; Noyes 2006). ITS and ETS were amplified and sequenced separately

using a modification of the PCR conditions of Urbatsch et al. (2003). The program we used (for both ITS and

ETS) consisted of a 3-min 95°C initial denaturation followed by 10 cycles of 30-sec denaturation at 95°C, 40-

sec annealing at 55°C, and 40-sec extension at 72°C with a 4-sec per cycle increase. The subsequent 20 cycles

proceeded with 30-sec denaturation at 95°C, 40-sec annealing at 50°C, and 40-sec extension at 72°C (with

4-sec per cycle increase), followed by a final extension of 7-min at 72°C. We employed relatively simple PCR

components including (in ~50 ul reactions) 41.3 pi PCR grade H 2 0, 5 pi lOx ThermoPol Reaction Buffer (New

England Biolabs, Beverly, MA, USA), 0.2 pmol/L each primer (1.0 pi of lOuM stock), 200 pmol/L each dNTP

(0.4 pi of mixture of 25 mmol/L each dNTP; #N0446S, New England Biolabs), 1.5 U (0.3 pi) ThermoPol Taq

polymerase (#M0267X, 5,000U/ml, New England Biolabs), and 0.5 to 2.0 pi of undiluted genomic DNA (rang¬

ing from 2 to 44 ng/pl).

We successfully amplified and sequenced ITS and ETS for five of the eight samples using a suite of internal

and external primers (Table 2; Fig. 2). The ITS region was amplified using primers ITSI(f) and ITS4(r) while

ETS was amplified using ETS-Ast8(f) and ETS-18S(r). For three samples that yielded strong amplifications for

both regions, PCR products were purified for sequencing using MinElute PCR purification columns (#28006,

QIAGEN Inc.). Products were eluted in a final volume of 18 pi, quantified by fluorometry, and sequenced using

separate forward and reverse reactions. For the ITS region, the primer ITS5 was used for the forward sequence

and ITS4 was used for the reverse sequence. For the ETS region, PCR primers (ETS-Ast8 and ETS-18S) were

used for sequencing.

Two samples yielded faint initial ITS and ETS PCR bands. For these we increased yields by internal

amplification using 1 pi of undiluted initial PCR product as a template using identical cycling conditions as

described above. For the ITS1 region we used primers ITSI(f) and ITS2(r) while for the ITS2 region we used

primers ITS3(f) and ITS4(r). For internal amplification of ETS we identified a centrally located conserved

Noyes et al., Phylogenetic affinities of Erigeron geiseri

Table 2. Primers used in this study to amplify ITS and ETS18S-26S nrDNA spacer regions.

Primer Designation

Primer Sequence

Reference

ETS

ETS-Ast8(f)

TTC TCT TCG TAT CGT GCG GT

Markos & Baldwin (2001)

ETS-18S(r)

ACT TAC ACA TGC ATG GGT TAA TCT

Baldwin & Markos (1998)

ETS-iR(r)

GAG TGT GTT GCA TGG TTC

This study

ETS-iF(f)

GAA CCA TGC AAC ACA CTC

This study

ITS

ITSI(f)

GTC-CAC-TGA-ACC-TTA-TCA-TTT-AG

Urbatsch et al. (2000)

ITS5(f)

GGA-AGG-AGA-AGT-CGT-AAC-AAG-G

White et al. (1990)

ITS4(r)

TCC-TCC-GCT-TAT-TGA-TAT-GC

White et al. (1990)

ITS2(r)

GCT-ACG-TTC-TTC-ATC-GAT-GC

White et al. (1990)

ITS3(f)

GCA-TCG-ATG-AAG-AAC-GTA-GC

White et al. (1990)

Fig. 2. Approximate locations of primers used to amplify the ITS and ETS 18S-26S nrDNA spacer regions in this study.

sequence among a diverse sample of Erigeron taxa (Noyes, 2006) and designed internal forward and reverse

primers (Table 2). Internal amplification of the 5’ETS region thereby employed primers ETS-Ast8(f) and

ETS-iR(r) and the 3’ETS region employed primers ETS-iF(f) and ETS-18s(r). PCR products were purified

and quantified as above. Sequencing of the ITS1 fragment employed, separately, ITS5 and ITS2 primers.

Sequencing of the other three products (ITS2, 5’ETS, 3’ETS) employed the primers used for internal PCR.

Samples for sequencing consisted of 150 ng purified PCR product plus 10 pmol (1 pi x 10 pM stock) se¬

quencing primer. Samples were dried using an Eppendorf Speedvac for 20 min at 65°C and shipped via First

Class USPS to the University of Missouri, Columbia, DNA Core Facility, which employs an Applied Biosystems

(Foster City, CA, USA) 3730x196-capillary DNA Analyzer and Big Dye Terminator cycle sequencing chemis¬

try. Forward and reverse chromatograms, downloaded by FTP, were merged into contigs and edited using Se-

quencher 4.2 (Gene Codes Corp., Ann Arbor, MI, USA). Care was taken to score bases as polymorphic when

there were obvious double peaks in both forward and reverse sequences.

Phylogenetic analyses

Analysis was first conducted using 81 ITS sequences to broadly evaluate the relationship of Erigeron geiseri

to other Erigeron species (Appendix 1). The data matrix included five sequences for F. geiseri, 75 published

ITS sequences for Astereae sect. Conyzinae (Erigeron and allies; Noyes 2000), plus the ITS sequence for F.

hyssopifolius Michx. (Noyes 2006). In addition, we substituted three sequences from the Noyes (2000) matrix

representing F. sect. Phalacroloma (F. strigosus (GenBank #AF118490), F. annuus (AF118489), and F. tenuis

(AF118488)), with those published for F. strigosus var. calcicola J. Allison (AY871318), F. strigosus var. dolomiti-

colaj. Allison (AY871323), and F. strigosus var. strigosus (AY871333) that study showed better represented

genetic diversity in the group (Noyes 2006). Erigeron neomexicanus A. Gray (AF118544) served as the

Outgroup.

Initial results indicated that further analysis with more comprehensive representation of Erigeron sect.

Phalacroloma was warranted. We therefore conducted a second analysis with concatenated ITS and ETS

Journal of the Botanical Research Institute of Texas 9(1)

sequences for a representative sample of 14 sequences for E. sect Phalacroloma, the five sequences obtained

here for E. geiseri, plus five additional Erigeron for which both ITS and ETS were available (Noyes 2006;

Appendix 2). The 14 sequences for E. sect. Phalacroloma include those for E. strigosus var. calcicola, E. strigosus

var. dolomiticola, plus 12 sequences representing the three divergent rDNA haplotypes (I, II, III) found for

sexual diploid E. strigosus var. strigosus (Noyes 2006). Sequences for E. acris L., E. hyssopifolius, E. pinnatisectus

(A. Gray) A. Nelson, and E. speciosus (Lindl.) DC. served as the Outgroup.

Sequences for analyses were compiled and aligned using ClustalW as implemented in MEGA6 (Tamura

et al. 2013). Genetic distances between subsets of sequences were explored in MEGA6 and phylogenetic analy¬

sis was conducted using Maximum Likelihood and Bayesian approaches. MrModelTest (2.3) (Nylander 2004)

was employed to select appropriate models of sequence evolution. This was accomplished by exporting respec¬

tive Nexus hies (ITS vs. ITS plus ETS matrices) from MEGA6, appending the MrModelTest command block,

and then implementing PAUP* (version 4.0b. 10; Swofford 2002). This yielded “.scores” hies that were em¬

ployed by the MrModelTest executable to produce a “.out” hie containing test scores and recommended

command lines to implement the optimal model of molecular evolution (based on the Akaike Information

Criterion (AIC)) for phylogenetic analysis.

Maximum likelihood analyses were executed in MEGA6 using the nearest-neighbor-interchange (NNI)

heuristic method with the initial tree generated automatically by neighbor joining. Analyses used the default

settings of “branch swap hlter = very strong”, and “number of threads = 1”. Gaps and missing data were subject

to complete deletion and analyses were run with 500 bootstrap replicates. Bayesian analyses were implement¬

ed with MrBayes 3 (Ronquist & Huelsenbeck 2003) by appending the prescribed commands from MrModelT¬

est to the respective Nexus hies. The program was executed using the Markov chain Monte Carlo (MCMC)

method with four chains, 3 million generations, print frequency of 1000, sample frequency of 100, and with

branch lengths saved. The number of initial trees discarded Churnin’) was determined by visually approximat¬

ing the point at which the average standard deviation of split frequencies appeared to stabilize. Bayesian prob¬

abilities for nodes were obtained from the resulting 50% compatibility consensus tree. Tree topologies sum¬

marizing analyses were those obtained from MEGA6 maximum likelihood analyses, to which maximum

likelihood and Bayesian probabilities were added. Topologies, as necessary, were edited with TreeView 1.6.6

(Page 1996).

RESULTS

All 67 specimens of Erigeron geiseri yielded pollen from which we were able to unequivocally classify mode of

reproduction. A total of 61 of the collections were unique; six duplicates were identihed. A total of 12 (19.7%)

of the unique collections yielded pollen consistent with the sexual diploid condition and 49 (80.3%) produced

pollen characteristic of apomicts (Fig. 3). All duplicates of a collection exhibited the same pollen characteristics

and therefore no duplicate sets provided evidence that populations included both sexual and apomictic plants.

The putative sexual diploid populations occur in two geographically adjacent but distinct areas (Fig. 4). The

first group (six collections) occurs along the Gulf Coast in Texas from Galveston Co. south to San Patricio Co.

The second group (also six collections) is approximately 200 km inland scattered across five counties in Texas

in the vicinity of Austin. Putative apomictic E. geiseri is sympatric with but more broadly distributed than

sexual populations with collections extending from Kleberg Co., along the southern Gulf Coast, north to

south-central Oklahoma, and west in Texas to Shackelford, Callahan, and McCulloch Cos. Reflecting the dif¬

ference in distribution, the mean latitude for apomicts (30.785°N) is significantly (one-tailed t-test, P = 0.0049)

more northerly than that for sexual diploids (29.253°N). The overall map is nearly identical to that of Nesom

(1978), but with the added reproductive data.

The map indicates the strong possibility of co-occurrence of the two reproductive modes because four of

the ten counties where sexual collections are recorded also exhibit apomictic collections. In addition, collec¬

tion dates, as a proxy for flowering time, document that putative sexual diploid and apomictic polyploid popu¬

lations flower at similar times (from mid-March to early June) intimating the possibility for gene flow between

Noyes et al., Phylogenetic affinities of Erigeron geiseri

Fig. 3. Pollen of herbarium specimens of Erigeron geiseri stained in cotton blue in lactophenol. A. Pollen for Fleetwood #9049 (TEX) shows small grains

(~15 pm diameter) with a high proportion staining darkly consistent with plants of the sexual diploid condition. B. Pollen for Tharp #7448 (TEX) shows a

mixture of staining grains that vary in size, plus a high proportion of grains that are unstained or lightly stained, consistent with the polyploid apomictic

condition. The largest staining grains are ~21 pm diameter and possibly triploid. Scale bar = 40 pm.

the two types. The overall average date of collection for apomictic plants is significantly later than for sexual

plants (30 April vs. 14 April; one-tailed t-test, P = 0.026), but this is likely a consequence of the more northerly

range of apomictic plants. When 29 apomictic plants are excluded that occur further north than the northern¬

most sexual collection, the mean flowering date for putative apomicts shifts to 25 April and the difference be¬

tween sexuals and apomicts is not significantly different (one-tailed t-test, P = 0.11).

We obtained ITS and ETS sequences for five of the eight assayed herbarium specimens, including two

putative sexual diploids (‘S3’ and ‘S4’) and three putative apomicts (AT, A2’, and A4’). Successful sequencing of

‘S3’ and A4’ required second amplification from weak initial PCR products using internal primers. GenBank

numbers are provided in Table 1. The sequenced specimens represent well the geographic distribution of E.

geiseri, with ‘S3’ and ‘S4’ representing coastal and interior sexual diploid populations, respectively, and AT,

A2’, and A4’ well separated by latitude across the range of apomicts. Extracts from ‘ST, ‘S2’, and A3’ did not

exhibit high molecular weight DNA and did not yield visible ITS or ETS PCR products.

The length of the ITS and ETS sequences were consistent with those for other Erigeron and allies within

sect. Conyzinae: 625-626 bp for ITS and 597 bp for ETS. The total number of polymorphisms per concatenated

sequence ranged from 3 for ‘S3’ to 8 for A2’, with an overall average of 5. Of these, 30% occurred at potentially

informative sites, while 70% were unique and therefore not informative. Three of the sequences exhibited indel

polymorphisms, which meant that complete sequence could be obtained only by reading a partial sequence

from the forward read (to the polymorphic site) and the remaining sequence from the reverse read (to the poly¬

morphic site). Single indel polymorphisms were inferred for A4’ in ETS, and for ‘S3’ in ITS2. Two indels were

detected in ETS for ‘S3’, one in the 5’ and one in the 3’ regions; these were resolvable because the region was

sequenced in two separate fragments. The nucleotide and indel polymorphisms did not preclude acquisition of

complete and robust nrDNA sequence. Overall the five sequences are very similar, differing by an average of only

0.08%, which corresponds to approximately a single base position (0.98) across the entire ITS plus ETS region.

The analysis of ITS for 76 members of Erigeron sect. Phalacroloma plus five Erigeron geiseri employed a

general time reversible (GTR) model of molecular evolution with proportion of invariable sites (I) and gamma

distribution of rate variation among sites (G). A partial topology showing the relationships of E. geiseri received

high level of support for several key nodes (Fig. 5). The five sequences for E. geiseri form a clade with 0.99

Bayesian probability, though demonstrating little internal structure. Erigeron geiseri, in turn, forms a clade

with E. sect Phalacroloma supported by 1.0 Bayesian probability and 98% bootstrap support from maximum

30°0'0"N

Journal of the Botanical Research Institute of Texas 9(1)

Fig. 4. Estimated geographic distribution of putative sexual diploid and polyploid apomictic Erigeron geiseri in Texas and Oklahoma, USA. 'SI -S4 7 and

'A1-A4'designate localities of specimens sampled for DNA study (Table 1).

30WN

Noyes et al., Phylogenetic affinities of Erigeron geiseri

i.o/6sr E - unjflorus ]

r E. borealis]

_ 0.99/87l - E. acrisj

E. uniflorus ] £ sect. Erigeron

E. sect. Trimorpha

0.65 / —

{

E. simplex

E. sect. Erigeron

1.0/99 L 5 gfra/?c//7/ori/s|

(to outgroup) - E. scopulinus ] £ sect. Scopuiincola

-E. /e/omeras] £ S ect. Asterigeron

-E. ursinus] £ sect. Rhizonexus

-E. karvinskianus^ E. sect. Karvinskia

0.71 / —

r t. religiosus I £ sect.

I— E. divergens I Olygotrichium

E. strigosus var. calcicola

E. strigosus var. dolomiticola

E. strigosus var. strigosus

— E. geiseri S3"

-E. geiseri S4

0.99 / — -E. geiseri A4

^ E. geiseri Al

£. sect.

Phalacroloma

0.55 / --T. £ geiseri A2

Fig. 5. Portion of ITS phylogeny for Astereae sect. Conyzineae. Sample designations for E geiseri according to Table 1. Values at nodes are Bayesian prob¬

abilities and bootstrap percentages respectively. Phylogeny for 81 sequences (Appendix 1) generated using a GTR+I+G evolutionary model. Taxonomic

sections according to Nesom (2008).

likelihood analysis. Together, E. sect. Phalacroloma and E. geiseri are sister to E. divergens Torr. & A. Gray and

E. religious Cronquist (E. sect. Olygotrichium). Erigeron sect. Phalacroloma, E. geiseri, E. sect. Olygotrichium, and

E. sect. Karvinskia (represented by E. karvinskianus DC.), in turn, form a well-supported clade distinct from all

other Erigeron taxa included in the analysis. There is no evidence that E. geiseri has taxonomic affinity with

members of E. sect Quercifolium or E. sect. Geniculactis. Those sections are so distant so as to preclude represen¬

tation in Figure 5.

Alignment for the ITS plus ETS concatenated analysis for 24 Erigeron sequences resulted in a matrix 1254

bp in length. The phylogenetic analyses employed the GTR plus G model of molecular evolution and yielded

trees similar in topology to the ITS-only analysis. The tree (Fig. 6) shows strong support for a monophyletic E.

geiseri (1.0 Bayesian probability, 63% maximum likelihood bootstrap support), but also shows support for in¬

ternal structure not present in the ITS-only analysis. Specifically, the ‘S3’ sample of coastal E. geiseri is shown

as sister to the remaining four samples. The combined sequence matrix shows that the ‘S3’ sequence differs

from ‘S4’ at 6 base positions, resulting in a genetic distance of 0.006. Sequences for ‘S4’, AT, A2’, and A4’ are

virtually identical (mean genetic distance 0.0001).

The remainder of the topology (Fig. 6) is consistent with the ITS-only topology (Fig. 5). Internal structure

of E. sect. Phalacroloma is supported with a basal E. strigosus var. calcicola, next divergent E. strigosus var. dolo¬

miticola, then three haplotype clades within E. strigosus var. strigosus as previously described (Noyes, 2006).

Support for E. sect. Phalacroloma plus E. geiseri increases with the addition of ETS sequence to 1.0 Bayesian

Journal of the Botanical Research Institute of Texas 9(1)

- E.hyssopifolius

- E.speciosus

- E.pinnatisectus

- E.acer

Outgroup

E.divergens^ £ sect. Olygotrichium —

var. calcicola -1543

— var. dolomiticola-'\ 545

var. strigosus HI-1316

i.o / ioo I - var. strigosus HI-1631

var. strigosus HI-1628

var. strigosus HI-1225

r var. strigosus HII-1608

1.0/ 941

I - var. strigosus HII-1610

var. strigosus HI 1-1616

var. strigosus HII-1617

■ var. strigosus Hill-1612

“var. strigosus Hill-1618

1 var. strigosus Hill-1614

^ var. strigosus Hill-1630

— E. geiseri S3

E. geiseri S4

“ E. geiseri A1

E. geiseri A2

E. geiseri A4

E. strigosus (sect.

Phalacroloma)

Fig. 6. ITS plus ETS phytogeny for Erigeron sect. Phalacroloma and Erigeron geiseri. Phytogeny for 24 sequences (Appendix 2) generated using a GTR+G

evolutionary model. Values at nodes are Bayesian probabilities and bootstrap percentages respectively. Sample designations for E. geiseri according to

Table 1. Sample designations for E. strigosus follow Noyes (2006).

probability and 100% maximum likelihood bootstrap support. In addition, support for the alliance between E.

sect. Phalacroloma (plus E. geiseri ) and E. sect. Olygotrichium is retained.

DISCUSSION

Erigeron geiseri likely includes sexual diploid and polyploid apomictic populations

The pollen analysis and distribution map support the hypothesis that E. geiseri includes both sexual diploid

and apomictic polyploid populations. The genetic similarity between samples is consistent with an evolution-

Noyes et al.. Phylogenetic affinities of Erigeron geiseri

ary model in which apomixis and polyploidy originated from within E. geiseri (via auto- or segmental poly¬

ploid origin) and not via hybridization involving other taxa. Alternatively, apomixis has introgressed into E.

geiseri from a related species, leaving no rDNA evidence of past ancestry Sequence analysis of apomixis genes

in Erigeron would be key for distinguishing these alternative hypotheses. However, this goal, for Erigeron or

any other natural apomict, remains elusive (Hand & Koltunow 2014).

The distribution of putative diploid sexuals versus polyploid apomicts in E. geiseri shows a pattern typical

for geographic parthenogenesis (Bierzychudek 1985) with sexual diploid populations with limited distribu¬

tion occurring to the south, and with apomicts with much wider distribution extending northward. Erigeron

geiseri is thus apparently similar to E. strigosus and E. tenuis (both E. sect. Phalacroloma) that feature narrowly

distributed sexual diploid populations in the southeastern US, and widespread triploid and tetraploid apo¬

micts extending northward (Noyes 2006; Groff 2010; Noyes & Givens 2013). Apomixis has also been docu¬

mented for E. annuus (E. sect. Phalacroloma; Noyes & Givens 2013), as well as E. divergens (E. sect. Olygotrichi-

um) of the western US, and also in E. karvinskianus (E. sect. Karvinskia) and so the trait is evidently common

among species in these related sections.

Sexual diploid plants in E. sect. Phalacroloma thus far investigated have a relatively rare form of reproduc¬

tive development, tetraspory, whereby meiosis of the megasporocyte yields a coenospore with four nuclei that

each undergo mitotic divisions to yield a genetically mosaic female gametophyte (Noyes & Allison 2005).

Tetraspory is thought to represent a temporal developmental shift in the timing of key meiotic events and may

represent a predisposition toward the evolution of apomictic development (Carman 1997). Tetraspory is pos¬

sibly common in Erigeron, perhaps more frequent than the classic monosporic ‘Polygonum’ type development

that is characteristic of most Asteraceae (Noyes 2007). We would therefore predict that sexual diploid popula¬

tions of E. geiseri would exhibit tetrasporic development and that apomicts would likely exhibit diplospory as

is found in apomictic relatives. Chromosome number and reproductive developmental studies are warranted;

the distribution map provided by this study will be a useful guide for locating populations. Further held study

also may be warranted to investigate genetic and morphological differentiation between coastal and interior

sexual diploid populations.

Erigeron geiseri is closely allied with E. sect. Phalacroloma

Our analysis of nrDNA sequence shows that samples of Erigeron geiseri form a monophyletic group that is sister

to E. strigosus and allies that constitute E. sect. Phalacroloma. Shinners (1947) hypothesized that E. geiseri was

related to E. coronarius based on the similarities including a biseriate few-bristled pappus, conical receptacle,

and annual habit. The later species presently serves as the type of Erigeron sect. Geniculactis, a group of six spe¬

cies restricted to Mexico and adjacent southern Arizona (Nesom 2008). Our ITS analysis included a sample for

E. coronarius, however, and no affinity between the two species was supported (Fig. 5). The classification of E.

geiseri in E. sect. Quercifolium (Nesom 2008) is also not supported. Our ITS sample included E. quercifolius (the

type species of the section), E. philadelphicus, E. dryophyllus, and E. veracruzensis of the section. While these

four species form a monophyletic clade, they are not closely related to E. geiseri (Fig. 5; Noyes 2000). Instead,

an affinity of E. geiseri with E. sect. Phalacroloma is supported (Fig. 6), a finding foreshadowed by earlier dis¬

cussion noting morphological similarities between E. geiseri and E. tenuis (Nesom 1987; Nesom 2008). Based

on nrDNA sequence, sexual diploid E. tenuis resides as a clade within E. strigosus with some individuals exhib¬

iting both ‘tenuis-like’ and ‘strigosus-like’ nrDNA haplotypes consistent with recent hybridization between the

two taxa (Noyes 2000; Groff 2010). Erigeron geiseri and E. tenuis, along with E. turnerorum (of Nuevo Leon,

Mexico) and E. tenellus (of extreme southern Texas and adjacent Mexico) share a suite of floral and vegetative

characters, which together with geographic parapatry, is consistent with close evolutionary relationship (Ne¬

som 1987). Erigeron geiseri is distinct from E. tenuis, and more similar to E. turnerorum and E. tenellus, because

of its tap-rooted, annual habit. Erigeron tenuis and other E. sect. Phalacroloma (E. strigosus and E. annuus ) are

fibrous-rooted and longer lived. Erigeron geiseri, E. tenellus, and E. turnerorum are distinct from the other six¬

teen species classified as E. sect. Quercifolium (Nesom 2008) in being the only tap-rooted taxa. We recommend

based on our results that E. geiseri be reclassified as a member of E. sect. Phalacroloma. By extension, based on

Journal of the Botanical Research Institute of Texas 9(1)

morphological considerations, we would predict that DNA study will show that E. tenellus and E. turnerorum

are also members of E. sect. Phalacroloma.

While our conclusion regarding the taxonomic reclassification of E. geiseri is well supported by the ITS

and ETS sequence, we must add the caveat that our phylogeny is nonetheless based on data from a single gene

region. Although ITS has been a mainstay in lower level plant systematics for nearly two decades (Feliner &

Rosello 2007) and has been proposed to be included, along with select chloroplast gene regions, as a compo¬

nent of a universal plant barcode (China Plant BOL Group 2011), it nonetheless potentially reflects a biased

portrait of relationship. It has long been appreciated that because of phenomena such as recombination, hy¬

bridization, lineage sorting, and horizontal gene transfer, different genes may yield topologically incongruent

phylogenies (Wendel & Doyle 1998). The problem is exemplified by recent work in Symphyotrichum (also tribe

Astereae) in which study of ITS plus ETS on the one hand, and the unlinked 5S nrDNA nontranscribed spacer

on the other, revealed seven phylogenetic disagreements attributed to three reticulate evolutionary events in¬

volving six different species (Morgan & Holland 2012). Multigene, genome wide phylogenetic reconstruction,

successfully implemented for study of the highest levels of plant diversity (e.g., Angiosperm Phylogeny Group

2009; Finet et al. 2010), remains elusive at lower taxonomic levels for many groups owing to the difficulty of

identifying strictly orthologous single copy nuclear genes (Feliner & Rosello 2007). Nonetheless, we anticipate

advances from the study of a small number of nuclear markers plus the further refinement of enrichment tech¬

niques coupled with Next Generation Sequencing methodologies (Mandel et al. 2014) to soon provide a means

for generating robust phylogenies for species rich complex groups as represented by North American Erigeron.

APPENDIX 1

Taxa and GenBank numbers for 76 ITS sequences of Astereae sect. Conyzinae used for phylogenetic analysis (AF046986:AF046992;

AF118478:AF118487; AF118491 :AF118545; AY871316; AY871318; AY871323; AY871333). GenBankaccession numbers for five ITS sequences

for Erigeron geiseri given in Table 1.

Aphanostephus ramossissimus [AF046990];Aph.skirrhobasis [AF118521 ];Apopyroswarmingii [AF118511];Conyzaapurensis[AF118482];

C. bonariensis [AF118513]; C. canadensis [AF046987]; C. chilensis [AF118510]; C. floribunda [AF118514]; C. ramosissima [AF118484];

Erigeron acris [AF118496]; E. aliceae [AF118541]; E. aphanactis [AF118531]; E. arenarioides [AF118528]; E. argentatus [AF118506]; E.

barbellatus [AF118536]; E. bellioides [AF118522]; E. borealis [AF118495]; E. caespitosus [AF118481 ]; E. cinereus [AF118502]; E. compositus

[AF118537]; E. coronarius [AF118520]; E. divergens [AF118485]; E. dryophyllus [AF118524]; E. eatonii [AF118540]; E. eruptens [AF118543];

E.fernandezianus [AF118515]; E.flettii [AF118503]; E.foliosus [AF118532]; E.formosissimus [AF118478]; E.fraternus [AF118518]; E. galeotii

[AF118519]; E. g la bell us [AF118498]; E.glaucus [AF118499]; E. grand iflorus [AF118494]; E. humilus [AF118504]; E. hyssopifolius [AY871316];

E. inornatus [AF118533]; E. kachinensis [AF118480]; E. karvinskianus [AF118487]; E. leiomerus [AF118492]; E. lepidopodus [AF118545];

E. leptorhizon [AF118483]; E. linearis [AF118534]; E. lonchophyllus [AF118505]; E. maximus [AF118509]; E. neomexicanus [AF118544]; E.

ochroleucus [AF118539]; E. philadelphicus [AF046989]; E. pinnatisectus [AF118501]; E. pinnatus [AF118517]; E. podophyllus [AF118542];

E. pulchellus [AF118500]; E. pumilus [AF118530]; E. pygmaeus [AF118526]; E. quercifolius [AF118525]; E. religiosus [AF118486]; E. rhizoma-

tus [AF046992]; E. rosulatus-1 [AF118512]; E. rosulatus-2 [AF118516]; E. scoparioides [AF118527]; E. scopulinus [AF118497]; E. simplex

[AF118493]; E. speciosus [AF118479]; E. strigosus var. calcicola [AY871318]; E. strigosus var. dolomiticola [AY871323]; E. strigosus var. stri-

gosus [AY871333]; E. tener [AF118535]; E.tweedyi [AF118529]; E. uniflorus [AF046988]; E. ursinus [AF118491]; E. utahensis [AF118507]; E.

vagus [AF118538]; E. veracruzensis [AF118523]; E. wislizenii [AF046991]; Hysterionicajasionoides [AF046986]; Nejafiliformis [AF118508].

APPENDIX 2

Taxa and GenBank numbers for 19 ITS plus ETS sequences used for phylogenetic analysis of Erigeron sect. Phalacroloma and E. geiseri

(ITS: AF118479, AF118496, AF118501, AY871316:AY871318, AY871323:AY871330, AY871333, AY871334, AY871340, AY871341, AY871553;

ETS: AY871355:AY871360, AY871365:AY871372, AY871375, AY871376, AY871382, AY871383, AY871744 ). GenBank accession numbers

for five ITS plus ETS sequences for E. geiseri given in Table 1.

Outgroups:

Erigeron acris [AF118496 (ITS), AY871356 (ETS)]; E. hyssopifoiius [AY871316, AY871355]; E. pinnatisectus [AF118501, AY871357]; E.

speciosus [AF118479, AY871358]

Erigeron sect. Olygotrichium:

Erigeron divergens [AY871317, AY871359]

Erigeron sect. Phalacroloma:

Erigeron strigosus var. calcicola [AY871318, AY871360]; E. strigosus var. dolomiticola [AY871323, AY871365]; E. strigosus var. strigosus

rDNA haplotype I: [AY871325, AY871367], [AY871326, AY871368], [AY871327, AY871369], [AY871324, AY871366]; rDNA haplotype II:

[AY871328,AY871370], [AY871553, AY871744], [AY871329,AY871371], [AY871330,AY871372];rDNAhaplotypelll [AY871333,AY871375],

[AY871334, AY871376], [AY871340, AY871382], [AY871341, AY871383].

Noyes et al., Phylogenetic affinities of Erigeron geiseri

ACKNOWLEDGMENTS

We would like to thank curators at the University of Texas and the Botanical Research Institute of Texas for

loans of specimens of E. geiseri and permission to sample material, the College of Natural Sciences and Math¬

ematics, UCA, for an undergraduate research award to Will Caraway, helpful comments from an anonymous

reviewer and Lowell Urbatsch, Louisiana State University, and the Department of Biology, UCA, for contribu¬

tion to resources used for this study

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NEW COMBINATIONS IN CORYPHANTHA AND ESCOBARIA (CACTACEAE)

Root Gorelick

Department of Biology, School of Mathematics & Statistics, and

Institute of Interdisciplinary Studies

Carleton University, 1125 Raven Road

Ottawa, Ontario K1S 5B6, CANADA

Root. Gorelick@carleton.ca

ABSTRACT

Most cactus taxonomists accept Coryphantha organensis as a synonym of Coryphantha orcuttii. Therefore, at infraspecific rank, Coryphantha

orcuttii needs to be considered a variety (not a subspecies) of C. sneedii because C. orcuttii is partially sympatric with the type subspecies/

variety of C. sneedii at Anthony’s Nose, Dona Ana County, New Mexico. This requires a new combination for Coryphantha orcuttii and

Escobarla orcuttii at the rank of variety, depending on whether you consider Escobaria to be a separate genus versus subgenus/section of

Coryphantha. The two new combinations are: Coryphantha sneedii var. orcuttii comb, et stat. nov. and Escobaria sneedii var. orcuttii

comb, et stat. nov.

RESUMEN

La mayoria de los taxonomos de cactus aceptan Coryphantha organensis como sinonimo de Coryphantha orcuttii. Sin embargo, a nivel infra-

specifico, Coryphantha orcuttii necesita ser considerado una variedad (no una subespecie) de C. sneedii porque C. orcuttii es parcialmente

simpatrico con la subespecie/variedad tipo de C. sneedii en Anthony’s Nose, Dona Ana County, Nuevo Mexico. Esto requiere una nueva

combinacion para Coryphantha orcuttii y Escobaria orcuttii en el rango de variedad, dependiendo si se considera o no que Escobaria sea un

genero separado del subgenero/seccion Coryphantha. Las dos nuevas combinaciones son: Coryphantha sneedii var. orcuttii comb, et stat.

nov. y Escobaria sneedii var. orcuttii comb, et stat. nov.

INTRODUCTION

While drafting a manuscript on cacti that survive unprotected outdoors in Canada (Gorelick et al. 2015), I

wanted to use a combination for a plant taxon in the Coryphantha sneedii complex that has not been published.

A few of the varieties/subspecies of C. sneedii (Britton & Rose) A. Berger, including C. orcuttii (Rose ex Orcutt)

Zimmerman and the type variety/subspecies of C. sneedii Britton & Rose, are the only cacti that I have success¬

fully grown unprotected in both Tempe, Arizona, USA where summer temperatures occasionally hit +50°C,

and Ottawa, Ontario, Canada where winter temperatures often hit -30°C, and that is without wind chill.

The type subspecies/variety of Coryphantha sneedii appears to be ancestral (Baker 2004) to all other sub¬

species, varieties, and forms, which are spread out radially in all directions (Zimmerman 1985): ‘leei’ and

‘guadalupensis ’ to the east; ‘ villardii ’ to the northeast; ‘ sandbergii’ to the north; ‘ orcuttii ’ to the west; two of Allan

Zimmerman’s unnamed varieties to the south; and ‘ albicolumnaria’ to the southeast. But, when first described,

it was not obvious whether these infraspecific taxa in the C. sneedii complex were partially sympatric or allo-

patric with one another hence it was not obvious whether these were subspecies or varieties (or merely just

forms).

Subspecies versus variety

Usage of ‘subspecies’ and ‘variety’ has at times been confused and has had a changing history in the more than

two centuries since both terms have been in existence (Clausen 1941; Stuessy 2009). Meanings of these two

infraspecific categories gradually stabilized by the mid 1900s (Clausen 1941), but there still is some contention

about their usage even to this day (Stuessy 2009). I therefore begin by describing Clausen’s (1941) definitions

given in a paper titled On the use of the terms ‘subspecies’ and ‘variety’, followed by Hamilton and Reichard’s

(1992) paper titled Current practice in the use of subspecies, variety, and forma in the classification of wild plants,

J. Bot. Res. Inst. Texas 9(1): 25 - 30.2015

Journal of the Botanical Research Institute of Texas 9(1)

followed by Turner and Nesom’s (2000) paper whose title begins Use of variety and subspecies, and finally

Stuessy’s (2009) definitions of subspecies and variety from his widely used text titled Plant taxonomy.

Subspecies are comprised of individuals with a consistent suite of morphological traits that differ from

other individuals of the species due to geographic, ecological, or physiological isolation (Clausen 1941). Geo¬

graphic and ecological isolation are easier for taxonomists to discern than physiological isolation, so I ignore

physiology herein. Both varieties and subspecies have consistent morphological features. If that morphology is

due to geographic or ecological variation, then Clausen (1941) called that taxon a subspecies, otherwise he

called it a variety.

Hamilton and Reichard (1992:491) “recogniz[ed] subspecies for groups primarily with greater geograph-

ic/ecologic distinctness than varieties...” Thus, like Clausen (1941), they required some modicum of geograph¬

ic or ecological isolation. Hamilton and Reichard (1992) also noted that European plant taxonomists tend to

use ‘subspecies’ more often, while North American plant taxonomist tend to use ‘variety’ more often.

Turner and Nesom (2000: 258) stated that, “use of the subspecies rank may point to larger patterns of

variation and/or coherence within the species. This use of infraspecific categories Ends support in the ICBN,

which implies that the term subspecies is used for clustering varieties.”

Stuessy (2009) did not so much define subspecies and variety, but provided a list of characters that are

useful in distinguishing the two. These characters are in the following categories: morphology, geographic

overlap, genetic divergence, likelihood of naturally hybridizing, and fertility of hybrids. For the Coryphantha

sneedii complex, we really only know about geographic overlap and morphology, so can ignore the other char¬

acteristics, meaning that difference between subspecies and variety still depend on both morphology and

geographical isolation, as was true three-quarters of a century ago (Clausen 1941). For Stuessy (2009), subspe¬

cies have a larger number of traits with morphological differences than do varieties. For Stuessy (2009), sub¬

species are largely allopatric or peripatric, while varieties can have somewhat overlapping distributions.

Some authors (e.g., Corogin & Judd 2014, in this journal) even seem to require that subspecies be allopat¬

ric. By contrast, “In American taxonomy, possibly the most widely accepted concept regarding varieties is that

... varieties are expected to overlap in distribution ...” (A. Michael Powell, pers. comm., 27 Dec 2014). Several

authors outside of North America also require varieties, but not subspecies, to have overlapping distributions,

e.g., “It is generally held that the rank of subspecies should be used for populations of a species that have sev¬

eral morphological differences and are geographically and/or ecologically separated, whereas varieties, also

with some morphological differences, overlap in their distribution” (Newton & Thiede 2015: 30). Therefore in

attempting to decide whether members of the Coryphantha sneedii complex are subspecies or varieties, we need

to focus on number of morphological differences and especially on geographic isolation.

Coryphantha versus Escobaria

Should Coryphantha sneedii be placed into the segregate genus Escobaria Britton & Rose? From a purely formal

view, this does not matter here because Coryphantha sneedii and Escobaria sneedii (Britton & Rose) A. Berger

have both been validly published. I leave both options open, even though I consider Escobaria to be a subgenus

[Coryphantha (Engelm.) Fern, subgenus Escobaria (Britton & Rose) A. Berger]. Other authors consider Esco¬

baria to be a section of Coryphantha [Coryphantha section Escobaria (Britton & Rose) H.E. Moore] This portion

of the paper should thus be considered dicta, i.e., this debate does not affect the conclusions of the rest of this

paper because I will make combinations in both genera. Debates over whether Escobaria is a genus or a subge¬

nus or a section of Coryphantha will probably continue for many years.

Any attempt to segregate Escobaria from Coryphantha will probably result in paraphyly. Thus the Flora of

North American (Parhtt & Gibson 2003; Zimmerman & Parhtt 2003) subsumes Escobaria under the earlier

valid genus epithet Coryphantha, as did Benson (1982), Powell and Weedin (2004), and most other North

American authors. Paraphyly provides a reason to dismiss Escobaria, even though there are morphological dif¬

ferences, albeit not consistent morphological differences, between taxa considered to be Escobaria versus Co¬

ryphantha. Subgenus Escobaria has pitted seeds, whereas subgenus Coryphantha does not (Anderson 2001;

Zimmerman & Parhtt 2003; Dicht & Futhy 2005; Hunt et al. 2006), except for C. gracilis L. Bremer & A.B. Fau

Gorelick, New combinations in Coryphantha and Escobaria

(Taylor 1986; Dicht & Luthy 2005). Most taxa in subgenus Coryphantha contain extra-floral nectary glands,

whereas all taxa in subgenus Escobaria lack these glands (Zimmerman & Parhtt 2003; Dicht & Luthy 2005).

Subgenus Escobaria generally has smaller flowers, fruits, and seeds than subgenus Coryphantha (Castetter et al.

1975). For these reasons, most European authors consider Escobaria to be a valid genus (Dicht & Luthy 2005),

even if many North Americans do not (Zimmerman & Parhtt 2003).

But, with the logic of trying to avoid paraphyly, there may also be reason to dismiss Coryphantha as a ge¬

nus and subsume it under Mammillaria. A phylogenetic analysis based solely on chloroplast markers showed

Coryphantha, Escobaria, Pelecyphora Ehrenb., Ortegocactus Alexander, and Neolloydia conoidea Britton & Rose

to all be embedded within Mammillaria Haw. (Butterworth & Wallace 2004), i.e., making both Escobaria and

Coryphantha paraphyletic. However, it is not obvious how much weight to place on that 2004 molecular study

because chloroplast inheritance is not always maternal in the Cactaceae (Corriveau & Coleman 1988; Gorelick

2014) and there is probably some alloploidy in this clade, e.g., the tetraploid Coryphantha missouriensis (Scheer)

Britton & Rose (Weedin & Powell 1980), indicating the need for reticulate phylogenies. I therefore lean to¬

wards retaining use of Coryphantha as a genus, subsuming all Escobaria therein, but could be persuaded other¬

wise after more systematic analyses are performed based on both genotype and phenotype, especially if nucle¬

ar markers are used in constructing molecular phylogenies.

Coryphantha orcuttii , C . organensis , C. sneedii

In New Mexico, Coryphantha orcuttii ranges from the Peloncillo Mountains to the Florida Mountains (Fergu¬

son 1998a). This taxon is also found in neighboring Arizona, Chihuahua, and Sonora. This range extends to

the Organ Mountains if C. organensis Zimmerman is considered a synonym of C. orcuttii (see below). C. orcuttii

and C. sneedii var. albicolumnaria (Hester) A.D. Zimmerman are by far the most widespread taxa in the Cory¬

phantha sneedii complex.

The type subspecies/variety of Coryphantha sneedii is endemic to the Franklin Mountains in El Paso

County, Texas, the southern end of the Organ Mountains in Dona Ana County, New Mexico, and some of the

mountains between the two, e.g., Bishop’s Cap and Anthony’s Nose.

Given the huge amount of morphological variation and geographic range in the Coryphantha sneedii com¬

plex (Zimmerman 1985), it makes sense to follow the modern consensus amongst cactus taxonomists that C.

orcuttii should be considered an infraspecific form of C. sneedii (Zimmerman & Parhtt 2003; Baker 2004; Pow¬

ell & Weedin 2004; Hunt et al. 2006). Thus, Coryphantha orcuttii is usually considered to be a synonym of Es¬

cobaria sneedii Britton & Rose subsp. orcuttii (Boed.) Luthy, but here I argue that a subspecies designation is

inappropriate for this taxon. Coryphantha orcuttii seems to be morphologically indistinguishable from Cory¬

phantha organensis (synonym: Escobaria organensis (Zimmerman) Castetter, P. Pierce, & K.H. Schwer.) (Fergu¬

son 1998 a,b). The only morphological trait that supposedly consistently distinguishes Coryphantha organensis

from the rest of the C. sneedii complex is a slight yellowish cast to spine color in C. organensis (Zimmerman

1972a; Taylor 1986; Ferguson 1998b). Color is typically a variable trait, often encoded by an allelic shift at a

single genetic locus, thus at most warranting designation as a form, not a variety, let alone a subspecies (Stuessy

2009). Therefore many authors consider C. organensis to be a synonym of C. orcuttii, itself possibly subsumed

under C. sneedii (e.g., Luthy 1999; Hunt et al. 2006; U.S. Geological Survey’s ‘Integrated Taxonomic Informa¬

tion System’ (ITIS; http://www.itis.gov/)). If C. organensis is a synonym of C. orcuttii, then the subspecies desig¬

nation of C. orcuttii as Escobaria sneedii subsp. orcuttii and the hitherto unpublished combination Coryphantha

sneedii subsp. orcuttii cannot be valid because C. organensis and C. sneedii subsp. sneedii have overlapping dis¬

tributions at Anthony’s Nose (Ferguson 1998 b,c; Gorelick 2006), which straddles the Texas-New Mexico

border, and probably also have overlapping distributions at North Anthony’s Nose (‘nose’ is an idiosyncratic

local term for these mountains). Anthony’s Nose and North Anthony’s Nose are separated by Anthony Gap,

through which runs New Mexico State Highway 404, at the northern end of the Franklin Mountains (Dona

Ana County, New Mexico and El Paso County, Texas), south of Fillmore Pass and the Organ Mountains. Clau¬

sen (1941), Hamilton and Reichard (1992), Stuessy (2009), and Corogin and Judd (2014) all require allopatry or

peripatry between two or more subspecies of a given species. Turner and Nesom (2000) do not require allopa-

Journal of the Botanical Research Institute of Texas 9(1)

try or peripatry between subspecies, but insist that a subspecies be an aggregate of two or more varieties. How¬

ever, that is not the case here because C. organensis is a synonym of C. orcuttii and there are no other obvious

varieties in the Coryphantha sneedii complex west of the franklin and organ Mountains. The only possible ex¬

ceptions are Escobaria orcuttii var. macraxina Castetter, P. Pierce & K.H. Schwer. from the Big Hatchet Moun¬

tains and Escobaria orcuttii var. koenigii Castetter, P. Pierce & K.H. Schwer. from the Florida Mountains

(Castetter et al. 1975; Taylor 1986), but nobody nowadays, including me, accepts these as valid varieties or even

consistent forms or morphs. Therefore a subspecies designation is inappropriate for Coryphantha orcuttii seem¬

ingly regardless of usage of the designation ‘subspecies’.

Instead of a subspecies designation, should Coryphantha orcuttii be considered a variety of C. sneedii? Co¬

ryphantha orcuttii is consistently morphologically different from C. sneedii subsp. sneedii, despite these two

taxa being partially sympatric at Anthony’s Nose. Coryphantha sneedii has many more shoots, sometimes

hundreds more, than C. orcuttii, which is often solitary (Ferguson 1998a,c; Anderson 2001; Hunt et al. 2006).

Coryphantha sneedii has smaller shoots (2.5-10 x 1.0-2.5 cm) than C. orcuttii (15 x 6-9 cm) (ibid). Coryphan¬

tha sneedii has shorter central spines (4.5-10 mm) than C. orcuttii (9-22 mm) (ibid). Seemingly the only viable

alternative is therefore to consider Coryphantha organensis to be a synonym of C. orcuttii and then to classify C.

orcuttii as a variety of either Coryphantha sneedii or Escobaria sneedii (not a variety of Coryphantha strobiliformis

(Poselg.) Moran, where the name orcuttii currently exists at the rank of variety). This requires one of the follow¬

ing two new combinations, the choice of which depends on whether or not you consider Escobaria to be a sepa¬

rate genus of subgenus/section of Coryphantha:

Coryphantha sneedii var. orcuttii (Boed.) Gorelick, comb, et stat. nov. Basionym: Escobaria orcuttii Bodecker, Ein Mam-

millarien Vergleichs-Schlussel 17.1933. Type: U.S.A. New Mexico. Hidalgo Co.: Peloncillo Mountains, near Granite Pass, Mar 1926,

J.N. Rose s.n. (lectotype, designated by Benson 1969: 26: DS).

Coryphantha strobiliformis (Poselg.) Moran var. orcuttii (Rose ex Orcutt) L.D. Benson, Cacti Ariz. ed. 3,26.1969.

Coryphantha orcuttii (Rose ex Orcutt) Zimmerman, Cact. Succ. J. (US) 44:156.1972.

Coryphantha organensis Zimmerman, Cact. Succ.J. (US) 44:114.1972.

Escobaria organensis (Zimmerman) Castetter, P. Pierce & K.H. Schwer., Cact. Succ.J. (US) 47:60.1975.

Escobaria sneedii Britton & Rose subsp. orcuttii (Boed.) Liithy, Kakt. and. Sukk. 50:278.1999.

Escobaria sneedii Britton & Rose subsp. organensis (Zimmerman) Liithy, Kakt. and. Sukk. 50:278.1999.

Escobaria sneedii var. orcuttii (Boed.) Gorelick, comb, et stat. nov. Basionym: Escobaria orcuttii Bodecker, Ein Mammil-

larien Vergleichs-Schliissel 17.1933. Type: U.S.A. New Mexico. Hidalgo Co.: Peloncillo Mountains, near Granite Pass, Mar 1926 J.N.

Rose s.n. (lectotype, designated by Benson 1969: 26: DS).

Coryphantha strobiliformis (Poselg.) Moran var. orcuttii (Rose ex Orcutt) L.D. Benson, Cacti Ariz. ed. 3,26.1969.

Coryphantha orcuttii (Rose ex Orcutt) Zimmerman, Cact. Succ.J. (US) 44:156.1972.

Coryphantha organensis Zimmerman, Cact. Succ.J. (US) 44:114.1972.

Escobaria organensis (Zimmerman) Castetter, P. Pierce & K.H. Schwer., Cact. Succ.J. (US) 47:60.1975.

Escobaria sneedii Britton & Rose subsp. orcuttii (Boed.) Liithy, Kakt. and. Sukk. 50:278.1999.

Escobaria sneedii Britton & Rose subsp. organensis (Zimmerman) Liithy, Kakt. and. Sukk. 50:278.1999.

According to Zimmerman (1972b), Joseph N. Rose in 1926 named this taxon ‘Neolloydia orcuttii ’ (nomen nu¬

dum) for a specimen collected by Charles Russell Orcutt from Granite Pass, near the New Mexico-Arizona

border, in the Peloncillo Mountains, and Rose did not designate a type. Granite Pass in the Peloncillo Moun¬

tains of New Mexico is almost certainly now known as “Granite Gap.” I have been unable to trace when this

wording change occurred other that it was on or before the mid 1950s (Gillerman 1957). The global unique

identifier for the lectotype is “CAS:DS:307410.” The California Academy of Sciences website (http://collections.

calacademy.org/bot/) gives a collection date of 28 March 1926 for this specimen. According to Roy Mottram

(pers. comm., 14 Feb 2014), Rose also provisionally published the combination ‘ Escobaria orcuttf for this plant

taxon in Cactography 4(1):5, 1926. Spelling of orcutti with a single ‘i’ also appears in the documentation for

Coryphantha strobiliformis in Benson (1982: 963), probably as a typographical error.

Gorelick, New combinations in Coryphantha and Escobaria

ACKNOWLEDGMENTS

Many thanks to A. Michael Powell, Barney Lipscomb, Roy Mottram, and an anonymous reviewer for greatly

improving this manuscript. This work was funded by a Natural Science and Engineering Research Council of

Canada (NSERC) Discovery Grant. I also belatedly thank the late Robert Clausen. As a first-year undergradu¬

ate mathematics major in 1976,1 stumbled into his research greenhouse around the time that he was retiring.

He not only shared his enthusiasm, but also a few small cuttings of Mexican Crassulaceae.

REFERENCES

Anderson, E.F. 2001. The cactus family. Timber Press, Portland, Oregon, U.S.A.

Baker, M.A. 2004. Further elucidation of the taxonomic relationships and geographic distribution of Escobaria sneedii

var. sneedii, E. sneedii var. leei, and E. guadalupensis (Cactaceae). In: P. Barlow-lrick, J. Anderson & C. McDonald, eds.

Southwestern rare and endangered plants: Proceedings of the Fourth Conference (22-26 March 2004). U.S. Depart¬

ment of Agriculture, Forest Service, Rocky Mountain Research Station (RMRS-P-48CD), Las Cruces, New Mexico, U.S.A.

Benson, L.D. 1982. The cacti of the United States and Canada. Stanford University Press, Stanford, California, U.S.A.

Butterworth, C.A. & R.S. Wallace. 2004. Phylogenetic studies of Mammillaria (Cactaceae) - insights from chloroplast se¬

quence variation and hypothesis testing using the parametric bootstrap. Amer. J. Bot. 91:1086-1098.

Castetter, E.F., P. Pierce, & K.H Schwerin. 1975 A reassessment of the genus Escobaria. Cactus Succ. J. (U.S.) 47:60-70.

Clausen, R.T. 1941. On the use of the terms "subspecies" and "variety." Rhodora 43:167-167.

Corogin, P.T. & W.S. Judd. 2014. New geographical and morphological data for Sideroxylon reclinatum subspecies austro-

floridense (Sapotaceae), a taxon endemic to southeastern peninsular Florida, U.S.A. J. Bot. Res. Inst.Texas 8:403-417.

Corriveau, J.L. & A.W. Coleman. 1988. Rapid screening method to detect potential biparental inheritance of plastid DNA

and results for over 200 angiosperm species. Amer. J. Bot. 75:1443-1458.

Dicht, R.F. & A.D. Luthy. 2005. Coryphantha: cacti of Mexico and southern USA. Springer-Verlag, Berlin, Germany.

Ferguson, DJ. 1998a. Escobaria orcuttii (Orcutt pincushion cactus) In: New Mexico rare plants. New Mexico Rare Plant

Technical Council: Albuquerque. Available at: http://nmrareplants.unm.edu. Accessed 18 July 2014.

Ferguson, D.J. 1998b. Escobaria organensis (Organ Mountains pincushion cactus) In: New Mexico Rare Plants. New Mex¬

ico Rare Plant Technical Council: Albuquerque. Available at: http://nmrareplants.unm.edu. Accessed 18 July 2014.

Ferguson, DJ. 1998c. Escobaria sneedii v ar. sneedii (Sneed's pincushion cactus) In: New Mexico Rare Plants. New Mexico

Rare Plant Technical Council: Albuquerque. Available at: http://nmrareplants.unm.edu. Accessed 18 July 2014.

Gillerman, E. 1957. Geology of the central Peloncillo Mountains, Hidalgo County, New Mexico and Cochise County,

Arizona. PhD dissertation. University of Texas, Austin, U.S.A.

Gorelick, R. 2006 Coryphantha dasyacantha found in New Mexico...and the cacti at Anthony Gap. Cactus Succ. J. (U.S.)

78:184-189.

Gorelick, R. 2014. Fishing for philosophical phylogenetic foibles. Ideas Ecol. Evol. 7:8-10.

Gorelick, R,T.D. Drezner, & K. Hancock. 2015. Freeze-tolerance of cacti (Cactaceae) in Ottawa, Ontario, Canada. Madrono

62:32-44.

Hamilton, C.W. & S.H. Reichard. 1992. Current practice in the use of subspecies, variety, and forma in the classification of

wild plants. Taxon 41:485-498.

Hunt, D.R., N.P. Taylor, & G. Charles. 2006. The new cactus lexicon. DH Books, Milborne Port, U.K.

Luthy, J.M. 1999. Escobaria sneedii subsp. orcuttii. Kakteen And. Sukk. 50:278.

Newton, L.E. & J. Thiede. 2015. Rank adjustments for the infraspecific taxa of Sansevieria pinguicula P.R.O. Bally (Asparaga-

ceae/Dracaenaceae). Cactus Succ. J. (U.S.) 87:28-32.

Parfitt, B.D. & A.C. Gibson. 2003. 37. Cactaceae Jussieu • cactus family. In: Flora of North America, volume 4. Oxford Uni¬

versity Press, New York, New York, U.S.A. Pp. 92-257.

Powell, A.M. & J.F. Weedin. 2004. Cacti of the Trans-Pecos and adjacent areas. Texas Tech University Press, Lubbock, Texas,

U.S.A.

Stuessy, T.F. 2009. Plant taxonomy: the systematic evaluation of comparative data (2nd edition). Columbia University

Press, New York, New York, U.S.A.

Taylor, N.P. 1986. The identification of Escobarias (Cactaceae). British Cact. Succ. J. 4:36-44.

Turner, B.L. & G.L. Nesom. 2000. Use of variety and subspecies and new varietal combinations for Stryaxplatanifolius

(Styracaceae). Sida 19:257-262.

SCLERIA BELLII (CYPERACEAE), A DISTINCTIVE AND UNCOMMON

NUTSEDGE FROM THE SOUTHERN U.S., CUBA, AND MEXICO

Richard J. LeBlond, Samantha M.Tessel, and Derick B. Poindexter

University of North Carolina Herbarium (NCU)

University of North Carolina at Chapel Hill

Chapel Hill, North Carolina, 27599-3280, US. A.

richardleblond@charter.net

ABSTRACT

A new species of Sderia (Cyperaceae) is described from the southeastern United States, Cuba, and Mexico. Although widespread, known

populations are few and scattered, with 14 in the United States from North Carolina to Texas, and one each in Cuba and Mexico. In the

United States, 12 populations are found on the outer Coastal Plain, and two in the Carolina Piedmont. At known sites, soils are wet, and

several populations occur in rare and unusual habitats with acidic pH and a calcareous influence.

RESUMEN

Se describe una nueva especie de Sderia (Cyperaceae) del sureste de Estados Unidos, Cuba, y Mexico. Aunque muy extendida, las poblacio-

nes conocidas son pocas y esparcidas, con 14 en los Estados Unidos desde Carolina del Norte a Texas, y una en Cuba y otra en Mexico. En los

Estados Unidos, 12 poblaciones se encuentran en la llanura costera externa, y dos en el piedemonte de Carolina. En los sitios conocidos, los

suelos son humedos, y varias poblaciones se dan en habitats raros e inusuales con pH acido y una influencia calcarea.

INTRODUCTION

A distinctive nutsedge sharing characteristics of the Scleria ciliata complex was found in southeastern North

Carolina during natural area inventories for the state’s Natural Heritage Program in the 1990s. Since that dis¬

covery, the new taxon was treated as “Scleria species 1” in Weakley (2013). Subsequently, more than 1,200

herbarium collections of taxa in the S. ciliata complex were examined from AUA, BRIT (including SMU and

VDB), DUKE, FLAS, FSU, GH, FSU, MO, NCSC, NCU, NY, US, and USCH. As a result, 16 populations of the

new nutsedge are now known from Florida, Georgia, Fouisiana, North Carolina, South Carolina, Texas, Cuba,

and Mexico (Fig. 1). Among the 14 populations known from the U.S., 12 are located on the outer Atlantic and

Gulf Coastal Plain, and two are from the Carolina Piedmont. This widespread but apparently rare nutsedge is

here described as a new species.

Scleria bellii FeBlond, sp. nov. (Fig. 2). Type: UNITED STATES. North Carolina. Onslow Co.: in moderately dense graminoid-

forb ground cover of wet savanna, formerly known as Haws Run Mitigation Site, now part of Maple Hill Savannas State Natural Area,

near Onslow/Pender county line between Sandy Run Swamp and Shelter Swamp Creek, 6 Jun 2013, R.J. LeBlond 6958 (holotype:

NCU; isotypes: AUA, BRIT, FLAS, FSU, GH, LSU, MO, NY, US, USCH).

Achene body smoothish with broad low rises, a few retrorse papillae at the very base; achene including hypogynium 2.5-3.2(-3.4) mm long,

2.3-2.7 mm wide; hypogynium basal rim 1.4-1.8(-2.1) mm wide; tubercles (4-)6.

Plants cespitose, few to several culms closely spaced along rhizomes 0.5-1 cm thick; culms (40-)60-100 cm

long, 2-5.5 mm wide at base (including sheaths); surface of culm and sheaths glabrous between ciliate angles

(culm sometimes puberulent between angles just below inflorescence); larger leaf blades 10-60 cm long,

(2.5-)3-7 mm wide, ciliate along margins and primary nerves, glabrous between nerves (sometimes pubes¬

cent with stiff trichomes between nerves on abaxial surface), lacking fine puberulent hairs; inflorescence of

terminal and axillary fascicles, the terminal fascicle of two closely-spaced clusters, the internode about 1 cm,

the proximal cluster smaller; axillary fascicle one (-two), comprised of a single cluster; internode between ter¬

minal and axillary fascicles (6-)18-48(-56) cm; primary bract of terminal fascicle 2-5 mm wide, the longer

cilia 1.1-2.2 mm; pistillate scales glabrous except for ciliate keel, the cilia 0.5-1 mm long; mature achenes in-

J. Bot. Res. Inst. Texas 9(1): 31 - 41.2015

Journal of the Botanical Research Institute of Texas 9(1)

eluding hypogynium 2.5-3.2(-3.4) mm long by 2.3-2.7 mm wide; achene body bony white, smoothish with

broad low rises, sometimes with faint transverse ridging, epapillate except for a few retrorse papillae at the very

base (these minutely setulose distally); hypogynium basal rim tan to light brown, 1.4-1.8(—2.1) mm wide,

0.1-0.2(-0.3) mm long; tubercles (4-)6.

Etymology .—The specific epithet honors C. Ritchie Bell (1921-2013), scientist, teacher, co-author of the

landmark Manual of the Vascular Flora of the Carolinas (Radford et al. 1968), and a founder and first director of

the North Carolina Botanical Garden.

Additional specimens examined (PARATYPES unless otherwise indicated), representing all known populations: CUBA. Santa Clara: 4 Sep

1903, Britton & Wilson 325 (NY!). MEXICO. Jalisco: abundante, 8 km al SE del Rancho El Mortero, municipio de Mezquitic, hab. bosque de

Pinus leiophyllay Quercusspp., 5 Nov 1963, Rzedowski 17703 (BRIT!). UNITED STATES. FLORIDA. Lake Co.: clay land, vicinity of Eustis,

1-15 Apr 1894, Nash s.n. (US!). Wakulla Co.: frequent in somewhat overgrown, but mowed area under powerlines along walking-bike trail

just S of hiway 98 and ca. 0.2 mi W of rte 363 (Woodville hiway), ca. 1 air mile NW of town of St. Marks, 18 May 1991, L.C. Anderson 13,398

(FSU!); frequent in moist, peaty sand with Stillingia aquatica (few low Ilex and Myrica ) in clearing under powerlines along walking-bike trail

just S of hiway 98 and W of rte 363 (Woodville hiway), ca. 1 air mile NNW of town of St. Marks, 3 Jun 1991, L.C. Anderson 13,434 (FSU!,

CLEMS). GEORGIA. Glynn Co.: replanted pine flatwoods 1.5 mi. w. of Altamaha River on co. rd. 620 to Everett City, 21 Jul 1966, Bozeman

6313 (NCU!). LOUISIANA. Calcasieu Par.: coastal prairie remnant dominated by Schizachyrium tenerum and S. scoparium, Rhynchospora

spp. conspicuous, Rudbeckia texana common, SW corner of Dennis Rd & Boudoin Rd., S of LA 108, E of LA 27,2.2 mi S of jet. w/I-10,14 May

2008, Reid et al. 6539 (LSU!). Jefferson Davis Par.: low prairies, Jennings, 15 May 1915, E.J. Palmer 7635 (MO!, US!). NORTH CAROLINA.

Granville Co.: Camp Butner, a collection made during a floristic study of Iredell soil in central Piedmont of N.C., 25 Aug 1951, Batson 543

(DUKE!, NY!). Onslow Co.: Sandy Run Swamp Powerline Savanna, Cooleys Meadowrue Powerline Survey Site, 4 Jun 1995, LeBlond 4254

(NCU!); formerly Haws Run Mitigation Site, now part of Maple Hill Savannas State Natural Area, near Onslow/Pender county line between

Sandy Run Swamp and Shelter Swamp Creek, 13 Jun 1996, LeBlond 6951, 17 June 1996, LeBlond 4572, 26 May 1998, LeBlond 4980, 20 Jul

1998, LeBlond 5023, 11 Jul 2000, LeBlond 5363, 14Jun 2003, LeBlond 5772 (TOPOTYPES: NCU!). Pender Co.: McLean Savanna, near south¬

western edge of Holly Shelter Game Land, about 0.7 mile north-northeast of confluence of Catskin Creek and Trumpeter Swamp, 18 Aug

1997, LeBlond &J. Taggart 4835 (NCU!); 20 Jun 2007, LeBlond 6420 et al. (NCU!). SOUTH CAROLINA. Berkeley Co.: Francis Marion Na-

LeBlond et al., A new species of Scleria

fs |i;l U

■%> >*» J ^

" 631814

fJCU bSlgiq-

United States of America

FLORA OF NORTH CAROLINA

County: Onslow

Scleria beIJii LeBlond Holotype

Sandy Run Swamp and Savannas Significant

Natural Heritage Area, in the portion formerly

knowm as 1 laws Run Mitigation Site, near

Onslow/Pendcr county line between Sandy Run

Swamp and Shelter Swamp Creek, off north side of

east/west trending road and ditch near cast side

access gate, lat. 34 l ’36'51"N, long. 77°37'54"W,

with Rli\nchospom tliornei. R. Uilifolia, Oenothera

fruticosa var. unguiculata, Aiuiropogon yjaucopsis,

Diehanthelium acuminatum var. acuminatum,

Symphyolrichum dumosum. SpiraiUlies vernalis.

R.J. LeBlond 695SL

6 J un 2013

Fig. 2. Holotype of Scleria bellii from Onlsow County, North Carolina {LeBlond 6958, NCU).

Journal of the Botanical Research Institute of Texas 9(1)

tional Forest, timber compartment 47, stand 03, with Sporobolus curtissii, Muhlenbergia capillaris, Ctenium aromaticum, Rhexia alijanus, R.

nashii, Helianthus angustifolius, Hypericum gymnanthum, Sabatia campanulata, Crotalaria purshiana, Eryngium yuccifolium, Bigelowia nudata

var. nudata, Scleria pauciflora var. pauciflora, 28 Jul 2012, L. Lee s.n. (NCU!); Francis Marion National Forest, in clumps on grassy, previ¬

ously scraped ground at savanna edge, east side of Forest Service Rd 212,640 m S of FSR 212F, 2.8 air mi E of downtown Homey Hill, 14 Sep

2012, Nelson 31381 (USCH!). Georgetown Co.: ca. 3 mi NE of Campfield along US 701, mown roadside through former pine flatwoods, with

Dichromena latifolia, Aster dumosus, Chaptalia tomentosa, Plantago sparsiflora, Bigelowia nudata, Rhexia lutea, R. alijanus, Buchnerafloridana,

Rhynchosporapusilla, R. pinetorum, 9 Jun 1994, Sorrie 8031 (USCH!). Jasper Co.: common in pine-toothache grass savannah along S-169,0.3

mile SW of jet. with U.S. 321,11 Aug 1983, Aulbach-Smith &Aulbach 2771 (USCH!). York Co.: Rock Hill Blackjacks Heritage Preserve, grassy

open spots near powerline right of way, edge of rocky woods over Iredell soil, just W of rock quarry on S-245, SE Rock Hill, 26 Jun 1987,

Nelson 5734 (USCH!, CLEMS). TEXAS. Harris Co.: 22 Jun 1877 Joors.n. (MO!); Houston, 1885, Nealley s.n. (NY!); a robust species among

prairie grasses along Underwood St. at intersection with North St. in LaPorte, 6 Aug 1983, Brown 6455 (BRIT!); sandy soil, coastal prairie W

of Underwood St., 0.3 mi. N of Spencer Hwy, 26 May 1984, Kessler & Brown 7467 (BRIT!, NY!).

DISCUSSION

Detailed morphometric analysis concentrated on the plants in eastern North Carolina; these were the only

known extant populations until discovery of the Berkeley Co., South Carolina, populations in 2012. Other

populations are known from one to a few herbarium specimens. Distinguishing characters established during

the analysis of the eastern North Carolina plants were applied to the vegetative and floral structures (including

at least one achene per sheet) of the populations represented only by herbarium specimens. Distinguishing

characters include the full range of measurements and qualities (e.g., indument density and location) gathered

from all specimens across the range of the species.

As with other Scleria species, the achene is critical in distinguishing S. bellii. Based on achene structure

(Fig. 3), it appears to be most closely related to taxa in the informal S. ciliata complex, but also showing some

morphological affinities with S. oligantha Michx. This treatment recognizes the following taxa in the S. ciliata

complex: S. ciliata Michaux var. ciliata, S. ciliata var. elliottii (Chapm.) Fernald, S. pauciflora Muhl. ex Willd.

var. pauciflora, and S. pauciflora var. caroliniana Alph. Wood. Other recent workers have recognized S. curtissii

Britton in J.K. Small and S. ciliata var. glabra (Chapm.) Fairey within the S. ciliata complex. They also are dis¬

cussed. Members of the S. ciliata complex together with S. bellii and S. oligantha are treated here as an informal

group based on shared features of the hypogynium. Scleria ciliata, “sensu lato,” is sometimes used for S. ciliata

var. ciliata and S. ciliata var. elliottii, including synonyms.

The need to include Scleria oligantha in this study gradually evolved as S. ciliata sensu lato collections

were examined. A few of the historical S. bellii collections had originally been identified as S. oligantha, and the

structure of the S. oligantha hypogynium bears a strong resemblance to those of S. bellii and the S. ciliata com¬

plex. Because S. oligantha is readily distinguished from the other members of the group, the focus of analysis

herein is on the morphological relationship of S. bellii with the more similar-appearing S. ciliata sensu lato.

Scleria oligantha herbarium collections need to be examined for the presence of S. bellii.

Scleria bellii, S. oligantha, and members of the S. ciliata complex share distinctive features of the achene

hypogynium, the structure that subtends the achene body in all Scleria species. This structure is rudimentary

in some species, and in others can remain attached to the pedicel after detachment of the achene body. In the

Scleria bellii-ciliata-oligantha group, the hypogynium consists of two parts visibile laterally: a broad and short

basal rim, and three to nine protuberances (tubercles) located between the basal rim and the achene body (Fig.

3). The basal rim and tubercles provide important diagnostic characters for separating the taxa in this group.

The rim is tan to dark brown and usually shiny. The tubercles are usually white or beige and surhcially mi¬

nutely papillose. Each papilla apex is rounded or pointed, with both apex types usually found on the same tu¬

bercle. The tubercle surface is similar to that of taxa in the S. triglomerata Michx. complex ( S.flaccida Steud., S.

minor W. Stone, S. nitida Willd.), which have an uninterrupted encircling hypogynium with a white papillose

surface, and lack a laterally visible basal rim.

Structure of the hypogynium tubercles in the Scleria bellii-ciliata-oligantha group is variable. Individual

tubercles can be isolated and distinct, or shallowly to deeply lobed, or confluent with an adjacent tubercle.

Confluent tubercles are indicated by a suture. Tubercle variation often occurs on the same achene. Size of indi¬

vidual tubercles is useful in distinguishing S. ciliata sensu lato from S. bellii and S. oligantha. The S. bellii

LeBlond et al., A new species of Scleria

Fig. 3. Representative achenes of the Scleria bellii-ciliata-oligantha group. Note differences in surface ornamentation and tubercle number/size on the

hypogynium. A) 5. bellii, B) 5. oligantha, C) 5. ciliata var. ciliata, D) 5. ciliata var. elliottii, and E) 5. pauciflora. Scale bar = 1 mm.

Journal of the Botanical Research Institute of Texas 9(1)

hypogynium is comprised of (four-) six tubercles, compared with three sometimes lobed tubercles of S. ciliata

sensu lato. Individual unlobed and nonconfluent tubercles in S. bellii average 0.3-0.5 mm wide, while those of

S. ciliata sensu lato are 0.5-0.7 mm wide.

The achene surface also provides important diagnostic features for the S. bellii-ciliata-oligantha group. In

all taxa, the mature achene is shiny and bony white (immature achenes are often darker and longitudinally

sulcate). The surface texture in the S. ciliata complex is variable, ranging from papillose to transversely ridged

to reticulate (including papillate ridging and reticulation). The mature achene body surface in S. oligantha is

uniformly smooth.

It should be noted there are two very different structures on the achene in the S. ciliata complex that are

described as papillate. The papillae on the achene body are readily visible at lOx and are about 0.1 mm in diam¬

eter or wider; their length can vary from <0.1 mm to >0.2 mm. By comparison, tubercle papillae are minute and

require stronger magnification to be seen.

It is achene body surface texture that most readily distinguishes S. bellii. Except for a few basal retrorse

papillae, the body is epapillate and essentially smooth. A closer examination (lOx) reveals broad low rises on

the body surface, including occasional faint low transverse ridging. These characteristics are comparatively

distinctive and occur throughout the range of the species. Achene dimensions, tubercle number, basal rim di¬

mensions, leaf and bract widths, and herbage pubescence clearly distinguish S. bellii.

The surface sculpturing of the Scleria bellii achene body is unique among North American Scleria, and

probably within Scleria as a whole. The few basally retrorse papillae and the character of the hypogynium,

along with vegetative characteristics, are likely what led to historical identification of most collections as S.

ciliata sensu lato (most frequently as S. elliottii or S. ciliata var. elliottii). Four collections were originally identi¬

fied as S. oligantha.

Achene and hypogynium measurements alone can differentiate fairly well between S. bellii, S. ciliata var.

ciliata, and S. ciliata var. elliottii. The Scleria bellii hypogynium basal rim is usually broader than that of S. ciliata

sensu lato. Achene width, although strongly correlated with hypogynium width (R = 0.81), is also generally

greater in S. bellii than in S. ciliata sensu lato (Fig. 4). Achene and hypogynium lengths in S. bellii overlap with

measurements of both varieties of S. ciliata, although the S. bellii lengths have a narrower range. Kruskall-Wal-

lis tests for one-way analysis of variance show that achene length, achene width, hypogynium length, and hy¬

pogynium width are each significantly different among the three taxa (a = 0.003 with Bonferroni correction),

although pairwise comparisons indicate that S. belli achene length and hypogynium length are not significant¬

ly different from S. ciliata var. elliottii and S. ciliata var. ciliata, respectively (Table 1). Of these metrics, achene

width and hypogynium width are the strongest predictors, showing clear distinctions among the three taxa.

Special mention must be made of the collection of Scleria bellii from Jalisco, Mexico. Only one specimen

has been seen ( Rzedowski 17703, BRIT), and it contains a single achene. That achene has the overall dimen¬

sions, basal rim dimensions, and the six tubercles of typical mature S. bellii achenes, but lacks the body’s basal

papillae. The tubercles are also more distinctly separated than in typical specimens. The body is longitudinally

sulcate, suggesting the achene may be immature. Culm and leaf dimensions and indument are the same as in

typical S. bellii. More specimens need to be examined from this disjunct population.

The Scleria bellii inflorescence usually produces a terminal fascicle and one axillary fascicle (rarely two

axillary, and rarely no axillary on what are likely late season culms). Scleria ciliata vars. ciliata and elliottii fre¬

quently produce culms with only terminal fascicles, as well as culms with both a terminal and one axillary

fascicle. Terminal primary bracts are on average wider in S. bellii, and their cilia longer, as are the cilia of pistal-

late scale keels. Pistillate scales are glabrous in S. bellii, and glabrous to puberulent in S. ciliata sensu lato.

Vegetatively, Scleria bellii most closely resemble S. ciliata var. elliottii. On average, S. bellii culms are taller

and the leaves wider. There are taxonomically important differences in culm and leaf indument. The S. bellii

culm has stiff trichomes on the angles and primary veins, but is glabrous between the veins and angles. (The

culm is sometimes puberulent between the angles and veins for 1-2 cm below the terminal fascicle.) The S.

ciliata var. elliottii culm is puberulent throughout with hairs 0.1-0.4 mm long and often has stiff trichomes on

LeBlond et al., A new species of Scleria

c\i

o S. bellii

a S. ciliata var. ciliata

+ S. ciliata var. elliottii

c ^

>> i-

£ CM

n -

1.8

2.0

2.2

2.4

2.6

achene width (mm)

Fig. 4. Achene and hypogynium width variation in 5. bellii 5. ciliata var. ciliata , and 5. ciliata var. elliottii.

the angles and primary veins. Similar to the culm, the S. bellii leaf adaxial surface has stiff trichomes on the

margins and primary veins, and is glabrous between the veins. In S. ciliata var. elliottii, the leaf adaxial surface

is puberulent throughout with hairs 0.1-0.4 mm long, and often has stiff trichomes along the margins and

primary veins.

Scleria oligantha readily separates from S. bellii and S. ciliata sensu lato by its eight or nine hypogynium

tubercles, and achenes 2.5-4 mm long with uniformly smooth surfaces. The inflorescence is either a terminal

fascicle, or terminal with one or two axillary fascicles. The axillary fascicles are borne on long filiform pedun¬

cles. The axillary fascicles in S. bellii and S. ciliata sensu lato are short to long-peduncled.

Plants in the Scleria ciliata complex with reticulate achenes have been recognized as S. curtissii, S. ciliata

var. curtissii (Britton) J.W. Kessler, andS. pauciflora var. curtissii (Britton) Fairey (Small 1903; Fairey 1967; Kes¬

sler 1987). Sculpturing of the achene surface is quite variable in the complex, ranging from papillose to trans¬

versely ridged to reticulate (including papillate ridging and reticulation). Reticulation and transverse ridging

sometimes occur in different areas on the same achene. Reticulate body surfaces can be found on achenes with

3 (S. ciliata) or 6 (S. pauciflora) tubercles, corresponding with the two varieties. Such variation argues against

recognition of curtissii as distinct.

Journal of the Botanical Research Institute of Texas 9(1)

Table 1 . Kruskall-Wallis chi-squared values from analysis of variance, p-values < 0.005 are indicated in bold.

Achene

Length

Achene

Width

Hypogynium

Length

Hypogynium

Width

Pairwise

5. bellii/S. ciliata var. cilioto

19.81

32.16

0.0261

33.87

5. ciliata var. ciliata/S. ciliata var. elliottii

16.03

9.46

15.58

5. ciliata var. elliottii/S. bellii

1.57

24.81

10.81

31.38

All three taxa

24.97

45.69

13.87

50.67

Essentially glabrous Scleria ciliata plants have been recognized as Scleria ciliata var. glabra. In his proto-

logue, Fairey (1967) describes the plants as “completely glabrous.” An examination of specimens found that

most plants identified as var. glabra have glabrous to glabrate culms and leaf blades. But sheaths, especially the

lower, are puberulent, and terminal fascicle primary bracts are usually ciliate or scabrous-margined. There are

many specimens identified as either var. glabra, var. ciliata, or S. ciliata sensu lato with few to some leaf margin

and/or culm angle cilia. Thus, varietal assignment seems arbitrary. Glabrate plants are essentially restricted to

the outer Coastal Plain, where they are sympatric with the more widespread ciliated plants. This difference in

indument maybe a response to environmental conditions, and in our opinion does not merit taxonomic recog¬

nition.

Herein, no attempt has been made to determine the validity of characters distinguishing Scleria pauciflora

sensu lato within the S. ciliata complex. That work needs to be done. It is possible characters used here for

other taxa in the group, including those of the hypogynium and herbage indument, may be of use. There were

several specimens on which the hypogynium tubercle count and achene size were ambiguous.

Putative Origin. —The shared morphological features with members of the S. ciliata-oligantha group, espe¬

cially in the achene, strongly suggest an evolutionary relationship. Regarding a hybrid origin, S. ciliata var. el-

liottii and S. oligantha appear to be the most likely candidates based on morphology, but several factors argue

against this. So far, S. bellii has not been found with either of these taxa. Scleria ciliata var. elliottii and S. oligan¬

tha are sympatric over an extensive region, but S. bellii is restricted to a small portion of that region. Scleria

bellii also occurs in the West Indies (Cuba), where S. oligantha is absent. Among the three, S. bellii has longer

culms, wider leaves, and longer pubescence. Both S. ciliata var. elliottii and S. oligantha are often pubescent be¬

tween the major veins on the leaf surface, while S. bellii is glabrous (rarely a few stiff trichomes between the

veins abaxially). The absence of S. ciliata var. elliottii and S. oligantha —and sometimes all other Scleria —from

known S. bellii sites strongly suggests an independently reproducing taxon.

The following key distinguishes the taxa in the S. bellii-ciliata-oligantha group. Within its range, the group

is distinguished from all other Scleria by a hypogynium consisting of a basal rim subtending a series of three to

nine tubercles. Characters used to distinguish S. bellii from S. ciliata varieties ciliata and elliottii are also shown

in Table 2.

1. Tubercles 8 or 9; mature achene 2.5-4 mm long, the surface uniformly smooth; inflorescence axillary fascicles, when

present, on elongate filiform peduncles_ S. oligantha

1. Tubercles 3 or 6; mature achene 1 -3.5 mm long, the surface either smoothish with broad low rises, or variously dense¬

ly papillose, transversely ridged, or reticulate; when present, inflorescence axillary fascicles short to long-peduncled.

2. Hypogynium with (5—)6 tubercles; achene 1 -2.5 mm long, the body papillate or reticulate-roughened; leaves 1 -2.5

mm wide.

3. Plants glabrous to hairy, but not villous-ciliate, the hairs to 0.4 m m long_ S. pauciflora var. pauciflora

3. Plants villous-ciliate with spreading hairs 0.5-1 mm long_ S. pauciflora var. caroliniana

2. Hypogynium with 3 to 6 tubercles; if more than 3 tubercles, then achene 2.5-3.2(-3.4) mm long, the body smooth¬

ish with low broad rises, and leaves (2.5-)3-7 mm wide.

4. Achene body smoothish with broad low rises, epapillate except for a few retrorse papillae at the very base,

sometimes with faint transverse ridging; achene including hypogynium 2.5-3.2(-3.4) mm long, 2.3-27 mm wide;

hypogynium basal rim 1.4—1.8(—2.1) mm wide, 0.1-0.2(-0.3) mm long; tubercles (4-)6, individual unlobed

tubercles 0.3-0.5 mm wide; longer pistillate scale keel cilia 0.5-1.0 mm, the scale surface glabrous; terminal

LeBlond et al., A new species of Scleria

Table 2. Key characters used to distinguish Scleria bellii, 5. ciliata var. ciliata, and 5. ciliata var. elliottii.

characters S. bellii S. ciliata var. elliottii S. ciliata var. ciliata

achene body surface

smoothish with broad low rises,

epapillate except at base,

sometimes with a faint

transverse ridge

full achene length (mm)

2.5-3.2(-3.4)

achene body width (mm)

23-2.7

hypogynium basal

1.4-1.8(-2.1)

rim width (mm)

hypogynium basal

0.1-0.2(-0.3)

rim length (mm)

tubercle number

6 (rarely 3 deeply lobed)

tubercle width (mm)

0.3-0.5

longer pistillate scale

0.5-1.0

keel cilia (mm)

maximum width primary

2.0-4.9

terminal bract (mm)

longer cilia on primary

1.1-2.2

terminal bract (mm)

axillary inflorescences

1(-2)

leaf blade width (mm)

(2.5-J3-7

blade margins and

ciliate with stiff trichomes

primary veins

leaf surface

glabrous between primary

veins (rarely a few

stiff trichomes)

culm height (cm)

40-100

papillose throughout,

sometimes also/or transversely

ridged and/or reticulate

2.3-3.1 (-3.3)

2.0-2.5(-2.6)

1.1 —1.4(—1.6)

papillose throughout,

sometimes also/or

transversely ridged

and/or reticulate

2.0-3.2(-3.5)

1.7-2.3(-2.5)

(0.6-)0.8-1.3(-1.5)

(0.1-)0.2-0.4(-0.5)

0.1-0.2(-0.3)

3 (sometimes deeply lobed)

0.5-0.7

(0.1-)0.2-0.5

3 (sometimes deeply lobed)

0.5-0.7

(0-)0.1-0.5

1.2-4.1

0.4-2.2

0.7-1.1 (-1.3)

(0-)0.1-0.7(-0.9)

0-1

(2.5-)3-6 mm

ciliate with stiff trichomes,

or puberulent

puberulent at least adaxially

0-1

1 —3(—3.5)

eciliate, ciliate, or puberulent

glabrous or puberulent

(15-)30-70

(15-)25-70(-75)

fascicle primary bract 2.0-4.9 mm wide at widest point, its longer cilia 1.1 -2.2 mm; inflorescence a terminal and

(0—)1 (—2) axillary fascicles; leaf blades (2.5-)3-7 mm wide, with stiff hairs on margins and primary veins, rarely

with a few stiff hairs between veins on abaxial surface, lacking fine puberulent hairs; culm height 40-100 cm

_S. bellii

4. Achene body papillose throughout, sometimes also transversely ridged or reticulate (infrequently papillae absent

in reticulate achenes); achene including hypogynium 2.0-3.3(-3.5) mm long, 1.7-2.4(-2.6) mm wide; hypogy-

nium basal rim (0.6-)0.8-1.4(-1.6) mm wide, 0.1-0.4(-0.5) mm long; tubercles 3, entire to deeply lobed, the

unlobed tubercles 0.5-0.7(-0.9) mm wide; longer pistillate scale keel cilia 0-0.5 mm, the scale surface glabrous

or puberulent; terminal fascicle primary bract 0.4-4.1 mm wide at widest point, its longer cilia (0-)0.1 -1.1 (-1.3)

mm; inflorescence a terminal and 0-1 axillary fascicles; leaf blades 1-6 mm wide, glabrous to ciliate on margins

and primary veins, to puberulent with fine hairs over entire surface, especially adaxially; culm height 15-75 cm.

5. Leaf blades (2.5-)3-6 mm wide, at least some adaxial surfaces puberulent throughout, often with stiff hairs

on margins and primary veins; terminal fascicle primary bract 1.2-4.1 mm wide at widest point, its longer

cilia 0.7-1.1 (-1.3) mm; hypogynium basal rim 1.1 -1.4(-1.6) mm wide, (0.1 -)0.2-0.4(-0.5) mm long_ S. ciliata

var. elliottii

5. Leaf blades 1 -3(-3.5) mm wide, adaxial surface glabrous and eciliate, or glabrous and ciliate on margins, or

puberulent and ciliate; terminal fascicle primary bract 0.4-2.2 mm wide at widest point, its longer cilia (0-)0.1 -

0.7(-0.9) mm long; hypogynium basal rim (0.6-)0.8-1.3(-1.5) mm wide, 0.1 -0.2(-0.3) mm long_ S. ciliata var. ciliata

DISTRIBUTION AND HABITAT

Although widespread, S. bellii populations are scattered to disjunct (Fig. 1). The population in Jalisco, Mexico,

is about 1,300 kilometers (800 miles) from the next nearest population, in Harris Co., Texas. The populations

in western Louisiana are about 800 kilometers (500 miles) disjunct from the population in Wakulla Co., Flor¬

ida. In the U.S., 12 of the 14 known populations are on or near the outer portion of the Atlantic and Gulf

Coastal Plain. It is possible the species was more common when sea level was lower, with a rising sea segment¬

ing distribution. The other two U.S. populations are in the Piedmont of North and South Carolina. This re-

Journal of the Botanical Research Institute of Texas 9(1)

stricted distribution, along with what is known of soil characteristics, suggest adaptation to restricted habitats

(though they may have been more common historically). Scleria bellii is a species of conservation concern, and

is ranked as imperiled globally (G1G2) by NatureServe (2015).

All of the known Coastal Plain sites occur on wet soils, primarily if not exclusively in pine savannas and

coastal prairies. At the three North Carolina Coastal Plain sites where the soil properties are known, there is a

calcareous influence even though the pH is usually acidic. These sites are fire-adapted wet acid pine and pine-

cypress savannas characterized by a subterranean limestone layer usually too deep to raise pH above acidic

levels, but close enough to contribute nutrients (LeBlond et al. 1994; LeBlond 2001). This natural community

has been described as the Very Wet Loamy Pine Savanna (Schafale 2011). According to Schafale, “These com¬

munities are very rare with a limited and patchy geographic range. They often, maybe always, have inclusions

where soils are high in calcium and have a higher pH (5.5 to 7.2), but the majority of their soil is similar to

other pine savannas (pH 3.8-4.1).”

Associated species at the Berkeley Co., Georgetown Co., andjasper Co. sites in South Carolina indicate a

fire-adapted rich wet savanna habitat. Associates at the Georgetown Co. site include Dichromena latifolia, Plan-

tago sparsiflora, and Rhynchospora pinetorum, species suggesting a calcareous influence.

The Wakulla Co., Florida, occurrence “could be described as a calcareously-influenced wet acid savanna

or coastal prairie.. .Limestone is at or near the surface” (Anderson 2013). The two sites in Louisiana are de¬

scribed as coastal prairie ( Reid 6539, LSU) or low prairie ( Palmer 7635, MO, US). One of the collections from

Harris Co., Texas, ( Kessler & Brown 3619, BRIT, NY) refers to coastal prairie. The Reid collection from Calca¬

sieu Parish, Louisiana, cites Schizachyrium tenerum and S. scoparium as dominants, suggesting that that site

might be an occurrence of the West Gulf Coastal Plain Southern Calcareous Prairie (NatureServe 2015).

The two Piedmont sites, one each in North and South Carolina, occur on acid to neutral soils with a high

ferro-magnesium content. Although the moisture regime at the Piedmont microsites is not known, the associ¬

ated soil series is moderately well drained.

Habitat data for other sites are ambiguous, with “Clay land” for the Lake Co. site in Florida and “Replant¬

ed pine flatwoods” in Glynn Co., Georgia. There are no data for the site in west-central Cuba. The label for the

collection from Jalisco, Mexico, describes the habitat as a Pinus leiophylla and Quercus spp. forest, which may

be fire-adapted. It is not known whether the microsite is wet, mesic, or dry.

ACKNOWLEDGMENTS

We are grateful to Carol Ann McCormick, curator of the UNC-CH Herbarium (NCU), for managing the exten¬

sive gathering of collections. We are also indebted to the curators and staff of the many herbaria who sent those

collections. Support and insight were also provided by colleagues Alan Weakley, Bruce Sorrie, and Brenda

Wichmann. Appreciation is given to DavidJ. Rosen and several other reviewers for their constructive contribu¬

tions to the manuscript.

In this era of declining support for herbaria, at least in the U.S., it is important to note the critical role

herbaria played in our understanding of the distribution, ecology, morphology, and phenology of this new spe¬

cies. These data are critical in guiding research and population protection efforts.

REFERENCES

Anderson, L.C. 2013. Personal communication: e-mail of April 15.

Fairey, J.E., III. 1967. The genus Scleria in the southeastern United States. Castanea 32:37-71.

Kessler, J.W. 1987. A treatment of Scleria (Cyperaceae) for North America north of Mexico. Sida 12:391-407.

LeBlond, R.J. 2001. Endemic plants of the Cape Fear Arch region. Castanea 66:83-97.

LeBlond, R.J., A.S. Weakley, A.A. Reznicek, & W.J. Crins. 1994. Carexlutea (Cyperaceae), a rare new Coastal Plain endemic from

North Carolina. Sida 16:153-161.

NatureServe. 2015. NatureServe Explorer. An online encyclopedia of life [web application]. Arlington, Virginia, U.S.A.

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.

Journal of the Botanical Research Institute of Texas 9(1)

BOOK REVIEW

Emmet J.Judziewicz, Robert W. Freckmann, Lynn G. Clark, & Merel R. Black. 2014. Field Guide to Wisconsin

Grasses. (ISBN-978-0-299-30134-7, pbk). The University of Wisconsin Press, Madison. (Orders: http://

uwpress.wisc.edu/order.html#individuals, 1-800-621-2736). $29.95 (eBook $24.95), 288 pp., 6" x 9".

This slender but information- and illustration-rich book is a winner. It is based on many years of effort and

experience by a group of authors highly knowledgeable about grasses.

The volume covers 232 species of grasses in Wisconsin. With over 1,000 figures, it is no wonder that most

species include several illustrations or photos, which typically include habitat shots, close-ups of spikelets and

their constituent parts, inflorescences, and photos of growth habit. A tremendous amount of time, effort, and

care clearly went into obtaining and selecting the illustrations and photos.

Most species are allotted one page, but others (e.g., Elymus lanceolatus, Calamagrostis canadensis) receive

two. A county-wide distribution map for Wisconsin accompanies each species at the top. Beyond the treat¬

ments for species, the book includes chapters on Morphology, Agrostology (more generally and including

prominent agrostologists of Wisconsin), Grasses in Wisconsin Plant Communities, and Keys to the Grass Genera

of Wisconsin. Wrapping up the volume is the Glossary, References, Illustration Credits, and Taxonomic Index.

For the tax-paying public, from whom state-supported universities ultimately derive most of their fund¬

ing, a volume like Field Guide to Wisconsin Grasses resonates more than the research that systematists carry out

(myself included) on taxa from far-distant lands. Botanists in the state-supported schools should remember

this reality and continue to publish relevant, useful information regionally.

The generally broad overlap in the distribution of grass species with Minnesota, Illinois, and Michigan

suggests that this volume could be used productively in those states as well. Field Guide to Wisconsin Grasses,

given its compact size, reasonable pricing, and quality of content, is an excellent model for future treatments.—

Neil Snow, PhD, T.M. Sperry Herbarium, Department of Biology, Pittsburg State University, Pittsburg, Kansas

66762, U.S.A.

J. Bot. Res. Inst. Texas 9(1): 42.2015

STYRAX PELTATUS (STYRACACEAE), A NEW SPECIES FROM OAXACA, MEXICO

Peter W. Fritsch

Department of Botany

California Academy of Sciences

55 Music Concourse Drive

San Francisco, California 94118-4503, U.S.A.

pfritsch@calacademy. org

ABSTRACT

A new species of Styrax T. series Valvatae Perkins, Styrax peltatus from the Isthmus of Tehuantepec in southeastern Mexico, is described

and illustrated. It is similar to the Mesoamerican species S. conterminus Donn. Sm., but differs in its shorter petioles, smaller chartaceous and

nearly glabrous leaf blades with six to eight secondary veins on each side of the midvein, the presence of leaf domatia in the abaxial axils of

the secondary veins, fewer-flowered inflorescences with more delicate rachises, and generally smaller flowers. The species is known only

from the type collection, made from the Uxpanapa-Chimalapa Region of the Selva Zoque in far eastern Oaxaca.

Key Words: Endemic, Isthmus of Tehuantepec, IUCN Red Tist, Mexico, new species, Oaxaca, Selva Zoque, Styrax, Uxpanapa-Chimalapa

Region

RESUMEN

Se describe e ilustra una especie nueva de Styrax L. series Valvatae Perkins, Styrax peltatus del istmo de Tehuantepec en el sureste de

Mexico. Es similar a la especie Mesoamericana S. conterminus Donn. Sm., pero difiere es sus peciolos mas cortos, laminas foliares mas

pequenas cartaceas y casi glabras con seis a diez venas secundarias a cada lado del nervio principal, la presencia de domacios en las axilas

abaxiales de las venas secundarias, inflorescencias con menos flores y raquis mas delicados, y flores generalmente mas pequenas. La especie

solo se conoce de la coleccion tipo, hecha en la region Uxpanapa-Chimalapa de la Selva Zoque en el extremo este de Oaxaca.

Styrax series Valvatae Perkins, with about 90 species, is endemic to the Neotropics and contains most of the

Neotropical species of Styrax. The species of this clade are distinguished from other members of the genus by

the combination of an evergreen habit, a fleshy to juicy indehiscent mesocarp, and an ellipsoid seed (Wallnofer

1997; Fritsch 1999, 2001). Other distinguishing features are: bases of young shoots without stalked ferrugine-

ous or fulvous stellate trichomes, unless these accompanied by a dense tomentum of trichomes of the same

general color and type; a subcoriaceous corolla with valvate floral aestivation and straight sides; and a smooth

(i.e., “non-crackled”) seed coat (Fritsch 1999).

The species taxonomy of Styrax for western Texas, Mexico, and Mesoamerica, including that of the spe¬

cies of S. series Valvatae in these areas, has been recently revised (Fritsch 1997). After the revision, I examined

a specimen of S. series Valvatae collected from the Uxpanapa-Chimalapa Region of southwestern Mexico that

does not fit the key to species in Fritsch (1997) and whose combination of characters agrees with no other spe¬

cies in the genus. This species is described and illustrated here as new to science.

Styrax peltatus P.W. Fritsch, sp. nov. (Figs. 1-2). Type: MEXICO. Oaxaca. Mpio. San Miguel Chimalapa: Loma Larga, en la

cabecera de la rama principal (S) del Rio del Cafe al O de Cerro Guayabitas, ca. 40-45 km al N de San Pedro Tapanatepec, 16°44'N,

94°14'W, 1500 m, 6 Nov 1985 (fl), Salomon MayaJ. 2450 (holotype: MEXU not seen, digital image!; isotype: CAS!).

Haec species Styrax conterminus Donn. Sm. simillima, sed ab eo petiolis 7-10 mm long, laminis 7.7-9.8 x 2.1-2.5 cm chartaceis, paginae

abaxialium laminae fere glabris, nervis laminae utroque costae latere 6-8, domatiis praesantibus, inflorescentiis 4-8-floris compositis,

rhachidibus inflorescentiae 0.6-1 mm latis, floribus ubique parvioribus, calycibus 2.4-3 x 3.3-4 mm, squamis ad medium calycis 0.2-0.3

mm diametro differt.

Trees, evergreen, 8 m tall. Young branchlets and vegetative buds densely covered with overlapping light ferru-

gineous to tawny lacerate-margined peltate scales. Petioles 7-10 mm long; leaf blades narrowly oblong, 7.7-9.8

x 2.1-2.5 cm, 2.9-3.9 times as long as wide, chartaceous, adaxially olive brown and glabrous except for sparse¬

ly scattered lacerate-margined peltate scales on midvein, abaxially olive green and nearly glabrous except for

J. Bot. Res. Inst. Texas 9(1): 43 - 47.2015

Journal of the Botanical Research Institute of Texas 9(1)

Fig. 1 . Styraxpeltatus. A. Branchlet with flower buds. B. Leaf, abaxial side. C. Close-up of abaxial side of leaf blade at base, showing lacerate-margined

peltate scales and (shaded) one domatium. D. Pedicel and flower bud. E. Flower. F. Stamen, ventral view. Two adjacent stamens are incompletely

drawn. G. Stamen, lateral view. H. Gynoecium and calyx, the latter cut away in front to show ovary. All drawn from the isotype 5. MayaJ. 2450 (CAS).

Fritsch, A new species of Styrax from Oaxaca, Mexico

Fig. 2. Styraxpeltatus, abaxial side of leaf blade. The leaf is oriented such that the proximal-to-distal direction is lower left to upper right. A. Midvein

near base, showing lacerate-margined peltate scales. B. Surface, showing marsupiform domatium at the junction of the midvein and a secondary vein.

Scale bars: A, 0.2 mm; B, 0.5 mm. From the isotype 5. MayaJ. 2450 (CAS).

scattered lacerate-margined peltate scales on midvein and rarely on blade surface, scales 0.2-0.3 mm diam.;

midvein raised abaxially, slightly depressed adaxially, secondary veins 6 to 8 on each side of midvein, often

strongly upcurved near margin, tawny yellow, raised on both sides, tertiary veins generally perpendicular to

midvein, slightly raised on both sides; base broadly cuneate to subrounded, margin entire, slightly revolute,

apex abruptly acuminate; leaf domatia present abaxially in axils of secondary veins, marsupiform, deltoid, side

along the midvein 1.6-2.2 mm long, glabrous or with sparsely scattered lacerate-margined peltate scales. Inflo¬

rescences axillary, racemose or paniculate, 2.5-4 cm long, 4- to 8-flowered, rachis 0.6-1 mm wide, rachis and

pedicels covered with dense lacerate-margined peltate scales. Flowers hermaphroditic, ca. 10 mm long. Pedi¬

cels 5.5-7 mm long. Calyx obconic, 2.4-3 x 3.3-3.9 mm, densely covered with olive green to scattered pale

orange peltate or lacerate-margined scales, margin truncate between the 5 minute teeth, margin toward inner

(adaxial) side glandular, at mid-calyx the scales 0.2-0.3 mm diam., with 30-45 radiate arms. Corolla white, ca.

9 mm long, petals connate ca. 3 mm beyond the calyx margin; lobes 5, valvate in bud, thickened, ca. 6.5 x 1.2

mm, linear, abaxially covered with lacerate-margined peltate scales. Stamens 10; stamen tube diverging from

corolla at ca. 2.5 mm, free portion ca. 3 mm long, dorsally covered with lacerate-margined peltate scales; dis¬

tinct portion of filaments ca. 0.5-0.8 mm long, of equal width throughout, ventrally without auricles but bear¬

ing mixed radiate and lacerate-margined peltate scales at the somewhat incurved filament margins, more

sparsely so on the inner surface, scales with arms to 0.18-0.24 mm long, the arms radiating from the central

point in various directions, not primarily oriented parallel to the filament; anthers ca. 2.8-3.2 mm long, con¬

nectives glabrous; thecae linear, non-tapered, slightly to distinctly exceeding connective, with very sparsely

scattered radiate scales on margins. Free portion of ovary rounded-pyramidal, densely covered with greenish

white lacerate-margined peltate scales. Style filiform, ca. 7 mm long, glabrous; stigma ca. 0.3 mm wide. Fruit

unknown.

Distribution, Habitat, and Phenology .—The new species is apparently endemic to the Atlantic slope rain

forest of the Uxpanapa-Chimalapa Region, part of the Isthmus of Tehuantepec in southeastern Mexico. This

region is in the Selva Zoque zone, known for its high species endemism (Wendt 1987, 1993). The type was

collected in the Chimalapa part of the region, in far eastern Oaxaca near the border with Chiapas. It has

been found only in the headwaters of the principal (south) branch of the Rio del Cafe to the west of Cerro

Journal of the Botanical Research Institute of Texas 9(1)

Guayabitas, growing in pine-oak forest and in canyons in bosque mesohlo de montana (montane mesophyll

forest) of Cedrela P. Browne, Liquidambar L., and Oecopetcdum Greenm. & C.H. Thomps. at 1500 m elevation.

The only known collection was flowering in November.

Etymology. —The epithet refers to the peltate scales of this species, which occur nearly throughout the

external surfaces of the plant, including the branchlets and vegetative buds, leaves, and inflorescences.

Conservation Status. —Because Styrax peltatus is known only from the type collection, we categorize this

species as Data Deficient (DD; IUCN 2014). The label information denotes the species as abundant at the time

of collection.

The morphological characters of Styrax peltatus are consistent with its placement among a phylogenetic

grade of species within S. series Valvatae. These species all possess exclusively bisexual flowers (versus female

and bisexual, i.e., plants gynodioecious), stamen filaments that are smooth ventrally (versus auriculate), tri-

chomes with arms that radiate from the central point in various directions and typically do not exceed 0.5 mm

long (versus primarily oriented parallel to the filament and are generally 0.8-2 mm long), and stamen thecae

that are not tapered apically (versus tapered) and that equal or slightly exceed the connective (versus do not

exceed the connective). The ten other species that comprise this grade (S. austromexicanus P.W. Fritsch, S. con-

terminus Donn. Sm., S. gentryi P.W. Fritsch, S. incarnatus RW. Fritsch, S. lanceolatus P.W. Fritsch, S. magnus

Lundell, S. ochraceus Urb., S. radians P.W. Fritsch, S. ramirezii Greenm., and S. warscewiczii Perkins) are to¬

gether distributed in Mexico, Mesoamerica, and the Antilles (Fritsch 1999).

In the key to the species of Styrax of Mexico and Mesoamerica in Fritsch (1997), the first lead of couplet 4

(page 712) states “Abaxial laminar surface (and calyx) with vesture of peltate or lacerate-margined scales;

scales at mid-calyx 0.72-0.44 mm diam.” and the other lead of the couplet states “Abaxial laminar surface with

vesture of stellate hairs, or rarely radiate scales or nearly glabrous; scales at mid-calyx 0.05-0.33 mm diam., or

else lacking.” Styrax peltatus does not key out in this key because its calyx trichomes are lacerate-margined

scales with a diameter at mid-calyx of 0.2-0.3 mm. If one followed the first lead, the new species would key to

S. conterminus because of the shared characters of non-auriculate stamen filaments (versus strongly auriculate

in S. peruvianus Zahlbr., the terminus of the alternative lead in couplet 5), leaves 2.8-2.9 times as long as wide

(versus 2.0-3.1 times), and non-tapered anther thecae (versus tapered). Styrax conterminus is a Mesoamerican

species of pine-oak and broad-leaved cloud forests that ranges from extreme southeastern Chiapas through

Guatemala to northern El Salvador and western Honduras (Fritsch 1997). In addition to the differences in tri-

chome diameter and geographic distribution, the new species is distinguishable from S. conterminus by the

following characters: petioles 7-10 mm long (versus 13-25 mm); leaf blades 7.7-9.8 x 2.1-2.5 cm (versus 8.5-

16.5 x 2.7-5.0 cm), chartaceous (versus thick-chartaceous), nearly glabrous on the abaxial surface (versus

densely lepidote), secondary veins 6 to 8 on each side of midvein (versus 9 to 13), domatia present (versus ab¬

sent); inflorescences 4- to 8-flowered (versus 4- to 20-flowered) with rachises 0.6-1 mm wide (versus 1.2-1.3

mm); calyces 2.4-3 x 3.3-4 mm (versus 3-4 x 4-5.5 mm); and generally smaller flowers.

Styrax peltatus could potentially be confused with S. ramirezii Greenm., a species also known to occur in

the Uxpanapa-Chimalapa region (e.g., S. MayaJ. 3102, CAS), because of the leaf and calyx vestiture in S. ramire¬

zii that occasionally consists of scales. In this region, however, the abaxial leaf surfaces in plants of S. ramirezii

are consistently covered by at least a thin tomentum, whereas in those of S. peltatus this surface is nearly gla¬

brous. The leaves of the latter are also generally smaller, and the domatia are much larger (domatia are usually

entirely lacking in S. ramirezii). Moreover, in S. ramirezii the calyx trichomes (and often those of the branchlets

and leaves) are either stellate hairs, or radiate scales with the distal part of the arms often distinct, whereas in

S. peltatus they are lacerate-margined peltate scales.

The only other species of Styrax from Mexico (and Mesoamerica) to be described after the revision of

Fritsch (1997) is S. uxpanapensis P.W. Fritsch, also apparently endemic to the Uxpanapa-Chimalapa Region. It

is easily distinguished from S. peltatus by a vestiture of stellate trichomes (versus scales) on most parts of the

plant (e.g., branchlet, vegetative bud, leaf, inflorescence rachis, and calyx), and also by larger leaves 13-21.5 x

5.4-8.2 cm (versus 7.7-9.8 x 2.1-2.5 cm) and prominently auriculate stamen filaments (versus non-auriculate).

Fritsch, A new species of Styrax from Oaxaca, Mexico

Styrax uxpanapensis is also distributed farther north than S. peltatus, i.e., in Oaxaca and Veracruz near the

border shared between these states, and occurs at lower elevations (225-250 m versus 1500 m; Fritsch 2005).

ACKNOWLEDGMENTS

I thank Fernando Chiang, Alberto Reyes, and Alfredo Ortiz for help in finding and imaging the holotype at

MEXU, and Tom Wendt for helping me access the digital image; T.W. and Ross C. Clark for helpful comments

on the original manuscript; Boni Cruz for assistance with leaf imaging; and Alan Chou for the illustration.

REFERENCES

Fritsch, P.W. 1997. A revision of Styrax (Styracaceae) for western Texas, Mexico, and Mesoamerica. Ann. Missouri Bot.

Gard. 84:705-761.

Fritsch, P.W. 1999. Phylogeny of Styrax based on morphological characters, with implications for biogeography and

infrageneric classification. Syst. Bot. 24:355-378.

Fritsch, P.W. 2001. Phylogeny and biogeography of the flowering plant genus Styrax (Styracaceae) based on chloro-

plast DNA restriction sites and DNA sequences of the internal transcribed spacer region. Molec. Phylogen. Evol.

19:387-408.

Fritsch, P.W. 2005. A new species of Styrax (Styracaceae) from southern Mexico. Novon 15:421-424.

IUCN Standards and Petitions Subcommittee. 2014. Guidelines for using the IUCN Red List categories and criteria, Version

11. Prepared by the Standards and Petitions Subcommittee. Available at www.iucnredlist.org/documents/RedList-

Guidelines.pdf. Accessed November 2014.

WallnOfer, B. 1997. A revision of Styrax L. section Pamphilia (Mart, ex A.DC.) B. Walln. (Styracaceae). Ann. Naturhist. Mus.

Wien, B, 99:681-720.

Wendt, T. 1987. Las selvas de Uxpanapa, Veracruz-Oaxaca, Mexico: Evidencia de refugios floristicos cenozoicos. Anales

Inst. Biol. UNAM 58 (1987), Ser. Bot. 1:29-54.

