VOLUME 1, NUMBER 2, 12 DECEMBER 2007

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JOURNAL OF THE \ i

BOTANICAL

RESEARCH

TEXAS

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JOURNAL OF THE BOTANICAL RESEARCH INSTITUTE OF TEXAS

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

nd iaht

nvri

FED

1993—BRIT becomes publisher/copyright holder.

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

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BIBLIOGRAPHICAL

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Journal of the Botanical Research Institute of Texas is

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

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J. Bot. Res. Inst. Texas

VOLUME 1 NUMBER 2

ISSN 1934-5259

12 DECEMBER 2007

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Barney L Lipscomb

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TABLE OF CONTENTS

SYSTEMATICS

Revision of Lobelia sect. Galeatella (Campanulaceae: Lobelioideae)

Thomas G. LAMMERS

A new species and two new combinations in Potentilla sect. Niveae (Rosaceae)

David F. MURRAY AND REIDAR ELVEN

A new species of Alocasia (Araceae: Colocasieae) from Panay Island, Philippines

MELANIE P. MEDECILO, GEORGE C. YAO, AND DOMINGO A. MADULID

Folia taxonomica 2. New species of Passiflora subgenus Passiflora

(Passifloraceae) from the Guianas

CHRISTIAN FEUILLET

Contributions to the genus Festuca (Poaceae: Poeae) in Mexico and a key to

the Central American species

STEPHEN J. DARBYSHIRE

Aphanelytrum peruvianum (Poaceae: Poinae): a new species from Peru

IsIDORO SÁNCHEZ VEGA, PAUL M. PETERSON, ROBERT J. SORENG, AND SIMON L44GAARD

A new species of Chusquea sect. Verticillatae (Poaceae: Bambusoideae)

from Ecuador

LYNN G. CLARK, CHRISTOPHER D. TYRRELL, Jimmy K. TRIPLETT, AND AMANDA E. FISHER

Estudios en las Apocynaceae Neotropicales XXX: tres nuevas especies andinas

de Mandevilla (Apocynoideae: Mesechiteae)

J. FRANCISCO MORALES

Estudios en las Apocynaceae Neotropicales XXXI: el complejo de Mandevilla

hirsuta y cuatro nuevas especies

J. FRANCISCO MORALES

Boerhavia triquetra var. intermedia (Nyctaginaceae); a new combination and

varietal status for the widespread southwestern North American B. intermedia

RICHARD SPELLENBERG

e be se a 1 fel 1 > 4 1 yperaceae)

L

Davip J. ROSEN, STEPHAN L. HATCH, AND RICHARD CARTER

Fourteen new combinations in Sedum (Crassulaceae)

HIDEAKI OHBA

A new gypsophilous species of Erigeron (Asteraceae: Astereae) from northeastern

Mexico

Guy L. Nesom

Folia taxonomica 3. Passiflora davidii (Passifloraceae), a new species in sub-

genus Passiflora and a key to the sections of supersection Stipulata

CHRISTIAN FEUILLET

Four new combinations in Mosiera (Myrtaceae) from the Caribbean

ANDREW SALYWON

New combinations in Philadelphus (Philadelphaceae)

JAMES HENRICKSON

Five new varieties of Agapetes (Ericaceae) from India

D. BANIK AND M. SANJAPPA

789

815

819

Phylogenetic analyses of the genus Baptistonia (Orchidaceae: Oncidiinae)

sensu lato based on morphological characters

Guy R. CHIRON

Muhlenbergiinae (Poaceae: Chloridoideae: Cynodonteae): from northeastern

México

PauL M. PETERSON, Jesus VALDÉS-REYNA, AND YOLANDA HERRERA ARRIETA

A second species of Cyclopogon s.s. (Orchidaceae: Spiranthinae)

Eric A. CHRISTENSON

Lectotypification of Maxillaria heteroclita (Orchidaceae)

Eric A. CHRISTENSON

Miscellaneous typifications, new combinations and one new variety in North

American Crataegus (Rosaceae)

J.B. PHIPPS

Kruschke names in North American Crataegus (Rosaceae)

J.B. PHIPPS

New combinations in Dudleya (Crassulaceae) and Ribes (Grossulariaceae)

Nancy R. Morin

New combinations in Limnanthes (Limnanthaceae)

Nancy R. Morin

New combinations in Micranthes (a segregate of Saxifraga, Saxifragaceae)

in North America

Luc BROUILLET AND RICHARD GORNALL

es

New | | new iesin the genus Spermacoce (Rubiaceae)

for the Flora of Goiás and Tocantins (Brazil) and the Flora of the Guianas

PIERO G. DELPRETE

Hawthorns (Crataegus: Rosaceae) of the Cypress Hills, Alberta and Saskatchewan

J.B. Puipps AND R.J. O'KENNON

Thomas Walter Typification Project, IV: neotypes and epitypes for 43 Walter

names, of genera A through C

DanteL B. WARD

ANATOMY AND MORPHOLOGY

Cotyledon areoles in the Fabaceae subtribe Clitoriinae

JAMES A. LACKEY

FLORISTICS, ECOLOGY, AND CONSERVATION

Rediscovery of Eleocharis kleinii (Cyperaceae), an overlooked species from

the highlands of South Brazil

R. Trevisan, D.J. ROSEN, AND I.I. BOLDRINI

Distribution of Gamochaeta (Asteraceae: Gnaphalieae) in Texas, Oklahoma,

Arkansas, and Louisiana

Guy L. Nesom

Flow diagrams for plant succession in the Middle Tennessee cedar glades

J.M. BASKIN, E. QUARTERMAN, AND C.C. BASKIN

913

1005

1011

1015

1017

1019

1023

1031

1091

1101

1119

1125

1131

Observation on vegetation changes in Cajun Prairie, a coastal prairie flora in

southwest Louisiana

CHARLES ÁLLEN AND SARA THAMES

Phytogeography of the Big Thicket, East Texas

MicHAEL H. MACROBERTS AND BARBARA R. MACROBERTS

A floristic survey of National Park Service areas of Timucuan Ecological and

Historic Preserve (including Fort Caroline National Memorial), Duval County,

Florida

Wenny B. ZOMLEFER, DaviD E. GIANNASI, AND WALTER S. JUDD

The vascular flora of the Chunky River (Mississippi)

Lucas C. MAJURE

New plant discoveries for Sonoran Islands, Gulf of California, Mexico

BENJAMIN T. WILDER, RICHARD S. FELGER, HUMBERTO ROMERO-MORALES, AND ADRIAN QUIJADA-MASCAREÑAS

Checklist of the vascular plants of Washington County, Pennsylvania

CYNTHIA M. MORTON AND LOREE SPEEDY

Saxifraga texana (Saxifragaceae) new to Louisiana

CHRISTOPHER S. REID, PATRICIA L. FAULKNER, BARBARA R. MACROBERTS, AND MICHAEL H. MACROBERTS

Vicia lathyroides (Fabaceae): new to the flora of Texas

SONNIA HILL, RUTH LOPER, JASON R. SINGHURST, AND WALTER C. HOLMES

Notes on the flora of Texas with additions and other significant records. III

Larry E. Brown, Eric L. Kern, Davin J. ROSEN, AND JOE LIGGIO

Notes on Texas orchids (Orchidaceae)

PauL MARTIN BROWN

Galium anglicum (Rubiaceae) new for Texas and notes on the taxonomy of

the G. parisiense/divaricatum complex

BARNEY L. LIPSCOMB AND Guy L. Nesom

IN MEMORIAM

James Hinton (1915-2006)

GEORGE S. HINTON AND BILLIE L. TURNER

Book Reviews AND Notices 826, 846, 852, 858, 870, 902, 932, 1018, 1148, 1156, 1228, 1250

Announcements 1281

1141

1149

1157

1179

1203

1229

1251

1253

1255

1265

1269

1277

INDEX to new names and new combinations in J. Bot. Res. Inst. Texas 1(2), 2007

Agapetes acuminata var. tipiensis Banik € Sanjappa, var. nov.—903

Agapetes flava var. nagensis Banik € Sanjappa, var. nov.—905

Agapetes megacarpa var. lohitensis Banik € Sanjappa, var. nov.—905

Agapetes odontocera var. mizoramensis Banik € Sanjappa, var. nov.—908

Agapetes salicifolia var. glanduliflora Banik € Sanjappa, var. nov.—910

Alocasia nycteris Medecilo, Yao & Madulid, sp. nov.—817

Aphanelytrum peruvianum Sánchez Vega, PM. Peterson, Soreng & Leegaard, sp. nov.—842

Boerhavia triquetra S. Wats. var. intermedia (M.E. Jones) Spellenb., comb. nov.—874

Chusquea annagardneriae L.G. Clark, C.D. Tyrrell, Triplett & A.E. Fisher, sp. nov.—847

Crataegus aquacervensis J.B. Phipps & O'Kennon, sp. nov.—1061

Crataegus brazoria var. viburnifolia (Sarg.) J.B. Phipps, comb. nov.—1007

Crataegus chrysocarpa var. blanchardii (Sarg.) J.B. Phipps, comb. et stat. nov.—1009

Crataegus chrysocarpa var. praecox (Sarg.) J.B. Phipps, comb. et stat. nov.—1009

Crataegus chrysocarpa var. vigintistamina J.B. Phipps, var. nov.—1008

Crataegus coccinea var. pringlei (Sarg.) J.A. Macklin & J.B. Phipps, comb. et stat. nov.—1005

Crataegus cupressocollina J.B. Phipps & O'Kennon, sp. nov.—1056

Crataegus series Cupressocollinae J.B. Phipps & O'Kennon, ser. nov.—1056

Crataegus series Montaninsulae J.B. Phipps & O'Kennon, ser. nov.—1065

Crataegus pruinosa var. dissona (Sarg.) J.B. Phipps, comb. et stat. nov.—1007

Crataegus purpurella J.B. Phipps & O'Kennon, sp. nov.—1066

Crataegus reverchonii var. mohrii (Beadle) J.B. Phipps, comb. et stat. nov.—1006

Crataegus rivuloadamensis J.B. Phipps & O'Kennon, sp. nov.—1065

Crataegus rivulopugnensis J.B. Phipps & O'Kennon, sp. nov.—1070

Crataegus rubribracteolata J.B. Phipps & O'Kennon, sp. nov.—1073

Crataegus sheila-phippsiae var. saskatchewanensis J.B. Phipps & O'Kennon, var. nov.—1079

Crataegus ursopedensis J.B.Phipps & O'Kennon, sp. nov.—1081

Crataegus viridis var. velutina (Sarg.) J.B. Phipps, comb. et stat. nov.—1009

Cyclopogon secundum Christenson, sp. nov.—1001

Dudleya abramsii subsp. costifolia (Bartel & Shevock) Moran, comb. nov.—1015

Dudleya abramsii subsp. setchellii (Jeps.) Moran, comb. nov.—1015

Eleocharis acutangula subsp. breviseta D.J. Rosen, subsp. nov.—885

Eleocharis acutangula subsp. neotropica D.J. Rosen, subsp. nov.—886

Erigeron heleniae Nesom, sp. nov.—891

Festuca xgonzalez-ledesmae S.J. Darbyshire, nothosp. nov.—832

Festuca longiligula S.J. Darbyshire, sp. nov.—827

Festuca xmiscella S.J. Darbyshire, nothosp. nov.—835

Limnanthes alba subsp. gracilis (Howell) Morin, comb. et stat. nov.—1017

Limnanthes alba subsp. parishii (Jeps.) Morin, comb. et stat. nov.—1017

Limnanthes douglasii subsp. striata (Jeps.) Morin, comb. et stat. nov.—1017

Lobelia gloria-montis subsp. longibracteata (Rock) Lammers, stat. nov.—806

Lobelia xkauaiensis (A. Gray) A. Heller, stat. nov.—801

Lobelia koolauensis (Hosaka & Fosberg) Lammers, comb. et stat. nov.—803

Lobelia wahiawa Lammers, sp. nov.—802

Mandevilla aequatorialis J.F Morales, sp. nov.—853

Mandevilla frigida J.E Morales, sp. nov.—855

Mandevilla horrida J.E Morales, sp. nov.—863

Mandevilla inexperata J.F Morales, sp. nov.—865

Mandevilla lojana J.E Morales, sp. nov.—855

Mandevilla megabracteata J.E Morales, sp. nov.—866

Mandevilla similaris J.E Morales, sp. nov.—867

Micranthes bryophora (A. Gray) Brouillet & Gornall, comb. nov.—1020

Micranthes calycina (Sternb.) Gornall & H. Ohba, comb. nov.—1020

Micranthes ferruginea (Graham) Brouillet & Gornall, comb. nov.—1020

Micranthes foliolosa (R. Br.) Gornall, comb. nov.—1020

Micranthes gormanii (Suksd.) Brouillet & Gornall, comb. nov.—1020

Micranthes idahoensis (Piper) Brouillet & Gornall, comb. nov.—1020

Micranthes mexicana (Engl. & Irmsch.) Brouillet & Gornall, comb. nov.—1020

Micranthes nelsoniana var. aestivalis (Fisch. & C.A. Mey.) Gornall & H. Ohba, comb. et stat. nov.—1020

Micranthes nelsoniana var. carlottae (Calder & Savile) Gornall & H. Ohba, comb. et stat. nov.—1020

Micranthes nelsoniana var. cascadensis (Calder & Savile) Gornall & H. Ohba, comb. et stat. nov.—1020

Micranthes nelsoniana var. insularis (Hultén) Gornall & H. Ohba, comb. et stat. nov.—1020

Micranthes nelsoniana var. pacifica (Hultén) Gornall & H. Ohba, comb. et stat. nov.—1020

Micranthes nudicaulis (D. Don) Gornall & H. Ohba, comb. nov.—1021

Micranthes petiolaris (Raf.) Brouillet & Gornall, comb. nov.—1021

Micranthes razshivinii (P. Yu. Zhmylev) Brouillet & Gornall, comb. nov.—1021

Micranthes tempestiva (Elvander & Denton) Brouillet & Gornall, comb. nov.—1021

Micranthes tischii (Skelly) Brouillet & Gornall, comb. nov.—1021

Micranthes tolmiei (Torrey & A. Gray) Brouillet & Gornall, comb. nov.—1022

Micranthes unalaschcensis (Sternb.) Gornall & H. Ohba, comb. nov.—1022

Mosiera androsiana (Urban) Salywon, comb. nov.—899

Mosiera cuspidata (Alain) Salywon, comb. nov.—899

Mosiera gracilipes (Alain) Salywon, comb. nov.—899

Mosiera xerophytica (Britton) Salywon, comb. nov.—900

Passiflora arta Feuillet, sp. nov.—819

Passiflora compar Feuillet, sp. nov.—821

Passiflora davidii Feuillet, sp. nov.—896

Philadelphus microphyllus var. argyrocalyx (Wooton) Henrickson, comb. & stat. nov.—901

Philadelphus microphyllus var. madrensis (Hemsley) Henrickson, comb. & stat. nov.—901

Philadelphus microphyllus var. stramineus (Rydb.) Henrickson, comb. & stat. nov.—901

Potentilla arenosa subsp. chamissonis (Hultén) Elven & D.E Murray, comb. nov.—813

Potentilla holmgrenii D.E Murray & Elven, sp. nov. —811

Potentilla hyparctica Malte subsp. elatior (Abrom.) Elven & D.E Murray, comb. nov.—813

Ribes oxyacanthoides var. cognatum (Greene) Morin, comb. et stat. nov.—1015

Sedum lanceolatum var. subalpinum (Fród.) H. Ohba, comb. & stat. nov.—890

Sedum laxum var. eastwoodiae (Britton) H. Ohba, comb. & stat. nov.—889

Sedum laxum var. flavidum (Denton) H. Ohba, comb. & stat. nov.—889

Sedum laxum var. heckneri (M. Peck) H. Ohba, comb. & stat. nov.—889

Sedum laxum var. latifolium (R.T. Clausen) H. Ohba, comb. & stat. nov.—890

Sedum obtusatum var. boreale (R.T. Clausen) H. Ohba, comb. & stat. nov.—889

Sedum obtusatum var. paradisum (Denton) H. Ohba, comb. & stat. nov.—889

Sedum obtusatum var. retusum (Rose) H. Ohba, comb. & stat. nov.—889

Sedum oreganum var. tenue (R.T. Clausen) H. Ohba, comb. & stat. nov.—890

Sedum radiatum var. ciliosum (Howell) H. Ohba, comb. & stat. nov.—889

Sedum radiatum var. depauperatum (R.T. Clausen) H. Ohba, comb. & stat. nov.—889

Sedum stenopetalum var. monanthum (Suksd.) H. Ohba, comb. & stat. nov.—890

Sedum wrightii var. densiflorum (R.T. Clausen) H. Ohba, comb. & stat. nov.—889

Sedum wrightii var. priscum (R.T. Clausen) H. Ohba, comb. & stat. nov.—889

Spermacoce burchellii (E.L. Cabral & Bacigalupo) Delprete, comb. nov.—1025

Spermacoce crispata (K. Schum.) Delprete, comb. nov.—1025

Spermacoce delicatula (E.L. Cabral) Delprete, comb. nov.—1025

Spermacoce dimorpha (J.H. Kirkbr.) Delprete, comb. nov.—1025

Spermacoce incognita (E.L. Cabral) Delprete, comb. nov.—1025

Spermacoce irwiniana (E.L. Cabral), Delprete, comb. nov.—1025

Spermacoce multiflora (DC.) Delprete, comb. nov.—1026

Spermacoce paraensis (E.L. Cabral & Bacigalupo) Delprete, comb. nov.—1027

Spermacoce pulchristipula (Bremek.) Delprete, comb. nov.—1027

Spermacoce semiamplexicale (E.L. Cabral) Delprete, comb. nov.—1028

Spermacoce spicata (Miq.) Delprete, comb. nov.—1028

Spermacoce tocantinsiana (E.L. Cabral € Bacigalupo) Delprete, comb. nov. 1028

REVISION OF LOBELIA SECT. GALEATEELA

(CAMPANULACEAE: LOBELIOIDEAE)

Thomas G. Lammers

dolia of Biology and eee

University of Wisconsin Osh

E Wisconsin 54901, po

ABSTRACT

Pew: 1 T: 1

Lobelia sect. Galeatella is a group of pachycaul treelets distinguished by a solitary pliestesial |

stock; a dense apical rosette of sessile coriaceous leaves leaving a helical pattern of raised ee cicatrices on the sida stem; a

on. or d MR short- pet teat f nie rnithophilous flowers on o let pedicels; calyx

hli hic lo! hed l d horizontally

about as n as a stout ‘ube to 3 times d the ventral connate for 7 their rM or more, e, forming a ilaa inrolled trifid lip

ee th rted, all five anthers with RUN HN Rd niu

E L

1 à 1 1 [xe 1! 11 2 s 21 0]

ue Hawaiian Islands, with two pn each on Kauai i. villosa D L. wahiawa, sp. nov.) and Oʻahu

(E; al and L. koolauensis, comb. et stat. nov.), an g L. gloria-montis) on Maui and Molokai, divided into two

subspecies: subsp. gloria-montis (West Maui and Moloka‘i) and bus longibracte ata stat. nov. (West Maui and East Maui). The two

species on Kauai hybridize, producing L. xkauaiensis, stat. n

RESUMEN

: “J T2211 : 1 A E 4 1; A qus 4 :

Lobelia sect. Galeatella es un grup paq I g p vástago solitari p pico qu

J > J a Tuo s pe 1 A 1 a s i pia N 3 1 1

o dE J É J E

T TT s o 1 5) 4 T J 1 J pa Ti A | 1 SES ii

; Į g Į

ebracteolados; lóbulos del cáliz cad el fruto joven; corola no azul con lóbulos lineares monomórficos, el par dorsal extendido o

recurvado pM dr tan d como el tubo rígido s tres veces más cd el n e en ’/, de su nud o

más, pu un labi j q p p

A 5 t ES RATE 1 1 Tae 1 JA] 1

bl 1 l l l d b la madure pol

y semillas oblongas alad testa débil te estriada. La ión ndémica de las Islas ns con dos especies en ds una

de las Kauai (L. villosa y L. wahiawa, sp. nov) y O'ahu (L. gaudichaudii y L. koolauensis, comb. et stat. nov.), y una sola especie (L.

gloria-montis) en Maui y Molokai, dividida en dos subespecies: subsp. gloria-montis (West Maui and HS y subsp. longibracteata,

stat. nov. (West Maui y East Maui). Las d pecies de Kauai hibridan produciendo L. xkauaiensis, stat. nov.

The finest of our lobelias is Lobelia gloria-montis, a truly royal and superb plant ...

To see this species growing is one of the delights of the botanist.—Rock (1919: 35, 117)

The Hawaiian archipelago is renowned among botanists for the many compositional peculiarities of its flora

(MacCaughey 1917; Hitchcock 1919; Carlquist 1974, 1980; Wagner et al. 1990). Among the most peculiar

is the fact that the family with the greatest number of native species is not Gramineae Juss. or Compositae

Giseke, but Campanulaceae Juss.; at no other spot on the globe does this family predominate statistically.

In a flora of not quite 1000 indigenous species (Wagner et al. 1990), 134 of them (all endemic) belong to

Campanulaceae (Lammers 20072). Furthermore, these 134 species represent nearly 696 of all species in

this cosmopolitan family, although the Hawaiian Islands only comprise 0.00496 of Earth's land surface.

Six genera are present, all members of subfamily Lobelioideae Burnett. Five are endemic and assigned to

the tribe Delisseeae C. Presl: Brighamia A. Gray, Clermontia Gaudich., Cyanea Gaudich., Delissea Gaudich.,

and Trematolobelia Zahlbr. ex Rock. The only one of the six not endemic to the archipelago is Lobelia L., a

member of tribe Lobelieae Rchb., which is virtually cosmopolitan in its distribution.

Lobelia is the largest of the 29 genera in the Lobelioideae, comprising 405 species of annual and peren-

nial herbs, shrubs, and trees (Lammers 20072). The genus is characterized by solitary or racemose flowers on

usually bibracteolate pedicels; corollas with the tube dorsally cleft to the base and the lobes monomorphic,

J. Bot. Res. Inst. Texas 1(2): 789 — 810. 2007

790 Journal of the Botanical R h Institute of Texas 1(2)

docs

or dimorphic and the ventral larger; and apically capsules or rarely berries (Murata 1995; Lammers

2004a, 2007b).

Because of its large number of species, Lobelia has been divided into a number of sections and other

infrageneric taxa over the years. The first species described from the Hawaiian Islands, L. gaudichaudii, was

assigned to sect. Homochilus A. DC. by Candolle (1839). As additional Hawaiian species of Lobelia were

discovered during the Nineteenth Century, they all were assumed to be related to L. gaudichaudii. Bentham

(1876) removed these Hawaiian species from sect. Homochilus to sect. Rhynchopetalum (Fresen.) Benth. (cf.,

Lammers 20045); the latter subsequently was subsumed into sect. Tylomium (C. Presl) Benth. by Schonland

(1889).

In contrast to this single-taxon view of Hawaiian Lobelia, Rock (1919) believed that the Hawaiian species

of Lobelia actually formed two taxa that were not each other's closest relatives: (1) L. gaudichaudii and allied

species, characterized by a large white, yellow, red, or purple-striped corolla with stout tube and spreading

or horizontally recurved dorsal lobes, and all five anthers apically tufted; and (2) L. yuccoides Hillebr. and

allied species, characterized by a small blue or magenta corolla with slender tube and recoiled dorsal lobes,

and only the ventral pair of anthers apically tufted. This two-taxon hypothesis supported by Skottsberg

(1928) and Carlquist (1980), and formalized by Wimmer (1948), who erected two sections within Lobelia

subg. Tupa (G. Don) E. Wimm. to accomodate the Hawaiian endemics: sect. Galeatella E. Wimm. for L.

gaudichaudii and its allies, and sect. Revolutella E. Wimm. for L. yuccoides Hillebr. and its allies (Wimmer

1953, 1968). Although Mabberley (1974, 1975) suggested returning both groups to different parts of sect.

Rhynchopetalum, he did not effect these changes and Wimmer's arrangement is the classification in current

use (Lammers 1990, 20072, 2007b; Murata 1995).

Over the past 15 years, the summary of Hawaiian Campanulaceae that I prepared (Lammers 1990)

for the Manual of the flowering plants of Hawai'i (Wagner et al. 1990) has been expanded into a series of

monographs. Previous entries covered Brighamia (Lammers 1989), Clermontia (Lammers 1991), and Delissea

(Lammers 2005). The present effort covers Lobelia sect. Galeatella. This treatment will also form part of a

series of monographs of the sections of Lobelia subg. Tupa; previous entries in that series accounted for sect.

Tupa (G. Don) Benth. (Lammers 2000) and sect. Homochilus (Lammers 2004b).

TAXONOMIC HISTORY

As might be expected from their predominance in the Hawaiian flora, Lobelioideae were among the first

botanical specimens collected in the islands at the time of their discovery by Europeans in 1778 (St. John

19772, 197 Tb, 1979). However, it was another half century before botanists visiting the archipelago obtained

specimens referable to Lobelia sect. Galeatella. During an October 1836 visit to O'ahu by the French corvette

La Bonite (cf., Lasegue 1845; Kuykendall 1947), ship's naturalist Charles Gaudichaud-Beaupré reached the

summit of the Ko'olau Range above Honolulu harbor (St. John & Titcomb 1983). Here he collected a pachy-

caul species of Lobelia a meter tall with a thick pithy stem densely covered in transversely rhombic leaf scars,

a dense apical rosette of coriaceous leaves, a terminal raceme of large red flowers, and capsules filled with

flat winged seeds. Not long after, Swiss botanist Alphonse de Candolle saw the specimen in the herbarium

at Paris and named it in Gaudichaud's honor (Candolle 1839). Although Gaudichaud never published a

description of Lobelia gaudichaudii himself, the atlas of La Bonite's voyage (Gaudichaud s.d.) did include an

illustration, labeled with Candolle's binomial; the fascicle of the book containing this plate is believed to

have appeared in 1842 (Stafleu & Cowan 1976).

The United States Exploring Expedition (cf., Wilkes 1845; Jackson 1985) called at the Hawaiian Islands

during 1840-41 and collected specimens of Lobelia on Kauai that were very similar to L. gaudichaudii of

O'ahu. Harvard botanist Asa Gray (cf., Dupree 1959; St. John 1985) segregated these plants from the species

as L. gaudichaudii var. kauaiensis (Gray 1861; Mann 1867). However, Austrian botanist Heinrich Wawra, who

visited the archipelago aboard the frigate Donau during 1869—70, did not distinguish the specimens he had

gathered on O'ahu and Kauai, calling them all L. gaudichaudii (Wawra 1873). This single-taxon approach

Lammers, Revision of Lobelia sect. Galeatella 791

was also adopted by the islands' first resident botanist, Wilhelm Hillebrand (cf., Degener 1957; Lammers

1994), who broadened the ci iption of the species even further by including plants he had discovered

on West Maui (Hillebrand 1888).

American Amos Arthur Heller, who botanized on O'ahu and Kauai in 1895 (cf., Wagner & Shannon

1999), not only disagreed with this single-taxon view, but further disagreed with their original treatment as

conspecific varieties. He segregated the Kauai populations from L. gaudichaudii as L. kauaiensis, commenting

(Heller 1897: 911), “Had Dr. Gray seen [plants of O'ahu and Kauai] in the living state, he certainly would

not have considered the one a mere form of the other, worthy of varietal rank only."

The next botanist to deal with these plants was Austrian expatriate Joseph Rock, who resided in the

islands from 1907 to 1920 and again from 1953 until his death in 1962 (cf., Chock 1963; Sutton 1974). At

first, he accepted Heller's two-species treatment, but modified it slightly by describing three new varieties.

Thinking that the white-flowered plants on West Maui were the original nomenclatural type of L. gaudi-

chaudii, he segregated the red-flowered O'ahu populations as L. gaudichaudii var. coccinea (Rock 1917); only

later did he realize that this was an error and that the type of the species was actually a red-flowered plant

from O'ahu. Tall plants with very long floral bracts and calyx lobes that he discovered on West Maui were

distinguished as L. gaudichaudii var. longibracteata (Rock 1913a, 1913b), while plants with densely pubes-

cent inflorescences that he discovered on Kauai were segregated as L. kauaiensis var. villosa (Rock 1917).

However, in his subsequent monograph of the Hawaiian Lobelioideae (Rock 1919), he treated the plants of

each island as a discrete species: L. kauaiensis and its var. villosa on Kauai, L. gaudichaudii (with var. coccinea

in synonymy) on O'ahu; and the newly described L. gloria-montis and its var. longibracteata on West Maui.

When plants referable to sect. Galeatella were found on Molokai, Degener (19382) named them L. glo-

ria-montis var. molokaiensis, though a paucity of good material precluded an adequate description. Similarly,

when white-flowered plants with a branched inflorescence was discovered in the northwestern Ko'olau

Range, Fosberg and Hosaka (1938) described them as a variety of the O'ahu species, L. gaudichaudii var.

koolauensis.

The first dedicated taxonomic study of these plants was by University of Hawaii professor Harold St.

John (cf., Anonymous 1980, 1991) and his student Edward Hosaka. Their revision (St. John & Hosaka 1938)

resurrected Hillebrand's (1888) broad circumscription of L. gaudichaudii, including in it plants of O'ahu,

Kauai, and West Maui. Within this broad circumscription, they distinguished five geographic varieties:

var. gaudichaudii (as *var. typica" and including var. coccinea as a synonym), var. gloria-montis (with three

heterotypic formae), var. kauaiensis (with one heterotypic forma), var. koolauensis, and var. longibracteata; var.

molokaiensis was excluded for want of adequate material. However, not all Kauai plants were included in L.

gaudichaudii var. kauaiensis; those recognized previously as L. kauaiensis var. villosa were accorded specific

rank as L. villosa. Years later, a population discovered on East Maui was described (St. John & Medeiros

1987) as a sixth geographical variety, L. gaudichaudii var. albiflora.

When Viennese botanist Franz Elfried Wimmer (cf., Rechinger 1961; Degener & Degener 1962a)

prepared his monograph of Lobelia for Das Pflanzenreich (Wimmer 1953), his treatment of sect. Galeatella

was identical to that of St. John and Hosaka (1938). In subsequent years, however, his view of the group

changed. In a letter to Otto Degener dated 20 November 1958 (quoted by Degener & Degener 1974), he

indicated his intention to accord the group generic status under the name “Schiexelia,” and to recognize five

species therein. His posthumously published supplement to the Pflanzenreich monograph (Wimmer 1968)

did recognize five species: L. kauaiensis and L. villosa on Kauai, L. gaudichaudii on O'ahu; and L. gloria-montis

and L. longibracteata (newly elevated from varietal rank) on West Maui. However, no mention was made of

elevating the section to generic rank and it remained a section of Lobelia subg. Tupa. Degener and Degener

(1962b, 1974, 1983) carried through on the elevation to generic rank, though they disregarded Wimmer's

provisional name and took up the sectional epithet at the new rank. Their synopsis of the group (Degener &

Degener 1974) maintained Wimmer's (1968) five-species classification. My treatment in the Manual (Lam-

mers 1990) was essentially the same, except for including L. longibracteata within L. gloria-montis.

792 Journal of the Botanical R h Institute of Texas 1(2)

MATERIALS AND METHODS

Revision of the classification of Lobelia sect. Galeatella was based upon morphological data; definitions of

qualitative character states follow Harris and Harris (1994). These data were gathered from almost 300

specimens (including the types of all heterotypic names) deposited in 31 herbaria (see Acknowledgments

for a complete list).

The data obtained from these specimens were analyzed via traditional taxonomic methodology (Leen-

houts 1968; Qualls 1986; Vogel 1987; Maxted 1992; Watson 1997; Winston 1999) within a geographic

framework. Populations referable to sect. Galeatella are known from just six discrete areas on four islands:

(1) the Wahiawa drainage of south-central Kauai at 570-800 m; (2) the Alaka‘i Swamp of north-central

Kauai, from Pihea to Wai'aleale, at 1100-1590 m; (3) the Ko'olau Range of O'ahu, from Pu'u Ka'inapuaa to

Mt. Olympus, at 670-960 m; (4) the mountains of West Maui at 915-1760 m; (5) the northwestern slopes of

Haleakala on East Maui at 1280—1700 m; and (6) eastern Moloka'i from Papa‘ala to Kalua'aha at 1165-1350 m.

Because of the extreme precinctiveness of the Hawaiian flora (MacCaughey 1917; Hitchcock 1919; Carlquist

1980; Wagner et al. 1990), I hypothesized that each of these six areas would harbor a single taxon unique to

that locale, or at least to that island. Therefore, when inspecting the data, I asked two questions. First, were

all specimens from a given area relatively homogeneous, with most characters evincing a continuous pattern

of variation? If so, the hypothesis would be supported; if instead, several characters consistently showed

correlated gaps in their patterns of variation within an area, the hypothesis would be refuted. Second, are

the plants that evince a given correlated suite of morphological features restricted to a single area or island?

If so, the hypothesis would be supported; if not, the hypothesis would be refuted.

Once taxa had been discerned in this fashion, they were compared to nomenclatural type specimens

to determine the correct name under the International Code of Botanical Nomenclature (ICBN; McNeill et al.

2006). Decisions on rank for the taxa were made in light of the definitions of species and subspecies that I

have employed previously (Lammers 1991, 20073).

RESULTS AND DISCUSSION

In revising Lobelia sect. Galeatella, three aspects of its classification were evaluated in light of the assembled

data: (1) the circumscription of the group, (2) its position and rank, and (3) its division into component

taxa.

Circumscription.—The circumscription and characterization of Lobelia sect. Galeatella have been invari-

ant since its publication by Wimmer (1948). Overall similarity among populations is quite high; recall that

the entire section has been treated as just one or two species until recently, with most variation expressed

by distinguishing geographic varieties. Furthermore, much of this overall similarity involves characters that

are quite unusual among Lobelioideae.

Most species of Lobelia are hemicryptophytes or therophytes (Lammers 20072); woodiness is less com-

mon, confined to certain groups within subg. Tupa (Carlquist 1969; Murata 1995). Even among those species

that are woody, the growth form of sect. Galeatella is unusual. In sect. Tylomium (Wilbur 1991), sect. Tupa

(Lammers 2000), and sect. Homochilus (Lammers 2004b), the woody species are iteroparous (polycarpic)

shrubs with several leptocaul stems that persist, branch, and flower each growing season. In sect. Galeatella,

in contrast, the stem is pachycaul, unbranched, pliestesial (i.e., requiring several to many years of growth

before it achieves sufficient mass to flower), and hapaxanthic (i.e., dying after flowering). Although Truman

and Augspurger (1991) described these species as semelparous (monocarpic), this is incorrect. An indi-

vidual shoot does indeed die by the time the capsules dehisce and disperse ripe seed. However, a new shoot

emerges around the base of the plant (presumably from a root bud) to replace it. This sucker, which is far

more pubescent than a mature shoot (cf., the “juvenile plant” of Degener 19382), will eventually grow into

a new flowering shoot (Hillebrand 1888; pers. observ.). As such, the individual plant is in fact iteroparous

(polycarpic), conforming to the *Tomlinson" architectural model (cf., Hallé & Oldeman 1970; Hallé et al.

Lammers, Revision of Lobelia sect. Galeatella 793

Similarly, the winged seeds of sect. Galeatella (Fig. 1) are unusual in Lobelia, occurring elsewhere only

in some species of sect. Rhynchopetalum and sect. Trimeris (C. Presl) A. DC. [the earliest available name for

sect. Colensoa (Hook. f.) J. Murata] (Mabberley 1975; Murata 1995; Buss et al. 2001). The possession of apical

tufts of trichomes on all five anthers rather than just the ventral pair is otherwise unknown in subg. Tupa,

though it does characterize sect. Cryptostemon (E. Wimm.) J. Murata and sect. Delostemon (E. Wimm.) J.

Murata in herbaceous subg. Lobelia (Murata 1995).

In summary, the assemblage of morphological features that characterize these Hawaiian species is

found in no other species of Lobelioideae and the circumscription adopted here is confidently believed to

create a natural group.

Position and rank.—In contrast to circumscription, there has been a diversity of opinion regarding

the relationships of Lobelia sect. Galeatella to other taxa of Lobelioideae and the appropriate rank at which to

recognize the taxon. Most authors who have commented on relationships agree in identifying as the group's

closest relatives certain species assigned to another section in Lobelia subg. Tupa, sect. Trimeris. However,

they disagree on specifics. Rock (1919) allied the group to tropical Asian members of the group, such as

L. boninensis Koidz. and L. pyramidalis Wall., while Mabberley (1974, 1975) saw a relationship to Brazilian

species such as L. hassleri Zahlbr. and L. organensis Gardner. The endemic Hawaiian genus Trematolobelia

has also been suggested as a close relative by some (Candolle 1839; Stapf 1893; Mabberley 1975). As for

rank, the group has been treated as either a section within Lobelia subg. Tupa (Wimmer 1948, 1953, 1968;

Lammers 1990, 2007b; Murata 1995) or as a genus distinct from Lobelia (Degener & Degener 1962b, 1974,

1983).

Because of the geological youth and de novo volcanic origin of the Hawaiian Islands, it is assumed

that its flora developed from continental species that in some way were able to colonize the archipelago

via long-distance dispersal (Guppy 1906; Brown 1921; Carlquist 1974; Funk & Wagner 1995). For many

years, it was hypothesized that multiple colonizations were required to explain the 134 endemic species of

Lobelioideae: a common one for Clermontia, Cyanea, and Delissea, and separate ones for each of the remain-

ing taxa, including Lobelia sect. Galeatella (Rock 1919; Stone 1967; Carlquist 1980; Mabberley 1974, 1975;

Lammers 1990).

However, recent analyses of molecular data (Knox et al. 1993; Givnish et al. 1995; E. Knox, Indiana

University, pers. comm.; A. Antonelli, Góteborg University, pers. comm.) have dealt a devastating blow to

this polyphyletic hypothesis. All these studies indicate that the endemic Hawaiian Lobelioideae are mono-

phyletic, i.e., that all 134 species in six genera evolved from a single ancestral colonization of the archipelago.

The Hawaiian species (as well as Apetahia Baill. and Sclerotheca A. DC., genera endemic to islands of the

southwestern Pacific) form a well supported clade in these analyses. Its sister-group is a clade comprising

the African and Brazilian species of sect. Rhynchopetalum and sect. Trimeris, and the outgroup of this larger

Hawaiian-African-Brazilian clade is a group of tropical Asian species of sect. Trimeris. Consequently, the

species that Rock (1919) and Mabberley (1974, 1975) believed were the immediate ancestors of sect. Galeatella

are more distantly related, and the groups closest relatives actually are to be found among the Hawaiian

endemics.

The molecular analyses cited above indicate that the Hawaiian clade comprises two subordinate clades:

one comprising the genera with axillary inflorescences (Brighamia, Clermontia, Cyanea, Delissea) or solitary

axillary flowers (Apetahia, Sclerotheca) and the other comprising the taxa with terminal inflorescences (Lo-

belia sect. Galeatella, Lobelia sect. Revolutella, and Trematolobelia). Within the latter subclade, sect. Galeatella

is sister to Trematolobelia while sect. Revolutella is their outgroup.

The sister-group relationship between sect. Galeatella andTrematolobelia is supported by morphology.

Overall, the two are very similar in structure and appearance. Both are pachycaul treelets with a solitary

pliestesial hapaxanthic shoot; a dense apical rosette of coriaceous leaves leaving a helical pattern of raised

corky cicatrices on the unbranched stem; a terminal bracteate raceme of large ornithophilous flowers; calyx

lobes caducous from the young fruit; non-blue corolla with linear monomorphic lobes; and exserted staminal

794 Journal of the Botanical R h Institute of Texas 1(2)

Is of Lobeli t. Galeatella prey 1 following B t al. (2001). Scale b 625 um. A. L villosa (Flynn 685, PTBG). B. L. xkauaiensis

(Fost 14224, K). E. 1 lichaudii (Gart

Fic. 1. S g

(Hitchcock 15437, US). C. L. wahiawa (Forbes 294, K, US). D. L. kool j

subsp. gloria-montis (Hitchcock 14735, US).

( ( EL gloria-montis

column. Both groups also have relatively large winged seeds and an irregular network of sclerenchymatous

bundles in the pericarp, character states unusual among Lobelioideae, which may serve as synapomorphies

uniting them.

In attempting to distinguish Lobelia sect. Galeatella from Trematolobelia, 1 could identify just five morpho-

logical characters that were consistently useful. Using the horizontal posture of the inflorescence branches

as a distinguishing character for Trematolobelia is compromised by the erect unbranched inflorescence of

T. singularis H. St. John; cf., Carlquist 1980, pg. 212D [top]. Similarly, use of corolla color is precluded by

overlap; red and white flowers are found in species of both groups.

Lammers, Revision of Lobelia sect. Galeatella 795

Although both taxa are characterized by a woody stem that is pachycaul, unbranched, pliestesial, and

hapaxanthic, there is a fundamental difference in longevity. While individual plants of sect. Galeatella are

iteroparous (polycarpic) via basal offshoots (the *Tomlinson" architectural model), individuals of Trema-

tolobelia are semelparous (monocarpic), dying by the time the capsules dehisce and disperse ripe seed (the

^Holttum" architectural model; Hallé & Oldeman 1970; Hallé et al. 1978). The latter plan may have evolved

from the former through the simple suppression of the basal offshoots.

Lobelia sect. Galeatella further differs from Trematolobelia, and is unusual among species of Lobelia

subg. Tupa, in having ebracteolate (vs. bibracteolate) pedicels and all five anthers (vs. just the ventral pair)

bearded at the apex with tufts of stiff white trichomes; these two characters are potential synapomorphies

for the species of sect. Galeatella. Conversely, Trematolobelia differs from sect. Galetella and is unique among

Lobelioideae in its strongly secund (vs. ample) inflorescences and laterally poricidal (vs. apically loculicidal)

capsule; these characters are potential synapomorphies for the species of Trematolobelia.

Although the fruit of Trematolobelia (described in detail by Carlquist 1962) is unique in the subfamily,

there is some similarity to the fruit of sect. Galeatella, as noted by Hillebrand (1888) and Stapf (1893). In

all Lobelioideae, the ovary is adnate to a hypanthium of appendicular origin (Kaplan 1967). As a result, the

pericarp of the fruit is actually a combination of hypanthial and gynoecial tissue. In both Trematolobelia and

sect. Galeatella, the calyx lobes are caducous early on; the exocarp and the parenchymatous portions of the

mesocarp deliquesce and are washed away by the frequent rains, leaving behind a series of sclerenchymatous

bundles surrounding the endocarp. In sect. Galeatella, this mesocarpal network is open distally, forming an

open basket or cup. The endocarp it loosely encloses is thick and woody, and dehisces apically through a pair

of triangular loculicidal valves. In Trematolobelia in contrast, this mesocarpal network t distally,

forming a laterally f i sphere. The endocarp is tightly enclosed and adnate to the sclerenchymatous

mesocarp at its summit, precluding the formation of dehiscent valves. Instead, the wall is thin and papery,

rupturing irregularly at its base. This releases the seeds, which accumulate in the bottom of the mesocarpal

network. When the plant is flailed by the strong winds characteristic of its montane habitat, the winged

seeds are shaken out through the lateral foramina in the sclerenchymatous mesocarp.

These similarities suggest that the fruit of sect. Galeatella represents an intermediate step in the evo-

lution of the unique poricidal capsule of Trematolobelia from the loculicidal capsules with intact exocarp

and mesocarp that are more typical of the Lobelioideae. All of the necessary components are in place: the

caducous calyx lobes, the mesocarpal bundles, the deliquescent exocarp and mesocarp parenchyma, and the

winged seeds. All that was required was (1) the apical closure and contraction of the mesocarpal network,

(2) cessation of apical dehiscence via adnation of the endocarp to the mesocarpal sclerenchyma, and (3) the

thinning of the endocarp.

In light of the well supported phylogenetic relationship between sect. Galeatella and Trematolobelia, the

current classification at these ranks appears suboptimal. From a cladistic perspective, sister-groups should

have the same rank. This could be achieved in the present case by elevating sect. Galeatella to generic rank,

following Degener and Degener (1962b, 1974, 1983); by including Trematolobelia in Lobelia as another sec-

tion, following Post and Kuntze (1903); or by expanding Trematolobelia to encompass sect. Galeatella.

However, I have elected to do none of these at present but rather to maintain the status quo and con-

tinue pro tempore to treat this taxon as a section within Lobelia subg. Tupa. First, it seems best to complete

projected studies of sect. Revolutella and Trematolobelia before making changes in the position and rank of

sect. Galeatella. These three taxa are intimately connected and any necessary changes would best be made in

concert. Second, the molecular analyses cited above raise issues that extend beyond far beyond the Hawai-

ian Islands. The sampled species of Lobelia in these studies fall throughout the phylogeny of the subfamily,

from the very first branch to the last; all other genera sampled thus far are embedded among them. Thus,

as currently circumscribed, Lobelia is one of the most unnatural genera imaginable. It not only violates key

principles of cladistic classification, but of traditional taxonomy as well. If we are to avoid treating the entire

subfamily as a single genus comprising 1192 species (Lammers 2007a), it will be necessary to segregate

796 Journal of the Botanical R h Institute of Texas 1(2)

a number of genera from Lobelia. However, the studies cited above have not yet reached the point where

itis feasible to do so. Although the overall problem is obvious, the best resolution is not yet evident. Any

changes in the position and rank of the Hawaiian taxa of Lobelia would best be done as part of the overall

realignment of the genus.

Component taxa.—My original hypothesis, that each of the six areas in which sect. Galeatella occurs

would possess a taxon unique to that locale, was not supported. Analyses of the data revealed a total of seven

taxa. Half the sites harbored two or even three taxa, and only four of the seven taxa were confined to a single

site. On the other hand, all taxa but one were confined to a single island, and even that one exception may

prove to be spurious once additional specimens have been studied.

The specimens examined from the Alaka‘i Swamp of north-central Kauai formed three groups on

the basis of morphology. The most widely distributed, occurring throughout the length of the region from

Pihea to Wai'aleale, is also the most distinctive member of the entire section, differing from all others by

its densely pubescent inflorescence on a conspicuously bracteate peduncle, pale yellow flowers that are the

smallest in the section, floral bracts rounded or subcordate at base, and calyx lobes rounded or obtuse at

apex. The type of L. kauaiensis var. villosa is a representative of this taxon.

At the southeastern terminus of the Alakai Swamp, on the summit of Wai'aleale, are found plants that

differ considerably from the last. This taxon is characterized by depressed obtrullate or depressed obovate

cicatrices on the stems, linear or oblanceolate leaves, a tall (1.5—2.5 m) glabrous branched inflorescence with

blackish purple axes and a largely naked peduncle, relatively narrow floral bracts, relatively large flowers,

calyx lobes acute or acuminate at apex, a pale purple corolla with dark purple longitudinal veins, and large

(1.7-2.5 mm long) seeds. This glabrous taxon corresponds to St. John and Hosaka's (1938) concept of L.

gaudichaudii var. kauaiensis; however, as pointed out by Skottsberg (1944), it does not match the type of that

name: the inflorescence in the type specimen is somewhat pubescent, while the apex of the calyx lobes is

rounded. In fact, none of the type specimens referable to sect. Galeatella match these plants, meaning that

none of the available names apply to the taxon; though known for well over a century, it has no name.

The third taxon is likewise found on Wai'aleale, but also has been collected at the opposite end of

Alakai Swamp, at Pihea. It resembles the unnamed glabrous taxon in its depressed obtrullate or depressed

obovate cicatrices on the stems, linear or oblanceolate leaves, branched inflorescence, relatively large flowers,

longitudinally striped pale purple corolla, and relatively large (2-3 mm long) seeds. However, it differs in

its somewhat pubescent inflorescence with green axes, broader floral bracts, rounded calyx lobes, and pink

corolla venation; the inflorescence is also shorter on average (1-2 m). The types of L. gaudichaudii f. hirsuta

and L. gaudichaudii var. kauaiensis represent this taxon.

As noted above, the densely pubescent taxon that corresponds to the type of L. kauaiensis var. villosa is

found throughout the Alaka'i Swamp and is the most distinctive member of the section. The totally glabrous

taxon occurs at just one place within its range, and at that place, the somewhat pubescent taxon is also

found. The densely pubescent taxon and the glabrous taxon exhibit pronounced differences in flower color

and size. Especially noteworthy is the gap in the lengths of their staminal columns: 45—57 mm long in the

densely pubescent plants vs. 70-80 mm long in the glabrous ones. All this suggests that they are isolated

reproductively under normal circumstances and merit distinction at specific rank. The name L. villosa is

available for the densely pubescent species; the glabrous species is christened L. wahiawa below.

The somewhat pubescent plants at Pihea and Wai'ale'ale resemble L. wahiawa in several characters, as

noted above; the characters by which they differ may be interpreted as due to the genetic influence of L.

villosa. The inflorescence is shorter, more pubescent, and not darkly pigmented; the floral bracts are wider

and the calyx lobes more rounded; and the pigmentation of the corolla veins is diluted. For these reasons,

osa and L. wahiawa. It is this taxon

==

this taxon is here interpreted as an interspecific hybrid between L. vil

to which the name L. kauaiensis properly applies. I generally have not endorsed the practice of christening

occasional interpecific hybrids with binomials. In this case, however, retaining a well known name in a new

status may result in | nfusion than if it simply disappeared. Therefore, L. villosa x L. wahiawa, which has

Lammers, Revision of Lobelia sect. Galeatella 797

been called L. gaudichaudii var. kauaiensis f. hirsuta (St. John & Hosaka 1938; Wimmer 1953) or Galeatella

kauaiensis var. hirsuta (Degener & Degener 1962b, 1974), now will be called L. xkauaiensis.

In the Wahiawa drainage of south-central Kauai, only a single taxon is found and it is L. wahiawa. Thus,

the range of this species is largely allopatric from L. villosa. The latter is confined to the high elevations of

the Alakai Swamp while L. wahiawa occurs in the low elevations of Wahiawa Swamp, except for the single

population in the Alaka'i Swamp at Wai‘ale‘ale. It is possible that the sympatry of the two on Wai'aleale is

a relatively recent phenomenon: the winged seeds of lowland L. wahiawa may have been carried the 10 km

to the mountaintop and successfully established there. Hybridization then occurred from time to time. The

presence of the hybrid without L. wahiawa at Pihea (15 km northwest of Wai'aleale) could be explained by

the dispersal of pollen of the absent parent via nectarivorous bird (cf., Lammers & Freeman 1986; Lammers

et al. 1988) or by dispersal of hybrid seed from Wai‘ale‘ale. It could also be evidence of a second montane

colonization, as yet undocumented or now extirpated, by L. wahiawa.

The specimens examined from O'ahu differ from all others in the section by their relatively short (0.4—1

m) cylindrical inflorescences, distinctly revolute leaves typically pubescent on the abaxial midrib, and small

(1-1.8 mm long) seeds. Other characters, however, vary among these specimens; furthermore, this variation

is correlated with geography. Specimens from the northwestern portion of the Ko'olau Range, between Pu'u

Ka'inapua'a and ‘Eleao at elevations of 670—860 m, are characterized by a 2-6-branched inflorescence, white

corolla, dorsal anthers 14-16 mm long, capsules 18-21 mm long and 9-10 mm in diameter, and seeds 1.5-1.8

mm long; the type of L. gaudichaudii var. koolauensis represents this taxon. Specimens from the southeastern

Ko'olau Range, between Pu'u Pauao and Mt. Olympus at 825-960 m, differ by their usually unbranched

inflorescence, red corolla, shorter (10-13 mm) dorsal anthers, larger (23-30 x 11-10 mm) capsules, and

smaller (1-1.5 mm) seeds; the types of L. gaudichaudii and L. gaudichaudii var. coccinea are representatives of

this taxon. Note that the ranges of these two taxa overlap between Pu'u Pauao and ‘Eleao.

These two taxa were treated previously as conspecific and distinguished at either varietal (St. John &

Hosaka 1938; Wimmer 1953, 1968; Degener & Degener 1962b, 1974) or subspecific (Lammers 1988, 1990)

rank. However, as noted by St. John and Hosaka (1938), the northwestern Ko'olau plants seem almost as

closely related to the plants of Kauai here called L. wahiawa as they do to the southeastern Ko'olau plants.

Characters shared by that species and the northwestern Ko'olau plants (but not the southeastern) include

the branched (vs. unbranched) inflorescence, light-colored (vs. red) corolla, dorsal anthers 14-16 (vs. 10-13)

mm long, and seeds 1.5-2.5 (vs. 1-1.5) mm long.

While the unique characters shared by the O'ahu plants may suggest their derivation from a common

ancestor, the characters that differ, particularly corolla color and anther length, are the sort that might be

indicative of reproductive isolation. This hypothesis is supported by the absence of morphologically inter-

mediate specimens in the zone of sympatry between Pu'u Pauao and 'Eleao. Because the evidence suggests

that these two taxa are unable to interbreed in nature, they are accorded specific rank. The name L. gaudi-

chaudii applies to the southeastern taxon, and the new combination L. koolauensis will be effected below for

the northwestern one.

Specimens seen from West Maui share a distinctive suite of features that at once distinguishes them from

the plants of Kauai and O'ahu: a pyramidal outline to the inflorescence, created by a progressive shortening

of the pedicels from base to apex; lamina with a thickened or subrevolute margin typically ciliate toward

base; the largest flowers and fruits in the group; and seeds of moderate size (2-2.3 mm long). Certain other

characters, however, evince considerable variation.

In the majority of plants from West Maui (including the type specimens of L. gaudichaudii f. kukuiensis

and L. gloria-montis), the inflorescence is glabrous and the corolla white. However, occasional specimens (such

as the type of L. gaudichaudii f. bryanii) have a densely pubescent inflorescence, while in others (including

the type of L. gaudichaudii f. sanguinea) the corolla is red. In neither case could I detect any correlation of

the unusual character state with other morphological characters, or with geography, elevation, or habitat;

the red-flowered variants at least are definitely reported to grow side-by-side with white-flowered plants

798 Journal of the Botanical R h Institute of Texas 1(2)

(Skottsberg 1944; Selling 1947). As such, it appears that each character—pubescent inflorescence and red

corolla—is a simple genetic variant occurring sporadically within a population.

There are other specimens from West Maui that resemble the dominant morphotype in their glabrous

inflorescence and white corolla, but that differ in other characters; furthermore, they are restricted to a dif-

ferent habitat. The plants mentioned in the preceding paragraph occur in open montane bogs on Pu'u Kukui,

'Eke, and other summits. All have a stem less than 1.5 m tall; an unbranched inflorescence; and relatively

short floral bracts, pedicels, and calyx lobes. Specimens collected in forests at the head of Tao Valley (cf.,

Rock 1913a, 1919) differ in their much taller (2.5—4 m) stems, 5-7-branched inflorescence, floral bracts

60-80 (vs. 20-47) mm long, pedicels 50-60 (vs. 15-45) mm long, and calyx lobes 30-35 (vs. 7.5225) mm

long. The type of L. gaudichaudii var. longibracteata is an example of these plants.

In my previous treatment (Lammers 1990), I did not distinguish these larger forest plants from the

smaller bog plants at any rank, primarily because of the close proximity of their distributions. I certainly do

not favor according them each specific rank as Wimmer (1968) and Degener and Degener (1974) have done.

The characters that distinguish the two taxa are not the sort that might be expected to confer reproductive

isolation, and in any case, evince some overlap in their ranges of values. As such, it is deemed best to treat

them as conspecific ecological subspecies. The name for the species is L. gloria-montis. The small bog plants

constitute autonymic subsp. gloria-montis, while the new combination subsp. longibracteata is effected below

for the large forest plants.

Specimens from East Maui (including the type of L. gaudichaudii var. albiflora) are very similar to the

tall forest plants of West Maui, with stem 2-4 m tall, floral bracts 62-65 mm long, pedicels 38-55 mm

long, and calyx lobes 25-30 mm long; their habitat, described as forested margins of bogs, also seems quite

similar. The only substantive difference is that the inflorescence of the East Maui plants, so far as known,

does not branch. Given the small number of specimens representing the tall forest plants of East and West

Maui, it seems best at present to assign them all to a single taxon. Consequently, the specimens from East

Maui are here assigned to L. gloria-montis subsp. longibracteata.

The optimal disposition of the populations on Moloka'i is uncertain. All known specimens (including

the type of L. gloria-montis var. molokaiensis) are sterile. Although one would predict that a geographically

isolated population might at least merit recognition at subspecific rank, the specimens at hand are indistin-

guishable from the typical morphotype of L. gloria-montis subsp. gloria-montis. Until more adequate material

is available for study, these plants are assigned to that taxon.

In summary, sect. Galeatella is treated here as comprising five species. Kauai and O'ahu each have two

endemic species: L. villosa and L. wahiawa on the former, L. koolauensis and L. gaudichaudii on the latter. The

Kauai pair hybridize where sympatric (L. xkauaiensis), while the O'ahu pair do not. The fifth species (L. gloria-

montis) is represented by two ecological races on West Maui: subsp. gloria-montis and subsp. longibracteata.

The populations on East Maui and Molokai are rather poorly known, but are assigned to L. gloria-montis

subsp. longibracteata and L. gloria-montis subsp. gloria-montis, respectively.

TAXONOMIC TREATMENT

Toc sect. Galeatella E. Wimm., Ann. Naturhist. Mus. Wien 56:369. 1948. Galeatella (E. Wimm.) O. Deg. € I.

., Fl. Hawaiiensis, Fam. 339. 1962. Tre (designated by Degener & Degener 1962b): L. kauaiensis var. villosa Rock

Pachycaul treelets of the *Tomlinson" architectural model (cf., Hallé & Oldeman 1970; Hallé et al. 1978), 0.7—5

m tall; plant polycarpic but the shoot pliestesial, hapaxanthic, and replaced by a sucker from the rootstock

after fruit maturation. Stem 0.3-4 m tall, 0.8-10 cm diam., solitary and unbranched, erect or ascending,

leaty only at apex; cicatrices raised, corky, transversely rhombic, depressed obtrullate, or depressed obovate

in outline with a single central vein scar, closely packed in a helical pattern and long persistent; pith soft,

broader than the lignified cortex toward apex but narrowing toward base as the cortex increases in thickness,

becoming hollow with age; latex white, viscous. Leaves alternate, simple, exstipulate, sessile, glabrous or

pubescent, forming a dense spherical or oblate apical rosette; lamina oblong, elliptic, lanceolate, oblanceo-

late, or linear, coriaceous, stiff, flat; margin entire, commonly revolute, with few to several intramarginal

Lammers, Revision of Lobelia sect. Galeatella 799

callosities distally; apex acuminate, acute, or rarely obtuse; base attenuate or cuneate. Flowers tetracyclic,

perfect, proterandrous, zygomorphic, epigynous, pedicellate, resupinate, ornithophilous, large; inflorescence

an erect terminal pedunculate bracteate raceme, 0.4—2.5 m tall, unbranched or 2-7-branched from base,

up to 100-flowered over its life, but seldom more than 4—8 open at any one time, glabrous or pubescent;

peduncle usually shorter than rachis, densely to sparsely covered with reduced leaves (sterile bracts) that

rapidly or gradually decrease in size acropetally; floral bracts conspicuous, equaling or shorter than the

pedicels; pedicels ascending, stiff, ebracteolate. Calyx synsepalous, glabrous or pubescent; tube adnate to

the basal '/,—/, of the ovary, forming a broadly obconic (rarely campanulate or hemispherical) appendicular

hypanthium //,—,, as long as the corolla; lobes 5, valvate, triangular, lanceolate, ovate, or oblong, shorter than

the corolla tube, a little shorter than the hypanthium to over 4 times longer, the dorsal slightly longer than

the ventral, caducous from the young fruit, the margin entire, the apex acuminate, acute, obtuse, or rounded

and sometimes apiculate. Corolla sympetalous, zygomorphic, subbilabiate, of various colors (white, pale yel-

low, red, or pale purple with darker longitudinal veins) but never blue, glabrous (the lobes rarely pubescent

at apex); tube straight or slightly curved, cylindric or expanding slightly towards the mouth, 1.3—5 times

longer than broad, dorsally cleft nearly to base but never fenestrate; lobes 5, valvate, linear, monomorphic,

acuminate or acute at apex, the dorsal pair spreading or recurved horizontally, about as long as the tube to

3 times longer, the three ventral lobes connate for ?/, their length or more, forming a deflexed inrolled trifid

lip shorter than the dorsal lobes. Stamens 5, antisepalous, connate distally for most of their length, strongly

exserted, emerging from the dorsal corolla slit above or between the dorsal lobes; filament tube suberect or

slightly deflexed, 2.8—6.3 times longer than anther tube, glabrous (rarely short pubescent ventrally); dorsal

anthers longer than the ventral, overhanging the orifice of the tube and somewhat occluding it; all 5 anthers

with tufts of white trichomes at apex; pollen 52-54 um polar diam. x 35-41 pm equatorial diam., prolate,

tricolporate, 3-celled at anthesis, the exine shallowly and minutely reticulate (Selling 1947; Erdtman 1952;

Pandey et al. 1993). Ovary 2-loculed, inferior, adnate to the hypanthium for /,—/, its length, the conic apex

free, forming a pronounced beak on the young fruit; placentae axile; ovules numerous; style 1, slender,

terete, with a ring of stiff white hairs near the apex; stigma 2-lobed, the lobes appressed and non-receptive

as the style grows through the anther tube, pushing out pollen, after which the stigmas spread and become

receptive. Fruit an ovoid or ellipsoid capsule; exocarp and parenchymatous portions of the mesocarp deli-

quescent, leaving a loosely enshrouding series of sclerenchymatous bundles, open at top; endocarp rigid

and durable, its conical apex splitting loculicidally into a pair of triangular valves; seeds (Fig. 1) winged,

oblong, compressed, honey- or chestnut-brown, the narrow wing paler, the testa faintly striate (Type C of

Murata 1992). Chromosome number unknown (Lammers 1993).

These species are restricted to wet habitats at elevations of 570-1760 m on four of the Hawaiian Islands.

On Kauai and Maui, they are found in level saturated bogs, while on O'ahu and Molokai, they occur on

windswept ridge crests and summit slopes. The plant communities that develop in all these areas (cf., Gagné

& Cuddihy 1990) typically are dominated by dwarfed shrubs and various graminoids, and the species of

Lobelia sect. Galeatella are typically among the tallest and most conspicuous elements in the vegetation (cf.,

Carlquist 1980, pp. 346, 350, 352 [bottom]). Flowering begins in mid-summer and is generally finished by

October, with most seed maturing by January.

The flowers of these species, like most Hawaiian Lobelioideae, are presumed to be ornithophilous

(Lammers & Freeman 1986; Lammers et al. 1988; Smith et al. 1995). Peale's (1848: 152) illustration of the

endemic nectarivorous bird Vestiaria coccinea (Forster) of the Drepanidini (Aves: Fringillidae) includes an

unequivocal depiction of Lobelia gaudichaudii; he commented that the bird's bill was *admirably adapted" for

extracting nectar from its corolla. The flat winged seeds (Fig. 1) are presumably suited for a limited degree

of anemochory.

Although none of the species has been examined cytologically, the group is hypothesized to be tetra-

ploid with 2n = 28, in light of the fact that all counted species of its sister-group (Trematolobelia) and their

outgroup (Lobelia sect. Revolutella) have that number (Lammers 1993).

800 Journal of the Botanical R h Institute of Texas 1(2)

KEY TO THE SPECIES AND SUBSPECIES

1. Inflorescense densely pul the peduncle d ly covered with sterile bract lually d g

l H | JI AA P | | | 13 s | d ; l | Ii y 24mm

long; fi filament tube EE 45 mm long; capsule 1 13- 18 mm "d de Swamp ofKaual) — — 1. Lobelia villosa

1. Inflorescense glabrous (rarely pubescent), th covered with sterile bracts rapidly decreas-

ing in size acropetaly, the floral Pine cuneate at base; I ee ithium SE. e) en eax lobes

acuminate or acute (rar ly rouna obtuse) a | with darker lor \gitudinal veins, red,

or white, 53-103 mm long, the Abe 2 5 times uid than broad, the ventral lip 24-50 mm long; filament

tube 57-80 mm long; capsule 19-37 mm lon

2. Cicatrices depressed obtrullate or depressed obovate, the corners rounded; lamina 17-37 cm long, com-

monly linear; inflorescence 1-2.5 m tall; corolla pale purple with dark purple or pink longitudinal veins;

seeds (1.7-)2-3 mm long (Kaua'i).

3. Inflorescence 1-2 m tall, ee 35-flowered, its axi | | | | urled

trichomes; floral bracts elliptic, oblong, or obovate; calyx lobes eed ae COS yth DOE

(Alakaʻi Swamp) 2.L jen xkauaiensis

3. Inflorescence 1.5-2.5 m tall, 20-85-flowered, its axis blackish purple and glabrous; floral bract

narrowly elliptic, narrowly oblong, or lanceolate; calyx lobes acuminate or acute (rarely obtuse) a

apex; corolla with dark purple veins (Wahiawa & Alaka'i Swamps) 3. Lobelia wahiawa

2. Cicatrices transversely rhombic, the corners acute; lamina 9-24 cm long, commonly oblanceolate; inflo-

rescence 0.4-1.5 m tall; corolla red or white; seeds 1-2(-2.3) mm long.

4. Inflorescence 0.4-1 m tall, cylindrical in outline; lamina di tly revolute at margin, pubescence (if any

confined to abaxial midrib; corolla 58-78 mm long, the tube 19-31 mm long, the dorsal lobes 27-45

a

mm long; capsule 18-30 mm long; seeds 1-1.8 mm long (O'ahu).

. Inflorescence 2-7- E corolla ane dorsal anthers 14-16 mm long; capsule 2 mm

long x 9-10 mm diameter; seeds 1.5-1.8 mm long (NW Ko'olau Range) Lobelia koolauensis

. Inflorescence unbranched; corolla red; dorsal anthers 10-13 mm long; capsule 2: 33430. "y mm lon

11-19 mm diameter; id 15 mm TOO GE Galan Range 5. Lobelia T

—

4. mc 15mtall, tel in, pubescence

, EN SOLES D E

long; capsule 27-37 mm long; seeds 2-2. s mm eng LOU MED 6. Lobelia ions montis

6. Stems0.4-1.5 mtallifloralbracts 20-47 lat licels 15-45mmlong;cal

lobes 7.5-25 mm long (Moloka'i, West Maui) 6a. Lobelia gloria-montis subsp. gloria-montis

6. Stems2-4m tall floral bracts 60-90 mm long, oblong or linear; pedicels 38-60 mm long; calyx lobes

25-35 mm long (West and East Maui) 6b. Lobelia gloria-montis subsp. longibracteata

1. Lobelia villosa (Rock) H. St. John & Hosaka, Occas. Pap. Bernice Pauahi Bishop Mus. 14:124. 1938.

Lobelia kauaiensis var. villosa Rock, Bull. Torrey Bot. Club 44:237. 1917. Galeatella villosa (Rock) O. Deg. & I. Deg., Fl. Hawaiiensis,

Fam. 339. 1962. Type: HAWAIIAN ISLANDS. Kauas: Wai‘ale‘ale, 21 Oct 1916, Rock 12844 (HoLoTYPE: BISH!; isotypes: BISH! K!). The

collection number is given as “12741” in the protologue and subsequent monograph (Rock 1919), but the figured specimen in the

latter (pl. 61; called “Type” in the caption) shows this sheet with this number.

Plant 1.2-2.2 m tall. Stem 0.6-2 m tall x1-6 cm diam.; cicatrices 3-3.5 mm tall x 6-7 mm wide, transversely

rhombic, the corners acute. Lamina 14-22 cm long x 1.73 cm wide, oblong or oblanceolate, densely pubescent

on margin, sometimes also pubescent on the dorsal and/or ventral midrib; margin subrevolute; apex obtuse

or acute; base cuneate or attenuate. Inflorescence 0.5-1 m tall, 2-5-branched (rarely unbranched in small

plants), densely pubescent, each branch 13-45-flowered and cylindrical in outline, its axis green; peduncle

densely covered with sterile bracts that gradually decrease in size acropetally; sterile bracts 25-120 mm

long x 10-30 mm wide, elliptic, oblong, lanceolate, ovate, or obovate, the apex acute or obtuse, the base

rounded; floral bracts 17-30 mm long x 8-18 mm wide, elliptic, ovate or obovate, the apex acute or obtuse,

the base rounded or subcordate; pedicels 15-35 mm long, of more-or-less equal length throughout. Hypan-

thium 5-9 mm long x 7-11 mm diam., broadly obconic, '/,—'/,, as long as the corolla, densely pubescent;

calyx lobes 8.5-12 mm long x 4-5 mm wide, triangular or oblong, equaling the hypanthium to 2.2 times

as long, the apex obtuse or rounded. Corolla 44-57 mm long, pale yellow, glabrous; tube 15-18 mm long x

8-13 mm diam. at middle, straight, cylindric or expanding slightly towards the mouth, 1.3-2 times longer

Lammers, Revision of Lobelia sect. Galeatella 801

than broad; dorsal lobes 29-39 mm long x 3.5-5 mm wide, 1.6-2.2 times longer than the tube; ventral lip

22-24 mm long x 12-17 mm wide. Filament tube 36-45 mm long, slightly deflexed, 2.8—4.5 times longer

than anther tube, glabrous; anther tube 4 mm diam.; dorsal anthers 9-13 mm long; ventral anthers 8-9.5

mm long. Capsule 13-18 mm long x 10-12 mm diam.; seeds (Fig. 14; Wimmer 1953, fig. 104c) 1.5-2.5

mm long x 0.6-0.8 mm wide.

Icon.—Rock (1919), pl. 43. Figured specimen.—Rock (1919), pl. 61 [holotype].

Distribution, habitat, and phenology.—Endemic to north-central Kauai, known only from Montane

Wet Mixed Communities (Gagné & Cuddihy 1990) of the Alaka‘i Swamp (cf., Carlquist 1980, p. 350) from

Pihea to Wai'ale'ale, at 1100-1560 m. Flowering from August to October. Considered “rare” by Wagner et

al. (1999).

Representative specimens: KAUAI. Waimea: Alakai, bogs between Lel | d Kilohana, Cranwell et al. 2984 (BISH, GB); Alaka‘i

Swamp, in 2"™ large bog on trail to old USGS cabin on way to Wai‘ale‘ale, Davis & Kores 131 (BISH); Alaka'i Swamp, Flynn 685 (PTBG),

Flynn 686 (PTBG); Alaka'i Swamp, Forbes 1122.K (BISH); Wai‘ale‘ale, Hitchcock 15450 (US); Alaka'i Swamp, along trail between Pihea and

Kilohana, Lammers et al. 5376 (DUKE, F, OS); Wai‘ale‘ale, Rock 5823 pro parte (GH), Rock 8856 (BISH); Wai'ale'ale, 23 Sep 1909, Rock s.n.

(BISH), Oct 1911, Rock s.n. (BISH); Alaka'i Swamp, Yuncker 3524 (DPU, US).

2. Lobelia xkauaiensis (A. Gray) A. Heller, pM. Bot. Stud. 1:911. 1897 (as ‘kauaensis’, pro sp.) stat.

NOV. Lobelia gaudichaudii var. kauaiensis A. Gray, Proc. Amer. Acad. Arts 5:150. 1861 (as “kauaensis’). Galeatella kauaiensis (A. Gray)

O. Deg. & I. Deg., Fl. Hawaiiensis, Fam. 339. 1962 d p Type: HAWAIIAN ISLANDS. Kauai: [Alaka‘i Swamp, 30 Oct 1840,

leg. W. | & C. Pickering (fide Wilkes 1845),] U.S. Exploring p s.n. is OTYPE: US!; ISOTYPE: PU i: d is

Es on m fable Hawaiian proper noun “Kaua'i;” Grays (1861

error to be corrected, following Skottsberg (1944), Wimmer (1968), and Degen and egerat (1974)

du

Lobelia gaudichaudii f. hirsuta H. St. John & Hosaka, Occas. Pap. Bernice Pauahi Bishop Mus. 14:121. 1938. Galeatella kauaiensis var.

hirsuta (H. St. John & Hosaka) O. Deg. & I. Deg., Fl. Hawaiiensis, Fam. 339. 1962. Type: HAWAIIAN ISLANDS. Kaua‘t: Wai‘ale‘ale,

Oct 1916, Rock 12845a (HOLOTYPE: BISH!; IsoTYPE: BISH!).

