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 and copyright.
1993— BRIT becomes publisher/copyright holder.
2007—First issue of J. Bot. Res. Inst. Texas.
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of botanical research and scientific discoveries
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The BRIT Press—bringing out the best in botanical
science for plant conservation and education.
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concerned with classical and modern systematic
botany, sensu lato, for publication in J. Bot. Res. Inst. Texas.
All submissions are peer-reviewed.
Guidelines for submissions are available
from the BRIT Press website, http://www.britpress.org.
BIBLIOGRAPHICAL
Citation abbreviation for the
Journal of the Botanical Research Institute of Texas is
J. Bot. Res. Inst. Texas following the principles
of B.P.H. (informally JBRIT).
International Standard Serial No. (ISSN 1934-5259)
FREQUENCY OF PUBLICATION
J. Bot. Res. Inst. Texas is published semiannually
(summer/fall) as one volume
by the Botanical Research Institute of Texas.
J. Bot. Res. Inst. Texas ISSN 1934-5259
VOLUME 2 NUMBER2 9 DECEMBER 2008
COPYRIGHT 2008
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TABLE OF CONTENTS
SYSTEMATICS
A new hedge-nettle (Stachys: Lamiaceae) from the Interior Highlands of the United States,
and keys to the southeastern species
JoHN B. NELSON
Gutierrezia elegans sp. nov. (Asteraceae: Astereae), a shale barren endemic of southwestern
Colorado
AL SCHNEIDER, PEGGY LYON, AND GUY NESOM
Juncus digitatus (Juncaceae), a new annual rush from Shasta County, California, U.S.A.
CAROL W. WITHAM AND PETER F. ZIKA
Two new subspecies of Sidalcea hickmanii (Malvaceae) in California
STEVEN R. HILL
Ipomoea tehuantepecensis (Convolvulaceae): a new species from the Isthmus of
Tehuantepec, Mexico
LETICIA TORRES-COLÍN, RAFAEL TORRES-COLÍN, MIRIAM P. RAMÍREZ DE ANDA, AND J. ANDREW MCDONALD
Illicium guajaibonense, elevated to species rank and compared with the subspecies
of Illicium cubense (Illiciaceae)
WALTER S. JUDD, J. RICHARD ABBOTT, AND ASHLEY MORRIS
Neocalyptrocalyx morii (Capparaceae), a new species from central French Guiana
XAVIER CORNEJO AND HUGH H. ITIS
A new species of Lasiolaena (Asteraceae: Eupatorieae: Gyptidinae) from Bahia, Brazil
NADIA ROQUE, SILVANA C. FERREIRA, AND HAROLD ROBINSON
Folia taxonomica 9. New species of Passiflora subg. Decaloba (Passifloraceae)
from northern South America
CHRISTIAN FEUILLET AND JOHN M. MacDoucar
Folia taxonomica 10. New species of Nautilocalyx (Gesneriaceae: Episcieae)
from the Venezuelan Guayana
CHRISTIAN FEUILLET
Folia taxonomica 11. Conspectus of Varronia (Cordiaceae: Boraginales) in the Guiana
Shield with three new combinations
CHRISTIAN FEUILLET
A new species of the fern genus Doryopteris (Pteridaceae) from southeastern Brazil
ALEXANDRE SALINO
New records of Poa (Poaceae) and Poa pfisteri: a new species endemic to Chile
ROBERT J. SORENG AND PAUL M. PETERSON
Leptochloa (Poaceae: Chloridoideae) in Colombia
NEIL SNOW, PAUL M. PETERSON, AND DIEGO GIRALDO-CANAS
Eragrostis (Poaceae: Chloridoideae: Eragrostideae) in Colombia
PAUL M. PETERSON AND DIEGO GIRALDO- CANAS
Bouteloua (Poaceae: Chloridoideae: Cynodonteae: Boutelouinae) del noreste de México
YOLANDA HERRERA ARRIETA, PAUL M. PETERSON Y Jesus VALDÉS-REYNA
Revision of Tocoyena (Rubiaceae: Gardenieae) from the states of Goiás and Tocantins
and a new species endemic to white-sand areas in the Brazilian Cerrado
PIERO G. DELPRETE
761
771
775
783
793
799
807
811
817
825
837
843
847
861
875
917
983
Cinco nuevas especies de Pilea (Urticaceae) de Costa Rica
ALEXANDER RODRÍGUEZ Y ÁLEXANDRE K. MONRO
The Diospyros salicifolia complex (Ebenaceae) in Mesoamerica
MITCHELL C. PROVANCE, IGNACIO GARCÍA RUIZ, AND ANDREW C. SANDERS
A taxonomic revision of Crataegus series Lacrimatae (Rosaceae)
J.B. PuiPPs AND K.A. DvorskY
Dicerandra modesta (Lamiaceae): raise in rank for a disjunct perennial in a new
coastal clade in Florida
Rosin B. Huck
Two new names in the Caribbean flora
ALAIN H. LIOGIER
Trochanthera lepidota gen. and sp. nov., a fossil angiosperm inflorescence in Dominican amber
GEORGE O. POINAR, JR., KENTON L. CHAMBERS, AND ALEX E. BROWN
Cretacifilix fungiformis gen. and sp. nov., an eupolypod fern (Polypodiales) in Early
Cretaceous Burmese amber
GEORGE O. POINAR, JR. AND RON BUCKLEY
An Early Cretaceous angiosperm fossil of possible significance in rosid floral diversification
GEORGE O. POINAR, JR., KENTON L. CHAMBERS, AND RON BUCKLEY
A phylogeny of Bejaria (Ericaceae: Ericoideae) based on molecular data
CATHERINE M. BUSH AND KATHLEEN A. KRON
Understanding Oligactis (Asteraceae: Liabeae): the true identity of O. sessiliflora
and O. volubilis
DIEGO G. GUTIÉRREZ
Senecio cymbalarioides, S. subnudus, and S. subpeltatus, history of a muddle
(Asteraceae: Senecioneae)
J.F. VELDKAMP
2C DNA content values in Amaranthus (Amaranthaceae)
DONALD B. PRATT, SHALINI N. JHANGIANI, AND ROBERT J. WIGGERS
Minuartia macrantha (Alsinoideae: Caryophyllaceae): morphological circumscription,
geographical range, and phylogenetic affinities
RONALD L. HARTMAN AND RICHARD K. RABELER
Correct basionym and combination for Pappostipa barrancaensis (Poaceae)
KONSTANTYN ROMASCHENKO
Los encinos (Quercus: Fagaceae) de Coahuila, Mexico
José A. VILLARREAL Q., JUAN A. ENCINA D. Y MIGUEL A. CARRANZA P.
Thomas Walter Typification Project, VI: neotypes for an additional 18 Walter names
DaNiEL B. WARD :
BOTANICAL HISTORY
Caryophyllaceae in North America: change over the last 110 years
RicHARD K. RABELER AND RONALD L. HARTMAN
The Big Thicket: pristine wilderness or archeologically understudied?