Wendt, T. 1993. Composition, floristic affinities, and origins of the canopy tree flora of the Mexican Atlantic Slope rain

forests. ln:T.P. Ramamoorthy, R. Bye, A. Lot, & J. Fa, eds. Biological diversity of Mexico: Origins and distribution. Oxford

Univ. Press, New York, U.S.A. Pp. 595-680.

Journal of the Botanical Research Institute of Texas 9(1)

BOOK REVIEW

H.CE. Hopkins, Y. Pillon, & R. Hoogland. 2014. Cunoniaceae: Flore de la Nouvelle-Caledonie. Vol. 26.

(ISBN: 978-2-85653-764-0, pbk). Publications scinetifiques du Museum, IRD Editions. Case postale 41,

MNHN, 57 rue Cuvier, F-75231 Paris cedex 05, FRANCE. (Orders: diff.pub@mnhn.fr, 01-40-79-48-05).

€59.00,456 pp., 520 b/w illustrations & color photos, 25.5 cm x 16.5 cm.

Commencing with Volume 26, Cunoniaceae, the Flore de la Nouvelle-Caledonie has undergone a complete over¬

haul in presentation and style. I applaud the changes without hesitation, which are too numerous to fully

enumerate.

The covers are of stiffer cardboard compared to earlier volumes and now are in color. In fact, color images

are abundant in this volume of Cunoniaceae, which included treatments authored in parts by Jason C. Brad¬

ford, Gordon McPherson, Porter P. Fowry II, Bruno Fogliani, and Gildas Gateble.

It is entirely in French, apart from English translations for keys to genera, keys to genera using vegetative

characters (to be used in conjunction with the illustrations), and keys to the species within genera. Seven gen¬

era are treated, which are easily located by colored bands (one per genus) on the outer margins of the page.

The volume covers 7 genera and 91 species, the richest being Pancheria and Cunonia. The distribution maps

(one per species) now highlight the extensive ultramahc areas in color, unlike maps used in previous volumes.

Cunoniaceae is a photogenic family, and numerous color photos and black and white illustrations (all of them

excellent) occur throughout.

The family introduction is thorough and includes illustrations of key diagnostic characters (leaves, stip¬

ules, inflorescence structure, and fruits), separate comparative tables for reproductive and vegetative charac¬

ters among the genera, and a table summarizing species number per genus and their distribution across ultra¬

mahc, non-ultramahc, and maquee substrates. Also included are sections on reproductive biology, dispersal

types, and chemical properties. Additional material is provided regarding the methodology underlying the

taxonomic treatments.

Each species has iconic summaries at the top of whether it occurs on ultramahc and/or non-ultramahc

rocks, its endemism status (all of them it appears), and its provisional IUCN conservation status. To reiterate,

excellent line drawings and color photos abound. In some cases tabular comparisons are provided for morpho¬

logically similar species (e.g., in Pancheria and Weinmannia).

One new species, Codiaxerophila Pillion, H.C. Hopkins & Gateble, is treated. A list of material examined by

genus will be greatly appreciated by collections managers.

Reaching beyond taxonomy, the introduction also covers traditional medicinal uses, pharmacological and

cosmetic uses, ornamental potential and uses, forestry and restoration applications, and information on seed

germination and asexual reproduction in vitro.

My one minor complaint is that the font size on the “quick key to the species of Cunonia ” is quite small.

Aside from this, however, the authors and editors of Cunoniaceae have created a higher standard for future

volumes of the Flore.—Neil Snow, PhD, F.M. Sperry Herbarium, Pittsburg State University, Pittsburg, Kansas,

66762 USA.

J. Bot. Res. Inst. Texas 9(1): 48.2015

NUEVAS ESPECIES DE MACHERIUM (LEGUMINOSAE: PAPILIONOIDEAE:

DALBERGIAE) EN MEXICO Y CENTROAMERICA

Jose L. Linares

Herbario Centro Universitario Regional

del Litoral Atlantico (CURLA)

Universidad Nacional Autonoma de Honduras (UNAH)

La Ceiba, HONDURAS

jose.linares@unah.edu.hn

RESUMEN

Se describen e ilustran cinco especies nuevas de Machaerium para Mexico y Centroamerica. Asimismo, se discuten sus afinidades taxonomi-

cas con otras especies.

Palabras Clave: Leguminosae, M achaerium, especies nuevas, Mexico

ABSTRACT

Five new species of M achaerium from Mexico and Central America are described and illustrated, and their taxonomic affinities to other spe¬

cies are discussed.

Keywords: Leguminosae, M achaerium, new species, Mexico

INTRODUCCION

Durante la preparacion del manuscrito del tratamiento del genero Machaerium para la Flora Mesoamericana, se

examinaron las colectas de la region depositadas en los herbarios Nacional de Mexico (MEXU), California

Academy of Sciences (CAS), Missouri Botanical Garden (MO), New York Botanical Garden (NY) y Field Mu¬

seum of Natural History (F), Duddley Herbarium (DS), Fundell Herbarium (FF) y material recientemente col-

ectado en el sureste de Mexico y todavla no distribuido. Ademas, se revisaron las descripciones recopiladas en

los tratamientos existentes para las especies de la zona, como las de Pittier (1922), Standley y Steyermark

(1946) y Rudd (1972, 1973, 1977, 1987) y el material tipo de la mayorla de las especies presentes en la zona

hasta el momento de realizar este estudio. Con base en los ejemplares examinados y en el manuscrito de Fin-

ares y Rudd del genero para la Flora Mesoamericana, se elaboro una clave dicotoma para todas las especies

conocidas hasta el momento de Mexico y Centroamerica. Como resultado del analisis del material disponible,

se detecto la presencia de cinco nuevas para la ciencia, las que se proponen a continuacion:

Machaerium ramosiae J. Finares, sp. nov. (Fig. 1). Tipo: MEXICO. Campeche: Municipio Calakmul, A 1.1 km al E de Conhuas

lat 18°32 , 36"N, 89°54 , 46 ,l O, 195 msnm Veg. Selva mediana subcaducifolia, Bejuco de flores blancas, 20 jul 2002, D. Alvarez 1726

(holotipo: MEXU; isotipo: MEXU).

Vegetative haec species M achaerio semanii similis sed inflorescentiis brevioribus et robustioribus, partibus subglabris, 5 (exceptus 3) foliolis

differt.

Bejucos grandes de hasta 20 m de largo, trepadores de otros arboles; ramitas fertiles grisaceas a blanquecinas,

serlceas cuando jovenes, glabras a muy esparcidamente pubescentes al madurar, con pelos subadpresos muy

esparcidos, el indumento cuando presente cafe oscuro, estlpulas de 1.5-2 mm de largo, triangulares a ligera-

mente falcadas, escariosas, deciduas; yemas vegetativas 0.9 mm, en las bases de las ramitas florlferas, ovoides;

hojas 8-14 cm de largo; pulvino glabro o con unos cuantos pelos subadpresos luego glabrescente; peclolo 2-2.5

cm de largo, glabro o glabrescente; raquis 2.5-4.5 cm de largo, terete, flexuoso, glabro o glabrescente; peciolu-

los 2-3 mm de largo, estrigosos a puberulentos cuando jovenes, glabros o glabrescentes cuando maduros;

follolos 5, alternos; follolos laterales 2.6-5 x (1.2-)1.5-2.2 cm, ovados, a veces oblicuamente ovados, la base

J. Bot. Res. Inst. Texas 9(1): 49 - 61.2015

Journal of the Botanical Research Institute of Texas 9(1)

Fig. \Macheriumramosiae J. Linares. A) Rama floral. B) Inflorescencia. C) Caliz. D) Estandarte. E) Ala. F) Quilla. G) Pistilo. H) Androceo. I) Rama con

frutos, detalle de fruto. A-H tornados de Alvarez 1726 (MEXU); I tornado de Alvarez6664 (MEXU).

Linares, Nuevas especies de Macherium en Mexico y Centroamerica

obtusa a aguda, el apice agudo a ligeramente acuminado, acumen 0.3 a 0.5 cm, el follolo terminal 3.2-6.1 x

1.4-2.4(-3) cm, ellptico, base aguda o cuneada, apice agudo a acuminado, todos los follolos glabros o glabres-

centes en el haz y en el enves cuando maduros, muy esparcidamente pubescentes cuando jovenes, la nerva-

dura central prominente por el enves; nervacion broquidodroma, con nervaduras conspicuas entre las nerva-

duras secundarias; nervaduras terciarias reticuladas y notorias por el haz y enves. Inflorescencias axilares y

terminales en ramitas del ano de crecimiento, 1.5-3.5 cm de largo, muy cortamente pedunculadas el pedun-

culo de 0.5-3.5 mm de largo, paniculadas, pedunculo y raquis serlceos o velutinos, la pubescencia de tricomas

cafe a cafe dorado, bracteas de las inflorescencias cuculiformes a cimbiformes, sesiles de 1-2 x 0.5-1 mm, pa-

piraceas, serlceas, ciliadas en los bordes; Flores ca. 6.2 mm de largo, pediceladas, el pedicelo 0.5 mm de largo,

las bracteolas 0.4 x 0.4 mm, deltoides cuando aplanadas, adpresas al caliz, cuculiformes; caliz 2.5 x 2.3 mm,

campanulado, conspicuamente nervado, las nervaduras prominentes, esparcidamente pubescente, lobos ad-

axiales 0.3 x 0.5 mm, deltoides, los lobos laterales 0.2-0.4 x 0.4-0.5 mm, el abaxial 0.4 x 0.6 mm, triangular. El

estandarte de 5 x 4 mm, incluyendo la una de 1 x 1.8 mm, deltoide, la lamina del estandarte deltoide, pelosa en

la parte adaxial, alas 7x2 mm (incluyendo la una de 2 mm), lamina 5x2 mm, oblonga, casi glabra, quillas 5.5

x 1.5-1.7 mm, (incluyendo la una de 1.5 a 1.8 mm), la lamina 2.5 xl.5-1.7 mm, diminutamente pelosa hacia el

lado abaxial, los estambres 10, diadelfos, el adaxial separado y los restantes unidos, 4.5-6 mm de largo, unidos

en los 2.5 mm basales, anteras 0.2 x 0.3 mm, ovoides, disco nectarlfero 1 x 0.5 mm, tubular, pistilo 5-5.5 mm

de largo, pie 1.5 mm de largo, estilo 1.5 mm de largo, cuerpo 2-2.5 x 1 mm, peloso, el pedicelo y el estilo gla¬

bros. Frutos 6.5-7 x 1.7 cm, sigmoides, a veces doblandose casi en angulo recto a la altura de la camara semi-

nlfera, estlpite 4-7 mm de largo, camara seminlfera 2 cm x 1.2 cm, ala 4.5-5 x 1.6-1.7 cm, margen adaxial

marcadamente convexo, el abaxial ligeramente achatado, la nervacion del fruto prominente. Semillas maduras

no vistas.

Distribution, habitat y fenologia. —Conocida hasta ahora solo de la region de Calakmul en Campeche, en

selva mediana subperennifolia, selva baja subcaducifolia, selva mediana caducifolia y selva mediana subcadu-

cifolia o vegetacion perturbada [datos tornados de las etiquetas] derivada de estas entre 150 y 250 msnm.

Etimologia. —El nombre especlhco honra a la botanica mexicana Clara Hilda Ramos (1941-), dedicada

estudiosa de la flora del sureste mexicano.

Florece en julio y fructihca probablemente de noviembre a diciembre.

Material examinado.— MEXICO. Campeche: Mpio. Champoton, 18°3r40"N, 089 0 54'10"0,150 msnm, selva mediana subperennifolia, se¬

cundaria, ruderal, suelo negro. Arbusto perenne, 4 m, regular, flor blanca, 4 abr 1995, C. Gutierrez B. 4637( MEXU). Mpio. Calakmul, A 1.1

km al NEE de Conhuas, camino a la Zona Arqueologica Nadzcaan, 18 0 32'35"N, 89 0 54'45"0, 193 msnm. Veg. Selva mediana caducifolia,

bejuco lenoso con frutos verdes, 20 ago 2002, D. Alvarez, et al. 1873(MEXU); A 5 km al NE de Calakmul. 18 0 8'35"N 89 0 47'2"0, 230 msnm.

Veg. Selva baja subcaducifolia, arbusto voluble con fruto, 9 die 1998, E. Martinez et S. Ramirez A. 31647 (MEXU); 3 km al Sur de Conhuas,

18°31 , 30"N, 089°56 , 00"0,150 msnm, suelosomero, hierbaperenne, 5 m, regular, fruto verde, 5jul 1995, C. Gutierrez4462 (MEXU); A3.35

km al SE del poblado de Conhuas, camino a Zona Arquelogica de Calakmul lat 18 o 31'0"N, 89°54 , 8"0,165 msnm, Veg. Secundaria, bejuco,

flores amarillas y latex bianco, 30 sep 2003, D. Alvarezy C. Jimenez 6664 (MEXU).

Discusion. —Vegetativamente esta especie es similar a Machaerium seemannii, pero se distingue de ella por las

inflorescencias las cuales son mas cortas y robustas en M. semmannii. Asimismo, M. seemmanii tiene inflores¬

cencias con raquis, flores y frutos serlceos a velutinos, versus las partes casi glabras de la especie nueva. Por otra

parte M. semmannii tiene de 9 a 13 follolos, versus los 5 (excepcionalmente 3) de la nueva especie.

Machaerium excavatum J. Linares, sp. nov. (Fig. 2). Tipo: COSTA RICA. San Jose: Mpio. San Isidro del General, Basin of El

General, 675-900 m, Feb 1940, A.E. Skutch 4705 (holotipo: NY; isotipos: NY, US).

Haec species foliolis magnis et grande caudatis atque habitu volubili giganteo, fructibus incisura notabili ad altitudinem loci seminalis ad

partem adaxialem, sicut Machaerium darienense, distinguitur; a M achaerio darienensi foliolorum numero, forma et magnitudine atque cavi-

tatis magnitudine margine abaxiali differt.

Bejucos gigantes de hasta 20 m de largo, trepadores en otros arboles, a veces reportado como arbolito de 3-5 m,

dentro del sotobosque; ramitas florlferas blanquecinas a grisaceas, conspicuamente lenticeladas, glabras; hojas

con estlpulas de 5-8 x 2-3 mm, glabras, triangulares, cocleadas, algo falcadas, papiraceas o pajizas, persis-

Journal of the Botanical Research Institute of Texas 9(1)

Fig. 2. Macherium excavatum J. Linares. A) Rama con hojas. B) Inflorescencia. C) Caliz y bracteolas. D) Estandarte. E) Ala. F) Quilla. G) Androceo. H)

Gineceo. I) Rama con frutos. J) Semilla. A tornado de Croat 14994 (US); B-H tornados de Skutch 4316 (US); I detalle de fruto. A-H tornados de Alvarez

1726 (MEXU); l-J tornado de Skutch 4705 (US).

Linares, Nuevas especies de Macherium en Mexico y Centroamerica

tentes; yemas vegetativas axilares, presentes solo en ramas esteriles, ovoides, de 2-7 x 1-2.5 mm, cubiertas de

catahlas imbricadas; hojas (17)22-30 cm de largo (probablemente mucho mas grandes), pulvino glabro o gla-

brescente, peclolo (2)4-6 cm de largo, glabro; raquis 5-10.5 cm de largo, terete, recto, glabro o glabrescente;

peciolulos 2-5 mm de largo, glabros o glabrescentes; follolos 5, alternos, follolos laterales 7-15 x 3-6.2 cm,

ampliamente ellpticos a obovados, la base aguda a obtusa, el margen liso y apice caudado, follolo terminal de

10-15 x 4-9 cm, ampliamente ellptico a obovados, la base atenuada a cuneada, el margen liso y el apice cau¬

dado, el acumen en todos los follolos 1-3.5 x 0.2-1 cm, todos los follolos glabros a estrigulosos en el haz y en el

enves (muy diminutamente), la nervadura central prominente por el enves; nervacion broquidodroma, con

nervaduras conspicuas entre las nervaduras secundarias; nervaduras terciarias reticuladas y muy notorias por

el haz y enves. Inflorescencias axilares paniculadas en ramitas del ano y en las partes desnudas adyacentes,

8-12 cm de largo, pedunculo de 2 mm de largo, bracteas de la inflorescencia 2.5-3 x 1.5-2 mm, cuculadas a

concavas, papiraceas, el pedunculo y el raquis estrigoso a serlceo, la pubescencia de tricomas cafe a cafe-dora¬

do, bracteas de las inflorescencias; Flores 8-9 mm de largo, pediceladas, el pedicelo 0.6 mm de largo, las brac-

teolas 1.9 x 2 mm(2.4 x 2.3 mm ya abiertas), deltoides cuando aplanadas, algo cuculiformes, adpresas al caliz;

caliz 23-3.2 x 3 mm, campanulado, marcadamente nervado, estrigoso a serlceo conspicuamente nervados, las

nervaduras prominentes, los lobos del caliz irregulares, los adaxiales 2 x 0.8 mm, los laterales 0.9 x 0.9 mm y el

abaxial 0.8 x 0.7 mm, deltoides. El estandarte 6-8 x 6.2-6.5 mm, incluyendo la una 2x2 mm, la lamina flabe-

lada, con el apice retuso, serlceo hacia la nervadura media por la cara adaxial; alas 7x3 mm (incluyendo la una

2.5 mm de largo), lamina 4.9 x 3 mm, obovada, casi glabras, quillas 6 x 2.9 mm, (incluyendo la una de 2 mm),

la lamina 3.9-4 x 2.8-2.9 mm, auricular, serlcea en la parte media y glabra hacia los bordes. Estambres 10,

unidos todos, excepto por el lado adaxial, 5-7 mm de largo, con la parte unida 2.5-3 mm de largo, anteras 0.6

x 0.5 mm, ovadas, el disco nectarlfero de 1.2 x 1 mm, urceolado a tubular; pistilo 5x1 mm, pie 2 mm de largo,

estilo 1.2 mm de largo, cuerpo 1.7 x 1 mm, oblongo, serlceo, el pedicelo y el estilo glabros. Frutos 7.5-12 x 2-2.8

cm, camara seminlfera 1.8-3 x 1.8 cm, el ala 4.7-7 x 1.9-2.5 cm, ambas con nervaduras muy marcadas, frutos

oblongos, el margen adaxial recto o casi recto y conspicuamente engrosado, el abaxial ligeramente sigmoide

achatado y muy expandido a la altura de la camara seminlfera formando una cavidad de hasta 4 mm de ancho

en la parte mas ancha. Semillas maduras no vistas.

Distribution, habitat y fenologia .—Conocida hasta ahora solo de la region sur de Costa Rica, del bosque

lluvioso de Golfo Dulce y de San Isidro del General, en bosques premontanos de 10-1000 msnm.

Etimologia .—El eplteto especlhco alude a la depresion de la camara seminlfera en el margen abaxial del

fruto que distingue a esta especie.

Florece en junio y fructihca de enero a marzo.

Material examinado.—COSTA RICA. San Jose: Mpio. San Isidro del General, Basin of El General, alt. 675-900 m, Jun 1940, A.F. Skutch

4954 (NY, US); Canton Puriscal, Zapoton de Puriscal, Arbolito creciendo en un paredon a orillas del camino, inclinado, frutos abundantes

de color verde-limon, 31 ene 1986, N. Zamora etal. 1161 (CAS); San Martin, Camino a Parrita, 9 o 44'10"N, 84°24 , 50"0,800 m, arbolito de 5 m

x 13 cm de DAP, ubicado en paredon cercano a la carretera, frutos muy abundantes verde-amarillentos, 29 nov 1989, Q. Jimenez et al. 750

(MEXU); vicinity of El General, 640 m, Riverside cliff, Jun 1939, A.F. Skutch 4316 (US); cutover hills, ca. 8 km SW of San Isidro del General,

1000 m, small tree 3-5 m tall, in forest, 27 Jan 1965, L.O. Williams et al. 28383 (NY, US). Puntarenas. Sin Mpio.: along N fork (known lo¬

cally as “Quebrada Mona”) of Quebrada Bonita, Carara reserve, 9°47'N, 84°36'0, 35-40 m, giant liana, hanging low along creek and in as¬

sociation with major treefall in primary forest, samaras light green with fine, darker veins (uniformly dark at base), 31 Aug 1985, M.H. Gra-

yum et al. 5952 (MEXU, MO), Reserva Biologica Carara Sector Quebrada Bonita, Sitio Sendero Valentin, 9°46 , 20"N, 84 0 36'35"0, 70 m, 11 ene

1990, R. Zuniga 75 (MEXU); Canton (?), Lomas de Cerro Esquivel, entre Santiago de San Ramon y Macacona de Esparza. 10°02 , 55"N,

84°36’20”O, 300-500 m, bejuco trepador, frutos inmaduros verde-amarillo, 15 sep 1986, G. Herrera et al. 3 (CAS, MEXU); Canton Golfito,

Peninsula de Osa. Puerto Jimenez. Cabecera Corozal 08°32 , 30"N, 83 o 18'30"O, 1-10 m, Bejuco, muy comun. Frutos verdes, 4 feb 1995, R.

Aguilar 3745 (NY); entre Puerto Jimenez y Agujas, entre los Rios Agujas y Terrones. 08 o 33'55"N, 83 o 23'20"O, 50 m, Arbusto-bejuco, frutos

verdes, 19 oct 1994, R. Aguilar y J. Gonzdlez 3614 (NY); Santo Domingo de Golfo Dulce, Fonduz 7018, Forets de Santo Domingo de Golfo

Dulce, mar 1896, Fonduz 7242 (US).

Discusion .—Esta es una de las especies mas distintivas, vegetativamente y resalta por sus follolos de gran

tamano y grandemente caudados, as! como su habito de liana gigante. Sus frutos son muy caracterlsticos por

tener una muesca muy notoria a la altura de la camara seminal por el lado abaxial, caracterlstica solo compar-

Journal of the Botanical Research Institute of Texas 9(1)

tida por M. darienense. Se distingue de esta ultima especie por el numero, forma y tamano de los follolos, asi

como el tamano de la cavidad en el margen abaxial.

Machaerium paucifoliolatum J. Linares, sp. nov. (Fig. 3). Tipo: Nicaragua. Chontales: Municipio La Gateada O, ca. 4.5

km S of Hwy 7 (from ca. 3.6 km E of La Gateada) on road to Nueva Guinea, ca. 12°57'N, 85°45'0, ca. 200 m, cut-over tall wet forest,

disturbed roadside and pastures with scattered individual trees and small remnant patches of forest, one seen, large woody vine,

climbing to canopy of large tree and forming dense masses, corolla white except wings brown, 17 Jul 1977, WD. Stevens 2847 (holo-

tipo: MEXU).

Vegetative haec species Machaerio pittieri similis, praeter 3 foliola, fructum angustiorem et saepe forte sigmoideum versus flexum in “L.”

Bejucos grandes de hasta 20 m de alto, trepadores de otros arboles; ramitas fertiles grisaceas a blanquecinas,

glabras, estriadas, lenticeladas, las del ano anterior similares pero mas blanquecinas. Estlpulas de 1.6 x 1.2

mm, triangulares a ligeramente falcadas, prontamente deciduas; yemas vegetativas axilares 0.7-2 x 0.5-1.2

mm, ovoides, presentes en las hojas y en los apices de las ramas, yemas seriales presentes y en zigzag; Hojas

7-17 cm de largo; pulvino glabro o con unos cuantos pelos subadpresos, glabrescente; peclolo 2-3.5 cm de

largo, glabro o glabrescente; raquis 1.1-2.5 cm de largo, terete, recto, glabro o glabrescente; peciolulos 2-3 mm

de largo, glabros o muy esparcidamente pubescentes; follolos 3, alternos; follolos laterales 3-8 x 1.9-4.9 cm,

ovados a anchamente ellpticos, la base atenuada a obtusa, el apice, acuminado o caudado, con un acumen de

ca. 1.5 x 0.4 cm, el borde entero, sin pelos, el follolo terminal 5-11 x 2.3-5 cm, ellptico, anchamente ellptico,

ovado u obovado, base obtusa a aguda, apice agudo a acuminado, todos los follolos glabros o glabrescentes en

el haz y en el enves, la nervadura central prominente por el enves; nervacion broquidodroma, con nervaduras

conspicuas entre las nervaduras secundarias; nervaduras terciarias reticuladas y notorias por el haz y enves,

nervaduras secundarias 6-8 pares. Inflorescencias axilares y terminales en ramitas del ano 1.5-3.5 cm de lar¬

go, muy cortamente pedunculadas el pedunculo de 0.5-3.5 mm de largo, paniculadas, el pedunculo y el raquis

serlceo o velutino, la pubescencia de tricomas cafe a cafe-dorado, bracteas de las inflorescencias cuculiformes

a cimbiformes, sesiles, 1-2 x 0.5-1 mm, papiraceas, serlceas, ciliadas en los bordes; flores 7 mm de largo,

pediceladas, el pedicelo 0.5 mm de largo, las bracteolas 2.5 x 1.2 mm, extendidas (1.5 x 1.5 mm en posicion

normal), deltoides cuando aplanadas, algo cuculiformes, adpresas al caliz; bracteolas 0.4-0.6 x 1-1.1 mm, cu¬

culiformes; caliz 3 x 2.5 mm, campanulado, conspicuamente nervados, las nervaduras prominentes, no llegan

hasta el borde del caliz, esparcidamente pubescente, lobos adaxiales 1.7 x 0.7 mm, deltoides, los tres restantes

mas pequenos 0.9 x 0.6 mm, deltoides. El estandarte 5.5 x 6 mm, incluyendo la una 1.2 x2 mm, deltoide,

lamina del estandarte obcordada (cuando aplanada), serlcea hacia el lado adaxial, alas 5.5 x 1.5 mm (incluy¬

endo la una 1.8 mm de largo), lamina 3.7 x 1.5 mm, oblonga, pelosas en la parte adaxial, quillas 5.5-5.8 x 2 mm

(incluyendo la una 2-2.2 mm de largo), la lamina 3.5-3.8 x 2 mm, oblonga, pelosa hacia el lado abaxial, corolas

blancas con las alas cafe. Estambres 10, diadelfos, el adaxial separado y los restantes unidos, 4-5 mm de largo

(incluyendo las anteras), hlamentos lateralmente comprimidos (0.3 mm de ancho), anteras 0.3 x 0.3 mm,

ovoides, disco nectarlfero 0.6 x 0.4 mm, tubular, crenado, con 10 crestas. Pistilo 5.5 mm de largo, pie 1 mm de

largo, estilo 1.2 mm de largo, cuerpo 3x1 mm, peloso, el pedicelo y el estilo glabros. Frutos 7-9 x 1.1-2 cm.

sigmoides, a veces doblados casi en “L” a la altura de la camara seminlfera, camara seminlfera 2.5 x 0.9 cm, es-

tlpite 3-5 mm, alas 4.5-7 x 1.1-2 cm, acuminada, obtusa o redondeada, glabra, margen adaxial marcadamente

convexo, el abaxial ligeramente achatado y canaliculado en la camara seminlfera, la nervacion del fruto

prominente. Semillas maduras no vistas.

Distribution, habitat y fenologia .—Conocida de selvas humedas (SAP), pluvioselvas, bosques premonta-

nos, de Honduras y Nicaragua donde crece en la vertiente Caribe y en Costa Rica donde crece en ambas verti-

ente oceanicas, florece en julio y fructihca en enero-marzo.

Etimologia .—El nombre especlhco alude al reducido numero de follolos caracterlstico de esta especie.

Material examinado.— HONDURAS. Olancho: Mpio. Santa Maria del Real, camino a represa del Rio Real, Arbol 15m, 14 o 50'78"N

85°56 , 76[sic] ,l O, 550 m, 20 marzo 2006, P.R. House et al. 3709 (MEXU). NICARAGUA. Chontales: Mpio. Cuapa, 0.8 km N of Cuapa, pasture

fencerows, 340 m, tree collected along edge of woody area, fruits dark green, 12-15 m, 22 Jan 1978, P.C. Vincelli 101 (MEXU); 3 km N of

Cuapa, premontane moist forest, elev. 500 m. Small tree 7 m, 3 Sep 1977, D. Neill 2501 (MEXU); ca. 2.8 km above (N of) Cuapa, ca. 12°17'N,

Linares, Nuevas especies de Macherium en Mexico y Centroamerica

Fig. 3. Macheriumpaucifoliolatum J. Linares. A) Rama con hojas. B) Flor. C) Caliz. D) Estandarte. E) Ala. F) Quilla. G) Androceo. H) Gineceo. I) Rama con

frutos. A-H tornados de Warren etal. 2847 (MEXU); I tornado de Vincelli 101 (MEXU).

Journal of the Botanical Research Institute of Texas 9(1)

85°23'0,400-500 m, roadside, pastures, disturbed evergreen forest on hillside, and along small stream, forest, large woody vine, climbing

by thick tendrils, wing of fruit pale yellow-green, W.D. Stevens 6055 (MEXU). Matagalpa: Mpio. Matagalpa, cut-over hills ca. 15 km NE of

Matagalpa along Rio Las Canas, ± 700-800 m, 14 Jan 1965, L.O.Williams et al. 27540 (DS, NY, US). Nueva Segovia: ca. k 233.5, ca. 6.2 km N

of N, edge of Ocotal, Quebrada El Nancital, ca. 13°41’N, 86°24’0, 700-760 m, mixed evergreen and deciduous forest in quebrada, cut-over

pine forest on slopes and ridge, large fallen tree along stream, leaves deep green, flowers white, fading to brown, W.D. Stevens 3036 (MEXU).

COSTA RICA. Alajuela: Canton Los Chiles, R.N.V.S. Cano Negro, Llanura de Guatuso. Cano Negro, Playuelas. 10°54 , 50"N 84°46 , 05"O 40

m, arbusto escandente de 4 m, frutos con parte seminal o semifera, verde y el ala amarillo-verdoso, Es muy comun en el lugar, 3 feb 1993, K.

Martinez et al. 54 (NY); Rio Chiquito, approx. 40 km Road to Upala, 800 m, L.D. Gomez 18616 (MEXU). Guanacaste: Mpio. Santa Rosa,

Camino del ICE de Santa Cruz a Vista de Mar, 900 m, bejuco de area en crecimiento secundario, flores aromaticas, cafe amarillento, 22-26

jul 1985, L.D. Gomez et G. Herrera 23672 (CAS, MEXU,

Discusion. —Vegetativamente esta especie es similar a Machaerium pittieri , excepto que siempre presenta tres

follolos versus M. pittieri que siempre tiene cinco o mas, ademas, el fruto es mas angosto y a menudo fuerte-

mente sigmoide a doblado en “L ”

Machaerium rubrinervumj. Linares, sp. nov. (Fig. 4). Tipo: Costarica, puntarenas: CantonOsa, r.f. GolfoDuke,

Cuenca Terraba-Sierpe, Bahia Chal., La Parcela, 8 o 43'50"N 83°27T7"0,150 m, bejuco lenoso, flores con el estandarte exteriormente

pubescente, amarillo brillante, internamente morado oscuro, quilla verdosa-amarillenta, 1 ago 1996, R. Aguilar 4635 (holotipo:

NY).

Haec species insigniter distinguibilis ramulis rubellis, foliolis nervibus centralibus rubellis et indumento pilis basi aurantiaca et apice albes-

centi, forma foliolorum praecipue elliptica et numero reducto.

Bejucos, trepadores de otros arboles; ramitas fertiles rojizas a cafes o cafe-grisaceas, esparcidamente hirsuto

pelosas, pelos rojizos. Hojas con estlpulas deciduas, no vistas; yemas vegetativas axilares 0.5 x 0.2 mm, ovadas.

Hojas 7.5-12 cm de largo; pulvino hirsuto; peclolo 1.2-1.7 cm de largo; raquis 1.8-4 cm de largo, terete, recto;

peciolulos 1.2-1.7 mm de largo, hirsutos. Follolos 5-7, alternos; follolos laterales 2.2-5.6 x 1.1-2.2 cm, ovados

a mayormente ellpticos, simetricos, la base obtusa a aguda, el apice acuminado con un acumen 0.5-1 x 0.4 cm,

mucronado, el borde entero, sin pelos, el follolo terminal 5-6 x 1.5-2 cm, ellptico, base aguda o cuneada, apice

acuminado, todos los follolos glabros por el enves, excepto por la nervadura media, la cual es hirsuta y estri-

gosa por el enves, la pubescencia del enves con pelos blanquecinos, pero con la base de color anaranjado a ro-

jizo, dando la apariencia de ser engrosado en la base, la nervadura central prominente por el enves, rojiza;

nervacion broquidodroma, con nervaduras poco conspicuas entre las nervaduras secundarias; nervaduras

terciarias reticuladas poco notorias por el haz y enves. Inflorescencias terminales o agrupandose hacia las axi-

las distales de las ramitas florlferas, paniculadas, 1.5-3 cm de largo, muy cortamente pedunculadas el pedun-

culo 0.5 mm de largo, el pedunculo y el raquis hirsuto pubescente, pelos cafe-rojizos, bracteas de las inflores¬

cencias 0.5 x 0.4 mm, triangulares, pajizas, escariosas, flores 5-6 mm de largo, con pubescencia cafe-rojiza,

adpresas al caliz, el pedicelo de menos de 1 mm, las bracteolas 1 x 0.8 mm, cocleadas, laciniadas en los bordes,

estrigulosas, casi glabras, caliz 2.8 x 2 mm, campanulado, estrigoso, lobos adaxiales de 0.4 x 1 mm, deltoides,

los laterales 0.4 x 0.8 mm, deltoides y el abaxial de 0.3 x 0.7 mm, triangular. Estandarte 5 x4 mm (incluyendo

la una 1.2 mm de largo), lamina suborbicular, alas 4.5-5 x 1.3 mm, quillas 4.5 x 1.3 mm (incluyendo el pie de

1.5 mm de largo), lamina 3 x 1.3 mm, oblonga, auriculada hacia el lado adaxial. Estambres 10, todos unidos

formando un collar que abraza al pistilo, este collar abierto por el lado adaxial, estambres 3.8-4.5 mm de largo,

la parte unida 3 mm de largo, anteras 0.3 x 0.2 mm, ovadas. Disco nectarlfero oblicuo, completamente abierto

por el lado abaxial con el extremo mas largo hacia el lado adaxial (de 0.6 mm de largo en su punto mas largo).

Pistilo de 4.4 x 0.7 mm, pie 1 mm de largo, glabrescente, estilo 1 mm de largo, glabro. Frutos no vistos. Semillas

maduras no vistas.

Distribucion, habitat y fenologia. —Conocida de una sola coleccion en la region de Golfo Dulce, en el sur de

Costa Rica, probablemente en bosques lluviosos. Florece en julio-agosto y fructihca probablemente en enero-

marzo.

Etimologia. —El nombre se rehere a las nervaduras marcadamente rojizas y contrastantes con la lamina

foliar.

Discusion. —Esta especie es muy distintiva por sus ramitas rojizas, hojas con nervaduras centrales rojizas

Linares, Nuevas especies de Macherium en Mexico y Centroamerica

Fam. LEGUMINOSAE

Machaerium

Det. Jose L. Linares 2003

hitvu -foL Coyk-y

FLORA DE COSTA RICg

COSTA RICA inbIO

mI- ^//r/w

;nca ierraba-bierpe

lia Chal. La Parcela.

8°43'50"M 83°27 J '1T'W

5uco lenoso. Flores co

unaicto Aguilar 4b3b t ® igog

INSTITUTO NACTCMAL DE BIODI VERS I DAD (]

Eli eolaboracion cob el Missouri Botanical Garden (HO)

< 5 ‘

IT 03

r+ w

Fig. 4. Machaerium rubrinervium J. Linares. Imagen escaneada del especimen tipo (/?. Aguilar 4635, NY)

Journal of the Botanical Research Institute of Texas 9(1)

y por el indumenta formado por pelos con la base anaranjada y el apice blanquecino. Ademas, la forma de los

follolos, mayormente ellpticos y en numero reducido es tambien muy caracterlstica.

Machaerium franksullyvaniij. Linares, sp. nov. (Figs. 5, 6). Tipo: el Salvador. Sonsonate: Mpio. izalco, Linca Marfa

Auxiliadora, campos de lava recientes y campos de lava arbolados, 13°47'05"N y 89 o 37'07"0,1060 m, 29 jul 2007J.L. Linares 12232

(holotipo: MEXU).

Species M. pittieri Macbride similis sed foliis margine repando (forte undulato) petalis carinae conduplicatis latere abaxiali, fructibusque

glandibus circularibus depressis, margine adaxiali insigniter incrassato, usque ad 2 mm crassum, habitatione campis lavae et silvis monta-

nis inter 1000 et 2000 m differt.

Bejucos muy grandes, de hasta 10-15 m de largo, trepadores de otros arboles, a veces reportado como arbolito

de 3-5 m, dentro del sotobosque, pero mayormente bejuco; ramitas florlferas pardas, grisaceas o cafe, con-

spicuamente lenticeladas, glabras; hojas con estlpulas de 5-8 x 2-3 mm, glabras, triangulares, cocleadas, algo

falcadas, papiraceas o pajizas, persistentes; yemas vegetativas axilares, presentes solo en ramas esteriles

jovenes, inconspicuas, con dos bracteas diminutas, de 1 mm o menos; hojas 10-21.5(-24) cm de largo, pulvino

glabro o glabrescente, peclolo (1.5-)2.3-3 cm de largo, glabro, comunmente el peciolo subtendido por una

bractea cimbiforme de 3 mm de largo y ancho, con pelos adpresos blanquecinos; raquis 4-8 cm de largo, terete,

algo flexuoso, glabro o glabrescente; peciolulos 5-7 mm de largo, glabros o glabrescentes; follolos 7(—9), alter-

nos, follolos laterales 2.6-7 x 1.4-4 cm, ovados a ellpticos, base aguda a obtusa, la base aguda a obtusa, el

margen repando u ondeando y apice de hasta 8 mm de largo, retuso y, en material de herbario comunmente

doblado, follolo terminal de 5-9 x 2.5-4 cm, ampliamente ellptico, obovados u oblongo, la base atenuada a

cuneada, el margen repando u ondeado y el apice acuminado, con un acumen de hasta 9 mm de largo, similar

al de los foliolos laterales, todos los follolos completamente glabros en el haz y en el enves, excepto por unos

cuantos pelos estrigosos en los pulvinos jovenes, las nervaduras primaria, secundarias y hasta de tercer orden,

aplanadas en el haz y prominente por el enves; nervacion broquidodroma, con nervaduras conspicuas entre las

nervaduras secundarias; nervaduras terciarias reticuladas y muy notorias por el enves. Inflorescencias mayor¬

mente terminales, si axilares restringidas a las ultimas 2-3 hojas de las ramas, generalmente simples o en fas-

clculos de 2-3 inflorescencias, a veces paniculadas, especialmente las terminales, naciendo en ramitas del ano,

1.5- 7 cm de largo, pedunculo de 0.5-1 mm de largo, generalmente con una bractea 4x1.5 mm, cimbiforme

emarginada, bracteas de la inflorescencia 1 x 0.5 mm, cuculadas, papiraceas, serlceas, el pedunculo y el raquis

estrigoso a serlceo, la pubescencia de tricomas ferruglneos a cafe-dorado; Flores 8-9 mm de largo, pediceladas,

el pedicelo 0.2-0.6 mm de largo, las bracteolas 1.5 x 2.5 mm, reniforme cuando aplanadas, traslapadas y for-

mando un involucro redondeado, subadpresas a patentes, caliz 2.3-4.5 x 4 mm, campanulado, estrigoso a

serlceo, los lobos del caliz irregulares, los adaxiales o vexilares 0.5 x 2 mm, en forma de triangulo, los laterales

0.5 x 1.5 mm, triangulares a obtusos y el abaxial 0.5 x 1 mm, redondeado. El estandarte 6-8 x 5-7.5 mm, in-

cluyendo la una 0.5 x 1-1.5 mm, la lamina reniforme a suborbicular, con el apice retuso, serlceo hacia la nerva-

dura media por la cara adaxial; alas 7x3 mm (incluyendo la una 1.5 mm de largo, curveada, casi sigmoide),

lamina 5.5 x 3 mm, obovada, glabras, quillas 7-7.5 x 2.5-3 mm, (incluyendo la una de 1.5 mm), la lamina

5.5- 6 x 2.5-3 mm, se semiorbicular, auriculada, glabra. Estambres 10, diadelfos (9:1), 6-7.5 mm, la parte unida

de 3.5 mm el vexilar de 5 mm, anteras 0.6 mm x 0.5 mm, ovadas, el disco nectarlfero de 0.5 x 0.6 mm, urceo-

lado a tubular; pistilo 6 x 1.5 mm, pie 1.5 mm de largo, estilo 1.3 mm de largo, cuerpo 2.5-3 x 1.5 mm, oblongo,

villoso, el pedicelo y el estilo glabros. Frutos 7-10.5 x 1.5-2.5 cm (incluyendo el pedicelo de 1-1-3 cm), camara

seminlfera 2.7-3 x 1.4 cm, el ala 4.7-5 x 1.9-2.2 cm, ambas con nervaduras muy marcadas, frutos oblongos, el

margen adaxial recto o casi recto y conspicuamente engrosado, hasta 2 mm de grueso, el abaxial ligeramente

sigmoide, obtuso, surcado en la camara seminlfera. Semillas maduras 19-20 x 10-11 x 4-5 mm, reniformes,

algo rugosas, hilo de 1 mm, linear.

Distribution, habitat y fenologia .—Conocida hasta ahora solo de la region suroccidental de El Salvador, en

la sierra de Apaneca-Lamatepec, de 1000-1900(-2000) msnm en selvas medianas subcaducifolias, bosques de

niebla, vegetacion derivada y cafetales, bordes de caminos.

Etimologia .—Es un placer dedicar esta especie al MSc. Frank Sullyvan Cardoza (1976-), joven apasiona-

Linares, Nuevas especies de Macherium en Mexico y Centroamerica

1265103

HERBARIO PAUL C. STANDLEY

EAP

PLANTAS DE EL SALVADOR

FAM. LEGUMINOSAE

Machaerium

cafetales, campos de lava recientes y campos de lava arbolados’

^ c b0 t? Ue !^ Cesionales sobre lavas ' “fetales y caminos. 13°

47 05 N y 89 37 07 W. Alt. 1060 m. Bejuco grande creciendo

en el bosque entre el cafetal y el campo de lava, pocos frutos.

Jose L. Linares 12232

29 de julio de 2007

Fig. 5. Especimen tipo de Machaerium franksullyvanii J. Linares [J.L Linares 12232, MEXU!).

Journal of the Botanical Research Institute of Texas 9(1)

Fig. 6. Machaerium franksullyvanii J. Linares. Hoja mostrando el haz (a), enves (b), notese los hordes de los foliolos daramente repandos; detalle de la

inflorescencia (c), flor ampliada (d).

Linares, Nuevas especies de Macherium en Mexico y Centroamerica

do de la botanica, que durante anos ha participado en la colecta, identihcacion y fotografla de la diversidad

vegetal en Mesoamerica. Su contribucion ha ayudado a documentar la biodiversidad alojada en esta region.

Frank ha participado como voluntario poniendo su destreza como fotografo y su patrocinio al servicio de algu-

nas expediciones botanicas. Estamos seguros que seguiremos contando con el en nuevas expediciones florlsti-

cas.

Florece en marzo y abril y fructihca de enero a marzo del siguiente ano.

Nombre vernacular .—Bejuco de una de gato, bejuco de flores negras.

Material examinado.—EL SALVADOR. Ahuachapan: Mpio. San Francisco Menendez, El Imposible, San Benito, en Cerro Campana,

13°49'N y 89°56'0, bejuco de ca. 23 m, flores negras, abr 1992, Sandoval y Chinchilla 375 (MEXU); Mpio. Apaneca, 5 km antes de Apaneca,

13°52'N y 89°45'0 1100 m, arbusto escandente con tallos lenosos, creciendo a orilla de calle, hojas compuestas imparipinnadas, frutos de

color verde claro, sep 1997, R. Villacortay M. Renderos (RV-02589 ) (MEXU); Mpio. Ataco, Las Oscuranas, oct 2006,J. Linares, J.F. Gutierrezy

J.E. Castillo s.n. (MEXU). Sonsonate: Mpio. Izalco, Campos de lava del volcan de Izalco, ca. 1 km al S de la Finca La Macarena, al N del canton

Cruz Grande, 13°59'N y 89°39'0, 1100-1200 msnm, arbusto achaparrado de copa densa, sin espinas, feb 2000, J. Linares 4889 (MEXU);

Mpio. Izalco, Campos de lava reciente y campos de lava arbolados, Veg. Bosques sucesionales sobre lavas, cafetales y caminos, 13 o 47'05"N y

89 o 37'07"O, 1060 msnm, bejuco grande creciendo en el bosque entre el cafetal y campo de lava, pocos frutos, jul 2007, J. Linares 12232

(MEXU); idem, arbol debil de 4 m, creciendo en el cafetal, muchos frutos, jul 2007J. Linares 12239 (MEXU); idem, bejuco, frutos verdes, ago

2007,J. Linares 12293 (MEXU).

Discusion .—Esta distintiva especie resalta por su habito vegetativo como bejuco o arbusto aparrado de gran

tamano con numerosas ramas escandentes cubriendo una extension relativamente grande, a veces trepando

sobre arboles cuando crece en bosques o creciendo como parras aisladas en campos de lava vieja; sus follolos

presentan margenes o bordes repandos, es decir, muy ondeados, con aspecto de haber sido secados en forma

defectuosa, pero es una caracterlstica claramente visible en vivo. Por otro lado, en cuanto a las flores, destaca el

caliz con los dientes o lobos inconspicuos, el estandarte casi orbicular, la quilla con uno o sus dos petalos clara¬

mente conduplicados en el margen abaxial. Ademas, sus frutos presentan, al menos cuando jovenes, pubescen-

cia estrigosa en todo el cuerpo, glandulas circulares depresas mas oscuras, a veces inconspicuas, y una sutura

vexilar muy engrosada de hasta 2 mm.

AGRADECIMIENTOS

A las autoridades de la UNAH y CURLA, especialmente a Jean O. Rivera y Eydi Yanina Guerrero que apoyaron

la hnalizacion de este trabajo, a Fernando Chiang Cabrera que facilito mis estancias en la Ciudad de Mexico

durante la realizacion de este trabajo, a Cyrill Nelson Hardy Sutherland (Cyril Hardy Nelson Sutherland) por

sus diagnosis en latln y sus valiosas sugerencias al manuscrito, a Gloria y Mario Sousa por su inestimable

ayuda en el herbario Nacional de Mexico (MEXU), a Marla del Rosario Garcia Pena (Maru) por su ayuda en el

manejo de los prestamos dentro y fuera de Mexico, a dos revisores anonimos que contribuyeron a mejorar no-

tablemente el manuscrito, a Frank Sully van Cardoza Ruiz y Ana Eugenia Aguilar por su ayuda en los viajes de

campos en El Salvador y Honduras, a los curadores de los Herbarios CAS, DS, F, LL, MEXU, MO, NY y US por

poner a mi disposicion gran cantidad de material para su estudio.

REFERENCIAS

Pittier, H. 1922. Fabaceae.The middle American species of Machaerium. Contr. U.S. Natl. Herb. 20(12):467-477.

Rudd, V.E. 1972. Newtaxa and combinations in Machaerium (Leguminosae) II. Phytologia 24:121-125.

Rudd, V.E. 1973. Newtaxa and combinations in Machaerium (Leguminosae). III. Phytologia 25:398-403.

Rudd, V.E. 1977. The genus Machaerium (Leguminosae) in Mexico. Bol. Soc. Bot. Mexico 37:119-146.

Rudd, V.E. 1987. Studies in Machaerium (Leguminosae) V. Phytologia 62:277-281.

Standley, P.C. & J.A. Steyermark. 1946. Leguminosae. Flora of Guatemala. Fieldiana, Bot. 24(5):1-368.

Journal of the Botanical Research Institute of Texas 9(1)

BOOK REVIEW

Susan Leopold, with illustrations by Nicky Staunton. 2014. Isabella’s Peppermint Flowers. (ISBN 978-0-692-

33302-0) (Orders: www.FloraForKids.org). $18.00 US, 36 pp., watercolor drawings, glossary, epilogue.

For the young aspiring botanist, this book will be a treasure. The story of two girls who go flower-hunting with

their mother becomes an introduction to the basics of botany, augmented by the excellent illustrations. The

author incorporates the historical context of botanical pioneers including an introduction to Linneaus, John

Clayton, and John Gronovius in the State of Virginia. Terms and names used in the context of the story are

clarified in the definitions at the end. This book is an excellent teaching text for beginning botanists.— Pamela

Edmondson, Volunteer, Botanical Research Institute of Texas, Fort Worth, Texas, USA.

What a fantastic and unique resource for both children and their caretakers! Author Susan Leopold uses the

story of two sisters and a mother walking in the woods to introduce the basics of taxonomy, floristics, and plant

collecting, all while dropping in asides about binomial nomenclature, rare and endangered species, and bo¬

tanical history. True, the story focuses in part on the history of the flora of Virginia, but the overall message is

universal: inspiring a sense of place and an appreciation of the natural world. For the sake of encouraging a new

generation of botanists, it would be a wonderful thing to see books like this for all 50 states.— Brooke Byerley

Best, PhD, Editor & Botanist, Botanical Research Institute of Texas, Fort Worth, Texas, USA.

BOOK NOTICE

Larry J. Littlefield & Pearl M. Burns. 2015. Wildflowers of the Northern and Central Mountains of New

Mexico: Sangre de Cristo, Jemez, Sandia, and Manzano. (ISBN-13: 978-0-82635-547-8, pbk.). Univer¬

sity of New Mexico Press, 1717 Roma Avenue NE, Albuquerque, New Mexico 87106, U.S.A. (Orders:

www.unmpress.com, 1-800-249-7737). $29.95 US, 408 pp., 419 color plates, 1 map, 5.5" x 8.5".

From the publisher: This unique reference work describes over 350 wildflowers and flowering shrubs that grow

in New Mexico’s Sangre de Cristo, Jemez, Sandia, and Manzano mountains, as well as neighboring ranges, in¬

cluding the Manzanita, San Pedro, Ortiz, and other lower-elevation mountains in central portions of the state.

With more than a thousand color photographs accompanied by visual descriptions, the easy-to-use guide or¬

ganizes plants first by flower color, then alphabetically by family common name, then by scientific name. The

authors also include information on traditional uses of the plants by indigenous peoples and an extensive glos¬

sary and bibliography. A brief geological history and description of the ranges examines the different life zones

and ecosystems and how these relate to elevation and microclimates. Wildflower enthusiasts and hikers will

welcome this useful book.

Larry J. Littlefield is a professor emeritus of plant pathology at Oklahoma State University. He has been a

volunteer with the Sandia Mountain Natural History Center and the trails maintenance crew for the U.S. Forest

Service since retiring in Albuquerque in 2005.

Pearl M. Burns, coauthor of the “Wildflowers” chapter of Field Guide to the Sandia Mountains (UNM Press), is

a member of the Friends of the Sandia Mountains and a leader of countless wildflower hikes for the U.S. Forest

Service, Albuquerque Open Space program, and other organizations.

J.Bot. Res. Inst. Texas 9(1): 62.2015

GNAPHALIOTHAMNUS NESOMII (ASTERACEAE: GNAPHALIEAE),

A NEW SPECIES FROM GUATEMALA AND NOMENCLATORIAL CHANGES

Michael O. Dillon

Federico Luebert

Botany Department

The Field Museum

1400 South Lake Shore Drive

Chicago , IL 60605 , USA.

mdillon@fieldmuseum.org

Universitat Bonn

Nees-lnstitut fur Biodiversitat der Pflanzen

MeckenheimerAllee 170

D-53115 Bonn, GERMANY

ABSTRACT

Gnaphaliothamnus nesomii M.O. Dillon & Tuebert (Asteraceae: Gnaphalieae) is a new species from the Sierra de los Cuchumatanes, De¬

partment Huehuetenango, Guatemala. The generic boundaries within the Gnaphalieae have been controversial and the genus Gnaphalio¬

thamnus has not been universally accepted. New molecular phylogenetic studies support the acceptance of Gnaphaliothamnus as distinct

from Chionolaena, which is congruent with cypsela trichome morphology. Two Gnaphalium species are transferred as Pseudognaphalium

stolonatum (S.F. Blake) M.O. Dillon and P. paramorum (S.F. Blake) M.O. Dillon.

Key Words: Chionolaena, Gamochaeta, Gnaphaliothamnus, Gnaphalium, Pseudognaphalium, Asteraceae, Gnaphalieae, Gnaphaliinae, Guate¬

mala, Lucilia- group, comb, nov., sp. nov., Sierra de los Cuchumatanes, taxonomy

RESUMEN

Gnaphaliothamnus nesomii M.O. Dillon & Tuebert (Asteraceae: Gnaphalieae), es una nueva especie proveniente de Sierra de los Cuchu¬

matanes, Departamento Huehuetenango, Guatemala. Eos limites genericos en Gnaphalieae son controvertidos y el genero Gnaphaliotham¬

nus no ha sido universalmente aceptado. Nuevos estudios filogeneticos moleculares apoyan la aceptacion de Gnaphaliothamnus como un

genero diferente de Chionolaena, lo que es congruente con la morfologia de los tricomas de las cipselas. Dos especies de Gnaphalium son

transferidas como Pseudognaphalium stolonatum (S.F. Blake) M.O. Dillon y P. paramorum (S.F. Blake) M.O. Dillon.

Palabras Clave: Chionolaena, Gamochaeta, Gnaphaliothamnus, Gnaphalium, Pseudognaphalium, Asteraceae, Gnaphalieae, Gnaphaliinae,

Guatemala, Lucilia- group, comb, nov., sp. nov., Sierra de los Cuchumatanes, taxonomy

INTRODUCTION

Gnaphaliothamnus Kirp. (Asteraceae, Gnaphalieae) comprises around 11 species distributed from Mexico to

Costa Rica. It has been accepted (Nesom 1990a,b; 1994) or subsumed in the South American Chionolaena DC.

(Anderberg & Freire 1989,1991; Freire 1993; Nesom 2001; Freire et al. 2015). Based upon differences in cypse-

lar (achenial) trichomes, Dillon (2003) argued that Gnaphaliothamnus was a monophyletic group not necessar¬

ily close to Chionolaena. The cypselar trichomes in Gnaphaliothamnus are described (Hess 1938) as short cla-

vate (zwillingshaares) with an enlarged adaxial basal cell ( schwellpolster ) and myxogenic apical cells 65-125 pm

long. The two apical cells in the cypselar trichomes in Chionolaena are much longer (250-850 pm long) and

not myxogenic (Dillon & Sagastegui 1991; Loeuille et al. 2011).

Pruski (2012) transferred a Guatemalan species, Gnaphalium stolonatum S.F.Blake, to Chionolaena, with the

comment that it resembled other Mexican and Central American members. He apparently did not examine the

type material of G. stolonatum, providing a new description drawn from a herbarium collection that he termed

a “topotype” (A. Molina et al. 16441, NY). From this voucher, he described the plants as reduced subshrubs to

30 cm tall with clusters of capitula each having 30-70+ pistillate florets and 11-25 hermaphroditic disc florets.

He described the cypselas as setose with elongate trichomes, a condition very different from that described for

the type by Blake, i.e., minutely hispidulous with conical, bluntish, few-celled, dark-based hairs (Blake 1937).

An examination of the holotype of Gnaphalium stolonatum (A.F. Skutch 1098, GH) has shown it to be distinct

from other collections from the same general region annotated as G. stolonatum. Gnaphalium stolonatum is here

treated as a Pseudognaphalium Kirp. and transferred to that genus. The other morphologically different collec-

J. Bot. Res. Inst. Texas 9(1): 63 - 73.2015

Journal of the Botanical Research Institute of Texas 9(1)

tions (e.g., A. Molina et al. 16441) are described here as a new species of Gnaphaliothamnus from the Sierra de los

Cuchumatanes.

In the ultimate offering from H. Robinson (2015), he relegates Pseudoligandra M.O. Dillon & Sagast. (1990)

and Gnaphaliothamnus to the synonymy of Chionolaena. He states that the publications by Freire (1993) and

Nesom (2001) have “totally resolved that problem by reducing all three genera to synonymy under the name

Chionolaena .” While we have been unsuccessful in extracting DNA from the samples representing Pseudoligan¬

dra at our disposal, we are confident that our nrDNA results have more definitively resolved the problem of the

recognition of Gnaphaliothamnus as distinct from Chionolaena.

TAXONOMY

Gnaphaliothamnus nesomii M.O. Dillon & Luebert, sp. nov. (Fig. 1). Type, guatemata. Huehuetenango: common on

moist bank along road to San Juan Ixcoy, Sierra Cuchumatanes, 12-23 Jan 1966, A. Molina R., W.C. Burger, and B. Wallenta 16441

(holotype: F1637117; isotype: NY, n.v.)

Similar to Gnaphaliothamnus salicifolius (Bertol.) G.T. Nesom but habit considerably smaller (to 15 cm tall), capitulescences glomerulate,

obscured in dense arachnoid-tomentose indumentum, cypselar trichomes ca. 100 pm long

Subshrubs, 5-10(-15) cm tall; stems ascending, basally-branched from lignihed base. Leaves sessile, oblan-

ceolate to spatulate, 10-42 mm long, 2-4 mm wide, abaxial surfaces white-tomentose, adaxial surfaces darker,

weakly arachnoid-tomentose, apices acute, apiculate, proximal leaves marcescent. Capitulescences densely

glomerulate with 7-ll(-20) tightly grouped capitula, pedicles obscured by dense arachnoid-tomentose indu¬

mentum. Capitula 5-7 mm long, 2-3 mm diam.; involucres narrowly campanulate, graduated, submerged in

dense arachnoid-tomentose indumentum; phyllaries 4-6-seriate, the outer with arachnoid-tomentose bases;

the inner with white-opaque, oblong apical lamina, 5-7 mm long, ca. 2 mm wide; outer florets 20-32, pistillate,

fertile; central disc florets 13-15, hermaphroditic, ovary sterile. Cypselas oblong, ca. 1 mm long, trichomes

4-celled, clavate, ca. 100 pm long, the apical cells myxogenic, not rupturing (Fig. 2A); pappus of ca. 20, setose

bristles, 2.8-3.2 mm long, apical cells of bristles of hermaphroditic florets expanded, apical cells of pistillate

florets obtuse, not expanded.

Etymology.—Gnaphaliothamnus nesomii is dedicated to Dr. Guy L. Nesom, noted synanthrologist and

systematist, one of the first botanists to accept Gnaphaliothamnus in modern usage, and first to provide a de¬

tailed revision for Mexican and Central American taxa. Over the years, Dr. Nesom has been generous with his

detailed knowledge of Mexican and Central American Asteraceae and particularly the Gnaphalieae. He also

commented on the uniqueness of some of the sheets examined within his loan of material from Field Museum

in 1990.

Distribution and Ecology. —The type locality is within the Sierra de los Cuchumatanes in the Department

of Huehuetenango, Guatemala [15°3TS, 91°32'W]. This is the highest non-volcanic mountain range in Central

America and has the most extensive highlands above 3000 m. The region is home to a variety of different bi-

omes, including montane pine-oak forest, intermittent shrublands, and grasslands. Gnaphaliothamnus nesomii

is found in alpine habitats at ca. 3700 m, notably different from the environments associated with the “llanos”

or plains where Gnaphalium stolonatum occurs (see below).

Conservation status.—Gnaphaliothamnus nesomii deserves a preliminary status of Critically Endangered

(CR) because total area of its known distribution is less than 100 km 2 and only three populations are known

(IUCN 2001).

Discussion. —Recent molecular phylogenetic studies (nrDNA) have provided results that suggest the great

majority of New World genera of Gnaphalieae are associated with the Eucilia- group within the FLAG clade

(Freire et al. 2015; Luebert et al., unpubl.). The Eucilia- group has been expanded to include Gnaphaliothamnus,

Chionolaena, and Antennaria Gaertn. from the subtribe Cassiniinae Anderb. (Anderberg 1991). These results

support the hypothesis that the gross morphological similarity between Gnaphaliothamnus and Chionolaena is

convergence; they are found in distant well-supported clades within the Eucilia- group (Fig. 3). Gnaphaliotham¬

nus is phylogenetically related to Antennaria, while Chionolaena appears to be sister to a group including the

type species of Eucilia Cass., which is in agreement with their similarity in cypselar trichomes.

Dillon and Luebert, A new species of Gnaphaliothamnus from Guatemala

GUATEMALA

Field Museum of Natural History

16441 Escuela Agricola Panamericana

Gnaphalium stolonsturn Blake

det, AmolinaK,

1637117

CHICAGO

NATURAL HISTORY

MUSEUM

HOLOTYPE

Gnaphaliothamnus nesomii M.O.Dillon & Luebert

sp.nov.

FIELD MUSEUM OF NATURAL HISTORY |F|

FIs. white, common on moist bank along

road to San Juan Ixcoy, Sierra Cuehumatanes,

Dept. Huehuetenango.

Alt. J?00 m.

Jan. 12-23, 1966

Fig. 1. Gnaphaliothamnus nesomii. Photograph of holotype collection, A. Molina R. etal. 16441 (FI 637117, imgV0093984F).

Journal of the Botanical Research Institute of Texas 9(1)

Fig. 2. Cypselar trichomes, white scale bar = 100 pm. A. Gnaphaliothamnus nesomii (FI637117), B. 6. salicifolius (FI639348), C. Pseudognaphalium

stolonatum (GH00008351), D. P. antennarioides (F1148157).

Nesom (1990a) provided a detailed taxonomic history for Gnaphaliothamnus and his rationales for its ac¬

ceptance (Nesom 1990b, 1994). Interestingly, Nesom also discussed the putative relationships for Gnaphalium

stolonatum as outside of Gnaphaliothamnus (Nesom 1990a, p. 367). He commented that this taxon did have

white-tipped phyllaries, but it had more florets per capitulum in general (+100) and fertile central disc florets,

Dillon and Luebert, A new species of Gnaphaliothamnus from Guatemala

0.93 1— Micropsis

■Lucilia E S. America

^ Facelis

0.99 1— Lucilia Andes

11 Belloa

Chionolaena

i— Gamochaeta

Gamochaetopsis

Stuckertiella

Lucilioclir

Mniodes

i|— Luciliocline

i P— Mnit

Loricaria

0 6 0.67 [ —4 Gnaphaliothamnus

' - Diaperia

-4 Chevreulia

Antennaria

-^Outgroups

Fig. 3. Simplified dadogram of the Ii/c/7/0-group derived from preliminary result of a Bayesian analysis of Gnaphalieae with nrDNA sequence d

(ITS+ETS) carried out in MrBayes v.3.1.2 (Ronquist & Huelsenbeck 2003). The numbers above branches are Bayesian posterior probabilities. The resi

are congruent with those reported by Freire et al. (2015). Chionolaena and Gnaphaliothamnus do not form a dade.

Journal of the Botanical Research Institute of Texas 9(1)

Table 1. Comparison of salient morphological characters in the species of Gnaphaliothamnus and Pseudognaphalium discussed here. Cypselar trichomes are

illustrated in Figure 2.

Character

G. nesomii

G. salicifolia

P. stolonatum

P. antennarioides

Habit

subshrubs to 15 cm

subshrubs to 100 cm

perennial herbs

Stolons

none

present

Capitula height

5-7 mm

6-7

4.5-5.5 mm

6-7 mm

Pistillate florets

20-32

(22-)34-55

44-92

(50-)75-190

Hermaphroditic florets

13-15

3-4(-7)

7-14

7-16

Cypselar trichomes

ca. 100 pm long, Fig. 2A

62-78 pm long, Fig. 2B

38-40 pm long, Fig. 2C

ca. 50 pm long, Fig. 2D

and suggested it was probably best maintained in Gnaphalium L. s.s. There has been acceptance of Pseudog¬

naphalium and the majority of New World species previously classified as Gnaphalium were transferred there

(Anderberg 1991).

Gnaphaliothamnus nesomii was originally treated as Gnaphalium stolonatum. In a note from Guy Nesom

(16 Feb 1990) accompanying the return of his Field Museum loan, he mentioned the presence of some un¬

usual or atypical specimens with fewer pistillate florets. When Pruski (2012) encountered what he considered

conspecihc material (A. Molina 16441, NY), he transferred the species to Chionolaena and described its floral

morphology exactly as it is in Gnaphaliothamnus. He related it to other Mexican and Central American species,

including C. eleagnoides Klatt, C. lavandulifolia (Kunth) Benth & Hook, f., C. concinna (A. Gray) Anderb. & S.

E. Freire (as C. mexicana S. E. Freire), and C. salicifolia (Bertol.) G.L. Nesom, all species considered here to be¬

long within Gnaphaliothamnus.

Gnaphaliothamnus nesomii is distinctive among members of Gnaphaliothamnus. It has a dwarf woody veg¬

etative habit with stems only 5-10(-15) cm long, the basal leaves are marcescent and cloaking the lower por¬

tions of the stems, and the glomerulate capitulescences with the heads immersed in dense, white, arachnoid-

tomentose indumentum, is a combination of characteristics unmatched in the genus. It has oblanceolate to

spathulate leaves to 42 mm long, white-opaque phyllary apices, and capitula with 20-32 pistillate florets, and

13-15 hermaphroditic florets, Nesom’s (1990a) key would lead to G. salicifolius; which is the only other mem¬

ber of Gnaphaliothamnus to be recorded from Guatemala, and it has a widespread distribution extending north

from Guatemala to central Mexico.

Gnaphaliothamnus salicifolius has a distribution quite unlike all other species, which tend to be narrowly

endemic and geographically restricted (Nesom 1990a,b, 1994). In Guatemala, it inhabits the edge of pine-oak

forests at about 3000 m and is also found in the Sierra de los Cuchumatanes, recorded on the road between

Paquix and San Juan Ixcoy (A. Molina R. 21293, F1661761; A. Molina R. et al. 30031, F1734420; A. Molina R. et al.

16550, F1639348) and N of Santa Eulalia (F. Almeda andJ.L. Luteyn 1686, F1733978).

Table 1 allows for comparison of the salient morphological characteristics of three of the Gnaphalieae

taxa known to occur in region around the Sierra de los Cuchumatanes.

Additional material examined: GUATEMALA. Huehuetenango: Sierra Cuchumatanes between Paquix and San Juan Ixcoy, 8 Jan 1974,

3000-3350 m, A. Molina R., A.R. Molina, andJ.A. Molina 30055 (F1734422); between Tojquia and Caxin bluff, summit of Sierra de los Cuchu¬

matanes, 3700 m, 6 Aug 1942,J.A. Steyermark 50159 (F1148150).

KEY TO GNAPHALIOTHAMNUS SPECIES

The following annotated key will allow for identification of Gnaphaliothamnus species (adapted from Nesom

1990a):

1. Inner phyllaries lacking prominent, white lamina; pappus bristles monomorphic (Chiapas - Mexico)_ G. cryptocephalus

G.L. Nesom

1. Inner phyllaries with prominent, white lamina.

2. Adaxial leaf surfaces with stipitate glandular trichomes beneath the arachnoid-tomentum.

3. Pistillate florets 5-10; pappus bristles strongly dimorphic, caducous; cypselas glabrous (Costa Rica)_ G. costaricensis

G.L. Nesom

Dillon and Luebert, A new species of Gnaphaliothamnus from Guatemala

3. Pistillate florets 12-24; pappus bristles monomorphic to weakly or strongly dimorphic, basally persistent; cypselas

with trichomes.

4. Pistillate florets 12-18, usually equal the number of hermaphroditic; pappus bristles strongly dimorphic (Oaxaca

- Mexico)_ G. macdonaldii G.L. Nesom

4. Pistillate florets 21-24, usually about twice as many as the hermaphroditic; pappus bristles monomorphic to very

weakly dimorphic (Veracruz, Puebla, Tlaxcala, Morelos - Mexico)_ G. lavandulifolius (Kunth) G.L Nesom

2. Adaxial leaf surfaces arachnoid-tomentose to glabrate, eglandular.

5. Leaves 7-8 mm long; phyllaries subequal (Oaxaca - Mexico)_ G. sartorii (Klatt) G.L. Nesom

5. Leaves shorter than 5 mm or longer than 10 mm long; phyllaries strongly graduated.

6. Heads few in tight clusters at tips of leafy stems.

7. Plants dioecious; leaves 2.5-5 mm long; phyllaries with reddish midregion (Oaxaca - Mexico)_ G. aecidiocephalus

(Grierson) G.L. Nesom

7. Plants polygamodioecious; leaves 10-20 mm long; phyllaries without a red midregion (San Luis Potosi -

Mexico)_ G. concinnus (A. Gray) G.L. Nesom

6. Heads numerous in corymbs above the leaves.

8. Leaves elliptic to elliptic-oblanceolate; pappus bristles weakly dimorphic.

9. Leaves elliptic to elliptic-oblanceolate, 15-42 mm long, 4-8 mm wide (Hidalgo, Oaxaca - Mexico)

_G. eleagnoides (Klatt) G.L. Nesom

9. Leaves elliptic-obovate, 10-20 mm long, 3-5 mm wide (Durango - Mexico)_ G. durangensis G.L. Nesom

8. Leaves linear to oblanceolate or spathulate; pappus bristles strongly dimorphic.

10. Leaves 20-80 mm long, 1 —3(—5) mm wide; pistillate florets (22-)34-55; hermaphroditic florets 3-4(-7)

(Guatemala, Mexico)_ G. salicifolius (Bertol.) G.L. Nesom

10. Leaves 10-42 mm long, 2-4 mm wide; pistillate florets 20-32; hermaphroditic florets 13-15 (Guatemala)

_G. nesomii M.O. Dillon & Luebert

NEW COMBINATIONS IN PSEUDOGNAPHALIUM

Pseudognaphalium stolonatum (S.F. Blake) M.O. Dillon, comb. nov. (Fig. 4). Gnaphalium stolonatum S.F. Blake, Brit-

tonia 2:341. 1937. Type: GUATEMATA. Huehuetenango: llanos of the Sierra Cuchumatanes, 10,500 ft [3200 m], 24 Aug 1934, A. E.

Skutch 1098 (holotype: GH00008351; isotype: TEX-LL00373732, n.v.).

Chionolaena stolonata (S.F. Blake) Pruski, Phytoneuron 2012-1:4. 2012.

Perennial herbs 10-25 mm tall, roots fibrous; stolons from the base, filiform; stems erect, usually simple,

rarely branched, thinly arachnoid-tomentose, purplish. Leaves oblanceolate to linear, 2-4.5 mm long, 0.5-1

mm wide, abaxial surfaces gray with dense tomentose indumentum, adaxial surfaces obscurely stipitate-

glandular under thinly arachnoid-tomentose indumentum, apices apiculate, bases slightly decurrent. Capit-

ulescences glomerulate, crowded at apices. Capitula campanulate, sessile, 4.5-5.5 mm tall, 4.5-5.5 mm diam.;

phyllaries ca. 4-seriate, graduated, thinly arachnoid-tomentose; outer ovate to oblong-lanceolate, obtuse to

acute, inner narrowly lanceolate, narrowed to an obtuse apex; pistillate florets 85-92, the corollas filiform,

white, ca. 2.5 mm long, pappus bristles ca. 3 mm long, apices acute; hermaphroditic florets (T—)11(—14), the

corollas cylindrical, white with rose-color lobes, the styles apically subtruncate, ovary sterile; pappus bristles

ca. 3 mm long, apically obtuse, slightly expanded. Cypselas ca. 1 mm long, minutely hispidulous, the tri¬

chomes clavate, 38-40 pm long (Fig. 2C).

Pseudognaphalium stolonatum has been collected within the Sierra de los Cuchumatanes from alpine

meadows on rock outcrops, 3200-3750 m. It was included in Gnaphalium s.s. by Anderberg (1991). Inspection

of the type has shown it to have floral morphology consistent with Pseudognaphalium (Anderberg 1991).

Blake (1937) specifically related Pseudognaphalium stolonatum to P. antennarioides (DC.) Anderb., a rosu-

late, stoloniferous herb distributed from Colombia to Bolivia with capitula having (50-)75-190 pistillate and

7-16 hermaphroditic florets. Phylogenetic studies have shown that Pseudognaphalium is a member of the HAP

clade (Smissen et al. 2011; Nie et al. 2013; Galbany-Casals et al. 2014), along with Achyrocline (Less.) DC., He-

lichrysum Mill., and Anaphalis DC.; Gnaphaliothamnus and Chionolaena are members of the FLAG clade, which

is only distantly related (Smissen et al. 2011).

Pseudognaphalium subsericeum (S.F. Blake) Anderb. (= Gnaphalium subsericeum S.F. Blake, 1927), a Costa

Rican endemic, was also mentioned as potentially related to Gnaphaliothamnus salicifolius by Blake (p. 62). An

examination of conspecihc material from Costa Rica (F1692005, F1692006) shows P. subsericeum more simi-

Journal of the Botanical Research Institute of Texas 9(1)

Holotype

Gnaphallurn stolonatum S, F, Blake

Brittonia 2i 341. 1937.

'■iSSiIfir 1983

HARVARD UNIVERSITY HERBARIA

Flora of Guatemala

HOLOTYPE

Gnaphalium stolonatum Blake

Dorothy L. Nash 1974

ctb&srvifi. tlx

j 0 t

ZH; II Jf.

PLANTS OF GUATEMALA y~

DEPT. OF HUEHU6TENANGO

FIs. white.

Xlanos of the Sierra Cuchumatanea

1098.

Coll. Alexander F. Skutch August 24,1934.

Fig. 4. Pseudognaphalium stolonatum. Photograph of holotype collection of Gnaphalium stolonatum S.F.BIake, AF Skutch 1098 (GH000008351).

Dillon and Luebert, A new species of Gnaphaliothamnus from Guatemala

Fig. 5. Pseudognaphaliumparamorum. Photograph of holotype collection of Gnaphaliumparamorum S.F.BIak e,A.Jahn 883 (US1186590, imgOOl 29548).

Journal of the Botanical Research Institute of Texas 9(1)

lar to P. stolonatum or P. antennarioides (DC.) Anderb., the latter an Andean species that has similar overall

morphology and cypselar trichomes (Fig. 2D). All these species have capitula in dense cymes or glomerules

and phyllaries with showy, white apices.

Additional specimens examined: GUATEMALA. Huehuetenango: vicinity of Chemal, summit of Sierra de los Cuchumatanes, 3700-3750

m, 8 Aug 1942,J. A. Steyermark 50253 (F1148157), 50273 (F1148180), 50275 (F1148179).

Pseudognaphalium paramorum (S.F. Blake) M.O. Dillon, comb. nov. (Fig. 5). Basionym: Gnaphalium paramorum S.F.

Blake, J. Wash. Acad. Sci. 21:328. 1931. Type: VENEZUELA. Estado Merida: Paramo Quirora, 2900 m, 24 Feb 1922, A.Jahn 883

(holotype: US1186590).

Gamochaeta paramora (S.F. Blake) Anderb., Opera Bot. 104:157.1991.

Blake (1931) commented that Pseudognaphalium paramorum had the appearance of Gnaphalium antennarioides

DC. (= P. antennarioides (DC.) Anderb.), and he related it to that taxon, which is a rosulate, stoloniferous herb

distributed from Colombia to Bolivia.

Anderberg (1991) transferred Gnaphalium paramorum to Gamochaeta Wedd.; an examination of the type

collection shows it to only superficially resemble Gamochaeta. It has a basal rosette of subspathulate leaves and

unbranched, ascending stems. The terminal capitulescences are not spicate; they are densely glomerulate cor¬

ymbs. The type collection has capitular and floral morphology consistent with Pseudognaphalium; the cypselas

are papillose and lack the 2-celled, sessile trichomes diagnostic for Gamochaeta.

ACKNOWLEDGMENTS

We thank the curators and staff of the herbaria at F, GH, MO, NY, TEX, and US for permitting access to their

material. Especially, we wish to acknowledge Ms. Emily Wood, Collections Manager at the Gray Herbarium,

who expedited loan of type material from that institution. We thank John Strother for many constructive sug¬

gestions in the review process. Andres Moreira-Munoz and Miguel Alvarez provided plant material for phylo¬

genetic studies. Digital images were obtained via the Internet in Figure 4 from GH and Figure 5 from US. We

thank Field Museum’s Christine Niezgoda, Anna Balia, Daniel Le, and Allie Stone for handling various aspects

of loans and digitizing collections.

REFERENCES

Anderberg, A. 1991. Taxonomy and phylogeny of the tribe Gnaphalieae (Asteraceae). Opera Bot. 104:1-195.

Anderberg, A. & S.E. Freire. 1989. Transfer of two species from Anaphalis DC. to Chionolaena DC. (Asteraceae, Gnapha¬

lieae). Notes Roy. Bot. Gard. Edinburgh 46(1 ):37—41.

Blake, S.F. 1927. New Asteraceae from Costa Rica. J. Wash. Acad. Sci. 17(3):59-65.

Blake, S.F. 1931. Nine new American Asteraceae. J. Wash. Acad. Sci. 21(14):325—336.

Blake, S.F. 1937. New Asteraceae from Guatemala and Costa Rica Collected by A.F. Skutch. Brittonia 2(4):329—361.

Dillon, M.O. 2003. New combinations in Luciliocline with notes on South American Gnaphalieae (Asteraceae). Arnaldoa

10(1 ):45—60.

Dillon, M.O. & A. SagAstegui-A. 1991. Sinopsis de los generos de Gnaphaliinae (Asteraceae-lnuleae) de Sudamerica. Ar¬

naldoa 1 (2):5—91.

Dillon, M.O. & A. SagAstegui A. 1990. Oligandra Less. Revisited and the need for a new genus, Pseudoligandra (Asteraceae:

Inuleae).Taxon 39:125-128.

Freire, S.E. 1993. A revision of Chionolaena (Compositae, Gnaphalieae). Ann. Missouri Bot. Gard. 80(2):397-438.

Freire, S.E., M.A. Chemisquy, A.A. Anderberg, S.G. Beck, R.l. Meneses, B. Loeuille, & E. Urtubey. 2015.The Lucilia group (Asteraceae,

Gnaphalieae): phylogenetic and taxonomic considerations based on molecular and morphological evidence. PI.

Syst. Evol. 301:1227-1248. doi: 10.1007/s00606-014-1147-0.

Galbany-Casals, M., M. Unwin, N. Garcia-Jacas, R.D. Smissen, A. Susanna, & R.J. Bayer. 2014. Phylogenetic relationships in He-

lichrysum (Compositae: Gnaphalieae) and related genera: Incongruence between nuclear and plastid phylogenies,

biogeographic and morphological patterns, and implications for generic delimitation. Taxon 63(3):608-624.

IUCN Species Survival Commission. 2001. IUCN Red List Categories and Criteria: Version 3.1. IUCN, Gland, Switzerland and

Cambridge, United Kingdom, 1-30.

Dillon and Luebert, A new species of Gnaphaliothamnus from Guatemala

Hess, R. 1938. Vergleichende Untersuchungen iiber die Zwillingshaare der Compositen. Bot. Jahrb. Syst. 68:435-496.

Loeuille, B., L. Beble, & J.N. Nakajima. 2011. Four new species of Chionolaena (Asteraceae: Gnaphalieae) from southeastern

Brazil. Kew Bull. 66(2):263-272.

Nesom, G.L. 1990a. Taxonomy of Gnaphaliothamnus (Asteraceae: Inuleae). Phytologia 68(5):366-381.

Nesom, G.L. 1990b. An additional species of Gnaphaliothamnus (Asteraceae: Inuleae) and further evidence for the integ¬

rity of the genus. Phytologia 68(6):1-3.

Nesom, G.L. 1994. Comments on Gnaphaliothamnus (Asteraceae: Inuleae). Phytologia 76:185-191.

Nesom, G.L. 2001. New combinations in Chionolaena (Asteraceae: Gnaphalieae). Sida 19(4):849-852.

Nie, Z.-L.,V. Funk, H. Sun,T. Deng,Y. Meng, &J. Wen. 2013. Molecular phylogeny of Anaphalis (Asteraceae, Gnaphalieae) with

biogeographic implications in the Northern Hemisphere. J. Plant Res. 126(1 ):17-32.

Pruski, J.F. 2012. Studies of Neotropical Compositae-IV. Pseudognaphalium leucostegium, a new species from Huehu-

etenango, Guatemala, and a new combination in Chionolaena (Gnaphalieae). Phytoneuron 2012-1:1 -5.

Robinson, H. 2015. Notes on the genus Chionolaena in Colombia with a new species Chionolaena barclayae (Asteraceae,

Gnaphalieae). PhytoKeys 46:67-71, doi: 10.3897/phytokeys.46.8976

Ronquist, F. & J.P. Huelsenbeck. 2003. MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics

19:1572-1574.

Smissen, R.D., M. Galbany-Casals, & I. Breitwieser. 2011. Ancient allopolyploidy in the everlasting daisies (Asteraceae:

Gnaphalieae): Complex relationships among extant clades. Taxon 60(3):649-662.

Journal of the Botanical Research Institute of Texas 9(1)

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J.Bot. Res. Inst. Texas 9(1): 74.2015

NEW COMBINATIONS IN EUMACHIA (RUBIACEAE)

FOR SPECIES OCCURRING ON THE GUIANA SHIELD

Piero G. Delprete

Joseph H. Kirkbride, Jr.

Herbier de Guy one, IRD - UMRAMAP

Boite Postale 165, 97323 Cayenne Cedex

Guyane Frangaise (French Guiana), FRANCE

piero. delprete@ird. fr

USDA-ARS, US. National Arboretum

Floral & Nursery Plants Research Unit

3501 New York Avenue NE

Washington, DC 20002-1958, US.A.

joseph.kirkbride@ars.usda.gov

ABSTRACT

Thirteen new combinations are made in the genus Eumachia for species formerly included in the genus M argaritopsis and occurring on the

Guiana Shield. For these 13 new combinations, their basionyms, other synonyms, types, and distributions are given. Three new combina¬

tions are also made for the types of the genera Chazaliella, Margaritopsis, and Readea, recognized synonyms of Eumachia.

Key Words: Chazaliella, Chytropsis, Margaritopsis, Guayana Shield, Neotropics, Readea, type designations

RESUMEN

Se hacen trece nuevas combinaciones en el genero Eumachia para especies anteriormente incluidas en el genero Margaritopsis , que ocurren

en el Escudo Guyanes. Eos basonimos, otros sinonimos, tipos y distribuciones se presentan para estas trece nuevas combinaciones, se pre-

sentan aqui. Se hacen tambien tres combinaciones nuevas para los tipos de los generos Chazaliella, Margaritopsis y Readea, sinonimos

reconocidos de Eumachia.

Palabras Claves: Chazaliella, Chytropsis, denominaciones de tipos, Margaritopsis, Escudo Guyanes, Neotropicos, Readea

Charles Wright (Sauvalle 1869) published the genus Margaritopsis C. Wright with a single species, M. acuifolia

C. Wright, from eastern Cuba. In the early 20th Century, Urban (1921, 1929) added two more species from

Haiti to the genus, M. lanceifolia Urb. and M. triflora Urb. Until the end of the 20th Century, it was considered

to be a small genus, endemic to the Greater Antilles. Andersson (2001, 2002a, 2002b), using molecular phylog-

enies, demonstrated that Margaritopsis is a pantropical genus, with species in the New World and Old World,

i.e. Africa, Madagascar, Asia, and the Pacific, and synonymized with Margaritopsis two Old World genera,

Chazaliella E.M.A. Petit & Verde, and Readea Gillespie, and one New World genus, Chytropsia Bremek. He es¬

timated that Margaritopsis would have approximately 50 species. Taylor (2005) cited 27 species of Margaritop¬

sis in the Neotropics, nearly three times as many as Andersson had estimated for the New World, and sug¬

gested that there are probably more species of Margaritopsis to be discovered there. Barrabe et al. (2012), using

molecular and morphological analyses, investigated more Asian, Australasian, and Pacific species, and con¬

cluded that more Old World species should be included in Margaritopsis, bringing the number of species to

more than 70 worldwide. Barrabe and Davis (2013) showed that Eumachia DC. is also synonymous with Mar¬

garitopsis, and has priority over it.

Barrabe and Davis (2013) proposed the conservation of Margaritopsis over Eumachia because the latter

has been either treated as a dubious genus or as a synonym of Psychotria s.l. (Robbrecht 1988:240, 1993:186),

and only has a single species, E. carnea (G. Forst.) DC. Margaritopsis was well known among rubiologists, and

at least 27 Margaritopsis species would have to be transferred to Eumachia. The Nomenclature Committee for

Vascular Plants felt that “in view of the small historical size of Margaritopsis and the fact that many species

from Asia and the Pacific, where Margaritopsis has not been considered to occur, must change their names in

any case, a majority of the Committee were not convinced that application of the principle of priority would

substantially reduce stability of nomenclature,” and so did not recommend conservation of Margaritopsis over

Eumachia (Applequist 2014). Since it is expected that this recommendation will be upheld by the General Com¬

mittee, Eumachia is the correct name for the combined genus and must be used instead of Margaritopsis. There-

J. Bot. Res. Inst. Texas 9(1): 75 - 79.2015

Journal of the Botanical Research Institute of Texas 9(1)

fore, thirteen new combinations are made here for the Margaritopsis species occurring on the Guiana Shield

(Funk & Berry 2005). These new combinations are needed for the forthcoming rubiaceous treatment in the

Flora of the Guianas.

SYSTEMATIC TREATMENT

Eumachia DC., Prodr. 4:478. 1830. Type: E. carnea (G. Forst.) DC. [Basionym: Petesia carnea G. Forst., Fl. Ins. Austr. 10.1786]

Margaritopsis C. Wright in Sauvalle, Anales Acad. Ci. Med. Habana 6:146. 1869. Type: M. acuijolia C. Wright in Sauvalle, Anales Acad.

Ci. Med. Habana 6:147.1869. [= Eumachia acuijolia (C. Wright) Delprete & J.H. Kirkbr., comb, nov.]

Readea Gillespie, Bernice R Bishop Mus. Bull. 74:35. 1930. Type: R. membrancea Gillespie, Bernice R Bishop Mus. Bull. 74:35. 1930. [=

Eumachia membranacea (Gillespie) Delprete &J.H. Kirkbr., comb, nov.]

Chytropsia Bremek., Recueil Trav. Bot. Neerl. 31:291. 1934. Psychotria sect. Chytropsia (Bremek.) Steyerm., Mem. New York Bot. Gard.

23:484.1972. Type: C. astrellantha (Wernham) Bremek. [= Eumachia astrellantha (Wernham) Delprete & J.H. Kirkbr.]

Chazaliella E.M.A. Petit & Verde., Kew Bull. 30:267.1975. Type: C. abrupta (Hiern) E.M.A. Petit & Verde. [Basionym: Psychotria abrupta

Hiern in Oliver, Fl. Trop. Afr. 3:205.1877; = Eumachia abrupta (Hiern) Delprete & J.H. Kirkbr., comb, nov.]

1. Eumachia albert-smithii (Standi.) Delprete &J.H. Kirkbr., comb. nov. Psychotria albert-smithii Standi., Publ. Field

Mus. Nat. Hist., Bot. Ser. 8:203. 1930. Margaritopsis albert-smithii (Standi.) C.M. Taylor, Syst. Geogr. Pi. 75:169. 2005. Type: PERU.

Foreto: Soledad, on the Rio Itaya, 110 m, 20-22 Sep 1929, E.P. Killip &A.C. Smith 29766 (holotype: F No. 607482; isotypes: fragments

G barcode G00300217, NY barcode 00132588, US No. 1463026 [barcode 00138632]).

Distribution. —Widespread in South America.

2. Eumachia astrellantha (Wernham) Delprete &J.H. Kirkbr., comb. nov. Psychotria astrellantha Wernham, j. Bot.

52:316. 1914. Chytropsia astrellantha (Wernham) Bremek., Recueil Trav. Bot. Neerl. 31:292. 1934. Margaritopsis astrellantha (Wer¬

nham) F. Andersson, Syst. Geogr. Pi. 72:230. 2002. Type: GUYANA: Potaro River, below Kaieteur Falls, Sep-Oct 1881 (fl), G.S.Jen-

man 959 (holotype: K barcode K000173648).

Rudgea obscura Sandw., Bull. Misc. Inform. Kew 1931:474. 1931. Type: GUYANA: Cuyuni River, below Akaio Falls, 22 Nov 1929 (fl, fr),

N.Y. Sandwith 647 (holotype: K barcode K000173646; isotypes: K barcode K000173647, NY barcode 00133225).

Distribution. —Southern Venezuela, Guyana, Suriname, French Guiana, and throughout Brazil.

3. Eumachia boliviana (Standi.) Delprete & J.H. Kirkbr., comb. nov. Psychotria boliviana Standi., Publ. Field Mus. Nat.

Hist., Bot. Ser. 7:302. 1931. Margaritopsis boliviana (Standi.) C.M. Taylor, Syst. Geogr. Pi. 75:170. 2005. Type: BOFIVIA. Fa Paz: San

Carlos, region of Mapiri, 850 m, 3 Nov 1926 (fl), O. Buchtien 1489 (holotype: F No. 609011; isotype: F No. 611732).

Psychotria lawrancei Standi., Publ. Field Mus. Nat. Hist., Bot. Ser. 17:281.1937. Type: COFOMBIA. BoyacA: region of El Humbro, 1050 m,

6 May 1933 (fl), A.E. Lawrance 788 (holotype: F No. 681079; isotypes: E barcode E00285008, F No. 1525171, G barcode G00300182,

G barcode G00300183, MO No. 1068126, S No. 05-1091).

In JSTOR Global Plants (2015) there is a specimen from WIS, barcode 00001028 MAD, with the label “Fawrance 788 - Psychotria

lawrancei, new species” and another label “Isotype Psychotria lawrancei Standi.” While the specimen label corresponds to that of

the holotype, the specimen belongs to the family Fauraceae, and is not an isotype.

Psychotriafoetidiflora Standi., Publ. Field Mus. Nat. Hist., Bot. Ser. 22:202.1940. Type: BRAZIF. Amazonas: Parintins, 22 Dec 1935 (fl), A.

Ducke s.n. (RB 34998) (holotype: F No. 932981; isotype: K barcode K000173649).

Psychotria kukenanensis Steyerm., Fieldiana, Bot. 28:601. 1953. Type: VENEZUEFA. Bolivar: Gran Sabana, Rio Kukenan, between wa¬

terfall at Rue-Meru (tributary of Rio Kukenan) and Divina Pastora, on Rio Kukenan N of Santa Helena, S of Mount Roraima, 1065

m, 3 Oct 1944 (fl),J.A. Steyermark 59227 (holotype: F No. 1181464; isotype: US No. 1908727 [barcode 00138821]).

Psychotria turboensis Standi, ex Steyerm., Acta Bot. Venez. 4:101, fig. 49. 1964. Type: COFOMBIA. Antioquia: N of Dabeiba, on road to

Turbo, 350 m, 26 Feb 1942 (fl), R.D. Metcalf &j Cuatrecasas 30119 (holotype: F No. 1124872; isotypes: MO No. 1272932, US No.

1833446 [barcode 00146645]).

Psychotriapuberulenta Steyerm., Mem. New York Bot. Gard. 23:488.1972. Type: GUYANA: Pakaraima Mountains, Upper Mazaruni River,

Sagaraimadai Savanna, 550 m, 16 Nov 1951 (fl), B. Maguire & D.B. Fanshawe 32618 (holotype: NY barcode 00132790; isotypes: US

No. 3035272 [barcode 00138942], VEN No. 82306).

Psychotria plowmanii Steyerm., Ann. Missouri Bot. Gard. 71:1178. 1984. Type: VENEZUEFA. Amazonas: Depto. Atures, along Rio Cat-

aniapo, 44-45 km SE of Puerto Ayacucho, 3 km downstream from damsite, 5°35'N, 67°15'W, 200-300 m, 9 May 1980 (fl), J.A.

Steyermark, G. Davidse&F. Guanchez 122132 (holotype: VEN No. 136287; isotype: MO No. 3100818).

Psychotria ronaldii Steyerm., Ann. Missouri Bot. Gard. 74:111, fig. 10.1987. Type: VENEZUEFA. Amazonas: Depto. Atabapo, Cerro Hua-

chamacari, 3°39'N, 65°42'W, 600-750 m, 4 Mar 1985 (fl), R.L. Liesner 18214 (holotype: MO No. 3327368; isotype: VEN No. 24833).

Delprete and Kirkbride, New combinations in Eumachia

Distribution. —Colombia, Ecuador, Peru, Bolivia, southern Venezuela, Guyana, Suriname, and French Guiana.

4. Eumachia cephalantha (Mull. Arg.) Delprete &J.H. Kirkbr., comb. nov. Mapouria cephalantha Mull.Arg., Flora

59:495,497.1876. Uragoga cephalantha (Mull. Arg.) Kuntze, Revis. Gen. Pi. 2:959.1891. Psychotria cephalantha (Mull. Arg.) Standi.,

Field Mus. Nat. Hist., Bot. Ser. 11:235.1936. M argaritopsis cephalantha (Mull. Arg.) C.M. Taylor, Syst. Geogr. Pi. 75:171. 2005. Type:

BRAZIT. Minas Gerais: in sylvis prope Tagoa Santa, s.d. ,J.E.B. Warming s.n. (lectotype (Andersson 1992:138): C n.v.; isolectotype: F

No. 667820).

Mapouria capituliflora Mull. Arg., Flora 59:495, 497. 1876. Uragoga capituliflora (Mull. Arg.) Kuntze, Revis. Gen. Pi. 2:959. 1891. Psy¬

chotria capituliflora (Mull. Arg.) Standi., Field Mus. Nat. Hist., Bot. Ser. 11:235.1936. Type: BRAZIT. Rio deJaneiro: without locality,

s.d., H.W. Schott s.n. (lectotype, here designated: BR barcode BR0000008257598).

Distribution. —Amazonian Colombia through Peru to Bolivia, and from northern (including Para) to southern

Brazil and Paraguay.

Muller (1876:495,497), along with the description of Mapouria capituliflora Mull. Arg., cited two syntypes,

Schotts.n.andSeIIows.n.,collectedintheBrazihanstateofRiodeJaneiro. Specimen BRbarcode 0000008257598,

has a label with “Brasilia, Schott”, author unknown, and is here selected as the lectotype of this name.

5. Eumachia deinocalyx (Sandwith) Delprete &J.H. Kirkbr., comb. nov. Psychotria deinocalyx Sandwith, Bull. Misc.

Inform. Kew 1939:554.1939. M argaritopsis deinocalyx (Sandwith) C.M. Taylor, Syst. Geogr. Pi. 75:172. 2005. Type: GUYANA: Kibi-

hiu Creek, Wiruni River, Berbice River, 9 Feb 1938 (fl), D.B. Fanshawe 39 (Forest Department of British Guiana 2683) (holotype: K

barcode K000173645).

Distribution. —Southern Venezuela, Guyana, Suriname, French Guiana, and Amazonian Brazil.

6. Eumachia guianensis (Bremek.) Delprete &J.H. Kirkbr., comb. nov. Chytropsia guianensis Bremek., Recueil Trav. Bot.

Neerl. 31:292. 1934. Psychotria moroidea Steyerm. (nom. nov. for Chytropsia guianensis), Mem. New York Bot. Gard. 23:485. 1972,

non Psychotria guianensis (Aubl.) Raeusch. 1797. M argaritopsis guianensis (Bremek.) C.M. Taylor, Syst. Geogr. Pi. 75:172.2005. Type:

SURINAME: Emma Range, 600 m, 15 Mar 1922 (fl), J.W. Gonggrijp & G. Stahel s.n. (B.W. 5667) (lectotype (Steyermark 1972: 485,

first-step; second-step lectotype, here designated): U!; isolectotype: BM barcode BM001009040).

Bremekamp (1934:292) cited the types of Chytropsia guianensis Bremek. as “Flab. Emma Range, Gonggrijp et

Stahel 5667. Upper Nickerie River, Tulleken 451 co-type:” Steyermark (1972:485) cited the type of Chytropsia

guianensis Bremek. as “Emma Range, Suriname, Gonggrijp & Stahel 56675 Although he did not cite a herbari¬

um, Steyermark’s citation is a first-step lectotypihcation, and the U specimen is here designated as the second-

step lectotype of this name.

Distribution. —Guyana, Suriname, French Guiana, and northern Brazil (Amapa, Para, Mato Grosso).

7. Eumachia kappleri (Miq.) Delprete & J.H. Kirkbr., comb. nov. Campichea kappleri Miq., Stirp. Surinam. Select. 181.

1851. Uragoga kappleri (Miq.) Pulle, Enum. Vase. Pi. Surinam. 446.1906. Psychotria kappleri (Miq.) Mull. Arg. exBenoist, Bull. Soc.

Bot. France 68:140.1921. Cephaelis kappleri (Miq.) Standi., Publ. Field Mus. Nat. Hist., Bot. Ser. 4:335.1929. M argaritopsis kappleri

(Miq.) C.M. Taylor, Syst. Geogr. Pi. 75:175. 2005. Type: SURINAME: Marowyne River, Sep 1847 (fr), A. Kappler 1871 (holotype: U!;

isotype: G barcode G00300293!, photo-MO).

Distribution. —Costa Rica, Colombia, Venezuela, Guyana, Suriname, French Guiana, and northern and north¬

eastern Brazil.

8. Eumachia microdon (DC.) Delprete & J.H. Kirkbr., comb. nov. Rondeletiamicrodon DC., Prodr. 4:408.1830. Psychotria

microdon (DC.) Urb., Symb. Antill. 9: 539.1928. Mapouria microdon (DC.) Bremek., Recueil Trav. Bot. Neerl. 31:287.1934. M argari¬

topsis microdon (DC.) C.M. Taylor, Syst. Geogr. Pi. 75:169. 2005. Type: CUBA: Fa Havana: Without locality, 1825 (fl),J.A. de la Ossa

s.n. (holotype: G-DC1; isotype: fragments F No. 635330).

Psychotria pinularis Sesse & Moq, Fl. Mex., ed. 2, 57. 1894. Type: PUERTO RICO: “in monticulo Porto de la Aguadilla, s.d., M. Sesse &

J.M. MoQiho s.n. (lectotype, here designated: MA barcode MA605231).

Psychotria microdon var. meridionalis Steyerm., Mem. New York Bot. Gard. 23:446. 1972. Type: VENEZUETA. Carabobo: Hacienda de

Cura, near Sanjoaquin, 480-1200 m, 4Jul 1916 (fl), H. Pittier 7911 (holotype: VEN; isotype: F No. 587678).

On MA barcode 605231 there is a handwritten label (author unknown), “S-l Psychotria annularis [sic], N. Ic.

1338” and a typewritten label, “Herbarium Horti Botanici Matritensis, Plantae Novae Hispaniae a Sesse,

Journal of the Botanical Research Institute of Texas 9(1)

Mocino, Castillo et Maldonado lectae (1787-1795-1804)” with a handwritten identification by Standley, “Psy-

chotria microdon (DC.) Urban.” It is here selected as lectotype of Psychotria pinularis Sesse & Mog.

Distribution. —Antilles, Mexico to Peru, Venezuela, and Suriname.

9. Eumachia nana (K. Krause) Delprete &J.H. Kirkbr., comb. nov. Psychotria nana K. Krause in ule, Verh. Bot. Vereins

Prov. Brandenburg 50:109.1908. M argaritopsis nana (K. Krause) C.M. Taylor, Syst. Geogr. Pi. 75:175.2005. Type: BRAZIT. Amazonas:

“Rio Jurua superioris, am Jurua Miry” [Upper Rio Jurua, on the Rio Jurua Mirim (affluent)], Aug 1901, E. Ule 5670 (holotype: B de¬

stroyed, photo [F neg. #467]; lectotype, here designated: K barcode K000173644; isolectotypes: F No. 895487, G barcode

G00300185, T barcode T0058073).

Psychotria nudiceps Standi, Publ. Field Mus. Nat. Hist., Bot. Ser. 8:378. 1931. Type: PERU: San Martin: Tarapoto, Alto Rio Huallaga,

360-900 m, 21 Feb 1930 (fl), Ll. Williams 6600 (holotype: F No. 614571; isotypes: G barcode G00300184, S No. 05-1103, US No.

1496734 [barcode 00138889]).

As the type specimen at B was destroyed, Taylor (2005:175) stated “MO holo-; ...” Taylor’s statement can be

interpreted as a lectotypibcation, but she did not include the expression “here designated.” According to article

7.10 of the ICNafp (McNeill et al., 2012), “For purposes of priority (Art. 9.19, 9.20, and 10.5), designation of a

type is achieved only if the type is definitely accepted as such by the typifying author, if the type element is

clearly indicated by direct citation including the term “type” (typus) or an equivalent, and, on or after 1 January

2001, if the typihcation statement includes the phrase “designated here” (hie designatus) or an equivalent.”

Therefore, Taylor’s lectotypibcation is not valid. The specimen at K with barcode K000173644 is here desig¬

nated as lectotype.

Distribution. —Peru, Bolivia, southern Venezuela (Amazonas, Apures), and Amazonian Brazil (Acre, Ron-

donia, Amazonas, Para).

10. Eumachia pallidinervia (Steyerm.) Delprete &J.H. Kirkbr., comb. nov. Psychotria pallidinervia Steyerm., Mem.

New York Bot. Gard. 23:492, fig. 68. 1972. Margaritopsis pallidinervia (Steyerm.) C.M. Taylor, Syst. Geogr. Pi. 75:175. 2005. Type:

VENEZUETA. Amazonas: trail between Camp 2 and Camp 3, Rio Yatua, Cerro de la Neblina, 500-700 m, 7 Nov 1957 (fl), B. Maguire,

J. Wurdack, & C.K. Maguire 41996 (holotype: NY barcode 00132763; isotypes: COT barcode 000004667, K barcode K000432835, U

No. 256671 [barcode U0006229], US No. 2575553 [barcode 00138905], VEN No. 81954).

Distribution. —Southern Venezuela.

11. Eumachia paupertina (Standi. & Steyerm.) Delprete &J.H. Kirkbr., comb. nov. Psychotriapaupertina Standi. &

Steyerm., Fieldiana, Bot. 28:603.1953. Margaritopsis paupertina (Standi. & Steyerm.) C.M. Taylor, Syst. Geogr. Pi. 75:175.2005. Type:

VENEZUETA. Amazonas: Near Base River (Cano Negro), at southeastern base of Cerro Duida, 215 m, 23 Aug 1944 (fl), J.A. Steyer-

mark 57919 (holotype: F No. 1181470; isotypes: NY barcode 00132773, VEN No. 15878).

Distribution. —Southern Venezuela.

12. Eumachia podocephala (Mull.Arg.) Delprete &J.H. Kirkbr., comb. nov. Mapouriapodocephala Mull.Arg., Flora 59:

460, 466. 1876. Psychotria podocephala (Mull. Arg.) Standi., Publ. Field Mus. Nat. Hist., Bot. Ser. 7:109. 1930. Margaritopsis podo¬

cephala (Mull.Arg.) C.M. Taylor, Syst. Geogr. Pi. 75:176. 2005. Type: VENEZUETA. Amazonas: San Carlos do Rio Negro, s.d. [1853—

1854], R. Spruce3076 (lectotype (Steyermark 1972:490, first-step; second-step lectotype, here designated): K barcode K00173642;

isolectotypes: G barcode G00300186, K barcodes K000173641 & K000173643, NY barcode 00132228 , P [2 sheets] barcodes

P02285223 & P 02428065, W barcode 0014055 n.v. [photo F neg. #31173]).

Cephaelis caruruensis Standi, ex Steyerm., Acta Biol. Venez. 4: 11. 1964. Type: COTOMBIA. Vaupes: Casa Alvarez, Bocas Caruru, 230 m,

26 Sep 1939 (fl), J- Cuatrecasas 7019 (holotype: US No. 1795039 [barcode 00129800]; isotype: COT No. 15768 [barcode 000004569]).

Taylor (2005: 176) stated “R. Spruce 3076 [presumably M n.v. lecto-; selected by Steyermark 1972 p. 490; ...”

Steyermark (1972: 490) cited the type gathering, but did not designate a single specimen as the lectotype. Tay¬

lor designated the specimen at M as lectotype, but failed to state “designated here”, so her second-step lecto-

typiheation is not valid. The specimen K barcode K00173642 is here designated as second-step lectotype be¬

cause it has three branches, two with complete inflorescences and one with a complete infructescence.

Distribution. —Colombia, Peru, southern Venezuela, and Amazonian Brazil.

Delprete and Kirkbride, New combinations in Eumachia

13. Eumachia wilhelminensis (Steyerm.) Delprete &J.H. Kirkbr., comb. nov. Psychotria wilhelminensis Steyerm.,

Mem. New York Bot. Gard. 23:487, fig. 67. 1972. M argaritopsis wilhelminensis (Steyerm.) C.M. Taylor (as “M argaritopsis wilhelmen-

sis ”), Syst. Geogr. Pi. 75:176. 2005. Type: SURINAME: 3 km S of Juliana Top, 12 km N of Tucie River, Wilhelmina Gebergte, 3°39'N,

56°32'W, 300 m, 24 Aug 1963 (fl, fr), H.S. Irwin, G.T. Prance, T.R. Soderstrom&N. Holmgren 55048 (holotype: NY barcode 00132865;

isotypes: F No. 1704829, US [2sheets] Nos. 2579088 & 2575552 [barcodes 00642657 & 00129578], VEN No. 82114).

Distribution. —Suriname and northern Brazil (Para).

ACKNOWLEDGMENTS

We are very grateful to John Wiersema (USDA) for revising the manuscript before submission, and Steven P.

Darwin and one anonymous reviewer for their comments and corrections.

REFERENCES

Andersson, L. 1992. A provisional checklist of neotropical Rubiaceae. Scripta Bot. Belg. 1:1—199.

Andersson, L. 2001. Margaritopsis (Rubiaceae, Psychotrieae) is a pantropical genus. Syst. Geogr. PI. 71:73-85.

Andersson, L. 2002a. Relationships and generic circumscription in the Psychotria complex (Rubiaceae, Psychotrieae). Syst.

Geogr. PI. 72:167-202.

Andersson, L. 2002b. Validation of three new combinations in Margaritopsis (Rubiaceae, Psychotrieae). Syst. Geogr. PI.

72:230.

Applequist, W.L. 2014. Report of the Nomenclature Committee for Vascular Plants: 66. Taxon 63:1358-1371.

BarrabE, L., S. Buerki, A. Mouly, A.P. Davis, J. Munzinger, & L. Maggia. 2012. Delimitation of the genus Margaritopsis

(Rubiaceae) in the Asian, Australasian and Pacific region, based on molecular phylogenetic inference and morphol¬

ogy. Taxon 61:1251-1268.

BarrabE, L. & A.P. Davis. 2013. (2207) Proposal to conserve the name Margaritopsis against Eumachia (Rubiaceae). Taxon

62:1069-1070.

Bremekamp, C.E.B. 1934. Notes on the Rubiaceae of Surinam. RecueilTrav. Bot. Neerl. 32:248-305.

Funk, V.A. & P.E. Berry. 2005. The Guiana Shield. In: G.A. Krupnick & WJ. Kress, eds. Plant Conservation: A natural history

approach. University of Chicago Press, Chicago, U.S.A. Pp. 76-79.