Plant 2.4—4 m tall. Stem 1-2 m tall x1-6 cm diam.; cicatrices 3-4.5 mm tall x 4.5-8 mm wide, depressed

obtrullate or depressed obovate, the corners rounded. Lamina 24-32 cm long x 2.2-3.3 cm wide, linear,

oblanceolate, or narrowly elliptic, glabrous or pubescent on ventral midrib; margin subrevolute; apex acu-

minate; base attenuate or cuneate. Inflorescence 1-2 m tall, 5-branched, sparsely pubescent with relatively

short curled trichomes, each branch 13-35-flowered and cylindrical in outline, its axis green; peduncle

sparsely covered with sterile bracts that rapidly decrease in size acropetally; sterile bracts 30-105 mm long

x 4-14 mm wide, linear or narrowly oblong, the apex acuminate, the base cuneate; floral bracts1 2-23 mm

long x 4-10 mm wide, elliptic, oblong, or obovate, the apex obtuse or rounded, the base cuneate; pedicels

25-45 mm long, of more-or-less equal length throughout. Hypanthium 6-10 mm long x 13-15 mm diam.,

broadly obconic, 1/7-1/11 as long as the corolla, glabrous or pubescent; calyx lobes 8-13 mm long x 3-6

mm wide, triangular or oblong, equaling to a bit longer than the hypanthium, the margin entire, the apex

rounded. Corolla 64-66 mm long, pale purple with pink longitudinal veins, glabrous; tube 20-22 mm long

x 10-12 mm diam. at middle, slightly curved, expanding slightly towards the mouth, twice as long as broad;

dorsal lobes 42-46 mm long x 4—5 mm wide, about twice as long as the tube; ventral lip 37-39 mm long x

15-17 mm wide. Filament tube 64—66 mm long, suberect, 4 times longer than anther tube, glabrous; anther

tube 4 mm diam.; dorsal anthers 14-16 mm long; ventral anthers 10-11 mm long. Capsule 19-30 mm long

x 8-20 mm diam.; seeds (Fig. 1B) 2-3 mm long x 0.7-0.9 mm wide, chestnut-brown.

Icones.—Rock (1919), pl. 42 [right]; Wimmer (1968), fig. 29.

Distribution, habitat, and phenology.—A putative hybrid of L. villosa and the next species, known only

from the Montane Wet Mixed C ities (Gagné & Cuddihy 1990) at the southeastern and northwestern

extremities of the Alakai Swamp on north-central Kaua‘i. At the former site (Wai'aleale, 1370-1570 m), it

occurs with both parents, as evinced by Roch 5823; the sheet at BISH represents the hybrid, while that at GH

is a mixed gathering of the two parents. At the latter site (Pihea, 1220-1350 m), L. villosa co-occurs but not

the other parent. Flowering in August, fruit ripening in October. This taxon is listed as “rare” by Wagner et

al. (1999), an assessment that also encompasses L. wahiawa.

802 Journal of the Botanical R h Institute of Texas 1(2)

Discussion.—Plants on Kauai that resemble the following species except for their puberulous raceme

have been treated as a minor variant of it by most authors (Rock 1919; St. John & Hosaka 1938; Wimmer

1953; Degener & Degener 1962b, 1974). In my earlier treatment (Lammer 1990), I suggested that they were

more likely interspecific hybrids, the pubescence indicating gene-flow from L. villosa. Unfortunately, I missed

clues (Gray 1861; Skottsberg 1944) that the type specimen of L. kauaiensis was an example of this same

hybrid. As a result, not only does the name L. gaudichaudii f. hirsuta apply to the hybrid, but L. kauaiensis

as well. The latter name is used here to denote the hybrid, with the addition of the multiplication sign to

denote its status as a nothotaxon.

R KAUA'I. Hanalei: ridge E of Pil | 1s Kilol Flynn & DeLappe 3198 (BISH, F, K, MO, PTBG, US);

Norn Bog one Wainiha pali, Wood & Perlman 3672 E PTBG). Waimea: Napali-Kona Forest AUR Wai'ale'ale trail, Alaka'i Swamp

Keáku River, Chock 994 (BISH, NSW); Wai‘ale‘ale, along trail from “Kauku” [Keakü?], Cranwell et al. 3047 pro parte (GB); Wai'aleale,

Hitchcock rd a (US); Wai'ale'ale, Rock 5109 (BISH); Wai'ale'ale, Sep 1909, Roch s.n. (BISH); Wai'ale'ale, Oct 1911, Rock s.n. (BISH);

Wai'ale'ale, R I BISH); ridge SW of Pihea, on east rim of Kalalau Valley, St. John et al. 22945 (DUKE).

E

3. Lobelia wahiawa Lammers, Sp. nov. Type: HAWAIIAN ISLANDS. Kauai: Koloa District, Wahiawa Swamp near headwaters of

Wahiawa Stream, 630 m, open swamp with some shrubby areas, frequent, 24 Apr 1964, Stauffer & Dehler 5910 (HoLotyeE: BISH!;

isotypes: A! DUKE! K! US).

leatellae | i ] i ] i imile L. xkauaiensi, cuj ,sed

o [2] E E in E a} L^

is vel acuminatis, et corollae venatione atro-purpurea differt.

Species Lobeliae sect.

Tai

dead

inflorescentia glabra calvcis

Plant 2-4 m tall. Stem 1-2.5 m tall x 1-10 cm diam.; cicatrices 3-5 mm tall x 4.5-9 mm wide, depressed

obtrullate or depressed obovate, the corners rounded. Lamina 17-37 cm long x 13.3 cm wide, linear,

oblanceolate, or narrowly elliptic, glabrous; margin subrevolute; apex acuminate; base attenuate. Inflores-

cence 1.5-2.5 m tall, 2-5-branched (rarely unbranched in small plants), each branch 20-85-flowered and

cylindrical in outline, its axis blackish purple, glabrous; peduncle sparsely covered with sterile bracts that

rapidly decrease in size acropetally; sterile bracts 12-160 mm long x 4-22 mm wide, linear, narrowly el-

liptic, or lanceolate, the apex acuminate, the base cuneate; floral bracts 16-27 mm long x 3-7.5 mm wide,

narrowly elliptic, narrowly oblong, or lanceolate, the apex acute or obtuse and sometimes apiculate, the base

cuneate; pedicels 17-35 mm long, of more-or-less equal length throughout. Hypanthium 5-9 mm long x

8-14 mm diam., broadly obconic or rarely almost hemispherical, 1/8-1/12 as long as the corolla, glabrous;

calyx lobes 9-15 mm long x 3-6.5 mm wide, triangular, narrowly oblong, or lanceolate, equaling to 3 times

longer than the hypanthium, the margin entire, the apex acuminate or acute (rarely obtuse) and sometimes

apiculate. Corolla 53-75 mm long, pale purple with dark purple longitudinal stripes, glabrous; tube 18-28

mm long x 6-10 mm diam. at middle, suberect or slightly curved, expanding slightly towards the mouth,

2—4.5 times longer than broad; dorsal lobes 30-48 mm long x 3-5 mm wide, 1.3-2.2 times as long as the

tube; ventral lip 25-38 mm long x 11-18 mm wide. Filament tube 57-65 mm long, decurved slightly, 4—5

times longer than anther tube, glabrous; anther tube 4-5 mm diam.; dorsal anthers 12-16 mm long; ventral

anthers 9-12 mm long. Capsule 20-28 mm long x 10-14 mm diam.; seeds (Fig. 1C) 1.7-2.5 mm long x

0.6-1.4 mm wide, honey- or chestnut-brown.

Icones.—Rock (1919), pl. 16; Carlquist (1980), p. 245 [left], p. 348 [top left]; Sohmer and Gustafson (1987),

fig. 114; Lammers (1990), pl. 57 [lower left]. Figured specimen.—Rock (1919), pl. 60 [Heller 2888, GH].

Distribution, habitat, and phenology.—Endemic to central Kauai, known primarily from the Lowland Wet

Mixed Communities (Gagné & Cuddihy 1990) at 570-800 m in the watershed of Wahiawa Stream west of

Kahili (cf., Carlquist 1980, p. 346), but also found in the unique Montane Wet Mixed Community (Gagné €

Cuddihy 1990) of the summit bog on Wai'aleale at 1100-1590 m. Flowering July to March, fruit ripening

December to April. Regarded as “rare” by Wagner et al. (1999), an assessment that includes its hybrid with

L. villosa (see previous taxon).

Etymology.—The specific epithet is the name of the type locality, used as a noun in apposition; it is

Hawaiian for “milkfish place" (Pukui et al. 1974), a reference to widespread Chanos chanos (Forsskál) of the

Chanidae (Actinopterygii).

Lammers, Revision of Lobelia sect. Galeatella 803

Vernacular name.—Pu'e (Rock 1919; Pukui & Elbert 1971).

Discussion.—This is the glabrous species that has been known for many years (Heller 1897; Rock 1919;

Wimmer 1968; Lammers 1990) as Lobelia kauaiensis. However, as noted by Skottsberg (1944), the original

description of the basionym of that name (Gray 1861) called for a plant *racemo puberulo; calycis viscosi"

and its typ indeed conforms to that diagnosis. As a result, this glabrous species has no name and

r

so is named here as a new species.

Representative specimens: KAUAT. [Kawaihau: Pohakupili (fide Wawra 1873), l Wawra 2044 (W). Koloa: Wahiawa Bog, Carlquist 1980

(RSA); Wahiawa Bog, 26 Nov 1972, Carlquist s.n. (RSA); Wahiawa Bog, Fagerlind & Skottsberg 6480 (GB, UPS, S); Hanapepe, in turfosis

Faurie 552 (A, BISH, P, W); Wahiawa Bog, Flynn 371 (PTBG); Wahiawa Bog, Flynn 668 (OS); Wahiawa Bog, Flynn 699 (OS); Wahiawa

Swamp, Forbes 294.K (BISH, US, W); Wahiawa Swamp, Aug 1909, Forbes s.n. (BISH); Wahiawa Bog, Gustafson 1049 (BISH, RSA); in and

near a bog at the head of the Wahiawa, 19 Oct 1895, Heller 2888 a GH, MO, NY, US); Wahiawa Bog, Kato LK-2 (BISH); Wahiawa Bog,

Lammers et al. 5861 (OS); Wahiawa Bog, 28 Dec 1980, Lucas s.n. (RS Wahiawa, MacDaniels 636 (BH, BISH); Wahiawa Swamp,

Sparre H.75 (S); Wahiawa Bog area west of Mt. Kahili, Stern & Carlquist 1331 (RSA, US); Wahiawa Bog, Stern & Herbst 2942 (PTBG, RSA);

kahili Swamp, Wahiawa, St. John et al. 10849 (A, K, BISH, NSW); Kahili Bog, Wahiawa, St. John & Fosberg 13561 (BISH, POM); west side

of Wahiawa drainage below ridge between Hulua and Pu'u'au'uka, Wood et al. 1059 (MO, PTBG). Waimea: Wai'aleale, along trail from

“Kauku” [Ke'akú?], Cranwell et al. 3047 pro parte (BISH); Wai'ale'ale summit, Davis & Kores 113 (A, BISH) & 113a (BISH); Wai'aleale, Hitch-

cock 15437 pro parte (US); Wai'ale'ale, Hitchcock 15499 (US); Wai‘ale‘ale, Rock 5115 (BISH); Wai‘ale‘ale, Rock 5823 pro parte (GH); Wai'ale'ale,

Rock 12845 (BISH, K); Wai‘ale‘ale, Sep 1909, Roch s.n. (NY, US, W); along Alaka'i Swamp Trail, Stone 1536 (BISH).

4. Lobelia koolauensis (Hosaka & Fosberg) Lammers, comb. et stat. nov. Lobelia gaudichaudii var. koolauensis Hosaka €

dde Occas. Pap. Bernice Pauahi Bishop Mus. 14:4. 1938. e NH CM var. aa auensis dd: & Fosberg) O. Deg. Y I.

Fl. Hawaiiensis, Fam. 339. 1962. Lobeli g Syst. Bot. 13:506. 1988

Deg.,

Type: HAWAIIAN ISLANDS. O'aHu: Ko'olau Range, divide | lof} i Nui and Kai lc]

sloping bog, 2600 ft, 31 May 1937, Hosaka & gn 1915 (uoLotyre: BISH [mounted on 5 pm ; ISOTYPES: BISH[2]! US! WD.

Plant 1-2.5 m tall. Stem 1-1.5 m tall x 1.8-3 cm diam.; cicatrices 3-4 mm tall x 6.5-10 mm wide, transversely

rhombic, the corners acute. Lamina 11.8-20 cm long x 0.9-2.8 cm wide, linear, oblanceolate, or narrowly

elliptic, glabrous or the abaxial midrib pubescent; margin distinctly revolute; apex acute or acuminate;

base attenuate. Inflorescence 0.5-1 m tall, 2-7-branched (rarely unbranched in small plants), each branch

12-25-flowered and cylindrical in outline, its axis green, glabrous; peduncle sparsely covered with sterile

bracts that rapidly decrease in size acropetally; sterile bracts 20-95 mm long x 4-13 mm wide, narrowly

oblong, narrowly elliptic, or lanceolate, the apex acute, the base cuneate; floral bracts 20-48 mm long x 5-10

mm wide, narrowly elliptic or lanceolate, the apex acuminate, the base cuneate; pedicels 22-60 mm long,

of more-or-less equal length throughout. Hypanthium 5-10 mm long x 7-11 mm diam., broadly obconic,

1/6-1/8 as long as the corolla, glabrous; calyx lobes 10-22 mm long x 3.5-6 mm wide, narrowly d

or lanceolate, equaling to 4.4 times longer than the hypanthium, the margin entire, the apex and

sometimes apiculate. Corolla 58-78 mm long, white, glabrous; tube 19-31 mm long x 8-12.5 mm diam. at

middle, suberect, expanding slightly towards the mouth, 2-3.5 times longer than broad; dorsal lobes 27-45

mm long x 4.5-6.5 mm wide, 0.9-2.4 times as long as the tube; ventral lip 24-40 mm long x 17-24 mm

wide. Filament tube 61-78 mm long, suberect, 4-5 times longer than anther tube, glabrous; anther tube

4—4.5 mm diam.; dorsal anthers 14-16 mm long; ventral anthers 9-10 mm long. Capsule 18-21 mm long

x 9-10 mm diam.; seeds (Fig. 1D) 1.5-1.8 mm long x 0.8-1 mm wide, honey-brown.

Icones.—Hosaka and Fosberg (1938), fig. 2; Wimmer (1953), fig. 104a.

Distribution, habitat, and phenology.—Endemic to O'ahu, known only from Montane Wet Shrublands

(Gagné € Cuddihy 1990) on the summit of the northwestern Ko'olau Range, between Puu Ka'inapua'a and

'Eleao, at 670—860 m. Flowering from late May to October, fruits ripening November to January. Considered

“endangered” by Wagner et al. (1999).

Representative specimens: O'AHU. Ko'oLau Mrs.: Poamoho Trail, Paaloa-Wahiawa, Cowan 252A (WAB); Poamoho Trail, Laie, Degener

et al. 11497 (MO, W); top of Poamoho Trail crest at head of Punalu'u Valley, Fosberg 13330 (BISH); main divide above Kaipapa'u Gulch,

Fosberg 14224 (BISH, K); Manana Trail, NE of Pacific Palisades, 12 Jul 1985, Obata s.n. (F, OS); Mánana Trail 1 km from summit, 12 Aug

1996, Obata et al. s.n. (BISH, OSH); Manana Trail 0.8 km from summit, Obata et al. 569 (BISH), Obata et al. 570 (BISH); Manana Trail,

below summit, Perlman & Obata 6484 (BISH, PTBG, US); Manana Valley, north-facing slope, Perlman et al. 17171 (PTBG); Manana Trail

to summit, next ridge north of Mànana, Wood et al. 8513 (PTBG).

804 Journal of the Botanical R h Institute of Texas 1(2)

5. Lobelia gaudichaudii A. DC. in DC., Prodr. 7:384. 1839. Dortmanna gaudichaudii (A. DC.) Kuntze, Revis. Gen. Pl

2:972. 1891. Galeatella gaudichaudii (A. DC.) O. Deg. & I. Deg., Fl. Hawaiiensis, Fam. 339. 1962. Type: HAWAIIAN ISLANDS. O'aHU:

sommet de la montagnes à gauche de là vallée de Onoruru [i.e., Honolulu], [8-24 Oct 1836, fide Kuykendall (1947)] Gaudichaud

s.n. (HOLOTYPE: P-00520806!; isotypes: GH! G-DC! P-00520807!)

Lobelia gaudichaudii var. coccinea Rock, Bull. Torrey Bot. Club 44:238. 1917. Type: HAWAIIAN ISLANDS. O‘anu: summit of Mt. Konahuanui,

1912, leg. G.W. Shaw, Rock 12742 (HoLotyPE: BISH!; isotypes: BISH[3]! GH! US).

Plant 1-3 m tall. Stem 0.8-2 m tall x 1.3-5 cm diam.; cicatrices 2.5—4.5 mm tall x 7-11 mm wide, trans-

versely rhombic, the corners acute. Lamina 9.5-19.5 cm long x 1.1-2.8 cm wide, oblanceolate, densely long

pubescent on the abaxial midrib (rarely glabrous); margin distinctly revolute; apex acute or acuminate; base

attenuate. Inflorescence 0.4-0.8 m tall, unbranched, 18-36-flowered and cylindrical in outline, its axis

green, glabrous; peduncle sparsely covered with sterile bracts that rapidly decrease in size acropetally; sterile

bracts 25-110 mm long x 10-18 mm wide, linear, narrowly elliptic, oblong, or ovate, the apex acuminate

or acute, the base cuneate; floral bracts 18-32 mm long x 6-8 mm wide, ovate or lanceolate, the apex acute

or acuminate, the base cuneate; pedicels 20-40 mm long, of more-or-less equal length throughout. Hypan-

thium 6-10 mm long x 7.5-12 mm diam., broadly obconic or campanulate, 1/8-1/9 as long as the corolla

glabrous; calyx lobes 13-18 mm long x 4.5-7 mm wide, ovate or lanceolate, 1.6-2.5 times longer than

the hypanthium, the margin entire, the apex acuminate or acute. Corolla 60-65 mm long, red, glabrous;

tube 20-30 mm long x 6-11 mm diam. at middle, slightly curved, expanding slightly towards the mouth

2—4.5 times longer than broad; dorsal lobes 30-40 mm long x 4-5 mm wide, equaling to twice as long as

the tube; ventral lip 24-30 mm long x 12-20 mm wide. Filament tube 58-66 mm long, decurved slightly,

4—6 times longer than anther tube, glabrous; anther tube 4-6 mm diam.; dorsal anthers 10-13 mm long;

ventral anthers 7.5-10 mm long. Capsule 23-30 mm long x 11-19 mm diam.; seeds (Fig. 1E) 1-1.5 mm

long x 0.5-0.7 mm wide, chestnut-brown.

Icones.—Gaudichaud (s.d.), pl. 45; Peale (1848), pg. 152; Rock (1919), pl. 56; Degener (1938b); Murata

(1995), figs. 56-57 [seed only]; Smith et al. (1995), fig. 1. Figured specimen.—Rock (1919), pl. 57 [holotype

of L. gaudichaudii var. coccinea].

Distribution, habitat, and phenology. —Endemic to O'ahu, known only from Montane Wet Shrublands

(Gagné & Cuddihy 1990) on the summit of the southeastern Ko'olau Range, between Pu'u Pauao and Mt.

Olympus, at 805-960 m. Flowering from July to October, fruits ripening November to January. Considered

“vulnerable” by Wagner et al. (1999)

Representative specimens: O'AHU. Ko'oLau Mrs.: windward pali of Puu Kónáhuanui, Bishop 1188 (BISH); Konahuanui, 10 Jan 1904, Bryan

s.n. (BISH); near summit Konahuanui (42), 30 Jul 1922, Bryan s.n. (BISH); main ridge between Konahuanui and Mt. Olympus, Degener

7965 (NY); Bowman Trail, at divide, leg. M. Kerr, i ud 19218 (NY); near summit ur im Trail, leg. M. Kerr, Degener 21674 (B, W);

mauka of Tripler Hospital, Degener 30174 (N Konahuanui 0.0 (B MO, NY); Forbes 2182.0 (BISH); between

Punalu'u and Kaipapa'u, 8-13 May 1909, Forbes & Thompson s.n. (BISH); ae of Mt. umm Garber 82 (BISH); Mt. Lanihuli peak

Garber 261 (BISH); Lanihuli, Garber 262 (BISH); Kipapa-Waiahole crest, Grant 7217 (BISH); summit of Konahuanui, Hillebrand 72 (K),

Kipapa M on main ie Hosaka 686 (BISH, K); Konahuanui, 100 yd E of East Peak, 13 Oct 1974, Obata s.n. (B, BISH); summit of

‘Aiea Ridge tr t, 3 Nov 1974, Obata s.n. (BISH); Kipapa Trail, right ridge of Kipapa Gulch, Ozaki 561 (BISH); Punalu‘u,

Rock 64 (BISH, GH), Rock 65 (BISH); Mt. Lanihuli, leg. Nelson & Stone, Rock 10003 (BISH); Konahuanui, Rock 10003 (BISH); Lanihuli,

leg. Daingerfiel

¡[añ

B, S); about Y2 hr Poamoho side of Castle Trail Summit along the Ko'olau Crest Trail, Stemmermann &

i pid 1207 (BISH); Waipi o-Waiahole divide, Kipapa Gulch, St. John 12077 (BISH, RSA); S of Pu'u Ka'aumakua, St. John 20252 (BISH);

u Konàhuanui, E. brink, St. John 20315 ee Konahuanui, Topping 3223 (NY); [above Honolulu, 25 Sep 1840 — 5 Apr 1841 (fide

cm 1845),] U.S. Exploring Expedition s.n. (GH

6. Lobelia gloria-montis Rock, Monogr. Stud. Haw. Lobelioid.: 17. 1919. Lobelia gaudichaudii var. gloria-montis (Rock)

H. St. John & Hosaka, Occas. Pap. Bernice Pauahi Bishop Mus. 14:121. 1938. Galeatella gloria-montis (Rock) O. Deg. & I. Deg., Fl.

Hawaiiensis, Fam. 339. 1962. Type: HAWAIIAN ISLANDS. West Maur: Pu'u Kukui Mt., 5700 i aed 1910, Rock & Hammond 8209

(HOLOTYPE: BISH!; isotypes: A[4]! BISH[4]! GH! K! NY! OSH! US! W!). Thi

Hammond is not mentioned as co-collector on many of the isotypes.

I f pl. 58 as “Type;”

Plant 1.5-5 m tall. Stem 0.4—4 m tall x 1.5—10 cm diam.; cicatrices 3-5 mm tall x 6-13 mm wide, transversely

rhombic, the corners acute. Lamina 10-24 cm long x 1.3-3.8 cm wide, oblanceolate or narrowly oblong,

Lammers, Revision of Lobelia sect. Galeatella 805

glabrous or margin ciliate toward base; margin subrevolute or merely thickened; apex acute or acuminate;

base attenuate, cuneate, or obtuse. Inflorescence 1-1.5 m tall, unbranched or 5-7-branched, each branch

18-100-flowered and pyramidal in outline, its axis green, glabrous or pubescent; peduncle sparsely covered

with sterile bracts that rapidly decrease in size acropetally; sterile bracts 30-120 mm long x 10-25 mm wide,

oblanceolate, narrowly oblong, oblong, or elliptic, the apex acuminate or acute, the base cuneate; floral bracts

20-90 mm long x 6-16 mm wide, obovate, elliptic, lanceolate, oblong, or linear, the apex acute or acumi-

nate, the base cuneate; pedicels 15-60 mm long. Hypanthium 8-15 mm long x 6-17 mm diam., broadly

obconic or campanulate, '/.—'/, as long as the corolla, glabrous or pubescent; calyx lobes 7.535 mm long x

3.5-9 mm wide, narrowly triangular or lanceolate, ?4 as long to 31⁄2 times longer than the hypanthium, the

margin entire and sometimes ciliate, the apex acuminate or acute and sometimes apiculate. Corolla 67-103

mm long, white or rarely red, glabrous (rarely sparsely pubescent at apex of lobes); tube 26-45 mm long x

7-15 mm diam. at middle, suberect or slightly curved, expanding slightly towards the mouth, 2.2-5 times

longer than broad; dorsal lobes 35-70 mm long x 4-6 mm wide, 0.9-3.1 times as long as the tube; ventral

lip 30-52 mm long x 12-28 mm wide. Filament tube 58-82 mm long, decurved slightly, 415—614 times

longer than anther tube, glabrous; anther tube 4—6.5 mm diam.; dorsal anthers 11-16 mm long; ventral

anthers 8-13 mm long. Capsule 27-37 mm long x 10-18 mm diam.; seeds [Fig. 1F] 2-2.3 mm long x 1-1.2

mm wide, honey- or chestnut-brown.

Distribution, habitat, and phenology.—Known from Montane Wet Mixed Communities and Montane Wet

Forest (Gagné & Cuddihy 1990) on Maui and Moloka‘i, where two subspecies occur; see below for details.

6a. Lobelia gloria-montis subsp. gloria-montis

Lobelia gloria-montis var. molokaiensis O. Deg., Fl. Hawaiiensis, Fam. 339. 1938. Galeatella i ea montis var. ee. ud p (2:

Deg. & I. Deg., Fl. Hawaiiensis, Fam. 339. 1962. Type: HAWAIIAN ISLANDS. ^ g Par pali, g g g- and

windswept pali, 6 Jun 1928, Degener 7778 (HoLotyPE: NY!).

Lobelia gaudichaudii f. bryanii H. St. John & Hosaka, Occas. Pap. Bernice Pauahi Bishop Mus. 14:123. 1938. Galeatella gloria-montis var.

bryanii (H. St. John & Hosaka) O. Deg. & I. Deg., Fl. Hawaiiensis, Fam. 339. 1962. Tre: HAWAIIAN ISLANDS. West Maur: Puu

Kukui, May 1910, Forbes 68.M (HOLoTYPE: BISH!; isotypes: BISH! NSW! P! US! WI).

Lobelia ed f. kukuiensis H. St. John & Hosaka, Occas. Pap. Bernice Pauahi Bishop Mus. 14:123. 1938. Galeatella gloria-montis

kuiensis (H. St. John & Hosaka) O. Deg. & ., Fl. Hawaiiensis, Fam. 339. 1962. Type: HAWAIIAN ISLANDS. West Maul:

em au-Pu'u Kukui trail, brushy opening in rain a 4400 ft, 19 Dec 1928, Ewart 89 (HoLoTYPE: BISH!; isotype: BISH!).

Lobelia gaudichaudii f. sanguinea H. St. John & Hosaka, Occas. Pap. Bernice Pauahi Bishop Mus. 14:123. 1938. Galeatel

sanguinea (H. St. John & Hosaka) O. Deg. & I. Deg., Fl. Hawaiiensis, Fam. 339. 1962. Type: HAWAIIAN ISLANDS. West Maut: Pu‘u

Kukui, in swamp, 4500 ft, 16 Aug 1933, Hartt s.n. (HoLoTYPE: BISH!; isotypes: BISH[4]! US!).

Plant 1.5-2.4 m tall. Stem 0.4-1.5 m tall x 1.5-4 cm diam. Lamina 10-24 cm long x 1.3-3.8 cm wide,

oblanceolate or narrowly oblong. Inflorescence 1-1.5 m tall, unbranched (rarely with 1 or two clearly sub-

a gloria-montis f.

ordinate basal branches), 18-100-flowered; floral bracts 20-47 mm long x 6-16 mm wide, obovate, elliptic,

or lanceolate; pedicels 15-45 mm long. Hypanthium 8-15 mm long x 6-17 mm diam., broadly obconic or

campanulate, '/.—'/,, as long as the corolla, glabrous or rarely pubescent; calyx lobes 7.5-25 mm long x 3.5-9

mm wide, narrowly triangular or lanceolate, ?4 as long to 342 times longer than the hypanthium. Corolla

67-96 mm long, white or rarely red, glabrous (rarely sparsely pubescent at apex of lobes); tube 26-45 mm

long x 7-13 mm diam. at middle, suberect, 2.2—5 times longer than broad; dorsal lobes 35-70 mm long x

4—6 mm wide, 0.9-3.1 times as long as the tube; ventral lip 30-48 mm long x 12-28 mm wide. Filament

tube 58-80 mm long, 415-64 times longer than anther tube; anther tube 4—6.5 mm diam.; dorsal anthers

11-16 mm long; ventral anthers 8-13 mm long. Capsule 27-37 mm long x 10-15 mm diam.; seeds [Fig.

1F] 222.3 mm long x 1 mm wide, honey-brown.

Icones.—Hitchcock (1919), pl. 25-1; Rock (1919), pl. 15 [right], 17, 18; Degener (193823); Wimmer (1943),

fig. 11d; Wimmer (1953), fig. 104b [seed only]; Degener and Degener (1974), p. 2; Carlquist (1980), pp. 212C

[bottom], 245 [right]; Degener and Degener (1983), p. 456; Kepler (1983), p. 80 [upper right]; Sohmer and

Gustafson (1987), fig. 121; Lammers (1990), pl. 57 [upper right]. Figured specimen.—Rock (1919), pl. 58

[holotype of L. gloria-montis].

806 Journal of the Botanical R h Institute of Texas 1(2)

Distribution, habitat, and phenology.—Known primarily from Montane Wet Mixed Communities (Gagné

& Cuddihy 1990) on Pu'u Kukui, West Maui, at 915-1760 m (cf., Carlquist 1980, p. 352); sterile plants from

Montane Wet Shrublands (Gagné & Cuddihy 1990) on eastern Molokai, at 1165-1350 m are placed here as

well. Flowering July through December, fruit ripening August through May. Considered “apparently secure”

by Wagner et al. (1999).

Representative specimens: Moloka‘i: Papa‘ala pali, Anderson 513 (BISH); Kalua'aha Ahu., ridge E of Manawai Gulch, Warshauer & McEl-

downey 2390 (BISH). West Maui: Mountains, 3 Aug 1961, Bonsey & Suthers s.n. (MSC); Pu'u Kukui, Bryan 632 (BISH); Pu'u Kukui, Dec

1928, Bryan s.n. (BISED; below summit, Pu'u exi Carlquist 553 (RSA); Pu'u Kukui, Carlquist 555 (RSA); Pu'u Kukui, Carlquist 2142

(BISH, MICH, RSA); S of Puu Kukui summit, Char & Arakawa PKWM-55-76 (PTBG); Puu Kukui, Cranwell et al. 2667 (BISH, GB, K, S),

Pu'u Kukui, on road to Nakalaloa, Cranwell 2668 (BISH, GB, K, S); just below summit of Mt. ‘Eke on northwest side, Degener 7937 (B,

n CAS, GH, K, MICH, MO, NY, WIS), Degener 7938a (B, CU, K, MASS, MO, NSW, NY, US, WELO, Degener 7938b (BISH, NY); Puu

Kukui summit, Degener et al. 25023 pun ‘Eke, Forbes 456.M (BISH); Puu Kukui, Gillett 1793 (BISH); trail up to Puu Kukui, Gillett et al.

2444 (BISH); Keahikauo Bog, NE of ‘Eke Crater ifson 2020 (OS, RSA); Pu'u Kukui, Higashino 76-29 (BISH); summit of Mt. ‘Eke, 1870,

Hillebrand s.n. (B); ‘Eke, s.d., Hillebrand s.n. (C, GH, K, NSW, S, US); Putu Kukui, Hitchcock 14735 (US); Pu'u Kukui, near summit, Hitch-

cock 14836 (US); Pu'u Kukui, Hitchcock 14861 (US); from Violet Lake to near the summit, Hobdy 665 (BISH); top of mt., Mann & Brigham

462 (BISH, GH, K); top of mts., Mann & Brigham s.n. (US); ‘Eke trail, base of ‘Eke Crater, Medeiros 341 (BISH); Puu Kukui, McAlpin 635

(DUKE); Mt. Kukui, Munro 418 (BISH); Pu‘u Kukui, Munro 419 (BISH); Pu'u Kukui, Munro 615 (BISH, K); Pu'u Kukui, 16 Aug 1933, Neal

& Hartt s.n. (BISH); western headwaters of Kahakuloa Str. N of ‘Eke, Oppenheimer & Kunna H119910 (BISH); Violet Lake, Perlman et al.

6884 (MO, PTBG); Pu'u Kukui, Aug 1910, Roch s.n. (A, NSW, NY); summit of Pu'u Kukui, Aug 1910, Rock s.n. (NSW, NY); summit of Mt

‘Eke, Sep 1918, Rock & Hashimoto s.n. (BISH); Puu Kukui, summit bog, Skottsberg 773 (BISH, GB, S); Pu'u Kukui, St. John 10274 (BISH);

Pu'u Kukui, Tessene 1786 (WIS); Pu'u Kukui, Wilbur & Webster 834 (DUKE); Pu'u Kukui summit bogs, Wood 11505 (OSH, PTBG).

6b. Lobelia gloria-montis subsp. longibracteata (Rock) Lammers, stat. nov. Lobelia gaudichaudii var. longibracteata

Rock, Indig. Trees Haw. Isl.: 78. 1913. Lobelia gloria-montis var. longibracteata (Rock) Rock, Monogr. Stud. Haw. Lobelioid.: 119.

1919. Lobelia longibracteata (Rock) E. Wimm., Pflanzenr. IV.276c: 889. 1968. Galeatella gloria-montis var. longibracteata (Rock) O.

Deg. & I. Deg., Fl. Hawaiiensis, Fam. 339. 1962. Galeatella longibracteata (Rock) O. Deg. & I. Deg., Prodr. Galeatella & Neowim-

meria: 6. 1974. Tyre (designated by Rock po HAWAIIAN ISLANDS. West Maur: Mt. Pu'u Kukui, 5780 ft, Aug 1910, Rock 8818

(LECTOTYPE: BISH!; ISOLECTOTYPES: A! BISH! GH

se ee var. albiflora H. St. John & A.C. Medeiros, Phytologia 63:366. 1987. Type: HAWAIIAN ISLANDS. East Maur: Haleakala

itt on bog margin above mid amp bog , 4200 m" [ft], 13 Jun 1982, Medeiros 4 (HOLOTYPE: BISH! ).

Plant 2.4—5 m tall. Stem 2-4 m tall x 2-10 cm diam. Lamina 18-23 cm long x 1.8-4.3 cm wide, oblanceo-

late. Inflorescence 1-1.5 m tall, unbranched or 5-7-branched, each branch 40-80-flowered; floral bracts

60-90 mm long x 6-10 mm wide, oblong or linear; pedicels 38-60 mm long. Hypanthium 10-13 mm long

x 11-18 mm diam., broadly campanulate, 1/8 as long as the corolla, pubescent; calyx lobes 25-35 mm long

x 5-9 mm wide, narrowly triangular, 2-342 times longer than the hypanthium. Corolla 78-103 mm long,

white, glabrous; tube 28-44 mm long x 12-15 mm diam. at middle, slightly curved, 2.5-3 times longer

than broad; dorsal lobes 45-70 mm long x 5-6 mm wide, half again as long as the tube; ventral lip 40-52

mm long x 22-26 mm wide. Filament tube 68-82 mm long, 4.5—5 times longer than anther tube; anther

tube 4.5-6 mm diam .; dorsal anthers 15-17 mm long; ventral anthers 11-13 mm long. Capsule 33-36 mm

long x 14-18 mm diam.; seeds 2 mm long x 1.2 mm wide, chestnut-brown.

Figured specimen.—Rock (1919), pl. 59 [lectotype].

Distribution, habitat, and phenology.—Known only from Montane Wet Forest (Gagné & Cuddihy 1990)

on the rim of Tao Valley, West Maui, at 1740-1760 m; and on the northwestern slopes of Haleakala, East

Maui, at 1280-1700 m. Flowering from June to August.

Representative specimens: West Maui: Pu'u Kukui summit and extending to east bog and Helu Peak along the ridge line, Hobdy 664

BISH). East Maui: NE outer rift on margin of Mid Camp bog, Medeiros 2076 (BISH); S rim of Kipahulu Valley, Kuiki, Haleakala, St.

John & Mitchell 21237 (BISH).

ACKNOWLEDGMENTS

The curators and staffs of the following institutions are gratefully acknowledged for allowing study of speci-

mens in their care, either via loans or during personal visits: A, B, BH, BISH, C, CAS, CU, DUKE, F, GB,

GH, K, MASS, MICH, MO, MSC, NSW, NY, OS, OSH, P, POM, PTBG, RSA, S, UPS, US, W, WAB, WELC,

Lammers, Revision of Lobelia sect. Galeatella 807

WIS. Figure 1 was expertly prepared by Dr. Todd Kostman, Director of the Electron Microscopy Facility

at the University of Wisconsin Oshkosh. The helpful suggestions of manuscript reviewers Eric Knox and

David Lorence were much appreciated.

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A NEW SPECIES AND TWO NEW COMBINATIONS IN

POTENTILLA SECT. NIVEAE (ROSACEAE)

David F. Murray Reidar Elven

University of Alaska Museum of the North Natural History M |B ical /

7 Yukon Driv University of Oslo

Fairbanks, Alaska 99775-6960, U.S.A. PO. Box 1172 Blindern

ffdfm@uafedu NO-0318 Oslo, Norway

reidar.elven@nhm.uio.no

ABSTRACT

Potentilla holmgrenii D.F. Murray & Elven, sp. nov. is described and illustrated. Two new combinations are made: Potentilla arenosa

(Turcz.) Juz. subsp. chamissonis (Hultén) Elven € D.F. Murray and P. hyparctica Malte subsp. elatior (Abrom.) Elven € D.F. Murray.

RESUMEN

Se describe e ilustra porcus uns dns D.F. Murray & Elven, sp. nov.. Se hacen nue mbinaciones: Potentilla arenosa

(Turcz.) Juz. subsp. Elven & D.F. Murray y P. hyparctica Malte subsp. elatior (Abrom.) Elven & D.F. Murray.

While preparing a treatment of boreal and arctic Potentilla for Flora of North America it became necessary

to assess how well the names of widespread arctic plants have been applied to Rocky Mountain species. A

case in point is Potentilla nivea L., in the sense of Eriksen et al. (1999) not Soják (1989).

Whereas P. nivea is indeed found in the Rocky Mountain region, some plants so named are morphologi-

cally well beyond the limits accepted for the species throughout its circumpolar and boreal-alpine ranges.

A densely tufted plant of mountain summits in Nevada and Utah is distinct by its small, compact stature,

its ternate leaves with densely floccose petioles and laminae, and leaflets with fewer teeth directed forward

and having rounded apices. This new species presented below is named P. holmgrenii. It can be distinguished

from P. nivea by the key provided.

Basal leaves densely hairy on both surfaces; central leaflets with teeth 2-4 per side, apices obtuse

to rounded, directed forward, terminal tooth often orbicular and frequently overlapped by ad-

jacent pair of lateral teeth; epicalyx bractlets elliptic, shorter and narrower than sepals; styles

1 - mm or more in OR without eue basal prs sags holmgrenii

ely hairy on upper itl h 3-5 per side, apices acute, not direct

forward, ten tooth not orbicular, not overlapped by adjacent pair of teeth; epicalyx bractlets eee or

lanceolate, as wide as sepals at the base; styles less than 1.2 mm long, evenly tubular, with distinct basal

papillae Potentilla nivea

Potentilla holmgrenii D.E Murray & Elven, sp. nov. (Figs. 1-2). Tyee: NEVADA. Wurre PINE Co.: Mt. Moriah, 6 Jul 1966,

Holmgren and Reveal 2818 (HoLoTYPE: NY; isotypes BRY, UTC).

Planta pulvinata. C ] il f. d v f. I : £A EET

LE vel idi apicibus obtusis vel qu: Ad Potentillae niveae L. E sed "armis foliorum supra adpressi-tomentosis,

Plants perennial, forming dense cushions to 10 cm in diameter (Fig. 1) and occasionally small mats gray-

ish-white or white, eglandular; caudex woody, stout, strongly ramified with branches 0.4-0.6 cm thick,

densely sheathed with persistent, blackish-brown, indurate remains of stipules, petioles and some whole

leaves, giving the branches a compact, columnar structure; stems ascending, 2-8 cm, densely white-floccose;

T:

leaves basal and cauline (0-1); stipules broadly ovate, obtuse, light brown, proximally glabrous, distally

and abaxially with sparse, long, soft hairs, becoming subglabrous; petioles 0.5-2 cm, densely floccose;

leaflets three, margins slightly revolute; central leaflet broadly elliptic or obovate, 0.7-1.3 x 0.4-0.6 cm,

short-stipitate, base cuneate, incised half way to midvein, teeth 2-4 per side, broadly elliptic to obovate,

J. Bot. Res. Inst. Texas 1(2): 811 — 814. 2007

812

Journal of the Botanical R h Institute of Texas 1(2)

Fe. 1.C NT £ D, Ha hal

Fic. 2. Shoot with and leaf cluster. Note shape and

orientation of the teeth (Tiehm 9862, BRY 305045).

Scale bar =

d Reveal 2819, UTC 115752 isotype). Scale bar = 2 cm.

apices obtuse to rounded Fig.2), abaxial surface with very dense

tomentum, white or grayish-white floccose with some straight,

soft, verrucose, subappressed hairs ca. 1 mm long on veins, adaxial

surface gray, densely hairy with straight, silky, subappressed hairs

ca. 1-1.5 mm long; inflorescence simple or open, 1-2) flowers;

bracts lanceolate or ovate; pedicels straight, 0.4—0.7 cm in flower,

1.5-2 cm in fruit, densely floccose and mixed with a few straight,

subappressed to subpatent hairs; hypanthium 1.8-2.2 mm. with

|

dense, straight, 1 hairs and shorter stiff hairs ca. 0.7 mm

long; epicalyx bractlets narrowly elliptic, obtuse, margins revolute,

gray-green or reddish, 2-2.5 x 0.5-0.9 mm, shorter and narrower

than the sepals; sepals broadly lanceolate or triangular, subacute or

acute, gray-green, 3-3.5 mm, petals pale yellow, broadly obovate

or obcordate, retuse, + overlapping, 4-5 mm, longer than sepals;

stamens ca. 20; filaments 1-2 mm; anthers ca. 0.5-0.4 mm; carpels

ca. 30-40; styles tubular, 1.2-1.4 mm, thickened at base and with a

few, indistinct papillae, distinctly dilated beneath stigma; achenes

pale olive-green, glabrous, 1.1-1.4 x 1 mm.

Habitat.—On wind-swept ridges, fellfields, rocky slopes;

3300-4500 m.

Distribution.—Potentilla holmgrenii is known presently from

the Schell Creek and Snake ranges of White Pine County in eastern

Nevada, and the Deep Creek Range in adjacent Juab County in

western Utah.

Etymology.—The name was chosen to honor Noel H. Holmgren,

who has long been a student of the intermountain flora. In his treat-

AA y IEI „A . J he 4° Dat PRTI 813

ment of Potentilla for the Intermountain Flora (Holmgren 1997), Holmgren called attention to the compact

nature of the populations in the Snake and Schell Creek ranges but refrained from treating them as distinct

from P. nivea.

Specim xamined: NEVADA. White Pine Co.: North Schell Peak, 31 Jul 1989, Mosely 1592A (BYU, NY, UC); North Schell Peak, 3

Ju al 1982, Lavin 4208 (NY); Shell [sic.], Creek Mts., 13 Jul 1938, Pennell and Schaeffer, Jr. 22959 (NY); Taft Peak area, 30 Jun 1985, Tiehm

9862 (BRY, COLO, NY, RM, UTC,); Mt. Wheeler, 20 Jun 1928, Cottam 3303 (BRY); Wheeler Peak, 20 Jul 1941, McVaugh 6032 (UTC); Mt.

Wheeler, 19 Jul 1954, Langenheim s.n. (UC); Mt. Wheeler, 5 Aug 1985, Holmgren and Vincent 10970 (NY, UTC, UC); Mt. Wheeler, 3 Jul

1980, White and Neese 323 (BYU); NW shoulder Wheeler Peak, 9 Jul 1969, Lewis 1698 (RM); Wheeler Peak, 29 Jul 1976, Albee 3319 (NY);

Mt. Moriah, 2 Aug 1999, Niles et al. 5817 (RENO). UTAH. Juab Co.: Deep Cr. Mts., 13 Jul 1983, Goodrich 19024 (BRY, NY).

Additionally, our treatment of arctic Potentilla requires two new combinations that are presented below.

otentilla arenosa (Turcz.) Juz. is the common, widespread arctic and boreal-alpine species to which the

name P. hookeriana has been largely misapplied, with P. hookeriana sensu str. now restricted to the mountains

that differ only,

but significantly, in one character: the petiole hairs. Subspecies arenosa is characterized by petioles with two

of Colorado, Nevada, and Utah. Potentilla arenosa comprises two largely allopatric sul

r

layers of hairs: long and straight and short and stiff or curly. Subspecies chamissonis has petioles with only

long, straight, verrucose hairs.

Potentilla arenosa (Turcz.) Juz. subsp. chamissonis (Hultén) Elven & D.E Murray, comb. nov. Basioxvw. Po-

a chamissonis Hultén, Bot. Not. ad 140. 1945; P hooheriana Lehmann subsp. chami is (Hultén) Hultén; P nivea sensu

tenti

Soják (1989), non L.; P nivea L. subsp. chamissonis (Hultén) Hiitonen.

Habitat.—On cliffs, dry heath tundra, ridge tops.

Distribution.—southern Greenland, Man., N.W.T., Nunavut, Que., n. Europe.

We have not been able to confirm reports by Hultén (1968) of this sul

É from Alaska or the Yukon

Territory.

Southern arctic plants of Potentilla hyparctica Malte differ from the high arctic ones in several presum-

ably independent features. The two forms are sympatric in Greenland, Baffin Island, northern Alaska, and

Yukon Territory and appear there to remain distinct. We propose for the southern taxon the combination

P hyparctica subsp elatior.

Potentilla hyparctica Malte subsp. elatior (Abrom.) Elven & D.E Murray, comb. nov. Basionwm. Potentilla emarginata

Pursh var. elatior Abrom., Biblioth. Bot. 8(Heft 42, part 2):8. 1899; P hyparctica var. elatior (Abrom.) Fernald

Plants mostly 4—20 cm, sparsely hairy; leaflets broadly obovate, lobes broadly oblong, rounded or obtuse,

central leaflet long-stipitate (2-3 mm), base broadly cuneate or rounded; inflorescence 1-3(-5)-flowered;

epicalyx bractlets broadly oblong-ovate, 2-5 mm wide; sepals rounded or obtuse.

Habitat. —In moist ericaceous heath, snowpatch meadows, moist arctic and alpine tundra, rocky out-

crops, solifluction slopes, and talus.

Distribution.—Greenland; Alta., B.C., Nfld. and Labr., N.W.T., Nunavut, Que, Yukon; Alaska; ne.

Asia.

ACKNOWLEDGMENTS

Authors wish to thank the curators at BRY, COLO, NY, RENO, RM, UTC, UC for loans of specimens and

Barbara Ertter and James Reveal for their careful reading of the manuscript and suggestions for improve-

ment. Arnold Tiehm and an anonymous reviewer provided helpful comments. Thanks to Guy Nesom for

the Latin diagnosis and to Barbara Murray for creating the figures.

REFERENCES

ERIKSEN, B, B. JonseLL, and O. NiLssow. 1999. Proposal to conserve the name Potentilla nivea (Rosaceae) with a con-

served type. Taxon 48:165-166.

HOLMGREN, N.H. 1997. Rosaceae. In: A. Cronquist, N.H. Holmgren, and PK. Holmgren. Intermountain flora: vascular

814 Journal of the Botanical R h Institute of Texas 1(2)

plants ofthe Intermountain West, U.S.A. Vol. 3, Part A: Subclass Rosidae (except Fabales). The New York Botani-

cal Garden, Bronx. Pp. 64-158.

HuLTÉN, E. 1968. Flora of Alaska and neighboring territories. Stanford Univ. Press, Stanford.

SoJAk, J. 1989. Notes on Potentilla (Rosaceae). VIII. P nivea L. aggr. Candollea 44:741-762.

A NEW SPECIES OF ALOCASIA (ARACEAE: COLOCASIEAR FROM

PANAY ISLAND, PHILIPPINES

Melanie P. Medecilo George C. Yao

n Sciences Department Philippine Horticultural Society

LIE Las lle University- asmarinas ila S edli g Bank

pom Cavite 4115 Quezon City

PHILIPPINES PHILIPPINES

Domingo A. Madulid

you Division

nal Museum

PHILIPPINES

ABSTRACT

Alocasia nycteris Medecilo, Y & Madulid is d ibed ies f Aklan and Antique Panay Island Philippines The spe-

NAM A Te

ts bat-wing shaped blade and deeply undulated t bpi

o E E E

d leaf margins. The new species is most closely

similar to A. sanderiana W. Bull.

Key WoRps: Araceae, Philippines, Panay Island, endemic plant, threatened plant

RESUMEN

Se describe un isa ia Yao & PE bos nueva v nS y e ne. dn oa La especie se

La

a nueva especie

eM Y O

es Eon más AR a A. sanderiana W. Bull.

There are 20 genera and about 130 species belonging to the family Araceae in the Philippines. The largest

genus in the Philippines is Alocasia (Schott) G. Don with 14 species recorded so far (Hay 1999) but based on

==

recent field studies, the list increases to at least 18 species (Medecilo, in prep.). All species of Alocasia in the

Philippines are endemic, except A. macrorrhizos (L.) G. Don which is widespread throughout the Asian tropics

and never found away from human disturbance anywhere in its range. Alocasia has more than 75 species in

the Malesian region, Sri Lanka, Australia, southern China and the southernmost parts of Japan.

The most recent taxonomic revision on the genus in the country was done by Hay (1999). Twelve species

enumerated by Merrill (1925) was reduced to ten and four new species were added, namely: A. boyceana A.

Hay, A. clypeolata A. Hay, A. ramosii A. Hay, and A. scalprum A. Hay. The center of diversity of the genus is

Borneo with at least 21 species (Hay 1999, 2000) and the country is the second most species-rich sub region,

next to Borneo in the Malesian region. Some of the Philippine species have West Malesian and continental

Asian affinity. The conservation status and local endemicity was discussed in the revisionary work of Hay

(1999). Six Philippine species, namely: A. atropurpurea Engl., A. sinuata N.E. Br., A. portei Becc. & Engl.,

A. scalprum A. Hay, A. sanderiana Hort. ex. Bull, and A. clypeolata A. Hay were known from two or three

herbarium collections taken in the wild but most of the species are known from cultivation. Most species

are known in horticulture except A. atropurpurea Engl., and systematic measures for protection are desired.

Several species are of ornamental interest because of the appearance of the foliage leaves. A. sanderiana

Hort. ex. Bullis being used for hybrid utilization and reported as extinct in the wild. Other species include

A. micholitziana Sander, A. scalprum A. Hay, A. zebrina K. Koch & Veitch, A. sinuata N.E. Br., A. heterophylla

Merr. and A. ramosii A. Hay. Some of these species are being tissue cultured for commercial purposes.

J. Bot. Res. Inst. Texas 1(2): 815 — 818. 2007

816

J Infl

¿VU Mal OF

Das P |

te of Texas 1(2)

Fic. 1. Al

Jecilo, Yao & Madulid Sp. Nov. A. Habit: B. venation: C. Inflorescence: D. Pistils: E. N

F. Sy

ice); G. Infructescence. Scale: A,B,C and G = 2 cm; D-F = 3 mm.

Medecilo et al., A pecies of Al ia f the Philippines 817

Wen £1 £cl r Lal LA Iul J L

162. Close-uj i idely sagi l i Fic 3. Close-up view of infructescence.

natifid margins.

Alocasia nycteris Medecilo, Yao & Madulid, sp. nov. (Figs. 1-3). Tw: PHILIPPINES: Nabas, Aklan, Panay Island,

11°54'18"N and 121°59'43"E, 3 Nov 2005, Medecilo 397 (HoLotype: PNH; isotypes: DLSU, DLSU-D)

Alocasia sanderiana arte similis in foliis late sagittatis profunde undulatis, sed spadicis appendice longa, interstitiis

sterilibus irregulariter factis differt. |

Terrestrial robust herb to 1.5 m tall; stem erect to decumbent, up to 12 cm x 2 cm; Leaves 2-4 together,

subtended by cataphylls; cataphylls papery, up to 12 cm long by 2 cm wide at base, narrowly ovate, acumi-

nate; petiole 45-100 cm x 1.5-2.0 cm wide, green with blackish green streaks, sheathed from one-third

to one-quarter its length, marginally marcescent; blade hastate to sagittate, 20-37 cm x 35 cm wide at the

middle, coriaceous, dark glossy green adaxially, paler green abaxially, membranous when dry, margin deeply

undulated to subpinnatifid along margin; anterior lobe widely triangular or deltoid, acuminate at apex, 3-4

pairs of lateral veins nearly opposite diverging at 60—65? angle on each side of the anterior costa; axillary

glands conspicuous, brownish green, 0.1 mm diam.; secondary veins very fine, 3 mm apart arising from

the anterior costa and primary lateral veins then abruptly deflected to the margin not forming interprimary

collective veins; posterior lobes diverging at 85?—95? angle, up to 24 cm long; posterior costae naked in the

sinus for 3—4 cm, tip rounded, in some leaves forked, with 3 primary veins arising from the posterior costa;

inner side of the posterior lobe up to 4 cm wide at the middle, lanceolate. Inflorescence 1—2 together, sub-

tended by cataphylls; cataphyll whitish green, narrowly ovate, 10-12cm x 3-4 cm at the middle, mucronate

to 2 mm long, thick when fresh; peduncle up to 24 cm long x 1 cm wide at base, light green with grooves

at the base; spathe up to 10 cm; lower spathe tube ovoid, light green, 1.5cm x lcm diam.; limb narrowly

lanceolate, cucullate, pale yellow, tip inate, separated from the limb by a short deep constriction; spadix

shorter than the spathe, partly adnate to the lower spathe at 1.5 mm; female zone 10 mm long, 7 mm wide

at the middle; ovaries light green, subglobose, 3 mm in diam.; stigma 3-lobed; lobes, erect, 1 mm diam.;

style 0.5 mm long, brownish green; pistillodes reduced, irregular shape with holes in the center, 1-2mm

wide, whitish; sterile interstice 5 mm long x 4 mm wide, cylindric; whorl comprised of diamond-shaped to

rhombo-hexagonal synandrodes, 2 mm wide; male 14 x 6 mm, cylindric, whitish green; synandria irregular

to rhombo-hexagonal, slightly wavy at the edges, 1.5-2 mm diam., apical pores capped by synconnective;

appendix 40 mm long x 5mm wide, yellow, tapering. Infructescence oblong. Fruit a berry, globose, 10 mm

diam, orange when mature, 1—3 seeded. Seeds, greenish-black, globular, 2 mm in diam.

Distribution. —PHILIPPINES: Panay Island, Nabas to Ibajay, Aklan and Antique, 0-20 masl. Endemic.

818 Journal of the Botanical R h Institute of Texas 1(2)

Habitat. —Remnant lowland forests and secondary forests. Common on rocky areas. Prefers shaded

places along roadsides. It also grows in disturbed secondary forest near ricefields.

Specimens examined: PHILIPPINES. Aklan Prov.: Nabas, living collections of Ray Ong in San Juan, Metro Manila, 3 Nov 2005,

Medecilo 397; Medecilo 398 (DLSU-DH 701-702

Notes.—Alocasia nycteris resembles A. sanderiana W. Bull and A. portei Schott in having deeply undulated

to subpinnatifid blade margins. It differs from A. sanderiana by its larger, taller habit, not having peltate

leaves, the broadly triangular, hastate-sagittate blade, longer posterior lobes (in some individuals) and the

absence of a silver white midrib and primary lateral veins. The color of the leaves of A. nycteris is green on

both lower and upper surfaces while A. sanderiana is shiny, deep blackish green in the adaxial side and

purple in the abaxial surface.

The species described here was first introduced to the public by an aroid enthusiast, Antonio Advin-

cula during a garden show of the Philippine Horticultural Society, Inc. (PHSI) in 2003. George Yao, former

President of the PHSI featured the species in the International Aroid Society Newsletter Vol.5, No. 3 (Aug 2003)

and named it Alocasia advincula. The species name, however, could not be considered as validly published

in accordance with the International Code of Botanical Nomenclature (Greuter et al. 2000) as it lacking a

Latin diagnosis, and there is no rank indicated and no citation of the holotype.

Etymology.—The specific epithet ‘nycteris’ alludes to the bat-shaped leaves of this plant. Nycteris is a

genus of African and Asiatic bats comprising the hollow-faced bats. The species is known by its horticultural

name “Bat Alocasia.” It was recently introduced in the horticultural trade and is now becoming popular as

an indoor or pot plant. It can be easily propagated vegetatively by separation of the suckers and division

of the rhizomes. It is best grown under shade with adequate water supply. The plant is not only known in

local markets but also in the international markets.

Conservation status.—Plants are still commonly found along roadsides and in private lands near second-

ary forests. At present the populations along accessible places are under threat because of over collection

for horticultural purposes and from habitat destruction. No legal protection is given to the plant at present.

Thus, it is highly recommended that appropriate conservation measures are imposed to prevent rapid decline

of population. Mass propagation of the plant by tissue culture is an alternative to collection of the plants in

the wild. No populational studies have been made and information is lacking about the biology, phenology,

pollination and reproductive biology of the species.

ACKNOWLEDGMENTS

The authors would like to thank the Philippine Horticultural Society, Inc. (PHSD for funding the collection

trip of Melanie Medecilo in Aklan. Thanks to Ray Ong for providing valuable information on the species

in Panay. The Abellar family provided accommodation and warm hospitality to Medecilo in Aklan. Peter

Boyce is acknowledged for his suggestion on the specific epithet and for commenting on the early draft of

the manuscript. Jeff Veldkamp is acknowledged for the Latin diagnosis. Nemesio Diego, Jr., artist from the

Botany Division, National Museum rendered the illustration. We are very grateful for the comments of Tom

Croat and Peter Boyce. We also wish to thank Marlon C. Pareja, Chairman of the Biological Sciences Depart-

ment, De La Salle University-Dasmariñas, Dasmariñas, Cavite and Director Corazon S. Alvina, National

Museum for their constant support and encouragement.

REFERENCES

GREUTER, W, J. McNeiLL, F.R. Barre, H.M. Bunper, V. DEMOULIN, T.S. FituEiRAs, D.H. Nicotson, P.C. Sit vA J.E. Skoc, P. TREHANE,

N.J. TurLAND, and D.L. HAwKswortH (eds). 2000. International code of botanical nomenclature (St. Louis Code).

Regnum Veg. 131. Koeltz Scientific Books, Konegstein. http://www.bgbm.fuberlin.de/iapt/nomenclature/

code/SaintLouis/0000St. Luistile.htm

Hay, A. 1999. The genus Alocasia (Araceae-Colocasieae) in the Philippines. Gard. Bull. Singapore 51:1-41.

Hay, A. 2000. Alocasia nebula. Bot. Mag., n.s. 17(1):14—18, pl. 381.

Mebecito, M.P in prep. Monograph of the Genus Alocasia in the Philippines.

FOLIA TAXONOMICA 2. NEW SPECIES OF PASSIFLORA

SUBGENUS PASSIFLORA (PASSIFLORACEAE) FROM THE GUIANAS

Christian Feuillet

Department of Botany, MRC-166

Smithsonian Institution, PO. Box 3712

Washington, D.C. 20013-7012, U.S.A.

feuillec@si.edu

ABSTRACT

Two new species of uoc re d with ps flowers are T Vids the Guianas. Du new an poe in subgenus

d fro

om P. coccinea

Į

row leaves and bracts. P compen from Guyana and French Guiana | j leaves wi ntinuoush

its

glandula dL ith P. coccinea

an 1 1 1 1

E o

B

A key to p persecti tone is edad:

RÉSUMÉ

D 11 peces de Passiflora (Passifl d à fl dé Les nouvelles espéces e au

sous-genre Passifl Coccinea. Passiflora arta de Guyana est facile à distinguer de P. coccinea grâce à ses feuilles et ses

bractées étroites. Passiflora compar de Guyana et de o a t feuilles p i in la marge n'est pas sandals de eo

continue. La couronne et operculum des d P. coccinea. U lé

E E

la supersection C

Passiflora L. is the largest genus of the Passifloraceae with around 530 described species (Feuillet & Mac-

Dougal 2007). Although members of the genus Passiflora L. are usually very visible and well collected, more

than 27 species of passionflowers have been added to the flora of the Guianas since Killip (1937 & 1938)

who mentioned the presence in the Guianas of 33 species. Including the two new species described below,

this is an 8296 increase. Although they have bright red flowers, the two species described herein have been

overlooked for a long time because they are uncommon and close enough to the most common species in the

Guianas, P. coccinea Aubl., to have been confused with it and rarely collected. The two new species belong

in subg. Passiflora supersect. Coccinea Feuillet & J.M. MacDougal.

Passiflora supersection Coccinea includes 13 South American species, two of them expanding into Central

America. Traditionally these species have been included in subg. Distephana (Juss. ex DC.) Killip (now a

supersection). They are here separated from Distephana on the base of a series of morphological characters.

The leaf blades are hairy, glandular at margin, dentate or serrate or crenate. The bracts are foliaceous, usu-

ally red. The erect and compact corona is much smaller than in supersect. Passiflora, Stipulata Feuillet €

J.M. MacDougal and Laurifolia (Cervi) Feuillet & J. M. MacDougal, but larger than the reduced corona of

supersect. Distephana (DC.) Feuillet & J. M. MacDougal and Tacsonia (Juss.) Feuillet & J[.M. MacDougal.

Passiflora arta Feuillet, sp. nov. (Fig. LA-C). Wee: GUYANA. U. Takutu - U. EsstQUIBO: Acarai M

of unnamed peak, 6 km S of Sipu River, 1?22'N, 58°56'W, 700 m, fl., 30 Aug 1998, H.D. Clarke 7230 roue US).

LES T T4 1

Passiflora arta in subg. xa ora supersect. i P. llipticis, corona uniseriata,

fer

operculo longiore dif

Liana probably reaching the canopy. Vegetative parts with short rufous trichomes. Young stems terete, stri-

ated. Tendrils long, base strong and straight. Vegetative bud although part of the axillary complex (leaves,

stipules, pedicel, tendril), clearly in apical position on the stem, 5-7 mm away from the petiole, prophylls

acute at apex. Stipule linear, 6 x 0.2 mm, soon deciduous. Leaves: petiole 5-13 mm long, with 2 lateral

glands at the base; glands sessile, round, ca. 1 mm in diam., glabrous; blade narrow-elliptic, 7-7.5 x 1.2-1.5

cm, acuminate, mucronate at the apex, rounded to acute at the base, the margin shortly glandular serrate

J. Bot. Res. Inst. Texas 1(2): 819 — 825. 2007

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from Feuillet 1423 (US).

section; D-G. P

Feuillet, N e £D Ora | è 4l Cart 821

to bi-serrate, drying olivaceous adaxially and yellow-brown abaxially, 4-6 main lateral veins on each side

of the midrib, tertiary venation reticulate. Inflorescence sessile, 1-flowered. Pedicels 11 cm long, with short

pubescence. Bracts three, verticillate at 2.5 mm below the flower, pink red, narrow-elliptic, 5-5.5 x 1-1.3

cm, free to base, acuminate at the apex, glandular crenate to loosely serrate at the margin, with short scat-

tered trichomes on both surfaces. Flowers pink red: hypanthium 9 mm long, round at the base, then short-

cylindric, short pubescent outside, glabrous inside; sepals narrow-oblong, 3-3.5 cm long plus a 9 mm long,

subapical awn, 3-7 mm broad, short pubescent in the parts exposed in bud, otherwise glabrous; petals

similar to the sepals, thinner, not awned, glabrous; corona in 1 row of filaments 8-12 mm long, subulate,

erect, slightly leaning against the androgynophore, linked ina 1.5 mm long membrane at the base, glabrous;

operculum borne at the top of the hypanthium, membranous, shortly laciniate at the margin, 9-10 mm

long, dependent to the base of the hypanthium, then shortly recurved inward, glabrous; disc annular at the

bottom of the hypanthium; limen tubular membranous, dentate at the margin, 2 mm long, surrounding the

base of the androgynophore, glabrous; androgynophore cylindric, ca. 2 cm long, glabrous; stamen glabrous,

filaments flat, 13-15 x 2 mm, joint for 2-3 mm in a membrane at base, anthers dorsifixed, rectangular, 7

x 2 mm; gynophore shorter than the membrane at the base of the stamens; ovary 3-3.5 mm long, 2-2.5

mm in diam., subglobose, densely pubescent; styles 10-12 mm long, glabrous; stigmas ca. 2.5 x 3.5 mm,

glabrous. Fruit not seen.

Distribution.—Passiflora arta is known only from the type locality, Acarai Mts., Upper Takutu - Upper

Essequibo region in Guyana. It inhabits a dense forest on brown sand at 700 m. The forest was dominated

by Clusia L., Oenocarpus Mart., and Palicourea Aubl. species.

Because of leaves with serrate margin, bracts with glandular-serrate margin, red bracts and perianth,

short corona filaments, straight, and forming a cone appressed to the androgynophore, Passiflora arta sp.

nov. clearly belong in P. supersect. Coccinea. In this group it is close to P. coccinea (figure 1 D-G; plate 105a

in Feuillet 2002b) which has a large Amazonian and Guianan distribution. The Guianan P. arta has nar-

row-elliptic rather than ovate leaves and bracts, a uniseriate rather than triseriate corona, and an operculum

dependent nearly to the bottom rather than to the middle of the hypanthium. Differences with other species

of Passiflora supersect. Coccinea are summed up in the key below. The flower measurements have been done

on the dried flower of the holotype.

Etymology.—The Latin specific epithet means *narrow" (street) or “thin” (person). It refers to the narrow

leaves and bracts of the new species as compared to the sympatric P. coccinea.

Passiflora compar Feuillet, Sp. nov. (Figs. 2, 3). Tee: GUYANA. PoTARO-SIPARUNI: 1-3 km NNE of Kato along trail to Para-

makatoi, 4?0'N, 59°49’W, 675 m, 19 Mar 1989, fl. & fr., LJ. Gillespie 873 (HoLotypE: US (f1.); isotypes: BRIT, MO, K, NY, US (fr.)).

Passiflora compar in subg. do dud EE Coccinea P P. coccineae affinis; foliis subtiliter biserratis, tubo floris breviore,

corona biseriata, prima serie | ti, operculo vix breviore quam tubum differt

Liana 8 m tall (de Granville 12498), m reaching the canopy. Vegetative parts with short white or rufous

trichomes. Young stems terete to subangular, striated. Tendrils long, base thin and straight. Vegetative bud

although part of the axillary complex (leaves, stipules, pedicel, tendril), clearly in apical position on the

stem, 5-7 mm away from the petiole, with 6 long acuminate prophylls. Stipule filiform, 8-12 x 0.2 mm,

dentate-glandular at margin, soon deciduous. Leaves: petiole 5-13 mm long, with 2 lateral glands at the

base, one petiole (of de Granville 12498) with a pair of smaller glands at the apex; glands sessile, round, ca.

1 mm in diam., glabrous; blade oblong, 3-7 x 1.5-3 cm, acuminate, cuspidate at the apex, rounded at the

base, the margin shortly glandular serrate to bi-serrate, drying olivaceous adaxially and yellow-brown ab-

axially, 3-5 main lateral veins on each side of the midrib, tertiary venation reticulate. Inflorescence sessile,

1-flowered. Pedicels 5-9 cm long, with short pubescence. Bracts three, verticillate at 2-3 mm below the

flower, pink red or orange red, oval, 4.5—5.5 x 2-3 cm, free to base, rounded at the apex, loosely glandular

crenate at the margin to glandular serrate at the apex, with short scattered trichomes on both surfaces. Flow-

ers: hypanthium 7-8 mm long, round at the base, then short-cylindric, short pubescent outside, glabrous

inside; sepals narrow-oblong, 2.5 cm long plus a 7 mm long, subapical awn, 8 mm broad, “pale red” (sic!)

822 Journal of the Botanical R h Institute of Texas 1(2)

Cathy faaqale-

2003

Fic. 2 P, 77) AC uhi 14

stipule, tendril, pedicel, and vegetative

| ain: C. Node with I tiole, a

ion. A-B from Jansen-Jacobs et al. 1262 (US); C: from de Granville 12498 (US); D-E from Jansen-

P

bud: D. Flower, face view: E. F

Jacobs et al. 1262 (US) & photograph by C. Feuillet.

Feuillet, N e £D Ora | è 4l Cart 823

Fic. 3. Passifl par Feuillet, Gillespie 873 (photo L. Gillespie).

adaxially, red abaxially, short pubescent in the parts exposed in bud, otherwise glabrous; petals similar to

the sepals, thinner, not awned, red, glabrous; corona in 2 rows of filaments, filaments 5-7 mm long, outer

subulate, inner filiform, red or pink, erect, slightly leaning against the androgynophore, free, glabrous;

operculum membranous, shortly laciniate at the margin, 8-9 mm long, borne at the top of the hypanthium,

dependent to the base of the hypanthium, then long recurved inward, glabrous; disc annular at the bottom

of the hypanthium; limen tubular membranous, upper third laciniate, 5 mm long, surrounding the base

of the androgynophore, glabrous; androgynophore cylindric, ca. 1.-2 cm long, glabrous; stamen glabrous,

filaments flat, 12-15 x 1 mm, joint for 3-4 mm in a membrane at base, anthers dorsifixed, rectangular, 5-6

x 2.5 mm, green, pollen yellow; gynophore shorter than the membrane at the base of the stamens; ovary 5

mm long, 3 mm in diam., subglobose, densely pubescent; styles 10-12 mm long, thin, red or rose with red

spots, with scattered trichomes; stigmas ca. 2.5 x 2 mm, yellow, glabrous. Fruit not seen.

Distribution.—Passiflora compar is known in Guyana from the Kanuku Mts. (Upper Takutu-Upper Es-

sequibo) and near Kato and in French Guiana from the upper Maroni basin, the Saúl area, and the hills SW

of Cayenne. It inhabits moist forest below 400 m of elevation, near rivers or on slopes.

Because of serrate leaf margins, bracts with glandular-serrate margin, red bracts and perianth, short

corona filaments, straight, and forming a cone appressed to the androgynophore, Passiflora compar sp. nov.

clearly belong in P. supersect. Coccinea. In this group it is close to P. coccinea (Fig. 1 D-G) which has a large

Amazonian and Guianan distribution. The Guianan P. compar has thin rather than leathery or rubbery leaves

and bracts, a corona in two rows rather than three, and an operculum dependent nearly to the bottom rather

than to the middle of the hypanthium. Differences with other species of Passiflora supersect. Coccinea are

summed up in the key below.

Etymology.—The Latin noun in apposition, “compar” means “equal, companion” and refers to the second

row of corona filaments nearly as long as the first row, and to the fact that this new species stands close to

P. coccinea.

824

£L Dag *..1D

Journal UI

h Institute of Texas 1(2)

PARATYPES: GUYANA. U. Takutu - U. Essequibo: Kanuku Mts., 3°08’N, 59°23’W, 280—400 m, fl. & fr., 20 Feb 1985, MJ. Jansen-Jacobs,

C. Feuillet, P. Hiepko, L.E. Skog & B.J.H. ter Welle 330 (B, BBS, BRG, CAY, K, P, NY, U, US); 3°21’N, 59?29"W, 140-320 m, 26 Nov 1987

Jansen-Jacobs et al. 1262 (BRG, U, US). FRENCH GUIANA: Bassin de la Litani (= upper Maroni), Saut Lavaud, 2?45'N, 54?14"W, 140m

15 Sep. 1994, de Granville 12498 (CAY, US).

KEY LOCTHE:SPEGIES-OFP.PASSTEELORA-SUBERSECTz ‘COCGINEA

1. Bracts linear, less than 5 mm wide.

2. Peduncle 10-15 cm long; petals white, corona 3 ranked, the 2 outmost of filaments free to ba

se

ea Wurdack

2. Peduncle ii to 5 cm long; petals pink or red, corona 2 ranked, both fused in a membrane half t

third of the w

. Bracts ues de cong lanceolate, mostly more than 5 mm wide.

3. Leaves always unlobe

HR Hu ER Rodschied

4. S nd crenate

5. Operculum erect

P. aimae Annonay & Feuillet

5. Operculum decumbent P. longicuspis Vanderplank

4. Leaf margin viable or serrate

6. Stipules fimbria P. callimorpha Harms

6. E dus serrate or linear.

7. Operculum filamentose.

8. Corona with 4 rows of filaments P. quadrifaria Vanderplank

8. Corona with 1 or 2 rows of filaments.

9. Corona with two rows of subulate filaments P. araujoi Sacco

9. Corona with one row of pad filaments P. tholozanii Sacco

7. Operculum membranous, fimbria

10. Corona with one row of filam P. arta Feuillet

10. Corona with two or three rows M ues

11. Corona with two rows of filaments subequal in length P. compar Feuillet

11. Corona with three rows of filaments.

12. Outermost corona dee erect, slightly diverging, dark red to violet-black

miniata Vanderplank

12. Outermost corona filaments erect, converging around the androgynophore, white

red.

13. Outermost corona filaments red

P. sp. (Guyana)

13. Outermost corona filaments white P. coccinea Aubl.

3. Leaves 3-5-lobed.

14. Bracts fimbriate P. involucrata (Mast) Gentry

14. Bracts entire, dentate, or serrate.

15. Bracts more than 10 mm wide.

16. Stipules 3-5 mm long; third row of the corona tubular

16. Stipules 6-15 mm long; third row of the corona filamentose

. P vitifolia var. bracteosa (Karsten) Killip

, margaritae Sacco

1:5: ae less than 10 mm wide.

7. Stipules incised-serrate P. buchtienii Killip

if p. cl serrate.