MicHAEL H. MACROBERTS AND BARBARA R. MACROBERTS
995
1009
1101
1163
1165
1167
1175
1183
1193
1207
1215
1219
1225
1233
1235
1279
1285
1291
ETHNOBOTANY
Diferencias en el uso de plantas entre hombres y mujeres en una comunidad de pie
de monte del norte del Ecuador
Davip SUÁREZ DUQUE
METHODS AND TECHNIQUES
Tree canopy research and student experiences using the doubled rope climbing method
CourTNEY M. Kircong, HAROLD W. KELLER, SYDNEY E. EVERHART, ANGELA R. SCARBOROUGH,
KENNETH L. SNELL, MELISSA S. SKRABAL, CHARLY POTTORFF, AND JOSEPH S. ELY
FLORISTICS, ECOLOGY, AND CONSERVATION
Aristolochia schultzeana (Aristolochiaceae) new to Ecuador
DoucLAs H. GOLDMAN
Asclepiadoideae (Apocynaceae) como maleza en el cultivo del agave tequilero
(Agave tequilana var. *azul")
IRMA LóPEz-MuRAIRA, RUBÉN IRUEGAS, IsAAC ANDRADE GONZÁLEZ Y JUAN FLORENCIO GÓMEZ LEYVA
The species of Fumaria (Papaveraceae) in Texas, including F. densiflora, first records
from North America
Monique REED, RICHARD V. LANSDOWN, AND Tim C.G. RICH
Noteworthy collections from the southeastern United States
BRUCE A. SORRIE AND RICHARD J. LEBLOND
Analysis of the completeness of vascular plant records in Florida
Justin K. WILLIAMS AND ANICA DEBELICA
Species richness of vegetational areas of Texas: a first approximation
MicHAEL H. MACROBERTS AND BARBARA R. MACROBERTS
Carex breviculmis (Cyperaceae), new to the flora of North America
Lucas C. MAJURE AND CHARLES T. BRYSON
Vascular flora of C. Bickham-Dickson/Red River Education and Research Park, Caddo Parish,
Louisiana: an oxbow lake community, with comments on exotic/native species ratios
MicHAEL H. MAcRoBERTS, BARBARA R. MACROBERTS, AND GARY M. HANSON
A floristic survey of vascular plants over parts of northeastern New Mexico
MICHAEL H. SCHIEBOUT, DONALD L. HAZLETT, AND NEIL SNOW
Checklist of the vascular plants of Fayette County, Pennsylvania
CYNTHIA M. MORTON AND LOREE SPEEDY
Rare and endemic plants of the Big Thicket: a phytogeographical analysis
MICHAEL H. MACROBERTS AND BARBARA R. MACROBERTS
Annotated checklist of the vascular flora of the Loblolly Unit of the Big Thicket National
Preserve, Liberty County, Texas
Larry E. Brown, BARBARA R. MACROBERTS, MICHAEL H. MacRobErTS, WARREN W. PRUESS,
I. SANDRA ELsIK, AND SUZANNE BIRMINGHAM WALKER
Neptunia plena (Fabaceae: Mimosoideae) rediscovered in Texas
ALFRED RICHARDSON AND KEN KING
Asclepias hirtella (Apocynaceae) newly documented for the flora of Texas
Matt WHITE
Lilaeopsis carolinensis (Apiaceae) a species new to Texas and a key to Lilaeopsis in Texas
STEPHAN L. HATCH AND A. TUCKER SLACK
1295
1309
1337
1341
1347
1353
1363
1373
1381
1389
1407
1449
1475
1481
1491
1495
1497
Book Reviews AND Notices 770, 782, 792, 798, 816, 860, 982, 994, 1008, 1166, 1174, 1206, 1214, 1224, 1232,
1234, 1284, 1290, 1294, 1340, 1346, 1352, 1362, 1372, 1380, 1388, 1448, 1480, 1490
ANNOUNCEMENTS 1494
REVIEWERS FOR VOLUME 1 (2007) ano Voume 2 (2008)—1499
INDEX TO VOLUME 1 (2007) AND VoLume 2 (2008)
Titles of Articles with Authors—1501
Authors—1507
Botanical Names and Subjects—1509
New Names and Combinations—1523
INDEX to new names and new binations in J. Bot. Res. Inst. Texas 2(2), 2008
Bouteloua tamaulipensis G.J. Pierce ex D. Pacheco & Columbus, sp. nov.—966
Crataegus lancei J.B Phipps, sp. nov.—1143
Cretacifilix G.O. Poinar & R. Buckley, gen. nov.—1176
Cretacifilix fungiformis G.O. Poinar & R. Buckley, sp. nov.—1176
Dicerandra modesta (Huck) Huck, comb. et stat. nov —1163
Diospyros acapulcensis subsp. chiquimulensis M.C. Provance, I. García & A.C. Sanders, subsp. nov—1025
UOS o acapulcensis r dwyeri M: C. Provance, I. García & A.C. Sanders, subsp. nov.—1027
is M.C. Provance, I. García & A.C. Sanders, subsp. nov.—1029
Diospyros acapulcensis subsp. mejocotensis M.C. Provance, I. García & A.C. Sanders, subsp. nov.—1031
Diospyros acapulcensis subsp. nicaraguensis (Standl.). M.C. Provance, I. García & A.C. Sanders, comb.
et stat. nov.—1033
Diospyros acapulcensis subsp. pedromorenoi M. C. Provance, I. García & A.C. Sanders, subsp. nov.—1036
Diospyros acapulcensis subsp. rivensis M.C. Provance, I. García & A.C. Sanders, subsp. nov.—1039
Diospyros acapulcensis subsp. veraecrucis (Standl.) M.C. Provance, I. García & A.C. Sanders, comb. et
stat. nov.—1044
Diospyros aequoris subsp. balsensis M.C. Provance, I. García & A.C. Sanders, subsp. nov.—1061
Diospyros aequoris subsp. chutlensis M.C. Provance, I. García & A.C. Sanders, subsp. nov.—1065
Diospyros aequoris subsp. martineziana (Standl. ex Leavenworth) M.C. Provance, I. García & A.C. Sand-
ers, comb. et stan nov.—1069
i i (Standl.) M.C. Provance, I. García & A.C. Sanders, comb. et stat. nov.—1071
Diospyros aequoris subsp. tehuantepecensis M.C. Provance, I. García & A.C. Sanders, subsp. nov.—1074
Diospyros yucatanensis subsp. spectabilis (Lundell) M.C. Provance, I. García & A.C. Sanders, comb. et
stat. nov.—1086
Doryopteris jequitinhonhensis Salino, sp. nov—-843
Gutierrezia elegans Al Schneider & P Lyon, sp. nov.—771
Hedyotis ekmanii Alain, nom. nov.—
Illicium guajaibonense (Imkhanitskaya) Judd & Abbott, comb. et stat. nov.—800
Ipomoea tehuantepecensis L. Torres-Colín, R. Torres-Colín, M.P Ramírez & J.A. McDonald, sp. nov. —793
Juncus digitatus C. Witham & Zika, sp. nov. —775
Lachnociona G.O. Poinar, K.L. Chambers & R. Buckley, gen. nov.—1184
Lachnociona terriae G.O. Poinar, K.L. Chambers & R. Buckley, sp. nov.—1184
Lasiolaena lychnophorioides Roque & H. Rob., sp. nov—812
Nautilocalyx crenatus Feuillet, sp. nov.—825
Nautilocalyx orinocensis Feuillet, sp. nov.—833
Nautilocalyx paujiensis Feuillet, sp. nov.—827
Nautlocalya pusillus Feuillet, sp. nov.—827
N roseus Feuillet, sp. nov.—830
Nautilocalyx ruber Feuillet, sp. nov. —830
Nautilocalyx vestitus Feuillet, sp. nov.—833
Neocalyptrocalyx morii X. Cornejo & H.H. Iltis, sp. nov.—807