JSTOR Global Plants. 2015.The JSTOR Global Plant Initiative, https://plants.jstor.org/. Accessed 02 Jan 2015.

Muller, J. 1876. Rubiaceae brasilienses novae. Flora 59:449-466.

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

Prado, W.F. Prud'homme van Reine, G.F. Smith, J.H. Wiersema, & NJ. Turland. 2012. International Code of Nomencla¬

ture for algae, fungi, and plants (Melbourne Code). Regnum Veg. 154:1-208.

Robbrecht, E. 1988. Tropical woody Rubiaceae. Characteristic features and progressions. Contributions to a new

subfamilial classification. Opera Bot. Belg. 1:1-271.

Robbrecht, E. 1993 [1994]. Supplement to the 1988 outline of the classification of the Rubiaceae. Index to Genera. In: E.

Robbrecht, ed. Advances in Rubiaceae macrosystematics. Opera Bot. Belg. 6:173-196.

Sauvalle, F.A. 1869. Flora Cubana: Revisio catalogi Grisebachiani vel index plantarum Cubensium. Anales Acad. Ci. Med.

Fis. Nat. Habana 6:146-150.

Steyermark, J.A. 1972. Rubiaceae. In: B. Maguire and collaborators. Botany of the Guayana Highland - Part IX. Mem. New

York Bot. Gard. 23:227-832.

Taylor, C.M. 2005. Margaritopsis (Rubiaceae) in the Neotropics. Syst. Geogr. PI. 75:161-177.

Urban, 1.1921. Plantae Haitiensis novae vel rariores a cl. Er. L. Ekman 1917 lectae. Ark. Bot. 17(7): 1-72.

Urban, I. 1929. Plantae Haitiensis et Domingensis novae vel rariores VI. a cl. E.L. Ekman 1924-1928 lectae. Ark. Bot.

22A(10):1—108.

Journal of the Botanical Research Institute of Texas 9(1)

BOOK NOTICES

Flora of North America Editorial Committee. 2015. Flora of North America North of Mexico. Vol. 9: Magno-

liophyta: Picramniaceae to Rosaceae. (ISBN-13: 9780195340297, hbk.). Oxford University Press, 198

Madison Avenue, New York, New York 10016, U.S.A. (Orders: oup.com/us, 1-800-445-9714). $95.00,

752 pp., 8.5" x 11".

From the publisher: Flora of North America North of Mexico Volume 9: Magnoliophyta: Picramniaceae to Rosaceae

includes treatments prepared by 54 authors covering 691 species in 74 genera classibed in four families, with

the economically important Rosaceae being by far the largest in the volume with 680 species in 68 genera.

Descriptions for all of the families, genera, and species are provided, plus occurrence maps for species are in¬

cluded and 28% of the species are illustrated. Keys are included to aid in the identification of genera in families

and species within the genera. Volume 9 is the eighteenth volume to be published in the planned 30-volume

Flora of North America North of Mexico series.

The thirty-volume ongoing publishing project Flora of North America is the first comprehensive taxo¬

nomic guide to the extraordinary diversity of plant life covering our continent north of Mexico. This ground¬

breaking scholarly series is a collaborative effort by researchers at more than 30 U.S. and Canadian botanical

institutions. The Flora provides revisions of many plant groups and synthesizes the results from studies pub¬

lished in hundreds of research papers of the last three centuries. With beautiful illustrations accompanying

many species, Flora of North America is concise, easy to use, and indispensable to botanists, conservationists,

ecologists, agronomists, foresters, range and land managers, and horticulturists.

Mark Gustafson. 2015. A Naturalist’s Guide to the Texas Hill Country. (ISBN-13: 978-1-62349-235-9, flex-

bound). Texas A&M University Press, John H. Lindsey Building, Lewis Street, 4354 TAMU, College

Station, Texas 77843-4354, U.S.A. (Orders: www.tamupress.com, 1-800-826-8911). $24.95 US, 360 pp.,

328 color photos, 3 maps, table, bib., index, 5.5" x 8.5".

From the publisher: In this guide, biologist Mark Gustafson introduces residents and visitors to the history, ge¬

ology, water resources, plants, and animals found in the nineteen counties occupying the eastern part of the

Edwards Plateau, the heart of the Hill Country.

He profiles three hundred of the most common and unique species from all of the major groups of plants

and animals: trees, shrubs, wildflowers, cacti, vines, grasses, ferns, fungi, lichens, birds, mammals, reptiles,

amphibians, fishes, and invertebrates. Color photographs are included for each species along with a brief de¬

scription.

He closes with a chapter on significant state parks and natural areas in the region as an invitation to visit

and explore the Texas Hill Country.

As large metropolitan areas continue to encroach on the Hill Country, newcomers are moving in and

more people are flocking to its many attractions. This guidebook will enrich the appreciation of the region’s

rich and unique biodiversity and encourage conservation of the natural world encountered.

Mark Gustafson is professor of biology at Texas Lutheran University in Seguin, Texas, where he directs the

environmental studies program and specializes in aquatic biology and ecology.

J.Bot. Res. Inst. Texas 9(1): 80.2015

NEW SPECIES OF ACANTHACEAE FROM ECUADOR

Dieter C. Wasshausen

Department of Botany

National Museum of Natural History, MRC-166

Smithsonian Institution

Washington, D.C. 20013-7012, USA .

dmwasshausen@atmc.net

ABSTRACT

Four new species of Acanthaceae are recognized for Ecuador. They are the following: Mendoncia hollenbergiae Wassh., M. sericea Teonard

ex Wassh., Dyschoriste ecuadoriana Wassh., and Stenostephanus holm-nielsenii Wassh.

RESUMEN

Se reconocen cuatro nuevas especies de Acanthaceae para Ecuador. Son las siguientes: Mendoncia hollenbergiae Wassh., M. sericea

Teonard ex Wassh., Dyschoriste ecuadoriana Wassh., y Stenostephanus holm-nielsenii Wassh.

While preparing the Acanthaceae treatment for the Flora of Ecuador, I had set aside a number of collections,

mostly from Aarhus (AAU) and Gothenburg (GB) which morphologically seemed to differ from the known

species of that region. Unfortunately, while vacating my office at US, these collections were misplaced and they

only resurfaced after the publication of the treatment last year. Hence, I would like to publish these novelties at

this time before returning these loaned collections to their respective institutions.

Mendoncia hollenbergiae Wassh., sp. nov. (Fig. 1 A-F). Type: ECUADOR. Napo: Rio Napo between Coca (Puerto Francisco

de Orellana) and Had. San Carlos, E of Coca, riverside vegetation, ca. 350 m, 9 Nov 1974, G. Hading & L. Andersson 11854 (holotype:

GB).

Differt a M. sericea Teonard ex Wassh. foliis parvis; bracteolis oblongis vel parce ovatis; floribus 1-4 axillaribus; corolla alba cum striis vio-

laceis.

Suffrutescent vine; stem terete, older glabrous, younger sparingly puberulent, nodes inflated, internodes to 10

cm long, slightly twisted. Leaves petiolate, the petioles 1.4-2 cm long, glabrous to minutely puberulous, the

blades elliptic to narrowly ovate, 11-13 cm long, 4.5-6 cm wide, apex gradually narrowed to an acuminate tip,

basally acute to rounded, firm, glabrous or nearly glabrous, mid- and lateral veins (ca. 3 pairs) more prominent

on lower surface than on upper. Flowers 1-4 in each axil; pedicels 1-1.5 cm long, densely appressed-pubescent

with white trichomes; bracteoles oblong to narrowly ovate, 20-23 mm long, 8-10 mm wide, densely white se¬

riceous-hirsute or appressed pubescent, conspicuously so along mid-vein and margin, reddish-brown and

glabrous within, apex rounded or obtuse, slightly and inconspicuously apiculate, base obtuse to rounded; ca¬

lyx annular, 0.5 mm long; corollas white with violet striae on lower lip and in throat, tubular, 35 mm long,

glabrous without, minutely and inconspicouosly puberulous within, the tube curved, 5 mm broad at base,

slightly constricted at 10 mm above base, thence distally expanded to 10 mm at mouth, the limb oblique, 12-15

mm in diameter, subequally 5-lobed, the lobes suborbicular, reflexed, 8-8.5 mm long and 6-8 mm wide; sta¬

mens attached 13 mm above the base of the corolla tube; filaments are 2 mm long; anthers sagittate, thecae

slightly dissimilar, 6 and 4 mm long respectively, thecae basally terminating in a pubescent disk; ovary gla¬

brous; style 30 mm long; stigma minutely bifid. Drupe not seen.

Mendoncia hollenbergiae is morphologically similar to M. sericea Leonard ex Wassh. from the provinces of

Orellana and Pastaza in Amazonian Ecuador. The new species is distinguished by having leaf blades 11-13 cm

long, 4.5-6 cm wide, flowers 1-4 in each axil, bracteoles oblong to narrowly ovate, 20-23 mm long, 8-10 mm

wide and corollas white with violet striae on lower lip and in throat, 35 mm long. In contrast, in M. sericea the

leaf blades are smaller, 7-11.5 cm long, 2.5-4.5 cm wide, flowers several to numerous, forming compact, axil-

J. Bot. Res. Inst. Texas 9(1): 81 - 88.2015

Journal of the Botanical Research Institute of Texas 9(1)

Fig. 1 . A-F. Mendoncia hollenbergiae {Harling& Andersson 11854). A. Habit. B. Bracteoles and pistil. C. Corolla. D. Corolla expanded and stamens. E.

Longer pair of stamens and basal pubescent disk. F. Shorter pair of stamens and basal pubescent disk.

Wasshausen, New species of Acanthaceae from Ecuador

lary clusters in each axil, bracteoles elliptic to ovate, 10 mm long, 6-8 mm wide and corollas 12-15 mm long,

these light green below median lobe.

Distribution. —This species is known only from the type locality.

Etymology. —This name honors Tinda A. Hollenberg, whose career was spent in energetic and passionate

care of the plant collections in the United States National Herbarium.

Mendoncia sericea Teonard ex Wassh., sp. nov. (Fig. 2 A-G). Type: Ecuador. Pastaza: Mera, in rastrojo, ca. lioo m, 12

Dec 1955, ErikAsplund 18815 (holotype: S; isotype: US).

Differt a M. glomerata Leonard follis ellipticus vel parce ovatis, firmus, rigidus, glabris; bracteolis ellipticus et anguste ovatis, sericeous; co¬

rolla 12-20 mm longa.

Slender, climbing shrub; stem subterete, older glabrous, younger sparingly puberulent, nodes inflated, inter¬

nodes 4-6 cm long, minutely puberulous. heaves petiolate, the petioles 2-4.5 cm long, appressed puberulous,

the blades elliptic to narrowly ovate, 7-11.5 cm long, 2.5-4.5 cm wide, apex gradually narrowed to an acumi¬

nate tip, basally acute to rounded, firm, glabrous or nearly so above, appressed pubescent below, especially on

mid- and lateral veins (ca. 3 pairs). Flowers several to numerous, forming compact, axillary clusters; pedicels

1-1.3 cm long, sericeous, the pubescence brownish, appressed and ascending; bracteoles elliptic to narrowly

ovate, 10 mm long, 6-8 mm wide, densely and softly sericeous, the trichomes subappressed, whitish to yellow¬

ish-brown, conspicuously so along mid-vein and margin, yellowish-brown and glabrous within, apex rounded

or obtuse, slightly apiculate, rounded at base. Calyx annular, minute, 0.5 mm long; corollas 12-20 mm long,

the tube pale green to white without, essentially reddish-violet within, light green below median lobe, glabrous

or subglabrous without, lengthwise minutely puberulous on one half of each lobe within, the tube erect or

curved, 5 mm broad at base, slightly constricted at 10 mm above base, thence distally expanded to 7 mm at

mouth, the limb oblique, 9 mm in diameter, subequally 5-lobed, the lobes suborbicular, reflexed, 6 mm long

and 5 mm wide; stamens attached 6 mm above the base of the corolla tube; filaments are 6 mm long; anthers

sagittate, thecae somewhat dissimilar 6 and 4 mm long respectively, thecae basally terminating in a pubescent

disk; ovary glabrous; style 10 mm long. Drupe not seen.

Mendoncia sericea is morphologically similar to M. glomerata Teonard from the Comisaria of Putumayo,

Colombia. The new species is distinguished by having leaf blades elliptic to narrowly ovate, 7-11.5 cm long,

2.5-4.5 cm wide, firm rigid, glabrous or nearly so above, bracteoles elliptic to narrowly ovate and corollas 12-

20 mm long. In contrast, plants in M. glomerata have leaf blades ovate, 5.5-6.5 cm long, 4-4.5 cm wide, thin,

veiny, sparingly strigose or glabrate above, bracteoles ovate and corollas 10 mm long.

Distribution. —Terra hrme, in primary humid forest in the provinces of Orellana and Pastaza at elevation

of between 200 and 300 meters.

Etymology. —The specific epithet describes the conspicuous sericeous pubescence evident on the bracte¬

oles of the species.

Material studied in addition to the type: ECUADOR. Orellana: Parque Nacional Yasuni, km 45 of Maxus Petroleum Road (=YPF), 00°40'S,

76°23'W, 200-300 m, 18 Jan 1998, Robyn J. Burnham w/Alexander Krings 1549 (MICH); Parque Nacional Yasuni, road construction to Maxus,

km 12-16,00°31'S, 76°32'W, 240 m, 6 Mar 1993 Jorge Zuleta 194 (US). Pastaza: Shiguacocha, ca. 5 km E of Puerto Sarayacu, 1 Oct 1974, Lugo

3833 (GB, US); Puerto Sarayacu, 30 Oct 1974 Lugo 3867 (GB, US); Rio Zupayacu, ca. 7 km S of Puerto Sarayacu, 11 Oct 1974, Lugo 4060 (GB,

US); Rio Capahuari, ca. 12 km N of Puerto Sarayacu, 13 Oct 1974, Lugo 4137 (GB, US).

Here the author wishes to note that earlier in his career while doing routine identifications of Acanthaceae he

determined a number of collections from Ecuador and Peru as ‘ Mendoncia sericea Teonard.” This was an un¬

published name and many of these collections were misidentihed. Subsequently these determinations were

listed in TROPICOS. In this present study of M. sericea Teonard ex Wassh., all of the Ecuadorean material at US

was critically examined and cited. No Peruvian collections were examined for this study.

Dyschoriste ecuadoriana Wassh., sp. nov. (Fig. 3 A—H). Type: ECUADOR. Guayas: Nobol, roadside vegetation, 14 May 1974,

G. Hailing & L. Andersson 14594 (holotype: GB; isotype: US).

A D. quitensis (Kunth) O. Kuntze differt habitu erecti; foliis corollisque parvis et calycibus brevis connatis valde distincta.

Journal of the Botanical Research Institute of Texas 9(1)

Fig. 2. A-G. Mendoncia sericea (Asplund 18815). A. Habit. B. Bracteoles and pistil. C. Corolla. D. Corolla expanded and stamens. E. Longer pair of

stamens and basal pubescent disk. F. Shorter pair of stamens and basal pubescent disk. G. Calyx, nectar disk, ovary, and pistil.

Wasshausen, New species of Acanthaceae from Ecuador

Fig. 3. A-H. Dyschoristeecuadoriana ( Harling&Andersson 14594). A. Habit. B. Bracts and calyx. C. Corolla. D. Corolla expanded and stamens. E. Calyx,

ovary, and pistil. F. Shorter stamen. G. Longer stamen. H. Capsule.

Journal of the Botanical Research Institute of Texas 9(1)

Branched or solitary, erect perennial herbs, from thickened woody base, 30-50 cm tall; young branches sub-

quadrangular, moderately puberulous, older stems subquadrangular, glabrescent. Leaves petiolate, the peti¬

oles (unwinged portion) 5-10 mm long, glabrous; the blades ovate to obovate, 4-6.5 x 1.5-3.5 cm, glabrous or

nearly so above, sparcely puberulent below, apex obtuse to acute, cuneate and decurrent on the petiole, margin

entire, ciliolate; leaves subtending inflorescence reduced, frequently distinctly more ovate andbasally rounded

rather than cauline leaves. Inflorescence of dichasia in leaf axils, often appearing as glomerules, dichasia op¬

posite or subopposite, 1-3-flowered, sessile to subsessile, peduncles (if present) to 1 mm long; bracts elliptic to

oblong, 7.25-8 mm long, 2-2.5 mm wide, glabrous or with a few scattered trichomes, minutely apiculate at

apex, margins ciliolate; bracteoles linear, minute, 1.5-2 mm long and 0.1 mm wide. Flowers sessile to subses¬

sile, pedicels to 1 mm long; calyx 12 mm long, the lobes basally fused for about one-third their length, subulate-

setaceous, hirsutulous, margins ciliolate; corolla pale violet, 10-12 mm long, sparingly puberulous, tube 9.5-

10 mm long, 2 mm broad at base, slightly constricted at 5 mm above base, thence distally expanded to 3.5 mm

at mouth, upper lip erect, deeply-bilobed, lobes oblong, rounded, 2.5 mm long and 2 mm wide, lower lip

spreading, 3.5 mm long, lobes elliptic, 3 mm long, 2.5 mm wide; stamens inserted just above base of throat, the

longer pair 3.5 mm long, the shorter pair 1.5 mm long; longer pair of blaments glabrous, shorter pair hispid;

anthers bithecous, thecae equal, 2.5 mm long, distinctly bicalcarate at base; ovay nitid, glabrous, 6 mm high;

style 7 mm long, included. Capsule narrowly elliptic to oblong, 9 mm long, 2.25 mm wide, glabrous, obtuse

and apiculate; seeds 2-4, oblong, about 2.5 mm long, 1.75 mm wide, notched and oblique at base.

Dyschoriste ecuadoriana is morphologically similar to D. quitensis (Kunth) O. Kuntze also from Ecuador

and Peru. This new species is distinguished by being a solitary or branched, erect perennial herbs 30-50 cm

tall, bracts elliptic to oblong, 7.25-8 mm long and 2-2.5 mm wide, calyx 12 mm long, basally fused for about

one-third their length, corollas 10-12 mm long, the tube 9.5-10 mm long and capsules narrowly elliptic to

oblong, 9 mm long and 2.25 mm wide. In contrast, D. quitensis has plants that are usually procumbent herbs

15-40 cm tall, the bracts are oblong to lanceolate, 8-10 mm long and 2 mm wide, calyx 7-11 mm long basally

fused for about one-half their length, corollas smaller, 9-10 mm long, the tube 6.5-7 mm long and capsules

narrowly oblong, 7-8 mm long and 2 mm wide.

Distribution. —Along roadsides and in waste places in the provinces of Manabl, Guayas and Loja at eleva¬

tions between 0 and 1900 meters.

Etymology. —The specibc epithet, D. ecuadoriana, denotes the western Andean region in which the new

species is known to occur.

Material studied in addition to the type: ECUADOR. Manabl: Portoviejo-Jipijapa Road, ca. 1 km S of La Pila, ca. 200 m, 4 May 1985, G.

Harling & L. Andersson 24823 (GB, US). Guayas: Guayaquil, at golf club, 23-30 Apr 1968, G. Hading, G. Storm & B. Strom 8803 (GB, US); Isla

Puna, May 1892, Eggers 14741 (US); Oil Camp, between Guayaquil and Salinas, 0-100 m, 21-24 Jun 1923, A.S. Hitchcock 19998 (US). Loja:

Road Catacocha-Macara, ca. 8 km S of Empalme, ca. 1900 m, 10 Nov 1977, G. Hading, U. Eliasson & E. Andersson 15267 (GB); Vilcabamba-

Yangana, 1700-1800 m, 15 Apr 1974, G. Hading & E. Andersson 13594 (GB).

Stenostephanus holm-nielsenii Wassh., sp. nov. (Fig. 4 A-E). Type: Ecuador. Napo: slopes of Guagra Urcu, on the loma

above upper Rio Borja, SE exposed montane forest, 2600 m, 0°28'S, 77°44'W, 25 Sep 1980, L. Holm-Nielsen et al. 26982 (holotype:

AAU).

Differt a S. luteynii (Wassh.) Wassh. foliorum lamina angustata; paniculae floriferae brevis, modice densae; pedicelli longiori; calycis seg-

menta anguste lanceolata, parce pilosa et minute bulbosa ad apicem, pilis glanduliferis intermixtis.

Suffrutescent plants probably 1-1.5 m tall or more (only tips available for study); stems subquadrangular, red¬

dish, glabrous or the upper portion minutely puberulous, trichomes appressed. Leaves petiolate, the petioles

1.5-5 cm long, moderately puberulous, the blades elliptic to ovate, 9-15 cm long and 3-4.5 cm wide, gradually

narrowed to a short-subacuminate tip, cuneate at base, rather thin and membranous, entire or undulate, both

surfaces minutely puberulous, lateral veins (6-12 pairs) more conspicuous on lower surface, this also drying a

lighter shade of green. Flower-bearing panicles terminal, 7-10 cm long and 5-6 cm broad, compact and mod¬

erately dense; mostly sterile cymes axillary to 3 cm long, branches of terminal inflorescence subhelicoid,

usually once-forked; bracts subtending branches of inflorescence cordate, 12 mm long, 7.5 mm wide, long-

Wasshausen, New species of Acanthaceae from Ecuador

Fig. 4. A-E. Stenostephanus holm-nielsenii. A-D: ( Holm-Nielsen etal. 26982). E: ( Holm-Nielsen etal. 2697\). A. Habit. B. Calyx and ovary. C. Corolla,

stamens, and pistil. D. Calyx lobes, nectar disk, and ovary. E. Calyx lobes, capsule, and seeds.

Journal of the Botanical Research Institute of Texas 9(1)

acuminate at apex, white cystoliths prominent; rachis and rachilla glandular-pilose, trichomes spreading,

whitish; pedicels erect, 4 mm long, sparingly glandular-pilose; bracts subtending pedicels green, subulate, 3 x

0.5 mm, sparingly glandular-pilose without; bracts subtending flowers linear, ca. 1.5 mm long, 0.5 mm wide

near base, glandular-pilose; calyx 5 mm long, lobes narrowly lanceolate sparingly glandular-pilose, bearing a

few acute trichomes to 0.16 mm long on outer and inner surface, especially towards tip, tip itself minutely bul¬

bous, calyces of sterile, lower axillary inflorescence 1-2.5 mm long; corolla wine-red, oblique, 25 mm long

from base to tip of upper lip, glabrous, the tube ca. 1.5 mm broad near base, slightly enlarged to 2 mm at 5 mm

above base, then geniculate and gradually enlarged to 4 mm at mouth, upper lip erect, 12 mm long, linear-

lanceolate, ca. 1 mm wide at base, gradually narrowed to an acute, recurved or curling tip, lower lip spreading,

8 mm long and 6 mm wide, conduplicate, tip 3-lobed, lobes ovate, ca. 1 mm long and wide, obtuse; stamens

exserted 4 mm beyond mouth of corolla tube, glabrous, anthers 1-1.5 mm long, densely glandular-pilose, mu-

ticous at both ends; style shorter than stamens by about 3 mm, stigma bilobed, lobes minute, rounded; ovary

glabrous. Capsule clavate, glabrous, 13 mm long, stipe 5 mm long, head 8 mm long; seeds yellowish, rugose, 2

x 2 mm.

Stenostephanus holm-nielsenii is morphologically similar to S. luteynii (Wassh.) Wassh. also from the prov¬

ince of Napo in Ecuador. This new species is distinguished by having narrower leaf blades, these 3-4.5 cm

wide, flower-bearing panicles 7-10 cm long, compact and moderately dense, bracts subtending branches of

inflorescence long-acuminate at apex, pedicels 4 mm long, calyx lobes narrowly lanceolate, apically sparingly

glandualar-pilose and minutely bulbous at tip and the corollas wine-red. In contrast, in S. luteynii the leaf

blades are broader, 3-5.5 cm wide, the flower-bearing panicles are less dense and compact, shorter, 6-8 cm

long, bracts subtending branches of inflorescence rounded and apiculate at apex, pedicels 1-2 mm long, calyx

lobes elliptic-oblong, glabrous, subtruncate and apiculate at apex and the corollas deep-maroon.

Distribution. —Montane forest, elbn forest, moist scrub forest at elevations between 2000 and 2900 m.

Etymology.—Stenostephanus holm-nielsenii is named after Lauritz Holm-Nielsen of Aarhus University in

recognition of the many interesting Acanthaceae he and his team collected during the course of their investiga¬

tion of plant diversity in Andean Ecuador.

Material studied in addition to the type: ECUADOR. Napo: slopes of Guagra Urcu, on the loma above Rio Bretania, 0°28'S, 77°41'W, 2000-

2200 m, 22 Sep 1980, L. Holm-Nielsen et al. 26807 (AAU); slopes of Guagra Urcu, on the loma above Rio Borja, 0°28'S, 77°44'W, 25 Sep 1980,

L. Holm-Nielsen et al. 26971 (AAU); upper slopes of Guagra Urcu, moist scrub forest, 0°28'S, 77°43'W, 2750 m, 26 Sep 1980, L. Holm-Nielsen

et al. 27183 (AAU); Guara Urcu, the pass between Rio Borja and Rio Suno, 0°28'S, 77°43'W, 2700 m, 27 Sep 1980, L. Holm-Nielsen et al. 27274

(AAU); upper Rio Suno, near Guagra Urcu, montane forest, 00°28'S, 77°42'W, 2900 m, L. Holm-Nielsen 27552 (AAU).

ACKNOWLEDGMENTS

My special thanks to Alice Tangerini for the skillfully prepared line drawings. The author further wishes to

thank David Neill (ECUAMZ) and an anonymous reviewer, and editors for providing critical suggestions and

corrections in their review of this manuscript.

REFERENCE

Wasshausen, D. C. 2013. Acanthaceae. In: C. Persson and B. Stahl, Flora of Ecuador. Department of Biological and Environ¬

mental Sciences, University of Gothenburg, Sweden. Pp. 1-329.

CUATRO NUEVAS ESPECIES Y UN NUEVO REGISTRO DE FREZIERA

(PENTAPHYLACACEAE) DE ECUADOR Y PERU

Daniel Santamarfa-Aguilar Laura P. Lagomarsino

Harvard University Herbaria

22 Divinity Avenue

Cambridge, Massachusetts 02138-2020 U.S.A.

daniel.santamaria366@gmail.com

Dept. ofOrganismic and Evolutionary Biology

Harvard University Herbaria

22 Divinity Avenue

Cambridge, Massachusetts 02138-2020 U.S.A.

lagomarsino.l@gmail.com

RESUMEN

Se describen cuatro especies de Freziera (Pentaphylacaceae) nuevas para la ciencia: Freziera humiriifolia y F. yanachagensis de Ecuador y

Peru; y F. neillii y F. tundaymensis de Ecuador. Se documenta por primera vez F. carinata para Ecuador.

ABSTRACT

Four new species of Freziera (Pentaphylacaceae) are described as new to science. Freziera humiriifolia and F. yanachagensis from Ecuador

and Peru; and F. neillii and F. tundaymensis from Ecuador. Freziera carinata is documented in Ecuador for the first time.

INTRODUCCION

La familia Pentaphylacaceae contiene 12 generos y 337 especies alrededor del mundo. En el Neotropico, esta

dividida en dos tribus, las cuales constan de cuatro generos y aproximadamente 127 especies (Every 2009). La

tribu Freziereae esta conformada por los generos Cleyera Thunb., Freziera Willd., y Symplococarpon Airy Shaw;

y la tribu Ternstroemieae por el genero Ternstroemia Mutis ex L.f. (Weitzman et al. 2004). Freziera es un genero

de arboles o arbustos con hojas dlsticas y flores funcionalmente dioicas con la corola urceolada, el estilo mas

corto que el ovario, y el estigma 3-5 lobulado (Weitzman et al. 2004; Every 2009). El genero, sin incluir las

especies aqul descritas, se encuentra representado por 11 especies en Ecuador (Weitzman 1999) y 12 especies

en Peru (van der Werff 1993). Los mismos autores comunican la presencia de otros taxones ineditos para los

palses mencionados.

Como resultado del estudio de ejemplares depositados en el herbario del Missouri Botanical Garden

(MO) y de los especlmenes recibidos para identihcacion en el Gray Herbarium (GH) of Harvard University, se

describen cuatro nuevas especies de Freziera para Ecuador y Peru. Ademas se registra por primera vez la pres¬

encia de F. carinata A.L. Weitzman en Ecuador, que se consideraba endemica de Venezuela, y restringida al

Escudo Guyanes en los estados Amazonas y Bolivar (Berry & Weitzman 2005, 2007, 2008).

Las cuatros nuevas especies de Freziera aqul descritas son provenientes de las mesetas de arenisca de la

Cordillera del Condor en Ecuador y habitats similares en Peru. La Cordillera del Condor, compartida entre

Ecuador y Peru, es una cadena montanosa ubicada al este de los Andes. Tiene una longitud aproximada de 150

km y una altura maxima de 2900 m. Los suelos estan formados principalmente por roca arenisca. Se calcula

que la Cordillera del Condor tiene una flora de alrededor de 4,000 especies (Schulenberg & Awbrey 1997; Neill

2005; Rodriguez et al. 2006). En los ultimos anos se han descrito una serie de novedades de la Cordillera del

Condor que demuestra su alta diversidad (Ulloa & Neill 2006; Clark et al. 2010; Stahl 2010; Daly et al. 2012;

Neill & Asanza 2012; Neill et al. 2012; Riina et al. 2014; Grant 2014). El hallazgo de Freziera carinata en la Cor¬

dillera del Condor demuestra una vez mas las abnidades florlsticas entre el Escudo Guyanes y esta cordillera,

que estan separados por mas de 3000 km (Ulloa Ulloa & Neill 2006). Otros ejemplos de generos o especies

disyuntos entre esta cordillera y el Escudo Guyanes han sido documentado por Berry et al. (1995) y Neill

(2007), incluyendo: Bonnetia Mart. (Bonnetiaceae), Digomphia Benth. (Bignoniaceae), Fverardia Ridl. (Cypera-

J. Bot. Res. Inst. Texas 9(1): 89 -102.2015

Journal of the Botanical Research Institute of Texas 9(1)

ceae), Phainantha Gleason (Melastomataceae), Podocarpus tepuiensisj. Buchholz & N.E. Gray (Podocarpace-

ae), Pterozonium Fee (Pteridaceae), y Retiniphyllum tepuiense Steyerm. (Rubiaceae).

TAXONOMI'A

Freziera humiriifolia D. Santam., sp. nov. (Fig. 1). Tipo: ECUADOR. Zamora-Chinchipe: ElPangui, Cordillera del Condor, near

Condor Mirador military post, on Ecuador-Peru border, cloud forest, disturbed, at low point of summit ridge, below sandstone

outcrops, 03°38 , 20"S, 78°23 , 29"W, 1760 m, 08 set 2003 (fl. d), DA. Neill, E. Rodriguez, W. QuizhpeJ. Homeier 14522 (holotipo:

QCNE-193416 [n.v]; isotipos: GH!, MO-6402480!).

This species is similar to Freziera dudleyi and F. sessiliflora, but differs from both species in its glabrous branches and large, ovate or oblong-

elliptic leaves that are densely pustulate on the upper surface and densely papillate on the lower surface.

Arboles o arbustos 1.5-5 [16-18] m o eplfita?; ramas maduras cillndricas, la corteza externa negruzca, longi-

tudinalmente estriada, glabras; ramitas cillndricas, glabras, [adpreso pubescentes, los tricomas blanquecinos

a amarillento-dorados, 0.3-0.5 mm de longitud], papiladas, sin lenticelas. Yema terminal conduplicada, 27-

70 mm de longitud, glabrescente, [finamente adpreso o densamente pubescente], los tricomas blanquecinos a

amarillento-dorados, 0.5-1.0 mm de longitud. Hojas con el peclolo 0.3-8.0 mm de longitud, caniculado o

piano adaxialmente, redondeado abaxialmente, glabro [finamente adpreso pubescentes, los tricomas cafe ro-

jizo claro, 0.2-0.5 mm de longitud], las alas peciolares presentes o ausentes, si presentes erectas y los margenes

enteros; coleteres ausentes; lamina 7.5-14 x 3.3-6.6 cm, ovada a oblongo-ellptica, la base obtusa, decurrente

sobre el peclolo, generalmente revoluta, ambos lados iguales, el apice acuminado y con una seta negra y cur-

vada, caduca; margen sinuado, 55-80 dientes por lado, con pequenas setas insertadas en los senos, uncinadas,

negras, caducas, las setas sin tricomas a su alrededor; haz glabro y densamente pustulado; enves glabrescente

o adpreso pubescente, los tricomas >0.3 mm [0.5-1.0 mm] de longitud y asociados con papilas, las papilas

densas y distribuidas sobre todo la lamina, cafe-rojizas claras o cafe-rojizas oscuras a casi negras; nervio cen¬

tral por la haz piano, glabro, densamente pustulado, redondeado en el enves, glabro [adpreso pubescente],

papilado-rugoso; nervios laterales 35-60 pares por lado, levemente impresos por la haz y pianos en el enves.

Inflorescencias de 1 flor solitaria, axilar o 3-6 flores fasciculadas, axilares y ramifloras; bractea 1.0-2.0 x

1.0-2.0 mm, en la base del pedicelo, persistente o caduca, triangular u obovada, generalmente carinada, el

apice agudo, que termina en una seta levemente curvada, caduca, el margen entero, la superhcie externa ad¬

preso pubescente, la superhcie interna glabra; pedicelo ca. 1.0 mm de longitud o ausente, cillndrico, erecto,

adpreso pubescente; bracteolas 2, 2.0-3.0 x 1.5-2.0 mm, opuestas o subopuestas, levemente desiguales, per-

sistentes, suborbiculares, triangular o en forma de ‘D’, carinada, la superhcie externa hnamente adpreso, la

superhcie interna glabra, el margen ciliado, el apice redondeado o retuso, terminando en una seta negra, ca¬

duca; botones florales 2.5-3.5 mm de ancho; sepalos 5, imbricados; sepalos externos 2, 2.5-3.5 x 2.0-3.5 mm,

ampliamente ovados a suborbiculares, apice redondeado y con una seta negra, conica, la superhcie externa

adpreso pubescentes, la superhcie interna glabra, el margen cartaceo o membranaceo, ciliado y con pequenas

setas cafe-rojizas en la base o arriba de la base; sepalos internos 3, casi del mismo tamano y similares en forma

a los externos, el apice redondeado con o sin una seta, adpreso pubescentes en la superhcie externa, glabros en

la superhcie interna, el margen membranaceo, ciliado y con pequenas setas en toda su longitud, incoloras o sin

setas; petalos 5, 2.5-4.5 x 2.5- 4.0 mm, blancos, amarillentos o rojos? (Wisum 13), unidos en la base, ovados,

glabros, el margen membranaceo y entero, el apice redondeado o agudo. Flores estaminadas con (19-)23-25

estambres, 1.0-2.5 mm de longitud, levemente unidos entre ellos y a la base de los petalos, ± desiguales; hla-

mentos 0.5-1.5 mm de longitud, levemente aplanados; anteras 0.5-0.9 mm de longitud, obovoides, oblicuas,

atenuadas en la base, apiculadas en el apice; gineceo 2.0-4.0 mm de longitud, mas o menos alargado y muy

deforme, glabro, 4-locular; estilo separado apicalmente por ca. 1 mm de longitud; estigma 2-4(-5) lobulado.

Flores pistiladas desconocidas. Frutos 4.0-5.0 x [3-]4.0-5.0 mm, oblatos o globosos, verdes y rojos, glabros;

paredes del fruto ca. 0.5 mm de grueso; semillas 20-35 por fruto, cafe-claro o cafe-negruzcas, brillantes, 1.5-

2.0 mm de longitud, de forma no muy dehnida o mas o menos cuneiformes, rugosas o rugoso foveoladas.

Habitat, distribucidnyfenologia.—Freziera humiriifolia ha sido recolectada en Ecuador en bosques enanos

sobre roca arenisca en la Cordillera del Condor en las provincias de Morona-Santiago y Zamora-Chinchipe,

Journal of the Botanical Research Institute of Texas 9(1)

entre los 1020-1760 m de elevation. En Peru ha sido recolectada en orillas de quebrada en la provincia de Ba-

gua, entre los 320-700 m de elevation. Flores estaminadas han sido recolectadas en setiembre y octubre y

frutos en enero, setiembre y diciembre. Lo demas de su fenologla se desconoce.

Etimologia. —El eplteto especlbco humiriifolia hace referencia a la similitud de las hojas de la nueva espe-

cie cuando secas, con las de algunos miembros de la familia Humiriaceae.

Discusidn.—Freziera humiriifolia se puede distinguir por la siguiente combinacion de caracteres mor-

fologicos: ramitas sin lenticelas, hojas cortamente pecioladas, ovadas a oblongo-ellpticas, numerosos nervios

laterales, levemente impresos por la haz y pianos en el enves, densamente pustulados por la haz, la base gener-

almente revoluta, y el enves glabrescente o adpreso pubescente y densamente cubierto por papilas cafe-rojizas.

Tambien es distintiva por sus inflorescencias sesiles o muy cortamente pediceladas, axilares o axilares y rami-

floras y por sus flores con el estilo separado apicalmente. Freziera humiriifolia se podrla relacionar con F. ses-

siliflora A.H. Gentry, especie muy rara de Colombia y con F. dudleyi de Peru y Bolivia. Estas especies comparten

hojas cortamente pecioladas, la base revoluta, el margen sinuado, y numerosos nervios no muy conspicuos e

inflorescencias axilares o axilares y ramifloras. Para diferencias entre estas especies, vease Tabla E

Las dos colecciones aqul citadas de Peru diberen del material tlpico de Ecuador por ser arboles mucho

mas grandes y que crecen en elevaciones inferiores. Esas colecciones diheren de las demas por las ramitas,

yema terminal, peclolos y hojas con el indumento mas denso y obvio, y los frutos mas angostos. Las medidas

entre corchetes corresponden a este material. Cuando mas material de esta especie se encuentre disponible,

sera posible determinar si corresponde a un taxon diferente o a variation a traves de su ambito geograhco.

Los signos de pregunta del habito y coloration de las flores corresponden a la coleccion A. Wisum 13 (GH,

MO). Es posible que los datos de la etiqueta no correspondan a esta especie.

Especimenes adicionales examinados (paratipos). ECUADOR. Zamora-Chinchipe: vicinity of EcuaCorriente copper mine concession, vi¬

cinity of mine site, along trail above parking area near end of road, 03 o 34'54"S, 78°26'06"W, 1330-1360 m, 21 set 2007 (fl. d), T.B. Croat &

G. Ferry 98957 (GH, MO, QCNE [n.v]); El Pangui, Cordillera del Condor, summit of sandstone plateau of Cordillera, southeast headwaters

of Rio Wawaime, above proposed EcuaCorriente copper mine area, 03°35 , 40"S, 78°25 , 11"W, 1930 m, 08 die 2005 (fl. d), D.A. Neill & W.

Quizhpe 15048 (GH, MO, QCNE [n.v]). Morona-Santiago: Limon Indanza, Cordillera del Condor, 2 Km al noreste del Centro Shuar Warint-

za, Meseta de roca arenisca, bosque densos y bajo, 03°09 , 16"S, 78°14 , 50"W, 1020 m, 05 oct 2005 (fl. d), A. Wisum 13 (GH, MO); Limon In¬

danza, Cordillera del Condor, Centro Shuar Warints, Cerro Mako Naint a 6 Km del Centro, 03°10 , 59"S, 78 0 17'57"W, 1600 m, 07 oct 2002 (fl.

d), C. Kajekai 23 (GH, MO, QCNE [n.v]). PERU. Bagua: Distrito Imaza, Comunidad Aguaruna Putuim, Anexo de Yamayakat, Zonas atlas

de Putuim, “campou” 240°SW de Putuim, 700-750 m, 23 ene 1996 (fr), C. Diaz et al. 7766A (GH, MO, USM); Region Nororiental del Mara-

non, Comunidad de Kampaenza, Ribera de la quebrada Shimutaz, rio Maranon, 04°55'S, 78°19'W, 320 m, 09 set 1994 (bot. fl. & fr), N.Jara-

millo 433 (GH, MO).

Freziera neillii D. Santam., sp. nov. (Fig. 2). Tipo: ECUADOR. Zamora-Chinchipe: Palanda Canton, Tapichalaca Reserve, S of

Podocarpus National Park, E of road between Yangana and Valladolid upper Rio Chinchipe watershed, near beginning of mule trail

to Quebrada Honda, exposed ridge with very dense montane montane scrub, 04 o 29'23"S, 79°08 , 11"W, 2550 m, 14 set 2007 (fl. $),

D.A. Neill, C. Davidson &S. Christoph 15999 (holotipo: QCNE-213282 [n.v]; isotipos: GH!, MO-6402471!).

This species differs from Freziera parva by the vilose to sericeous abaxial leaf surfaces (vs. glabrous) and 4-locular ovaries (vs. 3-locular).

Arboles o arbustos 3-4 m; ramas maduras cillndricas, la corteza externa negruzca, longitudinalmente estri-

ada, glabras; ramitas cillndricas o muy ligeramente aplanadas, vilosas a esparcidamente vilosas, los tricomas

blanquecinos a blanquecino grisaceos, tricomas 0.08-0.1 mm de longitud, papiladas, escasamente lenticela-

das, las lenticelas, si presentes, ellpticas y blanquecinas. Yema terminal conduplicada, 11-25 mm de longitud,

densamente hirsuta, los tricomas castano claro, 0.2-1.0 mm de longitud. Hojas con el peclolo 1.0-3.0 mm de

longitud, caniculado adaxialmente, redondeado abaxialmente, glabrescente o cortamente viloso, las alas peci-

olares presentes o ausentes, si presentes cortas y erectas con los margenes enteros, los tricomas castano claros,

de 0.5-1.0 mm longitud; coleteres 1, en la base del peclolo; lamina 3.5-6.1 x 2.7-3.7 cm, oblongo-ellptica, al-

gunas veces mas o menos auriculiforme, la base no revoluta, subtruncada a subcordada, algunas veces con un

lado asimetrico, el apice agudo o redondeado y levemente retuso, algunas veces con una seta negra y curvada,

caduca; margen crenado, 27-45 dientes por lado, con pequenas setas insertas en los senos, uncinadas, negras,

caducas, generalmente las setas rodeadas de tricomas; haz generalmente glabra, algunas veces diminuta

Santamaria-Aguilary Lagomarsino, Nuevas especies de Freziera

Tabla 1. Comparacion entre algunos caracteres diagnostics entre Freziera dudleyi, F. humiriifoliay F. sessiliflora.

Caracteres

F. humiriifolia

F. dudleyi

F. sessiliflora

Indumento en ramitas

Glabras [finamente adpreso]

Sericeas

Densamente pubescentes

Forma de hoja

Ovadas a oblongo elipticas

Angostamente elipticas a

casi lineares

Angostamente eliptica

Tamano de hojas

7.5-14x33-6.6 cm

7.8-93 x 1.2-2.2 cm

7.0-8.2 x 23-3.0 cm

Pustulas en el haz

Densas

Ausentes o escasas

Ausentes

Papilas en enves

Densas sobre todo la lamina

Principalmente sobre linea

de vernacion

Ausentes

Vena media por el haz

Plana

Levemente elevada, vista

como una cresta

No observada

Anteras

No moniliformes

Moniliformes

No observadas

Caracteres entre corchetes [ ] corresponden a las colecciones de Peru.

y muy esparcidamente pubescentes, densamente pustuladas; enves viloso a serlceo, los tricomas 0.5-1.0 mm

de longitud y asociados con papilas, las papilas distribuidas sobre todo la lamina, cafe-rojizas; nervio central

por la haz piano o levemente caniculado, glabro o esparcidamente pubescente, densamente papilado; redon-

deado o triangular en el enves, viloso a serlceo, densamente papilado; nervios laterales 14-27 por lado, im-

presos en la haz y elevados en el enves. Inflorescencias de 1 flor solitaria o 3 flores fasciculadas axilares;

bractea 1.5-3.3 x 0.8-1.5 mm, en la base del pedicelo, persistente o algunas veces caduca, triangular, carinada,

el apice agudo o acuminado, que termina en una seta negra, uncinada, levemente curvada y caduca, el margen

entero, la superficie externa adpreso pubescente o glabrescente, la superficie interna glabra; pedicelo 3.0-6.0

mm de longitud, cillndrico, erecto o alguna veces curvado, glabro; bracteolas 2, 1.5-3.0 x 2.0-3.3 mm, opues-

tas, levemente desiguales, persistentes, semiorbiculares, generalmente sin una carina, la superficie externa e

interna glabras, el margen ciliado, el apice redondeado y sin una seta negra; botones florales 2.5-4.0 mm de

ancho; sepalos 5, imbricados; sepalos externos 2,3.0-4.0 x 3.0-4.0 mm, ampliamente ovados, el apice redon¬

deado y sin una seta, glabros sobre ambas superficies, el margen membranaceo, esparcidamente ciliado o los

cilios ausentes, sin setas en el margen; sepalos internos 3, 3.0-4.0 x 3.0-3.5 mm, iguales a los externos, pero

con los cilios en los margenes generalmente mas notorios; petalos 5, 6.0-7.0 x 2.0- 3.0 mm, blancos o blanco-

rosados, levemente unidos en la base, lanceolados, glabros, el margen membranaceo y entero, el apice redon¬

deado o agudo. Flores estaminadas desconocidas. Flores pistiladas con 17-18 estaminodios (algunas veces

los estaminodios ausentes), 0.9-2.5 mm de longitud, unidos a la base de los petalos, lineares, pianos, apiculado

en el apice; gineceo 3.7-4.0 mm de longitud, ovado a conico, glabro, 4-locular; estilo no separado apicalmente;

estigma 4-lobulado. Frutos 5.0-6.0 x 3.0-5.0 mm, redondeados, amarillo-verdosos, glabros; paredes del fruto

ca. 0.3 mm de grueso; semillas 24-29 por fruto, cafe rojizas o negras, brillantes, 1.0-1.3 mm de longitud, mas

o menos cuneiformes o reniformes, foveoladas.

Habitat, distribucion y fenologia.—Freziera neillii es endemica de Ecuador. Hasta el momento se conoce

solo por tres colecciones, todas provenientes de la Reserva Tapichalaca. Ha sido recolectada en bosque mon-

tano en la provincia de Loja y Zamora-Chinchipe, entre los 2550-2630 m de elevacion. Flores pistiladas y fru¬

tos han sido recolectadas de setiembre a noviembre; lo demas de su fenologia se desconoce.

Etimologia. —El epiteto especlfico de E. neillii es en honor del Dr. David A. Neill (1953-), prolihco botanico

estadounidense y experto de la flora ecuatoriana. El colecto excelentes ejemplares de esta nueva especie. Tam-

bien lideraba el proyecto Inventario Botanico de la Region de la Cordillera del Condor, Ecuador-Peru, entre el

2004-2007, durante dicho proyecto se recolectaron varias de las especies aqul descritas.

Discusidn.—Freziera neillii se puede distinguir facilmente por varios caracteres en sus laminas foliares: la

textura cartacea, el tamano relativamente pequeno, su forma oblonga-ellptica o algunas veces mas o menos

auriculiforme, la base subtruncada a subcordada, el apice agudo o redondeado y levemente retuso, la haz

densamente pustulada, el enves viloso a serlceo y densamente papilado, los nervios reticulados, y el margen

Journal of the Botanical Research Institute of Texas 9(1)

MISSOURI

BOTANICAL GARDEN

HERBARIUM

N? 6402471

Zamora-Chinchipe: Palanda

Tapichalaca Reserve, south of

Podocarpus National Park, east of road

between Yangana and Valladolid, upper

Rio Chinchipe watershed, near

beginning of mule trail to Quebrada

Honda. Exposed ridge with very dense,

dwarf montane forest and low dense

montane scrub,.

04 0 29 1 23"S 79008’11 "W 2550 m

Small tree or shrub 4 m tall. Flowers

ivory-white or suffused pale pink.

Fruits yellow-green.

25 Sep 2007

David Neill,

C. Davidson & S. Christoph 15999

MISSOURI BOTANICAL GARDEN HERBARIUM (MO)

Fig. 2. Isotipo de Freziera neillii D. Santam. {D.A. Neill etal. 15999, MO-6402471).

Santamaria-Aguilary Lagomarsino, Nuevas especies de Freziera

con numerosos dientes y setas rodeadas de tricomas. Tambien es distintiva por sus bracteolas y sepalos relati-

vamente pequenos y completamente glabros (excepto en los margenes, que son ciliados). Una especie similar a

la nueva especie es Freziera parva Kobuski, del departamento de Amazonas, Peru. Ambas especies tienen

laminas foliares cartaceas, densamente pustuladas en la haz, los nervios reticulados, las setas en el margen de

las hojas cubiertas de tricomas y bracteolas y sepalos glabros. Freziera neillii se puede diferenciar de F. parva,

por los caracteres enumerado en la Tabla 2.

Especi'menes adicionales examinados (paratipos). ECUADOR. Loja: Tapichalaca Reserve, between Yangana and Valladolid, 04°29'S,

79°08'W, 2630 m, 24 oct 2004 (fl. $), J. Homeier et al. 144-3 (MO, QCNE [n.v]). Zamora-Chinchipe: Palanda Canton,Tapichalaca Reserve, S

of Podocarpus National Park, in upper Rio Chinchipe watershed, E of the Yangana-Valladolid road, at the head of the mule trail to Quebrada

Honda, 04°29 , 04"S, 79°07 , 59"W, 2570 m, 04 die 2006 (fl. $), DA. Neill 15361 (GH, MO, QCNE [n.v]).

Freziera tundaymensis D. Santam., sp. nov. (Fig. 3). Tipo: ECUADOR. Zamora-Chinchipe: Cordillera del Condor, vertiente

occidental, Parroquia Tundayme, Cuenca del Rio Tundayme, Camino al destacamento militar Condor Mirador, 03°37T5"S,

78°27 , 20"W, 1260 m, 14 feb 2008 (fl. $), W. Quizhpe, C.Juanga, L. Mayacu 2854 (holotipo: QCNE-223200 [n.v]; isotipos: GH!, MO-

6402469!).

This species differs from Freziera smithiana in its single trichome type (vs. two) and inflorescences of 5-12 flowers (vs. 5-7 flowers).

Arboles 10 m; ramas maduras cillndricas, la corteza externa negruzca, longitudinalmente estriada, gla-

brescentes; ramitas anguladas, velutinas o densamente adpreso pubescentes, los tricomas castano claros a

dorados, ca. 0.5 mm de longitud, sin papilas, escasamente lenticeladas, las lenticelas, si presentes, ellpticas o

orbiculares, blanquecina-grisaceas. Yema terminal conduplicada, 45-55 mm de longitud, densamente pu-

bescente, los tricomas castano claros a dorados, tricomas 1.0-1.5(-2.3) mm de longitud. Hojas con el peclolo

(27-)30-35 mm de longitud, caniculado, redondeado abaxialmente, densamente o esparcidamente pubescen-

te, papilado, las alas peciolares erectas, los margenes enteros, los tricomas castano claro a dorados, los tricomas

0.4-1.0 mm longitud y no asociados con papilas; coleteres ausentes; lamina 15-19.8 x 6.0-8.3 cm, ellptica, mas

o menos falcada, la base no revoluta, con un lado notablemente asimetrico, el lado largo de la lamina obtuso

a subtruncado, redondeado a decurrente en lado corto, el apice acuminado, seta negra no observada; margen

sinuado, 35-60 dientes por lado, con pequenas setas insertas en los senos, curvadas, negras o cafe rojizas, ca-

ducas, las setas rodeadas de tricomas; haz glabra y densamente pustulada; enves lanado, los tricomas 0.5-1.3

mm, sin papilas; nervio central por la haz por ca. 3.5 cm desde la base hacla el apice caniculado y glabrescente,

la parte restante como una cresta elevada de tricomas, sin pustulas; redon deado y densamente pubescente el

enves, esparcidamente pustalado, generalmente cubiertas por los tricomas; nervios laterales 45-50 por lado

(incluldos algunos intermedios), impresos y levemente surcados por la haz, elevados y redondeados por el en¬

ves. Inflorescencias de 5-12 flores fasciculadas, axilares o axilares y ramifloras; bractea 1.0-2.0 x 1.0 mm, en

la base del pedicelo, persistente y posiblemente caduca, triangular, aparentemente no carinada, el apice agudo,

la seta en el apice ausente, el margen entero, la superheie externa pubescente, la superheie interna glabra;

pedicelo 0.5-2.0(-4.0) mm de longitud, redondeado, erecto, pubescente; bracteolas 2, 2.0-3.0 x 2.0-2.2 mm,

opuestas o subopuestas, levemente desiguales, persistentes, suborbiculares, no carinada, la superheie externa

densamente pubescente, glabras en la superheie interna, el margen entero, el apice redondeado; botones flo-

rales 3.0-4.0 mm de ancho; sepalos 5, imbricados; sepalos externos 2, 2.8-4.0 x 3.0-3.5 mm, suborbiculares

a ampliamente ovados, apice redondeado y sin una seta negra, superheie externa densamente adpreso pubes¬

centes o el indumento principalmente hacla el centra del sepalo, superheie interna glabra, el margen cartaceo,

ciliado, sin setas en el margen; sepalos internos 3, 3.0-4.0 x 3.0, similares en forma a los externos, superheie

externa adpreso pubescentes, el indumento menos denso hacla el margen, superheie interna glabra, el apice

redondeado, generalmente quebradizo, el margen membranaceo, sin setas; petalos 5, 4.0-5.2 x 2.5-4.0 mm

en boton, amarillos, libres en la base, ovado, glabros, el margen membranaceo-cartaceo, el apice redondeado o

agudo. Flores estaminadas desconocidas. Flores pistiladas con 17-25 estaminodios, 0.03-0.05 mm de longi¬

tud, libres, lineares, pianos, agudos en el apice y con un lado oblicuo; gineceo ca. 3.0 mm de longitud, piriforme,

glabra, aparentemente 4-locular; estilo no separado apicalmente; estigma 4-lobulado. Frutos 4.0-7.0 x 4.0-5.0

mm, redondeados, de color desconocido, glabros; paredes del fruto ca. 0.3 mm de grueso; semillas ca. 21 por

fruto, cafe rojizas, de forma no muy dehnida o mas o menos cuneiformes, 1.0-1.5 mm de longitud, foveoladas.

Journal of the Botanical Research Institute of Texas 9(1)

Tabla 2. Comparacion entre algunos caracteres diagnostics entre Freziera neilli y F. parva.

Caracteres

F. neilli

F. parva

Tamano de hojas

3.5-6.1 x 2.7-3.7 cm

3.5-4.2X 1.7-2.1 cm

Indumento en enves

Viloso a sericeo

Glabro, o la pubescencia restringida

al nervio central

Longitud de peciolos

1.0-3.0 mm

5.0-7.0 mm

Base de hoja

Subtruncada a subcordada, algunas veces

con un lado asimetrico

Cuneada

Indumento en pedicelo

Glabro

Hirsuto

Numero de loculos

3 (segun el protologo)

Habitat, distribution y fenologia.—Freziera tundaymensis es endemica de Ecuador, conocida solo por la

coleccion tipo realizada en la region de la Cordillera del Condor, en la provincia de Zamora-Chinchipe. Ha sido

recolectada en bosque muy humedo a los 1260 m de elevacion. Flores pistiladas y frutos han sido recolectadas

en febrero; lo demas de su fenologia se desconoce.

Etimologia. —El eplteto especlfico, tundaymensis, hace referencia al rlo Tundayme, localidad donde fue

recolectado el tipo.

Discusidn.—Freziera tundaymensis se puede distinguir por sus laminas foliares densamente pubescentes

en el enves, mas o menos falcadas, con la base con un lado asimetrico y numerosos nervios laterales y peclolos

relativamente largos, y por las inflorescencias axilares o axilares y ramifloras con numerosas flores. Esta espe-

cie es similar a Freziera smithiana Kobuski de Colombia, de la cual se diferencia por tener peclolos mas cortos

(vs. 30-40 mm en F. smithiana), el nervio central por la haz densamente pubescente, al menos hacla la parte

distal (vs. glabro en todo la longitud), y pubescencia de un solo tipo (vs. rizada y recta); ademas F tundaymensis

tiene inflorescencias de 5-12 flores fasciculadas (vs. 5-7 flores fasciculadas). A razon de la base con un lado

asimetrico, tambien podrla ser comparada con F angulosa Tul. y F inaequilatera Britton, ambas endemicas de

Bolivia, y con Fforerorum A.H. Gentry de Panama. De la primera, se puede diferenciar por sus hojas sesiles o

casi as! (vs. largamente pecioladas en F. tundaymensis). Freziera inaequilatera se puede distinguir de F tunday¬

mensis por su indumento ferruglneo en las ramas, peclolos, pedicelos y bracteolas (vs. castano claro a dorados

en F tundaymensis), los peclolos 15-17 mm de longitud (vs. [27-] 30-35 mm), laminas foliares angostas de

4.0-6.0 cm de ancho (vs. 6.0-8.3 cm) y pedicelos de 3.0-4.0 mm de longitud (vs. 0.5-2.0[-4.0] mm). Final-

mente, se puede distinguir de Fforerorum por sus ramitas glabras y fuertemente anguladas (vs. ramitas veluti-

nas o densamente pubescentes y anguladas en F. tundaymensis) y laminas foliares sesiles y estrigosas en el en¬

ves (vs. pecioladas y lanadas en el enves).

Freziera yanachagensis D. Santam., sp. nov. (Fig. 4). Tipo: PERU. Pasco: Oxapampa, Distrito Huancabamba, Parque Nacional

Yanachaga-Chemillen, parte alta de la trocha Yanachaga-Palcazu, 10 o 22'42"S, 75°27'00"W, 2650 m, 01 die 2007 (fl. d), A. Monteg-

uado, A. Pena, V. Flores &R. Rivera 16076 (holotipo: USM-230940!; isotipos: GH!, MO-6402478!, USM-255371!).

Freziera yanachagensis differs from F. dudleyi in having small, elliptic or orbicular leaves with retuse apex and solitary, axillary inflores¬

cences.

Arboles o arbustos 1-6 m; ramas maduras cillndricas, la corteza externa negruzca y con manchas blanqueci-

nas, levemente estriadas longitudinalmente, glabrescentes; ramitas cillndricas, densamente pubescentes, los

tricomas blanquecinos a amarillento-dorados, 0.2-1.0(-1.5) mm de longitud, sin papilas, lenticelas ausentes.

Yema terminal conduplicada, 3.0-5.0 mm de longitud, velutina, los tricomas blanquecinos a amarillento-do¬

rados, 0.4-1.0 mm longitud. Hojas con el peclolo (0.5-)1.0-1.5 mm de longitud, piano o levemente caniculado

adaxialmente, redondeado abaxialmente, viloso o glabro, cuando pubescentes los tricomas cafe rojizos a dora¬

dos, tricomas 0.5-1.0 mm de longitud, las alas peciolares ausentes o cortamente aladas, aparentemente sin se-

tas en el margen; coleteres ausentes; lamina 0.5-0.9 x 0.55-0.6 cm, ellpticas o orbiculares, la base obtusa a re-

dondeada, igual o levemente desigual, no revoluta, el apice retuso y con una seta negra y curvada hacla el enves

de la lamina, caduca; margen entero en la mitad proximal, sinuado en la mitad distal, 3-6 dientes por lado

Santamaria-Aguilary Lagomarsino, Nuevas especies de Freziera

(levemente revoluto en toda su longitud), con pequenas setas insertas en los senos, conicas o levemente cur-

vadas, negras a cafe-rojizas, caducas, las setas sin tricomas alrededor; haz glabro, sin pustulas; enves adpreso

pubescente, los tricomas 0.8-1.5 mm de longitud y asociados con papilas, las papilas distribuidas sobre todo la

lamina, cafe-rojizas claro o cafe-rojizas oscuras a casi negras; nervio central por la haz piano o con una peque-

na cresta redondeada, glabro, levemente pustulado o las pustulas ausentes; semi-redondeado o aplanado en el

enves (algunas veces casi indistinto hacla el apice), adpreso pubescente, pustulado; nervios laterales 4-6 pares,

a menudo muy diflciles de distinguir o ausentes, casi inconspicuos en el enves. Inflorescencias de 1 flor soli-

taria, axilar (algunas veces las flores ocultas bajo las hojas); bractea 0.8-1.0 x 0.5-1.0 mm, en la base del

pedicelo, persistente o caduca, mas o menos triangular, carinada, el apice redondeado o agudo, que termina en

una seta conica, curvada, caduca, el margen ciliado, la superficie externa adpreso pubescente, la superficie in¬

terna glabra; pedicelo 0.5-3.2 mm de longitud, cillndrico, erecto o levemente curvado, glabro, puberulento o

adpreso pubescente; bracteolas 2, 0.6-1.0 x 0.7-1.0 mm, subopuestas, desiguales, persistentes, mas o menos

triangulares o en forma de ‘D’, no carinada, la superficie externa glabrescente o esparcidamente pubescentes,

rugosas, la superficie interna glabra, el margen ciliado, el apice redondeado y que termina en un seta negra o

cafe-rojiza, caduca; sepalos 5, imbricados; sepalos externos 2,1.2-1.3 x 1.1-2.8 mm, ampliamente ovados o en

forma de ‘D’, apice redondeado y con una seta negra, conica, superficie externa glabra o esparcidamente pubes¬

cente, superficie interna glabra, margen membranaceo, ciliado, algunas veces con diminutas setas en la base;

sepalos internos 3, 1.0-1.3 x 2.0-3.0 mm, iguales a los externos; petalos 5, (1.0-)3.0-3.5 x 2.5-2.8 mm, blan-

cos o blanco-rosados, levemente unidos en la base o libres, ovados o ampliamente ovados, glabros, el margen

membranaceo y entero, el apice obtuso y ligeramente curvado con unos diminutos dientes. Flores estamina-

das con (6—)10—16 estambres, 1.5-2.5 mm de longitud, libres o levemente unidos a la base de los petalos,

desiguales; filamentos 0.5-1.0 mm, mas o menos pianos o engrosados; anteras ca. 0.9-1.7 mm, irregularmente

moniliformes (no moniliformes en Montenegro 60), generalmente ovoides, la base subcordada o truncada,

apiculada en el apice; gineceo 0.7-2.0 mm de longitud, muy deforme o conico, glabro, 4-locular; estilo separado

por ca. 0.5 mm de longitud; estigma 2 o 3 lobulado. Flores pistiladas con (5-)10 estaminodios, 0.1-0.7 mm de

longitud, libres, lineares, pianos, agudos en el apice; gineceo 0.5-1.0 mm de longitud, redondeado, glabro, 4 o

3?-locular; estilo separado apicalmente por 0.3-0.5 mm de longitud; estigma 2, 3 o 4-lobulado. Frutos ca. 2.0

x 2.0 mm, redondeados, de color desconocido, glabros; paredes del fruto ca. 0.2 mm de grueso; semillas

desconocidas.

Habitat, distribucionyfenologia.—Frezierayanachagensis ha sido recolectada en Ecuador y Peru. En Ecua¬

dor, se conoce de bosque enano sobre roca arenisca entre los 1930-2000 m de elevacion en la Cordillera del

Condor en la provincia de Zamora-Chinchipe. En Peru, se conoce de bosque primario esclerohlo a los 2650 m

de elevacion en el Parque Nacional Yanachaga-Chemillen en la provincia de Oxapampa. Flores estaminadas

han sido recolectadas en diciembre; flores pistiladas y frutos han sido recolectadas en setiembre. Los frutos de

esta especie son poco conocidos; los dos duplicados estudiados tenlan pocos frutos y se encontraban agallados.

Por esa razon no fue posible observar las semillas.

Etimologia. —El eplteto especlhco hace referencia al Parque Nacional Yanachaga-Chemillen, la localidad

donde se recolecto el tipo.

Discusion.—Freziera yanachagensis se reconoce facilmente entre la mayorla de las especies del genero por

sus partes vegetativas y reproductivas muy pequenas. Tambien es unico en su genero por sus hojas que tienen

apice retuso, la venacion inconspicua, y el margen levemente revoluto en todo su longitud, entero en la mitad

proximal y sinuado en la mitad distal con 3-6 dientes pequenos, los cuales son poco visibles sin el uso de lupa.

Ademas, se caracteriza por tener el enves adpreso pubescente con tricomas asociados con papilas cafe-rojizas

y flores generalmente con las anteras irregularmente moniliformes y el estilo separado apicalmente. A razon de

estas dos ultimas caracterlsticas se podrla comparar con Freziera dudleyi A.H. Gentry de Peru y Bolivia y con F.

humiriifolia aqul descrita. Sin embargo, se puede distinguir por los caracteres enumerados en la Tabla 3. Tam¬

bien por tener laminas foliares pequenas, la nueva especie se podrla comparar Freziera minima A.L. Weitzman

de Ecuador y con F. stuebelii (Hieron.) A.L. Weitzman de Colombia, hasta el momento es conocida solamente

100

Journal of the Botanical Research Institute of Texas 9(1)

Tabla 3. Comparacion entre algunos caracteres diagnostics entre Freziera yanachagensis, F. dudleyiyF. humiriifolia.

Caracteres

F. yanachagensis

F. dudleyi

F. humiriifolia

Forma de la hoja

Elipticas o orbiculares

Angostamente elipticas

a casi lineares

Ovadas a oblongo elipticas

Tamano de las hojas

0.5-0.9 x 0.55-0.6 cm

7.8-9.3X 1.2-2.2 cm

7.5-14x3.3-6.6 cm

Apice de hoja

Retuso

Acuminado

Inflorescencia

Siempre axilares y solitarias

Axilares y ramifloras de 1 -3

flores fasciculadas

Axilares y ramifloras de 3-6 flores

fasciculadas

por la fotografla del holotipo (Weitzman, 1987). A1 comparar con F. yanachagensis, ambas de esas especies

tienen laminas foliares mas grandes (0.7-1.21 x 0.49-1.4 cm), los margenes foliares crenados, y los nervios

visibles en ambas superficies (mas conspicuos, ademas son bifurcados en F. stuebelii). La primera tambien

difiere por sus sepalos externos entre 3.0-3.6 x 2.5-3.3 mm (vs. 1.2-1.3 x 1.1-2.8 mm enF. yanachagensis) y por

sus frutos mas grandes (6.8-7.5 x 5.1-5.5 vs. ca. 2.0 x 2.0 mm). La segunda difiere por sus laminas foliares con

la base cordada. Freziera microphylla Sandwith y F. suberosa Tul. tambien tienen hojas pequenas y margenes

foliares con apariencia de no tener dientes y levemente revolutos en todo su longitud, sin embargo en estas dos

especies las laminas foliares son ellpticas a oblongo-ellpticas, entre 1.07-2.8 cm de longitud, el enves densam-

ente serlceas, y con los tricomas amarillos a amarillo-dorados y no relacionados con papilas.

Especimenes adicionales examinados (paratipos). ECUADOR. Zamora-Chinchipe: El Pangui, Cordillera del Condor, Cresta de la Cordil¬

lera, en la frontera Ecuador-Peru, 1 km al sur del destacamento militar Condor Mirador, bosque enano, sobre roca arenisca, 03°38’32”S,

78°23’36”W, 2000 m, 16 die 2000 (fl. S'), W. Montenegro & Grupo Post-Grado MO-QCNE 60 (MO, GH); El Pangui, Cordillera del Condor,

summit of sandstone plateau of Cordillera, SE of headwaters of Rio Wawaime, above proposed EcuaCorriente copper mine area, 03°35’40”S,

078°35’40”W, 1930 m, 19 set 2006 (fl. $ & fr), DA. Neill et al 15233 (GH, MO, QCNE [n.v]).

NUEVO REGISTRO PARA ECUADOR

Freziera carinata A.L. Weitzman, J. Arnold Arbor. 68 (3):325. 1987. Tipo: Venezuela. Bolivar: Auyan-tepui, cumbre de la

parte central occidental (division occidental del cerro), vecindad del “Drizzly Camp,” sobre piedra de arenisca, a lo largo de afluente

del Rio Churun, 1760 m, 04 may 1964, J. Steyermark 93366 (holotipo: GH!; isotipo: K, NY, U, US, VEN).

Habitat, distribucion y fenologia.—Freziera carinata se encuentra en Ecuador y Venezuela. En Ecuador, ocurre

en la provincia de Morona-Santiago, donde ha sido recolectada en bosque denso, bosque bajo, y sobre vege-

tacion expuesta en suelo rocoso, entre los 1220-2700 m de elevacion. En Ecuador, flores pistiladas han sido

recolectadas en diciembre y frutos en octubre; flores estaminadas todavla son desconocidas.

Discusidn.—Freziera carinata se reconoce por sus ramitas aplanadas y glabras, peclolos marcadamente

carinados abaxialmente, laminas foliares con el enves denso y diminutamente estrigoso pubescente y con

nervio central carinado, la base marcadamente revoluta y por sus flores con las bracteolas y sepalos glabros.

Freziera carinata en el lenguaje Shuar se le conoce con el nombre comun Tsempunim (A. Wisum 36).

Algunas colecciones todavla no identihcadas del Cerro Golondrinas, Carchi, Ecuador ( Palacios 12505,

MO; Palacios 12587, MO; Boyle et al 2338, MO; Boyle et al 3410, MO; y Bradford 146, MO) presentan similitudes

con F. carinata, pero podrla representar un nuevo taxon. Ambas poseen ramitas, bracteolas y sepalos glabros y

ramitas levemente surcadas. Sin embargo, todo el material mencionado posee laminas foliares un poco mas

pequenas (6.2-11 x 3.1-4.1 cm) con el nervio central por la haz redondeado y obviamente pubescente a simple

vista, los pedicelos gruesos, y las bracteolas muchas veces mas grandes (2.0-)3.5-5.0 mm de longitud que en F.

carinata.

Especimenes examinados. ECUADOR. Morona-Santiago: Limon Indanza, Cordillera del Condor, summit ridge of the Cordillera, quartz¬

bearing igneous rock substrate, headwaters of the Rio Warintza, SW of Shuar village of Warints, 03 o 15'37"S, 78 0 19'18"W, 2700 m, 16 die

2002 (fl. $ & fr), DA. Neill & Grupo Shuar de Conservacion 14152 (GH, MO, QCNE [n.v]); Cordillera del Condor, Cumbre del Cerro Paatin

Naint, 03°13 , 59"S, 78°16 , 15"W, 1220 m, 11 oct 2002 (fr), A. Wisum & Grupo Shuar de Conservacion 36 (GH, MO, QCNE [n.v]); Cordillera del

Condor, Centro Shuar Warints, recorrido desde el Cerro Chankinias hasta la Cordillera del Condor, 03 o 14'41"S, 78 0 17'29"W, 2200 m, 15 oct

2002 (bot. fl & fr), A. Wisum & Grupo Shuar de Conservacion 55 (GH, MO).

Santamaria-Aguilary Lagomarsino, Nuevas especies de Freziera

101

AGRADECIMIENTOS

Deseamos agradecer a los siguientes herbarios y su personal por las facilidades brindadas y por permitir el ac-

ceso y uso de sus colecciones: A, BM, CAY, CR, F, GH, INB, LPB, LSCR, MO, MOL, NY, PMA, SCZ, y USM.

Gracias al Missouri Botanical Garden (MO) por el envlo de material para identibcacion y por el apoyo

economico gracias a la beca Elizabeth E. Bascom para Botanicas(os) Latinoamericanas(os), que permitio una

pasantla en dicho herbario para D.S.A. Agradecemos a Barney Lipscomb, David A. Neill, y un revisor anonimo

por los comentarios que ayudaron a mejorar la presente contribucion; D.A. Neill tambien proporciono infor-

macion de los especlmenes depositados QCNE. D.S.A. tambien desea expresar su gratitud a R. Abbott, A.L.

Arbelaez, y R. Liesner por su apoyo durante las visitas a MO. Igualmente, las gracias al Harvard University

Herbaria y todo su personal por todas las facilidades brindadas y su hospitalidad. Kanchi Gandhi (GH) nos dio

ayuda en los aspectos nomenclaturales.

REFERENCIAS

Berry, P.E. &A.L. Weitzman. 2008.Theaceae. In: 0. Hokche, P.E. Berry &0. Huber, eds. Nuevo Cat. FI. Vase Venez. Fundacion

Instituto Botanico de Venezuela, Caracas. Pp. 640-641.

Berry, P.E. & A.L. Weitzman. 2007. Ternstroemiaceae. In: V.A.Funk, P.E. Berry, S. Alexander, T.H. Hollowell & C.L. Kelloff.

Checklist of the plants of the Guiana Shield (Venezuela: Amazonas, Bolivar, Delta Amacuro; Guyana, Surinam, French

Guiana). Contr. U.S. Natl. Herb. Pp. 535-536.

Berry, P.E. & A.L. Weitzman. 2005. Ternstroemiaceae. In: J.A. Steyermark, P.E. Berry & B.K. Holst, eds. FI. Venez. Guayana.

Missouri Botanical Garden Press, St. Louis. Pp. 300-308.

Berry, P.E., O. Huber, & B.K. Holst. 1995. Phytogeography of the Guayana Region. In: J.A. Steyermark, P.E. Berry & B.K. Holst,

eds. Flora of the Venezuelan Guayana. Introduction. Missouri Botanical Garden, St. Louis. Pp. 170-192.

Clark, J.L., D.A. Neill, A. Weber, J.A. Gruhn, &T. Katan. 2010. Shuaria (Gesneriaceae), an arborescent new genus from the

Cordillera del Condor and Amazonian Ecuador. Syst. Bot. 35:662-674.

Daly, D.C., D.A. Neill, & M.C. Martinez-Habibe. 2012. An ecologically significant new species of Dacryodes from the northern

Andes. Studies in Neotropical Burseraceae XV. Brittonia 64(1):49-56.

Every, J.L.R. 2009. Neotropical Pentaphylacaceae. In: W. Milliken, B. Klitgard. & A. Baracat (2009 onwards), Neotropikey

- Interactive key and information resources for flowering plants of the Neotropics, www.kew.org/science/tropa-

merica/neotropikey/families/Pentaphylacaceae.htm. Accesado Diciembre 2014.

Grant, J.R. 2014. De Macrocarpaeae Grisebach (ex Gentianaceis) speciebus novis XI: Five new species from the Andes of

Ecuador and Colombia. Harvard Pap. Bot. 19:227-239.

Neill, D.A. 2005. Cordillera del Condor: Botanical treasures between the Andes and the Amazon. PI. Talk 41:17-21.

Neill, D.A (coord.). 2007. Botanical Inventory of the Cordillera del Condor Region of Ecuador and Peru, Missouri Botani¬

cal Garden. Extended full report available at www.mobot.org/mobot/research/ecuador/cordillera/welcome.shtml.

Accesado die 2014.

Neill, D.A. & M. Asanza. 2012. Lozania nunkui (Lacistemataceae), a new species from the sandstone plateaus of the Cordil¬

lera del Condor in Ecuador and Peru. Novon 22:207-211.

Neill, D.A. H. Beltran, & W. Quizhpe. 2012. Clethra concordia (Clethraceae), a shrubby new species from the crest of the

Cordillera del Condor on the Peru-Ecuador border. Novon 22:212-216.

Riina, R., M.A. Vigo, & C.E. Ceron. 2014. Croton condorensis : an enigmatic new species of Euphorbiaceae from southern

Ecuador. Phytotaxa 164:154-158.

Rodriguez, E.F. (& 6 autores mas). 2006. Nuevas adiciones de angiospermas a la Flora del Peru procedentes de la Cordil¬

lera del Condor y areas adyacentes. Arnaldoa 13(2):318-322.

Schulenberg, T.S. & K. Awbrey (eds.). 1997. The Cordillera del Condor region of Ecuador and Peru: A biological assessment.

RAP Working Papers 7:1-231.

StAhl, B. 2010. Additions to the knowledge of the genus Symplocos (Symplocaceae) in Ecuador and Peru. Novon

20:84-94.

Ulloa Ulloa, C. & D.A. Neill. 2006. Phainantha shuariorum (Melastomataceae), una especie nueva de la Cordillera del

Condor, Ecuador, disyunta de un genero guayanes. Novon 16:281-285.

van derWerff, H.H. 1993.Theaceae. In: L. Brako&J.L. Zarucchi, eds. Catalogue of the flowering plants and gymnosperms

of Peru, Monogr. Syst. Bot. Missouri Bot. Gard. 45:1144-1146.

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Weitzman, A.L. 1987. Taxonomic studies in Freziera (Theaceae), with notes on reproductive biology. J. Arnold Arbor.

68(3):323-334.

Weitzman, A.L. 1999. Theaceae. In: P.M. Jorgensen & S. Leon-Yanez, eds. Catalogue of the vascular plants of Ecuador.

Monogr. Syst. Bot. Missouri Bot. Gard. Pp. 923-924.

Weitzman, A.L., S. Dressler, & P.F. Stevens. 2004.Ternstroemiaceae. In: K. Kubitzki, ed. Families and genera of vascular plants.

Vol. 6. Springer-Verlag, Berlin. Pp. 450-460.

NEW COMBINATIONS IN HEXASEPALUM (RUBIACEAE: SPERMACOCEAE)

Joseph H. Kirkbride, Jr.

Piero G. Delprete

USDA-ARS, US. National Arboretum

Floral & Nursery Plants Research Unit

3501 New York Avenue NE

Washington, DC 20002-1958, US.A.

joseph.kirkbride@ars.usda.gov

Herbierde Guyane, IRD - UMRAMAP

Boite Postale 165, 97323 Cayenne Cedex

Guyane Frangaise (French Guiana), FRANCE

piero. delprete@ird. fr

ABSTRACT

Ten new combinations in Hexasepalum (Rubiaceae: Spermacoceae) are here proposed, for species formerly in Diodella.

RESUMEN

Se proponen diez nuevas combinaciones en Hexasepalum (Rubiaceae: Spermacoceae), para las especies anteriormente posicionadas en

Diodella.

In April of 1913, Small (1913) published the new genus Diodella Small, a segregate from the genus Diodia L., and

transferred one species from Diodia to Diodella, Diodella rigida (Cham. & Schltdl.) Small. He distinguished

Diodella from Diodia by its unbranched styles, capitate or bilobed stigmas, and funnelform corollas versus Dio¬

dia with branched styles, elongate, slender stigmas, and salverform corollas. In September of the same year, he

(Small & Carter 1913) transferred a second Diodia species into Diodella, Diodella teres (Walter) Small. During

the following decades, other than Small himself, rubiaceous specialists considered Diodella to be synonymous

with Diodia.

Bacigalupo and Cabral (1999) revised the American species of Diodia, and altered the circumscription of

Diodia from approximately 43 to five species. They consigned 13 species to Borreria G. Mey. and nine species

to Galianthe Griseb., and suggested that 16 Diodia species belonged to Diodella. Of those 16 species, seven have

been reported to be junior synonyms of other Diodella species, and three additional species have been added to

Diodella, which leaves the genus with 12 accepted species at the present time. Bacigalupo and Cabral distin¬

guished Diodella by its isomorphic flowers, capitate-bilobed stigmas, fruits consisting of indehiscent meri-

carps that separate completely and are shed independently at maturity, seeds with an apical, transverse fold,

and echinulate pollen. Following the delimitation of Diodella proposed by Bacigalupo and Cabral (1999), Del¬

prete published a few new combinations in this genus, in the Rubiaceae treatments of Flora Ilustrada Catari-

nense. Flora dos Estados de Goias e Focantins, and in a synopsis of the Rubiaceae occurring in the states of Mato

Grosso and Mato Grosso do Sul, Brazil (Delprete et al. 2004; Delprete 2010; Delprete & Cortes-B. 2007).

Cabana Fader et al. (2012) examined the holotype of Hexasepalum Bartl. ex DC. and H. angustifolium

Bartl. ex DC. at PR and discovered that they are synonymous with Diodella and Diodella crassifolia (Benth.)

Borhidi, respectively. Since Hexasepalum has priority over Diodella this would require the transfer of 11 names

now accepted in Diodella into Hexasepalum. Cabana Fader et al. (2012) considered, “this would be most disrup¬

tive nomenclaturally because the name Diodella is, at present, extensively used, while Hexasepalum has rarely

been mentioned since its publication and generally treated as uncertain”. Therefore they proposed that Hexas¬

epalum and H. angustifolium both be rejected, which would allow the continued use of Diodella and its specific

names. The members of the Nomenclature Committee for Vascular Plants (Applequist 2013) voted one in fa¬

vor, 14 against, and three abstentions for rejection of Hexasepalum and H. angustifolium. The Committee felt

“that this is one of those cases in which implementation of the Principle of Priority at the generic level through

the adoption of Hexasepalum will not cause excessive disruption, so [did] not recommend rejection”, and “was

not convinced that a change of name for this species would be unusually disruptive, so [did] not recommend

the proposal” to reject H. angustifolium. This means that it is not possible to use the generic name Diodella or its

J. Bot. Res. Inst. Texas 9(1): 103 -106.2015

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Journal of the Botanical Research Institute of Texas 9(1)

specific names. To facilitate their use we are here transferring 10 specific names accepted in Diodella to the

genus Hexasepalum. The name Diodia teres Walter was already recently transferred to Hexasepalum by Kirk-

bride (2014).

SYSTEMATIC TREATMENT

Hexasepalum Bard, ex DC., Prodr. 4:561. 1830. Type: Hexasepalum angustifolium Bartl. ex DC.

Diodella Small, Fl. Miami 177.1913. Type: Diodella rigida (Cham. & Schltdl.) Small.

Hexasepalum angustatum (Steyerm.) J.H. Kirkbr. & Delprete, comb. nov. Basionym: Diodia angustata Steyerm., Los

Angeles County Mus. Contr. Sci. 21:26. 1958; Diodella angustata (Steyerm.) E.L. Cabral & Cabana Fader, Rodriguesia 61(1): 120.

2010. Type: BRAZIL. Goias: Chapada dos Veadeiros, 14 km S of Veadeiros [Alto Paraiso], 14°15'S, 47°30'W, 24 Apr 1956, E.Y. Dawson

14672 (holotype: R; isotypes: F, LAM, US!).

Distribution. —Endemic to the Chapada dos Veadeiros, Goias, Brazil.

Hexasepalum angustifolium Bartl. ex DC., Prodr. 4:561. 1830. Type: MEXICO. Hidalgo: Regio-Monte (Real del Monte)

Mexicanorum, s.d. [1791-1792], T.P.X. Haenkes.n. (holotype: PR; isotypes: G, MO!).

Diodia crassifolia Benth., Bot. Voy. Sulphur 108. 1845; Diodella crassifolia (Benth.) Borhidi, Rubiac. Mexico, ed. 1, 185. 2006. Type:

MEXICO. Nayarit: San Bias near Tepic, G.W Barklay s.n. (holotype: BM!).

Distribution. —Endemic to Mexico in the states of Nayarit and Sinaloa.

Hexasepalum apiculatum (Willd.) Delprete &J.H. Kirkbr., comb. nov. Basionym: Spermacoceapiculata Willd. inRoem.

& Schult., Syst. Veg. 3:531.1818; Diodella apiculata (Willd.) Delprete in A. Reis, Fl. Illustr. Catarinense RUBI, vol. 1:169. 2004. Type:

BRAZIL: without locality, s.d. [1804], F.W. Sieber in Hoffmannsegg s.n. (holotype, B-W 2626!).

Johann Centurius Graf von Hoffmannsegg never traveled outside of Europe (Lanjouw & Stafleu 1957). Franz

Wilhelm Sieber collected for Hoffmannsegg in Para, Brazil in 1804 (Lanjouw & Stafleu 1957; Vegter 1986), and

Hoffmannsegg distributed those specimens with labels bearing only his name. Therefore, Hoffmannsegg’s

Brazilian exsiccatae were actually collected by F.W. Sieber.

Distribution. —Ranges from Colombia, Venezuela, the Guianas, and Ecuador to Brazil and Paraguay.

Hexasepalum gardneri (K. Schum.) J.H. Kirkbr. & Delprete, COmb. nov. Basionym: Diodia gardneri K. Schum. in Mart.,

Fl. bras. 6(6):402. 1889; Diodella gardneri (K. Schum.) Bacigalupo & E.L. Cabral, Darwiniana 44:98. 2006. Type: BRAZIL. CearA:

Aracaty, G. Gardner 1705 (holotype: Bf ; lectotype, here designated: BM!).

Distribution. —Known from the Brazilian states of Bahia, Ceara, and Piaul.

Hexasepalum lippioides (Griseb.) J.H. Kirkbr. & Delprete, comb. nov. Basionym: Diodia lippioides Griseb., Cat. pi. Cub.

141. 1866; Diodella lippioides (Griseb.) Borhidi, Acta Bot. Hung. 51:275. 2009. Type: CUBA: C. Wright 2762 (lectotype, here desig¬

nated: GOET11312!; syntypes: G, GH, GOET 11313!, K, MO, MPU, NY).

There are two duplicates of W right 2762 at GOET, and they appear to be independent gatherings. GOET11312

has a specimen label with Grisebach’s handwritten diagnosis, so it is chosen here as lectotype.

Distribution. —Endemic to Cuba.

Hexasepalum mello-barretoi (Standi.) J.H. Kirkbr. & Delprete, comb. nov. Basionym: Diodia mello-barretoi Standi.,

Publ. Field Mus. Nat. Hist., Bot. Ser. 22:113. 1940; Diodella mello-barretoi (Standi.) Bacigalupo & E.L. Cabral, Darwiniana 44:100.

2006. Types: BRAZIL. Minas Gerais: Municipio Santa Luzia: Serra do Cipo, Km 116,13 Jan 1934, H.L. deMello Barreto 3541 (holotype:

F; isotype: BHCB!); 2 Feb 1938, H.L. de Mello Barreto 9274 (paratype: BHCB!).

Standley (1940) cited an isotype and paratype of Diodia mello-barretoi at the Jardim Botanico de Belo Hori¬

zonte, Minas Gerais, Brazil. Those specimens were deposited in the Museu de Historia Natural (BHMH), Uni-

versidade Federal de Minas Gerais, Belo Horizonte, Minas Gerais, Brazil. The specimens in BHMH were trans¬

ferred to the Departamento de Botanica (BHCB), Institute de Ciencias Biologicas, Universidade Federal de

Minas Gerais where they remain today (pers. comm.: Alexandre Salino 2014).

Distribution. —Apparently endemic to Serra do Cipo, Municipio Santa Luzia, Minas Gerais, Brazil.

Kirkbride and Delprete, New combinations in Hexasepalum

105

Hexasepalum radulum (Willd.) Delprete & J.H. Kirkbr., comb. nov. Basionym: Spermacoce radula Willd. in Roem. &

Schult., Syst. Veg. 3:531.1818; Diodella radula (Willd.) Delprete in A. Reis, Fl. Illustr. Catarinense RUBI, vol. 1:174. 2004. Type: BRA¬

ZIL. ParA: s.d. [1804], F.W. Sieber in Hoffmannsegg s.n. (holotype: B-W!).

Distribution. —Widespread in Central and South America; reported from the state of Virginia (USA) in 1959

(Reed 1964), but not persistent.

Hexasepalum rosmarinifolium (Pohl ex DC.) Delprete & J.H. Kirkbr., comb. nov. Basionym: Diodia rosmarinifolia

Pohl ex DC., Prodr. 4:564.1830; emend. K. Schum. in Mart., Fl. Bras. 6(6):18.1888; Diodella rosmarinifolia (Pohl ex DC.) Bacigalupo

& E.L. Cabral ex Delprete & Cortes-Ballen, Rev. Biol Neotrop. 3:34. 2006 [2007]. Type: BRAZIL: without locality, s.d. ,J.E. Pohl s.n.

(holotype G-DC1).

Distribution. —Occurring in southern Venezuela, and northern and central-western Brazil.

Hexasepalum sarmentosum (Sw.) Delprete & J.H. Kirkbr., comb. nov. Basionym: Diodia sarmentosa Sw., Prodr. 30. 1788;

Diodella sarmentosa (Sw.) Bacigalupo & Cabral, Darwiniana 44:100. [Jul] 2006. Type: JAMAICA: without locality, s.d., O.P. Swartz

s.n. (syntypes: S S-R-1510!, S S08-117!).

Molecular phylogenetic studies (Dessein 2003; Groeninckx et al. 2009) have suggested that this species is not

a member of the genus Hexasepalum. Pending further, more definitive phylogenetic studies we are maintaining

H. sarmentosa in the genus Hexasepalaum.

Distribution .—Widespread throughout the Neotropics, in Mexico, Central America, the Antilles, and

South America to Bolivia and Brazil; also present in tropical Africa, Malay Peninsula, and Pacific Islands.

Hexasepalum scandens (Sw.) J.H. Kirkbr. & Delprete, comb. nov. Basionym: Diodia scandens Sw., Prodr. 30.1788; Diodella

scandens (Sw.) Bacigalupo & E.L. Cabral, Darwiniana 44:104. 2006. Type: HISPANIOLA: without locality, s.d., O.P. Swartz s.n. (holo¬

type: SS-R-1511!).

Distribution .—Originally endemic to Hispaniola, and currently widespread and naturalized in tropical Africa.

Hexasepalum serrulatum (P. Beauv.) J.H. Kirkbr. & Delprete, COmb. nov. Basionym: Spermacoce serrulata P. Beauv., Fl.

Oware 1:39.1805; Dioda serrulata (P. Beauv.) G. Taylor in A.W. Exell, Cat. Vase. Pi. S. Tome 220.1944; Diodella serrulata (P. Beauv.)

Borhidi, Rubiac. Mexico, ed. 1,185. 2006. Type: NIGERIA: Oware, A.M.F.J. Palisot deBeauvois s.n. (holotype: G00014459!).

Distribution. —Coastal areas of western Africa, from Senegal to Angola, Macaronesia, southern Mexico to Co¬

lombia, and the Antilles.

Hexasepalum teres (Walter) J.H. Kirkbr., J. Bot. Res. Inst. Texas 8(1): 17. 2014. Basionym: Diodia teres Walter, Fl. carol.

87. 1788; Diodella teres (Walter) Small in Small & J.J. Carter, Fl. Lancaster Co. 271. 1913. Type: UNITED STATES. South Carolina:

Georgetown Co.: Georgetown, old field, 24 Aug 1939, R.K. Godfrey & R.M. Tryon 1682 (neotype, designated by Ward (2008: 467):

GH00277018!; isoneotypes: NY1163926!, US1838313!).

Distribution .—Its natural distribution is from North to South America and the Antilles. It is introduced and

naturalized in the Netherlands, many African countries, China, Japan, and Korea.

ACKNOWLEDGMENTS

We thank the directors and curators of the B, BM, G, GH, GOET, L, NY, R, and US herbaria for facilitating ac¬

cess to their collections and for loans of specimens and Alexandre Salino (BHCB) for the history of the isotype

and paratype of Diodia mello-barretoi and images of the specimens. We are also grateful to Steven Darwin (NO),

David Lorence (PTBG), and John Wiersema (BARC) for their helpful revisions of the manuscript.

REFERENCES

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

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

Darwiniana 37:153-165.

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

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

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Journal of the Botanical Research Institute of Texas 9(1)

Delprete, P.G. 2010. Rubiaceae - Parte 1: Introdugao, Generos A-H. In: J.A. Rizzo, coord. Flora dos Estados de Goias e To¬

cantins. IRD/UFG, Universidade Federal de Goias, Goiania, Brazil 40:1-580.

Delprete, P.G. & R. Cort£s-B. "2006" [2007]. A synopsis of the Rubiaceae of the states of Mato Grosso and Mato Grosso do Sul,

central-western Brazil, with a key to genera, and a preliminary species list. Rev. Biol. Neotr. 3:13-96.

Delprete, P.G., L.B. Smith & R.B. Klein. 2004. Rubiaceas, Volume 1 - Generos de A-G: 1 .Alseis ate 19. Galium. In: A. Reis, ed.

Flora ilustrada Catarinense. Herbario Barbosa Rodrigues, Itajai, Santa Catarina, Brazil. Pp. 1-344.

Dessein, S. 2003. Systematic studies in the Spermacoceae (Rubiaceae). Ph.D. dissertation, Katholicke Universiteit Leuven,

Leuven, Belgium.

Groeninckx, I., S. Dessein, H. Ochoterena, C. Persson,T.J. Motley, J. KArehed, B. Bremer, S. Huysmans & E. Smets. 2009. Phylogeny of

the herbaceous tribe Spermacoceae (Rubiaceae) based on plastid DNA data. Ann. Missouri Bot. Gard. 96:109-132.

Kirkbride, J.H., Jr. 2014. Hexasepalum teres (Rubiaceae), a new combination. J. Bot. Res. Inst. Texas 8:17-18.

Lanjouw, J. & Stafleu, F.A. 1957. Index Herbariorum: Collectors E-H, Part II (2). Regnum Veg. 9:280.

Reed, C.F. 1964. A flora of the chrome and manganese ore piles at Canton, in the port of Baltimore, Maryland and at

Newport News, Virginia, with descriptions of genera and species new to the flora of eastern United States. Phytolo-

gia 10:321-406.

Small, J.K. 26 Apr 1913. Flora of Miami. J.K. Small, New York, U.S.A.

Small, J.K. & J.J. Carter. 3 Sep 1913. Flora of Lancaster County. J.K. Small & JJ. Carter, New York, U.S.A.

Standley, P.C. 1940. Studies of American plants—X. Publ. Field Mus. Nat. Hist., Bot. Ser. 22:65-129.

Vegter, I.H. 1986. Index Herbariorum: Collectors S, Part II (6). Regnum Veg. 144:890.

Ward, D.B. 2008. Thomas Walter typification project, V: Neotypes and epitypes for 63 Walter names of genera D through

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

OBSERVATIONS ABOUT PHASEOLUS LIGNOSUS

(LEGUMINOSAE: PAPILIONOIDEAE: PHASEOLEAE),

A BEAN SPECIES FROM THE BERMUDA ISLANDS

D.G. Debouck

Centro Internacional deAgricultura Tropical

AA 6713, Cali, COLOMBIA

d. debouck@cgiar. org

ABSTRACT

This work is the validation of a bean species, Phaseolus lignosus Britton, endemic to the Bermuda archipelago. The plant grown out in Colom¬

bia from original seed collected on Tong Island, Bermuda, matches with the original description. Its morphological characteristics of vegeta¬

tive parts, raceme, flower and pod, being complete all suggest an affiliation with section Paniculati of this genus. An epitype has been desig¬

nated and distributed since any reference plant material was missing.

RESUMEN

Este trabajo es la validacion de una especie de frijol, Phaseolus lignosus Britton, endemica en el archipielago de Bermuda. Las plantas culti-

vadas en Colombia a partir de semillas originales colectadas en la isla Long Island de Bermuda, se conforman completamente con la descrip-

cion original. Sus caracteristicas morfologicas de las partes vegetativas, racimo, flor y vaina, ya completamente descritas sugieren una afili-

acion con la seccion Paniculati de este genero. Se designo y se distribuyo un epitipo por la falta de material vegetal de referencia.

Key Words: wild Bermuda bean, endemic species, Lima bean, type specimen

INTRODUCTION

On February 28, 1918 Nathaniel Lord Britton described Phaseolus lignosus, a new bean species, apparently

endemic to the Bermuda archipelago (SW North Atlantic Ocean) (Britton 1918, p. 183). This name is acknowl¬

edged as valid in the TROPICOS database (Tropicos 2013) and in the USDA germplasm system (Wiersema et

al. 1990). It however lacks several morphological details in its description and indications about plant type

specimens. For example, Figure 205 is mentioned in the text of the description but with no figure caption in

the ‘Flora of Bermuda’ (op. cit., p. 183), nor any indication that the illustration should be considered as the type,

leaving a doubt wheter the author meant something else. The description in effect ends with a bracket “[P.

semierectus ofReade]” (quoting).

Oswald A. Reade (1883, p. 21), possibly in one of the first plant inventories for that archipelago, described

in the non-cultivated Phaseolus species, only a P. semierectus, with no illustration nor any specimen mentioned.

A taxon P. semierectus was named by Linnaeus inl767, and listed in the ‘Flora Brasiliensis’ by Martius (1859-

62, p. 189-190) under section Macroptilium (and currently it is Macroptilium lathyroides var. semierectum (L.)

Urban (Marechal et al. 1978)). A careful reading of Reade’s description indicates that two legumes are indeed

involved in that text: the Bermuda bean of Britton (“stem twining and ascending among trees, etc, ten to twelve

feet; leaflets ovate pointed;... habitat, woods near caves (Joyce’s)”), and M. lathyroides (“pod narrowly cylindri¬

cal, three inches long. Distribution West Indies”). About the later, the readership is invited to consult the re¬

view by Barbosa-Fevereiro (1986-87, p. 133-136) about morphology and distribution. So, the protologue by

Britton includes elements that refer to two different legumes in two different genera, viz. Phaseolus L. and

Macroptilium (Bentham) Urban. This situation possibly explains why this name was still classified as ‘unre¬

solved’ in the Plant List with data supplied by March 23, 2012 (ThePlantList 2013).

By correcting this caption omission the illustration Fig. 205 (in Britton 1918, p. 183) could serve as a lec-

totype following the Articles 9.2., 9.3. and 9.11 of the Code (McNeill et al. 2012; Turland 2013). The date of

publication of this sketchy figure—after the deadline of 1 January 1908—would question the validity of the

J. Bot. Res. Inst. Texas 9(1): 107 -119.2015

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Journal of the Botanical Research Institute of Texas 9(1)

publication of this taxon along Art. 38.7 and 38.8. of the Code (McNeill et al. 2012; Turland 2013, p. 38 and p.

116). However, along Art. 38.9, an analysis is a figure or group of figures (where one would expect a text!) sepa¬

rate from the main illustration; therefore Art. 38.7 and 38.8 would not apply in this case. On the other hand, by

omitting the bracket “[P. semierectus of Reade],” a source of confusion would be eliminated, while it seems im¬

portant to indicate which parts of the original protologue actually apply to the Bermuda bean. For P. lignosus,

we thus have a description awaiting resolution and a figure (Fig. 205) that constitutes the lectotype.

Lacking a reference plant specimen may be a serious limiting factor in current (e.g. Delgado-Salinas et al.

1999, 2006; Lopez-Lopez et al. 2013) and future phylogenetic studies or any other requiring DNA, as stressed

elsewhere (Spencer et al. 2007, p. 17; Turland 2013, p. 71). Incidentally, is the lack of reference plant material the

explanation for the absence of P. lignosus in most previous reviews of the genus (e.g. Piper 1926; Lackey 1983;

Delgado-Salinas 1985; Freytag & Debouck 2002)? The purpose of this note is thus to fill in these missing data

in order to further validate this taxon—precisely along the need expressed by the later authors (p. 117)—and

to orient future botanical and ecological studies. Three points will be dealt with hereafter: i) the description of

a living bean plant received as P. lignosus from the Millennium Seed Bank (MSB, Kew, England), ii) the com¬

parison of it with the original description and subsequent validation, and iii) a discussion about findings and

relationships with other bean species.

MATERIALS AND METHODS

The plant material used in this study was obtained from MSB under serial number 515078. It is a collection

made by C. Watlington (as CW3) in Bermuda, Warwick (not Warick as indicated on the label at K), Cluster

Cottage Gardens, on 25 April 2008. This collection is also represented by two original herbarium specimens

kept in the Herbarium of the Royal Botanic Gardens, Kew, with bar codes K000650153 and K000650154 (De¬

bouck 2014). The material has been grown in the period April 2013 to August 2014 in closed glass-house at

CIAT Palmira, Valle and next in Popayan, Cauca, Colombia (lat. 02° 3T 02.8"N; long. 76° 38' 04.4"W; alt. 1765

masl). Observations were made on living plants in both stations, using calipers and color charts (Royal Horti¬

cultural Society (RHS) 1966). Voucher specimens were made out of the living plants by cutting flowering and

fruiting stems, and distributed later on to different Herbaria (see under Designation below). Small amounts of

seed, accessioned as G40876 (G40876A being the white-flowered variant, see below), can be obtained from the

genebank at CIAT (http://www.ciat.cgiar.org/urg). The wild Lima bean, P. lunatus L., used in the comparisons

is a collection from Santa Ana, El Salvador, Central America (DGD-3236; G27609 in CIAT genebank). As aptly

explained by Spencer and co-workers (Spencer et al. 2007, p. 58-59), because there are no genetic alterations

(first generation after original collecting, closed growing environment), these cuttings are given a single name

under the Botanical Code (viz. the Melbourne Code: McNeill et al. 2012) only.

RESULTS

Morphological observations

Plant a pluriannual herbaceous tall indeterminate vine 2-6 m long or more with 8-14 or more axillary in¬

florescences. Seedling 7-12 cm high, from epigeal germination (Fig. la). Hypocotyl 20-25 mm, purple (59A;

RHS 1966), terete, sparsely covered with minute uncinate hairs. Epicotyl 15 mm, green purple at nodes, terete,

glabrous but with few minute uncinate hairs. Eophylls petiolate opposite triangular acuminate cordate at base,

glabrous, central vein prominent; the blade 47-55x33-42 mm; the petiole terete canaliculate 28-33 mm long,

purplish (60A) with green pulvini; stipules entire or divided rectangular blunt at apex lxl mm. First true leaf

trifoliolate; terminal leaflet 53-65x32-37 mm, lateral leaflet 44-56x24-32 mm, petiole 36-43 mm, rachis

12-13 mm. Root herbaceous fibrous superficial for three months after germination then much ramified, base

turning thickened while ageing diam. 5 mm, up to 1 m long and more. Stems terete green with purplish dots

at nodes, glabrescent with short 1 mm sparsely distributed hairs and minute uncinate hairs mostly found at

nodes (Fig. lb); internodes 102-121 mm long 3-6 mm diam.; sprawling from the 3-4 lower nodes on main

stem; climbing from the upper nodes on main stem; after four months 3 m and more long; first internodes pro-

Debouck, Validation of Phaseolus lignosus from the Bermuda Islands

109

Fig. 1 . Drawing of Phaseolus lignosus Britton; a: seedling; b: flowering stem; c: flower; d: pod; e: seed; f: rooting internode. Scale bar: 5 mm.

110

Journal of the Botanical Research Institute of Texas 9(1)

during numerous adventicious roots if in contact with soil (Fig. 2 top left); first internodes becoming woody

after one year. Stipules triangular lanceolate, spreading horizontally, green glabrous basihx multiveined 3x1

mm. Leaves trifoliolate (Fig. lb) alternate in distichous succession, leaflets ovate rounded at base lengthily

acuminate entire lustrous glabrous with minute trichomes, lamina green 141 A, main veins prominent beneath

turning light purplish with age, terminal leaflets symmetrical 76-90 mm long 53-59 mm wide, lateral leaflets

asymmetrical 67-83 mm long 47-53 mm wide. Petiole terete canaliculate 58-107 mm long 1.5-2 mm diam.,

green sometimes with fine linearly distributed reddish (59A) dots, sparsely covered with short white hairs and

minute uncinate hairs. Rachis terete canaliculate green 15-20 mm long 1-1.2 mm diam.. Proximal and distal

pulvini green 3.5-5 mm long. Stipels narrowly triangular linear 1.5-2 mm long. Inflorescence a pseudora¬

ceme (Fig. lb; Fig. 2 lower part), ascending, appearing as a compact pyramidal cluster of floral buds intermixed

with bracts (9 mm high when peduncle is 20 mm long) (Fig. 2 top right). Peduncle at flowering 45-53 mm

long diam. 1.5-2 mm, curved or straight at base, terete, green with basal part towards first insertion turning

purplish 61A. Rachis erect or still curved, terete, green, 75-175 mm, with 8-17 floral insertions. Peduncle and

rachis glabrous with minute uncinate hairs (more densely covered in upper part of rachis). Primary bracts

lanceolate, 2-3 mm long 2 mm wide, green, 3(5)-nerved margin ciliate. Pedicelar bracts narrowly triangular

1-nerved, 1 mm long, green. Pedicel terete, purple, 7-12 mm long 1 mm diam., with minute uncinate hairs

(Fig. 3c). Anthesis sequence in secondary raceme was as follows: flower 2 one more day after flower 1, flower 3

four days after, flower 4 six days, and flower 5 (if not falling) eight days. Flower papilionaceous asymmetrical

(Fig. lc, Fig. 3a, Fig. 5 left). Bracteoles lanceolate, 2x1.5 mm up to 60% calyx length, greenish, 3(5)-nerved,

margin ciliate, central vein prominent, persistent up to one day after anthesis. Calyx cupped, finely reddish

speckled, sparsely covered with short white hairs and trichomes, tube 3 mm long, two upper lobes almost

united obtuse, three lower lobes subequal triangular 1 mm long 1-1.5 mm wide. Standard broadly half circular

markedly reflexed backwards at anthesis (Fig. 3c), external face glabrous cream with a greenish cast (145B)

and fine purple veining, internal face glabrous deep purple (72C) with two light magenta rose dots and fine

magenta veining, hooded over the keel, 3 mm long to point of flexure then 6 mm to upper margin, 11-12 mm

wide, upper margin obtuse-angled, tip slightly pointed, lateral edges unconspicuous, auricles 2-4 mm long,

claw 2 mm long whitish with erect narrow parallel callosities, flexure internally thickened. In white-flowered

variant (Fig. 5 right) external face pale green (142D). Wings glabrous, magenta (70B-67B) on anthesis fading

light purple (75A) the next day, parallel and forward spreading almost horizontally (Fig. 3a), the blade rounded

cupped 12 mm long 10 mm wide, left wing 1 mm larger in both directions, outward margin often irregularly

undulate, spur right-angled greenish adhering to the keel, claws white linear 3-3.5 mm long. In the white-

flowered variant wings pure white on anthesis fading cream yellow (13A) the next day. Keel tubular asym¬

metrical, greenish white at base pink purple in median part, with two close coils counterclockwise (facing the

keel) with a diam. of 3 mm, the terminal coil with a diam. of 2 mm, tube 5 mm long 2 mm wide to first curva¬

ture then 4 mm to the coils, terminal coil golden green. Stamens diadelphous (9+1), staminal tube white deli¬

cately veined 11 mm long ending into 9 free stamens 5 mm long, vexillar stamen white in free space of staminal

tube and on top of dorsal suture of ovary, claw terete 2 mm long with fusiform slightly unequal knob diam.

1.5 mm then 14 mm long, anthers yellow oval dithecal dorsihxed. Ovary green finely puberulent 4 mm long 1

mm wide, laterally compressed, 4-6 ovules, inserted on a yellowish veined basal disk 2 mm long. Style 15 mm

long terete coiled with 1.5 close coil at tip, covered by a short white pubescence then glabrous, ending with a

loose brush of white hairs in the last 2 mm below the terminal stigma; stigma internal narrow triangular 0.5

mm long. Pod laterally compressed (Fig. Id, Fig. 4a), usually 1-4 seeded (if one, only distal or near distal seed

develops); young pod green puberulent curved 18 mm long 3.8 mm wide; when mature 52-55 mm long 9-10

mm wide 7 mm thick, both sutures finely marked, green drying tan, puberulent with very small white hairs;

curvature smooth when all embryos develop, scimitar-like when 1-2 embryos develop; base of pod shortly

stipitate 3 mm long; pod beak 3.5 mm long delicately curved; spiral twisting of pod valves at dehiscence. Seed

oblong rounded slightly rhomboid, 8 mm long 5.5 mm wide 4.5 mm thick (Fig. le, Fig. 4c), 8.2 g/ 100 seed,

tan or grey finely black speckled ending into a black circle around the hilum, hilum 2 mm white sunken.