Outer rank of the corona red; operculum fimbriate

P. vitifolia Kunth

RM . P. speciosa Gardner

e Outer a corona white; operculum crenate or denticulate

ACKNOWLEDGMENTS

I would like to thank the curators of the herbaria CAY and U for lending the material in their care. I am

grateful to Cathy Pasquale-Johnson for the fine illustrations and to three reviewers who made this paper

better through their corrections and suggestions. As always, discussions with John MacDougal have been

stimulating. This is number 83 in the Smithsonian's Biological Diversity of the Guiana Shield Program

publication series.

REFERENCES

FeuiLLeT C. 2002b. Passifloraceae. In: S.A. Mori, G. Cremers, C. Gracie, J.-J. de Granville, S.V. Heald, M. Hoff, and J.D

Feuillet, New species of Passiflora from the Gui 825

Mitchell, eds. Guide to the vascular plants of central French Guiana. Part 2. Dicotyledons. Mem. New York

Bot. Gard. 76:566-570.

FeuiLLET, C. and J.M. MacDoucaL. 2003 [May 2004]. A new infrageneric classification of Passiflora L. (Passifloraceae).

Passiflora 13(2):34-38.

Feuittet, C. and J.M. MacDoucat. 2007 [Dec. 2006]. Passifloraceae. In: K. Kubitzki, ed. The families and genera of

vascular plants IX:270-281. Springer, Berlin.

HANSEN, A.K., L.E. GILBERT, B.B. Simpson, S.R. Downie, A.C. Cervi, and R.K. Jansen. 2006. Phylogenetic relashionships and

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

Kiur, E.P. 1937. Passifloraceae. In A.A. Pulle, ed. Flora of Suriname 3(1):306-327.

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

MacDouca. J.M. and C. Feuittet. 2004. Systematics. In: T. Ulmer and J.M. MacDougal, Passiflora: Passionflowers of

the World. Timber Press. Pp. 27-31.

826 Journal of the Botanical R h Institute of Texas 1(2)

BOOK REVIEWS

Thomas G. Lammers. 2007. World Checklist and Bibliography of Campanulaceae. (ISBN 978-1-84246-

186-0, pbk.). Royal Botanic Gardens, Kew, Richmond, Surrey, TWO 3AB, UK. (Orders: www.kew.org,

+44 (0)20-8332-5646 fax). $185.55, 675 pp., 8 1/4" x 11 5/8".

From the 67 species of Adenophora to the two species of Zeugandra, Thomas Lammers seems to cover all the known genera, species,

nde: and nee iu in the pu daa nu As science goes, no sooner s the oun is UE there are taxa

montis subsp. longibracteata, Lobeli 1 Lobelia wahiawa)

E

in this issue of J. Bot. Res. Inst. Texas d aE 5 789- d

1

n easily con. fth Campanulaceae “Eighty -four g d encompassing

e e

2319 2 391 subspecies and 27 pL Mad another 8135 names are ies as Mid j D. checklist is alphabetical by

the “checklist”

d SESS for the pm Not sure e what subfamily a eons bela dne t pe that m each genus ais n the number o

I g n known, and additional notes. The bibliography in and

of teal is quite a contribution The author says it is selective rather than ee a it’s pretty exhausting just to read through the

11 pages of references under the eae Campanula or the 12 pages provided for the B s ia. The checklist includes every validly

| Baa Tense PIRR |

at or below specific rank, as well as all validly published g place and date of

EM Nomenclatural types are provided for each generic name. The format of tl tire checklist i ht to

follow and find. Thank goodness!

Lord I didn't know there were so many genera in the Campanulaceae; I'll bet the Good Lord was surprised to see so many names

on the list as well. A useful work on an important worldwide family.— Barney Lipscomb, Botanical Research Institute of Texas, 509 Pecan

Street, Fort Worth, TX 76102-4060, U.S.A.

Rick Darke. 2007. The Encyclopedia of Grasses For Livable Landscapes. (ISBN 978-0-88192-817-4,

hbk.). Timber Press Inc, 133 S.W. Second Avenue, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders:

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J. Bot. Res. Inst. Texas 1(2): 826. 2007

CONTRIBUTIONS TO THE GENUS FESTUCA (POACEAE: POEAE) IN MEXICO ANI

A KEY TO THE CENTRAL AMERICAN SPECIES

Stephen J. Darbyshire

Agriculture and Argi-Food Canada

Eastern Cereal and Oilseed Research Centre

Central a Farm, Wm. Saunders Bldg. #49

wa, Ontario, KIA 0C6, CANADA.

ABSTRACT

new species, Festuca longiligula (Poaceae: Poeae), is described from the state of Coahuila, Mexico. Names and descriptions are

aA for two nothospecies from central Mexico, Festuca meonzaler: Lanne and F. crece putative RA OLE:

hil ] F. livida x F. tolucensis, respectively. O

Hel leria) TE are presented. A key to the |

ee x F.

pecies of Festuca in Central America T ——

Key Wonps: Festuca, new species, Mexico, Central America, hybridization, nothospecies, subgenus Helleria, keys

RESUMEN

Sed il ie, F P P delestado de Coahuila, México. Se d 1 1 p

dos nothospecies de exito central, A «gonzalez- Eme F. xmiscella, híbridos 1 de F. livida x F. hepl phila y F.

livida x F. tolucensi S t TUE. Pu ;

i i g

Helleria) livida. Se aport | de | peci idas de Festuca en América Central.

The pioneering work of E.B. Alexeev on the taxonomy of Festuca in Mexico and Central America (Alexeev

1980, 1984) set the ground work for further detailed investigations on the genus. Not all of his described taxa

have been widely accepted (cf. Soreng et al. 2003) and some await further study. A working list of species

is given in Table 1. This paper describes new species (including nothospecies), contributes some additional

observations and presents an updated key to the species known from Mexico and Central America.

Festuca longiligula S.J. Darbyshire, sp. nov. (Figs. 1, 2A, 2B, 3A) Tre: MEXICO. Coanuia: SO de Saltillo, ceja y ladera

S de Sierra La Viga, subalpina, 3700 msnm, 22 Aug 1986, [A.] McDonald 2102 (HoLotypE: TEX: isotype: WIS, XAL (non vidi)

Plantae perennes caespitosae, innovationibus intravaginalibus praeditae. Culmi 70-90 cm alti, leves. Vaginae foliorum ima basi tan-

tum integrae, leves vel scabrae; vaginae foliorum emortuorum in fibras longitudinales non solutae; ligulae scariosae, (2-)5-9.5 mm

longae, laceratae; laminae foliorum innovationum conduplicatae, 0.8-1.5 mm diametri, abaxialiter scabrae, adaxialiter hirsutae, cum

9-11 nervis, 7-11-costatae, ug qon de abaxialiter has continuo i d ao I fasciculis cum venas

g BOGE j KO

Paniculae 10-22 cm longae,

S laxae, axe distali et ramulis er Spiculae (6-)7.5-13 mm longae, ovatae ad ellipsiodeae, cum 4-7 flosculis; glumae

ovatae ad lanceolatae, scariosae vel margine late scariosae, glabrae vel pleurumque scabrosae ad hispidulae dimidi ti; glumae

inferiores 3.4—5.5(—5.7) mm longae, cu nervis; glumae superiores 4.5-6.5 mm longae, cum (123(-5) nervis; ice usque

1.5 mm longae, hispidulae unilateraliter; lemmatae (5-)6-7 mm longae, lanceolatae ad anguste ovatae, be vel apice scabra ad

hispidula, cum = a nervis Ms apie late scariosa, obtusa ad abrupte acuta, interdum emarginata

5m

ad plus minusve bidentata,

longa

I 1ga; paleae 5 m longae, apice bidentatae, inter venas et saepe in

marginalibus distalibus hispidulae, venae EN ciliatae prope apicem ad 1/2; T. 2.5-3.3 mm longae; ovarium apex sparsim

o

hispidulus. Caryopses 3.5-4.5 x

Plants perennial, caespitose, shoots intravaginal. Culms erect or slightly decumbent at the base, 70-90 cm

tall, smooth and glabrous. Leaf sheaths with the margins closed only at the base, smooth and glabrous or

sometimes scabrous, not decaying into longitudinal fibres; auricles absent; ligules scarious, (2-)5-9.5 mm

long, lacerate; leaf blades of vegetative shoots conduplicate, 0.8-1.5 mm in diameter, abaxially scabrous,

adaxially hirsute, with 9-11 veins and 7-11 adaxial ribs; sclerenchyma in a continuous abaxial band, fas-

cicles joining the major veins with abaxial epidermis and with adaxial fascicles opposite veins sometimes

joining veins to form girders (Fig. 2); bulliform cells absent or inconspicuous. Inflorescence paniculate,

J. Bot. Res. Inst. Texas 1(2): 827 — 840. 2007

£L Dag *..1D

hi titit

f Texas 1(2)

828 Journal of

TABLE 1. Sy f the genus A M | [A ll | ( ly from |

Species Mexico (States) Central America

(other countries)

F. aequipaleata E. Fourn., Biol. Centr-Amer. Veracruz

3(20):581. 1885

F. aguana E.B. Alexeev, Bot. Zhurn. (Moscow &

| d) 66:1493, f. 1.

L ningrad) 1981.

Rupr., Bull. Acad. Roy. Sci. Bruxelles

9236. 1842.

F. arundinacea Schreb, Spicil. Fl. Lips.:57, 1771

E aperea El p Alexeev, Bot. Zhurn. (Moscow &

1409

|

PCA

F. bajacalforniana Gonz.-Led. & S.D. Koch, Novon

4:28-30, f. 3. 1

F. beamanii 3 a Dec Bot. Zhurn. (Moscow &

0-1501, f. 2. 1981

12V, ft.

E.B. e Bot. Zhurn. (Moscow &

=>

Leningrad) 66:1496, f. 1.1981.

F. breviglumis Swallen, Contr. U.S. Natl. Herb.

29:398. 1

F. callosa (Piper) St. Yves, Candollea 2:291. 1925.

F. cartagana E.B. Alexeev, Bot. Zhurn. (Moscow &

Leningrad) 67:1291-1292, f. 1. 1982

F. a eat Swallen, Ann. Missouri Bot. Gard.

30:116. 1943.

F. is Gonz.-Led. & S.D. Koch, Novon 4:

27-27,£.2.1

F. diclina SEN Novon 5:129, f. 1, 2. 1995.

F. filiformis Pourret, Hist. & Mém. Rea Roy. Sci.

Toulouse 3:319. 1788.

k dab d Darbys

E.B. Alexeev, Bot

Ler :1498 B

[= EEG ?]

F. hephaestophila Steud., Syn. Pl. Glumac.

1:310. 1854

F. herrerae Davidse, Novon 2:322-324, f. 1. 1992.

F. hintoniana E.B. Alexeev, Bot. Zhurn. (Moscow &

Leningrad) 67:1292, f. 1. 1982.

F. jaliscana E.B. Alexeev, Bot. Zhurn. (Moscow &

Leningrad) 66:1493-1495, f. 1. 1981.

F. ligulata Swallen, Amer. J. Bot. 19:436, f. 1. 1932.

F. livida (Kunth) puta Syst. Veg. 1:353. 1825.

F. longiligula Darbys

F. ads (E. Fourn.) JA ol., Bol. Soc. Bot. Mexico

23:165. 1958

a (Moscow &

J

| 1 d)

Hl

F. xmiscella Darbysh.

is eps E.B. Alexeev, Bot. Zhur. 66:

—-1498, f. 2. 1981.

E ovina L., Sp. Pl 1:73-74. 1

F. panamica E.B. Alexeev, " Zhurn. (Moscow &

Leningrad) 67:1290-1291, f. 1. 1982.

Chiapas Distrito Federal Durango

Guerrero, Hidalgo, Jalisco, México,

Michoacán, Morelos, Nuevo León,

Puebla, Tlaxcala, Veracruz

México, Veracruz

México

Baja California

Veracruz

Jalisco, Michoacán, Morelos, Oaxaca

Oaxaca, Puebla

Coahuila

Chihuahua

México, Tlaxcala, Veracruz

México

México, Nuevo León, Puebla, Tamaulipas,

Tlaxcala, Veracruz

Coahuila, Nuevo León

Jalisco

oahuila

México, Puebla, Tlaxcala, Veracruz

Coahuila

Chiapas, Chihuahua, Distrito Federal,

Hidalgo, Jalisco, México, Michoacán,

Morelos, Oaxaca, Puebla, San Luis Potosí,

Tamaulipas, Veracruz

México

Distrito Federal, Durango, Jalisco, México

Puebla, Veracruz

Guatemala, Panama

Costa Rica, Guatemala,

Costa Rica, Guatemala

Guatemala

Costa Rica, Guatemala,

Panama

Costa Rica, Panama

Costa Rica, Panama

Costa Rica

Guatemala

Costa Rica, Guatemala

Costa Rica, Panama

Honduras

Costa Rica

Panama

Darbyshire, F t in Mavi Val to tl PS Y | . k 829

Taste 1. continued.

Species Mexico (States) Central America

(other countries)

F. pringlei St. Yves, Candollea 2:305-307, f. 53. 1925. Chihuahua, Durango, Jalisco,

San Luis Potosí

F. roblensis Gonz.-Led., Novon 8:147-149, f. 1. 1998. Guanajuato, Zacatecas

F. rosei Piper, Contr. U.S. Natl. Herb. 10:45. 1906. Distrito pa ee Jalisco, México, Guatemala

Michoacan, Morelos, Tlaxcala, Veracruz

E rubra L, Sp. Pl. 1 753. Chihuahua, México, Nuevo León Costa Rica

zedowskiana E E oim Bot. Zhurn. (Moscow & México

pros 66:1500, f. 2. 1981

F. subverticillata (Pers) E.B. Alexeev, Novosti Sist. based on a probable label error

cae t. 17:52, f. 2. 1980.

FUE Pr Alexeev, Bot. ps (Moscow & Panama

Leningrad) 66:1495-1496, f. 1. 1981.

F. talamancensis Davidse, oo 2:324, f. 1. 1992. Costa Rica

F. E Gonz.-Led. & S.D. Koch, Novon Michoacán

4:25-27, f. 1

F. nd qus Nov. Gen. Sp. 1:153. 1815. Chiapas, Distrito Federal, Jalisco, México, Costa Rica, Guatemala

Michoacán, Morelos, Oaxaca, Puebla,

San Luis Potosí, Tlaxcala, Veracru

F. valdesii Gonz.-Led. €: S.D. Koch, Novon 8: Coahuila, Nuevo León, Tamaulipas

149-151, f. 2.1998.

F. willdenowidia Schult. & Schult. f, Add. ad Mant. Distrito Federal, México, Michoacan Guatemala

3:650. 1827. Puebla, Tlaxcala, Veracruz

10-22 cm long, pyramidal, lax, rachis glabrous below and scabrous above; branches lax, 1(-2) per lower

node, scabrous. Spikelets (6-)7.5-13 mm long, ovate to ellipsoid, more or less distal on the branches, with

4-7 florets; glumes ovate to lanceolate, scarious or with broad scarious margins, glabrous or frequently

scabrous to hispidulous in the upper half; lower glumes 3.4—5.5(-5.7) mm long, with 1(-3) veins; upper

glumes 4.5-6.5 mm long, with (1-)3(-5) veins; rachilla up to 1.5 mm long, unilaterally hispidulous op-

posite to the floret; lemmas (56-7 mm long, lanceolate to narrowly ovate, glabrous or apically scabrous

or hispidulous, with 5(-7) obscure veins, the apex broadly scarious, obtuse to abruptly acute, sometimes

emarginate to more or less bidentate, awnless or with a short mucro arising between the lobes to 0.5 mm

long; paleas 5-7 mm long, bidentate, apically hispidulous between the veins and sometimes on the up-

per margins, veins scabrous-ciliate in the upper 1/2-2/3; anthers 2.5-3.3 mm long; ovary apex sparsely

hispidulous; styles 2. Caryopses 3.5-4.5 mm long, 0.8-1.5 mm wide.

Additional collection examined: MEXICO. Coahuila: Mpio. de Arteaga, Sierra la Viga 6 km al E de Jame, Puerto Maravillas, 25?22'N

100°34'W, bosque de Pinus rudis, P. ayacahuite, Pseudotsuga, Abies, Quercus, con arbustos de Ceanothus y Pinus culminicola, 3000-3150

m, 16 Sep 1989, J.A. Villarreal & et al. VR-1983 (TEX). Fig. 2B

Festuca longiligulata belongs to subgenus Festuca section Breviaristatae and is similar to F. thurberi Vasey of

the southern Rocky Mountains in the dense caespitose habit, glumes thinner in texture than the lemmas

and long ligules. It differs from the latter in the more ovate lemmas which are obtuse (sometimes abruptly

acute) to somewhat truncate or emarginate with a wide scarious margin, while in F. thurberi the lemmas are

more narrowly ovate to lanceolate and tapering to an entire acute apex with narrow scarious margins (Fig.

3). The anthers of F. longiligulata are usually shorter, 2.5-3.3 mm, versus 3-4.5 mm, and the ovary apex is

sparsely hispidulous, versus densely hispidulous. These characters and the isolated distribution in the Sierra

Madre Oriental suggest that F. longiligula is a distinct species. It differs from F. ligulata in the more densely

caespitose habit, usually longer ligules ((2-)5-9.5 mm, versus 2.5-3.5 mm), lemmas (5-7 mm, versus 4.5-6

mm) and anthers (2.5-3.3 mm, versus 1.5-2.6 mm):

£L Dat

Journal of

830

HERBARIUM

Fig. 1. Fi

Darbyshire, F t in Meyi A Rote A : 3 T

Fic. 2.1 tion shoot leaf blad ions of Fe longiligula. A — Holotype, McDonald 2102; B — Paratype, Villarreal et al. VR-1983.

Fic. 3. Lower lemmas. A — A longiligula, McDonald 2102 (hol ); B - Festuca thurberi, Swallen 6608 (DAO).

Festuca subgenus Helleria

In hi ; f Festuca Alexeev (1980) established F. subgenus Helleria (E. Fourn.) E.B. Alexeev (Novost. Sist.

Vyssh. Rast. 17:51. 1980; = Helleria E. Fourn., Mex. Pl. 2:128. 1886, non Nees & Martius, 1824; Hellerochloa

Rauschert, Taxon 31:561. 1982) in which he included F. livida as the sole species. The long glumes (longer

than the lemmas and Do uec the whole spikelet) and the unusual strongly drooping panicles were

J;

within the genus Festuca. Alexeev interpreted these characteristics

as a highly e. PUDE o dead to the alpine habitats in which the species occurs. Other

cited as uniquely

characteristics of F. livi in these rig climatic conditions. The small anthers

(0.8-1.5 mm long) and usually non- E ced stamens and NM are typical of a cleistogamous habit. The

anthers dehisce at an early stage of inflorescence development with pollen being released into the cavity

832 Journal of the Botanical R h Institute of Texas 1(2)

between the lemma and palea. The anthers dehisce at the level of the stigmas and remain entangled in the

stigmatic branches as the caryopsis matures. The leaf sheaths of shoots and culms are loose (inflated), trap-

ping air and providing insulation for the developing tissues within. Other Central American alpine species

of Festuca have loose flag leaf sheaths, but this characteristic is pronounced in F. livida.

The pendulous form of the panicle in F. livida is unusual in that the rachis is lax right to the lowest

inflorescence node. The lowest rachis internode is scarcely wider than the single branch at the lower node,

giving the inflorescence the appearance of consisting of two branches at the lower node. As in many spe-

cies of Festuca there is a ring of tissue associated with the lowest inflorescence node. In F. livida it is usually

well developed bearing a pointed tip and sometimes extending into short bract up to 25 mm long. In other

Central America species of Festuca, the ring of tissue may bear a pointed tip, but it is scarcely more than 1

mm long. Another unusual characteristic of F. livida is the distinctly swollen and slightly flattened upper

culm internode with a large inner cavity (Fig. 4).

Two chromosome counts have been reported for F. livida, n 2 14 (Nevado de Toluca; Beaman et al. 1962)

and n = 21 (Cofre de Perote; González-Ledesma 1991). Meiosis of plants from Cofre de Perote was, however,

disrupted (González-Ledesma 1991), indicating pollen fertility problems. Pollen from anthers of 9 herbarium

specimens (DAO) showed a considerable range of development and stainability in lactophenol cotton blue

preparations. Even though most anthers seemed to be normally dehisced, conservative estimates of pollen

1]-

F

viability ranged from 0-20%. When slightly less stained and/or non-spherical (but not d) grains were

deemed to be possibly fertile, calculations suggested that 2-48% of pollen grains might be viable. However,

low pollen viability does not necessarily present a fertility handicap for cleistogamous grasses and may in

fact be an adaptive conservation of resources. Field observations in 1997 at Cofre de Perote (19°29'42"N,

97°08'55"W) and La Malinche indicated that large numbers of plants bore inflorescences with abortive

caryopses, although most florets usually contained dehisced anthers. It is possible that unusual climatic

conditions in that season caused caryopsis abortion, although given the low proportion of stainable pollen,

lack of caryopsis development may have been the result of failed pollination.

The many specialized morphological and anatomical features of F. livida, unique or highly unusual in

Festuca, provide support for the recognition of the separate genus Hellerochloa. However the taxon is clearly

related to the members of Festuca subgenus Festuca, as indicated by the similarity of the vegetative shoot

structure, leaf blade anatomy (both in cross section and the abaxial epidermis) and ovary/caryopsis mor-

phology, as well as the spontaneous hybridization with at least F. hephaestophila and F. tolucensis (see below).

In spite of the fact that intergeneric hybrids have been reported between most genera in Loliinae (except

Leucopoa) and many genera in Poeae, and that F. livida exhibits several unique or unusual characters, the

recognition of Hellerochloa seems premature

Two spontaneous hybrids, F. livida x F. hephaestophila and F. livida x F. tolucensis are sterile, intermedi-

ate in many morphological characters and not uncommon in areas where the parental species come into

contact. How the largely cleistogamous F. livida is able to hybridize with sympatric species of Festuca is yet

to be determined.

Festuca xgonzalez-ledesmae S.J. Darbyshire, nothosp. nov. (Festuca livida (Kunth) Willd. ex Spreng. x Festuca

hephaestophila Nees ex Steudel). (Fig. 5). Type: MEXICO. Veracruz: Mpio. de Perote, Cotre de Perote, lado N en la base

del macizo rocoso que esta en la cima, pastizal de Festuca, Calamagrostis, Trisetum, Agrostis y Deschampsia, alt. 4050 m, 15 Sep 1986,

Gonzál desma & P Vera C. 196 (HoLoTYPE: DAO 683891; Isotype: CHAPA).

Inter Festuca livida et Festuca hephaestophila intermedia est. Plantae perennes caespitosae, interdum rhizomatibus brevibus praeditae,

innovationibus intravaginalibus praeditae. Culmi 4.5-22 cm alti, erecti, glabri, internodia superne interdum leviter tumida. Vaginae

foliorum ima basi tantum integrae, leves, plus minusve inflatae; vaginae foliorum emortuorum in fibras longitudinales plus minusve

solutae; ligulae 0.2-1(-1.5) mm longae; laminae foliorum conduplicatae, (0. a E —0.9 mm diametri, apice acuto ad obtuso, abaxialiter

glabrae, adaxialiter hirsutae, cum 5-7 nervis, 5-7-costatae, sclerenchyma li libus, cum venas majore abaxialiter, raro

adaxialiter ee cellulis Dor. destitutae. Paniculae 223-6 cm longae, ene minusve erectae, axe distali et ramulis

scabris. Spiculae 7-11(-13) mm longae, ellipsiodeae, cum 2-)3-4(-5) flosculis; glumae glabrae ad scabrae vel hispidulae, iru

ad lanceolatae ne tantem ellipsoideae, inermes vel arista usque ad 0.3 mm longa praeditae; glumae inferiores (5.8-)6.5-8.5-9) mm

longae, lanceolatae vel ovatae, cum 1-3 nervis; glumae superiores (6.5)7-9.5(-10) mm longae, cum 3-5 nervis; rachillae usque ad 1.5

Darbyshire, F t in Mavi Val to tl PS Y | . k 833

L J . £ Luis J L 4 f

Fic. 4. Festuca livida,

S -1 eera] 5-8.5(-9.5) mm longae, | ] d ellipsoid scabrae ad hirsutae, cum 5 nervis,

m longae

apice interdum eee inermia vel plerumque praedita arista (0.3-)0.8-2.5(-3.4) mm us paleae 5.5-7.5 mm longae, inter venas

et saepe in marginalibus distalibus hispidulae, venae prope apicem ad 2/3(-3/4) cum trichomatibus; antherae (1-)1.2-2 mm longae,

plerumque indehiscentes; ovarium apex glabrus vel sparsim hispidulus. Caryopses non praeditae

Intermediate between F. livida and F. hephaestophila. Plants perennial, densely caespitose, short rhizomes

sometimes present, shoots intravaginal. Culms 4.5-22 cm tall, erect, glabrous throughout, upper internode

sometimes slightly swollen. Leaf sheaths closed only at the base, glabrous, more or less loose and disin-

tegrating into fibres with age; ligules 0.2-1(-1.5) mm long; leaf blades conduplicate, (0.3—)0.4-0.9 mm

in diameter, apex abruptly acute to obtuse, abaxially glabrous (rarely sparsely scabrous), adaxially hirsute,

with 5-7 veins and 5-7 adaxial ribs, sclerenchyma in fascicles at the margins and abaxially at the veins,

rarely small adaxial fascicles, bulliform cells absent. Inflorescence paniculate, (2-2)3-6 cm long, erect,

stiffly open, axis distally scabrous, branches scabrous, a single branch at lower node. Spikelets 7-11(-13)

mm long, ellipsoid, with (22)3—4(—5) florets; glumes glabrous to scabrous or hispidulous (especially above),

subulate to lanceolate or sometimes narrowly ellipsoid, unawned or with an awn up to 0.3 mm long; lower

glumes (5.8—)6.5-8.5(-9) mm long, lanceolate or ovate, with 1-3 veins; upper glumes (6.5-)7-9.5-10)

mm long, with 3-5 veins; rachilla up to 1.5 mm long, unilaterally hispidulous opposite to the floret; lem-

mas 6.5-8.5-9.5) mm long, lanceolate to ellipsoid, scabrous to hirsute, with 5 veins, bidentate or entire,

unawned or usually with an awn (0.3-)0.8-2.5(-3.4) mm long; paleas 5.5-7.5 mm long, hispidulous between

the veins and sometimes on the upper margins, veins with trichomes from the apex to about 2/3(-3/4) to

the base; anthers (1-)1.2-2 mm long, mostly indehiscent; ovary apex glabrous or sparsely hispidulous.

Caryopses not formed.

Additional coll ined. MEXICO. México: Nevado de Toluca, talus slopes and rocky S. rim inside crater, 3 Jul 1964, G. Mick

& K. Roe 198. Tlaxcala: vertiente NO del volcán La Malinche, zacatonal de Festuca y Calamagrostis, suelo arenoso, alt. 4080 m, 16 Sep

1986, M. González-Ledesma & P. Vera C. 205; La Malinche volcano, SE of Tlaxcala, NE of Puebla, 19°14'02"N 98?01'56"W, alpine talus

t to F. livida and F. hephaestophila, mostly on windward side of ridges and peak, 4290 m,

22 Sep 1997, SJ. DI & M. Ganas ez-Ledesma 4824. Veracruz: summit of Cofre de Perote, 19°29'42"N 97?08'55"W, alpine talus

and solifluction sites, 4140 m, 20 Sep 1997, S.J. Darbyshire & M. González-Ledesma 47944.

The inflorescence of the F. xgonzalez-ledesmae has the erect appearance of F. hephaestophila. The branches

may be stiffly spreading, but the lower branch and lower rachis internode are not drooping to pendulous as

in F. livida. The glumes are broadly to narrowly lanceolate and almost as long as or longer than the adjacent

834

I Lnaft+h,D

¿VU Mal OF

HERBARIO-HORTORIO DEL COLEGIO DE POSTGRADUADOS (CHAPA)

CHAPINGO, MEXICO

Festuca hephaestoph le x [vida

MEXICO. VERACRUZ: Mpio. de Perote. Cofre

en la base del i t

de Perote, lado N

g la cima, Pastizal

Calamagrostis, is

Trisetum, Agrost

Alt. 4 050 a. 15-11-1986

Planta glauca, frecuente.

Fic. 5. Fest

M. Gonzalez-Ledesma 196 y P. Vera £.

40z INAN\

. Scale bar: left = 2 inches; right = 5 cm.

Darbyshire, F t in Mavi Val to tl PS Y | . P 835

lemmas, whereas they are broadly lanceolate to ellipsoid and distinctly shorter than the adjacent lemmas in

F. hephaestophila. The lemmas of the hybrid are scabrous to hispid at least in the upper third and sometimes

throughout, whereas they are glabrous or only slightly scabrous to hispidulous at the very apex in F. hepha-

estophila. A comparison of additional characters is given in Table 2. In lactophenol cotton blue preparations

from the type collection, 0-2% of pollen grains were evenly stained and/or regularly formed. This hybrid was

first recognized by Manuel González-Ledesma who described it in some detail (González-Ledesma 1991).

Festuca xmiscella S.J. Darbyshire, nothosp. nov. (Festuca livida (Kunth) Willd. ex Spreng. x Festuca tolucensis

. (Fi g. 6). Type: MEXICO. Mexico: E slope of Nevado de Toluca, near gate to crater, 19°15'N 99°45'W, alpine solifluction

sites, about ed m, 26 Sep 1997, SJ. Darbyshire & M. González-Ledesma 4851 (HoLoTYPE: DAO 811264).

Inter Festuca livida et Festuca tolucensis cue est. Plantae p

Bu: aeditae. Culmi

0—60 cm alti, erecti, scabri ad hispiduli eee tumida. V. leve

vel Mn plus minusve inflatae; vaginae foliorum emortuorum in fibras longitudinales non solutae; ligulae (0.5-)1-2.5 mm Vir

laminae foliorum conduplicatae, (0.530.7-1.522.1) mm diametri apice attenuato, abaxialiter glabrae vel scabrae, adaxialiter hirsutae,

cum 7-9 nervis, 7—9-costatae, d abaxialiter annulo continuo subepidermali disposito, fasciculis adaxialiter interdum

cum venas, cellulis bulliformibu utae. Paniculae 5-15(- n cm a ns et nutantes, axe dia: et d M sud

9-14(-16) mm longae, obovatae vel late ellipsiodae, cum 4-6(

inermae vel arista usque ad 1 mm longa praeditae; glumae inferiores (6.3 a 10€ i mm a cum 1-3 nervis; glumae superiores

(7.5-)8-11(-13.8) mm longae, cum 3-5 nervis; e usque ad 1.5 mm longae, hist lemmatae (7.5-)8-10(-12)

mm longae, lanceolatae, scabrae, papill

cum 5 nervis, pe nut bt inermia vel plerumque praedita arista

(021-2(-2.9) mm longa; paleae "E En mm longae inter hi pi ulae, venae prope apicem a

trichomatibus; antherae 1.5-4.5 mm longae, plerumque leed ovarium apex glabrus. Caryposes non praeditae.

Intermediate between F. livida and F. tolucensis. Plants perennial, densely caespitose, shoots intravaginal.

Culms 20—60 cm tall, erect, scabrous to hispidulous, upper internode sometimes slightly swollen. Leaf

sheaths closed only at the base, glabrous or scabrous, somewhat loose, not splitting into fibres with age;

ligules (0.5-)1-2.5 mm long; leaf blades conduplicate, (0.5—)0.7-1.5(-2.1) mm in diameter, apex attenu-

ate to a sharp point, abaxially glabrous or scabrous, adaxially hirsute, with 7-9 veins and 7-9 adaxial ribs,

sclerenchyma in a continuous abaxial band and sometimes with adaxial fascicles at the veins, bulliform

cells abscent. Inflorescence paniculate, 5-15(-20) cm long, lax and nodding, axis glabrous below and sca-

brous distally, branches scabrous, a single branch at lowest node. Spikelets 9—14(-16) mm long, obovate

or broadly ellipsoid, with 4—6(—7) florets; glumes glabrous, scabrous or hispidulous, subulate to lanceolate,

unawned or with an awn up to 1 mm long; lower glumes (6.3—)7—10(-11.5) mm long, with 13 veins; upper

glumes (7.5—)8—11(-13.8) mm long, with 3—5 veins; rachilla up to 1.5 mm long, unilaterally hispidulous

opposite to the floret; lemmas (7.5—8—-10(-12) mm long, lanceolate, scabrous, papillose or hirsute, with 5

veins, apex usually bidentate, unawned or usually with a short awn (O—)1—2(-2.9) mm long; paleas 6-7.5

mm long, hispidulous between the veins and sometimes on the upper margins, veins with trichomes from

the apex to about 3/4 to the base; anthers 1.5-4.5 mm long, mostly indehiscent; ovary apex glabrous.

puis not formed.

A dditi ined. MEXICO. México: E slope of Nevado de Toluca tet ter, 19°15'N 99°45'W, alpine solifluc-

tion sites, ca. 4000 m, 26 Sep 1997, S.J. Darbyshire & M. Gonzdlez-Ledesma 4849.

The open paniculate inflorescence of the F. xmiscella is similar in appearance to that of F. tolucensis, but with

fewer spikelets (usually less than 20 versus usually more than 20). The lower rachis internode is stiff or

somewhat lax, but not pendulous as in F. livida. The spikelets are obovate to broadly ellipsoid in F. xmiscella

with a length to width ratio usually 2 or less. In F. tolucensis the spikelets are usually lanceolate or narrowly

ellipsoid (occasionally broadly ellipsoid) with a length to width ratio usually 2.5 or greater (occasionally

as little as 1.5). The glumes are 3/4 as long as or about as long as the adjacent lemmas, whereas they are

usually 1/2 to 3/4 as long as the adjacent lemmas in F. tolucensis. A comparison of additional characters is

given in Table 2. In lactophenol cotton blue preparations from the type collection, 2-27% of pollen grains

were evenly stained and/or regularly formed. This hybrid was first identified and characterized by González-

Ledesma (1991).

£L Dag PRA D hi PEA

836 Journal of f Texas 1(2)

TABLE 2. A compari fd istics of A hepl hila, F livida, E tol

character F. hephaestophila F. xgonzalez- F. livida F. xmiscella F. tolucensis

ledesmae

spikelet length 45-84 7-11(-13) (9.4)10-17(21) | 9-14(-16) (6.5-)8-14(-18)

lower glume length — 2.8-4.5(-5.5) (5.8-)655-8.5(-9) — 9.2-15(-20) (6.3-)7-10(-11.5) Q7-)35-7.5(-8.6)

upper glume length 3.5-6.3 (6.5-)7-9.5(-10) 94-17(-21) (/5-)8-11(-13.8) | (3.6294.5-8.5(-9.3)

lemma length 4-5.5(-6.2) 6.5-8.5(-9.5) 6.5-12.5 (7.5-)8-10(-12) (4.6-)5.5-8.5(-10)

anther length 2.2-3.5 (1-)1.2-2 0.8-1.5 1.5-4.5 (2.6-)3-4.5(-5.2)

The following key to species of Festuca in Mexico and Central America is largely based on that by

Alexeev (1984), but has been updated with additional information and recently described taxa. Although

frequently treated in a different genus, Festuca arundinacea Schreb. [Lolium arundinaceum (Schreb.) Darbysh.;

Schedonorus arundinaceus (Schreb.) Dumort.] is included in the key and Table 1, because of its similarity to

and continued placement in Festuca by some authors. The inclusion of F. ovina L. is based on Godfrey 66649

report by Pohl (1980) for Costa Rica and accepted by Alexeev (1984). The specimen has not been examined

and its identity remains unclear.

KEY TO THE SPECIES OF FESTUCA IN MEXICO AND CENTRAL AMERICA

. Leaf blades of shoots usually more than 2 mm wide, flat in cross section, sometimes round in the upper part

(or becoming involute or convolute when dry), p ing distinct (wide) bulliform cells.

2. Margins of the blade-sheath junction with falcate auricles, more or less provided with cilia_______ F. arundinacea

2. Margins of the blade-sheath junction lacking auricles, without cilia.

3. Ligules of culm leaves (1.3-)2-10(-11) mm lon

4. Plants loosely caespitose; ligules of culm leaves less than 3.5 mm long; lemmas less than 7 mm long,

apex entire, awnless

5. Ligules of culm leaves 2.5-3.5 mm long; leaf blades up to 2 mm wide; lemmas 4-6 mm long;

plants of Mexico and Texas F. ligulata

5. Ligules of culm leaves less than 2.5 mm long; leaf blades 1-4 mm wide; lemmas 5.7-6.8 mm long;

BRE of e Rica ang E F. herrerae

4. Plant Im | lly 3-8( ue cuc e a

EUM lemmas greater than 7 mm lon bi ¿with an awn

or a mucro (sometimes absent) ea from NS the teeth.

6. Apex of ovaries densely hisp F. asperella

6. ign iu ies Pun

E

at most with a short mucro

less d 0. 5; mm jos ng F. tancitaroensis

7. Ligules of culm leaves (1.3-)3-8(-11) mm long; lemmas with an awn (0.5-)1-4 mm long.

8. Leaf blades stiff, 3-5 mm wide; upper glume (5.5-)6-8.5 mm long F. lugens

8. Leaf blades lax, 1.5-3.6(-6) mm wide; upper glume (3.8-)4-5.6(-6) mm long____________ Kroblensis

3. Ligules of culm leaves not more than 1.5 mm long

9. Lemmas 12-17.5 mm long.

10. Lower glumes (1-)2-6(-7.5) mm long; lemmas with 3-5 veins, awns 10-14(-16) mm long

F. breviglumis

10. Lower glumes (7.5-)8-9 mm long; lemmas with 5 veins, awns 1.5-3 mm long . .. F chiriquensis

9. Lemmas 3-9.5(-10) mm lon

11. Apex of lemmas bidentate awns (0.3-)0.5-1.8 mm long arising between the teeth.

12. Lemmas 5.5-6.2 mm long, scabrous throughout the back; paleas scabrous along the keels

to the base; anthers 2.4-2.7 mm long; apex of ovaries densely hispid; plants of Panama

Ligu 15

F. swallenii

12. Lemmas 6.5-7 mm long, scabrous on the upper half; paleas scabrous along the keels on

the upper third; anthers 3.5-4 mm long; apex of ovaries with a few hairs (1-4); plants of

F. bidenticulata

Mexico

11. Apex of lemmas without teeth or awns.

13. Lemmas 3-4.5 mm long, apex obtuse; anthers 1-1.3 mm long F. subverticillata

837

A y PRO J

Darbyshire,

pé AND AIF

¿E e

GS N^

= 3 81126 Y,

o >

awa, CASES.

2006

| LLLI

[ITI]

Plants of Mexico

Festuca livida x Festuca tolucensis

Mexico, Mexico, east slope of Nevado de Toluca; near gate to crater;

about 4000 m elev.

19?15'N, 99°45'W

alpine solifluction sites

J Darbvshi LM c+} Fak 4851

Coll.: S.

26 Sept. 1997

appears to be a hybrid swam with plants intermediate between F.

livida and F. tolucensis; 7 replicates

4851 (DAO). Scale bar: left — 2 inches; right — 5 cm.

Fic. 6. Fe

838 f the Bot

Journal UI

titute of Texas 1(2)

13. Lemmas 5.5-9.5(-10) mm "E apex acute or mucronate; anthers 1.6-4 mm long.

14. sag of ovaries glabrou

15. Leaf blades flat or T folded, 1-4 mm wide; spikelets 6.5-9 mm long _______ R herrerae

15: " blades i Du Or E 3 10 mm MuR du ie 17 mm long.

]le leaf blades

A or with shallow adaxial ribs; IE branches scabrous or ini

pilos .amplissima

16. Dead leaf sheaths not breaking up into longitudinal fibres; leaf blades ae

ong adaxial ribs; A branches mostly smooth F. jaliscana

14. Apex T E densely hisp

17. Anthers 2.8-4.5(-5) mm F plants of northeast Mexico F. valdesii

17. Anthers up to 2.5 mm long; plants not of Mexico (further south).

18. a branches smooth or almost smooth; spikelets 6.5-7.5 mm long, with

-3 florets; anthers 1.7-2 mm Heng) Sel s of Guatemala F. aguana

18. men brancl | long, with 4-5 florets; anthers

2-2.5 mm long; plants of Panama F. panamica

. Leaf blades of shoots usually less than 2 mm in diameter, folded (conduplicate) in cross section, lacking or

with indistinct narrow bulliform cells.

9. almost as long or longer than the florets (excluding awns) and longer than adjacent lemmas,

sometimes with a mucro up to 1.3 mm long; inflorescence nodding, lower rachis internode lax and more

or less pendulous F. livida

19. Glumes distinctly shorter than the florets and shorter than adjacent lemmas (except in sterile hybrids

with F. livida keyed below), without a mucro or with a mucro less than 0.3 mm long; panicles erect or

nodding, lower rachis internode stiff or lax but not pendulous

20. Plants sterile (anthers mostly indehiscent, caryopses not | ] usually more than 3/4

as long as spikelets and about as long as lemmas; plants alpine, o above tree-line; growing

with F livida (Central Mexico).

21. Plants 4.5-22 cm high, leaf blades with 5-7 veins, sclerenchyma fascicles about as wide as v

anthers 1-2 mm F- xgonzalez: -ledesmae

21. Plants 20-60 cm high, leaf blades with 7-11 veins, sclerenchyma fascicles much wider th

Si rs 1.5-4.5 mm lon

Plants usually fertile (anthers usually dehiscent, caryopses produced); glumes usually less than 3/4

as rae as spikelets and distinctly shorter than lemmas; plants montane or alpine; growing with F

livida or not.

22. Der blades of shoots with one adaxial rib.

23. Leaf blades of vegetative shoots with 3 veins, sclerenchyma in 7 small abaxial fascicles; apex

of ovaries hispid

23. Leaf blades of vegetative shoots with 5-7 veins, scl | ti

abaxial band; apex of ovaries glabrous.

24. Upper e usually 2.6-4 mm long; lemmas Ed more than 3.3 mm long, with

short awns 0.5-2 mm long; anthers (1.4-)2-2.6 m

©

=)

E xmiscella

N

e

ajacaliforniana

interrupted

F. ovina

24. aa Fae usually 2-3 mm long; lemmas EV less than 3.3 mm long, without

awns (rarely mucronate); anthers (1-)1.5-2 mm long F. filiformis

22. Leaf blades of shoots with 3-21 adaxial ribs.

25. Anthers (0.8-)1-1.4(-1.6) mm long.

26. Spikelets 8-10 mm long; lemmas 6-8.5 mm long, puberulent throughout —— F. willdenowiana

26. Spikelets 6.4-7 mm long; lemmas 5-6 mm long, glabrous or apically scaberulous

F. talamancensis

25. Anthers greater than 1.5 mm long.

27. Lemma aj bid i i |

or not at all; examine multiple lemmas),

awn (0.3-)0.7-3(-3.7) mm long, arising between the teeth or apex rarely emarginate

with the awn arising a little below the apex

28. Leaf sheaths usually glabrous; ligules of culm leaves (1-)1.5-3.2(-4.6) mm long;

abaxial surface of leaf blades Selects pal cule) ule oil eles ina eon ELON

or interrupted abaxial ~ usually g adaxial

fascicles often p

28. Leaf sheaths pen ionem scabrous; ligules of culm leaves 0.4-1 mm long;

abaxial surface of leaf blades glabrous or slightly scabrous; sclerenchyma of leaf

F. tolucensis

Darbyshire,

r

+

27,

I y to tl f. PS ee | i. | M 839

hlag i : | re | | E B HT

not or girders, adaxial fascicles

absent

F. aequipaleata

Lemma apex entire, without an awn or awned from the apex.

29, ligules o of culm is -)2.5 mm long or more.

aves 2.5-3.5 mm long; lemmas 4.5-6 mm long, apex narrowly

. Lig

scarious; sete 1.5-2.6 mm long F. ligulata

30.

Ligules of culm leaves (2-)5-9.5 mm long; lemmas 5-7 mm long, apex broadly

carious; anthers 2.5-3.3 mm long F. longiligula

29. aules fcum leaves ess than or equal 2 mm lon

sub

31

eaf blades of vegetative shoots (the widest) 1-2 mm in diameter, with 11-23

veins, with 9-21 adaxial ribs.

32. Panicle branches smooth; paleas scabrous along the keels on the upper

2/3 — 3/4; apex of ovaries glabrous; plants of Costa Rica and Panama

F. cartagana

32. Panicle! | | pal | | he keel the upp

1/3 — 1/2; apex of ovaries hispid; plants of Mexico F. orizabensis

Leaf blades of vegetative shoots always 0.3-1(-1.2) mm in diameter (some-

times wider in F. rubra), with 5-9 veins, with 3-7 adaxial ribs

33. Leaf blades Du se: shoots with abaxial sclerenchyma in a continu-

ous band, rarely ir Ens

34. Leafblades of g | 5 in di t ith 5(-7)

veins, adaxial surfaces with hairs 50- 80 um long; ies of blades

and sheaths swollen (callose auricles) F. callosa

34. Leaf blades of vegetative shoots 0.6-0.8 mm in diameter, with 7-9

veins, adaxial surfaces with hairs 20-30 um long; junction of blades

and sheaths not swollen (auricles not callose).

35. Margins of leaf sheaths of vegetative shoots closed up to 2/3;

ligules of culm leaves 0.2-0 ong; leaf blades usually

uco-pruinose; lemmas 6.5-7 mm long, not awned; apex of

ovaries densely hispid F. hintoniana

35. Margins of leaf sheaths of vegetative shoots closed up to 1/3;

ligules of culm leaves 0.5-1 mm long; leaf blades green, not

glaucous or pruinose; lemmas 5-5.2 mm long, awns 2-2.2 mm

long; apex of ovaries glabrous F. beamanii

33. Leaf blades of vegetative shoots with abaxial sclerenchyma in fascicles.

36. Leaf blades of vegetative shoots always with 5 veins, with 3 adaxial

ribs

37. Margins of leaf sheaths of vegetative shoots closed from the

ear the top; dead leaf sheaths not breaking up into

longitudinal fibres; leaf blades abaxially more or less scabrous

or rarely gata e E dense Meu Sel eee el

fascicles

the keels on the upper 1 V/;a apex KO CES with a few hairs, rae

glabrous; plants alpine F. hephaestophila

27. Margins of leaf sheatt is of tati | ts cl | fi the base

o 1/3; dead leaf sheaths more or less breaking up into longi-

tudinal fibres; leaf blades abaxially smooth (glabrous), adaxially

smooth or almost so, adaxial fascicles of sclerenchyma absent;

montane F. rzedowskiana

36. Widest E blades of vegetative shoots with 7-9 veins, with 3-7

adaxial ribs.

38. Plants laxly caespitose, often with short rhizomes; margins

of leaf sheaths of vegetative shoots closed from the base to

at least 1/2; lemmas with an awn greater than 1 mmlong________ F.rubra

38. Plants more or less densely caespitose, without rhizomes; mar-

gins of leaf sheaths of vegetative shoots closed from the base

840 Journal of the Botanical R Institute of Texas 1(2)

to not more than 1/3; lemmas awnless or sometimes with an

apical mucro up to 1 mm lon

39. Anthersinl r floret in tl ikel lly ak

of upper florets well-developed and 3-4.2 mm long_

39. Anthers in lower florets of the spikelet not abortive, all

anthers more or less the same length, 1.6-6 mm long.

, those

_F.diclina

40. Some ot g f f

tose); adaxial surface of leaf blades of vegetative shoots

with a few isolated trichomes 20-40 um long; anthers

1.6-2.3 mm lon

41. Ligules 0.1-0.2 mm long; spikelets 10.5-11.5 mm

long, with 5-6 florets; lemmas 6.5-7.5 mm long

F. guatemalica

41. Ligules 0.3-2 mm long; spikelets 6.5-9 mm

long, with 3-5 florets; lemmas 5.7-6.8 mm long

F. herrerae

40. All shoots intravaginal; adaxial surface of leaf blades of

vegetative shoots with dense trichomes 50-100 um

long; anthers greater than 2.5 mm long.

42. Blades of leaves always folded (conduplicate or

convolute), 0.3-0.6 mm in diameter, with 5-7(-9)

veins; panicles more or less dense, branches erect;

apex of ovaries glabrous (rarely with a few hairs)

F. rosei

42. Blades of culm leaves flat to loosely folded (con-

duplicate), (0.4-)0.6-1.2 mm in diameter, with

9-11 veins; panicles open, branches lax; apex

of ovaries more or less hispid (rarely glabrous)

F. pringlei

ACKNOWLEDGMENTS

I am grateful to Jacques Cayouette, Manuel González-Ledesma, Jochen Müller and Barbara Wilson for their

useful comments and discussions, to Jacques Cayouette and Jochen Müller for assistance with the Latin

descriptions, and to herbarium curators at ARIZ, DAO, TEX, US and WIS for making specimens available

for study. Manuel González-Ledesma was of invaluable assistance with field work.

REFERENCES

ALEXEEV, E.B. 1980. Festuca L. subgenera et sectiones novae ex America boreali et Mexica. Novosti Sist. Vyssh.

Rast. 17:42-53. [In Russian].

ALEXEEV, E.B. 1984. Genus Festuca L. (Poaceae) in Mexico et America Centrali. Novosti Sist. Vyssh. Rast. 21:25-59.

[In Russian].

Beaman, J.H., D.C.D. De Jone, and W.P. Stoutamire. 1962. Chromosome studies in the alpine and subalpine floras of

Mexico and Guatemala. Amer. J. Bot. 49:41—50.

GONZALEZ-LEDESMA, M. 1991. Révision del genera Festuca L. (Gramineae) en el Eje Volcánico Transversal. Ph.D. dis-

sertation. Colegio de Postgraduados, Chapingo, Mexico.

PoHL, RW, 1980. Family 15. Gramineae. In: W.C. Burger, ed. Flora Costaricensis. Fieldiana Bot. New Ser. 4:1-608.

SORENG, R.J., P.M. PETERSON, G. DAvipse, E.J. Jupziewicz, F.O. ZULOAGA, T.S. FILGUEIRAS, and O. Marone (eds.). 2003. Catalogue

of New World grasses (Poaceae): IV. Subfamily Pooideae. Contr. U.S. Natl. Herb. 48:1—730.

APHANELYTRUM PERUVIANUM (POACEAE: POINAE):

A NEW SPECIES FROM PERÚ

Isidoro Sánchez Vega Paul M. Peterson and Robert J. Soreng

Muss de Biología Department of Botany

Herbario CPUN National Museum of e eo

Universidad Nacional de Cajamarca Smithsonian Institu

Apartado 55, ACH PERÜ Washington, DC Ha v U.S.A.

svisidoro@yahoo.co peterson@si.edu sorengr@si.edu

Simon Leegaard

Department of Systematic Botany

DK-8000 Aarhus C, DENM,

simon.laegaard@biology.au. aa

ABSTRACT

Aphanelytrum peruvianum Sánchez Vega, P.M. Peterson, Soreng & Leegaard, sp. nov., is described and illustrated. The new species

occurs on rocky sites near Cerro Akumullca in Cajamarca, Perú and represents the second known species ascribed to Aphanelytrum.

Aphanelytrum peruvianum differs from A. procumbens by having nerved glumes (1-2 mm long), narrow leaf blades (0.2-1.2 mm wide),

shorter culms (14-24 cm a aus eS x 13mm a shorter lemmas (2.2-3.5 mm lone and shorter anthers 2-2.9 mm

long). A key and t g g g A. peruvianum and A. p l

E

RESUMEN

Sed RI :] ARES 1 : Caral TS

Vega, P.M. Peterson Soreng & Laegaard Sp. nov.

Į

en sitios rocosos cerca del ue MTM en Cajamarca, Perú. Esta nueva la segunda i ida atribuida

o

al género Aphanelytrum. Aphanelytrum Vi nb diri difiere de A. procumbens en ee tiene e gmas nervadas (1- 2 mm largo), láminas de

la hoja estrechas (0.2-1.2 mm MD rtos (14-24 cm alto), ent 3-13 mm largo), lemas más cortas

ee 2-3.5mm TN yant tas Q-2.9 mm largo). Se incluye una clave y una tabla de comparación de |

aA peruvianum de A. procumbens.

The genus Aphanelytrum (Hack.) Hack., established in 1902, was first named without a description by Sodiro

(1889) with a single species, “A. decumbens Hack.” (ex Sodiro, nom. nud.). Hackel (1887) initially described

it as a subgenus of Brachyelytrum P. Beauv.; B. subgen. Aphanelytrum Hack., including a single new species

B. procumbens Hack. Hackel (1902) subsequently recognized Aphanelytrum as a genus with the single spe-

cies A. procumbens (Hack.) Hack., all based on a single collection of Sodiro’s from Ecuador (Chase 1916).

The placement and evolutionary relationships of Aphanelytrum have been somewhat controversial since its

inception. It was originally placed in the subfamily “Festuceae” [Festucoideae], applied in the broad sense

of Bentham (1881), Hackel (1887), and Hitchcock (1935). Hackel mistook the individual florets for glume-

less single-flowered spikelets and placed Brachyelytrum within tribe Agrostideae, subtribe Stipinae. Agnes

Chase (1916) reinterpreted the spikelet morphology, and placed the genus between the subtribes Melicinae

and Centothecinae, which at that time were considered adjacent subtribes of the subfamily Festucoideae,

tribe Festuceae. More recently, after major realignments of the classification of the Poaceae family (e.g.

Clayton & Renvoize 1986), the genus was placed in the much more narrowly defined subfamily Pooideae

(syn. Festucoideae), tribe Poeae (syn. Festuceae) near Poa (Clayton & Renvoize 1986). Aphanelytrum and

Poa have very similar leaf anatomical characteristics, as well as multi-flowered, membranous spikelets with

5-nerved, keeled lemmas, glabrous ovaries, and caryopses with oval hilums (Clayton & Renvoize 1986). It

was specifically placed in subtribe Poinae (Soreng et al. 2003; Soreng et al. 2006) and seems to align near

J. Bot. Res. Inst. Texas 1(2): 841 — 845. 2007

842 Journal of the Botanical R h Institute of Texas 1(2)

[or on an unusually long branch within; 29 base pair differences in the internal transcribed spacer regions

(ITS) of nuclear ribosomal DNA] Poa in preliminary cpDNA and ITS sequence analyses (Gillespie et al. 2007;

Soreng et al., In press; Saarela pers. com.).

Aphanelytrum is currently recognized as a monotypic genus found at mid to high elevations 2000-4050

m) in humid to montane forests in the Andes of Bolivia, Columbia, Ecuador, and Perú (Hitchcock 1927;

Clayton & Renvoize 1986; Jorgensen & Ulloa Ulloa 1994; La Torre 2002; Soreng et al. 2003; Leegaard 2005).

Aphanelytrum procumbens is a peculiar grass that has unusual spikelets with minute and unnerved glumes,

two or three florets that are widely-spaced because of the long flexuous rachillas Y to % as long as the florets,

rachillas that are prolonged above the upper florets, and 5-nerved lemmas (Chase 1916; Hitchcock 1927;

Nicora & Rúgolo de Agrasar 1987; Watson & Dallwitz 1992; Leegaard 2005).

While collecting specimens for the floristic inventory of “jalca formations” in northern Perú, Isidoro

Sánchez Vega and others collected a specimen that was not easily assignable to Aphanelytrum procumbens.

Since this specimen morphologically resembles A. procumbens, we are describing it as a new species of

Aphanelytrum and are therefor ling the generic description to include additional morphological varia-

tion.

Aphanelytrum peruvianum Sánchez Vega, P.M. Peterson, Soreng & Leegaard, sp. nov. (Fig. 1A & B, D-L).

Tre: PERÚ. Cajamarca: Prov. Cajamarca, Dist. Cajamarca, Cerro Akumullca, al SO de Cajamarca, sobre la cima de la ladera occidental

del Valle de Cajamarca (7°14'15"S Lat, 78°29'24"W Long), 3300 msnm, 20 Mar 2003, I. Sánchez Vega 11718, M. Sánchez Montoya, R.

Cueva R. & J. Montoya (HoLotyee: CPUN!, isotypes: AAU!, Fl, HAO!, HUT!, LOJA!, MICH!, MOI, SI!, US-3472470!, USM).

Se ae procumbens Hack. culmis 14-24 cm longis, internodiis 3-13 mm longis, ligulis 2-3 mm longis, laminis 0.2-1.2 mm latis,

glumis 1-2 mm longis, inferioribus glumis 1-innervatus, superioribus glumis 3(4-)-innervatus, lemmatibus 2.2-3.5 mm longis 3- vel

5-innervatis, antheris 2-2.9 mm longis, recedit.

Caespitose perennials. Culms 14-24 cm tall, with many culms near base, primary and secondary culms

appressed, somewhat decumbent near base with extravaginal branching; internodes 3-13 mm long, nu-

merous. Leaf sheaths longer than the internodes, membranous to hyaline, open, slightly keeled; ligules 2-3

mm long, membranous to hyaline, decurrent, apex erose often lacerate; blades 3-7 cm long (flag leaf ca 1.6

mm long), 0.2-1.2 mm wide, flat to loosely involute, linear, apex naviculate. Panicles 1.7-2.5 cm long, few-

flowered with 5-10 spikelets; branches flexuous, the lower branches with two spikelets, the upper branches

with single spikelet. Spikelets 5-7 mm long, usually 3-flowered, purplish, disarticulating above the glumes

and between the florets; lower and middle florets usually staminate; upper florets usually pistillate; rachilla

joints 1.2-2 mm long, prolonged above the upper floret; glumes 1-2 mm long, subequal, apex acute, often

mucronate; lower glume linear, 1-nerved; upper glume oblanceolate, 3(4-)-nerved, often toothed or irregu-

larly lobed minutely bifid; lemmas 2.2-3.5 mm long, ovoid, 3- or 5-nerved, apex mucronate with two acute

lobes on each side of the mucro, the mucro 0.1-0.3 mm long; paleas 2-3.2 mm long, 2-keeled, apex bifid;

lodicules 7-8 mm long, lanceolate, membranous, glabrous; stamens 3; anthers 2-2.9 mm long, yellowish

to purplish; ovaries glabrous with two styles and two stigmas. Caryopses glabrous.

Phenology.—Flowering in March.

Distribution and habitat —Aphanelytrum peruvianum is known only from the type locality near the west-

ern highlands of the Cajamarca Valley and is found on rocky sites associated with jalca vegetation (humid

alpine grass ecosystems) at 3300 m

DISCUSSION

We are here emending the generic description of Aphanelytrum (Watson € Dallwitz 1992) to include: leaf

blades sometimes involute; spikelets with glumes 1-2 mm long, nerved; lower glumes 1-nerved; upper

glumes 3(4-)-nerved; lemmas 3- or 5-nerved.

Festuca reclinata Swallen superficially resembles Aphanelytrum since the former species from the Páramo

del Almorzadero in the Cordillera Oriental of Colombia exhibits similar morphologies (Stančík & Peterson

2007). The growth habit, panicles, and spikelet characteristics of F. reclinata are strikingly similar to those

<=

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Fic. 1. Aphanelytrum peruvianum [I. Sánchez Vega 11718, M. Sánchez Montoya, R. Cueva R. & J. Montoya (US)]. A. Habit. B. Sheath, ligule, and blade. D.

Panicle. E. Spikelet. F. Glumes, showing upper glume 4-nerved with ay hed

ifid. H. Lemma. I. Paleas. J. Palea, anthers, and rachilla. K. Lodicul ] y l Ovary Ar h ly p IPM.

Rodríguez (US)] C. Sheath, ligule and blade.

844 Journal of the Botanical R h Institute of Texas 1(2)

Taste 1. Salient fi paring Aphanelytrum peruvi ith A. procumbens.

Characters A. peruvianum A. procumbens

Culm height 14-24 cm 30-70 cm

Internode length 3-13 mm (15-)24-90 mm

Ligule length 2-3 mm 1-22 mm

Blade width 0.2-1.2 mm 1.5-4.2 mm

Glume length 1-2 mm 0.1-0.3 mm

Glume nerves present absent

Lemma length 2.2-3.5 mm 3.2-8.4 mm

Number of lemma nerves IONS 5

Anther length 2-29 mm 3.5-3.8 mm

found in A. peruvianum. Both species have weak, decumbent culms with extravaginal branching, narrow

few-spikeleted panicles, spikelets with flexuous rachillas, and small, nerved glumes (lower glumes 1-nerved;

upper glumes 3-nerved). In comparison with F. reclinata, A. peruvianum has smaller lemmas (2.2-3.5 mm

vs. 7-8.5 mm in F. reclinata), smaller anthers (2-2.9 mm vs. 3.5-3.8 mm), and shorter spikelets (5-7 mm vs.

10-13 mm). Moreover, the strongly keeled lemmas of F. reclinata do not agree with its placement in Festuca.

Further study is needed to confirm the connection of F. reclinata with Aphanelytrum.

Aphanelytrum peruvi differs from A. procumbens (see Table 1) by several characteristics, most notably:

culm height (14-24 cm long in the former versus 30-70 cm long in the latter), internode length [3-13 vs.

(15-)24-90 mm], ligule length (2-3 vs. 1-2 mm), leaf blade width (0.2-1.2 vs. 1.5-4.2 mm), glume length

(1-2 vs. 0.1-0.3 mm or absent), presence of glume nerves (lower glumes 1-nerved, upper glumes 3- or 4-

nerved vs. both glumes unnerved), lemma length (2.2-3.5 vs. 3.2-8.4 mm), number of lemma nerves (3 or

5 vs. 5), and anther length (2-2.9 vs. 3.5-3.8 mm). In addition to these salient characteristics, A. peruvianum

has leaf sheaths that are usually longer than the internodes (vs. shorter in A. procumbens), blades that are

flat to loosely involute (vs. flat in A. procumbens, see Fig. 1C), and shorter rachilla extensions (1.2-2 mm

vs. 1.6-4.2 mm in A. procumbens). A key is presented below to help differentiate between the two species of

Aphanelytrum.

KEY TO THE SPECIES OF APHANELYTRUM

—

. Glumes 1-2 mm long, the lower 1-nerved, the upper 3- or 4-nerved; leaf blades 0.2—1.2 mm wide; culms 14—24

cm long; internodes 3-13 mm long; lemmas 2.2-3.5 mm long; anthers 2-2.9 mm long . ____ A. peruvianum

1. Glumes 0.1—0.3 mm long, minute or absent, unnerved; leaf blades 1.5—4.2 mm wide; culms 30-70 cm long;

internodes (15—)24—90 mm long; lemmas 3.2—8.4 mm long; anthers 3.5-3.8 mm long ___ ____ A. procumbens

ACKNOWLEDGMENTS

We thank the National Science Foundation (NSF, Grant 0071506) to Michael O. Dillon and the The Field

Museum (Chicago) for financial support; Alice R. Tangerini at the Smithsonian Institution for preparing the

illustration; Patricia Gómez Bustamonte for help preparing the Spanish abstract; Alain Touwaide for help

correcting the Latin diagnosis; and Diego A. Giraldo Cañas and Jeffery M. Saarela for suggesting improve-

ments to the manuscript.

REFERENCES

BENTHAM, G. 1881. Supplemental papers to Bentham and Hooker's Genera Plantarum [Gramineae]. J. Linn. SOc.,

Bot. 19:14-134.

Chase, A. 1916. The structure of the spikelet of Aphanelytrum. Bot. Gaz. 61:340—343.

CLAYTON, W.D. and S.A. Renvoize. 1986. Genera Graminum. Grasses of the world. Kew Bull. Add. Ser. 13:1—389.

GILLESPIE, L.J., A. ARCHAMBAULT, and R.J. Sorens. 2007. Phylogeny of Poa (Poaceae) based on trnT-trnF sequence data:

major clades and basal relationships. Aliso 23: In press.

Ciakas V. + I,A | : f Apl ht r4 D 845

HackeL, E. 1887. Gramineae. In: A. Engler & K. Prantl, eds., Die Natürlichen Pflanzenfamilien 2.2:1-97. Wilhelm

Engelmann, Leipzig.

HackeL, E. 1902. Neue Graeser. Oesterr. Bot. Z. 52:8-15.

HircHcock, A.S. 1927. The grasses of Ecuador, Perú, and Bolivia. Contr. U.S. Natl. Herb. 24(8):201—519.

HitcHcock, A.S. 1935. Manual of the grasses of the United States. USDA Misc. Publ. 200. U.S. Govt. Printing Office,

Washington.

Jorcensen PM. and C. ULLOA ULLOA. 1994. Seed plants of the high Andes of Ecuador—A checklist. AAU Rep.

L&GAARD, S. 2005. Poaceae (Gramineae). In: P. Sklenář, J.L. Luteyn, C. Ulloa Ulloa, PM. Jorgensen, and M.O. Dillon,

eds. Flora genérica de los páramos. Mem. N.Y. Bot. Gard. 92. Pp. 358—391.

La Torre, M.I. 2002. Primer registro de Aphanelytrum (Poaceae: Poideae) para el Perú. Arnaldea 8(2):53—56.

Nicora, E.G. and Z.E. RUGOLO DE AGRASAR. 1987. Los Géneros de Gramíneas de América Austral Argentina, Chile, Uru-

guay y areas limítrofes de Bolivia, Paraguay y Brasil. Editoria Hemispherio Sur S.A., Buenos Aires, Argentina.

Sopiro, PL. 1889. Gramíneas ecuatorianas de la Provincia de Quito. Anales Univ. Centr. Ecuador 3(25):474-484.

SORENG, R.J., J.| Davis, and M.A. Voionmaa. In press. A phylogenetic analysis of Poaceae tribe Poeae s.l. based on

morphological characters and sequence data from three plastid-encoded genes: evidence for reticulation,

and a new classification for the tribe. Kew Bull.

SORENG, R.J., P.M. PETERSON, G. Davipse, E.J. Jubziewicz, F.O. ZULOAGA, T.S. FILGUEIRAS, and O. Morrone. 2003. Catalogue of

New World grasses (Poaceae): IV subfamily Pooideae. Contr. U.S. Natl. Herb. 48:1—730.

SORENG, R.J., PM. PETERSON, G. DAVIDSE, E.J. Jubziewicz, F.O. ZULOAGA, T.S. FILGUEIRAS, and O. Morrone. 2006. Suprageneric

Classification. http://mobot.mobot.org/W3T/Search/nwgclass.htm|

Stančík, D. and P.M. Peterson. 2007. A revision of Festuca (Poaceae: Loliinae) in South American paramos. Smith-

sonian Contr. U.S. Natl. Herb. 57: In press.

Watson, L. and M.J. Dallwitz. 1992. The grass genera of the world. CAB International University Press,

Cambridge.

846 Journal of the Botanical R h Institute of Texas 1(2)

DOOR REVIEWS

SYLVAN RAMSEY KAUFMAN and WALLACE KAUFMAN. 2007. Invasive Plants: Guide to Identification and the

Impacts and Control of Common North American Species. (ISBN 978-0-8117-3365-6, pbk).

Stackpole Books, 5067 Ritter Road, Mechanicsburg, PA 17055, U.S.A. (Orders: www.stackpolebooks.

com, orders@stackpolebooks.com, 717-796-0411, 717-796-0412 fax). $39.95, 458 pp., 503 color

photos, 8 1/45. 1/21.

Touted as “the first-ever field guide to invasive plants in North America”, this book is a thoughtful compendium of over 150 invasive

plant species of the U.S. and Canada and the larger issue of invasive species impact and control. The first 36 pages provide an adequate

line between "exotic" and “pest”, and the colored history of

overview of the e Re on what it means to be invasive, the fine

nd those D od sought me their ecological benefits. Suggestions are provided for various manage-

l ies, i ical, biological, and chemical methods, and are later revisited relative to each individual

des

ment and

species. All instructions are clear idi concise, as is the overview of common herbicides, their use, and their potential dangers. Natural

or eco-friendly herbicides are not mentioned.

TT 1

kseguesfromi | ial in Part I to species descriptions in Part II by way of a“ eo (s i Lus in the familiar

: Pet E ; : : E MEC

biology key format", which, though technically accurate, is hard 11 ] ld! g., bullets

numbers, letters). The individual ies d ipti ] l however lerful ff. Each plant is depicted i ]

color photograph, along with the specific epithet, various common names, family name, and any hybridizing species or species for which

n p may ES EUM In A DUE the id prove no for LE des MD M d features, habitat, and range, as

i erica. Species- M management

tips and a short list of a are Msc provided for each eine with special mention for sources that

bibliography of 25 articles, books, and pa a x credits section, and an index of categories from

d Ag i 1 layperson mco recommend

and with an annotated

key, Latin names

this book.—M. Brooke Byerley, Botanical Research Institute 2 Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.

Oscar E CLARKE, DANIELLE SVEHLA, GREG BALLMER, and ArLEE MONTALVO. 2007. Flora of the Santa Ana River

and Environs: With References to World Botany. (ISBN 978-1-59714-050-8, pbk.). Heyday Books,

PO. Box 9145, Berkeley, CA 94709, U.S.A. (Orders: www.heydaybooks.com). $29.95, 496 pp., 3200

images and illustrations, 6" x 9",

Casual observers of the botanical world and experts alike, located both a ix in: z ue Santa Ana River watershed, will enjoy

and bs uds this en new 2s The authors SB n: oud ] I I by not un. dd over 1,000 a

A } } 1 Spe [E

8 I

ecology iated pl ity. As noted by the authors, the Santa Ana River i is one of the largest rivers lying wholly within

Southern California, i dynamic plant communities found in three major segment alluvial scrub segment inland valley segment,

and dosi M Wand

1

E CNN S t I UNITE

Į I

species. The nro to the flora of the Santa Ana River is organized his basis and, therefore, [ ial for developing

a greater understanding of the pl p identification. Further, it is this very focus that makes the book so

broadly applicable to all levels of experti d to al a in UE pe do not padri reside ae the Santa Ana

River auras For tl hat do li ithi a River watershed, tl g k for identifying

f ] und n f species along the ri li ling envi While the

1 Es £T ape | PA Wa n pa E J Soe | J T J T 1 1-271

cies accounts are b cencralla Bue

to visually highlight these key features. To assist E dE in c cue E inu he s have also included a

pli d

"Botany Basics” section and glossary. E

Also of note are the numerous

Appendix 2—“Family Character Tables.” These summary tables, as well as the authors’ look at world phylogeny in Appendix 5 ar

useful and oe pe poe aa of ba book. This su is sure to not "o increase one's cun iur of e d of the

felinda

Melinda

ummary tables throughout e Book spotlighting family and genera B mq me

€ VCI

Santa Ana River

McCoy, Herbarium Volunteer, Botanical Research Institute of Texas, Fort Worth, Texas 76102-4060, USA.

J. Bot. Res. Inst. Texas 1(2): 846. 2007

A NEW SPECIES OF CHUSQUEA SECT. VERTICIEEATAE

(POACEAE: BAMBUSOIDEAE) FROM ECUADOR

Lynn G. Clark, Christopher D. Tyrrell, Jimmy K. Triplett Amanda E. Fisher

Department of Ecology, Evolution and Organismal Biology Department of Biological Sciences

lowa State University Idaho State University

Ames, lowa 50011-1020, U.S.A. Pocatello, Idaho 83209-8007, U.S.A.

ABSTRACT

Chusquea annagardneriae, a species of Chusquea sect. Verticillatae from Ecuador, is Md as new. It is ieee and compared

and contrasted ee Chusqued pd = o to Mo itis most pu CI ished from all other

L o O

species of Ch 1 1 a 5 :

ads ther ÉSTE UBER: 1 T cy 1 n

Į pen mpressed

spikelets with glumes I and II both greatly reduced and scalelike, and glumes III and IV extending about half of the spikelet length.

Chusquea annagardneriae is endemic to the province of Loja, Ecuador, and occurs in dry forest scrub and forest remnants at 2000 to

RESUMEN

Se describe Chusquea annagardneriae, una especie nueva de Ecuador que pertenece a Chusquea sección Verticillatae. Chusquea an-

MA se ilustra y se cds con C. dba la nM más A um Se goers Toen Me de todas las otras

J

lámina de la hoja

P más o menos pee is NUN de ramitas palate en el RM de ramas, una sinflorescencia abierta y

iramidal, gl I It I pd reducidas y escamosas, y glumas III y IV extendiéndose

por la mitad de if longitud de la espiguilla. CI a provincia de Loja, Ecuador, y g

secundaria de bosque seco y en restos de ne a2000a 12250 m.s.n.m.

INTRODUCTION

The native bamboo diversity of Ecuador is considerable and includes seven genera of woody bamboos (Bam-

buseae) and six genera of herbaceous bamboos (Olyreae). Among the woody bamboos of Ecuador, Chusquea

Kunth is the most diverse genus, with an estimated 30 pes of which several are undescribed. In this

paper we describe and illustrate Chusquea Ineriae, one of the Ecuadorian species new to science.

The 30 or so species of Chusquea in MN fall into either Chusquea subg. Swallenochloa (McClure) L.G.

Clark or Chusquea subg. Chusquea. The latter subgenus is subdivided into five sections and a few informal

groups, with representatives of all five of the formally recognized sections known from Ecuador. This new

species clearly belongs to sect. Verticillatae, based on its relatively thin foliage leaf blades, more or less dor-

sally compressed spikelets, and extremely reduced glumes I and I (Clark 1989). Chusquea annagardneriae

joins C. albilanata L.G. Clark, C. perligulata (Pilger) McClure, C. simpliciflora Munro, and C. uniflora Steud.,

the other members of this section in Ecuador. The new species most closely resembles C. albilanata, with

which it is compared and contrasted.