Pappostipa barrancaensis (EA. Roig) Romaschenko, comb. nov.—1233
Passiflora angusta Feuillet & J.M. MacDougal, sp. nov. —818
Passiflora rufa Feuillet & J.M. MacDougal, sp. nov.—822
Pilea alfaroana Al. Rodr. & A.K. Monro, sp. nov.—995
Pilea gamboana Al. Rodr. & A.K. Monro, sp. nov.—997
Pilea herrerae Al. Rodr. & A.K. Monro, sp. nov.—1000
Pilea longibracteolata Al. Rodr., A.K. Monro & L. Acosta, sp. nov.—1002
Pilea moragana Al. Rodr. & A.K. Monro, sp. nov. —1004
Poa pfisteri Soreng, sp. nov. —850
Rosaceae subseries Robustae j.B. Phipps, subser. nov. —1124
Rosaceae subseries Tenues J.B. Phipps, subser. nov.—1108
Salvia toaensis Alain, nom. nov.—1165
Sidalcea hickmanii subsp. napensis S.R. Hill, subsp. nov. —783
Sidalcea hickmanii subsp. pillsburiensis S.R. Hill, subsp. nov. —786
Stachys iltisii J. Nelson, sp. nov.—761
Tocoyena arenicola Delprete, sp. nov.—988
Trochanthera G.O. Poinar, K.L. Chambers & A.E. Brown, gen. nov.—1168
Trochanthera lepidota G.O. Poinar, K.L. Chambers & A.E. Brown, sp. nov.—1168
Varronia bullata subsp. humilis (Jacq.) Feuillet, comb. nov.—837
Varronia cremersii (Feuillet) Feuillet, comb. nov.—838
Varronia marioniae (Feuillet) Feuillet, comb. nov.—839
A NEW HEDGE-NETTLE (STACHYS: LAMIACEAE)
FROM THE INTERIOR HIGHLANDS OF THE UNITED STATES,
AND KEYS TO THE SOUTHEASTERN SPECIES
John B. Nelson
A. C. Moore Herbarium
Department of ar Sciences
niversity of South Carolina
Columbia, South Carolina MM U.S.A.
ABSTRACT
A new and geographically restricted species in the “hedge-nettle” genus, Stachys (Lamiaceae), is described from the Interior Highlands
f +1 AT Jc c 4 C E fra A +] ] Abla} es ro 1 re | Jç, Fa 1i Ales 1 Pag 1 1 1
r
nettles.
ZUSAMMENFASSUNG
Eine neue Art in der Ziest-G g (Stachys, Lamiaceae), mit kleinráumiger OREL. im Inneren re von Aa und Okla-
homa, wird | hriel S I y iltisii. Zwei Schlú
Stachys is one of the larger genera within the Lamiaceae, comprising nearly 300 species, and nearly cosmo-
politan, absent from Australasia. Its centers of species diversity include warm-temperate portions of western
Asia and the Mediterranean, southern Africa, and portions of North and South America (ilcim et al. 2008).
The genus in North America, north of Mexico, contains about 45 species, particularly concentrated west of
the Rocky Mountains (especially the Pacific states) and in the east, along the Atlantic seaboard states and
in the Appalachian region. Southeastern species occur in a wide variety of habitats, most often in mesic
sites, and at a broad range of elevations. The North and iban aC species were studied in detail by
Epling (1934); Mulligan and Munro (1989) provide a t ofthe North American taxa
north of Mexico. Ongoing studies in the genus show that an additional species should be recognized.
Stachys iltisii J. mal sp. nov. (Figs. 1-3). Te: UNITED STATES. Arkansas. Logan Co.: crowded population on rocky
, S side of Lodge Drive, 1.5 mi (2.4 km) W of AR Highway 309, near top of Magazine Mountain, 18
j
SIOUN 111 Ur
Jun 2007, J.B. Nelson 26650 (noLorvee: USCH; rsorvees: APCR, BRIT, E FSU, NCU, NY, OKL, OKLA, STAR, UARK, US, WIS).
Species propria, h ] bund landulif 1 in laterit li t guli jue folia pubentissi Caules
et folia, contusi, édsletit muschate. Est similis. entire sed differt quod habet mai liquant iora folia quae inferiore pagina
tam glandulifera non sunt, et quod habet flores paulo minores, haec p I pactiora i icillis aggregatis. Crescit in Okla-
oma et Árkansase.
Potentially robust perennial herbs to 1 m tall or taller from vigorous, pale, fragrant rhizomes, often in dense
colonies; stems erect, occasionally branched, older plants commonly branched; stem sides and angles
densely pubescent with a mixture of glandular and eglandular hairs, the glandular hairs capitate, the eg-
landular hairs with 3 or more cells, to 3 mm long, spreading and reflexed; leaves spreading, somewhat lax,
densely pubescent on both surfaces with glandular and eglandular hairs, the latter mostly appressed on
upper surface, and erect on lower surface, the lower surface equipped as well with scattered sessile (or
nearly sessile) capitate glands; the crushed leaves with a musky walnut-spice-fish scent; mid-stem leaves
4-6 cm broad, 10-17-19) cm long, commonly lyrate basally, featuring a somewhat narrowed waist, with
distally convex margins; inflorescences commonly elongated at maturity (fruiting), the lowest and oldest
several verticils usually slightly separated, the more distal and younger verticils more or less crowded to-
gether, thus nearly compact; cymules each with 6—8-flowers, the verticils thus 12-16-flowered, mostly
(turbinate)-hemispheric to nearly spherical; open flowers are visited by a variety of insects, including hon-
J. Bot. Res, Inst. Texas 2(2): 761 — 769. 2008
——— ———— 7 "NP T : i
both i LEFT: Stachys iltisii (J. Nelson 26653, LeFlore C ty OK, USCH) showing few glandular hairs. RIGHT: Stachys eplingii (H. Allard
19940, Fauquier C MAECEN OQ Kin HN DÉC DA pa eus
ic Di nae a Ef Ft AY b 2 J
eybees, small bumblebees, and wasps; calyces campanulate, densely pubescent with glandular and eglan-
dular hairs, tube (3.223.8—4 mm long, the lobes less that one-half the length of the tube, 1.5-2 mm long,
deltoid, minutely apiculate; corollas pink, 11-11.5 mm long from base to tip of galea, galeae frequently
notched or indented apically; the lower lip generally declined 90° at full anthesis, more so with age, 6 mm
long, featured prominent pink-purple blotches and spots on adaxial surface; corolla tube prominently sac-
cate toward base on lower side, internally glabrous, but with prominently slanting (oblique) annulus, this
copiously pubescent with soft, bulbous trichomes; mericarps 1.5 mm wide, 1.5-1.6 mm long, dark brown,
irregularly and minutely verrucose.