Debouck, Validation of Phaseolus lignosus from the Bermuda Islands

111

Fig. 2. Photographs of different parts of P . lignosus . Top left: rooting lower internode. Top right: young developing raceme. Lower part: blooming stems.

During growing-out the plant showed a dual growth habit with 2-4 m long guides sprouting from lower

nodes of main stem and spreading on the ground, and main stem and other guides from upper nodes continu¬

ing climbing upwards. The stems spreading on ground have a significant capacity to quickly produce adventi¬

tious roots (Fig. If, Fig. 2 top left). One plant in Popayan produced white flowers (see Fig. 5 for comparison);

white flowers were also noted at collection site, as indicated on the label of C Watlington CW3 at K (Debouck

2014), evidencing a good sampling by MSB for seed distribution. The original description by Britton reported

white flowers too; unless otherwise indicated the description above refers to the magenta-flowered plant. It is

112

Journal of the Botanical Research Institute of Texas 9(1)

Fig. 3. Close-up of flowers, top: front view (a,b), and below: lateral view (c,d). Left side (a,c): P. lignosus (C Watlington 3; G40876 ; epitype). Right side

(b,d): R lunatus, wild form [DGD-3236; G27609).

important to note that the white-flowered variant has the same black speckled seed just as described above for

the magenta-flowered variant. The material seems to be susceptible to spider mites and nematodes (Meloido-

gyne). Pulvini are active in sunshine spots with the blades of leaflets turning parallel to solar radiation by mid¬

day (Fig. 2 lower part).

Diagnosis and recognition

As indicated in the Discussion below the most relevant taxon with which P. lignosus is to be related to and

distinguished from is the wild form of P. lunatus present in Central America and the Caribbean.

Herba mox perennis cum caulibus inferioribus prostratis dum superioribus volubilibus. P. lunati L. affinis sed foliolis ovatis floribus majori-

bus vexilliique superficie externa glabra, leguminibus majoribus angustioribus. Habitat solum in sylvis propemodo destructis insularum

maximarum Bermudae archipelagi rarus.

Debouck, Validation of Phaseolus lignosus from the Bermuda Islands

Fig. 4. Close-up of pods (top: a,b) and seeds (below: c,d; scale bar = 1 mm each). Left side (a,c): P. lignosus (C Watlington 3; G40876 ; epitype). Right

side (b,d): P lunatus, wild form {DGD-3236; G27609).

114

Journal of the Botanical Research Institute of Texas 9(1)

Fig. 5. Close-up front photographs of flowers of P. lignosus at anthesis. Left: normal wild type. Right: white flowered variant. Both plants from original

collection CW3, increased and conserved as accessions G40876 and G40876A, respectively.

Flower twice as large as the one of wild Lima bean (‘Mesoamericari, see Discussion), with glabrous standard

and magenta wings (Fig. 3a,c) in contrast to the outer upper pubescent face of standard and lilac wings in the

latter (Fig. 3b,d), are some distinctive characteristics of P. lignosus. Leaflets in wild Lima bean are usually lan¬

ceolate while ovate in the Bermuda bean (lower half of lamina constantly more rounded in the latter as com¬

pared to wild P. lunatus). The longer and narrower pod (because it usually has one more seed; Fig. 4a) of P. lig¬

nosus as compared to wild P. lunatus (41 mm long, 12 mm wide, 5.5 mm thick; Fig. 4b) is worth mentioning, as

are the more rounded seeds and absence of seed ridges radiating from the hilum in the former taxon (Fig. 4c).

In CIAT experimental station at Popayan, the time elapsed between anthesis and physiological maturity of pod

is forty days or one additional week as compared to wild Lima bean planted in the same conditions.

Additional Specimens

Five specimens (although Brown 718 and Brown 1179 might be from the same population visited at different

dates) were seen by the author in 75 out of 82 Herbaria keeping Paniculati specimens and visited since 1977

(see list in Acknowledgments below; data as on the labels):

BERMUDA: Joyce Dork [or Dock?] Cave, 22 May-2 Jun 1909, Stewardson Brown 718 (US); thicket [indication of vegetation?], Joyce’s Dock,

29 Nov-14 Dec 1912, Stewardson Brown 1179, NLBritton, FredJSeaver, collectors (US); Hamilton Parish, Idwal Hughes Nature Reserve, along

trail east of west sink, Walsingham, 22 Feb 2007, MA Hamilton 610 (K); Castle Harbour Golf Course [32° 20' 30"N, 64° 43' 30" W; our esti¬

mate] , 5 May 1964, E Manuel 637 (A); Bermuda [sic!], Warwick, Cluster Cottage Gardens, 25 Apr 2008, C Watlington 3 with E Morbey (K; two

sheets).

The specimen Brown 680 mentioned by Delgado-Salinas et al. (1999, 2006) and Lopez-Lopez et al. (2013), and

claimed as deposited at PHILA has not been seen because the search for PHILA in Index Herbariorum (2013)

did not yield any result, nor did the Specimen Search in Tropicos (2013).

Designation of an epitype

As afore-mentioned in the original description (Britton 1918) there is no reference to any plant type specimen,

although Britton could have used any of his collections with Stewardson Brown, whom he acknowledged as

one of his collaborator (op. cit., p. xi). This situation should thus be corrected, either by using existing exsic-

cata (see above, Additional Specimens) or by forming new ones, hopefully numerous and complete. Along the

Debouck, Validation of Phaseolus lignosus from the Bermuda Islands

115

former possibility, the material of S Brown 1179 registered in the United States National Herbarium (US sheet

3413573), Smithsonian Institution in Washington, is not complete enough for a type: no flowers, and pods

harvested excessively late (borne on a single raceme pulled out of the main stem!). One of the collectors of this

specimen S Brown 1179 was Britton himself, and he (because it is the same handwriting as in the frontispiece of

the biography by E.D. Merril 1938) annotated the voucher prior to its transfer to US (Debouck 2014). Inciden¬

tally, this helped us to validate all fruit characteristics (dimensions, curvature, sutures, beak, base). The speci¬

men MA Hamilton 610 at K lacks fruits as does one duplicate of the C Watlington 3 collection (K000650153). The

specimens E Manuel 637 (in A) and C Watlington 3 (K000650154) lack mature pods and seeds, although they

are fairly complete but unique. Finally, the specimen Stewardson Brown 718 (US1341071) with flowers and

young pods, although identified by James A. Lackey as P. lunatus (Debouck 2014), could help but again is

unique. Incidentally, this last identification stresses further our point: the need for good plant specimens to

refer to! Following Art. 8, Art. 9.8 and 9.13 and Recommendation 9C.1 of the Melbourne Code (McNeill et al.

2012), using cuttings of the plant grown in Popayan (from seed of the original collection of C Watlington

(CW3), an epitype is designated and fourteen isoepitypes were made and deposited into the Herbaria indicated

below. The specimen in Kew consists of two sheets (for space reasons only): marked 1/2’ with leaves, racemes,

flowers, pods and seeds, and ‘2/2’ with root and seedling. Designating an epitype is the least that can be done

(because it is to classify plants not drawings!)—not a neotype as we thought at the beginning of this research

and distribution of vouchers—as there is a lectotype (an illustration; Art. 9.2).

Phaseolus lignosus Britton. Type: BERMUDA: plant seen in rocky woodlands between Castle Harbor and Harrington Sound, fig.

205, without citation of specimen, collector, or date (lectotype, here designated: Britton, Fl. Bermuda, Fig. 205 on page 183,1918).

(epitype, here designated: K with bar code K000556376 and K000556377; isoepitypes: BRIT, CAS, COT, F, FHO, FI, G, GH, MEXU,

MICH, MO, NCU, NY, P). Voucher specimens of the white flowered variant—so indicated—have been deposited into the Herbaria

BRIT, K, MICH, MO and P.

Geographic distribution, ecology and conservation status

Specimens studied so far indicate a distribution restricted to the major island of Bermuda: Long Island (above

data, and Debouck 2014), with no sure records for the other two large pieces of land: St. George’s Island and

Somerset Island. The record for Hungry Bay, Paget Parish on Long Island mentioned by Britton (1918), that we

have not seen, might be doubtful because it is anterior to the original publication. Christine Phillips-Watling-

ton (1996) reports the Bermuda bean from three areas of Long Island in the Walsingham Cave district only,

while Copeland and co-workers (2014) report it from four places. This vinelike legume is part of the under¬

story of the almost gone Upland Woodland with Juniperus bermudiana L., Sabal bermudana L.H. Bailey Celtis

laevigata Willd., Cassine laneana (A.H. Moore) J.W. Ingram and Zanthoxylumflavum Vahl. (Small 1913; Britton

1918; Phillips-Watlington 1996; Pettit et al. 2012). As the wild P. lunatus from the Peninsula of Yucatan

(Standley 1930; personal observations in February 1979), the Bermuda bean can thrive in karstic derived soils

too (Phillips-Watlington 1996). P. lignosus is listed as an endangered very rare endemic under the Protected

Species Act 2003 of the Government of Bermuda (Pettit et al. 2012), and as ‘critically endangered’ by the IUCN

(Copeland et al. 2014. It is this situation that incited us to use grown cuttings to make a large series of reference

specimens available to many Herbaria, rather than taking more reproductive parts from original plants.

DISCUSSION

The above facts elicit the following points for discussion. The first point—stressing further the purpose of this

note—relates to the question whether or not the plant described above is P. lignosus. We only have the original

description and a drawing to refer to (Britton 1918, p. 183, Fig. 205), and no reference specimens. The plant

described above is a wild bean plant because of pod dehiscence and seed characteristics. It is thus not any form

of P. coccineus L. (syn. P. multiflorus Willd.), P. lunatus L.and P. vulgaris L., all three grown as garden crops in the

archipelago of Bermuda, according to Britton (1918). The vegetative parts (stems, leaves, glabrousity) of C Wat¬

lington 3 grown in our conditions match with the original description (and with specimens MA Hamilton 610,

E Manuel 637 , and Stewardson Brown 1179). The minute uncinate hairs however are not mentioned in the origi-

116

Journal of the Botanical Research Institute of Texas 9(1)

nal paper; these will be noted later on (Doutt 1932), and their taxonomic value for Phaseolus sensu stricto iden¬

tification will be realized much later (Baudet 1973; Marechal et al. 1978). The raceme is a curved pseudoraceme

as in Fig. 205, and dimensions for peduncle and rachis fit within the range given by Britton. The calyx and wing

of the flower match with the original illustration even though most floral parts are wanting. The pods match

with the description (except the part in bracket “[P. semierectus of Reade]”) and with those of Stewardson Brown

1179 (seen by Britton), namely the beak, the base and fine sutures. The seed displays a sunken hilum as on the

illustration in the work of Britton (1918). From this we can conclude that the collection C Watlington 3 and its

afore-mentioned fourteen isoepitype specimens are a Phaseolus sensu stricto species (Marechal et al. 1978;

Lackey 1983), and belong to P. lignosus.

Nathaniel Lord Britton (op. cit., p. 183) indicated a (morphological) relationship of his new taxon with P.

polystachyus (L.) B.S.P., not with P. coccineus L. (syn. P. multiflorus Willd.), P. lunatus L. nor P. vulgaris L.. This

statement would mean that he has noted the differences between all these taxa. And this brings to the second

point about the nature of this taxon and its relationships with other Phaseolus species. We have to examine first

whether or not it belongs to the Paniculati section (Freytag & Debouck 2002). The size of the pluriannual

plant, its climbing growth habit from a woody lower stem, the leaf morphology, the pseudoraceme and size and

shape of the primary bracts and bracteoles, all point to this section. The capacity of the meristems of secondary

racemes to produce flowers beyond flower no. 3 is noteworthy. In addition, two phylogenetic analyses (Delga-

do-Salinas et al. 1999, 2006) have shown that P. lignosus belongs to the T. lunatus group’, distinct from the T.

polystachios group’, while both groups have been included into the Paniculati (Freytag & Debouck 2002). So, P.

lignosus would be an addition to the Lima bean gene pool, the richest in species in the genus (Porch et al 2013).

The third question might be: is P. lignosus a species distinct enough from P. lunatus and P. polystachyus as

to deserve the specific rank? Bearing in mind that the closest mainland is North Carolina of USA (Watson et al.

1970), one could also raise a similar question about the Bermuda bean towards P. sinuatus Nuttall ex Torrey &

Gray and P. smilacifolius Pollard, that are distributed in the southeastern USA (Isely 1990, and Small 1903, re¬

spectively). Differently lobed leaflets (small rounded lobes in P. sinuatus and large 1 cm long rounded lobes in

P. smilacifolius) of the latter two taxa clearly exclude an identity with the Bermuda bean. P. lignosus could not be

a variant of P. polystachyus either, because its leaflets are not rounded broadly ovate and delicately acuminate,

and its pod beak is longer and pointed. Growth of axes of secondary racemes is often seen in P. polystachyus,

giving a definite panicle aspect to the inflorescence, while not seen in P. lignosus. Another major difference is

ecologicaF P. polystachyus could experience frost in a significant part of its US continental range (from New

York to Missouri: Debouck 2014; Freytag & Debouck 2002), while P. lignosus thrives in almost frost-free habi¬

tat (Watson et al. 1970; Phillips-Watlington 1996).

The difference with wild P. lunatus is a relevant question—and purposely we alluded to it a couple of

times already, because the small-seeded wild form (or ‘Mesoamericari, not the Andean one: Serrano-Serrano et

al. 2010) has towards the northern part of its range a wide distribution from Islas Revillagigedo in the Pacific

Ocean, through Sonora and Tamaulipas in Mexico, and then the Greater and Lesser Antilles (Debouck 2014;

Freytag & Debouck 2002). The current floristic works for the Bahamas (Britton & Millspaugh 1920; Correll &

Correll 1982; Nickrent et al 1988) do not clearly conclude about the presence of P. lunatus as a wild plant or as

“spontaneous after cultivation.” Would the Bermuda bean be an expansion of wild or weedy P. lunatus into the

Bermuda archipelago? Although much alike a wild Lima bean during early vegetative growth, the plant differs

from the former by its flower, pod and seed characteristics (Figs. 3, 4). The flower of P. lignosus is bigger as

compared to the one of wild Lima bean; the wings are cupped and almost getting in contact instead of bending

forward spatulate and clasping on each other in the latter (Fig. 3b,d). The magenta color (the ‘quinacridone’

magenta used by botanical illustrators: Stevens 2007) is unknown in wild P. lunatus. The pod is not moon

crescent shaped as in wild Lima bean, and the pod beak is wider and longer (Fig. 4a) as compared to that of

wild Lima bean (Fig. 4b). The time from anthesis to pod maturity is longer in P. lignosus as compared to that of

wild P. lunatus. On the other hand, Pettit and co-workers (2012) indicated that P. lignosus would not thrive in

salinity affected areas, while wild P. lunatus seems to be salinity tolerant at least in some populations (Bayuelo-

Jimenez et al. 2002). For these reasons, the Bermuda bean can be considered as a different and valid species.

Debouck, Validation of Phaseolus lignosus from the Bermuda Islands

117

A fourth point is related to the distribution of this taxon, and it seems that definitely P. lignosus is endemic

to the Bermuda archipelago. All Phaseolus sensu stricto species as wild plants are distributed only in the New

World (Lackey 1983; Freytag & Debouck 2002), and thus these are the species the Bermuda bean should be

compared to in terms of geographic distribution. It has not been reported from the Carolinas (Radford et al.

1968; Debouck 2014). The floras of the Bahamas (Correl & Correl 1982; Nickrent et al. 1988) do not report

it either. P. lunatus exists as wild in the Greater Antilles (Cuba: Beyra & Reyes-Artiles 2004; Haiti and the

Dominican Republic: Debouck 2014; Puerto Rico: Perkins 1907) and Lesser Antilles (Perkins 1907), and

seems to be the only true wild Phaseolus species there, although P. dumosus Macfady. thrives as a weed in

mountainous Jamaica (Macfadyen 1837; Freytag & Debouck 2002). There is a report about the presence of P.

polystachyus in Puerto Rico (Liogier 1988, p. 194, “in thickets at lower elevations, Rincon”), but no specimens

are mentioned, and we have not seen so far any specimen of P. polystachyus reported from the Caribbean Is¬

lands (Debouck 2014). Delgado-Salinas (1985) did not either report any specimen of P. polystachyus from the

Caribbean. Finally, in addition to being an endemic plant, the low number of populations reported here (and

mentioned by Copeland and co-workers, 2014) is stricking: it might reflect a poor sampling of the Bermuda

archipelago (made of 180 islands and islets, although few covered originally by the Upland Woodland), or a

naturally or human induced low number of populations. This situation deserves additional held work and

frequent updating.

In conclusion, Nathaniel Lord Britton was right in identifying P. lignosus as a new bean species endemic to

the Bora of Bermuda. While some of its populations are nowadays at risk in original habitats (29 individuals,

and with population trend being evaluated as decreasing: Copeland et al. 2014), a positive note is an increased

capacity to produce ex situ seeds of this rare taxon through a cooperative effort of CIAT-MSB (at closing of

preparation of this note, 6,768 seeds have been harvested at CIAT, and harvests continue). Thanks to seed

availability some questions for future research such as: how, when and from where did P. lignosus land in the

Bermuda archipelago?, or which useful traits could it bring to bean breeding?, can now be addressed.

ACKNOWLEDGMENTS

This work is dedicated to Orlando Toro Chica who passed away during this research after contributing much

to the successful introduction and seed increase of Phaseolus lignosus in CIAT experimental stations.

The support of the Consultative Group on International Agricultural Research and the Global Crop Di¬

versity Trust for growing out the plant materials in the different CIAT stations is acknowledged. Special thanks

are due to Ruth Eastwood and Beverly Maynard at the Millennium Seed Bank for all the help provided for

procuring a few original seeds of the studied taxon. The Bermuda Department of Conservation Services gave

permission for the seeds to be used, and is fully acknowledged. Gratitude is warmly expressed to the following

persons at different steps of this research: Cesar Franco, Alina Freire-Fierro, Isabel Fernanda Gomez, Josehna

Martinez, Mariano Mejia, John R. Porter, Carlos Saa, Raul Urmendez and Eliana Urquijo. The author warmly

acknowledges the permission granted by the Curators of the following Herbaria (where specimens of the sec¬

tion Paniculati are kept): A, AGUAT, AHUC, ANSM, ARIZ, BA, BAA, BAB, BACP, BAFC, BM, BR, BRIT, CAS,

CHAPA, CICY, COL, CORD, CPUN, CR, CS, CUZ, DAV, DES, DS, DUKE, EBUM, ENCB, F, FI, G, GH, HAO,

HNMN, HUT, IA, IBUG, IEB, INB, ISC, ITIC, K, L, LAGU, LIL, LL, LPB, M, MA, MEXU, MICH, MIN, MO,

MOL, MSC, NA, NCU, NEBC, NY, O, P, PH, POM, PRG, QCA, RSA, SI, TCD, TEX, UC, US, USJ, USM, UVAL,

WIS, to study their collections in situ since 1977. Last but not least, the author warmly thanks Kanchi Gandhi

(Harvard University), Alfonso Delgado Salinas (Universidad Nacional Autonoma de Mexico), and two anony¬

mous Reviewers for very helpful suggestions on the nomenclatural issues.

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120

Journal of the Botanical Research Institute of Texas 9(1)

BOOK NOTICE

Jesse Vernon Trail. 2015. Quiver Trees, Phantom Orchids and Rock Splitters: The Remarkable Survival

Strategies of Plants. (ISBN-13: 978-1-77041-208-8, pbk). ECW Press, 2120 Queen Street East, Suite 200,

Toronto, Ontario M4E 1E2, CANADA. (Orders: ecwpress.com). $24.95 US/CAN, 300 pp., full color

throughout, 130 photos andillus., 5" x 7".

From the publisher: Plants around the world are often exposed to bitter cold, relentless winds, intense heat,

drought, fire, pollution, and many other adverse growing conditions. Yet they are still able to survive and often

even thrive.

Quiver Trees, Phantom Orchids and Rock Splitters: The Remarkable Survival Strategies of Plants is a handy

pocket guide to everything you could ever want to know about the tenacity and resilience of plants. Designed

with a beautiful full-color layout and peppered with over 130 eye-catching photographs and illustrations, [the

book] showcases the many weird and wonderful ways that plants adapt to survive and spread their progeny.

The book offers absorbing information that will inspire a new respect for nature’s innovation and resilience.

Jesse Vernon Trail is an author, instructor, and curriculum developer in environment, ecology, sustainability

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Making, Canadian Gardening, Plant and Garden, Fine Gardening, Gardenwise, Harrowsmith Country Fife, Alive,

and Outdoor Canada. Jesse lives in Vernon, B.C.

Contents:

Author’s Note

Pollination and Attracting Pollinators

Seeds and Their Dispersal

Vegetative Reproduction

Growing Challenges

The Vital Importance of Water and Air

The Arctic Example

Alpine Adaptations

Lessons from the Desert

A Mediterranean Perspective

An Enduring Champion

A Firm Footing

Chemistry

Fire and Plants

Rain Forests

Tropicals

Aquatic and Marginally Aquatic Plants

Defensive Strategies

Offensive Strategies

Other Fascinating Survival Strategies

Unusual Survival Strategies

Afterword

Selected Sources

Recommended Books and Websites

Additional Photo Credits

Index of Plants

J.Bot. Res. Inst. Texas 9(1): 120.2015

DASYLARYNX ANOMALUS GEN. ET SP. NOV., A TUBULAR

MONOCOTYLEDON-LIKE FLOWER IN MID-TERTIARY DOMINICAN AMBER

George O. Poinar, Jr.

Kenton L. Chambers

Department of Integrative Biology

Oregon State University

Corvallis, Oregon 97331, U.S.A.

poinarg@science.oregonstate.edu

Department of Botany and Plant Pathology

Oregon State University

Corvallis, Oregon 97331, U.S.A.

ABSTRACT

Dasylarynx anomalus is described as a new genus and species, based on two flowers preserved in Mid-Tertiary amber from the Dominican

Republic. The fossil is hypothesized as being adapted for pollination by hummingbirds or long-tongued insects, with a slender perianth tube

formed of 3 fused petals. The flower is bisexual, the stamens being positioned at the base of the corolla tube together with a single stipitate

pistil. The tube is densely hirsute within, bearing trichomes from base to apex. The trichomes, although appearing stiff, were evidently flex¬

ible enough to allow the bills or mouthparts of visiting pollinators to reach the source of food. Only part of the androecium is visible. The

fertile stamens are club-shaped, with introrse dehiscence. They are accompanied by several lattice-like staminodes that are perforated by

numerous openings of various sizes. The combination of unique androecial characteristics, together with a tubular, non-lobed corolla,

makes it difficult to place the fossil in any modern family of monocotyledons.

RESUMEN

Dasylarynx anomalus se describe como un Nuevo genero y especie, basados en dos flores conservadas en ambar del Terciario medio de la

Republica Dominicana. El fosil se supone estar adaptado para la polinizacion por colibries o insectos de lengua larga, con un tubo de peri-

anto delgado formado por tres petalos unidos. La flor es bisexual, los estambres estan posicionados en la base del tubo de la corola junto con

un pistilo estipitado simple. El tubo es densamente hirsuto en el interior, llevando tricomas de la base al apice. Los tricomas, aunque parecen

rigidos, eran suficientemente flexibles para permitir a los picos o partes bucales de los polinizadores visitantes alcanzar el recurso alimenti-

cio. Solo es visible una parte del androceo. Los estambres tienen forma de porra, con dehiscencia introrsa. Estan acompanados de varios

estaminodios en forma de rejilla que estan perforados por numerosas aberturas de varios tamanos. La combinacion caracteristicas unicas

del androceo, junto con una corola tubular, no lobulada, hace dificil situar el fosil en cualquier familia moderna de monocotiledoneas.

INTRODUCTION

The Mid-Tertiary Caribbean forests giving rise to Dominican amber were described by Poinar and Poinar

(1999), based on the known insect fauna and a limited number of available plant fossils. Since then, numerous

additional examples of the flora have been discovered, in the form of amber-embedded flowers of various for¬

est-dwelling taxa. The plant species producing the resin that trapped these fossils was Hymenaea protera

(Poinar 1991), a leguminous tree related to present-day Hymenaea courharil L. Dasylarynx anomalus, the sub¬

ject of this paper, is the 20th angiosperm species to be described from Dominican amber. Besides H. protera,

these include 2 species of Fabaceae (Dilcher et al. 1992, Poinar & Chambers 2015), 2 of Poaceae (Poinar &

Judziewicz 2005; Poinar & Columbus 2012), 3 of Arecaceae (Poinar 2002a, 2002b), 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

Myristicaceae (Poinar & Steeves 2013), 1 of Rhamnaceae (Chambers & Poinar 2014a), 1 of Ticodendraceae

(Chambers & Poinar 2014b), and 1 possibly of Moraceae (Poinar et al. 2008b).

Having 2 specimens of the Dasylarynx flower, it was possible to grind and polish the corolla tube of 1 to

expose the anthoecium and gynoecium, as well as to reveal the dense inner covering of trichomes. In the pro¬

cess, however, the stamens were disturbed, and some pollen was scattered that partially obscures the manner

of anther dehiscence and other structural features. The stipe, ovary, and style of the pistil, partly visible among

the trichomes, are indicated in Fig. 2. Nine stamens are assumed to be present, but only 5, including 3 stami-

J. Bot. Res. Inst. Texas 9(1): 121 -128.2015

122

Journal of the Botanical Research Institute of Texas 9(1)

nodes, are shown in the polished section (Fig. 3). The total number of fertile stamens is unknown, since only

part of the androecium is in view.

The assignment of Dasylarynx to the Monocotyledonae is consistent with its monocolpate pollen (Fig. 5)

and a perianth formed by 2 whorls of 3 parts, assuming our interpretation of the corolla is correct. Besides the

non-lobed corolla tube, other morphological features of the fossil are notable. Taking the place of 3 of the 5 vis¬

ible stamens are peculiar staminodes, which are thin and lattice-like in construction, variable in shape, and

perforated by different sized openings (Fig. 3). The manner of introrse anther dehiscence is not clear but is

perhaps by 4 pores (Fig. 3). The pistil is not completely exposed for analysis, but because of the single style and

lack of obvious lobing of the ovary, it most likely is composed of a single carpel. Our hypothesis concerning

details of the species’ pollination syndrome is discussed in a later section of the paper.

MATERIALS AND METHODS

Dominican amber has been assigned to the Mid-Tertiary period, based on planktonic fossils occupying the

marine strata in which it is deposited. Mines are located in the Central Mountain Ranges of Hispaniola, from

which the amber is extracted by workers who dig lengthy tunnels to reach the sought-after deposits. Samples

of amber, brought to the surface, are then sold at the mine entrance or are put on the market for buyers of this

semi-precious gemstone. Two differing dates have been proposed for the amber. Iturralde-Vinent & MacPhee

(1996) estimated an age of 20-15 Ma, based on foraminifera, whereas 45-30 Ma was the date given by Cepek

(in Schlee 1999), derived from studies of coccoliths. The strata containing the amber are turbiditic sandstones

of the Upper Eocene to Lower Miocene Mamey Group (Draper et al. 1994).

The flowers are contained in blocks of amber measuring 25 mm x 20 mm x 5 mm (holotype) and 25 mm

x 12 mm x 4 mm (paratype). Examination and photographs were made with a Nikon stereoscopic microscope

SMA-10-R at 80x and a Nikon Optiphot microscope at 800x.

DESCRIPTION

Dasylarynx Poinar & K.L. Chambers, gen. nov. Type Species: Dasylarynx anomalus Poinar & K.L. Chambers, sp. nov. (Figs.

1-5)

Diagnosis .—Plants hermaphroditic, flowers stout-pedicelled, perianth biseriate, sepals 3, petaloid, spreading,

glabrous, equal or unequal, ovate-lanceolate, with numerous parallel veins from the base, apex obtuse (Fig. 1),

petals probably 3, completely fused into an elongate, externally glabrous tube, lobes 0, interior of tube hirsute

nearly throughout (Fig. 2), stamens basal in the tube, 9 (?), probably in 3 whorls of 3, including an unknown

number of sterile staminodes, 2 fertile stamens visible, club-shaped, with broad, irregularly perforated fila¬

ments (Fig. 3F), the anther portion terminal, not much enlarged, introrsely dehiscent, perhaps with 4 pores

arranged as 2 pairs, 1 pair above the other (Fig. 3A), staminodes variable in shape, thin, perforated by small or

large openings (Fig. 3B, D, E), pistil 1, ovary elliptic-fusiform, glabrous, stipitate, style 1, terminal, elongate,

partly obscured by trichomes (Fig. 2), stigma not visible, pollen ellipsoidal, monocolpate (Figs. 4, 5), exine

minutely echinate, the spinules in +/- parallel rows, each spinule arising from a small basal cushion (Fig. 4).

Dasylarynx anomalus Poinar & K.L. Chambers, sp. nov. Type: HISPANIOLA. Dominican Republic: amber mine in the north¬

ern mountain ranges (Cordillera Septentrional), 1986, unknown amber miner s.n. (holotype: catalog number Sd-9-60A; paratype:

catalog number Sd-9-60B, both deposited in the Poinar amber collection maintained at Oregon State University, Corvallis, Oregon

97331, U.S.A.

Description .—Total length 15.6 mm (holotype), largest sepal of outer whorl 9.8 mm long, 4.1 mm wide, other 2

sepals shorter (lengths 7 mm and 6 mm), perhaps due to shrinkage during preservation in the resin (Fig. 1),

tube formed by petals of inner whorl 15 mm long, the opening 1.3 mm wide, diameter 2.5 mm below apex, 1.8

mm in center, 1.9 mm at base (Fig. 1), ca. 9-veined near base, hirsute trichomes within tube up to 1.0 mm long

(Fig. 2), fertile stamens 1.0 mm long, 0.33 mm wide (Fig. 3), staminodes 0.6-1.1 mm long, up to 0.22 mm wide

(Fig. 3), style slender, elongate, ovary 0.73 mm long, 0.36 mm wide, stipe 0.75 mm long (Fig. 2), dimensions of

pollen 86 pm x 65 pm (Fig. 4).

Poinar and Chambers, Dasylarynx anomalus, a monocotyledon-like flower in Mid-Tertiary amber

123

Fig. 1. Dasylarynx anomalus. Holotype flower, lateral view. A. Corolla tube. B. Sepal. Scale bar = 5.0 mm.

Etymology .—Genus name from Greek “dasyshairy, shaggy, and “larynx,” throat, gullet. Species name

from Greek “anomalos,” unusual, abnormal.

DISCUSSION

The floral morphology of Dasylarynx is so unusual that its placement in a modern family is problematic. We

believe that an androecium of 3 whorls can be identified, consisting of a mixture of fertile stamens and variably

perforated staminodes. Anther dehiscence by 4 pores in 2 superposed pairs is hypothesized (Fig. 3). Three

124

Journal of the Botanical Research Institute of Texas 9(1)

Fig. 2. Dasylarynxanomalus. Paratype flower, inside of perianth tube. A. Hirsute trichomes. B. Style. C. Ovary. D. Stipe. Scale bar = 1.0 mm.

Poinar and Chambers, Dasylarynx anomalus, a monocotyledon-like flower in Mid-Tertiary amber

125

Fig. 3. Dasylarynx anomalus. Paratype flower, enlarged view. A. Fertile stamen B. Staminode of outer androecial whorl. C. Fertile stamen of 2nd whorl.

D. Staminode of outer whorl. E. Staminode of 3rd whorl, interior to fertile stamen. F. Perforation in filament of fertile stamen. Scale bar = 0.2 mm.

visible staminodes are constructed of thin, lattice-like strands of differing sizes and are perforated by a variable

number of large and small openings. The function of such organs can hardly be guessed at. No nectar glands

are present, although a floral disc interior to the stamens may be present. Pollen grains were observed to be

monocolpate (Fig. 5), with the exine echinate and the spicules borne on basal cushions (Fig. 4). This type of

spicule was observed and illustrated independently in pollen of the Lauraceae (Raj & van der Werff 1988). It is

a question whether the 3 sepals were equal or of different sizes. Because the tissue of the left and right sepals

(Fig. 1) appears to have shrunken somewhat during preservation, we favor the interpretation that they were of

equal size in life.

126

Journal of the Botanical Research Institute of Texas 9(1)

In considering the reproduc¬

tive syndrome of this flower, we

propose that, in spite of its relatively

small size, it may have been attrac¬

tive to hummingbirds, especially if a

nectar disc was present at the base

of the pistil. In addition, it could

have been visited by long-tongued

insects, including species of large¬

bodied bees, flies, or lepidoptera,

particularly sphingid moths. Some

of the insect groups mentioned by

Renner and Fed (1993, Table 1)

as pollinators of tubular topical

flowers are known to have been

present in the Dominican Tertiary

forest. These include riodinid and

nymphalid butterflies (De Vries &

Poinar 1997; Hammond & Poinar

1998), tabanid horseflies (Lane et al.

1988), and euglossine bees (Poinar

1998). Pollinators attracted by the

petaloid sepals would have inserted

their bills or mouth parts into the

perianth tube in search of nectar or

pollen. Attached pollen would then

be transferred to the stigma of the

same or a different flower. Of further note is the hirsute inner epidermis of

the floral tube. How this might have played a role in pollination is not clear,

but at least it should not have prevented extraction of pollen by the pollina¬

tor’s bill or tongue. Possibly the corolla hairs functioned to protect against

the theft of nectar by small ants.

Bird and insect pollination is common in modern tropical forest

woody plants (Bawa et al. 1985). Stiles (1978, Appendix 1) provides a use¬

ful list of plant genera that he observed being pollinated by hermit and

non-hermit hummingbirds in Finca La Selva, Costa Rica. Among these is

Calathea, of the Marantaceae. Similarities between the fossil and this fam¬

ily are the presence of a corolla tube, which in Marantaceae is a 3-lobed

floral tube including androecium (Andersson 1998), and of 2-4 stamino-

dia in the androecium. These organs are petaloid and hooded or indurate

in Marantaceae (Andersson op. cit.), unlike those of Dasylarynx. Impor¬

tant differences are that the ovary of Marantaceae is inferior, there is 1 petaloid stamen with a single theca, and

the pollen is nonaperturate (Erdtman 1952).

It was noted in the Introduction that this newly recognized genus is one of some 20 taxa that have re¬

cently been described from Dominican amber. The environment in which these species grew was considered

by Hammel and coauthors (Hammel & Zamora 1990; Hammel & Burger 1991) to have been part of a once

widespread Laurasian warm climate forest flora of the Tertiary Period. This mild, humid, aseasonal climate

Fig. 4. Dasylarynx anomalus. Pollen grain. Arrow shows row of spicules, each with a small basal

cushion. Scale bar = 65 pm.

Fig. 5. Dasylarynx anomalus. Pollen grain, with

arrow on colpus. Scale bar = 46 pm.

Poinar and Chambers, Dasylarynx anomalus, a monocotyledon-like flower in Mid-Tertiary amber

127

was prevalent across the Northern Hemisphere in Paleogene times (65-25 Ma). Dominican amber, though of

uncertain age, can probably be dated to the late Oligocene or early Miocene.

ACKNOWLEDGMENTS

We are indebted to Jens Rohwer andjohn Dransheld for their helpful comments.

REFERENCES

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(Chrysobalanaceae). J. Bot. Res. Inst.Texas 4:217-218.

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minican amber. J. Bot. Res. Inst.Texas. 8:563-568.

Chambers, K.L., G.O. Poinar, Jr., & A.E. Brown. 2011a. A fossil flower of Persea (Lauraceae) in Tertiary Dominican amber. J.

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Res. Inst.Texas 5:463-468.

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Soc. London B. 264:1137-1140.

Dilcher, D.L., P.S. Herendeen, & F. Huber. 1992. Fossil Acacia flowers with attached anther glands from Dominican amber.

In: P.S. Herendeen and D.L. Dilcher, eds. Advances in legume systematics. Part 4. The fossil record. Royal Botanic

Gardens, Kew, UK.

Draper, G., P. Mann, & J.F. Lewis. 1994. Hispaniola. In: S. Donovan andT.A. Jackson, eds. Caribbean geology: An introduc¬

tion. The University of the West Indies Publishers'Association, Kingston, Jamaica. Pp. 129-150.

Erdtman, G. 1966. Pollen morphology and plant taxonomy. Angiosperms. Hafner Publishing Co., New York. 553 pp.

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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.

Hammond, P.C. & G.O. Poinar, Jr. 1998. A larval brush-footed butterfly (Lepidoptera: Nymphalidae) in Dominican amber,

with a summary of fossil Nymphalidae. Ent. Scand. 29:275-279.

Iturralde-Vinent, M.A. & R.D.E. MacPhee. 1966. Age and paleographic origin of Dominican amber. Science 273:1850-1852.

Lane, R.G., G.O. Poinar, Jr., & G.B. Fairchild. 1988. A fossil horsefly (Diptera: Tabanidae) in Dominican amber. Florida Ento¬

mologist 71:593-596.

Poinar, G.O., Jr. 1991. Hymenaea protera sp. n. (Leguminosae: Caesalpinioideae) from Dominican amber has African af¬

finities. Experientia 47:1075-1082.

Poinar, G.O., Jr. 1998. Paleoeuglossa melissiflora gen. n., sp. n. (Euglossinae: Apidae), fossil orchid bees in Dominican

amber. J. Kansas Entomol. Soc. 71:29-34.

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.

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128

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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 Domini¬

can 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.

Poinar, G.O., Jr. & K. L. Chambers. 2015. Prioria dominicana sp. nov. (Fabaceae: Caesalpinioideae), a fossil flower in Mid-

Tertiary Dominican amber. J. Bot. Res. Inst.Texas 9(1 ):129-134.

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Gard. 75:130-167.

Renner, S.R. & J.P. Feil. 1993. Pollinators of tropical dioecious angiosperms. Amer. J. Bot. 80:1100-1107.

Schlee, D. 1999. Das Bernstein-Kabinett. Stuttgarter Beitr. Naturk. Ser. C, 28.

Stiles, F.G. 1978. Temporal organization of flowering among the hummingbird foodplants of a tropical wet forest.

Biotropica 10:194-210.

PRIORIA DOMINICANA SP. NOV. (FABACEAE: CAESALPINIOIDEAE),

A FOSSIL FLOWER IN MID-TERTIARY DOMINICAN AMBER

George 0. Poinar, Jr.

Kenton L. Chambers

Department of Integrative Biology

Oregon State University

Corvallis, Oregon 97331, US.A.

poinarg@science.oregonstate.edu

Department of Botany and Plant Pathology

Oregon State University

Corvallis, Oregon 97331, US.A.

ABSTRACT

Prioria dominicana is a newly described species of Fabaceae from the Mid-Tertiary tropical forests of the Caribbean island of Hispaniola.

These forests were earlier described by Poinar and Poinar (1999) based on the flora and fauna preserved in amber from the fossilized resin

of Hymenaea protera (Poinar 1991). The single living neotropical species of Prioria, P. copaifera, is found in Jamaica, Nicaragua, Panama, and

Colombia. It differs from P. dominicana in the greater size of its bracts, absence of a pedicel, conspicuous anther connectives, and the villous

pubescence of its filaments and ovary. In comparing the fossil with Old World species of Prioria, P. dominicana shares important similarities

with the West African species P. mannii and P. joveri. It provides evidence, therefore, for a long-standing phylogenetic connection between

the Old and New World elements of the genus dating to the Middle or Early Tertiary. Tacking fruits or leaves, further comparison of P. do¬

minicana with its relatives is not possible at present.

RESUMEN

Prioria dominicana es una nueva especie descrita de Fabaceae de los bosques tropicales del Terciario medio de la isla caribena la Hispan¬

iola. Estos bosques fueron descritos previamente por Poinar and Poinar (1999) basados en la flora y fauna conservada en ambar de la resina

fosilizada de Hymenaea protera (Poinar 1991). Ta unica especie neotropical viviente de Prioria, P. copaifera, se encuentra en Jamaica, Nicara¬

gua, Panama, y Colombia. Se diferencia de P. dominicana en el mayor tamano de sus bracteas, ausencia de pedicelo, conectivos de las anteras

manifiestos, y la pubescencia villosa de sus filamentos y ovario. Comparando el fosil con especies del Viejo Mundo de Prioria, P. dominicana

comparte importantes similitudes con las especies del oeste africano P. mannii y P. joveri. Esto es la prueba, por tanto, de una conexion filo-

genetica de larga duracion entre elementos del Viejo y del Nuevo Mundo del genero desde el Terciario medio o temprano. Con la falta de

frutos u hojas, comparaciones posteriores de P. dominicana con sus parientes no es posible actualmente.

INTRODUCTION

The Mid-Tertiary moist, tropical forests of the Caribbean region have come to be better known floristically

from studies of fossil amber flowers from the Dominican Republic. Genera from the following families of an-

giosperms have been described from the amber: Fabaceae (Poinar 1991, Dilcher et al. 1992), Arecaceae (Poinar

2002a, 2002b), Poaceae (Poinar & Judziewicz 2005, Poinar & Columbus 2012), Chrysobalanaceae (Poinar et

al. 2008a, corrected by Chambers & Poinar 2010), Lauraceae (Chambers et al. 2011a, 2012a), Meliaceae

(Chambers et al. 2011b; Chambers & Poinar 2012a, 2012b), Burseraceae (Chambers & Poinar 2013), Myristi-

caceae (Poinar & Steeves 2013), Rhamnaceae (Chambers & Poinar 2014a), Ticodendraceae (Chambers &

Poinar 2014b), a possible Moraceae (Poinar et al. 2008b), and an unknown monocotyledon (Poinar & Cham¬

bers 2015).

The type species of Prioria, P. copaifera Griseb., is a resin-producing tree up to 40 m high found in low- to

medium-elevation forests of Nicaragua, Panama, Colombia, and Jamaica (Woodson & Schery 1951). The ge¬

nus was long assumed to be endemic to the New World. However, Breteler (1999) proposed an alternative cir¬

cumscription that expands Prioria to include the Old World Kingiodendron Harms 1897, Oxystigma Harms

1897, Pterygopodium Harms 1913, and Gossweilerodendron Harms 1925. Prioria is classibed in the Caesalpini-

oid tribe Detarieae (Lewis et al. 2005), the largest tribe in the subfamily, with around 84 genera (Fougere-

Danezan et al. 2010). In some previous studies, Prioria copaifera and its Old World relatives form a clade sepa¬

rate from, but sister to, Detarieae sensu lato (Bruneau et al. 2001; Wojciechowski et al. 2004; Lavin et al. 2005).

J. Bot. Res. Inst. Texas 9(1): 129 -134.2015

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Journal of the Botanical Research Institute of Texas 9(1)

The later work of Bruneau et al. (2008) places the Prioria clade as sister to the Deterieae sensu stricto, thus

supporting the resin-producing Deterieae clade as monophyletic. The dating of this clade will be discussed in

a later section.

MATERIALS AND METHODS

Dating of Dominican amber is still controversial, with the youngest proposed age of 20-15 Ma based on fora-

minifera (Iturralde-Vinent & MacPhee 1996) and the oldest of 45-30 Ma based on coccoliths (Cepek in Schlee

1990). Most of the amber is secondarily deposited in turbiditic sandstones of the Upper Eocene to Lower Mio¬

cene Mamey Group (Draper et al. 1994). Dilcher et al. (1992) suggested that “...the amber clasts, from all

physical characteristics, were already matured amber at the time of re-deposition into marine basins. There¬

fore, the age of the amber is greater than Miocene and quite likely is as early as late Eocene.” The issue is further

complicated by the discovery of Early Oligocene amber in Puerto Rico and Maastrichtian-Paleocene amber in

Jamaica (Iturralde-Vinent 2001), indicating that the age of Caribbean amber is still unresolved.

The flower is contained in a block of amber measuring 10 mm x 8 mm x 5 mm. Examination and photo¬

graphs were made with a Nikon stereoscopic microscope SMA-10-R at 80x and a Nikon Optiphot microscope

at 800x. Helicon Focus Pro X64 was used to stack photos for better clarity and depth of held.

DESCRIPTION

Prioria dominicana Poinar & K.L. Chambers, sp. nov. (Figs. 1-3). Type: HISPANIOLA. Dominican Republic: amber mine

in the northern mountain range (Cordillera Septentrional), 2013, unknown amber miner s.n. (holotype: catalogue number Sd-9-23,

deposited in the Poinar amber collection maintained at Oregon State University, Corvallis, Oregon 97331, U.S.A.).

Flower perfect, radially symmetrical, with well-developed pedicel, width at anthesis 3.8 mm (Fig. 1), basally

bracteate, bracts 2, separate, elliptic, length 0.6-0.7 mm, width 0.35-0.4 mm, glabrous (Figs. 2,3), disc appar¬

ently 0, calyx 5-merous, sepals free, boat-shaped, broadly ovate, 2.0-2.4 mm long, 1.5-1.9 mm wide, acute,

lightly ciliate near tip, otherwise glabrous, midnerve indistinct (Figs. 1,3), petals 0, stamens 10, free, filaments

linear, 3.0 mm long, attached around base of ovary, glabrous throughout (Fig. 2), anthers dorsihxed, ellipsoid,

0.7 mm long, 4-loculed, with narrow connective (Figs. 1, 2), pistil 1, stipe short, glabrous (Fig. 2), ovary ovoid,

1.5 mm long, 1.1 mm wide, glabrous (Fig. 2), style linear, 1.7 mm long, stigmatic tip scarcely enlarged (Figs. 1,

2), floral pedicel 1.1 mm, densely puberulent (Figs. 1,3).

Etymology. —Species epithet from the Dominican Republic.

DISCUSSION

Prioria dominicana differs from P. copaifera in the presence of a puberulent pedicel together with smaller, free,

subtending bracts, which in the modern species are orbicular, up to 1.5 mm long, and sheath the disciferous

part of the calyx (Woodson & Schery 1951, pers. observ.). The sepals of P. dominicana are apparently free and

imbricate at the base (Fig. 1) and are not at all scarious margined, as is described for P. copaifera. The fossil also

differs in the absence of villous pubescence on the filaments and ovary (Fig. 4) and in the less conspicuous

connective of the anthers.

The taxonomy of Prioria accepted in this paper is that of Breteler (1999, see above). In its similarities to

both the Old and New World species of the genus, P. dominicana apparently forms a link that helps to unify the

genus. Its closest living African relatives, deduced from Breteler’s key, descriptions, and illustrations (op. cit.),

are P. mannii (Baillon) Breteler and P. joveri (Aubrev.) Breteler, both native to Cameroun, Guinea, and Gabon.

The Indomalasian species of the genus can be excluded from consideration due to their unisexual flowers and

crateriform stigmas. Prioria mannii has dorsally pubescent sepals, with stamen filaments lightly puberulent at

the base, and ovary, together with style base, densely pubescent. Its pedicel is well-developed and puberulent,

and its anther connectives are inconspicuous. Prioria joveri has dorsally papillate sepals (per Breteler, Fig.

2[2]), puberulent stamen filaments, densely pubescent ovary and style base, and well-developed anther con¬

nectives. Its pedicel is densely puberulent. Prioria dominicana is similar to both African taxa, therefore, in its

Poinar and Chambers, Prioria dominica (Fabaceae), a fossil flower in Mid-Tertiary amber

131

• -*»

Fig. 1. Prioria dominicana. A. anther. B. style. C. sepal. D. pedicel.

well-developed, pubescent pedicel (Figs. ID, 3C) and small bracts, but differs in its glabrous sepals, pistil, and

style base.

Discovery a flower in Dominican amber that closely resembles certain African species was previously

noted with the fossil Hymcnacaprotcra, which has flowers most similar to H. verrucosa Gaertn., an extant spe¬

cies restricted to East Africa and adjacent islands (Poinar 1991). In view of the ambiguous age determinations

for Dominican amber (see Materials and Methods, above), the timing of its included fossils cannot be clearly

assigned at present. One can only speculate on how Prioria and Hymenaea were able to disperse from Africa to

the Caribbean or vice versa. Africa and South America were already separated by the developing South Atlantic

Ocean by 94 Ma (http://www.scotese.com/earth.htm); hence, a considerable water gap existed by the time

these genera evolved in the Early Tertiary. Various methods of fruit and seed dispersal might be hypothesized

that could have allowed elements of the New World tropical forests to overcome this barrier.

In studies pertinent to Prioria dominicana, the molecular phylogeny of the Fabaceae has been examined

using the chloroplast trnL intron (Bruneau et al. 2001), rbcL (Kajita et al. 2001), matK genes (Wojciechowski et

al. 2004), trnL intron and trnL-F spacer (Fougere-Danezan et al. 2007), and matK-trnK plus trnL genes (Bru¬

neau et al. 2008). The correlation between molecular and morphological data was examined by Fougere-

Danezan et al. (2010). Relationships among the first branching lineages of the legumes are not well supported

molecularly, with Cercideae, Detarieae, and the genus Duparquctia alternatively resolved as sister group to all

of the legumes (Bruneau et al. 2008). In this paper, the authors used 18 well-documented fossils as calibration

points, fixing the stem node of the legumes at 65 Ma. Among the fossils included was Prioria dominicana,

132

Journal of the Botanical Research Institute of Texas 9(1)

Fig. 2. Prioria dominicana. A. style. B. filament. C. ovary. D. stipe. E. bract.

which prior to its formal description had been illustrated in a book on the Tertiary amber forests of Hispaniola

by Poinar and Poinar (1999). As in other studies, Bruneau et al. (op. cit.) could readily define a monophyletic

Prioria clade, which by their results is sister to the resin-producing Detarieae sensu stricto. Within this clade,

Prioria, in the broad sense considered here, is assigned a divergence time of ca. 38 Ma (Bruneau et al. op. cit.,

Poinar and Chambers, Prioria dominica (Fabaceae), a fossil flower in Mid-Tertiary amber

133

Fig. 3. Prioria dominicana. A. bract. B. sepal. C. pedicel. A dark air bubble lies behind the bract.

Fig. 5). As mentioned above, present uncertainty as to the age of Dominican amber fossils implies that more

accurate dating to come, if this proves possible, will influence future studies of the age of major Detarieae

clades in the Caesalpinioideae.

ACKNOWLEDGMENTS

Useful suggestions by an anonymous reviewer are gratefully acknowledged. We thank the curator of the Mis¬

souri Botanical Garden herbarium for a loan of specimens important to this study.

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Chambers, K.L. & G.O. Poinar, Jr. 2012b. Additional fossils in Dominican amber give evidence of anther abortion in Mid-

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Chambers, K.L., G.O. Poinar, Jr., & A.S. Chanderbali. 2012. Treptostemon (Lauraceae), a new genus of fossil flower from Mid-

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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. 2014a. Distigouania irregularis (Rhamnaceae) gen. et sp. nov. in Mid-Tertiary amber from

the Dominican Republic. J. Bot. Res.Texas 8:551-557.

Chambers, K.L. & G.O. Poinar, Jr. 2014b. Ticodendron palaios sp. nov. (Ticodendraceae), a Mid-Tertiary fossil flower in

Dominican amber. J. Bot. Res. Inst.Texas 8:559-564.

Dilcher, D.L., P.S. Herendeen, & F. Huber. 1992. Fossil Acacia flowers with attached anther glands from Dominican Republic

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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. & E.J. Judziewicz. 2005. Pharusprimuncinatus (Poaceae: Pharoideae: Phareae) from Dominican amber. Sida

21:2095-2103.

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can 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¬

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Histor. Biol. 1-6.

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Missouri Bot. Gard. 38:27-29.

ADDENDUM: A GASTEROID FUNGUS, PALAEOGASTER MICROMORPHA

GEN. & SP. NOV. (BOLETALES) IN CRETACEOUS MYANMAR AMBER

George Poinar, Jr.

Donis da Silva Alfredo

Department of Zoology

Oregon State University

Corvallis, Oregon 97331, U.S.A.

poinarg@science.oregonstate.edu

Graduate Program in Systematics and Evolution

Department of Botany and Zoology Center of Biosciences

Universidade Federal do Rio Grande do Norte

BRAZIL

luri Goulart Baseia

Graduate Program in Systematics and Evolution

Department of Botany and Zoology

Center for Biosciences

Universidade Federal do Rio Grande do Norte

BRAZIL

In the original publication of “A gasteroid fungus, Palaeogaster micromorpha gen. & sp. nov. (Boletales) in Cre¬

taceous Myanmar amber” that appeared inj. Bot. Res. Inst. Texas 8(1): 139-143 (2014), the MycoBank number

of the genus was incorrect. The correct MycoBank numbers are:

Palaeogaster Poinar, Alfredo, & Baseia, gen. nov. (Figs. 1-8), MycoBank no.: MB 801126.

Palaeogaster micromorpha Poinar, Alfredo, & Baseia, sp. nov. (Figs. 1-8), MycoBank no.: MB 801127.

J. Bot. Res. Inst. Texas 9(1): 135.2015

136

Journal of the Botanical Research Institute of Texas 9(1)

BOOK NOTICE

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and eastern Ontario. The guide is formatted so that each page is arranged by the surface each lichen grows on

in the held, its shape or growth form, then by its color. Full color photographs and black and white drawings

for each species also aid in identification.

Troy McMullin, Ph.D., is a lichenologist and forest ecologist at the University of Guelph. His ecological inter¬

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consulting work on lichens in the public and private sector. He has worked closely with provincial and federal

governments in Nova Scotia, Prince Edward Island, and several other Canadian provinces. He completed his

doctoral work at the University of Guelph, where he studied the effects of forestry on lichen diversity in north¬

ern Ontario.

Frances Anderson, a Research Associate at the Nova Scotia Museum, has published a provisional checklist

of larger lichens for Nova Scotia and coauthored research reports into the risk status of two lichen species for

the Committee on the Status of Endangered Wildlife in Canada, among other scientific publications. She has

done lichen surveys for forestry companies and conservation groups, produced several lichen brochures re¬

lated to particular geographic areas or hiking trails, written a book chapter on lichens, and led lichen walks for

naturalists.

J.Bot. Res. Inst. Texas 9(1): 136.2015

FURROWED BLISTER PODS STRANDED ON NORTHERN ATLANTIC OCEAN

COASTS REPRESENT AN UNDESCRIBED SACOGLOTTIS (HUMIRIACEAE)

ENDOCARP MOST SIMILAR TO THE FOSSIL SACOGLOTTIS COSTATA

Raymond van der Ham

Natural is Biodiversity Center

P.O.Box 9517

2300 RA Leiden, NETHERLANDS

raymond.vanderham@naturalis.nl

Izumi Hanno

P.O. Box247

Post Office Shingaraja

81100 Bali, INDONESIA

Paul Mikkelsen

3705 Eleven Mile Road

Fort Pierce, Florida 34945, U.S.A.

Tinde van Andel

Natural is Biodiversity Center

P.O. Box 9517

2300 RA Leiden, NETHERLANDS

Theo Lambrechts

Leopold lei 96

2220 Heist-op-den-Berg

BELGIUM

Jenifer Mina

202 Joy Haven Drive

Sebastian, Florida 32958, U.S.A.

Linda Butcher

2616N20 1/2 Street

Harlingen, Texas 78550, U.S.A.

Kim Lincicome

883 Chickadee Drive

Port Orange, Florida 32127, U.S.A.

Bob Patterson

15 Fractious Street

Hamilton Parish, CR 04

BERMUDA, U.K.

Ed Perry

1770 Mason Terrace

Melbourne, Florida 32935, U.S.A.

Mike Romance

354 96th Street

Marathon, Florida 33050, U.S.A.

ABSTRACT

Ten distinctly furrowed endocarps stranded on coasts along the northern Atlantic Ocean (including Gulf of Mexico and North Sea) have so

far eluded identification. The endocarps resemble the blister pod, a common “sea-bean” originating from the fruit of Sacoglottis amazonica

(Humiriaceae), but they are larger and show deep furrows. The furrowed blister pods should be classified in the genus Sacoglottis sensu lato

(Sacoglottis and Schistostemon together). They represent an undescribed endocarp most similar to the fossil Sacoglottis costata Reid from

the Tertiary of Colombia, South America. The recent specimens are best denoted as Sacoglottis cf. costata. Presumably, they do not belong

to one of a few Sacoglottis species known only from flowering collections. Therefore, it is expected that an undescribed species, producing

costata-like endocarps, occurs somewhere in northeastern South America.

RESUMEN

Diez endocarpos claramente surcados de las costas del norte del oceano Atlantico (incluyendo el Golfo de Mexico y Mar del Norte)

habian eludido su identificacion. Eos endocarpos semejan a una vaina con ampollas, una “sea-bean” comun originada del fruto de

Sacoglottis amazonica (Humiriaceae), pero son mas grandes y muestran surcos profundos. Tas vainas sulcadas suelen clasificarse

en el genero Sacoglottis sensu lato ( Sacoglottis y Schistostemon juntos). Representan un endocarpo no descrito muy similar al del

fosil Sacoglottis costata Reid del Terciario de Colombia, Sudamerica. Eos recientes especimenes estan mejor denominados como

Sacoglottis cf. costata. Presumiblemente, no pertenecen a una o unas pocas especies de Sacoglottis conocidas solo de colecciones de

flores. Sin embargo, se espera que una especie no descrita, que produce endocarpos semejantes a los de S. costata, se de en alguna

parte de noreste de Sudamerica.

INTRODUCTION

Sea-beans are “seeds and fruits that are carried to the ocean, often by freshwater streams and rivers, then

drift with the ocean currents and (hopefully!) wash ashore” (wwwseabean.com 2014). About 2007, a

conspicuously furrowed sea-bean (Fig. 1A-D) was found among debris from the North Sea coast in the

Netherlands, and brought to the attention of the first author at the end of 2012. This sea-bean closely

matched a drawing in the World Guide to Tropical Drift Seeds and Fruits (Gunn & Dennis 1976, Fig. 71B;

J. Bot. Res. Inst. Texas 9(1): 137 -147.2015

138

Journal of the Botanical Research Institute of Texas 9(1)

Fig. 1. Endocarps of Sacoglottis cf. costata (Humiriaceae). First column (A, E, I, M): view of apex; second column (B, F, J, N): view of base; third column

(C, G, K, 0): view of median furrow on valve (C, G, 0) or seed remains (K); fourth column: view of septum (D, H) or section (L, P). Corresponding valves in

different photographs of the same specimen are indicated with an asterisk (*). a = apex, b = base, bs = basal scar, d = one of five depressions around

apex, I = locule, mf = median furrow on valve, r = one of two ridges on valve, c = cavity, s = septum, v = valve. Scale bar = 10 mm. A-D, Specimen #5,

from Yerseke, Netherlands. E-H, Specimen #4, from Palm Beach County, Florida, U.S.A. l-L, Specimen #7, from Bermuda, U.K. (photographed by Bob &

Helle Patterson). This specimen is cross-sectioned, with one valve removed from upper half of endocarp (Fig. 11) showing seed remains (Fig. 1K). Fig. 1L

represents the lower half, showing two adjacent fertile locules and many small cavities (compare with Fig. 1P, 3D). M-P, Specimen #9, from South Padre

Island, Texas, U.S.A. Fig. 1P shows a micro-CT cross-section showing two opposite fertile locules and many small cavities (compare with Fig. 1L, 3D).

van der Ham et al., Furrowed blister pods (Sacoglottis sp.) on coasts along the northern Atlantic Ocean

139

Fig. 2. Endocarps of Sacoglottis cf. costata (Humiriaceae). Scale bar = 10 mm. A-C, Specimen #2, painted by Izumi Hanno, from unknown location.

Corresponding valves are indicated with an asterisk (*). A, view of apex. B, view of base. C, view of septum (compare with Fig. 1D, 1H, 2D). D, Speci¬

men #1, from SE coast of Florida, U.S.A. (from Gunn & Dennis 1976; drawing by Pamela J. Paradine), view of septum (compare with Fig. 1D, 1H, 2C).

Fig. 2D in present paper). This drawing was included in a plate illustrating disseminules of mahogany

(Swietenia mahagoni (L.) Jacq., Meliaceae) from the southeastern coast of Florida, U.S.A. However, after

checking mahogany fruits it was learned that neither the drawing nor the sea-bean from the Netherlands

could possibly be this species or any other species of the family Meliaceae.

After some research, the sea-bean was identified as an endocarp 1 from a fruit of a Humiriaceae species. A

well-known sea-bean from this almost completely Neotropical family is the blister pod: Sacoglottis amazonica

Mart. (Fig. 3A-D). This endocarp, with crowded large globular cavities (blisters), is “one of the most common

tropical drift seeds found on Florida east coast beaches” (Perry & Dennis 2003:79). It is found on beaches all

around the North Atlantic Ocean and has occasionally also been collected in northwestern Europe (Devon, Ire¬

land, Scotland; Guppy 1917; Nelson 2000). However, the unknown, distinctly furrowed endocarp does not re¬

semble the endocarp of S. amazonica: the latter is small, its cavities are large, and its surface is bullate with only

shallow and irregular furrows (Fig. 3A-D). The endocarp of Vantanea guianensis Aubl. (Humiriaceae), mentioned

by Gunn and Dennis (1976) as being “not found too far beyond their northern South America homeland,” does

not compare either. The unknown endocarp was provisionally assigned to the genus Duckesia Cuatrec. (Hu¬

miriaceae), on the basis of the large size, the (supposed) presence of foramina (holes) in the septa near the apex

and medium-sized cavities visible at the surface of the valves, and comparison with material published by Her¬

rera et al. (2010). Fabiany Herrera (pers. comm. 2012) agreed, but advised to section the endocarp in order to

confirm this affinity. Consulting “sea-beaner” Ed Perry prompted the reply “This is Sacoglottis gabonensis! ... re¬

lated to blister pod ..., but much rarer in drift.” Following the accompanying link to Sacoglottis gabonensis on the

website www.seabean.com provided photographs of two furrowed, undoubtedly conspecihc endocarps from

Florida, one from Palm Beach County (Fig. 1E-H), the other from Lignumvitae Key. A photograph of an addi¬

tional specimen (Fig. 1I-L) from Bermuda was sent with Ed’s reply. The African Sacoglottis gabonensis (Baill.)

Urb. is the only non-American species of Humiriaceae. Endocarps of this locally common species are often found

floating in rivers and on sea shores in the western part of tropical Africa (Cuatrecasas 1961). S. gabonensis might

be closely related to S. amazonica (see also Renner 2004) and produces similar though smaller and more globular

endocarps, which differ from the unknown endocarp. A request by Ed Perry via Facebook (“Sea Beans” group)

generated donations of two more specimens of the unknown endocarp, one from Volusia County, Florida, and

one (Fig. 1M-P) found stranded on South Padre Island, Texas, U.S.A. In 2011 coauthor Izumi Hanno painted an

unidentified sea-bean in the Gunn/Dennis collection. This specimen (Fig. 2A-C) clearly represents the unknown

1 An endocarp is the woody ("stony") inner layer of the fruit wall of a drupe, together with the mesocarp (middle layer) and exocarp (outer layer, also called epicarp) making

up the entire fruit wall.

140

Journal of the Botanical Research Institute of Texas 9(1)

> Vpr « 1

. *

/ % i"

. a -

' X 0f'f ' CO

Fig. 3. Endocarp of Sacoglottis amazonica Mart. (Humiriaceae) from unknown location (#133, carpological collection of the Utrecht herbarium in Natu-

ralis Biodiversity Center, Leiden, Netherlands). Corresponding valves are indicated with an asterisk (*). I = locule, c = cavities, s = septum, v = valve.

Scale bar = 10 mm. A, view of apex. B, view of base. C, view of valve (compare with Fig. 1C, 1G, 10). D, cross-section showing one fertile locule and

relatively few large cavities (compare with Fig. 1L, 1P).

endocarp but deviates in size, shape, and details from the one depicted in Gunn and Dennis (1976, Fig. 71B; Fig.

2D in present paper). Therefore, it is assumed that two specimens might be or have been present in the Gunn/

Dennis collection 2 . In 2014 a specimen was reported byjenifer Mina. She found it (without data) on the ‘sharing

table’ during the Nineteenth Annual International Sea-Bean Symposium and Beachcombers’ Festival in Octo¬

ber 2014 at Cocoa Beach, Florida. According to Ed Perry, it might have been collected in Texas by George

Wolf (Pasadena, Texas), who donated so much to put on the sharing table. Correspondence with the collectors/

owners of the furrowed endocarps led to enthusiastic response and cooperation, so that (data of) ten speci¬

mens are now available. The purpose of the present paper is to describe them and to assess their status within

the Humiriaceae.

MATERIAL AND METHODS

Ten specimens of the furrowed endocarp are known (Table 1, Fig. 4), of which one (#3) is probably lost

(collection Cathie Katz, died 2001; seen by Ed Perry; no further data available). The present location of

the other specimens is cited in Table 1 in the column “collection.”

The CT image (Fig. IP) was made with a Bruker SkyScanll72 high-resolution microCT (Naturalis

Biodiversity Center, Leiden, Netherlands).

RESULTS

The furrowed endocarps available for further study (#4-10; Table 1, Fig. 1A-P) are 54-72 mm long,

40-62 mm wide (avg. 63.4 and 50.0 mm) and have an ovoid, ellipsoid to almost globular, or obovoid

shape (length/width ratio 1.02-1.42, avg. 1.28). Some specimens (especially #4 and #5) are slightly asym¬

metrical due to the larger size of the valves covering fertile locules. The base contains a depressed, circular

to pentagonal innervation scar, while the apex shows a bulging, 5-rayed star (= top of axis with radiating

septa) with a 2-6 mm deep, radially oblong depression in each septum. At the surface of the endocarp,

the five septa extend towards the base as 0.3 to ca. 1 mm wide irregular ridges, each at the bottom of an

oblong, up to 5 mm deep, and 15 mm wide furrow (Fig. 1C, D). Above the base, the septa diverge into

the area around the basal innervation scar. The valves range from oblong to elliptical in longitudinal view,

slightly shorter than the endocarp itself and up to 30 mm wide at the equator. Valves covering fertile loc¬

ules are slightly larger than those on the infertile ones. Only the basal part of each valve is connected with

2 The Gunn/Dennis collection contains the material that was the basis for the records in Gunn and Dennis (1976). At present, the collection is in the possession of coauthor

Izumi Hanno. It is in storage (Jacksonville, FL, USA) and temporarily inaccessible.

van der Ham et al., Furrowed blister pods (Sacoglottis sp.) on coasts along the northern Atlantic Ocean

141

Table 1. List of specimens, collection data and features of the furrowed endocarp ( Sacoglottis cf. costata).

locality

year

source/collector

collection

LxW(mm)

shape

Fig.

SE coast of Florida, U.S.A.

< 1976

Gunn & Dennis (1976)

Hanno

ca. 60 x 40

ovoid

Gunn/Dennis collection

Hanno

c. 64 x 49

ovoid

2A-C

<2000

Cathie Katz/Ed Perry

lost?

Palm Beach County, FL, U.S.A.

ca. 2003

Paul Mikkelsen

67x54

ellipsoid

1E-H

Yerseke*, Netherlands

ca. 2007

Theo Lambrechts

62x47

ovoid

1A-D

Lignumvitae Key, FL, U.S.A.

2009

Mike Romance

Romance

63x62

± globular

Bermuda, U.K.

"some y. ago"

Bob Patterson

Patterson

64x45

ellipsoid

1 l-L

Volusia County, FL, U.S.A.

2013

Kim Lincicome

FLAS

62x46

ellipsoid

South Padre Island, TX, U.S.A.

2013

Linda Butcher

BRIT

54x45

o bo vo id

1M-P

10 Texas?, U.S.A.

<2014

George Wolf? /

Jenifer Mina

72x51

o bo vo id

Note. The specimens are listed more or less in order of collecting. Collection: BRIT = Botanical Research Institute ofTexas, Fort Worth, Texas,

U.S.A.; FLAS = University of Florida Herbarium, Florida Museum of Natural History, Gainesville, Florida, U.S.A.; L = Naturalis Biodiversity

Center, Leiden, Netherlands. Collections Hanno, Patterson and Romance: see author addresses. L = length, W = Width. * Specimen #5 was

found in Yerseke, Netherlands, among material dredged from the Middeldiep, along the nearby North Sea coast.

Fig. 4. Map showing localities of stranded specimens of Sacoglottis cf. costata (Humiriaceae). #1: SE coast of Florida, U.S.A.; #4: Palm Beach County,

Florida, U.S.A.; #5: Yerseke, Netherlands; #6: Lignumvitae Key, Florida, U.S.A.; #7: Bermuda, U.K.; #8: Volusia County, Florida, U.S.A.; #9: South Padre

Island, Texas, U.S.A. Localities of #2, #3, and #10 unknown. The localities are plotted on a map originally compiled to show cold and warm sea currents

and the extent of sea ice in the North Atlantic Ocean (United States Army 1943).

the axis of the endocarp (Fig. 1A-C). Each valve shows a median furrow up to 10 mm deep and 10 mm

wide at the equator. Thus a total of ten distinct longitudinal furrows mark the surface of the endocarp

from base to apex. Alternatively, the surface may be described as beset with ten wide conspicuous ribs

(two on each valve). Cross-sections of three endocarps are available: a physical section of specimen #7

(Fig. 1L) and micro-CT scans of specimens #5 and #9 (Fig. IP). All sections show a 5-radiating axial

142

Journal of the Botanical Research Institute of Texas 9(1)

structure with septa narrowing outwards. Specimens #5 and #9 contain two opposite fertile locules, while

specimen #7 has two adjacent fertile locules, which are more or less obtusely triangular in cross-section,

each containing the remains of an oblong seed. Septa and valves contain many globular cavities, up to 4

mm in diameter. There are no traces remaining of the exocarp and mesocarp.

Some specimens are worn, especially #5. Specimens #2 and #4-10 are sparsely to densely covered

with marine epibionts, including barnacles, small bivalves, bryozoans, calcareous tube worms, foramini-

fers, hydroids, and coralline red algae.

DISCUSSION

Comparison with the family Humiriaceae

The family Humiriaceae belongs to the order Malpighiales and comprises eight genera ( Duckesia Cuatrec.,

Endoplcura Cuatrec., Humiria Aubl., Humiriastrum (Urb.) Cuatrec., Hylocarpa Cuatrec., Sacoglottis Mart.,

Schistostemon (Urb.) Cuatrec., and Vantanea Aubl.) with about 50 species. The gynoecium in Humiria¬

ceae is syncarpous and usually 5-merous. The fruit is a pea-sized to mango-sized drupe with a single,

multi-seeded woody endocarp, which is very distinctive and one of the outstanding features of the family

(Cuatrecasas 1961; Herrera et al. 2010). The endocarp shows five septa and five valves (up to 8 and 10 in

Vantanea and Humiria), but most with only one or two locules, in each of which a single seed develops,

except in the genus Humiria that has two seeds per locule. The septa radiate from the axis of the endocarp.

Together, the axis and septa make up a rigid and resistant frame from which the valves of the fertile loc¬

ules separate during germination. The endocarps of Duckesia, Humiriastrum (partially), Sacoglottis, and

Schistostemon contain large (up to 7 mm), globular, ± empty cavities in their septa and valves. Intergeneric

differences occur in the relative width of the septa and valves at the surface, the length of the valves, and

the surface sculpture of the septa and valves (see below). The features mentioned above and combined

with the resistant nature of the endocarps, make the fruits of Humiriaceae readily identifiable objects in

dispersed state (e.g., fossils, sea-beans). The woody endocarps of species that occur on river banks and in

temporarily flooded forests (e.g., S. amazonica, S. gabonensis) have a good chance of being dispersed by

water because the cavities provide them with extra buoyancy.

In most Humiriaceae endocarps, valve width at the surface (measured at the equator) is equal to or

smaller than that of the septa (Cuatrecasas 1961; Herrera et al. 2010). In Humiriastrum, valve width is

occasionally larger than septum width, but in this genus the valves are only up to half as long as the endo¬

carp and are located near the apex of the endocarp. In Sacoglottis and Schistostemon the valves are nearly

as long as the endocarp and the septa are very thin or hardly or not visible at the surface, so that the valves

are very wide, sometimes even adjacent. This exact condition is present in the furrowed endocarps ana¬

lyzed in this study. Sacoglottis and Schistostemon are differentiated from each other by a flower character

(stamen number: 10 and 20, respectively), but their endocarps are indistinguishable (Cuatrecasas 1961).

In their morphological cladistic analysis, Herrera et al. (2010) found that Sacoglottis and Schistostemon

are sister groups, and they consider that the genera should be reduced to subgenera of the genus Sacoglot¬

tis. Isolated endocarps with very thin septa, including the furrowed endocarps described in the present

study, might then be identified as belonging to Sacoglottis s.l. (sensu lato, in broad sense: Sacoglottis and

Schistostemon together). Additional features supporting the assignment of the furrowed endocarps to

Sacoglottis s.l. are the presence of up to 4 mm large cavities throughout the endocarp and the absence

of apical foramina (holes) in the septa. Cavities occur also in Duckesia and Humiriastrum (partially), but

they are only a little larger than 0.7 mm at most (pers. comm. Fabiany Herrera 2012). Apical foramina are

also absent in Hylocarpa and Vantanea, but endocarps of these genera are very different from the furrowed

endocarps (e.g., valves equal or narrower than septa). Sometimes, apical foramina seem to be present in

the furrowed endocarps (e.g., in specimen #4; Fig. IE), but actually these holes represent cavities surfac¬

ing in the depressions near the apex.

van der Ham et al., Furrowed blister pods (Sacoglottis sp.) on coasts along the northern Atlantic Ocean

143

Comparison with extant Sacoglottis s.l.

Sacoglottis s.l. comprises 20 species (Cuatrecasas 1961, 1972, 1990; Sabatier 1987; Burger & Zamora Villalobos

1991, Zamora Villalobos 2007). All known endocarps of Sacoglottis s.l. species are not or are only inconspicu¬

ously furrowed. Although inclusion of the furrowed endocarps in Sacoglottis s.l. is well-supported, matching

them with any of the known endocarps of the species in this group is therefore impossible. Furthermore, the

furrowed endocarps are relatively large: 54-72 mm (avg. 63.4) long. Only Schistostemon retusum (Ducke) Cuatr.

has larger endocarps: 70-80 mm long (Lorenzi 2002) 3 . Sacoglottis amazonica (Fig. 3A-D) produces the next

largest endocarps: 20-60 mm long (Gunn & Dennis 1976), but normal-sized specimens are about 40 mm long

(Perry & Dennis 2003).

A few rare species are known from flowering material only: Sacoglottis maguirei Cuatrec. and Schist¬

ostemon auyantepuiense Cuatrec. Sacoglottis maguirei is a xeromorphic endemic from Venezuela (Amazo¬

nas, frequent at top of Cerro Yapacana, 1200 m alt.; Cuatrecasas 1961). The single known specimen of S.

maguirei has flowers and very young fruits (see also Cuatrecasas & Huber 1999, Fig. 540). Theoretically

(by lack of data), the furrowed endocarps might belong to S. maguirei. Cerro Yapacana is adjacent to

the Orinoco, so transport of endocarps by that river to the Caribbean Sea might occur. Mature fruits of

S. maguirei might still be collected in the Yapacana National Park. For the time being, some evidence is

provided by Sacoglottis sp. A (Zamora Villalobos 2007), an endemic from Costa Rica (Cordillera de Tala-

manca, 440 m alt.). This species is known from one fruiting collection, bearing globular to ellipsoid, ca.

35 x 29 mm large fruits. Sacoglottis sp. A is possibly related to S. maguirei (Zamora Villalobos 2007). If

related, the small size of its fruits would separate the furrowed endocarps from S. maguirei.

Schistostemon auyantepuiense is an endemic from Venezuela (Bolivar, near Guayaraca, 900-1100 m

alt.). It is a xeromorphic species very similar to Schistostemon fernandezii Cuatrec. from the same area

(according to Cuatrecasas & Huber 1999 even questionably different) and closely related to Schistoste¬

mon reticulatum (Cuatrecasas 1961). Fruits of S. reticulatum measure about 30 mm (pers. comm., Pedro

Jordano 2014). If S. auyantepuiense is indeed closely related to S. fernandezii, it is expected to have fruits

like those of S. fernandezii (see Cuatrecasas & Huber 1999, Fig. 544, and Herrera et al. 2010, Fig. 20), i.e.,

with a small, hardly or not furrowed endocarp. Therefore, conspecihty of the furrowed endocarps with S.

auyantepuiense and/or S. reticulatum is less probable.

Comparison with fossil Sacoglottis s.l.

Herrera et al. (2010, 2014) have revised the fossil record of the family Humiriaceae, especially that of its

resistant endocarps. They combined several fossil species into Sacoglottis tertiaria Berry emend. Herrera,

which is now known from Oligocene, Miocene, Pliocene, and Pleistocene deposits in Bolivia, Colombia,

Costa Rica, and Panama (see also Lott et al. 2011). Most of the included fossils very much resemble the

Sacoglottis s.l. endocarps as described above. Their surface is described as slightly bullate. However, the

single specimen on which the included Sacoglottis costata Reid from the Tertiary of Colombia was based

shows five radial depressions around the apex and 15 longitudinal ridges between base and apex. Five

ridges are thin and represent the five septa. The remaining ten ridges are arranged in pairs, each pair

flanking a deep median furrow on a valve (Reid 1933, plate 14, Figs. 1, la, 2, 2a; Fig. 5A-D in present

paper). Herrera et al. (2010) interpreted the furrows as the result of abrasion or erosion, but originally

the fossil was described as a “beautiful specimen.” The wear noticed by Reid refers only to the lack of

cellular surface details (Reid 1933). The differences between S. costata and the endocarps of extant spe¬

cies of Sacoglottis s.l. (especially S. amazonica), mentioned and adequately illustrated by Reid (1933),

correspond very well with the unusual features of the furrowed endocarp (compare Fig. 1A-P, Fig. 3A-D,

3 The material described and illustrated by Lorenzi (2002) originates from the coastal region of southern Bahia (Brazil). Material from Amazonas (Brazil), Colombia and

Venezuela described by Ducke (1938, Fig. 2d), Cuatrecasas (1961, Fig. 31 j) and Herrera et al. (2010, Fig. 2D) contained much smaller, though apparently mature endocarps

(globular, 26-34 mm diameter). The endocarps illustrated by Lorenzi (2002) do not show any characteristics of Sacoglottis s.l. or even Humiriaceae (valves, septa, cavities),

but this may be due to their freshly peeled state.

Fig. 5. Only known specimen of the fossil endocarp Sacoglottis costata Reid (original figure from Reid 1933; symbols added in B). mf = median furrow

on valve, r = one of two ridges on valve, rc = (resin) cavity, s = septum, v = valve. Scale bar = 10 mm. A, oblique view of apex. B, as A, with lines

accentuating the position of septa (boundaries of valves) around the apex. C, oblique view of base. D, as C, with lines accentuating the position of

septa (boundaries of valves) around the base.

and Fig. 5). The only remaining difference is size: the fossil measures 30 x 26 mm, which is about half as

large. Incidentally, the size of S. amazonica endocarps shows a wide range: from 20 to 60 mm long (Gunn

& Dennis 1976). Of course, the state of preservation also distinguishes the fossil from the recent fur¬

rowed endocarps. The size difference may partly be explained by the fossilization process, e.g., shrinkage

by coalihcation. The state of the fossil was characterized as “highly carbonized, close, hard and glossy

in texture, and quite black” and said to “break with a conchoidal fracture.” This was regarded by Reid

(1933) as pointing to considerable antiquity, but the actual age of the fossil is unknown. Unfortunately,

van der Ham et al., Furrowed blister pods (Sacoglottis sp.) on coasts along the northern Atlantic Ocean

145

the specimen could not be traced in the Reid collections in the Natural History Museum in London (pers.

comm. Peta Hayes 2014).

The ribbed fossil endocarps from the Neogene of Colombia described by Wijninga (1996) as type

T295 ( Humiriastrum ) are small (10-21 x 8-18 mm) and their septa are wider than the valves and therefore

do not resemble the furrowed endocarp. According to Herrera et al. (2010) the fossils represent the genus

Duckesia.

Origin

The furrowed endocarp described here is not the first sea-bean known earlier than its producer. An endo¬

carp of S. amazonica (blister pod) was described and illustrated by Clusius (1605, libr. 11:45, as Fructus I.

lac. Plateau). It was given to him by Jacques Plateau, an apothecary from Tournai (Doornik) in Belgium,

who also provided Clusius with data of armadillos (Egmond 2009; Mason 2009). Thus, Plateau had links

with the Neotropics, and according to Nelson (2000) Clusius’ blister pod was not a European sea-bean.

Sloane (1696:214) did mention S. amazonica as a sea-bean ( Fructus exoticus cinereus, cum lineis & tubercu-

lis duris ) from Jamaica and also from Jamaica and Europe (Sloane 1725:186): “This is frequently cast up

on the Shores of this Island by the Waves, and is one of those Fruits thrown on the Northwest Islands of

Scotland, by the Seas”. However, S. amazonica as a botanical species was described much later by Martius

(1827), while its stranded endocarps from Jamaica were identified as S. amazonica only in 1889 (Guppy

1917; Cuatrecasas 1961).

It is possible to speculate about the origin of the furrowed endocarps. The five specimens stranded

in Florida indicate that the Caribbean Current might have been the way of transport (Fig. 4). This would

imply that the parent species of the furrowed endocarp, like Sacoglottis amazonica, grows somewhere in

northeastern South America, within the reach of river systems like the Amazon, Orinoco, Magdalena,

and Atrato Rivers. Transport from the Caribbean Current into the Gulf of Mexico to Bermuda and via the

Gulf Stream and the North Atlantic Drift to Europe is possible. Wherever occurring, the producer of the

furrowed endocarps might be rare and threatened. In earlier times, its fruits might have been dispersed

by a presently extinct megafauna, as described by Jordano (2004) and Guimaraes et al. (2008) for several

other large-fruited Neotropical species, including the humiriaceous Duckesia verrucosa (Ducke) Cuatr.

and Fndopleura uchi (Huber) Cuatr. In Africa, large mammals (e.g., elephants) are still known to forage

for fruits of Sacoglottis gabonensis (White 1994; Morgan 2009).

CONCLUSIONS

The furrowed endocarps described herein are allocated to Sacoglottis s.l. (sensu lato, in broad sense: Saco¬

glottis and Schistostemon together).

The furrowed endocarps show a unique feature (distinct furrows) and cannot be matched with any

described endocarp of an extant Sacoglottis s.l. species. Theoretically, they might belong to Sacoglottis ma-

guirei or Schistostemon auyantepuiense, two rare species of which fruits are unknown. However, endocarps

of (possibly) related species ( Sacoglottis sp. A and Schistostemon fernandezii, respectively) are different

from the furrowed endocarps. Therefore, the endocarps of S. maguirei and S. auyantepuiense are suspected

to be different from the furrowed endocarps too. This should be checked in the held.

The furrowed endocarps are most similar to the fossil Sacoglottis costata, which is based on a single

specimen of unspecified Tertiary age from Colombia. Sacoglottis costata is distinct from the fossil spe¬

cies S. tertiaria. Apart from the state of preservation and the size difference, it is difficult to define a

distinguishing characteristic difference between the extant furrowed endocarps and the fossil S. costata.

Therefore, we do not erect a new species, but prefer to indicate the furrowed endocarps as Sacoglottis cf.

costata. It is stressed that the resemblance between Sacoglottis cf. costata and S. costata is based on surface

characters; knowledge of the internal structure of the S. costata specimen might support affinity.

If the furrowed endocarps ( Sacoglottis cf. costata) do not belong to one of the incompletely known

146

Journal of the Botanical Research Institute of Texas 9(1)

species of Sacoglottis s.l., we believe that an undescribed species, producing costata -like endocarps, exists

somewhere in northern South America. In view of the unusual morphology of the endocarps, this species

is expected to deviate distinctly from the species of Sacoglottis s.l. known at present. As common name of

Sacoglottis cf. costata, “furrowed blister pod” is proposed.

ACKNOWLEDGMENTS

We thank Gerhard Cadee (Royal Netherlands Institute for Sea Research, NIOZ, Texel, Netherlands), Rene

Cappers (Universiteit of Groningen, Netherlands), Antoine Cleef (University of Amsterdam, Netherlands),

Nicky Desvoyes (Saint Nom La Breteche, France), Fabiany Herrera (Chicago Botanic Garden, Glencoe,

Illinois, U.S.A.), Steven Manchester (University of Florida, Gainesville, Florida, U.S.A.), Lisa Green and

Helle Patterson (Bermuda, U.K.), Peta Hayes (Natural History Museum, London, U.K.), Pedro Jordano

(Estacion Biologica de Donana CSIC, Sevilla, Spain), Paul & Hiltje Maas and Willem Prud’homme van

Reine (Naturalis Biodiversity Center, Leiden, Netherlands), Robbie Smith (Bermuda, U.K.), and Vincent

Wijninga (Amsterdam, Netherlands) for advice and help; George Wolf (Pasadena, Texas, U.S.A.), who

might have put furrowed blister pod #10 on the ‘sharing table’ during the Nineteenth Annual Interna¬

tional Sea-Bean Symposium and Beachcombers’ Festival at Cocoa Beach, Florida (see Introduction); Dirk

van der Marel and Willem Renema (Naturalis Biodiversity Center, Leiden, Netherlands) for micro-CT

scanning; Connie Baak and Niko Korenhof (Naturalis Biodiversity Center, Leiden, Netherlands) for edit¬

ing the figures, and Gerhard Cadee and Fabiany Herrera for reviewing and improving the manuscript.

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68:260-266.

Cuatrecasas, J. & O. Huber. 1999. Humiriaceae. In: P.E. Berry, K. Yatskievych & B.K. Holst, eds. Flora of the Venezuelan

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Ducke, A. 1938. Plantes nouvelles ou peu connues de la region amazonienne (Xe serie). Arch. Inst. Biol. Veg., Rio Janeiro

4:1-64.

Egmond, F. 2009. The exotic world of Carolus Clusius (1526-1609). In: K. van Ommen, ed. The exotic world of Carolus

Clusius (1526-1609):7-12. Leiden University Library, Leiden, Netherlands.

Guimaraes, P.R. Jr, M. Galetti, & P. Jordano. 2008. Seed dispersal anachronisms: rethinking the fruits extinct megafauna ate.

PLoS ONE 3(3):e1745. doi: 10.1371/journal.pone.0001745.

Gunn, C.R. & J.V. Dennis. 1976. World guide to tropical drift seeds and fruits. Quadrangle/New YorkTimes, New York, U.S.A.

Guppy, H.B. 1917. Plants, seeds, and currents in the West Indies and Azores. Williams and Norgate, London, U.K.

Herrera, F., S.R. Manchester, C. Jaramillo, B. MacFadden, & S.A. da Silva-Caminha. 2010. Phytogeographic history and phylog-

eny of the Humiriaceae. Int. J. Plant Sci. 171:392-408.

Herrera, F., S.R. Manchester, J. Velez-Juarbe, &C. Jaramillo. 2014. Phytogeographic history of the Humiriaceae (Part 2). Int. J.

PI. Sci. 175:828-840.

Jordano, P. 2004. The megafaunal fruits dataset page, http://ebd10.ebd.csic.es/mywork/frubase/bigfruits.html.

Accessed Jun 2014.

Lorenzi, H. 2002. Brazilian trees: a guide to the cultivation and identification of Brazilian native trees 2. Balogh Interna¬

tional, Instituto Plantarum de Estudios da Flora, Sao Paulo, Brazil.

Lott,T.A., D.L. Dilcher, S.P. Horn, O. Vargas, & R.L. Sanford. 2011. Pleistocene flora of Rio Puerto Viejo, Costa Rica. Palaeontol

Electron 14,1,5A:1-15. http://palaeo-electronica.org/2011_1/242/index.html. Accessed June 2014.

Martius, C.F.P. 1827. Nova genera et species plantarum brasiliensium 2:142-148.

van der Ham et al., Furrowed blister pods (Sacoglottis sp.) on coasts along the northern Atlantic Ocean

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Renner, S. 2004. Plant dispersal across the tropical Atlantic by wind and sea currents. Int. J. PI. Sci. 165(suppl):S23-S33.

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Sloane, H. 1696. Catalogus plantarum quae in insula Jamaica sponte proveniunt, vel vulgo coluntur cum earundem

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paleobotanical record of montane and lowland forests. Rev. Palaeobot. Palynol. 92:97-156.

White, L.J.T. 1994. Sacoglottis gabonensis fruiting and the seasonal movements of elephants in the Lope Reserve, Gabon.

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www.seabean.com. 2014. What's a Sea-Bean? Accessed June 2014.

Zamora Villalobos, N. 2007. Humiriaceae. In: B.E. Hammel, M.H. Grayum, C. Herrera, N. Zamora, eds. Manual de Plantas de

Costa Rica 6. Monogr. Syst. Bot. Missouri Bot. Gard. 111:10-15.

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Journal of the Botanical Research Institute of Texas 9(1)

BOOK NOTICE

E. Charles Nelson & DavidJ. Elliott. 2015. The Curious Mister Catesby. (ISBN-13: 978-0-8203-4726-4, hbk).

University of Georgia Press, Main Library, Third Floor, 320 South Jackson Street, Athens, Georgia

30602, U.S.A. (Orders: www.ugapress.org, 1-800-266-5842). $49.95 US, 456 pp., 238 paintings/illus./

photos/maps, 8" x 11".

From the publisher: In 1712, English naturalist Mark Catesby (1683-1749) crossed the Atlantic to Virginia.

After a seven-year stay, he returned to England with paintings of plants and animals he had studied. They suf¬

ficiently impressed other naturalists such that in 1722 several Fellows of the Royal Society sponsored his re¬

turn to North America. There Catesby cataloged the flora and fauna of the Carolinas and the Bahamas by

gathering seeds and specimens, compiling notes, and making watercolor sketches. Going home to England

after five years, he began the twenty-year task of writing, etching, and publishing his monumental The Natural

History of Carolina, Florida, and the Bahama Islands.

Mark Catesby was a man of exceptional courage and determination combined with insatiable curiosity

and multiple talents. Nevertheless no portrait of him is known. The international contributors to this volume

review Catesby’s biography alongside the historical and scientific significance of his work. Ultimately, this lav¬

ishly illustrated volume advances knowledge of Catesby’s explorations, collections, artwork, and publications

in order to reassess his importance within the pantheon of early naturalists.

E. Charles Nelson is a botanist who served for two decades as a Horticultural Taxonomist at the National

Botanic Gardens in Glasnevin, Dublin. He served as Honorary Editor of Archives of Natural History (1999-

2012) and has written or edited, singly or collaboratively, nearly forty books. His most recent title is Shadow

among Splendours: Fady Charlotte Wheeler-Cuffe’s Adventures among the Flowers of Burma, 1897-1921.

DavidJ. Elliott is founder, chairman, and now Honorary Trustee of the Kiawah Island Natural Habitat

Conservancy. He has been executive director of the Catesby Commemorative Trust since 2002.

J.Bot. Res. Inst. Texas 9(1): 148.2015

A SURVEY OF THE WOODY AND CLIMBING PLANTS OF THE

REFUGIO DE AVES DR. ALEXANDER SKUTCH, “LOS CUSINGOS

PEREZ ZELEDON CANTON, COSTA RICA

Ronald L. Jones Humberto Jimenez-Saa Allen C. Risk

Department of Biological Sciences

Eastern Kentucky University

Richmond, Kentucky 40475, U.S.A.

ron.jones@eku.edu

Tropical Science Center

P.O. Box8-3870-1000

San Jose, COSTA RICA

Department of Biology and Chemistry

Morehead State University

Morehead, Kentucky 40351, U.S.A.

ABSTRACT

Tos Cusingos is located in southern Costa Rica, near San Isidro de El General, and occupies an area of 77 ha ranging in elevation from 650 to

750 m. The refuge was established by the Tropical Science Center on property formerly owned by Alexander F. Skutch, who was well known

for his research in bird behavior and for his many books and journal articles on natural history and philosophical topics. Tos Cusingos

contains one of the largest remaining tracts of primary forest in the northern portion of the valley of the Rio General. A botanical survey of

the site between 2007 and 2010 resulted in the documentation of 314 species of woody and/or climbing plants in 217 genera and 84 families.

There were 7 species of pteridophytes, 18 species of monocots, and 289 species of dicots. Of the dicots, 57 species were classified as vines, 50

as terrestrial shrubs, 6 as epiphytic shrubs, 5 as parasitic shrubs, 154 as treelets or trees, and 17 as emergent trees. Included in the species list

are 16 species that were naturalized or with the potential to become naturalized. Additional non-native species were present in the Pamela

Tankester Garden. Of the native species, ten are listed as endangered or threatened in Costa Rica. The Tropical Premontane Wet forest at Tos

Cusingos is a remnant forest representing a mid-elevation community type that was once widespread in the Rio General valley, and contains

a number of species that are disjunct from populations found in forests of the Osa Peninsula region. Tos Cusingos is included within the

Alexander Skutch Biological Corridor, which was established by the Tropical Science Center. The Tos Cusingos preserve is a focal point of

work to develop this corridor project and to establish working relationships with the local people to encourage their assistance in protecting

the remaining resources of the area.

RESUMEN

Ta reserva Tos Cusingos se encuentra en la zona sur de Costa Rica, cerca de San Isidro de El General, y ocupa una superficie de 77 hectareas,

entre 650 y 750 m.s.n.m. Fue establecida por el Centro Cientifico Tropical en la propiedad que anteriormente pertenecio a Alexander F.

Skutch, bien conocido por sus investigaciones sobre el comportamiento de las aves y por sus numerosos libros y articulos de revistas acerca

de la historia natural y de temas filosoficos. En Tos Cusingos se encuentra una de las mayores extensiones de bosque primario aun presente

en la parte norte del valle del Rio General. El presente estudio se realizo entre 2007 y 2010 resultando la documentacion de las 314 especies

de plantas lenosas y/o trepadoras pertenecientes a 217 generos y 85 familias. Se documentan 7 especies de pteridofitas, 18 especies de mono-

cotiledoneas, y 289 especies de dicotiledoneas. De las dicotiledoneas, 57 especies fueron clasificadas como trepadoras, 50 como arbustos

terrestres, 6 como arbustos epifitos, 5 como arbustos parasitos, 154 como arbolitos o arboles, y 17 como arboles emergentes. Incluidas en la

lista hay 17 especies naturalizadas o con potencial para naturalizarse. Ademas, se incluyen algunas especies no nativas presentes en el jardin

Pamela Tankester. De las especies nativas, 10 figuran como amenazadas o en peligro de extincion para Costa Rica. El Bosque Humedo Pre-

montano Tropical (bhp-T) de los Tos Cusingos representa el tipo de comunidad de elevacion media que alguna vez cubrio la mayor parte del

valle del Rio General, y contiene especies que todavia forman parte de las comunidades boscosas de la Peninsula de Osa. Ta reserva Tos

Cusingos se incluye dentro del Corredor Biologico Alexander Skutch, establecido por el Centro Cientifico Tropical. Ta reserva Tos Cusingos

es un punto focal para desarrollar el corredor biologico y para establecer relaciones de trabajo con las comunidades locales buscando alentar

su apoyo a la proteccion de los restantes recursos de la zona.

INTRODUCTION

Refugio de Aves Dr. Alexander Skutch, “Los Cusingos,” hereafter referred to as Los Cusingos, is a 77 hectare

preserve in southern Costa Rica. It is managed by the Tropical Science Center (TSC), which is headquartered

in Sanjose. Numerous bird studies have been conducted (more than 300 bird species have been documented)

in the sanctuary, but there have been no comprehensive floristic studies (sporadic plant collections were made

by Alexander Skutch and others). Los Cusingos is one of the last remaining forest fragments in this now

J. Bot. Res. Inst. Texas 9(1): 149 -165.2015

150

Journal of the Botanical Research Institute of Texas 9(1)

largely agricultural region of the valley of the Rio General, an area that has been heavily impacted by conver¬

sion of forests for agriculture (especially coffee, sugar cane, and pineapple), as well as for housing and commer¬

cial enterprises. The site is also notable for the presence of several large rock formations with Native American

petroglyphs.

Visitors and researchers are welcome, and Los Cusingos is being actively promoted as a site for various

research and community activities. Researchers are encouraged to develop projects utilizing Los Cusingos,

and approval is required through the TSC. Los Cusingos is part of a network of preserves operated by the TSC.

This network was formally established in the year 2000 with the goal of conserving tropical forests as well as

providing environmental education, research opportunities, natural resources protection, environmental

services, and ecological tourism.

The goals of this study were to: 1) conduct a botanical survey focusing primarily on woody plants and

climbing plants (excluding Araceae, Bromeliaceae, and Orchidaceae); 2) prepare a species list for the preserve;

and 3) broadly describe the different kinds of plant groups present based on habit and community associa¬

tions.

Dr. Alexander Skutch and Los Cusingos

After receiving a doctorate in botany from John Hopkins University in 1928, Dr. Alexander Skutch worked on

a variety of projects in Central and South America, before settling in the 1940s on a farm near San Isidro de El

General (Skutch 1980,1992; Abarca Jimenez 2004; Lewis 2004). He named his farm “Los Cusingos,” the local

name of a species of toucan, the fiery-billed aracari (Pteroglossus frantzii Cabanis). In his early years he sup¬

ported himself by collecting plant specimens from Costa Rica and South America for other scientists. Many of

these plants were new to science, and his collaborators often named the new species in his honor, usually with

“skutchii,” as the epithet. Over his lifetime he produced some 200 scientific articles and 20 books on birds, and

another 50 articles and seven books on philosophical and autobiographical topics. His ground-breaking re¬

search into avian sociobiology and life history won him a number of national and international honors and

awards. His crowning achievement was A Guide to the Birds of Costa Rica (Stiles & Skutch 1989), published

when he was 85 years old. Toward the end of his life Dr. Skutch made arrangements with the Tropical Science

Center to maintain his farm as a bird sanctuary. He passed away in 2004, at the age of 99. Alexander Skutch is

buried in an unmarked grave behind his home at Los Cusingos.

Site Description

Los Cusingos is located near the small towns of Quizarra and Santa Elena, in the San Isidro de El General Dis¬

trict, Perez Zeledon Canton, San Jose Province, Costa Rica (Figs. 1, 2). It is 14 km (or about 20 minutes driving

time) from San Isidro de El General, which is about 3 hours by automobile from San Jose at a distance of 130

km. Los Cusingos is about 10 km southwest of the peak of Cerro Chirripo, the highest mountain in Costa Rica

at 3820 m.

The following climatic data was taken from World Weather Online (2014). In this region the dry season

extends from December through April, and the rainy season from May to late November. Rainfall averages are

less than 50 mm per month during the dry period, but increases to over 150 mm in May, and reaches highs of

around 200 mm or more per month in the rainy season. The annual precipitation at San Isidro de El General is

about 2700 mm. The average monthly daytime temperatures are relatively stable, ranging from 27°C during

the rainy period to 31°C in the dry period. Nighttime temperatures range from 18 to 19°C, averaging slightly

cooler during the dry period. The area of Los Cusingos closely corresponds to the Tropical Premontane Wet

Life Zone in the Holdridge Life Zone System (Holdridge 1967; Bolanos et al. 1999; Jimenez-Saa 2013).

The refuge is largely forested, with several streams, but has some open areas with buildings and gardens

at the north end of the refuge. The buildings include a visitor’s center (at 09°20'22"N; 83°37'43"W), a research¬

ers cabin, and Skutch’s house, which he began constructing in the early 1940s and is now restored and main¬

tained as a museum of his life’s work. A few other smaller buildings also exist on the property, and more are

planned. The area occupied by the buildings and demonstration areas is about one hectare. The remaining 76

hectares are chiefly primary forest. Elevations at Los Cusingos range from 650-750 meters, and several trails

Jones et al., Woody and climbing plants of Los Cusingos

151

Fig. 1. Location of Refugio de Aves Dr. Alexander Skutch, "Los Cusingos" in San Isidro de El General District, Costa Rica (Tropical Science Center, GIS

Department, Vladimir Jimenez).

are maintained (Fig. 2). A trail guide is available for the main loop trail, Sendero Naturalista; it provides color

photographs and descriptions of species for 21 stations that are permanently marked along the trail (Jones et

al. 2014). The main water course is the Rio Penas Blancas, which forms most of the eastern boundary of the

property. Other smaller streams include the Quebrada Llano, Quebrada Champulon, Quebrada Pejibaye, Que-

brada Cusingos, and Quebrada Mapache.

Hammel et al. (2004) noted that these forests in the Rio General valley have a floristic composition unique

in Costa Rica, often with species more characteristic of the southern Pacific Coast, especially Corcovado Na¬

tional Park. These authors further note that the forests were nearly completely destroyed over the last 60+ years

in the valley, but that the few remaining fragments provide evidence that a unique and phytogeographically

interesting forest once occupied these mid-range elevations. There have been few studies in Costa Rica of mid¬

elevation forests; examples include those by Fournier et al. (1985), Wattenberg and Breckle (1995), Homeier et

al. (2005), and Cascante Marin et al. (2012). Holdridge et al. (1971) reported studies on a number of forest sites

in Costa Rica, including several sites at similar elevation to Los Cusingos, and included information on soils,

vegetation features, quantitative vegetation data, and species lists.

152

Journal of the Botanical Research Institute of Texas 9(1)

Fig. 2. Map of Refugio de Aves Dr. Alexander Skutch, "Los Cusingos" in San Isidro de El General District, Costa Rica. (Tropical Science Center, GIS Depart¬

ment, Vladimir Jimenez).

Skutch (1980) described his first impressions of the Rio General valley on his first visit in 1935 as “an

isolated region, surrounded by vast, scarcely broken forests, and easily accessible only by air.” In this same

book he described his recollections upon first seeing the farm in 1941 that would become Los Cusingos: “What

interested me more was the forest... a large tract of unspoiled forest remained, with trees towering up to one

hundred and fifty feet, multitudes of palms with slender, towering trunks, orchids and many other epiphytes

on the trees, and beneath them many low palms and flowering shrubs, and great-leaved herbs.”

Jones et al., Woody and climbing plants of Los Cusingos

153

Skutch (1980, 1992) specifically mentions many of the species that grew on his property He noted the

larger trees such as palo de leche ( Brosimum utile), burlo ( Heliocarpus spp.), chumico ( Pourouma sp.), campana

(Gordoniafruticosa ), candela ( Virola spp.), cerillo ( Symphonia globulifera), gallinazo (Jacaranda sp.), iri chirica-

no ( Vantanea barbourii), mayo trees ( Vochysia spp.), and Goethalsia meiantha. In the understory were guarumo

(Cecropia spp.), cacique ( Myrciaria floribunda ), palms such as chonta ( Socratea sp.) and palmito ( Euterpe sp.)

and many species of Miconia. Gnarled sotacaballo trees ( Zygia longifolia) grew along the Rio Penas Blancas. He

noted his disgust for the two most aggressive vines in the forest: the gigantic leguminous vine Entada gigas with

stems looping high in the canopy from tree to tree like enormous cables, these sometimes 15 inches (38 cm)

thick, with pods over a meter long; and the slender vine Salpichlaena volubilis, a fern in the Blechnaceae, which

formed almost impenetrable thickets in the undergrowth. He also noted ongoing changes in the forest, such as

the sharp decrease in numbers of his palmito trees, cut by poachers for their edible “palm hearts,” and the aging

and loss of some canopy species, such as the campana, and the increase in numbers of the palo de leche. Once

settled in Los Cusingos, Skutch devoted his life to his bird studies and to protecting his property, but appar¬

ently did not collect many plant specimens from his farm, or compile a comprehensive list of the plant species.

METHODS

Prior to the study, a list of species documented in Perez Zeledon Canton was obtained from the databases of the

Herbario Nacional, San Jose, Costa Rica, but only about 30 specimens were actually from Los Cusingos. Addi¬

tional information on species records in the region was gathered from the INBio website (INBio 2007).

Collecting for this project was done by RLJ and HJS from February to May, 2007, in July 2008, and in

March 2009; collecting by ACR, HJS, and RLJ occurred in May, 2010. All sections of the refuge were regularly

accessed via trails, property boundaries, and stream corridors; roadsides and other areas adjacent to the refuge

were included in the survey. Typically specimens were not collected unless flowers or fruits were present; tele¬

scoping tree pruners that could extend to about eight meters were used to reach accessible specimens from the

lower canopy. Specimens from higher in the canopy were obtained through the use of weighted throwing

ropes, as well as from fallen branches. Collecting efforts were ground-based; no tree climbing was attempted.

If possible, specimens were collected in triplicate. Digital images were obtained both in the held and prior to

pressing as staged shots with a black backdrop. Detailed held notes were taken on habit (tree, shrub, or vine),

twig features (such as sap, stipules, odor), floral features (such as numbers of floral parts and colors, ovary posi¬

tion), and fruit features (color, odor, locules), as well as notes on habitat and GPS coordinates.

The plant drying method described by Blanco et al. (2006) was used; this method worked very well, usu¬

ally drying the specimens in one to three days. The specimens were divided into three sets, with the primary

set for the Herbario Nacional (CR), another set for Eastern Kentucky University (EKY), and a third set for Mis¬

souri Botanical Garden (MO). Duplicates of pteridophyte and bryophyte collections were designated for More-

head State University (MDKY).

Primary references used for the identihcation work were: 1) the Dendro-Matrix notebook prepared by

RLJ, based on participation in the Tropical Dendrology course organized by HJS in 2001 (Jimenez-Saa, 1999);

2) Gentry (1993); 3) Hammel et al. (2003a, 2003b, 2004, 2007, 2010); 4) Weber et al. (2001); and 5) the four

volume set on Arboles de Costa Rica (Holdridge et al. 1997, Zamora et al. 2000, 2004, 2011). Other useful refer¬

ences included Davidse et al. (1995), Krings and Braham (2005), Keller (2004), Mickel and Smith (2004),

Zamora and Pennington (2001), and the popular books with color photographs by Zuchowski (2005) and

Gargiullo et al. (2008). The listings and discussions of species at the site by Alexander Skutch in two of his

books (Skutch 1980, 1992) were also helpful. A number of specialists in various plant groups were consulted

(see Acknowledgements). Several internet sites also proved very helpful in identifying specimens, especially

Neotropical Herbarium Specimens (2014), Smithsonian Tropical Institute Herbarium (2014), and INBio

( 2012 ).

Angiosperm family classification follows Stevens (2001 onwards), and author abbreviations follow IPNI

(2014). Pteriodophyte family classification follows Smith et al. (2006). Binomial and trinomial nomenclature is

154

Journal of the Botanical Research Institute of Texas 9(1)

based on The Plant List (2013). In the few cases when the name used in the Manual de Plantes de Costa Rica

(Hammel et al. 2003b, 2004, 2007, 2010) differs from the accepted name in The Plant List, then that name is

given in parenthesis. No species authorities are given for native or naturalized species mentioned in the text, as

these are provided in Appendix 1. Authorities are included for non-native species discussed in the text but not

listed in Appendix 1.