Interestingly enough, the first collections of this species were made only in 1989, and the remaining

collections were made in 1996. Fortunately, both vegetative and flowering specimens are available so that we

have a relatively complete picture of the species, but it does appear to be uncommon. This area of Ecuador,

however, seems to be less well explored botanically so Chusquea annagardneriae may be more widespread

than currently available collections suggest.

TAXONOMIC TREATMENT

Chusquea annagardneriae L.G. Clark, C.D. Tyrrell, Triplett & A.E. Fisher, sp. nov. (Fig. 1). Tyee: ECUADOR.

Loja: road Velacruz-Catacocha, ca. km 6, 2040 m, ca. 4°S, 79°35'W, 11 Mar 1989 (fl), B. Ollgaard, L. Ellemann & B. Eriksen 90959

(HOLOTYPE: QCA!; ISOTYPE: AAUD).

J. Bot. Res. Inst. Texas 1(2): 847 — 851. 2007

848 Journal of the Botanical R h Institute of Texas 1(2)

SE

GEL S NA IN

uS :

J A Cul | f J 7

hes. D. Synflorescence. E. Detail of branch compl t base, showing fibrillar branchl | izes of subsidiary branches. F.

fl G. Detail of culm leaf pex showing pseudopetiolate blade, abaxial view. (A, C, E-G

based on dark & Asimbaya 1437; B, D based o on \Ollgaard et al. 90959).

Clark et al., A pecies of Cl from Ecuad 849

Culmi 2.5-4 m alti, 0.5-1.1 cm diametro, scandentes; Pacis 20-32 cm longa, MN ue Folia dex 24.5-28 cm

lam ntes

longa, ut videtur persistentia; vaginae 22-25cm ongae, 6-10- pl i ngi r quam la pubes

in quarta parte superiore, abaxialiter glabrae, tessellatae pro p | ingulum 0.2-1 mm latum, pubescens; ze. 2.2-4 cm

longae erectae, lanceolatae pl Į 1 laxiali brid abaxialitet glabrae g

; : : A IED ics) EEC Pees 1 D EE EN E : 1 m ES : dal |

HERI 3-5; v 1ae glabra l pilosa

I 3 b L E A TS >

maculatae; laminae 2.9-8.5 cm longae, 0.4-1.3 cm latae, ratio long.:lat. 5-8.5, lanceolatae, adaxialiter glabrae, abaxialiter glabrae,

non tessellatae vel interdum infirmissimus tessellatae, basibus auriculatis interdum cordatis. Synflorescentia 3-9 cm longa, aperta,

pyramidalis; rami primarii infimi 1.5-4.5 cm longi. Spiculae 5-5.7 mm longae, dorsaliter compressae; glumae I et II squamiformes,

minimae; glumae III et IV plerumque ad dimidium longitudem spiculae, acutae, plerumque breviter mucronatae; lemma 4.6-5

longum, subacutum vel breviter mucronatum.

Woody bamboo. Culms 2.5—4 m tall, 0.5-1.1 cm diam., scandent; internodes 20-32 cm long, tuberculate,

more or less terete, glabrous, a narrow band of pubescence often present just below the nodes in new shoots,

but absent from the mature internodes. Culm leaves 24.5-28 cm long (only two complete examples seen),

apparently persistent, juncture of the sheath and blade more or less horizontal; sheaths 22-25 cm long,

6-10 times as long as the blade, adaxially antrorsely scabrid-pubescent for at least the upper one-fourth,

abaxially glabrous, tessellate on the upper two-thirds, margins free, at least the overlapping one ciliate

toward the summit; girdle 0.2-1 mm wide, pubescent; inner ligule ca. 2 mm long, dark brown, abaxially

pubescent, ciliate; outer ligule sometimes visible as a ciliolate rim to 0.5 mm long; blades 2.2-4 cm long,

erect, lanceolate, adaxially antrorsely scabrid, abaxially glabrous, base rounded, narrower than the sheath

summit and thus more or less pseudopetiolate, apex subulate. Nodes at mid-culm with one triangular central

bud subtended by 40—50 subsidiary buds in several rows in a U-shaped crescent, nearly surrounding the

central bud; nodal line dipping slightly below the bud/branch complement; supranodal ridge more or less

pronounced. Branching um subsidiary prn: numerous, of two sizes, usually two larger and

J 1

rebranching, the several to many smaller subsidiary | ginto short, curly fibrillar branchlets

or longer leaf branches. Foliage leaves 3—5 per complement; sheath glabrous or pilose, mottled, only

To

weakly keeled toward the apex, the overlapping margin and summit ciliate; outer ligule a minute ciliolate

or ciliate rim 0.1-0.2 mm long; inner ligule 0.5-0.8 mm long, truncate to rounded or slanted, margin

ciliate; pseudopetiole 0.5-1 mm long, distinct, glabrous; blades 2.9-8.5 cm long, 0.4-1.3 cm wide, L:W

= 5-8.5, lanceolate, adaxially glabrous, abaxially glabrous, not tessellate or occasionally very weakly tes-

sellate in patches, midrib centric, margins glabrous to finely serrulate, base usually auriculate, sometimes

cordate, apex acuminate-subulate. Synflorescences 3-9 cm long, open, pyramidal, well exserted from the

subtending sheath; rachis flattened toward the base, becoming angular toward the apex, scabrous-pubescent;

branches flattened to angular, pubescent, the primary ones strongly spreading to slightly reflexed, pulvinate

at the base, the pulvinus area tomentose to villous, the lowermost primary branches 1.5-4.5 cm long, the

secondary branches and pedicels also spreading; pedicels 1-3 mm long, angular, scabrid-pubescent, often

sinuous. Spikelets 5-5.7 mm long, dorsally compressed; glumes I and II scalelike, no more than 1/25 the

spikelet length, obtuse, glabrous, nerveless, glume I ca. 0.1 mm long, glume II ca. 0.2 mm long; glumes

III and IV usually about Y the spikelet length, triangular, acute, usually shortly mucronate, margins cili-

olate apically, glume III 2-3 mm long, 3-nerved, glume IV 2.2-3.3 mm long, 3- or 5-nerved; lemma 4.6-5

mm long, navicular, subacute to shortly mucronate, pubescent toward the apex; palea 4.6-5.1 mm long,

bimucronate, 2-keeled, sulcate only toward the apex, 4- or 6-nerved, sulcus pubescent toward the apex.

Stamens 3; anthers ca. 2.7 mm long. Lodicules 3, the anterior pair 1-1.2 mm long, asymmetrical, ciliate,

the posterior one ca. 0.9 mm long, symmetrical, ciliate. Fruit unknown.

Etymology.—Chusquea annagardneriae is named for our friend and colleague Anna B. Gardner (1958-

2006) in honor of her contributions to agrostology, particularly to the Grasses of lowa project (Grasses of

lowa 2004) and to bambooillustration, in the form of both line drawings and website development (Bamboo

Denon o

listinguished from any other Chusquea by its auriculate foliage leaf blades (Fig. 1F).

It is ie NE by its more or less pseudopetiolate culm leaf blades, presence of fibrillar branchlets,

subsidiary branches of two sizes, and its delicate open synflorescences that bear relatively small, dorsally

850 Journal of the Botanical R h Institute of Texas 1(2)

Taste 1. A gi | f Ineriae and C. albilanata

Character C. annagardneriae C. albilanata

Culm diameter (cm) 0.5-1.1 1-1.5

Culm height (m) 2.5-4 -

Internode pubescence glabrous band of white, woolly pubescence

0.2-1 cm wide just below the nodal

lines

Culm leaf blade length (cm) 2.2-4 3-74

Culm leaf sheath: blade ratio 6-10 1.7-5.5

Foliage leaf blade length (cm) 2.9-8.5 8-17 (+22)

Foliage leaf blade width (cm) 0.4-1.3 1.8-3.7 (4

Foliage leaf blade base auriculate to rarely cordate rounded to od rounded

Foliage leaf sheath pubescence overlapping margin and summit ciliate glabrous

o length (cm) 3-9 11-15

ost branch length (cm) 1.5-4.5 7-9

Vd secondary dry forest scrub and montane forests, secondary

remnant forest patch vegetation

Distribution s. Ecuador c. and n. Ecuador, Colombia

compressed spikelets. Another unusual feature is the presence of an outer ligule in at least some of the culm

leaves, but since so few complete examples were seen, it is not clear how consistent this character might

be.

This species is known from only five collections from the province of Loja in southern Ecuador, where

it persists in secondary scrub from bosque seco (dry forest), around pastures and on rocky slopes, and at the

edges of forest remnants at elevations of 2000 to 2250 m. Based on current knowledge, C. annagardneriae

is endemic to Loja, of restricted distribution, and uncommon if not rare. It may, however, be more widely

distributed in Loja and neighboring El Oro province, and could perhaps be expected in northern Peru.

Among the Ecuadorian Chusqueas, C. annagardneriae is one of three species with fibrillar branchlets

present in the branch complement (Fig. 1E); Chusquea lehmannii Pilger and C. albilanata are the other two.

A fourth species, Chusquea scabra Soderstr. & C. Calderón from Costa Rica, also has fibrillar branchlets that

are caused by an ascomycetous fungus (Soderstrom & Calderón 1978) and we presume that this is true for

the three other species as well. Despite the presence of fibrillar branchlets, these species do not appear to

be closely related except for the presumed sister relationship between C. annagardneriae and C. albilanata.

Among the species of Chusquea sect. Verticillatae in Ecuador, C. annagardneriae and C. albilanata share

tuberculate internodes, the presence of fibrillar branchlets in the branch complements, open pyramidal

synflorescences bearing small delicate spikelets, and spikelets with glumes III and IV about half the spikelet

length. Besides the distinctive auriculate foliage leaf bases of C. annagardneriae, the two species differ in

several features summarized in Table 1. Most notably, mature internodes in C. albilanata retain a band of

hair just below the nodal line and the foliage leaf blades are larger. The synflorescences of C. albilanata are

also larger overall and the lowermost branches are longer than those of C. annagardneriae.

The two 1989 collections were made from flowering plants and one of the 1996 collections (Clark &

Asimbaya 143) was also made from a flowering plant, although the synflorescences were not mature. The

other two 1996 collections, which were from the same area, were completely vegetative. This suggests that

either C. annagardneriae has a short flowering cycle of approximately seven years, with one population simply

not flowering yet (or perhaps out of synchrony with the others) or this species is a sporadic bloomer.

Anu

Additional specimens examined. ECUADOR. Loja: 7.7 km SE of Velacruz and 11.3 km NE of Catacocha, 2150 m, 4%01'S, 79°32'W,

12 Feb 1996 (fD, L. Clark & P. Asimbaya 1436 (AAU, ISC, LOJA, MO, Q, QAP, QCA, QCNE, US); 2 km SW of Velacruz on the road to

Catacocha, 2170 m, 3°59'S, 79°35'W, 12 Feb 1996, L. Clark & P. Asimbaya 1437 (AAU, ISC, LOJA, MO, Q, QAP, QCA, QCNE, US); ca. 5

E of Velacruz, road Velacruz-Catamayo, 2250 m, 3?58'S, 79°32'W, 12 Feb 1996, L. Clark & P. Asimbaya 1438 (AAU, ISC, LOJA, QCA,

QCNE, US); carretero Velacruz-Olmedo, Km 18, 2050 m, 3°55-59'S, 79°34-38'W, 11 Mar 1989 (fl), A. Freire Fierro 1270 (QCA).

Clark etal., A pecies of CI from Ecuad 851

ACKNOWLEDGMENTS

Field work in Ecuador by Clark was supported by National Science Foundation (NSF) grant DEB-9218657

and was conducted under the auspices of the Pontifícia Universidad Católica del Ecuador (Herbario QCA).

Clark thanks the staff of QCA for their generous assistance and for access to the herbarium. Final preparation

of the manuscript was supported by NSF grants DEB-0515712 to Clark and DEB-0515828 to S. Kelchner

(Idaho State University). We thank Anna B. Gardner for the fine illustration of this species.

REFERENCES

BAMBOO Biopiversity. 2005. http;//www.eeob.iastate.edu/research/bamboo/index.html

CLARK, L.G. 1989. Systematics of Chusquea section Swallenochloa, section Verticillatae, section Serpentes, and

section Longifoliae (Poaceae: Bambusoideae). Syst. Bot. Monogr. 27:1—127

Grasses or lowa. 2004. http://www.eeob.iastate. edid

SODERSTROM, T.R. and C.E. CALDERÓN. 1978. Chusquea and Swall (Poac Bambusoid g icrelation-

ships and new species. Brittonia 30:297—312.

852 Journal of the Botanical R h Institute of Texas 1(2)

BOOK REVIEWS

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Once again Paul Martin Brown and partner artist Stan Folsum have produced a gem of a regional flora. Readers wh familiar with

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as you go searching for those plants in situ

both experienced and novice seekers of the wild orchid in its natural habitat, spotting and identifying the plant becomes

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1 1 T Ery pe es E: T 1 Es 1 TNS LI S | 1 : A 1 1 1 1

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necessary to separate and eee the plants. Even first time venturers into the field on peri gl d excit t

to become avid enthusiasts

With the upcoming holidays, this exquisite volume is an excellent gift choice.—Helen Jeude, Botanical Research Institute of Texas,

Fort Worth, TX 76102-4060, U.S.A.

ARTHUR M. SHAPIRO. 2007. Field Guide to the Butterflies of the San Francisco Bay and Sacramento

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One would expect excellence in a work by this well-known and accomplished lepidopterist, and, indeed, this field guide is among the

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is field guide can be recommended for any one interested in butterflies whether or not one frequents the regions covered. It is

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J. Bot. Res. Inst. Texas 1(2): 852. 2007

ESTUDIOS EN LAS ACOCTNACEAE NEOTROPICALES XXX:

TRESNUEVAS ESPECIES ANDINAS:DE

MANDEVILLA (APOCYNOIDEAE: MESECHITEAE)

J. Francisco Morales

Instituto Nacional de ud (INBio)

Apartado Posta

Santo Domingo, idm ee RICA

RESUMEN

Se describen Mandevilla mequatonaliss M. pieds y M. lejana: tres nuevas especies de los Andes de Ecuador y Perú. Se incluyen

descripciones, ilustracion las especies más afines.

PALABRAS CLAVE: Ápocynaceae, Apocynoideae, Mesechiteae, Mandevilla

ABSTRACT

Mandevilla aequatorialis, M. niam 2 M. lojan, ME new Vp from the Andes of Ecuador and Peru, are described and il-

lustrated and their relationshi

L

La concepción y circunscripción del género Mandevilla Lindl., el más grande de la subfamilia Apocynoi-

deae, ha variado notablemente desde la última monografía del género (Woodson 1933). En aquel entonces,

Mandevilla se trató como una unidad genérica distinta de Macrosiphonia Müll. Arg., con alrededor de 120

especies. Pichon (1948) no aceptó dicha propuesta taxonómica y propuso unir de nuevo ambos géneros,

lo cual prácticamente no fue seguido y paso desapercibo en afios subsecuentes. Zarucchi (1991) propuso

el género monotípico Quiotania, un grupo similar en varios caracteres morfológicos a Mandevilla, del que

difería principalmente por el tamaño de las corolas. En años posteriores, Henrickson (1996), elevó a nivel

genérico una de las dos secciones de Macrosiphonia, proponiendo el género Telosiphonia, grupo restringido

principalmente a México y el S de Estados Unidos. De esta manera, Mandevilla se volvió un género con alre-

dedor de 3 géneros satélites cercanamente relacionados (Macrosiphonia, Quiotania, Telosiphonia) y otros cuya

relación aparentemente no era tan estrecha (Allomarkgrafia Woodson, Mesechites Múll. Arg., Tintinnabularia

Woodson). Como resultado de estudios cladísticos derivados del análisis de varios indicadores molecula-

res y morfológicos, Simóes et al. (2004, 2006, 2007) redefinieron los límites genéricos de Mandevilla, de

manera que Macrosiphonia, Quiotania y Telosiphonia fueron reducidos a la sinonimia. Esta propuesta, elevó el

nümero de especies de Mandevilla a más de 170, convirtiéndolo en el género más grande de las Apocynaceae

neotropicales. La evidencia filogenética provista en los trabajos de Simões et al. (2004, 2006) demostró que

la clasificación infragenérica de Woodson (1933) es altamente parafilética, mientras que la propuesta por

Pichon (1948) es monofilética. Sin embargo, la mayoría de los taxones infragenéricos propuestas por ambos

taxonómos son polifiléticos, por lo que se necesita mayor evidencia y un estudio más detallado para poder

establecer una nueva clasificación intragenérica.

Continuando con el desarrollo de una nueva monografía de Mandevilla, se describen tres nuevas espe-

cies, restringidas a los Andes de Ecuador y Perú.

Mandevilla aequatorialis J.E Morales, sp. nov. (Fig. 1). Tiro. ECUADOR. Imsasura: Cotacachi, parroquia Plaza Gutiérrez,

Tabla Chupa, 12 May 1992 (fl), G. Tipaz et al. 976 (notoriro: MOI, fotografía, INB!; isoripo: QCNE!).

A Mandevilla jasminiflora Woodson cui affinis, foliis basi cordatis, sepala longiribus, et antheris auriculatis acuminatis differt.

Liana con tallos subleñosos, ramitas aplanadas cuando jóvenes, subcilíndricas en la madurez, moderada

y diminutamente puberulentas cuando jóvenes, el indumento más esparcido en tallos viejos, los coléteres

J. Bot. Res. Inst. Texas 1(2): 853 — 857. 2007

854 Journal of the Botanical R h Institute of Texas 1(2)

interpeciolares cortos e inconspicuos, cónicos, ca.

1 mm de largo, formando una línea interpeciolar

inconspicua. Hojas: pecíolos 10-23 mm de largo;

láminas 2,5-6,3 x 1,1-2,8 cm, ovadas a ovado-

elípticas, firmemente membranáceas, el ápice

gradualmente acuminado o agudo, la base leve o

conspicuamente cordada, glabras en la superficie

adaxial, glabrescentes a densa y diminutamente

puberulentas en la superficie abaxial, sin doma-

cios a lo largo del nervio central en la superficie

abaxial, coléteres del nervio central en grupos

3 6 4, ca. 1 mm de largo, agrupados en la base

del nervio, venas secundarias más o menos ar-

cuadas. Inflorescencias más larga o igualando

en longitud las hojas adyacentes, axilar, densa a

moderadamente puberulentas, con muchas flores,

pedúnculo 14-28 mm de largo, pedicelos 8-10

mm de largo, brácteas 1-3 x 0,5-1 mm, escariosas;

sépalos 3-5 x 1-1,5 mm, angostamente ovados,

acuminados, glabrescentes, a veces con unos cu-

antos pelos esparcidos cerca de la región apical y

a lo largo del nervio medio, con 2 6 3 coléteres en

la base de la cara adaxial, ubicados cerca de los

márgenes, ca. 0,5 mm de largo, irregularmente

lacerados apicalmente; corola hipocrateriforme,

verde amarillenta, glabra exteriormente (excepto

por la cara adaxial de los lóbulos), los cinco lóbu-

los con la aestivación dextrorsa, tubo 3-4 mm de

largo, 2 mm en diámetro en la boca, recto y no

inflado, con una corona anular rudimentaria y

con prolongaciones basales sobre cada lóbulo de la

Fic. ialis. A. Ramit fl las. B. Detallede corola, ca. 0,5 mm de largo o menos, lóbulos 2,5-4

las uina del nervio vio cent Vito. C C. oe dab dide x 2-2,5 mm, angostamente obovados, extendidos,

a adaxia con dos lobDulos

yd e pero algo reflexos distal y marginalmente, densa

Gineceo, most-

ando las anteras, disco y ovario. e Detalle de la cabeza estigmática (G. — Y diminutamente papilado-tomentulosos en la

mam 976, MO). superficie adaxial; estambres insertos en la mi-

Y

E

tad distal, anteras ca. 3 mm de largo, glabras, los

ápices exsertos, la base sagitada, ovario ca. 1 mm

de largo, glabro, estilo recto, cabeza estigmática 1,5-2 mm de largo, el apículo cerca de 1-1,4 mm de largo;

nectario variada e irregularmente pentalobulado, más corto que el ovario. Folículos desconocidos.

Distribución, habitat y ecología.—Esta especie es endémica a Ecuador, donde se conoce de los departa-

mentos de Pichincha e Imbabura, donde crece en bosques húmedos montanos, en elevaciones de 2300-3000

m. Floración ocurre en mayo y septiembre.

Mandevilla aequatorialis es morfológicamente algo similar a M. jasminiflora Woodson, pero se puede

separar fácilmente por sus hojas levemente cordadas basalmente (vs. redondeadas a obtusas), láminas foliares

con las venas secundarias arcuadas (vs. dispuestas ca. en ángulo recto en relación al nervio central, flores

con los sépalos más largos (3-5 x 1-1,5 mm vs. 1-1,5 x 1-1,5 mm) y anteras con las aurículas estériles

sagitadas, largas y cortamente acuminadas. El epiteto de este nuevo taxón hace referencia al hecho de que

la especie es conocida por el momento solo en Ecuador.

Morales, NM H A M a HII A E Al y Perú 855

Especímenes adicionales examinados. ECUADOR. Pichincha: Quito, carretera Nono-Guarumos-Tandayapa, ent y Cerro

Guarumos, 1 Sep 2001 (fD, A. Freire-Fierro et al. 3143 (QCNE)

Mende frigida J.E Morales, sp. nov. (Fig. 2). Tiro. PERÚ. Piura: Cerro Aypate, E de Ayabaca, 22 Sep 1991 (fl), A. Gentry

al. 74988 (HoLoTIro: MO}, fotografía, INB!; isoriros: INB!, USM!).

A Mandevilla assimilis (K. Schum.) J.F. Morales, cui affinis, foliis glabris, subtus in axillis nervi medii foveolatis et antheris 3,5-4 mm

longis differt.

Liana, ramitas inconspicusmente aplanadas cuando jóvenes, cilíndricas o subcilíndricas cuando viejas,

glabras; coléteres interpeciolares i | ca. 1 mm de largo. Hojas: pecíolos 5-8 mm de largo; láminas

2,2—5,4 x 1,2-2,4 cm, elípticas, e membranáceas, abruptamente corto-acuminadas apicalmente,

obtusas a anchamente cuneadas apicalmente, glabras, usualmente con domacios a lo largo del nercio central

en la superficie abaxial, coléteres del nervio central en grupos de 1 ó 2, ca. 1 mm de largo, agrupados en la

base del nervio, venas secundarias más o menos arcuadas. Inflorescencia igualando las hojas adyacentes,

axilar, algunas veces pareciendo terminal o subterminal tallos jóvenes, glabras, con pocas flores, pedún-

culo 35-47 mm de largo, pedicelos 10-14 mm de largo, brácteas 3,5—4,5 x 0,5 mm, escariosas; sépalos

2,5-3,5(-4) x 1-1,5 mm, angostamente ovados, agudos, muy diminuta e inconspicuamente puberulentos,

coléteres enteros, subenteros o apicalmente lacerados; corola hipocrateriforme, blanca, muy diminuta e

inconspicuamente puberulenta exteriormente, tubo 10-11 mm de largo, 2-2,5 mm en diámetro en la boca,

abultado sobre la posición de los estambres, lóbulos 8-9 x 2 mm, angostamente elípticos, erectos a suber-

ectos, diminutamente puberulentos por ambas caras; estambres insertos en ca. la mitad de la longitud del

tubo de la corola, anteras 3,5—4 mm de largo, glabras, incluidas, la base subauriculada; ovario ca. 1,5 mm

de largo, glabro, estilo levemente curvado, cabeza estigmática 2,5-3 mm de largo, el apículo ca. 2 mm de

largo, nectarios 2, ca. 1 mm de largo. Folículos desconocidos.

Distribución, habitat y ecología.—Endémica a Perú, donde es conocida únicamente de la localidad tipo, en

bosques montanos nubosos, en elevaciones de 2700-2800 m. Especímenes con flores han sido recolectados

en septiembre.

Mandevilla frigida se encuentra relacionada con M. assimilis (K. Schum.) J.F. Morales, conocida de Ec-

uador y Perú, de la que difiere por sus láminas foliares con la superficie abaxial glabra y con domacios a lo

largo del nervio central (vs. tomentulosa a densamente puberulenta y sin domacios) y anteras más cortas

(3,5-4 mm vs. 4,9—5,2 mm). El epiteto de este nuevo taxón hace referencia al hecho de que la especie crece

en sitios fríos de elevaciones altas de la Cordillera de los Andes.

Mandevilla lojana J.E Morales, sp. nov. (Fig. 3). Tro. ECUADOR. Loja: Cerro Villanaco, carretera Loja-La Toma, 17 Dic 1988

(fl, fr), J. Madsen 75938 (HoLotipo: QCA!, fotografía, INB!; isoriro: QCNE!, fotografía, INB!).

A Mandevilla ligustriflora Woodson, cui affinis, tubo longiribus et corollae lobis 5-7 mm longis differt.

Liana, ramitas tiernas aplanadas y anguladas, cilíndricas o subcilindricas cuando viejas, glabras; coléteres

interpeciolares inconspicuos, ca. 0,5 mm de largo. Hojas: pecíolos 4-8 mm de largo, láminas 3,5-5,6 x 1-1,8

cm, elípticas a obovado-elípticas, firmemente membranáceas, agudo-mucronuladas apicalmente, cuneadas a

obtusas basalmente, glabras en ambas superficies, usualmente con domacios a lo largo del nervio central en

la superficie abaxial, coléteres del nervio central en grupos de 2, ca. 0,5 mm de largo, agrupados en la base

del nervio, venas secundarias más o menos arcuadas. Inflorescencias más corta que las hojas adyacentes,

axilar, glabras, con muchas flores, pedúnculo 27-29 mm de largo, pedicelos 14-16 mm de largo, brácteas 1

x 0,5-1 mm, escariosas; sépalos 2 x x 1 mm, ovados, agudos, glabros a glabrescentes, coléteres irregular-

Jem Top.

mente lacerados; corola me, blanca, diminuta e inconspicuamente puberulenta

exteriormente, parte EM M del tubo de la corola 3-4 x 2 mm, parte superior angostamente cónica, 6-6,5

mm de largo, 5-6 mm en diámetro en la boca, corona reducida a vestigios coronales, leve y tenue, lóbulos

5-7 x 2,5-3,5 mm, ovados, erectos a suberectos, diminutamente puberulentos en ambas superficies; estam-

bres insertos en la base de la parte superior del tubo, anteras ca. 4,5 mm de largo, glabras, incluidas, la base

subauriculada, ovario ca. 1 mm de largo, glabro, estilo recto o levemente doblado, cabeza estigmática 33,2

856 Journal of the Botanical R h Institute of Texas 1(2)

mm de largo, el apículo 2-2,2 mm de

largo, nectarios dos, casi tan largos

como el ovario. Folículos 12,5-25 x

0,2-0,4 cm, levemente moniliformes,

glabros; semillas 10-11 mm de largo,

glabras, coma 7-14 mm de largo,

crema.

Distribución, habitat y ecología.—

Esta especie es endémica al S de

Ecuador, conocida por el momento

únicamente en la localidad tipo, en

bosques montanos nubosos y veg-

etación paramosa, en elevaciones

de 2500 m. Especímenes con flores

y frutos han sido recolectados en

diciembre.

Mandevilla lojana está cerca-

namente relacionada con M. ligus-

triflora, pero se puede separar por

sus flores con sus partes (sépalos,

tubo, lóbulos, anteras) más largas y

grandes, así como por sus hojas con

los domacios distribuidos a lo largo

del nervio central en la superficie

abaxial (vs. restringidos a la base

del nervio). El epiteto de este nuevo

taxón honra a la provincia de Loja,

sitio peculiar e interesante en Ecua-

dor de la que numerosos endemismos

son conocidos.

AGRADECIMIENTOS

Quiero agradecer a los curadores de

los herbarios COL CUVC- HUA,

Fic 2. Mandevilla frigida AR it infl ias. B. D | b i à HUQ, JAUM, K, MEDEL, MO, Q,

leni | |. C. Detalle delos d D. Cáliz, pedicelo, bráctea y E H i E

m uu uu OG OCNE TOPES SUS e

corola. E Sé| | vista adaxial. F. Tul

lla " E

7

| Icañalad flecha). G. Antera, por permitirme facilidades para el

estilo, ovario y nectario. H. Detalle de la cal tigmática (Gentry et al. 74988, MO). uso de colecciones. Asimismo, se

J

agradece al personal del herbario de

la Universidad de Antioquia (HUA),

por el apoyo logistico continuo brindado durante mis multiples visitas a Medellin, Colombia, asi como por

la asistencia en el trabajo de campo desarrollado. Se agradece también grandemente a Steffen Matezki, de la

Universidad de Bayreuth, Alemania, por suministrar y autorizar el uso de las fotografias de M. ligustriflora.

La asistencia de las siguientes personas en facilitar la visita de sus respectivos herbarios es grandemente

agradecida: Alberto Agudelo (HUQ) en Armenia, Quindio, Colombia, Homero Vargas (QCNE) en Quito,

Pichincha, Ecuador, Philip Silverstone-Sopkin (CUVC) en Cali, Valle del Cauca, Colombia y Ricardo Callejas

(HUA), en Medellin, Antioquia, Colombia.

M H A M PI HII A E Al

Morales

Fic. 3 Mandevillalojana A. Ramit infl ias. B. D

del i tral enl fi laxial. C. Detall | ios. D. Sépalo y coléter,

vista adaxial. E. Caliz, pedicelo, bractea y corola. F. Tubo dela corola abierto, mostrando las

Hu * H rj A Y J £ Ļ Y G n Hu Jal L : fet

H. Anteras, estilo, ovario y nectarios. I. Folículos. J. Semillas (Madsen 75938, QCA).

857

REFERENCIAS

HENRICKSON, J. 1996. Studies in Macro-

siphonia (Apocynaceae): generic

recognition of Telosiphonia. Aliso

14:179-195.

PicHoN, M. 1948. Classification des Apocy-

nacées. X. Genre Mandevilla. Bull. Mus.

Hist. Nat. (Paris), sér. 2, 20:211-216.

Simoes, A.O., M. ENDRESS, T. VAN DER NIET, L.

KINOSHITA & E. Conti. 2004. Tribal and

intergeneric relationships of Mes-

echiteae (Apocynoideae, Apocyna-

ceae): evidence from three not icoding

pastid DNA regions and morphology.

Amer. J. Bot. 91:1409-1418.

Simoes, A.O., M. ENDRESS, T. VAN DER NIET, L.S.

KINOSHITA & E. Conti. 2006. Is Mandevilla

(Apocynaceae, Mesechiteae) mono-

phyletic? Evidence from five plastid

DNA loci and morphology. Ann. Mis-

souri Bot. Gard. 93:565-591.

Simoes, A.O., L.S. KINOSHITA & M. ENDRESS.

2007. New combinations in Mandev-

illa Lindley (Apocynaceae). Novon

17:89-90

Woobson, R.E. 1933. Studies in the Apocy-

naceae. IV. The American genera of

Echitoideae XXVI. Ann. Missouri Bot.

Gard. 20:605-790.

ZARUCCHI, J.L. 1991. Quiotania: a new

genus of Apocynaceae-Apocynoi-

deae from Northern Colombia. Novon

1:33-36.

858 Journal of the Botanical R h Institut

f Texas 1(2)

BOOK REVIEW

GERHARD GOTTSBERGER and ILSE SILBERBAUER-GOTTSBERGER. 2006. Life in the Cerrado: a South American

Tropical Seasonal Vegetation Vol 1. Origin and Structure, Dynamics and Plant Use. (ISBN 3-

00-017928-3, hbk.). Reta Verlag, Ulm, Germany. (Orders: ilse.silberbauer@uni-ulm.de; Botanischer

Garten und Herbarium, Universitaet Ulm, Hans-Kreb-Weg o.n., D-89081 Ulm, Germany). $64.60, 280

pp., numerous color figures, maps, ques and tables, 29.5 x 22.5 em.

The book Life in the Cerrado: a South American Ti ER 1E o eu 1

P sd O.

the following: definition of a cerrado, location soil types, origin of ph g diversity of flora life history, and natural and human

influences. This book is full of very c olorful ima ges, maps and charts that cl l expla in the flora found in the cerrado.

The authors begin by defining B term as a Pd x um type that is drastically different from the vegetation found in a

typical savanna; these differ | ition, and fl

. This vegetation is mostly

found in central Brazil and has changed contac ened nS. The soil composition varies and has very poorly developed

mineral composition. Cerrado soils are well drained, and soil moisture is found to mace influence the ees of a

hera are five basi do f , distinguishable by vegetation height and density. In order from t

these are: cerradáo, cerrado sensu stricto, campo cerrado, canes sujo, and campo limpo. ee all five forms exhibit some of i

e domi

m nant species, floristic diversity of th varies o locality. Differences can occur over relatively short distances,

often abruptly changing from open to more closed canopy mt ed: of non- pod cad nd bs cerrado also occurs,

sometimes forming ecotones or transitional areas in the process. T

alc needed 11

order to fully understand t I iti and geographical r gi ns of th rrad

The noe of the cerrado consists of trees, lianas, palms, VA vines, ps au few vascular epiphytes. Trees often

appear stunted k, fissured, and corky bark and are adapted for rapid g lopment. piel ay appear quite

old, but longevity of fth I 30-40 years. A g | f other pl l hil d

life, many with thicl ] i Į water ODD and me high aluminum content of the soil

Ecological and cli f hat infl 1fl il ilability, fire, frost, and seasonality.

Moisture is of found until a soil depth of 2.0— de meters o iain, longer this water dur-

ing the dry season. To combat the nutrient-deficient soil, other pl I f f nitrog ] phospl

Fire pow a Pp 2 influence on the NA 2 deterring non-native or p tablish t ] fl

aiding in g g f dlings. Frost is very similar to fire in that i damage plant

but l l i getation with tl tion of new leaves. Fire is found throughout th do f

ti idered more of a local eS The region ane just two seasons: dry and wet. Flowering and fruiting of

ee dd die is determined by he parti and nati ue Sad on these seasonal patterns.

lly i I g There are many ful I l ] aluable vood, fruits ani

seeds, oils, cork, fiber, and pigments. However, th ] ion tl hi ded i s to peed understand the

medicinal uses of various plants and the potential economic resources available. Adding to its overall value and importance, this area

ee also been es asa is che ola dis au motel i eps has drastically decreased in size over the last decade. The

1

F population.

This was a very i ing ee o ighly ] i li ical

1

or pi 1 1 € 1 j 3l I 1 1 fuli charts

tables, and maps to help better explain the cerrado. The "d will take away a better understanding of the specific vegetation found

in this region and the natural and anthropogenic influences that have effected its growth.—Keri McNew, MS Biol

Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A

ogy, Project Manager,

J. Bot. Res. Inst. Texas 1(2): 858. 2007

ESTUDIOS EN LAS APOCYNACEAE NEO TROPICALES XXXI: EL COMPLEJO DE

MANDEVIELA HIRSUTA Y CUATRO NUEVAS ESPECIES

J. Francisco Morales

Instituto Nacional de o (INBio)

Apartado Postal 22

Santo Domingo, Heredia, a RICA

RESUMEN

a] Landevilla hi d ib orrida (Perú), M. inexperata

(Ecuador), M. megabracteata (Colombia y Guyana) y M. similaris (Venezuela). Se A ilustraciones, descripciones, datos de

distribución, especímenes examinados y se comentan las relaciones con los taxones más relacionados.

PALABRAS CLAVE: Apocynaceae, Apocynoideae, Mesechiteae, Mandevilla

ABSTRACT

i ENEA y P Pes 1 f,

Following a study of the M

w species are described: M. horrida (Perú), M. Mic Adm (pruden. M.

megabracteata (Colombia and Guyana) and M. Tu (Venezuela). Illustrations, descriptions, distributi

ined, as well as discussions about closely related taxa are included

Mandevilla Lindl. (Apocynoideae, Mesechiteae), es el género más grande de las Apocynaceae con más de

170 especies distribuidas desde México y las Antillas hasta el N de Argentina. La percepción del mismo ha

cambiado notablemente desde la última monografía (Woodson 1933), ya que varios de los géneros cercana-

mente relacionados y considerados como grupos distintos desde hace décadas (e.g., Macrosiphonia), no han

sido apoyados por estudios moleculares recientes (Simoes et al. 2004, 2006, 2007).

En términos general ndevilla es un grupo complejo por varias razones, principalmente por la gran

cantidad de taxones que lo conforman, la similaridad vegetativa de algunas especies, así como por la alta

variación intraespecífica presentes en algunos grupos, en cuanto a la densidad y distribución del indumento

en estructuras florales y vegetativas se refiere.

Mandevilla hirsuta (Rich.) K. Schum. es la especie con el mayor rango de distribución geográfica en el

género, el cual se extiende desde el S de México hasta Brasil, Bolivia y Paraguay (Morales 2005). Esta especie

presenta una alta variación intraespecifica en cuanto a la densidad del indumento de sus partes vegetativas,

de manera que existen especimenes con hojas densamente hirsutas en su superficie abaxial (Meneses et

al. 693, INB, LPB), hasta otros en los cuales el indumento esta casi ausente y las hojas son prácticamente

glabras (Melo et al. 827, INB, UB). Sin embargo, existen una serie de variaciones o estados intermedios que

no permiten reconocer variedades basado en la densidad del indumento. A pesar de esta variabilidad, los

caracteres florales son relativamente constantes, de manera que es posible reconocer fácilmente una sola

unidad taxonómica en todo el rango de distribución de la especie.

1 * a

A través del tiempo y basado en la variabilidad del indumento, otras esp con ese taxón,

han sido confundidas como M. hirsuta, grupo en el cual podemos distinguir a M. bracteata (Kunth) Kuntze,

M. krukovii Woodson, M. lancibracteata Woodson, M. moritziana (Múll. Arg.) Donn.Sm., M. sagittarii Woodson

y M. steyermarhii Woodson. Este grupo, el cual conoceremos como el complejo de M. hirsuta, ende en

ies Je] 186 (A beg al

común inflorescencias con brácteas foliáceas o subfoliáceas, corolas i mes y

moniliformes. En general, este complejo no ha sido estudiado con detalle y estudios preliminares recientes

han revelado varias novedades taxónomicas (e.g., Morales 2005, 2006)

Como resultado de una revisión detallada del complejo de M. hirsuta, se proponen cuatro nuevas espe-

a]

cies. Adicionalmente, se brinda la sinonimia completa de M. hirsuta, incluyend ipciones, ilustraciones

y datos de distribución para cada taxón.

J. Bot. Res. Inst. Texas 1(2): 859 — 869. 2007

860 Journal of the Botanical R h Institute of Texas 1(2)

Una clave para las especies del subgénero Exothostemon con brácteas florales foliáceas se brinda a con-

tinuación. Algunos datos de esta clave han sido tomados de una nueva monografía de Mandevilla (Morales,

datos sin publ.) por lo que pueden diferir de otros publicados anteriormente (e.g., Woodson 1933).

1. e de la corola hipocrateriforme

Lóbulos de | la bl la gargant illa; láminas foli 7-16 mm de ancho, las venas ebd

usualmente no evidente aridana J.F. Morales

Lóbulos de la a amarillos, la garganta usualmente ele a anaranjada; laminas foliares 15- - mm

anchor las venas terciarias evidentes al menos abaxialm

3. Corola muy diminuta y es id pul | lal glal te ext te, la pubescen-

E | | Al P | nen 15) a

os glabros M. rugellosa (Rich.) L. Allorge

Ps pilosa a pilosulosa externamente; inflorescencia densa a moderadamente pilosulosa; folículos

rcidamente hirsutos a da M. villosa (Miers) Woodson

1. Tubo de la po infundibulifo

4. Láminas foliares prem. con E base obtusa, cuneada a redondeada _M. javitensis (Kunth) K.Schum.

4. Láminas foliares usualmente con la base cordada a subcordada.

5. Hojas sésiles a subsésiles, con el peciolo 1-2 mm de largo M. tristis J.F. Morales

5 Hojas eee [eco neus eon el pue a " d mm de largo.

GA d a lo largo del nervio central y de las

venas secundarias en la superficie adaxial sépalos 1-2 mm de largo M. moritziana

(Müll.Arg.) Donn.Sm.

6. Láminas foliares no bulladas; coléteres distribuidos solo a lo e ac) nervio SO o si ocasional-

mente presentes en venas secundarias (M. lancibracteata) | del tub

de 9 mm de largo; sépalos 2,5-18 mm de largo

7. Parte inferior del tubo de la corola 7-14 mm.

8. Sépalos 7,5-9 mm de largo, parte inferior del tubo de la corola 12-14 mm de largo

M. ar

EM

rales

8. Sépalos 3-3,8 mm de largo, parte inferior del tubo de la corola 7-9 mm de largo ____M. E

7. Parte inferior del tubo de la corola 18-35 mm.

9, 7- lar mm de algo:

in tal s5 Q l | pall sl

puberulentos cuando jóvenes, glabrescentes con la aes M. similaris J.F. Morales

10. Venas terciarias evidentes; jeudi | hirsutos, híspidos

a hispidulosos

i Brácteas florales i 681 mm de Tu M. megabracteata J.F. Morales

. Brácteas florales 11-25 mm de largo.

12. Parte superior a tubo de la corola 15-21(-25) mm de largo, con un diámetro de

(1 6-21 mm en el orificio; folículos libres, solo unidos en el ápice — — M. hirsuta

(Rich) K. Schum.

12. Parte superior del tubo de la corola 27-31 mm de largo, con un diámetro de

7-9 mm en el orificio; folículos fusionados a lo largo de su entera longitud _ M. sagitarii

Woodson

9. Sépalos 2,5-6 mm de largo.

13. Tallos con una conspicua línea interpeciolar en cada nudo; ramitas jóvenes pelea

glabrescentes, el indumento no evidente steyermarkiiWoodson

13. Tallos careciendo de una conspicua línea interpeciolar; ramitas hirsutulas, psa pu-

berulentas a híspidas

14. Láminas foliares con el ápice obtuso-mucronado a redondeado-mucronado; tubo de

la corola con un diámetro de 14-22 mm en la fauce — — — M. bracteata (Kunth) Kuntze

14. Láminas foliares con el ápice caudado-acuminado, corto-acuminado o cortamente

cuspidado; tubo de la corola con un dpi. de En 3 mm en la fauce

15. Tallos esp | li lentos; brá ae 17-28

mm de largo; Mit inferior del bo 30- 357 mm de largo ___ la Woodson

15. Tallos hispidos; bracteas florales 32-46 mm de largo; parte inferior del tubo 2

5-23 mm de largo M. ais J.F. Morales

M H dal | PA pee | J sila hi + 861

Morales

Mandevilla hirsuta (Rich.) K. Schum., Nat. Pflanzenfam. 4(2):171. 1895. (Fig. 1). aa Du Rich., Actes Soc.

Hist. Nat. Paris 1:107. 1792. Echites richardi Roem. € Schult., Syst. Veg. 4:391. 1819, nom. illeg. Ambl ) Miers

Apocyn. S. Amer. 185. 1878. Tiro: GUYANA FRANCESA. Cayenne, 1792 (fl), Leblond s.n. (Lectotiro, designado por Allorge-Boiteau

(1998), P-LA [foto, INB]; isoLecToTIPOS, C, G-DC [foto F neg. 26863], P [2 cartulinas]).

Echites tomentosus Vahl, Symb. Bot. 3:44. 1794. Ed tomentosa du Miers, pun S. Amer. 213. 1878. Mandevilla tomentosa

(Vahl) Kuntze, Revis. Gen. Pl. 2:416. 1 M K. Schum., Nat. Pflanzenfam 4(2):171. 1895, non. inval. Tipo:

GUYANA FRANCESA: Cayenne, fecha ined (fl), von Rohr 63 (HoLotipo: C [2 usum isoriPOs: BM [2 cartulinas, fotos, INB]).

us UE Nov. Gen. 2: ee ed. e 3:218. 1818 [1819]. Exothostemon macrophyllum (Kunth) G. Don, Gent. Hist. 4:82.

Sc at. Pflanzenfam 4(2):171. 1895. Tiro. VENEZUELA. Amazonas: río Orinoco, cerca

Echites ibn e

1837.)

n Borja, Carchi Isla Panunamae, May año A fl), Humbol i ER 1214 (HoLotiro: P-HB; isotirO: B, destruido).

m campestris Vell., Fl. Flumin. 113. 1829. Amblyanthera campestris (Vell.) Müll.Arg., Fl. Bras. 6(1):149. 1860. Rhabdadenia camp-

estris (Vell.) Miers, Apocyn. S. Amer. 121. 1878. Tipo: BRASIL. Rio DE JANEIRO: Vell., Fl. Flumin., Icon 3:tab. 43. 1827 (uecroripo,

esignado aquí).

Echites hispidus Willd. ex Roem. & Schult., Syst. Veg. 4:795. 1819. Amblyanthera hispida (Willd. ex Roem. € Schult.) Müll.Arg., Fl.

1):147. 1860. Mandevilla hispida (Willd. ex Roem. & Schult.) Hemsl., Biol. Cent.-Amer., Bot. 2(10):316. 1881. Mandevill

tomentosa var. hispida (Willd. ex Roem. & Schult.) Kuntze, Revis. Gen. Pl. 2:416. 1891. Tiro: BRASIL. Datos perdidos, Hoffmannsegg

qu oriPo: B-W 5166 [foto F neg. 4490]).

Echites neg Pohl ex Stadelm., Flora 24(1 Beibl.):25. 1841. Mandevill iculata (Pohl ex Stadelm.) K. Schum., Nat. Pflanzenfam.

40 1. 1895. Tipo: BRASIL. DE JANEIRO: Serra de Macacú, fi dida (fl), Pohl 657 (HoLotipo: M; IsOTIPO: NY).

Echites Mu Ruiz & Pav. var. n. ius Stadelm., Flora 24(1 Beibl.): 27. 1841. Tipo: BRASIL. Bania: Datos perdidos (fl), Martius s.n.

(HoLoTiPo: M [fotocopia, INB

Echites hirsutus Ruiz & Pav. var. angustifolius Stadelm., Flora 24(1 Beibl.):28. 1841. Echites lasiocarpus A. DC. var. angustifolius (Stadelm.)

A. DC., Prodr. 8:463. 1844. Tipo: BRASIL. Rio DE Janeiro: Porto de Estrella, fecha perdida (fl), Martius s.n. (Lectotipo, designado

aquí, M [foto, INB]).

Echites almadensis Stadelm., Flora 24(1 Beibl.):28. 1841. Amblyanthera palustris Müll.Arg. var. almadensis (Stadelm.) Múll.Arg., Fl. Bras.

6(1):146. 1860. Tiro: BRASIL. Banu: Itahypé, Almada, fecha perdida (fl), Martius s.n. (HoLotipo: M, [fotocopia, INB, foto F neg.

20150]

hi lel ex Stadelm., Flora 24(1 Beibl.):29. 1841. Amb |i dapib (A. DC.) Müll.Arg. var. stadelmeyeri (Mart. ex

TIPO: no localizado).

Mart

Stadelm.) Müll.Arg., Fl. Bras. 6(1):149. 1860. Tipo: BRASIL. Minas Gerais: datos perdidos, Ackermann s.n. (HOLO

Minas Gerais: Lavras, Reserva Poco Bonito, 14 Nov 1987 (fl), Gavilanes & — designado aqui, INB; IsoNEoTIPO, ESAL)

Echites ciliatus Stadelm., Flora 24(1 Beibl.):32. 1841. Amblyanthera ciliata (Stadelm.) Müll.Arg., Fl. Bras. 6(1):145. 1860. Tiro: BRASIL.

Bania: Caiteté, fecha perdida (fl, fr), Martius s.n rbi ca M [fotocopia, INB, foto F neg. 20143])

Echites hirsutus Ruiz & Pav. var. angustifolius Stadelm., Flora 24(1):Beibl. 28. 1841. Echites lasiocarpus A. DC. var. angustifolius (Stadelm.)

DC., Prodr. 8:463. 1844. Tiro: BRASIL. Rio DE JANEIRO: Porto d'Estrella, fecha perdida (fl), Martius 162 (Lectotiro, designado por

Morales (2005), pr

Echites e Ruiz var. yu ius Stadelm., Flora 24(1):Beibl. 27. 1841. Tiro: BRASIL. Bania: Datos perdidos (fl), Martius s.n.

(HOLOTIPO: M HR INB])

Echites fluminensis A. DC., Prodr. 8:452. 1844. Amblyanthera fluminensis (A. DC.) Múll.Arg., Fl. Bras. 6(1):148. 1860. Mandevilla fluminensis

Donn.Sm., Enum. Pl. Guatem. 2:47. 1891. Tiro: BRASIL. Datos perdidos, Fl. Flumin. Icon 3:pl. 44. 1831 (1827) [LECTOTIPO,

designado aquí].

Echites fluminensis A. DC. var. claussenii A. DC., Prodr. 8:452. 1844. Amblyanthera claussenii (A. DC.) Miers, Apocyn. S. Amer. 187. 1878.

Tipo: BRASIL. Minas Gerais: Caxoeira do Campo, 1840 (fl), Claussen s.n. (HoLoTIPO: G-DC; Isotipo: K)

Echites tomentosus Vahl var. laticordatus A. DC., Prodr. 8:463. 1844. Amblyanthera palustris Müll.Arg., Fl. Bras. 6(1):145. 1860. Temnadenia

palustris (Mull.Arg.) Miers, Apocyn. S. Amer. 213. 1878. pa palustris (Müll.Arg.) Hemsl., Biol. Cent.-Amer., Bot. 2(10):317.

mann s.n. (LEcTOTIPO, designado aquí, G-DC [foto F neg. 26864];

1881. Tiro: BRASIL. Bania: localidad perdida, 1827-1830 (fl, fr), S

ISOLECTOTIPOS, CGE, HAL [2 cartulinas], K, MO, P [3 cartulinas]).

Echites lasiocarpus A. DC. Prodr. 8:463. 1844. Temnadenia lasiocarpa (A. DC.) Miers, Apoc. S. Amer. 210. 1878. Mandevilla lasiocarpa (A.

DC.) Malme, Bih. Kongl. Svenska Vetensk.-Akad. Handl. 24 (3/10):25. 1899. Tiro: BRASIL. Banta: Cuiabá (citado como Cuyabá),

fecha perdida (fl, fr), Manso 29 (HoLotipo: G-DC; isoripo: HAL).

hites lasiocarpa A. DC. var lobbiana A. DC., Prodr. 8:464. 1844. Temnadenia lobbiana (A. DC.) Miers, Apocyn. S. Amer. 209. 1878. Tipo:

BRASIL. Rio DE JANEIRO: montaña de Orgaos, fecha perdida (fl), Lobb s.n. (notorio: G-DC; isoriro: K)

ips hispida (Willd. ex Roem. & Schult.) Müll. Arg. var. tomentosa Müll.Arg., Fl. Bras. Ta pl. 44. f. 3. 1860. Tipo: BRASIL.

o Grosso: localidad perdida, 1833 (fl, fr), Gaudichaud 105 (LecrotiPO, designado aquí, P).

P ovata Miers, Apocyn. S. Amer. 188. 1878. Tiro: BOLIVIA. Datos perdidos E Miers s.n. (HoLoTIPO: BM [foto, INB]).

Temnadenia pallidiflora Miers, Apocyn. S. Amer. 211. 1878. Tiro: BRASIL. Rio DE JANEIRO: entre Magé y Freichal, sin fecha (fl), Miers s.n.

(HOLOTIPO: BM).

Mandevilla rusbyi Britton, Bull. New York Bot. Gard. 4:409. 1907. Tipo: BOLIVIA. Datos perdidos (fl, fr), Bang 2843 (HoLotipo: NY; ISOTIPOS:

F [foto F neg. 51185], K, MO, US [2 cartulinas]).

862 Journal of the Botanical R h Institute of Texas 1(2)

Mandevilla denticulata S.E Blake, Contr. Gray Herb.

7. Tiro: BELICE. New Haven, 8

Mar 1907 (fl), Peck 696 (HoLoTiPo: GH [foto,

INB]; isoripo: K).

os

a

Lianas; ramitas var iadamente RA

a hirsutas

o glabras; coléteres a in-

conspicuos, ca. 0,4 mm de largo. Hojas

opuestas; pecíolos 11-55 mm de largo;

láminas foliares 4,2-14,4 x 2,5-7,8 cm,

elípticas, ovado-elípticas a obovado-

elípticas, el ápice caudado-acuminado

a cuspidado, la base cordada a sub-

cordada, raramente redondeada, los

coléteres irregularmente distribuidos a

lo largo del nervio central, membraná-

ceas, esparcidamente estrigillosas a

glabrescentes o más raramente glabras

en la superficie adaxial, esparcida a

moderadamente hirsutulas a tomen-

tosas en la superficie abaxial, el indu-

mento más denso sobre los nervios,

más raramente glabras o glabrescentes,

no revolutas, las venas secundarias im-

presas en ambas caras, las venas terci-

arias impresas en la superficie abaxial,

evidentes o inconspicuas en la superfi-

cie adaxial. Inflorescencia más corta o

igualando las hojas subyacentes, axilar,

hispidulosas, estrigosas, puberulentas,

Fic. 1. Mandevilla hirsuta (A-H de Morales & Lápiz 2695, INB; I-J de Morales et al. 2411, NB)... tomentulosas a glabrescentes o glabras,

M it ; A fl bn la B in d fi ial de la hoja, mostrando los con muchas flores, pedúnculo 13-63

col A lo largo de a (se las cn feas € C. Cáliz y pedicelo, con la mm de largo, pedicelos 3-8 mm de

EE E OR UV PA BIS largo, brácteas 11-21 x 1,5-8 mm de

giboso. F. Antera, vista dorsal. G. Cabeza estigmática. H. Nectario y ovario. I.Foliculos.J. largo, ovadas, foliáceas a subfoliáceas;

Semilla. sépalos 7-12(-15) x 1-3 mm, angosta-

mente ovados, ovado-elípticos a linear-

ovados, esencialmente iguales, acuminados a acuminado- d M apicalmente, el ápice usualmente no

a} J 1

reflexo, foliáceos a subfoliáceos, densa a

, hirsutulos, tomentosos a glabrescentes

o glabros externamente, el coléter solitario, entero, peter ai | roso apicalmente; corola

infundibuliforme, el tubo crema a verde-crema externamente, algunas veces con tonos rojizos, los lobulos

amarillos, anaranjado-amarillentos, blanco-crema a blancos, la garganta usualmente roja internamente,

raramente blanca, variadamente hirsutas, el indumento más denso en botones florales, el tubo basalmente

giboso, la parte inferior 25—32(—35) x 2-5,8(-7) mm, la parte superior 15-21(225) mm de largo, cónica a

estrechamente cónica, con un diámetro de (11-)16-21 mm en el orificio, el ápice del botón floral acuminado a

cortamente apiculado; lóbulos (11—)22-29 x (92)18—25 mm, obovados, extendidos, e irregularmente reflexos

marginalmente; estambres insertos en la base de la parte superior del tubo, anteras 4,5—5,2 mm de largo,

glabras dorsalmente, la base auriculada, con las aurículas redondeadas a subtruncadas, cabeza estigmática

2-2,2 mm de largo; ovario 1,5-1,9 mm de largo, glabro o glabrescente; nectario anular, irregularmente

Morales, N iac dal SEPT RAR NEAR a

pentalobulado, ca. la mitad de la longitud total del ovario. Foliculos 10,8-19,3(-21,2) cm x 1,5-4,5 mm,

usualmente no fusionados longitudinalmente, usualmente solo unidos en su región apical, más raramente

fusionados a lo largo de su entera longitud, variadamente hispídulosos o puberulentos, raramente glabres-

centes, moniliformes; semillas 6,5-9(-13) mm de largo, usualmente diminuta y densa a moderadamente

puberulentas, más raramente glabrescentes, coma 1,3-2,5 cm de largo, amarillo-canela.

Distribución, hábitat y ecología.—N de México hasta Brasil, Paraguay, y Bolivia, así como en Trinidad y

en las Antillas menores (St. Thomas), creciendo en bosques, sabanas, vegetación asociada a afloramientos

rocosos y formaciones de cerrado, en elevaciones de 0-1800 m.

Mandevil

delind n partes vegetativas y florales, de manera que, aunque el indument á usualmente presente

==

a hirsuta es una de las especies más variables en el género, principalmente en el tipo y densidad

en prácticamente todas las partes de la planta, M con carencia total de indumento también son

ESTE

encontrados. Estos especimenes con hojas e infl mol j idénticos a la típica

variedad pubescente y no difieren en ningún otro carácter, mos lo que el ET aislado de estos rangos de

variación puede llevar a la errónea conclusión de que varias especies pueden ser reconocidas (a nivel espe-

cífico o varietal). Sin embargo, el análisis de la pubescencia en todo el rango de distribución geográfica, ha

demostrado que no existe una discontinuidad (ni siquiera a nivel geográfico) que justifique tal separación.

Además, antes de cualquier intento de segregación de este taxón, es importante tomar en cuenta la alta va-

riación en la presencia y densidad del indumento en otras especies del género (e.g., M. pohliana (Stadelm.)

A. H Gentry), así como la alta intergradación presente entre especímenes con indumento denso a aquellos

que carecen totalmente de este. En términos generales, Mandevilla hirsuta ha sido fácilmente confundida

con M. sagittarii Woodson otro taxón morfológicamente muy similar, del que difiere principalmente por la

forma del tubo de la corola.

Amblyanthera hispida var. tomentosa Müll.Arg. es lectotipificada y la colección Gaudichaud 105 selec-

cionada como el lectotipo, ya que este especimen tiene flores y frutos y se encuentra en buen estado de

preservación. El otro sintipo citado por Müller (1860) es la colección tipo de Echites lasiocarpus A. DC.

(Morales 2005).

Tres sintipos fueron citados en la descripción de Echites hirsutus Ruiz & Pav. var. angustifolius: dos cor-

responden a colecciones de Martius sin número de los estados de Bahia y Rio de Janeiro y la tercera, a una

colección de Pohl del estado de Minas Gerais, Brasil. Acá se selecciona la colección de Martius recolectada en

el estado de Rio de Janeiro como el lectotipo, pues es la que se encuentra en mejor estado de preservación.

En la lectotipificación de Echites tomentosus Vahl var. laticordatus A. DC. se selecciona la colección de

Salzmann s.n. como el lectotipo, ya que este especimen está representado en diferentes herbarios, mientras

que el otro sintipo, una colección realizada por Perrottet en Guyana, es conocido por un unico especimen

i nd: en el herbario de París (P).

Un i (Ackermann s.n.) fue citado en la descripción de Echites stadel i Mart. ex Stadelm.,

el cual no ud ser localizado en los T herbarios europeos. Por lo tanto, se procede a designar un

neotipo que concuerde con los caracteres mencionados en la descripción original.

Mandevilla horrida JE Morales, sp. nov. (Fig. 2): Tiro. PERÚ. Cajamarca: San Ignacio, San José de Lourdes, 23 nov 1999

), R. Vásquez & Flores 26349 (HoLoTIPO: INB; IsoTipo: MO).

A Mandevilla hirsuta (Rich) K. Schum., cui affinis, bracteis 32-46 x 10-19,5 mm (vs. 11-20 x 1,5-8 mm), calycis laciniis 4-6 mm longis

(vs. 7-12(-15) mm), et tubo proprio 20,5-23 mm longis (vs. 25-32(-35) mm) differt.

Lianas; ramitas densa a moderadamente híspidas; coléteres interpeciolares inconspicuos, ca. 0,6 mm de

largo. Hojas op ; pecíolos 14-48 mm de largo; láminas foliares (7,3-)8,6-15,9(-17,3) x G23,7-7,1C 7,8)

cm, elípticas, obovado-elípticas a obovadas, el ápice caudado-acuminado, la base cordada, los coléteres ir-

regularmente distribuidos a lo largo del nervio central, membranáceas, esparcidamente estrigillosas en la

superficie adaxial, densa a moderadamente hirsutas en la superficie abaxial, el indumento más denso a lo

largo de los nervios, no revolutas, las venas secundarias impresas en ambas caras, las venas terciarias impre-

sas en la superficie abaxial, apenas evidentes o inconspicuas en la superficie adaxial. Inflorescencia usual-

864 Journal of the Botanical R h Institute of Texas 1(2)

mente más corta que las hojas subyacentes,

axilar, densa a moderadamente hirsutas,

con muchas flores, pedúnculo 1,7-5,4 cm

de largo, pedicelos 1,5—3(-5) mm de largo,

brácteas 32-46 x 10-19,5 mm, ovadas a

ovado-elípticas, foliáceas; sépalos 4-6 x

1,4-2 mm, angostamente ovados a elíptico-

ovados, esencialmente iguales, acuminados

apicalmente, el ápice no reflexo, foliáceos,

esparcidamente hispídulosos, el coléter

solitario, subentero a levemente erosos api-

calmente; corola infundibuliforme, el tubo

rojizo, los lóbulos blancos, blanco-rosados,

amarillos o rojos, densa a esparcidamente

pilosa externamente, el indumento más

denso en botones florales, el tubo basal-

mente giboso, la parte inferior 20,5-23 x

1,9-4 mm, la parte superior 19-26 mm,

campanulada a urceolado-campanulada,

con un diámetro de 1-1,3 cm en el orificio,

el ápice del botón floral agudo a cortamente

acuminado; lóbulos 13-16 x 10,5-13 mm,

obovados, relativamente extendidos; estam-

ies}

bres insertos en la base de la parte superior

del tubo, anteras 3,9-4,1 mm de largo,

glabras dorsalmente, la base auriculada,

con las aurículas truncadas o subtrunca-

das, cabeza estigmática 1,1-1,3 mm de

largo; ovario 1,4-1,7 mm de largo, glabro;

nectario anular levemente pentalobulado,

Fic. 2. Mandevilla horrida (A-H de Vásquez & Flores 26349, INB; I-J de Gentry etal. ca. la mitad de la longitud total del ovario.

61230, INB). A. Ramita con infl B. Detalle de la superficie adaxial de | Folículos 16,8-23 cm x 1,5-5,2 mm, no

hoia J I Aa I I P p t ime Sal J £ L X 3 f :

) 3 ^ A fusionados longitudinalmente, usualmente

C. Cáliz y pedicelo, con la bráctea floral ida. D. Vista adaxial de dos sépalos, m : m

alee: la base. E. Botón floral parcial berto mostrando solo unidos en su región apical, híspidos o

eee Sane eee el tubo basalmente aiboso. F. Antera, vista dorsal. G. hirsutos, moniliformes; semillas 14-15,2

Cabeza estigmática. H. Nectario y ovario. I. Folículos. J. Semilla. mm de largo, glabras, coma 1,4-2,3 cm de

largo, amarillo-canela.

Distribución, habitat y ecología.—En-

démica al departamento de Cajamarca, en el N de Perú, donde crece en formaciones de selva andina, en

elevaciones de 1500-2050 m. Especímenes con flores y frutos han sido recolectados entre enero y marzo,

junio, julio y noviembre.

Mandevilla horrida ha sido confundida con la común M. hirsuta, taxón con una morfológia bastante

similar. En forma general, las especies relacionados con M. hirsuta (e.g., M. sagittari, M. steyermarkii) son mor-

fológicamente muy similares entre sí y es fácil confundirlos cuando sus caracteres florales no son estudiados

en detalle y su identificación se basa principalmente en los caracteres de las hojas y tallos. Sin embargo, M.

horrida se puede diferenciar con facilidad de M. hirsuta por sus inflorescencias con las brácteas florales mucho

más grandes (32-46 x 10-19,5 mm vs. 11-20 x 1,5-8 mm), sépalos ligeramente más pequeños (4-6 mm

vs. 7-12(-15) mm), así como corolas con la parte inferior del tubo de 20,5-23 mm de largo (vs. 25-325)

Morales, N iac dal BERT AAA NN TER -

mm) y la parte superior campanulada a urceolado-campanulada (vs. cónica a estrechamente cónica). En

forma general, estos caracteres han sido analizados cuidadosamente en el material de M. hirsuta disponible

ix en día y no se a localizar especimenes que representen existen estados intermedios.

E inados: PERÚ. Amazonas: Imaza, Yamayakat, río Marañón 28 jul 1994 (fD, Jaramillo et al. 183 (INB, MO).

PUE San Ignacio, Huarango, El Triunfo, 14 jul 1996 (fD, Campos & Díaz 2965 (INB, MO, USM); San José de Lourdes, Camana,

4 mar 1997 (fl), Campos & Corrales 3396 (INB, MO, USM); San Ignacio, La Copa, Vista Florida, 18 jun 1997 (fl, fr), Campos & García

4036 (INB, MO); San Ignacio, Huarango, 11 mar 2000 (fl, fr), Campos et al. 6565 (INB, MO, USM); San Ignacio, San José de Lourdes,

campamento Zural, 28 ene 1999 (fD, Díaz et al. 10550 (INB, MO); La Palma, NO de Chirinos, 5 feb 1988 (fD), Gentry et al. 61230 (INB,

MO, USM); San Ignacio, San José de Lourdes, 21 ene 1999 (fl, fr), Díaz et al. 10411 (INB, MO, USM).

Mandevilla inexperata J.E Morales, sp. nov. (Fig. 3). Tio: ECUADOR. Esmeraldas: San José, jun 1865 (fl), J. Isern 1110

(HoLoTIPO: MA [fotos, INB]).

A Mandevilla hirsuta (Rich) K. Schum., cui affinis, tubo proprio 12-14 mm longis (vs. 19-3 t i 9 mm diametro

(vs. 12-25 mm) differt.

Liana; ramitas diminuta y moderadamente estrigosas, el indumento algo uncinado apicalmente; coléteres

interpeciolares inconspicuos, hasta 0,8 mm de largo. Hojas opuestas; pecíolos 12-14 mm de largo; láminas

foliares 7-11 x 3,6-6,1 cm, elípticas, el ápice cuspidado-acuminado, la base subcordada, los coléteres irregu-

larmente distribuidos a lo largo del nervio central en la superficie adaxial, membranáceas, inconspicuamente

puberulentas a lo largo de las venas en ambas caras a glabrescentes, membranáceas, no revolutas, la venación

secundaria impresa en ambas caras, las venas terciarias usualmente solo impresas en la superficie abaxial,

a veces apenas visibles en la superficie adaxial. Inflorescencia más corta que las hojas subyacentes, axilar,

diminuta y moderadamente estrigosa, el indumento algo uncinado apicalmente, con muchas flores, pedún-

culo 8-32 mm de largo, pedicelos 2,5-4 mm de largo, brácteas 19-35 x 3-6,5 mm, dne elípticas a

ovado-elípticas, foliáceas; sépalos 7,5-9 x 1,7-2 mm, ang d l , el ápice

no reflexo, esparcida y diminutamente puberulentos, el coléter solitario, D a ula y levemente

lacerado apicalmente; corola infundibuliforme, el color desconocido, glabra o glabrescente externamente,

el tubo giboso basalmente, la parte inferior 12-14 x 1,9-3,1 mm, la parte superior 12,5-13,5 mm, angos-

tamente cónica, 8-9 mm de diámetro en la boca, el ápice del botón floral cortamente acuminado; lóbulos

10-12 x 9-10 mm, obovados; estambres insertos en la base de la parte superior de la corola, anteras 4-4,3

mm de largo, glabras dorsalmente, la base auriculada, con las aurículas obtusas, cabeza estigmática 1,9-2,3

mm de largo; ovario 1,8-2,2 mm de largo, glabro, nectario anular, levemente a moderada e irregularmente

pentalobulado, ca. la mitad de la longitud total del ovario. Folículos desconocidos.

Distribución, hábitat y ecología. —Endémica a la provincia de Esmeraldas, en el N de Ecuador, donde se

conoce ünicamente de la localidad tipo, en bosques muy hümedos bajo los 300 m de elevación. Flores han

sido recolectadas en junio y noviembre.

Mandevilla inexperata difiere de M. bracteata (Kunth) Kuntze M. hirsuta y M. steyermarkii Woodson

por sus corolas con un diámetro en la boca de 8-9 mm (vs. 12-25 mm) y tubo más pequeño, con la parte

inferior de 12-14 mm de largo (vs. 19-36 mm) y la superior de 12,5-13,5 mm de largo (vs. 14-24 mm).

Mandevilla krukovii Woodson, M. lancibracteata Woodson y M. moritziana (Múll.Arg.) Donn.Sm. difieren de

M. inexperata por sus sépalos mucho más cortos (2-5 mm vs. 7,5-9 mm). Finalmente, en M. sagittarii, otra

especie con corolas con una boca angosta, las hojas tienen un indumento persistentemente tomentoso y

glauco abaxialmente y corolas más largas en relación a M. inexperata.

Mandevil

uncinado de sus ramitas e inflorescencias (vs. hirsuto o hirsutulo y erecto o suberecto), láminas foliares incon-

c

a inexperata difiere de M. megabracteata (descrita acá) por el indumento estrigoso y apicalmente

spicuamente puberulentas a glabrescentes abaxialmente (vs. esparcidamente hirsutas), brácteas florales más

pequeñas (19-25 mm vs. 44-68 mm) y corolas con la parte inferior más corta (12-14 mm vs. 23-25 mm).

Es muy llamativo el hecho que esta especie fue recolectada hace más de 140 años y el espécimen tipo

paso inadvertido en las colecciones del Real Jardín Botánico en Madrid, Espafia, las cuales han sido estu-

diadas por varios especialistas en la familia.

866 Journal of the Botanical R h Institute of Texas 1(2)

ECIITANOD

Loja: entre Yangana y Valladolid, 5 nov 2002 (f1),

Grant et al. 02-4293 (MO).

Mandevilla els piod J.E Mo-

rales, sp. nov. (Fig. 4). Tiro: GUY-

ANA. Región cud montañas

Pakaraima, Mt. Wokomung, cabeceras de

Wusupubaru creek, 17 feb 1993 (fl, fr),

Henkel et al. 1447 (noLotipo: INB; IsoTIPOS:

A Mandevilla steyermarkii Woodson, cui affinis,

ramulis hirsutis (vs. glabris vel glabrescentiis),

petiolis 8-10 mm longis (vs. (1421-62 mm),

calyces laciniis 9-10 mm longis (vs. 2,5-3 mm), et

corollae faucibus 10-12 mm diametro (vs. 17-25

mm) differt.

Liana; ramitas moderada a esparcidam-

ente hirsutas; coléteres interpeciolares

inconspicuos, hasta 0,5 mm de largo.

Hojas opuestas; pecíolos 8-10 mm

de largo; láminas foliares 8,5-12 x

3,2-4,1 cm, elípticas, obovado-elípti-

cas a obovadas, el ápice abruptamente

corto-acuminado, la base cordada, los

coléteres irregularmente distribuidos a

lo largo del nervio central en la superfi-

cie adaxial, membranáceas, glabrescen-

tes en la superficie adaxial, con pelos

largos y espaciados en las nervaduras

únicamente, esparcidamente hirsutas

en la superficie abaxial, el indumento

Fic. 3. Mandevilla inexperata (Isern 110, MA). A. Ramitacon flores. B. Detalle de la pul denso a lo largo de los nervios, no rev-

= J 1 tall C Niatall J I E J E

| olutas, la venación secundaria impresa

lol lel i I (senalad fled ).D Cáliz y pedicelo, la brá fl |

removida, E Sépalo y coléter, vista adaxial F. Tubo de un botón parcialmente abierto, M fici

giboso. G. Antera, vista dorsal. H. Cabeza usualmente solo visibles en la superficie

en ambas superficies, las venas terciarias

estigmática. L Nectario y ovario. | abaxial. Inflorescencia conspicuamente

más larga que las hojas subyacentes,

axilar, el pedünculo hirsuto, el resto

moderadamente hirsutula, con muchas flores, pedúnculo 18-23 mm de largo, pedicelos 3-4 mm de largo,

brácteas 44-68 x 7-11 mm, angostamente elípticas, conspicuamente foliáceas, verdes; sépalos 9-10 x 0,9—1,1

mm, angostamente ovados, largamente acuminados apicalmente, el ápice a veces ligeramente reflexo, subfo-

liáceos, esparcidamente hirsutulos cerca de la base externamente, en el resto el indumento muy esparcido, el

coléter solitario, irregularmente lacerado apical te; corola infundibuliforme, el tubo verde externamente,

los lóbulos blancos a crema, densa o moderadamente hirsuta o hirsutula en botón, tornándose inconspicua

y muy esparcidamente hirsútula a glabrescente externamente en la antésis, el tubo giboso basalmente, la

parte inferior 23-25 x 2-3,5 mm, la parte superior 14-16 mm de largo, angostamente campanulada, con

un diámetro de 10-12 mm en la boca, el ápice del botón floral cortamente acuminado; lóbulos 11-15 x

9-13 mm, obovados, apenas extendidos; estambres insertos en la base de la parte superior del tubo, anteras

3,9-4,1 mm de largo, glabras dorsalmente, la base subtruncada y subauriculada, con las aurículas apenas

visibles, truncadas a subtruncadas, cabeza estigmática 2-2,2 mm de largo; ovario 1,8-2,4 mm de largo,

Morales, N iac dal PREC AAN NN TER -

glabro; nectario anular, leve e ir-

regularmente pentalobulado, 1-1,2

mm de largo. Folículos 11,6-16,1

cm x 1,2-4,1 mm, esparcidamente

hirsutulos o puberulentos, monili-

VW

formes; semillas inmaduras 10-11

W

mm de largo, diminuta y densamente EB wo 1

puberulentas, la coma 1,6-2,1 cm de

largo, canela.