Additional specimens seen: UNITED STATES. ARKANSAS: Crawford Co.: 5. Burgess 102, McMinn Gap, Forest Service Rd 1003
(UAM, UARK). Franklin Co.: L. Kirschberger 109, 14 Jun 1996, Ozark National Forest ddl: Garland Co.: D. Demaree 56064, 30 May
1967, rocky hillsides on Hickory Nut Mountain (BRIT); and d date, D 8 (ISC). Izard Co.: F. Pennell 10686,
1 Jun 1920 (NY). Logan Co.: D. Demaree 17743, 18 Jun 1938, Magazine Mountain (BRIT, FSU, WVA); H. Iltis 5048, 17 Jun 1954, Maga-
zine Mountain (UARK); H. Iltis 5374, 9 Jun 1955, ravine on N slope of Magazine Mountain (BRIT, GH, WIS); E. Palmer 23214, 11 Jun
1923, slopes of Magazine Mountain (MO, UARK); H. Pyle 796, 15 Jun 1937, MEER Mountain ED G. Tucker 15499, 23 Jul 1979,
Magazine Mountain, 1.5 mi W of Green Field (APCR); G. Tucker 22898, 10 Jul 1983, } UAM). Montgomery Co.
Demaree 45661, 10 Jun 1962, Hickory Nut Mountain (GH); J. Miller 187, 24 Jun 1970, iud on High Peak (BRIT, UARK); J. Roberts 273,
26 May 1977, Winding Stairs, E side of Little Missouri River (UARK); E. Sundell 2351, 28 May 1981, Little Missouri Falls (FLAS, NLU,
UAM, UARK); R. Thomas 100517, 27 Jun 1987, Collier Springs nuni Newton Co.: A. McWilliam 25, 19 Jun 1969, roadside between
Kingston and Boxley (UARK). Perry Co.: 5. Walker s.n., 8 Jun 1995, N f Ced (UARK). Polk Co.: D. Demaree 23183, 13 Jun
1942, Rich Mountain (BRIT, GH, MIN); D. Demaree 54519, 19 Sep 1966, Rich Mountain (BRIT); H. Iltis 5175, 20 Jun 1954, ae
sandstone, 8 mi W of Mena (UARK); A. McWilliam 482, 1 Jul 1955, 9 mi W of Mena (UARK); J. Nelson 26652, Rich Mountain, 0.65 m.
of Queen Wilhelmina State Park at AR Hwy#272 (NBYC, UARK, USCH); J. Nelson 26654, 19 Tune 2007, Rich Mountain at Pioneer Cem-
etery, 3 mi E of OK state line (CLEMS, STAR, USCH); J. Nelson 26655, 19 Jun 2007, Rich M , 2 mi E of AR Hwy#272
(APCR, FMUH, USCH); J. Nelson 26657, 20 Jun 2007, shady Creek, 2.5 mi E of Shady (town) (USCH); J. Nelson 26658
20 Jun 2007, shady d t , 4.1 mi E of Shady (town) on AR Hwy 425 (USCH). OKLAHOMA: LeFlore € Co.: M. Hopkins
574, 21 May 1944, meadow at base of Rich Mountain (OKL); J. Nelson 26653, 19 Jun 2007, 1.25 mi W of AR state line on OK Hwy #1
(OKL, OKLA, USCH); E. Palmer 20615, 23 Sep 1921, wooded slopes of Rich Mountain (F, NY); E. Palmer 21642, 1 Jun 1922, N slopes of
Rich Mountain (NY); U. Waterfall 17005, 31 May 1962, 10 mi SE of Stapp near Arkansas border (OKLA); F. Means, Jr. 3571, 15 Jun 1968,
N slope of Rich Mountain, 0.25 mi W of Arkansas border (OKLA); J. and C. Taylor 32462, 12 Aug 1984, State Hwy £1, 6.8 mi E of its
Nelson, A i FCtarl f Lx its: TTG l- afshan lic 763
junction with US 259 (BRIT). McCurtain Co.: G. Goodman 8360, 11 Jun 1973, S end of Beaver Bend State Park (OKL); M. Hopkins 380,
20 May 1944, Beaver Bend State Park (OKL).
Stachys iltisii is endemic to the Interior Highlands of Arkansas and Oklahoma. The oldest known collection
is that made by E. J. Palmer in 1920, from Izard County, Arkansas, which represents the eastern-most loca-
tion for this plant. Its first recognition in the literature was made by Epling (1934), who considered it, ad-
ditionally known by then from collections in Oklahoma, as a thin- Kar ed and unnamed variant of Stachys
nuttallii Shuttlew. ex Bentham. Fernald (1950), despit bl with Epling's assessments
on Stachys in general (1943), concurred that the plants of the Interior Bold belong in S. nuttallii, as did
Gleason (1952). Hugh Iltis (WIS) by way of annotation on the original label of his #5374 (BRIT) from
Magazine Mountain, Logan County, AR on 9 Jun 1955, suggested that these plants rightly constitute a
recognizable entity within S. nuttallii, offering "ssp. nov. arkansanus" for it. Nelson and Fairey (1979) es-
sentially concurred with Epling in recognizing these plants as distinctive within the S. nuttallii complex,
but maintained them as being a part of a distinctive and new taxon, S. eplingii; they treated the plants of the
Interior Highlands as forming the western (and disjunct) end of the range of S. eplingii (the eastern distribu-
tion consisting of primarily Appalachian portions of West Virginia, Virginia, Maryland, District of Colum-
bia, and historically, North Carolina). The somewhat conservative approach to the taxonomic status of these
plants taken by Nelson and Fairey at the time is evident within the protologue of their description of S.
eplingii: they list (1979, p. 493) three specimens from Arkansas as paratypes for S. eplingii (Demaree 17743
from Logan County; Miller 187 ... “Smith” was used for this collection, but in error... from Montgomery
County; Itis 5175 from Polk County). Their listing these paratypes for a different taxon has no bearing on
the matter at hand, and these three collections presently serve as paratypes for S. iltisii (see above).
This taxonomic position was maintained by Nelson (1981), and by Mulligan and Munro (1989; whose
formulation is the most recent treatment for all the North American species of Stachys), by Gleason and
Cronquist (1991), and finally by Smith (1994).
Recent field investigation in Arl d Oklahoma (during one of the hotter and stormier, lightning-
filled summers on record, 2007) and study of considerably more herbarium material since 1979 reveals that
the opinion expressed by Epling and Iltis was legitimate, at least in the sense that the plants of the Interior
Highlands are distinctive within the S. nuttallii “complex.” There is no question that these plants share many
features with S. eplingii, and the two entities are surely related. d differences between these two
species are apparent, and based on more recent collections, it is obvious that the two taxa can be consis-
tently separated, both from fresh material, and from herbarium specimens. Both species bear short-stalked,
capitate glands on the abaxial leaf surface, with unicellular, spheroidal heads, 25-30 um in diameter. The
lower leaf surface of S. iltisii is sparingly glandular, while that of S. eplingii is densely and prominently
atomiferous-glandular (Fig. 1), to the point of having a somewhat shiny, golden surface (with magnification)
below the non-glandular pubescence, as well as often leaving something of a yellowish, slightly oily stain
on newspapers when pressed. The leaf blades of S. iltisii are longer, broader, and somewhat thinner in texture
than those of S. eplingii. Additionally, the leaf blades of S. iltisii are frequently somewhat narrowed in the
lower third, thus somewhat lyrate, and widest toward the middle. Stachys eplingii has more ovate leaves,
usually widest in the lower third. In S. iltisii, the apex of the early blooming stem is relatively delicate and
somewhat flexuous, while that in S. eplingii is stiffer. The fruiting inflorescence of S. iltisii is commonly
compact, 1.5-2cm between the lowest verticils, whereas that of S. eplingii commonly exhibits more space,
(2223—4cm between the lowest verticils.