RESULTS

General Overview

A total of 314 species of woody plants and/or vines, belonging to 217 genera and 84 families was documented

as part of the naturally occurring (native or naturalized) flora of Los Cusingos (Appendix 1). All but ten of the

species records are based on specimens collected during this study; the additional ten records are based on Los

Cusingos specimens already deposited by other collectors at CR. The total of 314 includes 298 native species

and 16 non-native species. Families with ten or more species were: Melastomataceae (35 species), Fabaceae

(30), Rubiaceae (20), Malvaceae (13), Solanaceae (11), Moraceae (10), and Sapindaceae (10). The genera with

five or more species were: Miconia (14 species), Psychotria (10), Piper (6), Clidemia (5), Ficus (5), and Inga (5). Of

the 314 total, only about 100 of the species were listed in the database obtained from the Herbario Nacional for

Perez Zeledon Canton.

Twenty-five species of pteridophytes and monocots were documented. There were three species of climb¬

ing or scrambling ferns and four species of arborescent ferns. For monocots there were eight species of palms,

three species of cyclanths, and seven species of graminoids.

The following definitions were used to group the dicotyledonous species into major habit forms: a) vines

are those species that climb or scramble on other vegetation; b) shrubs are those woody species that are multi¬

stemmed from the base and known to reach heights of less than 5 m (shrubby epiphytes and shrubby parasites

are also distinguished); c) treelets are those woody species with a single main stem that are known to reach

heights of 5 to 15 m; d) trees are those woody species with a single main stem that are known to reach heights

of 15 to 30 m; and e) emergent trees are those woody species with a single main stem that are known to reach

heights greater than 30 m. Of the 289 dicotyledonous species, 57 were classified as vines, 50 as terrestrial

shrubs, 6 as epiphytic shrubs or cacti, 5 as parasitic shrubs, 154 as treelets or trees (species in Appendix 1 iden¬

tified as “shrub or treelet” or “treelet or tree,” are grouped in this category), and 17 as emergent trees.

Rare and Endemic Species

There are several sources of information on the status of threatened and endangered species in Costa Rica, in¬

cluding IUCN (2013), and the CITES and MINAE lists maintained at the INBio website, Species of Costa Rica

(INBio 2007). The following species documented at Los Cusingos are included on at least one of these refer¬

ences, and are listed below with their threat category, source of listing, and number of sites at Los Cusingos:

1) Cedrela odorata, IUCN Vulnerable (Americas Regional Workshop 1998)—few sites in gardens and open areas.

2) Cyathea bicrenata (trichiata ), Theatened or Reduced Populations (INBio 2007, CITES and MINAE)—two sites, near garden and along

trail.

3) Cyathea delgadii, Threatened or Reduced Populations (INBio 2007, CITES and MINAE)—two sites along trail and near gardens.

4) Cyathea multiflora , Threatened or Reduced Populations (INBio 2007, CITES and MINAE)—many sites on trails

5) Cecropia obtusifolia, IUCN Least Concern, (Mitre 1998)—several sites on trails.

6) Enterolobium schomburgkii, UCN Least Concern, (Groom 2012)—one site in field

7) Tabernaemontana (Stemmademia ) pauli, IUCN Vulnerable, (World Conservation Monitoring Centre 1998)—two sites on trails

8) Swartzia costaricensis (simplex), IUCN Least Concern, (Lopez Poveda 2012)—one site along river.

9) Tocoyena pittieri, IUCN Vulnerable, (Nelson 1998)—one site along river

10) Weberocereus imitans, IUCN Endangered (Hammel 2013)—one site at forest edge near gardens.

Based on a database provided by the Herbario Nacional and on listings maintained by INBio (2007), two of the

above species are considered as endemics of Costa Rica— Tabernaemontana pauli and Weberocereus imitans.

Other Costa Rica endemics at Los Cusingos include Aphelandra golfoducensis, Costus stenophyllus (planted),

Dendrophthora turrialbae, Eugenia grayumii, and Heliconia danielsiana.

Jones et al., Woody and climbing plants of Los Cusingos

155

Non-native Species

Several non-native species were documented from the forested or disturbed regions of Los Cusingos, but none

appeared to be particularly invasive. The following sixteen non-native species are either naturalized at Los

Cusingos or have the potential to become naturalized [see Lobo Cabezas (2012) and Vargas (2009)] and are

included in the species list in Appendix 1: Acacia mangium, Anacardium occidentale, Antigonon leptopus, Bam-

busa vulgaris, Cqffea arabica, Elaeis guineensis, Erythrina poeppigianajusticia betonica, Bryophyllum pinnatum,

Manihot esculenta, Morus alba, Pyrostegia venusta, Stachytarpheta mutabilis, Spathodea campanulata, Theobroma

cacao, and Syzygiumjambos.

The Pamela Lankester Garden

A botanical garden, named after Pamela Lankester, the wife of Alexander Skutch, has been developed around

the homesite, and features many plants originally obtained by Skutch. Many of these plants were mentioned in

his books, and he often commented on the types of birds attracted to the different species. These early plant¬

ings have been supplemented by additional species in recent years. Many of these are non-native, but some

native species can also be found in the gardens. Lehman et al. (2009) prepared a guide (with color images and

a map) for these gardens.

There was a variety of plant groups represented in the gardens. Large herbs and similar plants included

species of Alpinia, Calathea, Codiaeum, Cordyline, Costus, Etlingera, Hedychium, Heliconia, and Megaskepasma.

Fence-row plants included Dracaenafragrans (L.) Ker Gawl, Gliricidia sepium, and Yucca guatemalensis Baker.

Hedge-row plants included Eigustrum vulgare L. and Hibiscus rosa-sinensis L., with Arachis pintoi Krapov &

W.C. Greg, and Impatiens walleriana Hook.f. planted as a groundcover in some areas. Bambusa vulgaris formed

a large thicket in the woodland edge behind the Skutch home. Cycas revoluta Thunb. has been planted, and was

the only gymnosperm on the property. Many plants producing economically important fruits were present in

the gardens, including Annona muricata L., Artocarpus altilis (Parkinson) Fosberg, Averrhoa carambola L., Bac-

tris gasipaes Kunth, Citrus spp., Coffea arabica L., Mangifera indica L., Musa spp., Persea americana L., Psidium

guajava, Sambucus spp., Syzygium malaccense (L.) Merr. & L.M.Perry and Theobroma cacao. Other introduced

species documented from the gardens and around the buildings, but not considered as part of the native or

naturalized flora were: Allamanda cathartica L., Arundina graminifolia (D. Don) Hochr., Ardisia elliptica Thunb.,

Azadirachta indica A.Juss., Crotalaria micans Link, Eeea guineensis G. Don, Plumeria rubra L., and Thunbergia

mysorensis (Wight) T. Anderson. It should be noted that some of these non-native species have the potential to

become invasive, especially Alpinia spp., Arachis pintoi, Etlingera spp., Hedychium spp., and Impatiens walleri¬

ana. The following native species were documented from the gardens: Acnistus arborescens, Cecropia peltata,

Cedrela odorata, Costus stenophyllus, Clusia croatii, Crescentia cujete, Ficus spp., Albizia saman, and Vochysia

ferruginea.

Native and Naturalized Plants of Los Cusingos

Pteridophytes

Three woody pteridophyte species with viny or vinelike habit, as well as four species of tree ferns, were docu¬

mented. The viny species were Gleichenellapectinata, Polybotrya caudata, and Salpichlaenavolubilis. The climb¬

ing Salpichlaena was exceedingly abundant and weedy, as Skutch had described in his writings (Skutch 1992).

The species of Polybotrya lacked reproductive material, and the assigning of this fern to P. caudata is tenta¬

tive—another possibility is P. gomezii R.C.Moran. It was common along the less disturbed sections of the trails

through the primary forest and along streams, often found climbing on tree trunks. Gleichenellapectinata was

observed only in one open, trail-side area near the researcher’s cabin, where it formed a large colony scram¬

bling over other vegetation. The tree ferns were Alsophilafirma, Cyathea bicrenata, C. delgadii, and C. multiflora.

The last-mentioned species was, by far, the most common tree fern with C. delgadii and A.firma patchily dis¬

tributed within the refuge and C. bicrenata only observed as single individuals in two locations.

Additional work on the pteridophyte (as well as the bryophyte) flora has been initiated, but is yet incom¬

plete. A few common and conspicuous species have been identified: Pteridium aquilinum (L.) Kunth formed a

large population on the disturbed river bank in the southeastern section of the refuge; epiphytic ferns included

156

Journal of the Botanical Research Institute of Texas 9(1)

Microgramma lycopodioides (L.) Copeland, Niphidium crassifolium (L.) Lellinger, and Serpocaulon triseriale

(Sw.) A.R. Sm. Preliminary observations and collections suggest that the Polypodiaceae and Hymenophylla-

ceae are particularly species-rich at Los Cusingos. Other families with several species observed at the study site

were Blechnaceae, Dennstaedtiaceae, Lomariopsidaceae, and Selaginellaceae.

Palm and Palmlike Species (Arecaceae and Cyclanthaceae)

Palm and palmlike plants at Los Cusingos totalled 11 species, and including several tree-sized and dwarf

palms. The most conspicuous and abundant of the tall palms was Socratea exorrhiza, notable for its widely

spaced and spiny prop roots. Another tall palm was the palmito, Euterpe precatoria, with the prop roots dense¬

ly packed and nearly smooth. Several dwarf palms and palmlike species occur in the understory, including two

species of Bactris, two species of Geonoma , and one species of Chamaedorea. Three species in the palmlike

Cyclanthaceae were documented: Asplundia alata , A. euryspatha, and Carludovica drudei.

Woody and Climbing Graminoids (Cyperaceae and Poaceae)

Seven woody or climbing grasses and sedges were documented at Los Cusingos. These include two kinds of

bamboo, the non-native Bambusa vulgaris and the native Chusquea simpliciflora. A semi-woody grass, Lasiacis

procerrima, reaching over 3 m in height, dominated large openings on lower terraces near the river. Three spe¬

cies of climbing grasses were documented, Lasiacis rugelii, L. sorghoidea, and Olyra latifolia. These species of¬

ten climb into the lower branches along the trails and dangle from branches. A prickly sedge, Scleria secans,

was also abundant, dangling from branches and posing an entangling hazard for hikers.

Other observations on the monocot flora at Los Cusingos

A rich monocot flora exists at Los Cusingos, with the families Araceae, Bromeliaceae, and Orchidaceae appear¬

ing to be particularly species-rich. These non-woody groups were not targeted during this study, but many

were photographed, and some were documented with collections. The following taxa were noted at Los Cusin¬

gos: Araceae— Anthurium spp., Dieffenbachia spp., Dracontium spp., Monster a deliciosa Liebm., and Philoden¬

dron spp.; Costaceae— Costus guanaiensis Rusby, Costus scaber Ruiz & Pavon; and Costus stenophyllus Standi.

& L.O. Williams; Heliconiaceae— Heliconia danielsiana W.J. Kress, Heliconia irrasa Lane ex R.R. Sm., and other

Heliconia spp.; Marantaceae— Calathea spp.; and Orchidaceae— Brassia gireoudiana Rchb.f. & Warsz., Epiden-

drum spp., Gongora spp., Oncidium spp., and Xylobium spp. Many of these taxa can be found in and around the

gardens at Los Cusingos, either occurring naturally or having been transplanted from the surrounding forest.

Epiphytic and Parasitic Shrubs

Five species of woody parasites were documented, in two families: Loranthaceae— Psittacanthus schiedeanus,

Struthanthus leptostachyus, and S. orbicularis; and in the Santalaceae— Dendrophthora turrialbae and Phoraden-

dron crassifolium. The Psittacanthus formed red and yellow patches at the tops of some trees near the research¬

er’s cabin, and the other species were found in lower to upper branches of trees along the forest edges or in open

fields.

Six shrubby epiphytes in five families were documented, and these included two with dangling branches:

the red/white-flowered Satyria panurensis, and the orange-flowered Juanulloa mexicana, and two with more

upright habit, the white/orange-flowered Drymonia macrantha and white/red flowered Souroubea vallicola. At

least two species of epiphytic cacti occur in the area: Epiphyllum phyllanthus and the endangered Weberocereus

imitans.

The epiphytic and parasitic flora at Los Cusingos is obviously much richer than indicated by the docu¬

mentation of the 11 shrubby taxa listed above. Many tree or treelet species, especially in such genera as Clusia

and Ficus, often initiate their growth as epiphytes. Because they reach greater sizes than the shrubby species

above, they are categorized as treelets or trees for the purposes of this report.

Climbing Dicotyledonous species

The dicotyledenous vine flora is particularly rich at Los Cusingos, with 57 species documented in 20 families.

There were seven species in the Sapindaceae, six species in the Fabaceae, five species in the Bignoniaceae, and

four each in the Apocynaceae, Menispermaceae, Passifloraceae, and Vitaceae. Other families with numbers of

Jones et al., Woody and climbing plants of Los Cusingos

157

viny species indicated parenthetically include the following: Acanthaceae (2), Aristolochiaceae (1), Asteraceae

(2), Campanulaceae (1), Cucurbitaceae (2), Dilleniaceae (1), Loganiaceae (2), Malphigiaceae (2),

Melastomataceae (1), Polygalaceae (1), Polygonaceae (1), Rhamnaceae (2), and Schlegeliaceae (1). Davilla kun-

thii in the Dilleniaceae was very common, as were Entada gigas and Mucuna holtonii in the Fabaceae. Macrosce-

pis hirsuta (Apocynaceae) and Schlegelia parviflora (Schlegeliaceae) were rare. Introduced vines included Anti-

gonon leptopus and Pyrostegia venusta.

Shrubs

There were 50 species of terrestrial shrubs documented, with the major families and number of species as fol¬

lows: Melastomataceae (11), Rubiaceae (9), Solanaceae (9), and Asteraceae (4). All of the 9 shrub species in the

Rubiaceae were in the genus Psychotria, and there were 4 shrubby species in the genus Clidemia (Melastoma-

ceae). The endemic Aphelandra golfodulcensis was found at only a single site. Poikilacanthus macranthus was

common along trails in the forest understory. Psychotria data and P. poeppigiana , the famous “hot lips” plants,

were also very common, especially in more open places.

Treelets and Trees

There were 154 species classified as treelets or trees (woody plants commonly reaching heights of 5 to 30 m).

These species comprised almost half of the woody species at Los Cusingos. Families with five or more species

of treelets or trees are as follows, all except Rubiaceae with a majority of their species in this category: Clusia-

ceae (5/6), Euphorbiaceae (6/7), Fabaceae (17/25), Lauraceae (5/6), Melastomaceae (21/33), Moraceae (8/9),

Myrtaceae (5/5), Piperaceae (6/6), and Rubiaceae (8/18). Several of the tree species are non-native and natural¬

ized in the area, including Acacia mangium, Erythrina poeppigiana, and Syzygiumjambos. Genera with the larg¬

est number of species in this category were Miconia (13 species), Piper (6), Ficus (5), Inga (5), and Psychotria (5).

See the Discussion for further listings of these tree and treelet species associated with various habitats at Los

Cusingos.

Emergent Trees

Seventeen emergent species were identified, these being the trees capable of reaching the greatest heights and

with crowns projecting above the canopy formed by the majority of the trees. Two of the largest individuals in

diameter were a Brosimum utile along Quebrada Cusingos, and a Ceibapentandra along the Rio Penas Blancas,

north of the Toma de Agua, at the convergence of the two streams. Both trees were about 140 cm dbh. There

were numerous other trees close to 100 cm dbh. The following is a list of emergent species at Los Cusingos:

Brosimum utile, Buchenavia tetraphylla, Ceibapentandra, Dilodendron costaricense, Ficus tonduzii, Humiriastrum

diguense, Lafoensia punicifolia, Ocotea cernua, Pourouma bicolor, Pouteria torta subsp. tuberculata, Sloanea lauri-

folia, Sterculia recordiana, Symphonia globulifera, Terminalia oblonga, Virola surinamensis, Vochysia ferruginea,

and V guatemalensis.

DISCUSSION

This study has resulted in a substantial increase in the number of herbarium specimens from this northern

section of the Rio General valley. Prior to this study only about 30 woody and/or viny species had been re¬

corded from Los Cusingos; 314 have now been documented at Los Cusingos, and only about 100 of these spe¬

cies were listed in databases provided by the Herbario Nacional for Perez Zeledon Canton. These collections

also provided additional records for species and genera with only a few representative specimens in the Her¬

bario Nacional, as well as additional flowering and fruiting specimens of the less collected (and difficult to

press) species in the Arecaceae and other large-leaved groups. Several of the species documented were listed by

Hammel et al. (2004) as indicative of remnant populations in the Rio General valley of species more typical of

the Osa Peninsula, including the following: Annona amazonica, Brosimum utile, Buchenavia tetraphylla, Goua-

nia colombiana, Humiriastrum diguense, Sloanea laurifolia, Socratea exorrhiza, and Virola surinamensis.

As described in Hartshorn (1983), a typical mature forest community of the Tropical Premontane Wet

Forest life zone is a medium to tall, semi-evergreen forest with two or three strata. The canopy trees are 30 to

158

Journal of the Botanical Research Institute of Texas 9(1)

40 m tall, frequently covered by a dense layer of moss, and often buttressed. The understory trees are 10 to 20

m tall with deep crowns, including palms with stilt roots and occasional tree ferns. A dense shrub layer 2 to 3

m tall is also often present. The ground layer in this forest type is composed mostly of ferns, with epiphytes

being present but not conspicuous, and with abundant climbing vines. This description generally fits the for¬

ested sites at Los Cusingos.

A typical site within the relatively undisturbed forest community at Los Cusingos would include emer¬

gent species, some with extensive buttresses, such as Brosimum utile, Ceibapentandra, Ficus tonduzii, Humirias-

trum diguense, Lafoensia punicifolia, Mouriri gleasoniana, Sloanea laurifolia, Symphonia globulifera, Virola suri-

namensis, Vochysia ferruginea, and V guatemalensis. The understory of trees, treelets, and shrubs included

many kinds of dicotyledonous trees, with Myrciariafloribunda, Tovomita weddeliana, and Vismia macrophylla

being common. Other species in the understory included at least two species of tall palms with stilt roots

(Socratea exorrhiza, very common, and Euterpe precatoria, less common), with several species of shrubby

palms (species of Bactris, Chamaedorea, and Geonoma) being abundant as well. Tree ferns were also present

and mostly under five meters tall, with Alsophila firma and Cyathea multiflora most common. High-climbing

vines, especially Entada gigas and Mucuna holtonii, were conspicuous, and Salpichlaena volubilis was nearly

ubiquitous. Thick layers of moss were present on the tree trunks and branches, and epiphytes were common.

Tree species that tended to be more common in secondary forests and other areas of recent disturbance

include Alchornea latifolia, Bellucia pentamera, Byrsonima crassifolia, Cecropia peltata, Cedrela odorata, Clusia

spp., Cordia bicolor, Dendropanax arboreus, Erythrina spp., Gliricidia sepium, Goelthalsia meiantha, Guatteria

recurvisepala, Heliocarpus americanus, Inga spp. Jacaranda copaia, Maquira guianensis, Miconia spp. (and other

genera and species in the Melastomataceae), Piptocoma discolor, Pourouma bicolor, Protium spp., Schefflera moro-

totoni, Simarouba amara, Stryphnodendron microstachyum, Eurpinia occidentalis, and Vismia spp.

Zygia longifolia was probably the most important canopy tree along the major river corridor; Ficus spp.

and Dendropanax arboreus were also common. Other species noted predominantly along the Rio Penas Blancas

and the smaller streams included: Condaminea corymbosa, Marila laxiflora, Protium tenuifolium, Sorocea affinis,

Swartzia costaricensis, Eetragastris panamensis, and the rare Eocoyenapittieri.

Of the tree species mentioned by Skutch (1980, 1992) several were not observed during this study, in

particular the campana tree ( Gordoniafruticosa ), and the ira chiricano ( Vantanea barbourii ). Skutch had noted

that the campana trees seemed to be dying out, and replaced by Brosimum utile. The ira chiricano was also

searched for, but only its close relative, Humiriastrum diguense, also called chiricano, was located. It should be

noted that specimens could not be obtained from some tall trees at Los Cusingos, and their identity remains

unknown.

Studies of other mid-elevation sites in Costa Rica (Fournier et al. 1985; Wattenberg & Breckle 1995; Ho-

meier et al. 2005; Cascante Marin et al. 2012) reported forest compositions very different from that of Los

Cusingos. Most of the prominent species at Los Cusingos were not present at these other sites. Some sites re¬

ported by Holdridge et al. (1971) as Tropical Premontane Wet were generally similar to the Los Cusingos forest.

A site near San Isidro de El General at 620 meters elevation shared many species with Los Cusingos, including

Brosimum utile, Dendropanax arboreus, Sloanea laurifolia, Socratea exorhizza, Symphonia globulifera, and about a

dozen other species in common with Los Cusingos. Several other premontane wet sites were described but all

had very few species in common with those of Los Cusingos.

As noted by Hammel et al. (2004), such forest fragments as represented by Los Cusingos provide a

glimpse into the past, a suggestion of the types of forest communities that once covered this great river valley.

From a Google Earth view, Los Cusingos and a few nearby forested (but unprotected) tracts stand out in an

otherwise significantly altered landscape that extends northwest about 5 km to the lowermost forested ridges

of Cerro Chirripo. Hardly any other substantial forest tracts can be seen for some 15 km southeast—the entire

valley between Cerro Chirripo and the Fila Costena (Coastal Mountain Range) has essentially been deforested.

Such a view magnifies the significance of Los Cusingos, as it is currently the largest protected tract in an area

of about 35 sq km southwest of Chirripo National Park and northeast of Hermosa and Penas Blancas. This

Jones et al., Woody and climbing plants of Los Cusingos

159

rectangular region immediately south of Cerro Chirripo has now been designated by the Tropical Science

Center as the Alexander Skutch Biological Corridor; this corridor will link the remaining forest patches in the

vicinity with the extensive forests occupying the steep ridges extending south from Chirripo National Park.

Los Cusingos has become a focal point of work to develop this wildlife corridor project and to establish work¬

ing relationships with the local people to encourage their assistance in protecting the remaining resources of

the area.

Although widely known as a bird sanctuary, Los Cusingos can also be considered as a remnant biological

community, providing a haven for the many kinds of life associated with the long-vanished forests of the Rio

General valley. The refuge can thus serve as a living laboratory for a variety of studies on the plants, animals,

and their interactions, and provide a better understanding of what has happened to these ecosystems in the

past, and what may happen in the future.

In his A Naturalist in Costa Rica (1992), Skutch provides some vivid descriptions of what the countryside

was like, and what came to be. When he first entered the Rio General valley in 1935 there were “vast areas of

forest untouched by the ax, stretching in unbroken majesty from the seacoast to the open paramos of the Cor¬

dillera de Talamanca.” He later observed that the “destructions of the forests ... has been particularly striking

in this valley ... of the once magnificent wilderness only shreds and patches remain in the valley ... one of the

largest remaining tracts of forest in the valley is that of a 100 acres of so on this farm, which for nearly thirty

years I have tried to preserve in its pristine state.” He lived for 63 years at Los Cusingos, simply and close to the

earth. He witnessed the loss of the surrounding forests, but he continued to protect his land, and to study, and

to write: “And of the things that have gone, some have been preserved, albeit imperfectly, in memory and in

writings, in which I have tried to convey to others their interest and beauty.”

APPENDIX 1

Native and naturalized vascular plant species of Los Cusingos. Collection numbers for specimens collected by

RLJ and HJS are listed without initials; ACR refers to specimens collected by ACR, RLJ, and HJS. Other speci¬

mens from Los Cusingos already housed at CR are indicated by collector name and collection number. Non¬

native species are indicated by an asterisk (*). See text for definitions of habit forms.

I. PTERIDOPHYTES—woody and/or climbing ferns

Blechnaceae

Salpichlaena volubilis (Kaulf.) J. Sm. 10,136; ACR 17,418; climbing

fern.

Cyatheaceae

Alsophila firmo (Baker) D.S. Conant. 10,556; ACR 17,530, 17,585;

arborescent fern.

Cyathea bicrenata Liebm. ACR 17424,17,555; arborescent fern.

Cyathea delgadii Sternb. ACR 17,529,17,586; arborescent fern.

Cyathea multiflora Sm. 10,242; ACR 17,419, 17,440, 17,587; arbo¬

rescent fern.

Dryopteridaceae

Polybotryacaudata Kunze. 9886,10,240; ACR 17,453; climbing fern.

Gleicheniaceae

Gleichenellapectinata (Willd.) Ching 9868; ACR 17574; scrambling

fern.

II. ANGIOSPERMS—woody and/or climbing monocots

Arecaceae

Bactris glandulosa Oerst. 10,180; shrubby palm.

Bactris hondurensis Standi. 10,150; ACR 17,532; shrubby palm.

Chamaedorea tepejilote Liebm. 10,204; shrubby palm.

*Elaeis guineensis Jacq. 10,149; arborescent palm.

Euterpeprecatoria Mart. 10,151,10166,10212; arborescent palm.

Geonoma ferruginea H. Wendl. ex Spruce. 10,029; shrubby palm.

Geonoma pinnatifrons Willd. subsp. oxycarpa (Mart.) AJ. Hend.

10,213; shrubby palm.

Socratea exorrhiza (Mart.) H. Wendl.10,251; arborescent palm.

Cydanthaceae

Asplundia alata Harling. 10,224; shrubby cyclanth.

Asplundia euryspatha Harling. 10,162; climbing cyclanth.

Carludovica drudei Mast. 10,181; shrubby cyclanth.

Cyperaceae

Scleria secans (L.) Urb. 10,099; prickly climbing sedge.

Poaceae

*Bambusa vulgaris Schrad. 10,066; non-native bamboo.

Chusquea simpliciflora Munro. 10,370; native bamboo.

Lasiacis maculata (Aubl.) Urb. [ =L . sorghoidea (Ham.) Hitchc. &

Chase]. 10,131; climbing grass.

Lasiacisprocerrima (Hack.) Hitchc. 9856. semi-woody grass.

Lasiacis rugelii (Griseb.) Hitchc. 9940; climbing grass.

Olyra latifolia L. 10,036,10,154; bamboo-like grass.

III. ANGIOSPERMS—woody and/or climbing dicots

Acanthaceae

Aphelandra golfodulcensis McDade. 9950; shrub.

*Justicia betonica L. 9817; shrub.

Mendoncia gracilis Turrill. 10,207; woody vine.

Mendoncia lindavii Rusby. 10,390; woody vine.

Odontonema tubaeforme (Bertol.) Kuntze. 10,047,10197; shrub.

Poikilacanthus macranthus Lindau. 9841; shrub.

160

Thunbergio erecto (Benth.) T. Anderson. 9961,10,031; woody vine.

Achariaceae

Lindockerio lourino C. Presl. 9878,9921,10,534; shrub or treelet.

Actinidiaceae

Sourouio montono Seem. 10,394; tree.

Anacardiaceae

*Anacardium occidentale L. 9939; tree.

Spondias radlkoferi Donn. Sm. 10,210; tree.

Annonaceae

Annona omozonico R.E.Fr. Q. Jimenez 1591 (CR); tree.

Guotterio recurvisepolo R.E. Fr. 9823,9884; tree.

Xylopio frutescens Aubl. 10,248; treelet.

Apocynaceae

Mocroscepis hirsuta (Vahl) Schltr. 10,133; woody vine.

Mondevillo hirsuta (Rich.) K.Schum. 9844,9901; woody vine.

Peltastes isthmicus Woodson. 9946; woody vine.

Prestonia portobellensis (Beurl.) Woodson. 10,100; woody vine.

Tabemaemontana donnell-smithii Rose ex J.D. Sm. 10,205; tree.

Tabernaemontana pauli (Leeuwenb.) A.O. Simoes & M.E. Endress.

10,155,10,165; tree.

Araliaceae

Dendropanaxarboreus (L.) Decne. & Planch. 9888,9962,9987; tree.

Schefflera morototoni (Aubl.) Maguire, Steyerm. & Frodin. 10,008,

10085; tree.

Aristolochiaceae

Aristolochia pilosa Kunth. 10,057; herbaceous vine

Asteraceae

Eirmocephala brachiata H. Rob. 10,032; shrub.

Hidalgoa temata La Have. 10,178; woody vine.

Lasianthaea fruticosa (L.) K.M. Becker. 9827; shrub.

Mikania riparia Greenm. ex B.L. Rob. 9829. Semi-woody vine.

Piptocarphapoeppigiana (DC.) Baker. 10,079; shrub.

Piptocoma discolor (Kunth) Pruski. 9885; tree.

Zexmenia virgulta Klatt. 9857; shrub.

Begoniaceae

Begonia multinervia Liebm. 9965,10,033; shrubby begonia.

Bignoniaceae

Callichlamys latifolia (Rich.) K. Schum. 10,164; woody vine.

Crescentia cujete L. 10,065; treelet.

Fridericiapatellifera (Schltdl.) L.G. Lohman. 9914; woody vine.

Handroanthus ochraceus (Cham.) Mattos. 10,157; tree.

Jacaranda copaia (Aubl.) D. Don. 10,148; tree.

Lundia corymbifera (Vahl) Sandwith. 10,536; woody vine.

Pleonotoma variabilis (Jacq.) Miers. 9873; woody vine.

*Pyrostegia venusta (Ker Gawl.) Miers. 9990; woody vine.

*Spathodea campanulata P. Beauv. 10,123; tree.

Bixaceae

Bixa orellana L. 9947; tree.

Boraginaceae

Cordia bicolor A. DC. 10,161; tree.

Cordia curassavica (Jacq.) Roem. & Schult. 9974; shrub.

Tournefortia bicolor Sw. 10,101; shrub.

Burseraceae

Protium ravenii D.M. Porter. 10,092,10,363,9830; tree.

Protium tenuifolium (Engl.) Engl, subsp. sessiliflorum (Rose) D.M.

Porter. 10,027,10,374b; tree.

Tetragastrispanamensis (Engl.) Kuntze. 10374a; tree.

Journal of the Botanical Research Institute of Texas 9(1)

Cactaceae

Epiphyllum phyllanthus (L.) Haw. 10,082; epiphytic cactus.

Weberocereus imitans (Kimnach & Hutchinson) Buxb. 10,194;

epiphytic cactus.

Calophyllaceae

Marila laxiflora Rusby. 10,078,9942; tree.

Campanulaceae

Centropogon granulosus C. Presl. 10,392; woody vine.

Cannabaceae

Celtis iguanaea (Jacq.) Sarg. 10,114,10,217; shrub or treelet.

Trema micrantha (L.) Blume. 10,116; shrub or treelet.

Caricaceae

Jacaratia dolichaula (Donn. Sm.) Woodson. 9840; tree.

Chysobalanaceae

Licania platypus (Hemsl.) Fritsch. 10,062,10159; tree.

Clethraceae

Clethra lanata M. Martens & Galleotii. (C. costaricensis Britton).

10,107; D. Norby 365 (CR); tree.

Clusiaceae

Chrysochlamysglauca (Oerst., Planch., &Triana) Hemsl. 9983; shrub

or treelet.

Clusia croatii DArcy. 9955,10,045,10,063,10189; shrub or treelet,

usually epiphytic.

Clusia uvitana Pittier. 10,211; shrub or treelet, sometimes epiphytic.

Garcinia intermedia (Pittier) Hammel. 9988; treelet or tree.

Symphonia globulifera L.f. 9943,10,090; emergent tree.

Tovomita weddeliana Planch. &Triana. 9903; shrub or treelet.

Combretaceae

Buchenavia tetraphylla (Aubl.) R.A. Howard. 10,080; emergent tree

Terminalia oblonga (Ruiz & Pav.) Steud. 10,097; emergent tree.

Convolvulaceae

Jacquemontia ciliata Sandwith. 9815; herbaceous vine.

Maripa nicaraguensis Hemsl. 10,004,10,049,10,176,10,537; woody

vine.

Crassulaceae

*Bryophyllum pinnatum (Lam.) Oken. [=Kalanchoe pinnata (Lam.)

Pers.]. 9944; shrub.

Cucurbitaceae

Gurania coccinea Cogn. 9899,9915; woody vine.

Sechium pittieri (Cogn.) C. Jeffrey. 9995; woody vine.

Dichapetalaceae

Dichapetalum brenesii Standi. 9999; shrub or treelet.

Dilleniaceae

Davilla kunthii A.St.-Hil. 9863,9898; woody vine.

Elaeocarpaceae

Sloanea laurifolia (Benth.) Benth. 10,172,10019; emergent tree.

Ericaceae

Satyria panurensis (Benth. ex Meisn.) Hook.f. ex Nied. 10,127;

epiphytic shrub.

Erythroxylaceae

Erythroxylum citrifolium A. St.-Hil. 10,042; shrub or treelet.

Erythroxylum macrophyllum Cav. 10,025; shrub or treelet.

Euphorbiaceae

Acalypha diversifolia Jacq. 10,104; shrub or treelet.

Acalypha macrostachya Jacq. 9864; shrub or treelet.

Alchornea latifolia Sw. 9836,9929, 9948; tree.

Croton draco Schltdl. 9985,10,401; treelet or tree.

Jones et al., Woody and climbing plants of Los Cusingos

161

Croton smithianus Croizat. 10,028, 10371, 10043, 10220; treelet

or tree.

Croton tenuicoudatus Lundell. 9934,10,022; treelet or tree.

*Manihotesculenta Crantz. 9941; shrub.

Fabaceae

* Acacia mangium Willd. 10,198; tree.

Albizia saman (Jacq.) Merr. 10,560; tree.

Calliandra calothyrsus Meisn. 10,000; shrub or treelet.

Calliandra riparia Pittier. 9871; shrub or treelet.

Canavalia brasiliensis Mart, ex Benth. 9860; woody vine.

Diphysa americana (Mill.) M. Sousa. 10,076; shrub or treelet.

Entada gigas (L.) Fawc. & Rendle. 9838; woody vine.

Enterolobium schomburgkii (Benth.) Benth. 9930; tree.

Erythrina costaricensis Micheli. 10,017; shrub or treelet.

Erythrina fusca Lour. 10,002; tree.

^Erythrinapoeppigiana (Walp.) O.F. Cook. 9913,10,001; tree.

Gliricidiasepium (Jacq.) Kunth ex Walp. 9892,10109; treelet or tree.

Inga edulis Mart. 10,228; tree.

Inga oerstediana Benth. 10,106; tree.

Ingaspectabilis (Vahl) Willd. 10,229; tree.

Inga thibaudiana DC. 9920,9945; tree.

Inga umbellifera (Vahl) Steud. 9820,9994; tree.

Machaerium kegelii Meisn. 10,115; woody vine.

Machaerium seemannii Benth. ex Seem. 9993,9822; woody vine.

Mimosa myriadena (Benth.) Benth. 9923; woody vine.

Mucuna holtonii (Kuntze) Moldenke. 9874,10,056; woody vine.

Platymiscium pinnatum (Jacq.) Dugand. 10,040; tree.

Senna hayesiana (Britton & Rose) H.S. Irwin & Barneby. 9821,9971,

10,538; treelet or tree.

Senna spectabilis (DC.) H.S.Irwin & Barneby. 9839, 9991; treelet

or tree.

Stryphnodendron microstachyum Poepp. 10052,10,158; tree.

Swartzia costaricensis (Britton) N. Zamora. 9952; treelet or tree.

Zygia longifolia (Humb & Bonpl. ex Willd.) Britton & Rose. 9954,

10,369; tree.

Gesneriaceae

Drymonia macrantha (Donn. Sm.) D.N. Gibson. 10,072, 10,135;

epiphytic shrub.

Humiriaceae

Humiriastrum diguense (Cuatrec.) Cuatrec. ACR 17,533; emergent

tree.

Hypericaceae

Vismia baccifera (L.) Planch. &Triana. 10,026,9870; tree.

Vismia macrophylla Kunth. 10,102; treelet.

Lacistemataceae

Lacistema aggregatum (PJ. Bergius) Rusby. 10,050, 10,119; shrub

or treelet.

Lamiaceae

Cornutia pyramidata L. 10,054; shrub or treelet.

Scutellaria costaricana H. Wendl. 10,126; shrub.

Lauraceae

Beilschmiedia tovarensis (Klotzsch & H. Karst ex Meisn.) Sach. Nishida.

10,005a; tree.

Nectandra membranacea (Sw.) Griseb. 9968; tree.

Ocotea cernua (Nees) Mez. 9891; emergent tree.

Ocotea laetevirens Standi. & Steyerm. 9959. tree

Persea caerulea (Ruiz. & Pav.) Mez. 10,024; tree.

Loganiaceae

Strychnos chlorantha Progel. 10,223; woody vine.

Strychnos panamensis Seem. 9949; woody vine.

Loranthaceae

Psittacanthus schiedeanus (Schltdl. & Cham.) G. Don. 10,069;

parasitic shrub.

Struthanthus leptostachyus (Kunth) G. Don 10,061; parasitic shrub.

Struthanthus orbicularis (Kunth) Eichler. 10,014, 9910; parasitic

shrub.

Lythraceae

Lafoensia punicifolia DC. 10,111,10,359; emergent tree.

Malphigiaceae

Banisteriopsis muricata (Cav.) Cuatrec. 10,232; woody vine.

Byrsonima crassifolia (L.) Kunth. 10,089; tree.

Stigmaphyllon puberum (Rich.) A. Juss. 9963,9970; woody vine.

Malvaceae

Abutilon purpusii Standi. 9992; shrub or treelet.

Ceiba pentandra (L.) Gaertn. 10,252; emergent tree.

Goethalsia meiantha (Donn. Sm.) Burret. 10,039; tree.

Heliocarpus americanus L. 9861,10169; tree.

Hibiscus furcellatus Lam. 9843; shrub.

Malvaviscus penduliflorus Moc. & Sesse ex DC. 10,122; shrub.

Pachira aquatica Aubl. 10,558; tree.

Pachirasessilis Benth. 10,367; tree.

Pavonia dasypetala Turcz. 10,016; shrub or treelet.

Pseudobombaxseptenatum (Jacq.) Dugand. 10,353; tree.

Sida urens L. 9866; shrub.

Sterculia recordiana Standi. 10,399; emergent tree.

*Theobroma cacao L. 10,190; treelet.

Wissadula excelsior (Cav.) C. Presl. 9847; shrub.

Marcgravaceae

Souroubea vallicola\Noodson ex de Roon. 10,058; epiphytic shrub.

Melastomataceae

Aciotis indecora (Bonpl.) Triana. 9849; shrub.

Adelobotrys adscendens (Sw.) Triana. 9826; woody vine.

Belluciapentamera Naudin. 9889; treelet.

Blakea gracilis Hemsl. 9953.10,175; shrub, sometimes epiphytic.

Clidemia capitellata (Bonpl.) D. Don. 9912; shrub.

Clidemia dentata Pav. ex D. Don. 10,006; shrub.

Clidemia discolor (Triana) Cogn. 0,117,10,173,9852; shrub.

Clidemia hirta (L.) D. Don. 10,083; shrub.

Clidemia sericea D. Don. 10,095; shrub.

Conostegiasubcrustulata (Beurl.) Triana. 10,202; shrub.

Conostegia superba D. Don ex Naudin. 10,219; shrub or treelet.

Graffenrieda galeottii (Naudin) L.O. Williams. 10,130; treelet.

Henriettea succosa (Aubl.) DC. 10,108; shrub or treelet.

Henriettella fascicularis (Sw.) C. Wright. 10,067; shrub or treelet.

Leandra grandifolia Cogn. 10,357,10,364; shrub.

Miconia affinis DC. 10,064; shrub or treelet.

Miconia argentea (Sw.) DC. 9919; shrub or treelet.

Miconia bubalina Naudin. 10,146; shrub or treelet.

Miconia chrysophylla (Rich.) Urb. 9980,10,139; treelet or tree.

Miconia elata (Sw.) DC. 9835; tree.

Miconia gracilis Triana. 9853,10,177,10,351; shrub or treelet.

Miconia holosericea (L.) DC. 10,233; shrub or treelet.

Miconia lacera (Bonpl.) Naudin. 10,391,10,015,10,535; shrub.

Miconia matthaei Naudin. 9834,9848; shrub or treelet.

Miconia minutiflora (Bonpl.) DC. 10,554,10,187; shrub or treelet.

Miconia prasina (Sw.) DC. 10,218; shrub or treelet.

Miconia schlimii Triana. 10,129,10,203; shrub or treelet.

Miconia serrulata (DC.) Naudin. 9904; shrub or treelet.

Miconia trinervia (Sw.) D. Don ex Loudon. 9828,10,010; tree.

Mouririgleasoniana Standi. 9931, ACR 17,487; tree.

Ossaea macrophylla (Benth.) Cogn. 10,397; shrub or treelet.

Tibouchina longifolia (Vahl) Baill. D. Norby. 108 (CR); shrub.

162

Journal of the Botanical Research Institute of Texas 9(1)

Topobea maurofemandeziana Cogn. 9894; treelet.

Meliaceae

Cedrela odorata L. 10,055; tree.

Trichilio pallida Sw. S. Darwin 412 (CR); tree.

Menispermaceae

Abutapanamensis (Standi.) Krukoff&Barneby. 10,366; woody vine.

Cissampelos fasciculata Benth. 10,113; woody vine.

Cissampelos pareira L. 10,087; woody vine.

Odontocarya truncata Standi. 10,011; woody vine.

Monimiaceae

Mollinedia viridiflora Tul. 10,170; shrub or treelet.

Moraceae

Brosimum guianense (Aubl.) Huber ex Ducke. 10168; tree.

Brosimum utile (Kunth) Oken. 10,093,10,138; emergent tree.

Ficus brevibracteata W.C. Burger. 10,244; tree, often epiphytic.

Ficus citrifolia Mill. 10,188; treelet, often epiphytic.

Ficus colubrinae Standi. 9957; treelet, often epiphytic.

Ficus costaricana (Liebm.) Miq. 10,243,10,245,9986; treelet, often

epiphytic.

Ficus tonduzii Standi. 9960; emergent tree.

Maquira guianensis Aubl. 10,231,10,199,9938; tree.

*Morus alba L. 10,227; treelet.

Sorocea trophoides W.C. Burger. 10,112; shrub or treelet.

Myristicaceae

Compsoneura excelsa A.C. Smith. 9837; tree.

Virola sebifera Aubl. 9833,10532; tree.

Virola surinamensis (Rol. ex Rottb.) Warb. 10,132; ACR 17,492;

emergent tree.

Myrtaceae

Eugenia grayumii Barrie. 9980; treelet.

Myrcia splendens (Sw.) DC. 10,247; treelet or tree.

Myrciaria floribunda (H. West ex Willd.) O.Berg. 10,091,10,341,9881;

treelet or tree.

Psidium guajava L. 9989; treelet.

*Syzygium jambos (L.) Alston. 10,023; treelet.

Nyctaginaceae

Neea laetevirens Standi. 10,018; shrub or treelet.

Neeapsychotrioides Donn. Sm. 9922; shrub or treelet.

Passifloraceae

Passiflora ambigua Hemsl. 9858,10,153; woody vine.

Passiflora costaricensis Killip. 10,361; herbaceous or woody vine.

Passiflora quadrangularis L. 9879; woody vine.

Passiflora vitifolia Kunth. 9875; woody vine.

Phytolaccaceae

Phytolacca rivinoides Kunth & C.D. Bouche. 9846; shrub.

Piperaceae

Piper aduncum L. 9997; shrub or treelet.

Piper arboreum Aubl. 9832,10,005b; shrub or treelet.

Piper auritum Kunth. 9998; shrub or treelet.

Piper biseriatum C.DC. 10,209,9845; shrub or treelet.

Piper corrugatum Ku ntze. 1 0, 1 3 7; B u rger 4821 (CR); sh ru b or tree I et.

Piper curtispicum C.DC. 10,365,10,182; shrub or treelet.

Polygalaceae

Securidaca diversifolia (L.) S.F. Blake. 10,044; woody vine.

Polygonaceae

*Antigonon leptopus Hook. & Arn. 9979; woody vine.

Primulaceae

Ardisia opegrapha Oerst. subsp. wagneri (Mez) Pi poly & Ricketson.

10,215; shrub or treelet.

Ardisia guianensis (Aubl.) Mez. 9982; shrub or treelet.

Rhamnaceae

Gouania colombiana Suess. 10,160; woody vine.

Gouania lupuloides (L.) Urb. 9906; woody vine.

Rubiaceae

*Coffea arabica L. 10,530; shrub.

Condaminea corymbosa (Ruiz & Pav.) DC. 9981; treelet.

Flamelia patens Jacq. 10,355,10,038; treelet.

Palicourea guianensis Aubl. 10,553; treelet.

Palicourea triphylla DC. 10,084; treelet.

Palicourea tetragona (Donn. Sm.) C.M. Taylor. 10,360; treelet

Psychotria aurantibractea C.M. Taylor. 10,221; shrub.

Psychotria berteroana DC. 10,238; treelet.

Psychotria deflexa DC. 10,346; shrub.

Psychotria elata (Sw.) Hammel. 9877; treelet.

Psychotria gracilenta Mull.Arg. 9854,10140; shrub.

Psychotria marginata Sw. 9851; shrub.

Psychotria microbotrys Ruiz ex Standi. 10,171,10,222; shrub.

Psychotriapoeppigiana Mull.Arg. 9869; shrub.

Psychotria racemosa Rich. 10,009; shrub.

Psychotriasolitudinum Standi. 9905; shrub.

Rudgea cornifolia (Kunth) Standi. D. Norby 452 (CR); treelet.

Sabicea panamensis Wernham. 10,020; woody vine.

Tocoyenapittieri (Standi.) Standi. 10,105; treelet.

Salicaceae

Banara guianensis Aubl. 10,021; shrub or treelet.

Casearia sylvestris Sw. 9819; shrub or treelet.

Hasseitia floribunda Kunth. 10,103; shrub or treelet.

Santalaceae

Dendrophthora turrialbae Kuijt. 10,071; parasitic shrub.

Phoradendron crassifolium (Pohl ex DC.) Eichler. 10,013, 10,073;

parasitic shrub.

Sapindaceae

Allophyluspsilospermus Radik. 10,128,9956,9967; shrub or treelet.

Dilodendron costaricense (Radik.) A.H. Gentry & Steyerm. 10,088,

10,345; emergent tree.

Dilodendron elegans (Radik.) A. H. Gentry & Steyerm. 10,134; treelet.

Paullinia alata G. Don. 9969; woody vine.

Paullinia brenesii Croat. A. Weston 2473 (CR); woody vine.

Paullinia faginea (Triana & Planch.) Radik. J. Utley 4949 (CR);

woody vine.

Paullinia rugosa Benth. ex Radik. 10,533,10,156; woody vine.

Serjania caracasana (Jacq.) Willd. 9814; woody vine.

Serjania mexicana (L.) Willd. D. Norby 322 (CR); woody vine.

Serjania rhombea Radik. 9978; woody vine.

Sapotaceae

Pouteria durlandii (Standi.) Baehni. 10,529, T. Pennington 11369

(CR); tree.

Pouteria torta (Mart.) Radik, subsp. tuberculata (Sleumer)T.D.Penn.

10,400; emergent tree.

Schlegeliaceae

Schlegelia parviflora (Oerst.) Monach. 9936; woody vine.

Simaroubaceae

Simarouba amara Aubl. 10,003,9933; tree.

Siparunaceae

Siparuna gesnerioides A.DC. 9818,10,070; shrub or treelet.

Siparuna guianensis Aubl. 9918; shrub or treelet.

Siparuna thecaphora (Poepp. & Endl.) A.DC. 10,531; shrub or treelet.

Solanaceae

Acnistus arborescens Schltdl. 10,046; treelet.

Jones et al., Woody and climbing plants of Los Cusingos

163

Brugmansiasuaveolens (Humb. & Bonpl. ex Willd.) Bercht. & J. Presl.

10,179; shrub.

Cestrum racemosum Ruiz & Pavon. 10,216; treelet or tree.

Juanulloa mexicono (Schltdl.) Miers. 10,185, 9893, 9951; epiphytic

shrub.

Lycionthes beckneriana D'Arcy. 9831,10,041; shrub.

Lycionthes multiflora Bitter. 9902; shrub.

Solarium circinatum Bohs. 10,118; shrub.

Solarium jamaicense Mill. 9908; shrub.

Solarium rugosum Dunal. 9876,10372; shrub.

Solarium schlechtendalianum Walp. 9972,10,012,10,206; shrub.

Witheringia coccoloboides (Dammer) Hunz. 10,201; shrub.

Staphyleaceae

Turpinia occidentalis (Sw.) G. Don. 10,048,10358,10,068; tree.

Urticaceae

Cecropia obtusifolia Bertol. 10,250; treelet or tree.

Cecropia peltata L. 10,347,10,561,10086; treelet or tree.

Myriocarpa longipes Liebm. 9966; shrub or treelet.

Pourouma bicolor Mart. 9932,10,098; emergent tree.

Verbenaceae

Citharexylum cooperi Standi. 10,186; shrub or treelet.

Lantana camara L. 10,037; shrub.

*Stacinytarpineta mutabilis (Jacq.) Vahl var. violacea Moldenke.

9825; shrub.

Vitaceae

Cissus brevipes C. V. Morton and Standi. 9937; woody vine.

Cissus cacuminus Standi. 10,234; woody vine.

Cissus erosa Rich. 9909; woody vine.

Cissus fuliginea Kunth. 9911; woody vine.

Vochysiaceae

Vochysia ferruginea Mart. 10,053; emergent tree

Vochysia guatemalensis Donn. Sm. 10,249; emergent tree.

ACKNOWLEDGMENTS

This study was done in collaboration with Eastern Kentucky University, Morehead State University, the Tropi¬

cal Science Center, in San Jose, Costa Rica, and the National Herbarium of Costa Rica, in San Jose. Scientific

Passports for collecting specimens were granted by Ministerio del Ambiente y Energia (MINAE) for studies

focusing on Los Cusingos and the Rio General valley in 2007,2008,2009, and 2010. Eastern Kentucky Univer¬

sity granted a Spring Semester Sabbatical to RLJ in 2007, and also provided funding for the project through the

University Research Committee for held work in 2008, 2009, and 2010. Morehead State University helped to

defray travel costs for ACR in 2010. The Tropical Science Center (TSC) provided logistic support throughout

the project, including assistance in travel and lodging. This project overlapped the terms of three Executive

Directors of the TSC, Enrique Ramirez, Javier Espeleta, and Olivier Chassot, and all were fully supportive of

the project. Other staff of the TSC that were directly involved with the project were Rosa Elena Montero, Carlos

Hernandez, Carlos Cruz, and Eden Chinchilla. Guides from Los Cusingos, Andres Chinchilla and Byron Val-

verde, provided indispensable held assistance in locating and obtaining specimens. The staff of the Herbario

Nacional were also very helpful, with special thanks to Cecilia Pineda, the Director, for allowing the authors to

use the herbarium, and to curators Armando Estrada, Alonso Quesada, Armando Ruiz, and Joaquin Sanchez

for assisting in the identihcation of the plants. Armando Ruiz also provided much logistical help, especially in

the mailing of specimens back to the U.S. J. Richard Abbott and Jim Solomon were extremely helpful in organ¬

izing a visit (by RLJ) to the Missouri Botanical Garden (MO) Herbarium in March 2012. The following staff

members of MO provided much-needed assistance with identihcations: J. Richard Abbott, Tom Croat, Gerrit

Davidse, Shirley Graham, Michael Grayum, Peter Jorgensen, Ron Liesner, John MacDougal, Rosa Ortiz, Amy

Pool, John Pruski, Jon Ricketson, Peter Stevens, and Charlotte Taylor. We would also like to thank Barry Ham-

mel for his assistance with Clusiaceae and other families, Daniel Nickrent for his help with the woody para¬

sites, Susanne Renner for the Siparunaceae, and Dave Skinner for the Costaceae. We thank Michael Grayum

and one anonymous reviewer for their detailed reviews.

REFERENCES

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BOOK NOTICE

Ronald L. Jones & B. Eugene Wofford. 2013. Woody Plants of Kentucky & Tennessee: The Complete Winter

Guide to Their Identification and Use. (ISBN-13: 978-0-8131-4250-0, cloth; 978-0-8131-4309-5, epub;

978-0-8131-4310-1, web pdf). University Press of Kentucky, Editorial Offices, 663 South Limestone

Street, Lexington, Kentucky 40508-4008, U.S.A. (Orders: www.kentuckypress.com, 1-800-537-5487).

$45.00 US, 224 pp., 630 color photos, 2 maps, 57 figures, 7" x 10".

From the publisher: This comprehensive volume is the essential guide to woody plants in Kentucky, Tennessee,

and surrounding states during the winter season. Featuring color images of more than four hundred species,

this detailed botanical resource provides keys to the genera and species, as well as descriptions of the genera.

The species accounts include useful information on Latin meanings, common names, habitats and distribu¬

tions, and notes on toxicity, nativity, rarity, and wetland status. In addition, authors Ronald L. Jones and B.

Eugene Wofford provide notes on practical uses for the plants, including food, medicine, fiber, and weapons.

Winter identification of woody plants can be a daunting exercise, but Jones and Wofford present clear and

authoritative information that can help anyone spot these species in the wild. Whether taken into the held or

enjoyed at home, Woody Plants of Kentucky and Tennessee: The Complete Winter Guide to Their Identification and

Use is a comprehensive and accessible resource for professional and amateur botanists, students, commercial

landscapers, homeowners, and outdoor enthusiasts.

Ronald L. Jones is foundation professor of biological sciences and curator of the herbarium at Eastern Ken¬

tucky University. He is the author of Plant Fife of Kentucky: An Illustrated Guide to the Vascular Flora.

B. Eugene Wofford is research professor and director of the herbarium at the University of Tennessee. He is the

author of Guide to the Vascular Plants of the Blue Ridge and coauthor of Guide to the Trees, Shrubs, and Woody

Vines of Tennessee.

J.Bot. Res. Inst. Texas 9(1): 166.2015

CUSCATLANIA VULCANICOLA (NYCTAGINACEAE)

NUEVO REGISTRO DEL GENERO Y ESPECIE PARA LA FLORA DE HONDURAS

Jose Ledis Linares

Catedratico de Botanica

Centro Universitario Regional

del Litoral Atlantico (CURIA)

Universidad Nacional Autonoma

de Honduras (UNAH)

La Ceiba, HONDURAS

lin aresj_ 98@yahoo. com

Frank Sullyvan Cardoza Ruiz

Catholic Relief Services (CRS)

Alianza Cacao-El Salvador

Oficial de Medio Ambiente

MagisterScientiae en Manejo y Conservation de

Bosques Tropicales y Biodiversidad

Centro Tropical Agronomico de Investigacion y

Ensehanza (CATIE)

San Salvador, EL SALVADOR

fscardozan@gmail.com

Patricia Hernandez-Ledesma

Departamento de Botanica, Instituto de Biologia

Universidad Nacional Autonoma de Mexico (UNAM)

Mexico, D.F., MEXICO

phl@ibunam2. i biologia. unam. mx

RESUMEN

Se registra por primera vez a Cuscatlania vulcanicola Standi. (Nyctaginaceae) para la flora de Honduras en las localidades de Locomapa

(Departamento de Yoro) y Nuevo Celilac (Departamento de Santa Barbara). Se describe la especie tal como ocurre en el pais y se discuten

datos de su habitat y distribucion geografica.

Palabras clave: Cuscatlania, Nyctaginaceae, Locomapa, Yoro, Nuevo Celilac, flora de Honduras

ABSTRACT

Cuscatlania vulcanicola Standi. (Nyctaginaceae) is recorded for the first time for the flora of Honduras in the towns of Locomapa (Depart¬

ment of Yoro) and Nuevo Celilac (Department of Santa Barbara). The species is described as it occurs in the country, and data of its habitat

and geographical distribution are presented and discussed.

Key Words: Cuscatlania, Nyctaginaceae, Locomapa, Yoro, Nuevo Celilac, flora of Honduras

INTRODUCCION

Durante el ano 2010 se realizaron varias expediciones de colecta de plantas como parte de la consultorla de

fortalecimiento de las capacidades botanicas del personal del Jardln Botanico Lancetilla. Una de estas expedi¬

ciones tuvo lugar en la localidad de Locomapa, en el municipio y Departamento de Yoro. Mientras se haclan

recorridos por algunos remanentes de bosque tropical caducifolio, se pudo observar un parche de vegetacion

muy bien conservado, con abundante cantidad de humus y ausencia de incendios, caracterizado por la presen-

cia de especies que normalmente tar dan muchos anos en desarrollarse, como algunos cactus arborescentes del

genero Cephalocereus y Opuntia. Se realizaron algunas colectas de material fertil de diversas especies vegetales

y entre estas se colectaron algunos individuos de Cuscatlania vulcanicola Standi., de los cuales se prepararon

varios duplicados de muestras de herbario.

El genero Cuscatlania fue descrito por Standley (1923) de material procedente de la republica de El Salva¬

dor. En ese entonces fue colectada en una quebrada rocosa del departamento de San Vicente en las faldas del

volcan de San Vicente al oriente del pals. Todo el material conocido, hasta su redescubrimiento y nuevo registro

en 2014, pertenece a El Salvador (Linares et al. 2014).

J. Bot. Res. Inst. Texas 9(1): 167 -172.2015

168

Journal of the Botanical Research Institute of Texas 9(1)

El redescubrimiento

Fue durante estos recorridos, realizados por parte de esa consultorla en julio de 2010, que en la localidad de

Locomapa, Departamento de Yoro, se colectaron ejemplares de Cuscatlania vulcanicola. Asimismo, se pudo

localizar un ejemplar de herbario de ( House et al 5414, depositado en el Herbario CURLA, Fig. 3) que, indud-

ablemente representa a esta especie, con lo que se eleva a dos las localidades donde se presenta esta planta

en el pals. En Honduras, la familia Nyctaginaceae estaba representada solo por tres generos herbaceos,

Boerhavia, Salpianthus y Mirabilis (Molina 1975; Nelson Sutherland 2008), y, con la adicion de Cuscatlania vul¬

canicola, aumenta la diversidad de los generos a cuatro. Dada la proximidad geograhca, se esperarla encontrar

tambien a los generos Commicarpus, presente en Guatemala (Standley & Steyermark 1946), y Okenia, presente

en Nicaragua (Pool 2001). En este artlculo se da a conocer el hallazgo de esta interesante planta, la cual repre¬

senta un nuevo registro para la flora de Honduras, y se aumenta el area de distribucion del genero, conocido

hasta ahora solo en El Salvador (Fig. 1).

DESCRIPCION DE LA ESPECIE

Cuscatlania vulcanicola Standi., J. Wash. Acad. Sci. 13(20):437. 1923. Tipo: El Salvador. Departamento de San Vicente:

collected in a quebrada near the base of the Volcan de San Vicente, 500 m, mar 1922, Standley 21678 (holotipo: US - Imagen! ; isotipo:

MO).

Hierba perenne, ascendente o decumbente, ralces actinomorfas, ligeramente engrosadas, ca. 2-3(-4) mm de

grueso y cerca de 30 cm de largo; tallos ramibcados, las ramitas l-2mm, de grosor, glabras abajo y puberulen-

tas o vilosas arriba; los tallos engrosados en los nudos, 2-3.5 mm de grosor, entrenudos mas engrosados en la

parte basal que en la distal, comunmente de coloracion rojiza; peciolos delgados, teretes, mayormente de 1-1.5

cm de largo, glabros o esparcidamente puberulentos; hojas opuestas, en pares muy desiguales, la pequena

menos de la mitad del tamano de la mayor; la hoja mayor con margen entero, ovada a oblonga, ovada o lanceo-

lado-oblonga, 5-12 cm de largo 3-4.5 cm de ancho, acuminada o largamente acuminada, muy desigual en la

base, en un lado redondeada y en el otro aguda o atenuada, ligeramente carnosa, glabra o casi asl, con numero-

sos y conspicuos rahdios en ambas superficies.

Flores solo con el caliz como unico verticilo, pero semejando tener los dos verticilos debido a las bracteas

sepaloides que lo rodean y a que el caliz tiene una forma corolina o petaloide, es decir, semejando una corola

tlpica. Perianto infundibuliforme, rojo, purpura o magenta, casi 3 cm de largo, el tubo muy delgado, densam-

ente viscoso, viloso con pelos muy cortos, ligeramente constrenido por arriba del ovario y caedizo, dando asl la

apariencia de un ovario Infero, la limbo con forma de corola, 5-lobada; la garganta 3-4 mm de diametro; estam-

bres 3, los hlamentos hliformes, ligeramente exertos, insertos en la pared del perianto , las anteras dldimas,

blancas o amarillentas; ovario oblongo, estilo hliforme, exerto, el estigma capitado; antocarpo oblongo ovoide,

constrenido en la base y en el apice, casi 8 mm largo y 3 mm de diametro, compuesto por la base endurecida del

perianto, que encierra al utrlculo, indehiscente y hrmemente adherido a las paredes dando asl la apariencia de

un aquenio evidentemente 10-costillado, verruculoso con tricomas diminutos, el resto de la superhcie muy

esparcida y diminutamente hirtela, gris a casi negro. Inflorescencias cimosas, terminales o axilares, escasa-

mente ramihcadas o formando cimas compactas, con flores casmogamas y cleistogamas, las primeras termina¬

les, la parte distal de la rama que precede a la inflorescencia con 6-8(-10) hojas reducidas, 1-2 cm de largo y

0.8-1.3 cm ancho, opuestas, subiguales, ovadas a cordadas, densamente vlscido-vilosas, verde purpureo en el

haz y purpureas en el enves, las inflorescencias individuales rodeadas por un involucro de 4-8 bracteas libres,

lanceoladas u oblanceoladas a espatuladas, verde purpureas, 10-15 mm largo y 2-3(-5) mm de ancho, pubes-

centes, viscosas (Fig. 1). Las flores cleistogamas axilares, generalmente dando la apariencia de pequenos apen-

dices claviformes, a menudo en grupos de 2-3 o a veces solitarios sostenidos por 4-8 bracteas involucrales,

foliaceas, lanceoladas u oblanceoladas a oblongo-ellpticas, 8-10 mm largo y l-2(-3) mm de ancho, apice agudo

o acuminado, base atenuada, puberulenta.

Plantulas de germinacion epigea, cotiledones aplanados, reniformes, suborbiculares u oblados, 16-20

mm de largo, 20-22 mm de ancho, verde-oscuros por el haz y purpureos por el enves, primer par de eohlos

opuestos, ovados, base redondeada, apice agudo a acuminado, 13-15mm de largo por 9-11 mm de ancho.

Linares et al., Cuscatlania vulcanicola en Honduras

169

Fig. 1. A. Parche de vegetacion en la localidad de Locomapa, Municipio y Departamento de Yoro, donde se encontro Cuscatlania vulcanicola Standi. B.

Flores de Cuscatlania vulcanicola Standi, mostrando el perianto corolino y las anteras y estigma exerto. C. Inflorescencia lateral (presente todo el ano),

notese la ausencia de hojas reducidasy la coloracion verdosa de las bracteasy flores deistogamas que no abren. D. Inflorescencia de"invierno"en la parte

distal de la ramita. Notese las hojas reducidas y de color purpureo que preceden a la inflorescencia. La flor en el apice de la rama muestra el perianto

comenzando a tomar la forma y coloracion corolina. Fotografias: Frank Sullyvan Cardoza Ruiz.

Habitat .—Esta especie se conocla, como ya se menciono, solo de la coleccion tipo en el departamento

de San Vicente, y fue redescubierta en 2008 en el Departamento de Santa Ana, ambos en El Salvador. En

Honduras, C. vulcanicola se conoce solo de un pequeno fragmento de selva baja caducifolia ubicada en la

parte media de la cuenca del rlo Locomapa a ± 20 km al NO del municipio de Yoro y recientemente de otra

localidad del bosque seco en el municipio de Nuevo Celilac, Departamento de Santa Barbara. La localidad de

Locomapa presenta especies de arboles tlpicos de este habitat, como Acacia pennatula (Schltdl. & Cham.)

170

Journal of the Botanical Research Institute of Texas 9(1)

Benth., Astronium graveolens Jacq., Bakeridesia molinae D.M. Bates, Cochlospermum vitifolium (Willd.) Spreng.,

Luehea Candida (Moc. & Sesse ex DC.) Mart., Machaerium nicaraguense Rudd, Nopalea guatemalensis Rose,

Randia aculeata L., Rondeletia deamii (Donn. Sm.) Standi., Spondias radlkoferi Donn. Sm., Tabebuia ochracea

(Cham.) Standi, y Talisia oliviformis (Kunth) Radik. En el estrato herbaceo son comunes Achyranthes aspera L.,

Boldoa purpurascens Cav. ex Lag., Petiveria alliacea L. y Rivinia humilis L. La especie se desarrolla principal-

mente en suelos profundos, negros, ricos en materia organica y muy bien drenados, generalmente en lugares

sombreados.

Fenologia. —Los frutos originados de las flores casmogamas, aunque escasos, se pueden observar de hna-

les de diciembre a principios de marzo; mientras que los originados de las flores cleistogamas son mas

abundantes y se pueden observar casi todo el ano.

Material examinado.— HONDURAS. Santa Barbara: Municipio de Nuevo Celilac, San Jeronimo del Pinal, bosque seco, 14 0 58'59"N;

87°16 , 53 ,l O, 18 mar 2011, Paul House et al. 5414 (CURLA). Yoro: Municipio de Yoro, Parte alta del valle de Locomapa, ca. 13 km al NO (en

linea recta) de la ciudad de Yoro, cuenca del rio Locomapa, a la orilla de la carretera Yoro - Locomapa, 15 o 14'50"N y 87 o 14'08"N, 675 msnm,

10 jul 2010,J.L. Linares et al 14402 (JBL, MEXU) (Fig. 2 y 3).

Etimologia. —El nombre del genero, Cuscadania, acunado por Paul Carpenter Standley (1884-1963), viene de

Cuscatlan, nombre de origen nahuatl con que se conoce a El Salvador (Amaroli 1986), en alusion a que fue en-

contrado originalmente en esa republica; mientras que el nombre de la especie, vulcanicola, es una palabra de

Fig. 2. Distribution geografica y localidades en Honduras de Cuscatlania vulcanicola Standi..

Linares et al., Cuscatlania vulcanicola en Honduras

171

ACANTHACEAE

REPUBLICA DE HONDURAS

Departamento de Santa Barbara

Ruellia inundata Kunth

Hierba, flor rosada.

San Jeronimo del Pinal, Nuevo Celilac, Santa Barbara, 443msnm,

bosque seco, 14° 58' 59” N; 87° 16’ 53” 0.18 de marzo de 2011.

Paul House, Hermes Vega, lliam Rivera 5414

Herbario Universidad Naciona! Autonoma de Honduras (TEFH)

Fig. 3. Ejemplar de herbario de Cuscatlania vulcanicola Standi, determinada por el autor.

origen latino compuesta de vulcanus volcan, e incola habitante, en alusion a que fue encontrada originalmente

en las faldas del volcan San Vicente o Chinchontepec de El Salvador.

Comentarios. —La apariencia general es similar a la de algunas especies del genero Allionia (de Norte y

Sudamerica) o al de Mirabilis (de America tropical y subtropical) con los cuales indudablemente esta relacio-

nado (Standley 1923). Cuscatlania vulcanicola se distingue por la insercion de los estambres en la pared del tubo

del perianto (y no en la base), siendo esta una caracterlstica unica en toda la familia (Standley 1923). Esta espe-

cie es muy facilmente distinguible por la siguiente combinacion de caracterlsticas:

172

Journal of the Botanical Research Institute of Texas 9(1)

A. Hojas opuestas, en pares anisofilos, de los cuales la hoja mayor es mas de dos veces mas grande que la menor, base asimetrica.

B. Bracteas involucrales libres 4-8, sosteniendo 1-2 flores de perianto infundibuliforme, rojo, purpura o magenta, con los estambres

insertos en la pared del perianto.

CLAVE PARA LOS GENEROS DE LAS NYCTAGINACEAE HERBACEAS PRESENTES (o ESPERADAS) EN HONDURAS

1. Hojas alternas_ Boldoa

1. Hojas opuestas, sub opuestas, verticiladas o fasciculadas.

2. Antocarpo hipogeo; flores con (9—)14-19 estambres_ Okenia (presente en Nicaragua, esperadoen Honduras)

2. Antocarpo epigeo; flores con 1-5 estambres.

3. Bracteas fusionadas en la base en un involucro parecido a un caliz lobado; perianto ca 8-80 mm de largo_ Mirabilis

3. Bracteas libres, a veces muy proximas entre si, pero nunca fusionadas, perianto 1.5-30 mm de largo.

4. Perianto ca 30 mm de largo o mas, bracteas involucrales_ Cuscatlania

4. Perianto 1.5-3 mm de largo, bracteas no involucrales.

5. Plantas perennes, hierbas lenosaso arbustos, reclinadasotrepandosobre arbustos, muy ramificadasy lenosas

en la parte basal; inflorescencias umbeladas, largamente pedunculadas, perianto bianco o bianco verdoso

_Commicarpus (presente en Guatemala, esperado en Honduras)

5. Plantas herbaceas anuales o perennes, postradas o erectas, nunca trepadoras, poco ramificadas en la base,

inflorescencias cortamente umbeladas, caliz rojo purpura, blanquecino o rosado_ Boerhavia

CONCLUSIONES

En referencia al estado de conservacion de esta especie, el alto grado de deterioro y fragmentacion de su habitat

la hacen poco frecuente, y se considera extinta en otras zonas similares 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 la de Vulnerable, VU A 1(d).

AGRADECIMIENTOS

Esta publication, as! como el trabajo de campo y de herbario necesarios, fueron bnanciados por el Proyecto de

Gestion Sostenible de Recursos Naturales y Cuencas del Corredor Biologico Mesoamericano en el Atlantico

Hondureno. ALA/2006/018-324. Mision de ATI a corto plazo no. 34. Actualization del Herbario de Lancetilla y

Capacitacion en Estudios Botanicos en las Areas Protegidas. Ademas, durante el trabajo de campo se conto con

la valiosa ayuda del Ing. Luis Bejarano. En el trabajo de herbario fue muy valiosa la ayuda y colaboracion de

Ciro Navarro, director del jardln botanico de Lancetilla. Se agradece de manera especial tambien a Cyril Hardy

Nelson Sutherland, exdirector y cofundador del herbario TEFH, por sus valiosos y oportunos comentarios al

manuscrito y tambien a un revisor anonimo.

REFERENCIAS

Amaroli, P. 1986. En la Busqueda de Cuscatlan: Un proyecto etnohistorico y arqueologico. Manuscrito inedito, Direccion

del Patrimonio Cultural, San Salvador, El Salvador.

Douglas, N. & R. Spellenberg. 2010. A new tribal classification of Nyctaginaceae. Taxon 59(3):905—910.

Linares, J. L.; Cardoza, F.S Hernandez, P. 2014. Redescrubrimiento y nuevos registros de Cuscatlania vulcanicola (Nyctagina¬

ceae) para la flora de El Salvador. J. Bot. Res. Inst. Texas 8(2):603-609.

Molina R., A. 1975. Enumeracion de las plantas de Honduras. Ceiba 19(1 ):1 -118.

Nelson Sutherland, C.H. 2008. Catalogo de las plantas vasculares de Honduras, espermatofitas. Secretaria de Recursos

Naturales y Ambiente.Tegucigalpa: Editorial Guaymuras.

Pool, A. 2001. Nyctaginaceae. In: W.D. Stevens, C. Ulloa U., A. Pool, & O.M. Montiel, eds. Flora de Nicaragua. Monogr. Syst.

Bot. Missouri Bot. Gard. 85(3):1581 -1592.

Standley, P.C. 1923. New species of plants from Salvador. II. J. Wash. Acad. Sci. 13(20):36-443.

Standley, P.C. & J.A. Steyermark. 1946. Nyctaginaceae. In: P.C. Standley &J.A. Steyermark, eds. Flora of Guatemala - Part IV.

Fieldiana, Bot. 24(4):174-192.

Standley, P.C. & J.A. Steyermark. 1944. Studies of Central American plants - V. Publ. Field Mus. Nat. Hist., Bot. Ser. 23(3):

111-150.

STRUCTURAL ANALYSIS OF SHRUBLANDS ADJACENT TO THE

MONTERREY METROPOLITAN AREA, MEXICO

Pamela Anabel Canizales-Velazquez, Oscar Alberto Aguirre-Calderon,

Eduardo Alanfs-Rodriguez, Eduardo Javier Trevino-Garza

Facultad de Ciencias Forestales

Universidad Autonoma de Nuevo Leon

Carretera Linares-Cd. Victoria km 145.

Linares, Nuevo Leon, 67700, MEXICO

pamcanizaies@gmail.com

Jose Manuel Mata-Balderas

Gestion Estrategicay ManejoAmbientalS.C.

Carretera San Miguel - Fiuinala 935

Tercer Pi so, Local 38, Plaza Comercial Acanto

Apodaca, Nuevo Leon, 66647, MEXICO

ABSTRACT

Four communities of shrublands adjacent to the Monterrey Metropolitan Area of the state of Nuevo Teon, northeastern Mexico were stud¬

ied. The values of abundance, coverage, relative importance value ( R1V ), and dominance-diversity of all arboreal, shrubs, and succulent

species were recorded. An inventory of species was carried out with respect to height and diameter of individual specimens with basal stem

diameter of 1 cm or greater. A total of 84 species, 68 genera, and 30 families were recorded. The most representative family in three plant

communities was Fabaceae. The species with the highest RIV were Agave lecheguilla (Techuguilla agave, rosetophyllous desert shrubland),

Cordia boissieri (Texas Wild Olive, tamaulipan thornscrub and semithorn shrubland), and Acacia rigidula (Blackbush Acacia, piedmont

scrub). Plant communities were composed primarily of two strata: low and medium shrubs. Communities with higher aerial coverage were

tamaulipan thornscrub and piedmont scrub. According to the species dominance-diversity curve, it was determined that the communities

were at mature successional stage.

Key Words: Fabaceae, tamaulipan thornscrub, piedmont scrub

RESUMEN

Se estudiaron cuatro comunidades de matorrales adyacentes al Area Metropolitana de Monterrey del estado de Nuevo Teon en el noreste de

Mexico. Se registraron los valores de abundancia, cobertura, valor de importancia relativa ( VIR ) y la diversidad-dominancia de las especies

arboreas, arbustivas y suculentas presentes. Se llevo cabo un inventario de las especies considerando mediciones de altura y diametro de los

individuos mayores o iguales a 1 centimetro de diametro basal. Se registraron un total de 84 especies, 68 generos y 30 familias. Fa familia

mas representativa en tres de las comunidades fue Fabaceae. Fas especies con los valores de VIR mas altos fueron Agave lecheguilla (agave

lechuguilla, matorral desertico rosetofilo), Cordia boissieri (matorral espinoso tamaulipeco y matorral subinerme) y Acacia rigidula (mator-

ral submontano). Fas comunidades vegetales estan conformadas principalmente por dos estratos: arbustos bajos y medianos. Fas comuni¬

dades con mayor cobertura aerea fueron el matorral espinoso tamaulipeco y matorral submontano. De acuerdo con la curva de dominancia-

diversidad de las especies, se determino que las comunidades se encuentran en estadio sucesional maduro.

Palabras clave: Fabaceae, matorral espinoso tamaulipeco, matorral submontano

INTRODUCTION

Despite the vast territory occupied by desert areas, from year to year they are affected by changes in land use

which are the result of human activities such as rail and road infrastructures and urban development, agricul¬

ture, livestock, mining, and tourism (Ewing & Best 2004; Arriaga 2009; Alanis et al. 2013). These changes in

land use disturb the physical and biological environment by soil erosion, habitat modification, biological inter¬

actions of wild populations, animal behavior, ecosystem processes, and acceleration of the introduction of in¬

vasive species and the increase in the fragmentation of rural and urban areas (Arriaga 2009; Alix-Garcia et al.

2010; Krauss et al. 2010; Vranckx et al. 2012).

J. Bot. Res. Inst. Texas 9(1): 173 -185.2015

174

Journal of the Botanical Research Institute of Texas 9(1)

With respect to globally registered wilderness, arid and semi-arid areas of Mexico are at risk of decreasing

their coverage in the short-term future which will affect the most productive areas (Bonilla et al. 2013) with the

disadvantage that their biodiversity and ecosystem processes are poorly studied (Alanis et al. 2013) or un¬

known for some regions (Arriaga 2009).

The piedmont scrub, microphyllous desert scrub, and rosetophyllous desert shrubland are among the

vegetation types that have been recorded with a high species richness (Arriaga 2009; Canizales et al. 2009;

Encina et al. 2013; Alanis et al. in press). Despite their lesser abundance than in the humid and tropical regions,

plants in the arid and semi-arid areas of northern Mexico have evolved a rich and distinctive flora presenting

highly specialized growth forms that are generally unique (Rzedowski 1978).

According to the INEGI classification (2005), 14 types of scrub ecosystem are found in Mexico, most of

them widespread in the north of the country. In regard to the state of Nuevo Leon, northeastern Mexico, the

following five scrub communities are registered as: semithorn shrubland (SS), rosetophyllous desert shrubland

(RDS), piedmont scrub (PS), tamaulipan thornscrub (TT), and microphyllous desert scrub (MDS).

In this work, we studied four of the five most common types of shrublands found in the state of Nuevo

Leon (SS, RDS, PS, and TT) to determine their structure including their plant density (N/ha), plant coverage

(m 2 /ha), frequency, relative importance value (RlV)and dominance-diversity graph. The increase in the rate of

population growth, that of the peripheral municipalities of the Monterrey Metropolitan Area, has resulted in a

big impact on nature (Alanis 2005). Knowledge of the structure of these plant communities will contribute to

promoting a strategic planning with respect to conservation and sustainable management of these areas that

are currently heavily affected by anthropogenic activities.

MATERIALS AND METHODS

Study area. —The present study was carried out in an area of shrublands adjacent to the Monterrey Metropoli¬

tan Area of the state of Nuevo Leon, northeastern Mexico (25°09'-24°33'N, 99°07'W) which covers twelve

municipalities of the state of Nuevo Leon, has an area of 6,680 m 2 (SEDESOL et al. 2007), located between the

physiographic zones Gulf Coastal Plain and the Sierra Madre Oriental, with an altitude of 534 ma.s.l. The

characteristic climate is dry steppe, warm and extreme, with an annual average temperature of 22.1°C, with

irregular rainfall in late summer and a precipitation of 634 mm (Cervantes & Merla 1995). The study areas

were within the Monterrey Metropolitan Area in the municipalities of Monterrey (SS and PS), Santa Catarina

(RDS) and Salinas Victoria (TT) (Pig. 1).

Woody species with high abundance and cover are: Acacia rigidula, Acacia farnesiana, Havardia pollens,

Cordia boissieri, Karwinskia humboldtiana, and Prosopis glandulosa (Espinoza & Navar 2005; Alanis et al. 2008;

Jimenez et al. 2009). Table 1 presents some physical and climatic variables as a description of the studied areas.

Sampling methods. —In the years 2010 and 2011, 25 sampling plots with sizes of 100 m 2 (10 m x 10) were

established in each shrubland, resulting in a total of 100 plots for characterizing the vegetation. The distribu¬

tion of the sampling plots was systematic with a distance of 50 m between them. According to the species-area

curve, community SS presented a minimum of 25 sites, while the other communities displayed a fewer sam¬

pling plots. For the statistical analysis, we established the same number of sampling plots for each community.

A square shape was used for each plot because this shape made it easier to layout and measure structural at¬

tributes of vegetation in the studied sites (Canizales et al. 2009). In each sampling plot, an inventory of all of the

woody species was performed, including measurements such as determination of the plant coverage (m 2 ) and

stem diameter (cm) at 10 cm (d0.10 > 1 cm). Diameter measurements were performed 0.10 m above the soil

surface, which represents a standard measurement used for woody species in TT (Alanis et al. 2008; Jimenez et

al. 2009). The canopy cover was also measured based on the four cardinal points of the north-south and east-

west directions.

We analyzed relative abundance, dominance, relative importance value (RIV), and mean height of

woody individuals. The absolute abundance is Ai = Ni/S, where Ni is the number of individuals of species i and

S is the sample surface (in hectares). The relative abundance of species was calculated using the equation Ari =

Canizales et al., Shrublands adjacent to the Monterrey Metropolitan Area

175

Fig. 1 . Location of the study areas. A) North of Mexico and south of USA, B) Nuevo Leon state, northeastern Mexico, and C) municipalities of the Monter¬

rey Metropolitan Area, in addition to the study areas: TT, tamaulipan thornscrub, SS, semithorn shrubland, PS, piedmont scrub, RDS, rosetophyllous

desert shrubland.

Table 1. Description of assessed areas.

Vegetation

Tamaulipan

thornscrub

Semithorn

shrubland

Piedmont

scrub

Rosetophyllous desert

shrubland

Abbreviation

RDS

Coordinates (MTU)

368638 E

358755 E

369795 E

342616E

2866744 N

2845296 N

2832599 N

2842694 N

Altitude (meters above sea level)

480

850

660

640

Mean annual temperature (oC)

20 to 22

18 to 20

Mean annual precipitation (mm)

400 to 600

300 to 600

400 to 600

125 to 300

Soil type

Luvic Xerosols

Lithosols Rendzinas

Lithosols Vertisols

Regosols Xerosols

t(Ai/ZAi)/100, where EAi is the total abundance of species i (Jimenez et al. 2009). To estimate dominance, the

basal area for each individual was calculated. The absolute dominance of species is Di = Abi/S, where Abi is the

cover area (in square meters) of species i and S is the sample surface (in hectares). The relative dominance of

species was calculated using the equation Dri = (Di/EDi)/ 100, where Di is as aforementioned and EDi is the

total dominance of species. The relative frequency is Fri = (Fi/EFi)/100, where Fi is the number of sites that

present the species i and EEFi is the total frequency of all species (Mostacedo & Fredericksen 2000). The rela¬

tive importance value (RIV; Mueller-Dombois & Ellenberg 1974).) was calculated according to the following

formula:

RIV =

Relative abundance + relative dominance + relative frequency

The structure of the community was described in terms of the proportional abundance of species using a diver¬

sity-abundance graph which shows the graphical relationship between the importance value of species (on a

176

Journal of the Botanical Research Institute of Texas 9(1)

logarithmic scale) as a function of an array sequence of species ranging in importance from most to least (Mar-

tella et al. 2012).

RESULTS

According to the four plant communities studied, a total of 84 plant species were recorded including 68 genera

and 30 families (Table 2). Six forms of life were recorded: trees, shrubs, succulents, grasses, rosettes, and vines.

A number of 18 families were found in the Tamaulipan thornscrub. The most representative families were

Fabaceae and Cactaceae with 8 and 4 species, respectively. A total of 29 species was recorded including Cordia

boissieri (14.96%), Leucophyllum frutescens (13.29%), and Gymnosperma glutinosum (12.22%), which were the

most abundant, but it is noteworthy that none of these species belongs to the above-mentioned representative

families. The species with the highest frequency was Cordia boissieri (10.13%). The dominant life form was low

scrub (< 1.5 m) with 57% while the average recorded height was 1.86 m.

Eighteen families were recorded within the semithorn shrubland, and the most representative was

Fabaceae with 10 species. There was a total of 34 other species in this shrubland, and the most abundant were

Acacia rigidula (12.03%), Leucophyllum frutescens (11.05%), and Bcrnadia myricifolia (9.75%). The most preva¬

lent species in this community was Cordia boissieri (7.85%). The low and medium shrubs were the main form

of life, which accounted for 96% of individuals present in this community with an average height of 1.85 m.

Nineteen families were recorded in the piedmont scrub with over 34 species. The most representative was

Fabaceae with 9 species, and Acacia rigidula was the most abundant species with 45.50%. With respect to the

highest relative frequency, Acacia rigidula and Zanthozylumfagara both had 11.27%. The highest average height

recorded for this community was 2.54 m. Medium shrubs were the dominant life form with 69% of individuals.

Sixteen families and 48 species were recorded in the rosetophyllous desert shrubland. The most represen¬

tative was Cactaceae with 17 species, followed by Fabaceae with 6 species and Asteraceae and Asparagaceae

with 5 species, respectively. The most abundant and frequent species with 26.82% and 8.26%, respectively, was

Agave lecheguilla. In this community, the predominant categories were ground vegetation (41%) and low shrubs

(54%) with species types such as rosette and succulents, cacti being the most prevalent of the latter.

Frequencies evaluated for the RIV distribution of the ecosystem are shown in Figure 2. There is a decrease

in the number of species with respect to increasing percentage values of RIV, and classes of <2% of the RIV

showed the higher number of species, indicating a high number of species with little ecological weight and

conversely a small number of species with high ecological weight.

Tree covers for TT (13,718.68 m 2 /ha) and PS (17,267.78 m 2 /ha) communities were greater while covers for

SS (8,138.43 m 2 /ha) and RDS (8,710.46 m 2 /ha) were similar, but with lower values. This corresponds to the

predominant way of life in the communities as the shrub forms (low and medium) representing greater cover¬

age in communities such as TT and PS, while the forms of life which are predominantly semithorn shrubland

(SS) and rosetophyllous desert shrubland (RDS) represent less coverage (Fig. 3).

The shape of the dominance-diversity of species curve (Fig. 4) behaves as a typical log-normal curve. This

distribution is based on the number of individuals per species (on a logarithmic scale) which follows a normal

distribution whereby individuals of intermediate abundance are common and there are few species with very

high or low values of abundance (Magurran 1988).

DISCUSSION

The most representative family for the Tamaulipan thornscrub community was Fabaceae with eight species,

which agrees with previous studies developed Mora et al. (2013) and Molina et al (2013) who also recorded

eight species of Fabaceae, the former for a community with a history of agriculture/livestock use. Other re¬

searchers reporting the same information include Jimenez et al. (2012), Gonzalez et al. (2010), Alanis et al.

(2013), and Foroughbakch et al (2013). Cordia boissieri was frequently the most abundant species, in contrast

to what was reported by other authors who recorded Acacia rigidula as the most abundant species of TT (Gon¬

zalez et al 2010; Navar & Espinoza 2005). Meanwhile Jimenez et al (2012), Alanis et al (2013), and Mora

Canizales et al., Shrublands adjacent to the Monterrey Metropolitan Area

177

Table 2. Abundance, frequency, dominance and RIV in the communities assessed.

Abundance Dominance

(N/ha) Frequency (m 2 -ha)

Species No. Absolute Relative Absolute Relative Absolute Relative RIV (%)

TAMAULIPAN THORNSCRUB

Asparagaceae

Yucca filifera Chabaud

Asteraceae

Gymnosperma glutinosum (Spreng.) Less.

Berberidaceae

Berberis trifoliolata Moric.

Boraginaceae

Cordia boissieri A. DC.

Cactaceae

Coryphantha neglecta L. Brenner

Cylindropuntia leptocaulis (DC.) F.M. Knuth.

Echinocereusstramineus (Engelm.) F. Seitz

Opuntia engelmannii Salm-Dyck ex Engelm.

Euphorbiaceae

Croton torreyanus Mull.Arg.

Fabaceae

Acacia famesiana (L.) Willd.

Acacia greggii A. Gray

Acacia rigidula Benth.

Cercidium macrum I.M. Johnst.

Eysenhardtiapolystachya (Ortega) Sarg.

Havardiapallens (Benth.) Britton & Rose

Prosopis leavigata (Hu mb. & Bon Ip.

ex Willd.) M.C. Johnst.

Koeberliniaceae

Koeberlinia spinosa Zucc.

Oleaceae

Forestiera angustifolia Torr.

Rhamnaceae

Karwinskia humboldtiana (Schult.) Zucc.

Rutaceae

Zanthoxylum fagara (L.) Sarg.

Salicaceae

Neopringlea integrifolia (Hemsl.) S. Watson

Scrophulariaceae

Leucophyllum frutescens (Berland.)

I.M. Johnst.

Simaroubaceae

Castela erecta subsp. texana (Torr. & A.Gray)

Cronquist

Solanaceae

Capsicum annuum L.

Ulmaceae

Celtis leavigata Willd.

Celtis pallida Torr.

Verbenaceae

Aloysia sp.

Lantana camara L.

Zygophyllaceae

Guaiacum angustifolium Engelm.

0.39

125

500

12.22

228

5.57

153

612

14.96

0.20

2.05

0.20

1.76

0.59

0.10

0.20

1.47

0.59

140

3.42

0.29

0.20

0.78

2.35

372

9.09

0.20

136

544

13.29

104

2.54

0.10

212

5.18

244

5.96

1.76

117

468

11.44

124

3.03

1.32

165.88

1.21

0.97

6.61

253.01

1.84

6.89

3.96

42.41

0.31

3.28

10.13

4481.90

32.67

19.25

0.88

1.41

0.01

0.36

4.85

31.91

0.23

2.38

0.88

64.30

0.47

0.52

4.41

193.66

1.41

2.53

0.88

24.86

0.18

0.55

0.44

33.16

0.24

0.26

0.44

6.60

0.05

0.20

0.44

28.01

0.20

0.28

0.88

126.76

0.92

1.09

1.76

43.51

0.32

0.89

2.64

1653.61

12.05

6.04

0.88

356.28

2.60

1.26

0.88

26.45

0.19

0.42

2.64

115.79

0.84

1.42

4.85

94.85

0.69

2.63

8.81

1372.76

10.01

9.30

0.44

3.18

0.02

0.22

9.25

903.45

6.59

9.71

3.52

78.40

0.57

2.21

0.44

3.46

0.03

0.19

3.08

1597.97

11.65

6.64

7.05

1159.82

8.45

7.16

1.76

91.91

0.67

1.40

8.81

650.91

4.74

8.33

7.05

112.48

0.82

3.63

Total

1023 4092

100.00 227

100.00 13,718.68 100.00 100.00

178

Journal of the Botanical Research Institute of Texas 9(1)

Table 2. (continued)

Abundance Dominance

(N/ha) Frequency (m 2 -ha)

Species No. Absolute Relative Absolute Relative Absolute Relative RIV (%)

SEMITHORN SHRUBLAND

Asparagaceae

Agave lecheguilla Torr.

Yucca filifera Chabaud

Asteraceae

Gymnosperma glutinosum (Spreng.) Less.

Boraginaceae

Cordia boissieri A. DC.

Ehretia anacua (Teran & Berland.)

I. M. Johnston

Cactaceae

Opuntia engelmannii Salm-Dyck ex Engelm.

Ebenaceae

Diospyros texana Scheele

Euphorbiaceae

Bernardia myricifolia (Scheele) S. Watson

Croton torreyanus Mull.Arg.

Fabaceace

Acacia berlandieri Benth.

Acacia greggii A. Gray

Acacia rigidula Benth.

Caesalpinia mexicana A. Gray

Ebenopsis ebano (Berland.) Baneby &

J. W. Grimes

Eysenhardtiapolystachya (Ortega) Sarg.

Havardiapallens (Benth.) Britton & Rose

Prosopis glandulosa Torr.

Sophora secundiflora (Ortega) DC.

Mimosa malacophylla A. Gray

Fouquieriaceae

Fouquieriasplendens Engelm.

Oleaceae

Forestiera angustifolia Torr.

Bumelia lanuginosa (Michx.) Pers.

Rhamnaceae

Condalia hookeri M.C. Johnst.

Karwinskia humboldtiana (Schult.) Zucc.

Rubiaceae

Randia laetevirens Standi.

Rutaceae

Zanthoxylum fagara (L.) Sarg.

Salicaceae

Neopringlea integrifolia (Hemsl.) S. Watson

Scrophulariaceae

Leucophyllum frutescens (Berland.)

I.M. Johnst.

Ulmaceae

Celtis leavigata Willd.

Celtis pallida Torr.

Verbenaceae

Lantana camara L.

Zygophyllaceae

Guaiacum angustifolium Engelm.

Without identification

spl

sp2

0.65

108

2.93

196

5.31

268

7.26

0.54

1.41

100

2.71

360

9.75

288

7.80

120

3.25

0.98

111

444

12.03

0.54

0.33

240

6.50

0.54

0.11

0.87

0.65

1.19

1.84

0.65

104

2.82

1.84

0.33

212

5.74

1.19

102

408

11.05

1.19

1.08

176

4.77

1.08

0.76

0.33

1.37

15.99

0.20

0.74

5.46

952.10

11.70

6.69

5.12

166.77

2.05

4.16

7.85

1302.85

16.01

10.37

0.68

16.43

0.20

0.48

2.73

49.92

0.61

1.58

4.10

731.68

8.99

5.26

5.46

399.06

4.90

6.70

7.17

492.47

6.05

7.01

3.41

121.55

1.49

2.72

1.02

56.04

0.69

0.90

7.51

464.03

5.70

8.41

1.02

16.21

0.20

0.59

0.34

2.10

0.03

0.23

4.78

321.90

3.96

5.08

1.02

155.98

1.92

1.16

0.34

143.07

1.76

0.74

1.37

49.01

0.60

0.94

0.68

19.15

0.24

0.52

1.02

112.51

1.38

1.20

3.75

66.34

0.82

2.14

0.68

27.09

0.33

0.56

1.71

878.65

10.80

5.11

3.41

35.27

0.43

1.90

1.02

7.55

0.09

0.48

6.48

319.59

3.93

5.38

1.02

14.08

0.17

0.80

7.17

891.88

10.96

9.73

1.71

5.12

0.06

0.99

1.71

59.74

0.73

1.17

5.12

144.99

1.78

3.89

1.37

33.94

0.42

0.96

1.37

57.14

0.70

0.94

1.02

8.25

0.10

0.48

Total

923

3692

100.00 293

100.00 8138.43 100.00 100.00

Canizales et al., Shrublands adjacent to the Monterrey Metropolitan Area

179

Table 2. (continued)

Abundance Dominance

(N/ha) Frequency (m 2 -ha)

Species No. Absolute Relative Absolute Relative Absolute Relative RIV (%)

PIEDMONT SCRUB

Anacardiaceae

Rhus microphylla Engelm.

Boraginaceae

Cordia boissieri A. DC.

Ehretia anacua (Teran & Berland.)

I.M. Johnston

Cactaceae

Opuntia engelmannii Salm-Dyck

ex Engelm.

Ebenaceae

Diospyros texono Scheele

Diospyros virginiono L

Euphorbiaceae

Bernardia myricifolio (Scheele) S. Watson

Croton torreyonus Mull.Arg.

Fabaceae

Acocio berlondieri Benth.

Acocio fornesiono (L.) Willd.

Acocio rigidulo Benth.

Coesolpinio mexicono A. Gray

Eysenhordtio texono Scheele

Leucoeno sp.

Mimosa malacophylla A. Gray

Havardia pollens (Benth.) Britton & Rose

Sophora secundiflora (Ortega) DC.

Fagaceae

Quercuspolymorpha Schltdl. & Cham.

Lythraceae

Heimia salicifolia (Kunth) Link

Lauraceae

Litsea novoleontis Bartlett

Oleaceae

Forestiera angustifolia Torr.

Fraxinus americana L.

Rhamnaceae

Condalia hookeri M.C. Johnst.

Karwinskia humboldtiana (Schult.) Zucc.

Rubiaceae

Randia laetevirens Standi.

Rutaceae

Zanthoxylum fagara (L.) Sarg.

Sapindaceae

Sapindus saponaria L.

Sapotaceae

Bumelia lanuginosa (Michx.) Pers.

Scrophulariaceae

Leucophyllum frutescens (Berland.)

I.M. Johnst.

Solanaceae

Solanum erianthum D. Don

Ulmaceae

Celtis leavigata Willd.

Celtis pallida Torr.

Ulmus crassifolia Nutt.

Verbenaceae

Lontono Comoro L.

2.12

216

4.76

2.12

1.23

296

6.53

0.18

1.15

0.71

0.53

2.12

516

2064

45.50

204

4.50

0.44

0.35

0.53

1.41

0.26

120

2.65

0.18

1.76

128

2.82

0.35

0.71

1.06

127

508

11.20

0.18

1.06

0.79

0.18

0.88

0.53

0.71

0.18

0.98

65.36

0.38

1.16

6.37

1066.74

6.18

5.77

4.41

1005.51

5.82

4.12

3.92

44.17

0.26

1.80

9.31

1653.56

9.58

8.47

0.98

25.12

0.15

0.43

0.98

68.55

0.40

0.84

1.47

17.66

0.10

0.76

1.96

121.27

0.70

1.06

1.96

552.04

3.20

2.42

11.27

5420.23

31.39

29.39

6.37

227.93

1.32

4.06

1.96

44.10

0.26

0.89

0.98

0.85

0.00

0.45

1.96

27.62

0.16

0.88

0.98

140.05

0.81

1.07

0.49

3.31

0.02

0.26

2.94

140.27

0.81

2.13

0.98

4.54

0.03

0.39

1.96

1901.19

11.01

4.91

6.86

394.67

2.29

3.99

0.98

155.75

0.90

0.75

0.98

11.66

0.07

0.47

1.47

25.86

0.15

0.78

1.96

107.17

0.62

1.21

11.27

2429.38

14.07

12.18

0.98

33.22

0.19

0.45

2.45

103.91

0.60

1.37

1.47

62.72

0.36

0.88

0.98

2.65

0.02

0.39

2.94

398.51

2.31

2.04

2.94

63.19

0.37

1.28

1.47

947.76

5.49

2.55

0.98

1.27

0.01

0.39

Total 1134 4536 100.00 204 100.00 17,267.78100.00 100.00

180

Journal of the Botanical Research Institute of Texas 9(1)

Table 2. (continued)

Abundance Dominance

(N/ha) Frequency (m 2 -ha)

Species No. Absolute Relative Absolute Relative Absolute Relative RIV (%)

Amaranthaceae

Atriplexconescens (Pursh) Nutt.

Asparagaceae

Agave americana L.

Agave lecheguilla Torr.

Agave striata Zucc.

Dasylirion texanum Scheele

Yucca filifera Chabaud

Asteraceae

Artemisia ludoviciana Nutt.

Gutierrezia microcephala (DC.) A. Gray

Gymnosperma glutinosum (Spreng.) Less.

Parthenium incanum Kunth

Viguiera stenoloba S.F.BIake

Berberidaceae

Berberis trifoliolataMonc.

Cactaceae

Coryphantha compacta (Engelm.) Orcutt

Coryphantha difficilis (Quehl) Orcutt

Coryphantha neglecta L. Brenner

Cylindropuntia imbricata (Haw.) F.M.Knuth

Cylindropuntia leptocaulis (DC.) F.M. Knuth

Echinocactus horizonthalonius Lem.

Echinocereus enneacanthus Engelm.

Echinocereuspectinatus (Scheidw.) Engelm.

Echinocereusstramineus (Engelm.) F.Seitz

Ferocactus hamatacanthus (Muehlenpf.)

Britton & Rose

Lophophora williamsii (Lem. ex Salm-Dyck)

J.M. Coult.

Mammillaria melanocentra Poselger

Mammillariapottsii Scheer ex Salm-Dyck

Neolloydia conoidea (DC.) Britton & Rose

Opuntia engelmannii Salm-Dyck ex Engelm.

Sclerocactusscheeri (Salm-Dyck) N.P.Taylor

Thelocactus bicolor (Galeotti ex Pfeiff.)

Britton & Rose

Celastraceae

Mortonia greggii A. Gray

Ephedraceae

Ephedra antisyphilitica Berland. ex C.A. Mey.

Euphorbiaceae

Bernardia myricifolia (Scheele) S. Watson

Jatropha dioica Cerv.

Fabaceace

Acacia berlandieri Benth.

Acacia greggii A. Gray

Calliandra conferta Benth.

Eysenhardtia texana Scheele

Prosopis glandulosa Torr.

Sophora secundiflora (Ortega) Lag. ex DC.

Koeberliniaceae

Koeberlinia spinosa Zucc.

Oleaceae

Forestiera angustifolia Torr.

Pinaceae

Pinus catarinae Passiini

ROSETOPHYLLOUS DESERT SHRUBLAND

1.28

140

3.74

251

1004

26.82

2.56

2.35

0.11

0.75

208

5.56

0.11

0.32

1.60

1.71

0.11

0.64

1.92

0.11

2.46

0.53

1.28

2.35

0.21

2.46

0.11

168

4.49

0.11

0.75

0.32

0.53

132

3.53

0.43

0.64

1.92

124

3.31

2.46

140

3.74

0.21

2.35

1.28

0.11

0.32

0.64

1.24

20.39

0.23

0.92

3.72

53.53

0.61

2.69

8.26

7550.69

86.69

40.59

1.65

53.65

0.62

1.61

2.89

51.80

0.59

1.95

0.41

0.38

0.00

0.17

0.41

10.64

0.12

0.43

3.31

30.67

0.35

3.07

0.41

2.83

0.03

0.18

0.41

3.94

0.05

0.26

0.41

26.74

0.31

0.77

2.89

23.71

0.27

1.62

0.41

0.02

0.00

0.17

1.65

0.06

0.00

0.76

2.48

0.82

0.01

1.47

0.41

1.13

0.01

0.18

3.72

45.56

0.52

2.23

1.24

0.27

0.00

0.59

1.24

1.90

0.02

0.85

3.72

0.47

0.01

2.02

0.41

1.69

0.02

0.22

0.41

0.25

0.00

0.21

2.48

0.35

0.00

1.65

0.41

0.05

0.00

0.17

3.31

0.55

0.01

2.60

0.41

0.01

0.00

0.17

2.07

4.38

0.05

0.95

0.83

0.09

0.00

0.38

0.83

0.08

0.00

0.45

3.31

64.68

0.74

2.52

1.24

6.90

0.08

0.58

1.24

12.70

0.15

0.68

2.07

36.30

0.42

1.47

4.96

69.09

0.79

3.02

4.55

21.44

0.25

2.42

3.31

129.52

1.49

2.84

0.41

12.58

0.14

0.26

3.72

113.21

1.30

2.46

1.24

49.31

0.57

1.03

0.41

7.54

0.09

0.20

1.24

8.80

0.10

0.55

1.65

31.53

0.36

0.89

Canizales et al., Shrublands adjacent to the Monterrey Metropolitan Area

181

Table 2. (continued)

Species

No.

Abundance

(N/ha)

Absolute

Relative

Frequency

Absolute

Dominance

(m 2 -ha)

Relative

Absolute

Relative

RIV (%)

Rhamnaceae

Karwinskia humboldtiana (Schult.) Zucc.

0.43

1.24

14.96

0.17

0.61

Condalia hookeri M.C. Johnst.

1.82

3.31

49.79

0.57

1.90

Scrophulariaceae

Leucophyllum frutescens (Berland.)

120

3.21

4.55

29.80

0.34

2.70

I.M. Johnst.

Ulmaceae

Celtis pallida Torr.

0.11

0.41

8.55

0.10

0.21

Zygophyllaceae

Guaiacum angustifolium Engelm.

188

5.02

4.55

61.38

0.70

3.42

Larrea tridentata (Sesse & Moc. ex DC.)

112

2.99

4.55

95.76

1.10

2.88

Coville

Total

936

3744

100.00

242

100.00

8710.46

100.00

et al. (2013) recorded Cordia boissieri as part of the community but not as the most abundant. They recorded

Diospyros texana instead, as the most abundant species for communities located in Linares, Nuevo Leon. Fur¬

thermore Foroughbakch et al. (2013) recorded Prosopis glandulosa as most abundant. The studied community

presented geographic affinity with the medium semithorn shrubland the Altiplano and the Coastal Plain, to

have associations of Cordia-Leucophyllum-Zanthoxylum , although the genus Zanthoxylum is not predominant

in previous studies (Briones & Villarreal 2001). According to Muller (1947), the lower elevation and greater

precipitation correlate with vegetation characterized by thorny species and a growth more dense of shrubs and

low trees.

Fabaceae was the best represented family of the semithorn shrubland community with 10 species, which

is consistent with the findings of Estrada et al. (2005), who in turn reported to Acacia rigidula as the character¬

istic species of this community as well as species such as Acacia berlandieri and Havardia pollens. For the pres¬

ent study, A. rigidula was also abundant, but species like A. berlandieri and H. pollens were not. The most com¬

mon species was Cordia boissieri, which according to Garcia and Jurado (2008) is characteristic of semithorn

shrubland communities. High abundance of Leucophyllumfrutescens is also registered. This association Aca-

cia-Leucophyllum-Cordia has been registered by Rojas (1965) as community characteristic of the Coastal Plain

of Northeast Mexico; Briones and Villarreal (2001) reported similar associations in an ecotone between the

provinces of the Altiplano and Coastal Plain of northeast Mexico. This association is mainly due to humid

conditions where the development of these species is favorable. This humidity is carried by winds from the sea

of the Gulf of Mexico and is intercepted in the foothills of the Sierra Madre Oriental at the altitude of the com¬

munity studied (850 m a.s.l.; Briones & Villarreal 2001).

The most representative family of the piedmont scrub community was Fabaceae, with 9 species. This

family has been reported as the most representative in the region (Garcia & Jurado 2008; Canizales et al. 2009;

Estrada et al. 2012; Mata et al. 2014).. The most abundant species recorded in this study was A. rigidula; this

agrees with Canizales et al. (2009) who recorded A. rigidula as the most abundant species. This study differs

from that found by Garcia and Jurado (2008) and Estrada et al. (2012), who studied an area with non-distur¬

bance appearance with a high presence of Helietta parvifolia. In contrast, Mata et al (2014) reported a high

abundance of A. lecheguilla, an element that is not recognized as abundant within this community because the

area sampled was altered by anthropogenic activities.

The most representative family of the rosetophyllous desert shrubland was Cactaceae with 16 species.

The most abundant and frequent species was A. lecheguilla, referred by Alanis et al (1996), Rzedowski (2006),

and Mata et al. (2014) as the most characteristic species within of this community in the northeastern Mexico.

182 Journal of the Botanical Research Institute of Texas 9(1)

<2 2-4 4-6 6-8 8-10 >10

Relative Importance Value (%)

Fig. 2. Frequency distribution of RIV in the communities assessed: TT, tamaulipan thornscrub; SS, semithorn shrubland; PS, piedmont scrub; RDS,

rosetophyllous desert shrubland.

Shrubland

Fig. 3. Plant coverage of the communities assessed: TT, tamaulipan thornscrub; SS, semithorn shrubland; PS, piedmont scrub; RDS, rosetophyllous

desert shrubland.

Canizales et al., Shrublands adjacent to the Monterrey Metropolitan Area

183

Fig. 4. Dominance-diversity of species curve.