Distribución, hábitat y ecología.—

Restringida al N de Colombia (Depar-

tamento de Cesar y el E de Guyana,

donde se encuentra en bosques muy

hümedos y vegetación secundaria

relacionada, en elevaciones de 800—

1150 m. Material con flores ha sido

recolectado entre febrero y marzo.

El único espécimen con frutos fue

recolectado en febrero.

Mandevilla megabracteata es

morfológicamente similar a M. stey-

ermarkii, ya que ambas comparten

inflorescencias con brácteas florales

foliáceas y conspicuas, que ocultan

los botones florales. Sin embargo, M.

megabracteata puede ser fácilmente

distinguida por sus ramitas moderada

aesparcidamente hirsutas (vs. glabras

o glabrescentes), pecíolos 8-10 mm de

largo (vs. (14)21—62 mm de largo),

láminas glabrescentes en la superficie

adaxial y esparcidamente hirsutas ab-

Fic. 4. Mandevilla megabracteata (Henkel et al. 1447, INB). A. Ramita con flores y frutos. B.

~

OD lladal Erin ab

D. Cáliz y pedicelo, con la bráctea floral removida. E. Sépalo y coléter, vista adaxial. F. Tubo

d botón floral trando el tubo gib G. Antera, vista dorsal. H. Cab tiomáti

axialmente (vs. glabras), sépalos 9-10

mm de largo, angostamente ovados y I. Nectario y ovario. J. Semilla.

largamente acuminados apicalmente

(vs 2,5—3 mm, ovados y agudos apicalmente), corola con los lóbulos blancos a crema y con un diámetro de

10-12 mm en la boca (vs. lóbulos amarillos, anaranjado-amarillos o anaranjados y con un diámetro de 17-25

mm), así como anteras de 3,9-4,1 mm de largo (vs. 7,5-8,5 mm) y folículos esparcidamente hirsutulos o

puberulentos (vs. glabros).

Especímenes adicionales examinados: COLOMBIA. Cesar: La Jagua de Ibirico, vereda Alto de las Flores, 9 mar 1996 (fl), Fernández-

Alonso et al. 13186 (COL).

Mandevilla similaris J.E Morales, sp. nov. (Fig. 5). Tiro: VENEZUELA. BoLivar: Cumbre del Cerro Guaiquinima, 25 may

1978 (£D, Steyermarh et al. 117347 (HoLotipo: INB; isoripo: VEN).

A Mandevilla lancibracteata Woodson, cui affinis, calycis laciniis 8-10 mm longis (3-3,8 mm) et corollae faucibus 18-23 mm longis (vs.

7-9 mm) differt.

Liana; ramitas esparcidamente puberulentas cuando jóvenes, glabrescentes con la edad; coléteres interpe-

ciolares inconspicuos, hasta 0,5 mm de largo. Hojas opuestas; pecíolos 5-8 mm de largo; láminas foliares

4,5-10 x 2-4,8 cm, elípticas, obovado-elípticas a obovadas, el ápice abruptamente corto-acuminado, la

868 Journal of the Botanical R h Institute of Texas 1(2)

base cordada o subcordada, los coléteres

irregulamente distribuidos a lo largo del

nervio central en la superficie adaxial, mem-

branáceas, esparcidamente puberulenta a

glabrescentes en la superficie adaxial, densa

a moderadamente pilosulosas a glabres-

centes abaxialmente, no revolutas, venas

secundarias impresas en ambas caras, las

venas terciarias usualmente no evidentes.

3 Rer

más laroa aue lac hoias eanbva-

eo b J

centes, axilar, esparcidamente puberulenta

a glabrescente, con muchas flores, pedún-

culo 40—45 mm de largo, pedicelos 2,5

mm de largo, brácteas 25-48 x 4-9 mm

D de largo, angostamente elípticas, elípticas

a ovado-elípticas, foliáceas; sépalos 8-10

x 0,9-1,4 mm, angostamente ovados, acu-

minados apicalmente, el ápice no reflexo,

escariosos, i

r r

a glabrescentes externamente, el coléter

solitario, entero a subentero apicalmente;

corola infundibuliforme, el tubo verde-pá-

lido, los lóbulos amarillos o crema, esparcida

a moderadamente hirsutulas, sobretodo en

botones florales, el tubo basalmente giboso,

la parte inferior del tubo de la corola 18-23

x 2-3 mm, la parte superior 14-17 mm de

largo, angostamente campanulada, con un

diámetro de 10-13 mm en el orificio, el

Fic. 5. Mandevilla similaris (Steyermark et al. 104449, VEN). A. Ramita con flores. B. Spe del botón floral agudo; lóbulos 14-16

x 10-12 mm, obovados, extendidos; estam-

b J

). C. Cáliz y pedicelo, con la bráctea floral ida.D. bres insertos en la base de la parte superior

Lleawalad flanel

Sépalo y coléter, vista adaxial. E. Tubo de un botón trand

giboso. F. Antera, vista dorsal. G. Cabeza estigmática. H. Nectario y ovario.

del tubo, anteras 3,5-4 mm de largo, dor-

salmente glabras, la base inconspicuamente

J 1 J

auriculada, con las aurículas a

subtruncadas, cabeza estigmática 1,5-1,7

mm de largo; ovario 1,5—2 mm de largo, glabro a inconspicuamente hirsutulo en su región basal sobre

el nectario; nectario anular, irregularmente pentalobulado, 2/3 a la mitad de la longitud total del ovario.

Folículos desconocidos.

Distribución, hábitat y ecologta.—Endémica al Estado de Bolivar en Venezuela, donde crece formaciones de

Aq

sabana y vegetación arbustiva asociada a rocosos en elevaciones de 900-1600 m. Especimenes

con flores han sido recolectados en enero, mayo, agosto y diciembre Material con frutos fue recolectado en

septiembre.

Mandevilla similaris conforma un complejo junto con M. lancibracteata, ya que ambos taxones son

morfológicamente muy similares y fácilmente pueden ser confundidos entre sí, sobretodo por compartir

inflorescencias con las brácteas florales foliáceas y corolas con la parte superior del tubo angosta, con una

diámetro alrededor de 1 cm. Sin embargo, M. similaris se puede distinguir de M. lancibracteata por sus láminas

foliares con los coléteres restringidos al nervio central (vs. distribuidos también en las venas secundarias),

Morales, N iac dal BERE ARAN TN TER -

peer 1 d Y

con las venas terciarias no evidentes (vs. usua Imente sépalos de 8-10 mm de largo

(vs. 3-3,8 mm) y corolas con la parte inferior de 18-23 mm de longitud (vs. 7-9 mm de largo).

Especímenes adicionales pacas VENEZUELA. Bolivar: faldas del Auyan-tepuí, ene 1949 (fD, Cardona 2754 (VEN); Final de La

Escalera, 8 ago 1979 (fl), Morillo € 38 (VEN); La Escalera, al S de El Dorado, 13 feb 1980 (st), Morillo 8142 (VEN); parque nacional

Canaima, La Escalera, km 88 entre odo 8 sep 1987 (fr), Ramírez et al. 167 (VEN); drenaje del río Cuyuní, al S de El Dorado,

22-28 dic 1970 (fD, Steyermark et al. 104489 (\ ; drenaje del río Cuyuní, a lo largo del río Anawaray-parú, al S de El Dorado, 25 dic

1970 (fD, Steyermark et al. 84937 (VEN); Gran Sabana, entre El Dorado y p Elena, 30 Dic 1974 (fD, Steyermark 111289 (VEN).

AGRADECIMIENTOS

Quiero agradecer a los herbarios BM, B-W, C, CGE, COL, ESAL, F, G, G-DC, GH, HAL, K, M, MO, P, P-HB,

P-LA, UB, US, USM, VEN, W, Z por el envío de material en préstamo, así como por facilitar el acceso a sus

colecciones. Asimismo, agradezco la colaboración de Eliana Noguera, Shingo Nozawa y Sirli Leython que

permitió revisar material en el Herbario Nacional de Venezuela (VEN). Finalmente, Barry Hammel (MO)

realizó la traducción del resumen al inglés.

REFERENCIAS

ALLORGE-BorTEAU, L. 1998. New combinations in Odontadenia and Mandevilla (Apocynaceae). Phytologia 84:

Morales, J.F. 2005. Estudios en las Apocynaceae Neotropicales Xl: una nueva especie de Mandevilla (Apocynoi-

deae, Mesechiteae) para Sur América, con un nuevo reporte para las Apocynaceae de Paraguay. Sida

21:1549-1556

Morales, J.F. 2006. Estudios en las Apocynaceae Neotropicales XXV: novedades y nuevos reportes en las Apocy-

naceae (Apocynoideae, Rauvolfioideae) de Venezuela. Sida 22:355-365

MULLER ARGOVIENSIS, J. 1860. Apocynaceae. En: C.EP. von Martius, ed. Flora Brasiliensis 6(1):1-180.

Simoes, A.O., M. ENDRESS, T. VAN DER Niet, L.S. KinosHita, & E. Conti. 2004. Tribal and intergeneric relationships of Mes-

echiteae (Apocynoideae, Apocynaceae): evidence from three noncoding pastid DNA regions and morphol-

ogy. Amer. J. Bot. 1:1409-1418

Simoes, A.O., M. ENDRESS, T. VAN DER Niet, L.S. Kinoshita, & E. Conti. 2006. Is Mandevilla (Apocynaceae, Mesechiteae)

monophyletic? Evidence from five plastid DNA loci and morphology. Ann. Missouri Bot. Gard. 93:565-591.

Simoes, A.O., L.S. KinosHITA, & M. ENDRESS. 2007. New combinations in Mandevilla Lindley (Apocynaceae). Novon

17:89-90.

Woobson, R.E. 1933. Studies in the Apocynaceae. IV. The American genera of Echitoideae XXVI. Ann. Missouri

Bot. Gard. 20:605-790

—

WS

Journal of the Botanical R h Institute of Texas 1(2)

870

BOOK REVIEW

WiLLOw ZucHowskt. 2007. Tropical Plants of Costa Rica: A Guide to Native and Exotic Flora. (ISBN 978-

0-8014-7374-6, pbk.). Cornell University Press, Sage House, 512 East State Street, Ithaca, NY, 14850,

U.S.A. (Orders: www.cornellpress.cornell.edu). $35.00, 532 pp., 540 color photographs, 2 maps, 56

line la IE: PHAR 535-49

ts of Costa Rica i 11 f ling osta Rica

ph lOLOBI aph

Th;

or with a general interest in Wd) ans os descriptive species accounts, wonderful color

and conspicuous plants M in one Rica, the author certainly fulfills

d.

i :1 11

Turid Forsyth, and illustrations for 4

s both useful and dd small e to carry and utilize in the fiel

Perfect for the lay e or first-time visitor to Costa Rica, Spe-

her goal to create a guide th

O p E la

1 Tals

gnig 8

cies accounts in the ene are organized mus the g ] ings within which a traveler the plants. Each chapter of the

l setting id d by the Mn titles: Painted Treetops, Other Common

Trees, Roadside mi Garden OUS Fruits Er d ing F d Reforestation, Special Habitats, Typical pee Groups,

and Conspicuous Grasses. Individ le the followi bo b pan

ish and English), other a name to which the species bs physical description, fl ing/fruiting

distribution, identification of related species, and comments. Th I f the speci lay f. ginf

he natural history ial, medicinal, and ae d unique cl teristics of each sp

Sidel i a f1 1 A 1 1 pA

such titles as "Sex Life of Bs 2 ie Wasps, 5 “The ae xn in Costa Rica,” o does Chan come from?” and “Drift Seeds” to

a few A 1 a Ge A S

e aut t

g

both scientifi d common nam Melinda McCoy, Herbarium

name ju

a list m. by family, a complete bla and an index with

Research Institute of Texas, Fort Worth, Texas 76102-4060, U.S.A.

Volunteer, Botanical

BOOK NOTICES

Two Books on Florida Plants

RicHARD P. WUNDERLIN and Bruce E Hansen. 2003. Guide to the Vascular Plants of Florida (ed. 2). (ISBN

0-81302632-6; hbk.). Univ. Press of Florida, Gainesville. University Press of Florida, 15 Northwest 15th

Street, Gainesville, FL 32611-2079, U.S.A. (Orders: www.upf.com, 352-392-1351, 352-392-7302 fax,

1-800-226-3822). $39.95, 787 pp., 6" x 9".

We've all been using the book for several years, but this review gee perhap

rl the more than 4100 taxa of ferns, fern allies

companion to the multivolume Flora of Florida ceda & Hansen 2000 and in progress).”

ls botanists that it’s still for sale. It is “a means to

lized in Florida. It serves as the field guide

C. Rrrcuie Beit. and Bryan J. Tavior. 1982. Florida Wild Flowers and Roadside Plants (3" printing, 1998).

(ISBN 0-9608688-3-6, pbk.). Laurel Hill Press, 5000 Walnut Cove Rd., Chapel Hill, NC 27516-8168, U.S.A.

(Orders: www.laurelhillpress.com, 919-942-9533, 919-942-9533 fax). $19.95, numerous color photos, map,

D x B

500 species are illustrated by a color photo. For each, there is a brief di ic d iption, comments on habitat and distribution, and

a dot map (with the state divided into quarters). Useful book and a great stam

J. Bot. Res. Inst. Texas 1(2): 870. 2007

BOERHAVIA TRIQUETRA VAR. INTERMEDIA (NYCTAGINACEAE); A NEW

COMBINATION AND VARIETAL STATUS FOR THE WIDESPREAD

SOUTHWESTERN NORTH AMERICAN B. INTERMEDIA

Richard Spellenberg

rd of Biology, MSC 3AF

co State University

Las To wM 88003-8001, U.S.A.

AB OT edu

ABSTRACT

The name for the poorly known, narrowly distributed, Boerhavia triquetra S. Wats. predates the name B. intermedia M.E. Jones, a well-

T i 1 = £41 Sod A 1 E 1 A £1 1 i $ xj 1 11 J á E $

type locality of B. triquetra at Bahía de los Angeles in Baja California, shows the two taxa to thoroughly intergrade. The nomenclatural

combination Boerhavia triquetra S. Wats. var. intermedia (M.E. Jones) Spellenb. is proposed. A key distinguishing this variety and

the var. triquetra is presented.

RESUMEN

El nombre de la poca conocida Boerhava triquetra S. Wats., de distribución limitada, precede el nombre B. intermedia M.E. Jones, una

especie ios conocida y de o extensa en el suroeste árido de Norte América. Una revisión de muestras de Boerhavia en her-

barios y | g n localidad del ue en cae b los ao Baja California, demuestran qu los do

S. Wats. var. ee (M. E. Jones)

Pi Se p t ] disti variedad y la var. triquetra.

L o

In 1902 M.E. Jones described an annual Boerhavia from El Paso, Texas, U.S.A., as Boerhavia intermedia M.E.

Jones. Prior to that time this taxon was included in a broadly conceived B. erecta L., the name under which

Jones first distributed his collections of B. intermedia. For example, Gray (1853) concluded his review of

Wright’s collections of Boerhavia by noting that other species in his collection, apparently with annual roots,

had been distributed as B. erecta L. The outward similarity of this desert annual, B. intermedia, to the more

mesic and tropical B. erecta, is often noted (e.g., Kearney and Peebles 1960) and has resulted in a combination

of the former as a variety in the latter (B. erecta var. intermedia (M.E. Jones) Kearney & Peebles). Molecular

"

evidence suggests, however, that B. erecta and B. inter not particularly closely related (Norm Douglas,

pers. comm. 2007). Boerhavia intermedia as understood by Standley (1918) and Spellenberg (1993, 2003),

among other authors of regional floras, is widespread and variable, occurring throughout the warm, arid

regions of northwestern Mexico and southwestern United States. The only attempt to taxonomically address

the variation through taxonomic division of this taxon was B. universitatis Standl., from Tucson, Arizona

(Standley 1909), since shortly after its inception a name consistently considered a synonym of B. intermedia.

Though not explicitly stated by Jones (1902) or Standley (1909), B. intermedia has been understood to have

truncate fruits with five angles separated by rugose sulci.

In 1889 S. Watson described B. triquetra S. Wats. from a collection of plants made by E. Palmer, the

type originating from near Bahía de los Angeles, Baja California, Mexico. It was distinguished from other

Boerhavia by its 3-4-angled truncate fruits with rugose sulci. Since that time the distinction between B.

intermedia and B. triquetra in regional floras has not been clear. Both are distinguished from other Boerhavia,

and allied to one another, by their annual habit and narrowly obdeltate (in profile) ribbed (not winged) fruits

with pedicels that join in a terminal umbel. Those with umbels reduced to solitary fruits still differ from

other Boerhavia by the habit and fruit shape, though the distinction between B. intermedia and B. erecta is

then subtle.

Wiggins (1964) separated the two species in his key primarily by flowers solitary or in umbellate or

J. Bot. Res. Inst. Texas 1(2): 871 — 874. 2007

872 Journal of the Botanical R h Institute of Texas 1(2)

close cymules. He noted the fruit to be 3-5 angled in B. triquetra, 5 angled in B. intermedia, the fruit of the

former slightly broader. Annual Boerhavia of the very dry Mojave and western Sonoran deserts of south-

eastern California and adjacent Sonora often have one fruit per peduncle and have been called B. triquetra

on specimens. Spellenberg (1993) attempted to distinguish the two taxa by numbers of fruit per cluster,

and a subjective aspect of inflorescence structure, making no reference to rib number. In 2003 Spellenberg

maintained B. intermedia and B. triquetra, noting in discussion that the distinction between the two seemed

moot. The hesitancy to combine the two names in 2003 into one taxon was reinforced by Spellenberg not

having seen B. triquetra (which has priority over B. intermedia) in the field. In addition, the application of B.

triquetra, a name for a poorly known taxon from a very limited region, was not descriptive for the widespread

well-known species with usually 5-sided fruits. Nevertheless, in preparing the treatment of the Nyctaginaceae

for the Jepson Manual, 2*d edition, Andrew Murdock (at UC Berkeley) came to the conclusion that the two

names apply to one taxon in California (pers. comm. 2006).

In October, 2006, I spent several days in the vicinity of Bahía de los Angeles. Late summer rains had

been good, and small collections of Boerhavia were made, as were observations. Those collections have been

deposited at HCIB and NMC. Including these specimens and those deposited in herbaria, 71 collections

from throughout the range of both taxa were examined for numbers of fruit per terminal cluster, fruit size,

rib number per fruit, and pedicel length. The following observations were made

Pedicels vary from 0.2 to 3.2 mm throughout the range, the shortest occurring within the confines

of the Sonoran Desert, but longer (+ 1.7 mm) common within the region also. Watson (1889) noted the

pedicels of B. triquetra to be very short (ca. Y line [= 1/24” = ca. 1 mm]). My own collections from around

the Bahía de los Angeles had pedicels 0.271 mm long; to the north and to the south along the eastern side of

the peninsula in this region pedicels ranged to 1.3-1.9 mm long. Elsewhere they were as short as 0.3 mm.

On plants with sufficient fruiting peduncles the number of fruits in 10 umbels was counted. Numbers

of fruits per umbel ranged from 1-22. Those with single fruits per umbel occurred throughout the range, but

were particularly common in the deserts of southern California. Those with the highest number were from the

coast of Sonora. Number of fruits per umbel commonly was 1—5 in the vicinity of Bahía de los Angeles; one

plant which had consistently 1 fruit/umbel had only 5-sided fruits (Spellenberg 13790, below). No particular

difference between fruit number in this region versus other areas was obvious, except that in Chihuahua,

Coahuila and Durango fruit number commonly was 3-7 per umbel, with 1-fruited umbels less frequent.

Plants with primarily, or entirely, 1-fruited umbels from throughout the Southwestern warm deserts

are represented by the following collections (here and throughout all specimens at NMC unless otherwise

cited (herbarium citations from Holmgren and Holmgren, http://sciweb.nybg.org/science2/IndexHerbari-

orum.asp): AZ, Pinal Co., San Tan Mts. Regional Park, Damrel and Damrel 24716; BC, Mcpio. Ensenada,

on road to Bahia de los Angeles 37 km E of Hwy 1, Spellenberg 13790; Roos s.n., CA, San Bernardino Co.,

Big Morongo Reserve, in Big Morongo Canyon, Helmkamp B-19, UTEP; Riverside Co., Chuckawalla Mts.

2.5 mi SE of Desert Center, 23 Oct 1963, UTEP; NM, Dofia Ana Co., ca 10 mi E of Las Cruces, W base of

Organ Mts., Spellenberg 2921. All these had only five-sided fruits. Three- and four-sided fruits were found

on Sanders 051 and Van Devender & Reina G. 2000-939 (citations end of next paragraph), both of which had

few fruits per umber (1-2 fruits, av. 1.6; 1-4 fruits, av. 2.5, respectively). Of note is that the very earliest

inflorescences on many plants have 1 or few fruits, as often do very late inflorescences. The preponderance

of few-fruited umbellate plants on the Mojave and western Sonoran deserts may simply be selection for this

early-development feature in an area of unpredictable and reduced summer moisture.

Fruit size did not seem to differ much from one part of the range to the other. Within the Bahia de los

Angeles region fruits were 2.0-2.8 mm long; slightly to the south fruits reached 3 mm long. Elsewhere fruits

were 2.03.1 mm long. The range of fruit width is less in the vicinity of Bahía de los Angeles (the length/

width ratio equalling 1.5-2.0) than for plants from throughout the range of B. intermedia (l/w = 1.3-2.7).

The proportionately broadest fruits are from a plant that has only 3- and 4-sided fruits from Sonora (SON,

Mcpio. Hermosillo, Playa Esthela just N of Bahia de Kino, Van Devender & Reina G. 2000-939). A similar

D

intermedia 873

Spellenberg, ]

plant with only 4-sided fruits (and 1 fruit with a poorly developed 4" side, i.e., almost 3-sided) had fruits

with l/w = 1.9 (CA, Imperial Co., Julian Wash, E of Peter Kane Mt., Sanders 051).

Numbers of sides on the fruits varies throughout the range of Boerhavia intermedia. An attempt was made

to count all scorable fruits on a plant unless there were hundreds available, and then counting was haphazard,

i.e. without intentional bias. Outside the Sonoran Desert plants bear mostly 5-sided fruits, but there are oc-

casional fruits with 4 sides. In the following the numbers of fruits counted on any one plant are given in the

sequence 5-, 4-, and 3-sided as, for example, CHIH, 1 mi S of Camargo, (53, 10, 0) Henrickson 7735; CHIH, 7

mi S of Ojinaga, (74, 1, 0), Spellenberg & Spellenberg 3699; COAH, 4 km S of Las Margaritas, (38, 2, 0), Chiang

et al. 9519c; 14 mi NW of San Pedro, (56, 3, 0), Henrickson 5975; NM, Doña Ana Co., 15 mi N of Las Cruces,

(92, 7, 0), Spellenberg 2649; TX, Presidio Co., 24.9 mi E of Redford, (31, 1, 0), Spellenberg 3710.

In the Sonoran Desert, excluding central Baja California (reviewed below), plants have 5-sided fruits,

or mostly 5-sided fruits with some 3- or 4-sided fruits. Examples of those with some 3- and 4-sided fruits

are: AZ, La Paz Co., (49, 1, 1), Spellenberg 7864; Pinal Co., (152, 3, 0), Spellenberg 13271; SON, Mcpio. Em-

palme, 1.9 km NW of MEX 15 on Empalme-Guaymas bypass, (79, 8, 0), Reina G. et al. 2002-1055; 25 km

S of Empalme (98, 14, 1 [1 fruit with 9 ribs]), Spellenberg & Douglas 13317. Mcpio. Guaymas, 1 mi E of San

Carlos Bay, (45, 2, 0) Spellenberg et al. 2688; 4 mi. W of San Carlos, (35, 2, 0) Spellenberg et al 2693; 4 mi W of

San Carlos, (30, 2, 1), Spellenberg et al. 2694; Guaymas, Colonia La Peninsula, (71, 1, 0), Spellenberg & Willson

3632; 4.6 km W of San Carlos, (45, 4, 0), Spellenberg 13811. Mcpio. Hermosillo, ca.10 mi S of Hermosillo,

(90, 6, 0), Spellenberg et al. 2683. Mcpio. Moctezuma, 18.9 km SSE of jct with Moctezuma-Huásabas Hwy,

(43, 1, 0), Van Devender & Reina G. 2006-801. Plants with 3- and 4-sided fruits, usually among 5-sided fruits

on the same plant, are most common at low elevations on the Sonoran Desert around the northern portion of

the Gulf of California, and plants with the greatest proportion of 3- and 4-sided fruits are found here also.

To the north and south of Bahía de los Angeles, on the eastern side of Baja California, numbers of sides

per fruit also varies widely: Mcpio. Ensenada, ca 16 km S of Puertocitos, (22, 0, 0) Spellenberg 13783; ca.

30.5 km S of Puertocitos, (38, 0, 0) Spellenberg 13784; 13.5 km N of Punto Bufeo, (92, 4, 1) Spellenberg 13785.

Mcpio. Mulegé, 8 km W of Santa Rosalia, (29, 0, 0) Spellenberg 13803; ca. 1.5 km S of Santa Rosalia, (7, 14,

6), Spellenberg 13805; ca. 1.5 km S of Santa Rosalia, (41, 0, 0) Spellenberg 13806; 4.6 km N of Mulegé, (plant A

= 158, 0, 0; plant B = 56, 2, 0) Spellenberg 13807. It is noteworthy that 13805 and 13806 were within 100 m of

each other. Other collections from the general region: Mcpio. La Paz, Isla San José, León de la Luz 9409a, (25,

0, 0); Sierra la Giganta, Moran 18860 (60, 2, 0). Mcpio. Mulegé, Sierra la Giganta, Carter 4859, (47, 2, 0).

These results are compared with sampling from in the immediate Bahía de los Angeles (BLA) region

on the eastern side of Baja California: BC, Mcpio. Ensenada, road to BLA 37 km E of Hwy 1, (57, 0, 0)

Spellenberg 13790; 4.3 km W of BLA, (82, 51, 6) Spellenberg 13791; BLA, (30, 4, 0) Spellenberg 13792; 4.5 km

S of BLA (33, 95, 39) Spellenberg 13793; 1.6 km N of BLA, (70, 16, 0) Spellenberg 13795; 6.4 km W of BLA

(plant A = 4, 15, 21; plant B = 17, 8, 3) Spellenberg 13799. This last collection was taken from a crowded

population of plants in an arroyo bottom that did not differ noticeably among one another with regard to

habit or fruit number per umbel, but these were selected from among those that bore mostly 5-sided fruits.

Counts from the type specimens (Palmer 521, Lower California, Los Angeles Bay in 1887) are: (1, 18, 21)

US 933319; (8, 60, 89 — from a count of fruits in packet made in 2001) US 933313; (1, 24, 23) US 22954;

(0, 4, 6) UC 101261. Though I was able to find many plants that had a habit very similar to the types of B.

triquetra around Bahía de los Angeles, none that I found had proportions of 3- and 4-sided fruits as high as

in the types.

A broad survey of plants known as B. intermedia and B. triquetra revealed no differences that unequivo-

cally distinguish the two. Fruit size, rib number and rugosity are similar, and differences are variable and

overlapping. Stems may be more widely spreading in some populations from Baja California, but erect plants

occur there also. Plants may be more strongly ascending to the east, but widely spreading plants also occur.

It is proposed that the two names be considered to apply to a single species with two highly intergradient

varieties, requiring a new combination:

£L Dat PRA D hi PEA

874 Journal of f Texas 1(2)

Boerhavia triquetra S. Wats. var. intermedia (M.F. Jones) Spellenb., comb. nov. Boerhavia intermedia M.E. Jones, Contr.

W. Bot. 10:41. 1902. Boerhavia erecta L. var. intermedia (M.E. Jones) Kearney & Peebles, J. Wash. Acad. 29:475. 1939. Tyre: TEXAS:

El Paso, 10 Sep 1883, M.E. Jones 4173 (Lectotype, designated by Reed, Flora of Texas 211. 1969; US 223742! IsoLEcToTYPES: POM!).

Reed od this name in his treatment of Nyctaginaceae for Texas when he specifically cited the US specimen as the type.

Boerhavia universitatis Standl. Contr. U.S. Natl. Herb. 12:380. 1909. Type: ARIZONA: Tuscon, Campus Univ. of Arizona, alt. 740 m, 2 Sep

903, Thornber s.n. (HOLOTYPE: ARIZ! isotypes: MO! USD.

The two varieties may be distinguished as follows. The var. triquetra is restricted to low elevations (0-300 m)

in the Sonoran Desert around the Gulf of California. The var. intermedia is widespread in Mojave, Sonoran,

and Chihuahuan deserts, including low elevations around the Gulf of California. Though the key below

seems to allow for a potentially large number of intergradient plants, no plants in the sample that I observed

fall within the range of bearing 50-80% 5-sided fruits.

1. 50% or more of the fruits on a plant 4- or 3-sided B. triquetra var. triquetra

2. 80% or more of the fruits on a plant 5-sided B. triquetra var. intermedia

Four other named Boerhavia are either part of, or very closely related to, this complex. Two are narrow

endemics with the ribs expanded into broad wings, B. alata S. Wats., from the islands around Guaymas,

Sonora, and B. megaptera Standley, from south-central Arizona. Two others are poorly distinguished from B.

triquetra var. intermedia. One, B. maculata Standley, from the coast of southern Sonora and adjacent Sinaloa,

has larger and proportionately broader 5-ribbed fruits and comparatively large flowers in 1-2-flowered

umbels. Except for its larger flowers, a feature common among these annual Boerhavia that grow along the

Sonoran coast, it resembles other Boerhavia specimens with few-flowered umbels taken at low-elevation from

around the Gulf of California. The other, B. lateriflora Standley from the vicinity of Guaymas, Sonora, has

large inflorescences of dense head-like cymes or compound umbels. A suitable disposition of these names

may be revealed by critical study of molecular and population variation.

ACKNOWLEDGMENTS

I am grateful to Rob Soreng for counting ribs on fruits on the three types at US. Rob noted (pers. comm.),

“Could be a few more 5s but I did not want to pry the 4rs up to check, but most 4s were clearly 4s.” Andy

Murdock (UC) provided helpful comments during final preparation of the manuscript. Richard Felger and

an anonymous reviewer graciously reviewed the manuscript. Discussions with Norm Douglas (DUKE)

regarding annual species of Boerhavia in the Sonoran Desert have been very helpful.

REFERENCES

Gray, A. 1853. New genera and species of Nyctaginaceae, principally collected in Texas and New Mexico. Amer.

J. Sci. Arts, ser. 2, 15:259-263, 319-324.

Jones, M.E. 1902. Nyctaginaceae. Contr. W. Bot. 10:34-54.

Kearney, T.H. and RH. Peestes. 1964. Arizona flora, 2"4 ed., with supplement by J.T. Howell, E. McClintock and col-

laborators. Univ. Calif. Press, Berkeley.

Reep, C.F. 1969. Nyctaginaceae. Flora of Texas 2:151—220. Texas Research Foundation, Renner.

SPELLENBERG, R. 1993. Nyctaginaceae. In: The Jepson manual: higher plants of California, J.C. Hickman, ed. Univ.

Calif. Press, Berkeley. Pp. 768-774.

SPELLENBERG, R. 2003. Boerhavia. In: Flora of North America north of Mexico, vol. 4, Flora of North America Editorial

Committee, eds., Oxford Univ. Press, New York. Pp. 17-28.

STANDLEY, P.C. 1909. The Allioniaceae of the United States with notes on Mexican species. Contr. U.S. Natl. Herb.

12(8):303-389.

STANDLEY, PC. 1918. Allionaceae. N. Amer. Fl. 21(3):171-254.

WATSON, S. 1889. VI. Contributions to American botany. 1. Upon a collection of plants made by Dr. E. Palmer, in

1889, about Guaymas, Mexico, Muleja and Los Angeles Bay in Lower California, and on the island of San Pedro

Martin in the Gulf of California. Proc. Amer. Acad. Arts Sci. 24:36-87.

INERASPECIPIC TAXONOMY AND NOMENCLATURE OF

ELEOCHARIS ACUTANGULA (CYPERACEAE)

David J. en and Stephan L. Hatch Richard Carter

Tracy Herbarium Herbarium, D f Biolo

Department E ET Science & Management Valdosta State University

Texas A&M Universit) Valdosta, Georgia 31698, U.S.A.

4^

College Station, Texas 77843-2126, U.S.A.

ABSTRACT

A taxonomic study of Eleocharis acutangula (Roxb.) Schult. was conducted in order to better define this poorly understood and variable

po eee Multivariate statistical NE and ecological and ord data of worldwide collections of E. acutangula

E

acutangula, E tang bsp. breviseta D.J. Rosen, subsp. nov., and

E ] bsg eacop] Rosen seep nov. N ] I i ] tl I f E. acutangula and

a Bei synonym, E. fistulosa Schult. var. robusta Boeck. A i fE gula is provided that includes a key to

the subspecies, detailed P illustrations, and notes on habitat and distribution

RESUMEN

Se realizó un estudio taxonómico de Eleocharis acutangula n Schult. para definir mejor esta especie pantropical variable y pobre-

mente conocida. Un análisi adísti Mp yd lógicos y de distribución a nivel mundial de E. acutangula fueron la

iónen E bsp. acutangula, E. aene dnd pins ae Rosen, subsp. nov., y E. deutingum

saben necito D] Rosen SIS nov. Lai 5 totipifi E. ¢ ng de un sinónimo

heterotípico, E. fistulosa Schult. var. robusta Boeck. Se aporta un tratamiento taxonómico ide jn M que ed una clave de

subespecies, descripciones detalladas, ilustraciones, y notas sobre el hábitat y distribución.

Eleocharis R. Br. is a cosmopolitan genus of about 200 species and over 600 published names with a center

of diversity in the Neotropics (González-Elizondo & Tena-Flores 2000). Eleocharis subg. Limnochloa (P.

Beauv. ex Lestib.) Torr. (= Eleocharis ser. Mutatae Svenson) comprises over 35 species occurring in season-

ally wet to permanently flooded habitats from principally tropical regions, and is distinguished from other

Eleocharis by a combination of the following morphological characteristics: (1) cartilaginous, un-keeled

(rarely obscurely-keeled), many-veined floral scales; (2) generally large culms that are often as thick as the

cylindrical spikelet; and (3) biconvex (rarely trigonous) achenes usually with epidermis of large, conspicuous

polygonal cells (Svenson 1929; González-Elizondo & Peterson 1997). Five new species in subg. Limnochloa

have recently been described from the New World: E. eglerioides S. González & Reznicek and E. liesneri S.

González & Reznicek from Venezuela (S. González-Elizondo & Reznicek 1996), E. yecorensis Roalson from

Mexico (Roalson 1999), E. laeviglumis R. Trevis. & Boldrini from Brazil (Trevisan & Boldrini 2006), and

E. steinbachii D.J. Rosen from Bolivia (Rosen & Hatch in press). However, no comprehensive study of subg.

Limnochloa has been published since the seminal work of Svenson (1929, 1939).

Eleocharis acutangula (Roxb.) Schult. is the most widely distributed species of Eleocharis subg. Limno-

hloa (Svenson 1939 [as E. fistulosa Schult.]). In the New World it is reported from near sea level to eleva-

tions over 2200 m from various habitats including cloud forests, forest depressions, savannahs, grasslands,

palm swamps, lake margins, borrow pits, and roadside ditches. Old World habitats include swamps, forest

depressions, streams, savannahs, grasslands, borrow pits, lake margins, and rice paddies. Several authors

have reported considerable variation in E. acutangula (Svenson 1929 [as E. fistulosa], 1939; Haines & Lye

1983; Browning et al. 1997). Svenson (1929, 1939) indicated E. planiculmis Steud. and E. fistulosa Schult. var.

robusta Boeck. were potential segregates of E. fistulosa, which is treated herein as a synonym of E. acutangula.

Hess (1953) described Heleocharis pseudofistulosa H. Hess based on plants he collected in Angola, and stated

that they differed from E. fistulosa in surface characteristics of the achene. Hess (1957) later provisionally

J. Bot. Res. Inst. Texas 1(2): 875 — 888. 2007

876 Journal of the Botanical R h Institute of Texas 1(2)

reported H. cf. pseudofistulosa from South America (Brazil), which differed in having terete rather than sharply

three-angled culms; this is presumably E. obtusetrigona (Lindl. & Nees) Steud.

Svenson (1939) included E

ing et al. (1997) described variability among specimens of E. acutangula from different geographical areas in

=

eocharis fistulosa among five poorly defined tropical African taxa, and Brown-

southern Africa. Our research reported here, including observations of live plants in the field and a study of

herbarium specimens from a broad geographical area, shows considerable variability within E. acutangula,

thus confirming the work of Svenson (1939) and Browning et al. (1997). A critical examination of over 600

specimens of E. acutangula suggested sufficient variation existed to warrant recognition of three infraspecific

taxa: E. acutangula subsp. acutangula, E. acutangula subsp. breviseta, and E. acutangula subsp. neotropica. The

objectives of this research were: (1) to investigate the morphological variation within E. acutangula and (2)

to review all the apposite nomenclature in order to typify E. acutangula and its synonyms.

METHODS

Specimens were borrowed from herbaria that could provide loans yielding broad geographical representa-

—=

tion of Eleocharis acutangula including types and authentic specimens. Over 600 specimens were examined

from the following herbaria (acronyms follow Holmgren et al. 1990): BM, BRI, BRIT, C, CIIDIR, CM, E, F,

FTG, GA, GH, IBE, ICN, K, LL, M, MEXU, MICH, MO, NH, NU, NY, P, PH, PRE, RSA, TAES, TEX, US, USF,

VSC, WIS, Z, and ZT. Selected for multivariate analysis were 198 mature herbarium specimens (including

types) complete for all morphological characters measured. Specimens studied originated from Africa, Aus-

tralia, Bolivia, Brazil, China, Colombia, Cuba, Dominican Republic, Ecuador, El Salvador, Guyana, India,

Japan, Madagascar, Malaysia, Mexico, Panama, Peru, United States, Venezuela, and Vietnam. Because of

the limited number of specimens complete for all morphological characters, duplicate specimens collected

by the same collector were measured. A complete citation of all specimens examined during this research

can be found in Rosen (2006).

Quantitative and qualitative vegetative characters (e.g., culm height, width, texture, and cross-sec-

tional shape; leaf sheath texture and structure; rhizome length and diameter) are highly plastic. Although

these features are of some use in Eleocharis at the infrageneric level, they are of no value in distinguishing

infraspecific taxa. In Eleocharis subg. Limnochloa, significant variation in culm anatomy in response to en-

vironmental conditions has been reported (Edwards et al. 2003; Baksh and Richards 2006). Svenson (1929)

emphasized achene characters and perianth bristle texture in differentiating species of Eleocharis. A review

of the literature reveals a tendency of workers investigating closely related species and infraspecific varia-

tion in Eleocharis to rely primarily on characters associated with the achene (Hines 1975; Larson & Catling

1996; Gregor 2003). Indeed, achene-related characters are important in taxonomic limits in Eleocharis at all

levels (Menapace 1991).

oy ed E were selected for initial evaluation (Table 1). For each specimen a ma-

d from near the base of a spikelet. Each specimen measured

was complete for all rae so that the data matrix contained no missing values. One measurement per

character was taken from each specimen, and 198 specimens (114 of Eleocharis acutangula subsp. acutan-

gula, 67 of E. acutangula subsp. breviseta, and 17 of E. acutangula subsp. neotropica) were analyzed utilizing

principal component analysis (PCA). The raw morphometric data were standardized and analyzed using

NTSYSpc 2.11Q, and the principal components were generated using a correlation matrix (Rohlf 2000).

A final analysis comprising six characters (Table 2) was run, and a scatter plot of the first two principal

components was generated in an effort to depict morphological relationships.

RESULTS

The first three principal components represented 87.3% of the total variance (50.3%, 24.3%, and 12.7% for

PCI, PC2, and PC3 respectively; Table 2) of 198 specimens scored for six morphological characters. Prin-

cipal component 1 is most influenced by high positive loadings of LONBRSTL, TBRCL, BRSTLNACHNL

Rosen et al., T f Eleoci g | 877

TABLE 1. Initial 20 fi Fleact gul

Symbol Character

ACHNL achene length (from base to constriction at neck)

ACHNLW ratio of achene length to width (achene shape)

ACHNMAX ratio of achene length to distance from achene base to widest point (determines if achene is widest

above, at, or below middle)

ACHNSCAL ratio of achene length to floral scale length

ACHNW achene width (at widest point)

BRSTACH number of perianth bristles longer than summit of achene

BRSTLACHNL ratio of length of longest perianth bristle to achene length

BRSTLNACHNL ratio of number of perianth bristles longer than summit of achene to total number of perianth

bristles

BRSTN number of perianth bristles

LONBRSTL length of longest perianth bristle

LONROW number of longitudinal rows of cells on achene face

NECKWACHNW ratio ofachene neck width to achene width

NECKW achene neck width

SCALEL floral scale length

SCALELW ratio of floral scale length to width

SCALEW floral scale width

TBRACHW ratio of tubercle width to achene width

TBRCL tubercle length

TBRCLW ratio of tubercle length to width (tubercle outline shape)

TBRCW tubercle width

(Table 2). Principal component 2 is

most influenced by a high positive

loading of NECKWACHNW and a

high negative loading of TBRCIW

(Table 2). Although there are vary-

ing degrees of overlap among the

three taxa, specimens from each

1

subspecies

tinct groups (Fig. 1). Specimens of

Eleocharis acutangula subsp. breviseta

and E. acutangula subsp. neotropica

are almost completely separated

along principal component axes

l and 2 (Fig. 1). Specimens of E.

acutangula subsp. acutangula overlap

slightly with E. acutangula subsp.

breviseta along principal component

axis l and E. acutangula subsp. neo-

tropica along principal component

axis 2. The relatively small area of

the graph (Fig. 1) occupied by E.

acutangula subsp. breviseta and E.

subsp. neotropica compared to that

occupied by E. acutangula subsp.

acutangula is presumably the result

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iables from 198 specimens

ircles), E. acutangula subsp breviseta (close

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878 Journal of the Botanical R h Institute of Texas 1(2)

TABLE 2. El | ] | i |l h principl | loadi hefi i

used in PCA of 198 speci f Eleocharis gula subsp. acutangula, E. acutangula subsp. breviseta, and E. acutangula subsp. neotropica

Character PC1 PC2 PC3

Eigenvalue 3.018 1.459 0.760

Percent variance 50.302 24,317 12.675

LONBRSTL 0.862 0.365 0.030

TBRCL 0.832 -0.348 -0.290

BRSTLNACHNL 0.828 0.324 0.040

TB 0.695 -0.643 -0.259

ACHNL 0.644 0.220 0.576

NECKWACHNW 0.015 0.799 -0.524

El as I,

acutanaula. E acutangula subsp. breviseta and E acuranaula subsp. neotronica. M

Y Y | | |

TABLE 3. Select ct isons fi

+ 1 standard deviation) are provided for quantitative characters.

Character subsp. acutangula subsp. breviseta subsp. neotropica

achene length (mm) 1.8(1.6-2) 1.6(1.4-1.7) 1.7(1.6-1.8)

achene width (mm) 1.4(1.2-1.6) 1.3(1.2-1.4) 1.4(1.3-1.5)

achene color at maturity shiny dark amber shiny dark brown shiny yellow-green (tinged

(dark brown) with amber)

tubercle width (mm) 0.7(0.6-0.9) 0.7(0.6-0.8) 0.9(0.8-1)

chene neck width (mm) 0.6(0.4—0.7) 0.5(0.4—0.6) 0.8(0.7—0.8)

tubercle length (mm) 0.6(0.5—0.8) 0.4(0.3-0.5) 0.5(0.4—0.5)

ratio of achene neck width 0.4(0.3—0.5) 0.4(0.3-0.4) 0.5(0.6-0.7)

to achene width

description of perianth usually all overtopping usually few-none all overtopping tubercle;

bristles summit of achene and overtopping summit of coarsely retrorse nearly to

sometimes the tubercle; achene; only afew short, | the base

coarsely retrorse nearly salient retrorse spinules

to the base or less often near the tips

completely smooth

length of longest perianth 2.6(2-3.3) 1.2(1-1.5) 3.6(3.2-4)

bristle (mm)

ratio of length of longest 1.4(1.1-1.7) 0.8(0.6-0.9) 2(1.9-2.3)

perianth bristle to

achene length

ratio of tubercle length 0.8(0.6-1.1) 0.6(0.5-0.8) 0.5(0.4-0.6)

width

of less morphological variability in E. acutangula subsp. breviseta and E. subsp. neotropica, which perhaps

stems from their relatively limited geographical distributions when compared with the more morphologically

variable and widespread E. acutangula subsp. acutangula. This could also indicate active speciation (local

adaptation) in E. acutangula subsp. acutangula.

DISCUSSION AND CONCLUSION

Multivariate analysis and thorough examination of ca. 600 specimens, including types, warrants the

recognition of three infraspecific taxa within Eleocharis acutangula. The presence of several conspicuous

morphological differences between the three taxa along with a relatively cohesive geographic distribution

of E. acutangula subsp. breviseta and E. subsp. neotropica (Fig. 2) suggests subspecies is an appropriate rank

for classification (Stuessy 1990). A summary of the characters accounting for most of the variability in the

multivariate analysis and our observations of achene color and the length and texture of perianth bristle

Rosen et al., T f Eleochari t |

7 J 879

spinules indicate that features of the mature achene and perianth are essential for identification of the sub-

species of E. acutangula (Table 3). Eleocharis acutangula subsp. breviseta is distinguished from E. acutangula

subsp. acutangula by its shorter achenes and tubercles and short perianth bristles with only a few short

retrorse spinules near the tips. Eleocharis acutangula subsp. neotropica differs from E. acutangula subsp. acu-

tangula by its long, soft, flexuous perianth bristles, weakly constricted achene apex, and the tubercle being

usually as wide to wider than long. Differences between E. acutangula subsp. breviseta and E. acutangula

subsp. neotropica are summarized in Table 3. Greater variability was observed in specimens referable to

Eleocharis acutangula subsp. acutangu

pany

a for several of the parameters used in the multivariate analysis (Fig.

1; Table 3). Variation was also observed in achene epidermal cell shape, often in achenes from the same

herbarium specimen (Rosen 2006).

Svenson (1929, 1939) suggested that E. fistulosa var. robusta and E. planiculmis may represent taxa distinct

from E. acutangula. However, a critical examination of the types indicates they are only minor expressions

of highly variable E. acutangula subsp. acutangula. Specimens from Madagascar [DuPuy 2429 (MO, K, P);

Bathie 17929 (P, US); and Bathie 2722 (P)], including type material of E. fistulosa, exhibited obtusely trigonous

culms rather than sharply wing-angled triquetrous culms observed in all other specimens of E. acutangula

examined. Indeed, the protologue of S. fistulosus describes the culms as “subtriquetro”. These specimens are

otherwise referable to E. acutangula subsp. acutangula for the characters used in the multivariate analysis.

We do not propose segregation on the basis of a single, highly plastic vegetative character and with such a

limited number of specimens examined.

TAXONOMIC TREATMENT

KEY TO SUBSPECIES OF ELEOCHARIS ACUTANGULA

. Longest perianth bristle 3.2-4 mm long, soft, ee re spinulose to below the middle (nearly

to the base); achene neck weakly constricted, 0.6-0.7 times achene width; tubercle 0.4-0.6 times long as

wide; mature achenes yellow-green (tinged with les distribution limited to northwest South America

subsp. neotropica

. Longest Suid bristle 3.2 mm long or shorter, a Des leia only at the tips to near m base

Bern

or sometime Spoon eas neck aS constricted, 0.6 achene width or less; tubercle 0.5-1.1

times long a e; ! ch s dark amber to us brown distribution more aa

2. Perianth nice: ee achene or rarely few to all reac spinules

restricted to the distal half or more commonly only near the tip; achene 1 ie 7 E mm long, dark brown;

3 bercle 0.3-0.5 mm lon subsp. breviseta

erianth bristles | than ach | ,Spin ems

Adel to base or rarely HT omens, Piu MEAT 6-2 mm s qd abet or rarely dark

brown; tubercle 0.5-0.8 mm long

N

subsp. acutangula

jua

. Eleocharis acutangula (Roxb.) Schult. subsp. acutangula (Fig. 3 a—b). Scirpus acutangulus Roxb. Fl. Ind. 1:216.

1820. Eleocharis acutangula (Roxb.) Schult. Mant. 2:91. 1824. Limnochloa acutangula (Roxb.) Nees. Contr. Bot. India 114. 1834. Tver:

INDIA, Roxburgh s.n. (Lectotype here designated: BM [BM000847992]!).

Scirpus medius Roxb. Fl. Ind. 1:216. 1820. Limnochloa media (Roxb.) Nees. Contr. Bot. India 114. 1834. Tyre: INDIA, Roxburgh s.n. (not

found)

Scirpus fistulosus Poir. Encyclopédie Méthodique, Botanique 6:749. 1804. nom. illeg., non Scirpus fistulosus Forssk. 1775. Eleocharis

fistulosa Schult. Mant. 2:89. 1824. Type: MADAGASCAR, Poiret s.n. (HOLOTYPE: P [Herbier du Petit-Thouars., P00376392]!; ISOTYPE:

Herb. Poiret in Herb. Moquin-Tandon [P00370140]9.

Eleocharis pu osa Schult. var. robusta Boeck. Flora 62:563. 1876. Heleocharis jua o Hess. Ber. Schweiz. Bot. Ges. 63:331.

cum descr. ampl. Type: AFRICA, Africa centralis, Seriba Ghass g ep 1869, Schweinfurth 2326 (LECTOTYPE

here designated: GH!; ISOLECTOTYPES: Z [000006263, OI

Eleocharis planiculmis Steud. Syn. Pl. Glumac. 2:80. 1855. Tyre: Java, Zollinger 281 (notorvee: P [P00368895]!; isotypes: P [P00368896,

P00368897]!, K [K000290949, onto

Heleocharis pseudofistulosa H. Hess. Ber. Schweiz. Bot. Ges. 63:329. 1953. Type: ANGOLA: Provinz Huila, Guanhama, Tümpel, 15 km

südlich Cubango an der Strasse pu Cassinga, 14 Jan 1952, Hess 52/220 (HoLotyre: ZT photo! [based on Hesss designation of

“Typus”]; isorvre: BOL, K, TAES!, Z

880 Journal of the Botanical R h Institute of Texas 1(2)

Fic. 2. Distributi f Eleochari ti I b ti la ( irdes), E. acutanaula subsp breviseta (closed circles), and E. acutangula subsp

| \ g \ n g

Plants perennial. Roots coarse, fibrous, drab-brown to reddish, small storage structures present in care-

fully collected plants, cylindrical-reniform, brown; primary rhizomes caudex-like, thick, hard, ascending,

concealed by roots and persistent culm bases (occurring only in carefully collected specimens); secondary

rhizomes elongated, to 4 mm thick, scales to 17 mm long (few seen). Culms triquetrous (a few specimens from

Madagascar trigonous) distally, (25—)38-81(-135) cm long x (1.2-)2.1-4.4(-6.5) mm wide, soft, internally

spongy, with incomplete transverse septa, smooth, green, finely longitudinally striate when dry. Leaves 2,

reduced to sheaths, apically oblique, membranous, loose, friable, proximally pinkish to dark maroon (dark

purplish), distally drab, apex acute. Spikelets cylindric, (11-)21-39(-56) mm long x (2.5-) 3.2-4.8(-6)

mm wide, acute; proximal scale with flower, obtuse, amplexicaul-clasping, appearing as a continuation of

culm, remaining floral scales conspicuously spirally arranged, appressed to somewhat spreading at maturity,

ovate-oblong, 2.5—)3.8-5.3(-6) mm long x (1.7-)2.3-3.4(-4.8) mm wide, cartilaginous, abaxially greenish to

stramineous lly, j ginally, sparsely red-maculate and sometimes the veins or other areas

reddish or pinkish (purplish) tinged, usually with a fine dark band near apex, adaxially sparsely to copiously

red-maculate, apex acute (rounded), distal 0.1-0.5 mm translucent hyaline-erose, central area nearly flat,

coarsely many veined, only mid-vein conspicuous in adaxial view. Flowers with 6—7(-8) perianth bristles;

bristles sub-equal, usually 1.1-1.8 times the length of the achene (rarely one or few just reaching its summit

or slightly shorter), retrorsely spinulose nearly to base or rarely completely smooth (both conditions can

occur in same population), stramineous or pinkish to dark maroon; stamens 3; anthers (1.1-)1.3—2.2(-3.2)

mm long, stramineous; style 3-fid. Achenes biconvex, very broadly obovoid to obovoid, the shoulders and

sides near the apex usually straight and forming an obtuse angle, or sometimes rounded, (1.4-)1.6-2.0(-2.2)

mm long x (1.02)1.3-1.6(-1.8) mm wide, with (11212-15(-19) longitudinal rows of deeply concave trans-

versely oblong to linear polygonal cells visible through transparent periclinal layer on each achene face, dull

yellow-buff maturing to shiny dark amber (dark brown), apex constricted to a distinct neck about 0.3-0.5

times the width of achene. Tubercle dorsoventrally compressed, shallowly triangular-deltate (triangular),

0.5-0.8(-1.1) mm long x (0.5-)0.6-0.9-1.2) mm wide, stramineous, maturing to dark brown.

Rosen et al., T f Eleochari t | 881

Lectotypification of Eleocharis acutangula

Eleocharis acutangula was described by Roxburgh (1820) as Scirpus acutangulus based on plants from India. As

is the case with apparently all Roxburgh names, no type specimen was designated (Forman 1997). Schultes

transferred S. acutangulus to Eleocharis without indicating a type. A literature search revealed no reference to

a particular type specimen although a number of authors indicate the “type” is from India (e.g., Haines &

Lye 1983; Gordon-Gray 1995; Browning et al. 1997). Typifying Roxburgh names can be difficult since his

specimens were widely distributed, making locating specimens annotated by him or known to be associ-

ated with him challenging (Forman 1997). Almost all of Roxburgh’s nearly 2600 species were illustrated by

color drawings prepared by local Indian artists; the original set is at CAL, and a duplicate set at K (Sanjappa

et al. 1991). Forman (1997) indicated that the Flora Indica drawings were often superior to the correspond-

ing Roxburgh specimen (if one can be found), and in some instances make a better choice for a type. From

the set of drawings at Kew a high resolution digital photograph was obtained of the front and back of the

drawing of S. acutangulus. The drawing, a stylized depiction of an immature plant, was annotated in what

the first author interprets as Roxburgh’s hand.

We made queries to curators at key herbaria indicated by Forman (1997) in an effort to locate an au-

thentic Roxburgh specimen. Mark Spencer (BM) presented a specimen (BM-000847992) that he considers

to have been associated with Roxburgh, the most compelling evidence being annotations on the verso and

front of the specimen. The verso is annotated “Ind. Orient Roxburgh” in an unknown hand, indicating that

the specimen was received from Roxburgh, and the front was annotated “72” in what the first author inter-

prets to be Roxburgh’s hand. We selected this specimen as the lectotype of Scirpus acutangulus since it fits the

description in the protologue and is thought with reasonable certainty to have been used by Roxburgh.

Problems with Typification of Scirpus medius

Roxburgh described Scirpus medius as being similar to S. acutangulus but having shorter culms with smooth,

rounded angles. Nees (1842) transferred the name to the genus Limnochloa. Roxburgh’s description of the

culms as having rounded angles is troublesome, as all Asian specimens of Eleocharis acutangula examined

during this research had triquetrous culms. No specimens annotated as S. medius were seen by us, and no

specimens were located in herbaria where Roxburgh’s specimens were distributed. Thus, we follow Svenson

(1929) and others (Blake 1939; Koyama 1985; Gordon-Gray 1995) in placing S. medius in synonymy under

E. acutangula.

Clarification of the Authorship of Eleocharis fistulosa

Scirpus fistulosus Poir. is illegitimate because of an earlier homonym, S. fistulosus Forsskal. Eleocharis fistulosa

Link is also invalid because Link failed to associate the specific epithet with the name of the genus or species,

or with its abbreviation, as mandated by the Art. 33.1 of the ICBN (McNeill et al. 2006). Thus, E. fistulosa

Schult. is the correct author citation (See ICBN Articles 58.1; 7.5; and 33, Note 2.). Since the priority of E.

fistulosa does not date back to the publication of Poiret's illegitimate use, E. acutangula (Roxb.) Schult. has

priority as the oldest legitimate name for the species.

Lectotypification of Eleocharis fistulosa var. robusta

Eleocharis fistulosa var. robusta was described by Boeckeler based on Schweinfurth 2326 from Central Africa.

Boeckeler's types were at B, and, if the holotype of E. fistulosa var. robusta was ever extant at B, it was destroyed

by the fire of 1943 (Robert Vogt, B, pers. comm .). In this case, Schweinfurth 2326 from GH is designated as

lectotype, and two duplicates from Z become isolectotypes.

Excluded Name

Eleocharis fistulosa var. micrantha Chermezon was described from specimens from Senegal (Chermezon

1936). Attempts to locate type specimens cited in the protologue have been unsuccessful thus far. Svenson

(1939) relegated this name to synonymy under E. nupeensis Hutchinson & Dalziel based on the description,

a temporary solution adopted here.

882 Journal of the Botanical R h Institute of Texas 1(2)

Sa

SEED

ESOS

L F

+ 1 dicts] E E " f£. b LAM HI E 4 I I

Fic. 3. A-B Eleocharis acutangula subsp. acutangula: A abaxial view of act B apical view of achene. C-H E. acutangula subsp. breviseta: C habit;

| p ; ; | l-K E.

acutangula subsp pi [ iew of ad 1 K apical view of act A-B from Hooper € Gandhi 2373,

C-F from Rosen & Carter 3206, G-H from Howard & Howard 9862, and I-K from McDaniel & Rimachi 18552. Drawn by Neva Mikulicz.

P lecnibAl I Aer nf | A A

Representative Specimens Examined: NORTH AMERICA. MEXICO: Chiapas: E side of Pueblo Solistahuacan, Municipio of Pueblo

Nuevo Solistahuacan, elev. 1700 m, 26 Oct 1971, Breedlove 21527 (MO, NY). Hidalgo: Lake Atexca below Molango, 09 Nov 1946, Moore

1938 (GH). Jalisco: I Guadalajara, 1888, Pringle 2061 (NY). Nayarit: near Lake Labor, ca 15 mi SE of Tepic, 25 Sep 1960,

McVaugh 19426 (MICH). Tabasco: km 64 rumbo de Huimanguillo a Fransisco Rueda, 35 msnm, 06 Nov 1979, Orozco & Zamudio 2187

(MO). Veracruz-Llave: Mpio. Las Choapas, ca 5.4 km S of the town of Las Choapas, along rural road to El Chichon, 13 Jul 2006, Rosen

et al. 3870 (CIIDIR, GH, K, MICH, MO, TAES, TEX, US, VSC, WIS). CENTRAL AMERICA. GUATEMALA: Alta Verapaz: E of Tactic,

alt 1,300 m, 20 Feb 1942, Steyermark 43970 (F). Chiquimula: between Chiquimula and La Laguna, alt. 500-1000 m, 27 Oct 1939,

Steyermark 30713 (F). Huehuetenango: vicinity of Maxbal, ca. 17 mi N of Barillas, Sierra de los Cuchumatanes, alt. 1500 m, 15-16 Jul

1942, Steyermark 48770 (E). Izabal: near Puerto Barrios, sea level, 25 Apr 06 May 1939, Standley 72862 (F). Jutiapa: SE end of Potrero

Rosen et al., T

y of Eleocharis acutangul 883

Carrillo, 13 mi NE of Jalapa, alt. 1500-1700 m, 12 Dec 1939, Steyermark 33099 (F). Santa Rosa: 4 mi N of Barberena, 18 Feb 1951,

Fassett 28844 (F). BELIZE: Toledo District: near junction of Southern Highway and Pine Hill, 22 Nov 1998, Holst et al. 7064 (MO).

HONDURAS: Comayagua: beu of Siguatepeque, ca. 1050 m, 25 Mar 05-Apr 1947, Standley & Chacon 6595 (F). Copan: 14 Jul

1971, Harmon & Fuentes 6445 (MO, NY-2 sheets). Francisco Morazan: near Las Mesas, 900 m, 10 Sep 1950, Standley 26634 (GH).

Olancho: Santa Maria del BN 23 mi NE of San Esteban along road to Bonito Oriental, 03 Jul 1994, Davidse et al. 35564 (CIIDIR).

EL SALVADOR: Ahuachapan: Lagunita las Ninfas, Apaneca, 28 Jan 1951, Fassett 28721 (GH). NICARAGUA: Comarca del Cabo:

Bihmona, 7 Jul 1972, Seymor 5707 (CIIDIR, GH, MO-mixed with E. interstincta). Esteli: Reserva Natural Miraflor, Municipio de Esteli,

Comunidad los Volcancitos, 10 Jul 1999, Rueda et al. 11643 (MO). Zelaya: Cano Manso Awalka Tingni, reached by Geodesia turn on

road between Torre 7 and Bismuna Tara, ca. 11.9 km SW of Bismuna Tarra, 19 Apr 1978, Stevens 7704 (CIIDIR-2 sheets, MO). COSTA

RICA: Isla De Cocos: Macollas en suelo humedo, a la orilla del par tano, Bahia de Wafer, ca. nivel del mar, 31 Jul 1981, Gomez-Laurito

6915 (F, MO). Alajuela: 6 km W of Venicia, elev. 450 m, 15 Oct 1968, Davidse & Pohl 1307 (F, MO). Cartago: Laguna Dona Anacleta,

Canton Paraiso, Lago Crater, 22 Aug 1983, Novelo 1209 (MO). Guanacaste: upper N fork of Rio Sabalito, just N of San Joaquin de Coto

Brus, 13 Sep 1985, Grayum et al. 6011 (MO). Puntarenas: San Joachim de Sabosa, just N of San Vito, 22 Feb 1982, Barringer & Gomez

1688 (F). PANAMA: Chiriqui: S of El Boquete, 01 Mar 1918, Killip 4569 (NY). Cocle: El Valle de Anton and vicinity, 500—700 m, 23-27

Jul 1935, Seibert 476 (MO, NY, US). Veraguas: vicinity of La Mesa in sunny muddy bottom in pasture, 28 Dec 1968, Tyson 6054 (MO).

CARIBBEAN BASIN. JAMAICA: Clarendon Parish: Mason River Field Station, 4 mi W of Kellitts, 2300 m, 27-29 Jul 1979, Thomas

2146 (MICH). Saint Catherine Parish: Charlton, near Ewarton, 03 Apr 1903, Harris 8513 (NY). DOMINICAN REPUBLIC: San

Cristóbal Province: between Duarte Hwy. Kl. 28 and Haina, 11 Oct 1947, Allard 15976 Cee Santa Domingo Province: 8 km from

La Batata on road to Mata de Piedra and La Catalina, 09 Dec 1980, Mejia & Zar . DOMINICA: Lesser Antilles, St. George

Parish, vicinity of freshwater lake, NE of Laudat, E side of Morne Macaque, loccaly common, 2500’, 20 Mar 1991, Hill et al. 22119 (GH,

R

regimiento de Puerto Valdivia, km 11 de Pto a mina uc Oro "Canarias", colecciones en escombreras de 2 anos, 14 May 1987,

Callejas et al. 3486 (MO-—2 sheets, NY). Risaral La Virginia-Cerrito, extremo norte de parte ancha

del Valle del Rio Cauca, lomas bajas, 22 Aug 1989, Silverstone- ae 5504 (MO). Cauca: Chisquio, Finca Los Derrumbos, alt. c. 1700

m, 11 May 1940, Asplund 10577 (LL). Huila: 3 km W of Garzon, upper basin of Rio Magdalena, 17 Feb 1959, Mason 13888 (GH, US).

Meta: ca. 17 km SW of Puerto Lopez, along road between La Balsa and Bocas del Guayuriba, 17 Jan 1970, Schuyler 4165 (PH). Valle del

Cauca: Calima, on Rio Calima, 14-15 Sep 1922, Killip 11247 (GH, NY, PH). VENEZUELA: Apure: Guanare, Esteros y pantanos cerca

de los AA cano Mau 25 Oct 1980, Stergios 2387 (MO). Aragua: El Limon, near Maracay, in Morass, 29 Jan 1922, Pittier 10116

(GH). Bolivar: ato Sta. Teresa, Mar 1946, Tamayo 3211 (F, US). Guarico: Orituco, 25 km SW de la Estacion Biologice de

la Clanus (sic) Edo Gu 19 o 1982, Montes 1343 (MO). Portuguesa: terrenos de la Unellez, 06 Sep 1984, LU 7051 (MO).

GUYANA: Upper Takutu-Upper Essequibo Region: Rupununi Distr., Shea Village, 09 Feb 1994, Jansen-Jacobs et al. 3634 (NY, US).

SURINAM: in Maurisie swamp, W of 4-Gebroeders Mts., 27 Sep 1968, Oldenberger et al. 194 (NY). ECUADOR: ao Amazonica,

Archidona, Coca km 9.3 roadside, 15 Apr 1988, Laegaard & Renvoize 70909 (MO, NY). Pastaza: Amazonica, Hacienda San Antonio de

Baron von Humboldt, 2 km al Nede Mera, 20 Feb-20 Mar 1985, Palacios et al. 144 (MO). BRAZIL: Distrito Federal: Brasilia, 27 Jun

1979, Heringer 1652 (NY). Bahia: 37 km N from Correntina, on the Inhaumas road, 29 Apr 1980, Harley 21957 (NY). Mato Grosso: 17

Oct 1968, Harley et al. 10711 (NY). Minas Gerais: without location, 1816-1821, Catal 616 (K, P). Sao Paulo: Butantan, S. Paulo, with-

out date, Gehrt 5403 (GH, NY). Amazonas: lagoa permanente, 500 m ao Sul da BR 230 km 4, 15 Aug 1980, Janssen & Gemtchujnicov 514

(M). Maranhao: Brejo, Ets. Ecologica UFMG, 02 Apr 1991, Neto 461 (CIIDIR). Parana: Rolandia, Fazenda Conquista, area alagada,

dentro da lagoa, no. 14, 11 Mar 2003, Vanzela 35.42 (CIIDIR). Rio de Janeiro: Goias, Formosa, Bisual, 20 Oct 1965, Pereira & Duarte

9414 (NY). Rio Grande do Norte: near Bento Fernandes, 70 km W from Natal, shallow at pond’s edge, 28 Aug 1987, Tsugaru & Sano

B-1273 (GH). Rio Grande do Sul: M. Rio Pardo, Riniao Reserva, Feb 1923, Jurgens s.n. (US). Without location, 1844, Weddele 1195 (P).

BOLIVIA: La Paz: Iturralde, Luisita, sabana humeda, W del rio Beni, Palmar, 12 Sep 1984, Haase 540 (NY). Santa Cruz: Andres Ibanez,

NE side of Viru-Viru Pampa and property of Aeropuerto Internacional, along road to Chuchio, 4.5 km E of turnoff from highway from

Santa Cruz to Warnes on road to Chuchio, 15 May 1998, Nee 49365 (CIIDIR, TEX). PARAGUAY: Departamento Central: Estero del

Ypoa, Villeta, Puerto Guyrati, 4.5 km S of Villeta, 02 Dec 1992, Zardini & Aquino 34134 (CIIDIR, US). Caazapa: Tavai, Enrramadita, 05

Dec 1988, Mereles 2067, 2069, 2070, 2081 (MO). la Cordillera: 1 km E of Nueva Colombia on road to Atyra, 09 Jun 1990, Zardini &

Velazquez 20917 (MO). Misiones: Estancia La Soledad, Santiago, 30 Apr 1961, Pedersen 6029 (US, GH, MO, NY, TEX). Paraguari:

Estero Ypoa, between Nueva Italia and Yuquyty ona hill, 18 Mar 1992, Zardini & Aquino 31333 (CIIDIR). Amambay: Ao. Estrella, Prop.