Stachys eplingii is found in boggy places within the Appalachians. Its most common habitat from West
Virginia into northern Virginia includes wet, saturated meadows. Stachys iltisii is generally found in upland
forested habitats, commonly on thin soil of rocky places, in both the Ozark Plateau and the Ouachita Moun-
tains, thus on both sides of the Arkansas River Valley, though probably more widely distributed in the
Ouachitas. Although substantial differences in geology and soil types exist between the Ozark Plateau and
the Ouachitas, sites occupied by S. iltisii do not appear to be unusual or particularly special, and in fact,
" ] Vafého Rataniral D h Institute of Texas 2(2)
NEN "m"
LE AS I S
E RIS,
ERE RON gs
open roadsides tend to make excellent habitat for it, where it frequently grows with weedy plants such as
Coronilla varia, Daucus pusilla, and Ambrosia trifida. Common woody associates, in less disturbed sites include
Quercus alba, Acer rubrum, Robinia pseudo-acacia, Juglans nigra, Ostrya virginiana, and Sassafras albidum.
Nearly every population located during this field effort grew with Toxicodendron radicans, Parthenocissus
quinquefolia, and Rubus occidentalis.
An overview of biological endemism in the Interior Highlands is not the purpose of this paper. This
subject has been amply addressed for a considerable number of plants and animal species, and is discussed
in the recent literature; Robison and Allen (1995) provide a useful summary. Zollner et al. (2005) provide
an enumeration of the vascular plant taxa endemic to the Interior Highlands (24 species plus 12 subspecies
or varieties of more widely distributed species). Vicarious distributions of plant taxa between the Interior
Highlands and the Appalachians have been long recognized: Steyermark (1934) lists 87 vascular taxa of
the Ozark Plateau that also occur in the southern Appalachians. "Classic" examples include the distributions
of Magnolia tripetala, Cladrastis lutea, Robinia pseudo-acacia, and perhaps Halesia carolina (Little 1970). The
common notion regarding disjunction of the same species, or of two related species, presumes some prob-
able interconnection in the past (Hardin & Cooper 1967), followed by potential evolutionary divergence.
Vicariance and subsequent speciation has been demonstrated for fish species forming species “pairs” in the
Ozarks and the Appalachians (Strange & Burr 1997), and various examples exist for vascular plants, such
as Castanea ozarkensis and C. pumila. In the case of Stachys iltisii and S. eplingii, such potential commonality
of distribution no longer exists, and the two entities are now fully separated. Approximately 1000 km (640
mi) currently separate their nearest points of distribution. Based upon the isolation of the plants within the
Interior Highlands from all other related hedge-nettles, and upon the ease of separating plants based on
simple morphology, Stachys iltisii is here recognized and described at the level of species.
Personnel within the Arkansas Natural Heritage Commission (2005) list Stachys iltisii as S. eplingii (thus
following Nelson & Fairey 1979), and have assigned its state rank as “INV” (Inventory Element, “of conser-
vation concern”). The Oklahoma Natural Heritage Inventory (2003) similarly maintains 5. iltisii as 5. eplingii,
tracking it as a rare species. At least 40 known populations are verified for 13 counties in Arkansas and
Oklahoma. Reports of additional county occurrences in Arkansas (Howard and Scott Counties) but not on
specimens seen from them, is consistent with the known range (Fig. 2).
Reference to this species as an element of the flora of the Southeastern United States comes from its
presence in Arkansas, a politically delimited area generally recognized by botanists as a portion of the
Nelson, A new species of Stachys from the Interior Highlands of the US
Fic. 3. Stachys iltisii: living plants (population of Nelson 26653, LeFlore County OK) with unidentified pollinator (?) at lower left.
766 Journal of the Botanical R h Institute of Texas 2(2)
“Southeast” Although Oklahoma is not a Southeastern state, its eastern portions do indeed harbor many
"Southeastern" species.
*Ozark hedge-nettle" may be an appropriate common name. This is a handsome species, often robust
and conspicuous (Fig. 3), and fairly easy to spot from a moving vehicle. Due to their tendency toward a
somewhat np Ione Scene, especially in the sun, the plants do resemble Teucrium canadense...but
then, various Stach ly confused with “Canada germander,” frequently causing field-trip
r 4
turn-arounds.
Etymology.—Iltis unofficial differentiation of this plant as a new taxon deserves recognition. Its occur-
rence in Oklahoma as well as Arkansas, however, makes the choice of the epithet “arkansana” (Stachys as a
noun requires a feminine ending) unwarranted. As a way of commemorating Hugh Iltis’ interest in the
vegetation of the Interior Highlands, and in the genus Stachys, and as well acknowledging his tremendous
contributions to botany, this taxon is thus named in his honor.
Except that Stachys iltisii and S. eplingii are (were?) never really common, and that there is a relatively
low number of historic collections of each, it is somewhat remarkable that Epling (1934) did not consider
formally recognizing them in 1934. On the other hand, Stachys in the Southeastern United States has com-
monly been considered a troublesome genus. Its current taxonomy is fairly stable, although several of the
characters (e.g., stem pubescence, petiole length, and calyx lobe dimensions) useful in separating taxa are
variously continuous. The considerable variation in some species groups suggests future usefulness to be
attained from cytological and breeding system studies.
Nevertheless, practical taxonomy demands results. To this end, two keys are provided for distinguish-
ing Stachys iltisii from the other Southeastern species. The first uses relative petiole length as a primary
distinction, the second using shape of the calyx lobes. The different structuring of these keys is provided to
emphasize different characters; with a particular plant or specimen in hand, the user may find that one or
the other key provides the easier route to identification.
SOUTHEASTERN STACHYS: KEY #1
1. Annual; corolla scarcely exserted from calyx; LA-MS-AL-FL-SC S. agraria
1. Perennial; corolla prominently exserted from calyx, widespread.
2. Petioles obvious (at least some of those in the middle portion of the stem at least 1/5 as long as the leaf
blade).
3. Calyx tubes glandular.
4. Leaf blade d dentate or nearly so; stem a gles copiously pubescent with long (to 3 mm),
spreading hai
4. | xa blade margins mostly crenate; stem angles glabrate or pubescent with mostly short, retrorse
S. clingmanii
air
5. 9T blades «3 cm wide; plants perniciously weedy, commonly in dense patches, scarcely to .75
m p Lu white, idis tubers Termina ung rhizomes S. floridana
5. Lea cm | t weedy, never in dense patches, commonly 1m tall
of str rounded tubers dos sent.
eaf blades ovate, the bases cordate; margins crenulate; IN- on -WV-KY- VA S. cordata
E pes rd Es ic-oblong, the bases rounded to slightly ; g te to serrate;
A-AL- S. nuttallii
3. Calyx not ona or wey slightly SO.
7. Calyx glat tioles well-developed, especially in shade forms S. tenuifolia
7. Calyx variously haity, but: at ca hispidulous, oy strongly hispid; petioles short or long.
8. Petioles usually w g y pubescent spreading hairs; blade
margins commonly dentate S. clingmanii
8. p short to nearly pad stem angles commonly pubescent, with retrorse hairs; blade
argins serrate, but not den S. hispida
2. Petioles ne. (the midstem leaves is EE less than 15mm) or absent
9. Leaves linear-lanceolate to narrowly lanceolate, usually widest at or near T base; leaf margins entire
to crenulate, rarely serrulate, and then mostly toward the apex.