Relative importance Value (RIV). —Distribution of frequencies in the four regions showed that the number

of species with low ecological weight was greater than the number of species with high ecological weight. This

was most evident in the RDS community which presented 35 species with an importance value index < 2%,

which indicated a high diversity in this community where there is no high dominance of any large number of

species. The species that stands out with respect to its RIV was A. lecheguilla (40.59%), a result consistent with

that found for a community on the border between the states of Nuevo Leon and Coahuila (Mata et al. 2014)

which recorded as having the highest value of RIV for this species. Following this community is SS and TT with

21 and 16 species, respectively. Cordia boissieri had the highest RIV within the TT and SS communities (19.25%

and 10.37% respectively). However, other studies have reported A. rigidula as a species with a high value of

importance (Estrada et al. 2004; Gonzalez et al. 2010). Furthermore, Alanis et al. (2008) reported A. rigidula as

the most prevalent species in areas with a history of agriculture, livestock, and clear cutting. Jimenez et al.

(2009) also reported A. rigidula as the species of greatest ecological value for a clear cutting area in Linares,

while Mora et al. (2013) recorded Diospyros texana , Prosopis laevigata, and A.farnesiana with the highest RIVI

values for TT, in livestock areas, regeneration areas, and reference areas, respectively. Moreover, Jimenez et al.

(2009, 2012) noted a case of D. texana growing in an area with a history of agriculture. The high presence of A.

rigidula in areas with a history of agricultural use is because it is identified as an important component of the

diet of small grazing ruminants (Ramirez 2009). In the present study, the TT community did not have a his¬

tory of such use.

The lowest number of species was found in the piedmont scrub community (6 species). Nevertheless, it is

the best represented category which is consistent with studies reporting 3 species with the highest RIV values

(Canizales et al. 2009; Mata et al. 2014) as in the present work. However, the species were different between

studies, coinciding only in the genus Acacia, which can be due to a different usage history in the presented ar¬

eas. Meanwhile, Garcia and Jurado (2008) reported Helietta parvifolia as a species with the highest RIV

(52.14%) which was supported by the work of Estrada et al. (2012). This information differs from that found by

the present study because this species was not found in the sampled community, and piedmont scrub com¬

munities have undergone structural changes over time, which has brought a change in composition and struc¬

ture as noted by Canizales et al. (2009). These authors reported a low abundance attributed to the history of

184

Journal of the Botanical Research Institute of Texas 9(1)

extensive use which has reduced their populations to small groups in isolated communities which are largely

undisturbed (Garcia & Jurado 2008; Canizales et al. 2009).

Plant coverage .—The greatest coverage was recorded in the piedmont scrub community (17,267.78 m 2 /ha),

which was higher than that found by Canizales et al. (2009) at 9,724.27 m 2 /ha. Also, Mata et al. (2014) reported

the greatest coverage for a community of PS along the border of Nuevo Leon and Coahuila, while for the rose-

tophyllous desert shrubland the same authors reported a lower coverage (1,716 m 2 /ha) than that of the present

study. The semithorn shrubland (SS) had the lowest coverage (8,138.43 m 2 /ha) of the communities studied

composed predominantly of low-growing species such as Leucophyllumfrutescens and Croton torreyanus. Fol¬

lowing the SS community was the rosetophyllous desert shrubland (8,710.46 m 2 /ha) where predominate life

forms are rosettes and succulents.

The community with the second highest coverage was Tamaulipan thornscrub (TT), with 13,718.68 m 2 /

ha. Lower coverage for areas of shrubland, regeneration, and livestock in the southern Nuevo Leon were re¬

corded by Mora et al. (2013).

Distribution of abundance of species .—Distribution of species with respect to their proportional abun¬

dance demonstrated that species follow a normal distribution, since many species have intermediate values of

either very low or very high abundance (Martella et al. 2012), where these species are distributed towards the

tails of the distribution curve (Aguirre et al. 2008). This type of distribution is reported in northeastern Mexico

in plant communities at mature successional stage (Molina et al. 2014).

In conclusion, the results from this research lead us to note that there are constant changes in the species

composition of these plant communities. Therefore it is important to develop strategies to conserve these eco¬

systems as well as develop a sustainable management program.

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Journal of the Botanical Research Institute of Texas 9(1)

BOOK NOTICE

Mohamed A.F. Noor. 2012. You’re Hired! Now What? A Guide for New Science Faculty. (ISBN-13: 978-0-

87893-963-3, pbk). Sinauer Associates, Inc., Publishers, 23 Plumtree Road, P.O. Box 407, Sunderland,

Massachusetts 01375-0407, U.S.A. (Orders: www.sinauer.com, 1-413-549-4300). $14.41 US, 96 pp., 1

illustration, 6" x 9".

From the publisher: You’re Hired! Now What? is a short, conversationally written handbook describing chal¬

lenges and specific strategies to get ahead (or at least get by) in an academic setting. New college faculty are

well-versed in the scientific skills they’ll need for success in research, including design of projects, preparation

of manuscripts and grant proposals, and interactions during peer review. Yet typically they receive no training

in organization, management, or even basic college structure. This book is an attempt to begin to fill this void,

presenting thoughts and advice intended as a starting point for thinking about problems faced by new faculty.

Table of Contents:

Preface

Acknowledgments

SETTING UP: Your First Few Months (and Beyond)

ORGANIZING A RESEARCH TEAM

SUPERVISING PEOPLE: A General Overview

KEEPING UP: Communicating with Your Team

TEACHING CLASSES

SERVICE: The Time Sink

INTERACTIONS

STAYING COMPETITIVE: Building Your CV and Experience

KEEPING UP WITH THE “REST OF LIFE”

APPENDIX 1. Rules of the Laboratory

APPENDIX 2. Authorship Policy

APPENDIX 3. Graduate Student Expectations Handout

APPENDIX 4. Hypothetical One-on-One Graduate Student Weekly Meeting Agenda

APPENDIX 5. Laboratory Projects List

J.Bot. Res. Inst. Texas 9(1): 186.2015

MATELEA CHIHUAHUENSIS (APOCYNACEAE): AN ADDITION TO THE FLORA

OF THE UNITED STATES AND A SYNOPSIS OF THE SPECIES

Angela McDonnell & Mark Fishbein

Department of Botany

Oklahoma State University

301 Physical Sciences

Stillwater, Oklahoma, 74078, U.S.A.

angela.mcdonnell@okstate.edu

Meg Quinn & Trevor Hare

Borderlands Restoration L3C

2718 E Croyden St.

Tucson, Arizona, 85716, U.S.A.

Kevin Keith

P.O.Box 53

Gila, New Mexico, 88038, U.S.A.

ABSTRACT

M atelea chihuahuensis is reported as an addition to the floras of New Mexico and the United States with three occurrences in close proximity

documented from Hidalgo County. A full description of the species is presented, along with photos, a distribution map, and a review of its

taxonomic history. A lectotype is designated. M atelea chihuahuensis is compared to its close relatives and a key is provided to aid in distin¬

guishing similar M atelea species in Chihuahua.

RESUMEN

M atelea chihuahuensis se reporta como una adicion a las floras de Nuevo Mexico y Estados Unidos con tres ocurrencias en las proximidades

de las documentadas en el condado de Hidalgo. Se presenta una descripcion completa de la especie, junto con fotos, un mapa de la distribu-

cion, y una revision de su historia taxonomica. Se designa un lectotipo. M atelea chihuahuensis se compara con sus parientes cercanos y se

proporciona una clave para ayudar a distinguir las especies M atelea similares en Chihuahua.

Key Words: Apocynaceae, M atelea, Milkweed, North American Flora, Noteworthy Collection

INTRODUCTION

The Sky Island region of the southwestern United States and northwestern Mexico is a biologically diverse area

that is an ongoing source of botanical novelties. The Madrean ‘islands’ describe an archipelago of mountains

clad with evergreen oak and coniferous forests, interspersed with valleys of grasslands and scrub, and which

connect the Sierra Madre Occidental and the Rocky Mountain cordilleras (Warshall 1994). Some species with

widespread ranges in Mexico, in the Sierra Madre Occidental or beyond, have long been known to occur in the

U.S. from only one or a few sites in southeastern Arizona, e.g., Henry a insularis Nees (Acanthaceae; Daniel

2004), Cynanchum ligulatum (Benth.) Woodson (Apocynaceae; Sundell 1993), Dichondra sericea Sw.

(Convolvulaceae; Austin 2006), Coursetia glabella (A. Gray) Lavin (Fabaceae; Lavin 1988), and Quercus viminea

Trel. (Fagaceae; Van Devender et al. 2013). Among these are species that have never been re-collected following

their initial discovery in the U.S., e.g., Pherotrichis schaffneri A. Gray (Apocynaceae) and Diphysa thurberi (A.

Gray) Rydb. ex Standi. (Fabaceae) (Kearney & Peebles 1961). Such discoveries continue in the region: Mandev-

illafoliosa (Mull. Arg.) Hemsl. (Apocynaceae; Milsom & Chamberland 2013) and Echinochloa oplismenoides (E.

Fourn.) Hitchc. (Poaceae; Fishbein 1995) were reported as new to the United States and Eriogonum terrenatum

Reveal (Polygonaceae; Reveal 2004) was described as a new species. Similar to those discoveries in Arizona,

the smaller part of the Sky Island region in southwestern New Mexico recently has seen the discovery of a spe¬

cies new to the U.S., Grindelia oxylepis Greene (Asteraceae; Spellenberg 2014), and another new to science, Eu¬

phorbia rayturneri V.W. Steinm. & Jercinovic (Euphorbiaceae; Steinmann & Jercinovic 2013). To this list of

novelties in the New Mexico portion of the Sky Island region we add Matelea chihuahuensis (A. Gray) Woodson

J. Bot. Res. Inst. Texas 9(1): 187 -194.2015

188

Journal of the Botanical Research Institute of Texas 9(1)

(Apocynaceae). To our knowledge, there is no common name for this species, but we recommend using Chi-

huahuan hairy milkvine.

Matelea Aubl. is a genus of approximately 225 species in Apocynaceae subfamily Asclepiadoideae. While

vines are the predominant growth form in Matelea, other growth forms occur as apparent adaptations to grass¬

land and woodland environments. In the southern United States and northern Mexico, several Matelea species

are prostrate to decumbent, non-twining herbs. These species appear to be most closely related to Pherotrichis

Decne., a genus segregated from Matelea and characterized by an erect, non-twining habit, as well as several

species of microphyllous to moderately small-leaved vines, such as M. parvifolia (Torr.) Woodson and M. pro-

ducta (Torr.) Woodson (Krings et al. 2008; Parks 2008). Together, this group of ~30 arid-adapted species cor¬

responds loosely with Woodson’s (1941) Matelea subg. Chthamalia (Decne.) Woodson, which is under revi¬

sionary study by A. McDonnell (unpubl. data), and which includes the poorly known species Matelea chihua-

huensis (Fig. 1).

The type specimen of Matelea chihuahuensis was collected by Cyrus Guernsey Pringle during the early

part of the period during which he completed his most well known work: exploration and collection of plant

specimens throughout Mexico from 1885 to 1909 (Davis 1936; Knobloch 1979; Stafleu & Cowan 1983; Mauz

2011). The specimens were divided into numerous sets distributed to the major herbaria of the time, including

one sent to Harvard University. There, Asa Gray described many new species from Pringle’s specimens. Gray

described M. chihuahuensis as a species of Gonolobus Michx., where nearly all North American milkweed vines

of subtribe Gonolobinae were placed at the time. As syntypes of G. chihuahuensis A. Gray, he cited Pringle’s

collections #104 (with flowers, 1885) and #692 (with flowers and fruits, 22 August 1885) gathered from the

hills around Chihuahua. In his journal, Pringle describes the circumstances surrounding the collection of

#692. In the prior days, he had returned from the Santa Eulalia Mountains to collect in “the valley toward

Nombre de Dios” and the nearby cliffs (Davis 1936, p. 26). His #692 was collected “out around the railroad

shops” and must be from the area now part of the east side of Chihuahua City that has engulfed Nombre de

Dios, the site of caves that are presently a tourist attraction. A freight railroad still parallels Federal Highway 45

in this area.

Gray described Gonolobus chihuahuensis as procumbent, spreading, hairy on all sides, and with many

stems and leaves. He noted that the new species shared these characters with two other prostrate milkweeds

known to him, Mateleaparviflora (Torr.) Woodson (of Texas) andM. pedunculata (Decne.) Woodson (of Micho-

acan and adjoining states). He highlighted the floral differences between this species and all others known at

the time: “the crown [corona] is perfectly simple, with no internal crests or appendages, the margin abruptly

produced into the five narrow and entire lobes which overtop the anthers, these of the firm texture which is

common in the Chthamalia section, of which this species has wholly the aspect.” (Gray 1886). Thus, he hypoth¬

esized a close alliance with Chthamalia Decne., (Decaisne 1844). However, Gray did not recognize Chthamalia

as a genus distinct from Gonolobus (Engelmann & Gray 1845; McDonnell 2014).

When Woodson (1941) revised the circumscription of Matelea, reducing many formerly recognized gen¬

era to subgenera, he did not place M. chihuahuensis in Matelea subg. Chthamalia, but rather in Matelea subg.

Pherotrichis (Decne.) Woodson. Based on his key to the subgenera (1941), subg. Chthamalia, is distinct in hav¬

ing “subquadrate to oblong-reniform pollinia with a narrow hyaline margin” while subg. Pherotrichis has “fal¬

ciform or arcuate pollinia with no hyaline margin.” In contrast to the pollinia characteristics highlighted by

Woodson, the vegetative morphology, flowers, and fruits of M. chihuahuensis accord much more closely with

the prostrate species placed by Woodson in subgenus Chthamalia than with the erect herbs he placed in subge¬

nus Pherotrichis. As noted above, phylogenetic studies have shown that species in these subgenera are close

relatives (Krings et al. 2008; Parks 2008). Further study is required to revise generic circumscriptions and de¬

finitively place M. chihuahuensis.

The species of subg. Chthamalia that share the prostrate to ascending habit of Matelea chihuahuensis are

distributed from the southern U.S. to south-central Mexico. The most widespread species are M. biflora (Nutt,

ex Raf.) Woodson and M. cynanchoides (Engelm. & A. Gray) Woodson of the south-central U.S. (McDonnell

McDonnell et al., Synopsis of Matelea chihuahuensis and a new record for the U.S.A.

189

Fig. 1. Matelea chihuahuensis A. Habit, scale = 5 cm. B. Inflorescence, scale = 1 mm. C. Follicle, scale = 1 cm. D. Detail of flower, with 2 petals removed,

scale = 0.5 mm.

2014) and M. pedunculata of southwestern Mexico. The remaining species in the subgenus have narrow ranges,

including three species that until now were considered to be endemic to Chihuahua, Mexico. These are M.

chihuahuensis, M. lesueurii (Standi.) Woodson, and M. wootonii (Vail) Woodson, each of which is documented

by a small number of collections. Many of these collections have remained unidentified, occasionally others

have been misidentihed, and in the latter situation it is not unusual for the specimens to be identified as M.

chihuahuensis. For example, a vegetative specimen from Laguna Babicora ( Lebgue & Estrada 3206a [NMC]) was

collected and cited as M. chihuahuensis (Estrada-Castillon et al. 1997) but is actually M. lesueurii (see also Mc¬

Donnell 221 [OKLA] from the same locality). We provide here the first detailed morphological description of M.

chihuahuensis and document this species as a new member of the flora of the United States. We also provide a

key to distinguish this species from similar species in Chihuahua.

MATERIALS AND METHODS

Locality information was downloaded from online specimen repositories (including the Global Biodiversity

Information Facility: GBIF; www.gbif.org, TROPICOS: tropicos.org, and SEINet: http://swbiodiversity.org/

portal/index.php) and/or obtained directly from specimen labels. Specimen loans and images were obtained

from the Botanical Research Institute of Texas (BRIT), the California Academy of Sciences (CAS), Michigan

State University (MSC), the Missouri Botanical Garden (MO), the New York Botanical Garden (NY), Harvard

University (GH/HUH), the Rancho Santa Ana Botanic Garden (RSA/POM), University of Texas at Austin

(TEX/LL), the Smithsonian Institution (US), andJSTOR Global Plants (plants.jstor.org/). Herbarium abbrevia¬

tions follow Thiers (2014). Occurrence data were georeferenced using Google Earth (build 7.1.2.2041, 7 Oct

2013). New specimens were obtained through fieldwork in the states of New Mexico and Chihuahua. A distri¬

bution map was produced using ArcGIS® ArcMap™ v.10.2 (Esri, Redlands, CA) and Adobe® Illustrator CS4

190

Journal of the Botanical Research Institute of Texas 9(1)

(Adobe Systems Inc., San Jose, CA). Due to imprecise locality data of some older specimens, two of the 14 re¬

cords for M. chihuahuensis were excluded from the map. Measurements of floral and vegetative characters were

made from herbarium specimens as well as from fresh and liquid-preserved flowers using Olympus® cellSens

Entry 1.6 imaging software and an Olympus® SZX10 dissecting microscope outfitted with an Olympus®

SC30 CMOS color camera. Measurements of digital specimen images were made using ImageJ software 1.48v

(Rasband 1997-2014).

RESULTS AND DISCUSSION

Previous collections of Matelea chihuahuensis document a sparse but widespread distribution in Chihuahua

(Fig. 2). It has been collected several times in the vicinity of Chihuahua City ( Pringle 104, 692,880; Palmer 214;

Frey tag & Baxter 6; LeSueur 1182). It has also been found further west and northwest of the city in the foothills

of the Sierra Madre Occidental, near San Diego del Monte ( Hartman 735), and Colonia Juarez ( Jones s.n.; Pennell

19072). One collection was made at a disjunct site in the main cordillera of the Sierra Madre, at Mesa de Basase-

achic ( LeSueur 2009). This location is considered suspect given the more eastern and seemingly more arid

habitats of all other collection sites. The species has also been documented at disjunct locations in desert

mountains in far northeastern Chihuahua, south of Piramide (Johnston 8104) and in the Sierra Hechiceros

(Stewart 220). Fieldwork conducted by A. McDonnell and M. Fishbein of Oklahoma State University and S.

Gonzalez Elizondo and F. Ruacho Gonzalez of Centro Interdisciplinario de Investigacion para el Desarollo

Integral Regional Unidad Durango, Instituto Politecnico Nacional, was successful in locating a new Chihua-

huan population in the foothills of an eastern branch of the Sierra Madre, in the Sierra Fa Catarina ( McDonnell

211). It is noteworthy that M. lesueurii also inhabits these mountains, though at higher elevation and in more

mesic grasslands (McDonnell 203 [CIIDIR, MO, OKFA]).

Our research shows that, until now, Matelea chihuahuensis has not been documented outside of the state

of Chihuahua (Vega-Mares et al. 2014). However, during vegetation surveys in the fall of 2013, three new loca¬

tions were found in close proximity (< 8 km) of each other on the Diamond A Ranch (formerly Gray Ranch), in

the bootheel of New Mexico (Quinn s.n.). These plants were either non-reproductive or in fruit. Photos and a

specimen including viable seeds were sent to M. Fishbein and A. McDonnell for identification. Plants grown

from seed in January 2014 flowered approximately five months after germination and were positively identified

as Matelea chihuahuensis (McDonnell 199). The individuals from New Mexico represent the northernmost and

westernmost stations for this species and are additions to the floras of New Mexico and the United States.

Though there are relatively few collections of this species, we believe it to be most common on rocky sub¬

strates in drier grasslands and pastures in foothills of the eastern slope of the Sierra Madre Occidental. In

Chihuahua, we found Matelea chihuahuensis (McDonnell 211) growing in the Sierra Fa Catarina on a moderate,

rocky slope in grassland dominated by Bouteloua spp. with scattered Cylindropuntia imbricata (Haw.) F.M.

Knuth, Dasylirion wheeleri S. Watson ex Rothr., Fouquieiria splendens Engelm., Aloysia gratissima (Gillies &

Hook.) Tronc., and Mimosa biuncifera Benth. We were not able to locate historically documented populations at

several sites we visited in Chihuahua, including one northeast of the Sierra Fa Catarina (Yen & Estrada 7224)

where the vegetation was described as “matorral microhlo” on the specimen label. The vegetation at this site

was dominated by Vachellia vernicosa (Britton & Rose) Seigler & Ebinger, Prosopis glandulosa Torr., Mimosa

biuncifera, and Aloysia gratissima, suggesting the species occurs at least occasionally in desertscrub. The New

Mexican populations were found in Bouteloua- dominated grassland, in sandy loam overlying clay, on the west¬

ern piedmont of the Animas Mountains.

Matelea chihuahuensis perennates from a thickened taproot and its phenology is likely correlated with

patterns of rainfall. Most flowering specimens have been collected in July and August following the onset of

summer rains. Fruiting specimens have been collected between the months of July and October. Pollinators or

other fauna that might interact with M. chihuahuensis are unknown.

Closely Related Species

Occasionally, collectors have confused closely related species for Matelea chihuahuensis. Prostrate species of

McDonnell et al., Synopsis of Matelea chihuahuensis and a new record for the U.S.A.

191

Fig. 2. Distribution of Matelea chihuahuensis in Chihuahua, Mexico, and New Mexico, USA. Collection sites illustrated with black triangles.

Matelea have a similar overall appearance, including multiple stems and cordate to orbicular leaves. However,

the species are difficult to confuse in flower—M. chihuahuensis is especially distinct with cream and green-

striate corollas and pilose, terete, translucent-white corona segments that overtop the anthers (vs. purplish to

maroon corollas with white to maroon coronas). Matelea chihuahuensis can be distinguished from similar

Chihuahuan species by herbage that is densely hirsute throughout and leaves with undulate margins that

provide a ruffled appearance to the blades. Matelea lesueurii differs by having suborbicular leaves that are gla¬

brous and shiny adaxially and densely hirsute abaxially, the margins entire and planar. Matelea wootonii also

192

Journal of the Botanical Research Institute of Texas 9(1)

has suborbicular leaves with glabrous adaxial surfaces, sparsely hirsute abaxial surfaces and entire, planar

margins. These species can be distinguished with the key presented below. All three species are found in the

state of Chihuahua; M. lesueurii and M. wootonii generally grow at higher elevations and in more mesic habitats

than does M. chihuahuensis.

Other prostrate or procumbent species similar to Matelea chihuahuensis occur outside of Chihuahua. In

southern Texas, Coahuila, and Tamaulipas, two to three species occur: M. brevicoronata (B.L. Rob.) Woodson,

M. aff. brevicoronata, and M. parviflora. To the south of Chihuahua, Matelea aff. pedunculata (Decne.) Woodson

occurs in Durango and Zacatecas. All of these taxa share with M. chihuahuensis the dense vestiture of tri-

chomes and undulate leaf margins. Vegetatively, these species are difficult to distinguish, but in flower they are

rather distinct. While M. chihuahuensis has a green and white corolla, both M. brevicoronata and M. parviflora

have more or less uniformly bright green corollas with little or no white in them and M. aff. brevicoronata has a

green corolla when young that becomes maroon with age. At maturity, the corolla lobes of M. parviflora are

reflexed and in M. brevicoronata they are campanulate. In both M. chihuahuensis and M. aff. brevicoronata, the

corollas are rotate, but the species differ in color as noted above. The species that occur in Texas all have strap¬

shaped corona segments that vary in length but are not translucent and do not arch over the anthers as they do

in M. chihuahuensis. One specimen collected 8 miles south of Mirando City in Webb County, Texas ( Turner

80-61 [TEX]), was originally identified as M. chihuahuensis, but has been confirmed to be M. parviflora. M. aff.

pedunculata differs conspicuously from M. chihuahuensis by having tubular-campanulate corollas that are

muted green-brown with maroon striations. Its corona is much shorter, yellowish in color, and does not over¬

top the anthers.

TAXONOMIC TREATMENT

Matelea chihuahuensis (A. Gray) Woodson, Ann. Missouri Bot. Gard. 28:232. 1941. Gonolobus chihuahuensis A.

Gray. Proc. Amer. Acad. Arts 12:398-399.1886. Type: MEXICO. Chihuahua: rocky hills near Chihuahua, 22 Aug 1885, C.G. Pringle

692 (lectotype, designated here, GH; isolectotypes, AC digital image, BR digital image, CAS, CORD digital image, E digital image, F,

G, 2 sheets-digital images, GOET digital image, JE digital image, MICH digital image, NY, 3 sheets, RSA digital image, US, 2 sheets).

Plants: perennial, prostrate to procumbent, malodorous and hispid-hirsute throughout, usually with 5-20+

stems from a thickened taproot, stem length in flower 10-50 cm, lengthening in fruit. Leaves: largest with

petioles 0.2-1.6 cm long, blades deltoid to cordate, occasionally ovate, 0.7-3.3 cm long and 0.7-2.6 cm wide,

bases deeply to shallowly cordate, apices acute, margins undulate; leaf bases with 2-4 narrowly conical col-

leters that may break off on mature leaves. Inflorescences: axillary umbels of 2-5 flowers; peduncles 1-3 cm

long. Flowers: pedicels 3.5-5.0 mm; calyx lobes 5, elliptic, 1.9-3.5 mm long; corolla aestivation contorted, after

expansion shallowly campanulate-rotate, cream to white with fine green striations on adaxial surfaces, green

and white with green striations on abaxial surfaces, 3.8-7.9 mm in diameter, deeply 5-lobed; the lobes oblong

to spathulate, hirsute at base adaxially, glabrous abaxially, margins planar; corona consisting of 5 basally-

united lobes, arising at the junction of the gynostegial column and the corolla, incurved, incumbent on apex of

anthers and style, strap-like, approximately elliptic in cross section, densely hirsute, translucent cream to

white; style apex white with pink striations visible through corona. Fruit: follicle, muricate, ovate to lanceolate

or elliptic with an acuminate apex, green with dull green striations, 4.94-5.7 cm long, 1.7-2.4 cm wide, protu¬

berances numerous, 0.06-1.6 mm in height.

Specimens examined.—MEXICO. CHIHUAHUA. Mpio. de Buenaventura: Carretera Sueco-Flores Magon, 5-6 km, 7Jul 1997, C. Yen&E.

Estrada 7224 (BRIT); Sierra La Catarina, east slope, about 14 km SW of San Buenaventura, 7 Aug 2014, A. McDonnell 211 (CIIDIR, MO,

OKLA). Mpio. de Casas Grandes: plains near San Diego, ~5 mi S of Casas Grandes, 4 Aug 1891, C.V. Hartman 735 (GH); Colonia Juarez, 11

Sep 1903, M .E. Jones s.n. (POM); Colonia Juarez, 21 Sept 1934, F.W. Pennell 19072 (US). Mpio. de Chihuahua: Chihuahua, 1885, C.G. Pringle

104 (GH); vicinity of Chihuahua, May 1908, E. Palmer 214 (US); Ray Brown’s Canyon, Aug 1936, H. LeSueur 1182 (F); arroyos 5 mi W of

Chihuahua city on Hwy 260, 23 Jul 1975, G.E. Freytag & J.W. Baxter 6 (GH, MO, US). Mpio. de Manuel Benavides: Sierra Hechiceros, 9 km

S of Rancho Hechiceros, near the Coahuilan boundary, 1 Oct 1940, R.M. Stewart 220 (GH). Mpio. de Ocampo: Mesa de Basaseachic, 7 Jul

1936, H. EeSueur 2009 (GH). Mpio. de Ojinaga: 7.5 mi S of Piramide, 11 Aug 1941, 1.M. Johnston 8104 (GH, LL). U.S.A. NEW MEXICO. Hi¬

dalgo Co.: Gray Ranch (now Diamond A Ranch), rangeland monitoring site plot #5413, Cloverdale Valley, 13 Oct 2013, M. Quinn s.n.

(OKLA); 11 Jul 2014 A. McDonnell 199 (OKLA, grown from Quinn s.n. seed).

McDonnell et al., Synopsis of Matelea chihuahuensis and a new record for the U.S.A.

193

KEY TO THE PROSTRATE SPECIES OF MATELEA FOUND IN THE STATE OF CHIHUAHUA, MEXICO

1. Leaves deltoid to cordate, adaxial surface densely hirsute, margins undulate; corolla lobes white with green striations;

follicles densely hirsute with 4 or more protuberances per 4 cm length_ M. chihuahuensis

1. Leaves broadly ovate to orbicular, adaxial surface glabrous, margins planar; corolla lobes purple or maroon; follicles very

sparsely hirsute with 2 or fewer protuberances per 4 cm length.

2. Leaves densely hirsute abaxially; margins eciliate_ M. lesueurii

2. Leaves hirsute only along veins abaxially; margins ciliate_ M. wootonii

ACKNOWLEDGMENTS

Many thanks are due to Dra. Socorro Gonzalez Elizondo and Lizeth Ruacho Gonzalez, M.S., of Centro Inter-

disciplinario de Investigacion para el Desarollo Integral Regional Unidad Durango, Institute Politecnico Na-

cional, for their camaraderie and held assistance in Chihuahua. We also acknowledge and thank the curators

and staff of the following herbaria for their assistance in facilitating specimen loans and granting access to

high-resolution digital images: AC, ARIZ, ASU, BR, BRIT, CAS, CIIDIR, CORD, E, F, G, GH, GOET, JE, MICH,

MSC, MO, NMC, NY, GH/HUH, RSA/POM, TEX/LL, and US. Thank you to Richard Spellenberg and Eric

Sundell, who each provided comments and suggestions that improved the manuscript. This research was

funded in part by graduate student research awards from the Society of Herbarium Curators, the American

Society of Plant Taxonomists, and the Botanical Society of America made to A. McDonnell.

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Estrada-Castillon, E., R. Spellenberg, &T. Lebgue. 1997. Flora vascular de la Laguna de Babicora, Chihuahua, Mexico. Sida

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THE FIRST NATURALIZED OCCURRENCE OF THE GENUS FORSYTHIA

(OLEACEAE) IN ARKANSAS (U.S.A.), WITH ADDITIONAL NOTEWORTHY

ANGIOSPERM RECORDS FOR THE STATE

Brett E. Serviss, Tyler L. Childs, Sydney S. Grant, Tiffany A. Graves,

Ethan Holicer, Seth A. McBroom, and Logan Thomas

Department of Biology

Henderson State University

Arkadelphia, Arkansas 71999-0001, U.S.A.

servisb@hsu.edu

James H. Peck

Allen Leible

P.O.Box 705

Cedar Key, Florida 32625, U.S.A.

jhpeck@ualr.edu

Simonson Biological Field Station

Henderson State University

100 Simonson Road

Bismarck, Arkansas 71929, U.S.A.

Ieiblea@hsu.edu

ABSTRACT

Two species of Forsythia: F. suspensa (Thunb.) Vahl and F. viridissima Tindl., are reported as new for Arkansas. These records represent the

first voucher specimen-based documentation of the genus Forsythia in the state’s flora outside of cultivation. Two additional species of non¬

native angiosperms: Rhodotypos scandens (Thunb.) Makino and Frachelospermumjasminoides (Tindl.) Tern., are documented for only their

second occurrences in the state.

RESUMEN

Se citan dos especies de Forsythia: F. suspensa (Thunb.) Vahl y F. viridissima Lindl., como nuevas para Arkansas. Estas citas representan la

primera documentacion basada en especimenes testigo del genero Forsythia en la flora del estado fuera de cultivo. Se documentan dos espe¬

cies adicionales de angiospermas no nativas: Rhodotypos scandens (Thunb.) Makino y Frachelospermumjasminoides (Lindl.) Lem., como se-

gundas ocurrencias en el estado.

INTRODUCTION

Forsythia is a genus of about 11 species of deciduous or semi-evergreen shrubs distributed primarily across

Asia, with one species in Europe (Chang et al. 1996). Species of Forsythia are characterized by opposite, simple

(or rarely ternately compound) leaves with entire or coarsely serrate-toothed margins, perfect (bisexual) yel¬

low flowers, often with orange or red-colored nectar guides in the throat that are arranged in solitary or few-

flowered axillary inflorescences, and fruits that are small, flattened, few-seeded capsules (Chang et al. 1996).

Many species are important as ornamentals in temperate climates for their colorful and profuse display of

flowers in early spring, cold hardiness, and ease of cultivation (Bailey and Bailey 1976; Chang et al. 1996).

Two morphologically similar species: F. suspensa (Thunb.) Vahl (weeping forsythia) and F. viridissima

Lindl. (greenstem forsythia), are frequently cultivated in Arkansas, and one or both species have previously

been attributed to the Arkansas flora by various sources (Hardin 1974; Thompson 1977; Arkansas Vascular

Flora Committee 2006). Forsythia suspensa was included in the Checklist of the Vascular Plants of Arkansas (Ar¬

kansas Vascular Flora Committee 2006), although this record was apparently based on cultivated material

(Smith 1988; Gentry et al. 2013). Both species, however, were ultimately excluded from the state’s flora because

no voucher specimens were available to document the actual occurrence of naturalized plants (Gentry et al.

2013). Consequently, our discovery of F. suspensa and F. viridissima represents the first verifiable and docu¬

mented occurrence of these species, along with the genus Forsythia, outside of cultivation in Arkansas (Gentry

J. Bot. Res. Inst. Texas 9(1): 195 -199.2015

196

Journal of the Botanical Research Institute of Texas 9(1)

et al. 2013). Both species have previously been documented outside of cultivation in the U.S. in several other

states (NRCS 2014).

In Arkansas, naturalization and establishment of Forsythia species appears to be accomplished primarily

vegetatively via air layering of stems, where stems root when in contact with the ground; whole colonies of

plants may be produced in this manner. It is important to note, however, that both of the aforementioned spe¬

cies have also been observed in the fruiting condition in Arkansas.

The combination of opposite, coarsely toothed leaves, quadrangular stems with prominent, conical¬

shaped buds, yellow flowers that are produced in early spring prior to or simultaneously with leaf emergence,

and small, flattened, longitudinally furrowed capsules aid in distinguishing species of Forsythia from other

shrubs in the Arkansas flora.

ADDITIONS TO THE ARKANSAS FTORA

Forsythia suspensa (Thunb.) Vahl (Oleaceae), weeping forsythia (Fig. 1). Forsythia suspensa is a deciduous

shrub to about 3 m tall that is native to China (Chang et al. 1996). It is naturalized in several states in the east¬

ern U.S. and has also been documented from Kansas, Montana, Utah, and Washington (USDA, NRCS 2014).

Some forms of this species, in addition to having simple, unlobed leaves, may exhibit some leaves that are lobed

or ternately compound (Krussmann 1978; Chang et al. 1996).

Voucher specimens: ARKANSAS. Garland Co.: a single plant at edge of woods, Entergy Park, Lake Catherine, 16 Sep 2006, Peck 06-437

(HEND). Pulaski Co.: A few plants scattered in disturbed woods, not cultivated, Boyle Park, Little Rock, 11 Mar 2003, Peck 03011 (HEND).

Forsythia viridissima Lindl. (Oleaceae), greenstem forsythia (Fig. 2). Forsythia viridissima is a deciduous to

semi-evergreen shrub to about 3 m tall that is native to China (Chang et al. 1996). It is naturalized in several

states in the eastern U.S. and has also been documented from Kansas (USDA, NRCS 2014).

Voucher specimens: ARKANSAS. Clark Co.: colony of several plants along roadside on slope at edge of disturbed woods, 12th St. ca. one

block S of Mill Creek, Arkadelphia, 34.1302778, -093.0611111, 4 Apr 2013, Serviss 7930 (HEND). Hot Spring Co.: Small colony of plants

spreading asexually via air layering of stems and possibly also seed at edge of disturbed, upland, mixed-pine hardwood, adjacent to historic

Beaton Creek Road, ca. 500 m NW of the Henderson State University Simonson Biological Field Station, Bismarck, 34.2972222, -093.2675,

11 Dec 2014, Serviss 8160 (HEND).

KEY TO ARKANSAS SPECIES OF FORSYTHIA

Forsythia suspensa and F. viridissima are morphologically similar and easily confused with one another; how¬

ever, they may be distinguished using the following key (see Fig. 3 for internode sections showing hollow ver¬

sus lamellate pith).

1. Internodes internally hollow; leaves ovate to ovate-elliptic, margins serrate along most to nearly all of the leaf_ F. suspensa

1. Internodes internally lamellate; leaves oblong-elliptic, obovate-elliptic, or lanceolate, margins generally only serrate on

the distal one-third to one-half of the leaf, otherwise entire_ F. viridissima

SECOND OCCURRENCES IN THE ARKANSAS FLORA

Rhodotypos scandens (Thunb.) Makino (Rosaceae), black jetbead. Rhodotypos scandens is a scandent, decidu¬

ous shrub that is native to China, Japan, and Korea (Krussmann 1978; Li et al. 2003). It is naturalized in sev¬

eral eastern states, mostly in the north, but also documented in the Southeast from Alabama and South Caro¬

lina (USDA, NRCS 2014).

Rhodotypos scandens has the ability to asexually spread via air layering when its stems come in contact

with moist substrate; hence, both seed production and/or vegetative reproduction may establish naturalized

populations.

Vegetatively, in Arkansas, R. scandens closely resembles the rare native shrub, Neviusia alabamensis Gray

(Alabama snow wreath); however, it may be distinguished from N. alabamensis by its opposite leaves, solitary

flowers with four, large petals, and black fruits. Neviusia alabamensis, contrastingly, has alternate leaves, flowers

that lack petals and are generally clustered in groups of 3 to 8, and fruits that are whitish-tan to brown in color.

Serviss et al., Forsythia and other new floristic records for Arkansas

197

Fig. 1 . Photographs of Forsythia suspensa (from cultivated plants). A. Leaves. B. Stem and buds. C. Bark. D. Flowers. E. Young fruit.

Rhodotypos scandens has only previously been documented in Arkansas from Washington County

Voucher specimen: ARKANSAS. Garland Co.: several large and small shrubs along edge of small stream in open, semi-disturbed woods

along riparian zone, reproductive with mature fruits, Strauss Road, ca. 1.3 mi S of highway Arkansas 88, Spring Hill Camp area, 30 Oct 2014,

Serviss 8215 (HEND).

Trachelospermum jasminoides (Lindl.) Lem. (Apocynaceae), star jasmine. Trachelospermum jasminoides is

an aggressive, twining, evergreen liana to 10 m that is native to China (Bailey & Bailey 1976; Li et al. 1995). It

has been previously documented outside of cultivation in a few southeastern states, including Florida, Louisi¬

ana, and Texas (USDA, NRCS 2014).

Trachelospermum jasminoides is commonly cultivated for its ability to rapidly form a thick, evergreen

groundcover, tolerate full sun to nearly full shade conditions, and profuse production of fragrant flowers (Bai¬

ley and Bailey 1976). Stems root vigorously when exposed to bare soil or climbing substrate, such as trees or

fences. It is also tolerant of dry conditions once established. The above combination of characteristics facilitates

naturalization of T. jasminoides into novel habitats. In Arkansas, naturalized plants of T. jasminoides will form

a dense carpet of trailing stems in the understory, but will also readily climb trees via twining and/or adventi¬

tious roots. When climbing, leaf morphology changes from the smaller, thinner leaves found on trailing stems

to the larger, thicker leaves characteristically produced on climbing stems.

Trachelospermum jasminoides will readily produce fruits in Arkansas and spontaneous seedlings are

sometimes encountered in the vicinity of reproductive-age plants. The seeds of T. jasminoides are distinctive:

cinnamon-brown in color with a conspicuous coma (tuft of hairs) attached to the seed body. All parts of the

plant are poisonous.

Trachelospermum jasminoides may potentially be confused with the morphologically similar, native liana,

198

Journal of the Botanical Research Institute of Texas 9(1)

Fig. 2. Photographs of Forsythia viridissima (from cultivated plants). A. Leaves. B. Stem and buds. C. Bark. D. Flowers. E. Seeds and dehisced capsule.

F. Mature fruit.

Gelsemium sempervirens Ait. (yellow jasmine, Carolina jasmine); however, it may be distinguished from G.

sempervirens by its white to pale yellow flowers, elongate follicles, pubescent (when young) leaves and stems,

and white, milky latex that is exuded when any vegetative portion of the plant is cut or damaged. In contrast,

G. sempervirens has bright lemon-yellow flowers, short, elliptical capsules, foliage that is completely glabrous,

and lacks milky latex.

Trachelospermumjasminoides has only previously been documented in Arkansas from Drew County.

Voucher specimen: ARKANSAS. Clark Co.: large expanse of plants covering ground and climbing trees, edge of stream and adjacent slope

in disturbed woods of riparian zone, plants also spreading to edge of woods, plants sterile without reproductive structures, Mill Creek adja¬

cent to the intersection of Cypress Road and Twin Rivers Drive, 34.1250000, -093.0875000, Arkadelphia, 4 Sep 2014, Serviss 8176 (HEND).

ACKNOWLEDGMENTS

We sincerely thank Theo Witsell (Arkansas Natural Heritage Commission—ANHC) and one anonymous re¬

viewer for their helpful comments and suggestions regarding this paper. We also thank the Henderson State

University Biology Department and the University of Arkansas at Little Rock Biology Department for support¬

ing this work.

REFERENCES

Arkansas Vascular Flora Committee. 2006. Checklist of the vascular plants of Arkansas. Arkansas Vascular Flora Committee.

Fayetteville, Arkansas, U.S.A.

Serviss et al., Forsythia and other new floristic records for Arkansas

199

Fig. 3. Photographs of pith sections of F suspensa and F viridissima for comparison (from cultivated plants). A. Hollow pith of F suspensa. B. Lamellate

pith of F viridissima.

Bailey, L.H. & E.Z. Bailey. 1976. Hortus Third. A concise dictionary of plants cultivated in the United States and Canada.

Vol. 2. MacMillan, New York, U.S.A.

Chang, M., L. Chiu, Z. Wei, & P.S. Green. 1996. Forsythia. In: Flora of China Editorial Committee, eds. Flora of China. Vol. 15

(Myrsinaceae through Loganiaceae). Science Press, Beijing, China, and Missouri Botanical Garden Press, St. Louis,

Missouri, U.S.A. Pp. 279-280.

Gentry, J.L., G.P. Johnson, B.T. Baker, C.T. Witsell, & J.D. Ogle. 2013. Atlas of the vascular plants of Arkansas. University of

Arkansas Herbarium. Fayetteville, Arkansas, U.S.A.

Hardin, J.W. 1974. Studies of the southeastern United States flora. IV. Oleaceae. Sida 5:274-285.

KrOssmann, G. 1978. Manual of cultivated broad-leaved trees and shrubs. Vols. 2 and 3. Timber Press. Portland, Oregon,

U.S.A.

Li, B., A. Leeuwenberg, & DJ. Middleton. 1995. Trachelospermum. In: Flora of China Editorial Committee, eds. Flora of China.

Vol. 16 (Gentianaceae through Boraginaceae). Science Press, Beijing, China, and Missouri Botanical Garden Press, St.

Louis, Missouri, U.S.A. Pp. 166-168.

Li, C., H. Ikeda, & H. Ohba. 2003. Rhodotypos. In: Flora of China Editorial Committee, eds. Flora of China. Vol. 9 (Pittospro-

raceae through Connaraceae). Science Press, Beijing, China, and Missouri Botanical Garden Press, St. Louis, Missouri,

U.S.A. P. 192.

Smith, E.B. 1988. An atlas and annotated list of the vascular plants of Arkansas, second edition. E.B. Smith. Fayetteville,

Arkansas, U.S.A.

Thompson, R.L. 1977. The vascular flora of Lost Valley, Newton County, Arkansas. Castanea 42:61 -94.

USDA, NRCS. 2014. The PLANTS Database (http://plants.usda.gov). National Plant Data Team, Greensboro, NC 27401-

4901 U.S.A. Accessed 20 Nov 2014.

200

Journal of the Botanical Research Institute of Texas 9(1)

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Errata

J.Bot. Res. Inst. Texas 9(1): 200.2015

NEPHROLEPIS CORDIFOLIA (NEPHROLEPIDACEAE) NATURALIZED IN

SOUTHERN CALIFORNIA (U.S.A.): WITH NOTES ON

UNINTENDED CONSEQUENCES OF ESCAPED GARDEN PLANTS

Richard E. Riefner, Jr.

Alan R. Smith

Research Associate, Rancho Santa Ana Botanic Garden

1500 North College Avenue

Claremont, California 91711-3157, U.S.A.

rriefner@earthlink. net

University Herbarium, University of California

1001 Valley Life Science Building #2465

Berkeley, California 94720-2465, U.S.A.

arsmith@berkeley. edu

ABSTRACT

Nephrolepis cordijolia occurs in many tropical, subtropical, mild-temperate, and Mediterranean regions of the world. Because it is widely

cultivated, and there are many cultivars, populations have escaped from gardens and naturalized; these escapes have likely obscured its

ancestral native distribution. Nephrolepis cordijolia is also frequently cultivated in southern California, especially for groundcover since it

adapts well to a variety of soil types, tolerates full sun, and is drought-tolerant. We report here that Nephrolepis cordijolia grows outside of

cultivation on concrete walls, trunks of palm trees, calcareous cliffs, and an ocean bluff in Tos Angeles and Orange counties, coastal south¬

ern California. A naturalized population documented previously in 2006 from a streamside habitat in San Diego County was not relocated

in 2014. Documentation of ornamental plants that have escaped cultivation, especially in urban habitats, is an important but often over¬

looked component of establishing time lag phases and early pathways of dispersal for potentially invasive plant species introduced to new

regions. We provide notes on identification of N. cordijolia, its native and introduced ranges, garden plants and invasive weed introductions,

and discussion of naturalized habitats in southern California. We also cite voucher specimens and photographic documentation of the oc¬

currence of this species in southern California.

RESUMEN

Nephrolepis cordijolia ocurre en muchas regiones tropicales, subtropicales, semitempladas y mediterraneas del mundo. Debido a que esta

ampliamente cultivada, y hay muchas variedades, las poblaciones se han escapado de jardines y naturalizado; estos escapes probablemente

han oscurecido su distribucion nativa ancestral. Nephrolepis cordijolia tambien se cultiva con frecuencia en el sur de California, especial-

mente para cubierta vegetal ya que se adapta bien a una variedad de tipos de suelo, tolera pleno sol, y es tolerante a la sequia. Presentamos

aqui que Nephrolepis cordijolia crece fuera del cultivo en las paredes de hormigon, troncos de palmeras, acantilados calcareos, y un acanti-

lado al mar en los condados de Los Angeles y Orange, en el sur de la California costera. Una poblacion naturalizada documentada previa-

mente en 2006 a partir de un habitat ribereno en el condado de San Diego no fue localizada en 2014. La documentacion de plantas ornamen-

tales que se han escapado de cultivo, sobre todo en habitats urbanos, es importante pero a menudo se pasa por alto el componente de estab-

lecimiento de fases de retraso en el tiempo y principios de las rutas de dispersion de las especies de plantas potencialmente invasoras intro-

ducidas en nuevas regiones. Proporcionamos notas sobre la identificacion de N. cordijolia, sus rangos nativos e introducidos, las plantas de

jardin y las introducciones de malezas invasoras, y la discusion de los habitats naturalizados en el sur de California. Tambien citamos espe-

cimenes testigo y la documentacion fotografica de la presencia de esta especie en el sur de California.

Keywords: Nephrolepis cordijolia, California, ferns, invasive garden plants, non-native plants, ornamental horticulture

INTRODUCTION

Nephrolepis (variously placed in Lomariopsidaceae or Nephrolepidaceae), a genus of ferns found mostly in

tropical and subtropical regions, comprises 19 species and three varieties based on the treatment by Hoven-

kamp and Miyamoto (2005). It is a distinctive and an easily recognized genus, which is identified primarily by

its wiry stolons, medial to often supramedial or submarginal soral position and shape, linear one-pinnate

fronds, and articulate pinnae (Mickel & Smith 2004). Nephrolepis is sometimes placed in its own family,

Nephrolepidaceae, or in the Lomariopsidaceae (Smith et al. 2006), but historically was included in Davallia-

ceae or Dryopteridaceae (e.g., Mickel & Smith 2004; Hovenkamp & Miyamoto 2005).

Species of Nephrolepis have been cultivated since the early 20th century and many remain popular today

(Hoshizaki & Moran 2001). Tough and easy to grow, they are the most widely cultivated of all ferns (Brenzel

2007). The most notable is the “Boston fern,” which originated in the late 1800s or early 1900s from a mutant

J. Bot. Res. Inst. Texas 9(1): 201 - 212.2015

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Journal of the Botanical Research Institute of Texas 9(1)

of N. exaltata (L.) Schott and thereafter propagated widely for its attractive, arching fronds (Benedict 1916;

Bailey 1924; Code 1996). Today, it is frequently grown in homes and gardens and is a commercially important

plant (Diggs & Lipscomb 2014). Nearly 80 named varieties and cultivars of N. exaltata are available from the

horticultural industry (Brenzel 2007; BGCI 2014a).

Nephrolepis cordifolia (L.) C. Presl (southern sword fern, tuberous sword fern, fishbone fern, herringbone

fern, ladder fern, or narrow sword fern) is also widely cultivated, and is grown frequently in pots, hanging

baskets, and as a ground cover (Coile 1996; Brenzel 2007). Over 25 cultivars are known and many are grown

in arid regions (Brenzel 2007; BGCI 2014b). In southern California, N. cordifolia is often grown for ground-

cover since it adapts well to a variety of soil types, tolerates full sun, is drought tolerant, and spreads rapidly in

moist garden conditions (Rushing 2006; SDFS 2014).

Nephrolepis cordifolia is terrestrial, epiphytic, or epilithic (on rock) in moist to wet shady places, rain for¬

ests, coastal shrublands, wetland and riparian habitats, on epiphyte perches, particularly palm trunks, lime¬

stone ledges, cliff and rock habitats, urban areas, and around old home sites, roadsides, or waste places in

Florida, Mexico, West Indies, Central America, South America, Africa, Southeast Asia, the Pacific Islands and

Hawaii, Australia, and New Zealand (Nauman 1993; Bell 1998; Hovenkamp & Miyamoto 2005; NZPCN 2013;

ISSG 2014). Because N. cordifolia is frequently cultivated, its nativity is often questionable. Nephrolepis cordifolia

is said to be “indigenous” (i.e., native) to Hawaii (Valier 1995; Palmer 2003; SI 2014), but this is suspect because

the species was not included in the comprehensive fern flora of Hawaii compiled by Hillebrand (1888; see also

Hovenkamp & Miyamoto 2005), and it also grows in portions of northern Australia, but it is widely regarded

as naturalized in the Sydney, Melbourne, and Perth regions (Bell 1998; Weeds of Australia 2014). Nephrolepis

cordifolia has also naturalized on the southern Cook Islands (McCormack 2007) and on Moorea, French Poly¬

nesia (Murdock & Smith 2003). Nephrolepis cordifolia is apparently not native to Florida (FLEPPC 2014), New

Zealand (NZPCN 2013; ISSG 2014), Africa (Crouch et al. 2011), temperate and tropical Asia (ISSG 2014; USDA,

GRIN 2014), and elsewhere (Weber 2003). Hoshizaki and Moran (2001), however, gave the native range of N.

cordifolia as tropical America, Africa, Asia, Australia, Japan, and New Zealand.

Nephrolepis cordifolia is also probably not native to Mexico or Central America because many specimens

cited in the literature are from gardens, confused with closely related species, and/or plants may have natural¬

ized at some localities (Mickel & Smith 2004). Therefore, N. cordifolia may be native only in the central portion

of its known range in the Americas, i.e., Greater Antilles and Venezuela (Hovenkamp & Miyamoto 2005). A

review of Old World specimens at UC (by A.R. Smith, December 2014) indicates the oldest collections come

from Asia and the Pacific Region, including: New Caledonia, 1870; China, Hainan, 1878; India, 1891; Australia,

1900; Java, 1904; New Zealand, 1904; Philippines, 1907; and Madagascar, 1915, but not the Neotropics. Ac¬

cordingly, the nativity of N. cordifolia throughout its known range, and the location of escaped garden plants

and date of introductions, which has contributed to this confusion, is in need of critical review.

In the continental United States and its territories, N. cordifolia is reported for Alabama, Florida, Georgia,

Hawaii, Puerto Rico, and the Virgin Islands (USDA, NRCS 2014). Nephrolepis cordifolia was not reported for

California in The Jepson Manual, Second Edition, or online by the Jepson Flora Project (Baldwin et al. 2012;

Jepson Flora Project 2014). Nephrolepis cordifolia also was not included in other California publications that

identify non-native species growing spontaneously outside of cultivation, including Hrusa et al. (2002), Rob¬

erts et al. (2004), Rebman and Simpson (2006), Clarke et al. (2007), DiTomaso and Healy (2007), Dean et al.

(2008), and Roberts (2008). The Consortium of California Herbaria (CCH 2014) posts several collections from

cultivated sources; however, it also cites one naturalized population (with tubers) for San Diego County ( Cain

et al 134, SD). This population was first documented in 2006 from a streamside habitat at Chollas Lake Com¬

munity Park (CCH 2014). It was not relocated by R.E.R in 2014.

Nephrolepis cordifolia is reported here as naturalized, but uncommon, on calcareous cliffs and an ocean

bluff in native plant communities, and it is epiphytic on the trunks of palm trees and grows in joints of rough-

grouted masonry or concrete walls in urban areas in coastal southern California. At the cited localities, it ap¬

pears to propagate, either through spores, tubers, or stolons, without the aid of human intervention and inten-

Riefner and Smith, Nephrolepis cordifolia naturalized in southern California

203

tional summer-watering. It is to be expected elsewhere in similar habitats at other coastal localities. We pro¬

vide notes on identification, native and introduced range, garden weeds and invasive plant introductions, and

discussion of naturalized habitats for this species in southern California, as well as cite voucher specimens and

photographic documentation.

Vouchers: U.S.A.: CALIFORNIA: Los Angeles Co.: City of Long Beach, N of E 2nd St. and W of Marine Stadium Park, Alamitos Bay, near

docks on Bay Shore Ave. at E Broadway St., 33 o 45'35.6400"N, 118°07 , 43.3609"W, elev. ca. 4 m, local on palm tree trunk, 27Jul 2014, Riefner

14-258 (CAS, CDA, RSA, UC, UCR); City of Long Beach, N of E 2nd St. and W of Marine Stadium Park, Alamitos Bay, near docks on Bay Shore

Ave. between Monroe Ave. and E Vista St., 33°45 , 37.0260"N, 118 o 07 , 30.2406"W, elev. ca. 6 m, local on palm tree trunks, 27Jul 2014, Riefner

14-261 (RSA, UC, UCR). Orange Co.: City of Laguna Beach, N side of North Coast Highway, N 0.25 mi from Jasmine St., 33 0 32'42.1764"N,

117°47 , 28.0201"W, elev. ca. 20 m, rare, crevice in cinder block wall along sidewalk, spreading from adjacent cultivated sources, 14 May 2010,

Riefner 10-35 (UC); City of Laguna Beach, Aliso Beach County Park, bluff ca. 0.15 mi SE of Aliso Way, NW ca. 0.25 mi of West St.,

33 o 30'28.079"N, 117°45 , 12.265"W, elev. ca. 10 m, uncommon, seepage area on ocean bluff beyond the salt-spray zone, 25 Aug 2011, Riefner

11-120 (RSA, UC); City of Yorba Linda, southbound 91-Freeway at 55-Freeway interchange ca. 0.6 mi W of Lakeview Ave., 33°55 , 33.9528"N,

117°49 , 33.1985"W (approximate), elev. ca. 80 m, locally common, expansion joints in north-facing rough-grouted concrete wall, 5 Apr 2013,

Riefner 13-59 (CAS, UC); City of Dana Point, W ca. 0.1 mi from Island Way along Dana Point Harbor Dr., cliff slope immediately S of Santa

Clara Ave. and Amber Lantern St., 33°27 , 48.3192"N, 117°42 , 05.4726"W, elev. ca. 3-11 m, scattered clumps on cliff, in crevices and on and

beneath ledges, 24 Feb 2014, Riefner 14-64 (CDA, RSA, UC, UCR); City of Newport Beach, Cliff Drive Park, along Cliff Dr. near Beacon St.,

33 0 37'13.4472"N, 117°55 , 26.8840"W, elev. ca. 44 m, large clump growing on palm tree trunk, 14 Jun 2014, Riefner 14-171 (CAS, CDA, RSA,

UC, UCR). San Diego Co.: City of Lemon Grove, Chollas Lake Community Park, E of Hidden Cove, 32°738N, 117°0637W, elev. ca. 139 m,

south-facing slight slope near stream, tubers present, 27 Mar 2006, Cain et al. 134 (SD; CCH 2014).

Identification

Nephrolepis cordifolia is a tuft-forming plant with erect to spreading rhizomes and stolons that produce long

runners, with or without 1-3 cm wide scaly tubers. The upright evergreen fronds are once pinnate and grow

30-120 cm tall and 3-7 cm wide. Fronds often arch, producing a graceful habit pleasing to fern enthusiasts and

gardeners.

With fertile mature fronds, most species of Nephrolepis are relatively easy to identify. The separation of

closely-related taxa, however, which include N. pectinata (Willd.) Schott, N. pendula (Raddi) J. Sm., and N. un-

dulata (Afzel. ex Sw.) J. Sm., can be problematic. Identification of N. cordifolia versus N. exaltata may also prove

troublesome. With very limited sampling, Hennequin et al. (2010) found that four samples of N. cordifolia did

not form a clade, with the sole sample of N. exaltata being sister to only one of the samples of N. cordifolia and

one sample of N. cordifolia falling well outside the other three; however, two of their samples, including the

most disparate one, were both cultivated specimens, obtained from GenBank (vouchers not seen by the au¬

thors). These preliminary results suggest that further sampling of both native and cultivated plants needs to be

done before there is a clear understanding of species circumscriptions and relationships. Distinguishing fea¬

tures that serve to separate N. cordifolia from similar taxa include: conspicuously auricled basiscopic bases of

pinnae; pinnae with blunt to rounded apices; dense scaly, pale rachis indument with bicolored scales (pale with

dark point of attachment); indusia that are reniform to lunate; and the common presence of scaly tubers (some¬

times absent; Coile 1996; Mickel & Smith 2004; Hovenkamp & Miyamoto 2005). Important characters are

shown in Figs. 1 & 2.

Southern California Naturalized Populations

On coastal cliffs (Fig. 3) and an ocean bluff, N. cordifolia grows on calcareous substrates outside the salt-spray

zone, but within the influence of moisture-laden ocean fogs and often seepage areas. Laboratory analysis (con¬

ducted by Wallace Labs, El Segundo, California) indicates these substrates are slightly to moderately alkaline

(7.4-7.9 pH) and high in lime (calcium carbonate). Another non-native fern that favors mesic calcareous out¬

crops and old masonry, Cyrtomium falcatum (L.f.) C. Presl (holly fern), grows on outcrops with, or in close

proximity to, N. cordifolia at the Dana Point and Laguna Beach locations.

In urban areas, N. cordifolia is epiphytic on the trunks of palm trees (Fig. 4), where moist ocean breezes

provide a year-round source of moisture. Ficus rubiginosa Desf. ex Vent, (rusty fig, Port Jackson fig), native to

eastern Australia and also widely cultivated, was reported recently for California by Dean et al. (2008). It co¬

occurs with N. cordifolia at several locations, including calcareous outcrops and cliff habitats, and it is also epi-

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Journal of the Botanical Research Institute of Texas 9(1)

Fig. 1. Abaxial surface of a frond showing indusiate sori and auriculate overlapping base of pinnae typical of N. cordifolia. Photograph was taken at Dana

Point population. Orange County, California.

phytic on the trunks of palm trees (Fig. 5). Metropolitan areas are particularly vulnerable to alien species intro¬

ductions, which serve as entry pathways for invasions from the urban environment to native ecosystems (van

Ham 2013). Nephrolepis cordifolia also grows in the joints of old masonry and concrete walls (Fig. 6), appar¬

ently where urban water runoff provides deep wetting within the concrete joints; this enables establishment

and survival of the species in arid southern California.

The naturalized population of N. cordifolia documented previously in 2006 from a streamside habitat in

San Diego County, although not relocated in 2014, may indicate that moist-soil or shady riparian situations

could represent suitable habitat for the species elsewhere in southern California. Therefore, additional popula¬

tions of N. cordifolia will likely be identified at new localities on calcareous outcrops and cliffs, concrete walls

and masonry, trunks of palm trees, and in alkaline-soil riparian communities in coastal southern California.

The presence or absence of tubers has long been a topic of interest for this species. They appear to be func¬

tioning for water storage and not for starch (Morton 1958). With the exception of rock and concrete crevices,

some plants within each of the naturalized populations of N. cordifolia documented herein support tubers,

which can be locally abundant on underground and aerial runners. New plantlets can form from tubers and

water reserves can be used during drought (Hovenkamp & Miyamoto 2005). Accordingly, tuber presence may

play an important role for the establishment and persistence of N. cordifolia when growing outside of cultiva¬

tion in summer-dry southern California.

Escaped Garden Plants and Invasive Weed Status

Escaped horticultural plants are an important source of invasive species introductions (Mack & Lonsdale

Riefner and Smith, Nephrolepis cordifolia naturalized in southern California

205

Fig. 2. Nephrolepis cordifolia rhizomes, stolons, and scaly tubers, and adaxial frond surface with pale indument along the rachis. This photograph was

taken of plants growing on the trunk of a palm tree in Long Beach, Los Angeles County, California.

2001; Reichard & White 2001; Mack & Erneberg 2002). Of the 117 non-native plants reported for California by

Dean et al. (2008), 49 of these species are available in the horticultural trade, were documented spreading from

home sites, and/or are intentionally cultivated, and another eight species were first documented in California

as contaminants in nurseries. Garden plant introductions are also an important source of naturalized weeds in

Australia; of the 2,779 introduced plant species established in the Australian environment, approximately

1,831 (66%) represent escaped garden plants (Groves at al. 2005). In Europe, it is estimated that 80% of the

currently known invasive alien plants were initially introduced as ornamental or agricultural plants (Hulme

2007). In fact, invasive garden plants comprise over half (56%) of the 36 land and aquatic plants included on

the list of the 100 World’s Worst Invasive Alien Species (Lowe et al. 2000; Groves et al. 2005).

Nephrolepis cordifolia is a known threat to native plant species. In Florida, it can form dense stands and by

aggressive spreading, it can displace native vegetation (Langeland et al. 2008; ISSG 2014). Accordingly, it is

listed as a ‘Category I’ invasive plant (FLEPPC 2014). In New Zealand, it is listed as an environmental weed

(Howell 2008), and is included on the ‘National Pest Plant Accord’ preventing the sale, cultivation, and distri¬

bution of the plant (NZPCN 2013; NPPA 2014). Nephrolepis cordifolia is also regarded as an environmental

weed outside its native range in Australia, declared noxious in New South Wales, and prohibited from entry

into Western Australia (Groves et al. 2005; SOWN 2014; Weeds of Australia 2014).

Invasive plants are always a concern when they become established in the natural ecosystems of new re¬

gions. It is widely recognized that there is often a lengthy lag phase between the time when a species becomes

naturalized and represents an innocuous introduction, to when it may become highly invasive and a pest in

206

Journal of the Botanical Research Institute of Texas 9(1)

Fig. 3. The calcareous sandstone habitat of Dana Point, Orange County, California. Red arrow points to one of the clumps of N. cordifolia discovered on

these cliffs and the inset shows a close-up of the population observed during the severe drought of 2014.

native ecosystems. These transitional stages (beginning with problematic garden weeds or urban waifs that

sometimes become pest plants in native ecosystems) have usually proved to be unpredictable and unexpected

(EBGC 2014).

To date, preliminary surveys indicate that N. cordifolia is an uncommon garden escape in southern Cali¬

fornia. However, observations by gardeners may provide insights to its potential problematic behavior. In

2005, a gardener in Acton, Los Angeles County, California, posted comments online indicating that N. cordifo-

Riefner and Smith, Nephrolepis cordifolia naturalized in southern California

207

Fig 4. Photograph of N. cordifolia that has successfully established and is persistent on the trunk of palm trees in Long Beach, near Alamitos Bay, Los

Angeles County, California.

208

Journal of the Botanical Research Institute of Texas 9(1)

Fig. 5. Ficusrubiginosa epiphytic on a palm tree trunk growing with N. cordifolia. Maritime breezes off Alamitos Bay, Long Beach, California, likely provide

the moisture needed for each of these species to successfully establish and persist in urban environments.

lia is “very hardy here in hot, dry southern California ... and quite invasive. ... It is grown very commonly

down here and sold all over as an indoor as well as an outdoor fern. ... Spreads like wild fire” (Dave’s Garden

2014). We do not know if N. cordifolia will become a pest plant in California, but it is expected to expand its

range and establish elsewhere because of high propagule pressure associated with widespread cultivation. Ri¬

parian communities in watersheds affected by perennial urban runoff may be particularly vulnerable to inva¬

sions. Once established, seasonal flood scour may wash fragments of stolons or tubers downstream to colonize

new habitats.

Ancestral Range, Cultivar Introductions, and Additional Research

The ancestral native range of N. cordifolia is obscure, in part because numerous cultivars have been developed

that are widely grown in gardens and escape. These escapes have expanded the known range of the species, but

have also led to ongoing confusion regarding nativity of the species in many regions (Weber 2003; Mickel &

Smith 2004; Hovenkamp & Miyamoto 2005). The distribution and native versus non-native habitat occur¬

rences documented in Australia are a good case-in-point. Nephrolepis cordifolia is thought to be native to parts

of northern Australia, i.e., coastal eastern Queensland and parts of northeastern New South Wales, but it is

regarded as naturalized in some habitats and areas outside its natural distribution in these districts (Bell 1998;

Weeds of Australia 2014). It has also naturalized in Victoria, coastal southwestern Western Australia, Norfolk

Island, and other areas beyond its native range such as the coastal districts of central New South Wales (Bell

1998; Weeds of Australia 2014). Although presumably the same species, the native Australian form of N. cordi¬

folia is not invasive like the non-native plants thought to have been imported from Asia (SOWN 2014).

Riefner and Smith, Nephrolepis cordifolia naturalized in southern California

209

Fig. 6. Nephrolepis cordifolia grows in the joints of old masonry and concrete walls in areas that receive urban water runoff. Photograph taken along

Pacific Coast Highway in Laguna Beach, Orange County, California.

In New Zealand, Hovenkamp and Miyamoto (2005) note the confusion between native occurrences ver¬

sus introduced forms of N. cordifolia, but treated the native forms within the broad circumscription of N. cordi¬

folia sens. str. De Lange et al. (2005), however, reinstated N. flexuosa Colenso, native to Fiji, Norfolk Island,

Raoul Island, Lord Howe Island, and New Zealand, to help protect the indigenous populations while identify¬

ing N. cordifolia as the widespread naturalized and invasive plant. Nephrolepis flexuosa differs by its graceful

stature, more narrow straight-sided fronds, is teraploid with larger spores, and lacks tubers, but may be diffi¬

cult to separate from depauperate forms of N. cordifolia, which usually lack tubers (De Lange et al. 2005). Hov¬

enkamp and Miyamoto (2005) coin the name N. cordifolia var. pseudolauterbachii Hovenkamp & Miyam. for

seemingly very similar, tuberless plants from Fiji and Samoa. Detailed morphological, molecular, and cyto-

logical studies and intensive surveys of escaped cultivars and naturally occurring populations are needed to

resolve these issues.

CONCLUSIONS

Species beginning to escape cultivation often receive little attention by botanists, largely because of the diffi¬

culty in identifying them before they are documented and published in local or regional floras. A time lag in

plant invasions, which is a phase when plants are difficult to find and identify during early dispersal, is an im¬

portant part of a typical invasion scenario (Schierenbeck et al. 2007). Often overlooked, documentation of

garden plants escaping in urban habitats could provide important information that can be used to track the

early dispersal and naturalization of these plants in new regions. In the case of Ficus rubiginosa, careful docu-

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Journal of the Botanical Research Institute of Texas 9(1)

mentation of its early detection on brick and stone walls and on other trees in urban areas of Auckland, New

Zealand, provided important information regarding its establishment and spread in the region (Gardner &

Early 1996); F. rubiginosa is often associated with N. cordifolia in southern California. Early detection is an im¬

portant and fundamental management objective when dealing with invasive plant species (Rejmanek 2000).

Accordingly, continuing documentation of either species could provide important information regarding early

detection, establishment, stepping-stone urban habitats, and frequency of occurrences in native ecosystems

needed to identify time lag phases of potentially invasive species in southern California.

Ornamental horticulture has unintentionally promoted the introduction of potentially invasive species to

new regions (Dehnen-Schmutz & Touza 2008). The selection and cultivation of easy-to-grow, drought hardy,

salt tolerant, soil- and climate-adapted, and self-sowing cultivars may facilitate dispersal and establishment of

non-native species inadvertently escaping from gardens to native ecosystems, which may start an invasion

process (Mack 2000; Anderson et al. 2006). In the case of N. cordifolia, its introduction and naturalization in

non-indigenous regions has had unintended consequences, including serious threats to native biodiversity,

and this process has also compounded confusion regarding its true ancestral range and taxonomic circum¬

scription.

Note. —After the manuscript was accepted for publication, we identified a population of Nephrolepis cordifolia

(Riefner 15-10F, UC) naturalized around a natural spring and in riparian woodlands at the La Cristianita His¬

torical Baptismal Site, Camp Pendleton, San Diego County, California. Previously, it was identified as N. exal-

tata (initial collection made by K. Stockwell in 1999) on the Consortium of California Herbaria database with

additional comments that it was “presumably planted and persisting” at the site. Although we do not know if

this population was intentionally planted, it is clearly naturalized in 2015.

ACKNOWLEDGMENTS

Our thanks to Michael Sundue (The Pringle Herbarium, University of Vermont) and an anonymous reviewer

for providing helpful comments and suggestions that greatly improved the manuscript. We also thank Gwen

Kenney (Environmental Security, US Marine Corps Base Camp Pendleton) for site access.

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BUDDLEJA DAVIDII (SCROPHULARIACEAE) NATURALIZED POPULATIONS

IN TENNESSEE (U.S.A.) AND ITS WOODY ASSOCIATES

Ralph L.Thompson

Katrina Rivers Thompson

Hancock Biological Station, Murray State University

Murray, Kentucky 40271, U.S.A.

Botanical Research Institute of Texas

1700 University Dr., Fort Worth, Texas 76107-3400, U.S.A.

Berea College Herbarium, Biology Program

Berea College, Berea, Kentucky 40404, U.S.A.

ralph_thompson@berea.edu

Child and Family Studies Program

Berea College

Berea, Kentucky 40404, U.S.A.

katrina_rivers_thompson@berea.edu

ABSTRACT

Buddleja davidii Franch. (orange-eye butterfly bush), an introduced Chinese shrub in the Scrophulariaceae, is categorically naturalized at

seven sites in four new Tennessee counties from field reconnaissance in 2012-2014. Buddleja davidii was previously documented by her¬

barium specimens as escaping from cultivation in Washington County (1952), Cumberland County (1975), and Hamilton County (2010).

Buddleja davidii has limited naturalization in Anderson, Davidson, DeKalb, and Knox counties in ruderal habitats from field observations

and relative abundance values. In Tennessee, orange-eye butterfly bush was confined to culturally-disturbed habitats and was not present in

natural secondary forest communities. A total of 51 woody associated species (32 native, 19 non-native) were also documented among these

seven populations to obtain descriptive site data. Fifteen of 19 non-native woody associates were classified as state-listed invasive taxa,

which affect the colonization of B. davidii populations.

Key Words: Buddleja davidii, woody associated species, distribution, naturalized, orange-eye butterfly bush, Scrophulariaceae, invasives,

Tennessee counties

RESUMEN

Buddleja davidii Franch. (arbusto mariposa de ojos naranja), un arbusto chino introducido de Scrophulariaceae, esta naturalizado categori-

camente en siete sitios en cuatro nuevos condados de Tennessee a partir de estudios de campo realizados en 2012-2014. Buddleja davidii fue

previamente documentada por especimenes de herbario como escapada de cultivo en los condados de Washington (1952), Cumberland

(1975) y Hamilton (2010). Buddleja davidii tiene una naturalizacion limitada en los condados de Anderson, Davidson, DeKalb y Knox en base

a observaciones de campo y valores de abundancia relativa. En Tennessee, el arbusto mariposa de ojos naranja se limitaba a los habitats

perturbados por cultivo y no estaba presente en las comunidades forestales secundarias. Tambien se documentaron un total de 51 especies

asociadas lenosas (32 nativas, 19 no nativas) entre estas siete poblaciones para obtener datos descriptivos de los sitios. Quince de las 19 espe¬

cies asociadas lenosas no nativas fueron clasificadas como taxones invasivos registrados en el estado, que afectan a la colonization de las

poblaciones de B. davidii.

Palabras clave: Buddleja davidii, especies lenosas asociadas, distribucion, naturalizado, arbusto mariposa de ojos naranja, Scrophulariaceae,

invasivos, condados de Tennessee

INTRODUCTION

Buddleja davidii Franch. (orange-eye butterfly bush, butterfly bush, or summer lilac) are unarmed, multiple¬

stemmed, deciduous to semi-deciduous, shade-intolerant, pioneer shrubs placed in the Scrophulariaceae of the

Lamiales (Norman 2012). Orange-eye butterfly bush is native to 13 mountainous provinces in southwest and

central China (Zheng et al. 2006; Tallent-Halsell & Watt 2009). Buddleja is comprised of 90-100 species world¬

wide (Stuart 2006; Dirr 2009). Buddleja davidii has been the most extensively planted ornamental of the ca. 25

cultivated species. At least 150 cultivars and hybrids of B. davidii have been recognized (Stuart 2006; Dirr

2009). Buddleja davidii has been selected for ornamental horticulture due to its growth hardiness in many an¬

thropogenic habitats, attractive dark green and grayish-white foliage, fragrant white, lilac-lavender, lilac-rose,

red-violet to purple flowers, and high nectar content for the attraction of butterfly, moth, bee, and wasp pollina¬

tors (Stuart 2006; Dirr 2009; Tallent-Halsell & Watt 2009).

J. Bot. Res. Inst. Texas 9(1): 213 - 227.2015

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Journal of the Botanical Research Institute of Texas 9(1)

Buddleja davidii (herein, B. davidii, butterfly bush, or orange-eye butterfly bush), has a broad ecological

amplitude for propagation, establishment, colonization, and naturalization in a diversity of open, insolated and

weedy disturbed habitats through means of wind-dispersed seeds (Tallent-Halsell & Watt 2009; Trueblood

2009; Thompson & Abbott 2013). Butterfly bush is a fast growing, semi-cold-tolerant shrub that readily estab¬

lishes populations in full sun in a wide range of nutrient poor soil types (Tallent-Halsell & Watt 2009; True¬

blood 2009). Buddleja davidii thrives best in well-drained alkaline soils but regardless of adaptations, it is

rather a short-lived shrub mostly of less than 30 years (Smales 1990; Brunei 2006; Tallent-Halsell & Watt 2009;

Trueblood 2009). The presence of endophytic arbuscular mycorhizzae in B. davidii enhances its establishment

and survival (Camargo-Ricalde et al. 2003; Dickie et al. 2007).

In Kentucky, Buddleja davidii was reported naturalized in five counties from active and abandoned rail¬

road right of ways, weedy disturbed urban areas, roadside ditch thickets, lake floodplains, and severely-burned

national forest lands (Thompson & Abbott 2013). In 2014, butterfly bush was documented as naturalized for a

sixth Kentucky county distribution along a grassy road shoulder and guard rail in Leslie County within the

Cumberland Plateau [R.L.Thompson & K. Rivers Thompson 14-377 (BEREA, EKY)].

Our definitions of non-native “naturalized” species and “invasive” species are modified from Nesom

(2000) and Richardson et al. (2000) by subjective degree of colonization, establishment, and migration. A

“naturalized” plant is a non-native plant that grows naturally in ruderal or culturally-derived habitats, and is

able to maintain itself for at least 10 years without anthropogenic assistance. An “invasive” plant is an aggres¬

sive, non-native naturalized plant that tends to readily spread and displace the native flora and vegetation over

time. Thompson & Abbott (2013) described orange-eye butterfly bush as naturalized in Kentucky from held

reconnaissance, herbarium searches, and pertinent literature. Furthermore, Thompson and Abbott (2013)

determined that B. davidii should not be classified as an invasive species in Kentucky. Butterfly bush in six

Kentucky counties was closely restricted to those ruderal habitats where it was found with little evidence of

additional propagation, spread, or migration into contiguous natural vegetation.

Our major objectives were to conduct held reconnaissance searches for Tennessee populations of Buddle¬

ja davidii, pursue an in-depth herbarium specimen examination, review relevant literature, and compile de¬

scriptive data of the woody associated species at naturalized population sites.

BUTTERFLY BUSH NON-NATIVE STATUS

Regarding non-native origin status, Buddleja davidii has been classihed as naturalized in Australia, Canada,

Central America, Europe, New Zealand, Puerto Rico, South America, and the United States (Tallent-Halsell &

Watt 2009; Norman 2012). It has been recorded as an invasive shrub in Australia, England, France, New Zea¬

land, and in the USA states of Hawaii, Oregon, and Washington (Tallent-Halsell & Watt 2009; Young-Mathews

2011; Norman 2012; USDA, ARS 2014). In Oregon, B. davidii is designated as a Class “B” noxious weed; and, it

is listed as a Class “C” noxious weed along riparian floodplains in Washington (Young-Mathews 2011; USDA,

NRCS 2014). Buddleja davidii grows without cultivation in a variety of temperate habitats that include lime¬

stone quarries, coal surface-mined lands, active and abandoned railways, fallow helds, successional woodland

ecotones, roadside thickets and ditch margins, riparian corridors, streambeds, floodplains, and lake shores,

among numerous other disturbed habitats (Tallent-Halsell & Watt 2009; Norman 2012).

Orange-eye butterfly bush has been reported from 21 states (Kartesz 2014; USDA, NRCS 2014) to 24 states

in the United States (EDDMapS 2014). Distribution maps for these states do not clarify whether the non-native

status of B. davidii is cultivated, introduced, persisting, a waif, naturalized, or invasive among the non-native

classifications of Nesom (2000). Map distributions likely represent a grouping of all those non-native catego¬

ries from herbarium vouchers and literature reports.

Recent literature contains several discrepancies and inconsistencies on the invasiveness of butterfly bush

within the eastern United States. Buddleja davidii was not listed as an invasive species in AL, FL, GA, KY, MS,

NC, SC, nor TN by the SE-EPPC (2014). In contrast, the MA-EPPC (2008) reported butterfly bush as an inva¬

sive in CA, KY, NC, NJ, OR, PA, WA, and WV. Swearingen et al. (2010) listed it as a “Plant to Watch” in DE, MD,

Thompson and Thompson, Buddleja davidii in Tennessee

215

NJ, PA, VA, and WV. The KY-EPPC (2013) listed B. davidii for the first time, as a “Moderate Threat,” an exotic

plant that can spread into disturbed corridors although not readily invading natural areas. Butterfly bush has

been updated to an “Alert” rank for Tennessee, an exotic plant that possesses invasive characteristics in habi¬

tats similar to those found in Tennessee (TN-EPPC 2009). Jones and Wofford (2013) indicated B. davidii to be

a non-native escaped species of disturbed sites in Kentucky and Tennessee; and, it was expected to become

more problematic in the future. Chester et al. (2015) noted that B. davidii is “a landscaping species, rarely natu¬

ralized in ruderal areas statewide...shows strong invasive potential.”

Nevertheless, Trueblood (2009) developed an Invasive Species Assessment System data form and score

sheet for Buddleja davidii in North Carolina. She determined B. davidii to have limited ecological impact, distri¬

bution, and invasive potential in natural areas. In her assessment summary, orange-eye butterfly bush was

considered to be a non-invasive, shade-intolerant, short-lived pioneer shrub that tended to be controlled or

eliminated mainly through processes of natural plant succession by native woody vegetation. In brief, Buddleja

davidii has not been shown to have negative ecological impact in North Carolina natural areas.

METHODS

Field reconnaissance was conducted from 2012-2014 toward discovering naturalized populations of Buddleja

davidii in Tennessee. Descriptive data were also recorded for woody associated species for seven populations in

Knox, Anderson, DeKalb, and Davidson counties. “Associated woody species” are those characteristic or indi¬

cator taxa often found with certain other non-native and/or native species that have similar phenological life

cycles, plant durations, ecological adaptations, and habitat requirements.

Relative abundance values (frequency of occurrence) for orange-eye butterfly bush and interspersed

woody associates were based on the scale of Thompson (2007): Rare (R) = 1-4 individuals; Scarce (S) = 5-10

individuals; Infrequent (I) = 11-30 individuals; Occasional (O) = 31-100 individuals; Frequent (F) = 101-1000

individuals; and Abundant (A) = 1000s of individuals.

Relative frequency percentages were determined from absolute frequency by the presence of a native or

non-native woody associate within each Buddleja davidii population site (Table 1). Relative frequency is a

qualitative measure of the abundance and distribution of each associated species’ occurrence within the seven

B. davidii population sites.

Nomenclature, classification, and identification for associated woody taxa follow Jones and Wofford

(2013). Herbarium acronyms for voucher specimens follow Index Herbariorum of Thiers (2014). Herbarium

searches were made of the regional Tennessee university herbaria (APSC, ETSU, HTTU, MTSU, TENN, and

UCHT) and other state and national herbaria (BRIT, EKY, KY, MO, NCU, and VDB).

RESULTS AND DISCUSSION

Tennessee Butterfly Bush Herbarium Collections

From herbarium specimen label data, Buddleja davidii was first documented growing without cultivation in

Washington County on July 28, 1952 [R.L. James 16885 (TENN)]; and, it was also documented as an escape at

another site in Washington County in 1999 [Marie Dugger 21 (ETSU)]. In 1975, B. davidii was reported as es¬

caped at two locations in Cumberland County [B. Eugene Wofford 51846 (NCU, TENN); R. Krai 56489 (MO,

VDB)]. The Washington and Cumberland counties “escaped” populations are likely the basis of B. davidii being

classified as “naturalized” in A Fifth Checklist of Tennessee Vascular Flora by Chester et al. (2009). In Hamilton

County, a 2010 specimen was also labeled as “commonly escapes from cultivation” [ Gary H. Morton 9203

(UCHT)]. Orange-eye butterfly bush distribution was initially mapped for Tennessee in Cumberland, Knox,

and Washington counties (Chester et al. 1997; USDA, NRCS 2014), while Cumberland, Grainger, Knox, Shelby,

and Washington counties were mapped by TENN (2014) and EDDMapS (2014). Buddleja davidii has been

documented as a planted ornamental from herbarium specimens and held work in seven Tennessee counties

as listed: Blount (BEREA), Bradley (UCHT), Grainger (TENN), Knox (BEREA, TENN), Putnam (BEREA), Se¬

vier (TENN), and Shelby (TENN). Thirteen Tennessee counties have been documented by naturalized or

planted butterfly bush specimens (Fig. 1).

216

Journal of the Botanical Research Institute of Texas 9(1)

Table 1. Woody associates for seven Buddleja davidii populations in four Tennessee counties.

County

Species

Knox

Site 1

Knox

Site 2

Knox

Site 3

Ander.

Site 4

Ander.

Site 5

DeK.

Site 6

Dav.

Site 7

No.

Sites

Frequency

Acer negundo

100

**AHanthus oltissimo

100

**Ligustrum sinense

100

**Lonicera joponico

100

**Lonicera maackii

100

Parthenocissus quinquefolio

100

Plotonus occidental is

100

**Albizia julibrissin

85.71

**Pyrus calleryana

85.71

**Rosa multi flora

85.71

Rubus occidentalis

85.71

Toxicodendron radicans

85.71

**Ce!astrus orbiculatus

71.43

Cercis canadensis

71.43

Juniperus virginiana

71.43

Liriodendron tulipifera

71.43

**Paulownia tomentosa

71.43

Robinia pseudoacacia

71.43

Ulmus rubra

71.43

Vitis vulpina

71.43

Celtis occidentalis

57.14

**E!aeagnus umbellata

57.14

**Euonymus fortunei

57.14

Fraxinus americana

57.14

Salix nigra

57.14

Acer saccharum

42.86

Ampelopsis cordata

42.86

Campsis radicans

42.86

Fraxinus pennsylvanica

42.86

Rhus glabra

42.86

Rubus pensilvanicus

42.86

Acer saccharinum

28.57

Carya cordiformis

28.57

Clematis virginiana

28.57

Cornus amomum

28.57

Frangula caroliniana

28.57

Gleditsia triacanthos

28.57

**Morus alba

28.57

Quercus muhlenbergii

28.57

*U!mus pumila

28.57

Catalpa bignonioides

14.29

**Elaeagnus pungens

14.29

**Ligustrum vulgare

14.29

**Nandina domestica

14.29

*Prunus persica

14.29

**Pueraria montana

14.29

Rhus copallinum

14.29

Sambucus canadensis

14.29

Smilax rotundifolia

14.29

Ulmus americana

14.29

*Yucca filamentosa

14.29

Totals: 51 taxa

Ander. = Anderson County; DeK. = DeKalb County; Dav. = Davidson County.

(*) = a naturalized taxon, (**) = an invasive pest plant for Tennessee (TN-EPPC 2009).

The four new counties with naturalized butterfly bush are discussed separately. Tennessee specimen la¬

bel data on naturalized or planted shrubs from Tennessee counties are as follows:

Thompson and Thompson, Buddleja davidii in Tennessee

217

Fig. 1. Tennessee counties documented with Buddleja davidii specimens by collection year(s). Gray-numbered counties indicate naturalized popula¬

tions; white-numbered counties represent cultivated ornamental individuals or populations: 1 = Washington (1952,1999), 2 = Shelby (1972), 3 =

Cumberland (1975), 4 = Knox (1991,2012,2013,2014), 5 = Sevier (1998), 6 = Grainger (1999), 7 = Bradley (2002), 8 = Hamilton (2010), 9 = Anderson

(2014), 10 = Putnam (2014), 11 = DeKalb (2014), 12 = Davidson (2014), 13 = Blount (2014). (Tennessee map adapted by Melanie G. Bentley, Eastern

Kentucky University)

TENNESSEE. Blount Co.: Maryville, Maryville College, E side of Bartlet Hall Student Center lawn off Circle Drive and Morningside Lane,

cultivated shrub with lilac-lavender flowers; 35.75053°N, 83.96429°W, 448 m, 29 July 2014, R.L. Thompson & K. Rivers Thompson 14-341

(BEREA, TENN); Townsend, Great Smoky Mountain Heritage Center, 123 Cromwell Drive, off TN 73, one surviving shrub with purple

flowers planted by flagpoles in front, one dead from April freeze, 35.67472°N, 83.722292°W, 344 m, 29 July 2014, R.L. Thompson & K. Rivers

Thompson 14-342 (BEREA, TENN). Bradley Co.: Cleveland, growing at the S side of gymnasium at Cleveland State Community College,

adjacent to parking lot approximately 9 m from Adkisson Drive; shrub about 1.5 m tall, mint green color, few dark purple flowers, 220 m, 17

Mar 2002 , Anna Lea Griffith 15 (UCHT). Cumberland Co.: Crab Orchard, roadside along Highway 70, just W ofl-40 E, Exit329,30Jul 1975,

B. Eugene Wofford 51846 with W. Michael Dennis (TENN); escape by roadside rip-rap near creek, 28 Aug 1975, R. Krai 56489 (MO, VDB).

Grainger Co.: Rutledge, planted on Tampico Road, 6 Sep 1999, H.R. DeSelm s.n. (TENN). Hamilton Co.: Apison, along fence row at 4312 Bill

Jones Road, commonly escapes from cultivation, 9Jul 2010, Gary H. Morton 9203 (UCHT). Knox Co.: Knoxville, planted at Woodland Drive,

Sequoyah Hills, 20 Oct 1991, H.R. DeSelm s.n. (TENN); Knoxville, Ijams Nature Center, 2915 Island Home Avenue at NW corner of William

O. Miller Education Building (the Home Site), a single shrub planted in 1983, not spreading, 35.95758°N, 83.8696TW, 282 m, 29 Nov 2012,

R.L. Thompson & S. Brobst 12-1157 (BEREA, TENN); 18 May 2014, R.L. Thompson & K. Rivers Thompson 14-120 (BEREA, TENN). Putman

Co.: Cookeville, two shrubs planted at 2675 Lakeland Drive off 1-40 E at Exit 290, US 70 (TN 24), 36.13151°N, 85.44244°W, 291 m, 15 Apr

2014, R.L. Thompson & S.J. Stedman 14-25 (BEREA, TENN); white flowers, evident branch die-back freeze damage, 12 Jul 2014, R.L. Thomp¬

son & G.N. Douglas 14-340 (BEREA, TENN). Sevier Co.: Sevierville, planted on Section Springs Road, 17 Aug 1998, H.R. DeSelm s.n. (TENN).

Shelby Co.: Memphis, cultivated, Helene Griffith’s home, 568 St. Nick Drive, corolla white, 21 Jul 1972, Edward T. Browne, Jr. & Elizabeth M.

Browne 72 EH.3 (TENN). Washington Co.: Johnson City, escaped from cultivation off Knob Creek Road near Snow Chapel Memorial Baptist

Church, 28 Jul 1952, R.L. James 16885 (TENN); Johnson City, East Tennessee State University, escaped at forest edge of the hiking trail across

from Central Receiving, 20 Nov 1999, Mark Dugger 27 (ETSU).

Reconnaissance of Tennessee Butterfly Bush Populations

Field investigations for seven naturalized Buddleja davidii populations and their woody associates were expe¬

dited through the assistance of six Tennessee naturalists (S. Brobst & D. Estes, pers. comm. 2012; D. Bruce, T.

Crabtree, M. Smith, & S. Stedman, pers. comm. 2014). The seven population sites were discovered in Knox

County (3), Anderson (2), DeKalb (1), and Davidson (1) counties; these are new county naturalized Tennessee

populations. Orange-eye butterfly bush and the woody associates at each of seven sites are discussed in regards

to their location, habitat, physical site characteristics, relative abundance, relative frequency, woody associated

species status, and effects of B. davidii toward the native flora and vegetation.

Buddleja davidii naturalized in Knox County

Three naturalized populations of Buddleja davidii were discovered and studied in Knoxville, Knox County,

Tennessee, during 2012-2014 (Fig. 2; Fig. 3). Knoxville and vicinity are situated within the Southern Lime¬

stone/Dolomite Valleys and Rolling Hills Ecoregion of the Ridge and Valley Province. Potential vegetation is a

mosaic of Appalachian Oak and Mesophytic Forest often with Juniperus virginiana L. (Griffith et al. 1997).

Forested terrain typically consists of undulating to rolling, rounded hills comprised of underlying bedrock

strata from the Ordovician Holston Limestone Formation (Griffith et al. 1997).

Thompson and Thompson, Buddleja davidii in Tennessee

219

Fig. 3. Lakeshore Park levee (Site 3) designated by narrow white line adjacent to the west side of theTennessee River, Knox County, Knoxville. [Map

modified from Knoxville 2013 Master Plan, (www.cityofknoxville.org/lakeshore/masterplanupdate121613.pdf)

Ijams Nature Center, a 111.3-ha (275-acre) wildlife sanctuary and wildlife environmental learning center,

under the management of the Knox County Department of Parks and Recreation, Knoxville, is comprised of

lands mainly from two abandoned marble quarries, Ross Quarry and Mead’s Quarry (James 2010). From these

two quarries, Tennessee marble, a pink to cedar-red coarse crystalline limestone rock from the Ordovician

Holston Limestone Formation was excavated Games 2010). Mead’s Quarry, the larger quarry, was in operation

from 1881-1945 for large marble stone blocks. After 1945-1978, limestone was crushed, kiln-extracted and

transported as agriculture lime for commerce by the Knoxville and Holston Railway (KXHR) K Line. Mead’s

Quarry Lake, a 10.1-ha (25-acre lake) subsequently was formed when water pooled in the quarry pit after

Mead’s Quarry closed for the last time in 1978 (Fig. 2). The KXHR railway is situated parallel to Mead’s Quarry

and is still in operation as a short-line spur twice a month for transporting commerce to the Tennessee River.

The quarry lake and its environs became a part of Ijams Nature Center in 2001 Q ames 2010). In 2005, three

limestone kilns were demolished with a stone wall and three lime chute openings as the only remnants today

of the agricultural lime operation (Paul James, Director, Ijams Nature Center, pers. comm. 2012).

Site 1: Ijams Nature Center, Meads Quarry and Railroad Population

On November 21, 2012, a preliminary reconnaissance located the first naturalized Buddleja davidii population

along the KXHR tracks adjacent to Mead’s Quarry Lake (Site 1); 12 herbaceous and 13 woody associates were

recorded (Thompson & Abbott 2013). This butterfly bush population was noted in 2003 by Dwayne Estes,

220

Journal of the Botanical Research Institute of Texas 9(1)

Austin Peay State University, but vouchers were not taken (D. Estes, pers. comm. 2012). At this location, or¬

ange-eye butterfly bush is infrequent in relative abundance. Buddie]a davidii consisted of 20-25 plants com¬

prised of a few mature shrubs, smaller volunteering shrubs, and seedlings that colonized a 75 m extension of

the KXHR railroad tracks parallel to the limestone wall and kiln chutes. Butterfly bushes were embedded in a

thick crust of slag agriculture lime with typical seasonal vegetation cover of non-native and native woody and

herbaceous species. Unseasonally late April and early May freezes in conjunction with herbicide spraying by

the KXHR railway personnel between May and June 2014, killed much of the right-of-way vegetation including

two adjacent large and several smaller orange-eye butterfly bushes.

The original seed source was undoubtedly derived from a few Buddleja davidii between the limestone

kilns and the railroad tracks initially planted by a troop of Boy Scouts in 1983 as part of an ornamental planting

project near Mead’s Quarry (Sally Mirick, Former Director, Ijams Nature Center, pers. comm. 2012). The dis¬

persal of the tiny wind-carried seeds has probably been partly enhanced by the “slipstreaming” wake action by

the short-spur railroad freight cars and locomotives. Slipstreaming of B. davidii seeds was discussed in a recent

descriptive study of butterfly bush dispersal, colonization, and naturalization in Kentucky (Thompson & Ab¬

bott 2013).

Twenty-seven tree, shrub, and woody vine associated species (16 native, 11 non-native) were interspersed

among 68 herbaceous (28 native, 40 non-native) taxa. Mead’s Quarry habitats were open, sunny, ruderal areas

(hiking and biking trails, mowed and unmowed areas, parking lots, railroad track right-of-way) where many

opportunistic native and non-native naturalized taxa have thrived.

The 16 native trees, shrubs, and vines were documented. Scarce native woody taxa were Frangula carolin-

iana (Walter) A. Gray and Sambucus canadensis L. Infrequent to occasional native woody species were Acer

negundo L., Celtis occidentalis L., Cercis canadensis L., Gleditsia triacanthos L.Juniperus virginiana L., Lirioden-

dron tulipifera L., Parthenocissus quinquefolia (L.) Planch., Platanus occidentalis L., Rhus glabra L., Robinia pseu¬

doacacia L., Rubus occidentalis L., R. pensilvanicus Poir., Toxicodendron radicans (L.) Kuntze, and Ulmus rubra

Muhl. (Table 1). Eleven naturalized woody taxa documented were infrequent Celastrus orbiculatus Thunb.,

Ulmus pumila L.; occasional Ailanthus altissima (Mill.) Swingle, Albiziajulibrissin Durazz., Elaeagnus umbellata

Thunb., Paulownia tomentosa (Thunb.) Sieb. & Zucc. ex Steud., Pyrus callcryana Decne, Rosa multiflora Thunb

ex Murray; and frequent Ligustrum sinense Lour., Loniccra japonica Thunb., L. maackii (Rupr.) Maxim. (Table

1). All naturalized species, except Ulmus pumila, are classified as woody invasives by the Tennessee Exotic Pest

Plant Council (TN-EPPC 2009).

Label data from the Mead’s Quarry kiln wall and railroad site, Knox County, are summarized as follows: TENNESSEE. Knox Co.: Knox¬

ville, Ijams Nature Center, adjacent to Mead’s Quarry Lake, between the remaining limestone kilns wall and the railroad track bed; natural¬

ized Chinese shrubs, infrequent relative abundance, 15-30 shrubs, 35.95066°N, 83.86744°W, 272 m, 20 Nov 2012, R.L. Thompson & K.

Rivers Thompson 12-1153 (APSC, BEREA, MO, NCU, TENN); 8 Jul 2013, R.L. Thompson & K. Rivers Thompson 13-375 (APSC, BEREA, NCU);

5 Oct 2013, R.L. Thompson, K. Rivers Thompson, &P.F. Threadgill 13-708 (BEREA, TENN).

Site 2. Ijams Nature Center, Imery’s Trail Population

On November 29, 2012, a larger second naturalized population of Buddleja davidii was discovered at the ex¬

treme south-southwest portion of Ijams Nature Center near the end of Imery’s Trail (Site 2) near Aberdeen

Lane (Fig. 2). Orange-eye butterfly bush had occasionally spread into an open sunny area where old limestone

kilns and concrete block foundations were dismantled in 2008 (Paul James, Director, Ijams Nature Center,

pers. comm. 2012). Butterfly bush in various phenological stages was growing among brick and concrete block

debris and limestone slag wastes by Imery’s Trail. The butterfly bush population was enclosed by a secondary

successional mesophytic hardwood forest of frequent codominant Acer saccharum Marshall and Liriodcndron

tulipifera L. with a dense shrub and vine thicket. During May 2014, active bulldozing destroyed many butterfly

bush shrubs when Imery’s Trail to Aberdeen Lane was widened. Furthermore, the atypical late April-early May

freezes caused severe die-back of butterfly bush branches.

Thirty woody associates (18 native, 12 naturalized) were recorded from this open disturbed forest edge

(Table 1) scattered among 60 herbaceous (36 native, 24 non-native) taxa. Twenty-three of these woody taxa

were also recorded at the Mead’s Quarry (Site 1), which included frequent invasive Ligustrum sinense, Lonicera

Thompson and Thompson, Buddleja davidii in Tennessee

221

japonica, and L. maackii (Table 1). Six additional woody species (four native, two non-native) at Imery’s Trail

were infrequent to occasional Campsis radicans (L.) Seem, ex Bureau, Clematis virginiana L., Euonymusfortunei

(Turcz.) Hand.-Mazz., Fraxinus americana L., Pueraria montana (Lour.) Merr. var. lobata (Willd.) Maesen &

Almeida, and Vitis vulpina L. Four woody plants present at Mead’s Quarry, but not recorded at Imery’s Trail

(Site 2) were Frangula caroliniana, Gleditsia triacanthos, Sambucus canadensis, and Ulmus pumila (Table 1). Of

these 12 naturalized woody taxa, all except U. pumila, are listed as invasives by the TN-EPPC (2009).

The Acer saccharum-Firiodendron tulipifera woodland has progressed to a later serai stage of secondary

succession than the Mead’s Quarry site with fewer ruderal and more native herbaceous and woody taxa. The

encroachment of a dense understory shrub thicket and tree canopy confines the expansion of butterfly bush

colonization at the Imery’s Trail site. This observation supports further the conclusions of Trueblood (2009):

the migration of Buddleja davidii, a shade-intolerant pioneer shrub, is being controlled by competition through

natural secondary plant succession.

A total of 125 herbaceous and woody associated species (91 herbaceous, 34 woody) were recorded from

the two naturalized Buddleja davidii populations at Ijams Nature Center (Sites 1, 2). The woody species were

composed of 22 native and 12 naturalized taxa. Twenty-four (14 native, 10 naturalized) of the 34 woody associ¬

ates were common to both B. davidii population sites (Table 1). The 34 woody taxa (17 trees, 8 shrubs, 9 vines)

at the two Ijams Nature Center sites were comparable to the 34 woody taxa (19 trees, 7 shrubs, 8 vines) docu¬

mented at a drastically disturbed 13-year-old abandoned limestone quarry in central Kentucky (Thompson &

Green 2010).

The Ijams Nature Center Imery’s population (Site 2) label data are as follows: TENNESSEE. Knox Co.: Knoxville, Ijams Nature Center

property, 230 m S of 2515 Aberdeen Lane SE beside Imery’s Trail, occasional relative abundance, 60-80 naturalized shrubs among rubble,

35.94627°N, 83.87642°W, 286 m, 29 Nov 2012, R.L. Thompson &S. Brobst 12-1160 (APSC, BEREA, MO, TENN); 8Jul 2013, R.L. Thompson &

K. Rivers Thompson 13-376 (APSC, BEREA, BRIT, NCU, TENN); 5 Oct 2013, R.L. Thompson, K. Rivers Thompson, & P.F. Threadgill 13-701; 13-

702 (APSC, BEREA, TENN)

Site 3: Lakeshore Park Tennessee River Rip-rap Levee Population

On January 10, 2014, a third population of Buddleja davidii was investigated at Lakeshore Park, Knoxville, a

78.9-ha (195-acre) park also managed by the Knox County Department of Parks and Recreation. Forty-eight

orange-eye butterfly bush plants occasional in relative abundance, were established on the rounded apex of a

0.53 km limestone rip-rap levee (Site 3). The levee adjoins Lakeshore Park on the riparian west shoreline of the

Tennessee River, 17.2 km downstream from Ijams Nature Center (Fig. 3). Orange-eye butterfly bush coloniza¬

tion on the levee was probably due to the combination of wind-and-water seed dispersal. Twenty-eight woody

associates (17 native, 11 non-native) were documented. Infrequent to occasional associated species on the le¬

vee crest were Ailanthus altissima, Albiziajulibrissin, Figustrum sinense, Fonicera maackii, Paulownia tomentosa,

Platanus occidentalis, Rosa multiflora, and Salix nigra Marshall. Eight woody associates (7 native, 1 non-native)

not present at Ijams Nature Center (Sites 1, 2) were the infrequent Acer saccharinum L., Carya cordiformis

(Wangenh.) K. Koch, Cornus amomum Mill., Euonymus pungens Thunb., Fraxinus pennsylvanica Marshall,

Quercus muhlenbergii Engelm., Salix nigra, and Ulmus americana L. (Table 1).

Label data for Lakeshore Park (Site 4), Knoxville are as follows: TENNESSEE. Knox Co.: Knoxville, Lakeshore Park, along the top of a lime¬

stone boulder rip-rap levee, 48 naturalized shrubs, occasional relative abundance, 35.92372°N, 83.98660°W, 253 m at apex, 249 m at shore¬

line, 10 Jan 2014, R.L. Thompson & DM. Bruce 14-02 (APSC, BEREA, TENN).

Buddleja davidii naturalized in Anderson County

Orange-eye butterfly bush was documented from two naturalized populations in Anderson County on April 5,

2014. Both population sites are entirely located within the Southern Limestone/Dolomite Valleys and Low

Rolling Hills Ecoregion within the Ridge and Valley Province of potential Appalachian Oak and Mesophytic

Forest (Griffith et al. 1997).

Site 4: Grassy Road Shoulder and Highwall Population

A Buddleja davidii population was situated within the city limits of Clinton, Tennessee, beside the Charles G.

Seivers Boulevard (TN 61W). A mature butterfly bush, 4 m x 3 m, was growing within a mixed tall fescue (Fes-

222

Journal of the Botanical Research Institute of Texas 9(1)

tuca arundinacea Schreb.) road shoulder adjacent to an Ordovician limestone highwall road cut (Site 4). This

single shrub has undoubtedly served as the seed source to the 8-9 scarce volunteering shrubs embedded

within limestone fractures of a 7-9 m vertical highwall.

Twenty-one woody associates (10 native, 11 non-native) on the road shoulder and highwall were recorded

(Table 1). No new native or non-native woody taxa were recorded for Site 4. Among 11 state-listed invasive spe¬

cies, Ligustrum sinense and Lonicera maackii were frequent codominant shrubs. The unusual freezes of late

April and early May killed most of the new leafing branches of the large “parent” butterfly bush, and three

small butterfly bushes embedded in the limestone highwall crevices died from the spring freezes as well. Little

or no freeze effects were noted on the 11 invasive taxa. Some root collar sprouting was evident on May 18,2014,

and by late June and July, lilac-lavender inflorescences appeared on the surviving parent and surviving high¬

wall plants. Senescence occurred by late September.

Label data for the Anderson County roadside and highwall (Site 4) are as follows: TENNESSEE. Anderson Co.: Clinton, TN 61W adjacent

to Clinch River, tall fescue road shoulder between limestone highwall talus, scarce relative abundance, ca. 8-9 small shrubs established in

crevices of a 5-7 m highwall from a large parent shrub, infrequent relative abundance, 36.08231°N, 84.161154°W, 211 m, 5 Apr 2014, R.L.

Thompson & D.R. Bruce 14-23 (APSC, BEREA, TENN); severe leaf frost damage, 18 May 2014, R.L. Thompson & K. Rivers Thompson 14-111

(BEREA, TENN); lilac-lavender flowers on new or undamaged branches of parent shrub, 21 Aug 2014, R.L. Thompson & G.N. Douglas 14-520

(APSC, BEREA, TENN).

Site 5: Abandoned Asphalt, Concrete, and Gravel Landfill

The second Anderson County population was located in Oak Ridge east of Southern Illinois Avenue (TN 62) on

Union Valley Road contiguous to the east side of the Oak Ridge Golf Center, 119 Union Valley Road (Site 5).

Two rare orange-eye butterfly bushes were growing in different areas 40 m apart in an open, nearly level aban¬

doned asphalt, concrete, and aggregate gravel landfill. A smaller shrub had only a single shoot regenerating

from the rootstock, while a second larger shrub had several basal shoots rejuvenating from the rootstock al¬

though most of the shrub branches exhibited severe dieback.

This abandoned landfill (Site 5) had the highest species richness of the seven sites with 34 associated

woody plants (20 native, 14 non-native) within the immediate vicinity of these two shrubs (Table 1). This land¬

fill ruderal habitat site promoted wide-spread seed dispersal and establishment of both native and non-native

woody taxa from various anthropogenic refuse sources. Ligustrum sinense, Lonicera japonica, L. maackii, and

Pyrus calleryana were the frequent dominant taxa. Three additional native taxa at the landfill site not present

at the other four sites were occasional in relative abundance: Ampelopsis cordata Michx., Fraxinus pennsylvanica

Marshall, and Rhus copallinum L. Four non-native scarce naturalized taxa present were Morus alba L., Nandina

domestica Thunb., Prunus persica (L.) Batsch, and Yucca filamentosa L. (Table 1).

Label data for the abandoned landfill (Site 5) in Anderson County are as follows: TENNESSEE. Anderson Co.: Oak Ridge, Oak Ridge High¬

way (TN 170) to Southern Illinois Avenue (TN 62) to Union Valley Road east to a disused city landfill adjacent to Oak Ridge Golf Center, 119

Union Valley Road, rare relative abundance, small solitary shrub embedded in fractured asphalt-concrete layer, 36.00203°N, 84.22006°W,

284 m, 5 Apr 2014, R.L. Thompson & D.R. Bruce 14-24 (APSC, BEREA, TENN); small shrub with a single surviving basal branch and lilac-

lavender flowers, 21 Aug 2014, R.L. Thompson & G.N. Douglas 14-537 (BEREA); larger shrub with lilac-lavender inflorescences from several

basal shoots from rootstock, remainder of shrub dead, 36.00166°N, 84.21981°W, 282 m, 21 Aug 2014, R.L. Thompson & G.N. Douglas 14-538

(APSC, BEREA, TENN).