De Heisecke, 08 May 1989, Soria 3749 (MO). ARGEN oncepcion, Carambola, Estancis “Buena Vista”, 19 Feb 1985,

Pedersen 14072 (MO, NY). Misi Aira, 1913, Rodrig 3 (GH). Chaco: Dep lo de Mayo, Colonia Benite balsado burger,

16 Dec 1943, Schulz 4118 (F, GH). AFRICA. ANGOLA: Benguela: Gebirge sudlich Ganda, Tumpel bei Calusipa, 30 km sudlich Chicu-

ma, 1580 m, 24 Dec 1951, Hess 51/419 (Z). Bie Bie: Baixo Cubango, 28 km nordlich Caiundo in der Umgebung der Missao cat. Capico,

flacher sumpf, 31 Jan 1952, Hess 52/525 (Z). BOTSWANA: Ngamiland District: Moremi Wildlife Reserve, N Okavango swamp, Kwani

River floodplain, Jul 1964, Tinley 1057 (NU). BURKINA FASO: I’ ene Mare de Bidi, 20 Sep 1996, Madsen 5749 (NY). Boulgou:

some km SE of Tenkodogo, 31 Aug 1996, Madsen 5455 (NY). B JNDI: B Plaine Rusizi km 14, 800 m, 16 Mar 1975, Reekmans

4390 (MO). Bujumbura: Bujumbura, plaine Rusizi km 14, 780 m, 13 Feb 1972, Reekmans 1539 (MO). Provense ya Bururi: Giho

Gihara, 20 May 1980, Reel 205 (MO). CAMEROON: Nord: ca. 15 km NE of Maroua, along road to Waza, 12 Sep 1964, Wilde 3215

(K). CENTRAL AFRICAN REPUBLIC: Prefecture de la Sangha-Mbaere: Sangha Economique, Dzanga-Sangha Reserve, 40 km S of

884 Journal of the Botanical R h Institute of Texas 1(2)

Lidjombo on tributary of Keine, 26 Oct 1988, Harris & Fay 1488 (MO, PRE). COMORO ISLANDS: Mayotte, Grande Terre, Ouangani,

Coconi, Valarana, 26 Fev 2002, Barthelat 6 Sifari 708 (P). CONGO: Vallie Uruanda, 26 Oct 1953, Liben 852 (K, PRE). ETHIOPIA:

Kaffa Province: d ca. 5 km E of Jimma, along bi b to Addis Ababa, 02 Nov 1970, Friis et al. 38 (C, K). GABON: Nyanga: à

plus ou moins 7km sur la route de Doussala vers B direction Nord-Ouest, Petit nds 25 Mar 2000, Sosef 1016 (MO). Haut-

yon Bateke Plateau, Mpassa River watershed, 4. 2 km N of station of the Project de Protection Gorilles, 27 Nov 2001, Walters et

1. 982 (MO). GHANA: Brong-Ahafo Region: 1 m S of Atebubu, 16 Nov 1970, Hall & Duodu 42128 (MO). Guinea-Bissau: 08 Dec 1944,

o 1594 (MO). Ivory Coast: d’ Abidj f Abidjan, 21 Oct 1963, Wilde 1109 (Z). LIBERIA: Grand

Bassa County: Sanokwele Dist., Ganta, 02 Dec 1935, Harley 781 (NY, US. Nimba County: Mt. Nimba, Crete, Marc a’, 04 Jul 1974,

Adam 28878 (MO). MADAGASCAR: Antananarivo Province: Ankazobe, Jun 1927, Bathie 17929 (P, US). Fianarantsoa Province:

Ambatofinandrahana, Itremo, petite vallee a l'ouest du Massif de l'itremo, 1680 m, 26 Nov 1993, Du Puy & Andriantiana 2429 (MO, K,

anga Province: 10 km E Antsalova, 20 Mar 1993, Villiers et al. 4855 (K). Toamasina province, Marais de Didy, voir Joncacee

ef aa2m, 21 Feb LE Cours 1758 (P). Malawi: C l Region: Kasungu National Park, Angombe Hill, 03 Sep 1970, Hall-Martin 1712

(PRE). Nigeria: Kano: 12 Sep 1973, Jackson & Apcjoye 10-12973 (MO). Plateau: in vicinity of Bukuru, near Jos, 29 Jun 1970, Blum 2488

(WIS). Rhodesia: Hartley: Avondale farm dam, 25 Feb 1969, Mavi 983 (NU). S.W.A. (Southwest Africa?): 8 km S of Makuri vlei on

road to Gimsa, 03 Mar 1985, Hines 361 (PRE). Senegal: Kaolak Region: Kaolak, Nov 1824, Berhaut s.n. (Z). Sierra Leone: Elliot 4453

(GH). SOUTH AFRIC Province: Transvaal, Witklip S bos, Nelspruit Dist., in water in Witklipdam, Kruid, 27 Jan

1976, Kluge 862 (PRE). KwazalacNacal: North coast, Lake Nhlabane area, W corner of North Lake, 19 Sep 1991, Ward 11378 (PRE, NU,

NH red SUDAN: nan. Nyany, nr. Maar, 80 km N of Bor, 01 Feb 1981, Lock 81/10 (K). Swaziland: Malolotja Nature Reserve,

Yers dam, stream, 17 Dec 1985, Heath 406 (PRE). Tanzania: Dar es Salaam Region: Mbezi, 2km WNW of Dar es Salaam

SM (by cattle E to Tanzania packers), 12 Jun 1974, Wingfield 2752 (MO). Iringa District: T7, km 13 Ufinda-Sao Hill Rd., E

side of road, 10 Jun 1996, Faden et al. 96/130 (K, US). Ruvuma Region: Ruanda, Urundi, Vallie Uruanda, Oct 1953, Liben 852 (M). Zaire:

P Equateur: Bikoro, 01 Oct 1957, Thonet 7 (M). Kasai-Occidental: Kabinda, 26 Jul 1934, Becquaert 62 (GH). Katanga: River Kalule,

pres de la ferme Rostenne, Elisabethville, 27 Mar 1963, Symoens 10155 (K). Province Orientale: 1940, Germain 171 (M). Zambia:

Central Province: Mkushi Dist., David Moffat's farm, Munchiwemba dambo, 20 Sep 1995, Bingham & Nkhoma 9711 (PRE). Luapula

Province: Lake Bangweulu, S part, swamps between Ncheta Island and Chibambo Lagoon, 11 Feb 1996, Renvoize 5585 (K). Northern

Province: 8 km N of Kasama, 22 Jan 1961, Robinson 4296 (K, MO, NU). Western Province: ca. 10 mi NE of Mongu, 18 Nov 1959,

Drummond & Coohson 6597 (MO). Zimbabwe: Manicaland: Mare Dam, Rhodes Inyanga National Park, 06 Jan 1972, Gibbs Russell 1210

(M, MO-2 sheets, K). Matabeleland North: Wankie National Park, Ngamo Pans 54 mi SE of main camp, 17 Apr 1972, Russell 1645

(NU). Salisbury District: 6 mi spruit, 4800’, 10 Jan 1932. ASIA. CHINA: Huebi: Central China, 1885-1888, Henry 4102 (GH, US-2

sheets). Yunnan: 1530 m, May 1936, Wang 73552 (GH). INDIA: Bangladesh: East Bengal, 1863-64, Griffith 6235 (NY). Karnataka:

near Station, 10 Nov 1971, Hooper & Gandhi 2373 (MO, NY). Kerala: Malappuram Dist., between Tirurangadi and Parappanangadi,

almost sea level, 12 Nov 1993, Cook & Camenisch 5169 (Z). Maharashtra: Pashan, near Poona, lake margin, 30 Dec 1971, Hooper 112

(K). Nepaul: without date, Hook & Thomson s.n. (NY). Tamil Nadu: Dharmapuri, Denl luk to Jowalagiri, to Karareddy pond,

18 Dec 1978, Matthew & Venugopal 2041 . INDONESIA: Alor: 1938, Jaag? s.n. (ZT). Java: Meester Cornelis, 1991, unknown 23139

(K). Jawa: Barat, Banten, 1936, Hachenberg 1 (GH). JAPAN: Hondo: Shinjo in Kii, 11 Oct 1953, } i

Nagaitani valley, 15 Oct 1953, Koyama 5885 (MO, NY). Kyushu-chiho: Hondo, E in Kii, 11 Oct 1953, F 838 (BRIT, NY, US,

WIS). Yoron-jima: Liukiuensis, 30 Aug 1921, Uyehara s.n. (US). MALAY PENINSULA: Langkowi, ricefields near Kueh, 14 Nov 1941,

Comes? 37973 (K). MALAYSIA: Malacca: Kampong Bukit Piatu, 02 Apr 1955, es 40551 (K). PHILIPPINE ISLANDS: Lanao

Mindanao: in 6” of water pocket in grassland, 04 E 1938, Zwickey 50 (GH, US). SIAM: growing in open fields, 13 Jul 1968, Kerr 15798

(K). SRI LANKA: North Eastern Province: Amparai Dist., Helawe Piya ca. 7 miS m Panama, E of Helawe Lagoon, sea level, 08 Feb

1971, Koyama et al. 14026 (GH, NY). VIETNAM: Quang Nam-Dà Nang: T ( ): Annam, Mount Bani, in tl

ca. 25 km from Tourane (Da Nang), May—Jul 1927, Clemens & Clemens 4050 (E, X, MO, NY, PH, US, US, Z). OCEANIA. AUSTRALIA:

Queensland: Cook Dist., Abattoir swamp, 4.5 km N of Mount Molloy, 20 May 1995, Clarkson 10317 (BRD. PAPUA-NEW GUINEA:

Morobe District: vicinity of Kajabit Mission, n» 800-2000 ft, Aug-Dec 1939, Clemens 10600 (GH, US). National Capitol District:

Hohola Port Moresby, 14 Nov 1973, White 37815 (BRI, GH, K, M, US)

Distribution. —Pantropical; in México from the states of Chiapas, Hidalgo, Jalisco, Nayarit, Tabasco, and

Veracruz-Llave. In Central America known from Belize, Costa Rica, El Salvador, Guatemala, Honduras,

Nicaragua, and Panama. In the Caribbean Basin known from Dominica, Dominican Republic, Grenada, and

Jamaica. In South America known from records in Argentina, Brazil, Colombia, Ecuador, Guyana, Paraguay,

Surinam, and Venezuela. Some previous reports of Eleocharis acutangula from the Galápagos Archipelago are

based on misidentified specimens of E. obtusetrigona (Stewart 1911). Other reports were not verified (e.g.,

Jergensen & León-Yánez 1999), and no authentic specimens of E. acutangula from the Galá Archipelago

have been seen. Perhaps most widespread and occurring in more variety of habitats in Sd Africa, with

records from Angola, Botswana, Burkina Faso, Burundi, Cameroon, Central African Republic, Comoro Is-

lands, Congo, Ethiopia, Gabon, Ghana, Gunea-Bissau, Ivory Coast, Liberia, Madagascar, Malawi, Nigeria,

Rosen et al., T f Eleochari t | 885

Rhodesia, Senegal, Sierra Leone, South Africa, Sudan, Swaziland, Tanzania, Zaire, Zambia, and Zimbabwe.

Of sporadic distribution in Asia and Oceania with records from Australia, China, India, Indonesia, Japan,

Malaysia, Papua-New Guinea, Philippine Islands, Siam, Sri Lanka, and Vietnam.

—Various disturbed and natural freshwater herbaceous and forested wetlands including marshy

open grasslands, coastal savannas, and tropical forests from sea level to 2300 m. Reportedly forms expansive

stands on a variety of soil types usually associated with other aquatic plants. Weedy in rice and other crop rota-

tions and aquatic habitats, and used as a fiber crop in Borneo, Brazil, and Sumatra (Simpson & Inglis 2001).

Note.—Eleocharis acutangula subsp. acutangula as treated here remains a variable taxon and includes

forms meriting additional systematic study. Of particular interest are plants reviewed from Madagascar

(including the type of E. fistulosa) with obtusely trigonous culms.

2. Eleocharis acutangula (Roxb.) Schult. subsp. breviseta D.J. Rosen, subsp. nov. (Fig. 3 c-h). Tyee: DOMINI-

CAN REPUBLIC: El Seibo Province, 3-7 Nov 1946, Howard & Howard 9862 (HoLoTYPE: GH!; Isotype: NY-2 sheets!, P!, US).

A Eleochari la (Roxb.) Schult. subsp l ianthii setis | ioril inuli i i i acheniis

co) E E E E E E

parvibus et lona brevioribus recedit.

Plants perennial. Roots coarse, fibrous, mostly maroon (a few drab-brown), small storage structures pres-

ent in carefully collected plants, cylindrical-reniform, white; primary rhizomes caudex-like, thick, hard,

ascending, EU AME by roots and persistent culm bases (occurring only in carefully collected specimens);

secondary rl elongated, to 3 mm thick, scales to 9 mm long (few seen). Culms triquetrous, (19-)30—

712134) cm long x (1.12)1.5—3.7(-7) mm wide, soft, internally spongy, with incomplete transverse septa,

smooth, green when fresh, finely longitudinally striate when dry. Leaves 2, reduced to sheaths, apically

oblique, membranous, loose, friable (upper distil portion disintegrating when submerged), proximally dark

maroon, distally drab, apex acute. Spikelets cylindric, (10217-34(-49) mm long x Q.22)2.7-4.1075.5)

mm wide, acute; proximal scale with flower, obtuse, amplexicaul-clasping, appearing as a continuation of

culm; remaining floral scales conspicuously spirally arranged, appressed to somewhat spreading at maturity,

ovate-oblong, (3.1-)3.3—-4.5(-5.9) mm long x (1.4-)1.8-3(-4.0) mm wide, cartilaginous, abaxially greenish

centrally, stramineous marginally and sometimes reddish or pinkish tinged, with a fine dark band near apex,

adaxially sparsely to copiously red-maculate, apex acute (rounded), distal 0.1-0.4 mm translucent hyaline-

erose, central area nearly flat, coarsely many veined, only mid-vein conspicuous in adaxial view. Flowers

with (5—)6-7(-8) perianth bristles, bristles sub-equal, (0.4—)0.6-1(-1.2) times the length of achene (rarely

few-all bristles overtopping achene summit) with only a few short, salient retrorse spinules near tips (rarely

spinules present in distal half), stramineous or pinkish to dark maroon; stamens 3; anthers (0.921.1-1.9(-

2.7) mm long, stramineous; style 3-fid. Achenes biconvex, broadly obovoid, the shoulders and sides near

the apex usually straight and forming an obtuse angle, (1.3-)1.4-1.822.1) mm long x (1.121.2-1.4C1.6)

mm wide, with (9211-14(-16) longitudinal rows of deeply concave transversely oblong (linear) polygonal

cells visible through transparent periclinal layer on each achene face, dull yellow-green maturing through

amber to shiny dark brown, apex constricted to a distinct neck about 0.4 times width of achene, in the field

achenes sometimes persistent after the floral scales have shed giving spikelet a beaded appearance. Tubercle

dorsoventrally compressed, shallowly triangular, (0.22)0.3-0.5(-0.6) mm long x (0.5-J0.6-0.8=1) mm

wide, light brown tinged with green, maturing to dark brown.

Specimens examined: NORTH AMERICA. U.S.A: Florida: Lee Co.: 4 km SW of the intersection of Hwy. 82 and Green Meadows

Rd., SE of Fort Myers, 12 Nov 2004, Rosen 3206 & Carter (CIIDIR, GH, K, MEXU, MICH, MO, NY, P, TAES, TEX, US, VSC, WIS), S side

of Griffin Rd., just S of entrance to Pinewood Lakes in Gateway, 28 Oct 1993, Orzell and Bridges 22526 (BRIT, FTG, USF). MEXICO:

Campeche: a aprox. 10 km al sureste de la ciudad de Campeche, Mun. Campeche, alt. 80 m, 10 Nov 1980, Novelo & Zetina 721 (TEX),

14 km N of Ocozocoautla on road to Mal Paso, Municipio of Ocozocoautla de Espinosa, 07 Oct 1974, Breedlove 38254 (MEXU, MO).

Chiapas: 96 km S of Mexican Hwy. 190 on road to Nuevo Concordia, 10 Oct 1974, Breedlove 38516 (NY). Guerrero: 1.5 km al NW del

Rincon de la Via, 28 E 1988, Verduzco 389 (MEXU). CENTRAL AMERICA. PANAMA: vicinity of El Llano, 7-8 Sep 1962, Duke 5526

(MO, USF), Near the big he Rio Tecumen Province, 11 Dec 1923, Standley 26509 (MO). Canal Zone: Laguna de Portala,

near Chepo, province a Oct 1911, Pittier 4602 (NY). Panama: Sabanas near Chepo, 30 m, 20 Jan 1935, Hunter & Allen 87 (MO),

1.6 km W of Juan Diaz, 10 Oct 1917, Killip 4090 (PH, RSA, US), Camino del Boticario, near Chepo, altitude 30 to 50 meters, Oct 1911,

886 Journal of the Botanical R h Institute of Texas 1(2)

Pittier pr (GH, NY, US), near Matias Hernandez, wet ne 30 Dec 1923, Standley 28909 (US), near Matias Hernandez, wet field, 30 Dec

984 (US). CARIBBEAN BASIN. CUBA La Habana: Vedado-Habana, Sabana de Monasterio, 23 Jun 1920,

Leon 9215 (NY). a location, 1860-1864, Wright 3376 (GH, mixed with E. mutata, MO, NY, P). SOUTH AMERICA. COLOMBIA:

Magdalena: Rincon Hondo, Magdalena Valley, 10 Aug 1924, Allen 357 (MO). Vaupes: Rio Vaupes, Mitu y alrededores, 08 Sep 1951,

Schultes & Cabrera 13977 (GH), Rio Vaupes, Mitu and vicinity, 09 May 1953, Schultes & Cabrera 19257 (GH-2 sheets). VENEZUELA:

Bolivar: 27 km SW of Caicara along Hwy. 19 to Ciudad Bolivar, 22 Nov 1973, Davidse 4355 (MO). Tachira: between La Rochela and La

Espuma, SW of Santo Domingo, 31 Jul 1979, Steyermark & Liesner 119299 (MO). Zulia: Perija, carretera Calle Larga-San Felipe-Jaguacita,

km 25 al SE de San Felipe, 09 Oct 1977, Bunting 5656 (NY), ca 50 km SSW s pu by air, 19 km W of main road, 26 Mar 1982,

Liesner & Gonzalez 13183 (NY). GUYANA: Upper Takutu-U Baboon Hill ee Tau) 1.5 km S of Sand Creek

Village, 21 Jun 1989, Gillespie et al. 1803 (NY). ECUADOR ane Rios: p e San Juan, Vince, 07 Mar 1988, Laega

70652 (K, NY). BRAZIL: Mato Grosso do Sul: Pantanal do Miranda-Abobral, Passo do Lontra, Rodovia MS 122, e Sao Bento,

depois da 2a porteira, 11 Jul 1997, Rodrigues et al. 3 (K). Rondonia: Guapore, Porto Velho, 1952, Cordeiro & Silva 270 (US). Amapa:

Rio Macacoari, Municipio de Macapa, 05 Aug 1951, Froes & Black 27231 (US). Parana: Curitiba, Paso do Lontra O Miranda) Mato

Grosso, 13 Oct 1972, Hatschbach & Scherer 30441 (NY, Z). Rio de Janeiro: Rio de Janeiro, Jan 1914, Hoel 3 ). Rio Grande do

Sul: Jari, estrada do Caracuru, Jari, estrada do Caracuru, campo alagado, 09 Aug 1969, Silva 2636 (NY). Roraima: borrow pit close to

road from Furo do Maraca to SEMA research station, 10 Mar 1987, Edwards 2529 (K). BOLIVIA: Beni: Ballivian, la zona de influ

del rio Yacuma, 09 Mar 1980, Beck 3248 (NY), Ballivian, espiritu en la zona de influencia del rio Yacuma, 13 Apr 1980, Beck 3354 (NY).

AFRICA. SOUTH AFRICA: KwaZulu-Natal: Hlabisa, St. Lucia, E shores, 30 Nov 1959, Feely & Ward 15 (K, M, NU-2 sheets]), Near

Howick, 1990, Taylor 131 (NU), Ingivauvima distr., near Salumhlanga, Ndumu Game Reserve, 22 Dec 1972, Pooley 1624 (NU), Greater

Durban area, Mlazi Valley, 15 Mar 1992, Ward 11925 (NU, PRE), Greater Durban area, Mlazi Valley, 15 Mar 1992, Ward 11926 (NU, NH

photo), Transvaal, Waterberg, 13.2 m NW of Warmbaths, 19 Mar 1965, Acocks 23562 (K, PRE), near Maputa, D near Nyinyani, 29

Oct 1980, Cunningham s.n. (NU), Karkloof floodplain near junction with Kusane River, Jan 1977, Kotze s.n. (NU). TANZANIA: Singida

Dist.: T. 5, M. 12.7 from Issuna e Singida-Manyoni Road, 4,800 ft, 13 Mar 1964, Greenway & Polhill 11543 mum T4, Sumbawanga

Dist., goli Mbuga, 5 km S ga, 19 Jun 1996, Faden et al. 96/302 (US), T5, Manyoni Dist., Chaya Lake, S of Itigi-Tabora track,

16 km W of Kazikazi, 02 Jul 1996, Faden et al. 96/522 (US). ZIMBABWE: Gokwe, Sengwa Nature Resreve, Jan 1966, Jacobsen 73 (NU),

District Gokwe, Sengwa research station, 09 May 1966, Jacobsen 3218 (PRE).

Distribution.—In the U.S.A. known only from Lee County, Florida. In México known from the states of

Campeche, Chiapas, and Guerrero, and in Central America known only from Panama. In the Caribbean

Basin known only from Cuba and the Dominican Republic. Most widespread in South America with re-

cords from Bolivia, Brazil, Colombia, Ecuador, Guyana, and Venezuela. In Africa known from South Africa,

Tanzania, and Zimbabwe.

Habitat.—Disturbed and natural freshwater wetlands including marshy open grasslands and tropical

forests; reported from 0-1400 m

Etymology.—The subspecific epithet is indicative of the short perianth bristles of this taxon.

3. Eleocharis acutangula (Roxb.) Schult. subsp. neotropica D.J. Rosen, subsp. nov. (Fig. 3 i-k). Tee: PERU.

DEPARTAMENTO DE LoRETO: Maynas, Iquitos, prolongacion Yavari, Versailles-Paina, open annually burned grassland, 23 Mar 1974,

McDaniel & Rimachi 18552 (HOLOTYPE: MO!; isotypes: IBE-2 sheets [photos!], NY!

A Eleocharis acutangula (Roxb.) Schult. subsp. acutangula perianthii setis brevioribus mollibus flexuosis longissimis 1.8—2.4-plo lon-

gitudo achenii, achenii apicibus constrictis infirme usque ad 0.6—0.7-plo latitudem achenii et stylopodiis plerumque latioribus quam

dit.

longioribus rece

Plants perennial. Roots coarse, fibrous, drab-brown; primary rhizomes caudex-like, thick, hard, ascending,

concealed by roots and persistent culm bases; secondary rhizomes elongated, to 3.2 mm thick, scales to 14

mm (few seen). Culms triquetrous, (44-)56-86-106) cm tall x (2—)2.7-3.9(-4.3) mm wide, soft, internally

spongy, with incomplete transverse septa, smooth, green to drab gray-green and finely longitudinally stri-

ate when dry. Leaves 2, reduced to sheaths, apically oblique, membranous, loose, friable, proximally dark

maroon, distally drab, apex acute. Spikelets cylindric, (1.5—)2—3.1(23.5) cm long x (3823.2-4.2(-4.5) mm

wide, acute; proximal scale with flower, obtuse, clasping, appearing as a continuation of culm; remaining

floral scales conspicuously spirally arranged, appressed, ovate-widely ovate, (3.9-)4.1-4.8(25) mm long x

(2.3-)2.4-3.2(-3.8) mm wide, cartilaginous, stramineous (faintly greenish centrally), adaxially sparsely

red-maculate, apex acute (rounded), the distal 0.2-0.3 mm translucent hyaline-erose, central area nearly

flat, abaxially coarsely many veined, the veins raised and visible at 20x, only mid-vein distinguishable in

adaxial view. Flowers with (5-)6—7 perianth bristles, bristles sub-equal, (1.8—)1.9-2.3(-2.4) times achene

Rosen et al., T f Eleochari t | 887

length, coarsely retrorsely spinulose nearly to base, stramineous, sometimes becoming reddish-brown

distally; stamens 3; anthers 0.9-1.8(-2.3) mm long, stramineous; style trifid. Achenes biconvex, broadly

obovoid, (1.3-)1.6-1.8 mm long x (1.2-)1.3-1.5(-1.6) mm wide, with 12-14-16) longitudinal rows of

deeply concave transversely oblong polygonal cells visible through transparent periclinal layer on each

achene face, dull to shiny yellow-green (sometimes tinged with amber), apex constricted to a distinct neck

about 0.6—0.7 times achene width. Tubercle dorsoventrally compressed, wider than tall and appearing very

shallowly to shallowly triangular, sometimes apex appearing truncate or retuse, (0.3-)0.4-0.5(-0.6) mm

long x (0.620.8-1C-1.1) mm wide, light-dark brown.

Specimen mined: SOUTH AMERICA. ECUADOR: Pastaza: Villano, Compamento Base de Arco, Pantano al noreste de la pista,

jo 29'S, 77? 27’ W, Feb 1994, Palacios 12171 (CIIDIR, MO). Napo: Archidona, Reserva Ecologica Antisana, Comunidad Shamato, En-

trada por km 21, Shamato, 00? 43'S, 077? 49'W, 24 Apr 1998, Clark et al. 5122 (MO). PERU: Amazonas: Bagua, along roadside from

Chiriaco to Puente Venezuela (3.9 Km NE Chiriaco), elev 600—800 ft, 31 Oct1978, Barbour 4355 (F, IBE-photo, MO). Cusco: Paucar

Tawbo, Montaueza Choutachaca, 780-1000 m, 28 Nov 1965, Vargas 16887 (US). Loreto: Puerto Almendras on the Rio Nanay, 30 km

N of Iquitos, 600 m, 16 Aug 1981, Moore & Ruiz 114 (F); Prov. E ae de 80 cm, 03 Aug 1967, Torres 340 (GH-2 sheets); Prov.

Maynas, Iquitos, Carretera de Zungaro Cocha, cerca a la q 12 Aug 1983, Ri I 08 (IBE-photo, VSC); Maynas,

Dtto. Punchana, Rio 22 varadera de; caserio de Padre Cocha, 07 Jul 1994, Rimachi 11004 (IBE—photo, MICH-2 sheets, NY, VSC);

Maynas, Inmedia de la Guarnicion militar de Gueppi, sobre la margen izquierda del Rio Putumayo, borde con Ecuador, 26 May

1978, Diaz 368 (F, MO); Maynas, Distrito Iquitos, Caserio Nina Rumy, Rio Nanay, 73° 25’ W, 03° 48’ S, 22 Apr 1988, Ruiz 1262 (MO);

Iquitos Region, 26 Jul 1966, Martin & Lau-Cam 1164 (GH). Huanuco: Leoncio Prado, 3 km SE of Pucayacu, on road from Tingo Maria

to Tocache Nuevo, ca 75 km NW of Tingo Maria, 10 Dec 1981, Plowman & Rury 11288A (MO, NY). BOLIVIA: Pando: Provincia Nicolas

Suarez, Cobija 2 km hacia el Sur, 19 Oct 1988, Beck 17139 (K, US).

Distribution.—Known only from northwest South America from Ecuador, Bolivia, and Peru.

Habitat —Specimens examined are from various freshwater wetlands including marshy open grasslands,

tropical forests, and roadside ditches, reportedly from 0-1000 m.

Etymology.—The specific epithet indicates the decidedly neotropical distribution of this taxon.

ACKNOWLEDGMENTS

We are grateful to the staff of all of the herbaria cited for loan of specimens for study, assistance with questions

about the disposition of type specimens and literature, and assistance with nomenclatural questions. The

first author is grateful to Socorro González-Elizondo for sharing her expertise and for facilitating field work

in Mexico and Eric Roalson for bringing certain literature items to his attention. The helpful comments of J.

Wipff and two anonymous reviewers are gratefully acknowledged. We offer special thanks to Neva Mikulicz

for preparing the excellent illustration. Financial support for the first author was provided by the Frank

W. Gould Award for Graduate Student Research in Plant Systematics and the Dr. Harry Wayne Springfield

Graduate Student Endowment. Financial support for field work in Florida by R. Carter was provided by the

Valdosta State University Foundation.

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A. Morrison (eds.), Monocots: systematics and evolution. CSIRO Publishing, Australia. Pp. 637-643.

GORDON-GRAY, K.D. 1995. Cyperaceae in Natal. Strelitia 2. National Botanical Institute, Pretoria, South Africa.

GREGOR, T. 2003. Eleocharis mamillata - Distribution and infraspecific differentiation. Folia Geobot. 38:49-64.

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Hines, D.M. 1975. A monograph of the Eleocharis ovata complex (Cyperaceae) in North America. Ph.D. Disserta-

tion. University of Michigan, Ann Arbor.

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16:155-157.

FOURTEEN NEW COMBINATIONS IN SEDUM (CRASSULACEAE)

Hideaki Ohba

Department of Botany

Hongo 7-3-1, Tokyo 113-0033, JAPAN

ohbagum.u-tokyo.ac.jp

ABSTRACT

hi : ; lin Sedum L. for tl f, 1 i f+} in Flora of North A th of M.

s. Seins S. Watson var. ciliosuni (Howell) H. Ohba, Sradiatum var. denau nessun (R.T. Clausen) H. Ohba, S. wrightii A. Gray

var. priscum (R.T. Clausen) H. Ohba, S a var. dencia: rum (R.T. Clausen) H. Ohba, S. obtusatum A. Gray var. boreale RI

Clausen) H. Ohba, S. obtusatum var. paradisum (Denton) H. Ohba, S. obtusatum var. retusum (Rose) H. Ohba, S

A. Berger var. eastwoodiae (Britton) H. Ohba, S. laxum var. flavidum (Denton) H. Ohba, S. laxum var. heckneri (M. Peck) H. Ohba,

S. laxum var. latifolium (R.T. Clausen) H. Ohba, S. stenopetalum Pursh var. monanthum (Suksd.) H. Ohba, S. lanceolatum Torr.

var. subalpinum (Fröd.) H. Ohba, and S. oreganum Nutt. var. tenue (R.T. Clausen) H. Ohba

RESUMEN

Se necesitan catorce combinaciones nuevas en Sedum L. en el tratamiento del género para la Flora of North America north of Mexico: S.

radiatum S. Watson var. ciliosum (Howell) H. Ohba, S. radiatum var. depauperatum (R.T. Clausen) H. Ohba, S. wrightii A. Gray

var. priscum (R.T. Clausen) H. Ohba, S. wrightii var. densiflorum (R.T. Clausen) H. Ohba, S. obtusatum A. Gray var. boreale RI

Clausen) H. Ohba, S. obtusatum var. paradisum (Denton) H. Ohba, S. obtusatum var. retusum (Rose) H. Ohba, S

A. Berger var. eastwoodiae (Britton) H. Ohba, S. laxum var. flavidum (Denton) H. Ohba, S. laxum var. heckneri (M. Peck) H. Ohba,

S. laxum var. latifolium (R.T. Clausen) H. Ohba, S. stenopetalum Pursh var. monanthum (Suksd.) H. Ohba, S. lanceolatum Torr.

var. subalpinum (Fröd.) H. Ohba, y S. oreganum Nutt. var. tenue (R.T. Clausen) H. Ohba.

The following new combinations are required for the forthcoming treatment of Sedum (Crassulaceae) in

volume 8 of Flora of North America north of Mexico.

Sedum radiatum S. Watson var. ciliosum (Howell) H. Ohba, comb. & stat. nov. Basionyu: Sedum ciliosum Howell,

Fl. N.W. Amer. 214. 1898

Sedum radiatum S. Watson var. depauperatum (R.T. Clausen) H. vue comb. & stat. nov. Basionym: Sedum

radiatum subsp. depauperatum R.T. Clausen, Sedum N. Amer. 258, figs. 59-60.

Sedum wrightii A. Gray var. priscum (R.T. Clausen) H. Ohba, comb. & stat. nov. Basioxvw: Sedum wrightii subsp.

priscum R.T. Clausen, Bull. Torrey Bot. Club 106:215. 1979

Sedum wrightii A. Gray var. densiflorum (R.T. Clausen) H. Ohba, comb. & stat. nov. Basionym: Sedum wrightii

subsp. densiflorum R.T. Clausen, Variat. Sp. Sedum Mex. Cordill. Plateau 13. 1981.

Sedum obtusatum A. Gray var. boreale (R.T. Clausen) H. Ohba, comb. & stat. nov. Basowvw: Sedum obtusatum

subsp. boreale R.T. Clausen, Bull. Torrey Bot. Club 69:32. 1942

Sedum obtusatum A. Gray var. paradisum (Denton) H. Ohba, comb. & stat. nov. Basionyu: Sedum obtusatum subsp.

paradisum Denton, Brittonia 30:256, fig. 2. 1978.

Sedum obtusatum A. Gray var. retusum (Rose) H. Ohba, comb. & stat. nov. Basionvw: Gormania retusa Rose in Britton

& Rose, New N. Amer. Crassul. 31. 1903

Sedum laxum (Britton) A. Berger var. eastwoodiae (Britton) H. Ohba, comb. & stat. nov. Basioxvm: Gormania

eastwoodiae Britton in Britton & Rose, New N. Amer. Crassul. 903.

Sedum laxum (Britton) A. Berger var. flavidum (Denton) H. Ohba, comb. & stat. nov. Basionym: Sedum laxum

subsp. flavidum Denton, Brittonia 30:233, fig. 1. 1978

Sedum laxum (Britton) A. Berger var. heckneri (M. Peck) H. Ohba, comb. & stat. nov. Basionvm: Sedum heckneri

M. Peck, Proc. Biol. Soc. Wash. 50:121. 1937

J. Bot. Res. Inst. Texas 1(2): 889 — 890. 2007

890 Journal of the Botanical R h Institute of Texas 1(2)

Sedum laxum (Britton) A. Berger var. latifolium (R.T. Clausen) H. Ohba, comb. é stat. nov. Basionvm: Sedum

laxum subsp. latifolium R.T. Clausen, Bull. Torrey Bot. Club 69:38. 1942.

Sedum stenopetalum Pursh var. monanthum (Suksd.) H. Ohba, comb. & stat. nov. Basionym: Sedum monanthum

Suksd., Werdenda 1:19. 1927, based on Sedum uniflorum Howell, Fl. N.W. Amer. 213. 1898 not Sedum uniflorum Hook. & Arn

Bot. Beechey Voy. 263. 1838.

Sedum lanceolatum Torr. var. subalpinum (Fród.) H. Ohba, comb. & stat. nov. Basionvm: Sedum stenopetalum Pursh

var. subalpinum Fród., Acta Horti Gothob. 10(App.):70, figs. 521-527, pl. 43, fig. 2. 1935 [1936]

Sedum oreganum Nutt. var. tenue (R.T. Clausen) H. Ohba, comb. & stat. nov. Basionvu: Sedum oreganum subsp.

tenue R.T. Clausen, Sedum N. Amer. 355, fig. 91. 1975.

ACKNOWLEDGMENTS

Ithank James Zarucchi, Craig Freeman, and Nancy Morin (Flora of North America) for assistance with get-

ting these new combinations published. This study was supported by a Grants-in-aid for Scientific Research

from the Japan Society for Promotion of Science (Nos. 17570073 and 18255004).

A NEW GYPSOPHILOUS SPECIES OF ERIGERON (ASTERACEAE: ASTEREAE)

FROM NORTHEASTERN MEXICO

Guy L. Nesom

Botanical Research Institute of Texas

509 Pecan Street

Fort Worth, Texas 76102-4060, U.S.A.

ABSTRACT

Erigeron heleniae Nesom, sp. nov., is described from a gypsum outcrop northwest of Monterrey in Nuevo Leon, Mexico. It is char-

acterized by its diminutive, suffruticose habit, thickened taproot, hirsute-glandular vestiture, linear and subclasping leaves, and tiny,

solitary heads with white, filiform rays

RESUMEN

Exigeron ADR. Nesom, T nov., se describe de un o das en el noroeste » MO en Nuevo León, México. Se

I raiz engrosada, i glandular, hoj subapretadas, y cabezuelas

ST aoa Js ; x 121 PL

Gypsum outcrops in northeastern Mexico have fostered the evolution of many endemic species. Yet another

is described here, known only from a single collection by George S. Hinton, whose botanical acuity and

persistence (along with that of his father, Jaime Hinton) have greatly expanded knowledge of the region's

plants.

Erigeron heleniae Nesom, sp. nov. (Figs. 1-2). Tree: MEXICO. Nuevo Leon: Mpio. Mina: N of Los Molina, 2690409.23" N,

100?4427.57" W, gypsum hillside, [open gypsum exposures, flats and gentle slopes, with scattered ocotillo and other low perenni-

als], 737 m elev, 4 Sep 2005, George S. Hinton 28412 (HoLotypE: TEX).

Erigeronti gypsovero N ilis sed diff li lari li itulis minoril llis radii brevioribus filiformibus, foliis sub-

amplectentibus, vestimento hirsuto-glanduloso, et pappo externo squamarum prominentium.

mE MT

Suffruticose[ athickened, woody taproot, individual plants up to 25 cm wide; stems intricately

branched proximally, 5-8 cm high, hirsute to hirsute-villous, hairs strongly deflexed on proximal third,

spreading above; leaves, and phyllaries hirsute with white, erect-arching hairs 0.4-1 mm long, hairs arising

from thickened, orange-resinous basal cells, minutely stipitate-glandular, the phyllaries densely so. Leaves

linear to linear-oblanceolate, flat, 7-10(-15) mm long, 0.8-1 mm wide, even-sized to immediately below the

heads, slightly but distinctly auriculate and subclasping. Heads solitary on bracteate peduncles 3-7 mm long;

involucres turbinate-campanulate, 3.5-5 mm wide; phyllaries in 3(-4) subequal series, elliptic-lanceolate,

2.53 mm long, with a dark orange-resinous midvein, densely minutely glandular. Ray florets 22-30, white

or slightly pink-tinged, 3-3.5 mm long, ligules 0.2-0.4 mm wide, apparently not reflexing or coiling. Disc

florets 1-1.6 mm long. Achenes obovate-oblong, 0.7 mm long, 2-nerved, sparsely strigose; pappus of 10-12

persistent bristles 0.8-1 mm long, outer pappus of prominent scales 0.1 mm long.

Etymology.—Erigeron heleniae is named for Helen Hinton, mother of George S. Hinton and wife of Jaime

Hinton. She has been close witness to the excitement of discovery by her family of so many of the region's

beautiful and unusual species.

The new species occurs on gypsum outcrops about 55 kilometers northwest of the city of Monterrey.

There are two large gypsum outcrops in the immediate area, about 5.5 kilometers apart and covering a total

of about 3.5 hectares. Essentially the same set of gypsophilic species occurs on both outcrops; Erigeron

heleniae is common on both and prefers slopes to flat areas. As with many other gypsophilic endemics from

this region of Mexico, this area of outcrops probably is the total extent of the range of the new species.

Erigeron heleniae is similar in its diminutive habit and narrow leaves to E. gypsoverus Nesom, which is

J Bot. Res Inst Texas 1(2): 201 — 894. 2007

892

A A

Journal of

p

MOST A

\ SAN \ N TN N) |

ESA

Xx (Sd A \ (t N

NAAA

j N

4 1

l \ M A Y

Nesom, À new gypsophilous species of Erigeron 893

894 Journal of the Botanical R h Institut

f Texas 1(2)

relatively common in the open pine woods in the gypseous region around Galeana, Nuevo León, and known

from numerous collections. Erigeron heleniae occurs about 140 kilometers north of the closest population of

E. gypsoverus and at a lower elevation. Both species apparently are obligate gypsophiles, but the two probably

are not even closely related within the genus. Contrasts below distinguish them.

1. Stems, leaves, and phyllaries closely strigillose, a taproot slender; proximal branches elongate,

stolon-like; leaves not at all basally auriculate or claspir ES involucres 5-7 mm wide; ray florets 5-6 mm

outer ofin nconspici Ious setae; 1050- n m ele

=

=

Erigeron gypsoverus

. Stems, pum and phyllaries hirsute, minutely m CERTES taproot thickened; proximal branches

intricately branched; leaves slightly but distinctly auriculate and subclasping; involucres 3.5-5 mm wide; ra

florets 3.5-5 mm long; outer pappus of prominent scales; ca. 750 m elev.

ray

rigeron heleniae

Morphological features shared by Erigeron heleniae and E. dryophyllus A. Gray (of central Nuevo León and

adjacent Tamaulipas) suggest that these two species may be closely related: subclasping leaves and hispid-

villous vestiture, the hairs sharply deflexed on proximal portions of the stem. Erigeron dryophyllus differs

most conspicuously in its rhizomatous habit, broad and lobed leaves, much larger heads on long peduncles,

and habitats primarily over limestone.

e

ACKNOWLEDGMENTS

George Hinton provided details regarding the locality and habitat of the new species as well as a review of

the manuscript. Tim Lowrey also reviewed the manuscript. The illustration is by Linny Heagy.

FOLIA TAXONOMICA 3. PASSIFLORA DAVIDII (PASSIFLORACEAE),

ANEW SPECIES IN SUBGENUS PASSIFLORAANDA KEY TO THE

SECTIONS OF SUPERSECTION SHIPULATA

Christian Feuillet

Department of Botany, MRC-166

Smiths

sonian Institution

PO. Box 3712

Washington, D.C. 20013-7012, U.S.A.

feuillec@si.edu

ABSTRACT

E

The first key t

Stipulata Feuillet =e M. dida is Phone Dt davidii Pn nov. is described from

French Guiana. Its foliaceous sup and b

I I g I Stipulata

section Granadillastrum Triana & Planch.

RÉSUMÉ

La premiere clé d de la supersection Stipulata Feuillet Bu M. cud UE est fournie. rod aii iu nov. est décrite

de Guyane francaise. En raison d tipul 1 | p le sous-genre

Passifl I ion Stipulat ti dillastrum Triana & Planch.

In the new classification of Passiflora (Feuillet & MacDougal 2003; MacDougal & Feuillet 2004) replacing

the system proposed by Killip (1938), Passiflora subg. Passiflora supersect. Stipulata Feuillet & J.M. MacDou-

gal is a large coherent group which includes more than 95 species sharing foliaceous, usually asymmetric

stipules.

KEY TO THE SECTIONS OF SUPERSECTION STIPULATA

. Flowers in pseudoracemose inflorescences, 25-75 cm long. (P racemosa Brot). sect. Calopathanthus Harms

: Flowers solitary or in pair in leaf axils.

2. Bracts setaceous or narrowly ovate, less than 2.5 mm wide, scattered along the pedicel. (4 species)

ermesinae (Cervi) Feuillet & J.M. MacDougal

2. Bracts foliaceous, usually more than 2.5 mm wide (but often deeply dissected in sect. Dysosmia), usually

verticillate, sometimes scattered or loosely grouped

3. Stipules and bracts lacerate-dentate or laciniate to 1-4-pinnatisect. (20 sp.) sect. Dysosmia DC.

3. Stipules and bracts entire to serrate.

4. Hypanthium cylindric, as long as the sepals. (4 sp.) sect. Tacsonioides DC.

4. Hypanthium campanulate or rarely short-tubular and then shorter than the sepals. (> 66 sp.)

sect. Granadillastrum Triana & Planch.

The members of Passiflora sect. Granadillastrum are herbaceous or woody climbers, with leaves petiolate

and with two or more glands on the petiole; the laminas are unlobed or lobed; the venation palmate or

sometimes pinnate with 1—2 pairs of minor submarginal veins at base. Their flowers are solitary or rarely

two in the leaf axils. The hypanthium is campanulate or rarely short-tubular. Triana and Planchon (1873)

described Passiflora section Granadillastrum for one species, P. semiciliosa Planch. & Linden published in

the same paper. In Killip (1938), Granadillastrum was raised to the rank of subgenus and some species were

added that all belong now in supersect. Tacsonia. In addition to P. semiciliosa, we place in section Granadil-

lastrum species formerly placed in subgenus Passiflora (subg. Granadilla of Killip) ser. Kermesinae Killip ex

Cervi (in part), Imbricatae Killip ex Cervi, Simplicifoliae (Harms) Killip ex Cervi, Lobatae Killip ex Cervi, and

Menispermifoliae Killip ex Cervi. Those series were not described by Killip. His key to the series suggested

a strong emphasis placed on leaf shape, separating species with lobed leaves and those with unlobed leaves

J Bot. Res Inst Texas 1(2): 205 — 898. 2007

896 Journal of the Botanical R h Institute of Texas 1(2)

into informal series. Since then the series were validated by Cervi (1997), but several species proved to be

variable in leaf shape. Although the species in this group should be separated in subcategories, at this point

itis premature to propose formal series.

It is in 1987 that I saw the new species, P. davidii, for the first time. The young plant had two stems

climbing on a rope in the camp at Montagne des Nouragues in French Guiana (Feuillet 4395). Its stipules

with an egg-mimic yellow swelling at the apex and the unlobed leaves with several yellow swellings on the

margin set it apart from the other species of subgenus Passiflora. One stem had young flower buds, the other

was sterile. I saw this species again twice, in French Guiana, with David Rignon on Montagnes Tortue and

near Cacao, but without flower or fruit. No specimens were collected. After nearly 20 years, David collected it

in bloom in February 2006, confirming it was a new species and providing enough information for a formal

description.

Passiflora davidii Feuillet, sp. nov. (Fig. 1). Tv: FRENCH GUIANA: cultivated 1 km from the village of Cacao, along the road

to Cayenne, 0-50 m, 52°25'W, 4°33'N, 23 Feb 2006, D. Rignon 16 (HoLoTYPE: US; isotypes: CAY, P).

4n

Passiflora davidii i be. Passifl Stipulata sect. Granadillastro per

H TESTIS EDS 1 ex

tinens. Passiflorae actiniae Hook. affinis; stipulis cum mu-

LE Al 1 1 PE E 1 $

5 E

I ZI ec pcc

[e] jT

o

Į E

dictincta. A specieb i il ipulis foliacei j d pice inflatum differt.

S F de

Vine, size unknown, climbing with tendrils, short-pubescent throughout except where noted; stems terete,

in some places striate. Stipules foliaceous, 1.8-2 x 1 cm, half-cordate at base, rounded and short mucronate

at apex, the awn or mucro 1.5-2 mm long, blunt, with the apical half to 2/3 swollen and yellow, margin

entire, paler abaxially when fresh, midrib slightly to clearly excentric, but straight, other veins reticulate.

Leaves: petiole 3.8-4.2 cm long, 6-9-glandular; glands sessile, swollen, orange-yellow, spread along the

petiole with 2 in subapical position; blade simple, unlobed, ovate, 10 x 6.5-7 cm, widely rounded, slightly

peltate at base, acute to right angled at apex, margin entire, briefly recurved, narrowly glandular with scat-

tered yellow small swellings, glabrous adaxially, paler abaxially when fresh, venation palmate at the very

base with 5-7 veins, forming a submarginal wavy vein, apical half pinnate, with 6-7 main lateral veins

ascending and toward the margin also included in the formation of a submarginal vein, minor lateral and

most secondary veins more or less perpendicular to the midrib, tertiary veins reticulate. Flowers axillary,

one per leaf axil, pedicel 272.8 cm long below the bracts, stipe 5-6 mm between the bracts and the flower;

bracts 3, one larger, verticillate, green, lanceolate, 2-3 x 1-1.3 cm, rounded, slightly convex even around the

bud, briefly cuneate at base, acute to narrowly-rounded at apex with a short swollen mucro less than 0.01 x

0.01 cm, paler abaxially when fresh; hypanthium campanulate, about 0.3 x 0.4 cm deep, sepals white and

glabrous adaxially, yellow-green abaxially in parts exposed in bud, narrowly oblong, 3 x 1.5 cm, with apical

awn 1-2 mm long, petals white, narrowly oblong, 3.5 x 1-1.3 cm, glabrous, corona with micro-trichomes

on the outer filaments, outer filaments spreading to cover the perianth, 2-3.5 cm long in the same flower,

white in their proximal half, blue violet in their apical half, thin, numerous, in 2-3 not clearly separated

rows, inner filaments 0.3-0.4 mm, white, about as thick as the previous ones, in several rows, operculum

membranous, limen membranous, androgynophore 1.2 cm long, glabrous, stamens filaments 0.8-1.0 cm

long, flat, green speckled with red, glabrous, anthers 0.7-0.8 x 0.25-0.30 cm, yellow, glabrous, gynophore

ca. 0.1 cm long, ovary obovoid, 0.4 x 0.25-0.27 cm, styles green speckled with red, glabrous, wider toward

apex, stigmas hemispherical, pale yellow, glabrous. Fruit unknown.

The description is based on the holotype that has two nodes, each with a leaf and a flower. Therefore

the variation in measurement is the variation between 2 consecutive nodes. The sterile specimen (Feuillet

4395) consists in the apex of a stem actively growing, the stipules 1.5-2.5 x 0.5-1 cm are already fully de-

veloped, some are even longer than on the holotype, but the petioles 0.4—1.5 cm, and the leaf blades 4.5-7

x 2.5—4 cm, are not fully grown. The early development of the stipules, displaying the egg-mimic mucros,

apparently acts as a deterrent for the females of the solitary egg-laying species of heliconiid butterflies, and

protects the apical meristem from grazing by their caterpillars.

In the key by Killip (1938), Passiflora davidii would be close to P. actinia, a species from southeastern

Feuillet, Passiflora davidii, ies in sul Passifl 897

Fic. 1. Passiflora davidii Feuillet. Top: Rignon 16 (photo M.-F. Prévost), Lower left:

Feuillet 4395 (photo C. Feuillet).

Brazil. It can be distinguished from P. actinia by the short pu-

bescence on all vegetative parts and the stipules only slightly

asymmetric with a mucro that is swollen, yellow, and act as an

egg-mimic. The specimen looks like P. loretensis in general ap-

pearance, but can be recognized by the 6—9 glandular petiole.

Contrary to P. actinia and P. loretensis, P. davidii has a pubescent

ovary. The flower is very similar to that of P. retipetala, that, in

les with a longer

contrast with P. davidii, is glabrous and has stipu

awn. In the Guianas, P. davidii can be separated from the other

members of supersection Stipulata with the characters used in

the key below.

The epithet of the new species honors David Rignon who

collected the type specimen. He lives in French Guiana near the

village of Cacao and is interested in passionflowers and butter-

flies. David and his family were great field companions in 2003,

when we saw P. davidii without flowers near the type locality.

lairiére du C lesN

52°42'W, 4°03'N, 100-120

a

qu SES

o

m, 28 Aug 1987, st., Feuillet 4395 (CAY).

898 Journal of the Botanical R IRI

f Texas 1(2)

KEY TO THE SPECIES OF SUPERSECTION STIPULATA IN THE GUIANAS & VENEZUELAN GUAYANA

1. Stipules laciniate, cleft to base, not deciduous. (sect. Dysosmia).

2. Ovary and fruit glabrous.

3. Stem glabrous; fruit red etida var. orinocensis (Killip) Feuillet

3. Stem hispid; fruit yellow orange P. foetida var. hispida (DC. ex Triana & Planch.) Killip

2. Ovary and fruit indumentum scarce to dense.

4. Stem stiffly hirsute; ovary and fruit hirsute

4. Stem softly pilosulous

5. Bracts 2-3-pinnatisect, 2-3 cm long

5. Bracts 1(-2)-pinnatisect, less than 2 cm long

1. Stipules entire or serrulate. (sect. Granadillastrum).

6. Stipules acute at apex

6. Stipules obtuse to rounded at a

y: T with an apical awn or arista more than 5 mm long.

8. Stipule awn swollen at tip

8. Stipule arista not swollen at ti

7. Stipules with an apical arista or mucro less than 5 mm long.

9. Leaf blade reddish beneath; bracts petiolate

9. Leaf blade pale green or glaucous beneath; y bracts sessile

10. Leaves 3-lobed, red- glandular at ma P. garckei Mast.

10. Leaves unlobed, margin glandular yellow swellings P. davidii Feuillet

P. foetida L. var. foetida

P. foetida var. dip in (Ham.) Mast.

P. moritziana Planch.

P. exura Feuillet

P. retipetala Mast.

P. stipulata Aubl.

P. picturata Ker

ACKNOWLEDGMENTS

I am indebted to my friend and colleague Fanchon Prévost (CAY) for processing the plant collected by David

Rignon into herbarium specimens and for the detailed labels and great photographs. I would like to thank

the curators and directors of the herbaria CAY, NY, P, U, US whose collections have been made available

for my work when writing this paper. This is number 128 in the Smithsonian's Biological Diversity of the

Guiana Shield Program publication series. John MacDougal and Peter M. Jorgensen provided critical reviews

for which Iam most appreciative.

REFERENCES

Cervi, A.C. 1997. Passifloraceae do Brasil. Estudo do género Passiflora L., subgénero Passiflora. Fontqueria

FeuiLLET, C. and J.M. MacDoucaL. 2003 [May 2004]. A new infrageneric classification of Passiflora L. (Passifloraceae).

Passiflora 13(2):34-38.

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

MacDouca,, J.M. and C. FeuiLLET. 2004. 2. Systematics. In: T. Ulmer and J.M. MacDougal, eds. Passiflora, Passionflow-

ers of the world. Timber Press, Portland. Pp. 27-31.

TRIANA, J.J. and J.E. PLANCHON. 1873. Prodromus florae Novo-Granatensis. XIX Passifloreae. Ann. Sci. Nat., Bot., sér

5, 17:121-186.

—

FOUR NEW COMBINATIONS IN MOSIERA (MYRTACEAE) FROM THE CARIBBEAN

Andrew Salywon

Desert Botanical Garden

1201 N. Galvin Parkway

Phoenix, Arizona 85008-3437, U.S.A.

asalywon@dbg.org

ABSTRACT

FM L

New combinations for four C ly assigned to Eugenia L. or Psidium L. are proposed: Mosiera andro-

i d for Mocier d i

siana, M. cuspidata, M. gracilipes, and M xerophytica. A |

de

RESUMEN

ee : : de TENA i i d Eugenia | o Psidium L.: Mosiera

a M. cuspidata, M. ales yM. xerophytica. Se PE un lectotipo para Moe androsiana.

Recent taxonomic and molecular studies of Mosiera (Landrum 1992; Salywon 2003; Salywon et al. 2004;

Salywon & Landrum in press) have clarified the boundaries of the genus. However there are no morpho-

logical synapomorphic characters that differentiate Mosiera from related genera. It is therefore separated

from other genera by a combination of characters including: tetramerous flowers, lustrous or glandular

seed coats one to six cells thick, and C-shaped embryos with cotyledons that are ca. 4 the length and

about the same width as the hypocotyls. Landrum and Kawasaki (1997) and Salywon and Landrum (in

press) provide keys to distinguish Mosiera from similar genera. The genus contains approximately 18

species distributed mainly in the Caribbean, with two species in Mexico and one of these in Guatemala.

New combinations are required for four Caribbean species previously placed in Eugenia or Psidium that

can be assigned to Mosiera, in order to make the names available for a manuscript in preparation on the

molecular systematics of the family (Salywon et al. in prep.).

Mosiera androsiana (Urban) Salywon, comb. nov. Basiosvw: Eugeni iana Urban, Fedde, Rep. Sp. Nov. 13:467. 1915.

Psidium androsianum (Urban) Correll, J. Arnold Arbor. 58:41. 1977. Tree: BAHAMAS: Andros, Mangrove Cay, coppice, near Lisbon

Creek, 16-19 Jan 1910, Small & Carter 8496 (LeEcTOTYPE, selected here: NY [vascular plant type image library # 84,503]; IsoLECTOTYPES:

E US [2 sheets]).

The holotype in Berlin is lost, and the NY specimen is here chosen as the lectotype. Mosiera androsiana is

morphologically most similar to M. longipes (O. Berg) Small but differs in having leaves 0.5-2.1 x 0.15-1

cm (vs. 1.1-5.2 x 0.2-3.8 cm) with the secondary venation not visible (vs. secondary venation visible),

and seeds ca. 3 mm long (vs. ca. 2 mm long). The leaf morphology of this species is among the most

variable in the genus.

Mosiera cuspidata (Alain) Salywon, comb. nov. B Psidi pidatum Alain, Brittonia 20:159. 1968. Tyee: HISPANOLA

[DOMINICAN REPUBLIC]: Boca del Infierno, Los Haitises. Samaná Prov., on limestone cliffs, near the edge of water, 24 Jun 1930,

Ekman H. 15427 (HOLOTYPE: US; isotypes: MICH, S)

Mosiera cuspidata is a shrub or small tree to 4 m tall with distinctively large leaves for the genus. It is

ecologically interesting because it grows on vertical limestone cliffs and on small emergent limestone

outcrops in the ocean with little or no soil.

Mosiera depu id (Alain) Salywon, comb. nov. B Psidi ilipes Alain, Phytologia 25:269. 1973. Tye: DOMINICAN

BLIC. [Cordillera Central]: [Prov. La Vega, Municipio de eee), T Redonda, Ciénaga de la Culata, Constanza, in cloud

AA 1700-2000 m, 30 Nov 1969, Alain 17138 (HoLotYPE: NY; isotypes: JBSD, US).

J. Bot. Res. Inst. Texas 1(2): 899 — 900. 2007

900 Journal of the Botanical R h Institute of Texas 1(2)

This species is known only from three collections, all from the same locale, and may be of conservation

concern as clearcutting for farmland and wood has deeply encroached into the cloud forest where this

species grows. Additional fieldwork is needed to determine if it is distributed on other mountain tops

nearby.

Mosiera xerophytica (Britton) Salywon, comb. nov. Basionw: Ms ee ee Britton, Bull. Torrey Bot. Club 51:11. 1924.

b, bar

ERTO RICO: Cayo Muertos, limestone rocks, shrub, 2 m, g , 9-12 Mar 1915, Britton, Cowell &

Brown 4982 (HOLOTYPE: NY; IsoTYPE: US).

In the early 1990s, Myrtaceae specialist Bruce Holst (presently at Selby Botanical Gardens), while work-

ing at Missouri Botanical Garden, first recognized that this species was out of place in Eugenia and sug-

gested it might belonged in Psidium. He therefore brought it to the attention of Les Landrum, at Arizona

State University, who is conducting revisionary work on the genus. Subsequently, Landrum and Bonilla

(1996) in a study of anther glandularity in the American Myrtinae determined that this species should

be transferred to Mosiera, but they did not make the new combination. Instead they deferred doing so

until the genus could be worked on as a whole.

ACKNOWLEDGMENTS

I express my appreciation to Leslie Landrum for his valuable insights and support in my studies in Myrta-

ceae. I am very indebted to Frank Axelrod (UPRRP), and the staff at JBSD, especially Daisy Castillo de

Vásquez and Martín de la Cruz, who helped make my herbarium work and fieldwork possible. I thank

the curators of the following herbaria for kindly providing access to specimens for this study: ASU, BM,

F, FLAS, JBSD, MICH, MO, NY, S, UPRRP and US. The comments of Bruce Holst and Donald Pinkava

helped improve this paper. I gratefully acknowledge funding support from the Friends of the Center for

Latin American Studies at Arizona State University, American Society of Plant Taxonomists Graduate

Student Research Award, National Council of State Garden Clubs, Arizona Federation of Garden Clubs,

Sigma-Xi, and the Explorers Club.

REFERENCES

LANDRUM, L.R. 1992. Mosiera (Myrtaceae) in Mexico and Mesoamerica. Novon. 2:26-29.

LANDRUM, L.R. and J. Bonita. 1996. Anther glandularity in the American Myrtinae (Myrtaceae). Madroño 43:

508-68

LANDRUM, L.R. and M.L. Kawasaki. 1997. The genera of Myrtaceae in Brazil: and illustrated synoptic treatment and

identification keys. Brittonia 49:508-536.

SALYWON, A. 2003. A monograph of Mosiera (Myrtaceae). PhD. Dissertation, Arizona State University, Tempe,

U.S.A.

SALYWON, A., N. Snow, M.F. WoJciecHowski, J. CsizwADI, and L. Lanbrum. 2004. Phylogenetic relationships of Myrtaceae

as inferred from nrDNA ITS sequences. Botany 2004, abstract 38-5.

SALYWON, A. and L.R. Lanbrum. Curitiba (Myrtaceae): a new genus from the Planalto of southern Brazil. Brittonia

(in press).

NEW COMBINATIONS IN PHILADELPHUS (PHILADELPHACEAE)

James Henrickson

Plant Resources Center

University of Texas

Austin, Texas 78712-0471, U.S.A.

ABSTRACT

T] T ; d T pq ESTE Philadolnt H hell

A. Gray var. madrensis (Hemsley)

Henr eos comb. & stat. nov.; " Philadelphus ropas A. Gray var. TET cto) Henrickson, comb. & stat. nov.; and

Philadelphus microphyllus A. Gray var. stramineus (Rydb.) Henrickson, comb. & sta

RESUMEN

e ; Dita ida] TT DPhiladolnl : hell

A. Gray var. madrensis (Hems-

les) o comb. & stat. nov. “Philadelphus microphyllus A. Gray var. argyroealya (V Weston) Henrickson, comb. & stat. nov.,

y Philadelphus microphyllus A. Gray var. stramineus (Rydb.) Henrickson, comb. & stat. n

Preparation of a treatment of Philadelphus for Flora North America Vol. 8 necessitates the following new

combinations.

Philadelphus microphyllus A. Gray var. madrensis (Hemsley) Henrickson, comb. & stat. nov. Basowvw:

Philadelphus madrensis Hemsley, Kew Misc. Inform. Bull. 1908:251. 1908. Type: MEXICO. Duranco: Sierra Madre, B.C. Seemann

2167 (HOLOTYPE: K!).

Philadelphus microphyllus A. Gray var. argyrocalyx (Wooton) Henrickson, comb. & stat. nov. Basionyw:

Philadelphus argyrocalyx Wooton, Bull. Torrey Bot. Club. 25:452. 1898. Tyre: U.S.A. New Mexico. Lincoln Co.: Eagle Creek, 14 Aug

1897, E.O. Wooton 524. (HOLOTYPE: US!).

Philadelphus microphyllus A. Gray var. stramineus (Rydb.) Henrickson, comb. & stat. nov. Basionym: Phila-

delphus stramineus Rydb., N. Amer. Fl. 22:172. 1905. Type: U.S.A. CALIFORNIA. Mono Co.: White Mts., Aug 1888, WH. Schockley s.n.

(HOLOTYPE: NY!; isotypes: A! F! US).

&

Justification of the new combinations will be published at a later time in this journal.

J. Bot. Res. Inst. Texas 1(2): 901. 2007

902 Journal of the Botanical R h Institute of Texas 1(2)

BOOK REVIEWS

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An Illustrated Manual (ed. 2). (ISBN 978-0-8122-4003-0, hbk.). Univ. of Pennsylvania Press, 3905

Spruce Street, Philadelphia, PA 19104-4011, U.S.A. (Orders: Univ. Penn. Press Warehouse, PO Box

5030, Baltimore, MD 21211, U.S.A.; custser@pobox.upenn.edu; 800-537-5487; 410-516-6998 fax).

Bae 1088 pp., 2,645 line d maps qe TOt

Co E [2] 1 s A 1 1 1 «1 Tahal xl Ttf iJi

In

and genera E oyed in the first edition with family Ads that best SUE our current desque of phylogenetic

relationships within the vascular plants. ... We drew Hes on m Flora of North America (published volumes and website) and the

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The text Plant S Phylogenetic Approach, Second Edition, by Judd et al. (2002) was also an important source." An exceptional

fo)

value for 1088 pages, m nearly 3400 species, and including numerous additions to the flora beyond the first edition of 2000.

Wu ZHENGYI, PETER H. Raven, and Howc Deyuan (eds.). 2007. Flora of China, Vol. 13. Clusiaceae through

Araliaceae. (ISBN-Vol. 13: 978-1-930723-59-7, hbk.). Science Press (Beijing) and Missouri Botanical

Garden Press (St. Louis), PO. Box 299, Saint Louis, MO 63166-0299, U.S.A. (Orders: www.mbgpress.

org, orders@mbgpress.org, 314-577-9547, 314-577-9594 fax). $125.00, 548 pp., 8 3/4" x 11 1/4".

This is “the 13'^ of a 25-volume work. It includes 33 families, 151 genera, and 1288 species, among which 11 genera and 649 species

(5096) are endemic to China, and three families, 14 genera, and 76 species are introduced to China." Alangiaceae, Ancistrocladaceae,

E

Ir

Araliaceae, Begoniaceae, Bixaceae, Cactaceae, Caricaceae, Cistaceae, Clusiaceae, Combretaceae, Crypteroniaceae, Cynomoriaceae,

Elaeagnaceae, Elatinaceae, Flacourtiaceae, Frankeniaceae, Haloragaceae, Hippuridaceae, Lecythidaceae, Lythraceae

Mere corneas Myrtaceae, Nyssaceae, Onagraceae, Passifloraceae, Rhizophoraceae, Stachyuraceae, Tamaricaceae, Tetramelaceae,

Thymelaeaceae, Trapaceae, Violaceae. Chinese and American authors collaborate.

WALTER S. JUDD, CHRISTOPHER S. CAMPBELL, ELIZABETH A. KELLOGG, PETER E STEVENS, and MICHAEL J. DONOGHUE. 2007.

Plant Systematics: A Phylogenetic Approach (ed. 3). (ISBN 978-0-87893-407-2, hbk.). Sinauer

Associates, Inc. 23 Plumtree Road Sunderland, MA 01375. (Orders: orders@sinauer.com, publish@

sinauer.com, www.sinauer.com). $94.95, 620 pp., ca. 325 illustrations, 25 color plates, 8 1/2" x 11".

This justifiably much-used book is noted to be "appropriate for any course e ] f angi vascular plants

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edition (beyond 2002 2"4 ed): “color throughout the book, including m more ae two dozen pl l variation

in the vee pa new ibaa f maximum likelihood and B ly revised treatments eo 17 families that

in ci ll cl d to the many cladograms, taking into account recent

taxonomic UR and M Án accompanying CD contains over 3,100 color photographs depicting a wide variety of plants

from over 185 families and over 1900 species. Many species are oda by pend Dn um showing different views of the

plant, its flowers, its fruits, and its habit. Also on s CD Gwith links to photos) and three

the text, according to HM APG, nas and mE

Cone: in Dil by chapters: The Science of Plant Systematics; Methods and Principles of Biological Systematics; Classification

and Systems in Flowering Plants-Historical Background; Taxonomic Evidence-Structural and Biochemical Characters; Molecular Sys-

tematics; The Evolution of Plant Diversity; An Overview of Green Plant Phylogeny; Lycophytes, Ferns, and Phylogenetic

Relationships of Angiosperms. Two appendices: Botanical Nomenclature; Specimen Preparation and deb aon.

J. Bot. Res. Inst. Texas 1(2): 902. 2007

FIVE NEW VARIETIES OF AGAPETES (ERICACEAE) FROM INDIA

D. Banik M. Sanjappa

Botanical Survey of India Botanical Survey of India

Central National Herbarium CGO Complex, 34 MSO Building

PO: Botanic Garden, Howrah Block E, 5% Floor, DF Block, Sector-1

71 a , INDIA Salt Lake City, Kolkata—700 064, INDIA

banikdipanwita@yahoo.com m_sanjappa@yahoo.co.in

ABSTRACT

Five new varieties of the genus Agapetes D. Don ex G. Don from Arunachal Pradesh, Mizoram and Nagaland, India are described and

illustrated

Key Wonps: Agapetes, new varieties, Arunachal Pradesh, Mizoram, Nagaland, India

RESUMEN

Se describen e ilustran cinco nuevas variedades del género Agapetes D. Don ex G. Don de Arunachal Pradesh, Mizoram y Nagaland,

India

The genus Agapetes D. Don ex G. Don (Ericaceae) comprises ca. 95 species (Mabberley 1997) or ca. 80 species

(Ruizheng & Stevens et al. 2005) and is distributed from the E Himalayas through SW China and Indochina

to SE Asia. In India, the genus comprised ca. 34 species (Santapau & Henry 1973). Presently in India there

are 58 species and 15 varieties under the genus Agapetes.

A revision of the genus Agapetes for the Flora of India project has involved extensive field surveys in

different states of North East India and studying herbarium specimens at various Indian herbaria. During

this critical study, five new varieties of Agapetes came to light; they are described and illustrated here.

1. Agapetes acuminata D. Don ex G. Don var. tipiensis Banik € Sanjappa, var. nov. v. (Fig. D. Tyee: INDIA.

ARUNACHAL PRADESH: West Kameng district, on the way from Tipi to Sessa, 1050 m, 23 Apr 2002

ISOTYPES: ASSAM, CAL).

A varietate typica inflorescentia subumbellata, pedunculo breviore 2-3 mm longo, pedicello breviore 5-5.5 mm longo differt.

Epiphytic straggling shrubs, sometimes terrestrial, 0.5-1 m long. Lignotubers basal, amorphous in

epiphytic ones. Stems robust, terete, obscurely striate, rugose, glabrous, densely lenticellate even when

young; perulae 5-9, closely to loosely alternate, 4-6 mm apart, deltoid to linear, 0.5-2 x 0.2-1 mm,

serrate at margins, acuminate at apex, scaly, caducous. Leaves loosely 2-3-stichous, 1.5-3 cm apart;

petioles 0.7-1.5 cm long, obscurely rugose, slightly pulvinus at base, glabrous; lamina 10.5-25.5 x 2.5-7

cm, lanceolate to ovate-lanceolate, broadly cuneate to rounded at base, bluntly serrate (serratura 10-18

pairs) at margins, acutely acuminate (acumen 1-1.5 cm long) at apex, chartaceous, glabrous; veins

depressed above, brochidodromus, secondary veins 10-18 pairs at 45°-60°, marginal vein 0.5-4 mm

away from margins. Inflorescences axillary or cauline, subumbellate, 7-9-flowered, sparsely puberu-

lous; peduncles 2-3 mm long; peduncular bracts 5-11, deltoid or boat-shaped, 0.5-1.5 x ca. 0.4 mm,

acute at apex, scaly, caducous; pedicels crimson red, 5-5.5 mm long, obconical, ca. 3 mm broad at apex,

fleshy, puberulous, articulate at the base of calyx; pedicellar bract 1, linear, 1-1.5 x 0.5-0.6 mm, serrate

at margins, acuminate at apex, midvein raised outside, scaly, caducous; bracteoles 2, opposite, 0.5-1 x

0.2-0.3 mm, similar to bracts. Calyx crimson red, 2.53 x ca. 3.1 mm, puberulous; lobes deltoid ca. 1.3

x 1.5 mm, acute at apex. Corolla crimson red, urceolate, 3.5-4 x 4-4.5 mm; lobes 1-1.3 x 1-1.5 mm.

Stamens whorled, 3-3.5 mm long; filaments linear, 1-1.5 mm long, slightly incurved, sparsely pilose;

anthers 2-2.5 mm long including 1-1.1 mm long thecae, sparsely granular, tail obscure and tubules

J Bot Res. Inst Texas 1(2): 003 — 912. 2007

904 Journal of the Botanical R h Institute of Texas 1(2)

Hw

di

TE

i

p

"d

A K

e

s

Del. D. Banik

Fic. 1. A t ji D. Don ex. G. Don var.

IR F 3 FF

b = 2 mm; c = 3 mm; d-e = 1 mm (a-e, D. Banik 27953).

habit; b. flower; c. calyx and style; d-e. stamens. Scale bars: a = 1 cm;

Banik and Sanjappa, New varieties of Agapetes from India 905

1.3-1.4 mm long. Style cylindric, 3.5-4 mm long, glabrous, slightly impressed inside ovary, caducous;

stigma glandular-truncate. Fruit not seen.

Notes.—This variety differs from the typical variety in having subumbellate inflorescences, shorter

peduncles (23 mm long) and pedicels (5-5.5 mm long) whereas the typical variety has corymbose in-

J 1

florescences, 3.5-7 mm long p and 1-1.6 cm long pedicels. Variety tipiensis is found in Arunachal

Pradesh whereas the type variety occurs in Meghalaya (India), Bangladesh and Myanmar.

Distribution and Habitat.—India: Arunachal Pradesh. Common in the subtropical forests at altitudes

ranging from 1050-1200 m.

Flowering.—April-June.

Etymology.—This variety is named after the type locality.

PARATYPE: INDIA. Arunachal Pradesh: West Kameng district, Aka hills, Jun 1934 (fl), Bor 15741 (ASSAM).

2. Agapetes flava (Hook. f.) Sleumer var. nagensis Banik & Sanjappa, var. nov. d 2). Tyee: INDIA. NAGALAND:

Naga (Barail) hills, above Konoma, 2100 m, Jul 1886 (fl), D. Prain s.n. (HoLoTYPE: CAL, 264391)

A varietate t lycis lobis multo breviore 4-6 mm longo, ovato; foliis serrato, basi cuneato differt.

E

Epiphytic shrubs. Lignotubers basal, amorphous, 7-10 x 5-6 cm. Stems slender to robust, terete, stri-

ate, lenticellate, glabrous but sparsely puberulous when young. Leaves: pseudowhorls 2.2—4.5 cm apart,

2-3-leaved; petioles 3-6 x 1.5-2.5 mm, puberulous; lamina lanceolate to broadly elliptic, 7-8.5 x 1.7-3.8

cm, obtuse to cuneate at base, serrate and recurved at margins, acutely acuminate (acumen 0.8-1.7 cm

long) at apex; veins depressed above, midvein puberulous at base, brochidodromus, 13-14 pairs of

secondary veins at 60-85". Inflorescences axillary or cauline, corymbose or 5 — 6 flowers in fascicles,

puberulous; peduncles 1-2.5 mm long; peduncular bracts 2-3, closely alternate, ovate to linear, 0.5-1 x

ca. 0.5 mm; pedicels slender, 1.2-1.8 cm long, puberulous; pedicellar bract 1, triangular 1-1.5 x ca. 0.5

mm, puberulous; bracteoles 2, linear, 2-2.5 x ca. 0.5 mm, acuminate at apex, puberulous. Calyx winged,

8-10 x 4—4.5 mm, puberulous; lobes ovate, 4—6 x 2.5—3.5 mm, acute at apex. Corolla yellow, tubular-

urceolate, (1.4—1.5)20—22 x 0.5-0.6 cm, puberulous; lobes broadly triangular, obscure or ca.1 mm long.

Stamens (1.4)1.8-2 cm long; filaments 1-2 mm long, ellipsoid, glabrous to sparsely puberulous outside;

anthers (1.3)1.7-1.8 cm long including 4-4.5 mm long thecae, granular, tail ca. 0.5 mm long, slightly

recurved, granular and tubules (0.9)1.3-1.4 cm long, obscurely calcarate, spurs obscure, basal to tubule.

r

Ovary 3-4.5 mm in diameter; style slender, 1.4-1.6 cm long; stigma truncate. Fruit not seen.

Distribution and Habitat.—India: Nagaland. Grows in subtropical forests at ca. 2100 m.

Flowering.—June—August.

Etymology.—This variety is named after the type locality.

Notes.—This variety differs from the typical variety in having much shorter calyx lobes 4-6 mm

long, ovate and leaves serrate, cuneate at base whereas the typical variety has calyx lobes 8-11 mm long,

elliptic-lanceolate and leaves obscurely crenate, rounded at base. In typical variety the corolla tube al-

most is covered by calyx. Variety nagensis is found in Nagaland whereas the type variety is known from

Arunachal Pradesh.

PARATYPES: INDIA. Nagaland: Naga (Barail) hills, above Konoma, Aug 1886 (fl), D. Prain s.n. (BSIS, 17874); D. Prain s.n. (£D, (CAL,

264390); D. Prain s.n. (f1) (CAL, 264392).

3. aD megacarpa WW. Sm. var. lohitensis Banik & Sanjappa, var. nov. Duis 3). Tyre: INDIA. ARUNACHAL

10 )

ESH: Lohit district, Mailiang to Chipra, ca. 1400 m, 16 May 2003 (fl), D. Banik 27 G AL; isotypes: ASSAM

A varietate typica calycis lobis multo longioribus lineari-ellipticis 1.6-2.1 cm longis 3-5 mm latis differt.

Epiphytic or lithophytic shrubs, erect, 2-3 m high. Lignotubers basal, amorphous. Stems robust,

terete, striate, lenticellate, glabrous; perulae 6-12, close to loosely alternate, ovate to linear-lanceolate,

5-10 x 2.5-6 mm, serrate at margins, acuminate at apex, caducous to persistent. Leaves: pseudowhorls

2.5-5 cm apart, 3-6-leaved; lamina sessile, elliptic or broadly elliptic, 12-18 x 2.7-6 cm, auriculate to

906 Journal of the Botanical R h Institute of Texas 1(2)

| -

1

2"

A.

{ —

AA

"

Del. D. Banik

Fic. 2. Agapetes flava (Hook. f.) Sleumer var. nag jap

5 mm; d-e = 3 mm [a-e, D. Prain s.n. (Herb. Acc. No. 264392)].

907

Banik and Sanjappa, New varieties of Agapetes from India

Del. D. Banik

e. stamens; f. basal portion of stamens;

ry and stula: d—

yle;

7

habit; b flower; c caly

5 mm (a-g, D. Banik 27534).

=1 cm; d, e, 9 =

N. Sm. var. lof

F

2

Jr

g. apical portion of tubules. Scale bars: a-c, f

Fic. 3. A

908 Journal of the Botanical R h Institute of Texas 1(2)

cordate at base, entire to serrate at margins and obscurely recurved towards apex, acuminate to acutely

acuminate (acumen 1-1.5 cm long) at apex, coriaceous, glaucous above, dull below; veins raised above,

brochidodromus, 10-12 pairs of laterals at 45°-80°. Inflorescences axillary or pseudoterminal, corym-

bose, 6-15-flowered, glabrous, maroon, dealbatus; peduncles maroon, 1-1.5 cm long, 4-5 mm broad,

stout, glabrous, dealbatus; peduncular bracts maroon, 5-6, linear, 1-1.5 x 0.5-0.6 cm, prominently

veined, glabrous, dealbatus, caducous; rachis 3-3.5 cm long; pedicels 2-3-stichous, maroon, obconical,

23.2 cm long, 0.7-1 cm broad at apex, stout, glabrous, dealbatus; pedicellar bract 1, triangular, 3.5-4

x 1.5-2 mm, serrate, acute; bracteoles 2, opposite, linear, 2-3 x ca. 1 mm, serrate, acuminate, caducous.