Nelson
10. Leaf blades abundantly pubescent below with appressed hairs, thus closely tomentose, the lower
surface felty or velvety; corolla white, pink, or purplish.
11. Stems angles and sides abundantly soft pubescent, with mostly soft, spreading hairs, corolla
white; rare adventive (SC)
767
S. pilosa
11. Stem angles and sides abundantly pubescent, the angles equipped with at least some stiff,
retrorse hairs, corolla purple; rare adventive (AL)
S. palustris
10. Leaf blades variously pubescent or glabrate, but never felt rolla pink.
12. Leaf blades narrow, 3-6 mm wide; plants generally aes to a pubescent; blade
margins entire to obscurely Ec S.
12, Leaf blades broader, E G mm x APIAU US or at least moderately pube-
cent, or abundantiy late t
hyssopifolia
S. aspera
9. Leaves ovate to elliptic, MESA near the center or toward ne apes ee leaf margins crenate to
sharply serrate for nearly the entire length.
13. Stem sides commonly pubescent above.
14. Calyx lobes lanceolate; flower 6 per vertical; leaf margins serrate-crenate.
15. Leaves generally subsessile, the petioles thick, not longer than 6mmy blades densely pubes-
cent, frequently soft-pilose to scaberulous; leaf margins serrate; introduction from farther
north and west
15. Leaves generally petioled up to 15mm long, although commonly shorter, and relatively
aie blades pubescent on both surfaces, but never to the point of being pilose or
rulous; leaf margins crenate-serrulate; KY-TN-GA-AL-SC
S. arenicola
S. nuttallii
14. es robes deltoid; flowers s or more per verticil; leaf margins crenate-crenulate; native s with
bipolar distribution, OK-AR, WV-MD.
16. Lower leaf surface E sessile- and stipitate-glandular, blades ovate, widest in lower
third of leaf; adjacent upper verticils separated in fruit; WV-VA-MD-DC (NC)
S. eplingii
16. Lower leaf surface on but not densely so; blades elliptic-ovate, widest at middle;
adjacent upper ils commonly crowded in fruit; OK-AR
S. iltisii
13. Stem sides glabrous, the angles uem
17. Fruiting calyx lobes lanceolate, hispidulous to hispid, about half as long as the calyx tube
17. Fruiting calyx lobes deltoid or triangular, shorter than half the length of the calyx tube.
18 | I 1 | | | | P | h.l J H J J
2 D rd + t A $ d mi Á/ i)
from lowest flowering node, leaf blades elliptic, rounded to truncate at base; Blue Ridge
(VA)
18. Leaf gi te; teeth oft | |
from lowest flowering node; leaf blades oblong-elliptic; Blue Ridge o: upper piedmont,
VA-NC-TN-SC-GA
S. hispida
S. subcordata
S. latidens
SOUTHEASTERN STACHYS: KEY #2
1. Fruiting calyx lobes deltoid to broadly triangular, mostly «1/2 tube length.
2. Stem sides pubescent; foliage glandular, often with musky scent when crushed.
3. Petioles short to Wen Rd masiy raungan ish truncate- cordaire.
4. Lower leaf surf. g g |
ingly ovate, broadest toward base; W- VA-MD (N
S. eplingii
4. Lower surface glandular, but not densely; ae prevailing elliptic, somewhat lyrate toward base,
broadest toward midblade; AR-OK
S. iltisii
3. Petioles well developed, blades commonly cordate at s pase
J ra
5. Top of blooming stem frequently flexuous, somewhat rounded, IN-OH-WV-KY-VA-
NC
S. cordata
Top of blooming stem stiffish, not lax; blades oblong-elliptic; KY-TN-GA-AL-SC
S. nuttallii
2. "e sides glabrous; crushed fresh foliage with grass-like scent, not musky.
6. Leaves sessile to subsessile; bracts rapidly reduced upward (Appalachians, GA-WV)
6. Leaves subsessile or petioles to 10mm long; bracts gradually reduced upward (Blue Ridge, VA) — . S.
—
. Fruiting calyx lobes lanceolate, narrowly triangular, or nearly subulate, half as long as tube (or > 1/2 as long).
7. Leaf blades linear to lanceolate.
Corolla white; stem sides with at least moderate pubescence on the highest sterile internode,
8.
velvety-pilose (SC
moderately to densely pubescent on higher i ; lower leaf surface abundantly pubescent to
S. latidens
subcordata
S. pilosa
768 | of the Botanical R h Institute of Texas 2(2)
8. Corolla pink; stem sides without pubescence, except for internodes within the inflorescence, which
may bear light villous or glandular Nails: oe leaf surface glabrous or pubescent, but not velvety.
9. Leaf blade margins enti lly glab S. hyssopifolia
9. Leaf blade margins serrulate with at leas a few teeth pans ense or pubesce
10. Stems strict or sparingly | barely petioled, the blades nee to serrate
with shallow tee th S. aspera
10. Stems frequently branched from the upper nodes; leaves obviously petioled, the blades
sharply toothed . tenuifolia
7. Leaf blades wider, rounded bd to pu
1 l. P etioles poorly tially GVSETIL
12. Upper stem sides bos S. hispida
12. Upper stem sides vidus pubescent, glandular and/or eglandular.
13. Corollas purple; rare adventive (AL S. palustris
13. Corollas pink; pate species.
14. Blades coy y pubescent, scabrous to merely felty; rare in SEUS, adventive in VA from
north and west S. arenicola
14. Blades pubescent, but never scabrous or felty; higher elevations, KY-TN-GA-AL-SC
S. nuttallii
. Petioles obvious, frequently 1/5 the length of the blade.
15. Leaf blades dentate or sharply dentate; stem angles abundantly pubescent with spreading or
somewhat retrorse, long (to 3 mm) hairs S. clingmanii
15. Leaf margins crenate to serrulate, but never dentate; stem angles glabrous or pubescent (if the
latter, then with scattered, stiffish, retrorse hairs).
16. Annual; corolla scarcely exserted from calyx; LA-MS-AL-FL- = S. agraria
16. EE corolia prominently exserted from calyx; widespre
Plants commonly glandular on stems, leaves, and within ieee
En Plant producing thick, segmented, tuber-like rhizomes; blades oblong; [weedy,
mostly of coastal plain S. floridana
18. Plant rhizomatous, but not producing thick, tuberous thickening of rhizomes; blades
elliptic, cordate; not weedy [higher elevations, mostly mountains, KY-TN-GA-AL-SC
S. nuttallii
17. Plants not glandular on stems, leaves, and within infloresce
Calyx usually abundantly pubescent with stiff, k hais: fruiting calyx lobes
straight, Termal T ate S. hispida
19. Calyx usually g | fruit lyx lobes freq ly curved
or curling/retrorse, not at all or only weakly apiculate . S. tenuifolia
ACKNOWLEDGMENTS
I am grateful to Theo Witsell and other personnel of the Arkansas Natural Heritage Commission for sharing
information from their files, and also personnel from the Oklahoma Natural Heritage Inventory, for similar
help. I thank the US National Forest Service (Ouachita and Ozark National Forest units) for providing col-
lection permits. Funding support for field work and for publication costs were provided through the W.T.