Buddleja davidii naturalized in DeKalb County

On April 15, 2014, a naturalized population of Buddleja davidii was discovered off TN 83 north of Smithville to

Holmes Creek Road for 3.8 km north across from the concrete remains of Hidden Harbor Marina at Center Hill

Lake. The abandoned roadside thicket (Site 6) is located entirely within the Outer Nashville Basin Ecoregion of

the Interior Plateau Province. Major vegetation is comprised of Oak-Hickory Forest (Griffith et al. 1997).

Site 6: Abandoned Gravel Road by Center Hill Lake

A total of 9-12 infrequent seed-producing shrubs in various growth stages were rooted in scattered limestone

aggregate near an exposed herbicide-sprayed power line cut. Twenty-two woody associates (14 native, 8 natu¬

ralized) were recorded at Site 6 (Table 1). This population of orange-eye butterfly bush has been heavily shaded

Thompson and Thompson, Buddleja davidii in Tennessee

223

by the dense forest edge thicket by the frequent Asian invasives, Ligustrum sinense, L. vulgare L., and Lonicera

maackii (Table 1). The health of the butterfly bush population was also severely affected by late April and early

May freezes with a majority of the larger shrub branches killed back to the rootstock. Likewise, this population

appeared to be restricted to the abandoned gravel road due to herbicide spraying of the nearby power line cor¬

ridor.

Label data at the DeKalb County disused roadside thicket (Site 6) are as follows: TENNESSEE. DeKalb Co.: Smithville, on Holmes Creek

Road, 3.8 km N of TN 83 along Center Hill Lake on an abandoned limestone aggregate road near power line cut near the Hidden Harbor

Marina foundation, scarce relative abundance, ca. 9-10 shrubs present, 36.700175°N, 85.26480°W, 203 m, 15 Apr 2014, R.L. Thompson & S.J.

Stedman 14-26 (APSU, BEREA, TENN); lilac-rose flowers, severe die-back from spring freezes, 11 Jul 2014, R.L. Thompson & G.N. Douglas

14-330 (APSU, BEREA, TENN)

Buddleja davidii naturalized in Davidson County

Shelby Park, a 136-ha (336-acre) urban multi-use park located in downtown Nashville adjacent to the Cumber¬

land River, is located in Davidson County (Nashville Parks & Recreation 2014). The city park lies entirely

within the Outer Nashville Basin Ecoregion of the Interior Plateau Province and consists of potential Oak-

Hickory Forest (Griffith et al. 1997).

Site 7: Steep Cumberland River Rip-rap Bank Population

Two mature Buddleja davidii shrubs were growing out of large limestone rip-rap along the steep embankment

thicket of the Cumberland River (Site 7) near a chain link fence separating the Shelby Park lawn from Davidson

Street (Music City Bikeway). The thrysoid inflorescence was composed of lilac-lavender perfect, 4-merous

flowers (Fig. 4) as all B. davidii shrubs were within the seven populations. Twenty-four woody associates (17

native, 7 naturalized) were recorded from this Cumberland River riparian thicket (Table 1). The two butterfly

bushes were totally surrounded by a dense thicket of Euonymus fortunei, Ligustrum sinense, Lonicera japonica,

L. maackii, and various occasional native woody vines, Ampelopsis cordata, Parthenocissus quinquefolia, Toxico¬

dendron radicans, and Vitis vulpina. Two additional rare native species reported for the seven sites were Catalpa

bignonioides Walter and Smilax rotundifolia L. (Table 1). Frost damage was not evident mainly due to the tem¬

perature buffer from the close proximity to the Cumberland River. No seedlings or evidence of butterfly bush

colonization were found.

Label data for these two shrubs on the bank of the Cumberland River are as follows: TENNESSEE. Davidson Co.: Nashville, Shelby Park,

Davidson Street (Music City Bikeway) across chain link fence on a steep open embankment in limestone rip-rap by Cumberland River; rare

relative abundance, first naturalized shrub, 36.16505°N, 86.73046°W, 130 m, 16 Apr 2014, R.L. Thompson 14-30 (APSC, BEREA, MO,

TENN); lilac-lavender flowers, slight branch frost damage, 11 Jul 2014, R.L. Thompson & G.N. Douglas 14-336 (APSU, BEREA, TENN); sec¬

ond shrub adjacent lower down, lilac-lavender inflorescence, 36.164972°N, 86.73070°W, 129 m, 11 Jul 2014, R.L. Thompson & G.N. Douglas

14-338 (BEREA, TENN).

Synopsis of Naturalized Buddleja davidii Populations and Woody Associates

Buddleja davidii was documented as locally naturalized from held reconnaissance at seven population sites in

four Tennessee counties as follows: Knox (3), Anderson (2), DeKalb (1), and Davidson (1) during 2012-2014

(Table 1).

Ruderal or culturally disturbed sites and relative abundance values for the seven B. davidii populations

were as follows: Knox County, Ijams Mead’s Quarry Railroad (Site 1) = Scarce; Knox County, Ijams Imery’s

Trail (Site 2) = Occasional; Knox County, Fakeshore Park Rip-rap Fevee, Tennessee River (Site 3) = Occasional;

Anderson County, Highway Road Shoulder and Highwall (Site 4) = Scarce; Anderson County Disused Fandhll

(Site 5) = Rare; DeKalb County, Abandoned Road Thicket at Center Hill Fake (Site 6) = Infrequent; and David¬

son County, Cumberland River rip-rap embankment (Site 7) = Rare. These naturalized B. davidii populations

locally confined to culturally-derived and disturbed habitats have not volunteered or colonized any natural

thickets or forest edges.

A summary of descriptive site characteristics and woody associated species from the seven B. davidii

population studies include the following:

224

Journal of the Botanical Research Institute of Texas 9(1)

Fig. 4. Buddleja davidii inflorescence of lilac-lavender perfect flowers from Cumberland River rip-rap embankment population (Site 7), by Shelby Park,

Nashville. (Photo taken by Michael R. Smith, 8 July 2013).

1) Fifty-one woody species [32 native (62.75%), 19 non-native (37.25%)] in 41 genera from 26 plant families

were documented among the study sites.

2) These 51 woody taxa consisted of 26 trees (19 native, 7 exotic), 14 shrubs (7 native, 7 exotic), and 11 woody

vines (7 native, 4 exotic).

3) Fifteen invasives of the 19 naturalized woody taxa (78.95%) classified by the Tennessee Exotic Pest Plant

Council (TN-EPPC 2009) are listed as follows: Severe Threat (10), Significant Threat (2), Lesser threat

(1), and Alert (2).

4) Eleven woody non-native invasives with a Relative Frequency (frequency of occurrence) of 57% or greater (4

of 7 sites) are listed in order: Ailanthus altissima, Ligustrum sinense, Lonicerajaponica, L. maackii, Albizia

julibrissin, Pyrus calleryana, Celastrus orbiculatus, Paulownia tomentosa, Rosa multiflora, Elaeagnus um-

bellata, and Euonymusfortunei (Table 1).

5) These 11 opportunistic woody invasives are inclined to have a greater ecological amplitude and tolerance to

volunteer, colonize, compete, thrive, and invade natural habitats in early serai stages of progressive sec¬

ondary succession than Buddleja davidii.

6) Most of these invasives have greater relative abundance values than Buddleja davidii populations.

7) Some non-native invasive adaptations include direct and indirect insolation, variation in moisture-soil re¬

quirements and varied pH, strong intraspecific and interspecific competition, better acclimation to cold

and herbicides, and a length of establishment over 10 years in the case of typical woody non-native spe¬

cies (Pysek et al. 2004).

8) No Tennessee special concern, threatened, or endangered herbaceous or woody taxa on the Rare Plant List

(Crabtree 2014) were recorded within or near proximity to the seven study sites.

Thompson and Thompson, Buddleja davidii in Tennessee

225

CONCLUSION

We determined Buddleja davidii to be locally naturalized in Tennessee based on observations from field recon¬

naissance, voucher documentation from extant populations in Knox, Anderson, DeKalb, and Davidson coun¬

ties, and by label information from examination of Tennessee herbarium specimens from Cumberland, Ham¬

ilton, and Washington counties. Buddleja davidii definitely did not exhibit the widespread invasive adaptations

displayed by most non-native woody invasives in this study Butterfly bush relative abundance values for the

seven populations ranged from rare, scarce, infrequent, to occasional. These seven populations have not ex¬

panded their range beyond ruderal and culturally-derived areas. At an optimal classification, B. davidii popula¬

tions observed in this study are sparingly and locally naturalized. Similarly in a Kentucky study, Buddleja da¬

vidii was reported naturalized, but was not determined to be invasive (Thompson & Abbott 2013).

As observed from this study, Buddleja davidii is a shade intolerant, short-lived pioneer species established

in ruderal habitats of Tennessee that is eliminated through natural plant succession in absence of disturbance.

Butterfly bush continues to be restricted to those disturbed habitats rather than secondary forested areas.

Other contributing factors observed toward limited naturalization include poor competition among native and

especially aggressive invasive woody taxa, low adaptability to unseasonably cold temperatures, and a high

sensitivity to herbicides.

Trueblood (2009) evaluated the impact of Buddleja davidii on natural vegetation in North Carolina and

reported that it has a tendency to be eliminated through natural plant succession. Moreover, orange-eye but¬

terfly bush exhibited little ecological impact, distribution expansion, or invasive potential. These premises also

appear to be applicable to Tennessee and Kentucky naturalized B. davidii populations. We propose the limited

naturalization of Buddleja davidii in Tennessee at this present time clearly does not imply any invasive status

beyond its current “Alert” rank from the TN-EPPC (2009). Exotic pest plant council personnel in Tennessee,

Kentucky, and other southeastern states could benefit from evaluating B. davidii with a rational assessment

system, such as developed for North Carolina.

ACKNOWLEDGMENTS

We express our appreciation to J. Richard Abbott, New York Botanical Garden, Bronx, NY, Keith M. Clancy,

USDA, APHIS, Jamaica, NY, and Eliane M. Norman, Stetson University, DeLand, Florida, for constructive

manuscript reviews. For held locations and/or photographs of Buddleja davidii populations, we are grateful to

SarahJ. Brobst, Ijams Nature Center and 2003 Berea College biology graduate; Douglas R. Bruce, Clinton; Todd

Crabtree, Tennessee Department of Environment and Conservation; G. Neil Douglas, Berea College; Dwayne

Estes, Austin Peay State University and Botanical Research Institute of Texas; Michael R. Smith, Henderson¬

ville; StephenJ. Stedman, Tennessee Tech University; and Paul F. Threadgill, Maryville College. Special thanks

are extended to Melanie G. Bentley, Eastern Kentucky University, for three map figures. We recognize the cura¬

tors of APSC, BRIT, EKY, ETSU, HTTU, KY, MO, MTSU, NCU, TENN, UCHT, and VDB for their courtesy dur¬

ing herbarium visits and/or confirming their Buddleja davidii specimen holdings.

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Journal of the Botanical Research Institute of Texas 9(1)

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Petron

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Concepts and Methodologies for Measuring the Sustainability of Cities—M. Yetano Roche, S. Lechtenbohmer, M. Fischedick, M.-C. Grone,

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Networks and the Challenge of Sustainable Development—A. Douglas Henry & B. Vollan

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Cumulative Index of Contributing Authors, Volumes 30-39

Cumulative Index of Article Titles, Volumes 30-39

Errata

J.Bot. Res. Inst. Texas 9(1): 228.2015

ADDENDUM TO THE VASCULAR FLORA OF THE HANCOCK BIOLOGICAL

STATION, MURRAY STATE UNIVERSITY, CALLOWAY COUNTY, KENTUCKY, U.S.A.

J. Richard Abbott

Ralph L.Thompson

Hancock Biological Station, Murray State University

Murray, Kentucky 40271, USA.

University of Florida Herbarium, Gainesville, Florida 32611, U.S.A.

Missouri Botanical Garden, P.O. Box299

Saint Louis, Missouri 63166-0299, U.S.A.

New York Botanical Garden, Bronx, New York 10458, U.S.A.

rabbott@nybg.org

Hancock Biological Station, Murray State University

Murray, Kentucky 40271, U.S.A.

Botanical Research Institute of Texas

1700 University Dr., Fort Worth, Texas 76107-3400, U.S.A.

Berea College Herbarium, Biology Program

Berea College, Berea, Kentucky 40404, U.S.A.

ralph_thompson@berea.edu

ABSTRACT

A vascular plant reconnaissance was conducted at the Hancock Biological Station (HBS) of Murray State University during the summers of

2012-2015. The HBS a 37.5-ha tract of mainly upland oak-hickory forest, is located 23 km from Murray, Kentucky, in northeastern Calloway

County within the Jackson Purchase Region. In 2007, a floristic study of HBS reported 573 specific and infraspecific taxa in 334 genera from

121 families. Twenty-one additional taxa (12 native, 9 non-native) have been documented from incidental collections. Thirteen taxa were

new Calloway County distribution records including five Kentucky-listed invasive species. Ulmus alata was also documented as a new host

tree for Phoradendron leucarpum ssp. leucarpum in Calloway County. The documented native and exotic flora of HBS presently consists of

594 taxa, 345 genera, and 126 families.

Keywords: Hancock Biological Station, Calloway County, invasive plants, Kentucky vascular flora

RESUMEN

Un reconocimiento de plantas vasculares se llevo a cabo en la Estacion Biologica Hancock (HBS) de la Universidad Estatal de Murray du¬

rante los veranos de 2012 a 2015. El HBS, un tramo de 37,5 hectareas principalmente de bosque de roble-nogal de tierras altas, se encuentra

a 23 km de Murray, Kentucky, en el noreste del Condado de Calloway en la Region Compra Jackson. En 2007, un estudio floristico de HBS

reporto 573 taxones especificos e infraespecificos de 334 generos de 121 familias. Veintiun nuevos taxones (12 nativos, 9 no nativos) han

sido documentados en colecciones incidentales. Trece taxones fueron nuevos registros de distribucion del Condado de Calloway, incluyen-

do cinco especies invasoras listadas en Kentucky. Ulmus alata tambien fue documentado como un nuevo arbol huesped de Phoradendron

leucarpum ssp. leucarpum en el Condado de Calloway. La flora nativa y exotica documentada de HBS actualmente consta de 594 taxones, 345

generos y 126 familias.

Hancock Biological Station (HBS) was founded in 1966 by Hunter M. Hancock, Professor of Biology at Murray

State University from a 16.2-ha tract of old fields, pastures, and secondary successional upland oak-hickory

forest in northeastern Calloway County (White 2002). The HBS now consists of a 37.5-ha tract that adjoins

Kenlake State Resort Park to the north, impounded Kentucky Lake shoreline of the Tennessee River on the

east, and Pacer Point to the southeast off Watersport Road. The HBS is located 24 km from Murray, the county

seat of Calloway County, the southeastern-most county of the eight countyjackson Purchase Region.

Thompson (2003) recorded 560 taxa in 320 genera from 110 families from 1998-2002 in a checklist of the

vascular plants of HBS. Thompson (2007) designated 10 habitats at HBS in an on-going floristic study con¬

ducted during the summers of 2003-2006. Forest vegetation is predominately upland stands of dry or dry-

mesic oak-hickory forest, and mid-successional oak-hickory woodlands, which border various culturally de¬

rived ruderal areas, wetland habitats, and an old held burnt warm-season restored prairie. Thompson (2007)

reported the known vascular Bora from Hancock Biological Station as 573 specific and infraspecific taxa, 334

genera, and 121 families. Woods and Fuller (1988) listed 1018 taxa, 462 genera, and 129 families for the vascu¬

lar Bora of Calloway County.

In conjunction with teaching an undergraduate/graduate field botany course at HBS during the summers

of 2012-2015, we conducted a floristic reconnaissance within the boundaries of HBS and collected representa-

J. Bot. Res. Inst. Texas 9(1): 229 - 233.2015

230

Journal of the Botanical Research Institute of Texas 9(1)

tive vouchers for taxa new to the station. Several newly discovered plants have volunteered and colonized ex¬

isting HBS habitats since the 2007 survey report.

Nomenclature and classification follow Jones (2005), with three exceptions that reflect recent famil¬

ial recircumscriptions, i.e., Paulownia is transferred from Bignoniaceae to Paulowniaceae, Mazus from

Scrophulariaceae to Mazaceae, and Celtis from Ulmaceae to Cannabaceae (Stevens 2001). Herbarium acro¬

nyms follow Index Herbariorum from Thiers (2014), i.e., Berea College Herbarium (BEREA), University of

Florida Herbarium (FLAS), and Murray State University Herbarium (MUR). This addendum to the HBS anno¬

tated list mainly follows the format of Thompson (2007). Symbols preceding the scientific name of a taxon may

include: an asterisk (*) for a non-native taxon; a double asterisk (**) for a state-listed invasive species by the

KY-EPPC (2013); a dagger (t) for a native or exotic planted taxon; and a diesis ft) for a Calloway County distri¬

bution record based on the maps of Campbell and Medley (2012) and USDA, NRCS (2014). Relative abundance

values are abbreviated as Rare (R); Scarce (S); Infrequent (I); Occasional (O); Frequent (F); and Abundant (A)

ALISMATACEAE

Echinodorus cordifolius (F.) Griseb., Creeping Burhead, emergent at Kentucky Fake shoreline east of Watersport

Road; R. Associate: Carex crus-corvi; 27 Jun 2014, J.R. Abbott & R.L. Thompson 26525 (BEREA). This was in very

young bud when found and was transplanted to await flowering. The original inflorescence died, but the plant

reflowered and was pressed 20 Aug 2014.

ASTERACEAE

$*Senecio vulgaris F., Common Groundsel, aggregate gravel at the side border of the Mesocosm Building; R, 30

Jun 2013, R.L. Thompson & J.R. Abbott 13-252 (BEREA). Campbell and Medley (2012) mapped six Kentucky

counties for this European annual weed.

CANNABACEAE

Celtis laevigata Willd., Sugarberry, throughout oak-hickory forest near Kentucky Fake; F, 20 Jun 2013, R.L.

Thompson & J.R. Abbott 13-325 (BEREA, FFAS, MUR). The key features used for differentiating Celtis laevigata

and C. occidentalis F., vary from author to author, e.g., Jones (2005), Whittemore (2013). Species circumscrip¬

tion, obviously, involves taxonomic judgment, and the treatment provided by Alan Whittemore in the Flora of

Missouri seems to be the best for reflecting biological reality in the landscape. Thus, many of the plants tradi¬

tionally seen as C. occidentalis at HBS are now understood to be C. laevigata.

CARYOPHYLLACEAE

ZSagina decumbens (Elliott) Torrey & A. Gray, Trailing Pearlwort, wet lawn beside three 1200-gallon stock

water tanks; S. Associates: Callitriche terrestris, Centunculus minimus, Isolepis carinata; 5 Jun 2013, R.L. Thomp¬

son & J.R. Abbott 13-261 (BEREA, MUR). Campbell and Medley (2012) mapped five counties for this often

overlooked taxon; the USDA, NRCS (2014) mapped 10 Kentucky counties.

CELASTRACEAE

$**Celastrus orbiculatus Thunb., Oriental bittersweet, dry-oak-hickory forest ecotone at back of Resource

Building, a severe threat invasive colonizer; R, 5 Jun 2012, R.L. Thompson 12-653 (BEREA); roadside edge of

woods, S side of Emma Road W of junction with Wolfson Road, 28 Jun 2014, J.R. Abbott 26527 (FLAS).

CORNACEAE

f*Cornus kousa F. Buerger ex Miq., Pagoda Dogwood, an introduced Asian tree persisting as an ornamental in

lawn at the front of the Resource Building; R, 26 Jun 2013, R.L. Thompson & J.R. Abbott 13-253 (BEREA, MUR).

CYPERACEAE

4Carex crus-corvi Shuttlew. ex Kunze, Crowfoot Sedge, emergent at Kentucky Lake shoreline east of Waters¬

port Road; R. Associate: Echinodorus cordifolius ; 27 Jun 2014, R.L. Thompson & J.R. Abbott 14-248 (BEREA,

FLAS, MUR).

Abbott and Thompson, Flora of the Hancock Biological Station, Kentucky

231

HYDROCHARITACEAE

$**Hydrilla verticillata (L.f.) Royle, Hydrilla, an Old World submergent in Kentucky Lake near HBS Boat Dock;

R, 27 Jun 2013, R.L. Thompson & J.R. Abbott 13-341 (BEREA, MUR). Jones (2005) did not list Hydrilla verticil¬

lata as a current taxon for Kentucky Campbell and Medley (2012) mapped only Jefferson and Trigg counties;

USDA, NRCS (2014) noted Floyd, Jefferson, Johnson, Knott, and Trigg counties.

LENTIBULARIACEAE

ZUtricularia gibba L., Longspur Creeping Bladderwort, submergent in 14 year-old pond; I. Associate: Potamo-

geton nodosus; 27 Jun 2014, R.L. Thompson & J.R. Abbott 14-251 (BEREA, FLAS, MUR).

MAGNOLIACEAE

t Magnolia grandiflora L., Southern Magnolia, a planted native tree persisting as an ornamental in lawn circle

near Main Building; R, 25 Jun 2013, R.L. Thompson &J.R. Abbott 13-252 (BEREA, MUR).

MAZACEAE

$*Mazus pumilus (Bernm. f.) Steenis, Japanese Mazus, near rail fence in front lawn of Main Building; R, 2 Jun

2013, R.L. Thompson & J.R. Abbott 13-257 (BEREA, MUR). We also documented this often passed over Euro¬

pean taxon at the Land Between The Lakes as a distribution record for Lyon County at the Woodlands Nature

Center, 19 Jun 2013, R.L. Thompson & J.R. Abbott 13-310 (BEREA). Campbell and Medley (2012) mapped

Jefferson, Lewis, and McCracken counties; the USDA, NRCS (2014) listed Fayette, Henderson, Jefferson, Lewis,

and McCracken counties.

MORACEAE

$*Fatoua villosa (Thunb.) Nakai, Mulberry Weed, wet gravel floor inside of Greenhouse Building; O, 5 Jun

2012, R.L. Thompson 12-656 (BEREA); 2 Jun 2013, R.L. Thompson&J.R. Abbott 13-251 (BEREA, FLAS, MUR).

Campbell and Medley (2012) mapped six Kentucky counties mainly from voucher specimens reported by

Vincent (2004). The USDA, NRCS (2014) mapped seven Kentucky counties.

OPHIOGLOSSACEAE

Ophioglossum pycnostichum (Fern.) A. & D. Love, Southern Adder’s-tongue, 22 plants (6 fertile) found along an

off-trail mesic drainage area below Dry-Mesic Oak-Hickory Forest. Associates: Botrychium dissectum, Carex

grayi, Chasmanthium latifolium, Dioscorea villosa, Polystichum acrostichoides, Trepocarpus aethusae ; I, 5 Jun

2015, J.R. Abbott 26814 (BEREA). Woods and Fuller (1988) first reported this fern from Calloway County based

on Hunter 803 (MUR).

OXALIDACEAE

Oxalis dillenii Jacq., Southern Yellow Wood Sorrel, scattered in mowed and unmowed lawn; O, 2 Jun 2013, R.L.

Thompson &J.R. Abbott 13-361 (BEREA, MUR).

PAULOWNIACEAE

**Paulownia tomentosa (Thunb.) Sieb. & Zucc. ex Steud., Chinese Empress Tree, along trail in oak-hickory for¬

est ecotone, a recent Asian colonizer at HBS; S, 20 Jun 2013, R.L. Thompson & J.R. Abbott 13-328 (BEREA, MUR);

27 Jun 2014, R.L. Thompson & J.R. Abbott 14-253 (BEREA). One sapling in a large water tank (dried out with

terrestrial vegetation) and two seedlings were in gravel adjacent to the Resources Building, 13 Jun 2014, J.R.

Abbott 26480 (FLAS).

POACEAE

Dichanthelium scoparium (Lam.) Gould, Velvety Panic Grass, seasonally wet ditch at Emma Drive east of en¬

trance gate; S, 5 Jun 2012, R.L. Thompson 12-654 (BEREA); 25 Jun 2012, R.L. Thompson & J.R. Abbott 12-791

(BEREA, MUR).

232

Journal of the Botanical Research Institute of Texas 9(1)

4*Vulpia myuros( L.) C.C. Gmel, Rat-tail Fescue, gravel along roadside and around stock water tanks on W side

of Greenhouse Building, S. Associates: Acalypha rhomboidea, Conyza canadensis, Euphorbia maculata; 12 Jun

2015, R.L. Thompson & J.R. Abbott 15-403 (BEREA). A naturalized European annual reported in six Kentucky

counties by Campbell and Medley (2012) and USDA, NRCS (2014).

POTAMOGETONACEAE

Wotamogeton nodosus Poiret, Leafy Pondweed, floating-leaved aquatic in 14 year-old pond; F. Associate: Utricu-

lariagibba; 25 Jun 2012, R.L. Thompson&J.R. Abbott 12-815 (BEREA, MUR); 27 Jun 2014, R.L. Thompson&J.R.

Abbott 14-250 (BEREA, FLAS).

PRIMULACEAE

$Centunculus minimus L., Chaffweed, wet lawn beside three 1200 gallon stock water tanks; O, 5 Jun 2013, R.L.

Thompson & J.R. Abbott 13-260 (BEREA, MUR). Abbott et al. (2001) documented the brst Kentucky collection

in Madison County [R.L. Thompson & J.R. Abbott 95-374 (BEREA)] since 1840 [Short s.n. (PH)] in Muhlenberg

County. Campbell and Medley (2012) also mapped Harlan County. The USDA, NRCS (2014) listed Harlan,

Madison, and Muhlenberg counties.

$**Tysimachia nummularia L., Moneywort, Kentucky Lake shoreline; O, 20 Jun 2013, R.L. Thompson &J.R.

Abbott 13-327 (BEREA, MUR).

VISCACEAE

Phoradendron leucarpum (Raf.) Rev. & Johnst. ssp. leucarpum, American Mistletoe, a single clump hemipara-

sitic in Ulmus alata diagonally from Wolfson Drive off Emma Drive, a new host tree for Calloway County; R, 30

Mar 2013, R.L. Thompson & W.W. Overbeck 13-20 (BEREA, MUR). Thompson and McKinney (1990) brst re¬

ported Ulmus alata as a mistletoe host tree for Kentucky in Lyon and Trigg counties at Land Between The

Lakes. Thompson (2007) previously documented mistletoe on Cary a glabra in dry oak-hickory forest at HBS

[13 Oct 2006, R.L. Thompson & M.K. Graves 06-757 (BEREA, MUR)]. Campbell and Medley (2012) mapped

mistletoe for Calloway County based upon the above 2006 collection.

VITACEAE

Vitis vulpina L., Frost Grape, in an oak-hickory woodland ecotone; I, 27 Jun 2014, R.L. Thompson & J.R. Abbott

14-252 (BEREA, MUR); also in shrub thicket at woodland edge W of the Mesocosm Building and observed at

three other scattered woodland sites, 4 Jun 2014, J.R. Abbott 26470 (FLAS).

We documented an additional 21 vascular plant species, 11 genera, and five families for the flora of Hancock

Biological Station. These 21 species (12 native and 9 non-native) comprise 13 new distribution records for

Calloway County. Five are listed as Kentucky state-listed invasive plants by the KY-EPPC (2013). The 594 taxa

account for 58.3% of the 1018 species listed by Woods and Fuller (1988) for the flora of Calloway County.

The documented vascular plants of the Hancock Biological Station now consist of 594 plant species, 345

genera, and 126 families. These 594 taxa account for nearly 23% of the 2600 recorded by Jones (2005) for the

Commonwealth of Kentucky. Concentrated collecting still needs to be conducted for additional vascular

plants in western Kentucky as 13 of 21 taxa were new Calloway County distribution records.

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BOOKS RECEIVED

Marco Madella, Carla Lancelotti, & Manon Savard. 2014. Ancient Plants and People: Contemporary

Trends in Archaeobotany. (ISBN-13: 978-0-8165-2710-6, hbk). The University of Arizona Press, Main

Library Building, 5th Floor, 1510 E. University Blvd., P.O. Box 210055, Tuscon, Arizona 85721-0055,

U.S.A. (Orders: www.uapress.arizona.edu, 1-800-621-2736). $70.00 US, 344 pp., 6" x 9".

Amy Eisenberg. 2013. Aymara Indian Perspectives on Development in the Andes. (ISBN-13: 978-0-8173-

1791-1, cloth). The University of Alabama Press, Box 870380, Tuscaloosa, Alabama 35487-0380, U.S.A.

(Orders: www.uapress.ua.edu, 1-800-621-2736). $49.95 US, 38 illustrations, 280 pp., 6 W x 9 W.

Wilhelm Barthlott, Kerstin Burstedde, Jan Laurens Geffert, Pierre L. Ibisch, Nadja Korotkova, Andrea Miebach,

M. Daud Rafiqpoor, Anke Stein, & Jens Mutke. 2015. Biogeography and Biodiversity of Cacti.

Schumannia 7. (ISSN: 1437-2517; ISBN-13: 978-3-7308-1144-3, hbk). Deutschen Kakteen-Gesellschaft

e.V. und der Gesellschaft Osterreichischer Kakteenfreunde (German Cactus Society and the Austrian

Cactus Friends). (Orders: www.dkg.eu, gs@dkg.eu). 205 pp., color figures and maps, English-German

text, 8 V 2 " x 12".

Richard B. Primack. 2012. (A Primer of) Conservation Biology, 5th edition. (ISBN-13: 978-0-87893-623-6,

pbk). Sinauer Associates, Inc., Publishers, 23 Plumtree Road, P.O. Box 407, Sunderland, Massachusetts

01375-0407, U.S.A. (Orders: www.sinauer.com, 1-413-549-4300). $74.95 US, 363 pp., 129 illustrations,

7" x 9 Yi".

Jan Salick, Katie Konchar, & Mark Nesbitt. 2014. Curating Biocultural Collections: A Handbook. (ISBN-13:

978-1-84246-498-4, pbk). The University of Chicago Press, 1427 East 60th Street, Chicago, Illinois

60637, U.S.A. (Orders: press.uchicago.edu, 1-800-621-2736). $50.00 US, 250 pp., 100 color plates, 7 Vi" x 10".

T.S. Nayar, A. Rasiya Beegam, & M. Sibi. 2014. The Flowering Plants of The Western Ghats, India. Vol. 1.

Dicots. (ISBN-13: 978-81-920098-2-7, hbk). St. Joseph’s Press, Thiruvananthapuram 695 014, Kerala,

INDIA (Orders: Jawaharlal Nehru Tropical Botanic Garden and Research Institute, Palode, Thiruvana¬

nthapuram 695 562, Kerala, INDIA). $200.00 US, 931 pp., 6 3 / 8 " x 10 W.

T.S. Nayar, M. Sibi, & A. Rasiya Beegam. 2014. The Flowering Plants of The Western Ghats, India. Vol. 2.

Monocots. (ISBN-13: 978-81-920098-3-4, hbk). St. Joseph’s Press, Thiruvananthapuram 695 014,

Kerala, INDIA (Orders: Jawaharlal Nehru Tropical Botanic Garden and Research Institute, Palode,

Thiruvananthapuram 695 562, Kerala, INDIA). $200.00 US, 752 pp., 6 %" x 10 V4".

J.Bot. Res. Inst. Texas 9(1): 234.2015

FLORISTICS AND COMMUNITY ECOLOGY OF

AQUATIC VEGETATION OCCURRING IN SEVEN LARGE SPRINGS AT

OZARK NATIONAL SCENIC RIVERWAYS, MISSOURI (U.S.A.), 2007-2012

David E. Bowles and Hope R. Dodd

National Park Service

Heartland Inventory & Monitoring Network

c/o Department of Biology

Missouri State University

901 South National Avenue

Springfield, Missouri 65897, U.S.A.

david_bowles@nps.gov; hope_dodd@nps.gov

ABSTRACT

We studied the aquatic vegetation communities of seven large springs located at Ozark National Scenic Riverways, Missouri, 2007-2012.

This study provides a baseline for assessing the impacts of anthropogenic and natural disturbances on the ecological integrity of springs.

Vegetation was assessed quantitatively along sample cells located on multiple transects in each spring. We list 69 distinct taxa with substan¬

tial overlap of species occurrences among springs, including 6 families, 6 genera, and 6 species of algae, and 9 families, 12 genera, and 19

species of mosses and liverworts. Among angiosperms, we report 10 families, 13 genera, and 20 species of monocots, and 16 families, 23

genera, and 24 species of dicots. Some species previously reported from the springs are no longer present, which may be due to long-term

changes in watershed condition. Individual sample cells typically contained four to six taxa, although Welch Spring generally had only two

to three taxa represented. Effective numbers of species were generally consistent among years for all springs, but the various species did not

occur in equal abundance in the community within or among sample years. Taxa richness was slightly higher than Simpson’s Diversity In¬

dex effective number (De) and Shannon’s Diversity Index effective number (He) for all years among springs. He ranged from 1.34 to 3.76

among sampling years and springs with values for Alley and Blue springs of approximately 3 while those for Big, Pulltite, Round, and Welch

springs were closer to 2. De ranged from 1.25 to 3.86 among sampling years and springs. Few non-native plant species occurred in the

springs, and they accounted for <15% of the foliar cover across transects.

RESUMEN

Estudiamos las comunidades de vegetacion acuatica de siete grandes manantiales ubicados en Ozark National Scenic Riverways, Missouri,

en 2007-2012. Este estudio proporciona una base para evaluar el impacto de las perturbaciones naturales y antropogenicas en la integridad

ecologica de los manantiales. La vegetacion se evaluo cuantitativamente a lo largo de cuadrantes de muestreo ubicados en varios transectos

en cada manantial Listamos 69 taxones diferentes con una considerable superposicion de presencia de especies entre los manantiales, entre

ellas 6 familias, 6 generos, y 6 especies de algas, y 9 familias, 12 generos, y 19 especies de musgos y hepaticas. Entre las angiospermas, regis-

tramos 10 familias, 13 generos, y 20 especies de monocotiledoneas, y 16 familias, 23 generos, y 24 especies de dicotiledoneas. Algunas espe¬

cies previamente reportadas en los manantiales ya no estan presentes, lo que puede deberse a cambios a largo plazo en la cuenca. Los

cuadrantes de muestreo individuales tipicamente contuvieron de cuatro a seis taxones, aunque Welch Spring en general solo tuvo de dos o

tres taxones representados. Los numeros efectivos de especies fueron generalmente consistentes entre anos para todos los manantiales, pero

las diferentes especies no tuvieron una abundancia igual en la comunidad en o entre anos de muestreo. La riqueza de taxones fue ligera-

mente superior al numero efectivo del Indice de Diversidad de Simpson (De) y al numero efectivo del Indice de Diversidad de Shannon (He)

para todos los anos entre los manantiales. He vario de 1.34 a 3.76 entre los anos de muestreo y los manantiales, con valores para los manan¬

tiales Alley y Blue de aproximadamente 3, mientras que los de los manantiales Big, Pulltite, Round, y Welch fueron cercanos a 2. De oscilo

entre de 1.25 a 3.86 entre anos de muestreo y manantiales. Pocas especies de plantas no nativas estuvieron presentes en los manantiales, las

cuales representaron < 15% de la cubierta foliar en los transectos.

Keywords: aquatic vegetation, floristics, community ecology, Ozarks, springs

INTRODUCTION

Mosses, algae, and higher plants are particularly important structural and biological constituents of springs

and aquatic systems in general (Hannan & Dorris 1970; Cushing & Wolf 1984; Carpenter & Lodge 1986;

Durarte & Canfield 1990; Stream Bryophyte Group 1999; Cantonati et al. 2006), and they often have complex

J. Bot. Res. Inst. Texas 9(1): 235 - 249.2015

236

Journal of the Botanical Research Institute of Texas 9(1)

relationships with some fish and aquatic invertebrates (Cyr & Downing 1988; Xie et al. 2005, 2006). Addition¬

ally, aquatic plants are important biological biters of a variety of chemical contaminants and nutrients (Demars

& Harper 1998; Cantonati et al. 2006; Vardanyan & Ingok 2006). Aquatic vegetation communities in spring

ecosystems are vulnerable to a broad variety of anthropogenic disturbances (Sanford 1979; Englund 1991;

Schutz 1995; Preston et al. 2003). Land use, particularly vegetation clearing practices with associated increases

in sediment and nutrient loads and other point and nonpoint inputs to surface water, has been reported as the

greatest long-term threat to streams and springs in the Ozark Highlands (Duchrow 1977; Jacobson & Primm

1997; Scott & Udouj 1999; Imes & Frederick 2002). Recent studies of the possible effects of global climate

change indicate that increases in annual average water temperature (Hogg & Williams 1996; Poff et al. 2002;

Johnson 2007), changes in discharge (Erman & Erman 1995), and non-native (i.e., exotic) species introduc¬

tions (Bowles & Bowles 2013) may impact spring communities as well. Therefore, aquatic vegetation surveys

are important to researchers and managers to establish inventories and aid in the diagnosis of ecosystem dis¬

turbances. Monitoring changes in aquatic vegetation has long been used as an indicator of anthropogenic dis¬

turbance throughout Europe. This approach has received little attention in the United States (Romero &

Onaindia 1995; Tremp & Kohler 1995; Small et al. 1996; Bartodziej & Ludlow 1997; Ali et al. 1999; Schorer et

al. 2000; Bernez et al. 2001, Bernez 2004; Haury et al. 2002; Scott et al. 2002; Daniel et al. 2005; Haslam 2006).

Documenting the occurrence and distributions of bora in springs provides a baseline for assessing the poten¬

tial for, or extent of, a variety of both anthropogenic and natural disturbances and is an integral step toward

their conservation and understanding their role in ecosystem function.

The extensive dolomitic karst topography in the area surrounding Ozark National Scenic Riverways

(OZAR), Missouri, is conducive to the formation of springs (Mugel et al. 2009). Within the karst terrain of

OZAR there are more than 425 springs. Although most of these springs are relatively small, several are debned

as 1 st and 2 nd magnitude, having minimum discharges of 2.8 m 3 /sec and 0.28 m 3 /sec, respectively (Meinzer

1927). The largest (Big Spring) has a maximum recorded discharge greater than 36 m 3 /sec and is ranked as one

of the bve largest springs in North America (Brune 1981). Springs at OZAR are highly vulnerable to biological

and chemical degradation and may represent the ultimate challenge in water quality protection (Gibert 1990;

Imes & Fredrick 2002). Despite the large amount of literature addressing groundwater contamination and its

complexities, there is little information on the physical and biological impacts of contaminated groundwater

within springs or within surface water fed by springs (Williams 1991; Notenboom et al. 1994; van der Kamp

1995).

Prior to this study, no long-term biomonitoring programs for aquatic vegetation were conducted for the

springs at OZAR, although there had been some short-term studies of algae and aquatic vegetation completed.

Drouet (1933) reported on the algae found in Big, Alley, and Round springs in addition to listing a few records

for vascular plants. Steyermark (1941) conducted the brst known inventory of aquatic plants in Missouri

springs, including those assessed in this study, and he further corrected several misidentibcations made by

Drouet (1933). Lipscomb (1969) conducted a boristic inventory of the aquatic plants at eight springs at OZAR,

and Currier (1990a, b) conducted boristic inventories at four springs. Redfearn et al. (without date) conducted

a botanical survey of OZAR and listed some aquatic species although they did not focus on the springs. Red¬

fearn (1981) addressed aquatic mosses and liverworts known to occur in Ozark springs, including those at

OZAR. Converse (1994) conducted a quantitative study of hydrophyte production at Big Spring, but this work

was never published. Most of these previous studies, described above, provided only qualitative data and no

effort was made to quantify species densities.

To address this lack of knowledge of anthropogenic and natural disturbance effects on springs, we began

a spring monitoring program in 2007 to provide a baseline for assessing the ecological integrity of springs at

OZAR, including the aquatic vegetation community (Bowles et al. 2008). This paper includes a boristic inven¬

tory of the aquatic vegetation of OZAR springs and provides an analysis of plant community diversity dynam¬

ics, 2007-2012.

Bowles and Dodd, Aquatic vegetation communities at Ozark National Scenic Riverways, Missouri

237

METHODS

Study sites .—Aquatic vegetation was assessed annually at each of six 1 st or 2 nd magnitude springs (Meinzer

1927; Vineyard et al. 1974) located at OZAR (Fig. 1), during late July-early August from 2007 to 2012. An ad¬

ditional 2 nd order spring (Phillips Spring) was sampled in 2011 and 2012. Banks immediately adjacent to the

spring branches generally are well-vegetated, undisturbed, and stable, although some of the springs have well-

worn walking paths along at least one bank and/or managed lawns in their general vicinity. The riparian can¬

opy for all the springs is generally less than 40%. Among springs, temperature ranged from 13.5 to 14.5 °C,

dissolved oxygen concentration ranged from 8.2 to 11.1 mg/L, specific conductance ranged between 200 and

350 pS/cm, and pH fell between 7.2 and 7.6 at time of sampling. Turbidity was typically at low levels in the

springs (<1 NTU) during sampling but was around 4 NTU at Big Spring in 2012 due to an apparent collapse

within the spring conduit that caused the water to appear milky. Run habitat, defined here as deep, fast water

with little or no turbulence, was dominant in all springs. Dominant substrate for all of the springs varied but

generally consisted of large pebble to small cobble (32-90 mm). The substrate of Big and Round springs con¬

tained smaller mean substrate sizes, including a higher percentage of sands, compared to the other springs.

Discharge among springs during sampling in 2007-2012 varied widely: Alley Spring (2.21-3.80 m 3 /sec), Big

Spring (9.85-13.93 m 3 /sec), Blue Spring (2.19-3.97 m 3 /sec), Pulltite Spring (0.37-2.21 m 3 /sec), Round Spring

(0.50-2.99 m 3 /sec), Welch Spring (4.01-8.41 m 3 /sec). Discharge for Phillips Spring was 0.58 m 3 /sec in 2011 and

0.61 m 3 /sec in 2012. A further assessment of the physical habitat is not included here.

Reach selection .—The portion of the spring run to be sampled was unique for each spring due to their re¬

spective sizes and other physical characteristics. One reach was sampled in each spring run and began as close

to the spring source as practical, taking into consideration depth, flow, and crew safety near the spring source.

Reach length was based on a weighting factor that accounted for variation among average widths to ensure a

uniform sampling effort that represented each spring’s total area, and facilitated inclusion of representative

macrohabitats (Bowles et al. 2008). Reach lengths were Welch Spring (36 m), Phillips and Pulltite springs (150

m each), Blue Spring (160 m), Alley Spring (190 m), Round Spring (240 m), and Big Spring (540 m). The sam¬

pling reach was divided into 11 equally spaced transects perpendicular to flow with three sample cells (1 m 2 )

per transect. The only exception to this was Welch Spring, where only three transects were established because

of the short spring run (36 m) from the source to the confluence with the Current River. Transects defined the

sample locations with physical habitat sampled at each transect (N = 33; 9 Welch Spring) and aquatic vegeta¬

tion sampled at odd numbered transects (six transects, N = 18; 9 Welch Spring) using a PVC sampling frame

and view bucket to observe vegetation. The sampling frame was divided into quadrats to facilitate assessing

vegetation abundance. Plant species foliar cover in each sample cell was recorded using a modified Dauben-

mire scale: 1 = 0-0.99%, 2 = 1-5%, 3 = 5-25%, 4 = 25-50%, 5 = 50-75%, 6 = 75-95%, 7 = 95-100% (Dauben-

mire 1959; Bowles et al. 2008).

Data summary and analysis .—Data collected from all sample cells were summarized by taxa for the area

sampled. Mean values along with a measure of variability (± 1 standard error of the mean) were calculated for

the community. Foliar cover estimates for each species were used to determine measures of species diversity

(McCune et al. 2002).

Foliar cover data were analyzed using the midpoints of the respective cover classes and were analyzed

under two broad categories: 1) diversity and 2) individual species frequency. Diversity analyses included taxa

richness, Shannon’s Diversity Index (Shannon 1948), and Simpson’s Diversity Index (Simpson 1949), which

were subsequently expressed as effective number of species (S, He, De) (Hill 1973; Joust 2006; Bowles et al.

2008). Effective number of species for each diversity measure reflects the number of species found in a similar

community when all species occur in equal density (Bowles et al. 2008). If all species occurred in equal abun¬

dance in the community within and among sample years, then taxa richness, Shannon’s Diversity Index, and

Simpson’s Diversity Index would all be equal (Washington 1984). The advantage of converting Shannon’s Di¬

versity Index and Simpson’s Diversity Index to effective number of species is that it makes their values linear

thus allowing for direct comparison.

238

Journal of the Botanical Research Institute of Texas 9(1)

MONTAUK

Metrics calculated for individual species included individual species frequency, percent foliar cover, and

species importance value (Bowles et al. 2008). Species importance values are a measure of the relative domi¬

nance of species in a spring community—the larger the importance value, the more important the species. We

also calculated percent exotic taxa richness and the ratio of percent foliar cover of exotic species to percent fo¬

liar cover of all species. This latter metric serves as a measure of the relative dominance of non-native species

and thus serves as means of assessing community disturbance.

Specimen processing and voucher specimens .—Most angiosperms were pressed and mounted on herbarium

sheets following guidance issued by the University of Florida Herbarium (2015). Representative Lemnaceae

were stored in 4% formalin. Bryophytes were air dried and stored in folded paper packets made from 100% cot¬

ton fiber paper. Identification keys used included Redfearn (1972), Godfrey and Wooten (1979, 1981), Flora of

North America (1993+), and Yatskievych (1999, 2006, 2013). Voucher specimens are deposited in the National

Park Service, Heartland Inventory & Monitoring Aquatic Program reference collection (HTLN) located at the

Bowles and Dodd, Aquatic vegetation communities at Ozark National Scenic Riverways, Missouri

239

Department of Biology, Missouri State University, Springfield, Missouri. Other reference species are deposited

in the Ozarks Regional Herbarium (SMS) located in the Department of Biology, Missouri State University,

Springfield, Missouri. Common names follow USDA, NRCS (2014).

RESULTS

Floristic study .—A broad assemblage of aquatic vegetation occurs among the springs at OZAR with 69 distinct

taxa reported from this study. A complete list of plant species recorded from transects in each spring is in the

Appendix. We recorded 5 families, 5 genera, and 5 species of algae. We also found modest amounts of filamen¬

tous green algae (Chlorophyta) at all of our sites, but it was not identified to a more specific taxonomic level. We

report 9 families, 13 genera, and 19 species of mosses and liverworts. Many of these were reported by Redfearn

(1972) and were not identified by us. Among angiosperms, we list 10 families, 13 genera, and 20 species of

monocots, and 16 families, 23 genera, and 24 species of dicots. Star duckweed ( Lemna trisulca) was found at all

springs except Pulltite and Welch springs. This species has a S2 state heritage ranking, indicating it is imper¬

iled because of rarity or other factors, thus making it vulnerable to extirpation in Missouri. The environmen¬

tally sensitive red alga, Batrachospermum, occurs in all of the springs (Barinova 2013).

We found examples of Elodea canadensis and Elodea nuttallii in most of the springs. It was not practical to

distinguish these two taxa in the vegetative state while doing the community assessment. Based on flowering

specimens we found, it appears that E. canadensis is the more abundant of the two species. Additionally, Yatski-

evych (1999) noted that specimens intermediate between these two species are known to occur in the region,

suggesting they may produce hybrids. Our observations support that notion, and we treat the two species at

the generic level for the community analysis.

Two closely related species, Veronica anagallis-aquatica and Veronica catenata, occur within the range of

the springs we sampled. These two species can be separated most easily on the basis of leaf size and shape.

Yatskievych (2013) considers V anagallis-aquatica to be non-native, whereas V catenata is native. Although V

catenata commonly occurs as a submerged aquatic, Yatskievych (2013) noted that the vegetative plants of V

anagallis-aquatica also may occur as submerged aquatics and would be morphologically indistinguishable

from those of V catenata. Vegetative forms of Veronica occurred in all of the springs we sampled, but based on

reproductive forms of Veronica we found, we have identified this species as V catenata. It is possible that hy¬

brids between these two species may exist (Yatskievych 2013). In contrast, USDA, NRCS (2014) treats these

two species as synonyms with V catenata being a junior synonym of V anagallis-aquatica, and further consid¬

ers this species to be native.

Few non-native (exotic) plants occurred in the springs. The most commonly occurring non-native plant

was water-cress ( Nasturtium officinale), but other introduced species included annual bluegrass ( Poa annua),

watermint ( Mentha aquatica), creeping bentgrass ( Agrostis stolonifera), creeping jenny ( Eysimachia nummu-

laria), and bitter dock ( Rumex obtusifolius). It is possible that the species we identify as M. aquatica may be a

hybrid of that species and Mentha spicata L. (=M. x piperita L.) as described by Yatskievych (2013), but we were

unable to make that determination. Although all of these species are considered terrestrial or wetland plants,

they grow in the spring runs at OZAR where some specimens are completely submersed. Invasive species, in¬

cluding Myriophyllum spicatum are present in the Current River basin (Padgett 2001, personal observations

both authors) and may eventually enter the springs, especially in areas having slow current velocities.

Unusual findings include examples of fully submersed terrestrial or wetland species including Eobelia

cardinalis, Eysimachia nummularia, Mentha aquatica, Physostegia virginiana, Poa annua, and Rumex obtusifolius.

We have found examples of R. obtusifolius at Big Spring near mid-channel (~20 m from bank) in water approxi¬

mately 1.5 m deep. Physostegia virginiana grows completely submersed at Blue Spring where plants are distrib¬

uted over a 15 m patch located on the right side of the channel downstream of the spring source in water ap¬

proximately 1 m deep. The present distribution of P. virginiana at Blue Spring is in the same general area as that

reported by Steyermark (1941). We also found a small clump of P. virginiana growing submersed at Alley

Spring, but that clump subsequently disappeared. At Phillips Spring, L. cardinalis produces inflorescences that

240

Journal of the Botanical Research Institute of Texas 9(1)

extend above the surface of the water at mid-channel while the stems and leaves remain submersed. Lysimachia

nummularia and M. aquatica are commonly encountered along the margins of the springs at OZAR and we

commonly find submersed plants of both species. Poa annua can be found at many springs in the Ozarks region

where it grows submersed.

Several species previously reported from some springs (Steyermark 1941; Currier 1990a, b) were not

found during this study. These species and the springs from which they were formerly known include: three

species of Juncus (Round, Welch); creeping bentgrass, Agrostis stolonifera (Blue); grassleaf mudplantain, Heter-

anthera dubia (Big, Blue); star duckweed, Lemna trisulca (Pulltite); horned pondweed, Zannichellia palustris

(Big, Round, Welch; now only found at Blue where it is rare); Illinois pondweed, Potamogeton illinoensis (Alley,

Big, and Welch); long beak buttercup, Ranunculus aquatilis (Alley, missing at Welch in 2008 and 2009); and

leafy pondweed, Potamogeton foliosus (Alley, Big, Pulltite). The community data we present are based on tran¬

sect data, but a thorough visual inspection of the areas between transects and areas above and below the sam¬

ple reaches did not locate these species. Numerous other species of plants occurred in the springs at OZAR, but

they were not present in sample cells (see Appendix).

Community diversity. —The highest richness among springs based on transect data from all years com¬

bined was at Round Spring, where 20 taxa were found, whereas Alley and Blue springs each had 19 taxa. Sev¬

enteen taxa were found across transects at Big Spring, whereas Pulltite, Welch, and Phillips had the fewest taxa

(15,13, and 11 taxa, respectively). Individual sample cells typically contained four to six taxa, although Welch

Spring generally had only two to three species represented. In general, Welch Spring shared the fewest species

in common with the other springs, which may be due to its short (36 m) spring run, smaller sample size (3

transects), and high current velocities that limit vegetation growth in the mid-channel. Because the Welch

spring run is short, we were able to visually examine most of the entire bottom, and based on those observa¬

tions, the low taxa richness estimates for that spring are accurate. Foliar coverage of aquatic vegetation was

extensive in all of the springs, with bare substrate occupying 20% or less of sample cells on average. Effective

numbers of species (S, He, and De) were generally consistent among years for all springs (Fig. 2 a-f), and the

various species did not occur in equal abundance in the community within and among sample years. Taxa

richness was higher than De and He for all years among springs, and the difference between those values was

around two fold or less in most cases. The Mean Shannon’s Index effective number (He) ranged from 1.34 to

3.76 among sampling years and springs, with values for Alley and Blue springs approximately 3 while those for

Big, Pulltite, Round, and Welch springs were generally closer to 2. Simpson’s Index effective numbers (De)

ranged from 1.25 to 3.86 among sampling years and springs. Similar to the values recorded for He, De values

were highest for Alley and Blue springs compared to the other springs. At Phillips Spring, taxa richness during

2011 and 2012 was 3.11 (±0.59) and 2.22 (±0.11); He was 2.22 (±0.32) and 1.73 (±0.02); and De wasl.96 (±0.28)

and 1.58 (±0.04).

Mosses and algae occurred most frequently and had the greatest percent foliar cover among all springs

except for Big and Round springs (Figs. 3-4). Nasturtium officinale and Cardamine bulbosa had the highest In¬

dividual Species Frequency (ISF) and Percent Foliar Cover (PFC) at Alley Spring (Fig. 3a). At Big Spring, Ra¬

nunculus aquatilis had the highest ISF and PFC (Fig. 3b). Lemna trisulca was more frequently encountered

compared to the other flowering species at Blue Spring, but the PFC for all angiosperms was relatively low (Fig.

3c). At Pulltite Spring, Persicaria hydropiperoides and N. officinale were the most common flowering plants (Fig.

3d), while several taxa were co-dominant at Round Spring (Fig. 4a). Myriophyllum heterophyllum and N. offici¬

nale were the most dominant angiosperms encountered at Welch Spring (Fig. 4b). Only six angiosperms were

recorded on transects at Phillips Spring and all had relatively low PFC values (Fig. 4c). Mean Species Impor¬

tance Values (SIV) among springs clearly showed that some taxa were more important than others, which is

not always reflected by their ISF and PFC values when considered independently (Table 1). Collectively,

mosses and algae had the highest SIVs at all springs. Among angiosperms, N. officinale and C. bulbosa had the

highest SIVs at Alley Spring (SIV = 14.24 and 13.96, respectively). Ranunculus aquatilis was the most important

species at Big Spring (SIV = 38.89) followed by N. officinale (SIV = 17.09). At Blue Spring, C. bulbosa, L. trisulca,

Bowles and Dodd, Aquatic vegetation communities at Ozark National Scenic Riverways, Missouri

241

♦S ♦He ♦De

2007 2008 2009 2010 2011 2012

♦S ♦He *De

-A-S-®-He-»-De

♦SR ♦He ♦De

♦S ♦He ^De

♦S ♦He *De

Fig. 2. Mean and standard error of effective numbers of species for aquatic vegetation occurring in six large springs at Ozark National Scenic Riverways,

Missouri. S = taxa richness. He and De = effective number of species for Shannon's Diversity Index and Simpson's Diversity Index, respectively: (a) Alley

Spring, (b) Big Spring, (c) Blue Spring, (d) Pulltite Spring, (e) Round Spring, (f) Welch Spring.

and Veronica catenata had the highest SI Vs (9.13, 7.00, and 4.41, respectively). Persicaria hydropiperoides was the

most important species at Pulltite Spring (SIV = 8.16) followed by Elodea spp. (SIV = 5.04). At Round Spring, M.

heterophyllum, Sparganium americanum and Potamogeton illinoensis shared similar importance values (SIV =

16.33, 15.34, and 14.82, respectively). Welch Spring had M. heterophyllum and N. officinale as the most impor¬

tant species (SIV = 20.70 and 12.49, respectively), and N. officinale also had the highest SIV at Phillips Spring

(SIV = 13.71).

Non-native species were poorly represented among all springs and their percent foliar cover was <15%

across transects. The most commonly encountered and abundant non-native species was Nasturtium officinale,

while Mentha aquatica , Poa annua , and Rumex obtusifolius were found less frequently. Non-native species gener¬

ally had low SIVs, but N. officinale was the most important species at Alley and Phillips springs and among the

most important species at Big, Welch, and Pulltite springs. Although N. officinale was commonly encountered

in the spring runs, the majority of the foliar coverage for this species was near the wetted margins out of the

main spring flows. The individual plants of N. officinale we encountered in the faster spring flows of the main

channels typically had low percent foliar cover.

242

Journal of the Botanical Research Institute of Texas 9(1)

Fig. 3. Individual Species Frequency (ISF) and Percent Foliar Cover (PFC) for aquatic vegetation occurring in (a) Alley, (b) Big, (c) Blue, and (d) Pulltite

springs at Ozark National Scenic Riverways, Missouri, 2007-2012.

DISCUSSION

Because of their unique and often independent sources, springs typically have a mosaic structure, a high de¬

gree of individuality, and typically an azonal character attributed to their physiochemical stability (Williams

& Danks 1991; Cantonati et al. 2006). The monitoring results presented here generally support that assess¬

ment. The aquatic plant communities of the springs we studied are diverse and dynamic, and they share many

species in common. In general, mosses and algae were the dominant taxa among most springs. The lack of

Bowles and Dodd, Aquatic vegetation communities at Ozark National Scenic Riverways, Missouri

243

Fig. 4. Individual Species Frequency (ISF) and Percent Foliar Cover (PFC) for aquatic vegetation occurring in (a) Round, (b) Welch, and (c) Phillips springs

at Ozark National Scenic Riverways, Missouri, 2007-2012.

dominance by mosses at Big and Round springs may be due to these systems having a predominance of sand

substrates compared to gravel and cobble substrates found in the other springs.

The diversity of aquatic vegetation in the springs reported here was not unexpected. The findings of Stey-

ermark (1941), Lipscomb (1969), and Currier (1990a, b) suggest aquatic plant community structure in these

springs is dynamic. Currier (1990a, b) reported that the plant community structure in several springs had

changed since Steyermark’s study (Steyermark 1941), with some species apparently disappearing and others

documented for the first time. For example, Steyermark (1941) reported horned pondweed ( Zannichelliapalus-

tris), leafy pondweed ( Potamogeton foliosus ), and submerged stands of the non-native annual bluegrass ( Poa

annua) at Big Spring, but Currier (1990b) did not find those species. In contrast, Lipscomb (1969) found these

three species were present at Big Spring. Our study found P. annua in Big Spring, but no representatives of leafy

pondweed were detected. Steyermark (1941) did not record star duckweed (Lemna trisulca) or longbeak but¬

tercup ( Ranunculus aquatilis) from Big Spring, but these species are now present and abundant there (Lipscomb

1969; Currier 1990b; this study). Our data show that some species, such as swamp smartweed ( Persicaria hy-

dropiperoides) at Pulltite Spring, have made substantial increases in their foliar coverage since monitoring was

initiated while others have decreased. Steyermark (1941) reported Heteranthera dubia from the lower portion of

Blue Spring, whereas Lipscomb (1969) reported this species from near the confluence of the spring with the

244

Journal of the Botanical Research Institute of Texas 9(1)

Table 1. Mean species importance values for aquatic vegetation at large springs located at Ozark National Scenic Riverways, Missouri.

Alley

Big

Blue

Pulltite

Round

Welch

Phillips

Botrochospermum spp.

7.18

6.87

1.60

4.06

2.17

4.70

8.83

Filamentous green algae

8.68

6.50

14.50

16.41

8.77

17.03

7.15

Filamentous blue-green algae

—

2.19

0.51

—

3.89

9.03

Nostoc sp.

5.34

4.13

5.50

7.24

1.82

1.81

21.93

Mosses

24.44

14.02

34.02

40.68

7.94

42.01

27.68

Riccia fluitans

—

0.28

—

Collitriche heterophyllo

7.59

3.49

2.89

1.44

1.14

1.12

—

Cardamine bulbosa

13.96

4.88

9.13

0.30

0.70

—

2.96

Cerotophyllum demersum

—

4.55

—

Eleocharis acicularis

0.76

0.20

—

0.15

—

Elodeo spp.

5.15

6.62

1.06

5.04

10.24

0.58

0.36

Glyceria striata

4.19

2.58

3.04

0.18

—

Lemna minor

2.55

1.23

—

0.12

0.46

—

Lemna trisculca

6.58

4.53

7.00

—

Lobelia cardinalis

—

1.83

Ludwigia palustris

—

0.61

—

Myriophyllum heterophyllum

—

16.33

20.70

—

Mentha aquatica

0.50

—

Nasturtium officinale

14.24

17.09

1.71

4.12

5.69

12.49

13.78

Physostegia virginiana

2.20

—

1.65

—

Poa annua

—

2.72

0.20

—

3.04

—

0.74

Persicaria hydropiperoides

—

0.45

—

8.16

0.15

—

Potamogeton illinoensis

0.97

—

0.85

—

14.82

—

Potamogeton nodosus

—

0.35

—

Ranunculus aquatilis

0.30

38.89

2.48

1.58

1.06

3.01

—

Rumex obtusifolius

0.59

—

Sparganium americanum

2.40

1.81

3.00

4.26

15.34

4.91

5.71

Veronica catenata

3.99

3.02

4.41

3.77

3.56

3.78

—

Zannichellia palustris

—

0.23

—

Current River. In contrast, Currier (1990a) did not report H. dubia from Blue Spring, and we only found a few

examples below its confluence with the Current River. The extant conditions of the watersheds of the springs

have changed appreciably since Steyermark began his floristic study of Ozark springs in 1928 (Steyermark

1941). The historic Ozark forest was largely deforested by the late 1920s resulting in heavy gravel and silt loads

entering waterways (Benac & Flader 2004), in addition to lessening recharge times into the source aquifer.

Such impacts could have altered their vegetation communities at the time Steyermark studied them. Forest

cover has now largely recovered, but it remains unclear how close the extant vegetation communities of the

springs are to their historic conditions. It is possible that “weedy” species such as P.foliosus and H. dubia may

no longer occur in the springs because those habitats have become more stable as the landscape has recovered.

Conversely, species such as L. trisulca and R. aquatilis now find those habitats favorable.

The data presented here reflect the broad natural habitat diversity, and physical and chemical stability in

these springs. Geographic location of the springs does not appear to be a primary contributing factor in the

composition of their respective vegetation communities. Rather, watershed-level characteristics (recharge

zone), local physical habitat factors including substrate size, canopy cover, and other unknown contributing

factors may play important yet undetermined roles in structuring these communities. However, collecting and

analyzing additional monitoring data is required to determine the importance and relative contribution of

these environmental factors. Additional monitoring also will allow insight into any patterns that may exist in

plant community structure among these springs. Furthermore, our data will be useful in gaging impacts to the

springs from anthropogenic disturbances, potentially ensuring their protection.

Bowles and Dodd, Aquatic vegetation communities at Ozark National Scenic Riverways, Missouri

245

APPENDIX

Springs: 1 =Alley Spring, 2=Big Spring, 3=Blue Spring, 4=Phillips Spring, 5=Pulltite Spring, 6=Round Spring, 7=Welch Spring.

Growth Habit Types: Emergent=E, Floating=F, Submersed=S, Wetted margin=W

Non-native species are designated by an asterisk (*)

Collector/ldentifier: D.E. Bowles, unless noted otherwise.

Collection number: (HTLN#)

ALGAE (5 families, 5 genera)

Batrachospermaceae

Botrochospermum spp.: 1, 2, 3,4, 5,6, 7. S (HTLN 072)

Characeae

Nitello ocuminoto A. Braun ex Wallman: 2,3. S (HTLN 083)

Nostocaceae

Nostoc sp.: 1,2, 3,4, 5,6, 7 (HTLN 078). S

MOSSES, LIVERWORTS

Amblystegiaceae

Hygroamblystegium fluviotile (Hedw.) Loeske (streamside hygroam-

blystegium moss): 2, 5,6, 7 (Redfearn 1972,1981).

Hygroamblystegium noterophilum (Sull. & Lesq.) Warnst. (hygroam¬

blystegium moss): 3 (Redfearn 1972,1981).

Hygroamblystegium tenax (Hedw.) Jenn. (hygroamblystegium

moss): 1,2, 3 (Redfearn 1972,1981).

Leptodictyum riparium (Hedw.) Warnst. (streamside leptodictyum

moss): 1,3, 5,6, 7 (Redfearn 1972,1981).

Brachytheciaceae

Brachythecium rivulare Schimp. (brachythecium moss): 1,2,3,5,6,

7 (Redfearn 1972,1981).

Platyhypnidium riparoides (Hedw.) Dix. (platyhypnidium moss): 2

(Redfearn 1972,1981).

Fontinalaceae

Fissidens bryoides Hedw. (bryoid fissidens moss): 1, 2, 3 (Redfearn

1972,1981).

Fissidens fontanus (Bach. Pyl.) Steud. (fissidens moss): 1,2, 3 (Red¬

fearn 1972,1981).

Fissidens filiformis Sull. & Lesq. (fontinalis moss): 3 (Redfearn 1972,

1981).

Fissidens grandifrons Brid. (largeleaf fissidens moss): 1, 2, 3, 5, 6

(Redfearn 1972,1981).

Oscillatoriaceae

Lyngbya sp.: 3,4, 5 (HTLN 081). S

Xanthophyceae

Vaucheria sp.: 1,3, 5, 6 (HTLN 080). S

Chlorophyta

unidentified filamentous green algae: 1,2,3,4,5,6,7 (HTLN 079).

(9 families, 13 genera)

Fontinalis hypnoides Hartm. var. duriaei (Schimp.) Husn. (fontinalis

moss): 1,2, 3, 5,6, 7 (Redfearn 1972,1981).

Fontinalis missourica Cardot (Missouri fontinalis moss): 3 (Redfearn

1972,1981).

Pottiaceae

Hyophila involuta (Hook.) A. Jaeger (hyophila moss): 1, 2, 5, 7

(Redfearn 1972,1981).

Thuidiaceae

Cyrto-hypnumpygmaeum (Schimp.) W.R. Buck& H.A. Crum (pygmy

cyrto-hypnum moss): 2, 5 (Redfearn 1972,1981).

Conocephalaceae

Conocephalum conicum (L.) Dumort. (liverwort): 2, 3,4, 5,6 (HTLN

085).

Marchantiaceae

Dumortiera hirsuta (Sw.) Nees (liverwort): 2, 3, 5 (Redfearn 1981,

Timme & Redfearn 2011).

Marchantiapolymorpha L. (liverwort): 1,2,3,6, 7 (HTLN 084).

Porellaceae

Porella pinnata L. (liverwort): 1, 2, 3,4, 7 (Redfearn 1981, Timme &

Redfearn 2011) (HTLN 086).

Ricciaceae

Riccia fluitans L. (liverwort): 4, 7 (HTLN 073). F, S

VASCULAR PLANTS

MONOCOTS (10 families, 13 genera)

Alismataceae

Alismasubcordatum Raf. (American water plantain): 2 (HTLN 063). E

SagittarialatifoliaWiWd. (broadleafarrowhead):2,4,6 (HTLN 004). E

Cyperaceae

Eleocharis acicularis (L.) Roem. & Schult (needle spikerush): 2, 3,4,

5,6 (HTLN 011). E, S

Hydrocharitaceae

Elodea canadensis Michx. (Canadian waterweed): 1, 2, 3, 4, 5, 6, 7

(HTLN 015). S

Elodeanuttallii (Planch.) H.St.John (western waterweed): 1,2,3,4,

5,6, 7 (HTLN 014,089). S

Juncaceae

Juncus acuminatus Michx. (tapertip rush): 6,7 (Steyermark 1941). E

Juncus c/ud/ey/Wiegand (Dudley's Rush): 7 (Steyermark 1941). E

Juncus tenuis Willd. (poverty rush): 7 (Steyermark 1941). E

Lemnaceae

Lemna minor L. (common duckweed): 1,2,3, 5,6 (HTLN 069). F

Lemna minuta Kunth (least duckweed): 6 (HTLN 076). F, S

Lemna trisulca L. (star duckweed): 1,2, 3,5 (HTLN 077). S

Poaceae

Agrostis stolonifera L. (creeping bentgrass)*: 3 (Steyermark 1941).

E, W

Glyceria striata (Lam.) Hitchc. (fowl manna grass): 1, 3, 4, 6 (HTLN

027). E, S,W

Poa annua L. (annual blue grass)*: 1,2,4,6, 7 (HTLN 028). E, S, W

Pontederiaceae

Heteranthera dubia (Jacq.) MacMill. (grassleaf mudplantain): 2, 3

(Steyermark 1941; HTLN 033). S

Potamogetonaceae

Potamogeton foliosus Raf. (leafy pondweed): 1,2,5,6 (HTLN 034). S

Potamogeton illinoensis Morong (Illinois pondweed): 1, 2, 3, 4, 6,

7 (HTLN 036). S

Potamogeton nodosus Poir. (longleaf pondweed): 6 (HTLN 095). S

Sparganiaceae

Sparganium americanum Nutt. (American bur-reed): 1, 2, 3,4, 5, 6

(HTLN 041). E, S

Zannichelliaceae

Zannichelliapalustris L. (horned pondweed): 2,3,6,7 (HTLN 082). S

246

DICOTS (16 families, 23 genera)

Acanthaceae

Justicio omericono (L.) Vahl (American water-willow): 6 (HTLN

087). E, S

Balsaminaceae

Impatiens capensis Meerb. (jewelweed): 1, 2, 3, 4, 5, 6, 7 (HTLN

008). W

Brassicaceae

Cardamine bulboso (Schreb. ex Muhl.) Britton, Sterns & Poggenb.

(bulbous bittercress): 1,2, 3,4, 5,6, (HTLN 006). S

Nasturtium officinale\N.J. Aiton (watercress)*: 1,2,3,4,5,6,7 (HTLN

007). E, F, S, W

Callitrichaceae

Callitriche heterophylla Pursh (twoheaded water-starwort): 1, 2, 3,

5,6, 7 (HTLN 009). S

Campanulaceae

Lobelia cardinalis L. (cardinalflower): 4 (HTLN 060). E, S, W

Ceratophyllaceae

Ceratophyllum demersum L. (coon's tail): 1,2,6 (HTLN 010). S

Crassulaceae

Penthorum sedoides L. (ditch stonecrop): 3 (HTLN 062). W

Haloragaceae

Myriophyllum heterophyllum Michx. (twoleaf water milfoil): 1, 2, 5,

6, 7 (HTLN 012). S

Myriophyllum spicatum L. (Eurasian watermilfoil)*: 1 (backwater

only), (HTLN 052). S

Lamiaceae

Lycopus rubellus Moench (taperleaf water horehound): 3 (HTLN

047). E, W

Journal of the Botanical Research Institute of Texas 9(1)

Mentha aquatica L. (watermint): 1,3,4,6, 7 (HTLN 017)*. E, S, W

Physostegia virginiana (L.) Benth. (obedient plant): 3 (HTLN 065).

E, S,W

Onagraceae

Ludwigia palustris (L.) Elliott (marsh seedbox): 2, 4, 6 (HTLN 021).

E, S,W

Polygonaceae

Persicaria hydropiperoides (Michx.) Small (swamp smartweed): 2,4,

5,6 (HTLN 045). E, S,W

Rumexobtusifolius L. (bitter dock)*: 1,2,3,4,5,6,7 (HTLN 032). E, S, W

Primulaceae

Lysimachia nummularia L. (creeping jenny)*: 2,6 (HTLN 037). S, W

Samolus valerandi L. ssp. parviflorius (Raf.) Hulten (seaside brook-

weed): 4,6 (HTLN 092). W

Ranunculaceae

Ranunculus aquatilis L. (longbeak buttercup, whitewater crowfoot):

1,2,3, 5, 7 (HTLN 037). S,W

Saururaceae

Saururus cernuus L. (lizard's tail): 2,4 (HTLN 039). W

Scrophulariaceae

Linderniadubia (L.) Pennell (false pimpernel): 7 (Steyermark 1941). W

Veronica catenata Pennell (water speedwell): 1, 2, 3, 5, 6, 7 (HTLN

025). E, S,W

Urticaceae

Pilea pumila (L.) A. Gray (Canadian clearweed): 1, 3, 4, 6 (HTLN

043). W

Boehemeria cylindrica (L.) Sw. (smallspike false nettle): 1, 2, 3,4, 5,

6, 7 (HTLN 088). W

ACKNOWLEDGMENTS

We thank Tyler Cribbs, Jan Hinsey, Catherine Ciak, Ryan Green, Myranda Clark, Beth Bailey, Victoria Grant,

Kevin Murray, Melanie Weber, Zach Morris, Chris Morris, and Josh DeLay for assisting us in the held. Victoria

Grant kindly reviewed an earlier draft of this paper. We also thank Michelle Bowe and the Ozarks Regional

Herbarium (SMS) for allowing us access to the collection to verify species. Atilano Contreras-Ramos, Institute

de Biologla, Universidad Nacional Autonoma de Mexico, kindly edited the Spanish abstract. We also thank

George Yatskievych and Chetta Owens who provided valuable, constructive criticism on an earlier draft of this

paper. Views, statements, findings, conclusions, recommendations, and data in this report are solely those of

the author(s) and do not necessarily reflect views and policies of the U.S. Department of Interior, National Park

Service. Mention of trade names or commercial products does not constitute endorsement or recommendation

for use by the National Park Service.

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BOOKS RECEIVED

Alan Graham. 2014. Academic Tapestries: Fashioning Teachers and Researchers Out of Events and Expe¬

riences. MSB 126. (ISBN-13: 9780915279968, pbk). U.S.A. (Orders: www.mbgpress.info, mbgpress@

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Karen V. Root. 2012. Environmental Science Online Textbook. (ISBN-13: 978-0-87893-962-6, pbk). Sinauer

Associates, Inc., Publishers, 23 Plumtree Road, P.O. Box 407, Sunderland, Massachusetts 01375-0407,

U.S.A. (Orders: www.sinauer.com, 1-413-549-4300). $65.00 US for 180-day subscription, online-only

resource.

Tennessee Flora Committee. 2015. Guide to the Vascular Plants of Tennessee. (ISBN-13:1-62190-100-9, hbk).

University of Tennessee Press, Chicago Distribution Center, 11030 South Langley Ave., Chicago, Illinois

60628, U.S.A. (Orders: utpress.org, 1-800-621-2736). $49.95 US, 813 pp., 7" x 10".

Mark D. Bertness, John F. Bruno, Brian R. Silliman, & John J. Stachowicz. 2014. Marine Community Ecology

and Conservation. (ISBN-13: 978-1-60535-228-2, hbk). Sinauer Associates, Inc., Publishers, 23 Plum-

tree Road, P.O. Box 407, Sunderland, Massachusetts 01375-0407, U.S.A. (Orders: www.sinauer.com,

1-413-549-4300). $96.01 US, 560 pp., 8 3 /4" x 11 YV.

John M. Marston, Jade dAlpoim Guedes, & Christina Warinner. 2015. Method and Theory in Paleoethnobota-

ny. (ISBN-13: 978-1-60732-315-0, pbk; ISBN-13: 978-1-60732-316-7, eBook). University Press of Colora¬

do, 5589 Arapahoe Avenue, Suite 206C, Boulder, Colorado 80303, U.S.A. (Orders: www.upcolorado.

com, 1-800-621-2736). $34.95 US (pbk), $27.95 (eBook), 480 pp., 87 illustrations, 6" x 9".

W.D. Stevens, O.M. Montiel, & PH. Raven. 2014. Paleobotany and Biography: A Festschrift for Alan Gra¬

ham in his 80 th Year. MSB 128. (ISBN-13: 9780915279975, hbk). U.S.A. (Orders: www.mbgpress.info,

mbgpress@mobot.org, 1-888-271-1930). $100.00 US, 404 pp., 7" x 10".

Peter A. Thomas. 2014. Trees: Their Natural History, 2nd ed. (ISBN-13: 978-0-52113-358-6, pbk.) Cambridge

University Press, 32 Avenue of the Americas, New York, New York 10013-2473, U.S.A. (Orders: www.

cambridge.org, 1-212-337-5000). $42.99 US, 408 pp., 10 b/w illus., 218 color illus., 7" x 9 U".

J.Bot. Res. Inst. Texas 9(1): 250.2015

CYPERUS GRANITOPHILUS (CYPERACEAE), A GRANITE OUTCROP ENDEMIC,

NEW FOR TEXAS AND OKLAHOMA (U.S.A.)

Robert J. O'Kennon and Kimberly Norton Taylor

Botanical Research Institute of Texas

1 700 University Dr.

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

okennon@brit.org, ktaylor@brit.org

ABSTRACT

Cyperus granitophilus McVaugh (Cyperaceae) has been discovered at Enchanted Rock State Natural Area in Gillespie and Tlano counties in

the Tlano Uplift of Central Texas. Examination of herbarium specimens of the closely related C. squarrosus L. revealed 18 additional collec¬

tions (total of 27) of C. granitophilus from the Llano Uplift of Texas, 10 collections from the Wichita Mountains in Comanche and Greer

counties, Oklahoma, and one collection from the Arbuckle Uplift in Johnston County, Oklahoma. All collections were from regions with

granitic outcrops. These collections represent the first documented occurrences of C. granitophilus in Texas and Oklahoma and are disjunct

approximately 1270 km from the nearest previously documented populations in the Piedmont of Alabama.

RESUMEN

Cyperus granitophilus McVaugh (Cyperaceae) ha sido descubierto en la Enchanted Rock State Natural Area en los condados Gillespie y Llano

en el Llano Uplift de Texas Central. El examen de especimenes de herbario del muy relacionado C. squarrosus L. revelo 18 colecciones adi-

cionales (total de 27) de C. granitophilus del Llano Uplift de Texas, 10 colecciones de las montanas Wichita en los condados Comanche y

Greer, Oklahoma, y una coleccion del Arbuckle Uplift en el condado de Johnston, Oklahoma. Todas las colecciones proceden de regiones

con afloramientos graniticos. Estas colecciones representan las primera ocurrencias documentadas de C. granitophilus en Texas y Oklahoma

y tienen una disyuncion de unos 1270 km desde las poblaciones documentadas mas proximas en el Piedmont de Alabama.

INTRODUCTION

Cyperus granitophilus McVaugh was first described by Rogers McVaugh in 1937 from a granite outcrop in Geor¬

gia (McVaugh 1937). This granite outcrop endemic is primarily restricted to the Piedmont region of the south¬

eastern United States (Ware et al. 2011) with records from Georgia (McVaugh 1937), South Carolina (Hill &

Horn 1997), Alabama, North Carolina, Tennessee, and Virginia (Estill & Cruzan 2001) (Fig. 1). Cyperus grani¬

tophilus is typically found in the dry, shallow soil zone bordering exposed granite bedrock and is often domi¬

nant in these areas (Ware et al. 2011). Burbanck and Platt (1964) refer to this zone as the “lichen-annual herb”

community with a maximum soil depth of 7 to 15 cm. The species is also found in “scattered tufts” in the shal¬

low soil areas of their “annual-perennial herb” communities which typically have deeper soils, up to 39 cm.

Cyperus granitophilus is often found growing in association with the closely related species C. squarrosus

L. (Murdy 1968). McVaugh (1937) distinguished C. granitophilus from C. squarrosus by its slightly longer and

wider spikelets, longer scales, 4 vs 3 lateral nerves on the scale, and non-stipitate achene (Fig. 2). McVaugh

(1937) indicates that C. granitophilus can also be distinguished by its “somewhat stiffer culms, its densely

crowded capitate inflorescence and somewhat coarser appearance, and by the awns of its spikelet-scales, which

are spreading instead of reflexed as they often are in C. inflexus (C. squarrosus ).” McVaugh indicates that while

there is some overlap in the size measurements of the two species there is no intergradation in the achene char¬

acter. Cyperus granitophilus is documented as an autotetraploid derivative of C. squarrosus (Garoni & Murdy

1964; Wynne 1964) with consistently higher chromosome numbers in C. granitophilus (2 n = ca. 80, 88, 96)

versus C. squarrosus (2 n = ca. 48, 56, 64) (Murdy 1968).

METHODS AND RESULTS

During a study of the flora of Enchanted Rock State Natural Area, Flano and Gillespie counties, Texas, during

the summer of 2014, Cyperus granitophilus was discovered at six locations, one in Gillespie County and five in

J. Bot. Res. Inst. Texas 9(1): 251 - 257.2015

252

Journal of the Botanical Research Institute of Texas 9(1)

Fig. 1. Map showing updated county level distribution for C . granitophilus in the Eastern United States (Kartesz 2014; US Environmental Protection

Agency 2013).

Llano County. The five main population centers within the park include 1) vernal pools on the summit of En¬

chanted Rock, 2) vernal pools on Little Rock, 3) seasonal seepage areas over granite rock west of Little Rock, 4)

granitic sand over granite bedrock on Sandy Creek, and 5) a granite dike approximately 30 cm wide intruding

in an area of Packsaddle Schist. This last population consisted of very few plants and is surrounded by habitat

that is not suitable for the species.

Plants at Enchanted Rock State Natural Area were found growing in shallow soil no more than 15 cm in

depth, but typically less than 5 cm. These shallow soil zones were found on the edges of exposed granite on the

margins of vernal pools, cracks in granite, and granitic sand along drainage ways (Lig. 3). The plant was typi¬

cally found in areas with slightly increased moisture relative to surrounding areas. Associated species include

Bulbostylis capillaris (L.) Kunth ex C.B. Clarke, Calibrachoa parviflora (Jussieu) DArcy, Cyperus haspan L., C.

squarrosus, Elatine brachysperma A. Gray, Hypericum drummondii (Grev. & Hook.) Torr. & A. Gray, Hypericum

gentianoides (L.) B.S.P., Ipomoea costellata v ar. edwardsensis R. O’Kennon & Nesom, Isoetes lithophila N.E. Pfei¬

ffer, Lepuropetalon spathulatum Ell., Lindernia dubia var. anagallidea (Michx.) Cooperrider, Phemeranthus par-

viflorus (Nutt.) Kiger, Portulaca pilosa L., Sedum nuttallii Torr. & James ex Eat., Selaginella corallina (Riddell)

Wilbur & Whitson, Steinchisma hians (Ell.) Nash, Tripogon spicatus (Nees) Ekman, and Utricularia cornuta

Michx.

The authors conducted a thorough examination of specimens of the closely related species Cyperus squar¬

rosus (= C. aristatus , C. inflexus) from the following herbaria: BRIT, OKL, SMU, TAMU, TEX-LL, VDB. During

this process, 18 additional specimens of C. granitophilus (total of 27) were discovered in four Texas counties:

O'Kennon and Taylor, New Texas and Oklahoma records for Cyperus granitophilus

253

C. squarrosus

1 Mini

C. granitophilus

1 mm

> \ 1

1 mm

t f (

Texas Oklahoma Piedr

)

nont

Fig. 2. Spikelets (top), floral scales (middle), and achenes (bottom) of C . squarrosus ( O'Kennon 8128 a , BRIT), and C . granitophilus from the Llano Uplift

of Texas ( O ' Kennon , Taylor & Rehman 26142 , BRIT), the Wichita Mountains of Oklahoma {Waterfall 2904 , OKL), and the Piedmont Granite of South

Carolina { Hill 20023 , BRIT).

254

Journal of the Botanical Research Institute of Texas 9(1)

Fig. 3. Images of C. granitophilus and its habitat at Enchanted Rock State Natural Area. Top left: vernal pool on summit of Enchanted Rock; top right:

image of plant; bottom: C. granitophilus growing in shallow soil near edge of granite rock.

O'Kennon and Taylor, New Texas and Oklahoma records for Cyperus granitophilus

255

Burnet, Gillespie, Llano, and Mason; and 11 specimens in three Oklahoma counties: Comanche, Greer, and

Johnston. All of the Texas collections are from the Llano Uplift while all of the Oklahoma specimens are from

the Wichita Mountains or Arbuckle Uplift (Fig. 1). These new populations in Texas and Oklahoma are disjunct

approximately 1270 km (790 mi) from the western most location in Alabama.

Since Cyperus granitophilus does not occur in any regional floras within Texas or Oklahoma it is not sur¬

prising that specimens from the region were misidentihed as C. squarrosus. Cyperus granitophilus is widely

distributed across the Llano Uplift and observations at Enchanted Rock State Natural Area suggest it is likely a

common member of the granitic flora in the region. Additional exploration of granite outcrops in both Texas

and Oklahoma will undoubtedly reveal additional populations.

DISCUSSION

The Llano Uplift of central Texas, and the Wichita Mountains and Arbuckle Uplift of Oklahoma are all regions

with large expanses of exposed granite, similar to that in the Piedmont of the southeastern United States where

Cyperus granitophilus is wide-spread. The Llano Uplift of Texas spans 10 counties and includes approximately

5180 km 2 on the eastern margin of the Edwards Plateau (Walters & Wyatt 1982). It is composed of Precam-

brian granites, gneisses, and schists surrounded by escarpments of Paleozoic and Mesozoic limestone and

sandstone. The region was once covered by Cretaceous limestone and sandstone with the granite basement

rock being exposed due to erosion during the Miocene and Pliocene (Walters & Wyatt 1982). Therefore the

Llano Uplift is a “topographical basin consisting of exposed Precambrian metamorphic rocks and large mas¬

sifs of Precambrian granite surrounded by a discontinuous rim of Cretaceous limestone” (Walters & Wyatt

1982).

The Wichita Mountains of Oklahoma are primarily in Comanche County, but enter four additional coun¬

ties, covering approximately 945 km 2 (US Environmental Protection Agency 2013). These rugged hills of igne¬

ous granite, granite-porphyry, diabase (dolerite), and gabbro rise above a relatively flat plain of Cambrian and

Ordovician sandstone and limestone (Taff 1928).

The Arbuckle Uplift of Oklahoma, including the Arbuckle Mountains, is a plateau moderately elevated

above the surrounding plain entering seven counties and covering approximately 3066 km 2 (US Environmen¬

tal Protection Agency 2013). The area is predominately Cambrian to Devonian limestone with Carboniferous

shales, sandstones, and conglomerates on the borders. A mass of granite, granite-porphyry, diabase, and asso¬

ciated crystalline rocks can be found in the central part of the uplift. Igneous rocks are exposed in three areas,

with the largest in the east composed mostly of granite and diabase dikes. The surface granite is nearly flat and

much of it is concealed by materials produced by its own disintegration (Taff 1928).

The granites of the Wichita Mountains, Arbuckle Uplift, and Llano Uplift are markedly similar in struc¬

ture (Taff 1928) and all possess similar plant communities, particularly near exposed granite rock. Burbanck

and Platt (1964) note the similarities between the Piedmont granite and the Llano Uplift of Texas stating “al¬

though the species are different, the types of plants and the stages of succession on exposed granite and accu¬

mulated gravel at Enchanted Rock, Texas.. .are similar to conditions in Georgia.” Walters and Wyatt (1982)

echo this sentiment by stating “that granite outcrops provide similar types of microenvironments despite ma¬

jor differences in the climate and surrounding vegetation of each region.”

While the granitic regions west of the Mississippi River (Llano Uplift, Wichita Mountains, and Arbuckle

Uplift) share similar geology and microenvironments to that of the Piedmont granite, their floras are quite dif¬

ferent (Walters & Wyatt 1982). This lack of similarity in characteristic and endemic species between the east¬

ern and western granitic outcrops “suggests that each region contains a unique group of plants independently

derived from the native plants of that region” (Walters & Wyatt 1982). When examined at the generic level this

is not necessarily the case though, with several genera having “highly specialized representatives in both re¬

gions” (Walters & Wyatt 1982).

These widely separated regions possess similar floras, including disjunct taxa such as Cyperus granitophi¬

lus, as a result of a shared flora during the Tertiary period. The Madro-Tertiary Geoflora of western and south-

256

Journal of the Botanical Research Institute of Texas 9(1)

western North America shows remarkable floristic similarities to that of the Llano Uplift, Wichita Mountains,

Arbuckle Uplift and Piedmont granite rock outcrop floras (Walters & Wyatt 1982). Warmer and drier climates

during the Late Tertiary allowed plants from western and southwestern North America to expand their ranges

into the southeastern United States (Braun 1955). Thus, it is likely that the Llano Uplift, Wichita Mountains,

Arbuckle Uplift, and Piedmont once had a shared flora and that flora was adapted to warmer and drier condi¬

tions than what exist today across most of the region. Cooler and wetter temperatures in the Pleistocene subse¬

quently forced xerophytes in the Southeast to take refuge on rock outcrops. Once isolated, subsequent specia-

tion would result in narrowly restricted endemics within each system, such as Isottes lithophila in the Llano

Uplift and I. melanospora Engelm. in the Piedmont.

Alternatively, the absence of species divergence after isolation would result in highly disjunct popula¬

tions. This distribution pattern is not atypical for ecological endemics with highly restrictive specificity to a

single substrate. This is seen with Cyperus granitophilus, Eriocaulon koernickianum, and Isottes piedmontana

(N.E. Pfeiffer) C.F. Reed on granite outcrops and Isoetes butleri Engelm. (Taylor et al. 2012), Gratiola quarterma-

niae D. Estes (Taylor & O’Kennon 2014), and Oenothera macrocarpa Nutt. ssp. macrocarpa (Kartesz 2014) on

limestone outcrops. Additionally, environmental differences between outcrop communities and the surround¬

ing habitats is markedly more intense in the Piedmont where the incident radiation, temperatures and high

levels of evapotranspiration are drastically more severe on the rock outcrops than the surrounding habitats.

Warmer and drier overall climates in Texas and Oklahoma result in a less extreme environmental difference

between outcrops and surrounding vegetation, less isolation from congeners, and a lower degree of endemism

(Walters & Wyatt 1982).

Voucher Specimens. U.S.A. OKLAHOMA. Comanche Co.: Elk Mountain, Wichita Mountain Wildlife Refuge, 34.725819 -96.721114,19Jun

2015, O’Kennon, Taylor, Jensen, &Rylander, 28463 (BRIT); Northwest Sunset Peak, 21 May 1983, Rose 85 (OKL); wet, sandy shoreline, Qua-

nah Parker Lake, Wichita Mts., 2Jun 1949, Penfound P-197 (OKL); sandbar, head of Lake Elmer Thomas, 9 Jul 1948, Penfound P-54 (OKL);

Wichita Mts. Wildlife Refuge, Sugar Creek, open prairie near North boundary, 29 Jul 1942, Rouse 405 (OKL); growing in soil collected

among granite boulders near top of Mt. Scott, Wichita Mts, 4 Jul 1941, Waterfall 2925 (OKL); clay and shale on granite with a mixed grass -

sedge association, N of Mt. Scott about 1 mi E of Meers, Wichita Mts., 4Jul 1941, Waterfall 2904 (OKL); Medicine Park, 20 Oct 1934, Bradbury

593 (OKL); s.d., Stevens 1330 (OKL); s.d., Stevens 1358 (OKL). Greer Co.: at edge of small mountain near granite, 17Jun 1913, Stevens 1027

(OKL). Johnston Co.: granite rockat 10 Acre Rock, 34.328989 -96.762469,13Jun 2015, O’Kennon28420 (BRIT); granite rockarea W oftown

of Mill Creek, 19 May 1967, Taylor & Taylor 3690 (OKL). TEXAS. Burnet Co.: Granite Mountain along Larm Road 1431, 1.8 mi W of L.M.

1431 andU.S, 281, W of Marble Lalls, 7Jun 1988, Urbatsch4791 (BRIT); growing in loose small granite rocks, Granite Mountain, 16Jun 1946,

Cory 12729 (SMU). Gillespie Co.: Enchanted Rock State Natural Area, in moist granitic glade south of Little rock, N30.495971, W98.829125,

26 Jun 2014, O’Kennon, Taylor, & Rehman 26146 (BRIT); Enchanted Rock State Natural Area, in moist granitic grus, S of Little Rock,

N30.4959566, W98.8291267,16 Jul 2014, Taylor, O’Kennon, & Rehman 2852 (BRIT); in granite gravel on outcrop near Coal Creek, N of Wil¬

low City, 10 Jul 1958, Correll & Johnston 19582 (TEX-LL); in shallow depressions in granite on Bear Mt., just N of Lredericksburg, 29 Jun

1957, Correll & Johnston 17258 (TEX-LL); 5 Aug 1940, Strandtmann s.n. (TEX-LL). Llano Co.: Enchanted Rock State Natural Area, vernal

pool on Enchanted Rock, N30.506892, W98.81962436, 21 Oct 2014, Taylor & O’Kennon 2935 (BRIT); granite dike E of Enchanted Rock,

N30.51247, W98.8026, 19 Sep 2014, Taylor & O’Kennon 2908 (BRIT); drainage over granite, N30.50604, W98.8348, 18 Sep 2014, Taylor &

O’Kennon 2885 (BRIT); vernal pool on West Rock, N30.50006, W98.8229, 17 Sep 2014, Taylor & O’Kennon 2860 (BRIT); Enchanted Rock

SNA, N30.500257, W98.819718,15 Jul 2014, Taylor, O’Kennon & Rehman 2776 (BRIT); Enchanted Rock State Natural Area, in moist granitic

sand along margin of Sandy Creek that runs near the southern and eastern base of Enchanted Rock, N30.510462, W98.804992,15 Jul 2014,

O’Kennon, Taylor, & Rehman 26142 (BRIT); Enchanted Rock SNA, Moss Lake, 14 Jul 2014, O’Kennon, Taylor, & Rehman 26145 (BRIT); En¬

chanted Rock SNA, along moist bank of Moss Lake, N of Enchanted Rock, N30.50888, W98.825086,4 Oct 1990, O’Kennon 8128g (BRIT); 9.6

mi SW on EM 2323 from its jet with TX16, then S ca 100 m on an unnamed paved rd, E side of rd, SW of Llano, granitic outcrop between the

road and fence, associated species other Cyperus, Teptochloa, Digitaria, Bouteloua, Tripogon, and Bothriochloa, 26 Sep 1987, Wipff563 (TEX-

LL); granite outcrop 1.4 km W on Lerguson Rd from intersection of Lerguson Rd and RR 2147 at Horseshoe bay, 11 Aug 1979, Walters 1000

(TAMU); granite outcrop 3.4 km N on W side of RM2241 from the intersection of RM 2241 and U.S. Hwy 21 at Bluffton, 18 Jun 1979, Walters

681 (TAMU); common sedge in many of the vernal pools on the summit of Enchanted Rock, 2 Oct 1976, Butterwick & Smith 3376 (TEX-LL);

common annual sedge in vernal pools on summit of Enchanted Rock, 24 Jul 1976, Butterwick & Tamb 3020 (TEX-LL); Watch Mountain,

granite outcrop 17.2 km W on N side of Inks Ranch Rd, from the intersection of Inks Ranch Rd and U.S. Hwy 16,13 Jun 1974, Walters 496

(TAMU); Enchanted Rock, 20 Aug 1936, Tharp s.n. (BRIT); granitic areas, 5 Aug 1931, Wolff 3157 (BRIT); Enchanted Rock, 11 Jun 1930,

Whitehouse & Tharp s.n. (TEX-LL); Enchanted Rock, Jul 1892, Nealley 80 (TEX-LL). Mason Co.: Mason Mountain WMA, in moist cracks in

granite, N30.826464 W99.220053,14 Aug. 2012, O’Kennon, Taylor, & Rehman 25348 (BRIT); granite outcrop and sandy creek bottom, 14.2

mi E of Mason by highway TX 29, local in thin sand over rock, 16 Aug 1989, Carter 8214 (SAT, VSC); granite outcrop 2.6 km E on S side of RM

O'Kennon and Taylor, New Texas and Oklahoma records for Cyperus granitophilus

257

1222 from intersection of U.S. Hwy 87 and RM 1222 at Camp Air, Walters 441 (TAMU); granite outcrop 2.7 km S on U.S. Hwy 87 from inter¬

section of U.S. Hwy 87 and RM 1222 at Camp Air, W 0.2 km, 11 Jun 1979, Walters 424 (TAMU).

ACKNOWLEDGMENTS

We thank David Riskind, Doug Cochran, and Scott Whitener from Texas Parks and Wildlife for permits and

assistance in accessing Enchanted Rock; Richard Carter for his advice and help with identification; and the

following herbaria for loans or use of facilities to examine specimens: Botanical Research Institute of Texas

(BRIT, SMU, VDB), University of Oklahoma (OKL), Texas A&M University (TAMU), and University of Texas

(TEX-LL). We also thank Gordon Tucker (EIU) and Bruce Hoagland (OKL) for their helpful reviews of the

manuscript.

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Journal of the Botanical Research Institute of Texas 9(1)

ANNOUNCEMENT

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 may be 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 applica¬

tions 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 ac¬

cepted before applying. The Systematics Symposium dates for 2015 are 8-10 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 9(1): 258.2015

PHYTOGEOGRAPHICAL RELATIONSHIPS AND ANALYSIS

OF THE FLORA OF SOUTH-CENTRAL TEXAS, U.S.A.

A.A. Saghatelyan

Department of Biology

McMurry University, McMurry Station

Abilene, Texas 79697, U.S.A.

saghatelyan@mcm.edu

ABSTRACT

A flora of the elevated portion of Edwards Plateau and western half of the Tlano Uplift in South-Central Texas (SC TX flora) was studied from

a biogeographical perspective. A checklist of the designated flora was extracted primarily from the “Floristic Synthesis of North America”

and data concerning general distribution of the taxa were obtained from the literature and online databases. Phylogenetic literature was

searched to find relationships, patterns of migrations, and geographical connections of the species of major clades. Taxonometric and geo¬

graphic spectra of the SC TX flora were obtained and compared with those for the flora of Big Bend Region (BB), Texas. There are 1619 native

species of 626 genera and 149 families in the SC TX flora. These species were classified into 21 geographic (floristic) elements. Herein is

presented a checklist accompanied by the geoelement descriptions and the taxonometric and geographic analyses of the flora. The flora,

being on the boundary of two floristic subkingdoms and two regions, has a complex pattern of connections. Among five most specious

families, the Asteraceae and Euphorbiaceae have half of their species in the local clades which diversified after transatlantic long distance

dispersals. The rest of the spurges and about 70% of the grasses and sedges have tropical/subtropical connections. Almost two thirds of the

Fabaceae have connections to SE N American/Madrean/Mesoamerican, Tethyan or S American radiations. The largest geoelement in the

flora is comprised of 236 E North American species. The entire North American Atlantic Region is represented by 679 species (42% of the

flora). Only 11% of the Big Bend flora species are in this group. The Holarctic Subkingdom in general is represented by 794 species (49% of

the flora). The Western Region/Madrean Subkingdom has 450 species (28% of the flora), followed by the Madrean/Neotropical Region with

212 species (14% of the flora). There are 132 differential genera in the SC TX flora not found in the Big Bend flora, including 54 tropic/sub¬

tropical, 34 temperate N Hemispheres, 14 E North American, and 12 SC/SW USA-N Mexico genera. Higher numbers in mesophyllous gen¬

era and more connections with the North American Atlantic Region are evident from the family and generic spectra of the SC Texas flora,

while the Big Bend flora spectrum has more species of the xeric genera native to southwestern North America. In general, southern and

eastern connections prevail in the SC TX flora. Most characteristic scenario in taxonomically significant groups shows local diversification

after transatlantic crossings of the ancestral species from the Old World, as well as S/N American bidirectional migrations, whereas most

oligotypic and monotypic genera include relictual species of tertiary Lauroasian genera, best preserved in the southeastern North America,

or species of tropical genera extending to the southern boundary of U.S.A.

RESUMEN

Se estudio la flora de una porcion elevada del Edwards Plateau y oeste de la elevacion del Llano en el centro-sur de Texas (SC TX flora) desde

una perspectiva biogeografica. Se extrajo primariamente un catalogo de dicha flora a partir de la “Floristic Synthesis of North America” y se

obtuvieron datos de la distribucion general de los taxa a partir de la bibliografia y de databases online. Se examine la bibliografia filogenetica

para hallar relaciones, patrones de migraciones y conexiones geograficas de las especies de los clados mayores. Los espectros taxonometri-

cos y geograficos de la flora SC TX se obtuvieron y se compararon con los de la flora de la region Big Bend (BB), Texas. Hay 1619 especies

nativas de 626 generos y 149 familias en la flora SC TX. Estas especies se clasificaron en 21 elementos geograficos (floristicos). Se presenta

aqui un catalogo acompanado de descripciones del geoelemento y de analisis taxonometricos y geograficos de la flora. La flora, situada en la

frontera de dos subreinos floristicos y dos regiones, tiene un patron complejo de conexiones. Entre las cinco familias con mas especies, las

Asteraceae y Euphorbiaceae tienen la mitad de sus especies en los clados locales que se diversificaron despues de las dispersiones a larga

distancia transatlanticas. El resto de las euforbias y sobre el 70% de las gramineas y ciperaceas tienen conexiones tropicales/subtropicales.

Casi dos tercios de las Fabaceae tienen conexiones con radiaciones con SE N Americanas/Madreanas/Mesoamericanas, Tethyanas o S

Americanas. El mayor geoelemento de la flora comprende 236 especies del E Norte Americano. La Region entera Norte Americana Atlantica

esta representada por 679 especies (42% de la flora). Solamente el 11% de las especies de la flora Big Bend estan en este grupo. El subreino

Holartico en general esta representado por 794 especies (49% de la flora). La Region Oeste /Subreino Madreano tiene 450 especies (28% de

la flora), seguida por la Region Madrean/Neotropical con 212 especies (14% de la flora). Hay 132 generos diferenciales en la flora SC TX que

no se encuentran en la Big Bend, incluyendo 54 generos tropicales/subtropicales, 34 del hemisferio N templado, 14 E Norte Americanas, 12

SC/SW USA-N Mexico. Los numeros mas altos en los generos mesofilos y mas conexiones con la Region Norte Americana Atlantica son evi-

dentes en los espectros de familias y generos de la flora SC Texas, mientras que el espectro de la flora Big Bend tiene mas especies de los

J. Bot. Res. Inst. Texas 9(1): 259 - 294.2015

260

Journal of the Botanical Research Institute of Texas 9(1)

generos xericos nativos del suroeste de Norte America. En general, las conexiones con el sur y este prevalecen en la flora SC TX. El escenario

mas caracteristico en los grupos mas significantes taxonomicamente muestra diversificacion local despues del paso transatlantico de las

especies ancestrales desde el Viejo Mundo, asi como las migraciones bidireccionales S/N Americanas migraciones, mientras que los generos

mas oligotipicos y monotipicos incluyen especies relicticas de generos del terciario Lauroasiatico, mejor preservados en el sureste de Norte

America, o especies de generos tropicales que se extienden hasta la frontera sur de Estados Unidos.

INTRODUCTION

Comparative floristic analysis has long been one of the major tasks for phytogeographers from different parts

of the world. Well defined and carefully analyzed regional floras can be used for comparison with other floras

and elucidation of the connections among them. The analysis is preceded by the grouping of the species of a

flora into diverse floristic elements: geographic, ecological, historic, etc, depending on the goal. A natural flora

can be viewed as a section of, or a probe into, the ever changing continuum of plant cover. It is a section of a

particular size in space and time and as such it can be used for the clarification of its composition and how it

changes through time. For chorionomic regionalization, which seeks to delineate natural geographic areas

with distinct floras, it should be delimited by natural barriers. Such floras are characterized by specific assem¬

blages of endemic taxa of particular rank, depending on the rank of the chorion (Takhtajan 1986; Kamelin

1973; 1990; 1998; 2010). To define biogeographical regions, different authors used cluster analysis, principal

component analysis, and other statistical techniques (McLaughlin 2007; Finnie at al. 2007; Born et al 2007;

Linder et al. 2012) or used the phylogenies of particular taxonomic groups for parsimony analysis of endemic-

ity (Morrone et al 1999; Katinas et al 2004). A different approach to floristic analysis utilizes more loosely

defined boundaries of a flora to focus not on its regionalization but on the presence of its extant taxa in the re¬

gion and their phylogenetic relationships with sister taxa in other floras (Galley & Linder 2006), especially on

the taxa with intercontinental disjunctive patterns (Thorn 2004; Renner 2004; Donoghue & Smith 2004; Wen

& Ickert-Bond 2009; Olmstead, 2013). With the wealth of now available published phylogenies and historical

biogeographical analyses, some repetitive patterns of disjunctions, dispersals, vicariance, and extinctions

emerge among many non-related taxonomic groups of a given flora. This paper focuses on the search of such

repetitive patterns of disjunctions in the phylogenetic literature for the taxa of a sample flora in South-Central

Texas. For this purpose, a flora in South-Central Texas, which includes more elevated parts of the Edwards

Plateau and the western Llano Uplift in the natural regions 7 and 8 (Fig. 1), is loosely delineated and will be

presented and analyzed in this work. It is not our intention to reconsider the natural regions in the area, as it is

a separate task.

Another purpose of this paper is to compare two almost adjacent floras in Texas: the floras of South-

Central Texas (SC TX), which is the subject of this study, and the flora of Big Bend Region (BB), in natural re¬

gion 11 (Fig. 1) analyzed in A. Saghatelyan (2009). Depending on the chorionomic system we use, SC TX falls

into two different regions. It is a part of the North American Prairies Province of the Atlantic North American

Region in the chorionomy of Cronquist (1982) and Takhtajan (1986) accepted in this paper (Fig. 2). According

to this system, SC TX falls into a different region than BB, which is a part of the Madrean Region. Alternatively,

it is a part of the Southwestern Region of the chorionomic system of S. McLaughlin (2007), which includes the

Great Plains Province. As such, the SC TX flora would be placed together with the BB flora in the Southwestern

Region. Thus, a detailed floristic analysis can support one of those placements, depending on its species con¬

nections and the degree of similarity of the two sample floras.

GEOGRAPHIC SETTING

The Edwards Plateau of SC Texas is a plateau east of the continental divide, sloping gently eastward, but

steeper and dryer at the western margin. Its elevation ranges from 450 ft to 3000 ft. The dissected southern and

eastern half of the Edwards Plateau is known as the Texas Hill Country. Its surface consists almost entirely of

outcrops of Cretaceous limestone. A relatively small area in the north central margin of the Edwards Plateau is

known as the Granitic Central Basin, or Llano Uplift. In this area the Cretaceous limestone has been removed

by erosion. The exposed rocks are largely granitic and gneissic. The Balcones Escarpment at the southeastern

Saghatelyan, Phytogeographical analysis of the South-Central Texas flora

261

Fig. 1. Texas county map with the study area of the South Central Texas flora (SC TX) and the Big Bend area (BB) outlined by a bold contour. The map is

courtesy of Texas Parks and Wildlife and was modified in Photoshop by Mike Karabegov.

edge of the Edwards plateau seems to be a floristic barrier (MacRoberts & MacRoberts 2008). The study area

covers the elevated central part of the mostly limestone Edwards Plateau east of the Pecos River and the west¬

ern part of the granitic Llano Uplift. Most of its land surface has been exposed continuously for occupation by

terrestrial biota for at least 65,000,000 years (Correll & Johnston 1970) and has subhumid subtropical climate.