Calyx maroon, 2.3-2.5 x 1.5-1.6 cm, glabrous, rugose at base, dealbatus; lobes linear-elliptic, 16-21 x

3-5 mm, slightly constricted at base, entire at margins, acuminate to obtusely acuminate at apex, thickly

coriaceous, 3-4 pairs of secondary veins at 15°-30°, raised above. Corolla maroon-red, tubular-urceolate,

5-6 x 1.5-2 cm, glabrous, dealbatus, 6-7 pairs of secondary veins at 30°-60°, dark maroon, directed

upwards forming closed reticulations; lobes green, ovate-caudate or triangular-caudate, 15-16 x 6—6.5

mm, glabrous, dealbatus. Stamens 10, 2-whorled; filaments spathulate, 4.5—7 mm long, incurved, sparsely

hirtellous outside; anthers 4.6—4.8 cm long including 1.1-1.5 cm long thecae, verrucate, tail 0.5-0.7 mm

long, verrucate, reflexed and tubules 3.2-3.7 cm long, sparsely verrucate to smooth, calcarate, spurs

apical on tubule, 0.5-0.7 mm long, verrucate, ascending and reflexed in adjacent anthers. Style slender,

ca. 5.7 cm long; stigma 5-lobed, 23 x 2-2.5 mm. Fruit a berry, maroon, obovoid, 2.5-2.8 x 1.5-1.8 cm,

glabrous, dealbatus. Seeds obovoid, 3-5 x 2-2.5 mm, pointed at base; seed coat reticulate.

Distribution and Habitat.—India: Arunachal Pradesh. Common in the subtropical forests at altitudes

ranging from 1300-1500 m.

Flowering and Fruiting.—March-May.

Etymology.—This variety is named after the type locality.

Uses.— Corolla used as vegetable.

Note.—This variety differs from the typical variety in having much longer (1.6-2.1 cm) linear-el-

liptic calyx lobes whereas the typical variety has shorter (6.5-11 mm) ovate to lanceolate calyx lobes.

Variety lohitensis is found in Arunachal Pradesh whereas the typical variety is distributed in Nagaland

(India), China, and Thailand.

PARATYPES: INDIA Arunachal Pradesh: Lohit district, Kuibang, 25 Mar 1986 (fD, K. Harid 2909 (APFH); Lailiang to Mailiang, near

Lailiang, 1400-1500 m, 16 May 2003 (£D, D. Banik 27533 (CAL).

4. Agapetes odontocera (Wight) Hook. f. var. mizoramensis Banik AD var. nov. (Fig. 4). Tee: INDIA:

Mizoram, Thaltlang phui, 16 May 1990 (fl), B.M. Wadhwa & K.P Singh 96834

A i Į fl ifi 1 la differt

E

Small trees. Stems robust, terete, obscurely striate, lenticellate, glabrous; perulae many, compactly al-

ternate to loosely alternate (5-8 mm apart), triangular to linear-triangular. Leaves: pseudowhorls 4-5.5

cm apart, 3-5-leaved; petioles 1-2 mm long, glabrous; lamina narrowly elliptic-oblanceolate, 14-15 x

2.3-3.8 cm, cuneate to obtuse at base, serrate at margins, acutely acuminate at apex, coriaceous; veins

raised above, brochidodromus, 15-16 pairs of secondaries at 45°—60°. Inflorescences axillary or cauline,

flowers 3-6 in fascicles, puberulous; inflorescence bracts closely alternate; pedicels slightly obconical,

1.8-2 cm long, puberulous; pedicellar bract 1, deltoid, ca.1 x 1 mm, scaly, caducous; bracteoles 2, op-

posite, linear, ca.1 x 0.5 mm, similar to bract. Calyx ca. 5 x 2.5 mm, puberulous; lobes ovate-triangular,

ca. 3.5 x 2 mm, acuminate at apex. Corolla tubular, 2.5-2.8 x 0.8-1 cm, puberulous; lobes triangular,

ca. 1 x 0.5 mm, acuminate at apex. Stamens 10, in 2 whorls, ca. 2.5 cm long; filaments spathulate, 2-2.5

mm long, slightly incurved, puberulous to pilose outside; anthers 1.8-2.2 cm long including 4.5-5.5 mm

long thecae, granular, tail obscure and tubules 1.4-1.6 cm long, sparsely granular, calcarate, spurs ca. 1

mm long, linear, reflexed. Style slender, 2.5-2.7 cm long; stigma truncate. Fruit not seen.

Distribution.—India: Mizoram.

Banik and Sanjappa, New varieties of Agapetes from India 909

Del. D. Banik

J Minha U I . D TOC

Fic. 4 Ag ji (Wight) F. var. j habit; b. flower; c calyx and style Scale bars: a = 1 cm; b-c = 5mm

(a-c, B.M. Wadhwa & D.K. Singh 96834).

910 Journal of the Botanical R h Institute of Texas 1(2)

Flowering.—May.

Etymology.—This variety is named after the type locality.

Notes.—This variety differs from the typical variety in having uniformly puberulous inflorescences

whereas the typical variety has totally glabrous inflorescences. Variety mizoramensis is recorded from

Mizoram whereas the typical variety is widely distributed in Arunachal Pradesh, Assam, Meghalaya,

Mizoram, Manipur and Nagaland.

5: sa ite pe C.B. Clarke var. glanduliflora Banik € Sanjappa, var. nov. (Fig. 5). Tyee: INDIA. ARU-

ESH: Lower Dibang valley district, Mehao lake to Roing, 27 Nov 2000 (fl), D.K. Singh & Party 9319 (HoLotTyPE: CAL;

ISOTYPE: Rum

A varietate typica inflorescentia glandulosopubescenti vel glandis subsessilibus in pedunculo, pedicellis, calyce et corollae costis dif-

fert

Epiphytic shrubs, erect, 0.5-1 m tall. Stems robust, terete, 5-8 mm in diameter, striate, glabrous,

lenticellate when mature; perulae 3-6, closely to loosely alternate, 1—4.5 x 1-1.2 mm, deltoid-ovate to

linear-elliptic, entire to glandular at margins, acute to acuminate at apex; midnerve raised, glabrous,

persistent. Leaves: pseudowhorls 5.5-8.5 cm apart, 8—9-leaved; petioles 4-8 mm long, glabrous, winged,

slightly rugose outside, slightly pulvinus at base; lamina 6-16 x 1-2.5 cm, linear-elliptic, attenuate at

base, entire at margins, acuminate at apex, coriaceous, glaucous above, pale below; veins slightly raised

above, brochidodromus, 12-16 pairs of secondary veins, alternate at 30-80”. Inflorescences axillary or

cauline, corymbose, erect, 9-16-flowered, glandular hairy; peduncles dark red, 3.5-5.5 mm long, glan-

dular hairy; peduncular bracts 3-6, closely alternate, broadly deltoid, obscure, glabrous, membranous,

caducous; pedicels dark red, 4.5-7 mm long, striate, glandular hairy; pedicellar bract 1, 1-1.5 mm long,

linear, persistent; bracteoles 2, linear, ca.1 mm long, similar to bracts. Calyx dark red, 5-6.5 x 3.5-4.2

mm, glandular hairy or with subsessile glands; lobes ovate, 2.83.5 x 1.3-2.5 mm, acuminate at apex,

membranous, veins raised above, glandular hairy. Corolla dark red to orange red, tubular-urceolate,

1.5-2 x 0.4-0.5 cm, angular, membranous, reticulate in dark red, secondary veins 5-6 pairs, at ca. 60°,

midveins glandular hairy; lobes green or greenish- yellow, 1.5-2 x ca. 1.5 mm, ovate, acute to obtuse at

apex, glandular hairy. Stamens 2-whorled, 1.5-1.8 cm long; filaments 2.5-3 mm long, linear, incurved,

puberulous outside, hyaline; anthers 1.2-1.5 cm long including 5-5.5 mm long thecae, verrucate, tail

bilobed, recurved and tubules ca. 10 mm long, glabrous. Ovary 33.5 mm in diameter, striate, glabrous;

style slender, 1.7-1.9 cm long, ca. 1 mm deeply impressed inside ovary; stigma glandular capitate, 5-

lobed, ca. 0.5 x 0.6 mm. Fruit a berry, globose, 4—6.5 x 4.5-5 mm, sparsely glandular hairy or with

sessile glands. Seeds obovoid, 1-1.2 x 0.5-0.7 mm; seed coat reticulate.

Distribution and Habitat.—India: Arunachal Pradesh. Common in the subtropical forests between

1300 and 1400 m altitude.

Flowering and Fruiting.— October-November.

Etymology.—This variety is named for the glandular pubescence of inflorescence.

Notes.—This variety differs from the typical variety in having glandular hairy inflorescence or with

subsessile glands on peduncles, pedicels, calyx and midveins of corolla whereas the typical variety has

glabrous and eglandular inflorescence. Both varieties are distributed in Arunachal Pradesh.

PARATYPES: INDIA. Arunachal Pradesh: Lower Dibang valley district, Tiwari gaon to Roing, 1300 m, 3 Nov 2002 (fl), D. Banik 27413

(CAL); Mehao lake to Baldi camp, 1400 m, 16 Nov 2002 (fD, D. Banik 27432 (CAL); Lohit district, Hawa camp to Udayak pass, 1400 m,

24 Apr 2003 (fr), D. Banik 27508 (CAL).

ACKNOWLEDGMENTS

The authors are thankful to J.F. Veldkamp, Nationaal Herbarium Nederland, Leiden, the Netherlands

for the Latin diagnoses of the new varieties. Pakshirajan Lakshminarasimhan and one anonymous re-

viewer provided helpful review comments. One of us (DB) is grateful to the Director, Botanical Survey

of India, for granting fellowship, to D.K. Singh, the Additional Director; Botanical Survey of India for

Banik and Sanjappa, New varieties of Agapetes from India 911

>

LESS >

=

(=

3

ea

2

=

o

a

d

IG. 5 C.B. Clarke var. g Banik & Sanjappa: a. habit; b. flower; c. caly yle; d. stamen; e-h. peduncular bracts; i. t.s.

of ov. 2an;b 5 d = 3 mm; e-h = 2 mm; I = 1 mm (a-i, D.K. Singh & Party 9319).

912 Journal of the Botanical R h Institute of Texas 1(2)

specimens; and to the Joint Director, Central National Herbarium, Botanical Survey of India, Howrah,

for facilities.

The acronym APFH is used in the text for The Herbarium, State Forest Research Institute, Arunachal

Pradesh, Van Vihar, Chimpu, Itanagar 791 111, Arunachal Pradesh, India.

REFERENCES

Massertey, D.G. 1997. The plant book.2™ ed. (Reprinted with corrections 1998, 2000). Cambridge University

Press.

RUIZHENG, F. and PF. Stevens. 2005. Agapetes. In: Editorial Committee, eds. Flora of China, vol. 14. Science Press,

Beijing and Missouri Botanical Garden Press, St. Louis. Pp. 504-516.

SANTAPAU, H. and A.N. Henry. 1973. A dictionary of the flowering plants in India. (Reprinted in 1998). National

Institute of Science Communication, New Delhi.

PHYLOGENETIC ANALYSES OF THE GENUS BAPTISTONIA (ORCHIDACEAE:

ONCIDIINAE) SENSU LATO BASED ON MORPHOLOGICAL CHARACTERS

Guy R. Chiron

biers, Université Claude Bernard - Lyon 1

9 rue roe Dubois, 69622 Villeurbanne Cedex, FRANCE

ofr

chiron@wanado

ABSTRACT

Pelar; 1 ip shin R l ; ; ; 11 1 pl ; Uem 1 £47 pl ] gi Tel The analyses support

monophyly of Baptistonia, but without strong | p support. S praspecifi pported by th l di d

RÉSUMÉ

On présente, sur la base d'une analyse Pune que et ona de 47 caracteres morphologiques, les relations de parenté entre les

espèces de Baptistonia. L'étude soutient 1 genre, avec cependant de faibles valeurs de bootstrap. Quelques

taxons supraspécifiques, suggérés par l'analyse, sont den

Key Worbs: Baptistonia, Brazil, Gomesa, Orchidaceae, Oncidiinae, phylogeny

INTRODUCTION

The genus Baptistonia was founded in 1877 by Barbosa Rodrigues based on a single species: B. echinata Barb.

Rodr. Soon after the concept was published, it was generally neglected. Kránzlin (1922), in his revision of

the entire subtribe Oncidiinae, placed the species in the genus Oncidium Swartz. However, Pabst and Dungs

(1977), in their work on the Orchidaceae of Brazil, conserved Barbosa Rodrigues' monospecific genus.

Chiron and Castro Neto (2004) noted that the characteristic vegetative and floral structures of the

plants of the section Waluewa (Regel) Schltr. of the genus Oncidium are close to those of Baptistonia echinata,

and that all species involved are endemic to the southeastern part of Brazil. Consequently they transfered

the species of section Waluewa to the genus Baptistonia. The thus enlarged genus now comprises twenty one

species and two natural hybrids.

DNA sequence data has shown that the species involved are distinct from those belonging to the true

genus Oncidium (Williams et al. 2001; Chase et al. 2005). This data also seems to indicate that these species

form a monophyletic group which is distinct from its neighboring groups (Faria 2004). It should however be

noted that the work of Chase et al. and Faria is based on a limited number of species and that the variations

observed in the analyzed sequences are extremely small.

At this time, a study based on the trnS-G region of a large number of members of the genus is in progress.

Parallel to this study, however, I also wanted to explore the utility of morphology and anatomy in respect to

phylogenetic relationships. Faria (2004) published morphological phylogenetic of seven Baptistonia species

(less than one-third of the group) but included representatives of various other groups of Brazilian oncidi-

ums. The objective of my study is to complement the work of that author and to estimate the interspecific

relationships of all naturally occurring Baptistonia taxa at the species level, the hybrids being excluded.

MATERIALS AND METHODS

The anatomical and morphological characters that are presented and discussed below have mainly been

obtained through the observation of several live plants of each species. Herbarium material was only utilized

whenever live material was not available in any sufficient quantity. The observations were supplemented by

data deduced from the original descriptions of the taxa.

The genus Baptistonia is closely related to the Gomesa alliance, a Brazilian Oncidiinae group compris-

ing the genera Gomesa R. Br., Rodrigueziella Kuntze and Rodriguesiopsis Schltr. In this study, I included four

J. Bot. Res. Inst. Texas 1(2): 913 — 931. 2007

914 Journal of the Botanical R h Institute of Texas 1(2)

Gomesa species as outgroup: G. alpina Porsch, G. crispa Klotzsch ex Rchb.f., G. recurva R. Br. and G. sessilis

Barb. Rodr. The trees were rooted with three species of the genus Oncidium: O. altissimum Swartz (the type

species of the genus, originating from the Antilles), O. nebulosum Lindl. (a mesoamerican species belonging

to Oncidium section Oblongata Kraenzlin) and O. baueri Lindl. (another species of the section Oblongata,

native of the Amazonian forests.) Furthermore, I have added two species that I suspect to be close to the

species of the genus Baptistonia:

— Carriella colorata (Kóniger & J.G. Weinm.) V.P. Castro & K.G. Lacerda, which was placed in Oncidium

section Waluewa by Senghas (1997) but elevated to a monospecific genus by Castro Neto and Lacerda (2006),

and

—Oncidium trulliferum Lindl., close to the plants of the genus Baptistonia in respect to its vegetative

characters as well as in respect to some of its floral structures, but generally placed in Oncidium section

Rostrata Rolfe.

In view of the work of Faria (2004), no specimens were included belonging to any of the other groups of

the Gomesa clade (a monophyletic group consisting of the Gomesa alliance, Baptistonia and many of the

Brazilian oncidiums).

All specimens that were studied, with the exception of those belonging to the external Oncidium group,

are epiphytic plants originating from the moist forests of the Mata Atlantica. A list of the 217 specimens

studied is given in Appendix 1.

The second important issue to discuss relates to the pertinence of the characters that have been se-

lected for the analysis. Chase (1986) stated that several morphological characters, such as the angle between

the labellum and the column, the fusion of the lateral sepals, the general shape of the floral segments, the

column arms, and the callus of the labellum, cannot be used for an objective phylogeny. Faria (2004), on

the other hand, disagrees with Chase's opinion on this matter. Burns-Balogh and Funk (1986) as well as

Freudenstein and Rasmussen (1999) discussed a great number of morphological characters in their cladistic

analysis of Orchidaceae. In these articles, however, the utility of the characters was evaluated only in respect

to generic relationships, not interspecific. Each of the observable characters may either be invariant within

al

(e)

the groups studied (and therefore are of no use for pl ic analysis), may vary among species, or—as a

third possibility—may even vary within a given species, thus being inconsistent and therefore unsuitable for

species-level phylogenetics. Thus, this pertinence of the 69 selected vegetative and floral characters, either

microscopic or macroscopic, needs to be discussed, in order to (a) eliminate the unsuitable ones, and (b)

decide how many different states of each character retained can be used in the present study. In accordance

with the recommendations given by Garcia-Cruz and Sosa (2006) for quantitative characters, I employ the

Gap-Weighting (GW) coding method as proposed by Thiele (1993), with a limited number of statistical

conditions. In this way, the danger of overweighting certain characters is eliminated. Polarity of character

coding is based upon pleisiomorphy of Oncidium.

According to van den Berg (unpublished), the Oncidiinae evolved, at the beginning of the Miocene,

about 23 million years ago, from an ancestor belonging to the tribe Maxillarieae, itself having diversified

from a Mesoamerican ancestor. At that time, South and North America were still separated from each other,

the Panamean isthmus having been formed during the Pleiocene. It is generally assumed (Por 1995) that

there was an inland sea from the east of what is now Colombia to the Pantanal, isolating the entire eastern

part of South America, the sea not being connected with the Atlantic Ocean until the Miocene. Therefore,

the Brazilian Oncidiinae, especially the Gomesa clade, would have diversified much later, on the basis of

an ancestor in Central America. The species of Oncidium sensu stricto have their center of distribution in

Central America, and they are probably much closer to the common ancestor than the species belonging to

the genus Baptistonia.

Morphological and anatomical characters considered

According to the above discussion, the following twenty characters were considered but excluded: spacing,

Chiron, Phylogenetic study of Baptistonia 915

diameter and surface structure of the pseudobulbs; length, shape, color of the leaves, consistency of the

lamina, distribution of the stomata, shape of the epidermal cells; length and diameter of the peduncle of the

inflorescence, length of sterile bracts, flower density; fusion of the lateral sepals of the flowers, outline and

apical shape of the lateral sepals, shape and position of the lateral lobes of the lip, curvature of the column,

shape of the stigmatic cavity. Some of them correspond to what Chase (1986) calls “unreliable characters.”

A. Pseudobulbs (mature but young)

* Height (character 01)

Within any given species, the height of the pseudobulbs may vary. Consequently, I used the maximum

height observed. The GW Method, applied with the number of states limited to 2, allows setting a limit of

7 cm to differentiate between species with small or large pseudobulbs.

01-maximum height of pseudobulbs: 1 up to 7 cm—2 more than 7 cm

Assumed evolution: 1 — 2

* Shape in vertical section (character 02)

The distinctly elongated shape of the pseudobulbs is characteristic for the genus, but certain species of

Baptistonia do have pseudobulbs that are elongated-ovoid as in Gomesa. For the evaluation of this character

I have chosen to use the ratio of height over diameter (H/D), the diameter being measured at the thickest

part of the pseudobulb. The scoring, according to the GW method, has three states:

02-Vertical section of the pseudobulbs: 1 ovate (H/D«2)—2 elongated ovate—3 cigar shaped (H/D»4)

Assumed evolution: 1 > 2 > 3

* Shape in horizontal section (character 03)

Generally, the horizontal section of the pseudobulbs is rounded, but, in Baptistonia, it is sometimes somewhat

flattened although never strongly flattened (elliptical). This character is evaluated by the ratio (D/d) of the

large diameter over the small diameter of the section, calculated according to the GW method accepting

two states:

03—horizontal section of the pseudobulbs: 1 strongly flattened (D/d > 1.5)—2 slightly flattened (D/d «

Assumed evolution: 1 > 2

* Margin of pseudobulb in horizontal section (character 04)

The pseudobulb in an horizontal section is generally ribbed to entire and this character seems to be a per-

tinent one at the generic level, although not at the specific level.

04—Margins of the horizontal section: 1 angular—2 intermediate—3 rounded

Assumed evolution: 1 > 2 3

* Color (character 05)

When observed in their natural state, the plants belonging to the genus Baptistonia are easily differentiated

from the plants of the genera Oncidium or Gomesa by the color of their pseudobulbs and leaves. The shade

of the green of the pseudobulbs and leaves varies distinctly among these genera. Under cultivation at the

same spot, this difference remains, showing that the various shades of green are not adaptations to chang-

ing environmental conditions.

05—color of the pseudobulbs and leaves: 1 bright green—2 dark green

Assumed evolution: 1 — 2

B. Leaves

Williams (1974) studied the morphology and the anatomy of the leaves of 80 species belonging to a total

of 22 genera of the Oncidiinae. Of those 80 species only 4 belonged to the Gomesa clade. He concluded

that the leaves offer very few interesting morphological characteristics, but that some anatomic features

can be very useful. For the evaluation of those characters in the genus Baptistonia, see Chiron and Guiard

(in preparation).

916 Journal of the Botanical R h Institute of Texas 1(2)

* Basal sheaths (character 06)

At the generic level this character is useful as, in Oncidium, the sheaths are long and leafy, hardly shorter

than the real leaves. On the other hand, in Gomesa, they are leafy but short and, in Baptistonia, they are

non-leafy.

06—Basal sheaths: 1 long and leafy—2 short and leafy—3 non-leafy

Assumed evolution: 1 > 2 > 3

* Number of leaves (character 07)

The majority of the species of Baptistonia show a varying number of leaves per pseudobulb. Therefore, for

the present study, I considered the maximum number of leaves for any given species.

07-Maximum number of leaves: 1 more than one—2 a single leaf.

Assumed evolution: 1 > 2

* Stomata density (character 08)

The density of the stomata on the abaxial surface of the leaves in the genera Baptistonia, Oncidium and Gomesa

varies from one species to the others (Chiron & Guiard, in preparation). Although this density also varies

within any given species, the intraspecific variation is much lower than the variation among the species.

The density of the stomata in most of the species is between 20 and 45 stomata per mm?; in the other spe-

cies studied, the density was higher than 55 stomata per mm". This character therefore is pertinent at the

generic level.

08-stomata density: 1 2 55 per mm?—2 « 45 per mn?

Assumed evolution: 1 — 2

* Shape of the stomata (character 09)

The shape of the stomata as observed in Gomesa and Baptistonia is sub-circular to elliptic, whereby the ratio

major axis/minor axis can attain 1.4. Notwithstanding the fact that this character can vary considerably within

any given species, the differences among the different species allow scoring for three different states.

09—Ellipticity of the stomata: 1 € 1.05—2 > 1.05 and < 1.23 > 1.2

Assumed evolution: 1 + 2 > 3

C. Inflorescence

* Total length (character 10)

The total length of the inflorescence varies from one species to another. This character must be measured

on adult plants during a normal flowering.

10-length of the inflorescence: 1 distinctly longer than the growths—2 as long as or shorter than the growths

Assumed evolution: 1 2

* Direction (character 11)

The inflorescences are generally erect, later becoming arched under the weight of the flowers. However, in

certain species of the genus Baptistonia, they are abruptly folded downward directly above the base and long

before the buds begin to form.

11—direction of the inflorescence: 1 upward, more or less arched—2 directly folded downward

Assumed evolution: 1 > 2

* Form (character 12)

Generally, the inflorescences are branched. They are, however, unbranched in Gomesa and in some Baptis-

tonia species.

12- inflorescence: 1 branched -2 unbranched

Assumed evolution: 1 — 2

* Formation of the buds (character 13)

In general, the buds start to develop after the inflorescence has more or less reached its final length. This is

not so in the genus Gomesa and in some other species of the Gomesa clade. In those cases, the buds start to

Chiron, Phylogenetic study of Baptistonia 917

develop as soon as the inflorescence starts to develop within the protection of the leafy sheaths.

13—Formation of the buds: 1 when the inflorescence has attained its length—2 as soon as the inflorescence

starts to develop

Assumed evolution: 1 > 2

D. Flowers

* Form (character 14)

Depending on the species that is under observation, the flowers of Baptistonia can be more or less spread or

more or less closed and as such globular. I consider this character potentially pertinent.

14—flower: 1 fully spread (more or less in one plane)—2 quasi spherical (petals and dorsal sepal directed

forward).

Assumed evolution: 1 > 2

e Outline (character 15)

The outline of the flowers can be distinctly circular or vertically ovate. This was determined by using the

ratio (H/W) between height and width of the flowers.

15—Flower outline: 1 circular (H/W < 1.1)—2 vertical (H/W > 1.1)

Assumed evolution: 1 — 2

* Color (character 16)

Originally, I had based my color evaluation of the flowers on the visual absence of yellow, red, and green

pigmentation. However, even when we attribute but two states to each color, meaning “visible” and “non-

visible", we obtain a total of 8 possible states. This is too many states for this type of character. Finally, I

have decided to retain three states.

16-Flower color: 1 mainly colored bright yellow and/or dark red—2 pale yellow to whitish with reddish-

violet spots—3 yellowish or greenish without any red spotting

Assumed evolution: 1 2 2 > 3

* Floral bracts (character 17)

Within the genus Baptistonia, this characteristic does not vary much, but long flower bracts are typical for

the species of the genus Gomesa. The GW coding allowing for 3 states gives:

17—Floral bracts: 1 very short («1/5 pedicel)—2 short—3 long (> 1/2 pedicel)

Assumed evolution: 1 — 2 3

D1. Tepals

* Outline of the dorsal sepal (character 18)

This character is partially pertinent within the genus Baptistonia.

18-Outline of the dorsal sepal: 1 obovate—2 spathulate

Assumed evolution: 1 > 2

* Degree of concaveness of the dorsal sepal (character 19)

Several species of Baptistonia have a spoon-shaped dorsal sepal. Therefore, this character seems to be per-

tinent within the genus Baptistonia as well as at the generic level.

19—dorsal sepal: 1 plane—2 cucullate

Assumed evolution: 1 — 2

* Outline of the petals (character 20)

In order not to overrate this character, I have coded for only two states.

20—Petals: 1 obovate or claviform—2 notched spathulate to panduriform.

Assumed evolution: 1 > 2

* Apex of the petals and of the dorsal sepal (character 21)

The form of the apices of the petals and of the dorsal sepal seems to be characteristic within Baptistonia.

918 Journal of the Botanical R h Institute of Texas 1(2)

21- Apex of the petals and of the dorsal sepal: 1 angular—2 rounded or retuse

Assumed evolution: 1 > 2

D2. Labellum

* Fused (character 22)

The labellum may be fused to the base of the column or not.

22-labellum: 1 fused—2 articulated

Assumed evolution: 1 > 2

e Claw (character 23)

The presence of a long claw is characteristic for the genus Baptistonia. This character, evaluated by the factor

R— being the ratio of the length of the claw over the length of the labellum—was GW coded for 3 states.

23-Labellum claw: 1 absent or nearly so (R< 1/20)—2 short (1/20<R< 1/5)—3 long (R> 1/5)

Assumed evolution: 1 => 2 3

* Shape of the lateral lobes (character 24)

24—Lateral lobes of the labellum: 1 somewhat elongated (rounded, triangular or square)—2 wider than

long-—3 non-existent—4 very elongated (linear, elongated triangular, or linguiform)

Assumed evolution: 1 >2 5 3 and 1 > 4

* Direction of the lateral lobes within the labellum plane (character 25)

25—Direction of the lateral labellum lobes: 1 to 90° away from the lip axis—2 toward the base of the lip—3

toward the apex

Assumed evolution: 1 2 and 1 3

* Direction of the lateral lobes in respect to the position of the labellum plane (character 26)

26-Lateral lobes: 1 more or less within the plane of the labellum or slightly curved forward—2 at least their

basal half distinctly directed backward

Assumed evolution: 1 — 2

* Width of the midlobe (character 27)

Evaluated in respect to the width of the labellum at the level of the lateral lobes, this character is variable

within the genus Baptistonia.

27-Width of the labellum at the level of the midlobe: 1 greater—2 similar—3 smaller in comparison to the

width measured at the level of the lateral lobes

Assumed evolution: 1 > 2 > 3

* Shape of the midlobe (character 28)

or of the labellum if it is entire. Besides being variable within the genus Baptistonia, this character seems to

have a certain degree of importance in respect to generic differentiation.

28-Shape: 1 wider than long—2 just about circular or square—3 longer than wide

Assumed evolution: 1 > 2 > 3

* Apex of the labellum (character 29)

29-Apex: 1 entire—2 split

Assumed evolution: 1 > 2

e Isthmus of the labellum (character 30)

I have accorded a great importance to this character as it is a potential differentiating character at the spe-

cies level in Baptistonia. In fact, the character is stable and specific for groups of species. It overlaps slightly

with character 24 (state 3 of character 24 is the same as state 5 of character 30).

30-Isthmus of the labellum: 1 elongate rectangular—2 trapezoidal—3 short and wide with rounded sinus—4

short with triangular sinus—5 absent (labellum undivided)

Assumed evolution: 1 > 2 > 3 > 45

Chiron, Phylogenetic study of Baptistonia 919

* Margins at the sinus (character 31)

A small number of species show a sinus with indented margins and this character appears to be stable and

characteristic.

31-Margins at the sinus: 1 entire—2 irregular

Assumed evolution: 1 2

* Callus (characters 32 to 34)

Within the genus Baptistonia, the callus of the labellum is generally made of three parts: a basal part (on

the claw), a center part (between the lateral lobes) and an apical part (on the isthmus and on the midlobe).

According to Chase (1986), this is a derived condition in the Oncidiinae. In the species that do not have a

claw, the callus has no basal part. In Oncidium, the callus does not extend onto the median labellum lobe. In

Gomesa, the callus is made up of two parallel, longitudinal lamellae, followed by two crests in the apical part

of the labellum. The diversity ofthe structures and their remarkable stability within each species have caused

me to retain three distinct characters and have made me accord special importance to the median part.

32—Basal part: 1 absent or not differentiated, not ending in two teeth—2 ending in two well differentiated teeth

Assumed evolution: 1 > 2

33-Median part made up of: 1 a compact construction of several well-developed teeth—2 some kind of

verrucose and/or sheeted plate—3 a thick, non-verrucose plate, depressed in the center—4 a thin, smooth,

inverted V-shaped plate—5 an inverted Y-shaped crest—6 a construction of small elongated wart-like struc-

tures, more or less spaced in two rows—7 two large, low crests—8 two thin, high crests

Assumed evolution: 1 > 2 => 3 > 4 —> 5and: 1 > 65758

Other character state polarities have been evaluated. This one has been retained because it is most congru-

ent with other characters.

34—Apical part of the callus: 1 absent—2 short (< 1/3 of the midlobe)—3 long (> 1/3 of the midlobe)

Assumed evolution: 1 > 2 > 3

D3. Column

e Slenderness (character 35)

This character is variable within the genera studied and seems to be pertinent for the genus Baptistonia. It is

quantified by the ratio H/W between the height of the column and its width measured below the stigmatic

cavity and coded according to the GW method for three states.

35-Column: 1 medium (2.5 € H/W «4.5)—2 stocky (H/W « 2.5)—3 slender (H/W > 4.5)

Assumed evolution: 1 > 2 and 1 => 3

* Positioning in relation to the labellum (character 36)

This character does not seem to be pertinent within the genus Baptistonia where it is remarkably stable, but

it is useful to differentiate among the genera.

36-column: 1 in the same line as the labellum—2 at an angle of 90? to the labellum—3 tight against the

basal part of the labellum

Assumed evolution: 1 — 2 3

* Pubescence (character 37)

Pubescence of the column is a character of all species within the genus Baptistonia, whereby the degree of pu-

bescence varies. Furthermore, this character is useful for the study of the relationships among the genera.

37-column: 1 glabrous—2 slightly pubescent—3 strongly pubescent

Assumed evolution: 1 > 2 3

* Tabula infrastigmatica (character 38)

The absence of the tabula beneath the stigmatic cavity is common to all species of Baptistonia (although with

a regression in B. uhlii). It marks an evolutionary step in relation to other genera.

920 Journal of the Botanical R h Institute of Texas 1(2)

38-tabula infrastigmatica: 1 well-developed—2 slightly developed—3 absent

Assumed evolution: 1 > 2 > 3

* Shape of the wings (character 39)

39-wings: 1 independently of their shape, wider than long—2 absent or reduced to two minuscule points—3

longer than wide (sometimes becoming wider at the apex)

Assumed evolution: 1 + 2 and 1 > 3

* Spreading of the wings (character 40)

40—wings: 1 opened to 90?—2 absent or opened to 180% —3 parallel or as an arch directed to the front

Assumed evolution: 1 > 2 and 1 > 3

* Hairiness of the stigmatic cavity (character 41)

The hairiness of the margins of the stigmatic cavity varies between nearly absent and very dense.

41—hairiness: 1 hairs nearly or fully absent—2 hairs rare and/or short—3 hairs long and dense

Assumed evolution: 1 2 2 > 3

* Presence and shape of an appendix to the anther (characters 42 to 44)

The apex of the anther can be rounded or elongated, more or less recurved upward and more or less split

into two parts. I have retained three characters to describe this appendix where its absence or presence is

incorporated into those three characters.

42—appendix: 1 absent to short—2 long

Assumed evolution: 1 > 2

43-appendix: 1 simple—2 bifid—3 double

Assumed evolution: 1 > 2 > 3

44—appendix: 1 straight —2 recurved

Assumed evolution: 1 > 2

* Viscidium (character 45)

The form of the viscidium, measured as the ratio D/d of the large and small diameter of the ellipse, seems

to be pertinent at the generic level.

45-viscidium: 1 circular or ovate (D/d«3)—2 elliptic (D/d23)

Assumed evolution: 1 — 2

* Tegula (character 46)

46-tegula of the pollinarium: 1 larger on the side of the viscidium or equally large on both sides—2 spoon-

shaped (larger on the side of the pollinia)

Assumed evolution: 1 > 2

* Margins of the clinandrium (character 47)

Well-developed, irregular margins are characteristic for Baptistonia but little intrageneric variation can be

observed.

47-margins of the clinandrium: 1 only slightly developed—2 extending beyond the anther

Assumed evolution: 1 > 2

The evaluation of these 47 characters has made it possible to develop the data matrix shown in Table 1. This

matrix was subsequently exploited in two different methods: parsimony and distances.

Maximum parsimony analysis.—For the first method the phylogeny for the group was obtained by us-

ing the software package PHYLIP (see Felsenstein 1989, 1993), which is composed of SEQBOOT version

3.570, MIX version 3.572c and CONSENSE version 3.56c. The coding of the characters corresponding to

the evolutionary hypothesis delineated above resulted in a data matrix (not shown here) of 0 and 1. The

software parameters were set as follows:

SEQBOOT: number of replications set to 1000— morphological data

Chiron, Phylogenetic study of Baptistonia 921

MIX: Successive parsimony method according to Wagner and to Camin-Sokal—set of 1000 replica-

tions

CONSENSE: Oncidium altissimum set as tree root.

Neither the introduction of a threshold within the algorithm nor the random selection of the species had

any significant impact on the results.

Distance analysis. —For the second approach, I have used the software ADE-4 (see Thioulouse et al. 1997)

to calculate a distance matrix on the basis of the data matrix given in Table 1 by means of the Manhattan

method as performed by the DMAUtil-Quantitative Variables utility, after which the minimal length tree was

computed by means of the UPGMA method as provided by the NGStat-Minimal Spanning Tree utility.

RESULTS

In respect to the two non-strict consensus phylogenetic trees obtained with the parsimony method (one for

the Wagner parsimony and one for the Camin-Sokal parsimony), it can be noted that: (a) as a general rule,

the bootstrap values are weak, some branches being supported by values below 5096; (b) there is a distinct

separation of the genera Gomesa and Baptistonia within the Gomesa clade; (c) there are, among the species

of Baptistonia, 6 small groups of species supported by relatively good bootstrap values (25096), groups that

are referred to as complexes; (d) these diagrams suggest a low level of reliability for the other relationships

(bootstrap «4096).

I have attempted to define these

1 $ L:

:u

of taxa was reduced by replacing! the species by the corresponding complexes. The characters attributed to

even further in a second step. For this purpose, the number

any given complex were reconstructed on the basis of the characters of the constituent species as well as on

the basis of the evolutionary analysis resulting out of these first diagrams. The topology of the initial trees

is respected and the relationships among the different parts are consolidated (higher bootstrap values).

Figure 1 represents the semi-strict consensus tree obtained by the combined operations described

above. The bootstrap values for the complexes are those obtained in the first step using the Camin-Sokal

method, and the bootstrap values of the higher groupings are those obtained in the second step, using the

Wagner method. They are labeled above the branches and relate to the node situated to the immediate right.

The nodes supported by bootstrap values below 4096 are not reproduced. The numbers related to a branch

by a dotted line represent the synamorphies of the node ending that branch.

Even though, as is usual when using a large data matrix, the consistency index remains low (0.29), the

retention index is relatively high (0.78). This suggests that, in spite of the fact that there is a great deal of

homoplasy in respect to the chosen characters, the synapomorphies are rather numerous (cf. Freudenstein

& Rasmussen 1999).

Figure 2 represents the distance tree obtained as described above. Its overall topology is close to that of

the tree shown in Figure 1: (a) there is a large distance between the Oncidium group and the Gomesa group;

(b) there is a great distance between these two groups and the rest of the species included in the study; (c)

mnlos

LS

among the latter, we find the group “GMT” as well as th brieniana, silvana, pubes, leinigii, pulchella

and truncata shown on Figure 1 (for all the specific names of Baptistonia, the taxonomic authorities are given

in Appendix 1); (d) Oncidium trulliferum and Carriella colorata are placed at the base of the genus Baptistonia.

The following discussion is based on the examination of Figure 1, but is overall compatible with Figure 2.

DISCUSSION

The genera Baptistonia and Gomesa share a number of characters which are not found in the members of

the genus Oncidium. However, the present study also stresses the numerous synapomorphies shown by the

members of each of the two genera.

1A | H £ J " n E J 3 | D £f

The consolidation of

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922

Tague 1. Evaluated states for the different characters of species studied.

sisuapueifou 'g

ejeupind g

seqnd *g

Insqed 'g

epniu ‘g

lozjal|'g

"brul “g

Vorsyney *g

eueipunojgnb *g

eyeuitpe 'g

louarewep 'g

eyennp ‘g

e1ofiuJ05 *g

eueiuauq 'g

Iouiqje *g

ENEC CON MS O UST CON OE SONISCN

ONE CG O Sere cn C

NI SS ONS USE

NTE ON M e e

CM ONG e > Ecce

SS Rr

CN ea NIS iA E

AECA RON er E

A e A ES BE a cie

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NON SS SS E SS r

CN EON er a ee a (N

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Chiron, Phylogenetic study of Baptistonia

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==

47

924 Journal of the Botanical R h Institute of Texas 1(2)

B. brieniana

B. albinoi

B. riograndensis

B. pubes

B. lietzei

14,18,24,26 | B. silvana

27,30,33 B. gutfreundiana

B. cruciata

B. pabstii

B. velteniana

i

I

I

I

I

I

I

I

I

I

B. damacenoi l=

l

l

|

l

l

|

|

|

I

l

I

|

B. leinigii

Baptistonia

B. cornigera

! 01,23,24

60 30,32,33

echinata

pulchella

uhlii

05,23,24,37

widgrenii

nitida

kautskyi

B.

B.

B.

B.

MEM B. sarcodes

B.

B.

B.

truncata

:

07,11,16,25,28,

>

O. trulliferum

—— 02,03,04,21,39

C. colorata

G. recurva

l fo G. alpina

09,10,12,13,15,16,17 i —Lr G. crispa

22,24,33,36,39,45 GL sessili

Gomesa

IC. 02,06,33,36,38 O. baueri

O. nebulosum

groupe

externe

O. altissimum

Cami

l . un

synapomorphies.

Chiron, Phylogenetic study of Baptistonia

55 B. brieniana

13 B. riograndensis

B. albinoi

B. pubes

B. lietzei

B. damacenoi

B. silvana

B. gutfreundiana

B. cruciata

41 88 B. velteniana

61 B. leinigii

20

B. pabstii

B. cornigera

B. nitida

B. widgrenii

B. sarcodes

31

B. echinata

21 B. uhlii

B. pulchella

= B. truncata

100 |

| B. kautskyi

O. trulliferum

C. colorata

G. sessilis

100 | » crispa

G. recurva

G. alpina

O. baueri

O. nebulosum

La altissimum

100

GMI

r:

Fic. 2. Distance tree

J

£L Dat PRA D hi PEA

926 Journal of f Texas 1(2)

Synapomorphies of the genus Gomesa:

* QO—elliptic stomata; * 17—floral bracts very long;

* 10-short inflorescence; * 22-articulated labellum;

* 12—inflorescence not branched; * 24—entire labellum;

* 13—formation of buds as soon as the inflorescence — * 33-callus consisting of two longitudinal crests;

has been generated; * 36-column pressed against the labellum;

* 15—vertical shape of the flowers ovate; * 30—wings almost non-existent

* ]6-flowers without reddish violet spots; * 45—viscidium elliptic.

Synapomorphies of the genus Baptistonia Gwith the e of Carriella colorata and Oncidium trulliferum):

* 03—pseudobulbs faintly flattened; 21-apex of the petals and of the dorsal sepal not

* 04—horizontal section of the pseudobulbsrounded; angular;

* 05—pseudobulbs and leaves dark green; * 37-column pubescent;

* 06—basal sheaths non-leafy; * 39—column wings much longer than wide;

* 08—density of stomata low; * 46—tegula spoon-shaped

* 47-margin of the clinandrium well-developed.

Other synapomorphies may also be relevant, but with reversals in certain taxa:

* 02—pseudobulbs cigar-shaped (except in B. uhlii, B. | * 42-appendix of the anther long (except in B. sar-

truncata, B. nitida and B. widgrenii); codes, B. nitida and B. widgrenii);

* |9—dorsal sepal cucullate (except in B. uhlii, B. * 44—appendix recurved (except in B. nitida and B.

widgrenii, B. albinoi and B. riograndensis); sarcodes).

* 23-clawed lip (except in the echinata complex);

Finally, we may note the evolutionary steps shown by the two genera in relation to the genus Oncidium.

However, it is not possible to consider them as synapomorphies of the Gomesa clade. To do so it would be

necessary to ascertain whether the other groups that are part of this clade would share these synapomor-

phies, a step that would go beyond the scope of this study:

* 02—elongation of the pseudobulbs; * 36-diminuation of the angle between the column

* 06—shortening of the basal sheaths; and the labellum;

* 33—modification of the median part of the callus; — * 38—disappearance of the tabula infrastigmatica.

Oncidium trulliferum shares 10 of the 16 synapomorphies of the genus Baptistonia and seems to be integrated

into this genus. This grouping, which could possibly be designated as Baptistonia sensu lato, is supported by

a bootstrap of 5896. This value is fairly low and is caused by the fact that Carriella colorata is found on some

of the most parsimonious trees together with one or another group of species belonging to Baptistonia sensu

lato. Carriella colorata is placed within the phylogenetic reconstruction as the sister species of this group,

the whole being supported by a bootstrap value of 9696.

I will now review the six complexes mentioned above:

The “truncata” complex [B. truncata and B. kautskyil: bootstrap value 99%. This complex is defined by five

synapomorphies that are found only in this group:

* 16-ground color of the flower is pale;

* 244+25—the lateral lobes of the labellum are short and directed toward the apex of the labellum;

* 30-the isthmus of the labellum is short with triangular sinuses;

* 40—the wings of the column are spread at 180?

Other characters are shared with some of the other groups:

* Ol-pseudobulbs not very tall (shared with the echinata complex, with B. widgrenii and with Carriella

colorata);

* qe

07—pseudobulbs unifoliate (shared with the pulchella complex and with Carriella colorata);

* 11-inflorescence growing downward (shared with B. pulchella and B. uhlii);

Chiron, Phylogenetic study of Baptistonia 927

* 28—width of labellum midlobe equal to its length (shared with the brieniana complex);

* 33—median part of the callus made of a thin, smooth, inverted V-shaped plate (shared with the brieniana

complex, with the pubes complex and with B. cruciata);

* 34—callus not extending onto the midlobe (shared with B. cornigera and with B. sarcodes).

According to my analysis, this complex is at the basis of the genus Baptistonia, and is separated from the other

members by a branch that is relatively well supported (6096) and defined by two synapomorphies:

* 41— stigmatic cavity with more or less long hairs;

* 43—appendix of the anther bifid or double.

The “silvana” complex [B. silvana and B. gutfreundianal: 8696. This group is defined by a combination of

shared characters each of which can also be found in other species:

* 18-dorsal sepal spathulate (shared with the pubes complex and with B. nitida);

* 24—lateral lobes of the labellum much longer than wide (shared with the pubes and brieniana complexes

as well as with three other species);

* 26—base of the lateral lobes distinctly folded backward (shared with the pubes and pulchella complexes,

and with B. cruciata);

* 27-labellum midlobe wider than the width of the labellum as measured over the lateral lobes (shared

with B. nitida);

* 30-isthmus of the labellum short and wide with rounded sinuses (shared with B. sarcodes, B. widgrenii

and with the echinata complex);

* 33—median part of the callus consisting of a thick, non-verrucose plate that is indented at the center

(shared with B. cornigera).

The “pulchella” complex [B. pulchella and B. uhlii]: 81%. One synapomorphy:

* 43—appendix of the anther is bifid;

and shared characters, also found in other species:

* O7—pseudobulbs unifoliate (shared with the truncata complex and with Carriella colorata);

* |O-inflorescence short (shared with B. truncata and with Carriella colorata);

* ]l-inflorescence growing downward (shared with the truncata complex);

* 26—base of the lateral lobes of the lip distinctly folded backward (shared with the silvana and pubes com-

plexes, and with B. cruciata);

B. echinata is positioned as a sister species to this complex (bootstrap of 4796) with one synapomorphy:

* 33—median part of the callus made up of two low but wide crests;

and several shared characters, that are, however, also shared with other species:

* Ol-pseudobulbs not very tall (shared with the truncata complex, with B. widgrenii and with Carriella

colorata);

* 23-claw very short (shared with B. sarcodes, Carriella colorata and Oncidium trulliferum),

* 24—width of the lateral lobes of the lip equal to their length (also found in B. sarcodes and Oncidium trul-

liferum);

* 30-isthmus short and wide with rounded sinuses (shared with B. sarcodes, B. widgrenii and the silvana

complex);

* 32—basal part of the callus without teeth.

It should be noted, however, that, in respect to the distances, B. echinata is the closest neighbor of B. sarcodes,

and this pair of species is found to be a sister group of the pulchella complex.

The “brieniana” complex [B. brieniana, B. riograndensis and B. albinoil: 79%. This group also ha

synapomorphies; it is defined by the following combination of characters:

* 09-circular stomata;

* 20—petals spathulate to panduriform (shared with B. widgrenii and the pubescomplex);

* 28—width of the labellum midlobe equal to its length (shared by the truncata complex);

928 Journal of the Botanical R h Institute of Texas 1(2)

* 30-isthmus of the labellum long and rectangular (shared with the pubes complex as well as with B. nitida

and B. cornigera);

* 33—median part of the callus made up of a thin, smooth, inverted V-shaped plate (shared with the pubes

complex, with the truncata complex and with B. cruciata);

* 54—apical part of the callus long (shared with the pubes complex and with B. widgrenii).

The “leinigii” complex [B. leinigii, B. pabstii and B. veltenianal: 78%. This group is characterized by two

synapomorphies:

* 30-isthmus trapezoidal

* 33—median part of the callus made up of a verrucose, sheeted plate.

The “pubes” complex [B. pubes, B. lietzei and B. damacenoil: 7496. Here again, the group is characterized

only by shared characters also found in other species:

* 20—petals spathulate to panduriform (shared with B. widgrenii, Oncidium trulliferum and with the brieniana

complex);

* 30-isthmus of the labellum long and rectangular (shared with the brieniana complex as well as with B.

nitida and B. cornigera);

* 33-median part of the callus consisting of a thin, smooth, inverted V-shaped plate (shared with the

brieniana and truncata complexes and with B. cruciata);

* 34—apical part of the callus long (shared with the brieniana complex and with B. widgrenii).

In my reconstruction, the pubes and silvana complexes are sister groups, although with a weak bootstrap

value (4496). The two groups are also close to each other in the distance tree. The four synapomorphies that

characterize this clade are all characters of low importance:

* 14—flower form;

* 18—dorsal sepal spathulate (shared with the silvana complex, as well as with B. nitida, Oncidium trulliferum

and Carriella colorata);

* 24—form of the lateral lobes of the labellum;

* 26—base of the lateral lobes distinctly folded backward (shared with the silvana and pulchella complexes,

and with B. cruciata).

Together, the pubes, brieniana, silvana, and leinigii complexes form a monophyletic group, here referred to as

« GMI ». This group, although weakly supported (bootstrap value: 4296), is characterized by the following

synapomorphies:

* 23-nail of the labellum long;

* 32—basal part of the callus ending in a pair of diverging teeth (shared with B. echinata).

Baptistonia cruciata is included in « GM1 » but the analyses did not allow a precise clarification of its position.

Baptistonia cornigera appears as a sister species of this GM1 group (bootstrap of 46%).

Finally, it should be noted that, on the initial consensus tree, B. widgrenii appears as a sister species to the

==

entity formed by B. echinata and the « pulchella » complex, with a very weak bootstrap value (26%) and the

shared characters 01 and 30. However, this grouping, besides not supported, would probably be artificial

because (a) the structure of the pseudobulbs is very different although the pseudobulbs of B. widgrenii are

short, and, like those of the other species in this group, much shorter than the pseudobulbs of the other

Baptistonia species; and (b) the lateral lobes of the labellum are very different although the isthmus is the

same. Furthermore, B. widgrenii does not share any real synapomorphies with this group. Therefore, the

parental relationships of B. widgrenii remain to be clarified.

CONCLUSIONS

The present study supports monophyly of Baptistonia as defined by Chiron and Castro Neto (2004) as well

as its distinctiveness from Gomesa based on numerous morphological and anatomical characters.

Chiron, Phylogenetic study of Baptistonia 929

The proper place of the two species added to the bulk of the specimens to be studied, Carriella colorata

and Oncidium trulliferum, is difficult to define on the basis of this study alone. Oncidium trulliferum is posi-

tioned at the base of the genus Baptistonia and should probably be integrated into that genus (bootstrap of

58%). Carriella colorata should rather be regarded as a sister species to the genus. It is, however, better to

wait for the results of molecular studies in progress that may clarify this issue.

The phylogenetic relationships within the genus Baptistonia are poorly resolved due to the low number

of synapomorphies defining the complexes and to homoplasy. However, some monophyletic groups can be

distinguished.

Baptistonia kautskyi and B. truncata are strongly supported as sister taxa, and are sister to all other spe-

cies of Baptistonia.

The majority of the complexes defined above have been discussed in detail by Chiron and Castro Neto

(2005a, 2005b, 2006a, 2006b). One could argue that each of these complexes represents nothing but the

expression of the existence of different populations of the same, rather variable species. As many of the

combinations of morphological characters that distinguish the members of a complex as well as their dis-

tribution zones do not form a continuum but rather disjointed conglomerates, I come, however, to another

conclusion.

APPENDIX 1: SPECIMENS EXAMINED

Living material (Vouchers have been deposited in LY (and/or SP when noted) for all plants).

Baptistonia albinoi (Schltr) Chiron & V.P. Castro: Chiron 2578, Chiron 0084, Chiron 0085, Chiron 0086. B. brieniana (Rchb.f)

V.P. Castro & Chiron: Chiron 04604, Castro Neto s.n., Paraguay s.n. B. cornigera (Lindl.) Chiron & V.P. Castro: Chiron 03017, Chiron

03035, Chiron 02549, Chiron 03069, Chiron 03008, Chiron 03066, Chiron 05206, Chiron 06576, Chiron 07075, Chiron 3051, Chiron

3052, Chiron 3062. B. cruciata (Rchb.f) V.P. Castro & Chiron: Chiron 05441, Chiron 05446, Chiron 05452, Zézé s.n., Uhl s.n., Jardin

botanique de Lyon 020539. B. damacenoi Chiron & V.P. Castro: Chiron 03208, Chiron 2273, Chiron 2589 (type, SP), Frey746. B.

echinata Barb. Rodr.: AOSP s.n. ex Chiron 0042, AOSP s.n. ex Chiron 0043, Jardin botanique de Sáo Paulo s.n. ex Chiron 0065 and

Chiron 0066, Chiron 2582, Chiron 3044, Chiron 3045, Chiron 3055, Chiron 3063. B. gutfreundiana (Chiron & V.P. Castro) Chiron

& V.P. Castro: Chiron 2240, Chiron 03414, Chiron 05844, Chiron 05848, Castro Neto s.n. ex Chiron 0058, Chiron 3060, Castro Neto

s.n. (type, SP). B. kautskyi (Pabst) V.P. Castro & Chiron: Chiron 03194, Chiron 04813, Chiron 04814, Chiron 04816, Chiron 05769,

Chiron 06503, Frey 741, Frey 1079, Caliman s.n. B. leinigii (Pabst) V.P. Castro & Chiron: Chiron 0101, Chiron 07009, Chiron 07011

Chiron 07012, Chiron 07015, Chiron 07017, Chiron 07018, Chiron 07019. B. lietzei (Regel) Chiron & V.P. Castro: AOSP s.n. ex Chiron

0041, AOSP s.n. ex Chiron 0044, AOSP s.n. ex Chiron 0045, Castro Neto s.n. ex Chiron 0046, Chiron 06504, Chiron 06505, Chiron 2273,

Chiron 2283, Chiron 07026, Campacci s.n. B. nitida (Barb. Rodr.) V.P. Castro & Chiron: Chiron 2591, Castro Neto s.n. ex Chiron 0039,

Castro Neto s.n. ex Chiron 0040, pou 2592, Chiron e Chiron 2596, Chiron 05532, Chiron 06540. B. pabstii (Campacci & C.

Espejo) V.P. Castro & ore Chiron 3042, Chiron 3059. B. pubes (Lindl.) Chiron & V.P. Castro: Chiron 3034, Chiron 3036, Chiron

3040, Chiron 3046, Chiron 3047, Chiron 3048, Chiron 3053, Chiron 3054, Chiron 06506, Chiron 2241, Castro Neto s.n. B. pulchella

(Regel) Chiron & V.P. Castro: Chiron 05466, Chiron 05487, Chiron 05490, Chiron 05498. B. riograndensis (Cogn.) V.P. Castro &

Chiron: Chiron 07060, Chiron 07061, Chiron 07062, Chiron 07063, Chiron 07069, i oe B. oe. leg) Chiron & V.P.

Castro: Chiron 04850, Chiron 04852, Chiron 05218, Chiron 06570, Chiron 2242, Chiron li

silvana (V.P. Castro & Campacci) V.P. Castro & Chiron: Chiron 2890, Chiron 3049, das os Chiron 05600 Chiron 05807, Chiron

06509, Castro Neto s.n. ex Chiron 0057. B. truncata (Pabst) Chiron & V.P. Castro: Chiron 05427, Chiron 2785, Chiron 2262, Chiron

2264, Chiron 2769, Chiron 2899A, Chiron 2899B, Chiron 2899C. B. uhlii Chiron ave dit Chiron Bur fees SP), Chiron 06511,

Chiron 06512, Chiron 06534. B. velteniana V.P. Castro & Chiron: Kautskyi s.n. (type, SP), Uhl s.n. ex Chiron 0028 and Chiron 0029. B.

widgrenii (Lindl.) V.P. Castro & Chiron: Chiron 0112, Chiron 2243, Chiron Fn Chiron ne Chiron o oli colorata

(Kóniger & J.G. Weinm.) V.P. Castro €: K.G. Lacerda: Caliman s.n., Castro Neto s.n., Chiron 2796, Chiron 06508. Gomesa alpina

Porsch: Jardin botanique de Sáo Paulo s.n. (SP), Chiron 05346, Alambari s.n. G. crispa Klot s ex ee Chiron ne Chiron

05481, Jardin botanique de São Paulo s.n. (SP). G. recurva R. Br.: Chiron um Chiron 04683, Chiron 04702, Chiron 04810, Chiron

04811, Jardin botanique de Sáo Paulo s.n. (SP). G. sessilis Barb. Rodr.: Chiron 04695, Chiron 05388, Chiron 05398, Chiron 05702,

Chiron 05715. O. altissimum Swartz: Chiron 0834, Chiron 1919, Jardin botanique de Lyon s.n. Oncidium baueri Lindl.: Chiron

02041. O. nebulosum Lindl.: Chiron 99041. O. trulliferum Lindl.: Jardin botanique de São Paulo 7028 (SP), Jardin botanique de

Sáo Paulo 11008, Chiron 05249, Chiron 0107, Chiron 0108, Chiron 0109.

D, IN A40? .B.

UUIO AT

a

Herbarium material

Baptistonia albinoi: MBM 209548, MBM 99608, MBM 6225, MBM 48898, MBM 48896, MBM 29392, MBM 22869, SP 25185,

SP 31577, W-R 35126. B. brieniana: W-R 16453 (type de Oncidium brienianum), W-R 25704, W-R 7031, P 00430186 (type de

Oncidium verrucosissimum). B. echinata: W-R 10660, W-R 20585, W-R 20586 (type de Oncidium brunleesianum). B. lietzei: LE

£L Dag PRA D hi PEA

930 Journal of f Texas 1(2)

s.n. (type de Oncidium lietzei), W-R 25721 (type de Oncidium hrubyanum). B. leinigii: SP 334938, SP 341393. B. pubes: W-R

13085, W-R 28099, W-R 25713. B. pulchella: W-R 28923. B. riograndensis: ICN-Dutra 925, SP 363181 (type de Oncidium

cassolanum). B. sarcodes: K-Lindley no.84 iud de Oncidium sarcodes), MBM 10148, MBM 218812, W-R 44709, WU s.n. B.

widgrenii: K- inue n?50 (type de Oncidium widgrenii), SP 46502, W-R 35291, W-R 10423. Gomesa crispa: W-R 45401, xm R

45361. G. re a: W-R 34429, W-R 14023, W-R 11818. Oncidium trulliferum: MBM 7758, MBM 10717, MBM 48903, MBM

226953, W-R pisce R 6817, W-R 3319.

=

ACKNOWLEDGMENTS

The author is indebted to Philippe Choler, Université Joseph Fourier—Grenoble I, for his cooperation in

the computation of the phylogenetic tree in figure 2; to Guido Braem, University of Maryland, University

College Europe, for his helpful discussions; and to Mark Whitten and Norris Williams for providing critical

comments on the manuscript. The study of the herbarium materials was made possible by the hospitality

demonstrated by the curators of ICN, K, LE, MBM, P, SP, W, and WU to whom I extend my sincere gratitude.

My stay at Vienna was supported by the SYNTHESYS Grant AT-TAF-2736. Access to live specimens was

made possible through the collaboration of my Brazilian friend Vitorino P. Castro Neto and through some

of his local contacts, especially: Carlos Régent, Sergio Gutfreund, Sydney Marcal, and Thomas Adamski. I

am deeply indebted to each and every one of them.

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932 Journal of the Botanical R h Institute of Texas 1(2)

BOOK NOTICES

Top E Stuessy, VERONIKA Mayer, and Elvira HORANDL (eds). 2003. Deep Morphology: Toward a Renaissance

of Morphology in Plant Systematics. (ISBN 3-906166-07-4, hbk.). Regnum Vegetabile: Vol. 141.

Gantner Verlag, Liechtenstein. (Orders: Koeltz Scientific Books, D-61453 Koenigstein, PO. Box 1360,

Germany; koeltz@t-online.de; www.koeltz.com). $156.00, 326 pages, illus. ca. illus., 6 1/4" x 9 1/4".

After an introduction (Chapter 1), the chapters are divided into three sections: (2-5) Genetics and Development, (6-8) Phylogenetic

Analysis, (9-13) Ecology and Adaptation, with a final overview (Chapter 14).

Table of Contents

1 MA 7] E T

I l g and why is it time for i i i dd ics? (Weber).

2.Th SE : TE RE T6 fd ES " nd z Gailing).

3. A m effects on floral M and function: a revie (Digg le).

4. Floral iR AM features ane med na in US M ag and p

5C l hyl

6. o and ee evolution. (Williams and di

7. What should a *complete" pasion Ee ds a dd

8. Beyond morphoclines and trends: f (Hufford and McMahon).

and Neinhui

10. Toward a deeper und li d df ion i llen grains: the sporoderm (Hesse).

11. Ecological adaptions and deep phylogenetic splits—evidence and ens n the Nn xylem (Baas, Jansen and Wheeler).

12. The potential of plant biomechanics in functional biology and systematics (Speck, Rowe, Civeyrel, ClaBen- Bockhoff, Neinhuis,

and Spatz).

9. Epicuticular waxes and vascular huh systematics: integrating micr NOTOS ood and TE data (Bart Theisen, Borsch

s).

13. How a better understanding of adaptions can yield better use of morphology in plant systematics: toward Eco-Evo-Devo

(Givnish).

14. Morphological data in pl ics (S

STEVEN R. RADOSEVICH, JODIE S. Hot, and CLaubio M. Guersa. 2007. Ecology of Weeds and Invasive Plants:

Relationship to Agriculture and Natural Resource Management (ed. 3). (ISBN 978-0-471-76779-4,

hbk.). John Wiley & Sons Inc., One Wiley Drive, Somerset, NJ 08875, U.S.A. (Orders: www.wiley.com,

877-762-2974, 1-800-597-3299 fax). $74.95, 454 pp., b/w photos, line drawings, 6 1/4" x 9 1/4".

Table of Contents

Weeds and Invasive Plants (definitions and classificati i ducti d wildl

F W ildlands).

. Principles of Weed and Invasive Plant Ecology.

Invasibility of Agricultural and Natural Ecosystems.

Evolution of Weeds and Invasive Plants.

Weed Demography and Population Dynamics.

Plant-Plant Association

Weed and Invasive Plant Management Approaches, Methods and Tools.

Herbicides

Systems Approaches for Weed and Invasive Plant Management.

e UU UR ev

J. Bot. Res. Inst. Texas 1(2): 932. 2007

MUHLENBERGIINAE (POACEAE: CHLORIDOIDEAE: E YNODONTEAE):

FROM NORTHEASTERN MÉXICO

Paul M. Peterson Jesus Valdés-Reyna

Department of Botany e de Botánica

National Museum of Natural History Univerisid ma Agraria “Antonio Narro”

mithsonian Institution Buenavista, Saltillo

diis DC 20013-7012, U.S.A. Coahuila 25315, MÉXICO

eterson@si.edu jvaldeseuaaan.mx

Yolanda Herrera Arrieta

Instituto Politécnico Nacional

CIIDIR Unidad Durango-COFAA

Durango 34220, MÉXICO

yherreragipn.mx

ABSTRACT

A taxonomic treatment of the subtribe ardid for northeastern México (Coahuila, Nuevo León, and Tamaulipas) is given. A

total o dy area in four genera: Aegopogon (1), Blepharoneuron (1), Lycurus Q), and Muhlenbergia

(35). The names, aa medía var. ramosa ees and Muhlenbergia spiciformis Trin., are lectotypified. A key for piacere

the ee x distributions illustrations, synonymies, and a

E

n M és 3

o

RESUMEN

Se presenta el estudio taxonómico la subtribu Muhlenbergiinae para el noreste de México (Coahuila, Nuevo León y Tamaulipas). En

total se reconocen 39 ti n el área de estudio pertenecientes a cuatro géneros: Aegopogon (1), Blepharoneuron (1), Lycurus

(2), y Muhlenbergia (35). n nombres, Muhlenbergia glomerata var. ramosa Vasey y Muhlenbergia spiciformis Trin., son lectotipificados.

Se incluyen una clave para determinar las especies así como descripciones, distribuciones, especimenes examinados, ilustraciones y

sinonimia para cada especie. Se incluye una breve discusión indicando las relaciones entre todas las especies de Muhlenbergia para el

noreste de México.

Coahuila, Nuevo León, and Tamaulipas or northeastern México covers an area of 291,955 km? or 15 96 of

the total land of México. This area includes portions of two natural regions known as the Chihuahuan and

Tamaulipan Deserts. These regions are considered a center of origin and diversification of arid and semi-arid

plant species (Davila-Aranda et. al. 2004; Peterson et al. 2007). As part of the current revision of the grass

flora of northeastern México, an examination of the taxonomy and distribution of the species of subtribe

Muhlenbergiinae, was begun to aid the agriculture and livestock industries. This study treats 39 species

currently placed in four genera: Aegopogon (1 sp.), Blepharoneuron (1 sp.); Lycurus (2 sppJ, and Muhlenbergia

(OD spp.).

ae Muhlenbergiinae is characterized by having spikelets perfect, staminate or sterile; occasionally

with cleistogenes in the leaf sheaths; inflorescence paniculate of spicate main branches or a single raceme;

spikelet bearing axis disarticulates falling entire or is persistent; spikelets solitary, paired, or in triplets,

occasionally second, 1—3(-6)-flowered; glumes awned or unawned, 1-4-nerved; lemmas 3-nerved (rarely

]-nerved), awned or unawned; and a base chromosome number of x = 8-10. An ongoing phylogenetic study

of the Muhlenbergiinae based on nuclear and chloroplast DNA sequences is currently in progress (Columbus

& Peterson, in prep). Ina preliminary analysis, Muhlenbergia appears to be paraphyletic since the remaining

nine genera in this subtribe are nested within its clade (Peterson et al. 2001a; 2004; 2007).

Subtribe Muhlenbergiinae (Duvall et al. 1994; Peterson 2000; Peterson et al. 1995, 1997, 2001b, 2007)

currently consists of 10 genera: Aegopogon with four species; Bealia mexicana Scribn., a monotypic genus

J. Bot. Res. Inst. Texas 1(2): 933 — 1000. 2007

934 Journal of the Botanical R h Institute of Texas 1(2)

from northern Mexico (Peterson 1989; Peterson et al. 1993); Blepharoneuron with two species (Peterson &

Annable 1990, 2003); Chaboissaea with four species, three from central México and C. atacamensis (Parodi)

P.M. Peterson € Annable from Argentina and Bolivia (Peterson & Annable 1992; Peterson & Herrera Ar-

rieta 1995; Sykes et al. 1997); Lycurus with three species, including the amphitropical disjuncts, L. setosus

(Peterson & Morrone 1997); Muhlenbergia with 147 species centered in northern Mexico and the southwest-

ern United States (Herrera Arrieta & Peterson 2007; Peterson 2000, 2003), containing the important range

grass M. montana and the amphitropical disjuncts M. arenicola and M. torreyi (Peterson & Ortíz-Diaz 1997);

Pereilema with four species, two from western, central, and southern México; Redfieldia flexuosa (Thurb.

ex A. Gray) Vasey, a monotypic genus and southwestern United States endemic of probable hybrid origin;

Schaffnerella gracilis (Benth.) Nash, a monotypic genus known only from San Luis Potosí, México (Columbus

et al. 2002); and Schedonnardus paniculatus (Nutt.) Trel., a monotypic genus and amphitropical disjunct with

spicate primary branches.

As currently understood for the New World Chloridoideae, tribes Cynodonteae and Zoysieae are sister,

and sister to this clade are members of the Eragrostideae (Columbus et al. 2007; Peterson et al. 2007). All

three tribes are supported by DNA sequence data from the chloroplast (matK, trnL—F region) and nuclear

(ITS) genomes (Hilu & Alice 2000, 2001; Columbus et al. 2007). Relationships among the ten subtribes

within Cynodonteae, as well as the 25 unplaced genera, are uncertain pending additional study. At this

point we have no clear idea as to the relationships among these ten Cynodonteae subtribes. However, we

do have good molecular support for maintaining the tribe Cynodonteae and morphological support for all

of the ten subtribes.

The following | contains a key for determining the species, descriptions, distribution,

specimens d, illustrations, synonymies, and a brief discussion indicating hypothesized relationships

among all native and adventive species of Muhlenbergiinae in northeastern México. This study is based

on the examination of herbarium specimens from ANSM, ARIZ, B, BA, BAA, BM, CIDIIR, COCA, ENCB,

GH, HINT (George S. Hinton's personal herbarium), IBUG, INEGI, K, LE, MEXU, MICH, MO, MSC, NMC,

NMSU, NY, P, RSA, SL SEPM, TAES, TEX, UAT, US UT, UTC VT, and W; including the type specimens

of most of the species studied.

KEY TO THE SPECIES OF MUHLENBERGIINAE IN NORTHEASTERN MÉXICO

1. Plants annual.

2. Lemmas unawned

3. Ligules with lateral lobes extended into auricles; leaf blades with margins and midveins whitish-

es basally, ia mu panicle branches diverging 80-1009 from the rachises; pedicels

m long, gl throughout or obscurely puberulent, the hairs ipd m lon

Muhlenbergia fragilis

Be ees ae Yout | Ho! tended int da NI. ith | | lol longer than the body;

fl i i le branches diverging 25- 80°

from the rachises; pedicels 2-7 mm long; ee a near Ks apex, the hairs 0.1-0.3 mm long

19. Muhlenbergia minutissima

DE s awned, the awns 3-40 mm lon

4. Spikelets in clusters of three, the 20 spikelet perfect and the two lateral spikelets ano and

staminate or sterile; lemmas usually with lateral nerves extending into aw . Aegopogon tenellus

4. DET. not in clusters of three, or if in clusters then in pairs; lemmas usum lateral n tendi

into a

5. ne glumes 2-nerved, minutely to deeply bifid, the teeth aristate or with awns to 1.3 mm long;

e often in sessile-pedicellate pairs; disarticulation at the base of the pedicels; MS with

s 6-15 mm long, sti 8. Muhlenbergia depauperata

5 p n 1 Da or eos [IDE eu MED. ot with a single mucro or awn; spikelets

| inal -40 mm long, delicate straight

Ung

to flexuous.

6. Glumes 1.2-2.8 mm long, acute to acuminate; upper glumes 1.5-2.8 mm long

__ 34. Muhlenbergia

tenuifolia

Peterson et al., Muhlenl ii f tl t M

935

6. Glumes 0.3-1.3 mm long, obtuse to acute; upper glumes 0.5-

7. Cleistogamous panicles with 1

open and not densely flowered, 1

the rachises; ligules truncate to o

1.3 mm long.

-3 pus present in the axils of the lower sheaths; puc

m wide; primary branches ir en to

hle naa microsperma

7. Cleistogamous panicles absent in i axils of the lower sheaths; panicles narrow, contrac

0.6-2.8 cm wide; primary branches spreading up to 30? from the rachises; ligul

acu

32 DUREE spiciformis

. Plants perennial.

8. Plants rhizomatous, rhizomes oft

9. Upper g

stout, scaly, and creepin

g.

lumes always 3-nerved, the apex usually with 2-4 small teeth; old sheaths flattened, ribbon-like

or papery, sometimes spirally coiled

lumes greenish-plumbeous, the surface scabrous, usually with a few short hairs below; pan-

icles 0.5-2 cm wide, the primary branches appressed to spreading up to un from the ra

Muhlenbergia quadridentata

10. Glumes whitish, stramineous or SUE green sometimes with EE M the surface

shiny, usually glabrous to rarel belo OW; panicles 0.5-6 cm wide, the branches appressed

to spreading up to 45? from the rachises Muhlenbergia virescens

per glumes edil ]-nerved or sometimes 2- or 3-nerved, but the apex never ms old sheaths

MAN flattened

11. T the awns 1-20(-25) mm

12. Lemma awns 0.1-3(-5) mm long, straight; eaei ein the internodes

— 13. Muhlenbergia

glauca

12. Lemma awns 10-20(-25) mm long, flexuous; leaf sheaths mostly shorter than internodes

21. Muhlenbergia polycaulis

11. Lemmas unawned or mucronate, the mucros to 1 mm long; anthers yellow, purple, greenish or

reddish.

13. Panicles narrow Ic Open 4- lo cm wide.

Panic le bran

to 40° fi

red, | not capillary, ightly appressed or loosely spreading up

|

UVES VLG

and along the margins on proximal 2/3 to % 15. Muhlenbergia jaime-hintonii

14. Panicle branches capillary, diverging 30-90(-100)* from the rachises at maturity; paleas

glabrous.