Batson Endowment for the A.C. Moore Herbarium, USC. Kathleen Ross (Department of Languages, Litera-
tures, and Cultures, USC) provided the Latin diagnosis. Johannes Stratmann (Department of Biological
Sciences, USC) and Hartmut Hilger (Systematische Botanik und Pflanzengeographie, Frei Universitat, Ber-
lin) helped with the abstract. Clint Cook (Department of Biological Sciences, USC) provided, as usual, his
constant help with the graphics and figures within this paper. Zack Murrell (BOON), Bert Pittman (South
Carolina Department of Natural Resources), and Alan Weakley (NCU) graciously provided reviews on the
manuscript.
I am especially grateful to George Johnson at the herbarium (APCR) of Arkansas Tech University, who
was so accommodating in helping me with the field aspect of this work, and in introducing me to a new
field pressing technique.
769
. £C. 4L 4 Li IE FPEM NI 4
Nelson, A [ Stact y fi | Highlands of the US
REFERENCES
ARKANSAS NATURAL HERITAGE COMMISSION. 2005. State species of special concern — Plants. List available from Depart-
ment of Arkansas Heritage, 1500 Tower Building, 323 Center Street, Little Rock, AR 72201. url: http://www.
naturalheritage.com/program/rare-species/. Accessed July 2008.
EPuwc, C.C. 1934. Preliminary revision of American Stachys. Feddes Repert. Spec. Nov. Regni Veg. 80:1-75.
FERNALD, M.L. 1943. Virginian botanizing under restrictions. Rhodora 45:357—516.
FERNALD, M.L. 1950. Gray's manual of botany, 8th Edition. American Book Company, New York.
GLEASON, H.A. 1952. Illustrated flora of the northeastern United States and adjacent Canada. New York Bot. Gard.
New York.
GLEASON, H.A. AND A. CRONQUIST. 1991. Manual of vascular plants of northeastern United States and Adjacent
Canada. Ed. 2. New York Bot. Gard., New York.
HARDIN, J.W. AND A.W. Cooper. 1967. Mountain disjuncts in the eastern Piedmont of North Carolina. J. Elisha Mitch-
ell Sci. Soc. 83:139-150.
ILcIM, A., C. MENDERES, AND M. Y. DADANDI, 2008. Stachys marashica (Lamiaceae), a new species from Turkey. Ann. Bot.
Fenn. 45:151-155.
Lite, E.L. Jk 1970. Endemic, disjunct and northern trees in the southern Appalachians. In: Holt, PC., ed. The
distributional history of the biota of the Southern Appalachians. Part 2: Flora. Virginia Polytechnic and State
University, Blacksburg. Pp. 249-290.
MULLIGAN G.A. AND D.B. Munro. 1989. Taxonomy of North American species of Stachys (Labiatae) found north of
Mexico. Naturaliste Canad. 116:35-51.
Netson, J.B. 1981. Stachys (Labiatae) in southeastern United States. Sida 9:104-123.
NELSON, J.B. AND J.E. Fairey. 1979. Misapplication of the name Stachys nuttallii (Lamiaceae) to a new southeastern
species. Brittonia 31:491—494.
OKLAHOMA NATURAL HERITAGE Inventory DATABASE, 2003. Working list of rare Oklahoma plants. List available from
Oklahoma Natural Heritage Inventory, 111 E. Chesapeake Street, University of Oklahoma, Norman, OK 73019.
url: http: //www.oknaturalheritage.ou.edu/, Accessed July 2008.
Rosison, H.W. AND R.T. Auen. 1995. Only in Arkansas: a study of the endemic plants and animals of the state. Uni-
versity of Arkansas Press, Fayetteville.
SMrrH, E.B. 1994. Keys to the flora of Arkansas. University of Arkansas Press, Fayetteville.
STEYERMARK, J.A. 1934. Some features of the flora of the Ozark region in Missouri. Rhodora 36:214-233.
STRANGE, R.M. AND B.M. Burr. 1997. Intraspecific phylogeny of North American highland fishes: a test of the Pleis-
tocene vicariance hypothesis. Evolution 51:885-897.
ZOLLNER, D., M.H. MacRoserts, B.R. MACROBERTS, AND D. Lapp. 2005. Endemic vascular plants of the Interior Highlands,
U.S.A. Sida 21:1781-1791.
" i Inftha Rataniral D h Institute of Texas 2(2)
BOOK REVIEWS
A. MICHAEL POWELL, James E WEEDIN, AND SHIRLEY A. PoweLL. 2008. Cacti of Texas: A Field Guide, with
Emphasis on the Trans Pecos Species. (ISBN-13: 978-0-89672-611-6, pbk.). Texas Tech University
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800-832-4042). $24.95, 383 pp., 314 color ede 124 maps, 9" x 6".
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ni species ee Te ous treatment o is a systemati umentation of the Trans Pecos species of Cactaceae and
biology, chemistry, and genetics. Cacti of Texas now provides an excellent reference for
North America.—Justin Allison, Research Associate,
+} th
a ae challenging subject
Botanical Research Institute 2 Texas, 500 East 4th Street, Fort Worth, Texas 76102-4025, U.S.A.
11
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that someone who knew nothing about cacti could quickly find the place nm tus is di d Beginning with Eu narrow
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the same time spinier and spinier. Finally, on pages 264-266, we come to some cacti that are so round and so spiny that i look like
nowmen—members of the genus Epilantha. Then the cacti grow longer, thinner and less spiny again, as well as becoming more like
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Trans-Pecos region they grow most n Itisa Nen dn guide for identifying wild cacti. Too bulky for a pocket, it will fit into most
purses and all backpacks. 1 Kalisi Waterman, Junior Intern, Botanical Research Institute of Texas, 500 East
4th Street, Fort Worth, Texas 76102-4025, U.S.A.
J. Bot. Res. Inst. Texas 2(2): 770. 2008
GUTIERREZIA ELEGANS SP. NOV. (ASTERACEAE: ASTEREAB),
A SHALE BARREN ENDEMIC OF SOUTHWESTERN COLORADO
Al Schneider Peggy Lyon
19049 Road V Colorado Natural E Program
Lewis, Colorado 81327, U.S.A. Colorado State University
Fort Collins, C E 80523, U.S.A.
G I Nesom
artwood Drive
Fort "n Texas 76109, USA,
ABSTRACT
1 f. I 1 1] NA Clg d
Gutierrezia elegans Al Schneider & P. Lyon, sp. nov.,
Colorado. The new species is known only from several populations in and e the developing Lone ie Park in Dolores
County, where it is represented by at least 4000 ptis Tue peus are low p bs with wood | ] e-
cumbent-ascending, evenly leafy stems, short-t gested corymboid clusters, and ne 3-nerved leaves.