The study area is outlined on the map (Fig. 1) and includes the following counties: Bandera, Concho,

Crockett, Edwards, Gillespie, Irion, Kendall, Kerr, Kimble, Kinney, Llano, Mason, Medina, Menard, Reagan,

Real, Schleicher, Sutton, Tom Green, Uvalde, and Val Verde. The Edwards Plateau environments and vegeta¬

tion are well known (Amos & Gehlbach 1988). The “original” vegetation was grassland or more commonly a

type of open temperate grassland, with shrubs and low trees along rocky slopes (Correll & Johnston 1970;

262

Journal of the Botanical Research Institute of Texas 9(1)

Arctic

ARC

Canadian

CAN

Appalachian

APP

Atlantic and Guif Coastal

ATL

North American Prairies

NAP

Chihuahuan

CHI

Tamaulipan

TAM

Rocky Mountain

ROC

Great Basin

GBA

Vancouverian

VAN

Californian

CAL

Mojavean

MOJ

Sonoran

SON

Sierra Madre Occidental

SMO

Mexican Alti piano

MAS

Trans Mexican Volcanic

TMV

Sierra Madre Oriental

SMR

Sierra Madre del Sur

SMS

Neotropical

Fig. 2. Floristic provinces/areas of endemism of North America. Regional schema adapted from Takhtajan (1986) as published in Thorne (1993). The areas

were drawn in Photoshop by Alice Tangerini on a base map from ArcMap8.2 with a North American Lambert conformal conic projection. The map was

published in Katinas et al. (2004) and is used with permission of Missouri Botanical Garden Press and the authors.

Stanford 1976; Hatch et al. 1990). The floristic geography of woody and endemic plants was analyzed by Amos

and Rowell (1988).

Here is presented a checklist and detailed analysis of the SC TX flora, including the study of general dis¬

tribution outlines of its species and genera as well as their classification into geographical elements. This infor¬

mation coupled with the phylogenetic literature data will help to reveal connections of the flora to other floras

and possible migration routes of its ancestral species.

MATERIALS AND METHODS

A checklist of the flora of the study area in SC Texas, including the Edwards Plateau and western part of the

Saghatelyan, Phytogeographical analysis of the South-Central Texas flora

263

Llano Uplift, was compiled using primarily the Synthesis of the North American Flora (Kartesz 2013). The

non-native species were excluded. Twenty-seven sandy land species reported outside of the Edwards Plateau

ecoregion in Medina County, only from the Carrizo Sand south of the Balcones Escarpment, were also ex¬

cluded. Taxonomic counts of the families, genera, and species were performed. Distributional and other data

was obtained from the Synthesis of the North American Flora (Kartesz 2013), Tropicos (Tropicos.org.), Flora of

North America and Digital Flora of Texas databases, literature on Texas flora (Correll & Johnston 1970; John¬

ston 1990; Turner et al. 2003; Diggs et al. 1999), many online databases on different taxonomic groups (like G.

Nesom, Astereae database), and other literature sources. After revealing general distribution outlines for each

species of the flora, congruent ranges of two or more species were classified into geographic elements using the

classification system of geographic elements (Saghatelyan 2009) developed on the example of the BB flora.

Each species was thus referred to and treated as a particular geographic element or geoelement (Saghatelyan

1997a, b). The distributional data on all the genera were retrieved from Wielgorskaya (1995), Mabberley (1997),

the Flora of North America, Flora of China, and online sources. The genera were classified into 19 groups based

on their distribution outlines. The species list (Appendix 1) for the above-mentioned counties was compiled

with the major objective of defining the geoelements and follows the nomenclature accepted in Kartezs (2013),

with the exception of Liliales and Asparagales, which follow APG. The unidentifiable ranges of a few species

are noted with a question mark in the checklist. After analyzing the SC TX flora, the taxonomic and geograph¬

ic compositions of the flora were compared with those of the BB flora. A special attention was given to phyloge¬

netic literature search to clarify the relationships of significant in the flora taxonomic groups and patterns of

vicariance and/or migration.

TAXONOMETRIC ANALYSIS OF MAJOR FAMILIES

There are 1619 species in 626 genera and 149 families in the SC TX flora (Appendix 1). Specific and generic

richness of the largest families is presented in Table 1. First ten largest families comprise 859 species or 53% of

the flora. The family spectrum is close to that of the BB flora. However, Fabaceae, Euphorbiaceae, Cyperaceae,

Rosaceae, and Apiaceae are more speciose in the SC TX flora, while Cactaceae, Boraginaceae, and Brassicaceae

have a higher percentage of species in BB.

The largest family Asteraceae has 232 species in 98 genera in the SC TX flora. Different clades of the com-

positae metatree (Funk et al. 2009) have the following numbers of species in the flora. The flora includes only

four species of the South American Mutiseae clade, which have the Sonoran Province/East Madrean ranges.

North American Cardueae have only eight species in the flora. This latter group diverged from Plio-Pleistocene

or later (Hellwig 2004) diversification in the West Asian-Mediterranean thistle clade and represents a recent

monophyletic radiation to North America (Barres et al. 2013; Funk et al. 2005; 2009).

Fourteen of the 15 species of the Lactuceae and Vernonieae in the flora have the East Prairie-Eastern

North American ranges. This concentration of the Lactuceae in the eastern North America supports a probable

trans-Atlantic crossing from the Mediterranean. While Lactuceae may have arrived from the Mediterranean,

the dispersal in Vernonieae was from tropical Africa to South America and then from South to North America,

with Meso/Central America playing an important role (Keeley et al. 2007; Keeley et al. 2013). The small tribe of

Moquinieae is in a clade with Vernonieae in the Atlantic SE Brazil, while North American members of Vernon¬

ieae are more highly nested in that clade (Funk et al. 2005; Funk et al. 2009).

Asteroideae, the largest subfamily in North America, starts with the global tribe Senecioninae, richly

represented (with 61 sp. of Senecio and 54 sp. of Packera) in western North America, but with only seven spe¬

cies in the SC TX flora. The Gnaphalieae (eight sp.) and Anthemideae (two sp.) are also scarce in SC TX while

being highly diverse in the western North America. For example, the North American endemic Artemisia

group with 58 species in western North America has only one species in the SC TX flora. The group diverged

from Asian ancestors by the Late Miocene (Gonzalez et al. 2011) and arrived via Beringia. The pattern of distri¬

bution of the species of Senecioninae in North America resembles that of the genus Artemisia and points to a

probability of the Beringia crossings.

264

Journal of the Botanical Research Institute of Texas 9(1)

Table 1. Edwards Plateau (EP) and Big Bend (BB) flora largest families. The list is ordered by the number of Edwards Plateau species in each family, and percentages

are relative to the entire flora.

Largest Families

EP species (%) / genera

BB species (%) / genera

Asteraceae

232 (14.3)/98

230 (14.5) /107

Poaceae

199 (12.2)/65

202 (12.7)/63

Fabaceae

120 (7.4)/41

107 (7.1) / 37

Cyperaceae

69 (4.2) / 11

40 (2.5)/10

Euphorbiaceae

68 (4.1)/12

59 (3.7)/10

Cactaceae

40 (2.5)/13

59 (3.7)/17

Brassicaceae

37 (2.3)/14

43 (2.7)/21

Lamiaceae

36 (2.2) / 11

38 (2.4) / 11

Plantaginaceae

34 (2.1)/11

29 (1.8)/13

Solanaceae

28 (1.7) / 10

32 (2.0)/10

Pteridaceae

27 (1.7)/7

33 (2.1)/8

Rosaceae

26 (1.6)/10

12 (0.8) / 9

Malvaceae

24 (1.5) / 12

Apiaceae

24 (1.5)/ 18

7 (0.3)/5

Onagraceae

24 (1.5)/3

28 (1.8)/4

Boraginaceae

22 (1.4)/11

33 (2.1)/9

Asclepiadaceae

21 (1.3) / 4

23 (1.5)/4

Astereae have 72 species in the SC TX flora. Conyzinae have 15 species and are a North American group

with the basal taxa being Erigeron species from southwestern N America (Nesom 1994; Noyes & Rieseberg

1999). The Solidagininae have 11 species all with ranges in the eastern North America. This is where the center

of diversity of the genus Solidago lies. Broadly North American genera Symphyotrichum and Heterotheca have 14

species in the flora. Remaining two thirds of the Astereae species have southern or western connections.

Among them are 25 species of the genera Baccharis, Gutierrezia, Laennecia, Grindelia, Isocoma, and Croptilon,

centered in Mexico, western or southern North America and South America. All North American Astereae are

members of a strongly supported clade, with basal Astereae being predominantly woody taxa from Africa,

Australia, and South America (Noyes & Rieseberg 1999). African genera form a basal clade in the tribe As¬

tereae along with the Chinese genus Nannoglottis and South American and New Zealand genera. The latest

diverging African clade, subtribe Grangeinae, is sister to the Eurasiatic subtribe Bellidinae, and together they

are sister to the Astereae crown lineages of Australasia-Asia and South and North America (Brouillet et al.

2009).

Inulea have only two species of the tropical-warm temperate genus Pluchea in the flora.

Heliantheae s.l. is the most speciose group of the Asteraceae in the flora. An eastern Cuban genus Feddea

was shown to be sister to the Heliantheae s.l., and the New World Teddea/Heliantheae s.l. group was shown to

be sister to the mostly Old World (predominantly in E Africa) tribe Athroismeae (Cariaga et al. 2008). Funk et

al. (2009) do not exclude the possibility of a direct Old World to New World dispersal of the American clade of

Heliantheae.

The core Heliantheae Alliance begins with Helenieae.The flora has 20 species of Helenieae with the south-

central or southeastern North American ranges, including one species of the genus Marshallia. Marshallia is

sister to the rest of Helenieae and grows in mostly mesic habitats in south-central/southeastern United States.

This supports the above mentioned proposed dispersal route for Heliantheae from the Old World.

The Coreopsideae have nine species with similar ranges.

The Eupatorieae are concentrated in Mexico and Central and South America; they have 14 species in the

flora. Eupatorium had initial divergence in North America and shares a common ancestor with Liatris (Schmidt

& Schilling 2000). The Liatrinae is essentially restricted to the eastern and south-eastern United States (Ne¬

som 2005). Remaining 91 species of the Heliantheae Alliance in the flora have the Chihuahuan-Tamaulipan,

Madrean, or Comanchian ranges, or broader south-central North American-E Prairie distribution.

Saghatelyan, Phytogeographical analysis of the South-Central Texas flora

265

Thus, a majority of the Asteraceae species of the SC TX flora belong to the largest North American Heli-

antheae Alliance. Together with the species of the Astereae, Inuleae, and Vernonieae they have mostly south¬

ern connections, especially in SE/SC North America or Central/South America. For present-day Asteraceae

composition of SC TX flora, long distance dispersals of the ancestral taxa from Mediterranean to Mesoameri-

ca/North America or from Africa to South America to North America were much more important than Berin-

gia crossings. The groups that have richly diversified in Western North America after a Beringia crossing, have

only a few species in the SC TX flora.

Poaceae, with 199 species of 65 genera in the flora, is the second largest family. Its species, as grasses in

general, have broad geographic ranges. The subfamilies characteristic to tropical/warm temperate latitudes,

especially Chloridoideae (71 sp.) and Panicoideae (61 sp.), have the majority of the species and eight largest

grass genera in the flora. Major centers of diversity of these genera are in Mexico and Mesoamerica. The ranges

of their species found in the flora are mostly in North American southwestern, south-central or, for a few, in

southeastern parts. To the contrary, the mostly north temperate subfamily Pooideae has only 29 species in the

flora. These Pooideae genera are small in the flora, monotypic or ditypic, with the largest one having only eight

species.

Fabaceae has 120 species of 41 genera and is in third place in the SC TX flora. Compared to the legumes of

the Big Bend flora (Saghatelyan 2009), there are seven more genera in SC TX: E Asian-E North American-South

American disjunctive genus Gleditsia, four tropical/subtropical genera, Bauhinia, Clitoria, Neptunia, and Zor-

nia, and two E North American genera of Papilionoideae, Baptisia and Apios. Caesalpinioideae of the SC TX

flora have eight genera and 15 species with mostly E North American, SC TX, or Chihuahuan-Tamaulipan

ranges. For the genus Gleditsia of E Asian origin both the mid-Miocene N Atlantic Land Bridge crossing (Tiff-

ney 1985; Schnabel & Wendel 1998; Lavin et al. 2000) and the Bering Land Bridge crossing (Schnabel et al.

2003) were proposed. The Atlantic crossing seems more plausible due to the strong macrofossil record for

Gleditsia in Europe. According to Mai (1995), Gleditsia was present in the savanna type vegetation in the Mio¬

cene of southern Europe, and its presence had decreased all over Europe in the Pleistocene. Other thermophyl-

lous genera, like Gymnocladus and Cercis, exhibit a similar pattern (Mai 1995). According to a molecular phy-

logeny of Cercis (Fritsch & Cruz 2012), North American species are sister to the western Eurasian species with

strictly east-to-west vicariance. It was inferred that the ancestor in which this divergence occurred in the

mid-Miocene was xerophytic and used the Miocene North Atlantic corridor for semi-arid plants. Basalmost

Caesalpinioideae ( Gymnocladus, Cercis, Ceratonia, Gleditsia, and Senna) had a strong participation in the Ter¬

tiary assemblages of subtropical floras of littoral type (Mai 1995). Schrire et al. (2005:405) also conclude: “Such

older northern temperate diversifications thus reflect a Tethyan-wide Tertiary tropical dry forest distribution

which existed prior to temperate conditions being superimposed on these areas.”

In North America, the genera Senna, Bauhinia, Chamaecrista, and Parkinsonia of the Succulent Biome of

Schrire et al. (2005), which has pantropical disjunct distribution, are restricted to south or southeastern United

States and Mexico (Kartesz 2013). This suggests transatlantic peri-Tethyan migrations for the pantropical gen¬

era ( Parkinsonia ) and south/north migrations for the mostly tropical American genera ( Chamaecrista ). The

Grass Biome (Schrire et al. 2005) genus Hoffmannseggia is amphitropically disjunct due to the long distance

dispersals from South to North America (Simpson et al. 2005). In the United States its species grow in the

southwest, which is common among other amphitropically disjunct genera. The genus Pomaria of the Grass

Biome of Schrire et al. (2005) also has amphitropical disjunction with extension to the western Prairie Region.

It had dispersal from southwestern United States and adjacent Mexico to southern Africa and second dispersal

from North America to temperate South America (Simpson et al. 2006).

Mimosoideae in the SC TX flora have 32 species in seven genera, all with southern connections. Many of

these species are common with the Big Bend flora (Saghatelyan 2009). However, the Madrean and Gulf Coastal

species of the Mimosoideae are more numerous in the SC TX than in the BB flora. The Succulent Biome species

comprise two thirds of the Mimosoideae in SC TX. One-third of the Mimosoideae represent the Grass Biome of

the Western hemisphere.

266

Journal of the Botanical Research Institute of Texas 9(1)

Papilionoideae have 84 species in the flora with the ranges mostly in the N American Atlantic Region: E

North American, Comanchian, E Madrean, Mesoamerican-Madrean, SC North American, E Prairie, and Tam-

aulipan. To the contrary, the species of papilionoids in the Big Bend flora have mostly western and southwest¬

ern North American ranges.

The basal Papilionoids in the SC TX flora are represented by two out of the three distinct lineages of the

Sophora Alliance. These two lineages have two monotypic genera in the flora. One of them, the Mexican-Me-

soamerican genus Styphnolobium, is related to E Asian-E North American disjunctive genus Cladrastis, while

the other one, a Madrean genus Calia (where Sophora secundiflora was moved), is related to the South American

genus Holocalyx (Schrire et al. 2005). The third lineage is comprised of Eurasian, pantropical, and south tem¬

perate representatives of the genus Sophora (Heenan et al. 2004) and is absent from the flora.

The pantropical dalbergioid clade is represented by only one species of the genus Zornia in the flora of SC

TX. Dalbergioids are sister to the Amorpheae clade, which is confined to the arid regions of temperate and

tropical North America (Wojciechowski 2003). The Amorpheae have three genera and 21 species in the SC TX

flora, including 18 species of a large and more widespread genus Dalea. Genus Lupinus (one sp.), whose ances¬

tor arrived from the Macaronesian-Mediterranean Region to the New World, represents South African based

temperate (Tethyan) radiation of the Genisteae clade of core genistoids (Schrire 2005). One more core genis-

toid genus in the flora, Baptisia (E North American), allies with the North American genus Thermopsis (mostly

western North American) in the Thermopsideae clade (Wang et al. 2006). Interestingly, the East Asian Ther¬

mopsis allies with the monotypic Canarian genus Anagyris and the Sino-Himalayan genus Piptanthus. Wang et

al. (2006) suspect that there was some intercontinental exchange of species in the genus Thermopsis around

the Bering Strait which started the divergence in that genus.

The Phaseoloids in the SC TX flora have eight genera of the Grass Biome affiliation, with pantropical or

amphi-atlantic disjunctions, and two genera of Psoraleae of the Succulent Biome affiliation and montane pan¬

tropical disjunction. The genus Apios has E Asia-E North American disjunction. Together the phaseoloids have

28 species in the flora with the ranges in E North America and belong to the Comanchian, E Prairie, E Madre¬

an, Mesoamerican-Madrean, or Tamaulipan geoelements.

Thus, Papilionoideae I to IV have 45 species of 13 genera in the SC TX flora (57% of all papilionoids of the

flora). All these genera with the exception of Baptisia and Apios have direct southern or Madro-Tethyan connec¬

tions and are affiliated with succulent or grass/rainforest biomes of Schrire et al. (2005).

Remaining 18 species of the papilionoids are in the Hologalegina clade of Schrire et al. (2005). Among

them are 4 species of non IRLC Hologalegina of the Succulent Biome affiliation with amphiatlantic and Tethy¬

an disjunctions. The IRLC clade has 13 species of Galegeae, with 12 species being of the genus Astragalus. Ac¬

cording to Scherson et al. (2008), Astragalus arrived from Asia via the Bering Strait. This genus is among the

few legumes in SC TX flora which represent large north temperate, mostly Old World, radiation in the IRLC

clade. The 13 th species, Lathyrus pusillus, found throughout temperate South America and disjunctly in the

south-central United States, is in the South American Notolathyrus clade. It was suggested that it originated by

long-distance dispersal of taxa to South America from Eurasia (Kenicer et al. 2005). It is outside of the rela¬

tively recent transberingian clade of these authors’ phylogeny which contains the eastern Eurasian species

(ancestral stock) and the majority of extant North American Lathyrus species.

Thus, the overwhelming majority of the native legumes in the SC TX flora has direct connections with

Madrean/Mesoamerican, Tethyan, or South American radiations. The Old World Tethyan genera (with the

exception of Astragalus ) as well as Southern Hemisphere genera present in the flora used the transatlantic

crossing in most cases.

The Cyperaceae have 69 species of 11 genera and are in fourth place in the flora of SC TX. Because they

occupy mostly wetland-aquatic habitats and have broad species ranges, they are not very informative biogeo-

graphically. However, unlike more temperate floras, e.g., flora Armenia (Saghatelyan 2006), the proportion of

temperate and subtropical species of Cyperaceae in SC TX is significantly shifted towards tropical/subtropical/

warm temperate ones. The largest genus of sedges in the flora is Cyperus (20 sp.) with Carex having only 14

Saghatelyan, Phytogeographical analysis of the South-Central Texas flora

267

species, while in more temperate latitudes Carex has more than a hundred species per flora of similar size.

Genus Eleocharis (14 sp. in SC TX) has centers of diversity in the Amazon rainforest and adjacent eastern slopes

of the South American Andes, eastern North America, California, Southern Africa, northern Australia, and

subtropical Asia. A majority of species of Cyperaceae in SC TX flora has southeastern North American-Meso-

american, E North American, and tropic/subtropical ranges. Five more Cyperaceae genera with 14 species in

the flora also have tropical to warm temperate distribution.

Fifth place in the SC TX flora is occupied by Euphorbiaceae with 68 species in 12 genera. Subfamily Aca-

lyphoideae has 4 genera and 21 species in the flora. Argythamnia and Bernardia are tropical/subtropical

American genera extending to Sonoran-Chihuahuan Subprovinces in the southwestern United States. Three

of their eight species in the flora are Edwards Plateau endemics. Tragia and Acalypha are broadly tropical/sub¬

tropical genera. Native to the Americas species of Acalypha (seven sp. total in the flora) comprise two-thirds of

the genus and have mostly eastern and southern distribution in the United States. Acalypha is sister to the

tropical West African genus Mareya and is nested among strictly Old World genera (Wurdack et al. 2005).

Subfamily Crotonoideae has two tropical/subtropical genera in the flora: Jatropha, with one Madrean and

one Chihuahuan species, and Croton. Two thirds of all species of the genus Croton are restricted to the Ameri¬

cas while one third is scattered in the Old World (Van Ee et al. 2011). Six of the nine of its species in the flora

have south-central North American ranges and belong to the two endemic North American/Mexican sections.

The remaining three species are in the two Mexico/Mesoamerica/South American sections of Van Ee et al.

( 2011 ).

Half of the species of Euphorbiaceae of the flora belong to the largest subfamily Euphorbioideae with the

majority of species in the Euphorbia subg. Chamaesyce. Subgenus Chamaesyce has the Old World origin and

includes the largest New World radiation within the Old World-centered genus Euphorbia (Yang et al. 2012).

The base of subgenus Chamaesyce consists of several clades distributed in Africa (Zimmermann et al. 2010).

New World groups in the subg. Chamaesyce had more than one origin from the Old World group. The section

Crossadenia became established in Brazil first, and later there was a separate introduction accounting for

the rest of the New World clade which includes 500 species (Yang et al. 2012). The Chamaesyce clade of that

subgenus, which originated in arid areas of North America diversified locally into three major clades: the

Acuta clade with 3 endemics in southwestern U.S.A. and N Mexico, the Peplis clade, mostly endemic to south¬

western U.S.A. and N Mexico, and Hypericifolia clade, which subsequently achieved worldwide distribution

through multiple long-distance dispersal events (Yang & Berry 2011). There are 20 species of Chamaesyce in

the SC TX flora from all three lineages with the ranges in various parts of warm temperate and subtropical

North America.

According to Zimmerman et al. (2010), Euphorbia evolved from progenitors of the subgenus Esula in

South Africa and had long distance dispersal from Africa to South America and farther to North America. All

remaining species of Euphorbia in SC TX flora belong to that subgenus Esula, members of which are mostly

herbaceous in the northern hemisphere. They fall in the Euphorbia spathulata (one sp.) group and Euphorbia

commutata (6 sp.) group. Euphorbia spathulata is one of the most widespread native species of Euphorbia in the

New World, occurring in most of North America. In the phylogeny of Zimmerman et al. (2010), the species are

nested among Eurasian and Frontal Asian species, and a southwestern North American E. alata. The remain¬

ing six species of Euphorbia of the flora are annuals of the Euphorbia commutata group of the section Peplus

(Geltman et al. 2011). It has diversified greatly from Western Asia to Europe and to North America and lacks

true succulents (Bryuns et al. 2011). Besides E. commutata (E/NE North American), the group has a SC U.S.A.,

an Edwards Plateau, and an E Comanchian species in SC TX flora. The concentration of the American species

of the subgenus Esula in the eastern and southeast-central North America coupled with wide Eurasian distri¬

bution of subgenus Esula resembles the Africa-Mediterranean/Eurasia- transatlantic crossing scenario for

some legumes and Asteraceae mentioned earlier.

The genus Stillingia (two sp. in the flora) of the Euphorbioideae is paraphyletic in an almost exclusively

Neotropical clade with three other genera (Wurdack et al. 2005).

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Journal of the Botanical Research Institute of Texas 9(1)

An example involving vicariance events comes from the former Euphorbiaceae tribe Poranthereae

(family Phyllanthaceae by APG 2009). An endemic North American clade of Andrachne section Phyllanthopsis

(Vorontsova et al. 2007) includes two species disjunctly distributed in Trans-Pecos-Coahuila, Edwards Pla¬

teau, and Ozarks Plateau. This clade is sister to the Leptopus + Actephila (Vorontsova et al. 2007) clade of meso-

phyllus shrubs. The latter clade is disjunctly distributed in the relictual humid forests of Euxinian and Hyrca-

nian Provinces of Takhtajaris Floristic System in Transcaucasia, in the Sino-Himalayan Province of Wu (1979),

and Indonesia plus a clade of Asia and Australasia. This is a striking example of subtropical Northern Hemi¬

sphere (Madro-Tethyan) radiation along the northern Tethys shore that is sister to the tropical Africa/Mada-

gascar-Australian radiation of the Andrachne s.s. clade. The latter clade is basally branching in the “clade A” of

the Porantereae phylogeny of Vorontsova et al. (2007) and consists of xerophytes from the Horn of Africa. It

started the first Madro-Tethyan (southern shore) radiation with two species in Mexico and Peru and a large

xerophytic radiation in the Tethyan Subkingdom. Thus, there might have been two transatlantic migrations

from North Africa and two single colonization events to North America, followed by vicariance events, and a

single long-distance dispersal to Peru.

Cactaceae with 40 species of 13 genera is in sixth place in the flora. The broadly American Opuntioideae

is represented by 13 species, while the remaining 27 species belong to the tribe Cacteae of Cactoideae. The spe¬

cies of Cacteae have mostly Chihuahuan-Tamaulipan ranges, while the Opuntioideae have a broader distribu¬

tion in the North American southwest. The cacti originated in northern South America in mid-Tertiary;

Opunioideae and Cactoideae occur mostly in South America while the tribe Cacteae is exclusively North

American (Nyffeler 2002).

Middle sized families of SC TX flora, Brassicaceae (37 sp. /14 gen.) and Lamiaceae (36 sp. /II gen.), are

more numerous in the Tethyan floras of the Old World where they have their major centers of diversity. For

example, flora Armenia has more than twice the number of species and a higher percentage of endemism in

these families than those of either the SC TX or Big Bend (Saghatelyan 2006). Salvia (12 sp. in SC TX/19 sp. in

Armenia) is the only large genus of mints in SC TX flora. The New World Salvia subgenus Calosphace (500 sp.

total) has centers of diversity in Mexico and South America (Walker & Sytsma 2007). Mexico is supported as

the geographic origin of Calosphace and dispersal events to South America account for its current disjunctive

distributions (Jenks et al. 2012). The close affinity of a group of South African species of Salvia to the SW

American species in Salvia sect. Heterosphace suggests a single dispersal from the Old World to the New World

with subsequent diversification (Walker et al. 2004).

For Brassicaceae, the following numbers were published in Koch & C. Kiefer (2006): ca. 900 sp. in the

Irano-Turanian region, ca. 630 sp. in the Mediterranean region, and ca. 778 sp. in North America. The family

Brassicaceae is thought to have originated in the Irano-Turanian region, where the highest species diversity is

found (Hedge 1976) and where Aethionema, the genus sister to the rest of the family, is also most diverse (Al-

Shehbaz et al. 2006; Couvreur et al. 2010). In the flora Armenia it is the fourth largest family with 195 species

(5.8% of all species of the flora), while Texas with the area 21 times larger than that of Armenia, has only 93

species (2% of the TX flora). The Bering hand Bridge is a probable route to North America from NE Asia since

most of the diversity of the family in North America is concentrated in its western half. The Beringian crossing

was suggested for Lepidium (Mummenhoff et al. 2001), Draba (Koch & Al-Shehbaz 2002), and for other Bras¬

sicaceae genera. The same Beringia crossing scenario is repeated for Tethyan groups from other families, like

Astragalus, Artemisia, Silene, Boraginoideae, and Betoideae. However, these groups are absent or not diverse in

the SC TX flora.

The Plantaginaceae has 34 species in 11 genera in SC TX flora. The discussion below is based upon the

phylogenetic tree of Albach et al. (2005). The mainly south and Central American tribe Gratioleae has five spe¬

cies of three Neotropical genera in the flora: Bacopa, Mecardonia, and Stemodia. South American Angelonia

clade is sister to the Gratioleae, and together these clades form the first major group of Plantaginaceae (Albach

et al. 2005). Cheloneae also have a New World ancestry with the basal genera in Mesoamerica and only one Old

World genus in Japan and eastern Russia (Wolfe 2002). North American members of Cheloneae diversified

Saghatelyan, Phytogeographical analysis of the South-Central Texas flora

269

initially in the Klamath Region with subsequent migration to the Rocky Mountains (Wolfe et al. 2002). An

endemic North American genus of this tribe Penstemon (271 sp. total, 10 sp. in the SC TX flora) originated in

the Rocky Mountains, radiated throughout western states, and then to the south and east (Wolfe et al. 2006). A

similar general pattern of north-to-south and west-to-east dispersal for two other genera of Cheloneae, Collin-

sia and Tonella, was found by Baldwin et al. (2011). Next tribe, mostly Old World Veroniceae (2 sp. in SC TX),

was shown to be sister to Plantago (10 sp. in the flora) which is sister to the South American paramo genus

Aragoa (Bello et al. 2002). Albach et al. (2005) suggested that an ancestor of Aragoa migrated from the Old

World to the New World and later adapted to the high Andean paramo environment.

The biogeography of the next tribe Antirrhineae (three sp. of Maurandella and Nuttallanthus in SC TX)

suggests its Old World origin with two subsequent dispersal events to the New World (Albach et al. 2005).

Based on their phylogenetic hypothesis, Albach et al. (2005) infer a New World ancestry of the Old World clade

because Russelia, Tetranema, and Cheloneae are sister to the rest of this clade, and together they are sister to the

New World Gratioleae-Angelonia clade. The disjunction of Plantaginaceae taxa between the New World and

Old World, especially the Mediterranean region, exemplifies the biogeographic pattern of Madro-Tethyan

disjunction.

The Chihuahuan-Tamaulipan desert genus Leucophyllum (2 sp.) is the only genus of the reorganized

Scrophulariaceae in the flora. Leucophyllum is in a clade with Myoporaceae and together they are sister to the

mostly South American and E Asian Buddlejaceae (Olmstead et al. 2001). The temperate to tropical family

Myoporaceae is mainly distributed in Australia, with a few representatives in South Africa, Eastern Asia, and

West Indies (Watson & Dallwitz 1992).

Solanaceae has 28 species in 10 genera in the flora. Olmstead et al. (2008) suggested the New World as the

most likely place of origin for Solanaceae. The SC TX flora has mostly large and broadly distributed genera, like

Solanum (7 sp.) and Physalis (7 sp.) with the species ranges in western and SWC North America, the Sonoran

and Madrean provinces. Lycium, which has a center of diversity in southwestern North America (Levin et al.

2011) has 5 species in the BB flora of the Madrean Region, but only one species in SC TX flora.

The cosmopolitan Pteridaceae (27 sp. in SC TX) has 24 species in common in the two Texas floras. In the

SC TX flora more species of Pteridaceae have eastern N American ranges, while the BB flora has six more

Madrean/Chihuahuan species.

The next family in the SC TX flora, mostly Laurasian temperate Rosaceae, has only 27 species (1.6%) in 10

genera. This is twice the number of Rosaceae in the BB flora, but only about one third of the Rosaceae in a typi¬

cal mountainous temperate Eurasian flora. The flora of Armenia, for example, has 163 species of Rosaceae

(4.9% of the flora). The bulk of the Rosaceae family in SC TX flora is comprised of largely distributed holarctical

genera, like Prunus (9 sp.), Crataegus (7 sp.), Rubus (3 sp.), and Rosa. Most of these genera are of the Old World

origin. There are also two species of western North American genera, Fallugia and Petrophytum, in both TX

floras.

Apiaceae has 24 species in 18 genera in the SC TX flora. Sanicula canadensis is the only species of this

widely distributed temperate genus in the flora. Southern African origins of subfamily Saniculoideae and of

both of its tribes are supported by a DIVA analysis (Calvino et al. 2008a). The genus Eryngium has four species

in the flora. The ancestor of Eryngium was inferred by Calvino et al. (2008b) to have occurred in the western

Mediterranean, whereupon the genus split to the two major clades, the Old World clade and the New World

clade. The basal lineages in the New World clade show semi-aquatic preferences, and one of them, E. nasturti-

ifolium of the “Eastern U.S.A. clade” of Calvino et al. (2008b), shows similarities with species from the western

Mediterranean. One of the biogeographical scenarios proposed by the authors for the “Eastern U.S.A. clade” is

that the dispersal from the western Mediterranean to the eastern U.S.A. or Mexico may have been by water.

Another species of Eryngium in the flora, E. yucciifolium, is in the “North American Monocotyledonous clade”

of these authors. For that clade a different scenario was proposed: it originated by dispersal from central-east

South America (SE Brazil), a center of diversification which, in its turn, originated after a separate trans-Atlan¬

tic dispersal from Africa.

270

Journal of the Botanical Research Institute of Texas 9(1)

The rest of the Apiaceae species of the flora are in the African based subfamily Apioideae which is widely

distributed in North temperate latitudes. In North America their ranges are mostly in eastern and south-east¬

ern parts. In SC TX flora seven of these species are in the four genera of the tribe Oenantheae of the “North

American Endemics clade.” It is a sister group to a clade composed primarily of Old World taxa (Downie et al.

2008). Six more genera are in the “Selineae clade” (Downie et al. 2010). Apioideae originated in southern Africa

and migrated north along two different tracks: the eastern and western. The radiation of euapioids occurred in

Eurasia with the trans-Atlantic connection being as important as the trans-Pacihc one. For both tracks, the

prevailing direction of dispersal was from the Old World to the New World (Banasiak et al. 2013). The apioids

found in SC TX flora, judging from their present distribution and connections, probably used the north tem¬

perate trans-Atlantic crossing.

Western hemisphere family Onagraceae has 24 species of three genera in the flora. There are four species

of the genus Ludwigia of one of the three basally branching Neotropical clades. The remaining 20 species are in

the tribe Onagreae which is endemic to southwestern North America (Katinas et al. 2004). The Big Bend flora

has 12 species in common with the Onagreae of the SC TX flora.

The Malvaceae has 24 species in 12 genera. All these genera are in the tribe Malveae which centers in the

Americas, but also contains South Pacific and European taxa (Tate et al. 2005). Half of these species belong to

pantropical or warm temperate to tropical/subtropical genera, with the rest in the Mexico/Texas or North/

South American genera. The Malveae of the flora have tropical/subtropical American connections.

SUMMARY OF TAXONOMIC GROUPS IN SC TX FTORA

Thus, taxonomic groups of SC TX flora exhibit several different patterns. The species of many woody genera

like Cary a, Juglans, Platanus, Acer, Quercus, Rhus, Cornus, Diospyros, Viburnum, and Lonicera, show tertiary

Laurasian flora vicariance pattern. This genera are olygotypic or monotypic in the flora. Among the larger

families mentioned in the discussion above, those with the Old World roots whose ancestors used transatlantic

crossing are much more numerous than those whose ancestors came from Asia via Beringia. These numbers

drop significantly for BB flora, exemplified by a four-fold decrease of the number of species of Apiaceae there.

Long distance dispersals of these ancestral taxa were occurring from the Mediterranean to Mesoamerica/

North America or from Africa to South America (SE Brazil) first, followed by dispersal to Mesoamerica/North

America. Madro-Tethyan vicariance and/or bidirectional migrations were important in the assembly of the

flora. Such examples are represented by Ephedra, Juniperus, Cesalpinioideae and some other legumes,

Plantaginaceae, Mahonia, Pistacia, Arbutus, Helianthemum, and Eryngium. Western Hemisphere groups have

south/north connections and are more mesophitic in SC TX than in BB, with the latter having higher numbers

of xerophytes, like Cactaceae. The xerophitic groups tend to have more western (Andean) south/north connec¬

tions. Cosmopolitan families, like Poaceae and Cyperaceae, have more tropical/subtropical representatives in

the flora than temperate ones. Mesophyllous tropical groups are more numerous in SC TX than in BB flora.

Southwestern North American centers of xerophytic speciation in many Asteraceae, the Chamaesyce clade,

Cactaceae, Agavaceae, and others, as well as the Mexican centers in Salvia, Quercus, and many Madrean genera

played an important role in the assembly of the SC TX flora.

MAJOR GENERA

The largest genera of the flora are listed in Table 2. First 40 genera of the spectrum include only 434 species

(27% of the flora). The generic diversity is high, but there are no speciose genera, with the two largest ones hav¬

ing only 20 species each. Compared to the generic spectrum of BB flora, more-speciose in SC TX are four major

Cyperaceae genera, as well as Prunus, Tradescantia, Symphiotrichum, and Yucca, while Chamaesyce and Muhlen-

bergia have more species in the BB flora.

Major geographical groups of widely distributed temperate genera in the SC TX (SC) and BB floras are as

follows.

Broadly temperate Old World-New World genera are exemplified by Salvia (12/12), Stachys (1/1),

Artemisia (1 E SC 5 BB), Solidago (7 SC 5 BB), as well as several genera of the Pooideae.

Saghatelyan, Phytogeographical analysis of the South-Central Texas flora

271

Table 2. Large and middle size genera of the SC TX flora.

Genus

No. of species in each genus

Chomoesyce, Cyperus

Dalea

Oenothera, Quercus

Juncus, Muhlenbergia

Bouteloua, Eleocharis

Carex

Salvia

Asclepias, Astragalus, Cheilanthes

Euphorbia, Erigeron, Panicum, Penstemon, Plantago, Symphiotrichum

Acacia, Aristida, Eragrostis, Physaria, Ruelia, Setaria, Sporobolus

Selaginella, Tradescantia, Yucca, Linum, Prunus

Desmanthus, Elymus, Galium, Opuntia, Polygala,

Mimosa, Solanum, Physalis

Echinocereus

North temperate Old World genera originated in the Tethyan Subkingdom and arrived to Western

North American via Beringia: Astragalus (10 SC 11BB), Silene (1 SC TX/2 BB), Allium (4 SC TX/6 BB), Lithosper-

mum (4 SC 5 BB), and Omphalodes (1/1). Some other genera, like Helianthemum (1/1) and Linum (9 SC/8 BB)

arrived to Eastern North America from Europe.

Cold temperate genera with major centers of diversity in Eastern Asia. In the United States this group

has mostly radiated in the western mountainous part and has few species in Texas: Arenaria (1 SC TX/5 BB),

Anemone (2 SC TX/3 BB), Clematis (2 SC 3 BB), Delphinium (2 SC 3 BB), Draba (3/3), and Aquilegia (2 SC 1 BB).

Widely and disjunctly distributed woody North temperate genera like Cercis (1/1), Crataegus (7 SC 1

BB), Prunus (9 SC 4 BB), Lespedeza (3 SC IBB), and Vitis (6 SC 1 BB). For Juglans (3 SC 2 BB), Fraxinus (6 SC 4

BB), and Acer (2 SC 1 BB) a North American origin was proposed (Manchester 1999), while for Lonicera (1/1)

and Viburnum (1 SC 2BB) an E Asian origin was proposed (Bell & Donoghue 2005).

Madro-Tethyan genera disjunctly distributed in the Tethyan and Madrean Suhkingdoms: Ephedra (5

SC 4 BB), Arbutus (1/1), Styrax (1 SC 0), Oligomeris (1/1), Maurandella (1/1) and Nuttallanthus (2 SC 1 BB). There

are two more genera of this group in BB, Cupressus and Peganum, absent from SC TX flora.

Comparison of the generic geographical spectra of the two floras reveals 132 different genera in the SC TX

flora not found in the BB flora (Table 3). Of these 54 (41%) are broadly tropical/subtropical genera, 14 are E

North American, eight are E Asia-E North American and eight are SC U.S.A. genera. The numbers of differen¬

tial temperate Northern Hemisphere genera are comparable: 18 genera in SC TX and 17 in BB flora. The BB

flora has total 100 differential genera 57 of which are West and SW North American genera, while 26 genera are

mostly American tropical/subtropical. These numbers illustrate that SC TX has more tropical/subtropical con¬

nections, including connections to the floras in the Old World. The BB spectrum exhibits closer ties with

western North American floras, has more species of the xeric genera of the Madrean Region and connections

to the mostly Western Neotropics.

Woody genera have more species in the SC TX flora. For example, in SC TX Crataegus and Prunus each

has 7 species; Vitis has 5 species, while in BB flora these genera have only one species each. In the SCTX flora

higher species numbers in mesophyllous, including relictual, genera and more connections with the N Ameri¬

can Atlantic Region are evident from the comparison of the generic spectra of the SC TX and BB floras. This

comparison supports their position in different chorionomic regions of the North American flora.

GEOGRAPHIC ELEMENTS OF THE FLORA

Detailed descriptions of the geographic elements are given in Saghatelyan (2009). Below are brief descriptions

and sample taxa for each geoelement. The geographic spectrum of the flora is illustrated by Tables 4 and 5. All

phytochoria are assumed within the Takhtajan (1986) System boundaries.

272

Journal of the Botanical Research Institute of Texas 9(1)

Table 3. South-Central TX genera not found in Big Bend.

Tropic/Subtropical: 54 genera

Temperate N Hemisphere: 18 genera

Temperate/Subtropical/ (Tropical) Asia (E) - E N America: 8 genera

N American: 8 genera

E N American: 14 genera

SC US (TX): 8 genera

Sw US-N Mexico: 4 genera

Cosmopolitan: 8 genera

Table 4. Geographical spectrum of the SCTX (SC) flora and Big Bend (BB) flora. Numbers of species attributed to major geographic elements are in bold; numbers of

species of the subelements are in regular typeface. The geoelements are listed by the higher chorionomic units in the systems ofTakhtajan (1986) and McLaughlin

(2007). Their grouping is somewhat modified based on the present data.

Geoelement

Of Broad Distribution-

-143 spp. (9% of flora)

Polichorous

26 / 1.6

14/0.8

Holarctical

13/.8

21/1.3

Tropical/Subtropical

30/1.8

32/2.0

American wide

74 / 4.5

101 / 6.3

Holarctic Subkingdom/N American Atlantic Region—794 spp. (49% of the flora)

North American

149/9.2

98 / 6.2

E North American / SE North American

236/14.6

37/2.3

Gulf Coast

1 7 / 1.0

Prairie

159/9.8

85 / 5.4

Comanchian

63 / 3.9

46 / 2.9

SC USA/SC North American

63 / 4.0

Tamaulipan

34/2.0

5/0.3

Edwards Plateau

71/4.3

Madrean Subkingdom/Western N America—450 spp. (28% of the flora)

W North American

58/3.6

114/7.2

SW North American

35/2.2

149/9.4

SWC North American

47/2.8

Amphitropical

10/ .6

19/1.2

Sonoran

35/2.2

82/5.2

Chihuahuan

194/12.0

253/16.0

Including Chihuahuan-Tamaulipan

Including Sonoran-Chihuahuan

South Texas /Coahuila Endemic

72 / 4.4

103/6.5

Including SCTX/SETX Endemic

Including South TX/SECTX /Coahuila

Madrean Subkingdom/Neotropical Kingdom—212 spp. (14 % of the flora)

Madrean

101/6.2 192/12.1

Including E Madrean

Mesoamerican

49/3.0

84/5.3

Caribbean

11/0.7

American Tropical/Subtropical

52/3.2

Not Established

18/1.1

25/1.6

Polichorus: wide distribution on several continents; 26 species. These are wetland, aquatic, and weedy

species of cosmopolitan herbaceous genera.

Holarctic: wide ranges in north temperate latitudes of the New and Old World; 13 species. These are hy¬

drophytes (Nuphar luted) and mesophytes (Sambucus nigra).

Tropical/Subtropical: wide distribution in tropical and subtropical latitudes; 30 species. Among them

are 8 species of grasses and 7 species of Cyperaceae. Trop/Subtr/Warm Temperate subelement: mostly in

Saghatelyan, Phytogeographical analysis of the South-Central Texas flora

273

subtropical and warm temperate regions ( Malvaviscus arboreus ) or extending from tropical to warm temperate

latitudes ( Cyperus flavescens ).

American Tropical/Subtropical: 52 species. These are American species of genera of different origin.

Some of them are species of Mexican genera extending to southern United States or of tropical Central and

northern South American genera, like Agave. Others are species of broadly tropical American genera, like Bac-

caris and Tillandsia, tropical/subtropical genera, like Acacia and Boerhavia, or cosmopolitan genera, like Ele-

ocharis. The Poaceae (15 sp.) and Cyperaceae (7 sp.) are important in this group.

American: wide distribution in the Americas; 74 species. Many of them grow in different types of habitats

(.Plantago patagonica) or are hydrophytes ( Lemna minuta, Najas guadalupensis). Species of different families are

in this group with many in Poaceae (16 sp.) and Asteraceae (6 sp.).

North American: wide ranges in temperate regions of North America; 149 species (9% of the flora). Many

hydrophytic and mesophytic species of cosmopolitan genera are in this group. They are mostly grasses (30 sp.),

Asteraceae (15 sp.), sedges and rashes (11 sp. each).

East North American: wide ranges in the Atlantic North American Region; 236 species (17% of the flora).

Among them are 52 species of SE North American subelement with more narrow southern ranges. This geo¬

element includes 60 mesophyllous woody species, like relictual Laurasian Platanus occidentals, Juglans nigra,

and Hamamelis virginiana. The SE North American subelement is exemplified by Taxodium distichum, Sabal

minor, and Smilax bona-nox. Other prominent groups in this geoelement are grasses (34 sp.), Asteraceae (32

sp.), Cyperaceae (15 sp.), Fabaceae (11 sp.), and Apiaceae (10 sp.). Connections across the Atlantic are prevalent

in many relictual amphiatlantically disjunct genera of this geoelement, for example, in Ccrcis and Gleditsia.

The species of East Asia-E North American disjunctive genera, like Lcspcdcza, are also in this group as are

representatives of tropical genera, like Bignonia, Diospyros, and Passiflora.

Gulf Coast: ranges in the Atlantic and Gulf Coastal Plain Province; 17 species. Some of them are in genera

with amphi Atlantic disjunctions, like Linum, Quercus, and Sesbania, while others are in genera with pantropi-

cal disjunctions (Zornia, Tridens).

Prairie: wide ranges in the North American Prairies Province; 159 species (9% of the flora). All but 7 spe¬

cies of this geoelement are herbaceous with the Asteraceae being most diverse (38 sp.), followed by the grasses

(18 sp.) and legumes (15 sp.). Among the Asteraceae the Astereae (14 sp.) and Heliantheae (9 sp.) are important,

while eight other clades have no more than one to three species in this geoelement. Half of the prairie grasses

are in the tropical to warm temperate subfamily Chloridoideae. Connections across the Atlantic (in Linum,

Mimosa, Rhynchosia, and Erythronium) and pantropical disjunction (in Desmodium) are important. The New

World genera, like Sisyrynchium and Triodanis, SW North American genera, like Dalea, and W North American

genera, like Penstcmon, each have several species in this geoelement.

Comanchian: ranges on the Edwards Plateau extending northward into central Oklahoma/Ozark Pla¬

teau, southward into northeastern Mexico and western Louisiana; 63 species. Asteraceae (15 sp.), Fabaceae (12

sp.), and Apiaceae (7 sp.) are important in this geoelement. Rosaceae, Lamiaceae and Agavoideae have 4 species

each. Tropical/subtropical genera, like Eryngium and Zanthoxylum, SW North American genera, like Guticrrc-

zia and Yucca, are well represented, while grasses only have two species of Chloridoideae in this geoelement.

SC USA/SC North American: from Central and South Texas extending northward to adjacent states and

southward to northern Mexican Plateau; 63 species. Among them are Asteraceae (11 sp.), Poaceae (10 sp.), and

Euphorbiaceae (6 sp.); remaining genera are from different families and have just 1-2 species each ( Plantago,

Lechea, Phlox, Linum, and Salvia).

Tamaulipan: from south-eastern and south-central Texas to Tamaulipas in Mexico; 34 species. The ma¬

jority of them are in tropical/subtropical ( Ehretia, Sesbania, Condalia) or tropical to warm temperate (Cynan-

chum) genera. A few species are in north temperate ( Crataegus ) and amphitropical ( Kramcria ) genera. All these

genera except Condalia (2 sp.) are monotypic in the flora.

W North American: wide ranging in western North America mostly north of Mexico or in its parts; 58

species. These species belong to mostly cosmopolitan ( Chenopodium ), largely Holarctic ( Corydalis, Draba),

274

Journal of the Botanical Research Institute of Texas 9(1)

North American ( Helianthus ), or tropical to warm temperate (Rhus) genera. Asteraceae with 12 species is the

largest family of this geoelement.

SW North American/SW USA: embraces southern part of the Rocky Mountain Province, Colorado Pla¬

teau, southern and eastern Great Basin, and the Sonoran Province eastward through New Mexico to south¬

western Texas and southward to adjacent northern Mexico; 35 species. These are species of W North American

genera (Fendlera, Sphaeralcea), broadly North American genera of western origin ( Penstemon ), or tropic/sub¬

tropical American genera ( Mentzelia ). The largest families in this geoelement are Asteraceae (7 sp.) and Cacta-

ceae (6 sp.).

SWC North American/SWC USA: more eastern than the previous geoelement including western part of

the Prairie Province; 471 species. These are species ofW North American ( Nama, Phacelia) genera, American-

African (Asclepias, Pomaria, Mimosa), tropical (Senna), tropical American (Bouteloua), and North American

(Monarda) genera.

Amphitropical: disjunctive ranges in warm temperate deserts of the western North and South America;

10 species of peculiar desert genera, like Koeberlinia, Aloysia, and Kallstroemia.

Madrean: ranges embracing the Madrean Region, or mostly the mountains of Mexico; 101 species. This

group has 21 species of woody genera (Quercus, Acer, and Rhus), 17 species of Asteraceae, 11 species each of the

legumes (Calliandra) and ferns (Notholaena), 10 species of grasses, 6 of Euphorbiaceae (Jatropa ), 6 species of

Oleaceae (Fraxinus), and 3 species of gymnosperms (Ephedra and Pinus). The Madrean element has different

subelements, North, South, West, and East. The largest, East Madrean subelement has 35 species: mostly in

the Sierra Madre Oriental Province as defined by Morrone et al. (1999) extending northward to the Edwards

Plateau and mountains in Trans-Pecos (Quercus laceyi).

Mesoamerican: wide ranges in Mesoamerica and the Caribbean Region extending to southern United

States and northern South America-49 species. Many of them are in pantropical (Cissus, Indigofera) or tropical

to temperate (Ruellia, Cynanchum, Ipomoea) genera; some, like Eustoma, are in Mega Mexico III genera (Saghat-

elyan 2009), or American-African genera (Sideroxylon and Fantana). There are also species of the Madro-

Tethyan genera (Arbutus and Pistacia), as well as temperate to subtropical American genera (Oenothera). The

largest in the Mesoamerican geoelement family Asteraceae has only 9 species, followed by Poaceae (5 sp.) and

Cyperaceae (4 sp.). Caribbean subelement is extending to SE North America and has 23 species of different

tropical/subtropical genera including six genera of grasses.

Sonoran: embracing mostly the Sonoran Sub-province or the entire Sonoran Province; 36 species. These

are xerophytes from different families, the largest being Fabaceae (5 sp.) and Cactaceae (3 sp.). Eleven of these

species are woody (Mahonia trifoliata, Chilopsis linearis, Quercus grisea).

Chihuahuan: ranges in the Chihuahuan Sub-province including Trans-Pecos and Edwards Plateau; 194

species. There are 38 species of Asteraceae, 18 of cacti, 19 of legumes, 12 of grasses, 10 of Euphorbiaceae, 8 spe¬

cies each of Solanaceae and Boraginaceae (mostly in tropical tribes, Heliotropium and Fiquilia), 5 species of

Acanthaceae, 4 each of Brassicaceae, Agavaceae, Nyctaginaceae, and Lamiaceae. Sonoran-Chihuahuan sub¬

element has ranges in the Sonoran and Chihuahuan Subprovinces; 21 species (Fycium berlandieri, Cheilanthes

villosa). Chihuahuan-Tamaulipan subelement has ranges in the Chihuahuan Subprovince extending east¬

ward to the Tamaulipan Subprovince; 66 species (Matelea reticulata, Acacia rigidula, Dasylirion texanum).

South Texas Endemic: 72 species. This element includes 35 species of SC Texas/SEC Texas Endemic

and 37 species of South Texas-Coahuila subelements. SC Texas Endemic subelement is mainly found on

Edwards Plateau extending to Trans-Pecos and along the Rio Grande valley to N Tamaulipan Subprovince.

This element is closest to the Chihuahuan or Tamaulipan geoelement with narrow ranges in different parts of

South Texas or SW Texas-Edwards Plateau, and Coahuila/Nuevo-Leon of northern Mexico. The species are

from different xerophytic genera of different families and mostly have southern connections (Opuntia edward-

sii, Hesperaloefunifera, Menodora heterophylla).

Edwards Plateau (Endemic): 71 species endemic or sub endemic to the Edwards Plateau. They are from

different families, mostly Asteraceae (10 sp.), Fabaceae (6 sp.), Poaceae (4 sp.), and Euphorbiaceae (4 sp.).

Saghatelyan, Phytogeographical analysis of the South-Central Texas flora

275

The proportions of different geographical elements in the two TX floras are presented in Table 4. In the SC

TX flora the largest geoelement, E North American, has 236 species, which, combined with 63 Comanchian,

64 SC USA, 33 Tamaulipan, 159 Prairie, 17 Gulf Coast species, 35 SC/SE Texas endemics, and 73 Edwards

Plateau endemics, make 680 species or 42% of the flora. They belong to the N American Atlantic Region of

North America. This group only has 11% in the Big Bend flora. The Chihuahuan element has 194 species (12%)

in SC TX, but more species, 253 (16%), in Big Bend. There are only 101 species (6.2%) of the Madrean element

in SC TX which has twice as much species in Big Bend (12.1%). The entire Madrean Region including the So¬

noran, Chihuahuan, and Tamaulipan subprovinces, has 457 species (28%) in SC TX and 798 species (50%) in

BB. The combined tropical element has 142 species (8%) in Big Bend and 142 (8.7%) in SC TX. Thus, the two

floras are comparable in their southern connections, both having almost the same weight of tropical species.

The influence of the Madrean Region is significantly stronger in the Big Bend flora, which has two times more

Madrean and Sonoran species, while SC TX has more gulf coastal and SC North American species than BB.

Thus, these two Texas floras, though in proximity to each other, represent two different chorionomic units: the

N American Atlantic Region (SC TX), and Madrean Region (BB) of the North American flora.

CONCLUSIONS

There are 1619 native species of 626 genera and 149 families in the SC TX flora. Half of the 232 species of the

largest family Asteraceae represent local diversification in the largest North American Heliantheae Alliance.

Together with the taxa of several other clades of the family found in the flora they are mostly native to south-

eastern/southwestern/southern North America or have Central/South American connections. The ancestors

of the basally branching taxa of many of these clades had either Mediterranean to Mesoamerica to North

America or Africa to South America to North America long distance dispersals. Second largest family Poaceae

(199 sp.) has 67% of its species in the tropical to warm temperate subfamilies. A majority (73%) of all the

Fabaceae (120 sp.) have connections with SE N American/Madrean/Mesoamerican, Tethyan, or South Ameri¬

can radiations. Old World Tethyan, as well as Southern Hemisphere legume genera found in the flora in most

cases used the transatlantic crossing. The Cyperaceae (69 sp.), unlike those in more temperate floras, have a

majority of species in tropical/subtropical/to warm temperate genera. Half of the Euphorbiaceae (68 sp.) spe¬

cies are in the Euphorbioideae, mostly in the Euphorbia subgenus Chamaesyce, which originated in arid North

America after an introduction from the Old World. The species of Acalyphoideae and Crotonoideae comprise

the other half the family and are in tropical/subtropical, mostly American genera. The middle sized families

Brassicaceae (37 sp.) and Lamiaceae (35 sp.), with the only large genus Salvia are more prominent in the Old

World Tethyan floras, where their major centers of diversity lie. To the contrary, the species of Plantaginaceae

(34 sp.) belong to the mainly South and Central American clades with the basal genera in Mesoamerica and a

pattern of Madro-Tethyan disjunction. The largest genera of the flora are Chamaesyce, Cyperus, Dalea, Oeno¬

thera, Quercusjuncus, Muhlenbergia, Bouteloua, Eleocharys, Carex, and Salvia. There are 132 differential genera

in the SC TX flora not found in the Big Bend flora. These include 54 tropic/subtropic genera, 34 temperate

Northern Hemisphere, 14 E North American, 12 SC/SW USA-N Mexico, and 8 cosmopolitan genera. In the

geographical spectrum of the SC TX flora the largest geoelement is comprised of 236 E North American spe¬

cies, which, combined with Comanchian, Tamaulipan, prairie, SC USA, gulf coast species, and local endemics,

make 680 species (42% of the flora) of the North American Atlantic Region. This group only has 11% in the Big

Bend flora. The Holarctic Subkingdom in general is represented by 793 species (49% of the flora). The Madrean

Subkingdom/Western N America have 449 species (28% of the flora), followed by the Madrean Region/Neo¬

tropical Kingdom with 212 species (14% of the flora).

In general, in the SC TX flora the southern and eastern connections prevail. Transatlantic crossings, as

well as S/N American bidirectional migrations were most important in the history of its assembly. This study

supports placement of the SC TX flora in the N American Prairies Province of the N American Atlantic Floristic

Region. In the McLaughlin (2007) classification it is a part of the Comanhian province. However results of this

study support placement of the Comanchian chorion, including the SC TX flora, in the Atlantic Region rather

276

Journal of the Botanical Research Institute of Texas 9(1)

the South Western Region of McLaughlin. The BB flora, which is just west of the continental divide from the SC

TX flora, has a placement in a different region, namely the Chihuahuan Province of the Madrean Region.

APPENDIX 1

Checklist and area diagnoses of the flora of SC Texas. The following list of species was extracted mostly from the Synthesis of the North

American Flora (Kartesz 2013) and it follows, with few exceptions, all the nomenclatural combinations as well as author citations of this

source. The families of vascular plants are arranged in alphabetical order as are genera and species. Numbers by each family name indicate

species/genera ratios in the family. The ranges of some species could not be referred to a particular geoelement. These are noted with a

question mark in the checklist and are omitted from the analysis.

Abbreviations:

adv els —adventive elsewhere

Afr —African

Amphitrop —Amphitropical

Amer —American

Apach —Apachian

AZ —Arizona

Carib —Caribbean

C —central

Chih —Chihuahuan

Coah —Coahuila

CO —Colorado

Comanch —Comanchian

disj —disjunctive

E— East

Edw Plat —Edwards Plateau

FL —Florida

Gulf —Atlantic & Gulf Coastal Plain

intr els —introduced elsewhere

Madr —Madrean

Mesoam —Mesoamerican

Mont —montane

N —North

NM —New Mexico

OK —Oklahoma

Polichor —polichorous

Prair —Prairie

Roc Mt —Rocky Mountains Province

S —South

SC —south-central

Son —Sonoran Province

Tam —Tamaulipan Province

Temp —temperate

Trop/Subtr —tropical/subtropical

TX— Texas

USA —United States of America

W— West

WO —Wyoming

Counties: Bandera, Concho, Crockett, Edwards, Gillespie, Irion, Kendall, Kerr, Kimble, Kinney, Llano, Mason, Medina, Menard, Reagan,

Real, Schleicher, Sutton, Tom Green, Uvalde, and Val Verde.

ANEMIACEAE 1/1

Anemia mexicana Mesoam

ACANTHACAE 7/19

Anisacanthus quadrifidus Chih

Carlowrightia texana Chih

Carlowrightia torreyana SC TX-Coah

Dicliptera brachiata E N Amer

Dyschoriste linearis SC USA

Justicia americana E N Amer

Justiciapilosella Chih-Tam

Justicia warnockii S TX-Coah (SWTX)

Justicia wrightii S TX

Ruellia caroliniensis E N Amer

Ruellia corzoi Chih

Ruellia drummondiana Edw Plat

Ruellia humilis E N Amer (+ Prair)

Ruellia metziae Edw Plat

Ruellia nudiflora Mesoam-Madr

Ruellia occidental is E Madr

Ruellia parryi Chih

Ruellia yucatana Ta m

Yeatesia platystegia Tam

ACERACEAE 1/2

Acer grandidentatum N Madr Mont

Acer negundo N Amer

AGAVACEAE 6/15

Agave lechuguilla Chih

Camassia scilloides E N Amer

Echeandia flavescens Son (W)

Hesperaloe funifera SC TX-Coah

Hesperaloeparviflora Comanch

Nolina lindheimeriana Edw Plat

Nolina texana SWC USA

Yucca arkansana Comanch

Yucca constricta SC TX (Edw Plat)

Yucca glauca Prair

Yucca pallida Comanch (S)

Yucca reverchonii Edw Plat (W)

Yucca rupicola Comanch (S)

Yucca torreyi Chih

Yucca treculeana Chih-Tam

AIZOACEAE 1/1

Sesuvium verrucosum Amer

ALISMATACEAE 2/5

Echinodorus berteroi Amer

Sagittaria brevirostra N Amer (N-C)

Sagittaria calycina N Amer

Sagittaria longiloba N Amer (W)

Sagittaria platyphylla N Amer (E)

AMARYLIDACEAE 3/7

Allium canadense E N Amer

Allium drummondii SC USA (Prair)

Allium kunthii Madr

Allium perdu Ice Prair

Cooperia chlorosolen SC USA

Cooperia pedunculata SEC USA

Nothoscordum bivalve E N Amer (+ Mesoam)

AMARANTHACEAE 9/17

Alternanthera caracasana Amer Trop/Subtr

Amaranthus albus Polichor

Amaranthus arenicola Prair wide

Amaranthus crassipes Carib

Amaranthus palmeri N Amer

Amaranthus polygonoides Carib

Amaranthus scleropoides Chih-Tam

Amaranthus tuberculatus N Amer

Celosia nitida Amer Subtr

Froelichia floridana N Amer (Prair-SE)

Saghatelyan, Phytogeographical analysis of the South-Central Texas flora

277

Froelichia gracilis N Amer

Gomphrena haageana S TX-Coah?

Gossypianthus lanuginosus N Amer (SC + Carib)

Guilleminea densa Madr

Iresine heterophylla Son wide

Iresine rhizomatosa E N Amer

Tidestromia lanuginosa SC USA

ANACARDIACEAE 3/7

Cotinus obovatus E N Amer disj

Pistacia mexicana Mesoam-E Madr

Rhus copallinum E N Amer

Rhus lanceolata SC N Amer

Rhus microphylla Son-Chih

Rhus trilobata W N Amer (+ W Prair)

Rhus virens Madr (E)

APIACEAE 19/25

Ammoselinum butleri E N Amer

Ammoselinum popei S Prair

Bern la erecta Holarctical

Bifora americana Comanch (E N Amer)

Bowlesia incana Amer (Warm Temp)

Centella erecta SE N Amer (+ Mesoam-Carib)

Chaerophyllum tainturieri E N Amer

Cicuta maculata N Amer

Cymopterus macrorhizus Comanch (NC TX)

Daucosma laciniata ?

Daucus pusillus N Amer

Eryngium diffusum Comanch

Eryngium leavenworthii Comanch

Eryngium nasturtiifolium Madr (Mesoam-Madr)

Eryngium yuccifolium E N Amer

Eurytaenia texana SC USA

Hydrocotyle prolifera N Amer

Hydrocotyle umbel lata Amer

Limnosciadium pinnatum SEC N Amer (Comanch?)

Polytaenia nuttallii Prair

Polytaenia texana Edw Plat?

Ptilimnium nuttallii SEC N Amer

Sanicula canadensis E N Amer

Spermolepis echinata N Amer (S)

Spermolepis inermis Prair

APOCYNACEAE 3/4

Amsonia ciliata SE N Amer (+ Comanch)

Amsonia longiflora Chih

Apocynum cannabinum N Amer

Telosiphonia macrosiphon S TX-Coah

AQUIFOLIACEAE 1/2

Ilex decidua E N Amer

llexvomitoria SE N Amer

ARACEAE 1/1

Arisaema dracontium E N Amer

ARECACEAE 1/1

SabaI minor SE N Amer

ARISTOLOCHIACEAE 2/3

Aristolochia coryi S TX Endemic

Aristolochia serpentaria E N Amer

Isotrema tomentosum E Prair-E N Amer

ASCLEPIADACEAE 4/21

Asclepias asperula SWC N Amer

Asclepias emoryi Tam

Asclepias engelmanniana SWC N Amer

Asclepias incarnata E N Amer

Asclepias latifolia SWC N Amer

Asclepias oenotheroides Mesoam (+ SC USA)

Asclepias texana S TX-Coah

Asclepias tuberosa N Amer (E + SW)

Asclepias verticillata E N Amer

Asclepias viridiflora E N Amer

Asclepias viridis SE N Amer (+ Prair)

Cynanchum barbigerum Tam

Cynanchum maccartii Edw Plat-Tarn

Cynanchum racemosum Mesoam-Madr

Funastrum crispum SWC USA

Funastrum cynanchoides Madr (N & W)

Matelea biflora Comanch (W)

Matelea edwardensis Edw Plat

Matelea gonocarpos E N Amer

Matelea reticulata Chih-Tam

Matelea sagittifolia S TX Endemic

ASPARAGACEAE 1/1

Dasylirion texanum Chih-Tam

ASPLENIACEAE 1/1

Asplenium resiliens N Amer

ASTERACEAE 98/232

Achillea millefolium Holarctical

Acourtia nana Son-Chih

Acourtia runcinata Chih-Tam

Acourtia wrightii Son-Chih

Ageratina altissima E N Amer

Ageratina havanensis E Madr (+ Carib)

Amblyolepis setigera Chih-Tam

Ambrosia artemisiifolia Amer (mostly E)

Ambrosia confertiflora SWC N Amer

Ambrosia monogyra Son-Chih

Ambrosia psilostachya N Amer wide (Polichor)

Ambrosia trifida N Amer (mostly E)

Amphiachyris amoena Comanch (C/NCTX Endemic)

Amphiachyris dracunculoides Prair

Aphanostephus ramosissimus Chih-Tam

Aphanostephus riddellii Chih

Aphanostephus skirrhobasis SC N Amer

Arnoglossum plantagineum E Prair

Artemisia ludoviciana N Amer

Astranthium ciliatum Prair (SE)

Baccharis neglecta Chih-Tam

Baccharis salicifolia Amer Trop/Subtr

Baccharis salicina SW N Amer

Baccharis texana Prair (S)

Bahia absinthifolia Son-Chih

Baileya multiradiata Madr

Berlandiera lyrata Madr

Bidens bipinnata E N Amer-E Asian

Brickellia cylindracea Edw Plat

Brickellia dentata Edw Plat (+ E TX)

Brickellia eupatorioides N Amer

Brickellia laciniata Chih wide

Chaetopappa bellidifolia Edw Plat Endemic

Chaetopappa bellioides Chih-Tam

Chaetopappa effusa Edw Plat Endemic

Chaetopappa ericoides SW N Amer

Chaetopappa parryi Chihun-Tam

Chaptalia texana E Madr

Chloracantha spinosa Mesoam-Madr

Chromolaena bigelovii Chih?

278

>9(1)

stlyS-C)

279

air (C-E)

■(S&C)

280

Journal of the Botanical Research Institute of Texas 9(1)

Physaria sessilis Edw Plat

Rorippa curvipes W N Amer

Rorippo sessilifloro E Prair (+ E N Amer)

Rorippa teres Gulf

Selenia jonesii S TX endemic (WC)

Sibara viereckii Tam

Sibara virginica E N Amer

Streptanthus bracteatus Edw Plat endemic

Streptanthus carinatus Son-Chih

Streptanthus platycarpus Edw Plat endemic (W)

BROMELIACEAE 1/2

Tillandsia recurvata Amer Trop/Subtr

Tillandsia usneoides Amer Trop/Subtr

BUDDLEJACEAE 2/2

Buddleja racemosa Edw Plat endemic (S)

Polypremum procumbens Amer Trop/Subtr

CACTACEAE 13/40

Ancistrocactus brevihamatus S TX (SW-SC)

Ancistrocactus scheeri Ta m

Ariocarpus fissuratus Chih

Coryphantha echinus Chih

Coryphantha macromeris Chih

Coryphantha sulcata SC N Amer

Cylindropuntia davisii SW N Amer

Cylindropuntia imbricata SW N Amer

Cylindropuntia kleiniae Son-Chih

Cylindropuntia leptocaulis Chih-Tam

Cylindropuntia tunicata Madr (+ S Amer)

Echinocactus horizonthalonius Chih

Echinocactus texensis SC N Amer

Echinocereus coccineus SW N Amer

Echinocereus enneacanthus Chih-Tam

Echinocereus pectinatus Chih-Tam

Echinocereus reichenbachii SC N Amer

Echinocereus stramineus Chih

Echinocereus triglochidiatus SW N Amer

Epithelantha micromeris Son-Chih

Escobaria emskoetteriana E Chih-Tam

Escobaria missouriensis W N Amer (WC)

Escobaria vivipara W N Amer

Ferocactus hamatacanthus Chih-Tam

Glandulicactus uncinatus var. wrightii Ch\h

Grusonia schottii Chih-Tam

Hamatocactus bicolor Tam

Mammillaria heyderi Son (+ Tam)

Mammillaria lasiacantha Son-Chih

Mammillaria prolifera var.texana Chih-Tam

Mammillaria sphaerica Tam

Neolloydia conoidea Chih-Tam

Opuntia atrispina S TX endemic

Opuntia edwardsii S TX endemic

Opuntia engelmannii Son wide

Opuntia humifusa E N Amer

Opuntia macrocentra Son wide

Opuntia macrorhiza SW N Amer (+ Prair)

Opuntia phaeacantha SW N Amer

Opuntia strigil S TX endemic

CALLITRICHACEAE 1/2

Callitriche heterophylla Amer (C & N)

Callitrichepeploides SE N Amer

CAMPANULACEAE 3/9

Campanula reverchonii Edw Plat endemic

Lobelia berlandieri Chih

Lobelia cardinalis N Amer

Triodanis biflora N Amer

Triodanis coloradoensis Edw Plat

Triodanis holzingeri Prair (CS)

Triodanis leptocarpa Prair (E & N)

Triodanis perfoliata N Amer

Triodanis texana Comanch (CE/NC TX endemic)

CAPPARACEAE 3/4

Cleomella angustifolia Prair

Koeberlinia spinosa Amphitrop (Son-Chih)

Polanisia dodecandra N Amer

Polanisia erosa Comanch?

CAPRIFOLIACEAE 4/4

Lonicera albiflora SWC N Amer disj Mont

Sambucus nigra Holarctical

Symphoricarpos orbiculatus E N Amer

Viburnum rufidulum E N Amer

CARYOPHYLLACEAE 7/12

Arenaria benthamii SC N Amer (C-STX)

Cerastium brachypodum N Amer

Loeflingia squarrosa W N Amer

Paronychia drummondii Comanch

Paronychiajamesii Prairie (W)

Paronychia lindheimeri Edw Plat

Paronychia monticola S TX-Coah

Paronychia setacea CS TX endemic

Paronychia virginica E N Amer disj

Sagina decumbens N Amer

Silene antirrhina N Amer

Stellaria cuspidata Mesoam (+ Andean)

CELASTRACEAE 3/3

Celastrus scandens N Amer (NE-NC)

Mortonia sempervirens Chih

Schaefferia cuneifolia Chih-Tam

CERATOPHYLLACEAE 1/1

Ceratophyllum demersum Pol ichor

CHENOPODIACEAE 2/8

Atriplex argentea W N Amer

Atriplex canescens W N Amer

Chenopodium berlandieri N Amer

Chenopodium desiccatum W N Amer

Chenopodium incanum W N Amer

Chenopodium leptophyllum W N Amer

Chenopodium pratericola N Amer

Chenopodium simplex N Amer

CISTACEAE 2/4

Helianthemum georgianum SE N Amer (SC-SE)

Lechea mucronata E N Amer

Lechea san-sabeana Comanch (ETX)

Lechea tenuifolia E N Amer

CLUSIACEAE 1/3

Hypericum drummondii E N Amer

Hypericum gentianoides E N Amer

Hypericum mutilum E N Amer

COMMELINACEAE 3/10

Commelina erecta E N Amer

Tinantia anomala Edw Plat endemic

Tradescantia brevifolia STX (Trans-Pecos)-Coah

Tradescantia edwardsiana C TX endemic

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Tradescantia gigantea Comanch(CNE TX)

Tradescantia hirsutifloro E N Amer

Trodescontio humilis SE TX

Tradescantia occidentalis Prair wide

Tradescantia ohiensis E N Amer

Tradescantiapedicellata Edw Plat endemic

CONVOLVULACEAE 6/13

Bonamia repens N Chih

Convolvulus equitans SWC N Amer

Cressa truxillensis Madr

Dichondra carolinensis E N Amer (+ Japan)

Amer-Asian?

Dichondra recurvata E Edw Plat endemic

Evolvulus alsinoides Madr

Evolvulus nuttallianus WC N Amer

Evolvulus sericeus Amer Trop/Subtr

Ipomoea cordatotriloba Amer Trop/Subtr

Ipomoea costellata Son

Ipomoea leptophylla Prair

Ipomoea lindheimeri Chih

Ipomoea pandurata E N Amer

CORN ACEAE 1/1

Cornus drummondii E Prair

CRASSULACEAE 2/3

Crassula aquatica Amer (N & C) Coastal

Sedum nuttallianum Comanch

Sedum wrightiiSVJC N Amer (Apach)

CROSSOSOMATACEAE 1/1

Glossopetalon texense Edw Plat endemic

CUCURBITACEAE 5/6

Cucurbita foetidissima Madr wide

Cyclanthera dissecta Mesoam-Madr

Ibervillea lindheimeri SC N Amer-Gulf

Ibervillea tenuisecta Chih

Melothriapendula E N Amer (Gulf)

Sicyos angulatus N Amer (C-NE)

CUPRESSACEAE 2/4

Juniperus ashei Comanch

Juniperus pinchotii SWC USA

Juniperus virginiana E N Amer

Taxodium distichum SE N Amer

CUSCUTACEAE1/6

Cuscuta cuspidata Prair

Cuscuta indecora N Amer

Cuscuta pentagona Amer

Cuscuta squamata Chih

CYPERACEAE 11/69

Bulbostylis capillaris Amer (Amphipacific)

Bulbostylis juncoides S Mesoam

Carexalata E N Amer (Gulf)

Carexamphibola E N Amer

Carexaureolensis SC US

Carexaustrina Prair

Carexbrevior N Amer (N & C Prair)

Carex cephalophora E N Amer (NE-NC)

Carexcherokeensis SE N Amer

Carexedwardsiana Edw Plat endemic

Carex emoryi N Amer (C & NE)

Carexmicrodonta Prair (CS)

Carex muehlenbergii E Prair-E N Amer

Carexperdentata Comanch

Carexplanostachys SC N Amer

Cladium mariscus Subtr

Cyperus acuminatus N Amer

Cyperus croceus SE N Amer-C Amer

Cyperus echinatus E N Amer

Cyperus elegans Mesoam-S Amer

Cyperus erythrorhizos N Amer

Cyperus esculentus Amer-Afr (Polichor)

Cyperus flavescens Trop/Subtr/Warm

Cyperus haspan Trop/Subtr

Cyperus hermaphrodites Amer Trop/Subtr

Cyperus lupulinus E N Amer

Cyperus odoratus Amer

Cyperus pseudothyrsiflorus SC USA

Cyperus pseudovegetus E N Amer-C Amer

Cyperus reflexus S Amer-Mesoam

Cyperus retroflexus S N Amer

Cyperus retrorsus SE N Amer-C Amer

Cyperus setigerus S Prair

Cyperus squarrosus Amer-Afr

Cyperus strigosus N Amer

Cyperus thyrsiflorus Amer Trop/Subtr

Eleocharis acicularis Amer

Eleocharis atropurpurea Trop/Subtr/Warm

Eleocharis cellulosa Amer Trop/Subtr

Eleocharis compressa E N Amer (+ Prair)

Eleocharis flavescens Amer Trop/Subtr

Eleocharis geniculata Amer-Afr

Eleocharis interstincta Amer Trop/Subtr

Eleocharis lanceolata SE Prair

Eleocharis montevidensis Amer (N Amer-C Amer)

Eleocharis obtusa N Amer

Eleocharis occulta Comanch

Eleocharispalustris Holarctical

Eleocharisparvula Holarctical

Eleocharis quadrangulata N Amer (E)

Eleocharis rostellata N Amer-C Amer

Fimbristylis autumnalis Amer

Fimbristylis dichotoma Trop/Subtr

Fimbristylispuberula E N Amer (Prair-SE N Amer)

Fimbristylis vahlii Amer Trop/Subtr

Fuirena simplex N Amer-Mesoam

Fuirena squarrosa SE N Amer

Isolepis carinata SE N Amer

Kyllinga brevifolia Trop/Subtr

Lipocarpha aristulata N Amer

Lipocarpha micrantha Amer (+ Afr)

Rhynchospora colorata Carib (Gulf-C Amer-Carib)

Rhynchospora nivea Comanch

Schoenoplectus californicus Amer Trop/Subtr/Warm

Schoenoplectus hallii Prair

Schoenoplectus maritimus W N Amer

Schoenoplectuspungens Polichor

Schoenoplectus saximontanus N Amer (W & C)

Schoenoplectus tabernaemontani Polichor

DENNSTAEDTIACEAE 1/1

Dennstaedtia globulifera Amer Trop/Subtr

DRYOPTERIDACEAE 5/5

Cystopteris utahensis ?

Dryopteris cinnamomea Madr (S & W)

Onoclea sensibilis E N Amer

Tectaria heracleifolia Mesoam (+ N S Amer)

Woodsia obtusa E N Amer

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Journal of the Botanical Research Institute of Texas 9(1)

EBENACEAE 1/2

Diospyros texana Chih-Tam

Diospyros virginiono E N Amer

ELATINACEAE 2/2

Bergia texana ? (W N Amer-Prair)

Elatine brachysperma ? (N Amer)

EPHEDRACEAE 1/5

Ephedra antisyphilitica Chih-Tam (+ NE TX, OK)

Ephedra aspera Madr disj

Ephedrapedunculata Chih-Tam

Ephedra torreyana SW N Amer

Ephedra trifurca N Madr

EQUISETACEAE 1/3

Equisetum xferrissii ?

Equisetum hyemale Holarctical

Equisetum laevigatum N Amer

ERICACEAE 1/1

Arbutus xalapensis Mesoam

ERIOCAULACEAE 1/1

Eriocaulon koemickianum E N Amer disj

EUPHORBIACEAE 12/68

Acalypha gracilens E N Amer

Acalypha monococca Prair (E)

Acalypha monostachya Madr

Acalypha ostryifolia Amer (C & N Amer)

Acalypha phleoides Madr wide

Acalypha radians S TX (SCE TX)

Acalypha virginica E N Amer

Argythamnia aphoroides Edw Plat

Argythamnia argyraea S TX endemic

Argythamnia humilis S Prair

Argythamnia mercurialina Prair-W Amer

Argythamnia neomexicana N Madr

Argythamnia simulans Edw Plat endemic

Bernardia myricifolia ? Son disj

Bernardia obovata Chih (N)

Chamaesyce acuta Chih (N)

Chamaesyce albomarginata Madr (N)

Chamaesyce angusta S TX (Trans-Pecos-Edw Plat)

Chamaesyce bombensis Gulf (C Amer-Gulf)

Chamaesyce chaetocalyx SW N Amer (Apach)

Chamaesyce cinerascens Chih-Tam

Chamaesyce fendleri WC USA

Chamaesyce glyptosperma N Amer

Chamaesyce hypericifolia Amer Trop/Subtr

Chamaesyce hyssopifolia Amer Trop/Subtr

Chamaesyce jejuna Chih (N)

Chamaesyce lata SWC USA

Chamaesyce maculata E N Amer (+ E Asia)

Chamaesyce missurica Prair

Chamaesyce nutans N Amer (C & E)

Chamaesyce prostrata N Amer (C & N)

Chamaesyce serpens Amer

Chamaesyce serrula Son-Chih

Chamaesyce stictospora N Amer

Chamaesyce villifera Mesoam-Madr

Cnidoscolus texanus SC USA

Croton capitatus E Prair

Croton dioicus Chih-Tam

Croton fruticulosus Chih-Tam

Croton glandulosus Amer Trop/Subtr

Croton incanus E Madr (+ Tam)

Croton lindheimerianus S Prair

Croton monanthogynus E N Amer

Croton pottsii Son-Chih

Croton texensis SWC N Amer

Euphorbia brachycera W N Amer

Euphorbia commutata E N Amer

Euphorbia cyathophora N Amer

Euphorbia dentata N Amer (+ C Amer)

Euphorbia heterophylla N Amer

Euphorbia longicruris SC USA (Comanch?)

Euphorbia marginata Amer

Euphorbia roemeriana Edw Plat endemic

Euphorbia spathulata N Amer

Euphorbia wrightii ? S TX-Coah

Jatropha cathartica Chih-Tam

Jatropha dioica Madr

Leptopusphyllanthoides Comanch

Phyllanthus polygonoides SC N Amer

Stillingia sylvatica SEC N Amer

Stillingia texana SC USA (+ Coah)

Stillingia treculiana E Madr-Tam

Tragia amblyodonta Son Wide

Tragia betonicifolia S Prair

Tragia brevis pica Comanch

Tragia leptophylla STX endemic

Tragia nigricans Edw Plat endemic

Tragia ramosa SWC USA

FABACEAE 41/121

Acacia angustissima Amer Trop/Subtr

Acacia berlandieri Chih-Tam

Acacia constricta Madr wide

Acacia farnesiana Trop

Acacia greggii Madr

Acacia neovernicosa Son

Acacia rigidula Chih-Tam

Acacia roemeriana Chih

Acacia schottii N Chih

Amorpha fruticosa N Amer

Amorpha roemeriana S Edw Plat endemic

Apios americana E N Amer

Astragalus brazoensis S TX-Coah (SCE TX)

Astragalus canadensis N Amer

Astragalus crassicarpus Prair wide

Astragalus lindheimeri Comanch (C TX-Comanch)

Astragalus lotiflorus Prair

Astragalus mollissimus WC N Amer

Astragalus nuttallianus WC N Amer

Astragalusplattensis Prair (W)

Astragalus pleianthus Edw Plat endemic

Astragalus waterfallii Chih

Astragalus wrightii S TX endemic

Baptisia bracteata E Prair

Bauhinia lunarioides Chih

Calliandra conferta Madr (S, C, & E)

Cal Hand ra eriophylla Madr

Centrosema virginianum Mesoam-S Amer

Cercis canadensis E N Amer

Chamaecrista fasciculata E N Amer

Clitoria mariana E N Amer

Da lea a urea Prair

Dalea bicolor Son

Da lea compacta Comanch

Dalea emarginata Tam

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Da lea enneandra Prair

Dalea formosa SWC N Amer

Da lea frutescens E Madr

Dalea greggii E Madr

Dalea hallii Comanch (TX endemic)

Dalea Ianiceps N Chih

Dalea lasiathera Edw Plat (+ S TX-Coah)

Dalea multiflora Prair

Dalea nana Son wide

Dalea pogonathera Son

Daleapolygonoides SW N Amer (disj)

Dalea purpurea N Amer (NC)

Dalea tenuis Comanch

Dalea wr/ghf/V Son-Chih

Desmanthus acuminatus SE TX endemic

Desmanthus glandulosus Son-Chih (N)

Desmanthus illinoensis Prair

Desmanthus leptolobus S Prair

Desmanthus obtusus Chih

Desmanthus reticulatus Edw Plat endemic

Desmanthus velutinus Chih

Desmanthus virgatus Amer (except N)

Desmodium lindheimeri Tam

Desmodium paniculatum E N Amer

Desmodium psilophyllum Mesoam (+ E Madr)

Desmodium sessilifolium E Prair

Desmodium tweedy! Comanch

Eysenhardtia texana E Chih-Tam

Galactia canescens Chih-Tam

Galactia heterophylla S TX endemic (CS TX)

Galactia marginalis AmerTrop/Subtr

Galactia texana Edw Plat endemic

Gleditsia triacanthos E N Amer

Hoffmannseggia drepanocarpa SW USA

Hoffmannseggia glauca Amphitrop

Hoffmannseggia oxycarpa Chih

Indigofera lindheimeriana Chih

Indigofera miniata Mesoam-Carib

Lathyrus pusillus ?

Lespedeza stuevei E N Amer (S)

Lespedeza texana Comanch (+ Trans Pecos)

Lespedeza virginica E N Amer

Leucaena retusa Chih

Lotus unifoliolatus N Amer

Lupinus texensis Comanch

Mimosa aculeaticarpa Madr wide

Mimosa borealis SWC USA

Mimosa latidens SE TX (W Gulf)

Mimosa malacophylla Tam

Mimosa nuttallii Prair

Mimosa roemeriana Comanch

Mimosa strigillosa Amphitrop (Gulf-S S Amer)

Mimosa texana Chih

Neptunia lutea SE Prair (+ Gulf)

Neptuniapubescens Mesoam (+ Gulf)

Oxytropis lambertii Prair (+ Roc Mt)

Parkinsonia texana S TX (Tam?)

Pediomelum cuspidatum Prair

Pediomelum cyphocalyx Comanch (NC TX endemic)

Pediomelum humile ? (Val Verde c)

Pediomelum hypogaeum Prair (+ Roc Mt)

Pediomelum latestipulatum Comanch (CN TX)

Pediomelum linearifolium W Prair (+ Roc Mt)

Pediomelum rhombifolium SC US

Pomaria brachycarpa S TX endemic (SC)

Pomaria jamesii S WC USA

Prosopis glandulosa W N Amer (mostly S)

Psoralidium tenuiflorum WC N Amer (Prair-WC)

Rhynchosia minima Trop/Subtr

Rhynchosia senna Amphitrop (Madr-Temp S Amer)

Senna bauhinioides Madr

Senna lindheimeriana Chih-Tam

Senna pumilio Chih

Senna roemeriana SWC USA (NM & WTX)

Senna wislizeni Madr (N & E, disj)

Sesbania drummondii Gulf-Tarn

Sesbania herbacea Mesoam

Sesbania vesicaria Gulf

Sophora secundiflora Madr (E)

Strophostyles helvola E N Amer

Strophostyles leiosperma E N Amer (+ W Madr)

Styphnolobium affine Comanch

Tephrosia lindheimeri S TX endemic

Tephrosia potosina Tam

Vida ludoviciana N Amer (S)

Zornia bracteata Gulf

FAGACEAE 1/17

Quercus buckleyi Comanch

Quercus falcata E N Amer

Quercus fusiformis SC N Amer

Quercus gravesii Chih

Quercus grisea Son wide

Quercus havardii SW Prair

Quercus laceyi STX endemic

Quercus macrocarpa E N Amer (NE-Prair)

Quercus marilandica E N Amer

Quercus mohriana SWC USA

Quercus muehlenbergii E N Amer

Quercuspolymorpha E Madr (+ Mesoam)

Quercuspungens Madr (N & E)

Quercus shumardii E N Amer

Quercus sinuata E N Amer disj

Quercus stellata E N Amer

Quercus vaseyana Chih

FOUQUIERIACEAE 1/1

Fouquieria splendens Son (Son-Chih)

FUMARIACEAE1/2

Corydalis curvisiliqua Prair (WC)

Corydalis micrantha Prair (E + SE N Amer)

GARRYACEAE 1/1

Garrya ovata S & E Madr

GENTIANACEAE 3/5

Centaurium beyrichii Comanch

Centaurium calycosum E Madr

Centaurium texense Comanch? SE Prair

Eustoma exaltatum Mesoam (+ N Amer)

Sabatia cam pest ris E Prair

GERANIACEAE 2/3

Erodium texanum SWC USA

Geranium carolinianum Amer

Geranium texanum SC US (CE TX-SW LA)

GROSSULARIACEAE 1/1

Ribes aureum N Amer (+ W Europe)

HAMAMELIDACEAE 1/1

Hamamelis virginiana E N Amer

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Mentzelia multiflora SW N Amer

Mentzelia nuda W Prair

Mentzelia oligosperma Prair

Mentzelia reverchonii SC USA

LOGANIACEAE 2/3

Mitreolapetiolata Trop/Subtr (Mesoam)

Spigelia hedyotidea Mesoam (+ SCTX)

Spigelia texana SE TX

LYTHRACEAE 4/6

Ammannia auriculata Trop/Subtr-Warm

Ammannia coccinea Mesoam (+ N Amer)

Lythrum californicum W N Amer

Lythrum ovalifolium Edw Plat

Nesaea longipes N Chih

Rotala ramosior Amer

MALPIGHIACEAE 2/2

Aspicarpa hyssop!folia S TX-Chih

Galphimia angustifolia E & W Madr

MALVACEAE 12/24

Abutilon fruticosum Trop

Abutilon parvulum Son

Abutilon wrightii Chih

Allowissadula holosericea Chih-Tam

Callirhoe involucrata Prair

Callirhoe leiocarpa Prair (S, Comanch?)

Callirhoepedata Comanch

Herissantia crispa Trop/Subtr

Hibiscus coulteri Son wide

Hibiscus laevis E N Amer

Hibiscus martian us Madr (E + Tam)

Hibiscus moscheutos E N Amer

Malvaviscus arboreus Trop/Subtr (Amer)

Meximalva filipes Tam

Modiola caroliniana Amer Trop/Subtr

Pavonia lasiopetala Chih (E)-Tam

Rhynchosida physocalyx Amer Trop/Subtr

Si da lindheimeri SE TX

Sida longipes Chih-Tam

Sida spinosa Amer

Sphaeralcea angustifolia Madr

Sphaeralcea coccinea W N Amer

Sphaeralcea hastulata SW N Amer

Sphaeralcea incana SW N Amer

MARSILEACEAE 2/3

Marsilea macropoda SE N Amer disj

Marsilea vestita N Amer (+ Peru)

Pilularia americana Amer (Warm)

MELANTHIACEAE 2/2

Schoenocaulon texanum Chih

Toxicoscordion nuttallii E Prair

MENISPERMACEAE 2/2

Cocculus carolinus E N Amer

Menispermum canadense E N Amer (NE/EC)

MORACEAE 2/3

Madura pomifera E N Amer

Morus microphylla SW N Amer (+ SC)

Morus rubra E N Amer

MYRICACEAE 1/1

Morelia cerifera Mesoam (+ Gulf)

NAJADACEAE 1/2

Najas guadalupensis Amer

Najas marina Pol ichor

NELUMBONACEAE 1/1

Nelumbo lutea E N Amer (+ Mesoam)

NYCTAGINACEAE 6/17

Acleisanthes anisophylla Tam

Acleisanthes crassifolia S TX-Coah

Acleisanthes longiflora Son wide

Acleisanthes obtusa E Madr (+ Tam)

Acleisanthes wrightii S TX (SWTX endemic)

Allionia incarnata Amer (mostly SW N & C Amer)

Boerhavia coccinea Trop/Subtr

Boerhavia diffusa Trop/Subtr

Boerhavia erecta Amer Trop/Subtr

Boerhavia linearifolia Chih (+ N NM)

Cyphomeris crassifolia E Chih

Cyphomeris gypsophiloides Chih

Mirabilis albida N Amer

Mirabilis linearis W N Amer (+ Prair)

Mirabilis nyctaginea N Amer-Mesoam

Mirabilis texensis SW TX

Nyctaginia capitata Chih

NYMPH AEACEAE 1/2

Nuphar lutea Holarctical

Nymphaea elegans Mesoam (+ Gulf)

OLEACEAE 3/12

Forestiera angustifolia Chih-Tam

Forestiera pubescens SWC N Amer

Forestiera reticulata E Madr

Fraxinus americana E N Amer

Fraxinus berlandieriana Madr

Fraxinus cuspidata Madr (N & E)

Fraxinus greggii Madr (E Mont)

Fraxinus pennsylvanica E N Amer

Fraxinus texensis Comanch

Menodora heterophylla SE TX

Menodora longiflora SC N Amer

Menodora scabra Madr

ONAGRACEAE 3/24

Calylophus berlandieri SC USA

Calylophusserrulatus Prair

Ludwigia glandulosa E N Amer

Ludwigia octovalvis Polichor

Ludwigiapalustris Polichor

Ludwigia peploides Amer

Ludwigia repens E N Amer

Oenothera brachycarpa Madr

Oenothera calcicola E Madr

Oenothera hartwegii SWC USA

Oenothera grand is Prair

Oenothera jamesii SC N Amer

Oenothera kunthiana Mesoam (+ Madr)

Oenothera laciniata Amer, adv els

Oenothera lavandulifolius W N Amer

Oenothera macrocarpa Prair

Oenothera rhombipetala Prair

Oenothera rosea Trop/Subtr (mostly Amer)

Oenothera speciosa N Amer (+ Mesoam)

Oenotherasuffulta N Chih

Oenothera suffrutescens W N Amer

Oenothera triloba Prair

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Bouteloua ramosa N Chih

Bouteloua repens American Trop/Subtr

Bouteloua rigidiseto S Prair

Bouteloua trifida N Madr

Bouteloua uniflora Chih

Bromus pubescens N Amer (mostly E)

Bromus texensis SC TX endemic

Cenchrus echinatus Trop/Subtr

Cenchrus longispinus N Amer

Cenchrus myosuroides Amer Trop/Subtr

Cenchrus spinifex ?

Chasmanthium latifolium E N Amer

Chloris andropogonoides SC TX

Chloris barbata Trop/Subtr

Chloris cucullata SC N Amer

Chloris verticillata Prair (+ W N Amer)

Coelorachis cylindrica SE N Amer

Danthonia spicata N Amer

Dasyochloa pulchella Madr wide

Dichanthelium aciculare SE N Amer (+ Carib)

Dichanthelium acuminatum N Amer (+ Mesoam)

Dichanthelium dichotomum E N Amer (+ C Amer)

Dichanthelium linearifolium E N Amer (NE + Prair)

Dichanthelium oligosanthes N Amer

Dichanthelium pedicel latum Mesoam

Dichanthelium sphaerocarpon E N Amer (+ C Amer)

Digitaria californica Amphitrop

Digitaria cognata E N Amer

Digitaria hitchcockii S TX-Coah

Digitaria insularis Amer Trop/Subtr

Digitaria patens S TX (SW-SC) endemic

Digitaria pubiflora SWC N Amer

Disakisperma dubium SWC N Amer

Distichlis spicata Amer

Echinochloa crus-pavonis Trop/Subtr/Warm

Echinochloa muricata N Amer

Echinochloa waited E N Amer (+ C Amer)

Elionurus tripsacoides Amer Trop/Subtr

Elymus canadensis N Amer

Elymus elymoides W N Amer

Elymus glabriftorus E N Amer

Elymus interruptus ?

Elymus macgregorii E N Amer

Elymus texensis Edw Plat

Elymus villosus E N Amer (mostly NE)

Elymus virginicus E N Amer

Enneapogon desvauxii Polichor

Eragrostis curtipedicellata SC USA

Eragrostis hirsuta E N Amer

Eragrostis intermedia SWC N Amer

Eragrostis palmeri Chih

Eragrostis pectinacea Amer

Eragrostis secundiflora Amer Trop/Subtr/Warm

Eragrostis sessilispica S Prair

Eragrostis spectabilis N Amer

Eragrostis trichodes N Amer (mostly Prair)

Eriochloa contracta N Amer

Eriochloa sericea Prair (S)

Erioneuron pilosum WC N Amer

Festuca versuta ?

Glyceria striata N Amer

Hesperostipa comata W N Amer

Hesperostipa neomexicana SW USA

Heteropogon contortus Trop/Subtr

Hilaria belangeri Chih

Hopia obtusa SC N Amer (S USA-Madr)

Hordeum jubatum Holarctical

Hordeum pusillum N Amer

Leersia oryzoides Holarctical

Leersia virginica E N Amer (+ Brazil)

Leptochloa fusca Polichor

Leptochloapanicea Trop/Subtr (Amer-Asian)

Limnodea arkansana S Prair (+ Gulf)

Melica montezuma Chih

Melica mutica E N Amer

Melica nitens E N Amer (Appalachian)

Muhlenbergia arenacea Son-Chih

Muhlenbergia arenicola SWC USA (+ Chih)

Muhlenbergia asperifolia Amer (WTemp)

Muhlenbergia dubia Madr Mont

Muhlenbergia xinvoluta SC TX (SWC)

Muhlenbergia lindheimeri Edw Plat-Coah

Muhlenbergia ported Madr (N & WC)

Muhlenbergia pungens SW N Amer (+ W Prair)

Muhlenbergia reverchonii Comanch

Muhlenbergia rigida W Amer Mont

Muhlenbergia schreberi E N Amer (+ S Amer)

Muhlenbergia setifolia Chih

Muhlenbergia sobolifera E Prair (+ NE N Amer)

Muhlenbergia spiciformis Carib

Muhlenbergia tenuifolia Madr

Muhlenbergia utilis Madr

Nassella leucotricha SC N Amer

Neeragrostis reptans Prair (E)

Panicum anceps E N Amer

Panicum brachyanthum SE Prair (+ E Gulf)

Panicum bulbosum Amer

Panicum capillare N Amer (+ Temp S Amer)

Panicum dichotomiflorum Amer

Panicum diffusum Gulf (ETX-Gulf-Carib)

Panicum hallii SWC USA

Panicum hirticaule Amer

Panicum rigidulum E N Amer (+ C Amer-Carib)

Panicum virgatum N Amer

Pappophorum bicolor Madr (E)

Pappophorum vaginatum Amphitrop

Pascopyrum smithii W USA

Paspalidium geminatum Trop/Subtr

Paspalum botterii Gulf (+ Meso Amer)

Paspalum distichum Trop/Subtr/Warm

Paspalum plicatulum Amer Trop/Subtr

Paspalum pubiflorum E N Amer (+ Mesoam)

Paspalum setaceum N Amer (+ C Amer)

Phalaris angusta Amer (coastal)

Phalaris caroliniana N Amer (coastal)

Phragmites australis Polichor

Piptochaetium avenaceum SE N Amer

Pleuraphis jamesii SW USA

Pleuraphis mutica Son wide

Poa arachnifera S Prair

Poa bigelovii N Madr (SWC USA)

Polypogon interruptus Amer (W Amer)

Schedonnarduspaniculatus Prair

Schizachyrium scoparium N Amer

Setaria grisebachii Madr-Mesoam

Setaria leucopila SWC N Amer (+ E Madr)

Setaria macrostachya Amer Trop/Subtr

Setaria magna Carib (Amer Subtr)

Saghatelyan, Phytogeographical analysis of the South-Central Texas flora

289

RANUNCULACEAE 5/11

Anemone berlandieri Amer (SE N Amer/ S S Amer)

Anemone edwordsiono Edw Plat

Anemone tuberosa N Madr

Aquilegio canadensis E N Amer (N)

Clematis drummondii Son

Clematis pitched E N Amer

Clematis texensis Edw Plat endemic

Delphinium carolinianum Prair (+ SE N Amer)

Delphinium madrense Chih-Tam

Delphinium wootonii SW N Amer

Ranunculus macranthus SEC TX

RESEDACEAE 1/1

Oligomeris linifolia Madro (N)-Tethyan

RHAMNACEAE7/13

Berchemia scandens SE N Amer (+ C Amer)

Ceanothus amedcanus E N Amer

Ceanothus herbaceus Prair

Colubdna texensis Chih

Condalia edcoides Son Mont

Condalia hooked Tam

Condalia spathulata Tam

Condalia viddis Chih (+ Sonora)

Condalia wamockii Son

Frangula betulifolia Madr wide Mont

Frangula caroliniana SE N Amer

Karwinskia humboldtiana Carib-Son

Ziziphus obtusifolia Madr

ROSACEAE 10/26

Cercocarpus montanus W N Amer

Crataegus crus-galli_E N Amer

Crataegus greggiana Tam

Crataegus reverchonii Comanch ?

Crataegus tracyi S TX-Coah

Crataegus turnerorum S TX endemic

Crataegus uvaldensis Edw Plat endemic

Crataegus viridis SE N Amer

Fallugiaparadoxa Madr (NC)

Geum canadense E N Amer

Malus ioensis E Prair (mostly NE)

Petrophyton caespitosum W N Amer

Potentilla rivalis W N Amer (+ WC)

Prunus angustifolia SE N Amer

PrunushavardiiSIX (Trans-Pecos) endemic Mont

Prunus mexicana E Prair (+ E N Amer)

Prunus minutiflora Edw Plat endemic

Prunus munsoniana E N Amer (+ E Prair)

Prunus rivularis Comanch

Prunus serotina Amer (+ Europe)

Prunus texana SC TX endemic

Rosa Carolina E N Amer

Rosa foliolosa Comanch ?

Rubus aboriginum ?

Rubus oklahomus Comanch ?

Rubus trivialis SE N Amer

RUBIACEAE 6/17

Cephalanthus occidentalis N Amer (not C)

Diodia teres Amer

Diodia virginiana E N Amer

Galium aparine Holarctical

Galium circaezans E N Amer

Galium correllii Chih

Galium microphyllum Madr

Galium proliferum N Madr

Galium texense Comanch?

Galium uncinulatum Mesoam

Galium virgatum SC USA (Comanch)

Floustonia acerosa SW N Amer

Floustonia humifusa SC USA

Floustoniapusilla E Prair (+ E N Amer)

Richardia tricocca Madr (mostly E)

Stenaria nigricans E N Amer

Stenaria rupicola N Chih (Trans-Pecos?)

RUTACEAE 3/3

Ptelea trifoliata N Amer (+ Europe)

Thamnosma texana N Son

Zanthoxylum hirsutum Comanch

SALICACEAE 1/1

Populus deltoides Holarctical

SAPINDACEAE 4/4

Cardiospermum halicacabum Trop/Subtr

Sapindus saponaria Trop/Subtr

Serjania brachycarpa Madr (S & E)

Ungnadia speciosa Chih wide?

SAPOTACEAE 1/1

Sideroxylon lanuginosum SE N Amer

SAXIFRAGACEAE 1/1

Lepuropetalon spathulatum SE N (+ S Amer)

SCROPHULARIACEAE 1/2

Leucophyllum frutescens Chih-Tam

Leucophyllum minus Chih

SELAGINELLACEAE 1/8

Selaginella apoda SE N Amer

Selaginella arenicola SE N Amer

Selaginella arizonica Son

Selaginella lepidophylla Madr (E & S)

Selaginella mutica SWC N Amer (Roc Mt)

Selaginella peruviana W Amer

Selaginella underwoodii\N N Amer

Selaginella wrightii Chih-Tam

SIMAROUBACEAE 1/1

Castela erecta C Amer (Carib)

SMILACACEAE 1/1

Smilax bona-nox SE N Amer (+ E Madr)

SOLANACEAE 10/28

Bouchetia erecta Madr (E & S)

Chamaesaracha coniodes SC N Amer

Chamaesaracha coronopus SWN Amer

Chamaesaracha edwardsiana ?

Chamaesaracha pallida Chih

Chamaesaracha sordida SWC N Amer

Chamaesaracha villosa Chih

Datura wrightii N Amer (mostly W)

Flunzikeria texana Chih-Tam (E)

Lycium berlandieri Son-Chih

Margaranthus solanaceus Madr

Nicotiana obtusifolia Madr (N + W)

Nicotiana repanda Chih ?

Physalis angulata Amer Trop/Subtr

Physalis cinerascens SC N Amer (+ Mesoam)

Physalis hederifolia SWC N Amer

Physalis longifolia N Amer

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Journal of the Botanical Research Institute of Texas 9(1)

Physalis mollis SE N Amer

Physalis pubescens Amer (Polichor)

Physolis virginiono N Amer (E & C)

Quincula lobata SWC N Amer

Solarium citrullifolium Chih?

Solarium dimidiatum S Prair

Solarium elaeagnifolium Amer

Solarium ptycanthum N Amer (mostly E)

Solarium rostratum Prair (+ W N Amer)

Solarium tenuipes Chih

Solarium triquetrum Chih-Tam

STERCULIACEAE 3/3

Ayeniapilosa Chih-Tam

Hermannia texana Jam (+ SC TX)

Melochia pyramidata Amer Trop/Subtr

STYRACACEAE 1/1

Styraxplatanifolius S TX-Coah

THELYPTERIDACEAE 1/1

Thelypteris ovata Carib (Gulf-Mesoam-Carib)

THEMIDACEAE 1/1

Androstephium caeruleum Prair

TILIACEAE 1/1

Tilia americana E N Amer

TYPHACEAE 1/2

Typha domingensis Trop/Subtr

Typha latifolia Pol ichor

ULMACEAE 2/7

Celtis ehrenbergiana Amer Trop/Subtr

Celtis laevigata E N Amer

Celtis lindheimeri Edw Plat endemic

Celtis reticulata W N Amer

Ulmus americana E N Amer

Ulmus crassifolia SC USA

Ulmus rubra E N Amer

URTICACEAE 3/3

Boehmeria cylindrica Amer

Parietaria pensylvanica N Amer

Urtica chamaedryoides Amer

VALERIANACEAE 1/5

Valerianella amarella Comanch

Valerianella florifera SE TX endemic

Valerianella radiata SE N Amer

Valerianella stenocarpa Edw Plat endemic

Valerianella texana Edw Plat endemic

VERBENACEAE 6/13

Aloysia gratissima Amphitrop

Aloysia wrightii Son

Bouchea linifolia Chih (S TX-Coah)

Callicarpa americana SE N Amer

Glandularia bipinnatifida Prair

Glandularia pumila SC USA

Glandularia quandrangulata Chih-Tam

Glandularia tumidula S Edw Plat

Lantana achyranthifolia Mesoam-S Amer

Phyla cuneifolia Prair (WC USA)

Phyla fruticosa Amer Trop/Subtr

Phyla lanceolata Amer Trop/Subtr

Phyla nodiflora Amer Trop/Subtr

VIOLACEAE 2/3

Hybanthus verticillatus SWC USA (+ W Prair)

Viola langloisii ?

Viola sororia E N Amer

VISCACEAE 1/2

Phoradendron hawksworthii N Chih

Phoradendron leucarpum ?

VITACEAE 4/11

Ampelopsis arborea SE N Amer (+ Carib)

Ampelopsis cordata E N Amer

Cissus trifoliata Mesoam (+ S N Amer)

Parthenocissus heptaphylla Edw Plat

Parthenocissus quinquefolia E N Amer

Parthenocissus vitacea N Amer (W & N)

Vitis cinerea E N Amer

Vitis monticola Edw Plat

Vitis mustangensis SC N Amer

Vitis palmata EC N Amer

Vitis rupestris E Prair (E Prair-EC N Amer)

ZANNICHELLIACEAE 1/1

Zannichellia palustris Pol ichor

ZYGOPHYLLACEAE 3/5

Guaiacum angustifolium Chih-Tam

Kallstroemia hirsutissima Son-Chih

Kallstroemiaparviflora Madr (Amphitrop)

Kallstroemia perennans SWTX endemic

Larrea tridentata Madr wide (W)

ACKNOWLEDGMENTS

I am deeply thankful to an anonymous referee for very thorough reviews of the manuscript and valuable com¬

ments; M.H. MacRoberts and B. Lipscomb for valuable suggestions and helpful reviews. I also thank A. Kara-

begov for the help with wording and K. Merrit for technical help. I would like to acknowledge funding for this

work provided by 2012 grant from Sam Taylor Foundation, 2013 KIVA Award, and McMurry University during

my sabbatical in 2013.

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