1

5. Ligules 0.5-2 mm long, hyaline, with well-developed lateral lobes or P bla

margins and midveins prominent, whitish, thicken aa arenacea

15. Ligules 0.2-1 mm long, ciliate, without lateral lobes or ‘or auricles; sera margins an

idveins not conspicuously thickened, greenish

13. Panicles contracted, narrow, 0.1-3.9 cm

16. Spikelets 1.4-2.4 mm lon

17. Culms 44-82 cm Ms leaf blades 5-22 cm long, 1

long 15. Muhlenbergia jaime-hintonii

17. Culms 4-30 cm tall; leaf blades 0.2-1.8 mm wide; panicles 1-5 cm long

18. Lemmas and paleas glabrous or with minute appressed pubescence, the hairs

about 0.1 mm long; leaf sheaths 0.3-2.4 cm long; panicles partially included i

the upper sheaths; ligule apices truncate Muhlenbergia utilis

18. Lemmas and paleas densely villous, the hairs less 0.4-1 mm long; leaf sheaths

15 cm long; panicles usually on an exserted peduncle; ligules apices acute

37. Muhlenbergia villiflora var. villiflora

7. Muhlenbergia asperifolia

wide

8-5 mm wide; panicles 13-34 cm

16. Spikelets 2.4-8 mm long.

19. Glumes 3-8 mm long anang the awns), 1.3-2 times longer than us lemma; lea

blades 2- UN vide, fl 0.4-0.8 mm lon

19. Glumes 1

Muhlenbergia racemosa

-36mm Io Y to as long as the lemma; leaf blades 0.5- 6 mm wide, flat

to mou anthers 0.7- ib mm eng

mas with ta 0

m long on the lower 1⁄2 along the midveins

Pa margins; leaf Ps m 42 cm AAA not arcuate; anthers 1.8- ri mm lo

orange

20. Lemmas glabrous or

3. Muhlenbergia glauca

the hairs less than 0.1 mm long; leaf blades 0.4-6 cm long, somewhat arcuate

spreading; anthers 0.7-1.4 mm long, yellow to purplish _ 27. Muhlenbergia repens

936

Journal of the Botanical R h Institute of Texas 1(2)

8. Rhizomes absent

2] T glumes 3-nerved; old sheaths flattened and sometimes spirally twisted near base

No

2. Upper glumes 3-toothed and 3-awned, the teeth (including the awns) 1/3 to Y p en of the

glume, and the awns up to 1.7 mm long pets montana

22. Upper glumes 2-4-toothed, the teeth small and unawned, less than 1/6 T length oft

glume.

23. Glumes a plumbeous, the surface scabrous, usually with a few short hairs below;

panicles 0.5-2 cm wide, the primary branches appressed to spreading up to 30° from the

n 25. Mahlenbergia quadridentata

23. Glumes whitish, stramineous or grayish ith pluml the surface

shiny, usually glabrous below; panicles 0.5-6 cm wide, the branches a: " ps

° from the rachises hlenbergia virescens

. Upper glumes usually 1-nerved (rarely 2- or 3-nerved); old sheaths usually not T or spirally

twisted near base.

24, Panicles wide, open, oy contracted to diffuse, (2-)3-20 cm wide; panicle branches usually

widely spreadir «M LU loosely aov ci dit Y.

25. Lemmas unawned, mucronate (mucros < 1 mm long), or awned, with awns 1-4 mm long.

26. Culms (65-)80-230(-300) cm tall; leaf blades 20-100 cm long, 2-7 mm wide, flat or

ded.

Sos

O

. Leaf sheath auricles absent; ligules 10-25 mm long; lemmas eL along the

midvein and margins on lower 5-34 hlenbergia emersleyi

. Leaf p auricles present (-1)2-4(-10) mm long, linear EE to phe tri-

Susp du above, straight or twisted, firm below; ligules 2-10(-12) mm long;

| pubescent with scattered hairs on lower Y __ 30. Muhlenbergia

busta

N

^

26. Culms 10-70 cm tall; leaf blades 1-16 cm long, 0.3-2.5 mm wide, filiform, involute, flat or

olded.

28 Boma Mende SERE inn no vous meee to ee lemmas ie a

densely villous nes en nerves 2. Blepharoneuron tricholepis

28. Lemmas with awns 0.5-4 mm long; spikelet pedicels erect, not flexuous nodding to

reflexed; lemmas without a thick covering of tawny to shiny silky hairs on the midvein

and margins but with appressed pubescence, the hairs scattered; paleas sparsely

pubescent between the nerves.

29. Leaf blades not arcuate, 1-2.2 mm wide, 4-10(-16) cm long; 1 or more culm nodes

exposed; leaf blades reaching 14—12 of the plant height______ 6. Muhlenbergia arenicola

29. Leaf blades arcuate, 0.3-0.9 mm wide, 1-3(-5) cm long; usually no culm no

exposed; most leaf blades reaching no more than 1/5 of the Ta height

5. Muhlenbergia torreyi

25. Lemmas awned, with awns 5-30(-40) mm long.

30. Plants conspicuously branched, especially from the middle and lower nodes, loosely

caespitose; culms O geniculate, I

31. Plants distinctly bushy with ase;

panicles 6-15 cm wide; anthers 1.5- 25 mm | ng; caryop ceeds compresse ed,

Xo sh brown 22. Muhlenbergia porteri

. Plants not distinctly bushy in appearance; plants short-lived perennials to annuals

with delicate bases; panicles 1-6.5 cm wide; anthers 0.9-1.5 mm ds Car

narrowly fusiform, terete, brownish 34. Müller gs tenuifolia

30. Plants not Sour branched, caespitose; culms erect to rarely slightly decumbent

ar bas

2 n eae heaths compressed-keeled or flattened; glumes longer, as long or a little

dud the lemm

eaf sheath eae present 0.4-2.6 cm long on lower portions and up to 6.4 cm

d above; lemmas glabrous or margins pubescent on the lower 1/3, LE

lower 1/3 with scattered hairs; glumes usually with faint, pios, scattered hairs,

the hairs less en 0. l mm long aa rd

33. Leafsheath aur pul along the midvein and m

on the mon 34; glumes without widely scattered hairs — 11. MUERE UE emersleyi

Peterson eta

| MA ll L H4 £, 41 + M

32. Basal leaf sheaths ; glumes usually much shorter than the lemma.

34. Spikelets 3.5-5.1 mm long; lemmas 3.5-5.1 mm long; paleas ki mm a

35. | | | | not +l Pehia

1-3 mm wide, flat or involute, not falcate; culms usually with la Aes

937

node;

anthers purplis Muhlenbergia rigida

35. Lemmas glabrous, smooth and shining, stramineous; leaf Tn 0.2-1.2

mm wide, tightly involute, falcate; culms usually with 2-4 bon, anther

ers

reenish 31. n E setifolia

34. Uu -)2.4-3.6 mm long; lemmas 2.3-3.6 mm long; paleas (2-)2.4-3.6 m

lon

36. Cums25- n call el MAU ju guid and

m long;

caryopse s1- | 2mm long, fusiform to ovoid __

36. Culms 74-150 cm tall; leaf blades 20-65 cm long, r not aen lemmas

green and glabrous; an glabrous between the nerves; ur

mm longi ONORE I -1.7 mm long, fusiform___

24. Panicles narrow, contract ted, and or spikelike 0.2-3 cm wide; m branches

uoud

lly CIU CU UU loosely spreadit |

34. cen Pr 0.2-1.2 mm wide, densely cios sometimes interrupted near base;

AM branches tightly appressed, the branches 0.1 m lon

. Culms (3550-200 cm tall; panicles 15-60 cm E e rounded near base.

39. Lemmas awned, the awns 10-20 mm long; leaf blades 0.5-1.5 mm wide, tightly

involute, falcate, and densely pubescent above; glumes puberulent on upper 2/3

14.

39. Lemmas mucronate, the mucro less than 1 mm long; leaf blades 1.5-6 mm wide, flat

or involute, never falcate, and without hairs above; glumes without hairs, scabrous.

40. Glumes 3.4-5.6 mm long, usually longer than the lemma, apex unawned and

not mucronate; ligules (5-)8-40(-50) mm long; anthers 1.5-2. a pls greenish

40. Glumes 1.8-3.2 mm long, shorter than the lemma, vae ind mucronate

or short-awned, the awn up to 1.7 mm long; ligules 0.5-2(-3) mm long; anthe

ER purpusii

dark

MS e stricta

Muhlenbergia gypsophila

Muhlenbergia macroura

-1.8 yellow to purple 28. aos rigens

09)

CO

. Culms 15-60 cm tall; panicles (2-)4-16 cm long; leaf sheaths compressed-keeled near

41. Lower glumes 2- or 3-nerved and awned, the awns 1-5 mm long; spikelets paired on

a branch, the lower spikelet staminate or zs and De Men Rond b

42. Upper leaf blades a

SU V.

1.5- 3m usu ly with triangular | lol . |

to vee 3. os phleoides

42. Upper leaf blades acuminate with slender awn-like seta 3-8(-10) mm long;

ligules (2-)3-12 mm long, sometimes with lateral lobes the same length as the

ligule; culms erect 4. Lycurus setosus

41. Lower glumes all 1-nerved, unawned or mucronate, the mucros 0.5-1 mm long; spike-

lets borne singly on a branch, or if in pairs, the upper and lower spikelets both bisexual.

43. Lemmas awned, the awns 10-20 mm long; spikelets 3.5-4.3 mm long, stra-

mineous to purplish; glumes 1.2-2.5 mm, the apex a ks erose, not

K

Qo

. Lemmas mucronate, the mucros 0.5-1 mm long; spikelets ‘i p mm lon

green or plumbeous; glumes Du mm long, apex acute to obtuse 2

reos the mucros 0.5-1 mm long; lemmas without distinct, raised lateral

nerv

37. Panicles narrow, os- )1-3 cm wide, loosely contracted to narrowly open; primary branches

loosely appressed to spreading up to 70° from the rachises, the branches 0.4-8.5 cm lon

44. Lemmas unawned, mucronate (mucros < 1 mm long), or awned, the awns 1-6 mm long.

45. Spikelet pedicels flexuous nodding to reflexed, capillary; lemmas with a thick

covering of tawny to shiny silky hairs located on the midvein and margins; ligules

2. Ble

mucronate; lemmas with distinct, lateral nerves — ossis gypsophila

g, dar

Muhlenbergia wrightii

(0.3-)0.7-2(2-2.7) mm long, hyaline to opaque throughout == pharoneuron

tricholepis

938 Journal of the Botanical R h Institute of Texas 1(2)

45. Spikelet pedicels erect, not fl

a thick covering of tawny to sl tl lvei | ins but without

hairs, puberulent throughout or X dense villous on the margins b lo ;ligules 4-35

mm long, E gua ag or SIME

46. Spikelets «anti 1.5-2.2 mm lona, gree-

ish; T not decurrent and brownish below alien dubia

46. Spikelets 2.4-3.5 mm long; basal leaf sheaths compressed ea or flattene

s 1.1-1.6 mm long, yellowish or purplish; ligules deccurent, bou

below.

47. Glumes longer than the florets, glabrous and without hairs, scabrous or

smooth; lemmas glabrous, scabrous or smooth, rarely uM ligules

10-35 mm long hlenbergia lindheimeri

47. Glumes shorter than the florets, pubescent manm near m base; lem-

mas villous on the lower Y, and margins below; ligules 5-13 mm long

23. Muhlenbergia pubigluma

44. Lemmas d the awns 10-40 mm long.

. Glumes 0.3-1.1 mm long, < % as long as the lemma, apices obtuse to acute.

49. Cums udis ub a 1 a none ligules 4-10 mm long, decurrent, firm

i on the upper 2/3; lemmas short pilose

oed: 2.3-3.1 mm long, apices acute; lemma awns 10-20 mm d anther

mm long; caryopses 1-1.2 mm long 4. Mublenbergia purpusii

49. ree 8 nodes; ligules 1-3 mm long, not decurrent

ish below; glumes glabrous; lemmas sparsely appressed-pubescent on the

lower 1⁄4, 2.8-4 mm long, apices acuminate; lemma awns (10-)20-40 mm lon

anthers 0.9-1.6 mm long; caryopses 2-26 mm long _____ 32 MURIS Bere spiciformis

48. Glumes (1-)1.2-2.8 mm long, > Y as long as the lemma (at least the upper glume),

apices acute to acuminate.

50. Culms stigulose below the nodes; anthers 1.5-2 mm a E lemmas

elliptic, widest near the middle Meier a polycaulis

50. Culms scaberulous below the nodes; anthers 0.9-1.5 mm > yellowish; le-

mas lanceolate, widest near base 34. Muhlenbergia tenuifolia

Aegopogon Humb. & Bonpl. ex Willd., Sp. Pl. 4(2):899. 1806. Tr: Aegopog hroides Humb. & Bonpl. ex Willd.

Slender annuals or perennials, caespitose to sprawling, occasionally with stolons; hermaphrodites or func-

tionally andromonoecious. Culms 2-55 cm tall, erect, geniculate or decumbent, herbaceous; internodes

hollow. Leaf sheaths open; ligules membranous; blades flat. Inflorescences terminal, open to somewhat

contracted, racemelike, l-sided panicles; each branch with 3 spikelets; disarticulation at the base of the

branches. Spikelets 1-flowered; lateral spikelets pedicellate, staminate or sterile, sometimes rudimentary;

central spikelets nearly sessile or short-pedicellate, laterally compressed, perfect; glumes shorter than the

florets, equal, 1-nerved; lemmas membranous, 3-nerved, the central and lateral nerves usually extending

into awns; paleas 2-nerved, the nerves usually extending into awns; lodicules 2; ovary glabrous, styles free

to their bases, stigmas 2; stamens 3. Caryopses fusiform to ellipsoid, laterally compressed. Chromosome

number x = 10. Four species, three in México and one endemic to South America.

Etymology.—Name from the Greek aix, ‘goat’, and pogon, ‘beard’, alluding to the many awns of the

spikelets.

Comments.—In a preliminary molecular analyses all species a placed in Aegopogon fall into a

“Muhlenbergia subg. Muhlenbergia” clade that has anatomical characteristics consistent with PCK (phospho-

enolpyruvate carboxykinase) subtype of C, photosynthesis (Peterson 2000; Peterson et al. 2001b, 2004;

Peterson & Herrera Arrieta 2001; Columbus & Peterson, in prep.). Anatomical features of this clade include:

loosely arranged chlorenchyma with tabular cells that are indistinctly radiate and continuous between the

vascular bundles [PCK subtype, defined as centrifugal/evenly distributed photosynthetic carbon reduction

(PCR) cell chloroplasts (with grana), XyMS+ and presence of PCR cell wall suberized lamella, in Hattersley

and Watson's (1992) sense], shield-shaped (narrower than deep) central bulliform cells, and primary vascular

bundles with non-sclerosed phloem (Peterson & Herrera Arrieta 2001).

Peterson et al., Muhlenl ii f T í Mavi 35

1. Aegopogon tenellus (DC.) Trin., Gram. Unifl. Sesquifl. 164. 1824. (Fig. 1, A & B). Lamarchia tenella DC., Cat.

Pl. Horti Monsp. 120. 1813. Tyee: Cultivated at Montpellier, De Candolle s.n. (HoLotyPe: MPU?; Isotype: US-75926 fragm!.).

Hymenothecium unisetum Lag., Gen. Sp. Pl. 4. 1816. Aegopogon unisetus (Lag.) Roem. & Schult., Syst. Veg. 2:805. 1817. ibo gemini-

florus var. unisetus (Lag.) E. Fourn., Mexic. Pl. 2:71. 1886. Tree: MÉXICO: cultivated from seed, D. Sessé s.n.(HOLOTYPE: MA).

Aegopogon gracilis Vasey, Bull. Torrey Bot. Club 13(12):230. 1886. Tyee: MÉXICO. Chinuanua: Aug 1885-Nov 1885, E. Palmer 28 (uo-

53050).

Aegopogon geminiflorus var. abortivus E. Fourn., Mexic. Pl. 2:71. 1886. Tree: MÉXICO. Veracruz: Orizaba, 1864, Weber & Thomas s.n.

(SYNTYPE: P; IsosyNTYPE: US-75961 fragm. ex P!). Escamela, M. Bourgeau 750 (sYNTYPE: P; IsosyNtyPE: US-75303 fragm!). VERACRUZ:

Orizaba, Botteri 41 (syNtypE: P). Orizaba, M. Botteri & A.L. Sumicrast 84 (syntypE: P). San Luis Porosi: Sep 1877, Schaffner 7 (SYNTYPE:

P; ISOSYNTYPE: GH)

>

Aegopogon geminiflorus subvar. purpureus Griseb. ex E. Fourn., Mexic. Pl. 2:71. 1886. Type: MEXICO. Distrito Feperat: Tacubaya, Schaffner

168a (SYNTYPE: P, IsOSYNTYPE: US-75962!). PANAMA. Seemann s.n. (SYNTYPE: K).

Aegopogon cenchroides var. abortivus E. Fourn., Mexic. Pl. 2:72. 1886. Aegopogon tenellus var. abortivus (E. Fourn.) Beetle, Univ. Wyoming

19. 1948. Type: MÉXICO. Veracruz: M. Botteri & A.L. Sumicrast 1187 (HOLOTYPE: P!; isotype: US-75949 fragm. ex P!).

Aegopogon imperfectus Nash, N. Amer. Fl. 17(2):138. 1912. Tyee: MÉXICO. Cutnuanua: Arroyo Aucho, Sierra Madre, cool mossy ledges,

15 Oct 1887, C.G. Pringle 1408 (HOLOTYPE: NY; isotype: US-740860!).

Slender often sprawling, caespitose annuals. Culms (2—) 6-30 cm tall, glabrous below the nodes; internodes

0.6-6 cm long, glabrous to pilose. Leaf sheaths mostly 0.5-4.8 cm long, shorter than the internodes, gla-

brous to sparingly pilose; ligules 0.6-1.5 mm long, apex mostly truncate, lacerate; blades 1.5-6 cm long,

0.5-1.5(C1.7) mm wide, flat, scaberulent and pubescent above, smooth beneath. Panicles 2-6 cm long,

0.5-1.2 cm wide, open, loosely-flowered, racemose primary branches 2-4 mm long, excluding the awns,

one per node. Spikelets 1.5-3.2 mm long, often greenish or purplish, the clusters with one short-pedicelled

spikelet (bisexual), the pedicels 0.2-0.6 mm long and the other two spikelets (staminate or sterile) short-

pedicelled, the pedicels about 0.7-1.5 mm long; glumes (1-)1.3-2 mm long, oblong and wider distally, apex

deeply notched, entire or mucronate, the mucro 0.2-1 mm long, lobes obtuse; lemmas 2.5-3.2 mm long,

3-awned, the central awns 3-8(-11) mm long, lateral awns usually 0.2-2 mm long or missing; paleas 2.2-3

mm long, puberulent, apex 2-mucronate, the mucros less than 0.8 mm long; anthers 0.5-0.8 mm long,

yellowish. Caryopses 1.1-1.3 mm long, obovoid, light brownish. 2n = 20, 60.

Phenology.—Flowering August through November.

Distribution and habitat.—Moist slopes, cliffs, barrancas, canyons, roadsides, and along or near springs

usually in shaded areas associated with Pinus spp., Quercus spp., Salvia sp., Ceanothus sp., Juniperus spp.,

Arbutus sp., Holodiscus sp., and Acacia spp.; 1300-2860 m. Aegopogon tenellus ranges from southern Arizona

throughout most of México and Central America.

Specimens examined. MEXICO. Tamaulipas: Municipio de Gémez Farias, ee San José, M.E. Crespo-Ovalle 523 (ANSM). Nuevo

Leon: Municipio de Galeana, km 12 carr. Galeana-Zaragoza, G.

E

Blepharoneuron Nash, Bull. Torrey Bot. Club 25: 88. 1898. Tee: Blepharoneuron tricholepis (Torr.) Nash.

Caespitose perennials and slender annuals, much branched near base, rhizomes absent. Culms 10-70 cm

1

tall, hollow, occasionally pithy, to strigose below the nodes. Leaf sheaths open, glabrous, usually

longer than the internodes; ligules membranous or hyaline, truncate to obtuse often lacerate, the margins

entire, decurrent; blades flat to involute, short pubescent above, glabrous to scabrous below. Inflorescences

open or often somewhat contracted panicles with ascending to spreading branches; the pedicels often capil-

lary, wiry and flexuous or nodding to reflexed, minutely granular just below the spikelet. Spikelets 1-flow-

ered, slightly compressed, disarticulating above the glumes; glumes subequal, ovate to obtuse, occasionally

lanceolate, faintly 1-nerved, grayish-green (olivaceous), glabrous; lemmas 3-nerved, firmer than the glumes

and slightly longer, lanceolate, grayish-green, with densely appressed, tawny to shiny silky hairs located on

the midvein and margins, the apex acute to obtuse, occasionally mucronate; paleas about same length as

lemmas or slightly longer, 2-nerved, densely villous between the nerves; lodicules 2, club-shaped, truncate,

the lateral margins thin; stamens 3; styles not at all united at the base; stigmas plumose. Caryopses fusiform

to ellipsoid, brownish. Chromosome number x = 8. Two species in México, one of these endemic.

940 Journal of the Botanical R h Institute of Texas 1(2)

Fic. 1. Aegopogon tenellus [FW. Gould 10391 (TAES, US)]. A. Habit B. Panicle branch with tl pikelets, central spikelet bi | (perfect) and the

two | | spikel i terile. Muhlent j [PM. Peterson 5703 & CR. Annable (US)]. C. Habit. D Ligule E. Glumes. F. Florets. G

J

Lodicules, stamens, and pistil.

Peterson et al., Muhlenl ii f +1 vd Mavi 25

Name from the Greek blepharis, ‘eyelash’, and neuron, ‘nerve’, a reference to the silky hairs located on the

margins and midvein.

Comments.—In a preliminary molecular analyses Blepharoneuron tricholepis and B. shepherdii are sister,

or are within a grade that includes Muhlenbergia filiformis (Thurb. ex S. Watson) Rydb., M. ligularis (Hack.),

and M. vaginata Swallen (Peterson et al. 2001b, 2004; Columbus & Peterson, in prep.).

2. Blepharoneuron tricholepis (Torr.) Nash, Bull. Torrey Bot. Club 25:88. 1898 (Fig. 2, A-J). vilfa tricholepis

Torr., Pacif. Railr. Rep. 4(5):155. 1857. Sporobolus tricholepis (Torr.) J.M. Coult., Man. Bot. Rocky Mt. 411. 1885. Type: U.S.A. New

Mexico. Bernadillo Co.: Sandia Mountains, 10 Oct 1853, Bigelow s.n. (LecrotyPE: NY! designated by Peterson & Annable, Syst. Bot.

15:522. 1990; isoLecTroTYPE: MO!)

Densely tufted, caespitose perennials. Culms 10-70 cm tall, 0.3-0.7 mm diameter just below the inflores-

cence, glabrous to scabrous just above and below the nodes, slender, erect; internodes 1.2-10 cm long. Leaf

sheaths 1.8-9.5 cm long, usually glabrous, crowded near base, the margins glabrous, occasionally scabrous,

whitish, shorter or longer than the internodes; ligules (0.3—)0.7-2(-2.7) mm long, hyaline to opaque, truncate

to obtuse, the margins entire, decurrent, the apex erose-dentate to finely ciliate and irregularly toothed;

blades 1-15 cm long, 0.6-2.5 mm wide, filiform, involute, arcuate, scabrous above and below, the midvein

evident, raised on the abaxial surface. Panicles 3-25 cm long, 1-10 cm wide, a narrow or open panicle with

ascending primary branches spreading 0—50? from the rachises; primary branches 1-6 cm long, 1-3 per

node; pedicels 2-9 mm long, slender, capillary, minutely granular just below the spikelet, flexuous nod-

ding to reflexed; nodes per panicle 5-25. Spikelets 2.3-3.4(-3.8) mm long, grayish-green; glumes nearly

equal to subequal in length, grayish-green (olivaceous) and occasionally purplish tinged, glabrous, ovate

to oblong-elliptical, the apex obtuse to acute often irregularly toothed; lower glumes (1.5-)1.8-2.6(3) mm

long; upper glumes (1.7—)2-3.2(-3.7) mm long, broader and often appearing 3-nerved by its characteristic

folding, however, only the midvein contains a vascular trace; lemmas (2-)2.33.4(3.8) mm long, lanceolate,

grayish-green (olivaceous), often purplish tinged; apex acute to obtuse with a thick covering of appressed

to spreading tawny to shining silky hairs on the midvein and margins, the hairs up to 1.2 mm long; pa-

leas (22)2.2-3.5(-3.9) mm long, densely villous between the nerves; anthers 1.2-2.1 mm long, brownish.

Caryopses 1.2-1.4 mm long, fusiform to elliptic, the embryo with a dorsal ridge extending 2/3 of the grain,

light brownish. 2n = 16.

Phenology.—Flowering mid August through November.

Distribution and habitat.—Dry rocky to sandy slopes, canyon walls, rock outcrops, dry meadows, and

open woods in pine-oak-madrone forests with Arctostaphylos spp., pinyon-juniper woodlands, spruce-fir

forests with Ceanothus sp., Cupressus sp., and Cornus stolonifera, aspens groves, and pine woodlands with

Artemisia and Garrya; central México north throughout the Sierra Madre Occidental, Sierra Madre Oriental,

and through the Rockies to Utah and Colorado, U.S.A. (Peterson & Annable 1990); 700-3660 m.

Specimens examined. MEXICO. Coahuila: Municipio de Arteaga, 4 km N de Las Vigas, R. Banda-Silva s.n. (ANSM); Las Vigas, Cañón de

ee Sierra de Arteaga, J.A. Villarreal Q. 1794, M.A. Carranza P. & J. Valdés-Reyna (ANSM); Municipio de Múzquiz, Madera del Carmen,

ooded canyon above Campo El Dos, P.M. Peterson 18908 & J. Valdés-Reyna (US). Nuevo Leon: 11.4 mi W of Dieciocho de Marzo up

n towards Cerro Potosi, P.M. Peterson 13335 & M.B. Knowles (US); 17.5 mi W of Dieciocho de ^ f Cerro Potosi, P.M.

Peterson 13355 & M.B. Knowles (US); Municipio de Aramberri, Cerro El Viejo, G.S. Hinton 23968 (ANSM, MEXU, TEX); Municipio de

Galeana, Sierra Madre Oriental: ascent of Infernillo, 15 mi SW of Galeana, C.H. Mueller & M. T. Mueller 926 (MEXU, TEX); Municipio

de General Zaragoza, Peña Nevada, M.H. Cervera-Rosado 336 (COCA).

Lycurus Kunth, Nov. Gen. Sp. 1:141. 1816. Wee: Lycurus phleoides Kunth, (Lecrorvee: designated by Hitchc. U.S.D.A. Bull.

722:139. 1920).

Caespitose perennials; rhizomes absent. Culms 10—60 cm tall, erect or decumbent, usually branched. Leaf

sheaths open above, laterally compressed; ligules membranous to hyaline, decurrent; blades flat or folded,

margins prominent. Inflorescences contracted spikelike panicles, terminal and axillary; each branch with

a pair of unequally pedicellate spikelets or a pedicellate spikelet and a short secondary branch bearing two

spikelets; disarticulation at the base of the pedicels or branch, the paired spikelets falling as a unit. Spikelets

Botanical R h Institut f Texas 1(2)

942 Journal of tl

Fic. 2. Blepharoneuron tricholepis [PM. Peterson 5567 & C.R. Annable (US)]. A. Habit. B. Ligule. C. Panicle. D. Spikelet. E. Glumes. F. Floret. G. Floret. H.

Palea. I. Lemma, ventral view. J. Lodicules, stamens, and pistil.

Peterson et al., Muhlenl ii f T í Mavi T

1-flowered; usually the lower spikelet in each pair staminate or sterile and the upper spikelet bisexual; glumes

subequal, awned, 1-or 2(3-)-nerved; lower glumes usually 2-nerved, 2-awned; upper glumes 1(2-)-nerved,

1(22-awned; lemmas membranous, 3-nerved, lanceolate, puberulent along the margins, apices awned, the

awn usually shorter than the lemma; paleas about as long as the lemmas, apex acute, sometimes the nerves

extending as two mucros, margins hyaline; stamens 3. Caryopses fusiform. Chromosome number x = 10.

Name from the Greek lykos, ‘wolf’, and oura, ‘tail’, a reference the the spikelike inflorescence.

Comments.—Possession of paired spikelets and 2-nerved lower glumes that are 2-awned, although

not common within the Muhlenbergiinae, is not unique to Lycurus. Muhlenbergia brevis and M. depauperata

exhibit morphological features that suggest a close relationship with Lycurus. Mez (1921) indicated a rela-

tionship with Lycurus when he transferred M. shaffneri E. Fourn., considered a synonym of M. depauperata,

to Lycurus. Muhlenbergia brevis and M. depauperata share many morphological features with Lycurus, most

importantly: spikelets borne in pairs and lower glumes that are 2-nerved and 2-awned (Peterson 2000).

Schaffnerella gracilis is apparently sister to the three species of Lycurus for trnL-F and ITS sequence analyses,

and this Lycurus-Schaffnerella clade is found in a well-supported trnL-F-derived clade that includes M. wrightii,

all four species of Chaboissaea, and Schedonnardus paniculatus (Peterson et al. 2004; Columbus & Peterson,

in prep.).

Allozyme studies were useful is determining the biogeographical history of the amphitropical Lycurus

setosus and it seems likely that this species has recently dispersed to South America because populations

there contain less genetic variation (Peterson & Morrone 1997; Peterson 2000).

3. Lycurus palon Kunth, Nov. oo: Sp. No ed. ) 1:142, pl. 45. 1816. (Fig. 3, A & B). Wee: MÉXICO:

inter Guanaxuato et Ti a Buffa, alt. 1030 hexap., Sep, EWH.A. Humboldt & A.J.A. Bonpland

s.n. (HOLOTYPE: P!; isotypes: BAA-fragm.!, US-610840 fragm. ex P-Bonpl!, US-610841 fragm.!).

A

Loosely caespitose perennials. Culms 20-60 cm tall, erect t umbent at base, scabrous to puberulent just

above or below the nodes. Leaf sheaths usually 0.5—3 cm long, glabrous to pubescent, compressed-keeled near

base; ligules 1.5-3 mm long, acuminate and erose, usually with triangular lateral lobes 0.5-2.5 mm long,

the lateral lobes decurrent below; blades 2-8 cm long, 1-1.5 mm wide, flat or folded, glabrous to puberulent

below and puberulent to hispidulous above, margins and midvein whitish-thickened, apex acute or some-

times with a narrow bristle 0.5-3 mm long on the upper blades. Panicles (2—)4-10 cm long, 4-8 mm wide,

contracted, narrow, densely flowered, and spikelike; primary branches 0.1-0.3 cm long, tightly appressed.

Spikelets 3-4 mm long, 1-flowered; glumes 1-2 mm long, scaberulous near apex, awned, the awns 1-5

mm long; lower glumes usually 2(3-)-nerved; upper glumes 1(2-)-nerved, awns 2-5 mm long; lemmas 3-4

mm long, puberulent along the margins, awned, the awns 1-3 mm long; paleas 3-4 mm long, puberulent

between the nerves; anthers 1.5-2 mm long, yellow. Caryopses 1.8-2.2 mm long, brownish. 2n = 40.

Phenology.—Flowering July through October.

Distribution and habitat. —Rocky slopes, canyons, bajadas, and flats often occurring in calcareous soils

and associated with Quercus gregii, Larrea tridentata, Pinus cembroides, Juniperus flaccida, Salvia regla, Cowania

plicata, Bouteloua gracilis, B. uniflora, Dasylirion longissimum, Y. carnerosana, Agave lecheguilla, Jatropa dioica,

Muhlenbergia spp., Berberis sp., Opuntia spp, Selaginella sp, Hechtia sp., Brahea sp., and Mimosa sp.; northern

and central México in Aguascalientes, Chihuahua, Distrito Federal, Durango, Guanajuato, Hidalgo, Jalisco,

México, Michoacán, Morelos, Nayarit, Oaxaca, Puebla, Querétaro, San Luis Potosí, Sonora, Tlaxcala, and

Zacatecas to southwestern U.S.A. in Texas and New Mexico (Espejo Serna et al. 2000; Peterson et al. 2001b;

Powell 1994; Reeder 1985; Sánchez & Rügolo de Agrasar 1986); 1200-2500 m.

Specimens examined. MÉXICO. Coahuila: Sierra El Pino, 9.6 km SW of Rancho El Cimarron, P.M. Peterson 10635 & C.R. Annable (US);

33.5 km W of Rancho El Cimarron, P.M. Peterson 10656 & C.R. Annable n 5 mi W of Chapultepec, P.M. Peterson 13278 & M.B. Knowles

(US); 30.6 km SE of Saltillo on HWY 57 towards San Luis Potosí, P. M. Pi 240 & C.R. Annable (US); 6 km S of Saltillo, P.M. Peterson

8347 & J. Valdés-Reyna (US); 35.4 km W of Ocampo at Cuesta Zozaya, P.M. Peterson 8367 & J. Valdés-Reyna (US); western base of Picacho

del Fuste, I.M. Johnston 8444 (US); du of La Noria, R.M. pL Municipio de Acuña, del Carmen Mountains, E.G. M

Jr. 658 (MEXU); L li XU); Municipio de Arteaga, 2 km NW of Ejido Sierra Hermosa,

R. ees n. (ANSM); Sierra Zapaliname, at 2 a E of Saltillo up Camino de Quatro (Las Palapas), P.M. Peterson 17858, J. Valdés-

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Peterson et al., Muhlenl ii f T í Mavi "T

Reyna & R.H. Cardenas (US), P.M. Peterson 18805 & J. Valdés-Reyna (US); Cañón San Lorenzo, Sierra de Zapalinamé, 8 km S of Saltillo

& 3.2 km E de Buenavista, R. López Aguillón s.n. (ANSM); 3 mi SE of Saltillo, I.M. Johnston 7250 (US); road to Canyon San Lorenzo, P.M.

Peterson 10545 & C.R. Annable (US); Las Vigas, Cañón de la Carbonera, Sierra de Arteaga, J. Valdés-Reyna 1797, J.A. Villarreal Q. & M.A.

Carranza P. (ANSM); Predio El Cristal, E. Pérez Torres 61 (COCA): Municipio de Castaños, Paso de San Lázaro, Sierra de la Gavia, 37.6

nclova on Hwy 57, P.M. Peterson 9990, C.R. Annable & J. Valdés-Reyna (ANSM, US); Municipio de Cuatrociénegas, Sierra de la

Madera, Cañón Charreteras, Rancho Charreteras, J.A. Villarreal Q. 7350, M.A. Carranza P. & R. Rodríguez L. (ANSM); Cañada Los Pozos,

8 mi SW of Tanque El Tropical along road to Carión del Desiderio, T.L. Wendt s.n. & J. Valdés-Reyna (MEXU); Municipio de Müzquiz,

near Santo Domingo, F.W. Lyle 454, C.H. Muller (ANSM, US); Sierra Maderas del Carmén, E. García Aguilera s.n. (ANSM); Municipio de

Ocampo, Mesa Grande, 40 km NW of Hacienda de La Encantada, R.M. Stewart 1631 (MEXU); W base of Picacho del Fuste, NE of Tanque

Vaionetta, I. M. Johnston 8444 (MEXU); Municipio de Parras, Sierra de Parras, 2-4 km S of Ejido Colorado, 8.8 km al W of Parras, C.P.

Cowan 3610 (TEX- m LN de Ramos Arizpe, Sierra de la Paila, Ejido El Cedral, J.A. Villarreal Q. 5330, M.A. Carranza P. & A.

Rodriguez G. (ANSM de Saltillo, 20 km E of Saltillo, Carretera Saltillo-Torreón, R. Almeida 1642 (ANSM); 4 km W of Saltillo,

Carretera Saltillo- General Ea J. Valdés-Reyna 1553, L.E. Rodríguez G. & R. Vásquez A. (ANSM); 6 mi W of Saltillo along highway

40 towards Torreón, S.L. Hatch 5040 & J. Valdés-Reyna (ANSM); Buenavista, Brigada Coahuila 102, 103, 104 (COCA); Buenavista, 7 km S

de Saltillo, Carretera 54 Saltillo- Concepción del Oro, A. Rodríguez-Guillén 101, R.E. Rodríguez-Charúa 107, M. Jiménez-F. s.n., N. Ochoa-

R. s.n., R. Palomo-Garza & P. García-Solís s.n. (ANSM); Cañón San Lorenzo, 1.5 km E of Pozo, R. López Aguillón s.n. (ANSM); Cerro del

Pueblo, W of La Ciudad de Saltillo, J. Valdés-Reyna 2045, M. Martínez M. & M.A. Carranza P. (ANSM); Hwy 54 towards Concepción del

Oro, 6 km S of Saltillo, M.E. Barkworth 5108, P.S. Hoge, J. Valdés-Reyna, & M.A. Carranza P. (ANSM); Rancho Experimental Los Angeles,

48 km S of Saltillo, Carretera 54 Saltillo- Concepción del Oro, J.A. Villarreal Q. 6710, S.L. Hatch 4538, C.W. Morden, J. Valdés- Reyna, M.A.

Carranza P. & D.E. Lozano (ANSM); Saltillo, G. Fisher s.n. (UAT); Sierra Madre Oriental, Terrenos de Tranquitas, R.M. Francois 5827 &

J. Passini (ANSM); Sierras adyacentes al Paso de Carneros, J.S. Marroquín-de la Fuente 3058 (ANSM); 29 km S of Saltillo on MEX 54 just

W of Estacion Caranaros, P.M. Peterson 8413 & M.A. Carranza P. (US); RS de San Pedro, entre Cuatrociénegas & San Pedro, A.

Rodríguez- Gámez 945, J J. López & J. L. Elizondo E. (ANSM, TEX); Muni de Si Mojada, Cañon de Tinaja Blanca, W of Santa Elena

Mines, R.M. Stewart 1950 (US); Municipio de Torreón, Sierra de Jimulco, 1L 7 km E of Flor de Jimulco, P.M. Peterson 8453, J. Valdés-Reyna,

P.A. Fryxell & J.A. Villarreal Q. (ANSM, US); Sierra de Jimulco, Mina San José, J.A. Villarreal Q. 7791 & M.A. Carranza P (ANSM, TEX).

Nuevo León: 12.3 mi S of Milagro, P.M. Peterson 17810 ae Valdés- oe & G.S. Hinton ide 16 km NE of Sandia on road to La Ascensión,

P.M. Peterson 15826 & J. Valdés-Reyna (US); i Ejido La Encantada, P.M. Peterson 16733, J. Valdés-Reyna

& M. Sosa Morales (US); 9.4 mi W of San Antonia de Peña Neda P.M. Peterson 16791, J. Valdés-Reyna & M. Sosa Morales (US); 75.8 km N

of Dr. Arroyo on Hwy 61, P.M. Peterson 11144 & C.R. Annable (US); 6.5 mi S of Border of Coahuila and Nuevo Leon on Hwy 57 towards

Matehuala, P.M. Peterson 13286 & M.B. Knowles (US); 2 mi W of Dieciocho de Marzo up road towards Cerro Potosí, P.M. Peterson 13325 &

M.B. Knowles (US); 36 mi NE of Dr. Arroyo on Hwy 61 towards Linares, P.M. Peterson 13366 & M.B. Knowles (US); N

10 mi E of San Roberto along highway 58, S.L. Hatch 4978, J. Valdés-Reyna 6 P. Dávila A. (ANSM); 12 km SE of San Roberto on eee

58, S.L. Hatch 4580, C.W. Morden & J. Valdés-Reyna (ANSM); Alrededor de Galeana, J.A. Ochoa-Guillemar 1258 (COCA); Hacienda Pablillo,

M. Taylor-Edwards 134 (MEXU); Rancho Aguililla, G.B. Hinton 19863 (MEXU, TEX); alrededores del poblado Santa Clara de González,

N. Bazaldua- poesia 110 ee Tamaulipas: Municipio de Bustamante, Camino a La Joya de San Francisco, J.L. Ramos-Delgado 99

(COCA); M ] 15 km SW of Miquihuana, P. Hiriart 301, V. Juárez & R. Molczadzki (MEXU); 7 km W of Miquihuana,

P. Hiriart 899,V. Tuáre R. Molczadzhi, J.L. López, F. Ojeda & J.A. Gutiérrez (MEXU); 9 km N of Miquihuana, P. Hiriart 335, V. Juárez & R.

Molczadzhi (MEXU); 3 mi N of Miquihuana, Stanford 2481, Lauber & Taylor (US); Colonia La Peria, J.F. Iribe-Duarte 106 (COCA); Ejido

El Aserradero, J.G. Galván-Infante 37 & R.A. Carranco-Rendon 70 (COCA); Ejido Valle Hermoso, Cañón de La Sierra Miquihuana, R.A.

Carranco-Rendon (COCA); Municipio de Palmillas, Ejido Palmillas, R.A. Carranco-Rendon 213 (COCA); Municipio de Tula, La Tapona,

M.H. Cervera-Rosado 133 (COCA)

4. Lycurus setosus (Nutt.) C. Reeder, Phytologia 57:287. 1985. (Fig. 3, C—E). Pleopogon setosum Nutt., Proc. Acad.

Nat. Sci. Philadelphia 4:25. 1848. Tyre: U.S.A. New Mexico: Santa Fe Co.: mountains near Santa Fe, 1841 or 1842, William Gambel

s.n. (HOLOTYPE: K!; isotypes: PH, US-610839 fragm. ex K!).

Lycurus alopecuroides Griseb., Abh. Kónigl. Ges. Wiss. Gottingen 19: 255-256. 1874. Type: ARGENTINA: Prov. CATAMARCA: ca. Belén,

en el altivalle de las Granadillas, Feb 1872, P G. Lorentz 548 (HoLoTYPE: GOET; IsoTYPES: BA!, BAA!, CORD, SI!, US-996080 fragm.

ex GOET!).

Ly phleoid glaucifolius Beal, Grass. N. Amer. 2:271. 1896. Type: MEXICO. C ky hills near Chihuahua, 28 May 1885,

C.G. Pringle 426 (Lectotype: MSC designated by Hitchcock, Contr. U.S. Natl. Herb. 17:305. 1913, without explicitly citing a specific

DI

et in a specific herbarium, fully lectotypified by C. Reeder

isolectotypes: MO-3701765!, NY, P!, SI!, US-996079!).

.1985 , who cited the MSC specimen as “holot otype”;

>

Caespitose perennials. Culms 30-60 cm tall, erect, scabrous to puberulent just above or below the nodes.

Leaf sheaths usually 1-5 cm long, glabrous to pubescent, compressed-keeled near base; ligules (22)3-12

mm long, acuminate to erose, sometimes with lateral lobes the same length as the ligule, the lateral lobes

decurrent below; blades 4-10(22) cm long, 1-2.5 mm wide, flat or folded, glabrous to puberulent below

946 Journal of the Botanical R h Institute of Texas 1(2)

and puberulent to hispidulous above, margins and midvein whitish-thickened, apex acuminate with slender

awn-like seta 3-8(-10) mm long on the upper blades. Panicles 2-)4-10 cm long, 4-8 mm wide, contracted,

narrow, densely flowered, and spikelike; primary branches 0.1-0.3 cm long, tightly appressed. Spikelets 3-4

mm long, 1-flowered; glumes 1-2 mm long, scaberulous near apex, awned, the awns 1-5 mm long; lower

glumes usually 2(3-)-nerved; upper glumes 1-nerved, awns 2-5 mm long; lemmas 3-4 mm long, puberulent

along the margins, awned, the awns 1-3 mm long; paleas 3-4 mm long, puberulent between the nerves;

anthers 1.5-2 mm long, yellow. Caryopses 1.8-2.2 mm long, brownish. 2n = 40

Phenology.—Flowering July through October.

Distribution and habitat. —Wooded canyons, rocky slopes, desert grasslands, and rock outcrops with

Pinus spp. Quercus spp., Bouteloua hirsuta, Chrysothamnus spp., Pseudostuga menziesii, Cupressus spp., Abies

spp., Cornus stolonifera, Ceanothus spp., and Yucca spp.; northwestern México in Baja California, Chihuahua,

1

Durango, Sonora, and extending into the southwestern U.S.A., then disjunct in

Bolivia (Espejo Serna et al. 2000; Peterson et al. 2001b; Reeder 1985; Sánchez & Rúgolo de Agrasar 1986);

1400-2500 m

Specimens examined. MÉXICO. Coahuila: Sierra El Pino, 39.5 km W of Rancho El Cimarron, P.M. Peterson 10672 & C.R. Annable (US);

1219 A NIC

Municipio de Acuña, Serranías del Burro, Rancho El Bonito, Cañón El Bonito, 7 mi W de la casa principal, D.H. Riskind 2185 (ANSM

Argentina and

Municipio de Cuatrociénegas, Sierra de la Madera, Cañada Los Posos, 0.8 mi by road SW from Tanque El Tropical along road to Cañón

del Desiderio us Valdés- a 1053 & T.L. Wendt (ANSM), Sierra de la Madera, Cañón del Agua, mouth of canyon un Valdés- NAE 1000,

10126 T.L Orientale, Ejido el Cinco, R.M. F 92 & J. Passini (ANSM); M :

Sierra Madre Oriental Ejido El Cinco, R.M. F i 83 & J. Passini (ANSM); Municipio de M quiz, Cuesta de la mE J.A a

L. s.n. (ANSM); Sierra del Carmén, R.M. F is 5514 & J. Passini (ANSM); Ejido San men P.M. Peterson 18842, J. Valdés-Reyna & C.

Sifuentes (US); Madera del Carmen, 7.2 mi NW of Pilares, P.M. Peterson 18876 & J. Valdés-Reyna (US); Campo El Dos, P.M. Peterson 18910

& J. Valdés-Reyna (US); Municipio de Sierra Mojada, San Antonio de los Alamos, I.M. Johnston 8252a (US).

Muhlenbergia Schreb., Gen. Pl. (ed.8) 1:44. 1789. Wee: Muhlenbergia schreberi J.E Gmel., (LecroTvrE: designated by Nash in

Britton & Brown, Ill. Fl. N. U.S., ed. 2, 1:184. 1913).

Plants annual or perennial; hermaphrodites or occasionally monoecious; often caespitose, sometimes mat-

forming, usually rhizomatous, rarely stoloniferous. Culms 2-300 cm tall, erect, geniculate, or decumbent,

usually herbaceous, sometimes becoming woody; internodes solid or hollow. Leaf sheaths open, auricles

absent; ligules membranous or hyaline, sometimes firm or coriaceous, acuminate to truncate, sometimes

minutely ciliolate, sometimes with lateral lobes longer than the central portion; blades narrow, flat, folded,

or involute, sometimes arcuate. Inflorescences terminal, sometimes also axillary, open to contracted or

spikelike panicles; disarticulation usually above the glumes, occasionally below the pedicels. Spikelets

1(2—3)- flowered, laterally compressed or terete; glumes subequal in length, mostly shorter than the lemmas,

or as long or exceeding the lemmas, usually (0)1(2, 3)-nerved, apices entire, erose, or toothed, truncate to

acuminate, sometimes mucronate or awned; lower glumes sometimes rudimentary or absent, occasionally

bifid; upper glumes shorter than to longer than the florets; lemmas membranous or rather firm, glabrous,

scabrous, or with short hairs, 3-nerved (occasionally appearing 5-nerved), callus glabrous or hairy, apices

awned, mucronate, or unawned; awns, if present, straight, flexuous, sinuous, or curled, sometimes borne

between 2 minute teeth; paleas membranous or rather firm, well developed, about as long as the or shorter

than the lemma, 2-nerved, rounded on the back; lodicules 2, short, fleshy, truncate; lateral margins thin;

ovary glabrous, styles free to their bases, stigmas 2; stamens (1, 2)3, anthers purple, orange, yellow, or oli-

vaceous. Caryopses elongate, fusiform or elliptic, slightly dorsally compressed, usually not falling free from

the lemma and palea. Cleistogamous panicles sometimes present in the axils of the lower cauline leaves,

enclosed by a tightly rolled, somewhat indurate sheath. Embryo large, with an epiblast, scutellar tail, and

elongated mesocotyl; embryonic leaf margins meeting, endosperm hard. Chromosome number x = 10.

Etymology.—Named for Gotthilf Henry Ernest Muhlenberg (1753-1815), a Lutheran minister and

pioneer botanist of Pennsylvania.

Comments.—A good historical account, biogeographical summary, and subgeneric classification of

Muhlenbergia was given in Peterson (2000) and Peterson and Herrera Arrieta (2001). Based on population

Peterson et al., Muhlenl ii f T í Mavi "

genetic studies of the amphitropical disjunct, Muhlenbergia torreyii (Peterson & Ortíz-Diaz 1998), it is clear

that migration predominantly occurred from a center of diversity (for Muhlenbergia & Muhlenbergiinae) in

southwestern North America (centered in northern Mexico) to northern Argentina-southwestern Bolivia

(Peterson & Herrera Arrieta 1995; Peterson & Columbus 1997; Sykes et. al. 1997; Peterson & Morrone 1997;

Peterson & Ortíz-Diaz 1998; Peterson 2000). Chihuahua apparently has the highest number of species

of Muhlenbergia (59 spp.), followed by Durango with 55 species (Herrera Arrieta 2001; Herrera Arrieta &

Peterson 2007). In northeastern México, M. jaime-hintonii is endemic to the region and is known only from

Nuevo León.

Even though preliminary phylogenetic results of trnL-F and ITS DNA sequences indicate that Muhlen-

bergia is polyphyletic, there are monophyletic lineages that correspond to previous delineated taxa (Peterson

et al. 2001a, 2004; Columbus & Peterson, in prep.). In addition to all species of Aegopogon and Pereilema,

the following seven species placed in Muhlenbergia subg. Muhlenbergia are well supported in the molecular

analysis and exhibit PCK-like anatomical characteristics: M. glauca, M. microsperma, M. polycaulis, M. porteri,

NAGEL 1

M. racemosa, M. spiciformis, and M. tenuifolia ia distichophylla, M. dubia, M. emersleyi, M. gypsophila,

M. jaime-hintonii, M. lindheimeri, M. macroura, M. pubigluma, M. purpusii, M. rigens, M. rigida, M. robusta, M.

setifolia, and M. stricta (Muhlenbergia subg. Trichochloa) have anatomical characters of deep adaxial furrows,

vascular bundles positioned in two or three levels, and inflated cells located below (abaxial to) the primary

vascular bundles; and are found in a well supported molecular clade (Peterson et al. 2001a, 2004; Peterson

& Herrera Arrieta 2001). There is good molecular support for M. montana, M. quadridentata, and M. vire-

scens, all members of a group referred to as the “Muhlenbergia montana complex” that have 3-nerved upper

glumes and sclerosed phloem (Peterson et al. 2001a, 2004; Peterson & Herrera Arrieta 2001). The annuals,

M. depauperata, M. fragilis, and M. minutissima, with M. arenicola and M. torreyii form a fairly well supported

molecular clade; and Muhlenbergia arenacea, M. asperifolia, M. repens, M. utilis, M. villiflora, and M. wrightii

are all members of a fairly well supported molecular clade that includes the M. montana complex, Lycurus-

Schaffnerella, Chaboissaea spp., and Schedonnardus paniculatus (Columbus & Peterson, in prep).

5. Muhlenbergia arenacea (Buckley) Hitchc., Proc. Biol. Soc. Wash. 41:161. 1928. (Fig. 1, C-G). sporobolus

arenaceus Buckley, Proc. Acad. Nat. Sci. Philadelphia 14:89. 1862. Type: U.S.A. New Mexico/Texas: May-Oct 1849, C.H. Wright 737

(LECTOTYPE: GH designated by Hitchcock, Man. Grass. U.S. 886. 1935, but without citing a specific sheet in a specific herbarium;

ISOLECTOTYPE: US-997371!

Sporobolus auriculatus Vasey, Contr. U.S. Natl. Herb. 3(1):64. 1892. Tyre: U.S.A. Texas: Duval Co.: Pena, 1890, G.C. Nealley 492 (LECTOTYPE:

US-997369! designated by Hitchcock, N. Amer. Fl. 17(6):447. 1935, but witl t in a specific herbarium)

o

Lat

Rhizomatous perennials. Culms 10—30(-40) cm tall, decumbent, terete to compressed-keeled near

the base; internodes scaberulous below the nodes. Leaf sheaths about 1⁄2 as long as the internodes, margins

hyaline; ligules 0.5-2 mm long, hyaline, with lateral lobes or auricles 1-2 mm long; blades 0.7-4(-6) cm long,

0.5-1.7 mm wide, flat, occasionally folded, tapering, scabrous abaxially, strigulose adaxially, margins and

midveins thickened, whitish, apices narrow, often sharp, erose. Panicles 5-15 cm long, 4-14 cm wide, broadly

ovoid, open; primary branches 2-8 cm long, capillary, straight to slightly flexuous, diverging 45-80(-100)?

from the rachises, never appearing fascicled, naked proximally; pedicels 1-11 mm long, usually longer than

the spikelets. Spikelets 1.5-2.5 mm long, occasionally with 2 florets; glumes 0.9-2 mm long, equal, 1-nerved,

usually acute to acuminate, occasionally erose and mucronate, mucros to 0.2 mm long; lemmas 1.5-2.5

mm long, lanceolate to oblong-elliptic, plumbeous to purplish, sparsely appressed-pubescent on the lower

15 of the margins and midveins, hairs to 0.3 mm long, apices acute to obtuse, sometimes shallowly bilobed,

mucronate, mucros to 0.3 mm long; paleas 1.5-2.6 mm long, lanceolate, glabrous, obtuse to acute; anthers

1-1.5 mm long, yellowish to purplish. Caryopses 1-1.3 mm long, elliptic, brownish.

Distribution and habitat—Muhlenbergia arenacea grows in sandy flats, plains, alluvial fans, washes,

depressions, and alkaline mesas in open grasslands, at elevations of 1000-2200 m. It ranges from the

southwestern U.S.A. to northern México in Chihuahua, San Luis Potosi, and Zacatecas (Espejo Serna et al.

2000; Peterson 2003).

948 Journal of the Botanical R h Institute of Texas 1(2)

Specimens examined. MÉXICO. Coahuila: 153.2 km NW of Muzquiz on Hwy 53 towards Boquilla del Carmen, P.M. Peterson 10602

& C.R. Annable (US); Sierra El Pino, 9.2 km SW of Rancho El Cimarron, P.M. Peterson 10624 & C.R. Annable (US); Municipio de Acuña,

Sierra del Carmen, E.G. Marsh Jr. 907 (MEXU); Municipio de Ocampo, Rancho el Barranquito, 50 Km de S rumbo a Sierra pde

M.A. Carranza P. 576 & F.J. Carranza P. (ANSM); Rancho La Rueda, 87 km NW of Ocampo, D. Ibarra s.n. (ANSM

12 km N of the Coahuila-Zacatecas state line along highway 54 between ed and Concepción del Oro, Zacatecas, S. L Hatch 4516,

C. W. Morden & J. Valdés-Reyna (ANSM); 29.2 mi S of Saltillo, carretera 54 Saltillo- Concepción del Oro, Zacatecas, P.M. Peterson 10033,

C.R. Annable & J. Valdés-Reyna (ANSM, US); 3 mi E of San Jose, I.M. Johnston 8217 (MEXU); Municipio de Sierra Mojada, Reserva de la

Biosfera de Mapimí limite del estado de Coahuila, Chihuahua & Durango, R. Ruíz Esparza R. 552 (ANSM)

6. Muhlenbergia arenicola Buckley, Proc. Acad. Nat. Sci. Philadelphia 14:91. 1862. (Fig. 4, A-E). Podose-

mum arenicola (Buckley) Bush, Amer. Midl. Naturalist 7(2):40. 1921. Type: U.S.A. Texas: arid places in western Texas, C. Wright 735

(LecroTYPE: PH designated by Hitchcock, Man. Grass. U.S. 886 (1935), but without citing a specific sheet in a specific herbarium,

ISOLECTOTYPE: US-87239 fragm).

Caespitose perennials. Culms (15—)20—60(-70) cm tall, somewhat decumbent, 1 or more nodes exposed;

internodes hispidulous below the nodes. Leaves somewhat basally concentrated, most blades not reaching

more than 34-15 of the plant height; leaf sheaths usually a little shorter than the internodes, not keeled,

scaberulous, margins hyaline, basal sheaths rounded, not becoming spirally coiled when old; ligules 2-9

mm long, hyaline, acute, lacerate, often with lateral lobes; blades 4—10(—16) cm long, 1-2.2 mm wide, not

arcuate, flat, folded, or involute, scabrous, often glaucous, midveins and margins not thickened, green.

Panicles 12-30 cm long, 5-20 cm wide, diffuse; primary branches 1-10 cm long, diverging 30-80? from

the rachises, naked basally; pedicels 1-4(-6) mm long, erect. Spikelets 2.5-4.2 mm long; glumes 1.4-2.5

mm long, equal, 1-nerved, apices scaberulous, acute to acuminate, minutely erose, sometimes mucronate,

the mucro to 1 mm long; lemmas 2.5-4.2 mm long, narrowly elliptic, usually purplish, scabrous distally,

appressed-pubescent on the lower !5—34 of the margins and midveins, apices acuminate, awned, awns 0.5-4

mm long; paleas 2.5-3.5 mm, narrowly elliptic, intercostal region sparsely pubescent, apices acuminate,

with 2 short mucros 0.1-0.2 mm long; anthers 1.5-2.1 mm, greenish. Caryopses 1.9-2.3 mm long, fusiform,

brownish. 2n = 80, 82.

Distribution and habitat—Muhlenbergia arenicola grows on sandy mesas, limestone benches, and in val-

leys and open desert grasslands, associated with Bouteloua gracilis, B. uniflora, Larrea tridentata, and Yucca

carnerosana at elevations of 600-2135 m. The range of M. arenicola extends to southwestern U.S.A. and the

species is a disjunct in northwestern Argentina (Peterson & Ortiz-Diaz 1998).

Specimens examined. MEXICO. Coahuila: Sierra El Pino, 9.2 km SW of Rancho El Cimarron; P.M. Peterson 10619 & C.R. Annable

(US); 153.2 km NW of Muzquiz on Hwy 53 towards Boquilla del Carmen, P.M. Peterson 10603 & C.R. Annable (US); 0.3 km E of Hwy

to Zacatecas up road to Canyon San Lorenzo, P.M. Peterson 10550, C.R. Annable & J. Valdés-Reyna (US); Municipio de Acuña, 2 km N of

El Conejo on the winding road to La Huerfanita, M.C. Johnston 9192, F. Chiang C. & T.L. Wendt (ANSM); Municipio de Arteaga, Sierra

de Arteaga, Bosque de la Montaña, 10 km del entronque Los Lirios & carretera 57 a Matehuala, rumbo a los Lirios, P.S. Hoge. 257, M.E.

Barkworth, J.A. Villarreal Q. € J. Valdés-Reyna (ANSMD; Municipio de Francisco I. Madero, Valle de Buenavista, S de los Remedios, T.L.

Wendt 1387 & EJ. Lott (ANSM); Municipio de Múzquiz, La Cuesta del Plomo on the Müzquiz-Boquillas highway, F. Chiang-Cabrera

9207 (MEXU), T.L. Wendt & M.C. Johnston 9207 (ANSM); Municipio de Ocampo, Rancho experimental Santa Teresa de La Rueda, 87

km NE of Ocampo, brecha Ocampo-Boquillas del Carmen, R. Vásquez-Aldape s.n. (ANSM); Rancho experimental Santa Teresa de La

Rueda, 87 km NE of Ocampo, brecha Ocampo-Boquillas del Carmen, M. Vásquez-Rodríguez s.n. (ANSM); Rancho La Rueda, 87 km NE

of Ocampo, D. Ibarra s.n. (ANSM); Municipio de Parras, Rancho el Tunal, 25 "m ESE of Parras de la Fuente, A. Rodríguez- Gámez 1181 &

M.A. Carranza P. (ANSM); ^ Saltillo, 28.3 mi S of Saltillo, carretera 54 a Concepción del Oro, Zacatecas, P.M. Peterson 10026

C.R. Annable & J. Valdés-Reyna (ANSM, US); 47 km S of Saltillo on MEX Hwy 54 to Concepcion del Oro near entrance road to El Salitre

Randiera, P.M. Peterson 10032, C.R. Annable & J. Valdés-Reyna (ANSM, m 29.2 mi S of Saltillo, carretera 54 Saltillo- Concepción del

Oro, Zacatecas, P.M. Peterson.10032, C.R. Annable & J. Valdés-Reyna (ANSM, US); 30.6 km SE of Saltillo on Hwy 57 towards San Luis

Potosi, P. M. Peterson 6239 & C.R. Annable (US); 50 km S of Saltillo, carretera 54 a Concepción del Oro, Zacatecas, entronque El Colorado,

J.A. Villarreal 2 2061, M.A. Carranza P. & J. Valdés-Reyna (ANSM, MEXU); 3 mi E of San Jose, I.M. Johnston 8219 (MEXU); Entronque

Derramadero, a 20 km S de Saltillo por la carretera Saltillo-Zacatecas, carretera 54, J. Valdés-Reyna & M.A. Carranza P. (ANSM); Rancho

experimental Los Angeles, 48 km S de Saltillo, carretera 54 Saltillo- Concepción del Oro, Zacatecas D pr -Aburtor 56, S. L. Hatch.

5019, G.A. Puente-Tristán, J. S. Sierra-Tristán & J. Valdés-Reyna (ANSM); Municipio de Sierra Mojada, 15 km E of San Antonio de los

Alamos, I.M. Johnston 8288 (MEXU). Nuevo León: 64.4 km S of San Rafael on d 57, P.M. Peterson 6236 & C.R. Annable (US); 12.3

mi S of Milagro, P.M. Peterson 17815, J. Valdés-Reyna 6 G.S. Hinton (US); ¡Municipio de Galeana , 2 km SE of El Canelito, on road to San

Peterson et al., Muhlenl ii f T í Mavi T

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Fic. 4. Muhlenbergia arenicola [PM. Peterson 6239 & C.R. Annable (US)]. A. Habit B. Ligule. C. Glumes. D. Floret. E. Lodicules, stamens, and pistil. Muhlen-

bergia torreyi [PM. Peterson 5608 & C.R. Annable (US)]. F. Habit. G. Ligule. H. Glumes. I. Lemma.

950 Journal of the Botanical R h Institute of Texas 1(2)

Juan de Dios, M.C. Johnston 7984, T.L. Wendt (ANSM); Ejido El Tokio, M.L. Avalos-Marín s.n. (UAT, ANSM). Tamaulipas: Municipio de

Bustamante, Ejido Felipe Ángeles 2 km NO rumbo a Bustamante, R. Díaz-Pérez s.n. (UAT)

7. Muhlenbergia asperifolia (Nees & Meyen ex Trin.) Parodi, Revista Fac. Agron. Veterin. (Buenos Aires)

Osee . (F ig. 5, A-D). vi lfa UA ia Nees & Mon ex Trin., Mém. Acad. Imp. Sci. Saint-Pétersbourg, Sér. 6, Sci.

Math., Seconde Pt. Sci. Nat. 6,4(1-2):95. 1840. ifolius (Nees & Meyen ex Trin.) Nees & Meyen, Gramineae 9-10.

1841. Tyee: Chile. Río Mayno, 1000' alt, Mar 1831, , Meyen s.n. coe B, BAA-3464 fragm. ex B!, LE-TRIN-1679.01b!); Copiapó,

Meyen s.n. (SYNTYPE: LE-TRIN-1679.01a!).

Agrostis eremophila Speg., Anales Mus. Nac. Hist. Nat. Buenos Aires 7:190. 1902. Agrostis distichophylla Phil., Fl. Atacam. 54. 1860. nom.

illeg. hom. Tyre: Chile. prope Tilopozo ad marginem paludos, 23.20 lat., 7000 ft (isotypes: BAA-4153 fragm. ex SGO!, SGO-63100,

SGO-37135, US-1939353 fragm. ex SGO-37135!)

Rhizomatous perennials, occasionally stoloniferous. Culms 10-60(-100) cm tall, decumbent-ascending,

bases somewhat compressed-keeled; internodes glabrous, shiny below the nodes. Leaf sheaths glabrous,

margins hyaline; ligules 0.2-1 mm long, firm, truncate, ciliate, without lateral lobes; blades 2-7(-11) cm

long, 1-2.8(-4) mm wide, flat, occasionally conduplicate, smooth or scaberulous abaxially, scaberulous

adaxially, margins and midveins not conspicuously thickened, greenish, apices acute, not sharp. Panicles

6-21 cm long, 4-16 cm wide, broadly ovoid, open; primary branches 3-12 cm, capillary, lower branches

spreading 30—90? from the rachises, never appearing fascicled; pedicels 3-14 mm long, longer than the

spikelets. Spikelets 1.2-2.1 mm long, occasionally with 2 or 3 florets; glumes 0.6-1.7 mm long, equal, pur-

plish, scaberulous, particularly on the veins, 1-nerved, apices acute; lemmas 1.2-2.1 mm long, lanceolate to

oblong-elliptic, somewhat plumbeous, glabrous, usually smooth, occasionally scaberulous near the apices,

apices acute, unawned or mucronate, mucros to 0.3 mm long; paleas 1.2-2.1 mm long, lanceolate, glabrous,

acute; anthers 1-1.3 mm long, greenish-yellow to purplish at maturity. Caryopses 0.8-1 mm long, fusiform,

brownish. 2n = 20, 22, 28.

Distribution and habitat.—Muhlenbergia asperifolia grows in moist, often alkaline meadows, beach margins,

and sandy washes, on grassy slopes, and around seeps and hot springs, at elevations of 55-3000 m. This

species is common in western U.S.A. and Canada (Peterson 2003), and is also found in Argentina, Bolivia,

and Chile (Zuloaga et al. 1994). In México M. asperifolia occurs in Baja California, Chihuahua, Durango,

Jalisco, and San Luis Potosí (Espejo Serna et al. 2000).

Comments.—The caryopses of Muhlenbergia asperifolia are frequently infected by a smut, Tilletia asperifolia

Ellis & Everhart, which produces a globose body filled with blackish-brown spores.

Specimen examined. MÉXICO. Coahuila: Municipio de Cuatrociénegas, D la Poza de la Becerra en Cuatrociénegas, J.A

Villarreal Q. 3200, J. Valdés-Reyna & M.A. Carranza P. (ANSM

8. Muhlenbergia depauperata Scribn., Bot. Gaz. 9:187, f. 1-2. 1884. (Fig. 6, A-C). Tre: U.S.A. Arizona: Cochise

Co. or Santa Cruz Co.: M. Mountains, 13 P E n € s.n. mu TOTYPE: US-994221! designated by McVaugh, Fl. Novo-

Galiciana 14:236. 1983, b dicated by Peterson & Annable, Syst. Bot. Monogr.

31:35. 1991; IsSOLECTOTYPES: K!, MO!, MSCII, NYI, PI, USE VAN, W-1916- 27671.

Muhlenl haffneri E. Fourn., Mexic. Pl. 2:85. 1886. Type: MÉXICO. Tacubaya, W. Schaffner s.n. (Lectotype: P! up Hitchcock,

MN US: Natl. Herb. 17:294. 1913; ISOLECTOTYPES: LE (2 sheets)!, MO!, P!, US-3412353!, US-2312354 fragm. ex P

Caespitose annuals. Culms 3-15 cm tall; internodes mostly scaberulous or pubescent, pubescent or strigose

below the nodes, much branched at the lower nodes. Leaf sheaths often longer than the internodes, some-

what inflated, smooth or scabrous, keeled, margins scarious; ligules 1.4-2.5 mm long, membranous, acute,

with lateral lobes; blades 1-3 cm long, 0.6-1.5 mm wide, flat or involute, scabrous to strigose, midveins

and margins thickened, whitish. Panicles 2.5-8.5 cm long, 0.5-0.7 cm wide, contracted; primary branches

1-2.2 cm long, appressed, spikelet-bearing to the base, spikelets borne in subsessile-pedicellate pairs; lon-

ger pedicels 3-6 mm long, scabrous; disarticulation beneath the spikelet pairs at base of pedicel. Spikelets

2.5-4.5 mm long, appressed; glumes 2.3-5.1 mm long, equaling or exceeding the florets; lower glumes

2.3-4 mm long, subulate, 2-nerved, minutely to deeply bifid, teeth aristate or with awns to 1.3 mm long;

upper glumes 3-5.1 mm long, lanceolate, 1-nerved, entire, acuminate; lemmas 2.5-4.5 mm long, narrowly

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951

Fi. 5. Muhlenbergia asperifolia [P.M. Peterson 1703, C.R. Annable, R.F. Thorne & K. Kubitskie (US)]. A. Habit. B. Ligule. C. Glumes. D. Lemma. Muhlenbergia

tenuifolia [PM. Peterson 9705 & A. Campos-Vill (US, ANSM)]. E. Habit. F. Ligule. G. Glumes. H. Floret.

952 Journal of the Botanical R h Institute of Texas 1(2)

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Fic. 6. Muhlenbergia depauperata [PM. Peterson 4082 & C.R. Annable (ARIZ, ENCB, GH, MEXU, MICH, MO, NMC, NY, RSA, TAES, UC, UNLV, UTC, US, WIS, WS)].

A. Habit. B. Glumes. C. Florets. Muhlenbergia fragilis [PM. Peterson 5456 & C.R. Annable (US)]. D. Habit. E. Ligule. F. Glumes. G. Lemma. Muhlenbergia

minutissima [PM. Peterson 5601 & C.R. Annable (US)]. H. Habit. I. Ligule. J. Spikelet. K. Floret.

Peterson et al., Muhlenl ii f T a Mavi T

lanceolate, light greenish-brown to purplish, scabrous, appressed-pubescent on the margins and midveins,

apex acuminate, awned, the awns 6-15 mm long, stiff; paleas 2.4—3.6 mm long, lanceolate, intercostal region

appressed-pubescent, apex acuminate; anthers 0.5-0.8 mm long, purplish to yellowish. Caryopses 1.5-2.3

mm long, narrowly fusiform, brownish. 2n = 20.

Distribution and habitat. —Muhlenbergia depauperata grows in gravelly flats, rock outcrops, exposed

bedrock, and sandy banks, in grama grassland associations, usually on soils derived from calcareous parent

materials, associated with species of Selaginella, Hechtia, Opuntia, Salvia, Brahea, and Juniperus; 1530-2400

m. The range of M. depauperata extends into the southwestern U.S.A., and in México this species occurs in

Chihuahua, Durango, Aguascalientes, México, Distrito Federal, Guanajuato, Jalisco, San Luis Potosí, Puebla,

Veracruz, and Oaxaca (Espejo Serna et al. 2000

Comments.—Muhlenbergia depauperata and M. brevis C.O. Goodd., found commonly in the Sierra Madre

Occidental, share several features with Lycurus: spikelets borne in pairs, 2-nerved and 2-awned lower glumes,

]-nerved and awned upper glumes, acuminate, awned lemmas with short pubescence along the margins,

and pubescent paleas.

Specimens examined. MÉXICO. Coahuila: Municipio de Ocampo, W side of Potrero de la Mula, 20 km NW of Ocampo, I.M. Johnston

9250 (MEXU); Municipio de Saltillo, Sierra Madre Oriental, 18 mi S of Saltillo on México 54 and 1.3 mi W on road to microondas,

= W of Estación Carneros, P.M. Peterson 8425 & M.A. Carranza P. (ANSM, MEXU, US); Municipio de Torreón, Sierra de Jimulco, 3

f Mina San José and 8 km NE of Estación Otto, M.C. Johnston 9552, F. Chiang C.& T.L. Wendt (ANSM). Nuevo León: 6.5 mi S of

en of CHE and Nuevo Leon on Hwy 57 towards Matehuala, P.M. Peterson 13282 & M.B. Knowles (US); 5.2 mi S of Zaragoza on

road towards Ejido La Encantada, P.M. Peterson 16741, J. Valdés-Reyna & M. Sosa Morales (US); Municipio de Santiago, La Escondida, I.

Cabral- Cord 98 (ANS

9. Muhlenbergia distichophylla (J. Presl) Kunth, Enum. Pl. 1:202. 1833. (Fig. 7, A-C). Podosemum distichophyllum

J. Presl, Reliq. Haenk. 1(4—5):231. 1830. Epicampes stricta var. distichophylla J. Presl) M.E. Jones, Contr. W. Bot. 14:6. 1912. Type:

MÉXICO: T. Haenke 173 (HoLotype: PR, photo US!; isotypes: MO-1837831!, US-90711 fragm. ex PRI, US fragm. ex W!, WU).

Meu Mi ia Swallen, N. Amer. Fl. 17(6):457. 1935. Type: MEXICO. JaLisco: rocky hills near Guadalajara, 11 Nov 1889, C.G.

US-822882!; isotypes: LE!, MO-1837815!, US-995828!, US-3274342 fragm!).

Caespitose ME Culms 100-180 cm tall, erect, glabrous to pubescent below the nodes; internodes

glabrous. Leaf sheaths 8-42 cm long, longer than the internodes below, glabrous, the keels prominent,

sometimes coiled to shredded below, basal sheaths compressed-keeled; sheath auricles 0.4-2.6 cm long on

lower portions and up to 6.4 cm long above; ligules 4-15 mm long, membranous, apex finely lacerate some-

times almost to base; blades 18-90 cm long, 2-7 mm wide, flat or folded, scaberulous to scabrous above and

below, the margins and keel saw-toothed. Panicles 35-70 cm long, 4-15 cm wide, densely-flowered, oblong,