RESUMEN
Se describe os elegan Al Schneider E P. nis on, sp. nov., de los afl i ill grisá de M Shal el su-
roeste de Colorado. I 11 Mesa S Parl lreded , enel condado
de Dolores, donde está representada An al menos iid plantas, Las eee son siete bajos, compactos con ramas caulinares
leñosas, y tallo jas decumbente-ascen endentes, I racim gestos, y hojas
cortas dius
Biological exploration of the developing Lone Mesa State Park in Dolores County, Colorado, has brought to
light a previously unknown species of Gutierrezia.
Vip elegans Al Schneider & P Lyon, sp. nov. (Figs. 1, 2). Tee UNITED STATES. CoLorapo. Dolores Co.: Lone
ate Park, S end of the park, ca. 23 mi N of the town of Dolores, T39N, R14 W, Section 35, ca. 108" 28'W, 37° 41'N, exposure
oth Ms us ca. i Supe. s ft, with DE Penante A Tetraneuris ui and vul ied ie ib
amphibolus and
nuttallii, Delphinium E hoedi i aiee dalini, Physaria pulvinata, Packera oödes, Solidago singlcs. and others, Pinus
ponderosa with pinyon-juniper on nearby surrounding slopes, 23 Aug 2008, A. Schneider G-1, with members of the Colorado Native
Plant Society (HoLoTyPE: COLO; isotypes: ASC, ASU, BRY, KHD, MO, NMC, NY, RM, TEX, US, UNM, UVSC)
Differt a Gutierrezia pomariensi (S.L. Welsh) S.L. Welsh caulibus brevioribus decumbentibus vel ascendentibus, capitulis minoribus in
inflorescentiis congestis dispositis, et foliis trinerviis.
Subshrubs from a woody taproot with short-branched woody caudex branches, plants growing vertically
7.5—12.5(-15) cm high. Stems decumbent-ascending, 8-15 cm long, moderately hirtellous. Leaves alternate,
sessile, basal absent at flowering, cauline evenly distributed, blades linear-lanceolate to linear-oblanceolate,
mostly 10-16 mm x 2-3 mm, primarily 1-nerved but with a distinct pair of narrower lateral nerves, slight-
ly or not all reduced in size distally, margins entire, surfaces and margins hirtellous. Heads borne mostly
in congested corymboid clusters of 2- a sometimes i peduncles 1-6 mm or rarely to 20 mm from
proximal branches of the capitul late, 3-4 mm long, 2.5-3 mm diam.
Phyllaries lanceolate-oblong, proximal 4/5 white-indurate, apices ranguar to ovate with acute to obtuse
tips, green, thickened, gland-dotted. Ray florets 6-8, fertile, corollas yellow, 3-5 mm long, laminae coiling
at maturity. Disc florets 6—9, yellow, fertile. Cypselae cylindric, 1-2 mm long, faces without oil cavities,
sparsely short-strigose; pappus of 1-2 series of persistent, oblong-lanceolate scales 1-1.5 mm long. Chromo-
some number unknown.
J. Bot. Res. inst. Texas 2(2): 771 — 774. 2008
772 Journal of the Botanical R h Institute of Texas 2(2)
Fic. 1.6 bh habi TET Er,
Flowering late July through early September. Bare Mancos shale outcrops and thin soil over shale,
Gutierrezia elegans scattered to abundant in the barrens and also occurring with Artemisia nova and other
species in sites with deeper soil over the shale, Pinus ponderosa and pinyon-juniper on surrounding slopes;
7575-7600 ft (ca. 2500-2550 m). Apparently endemic to southwestern Colorado.
Additional collections examined: Colorado. Dolores Co.: Lone Mesa State Park, type locality, 4 Aug 2008, Schneider and Lyon s.n. (TEX);
e Mesa State Park, S end of the park, ca. 23 mi N of the town of Dolores, m R14 W, Section 23, ca. 108” 28'W, 37° 42'N, sandy
sol over shale, 80% veg dominant A dA I leni ith Artemisia dica Eriogonum alatum,
Penst A lus mi iensis val aÈ H heca villosa, Pascopyrum smithii, and Chryo-
has depressus, slopes 8%, SSE- facing Pinus pan densa on surrounding slopes, 7600 ft, 23 Aug 2008, Schneider G-2, with members
of the Colorado Native Plant Society (COLO, TEX).
Common name.—Lone Mesa Snakeweed.
Etymology.—We have chosen the specific epithet *elegans" because it summarizes so many of the most
obvious visual characteristics of this new species. Gutierrezia elegans is delicate with masses of brilliant
yellow flowers topping gracefully arching stems that form into a low, domed symmetry. In short, the plant
is elegant.
Gutierrezia elegans is highly distinctive among all its congeners (Lane 1985), especially in its low, sub-
shrubby habit with decumbent-ascending, evenly leafy stems, short-pedunculate heads in congested corym-
boid clusters, and short, 3-nerved leaves. It presumably ud be most bal related to the group of species
with a subshrubby habit, decid basal leaves, and heads mostly p late and in relatively open arrays
(vs. the subshrubby G. sarothrae (Pursh) Britt. & Rusby and G. microcephala (DC.) A. Gray, which have heads
mostly sessile or subsessile, in glomerules, often in broad, dense, flat-topped arrays). Additional color pho-
Schneider et al., À
Fic. 2. Cluster of heads, Guti
tos of the new species and its habitat are posted on Southwest Colorado Wildflowers, Ferns, & Trees (Sch-
neider 2008). The following key, which identifies G. elegans in the context of similar species, is adapted from
the FNANM treatment of Gutierrezia (Nesom 2006)
1. Perennial herbs; basal leaves
M
ict, VUE
y corollas 5-8(-10) mmy disc florets 15-23; Utah rrezia
tradoria (Welsh & Ems E E Welsh
1. o basal leaves not persistent; ray corollas 2-7 mm; disc florets 5-15(- Je Arizona, Colorado, Utah,
Califo
Les |
» Pryilary
tri Colorado and Utah.
3. Stems 8- 15 cm long, decumbent ascending; jio masy in ongea conmbala cas of 2-8,
sometimes solitary, peduncles 1-6 mm or rarely to 2
leaves mostly 10-16 mm long, 3-nerved; involucres 3-4 mm long; Colorado
Gutierrezia elegans
Al Schneider & P. Lyon
3. Stems mostly 20-50 cm long, erect to ascending-erect; heads in open corymbs, peduncles ko
10-40 mm long; leaves mostly 20-40(-60) mm long, 1-nerved; involucres 5-8 mm long; Utah.
utierrezia pomariensis (S. E m S.L. Welsh
2. | hyllary apices not thich j; |
4 Ce I} i |
yp y sti Oe A Arizona and California.
late (as long as wide); cypsel 1.2 mm; Arizona, Gutierrezia serotina
Greene
4. St =o minutely hispidul Pee is "M E "
cypselae 2-2.8 mm; California
( than wide);
Gutierrezia californica (DC.) Torrey & A. Gray
Lone Mesa State Park is situated in the Dolores River drainage, immediately north of the San Juan River
drainage and thus was not included in the Four Corners Flora (Heil et al. 2008), although large areas of the
" : LV e£sha Dataniral D h Institute of Texas 2(2)
Mancos Shale also occur in the San Juan drainage. Gutierrezia sarothrae and G. microcephala are the only
species of the genus treated in the Four Corners flora (Nesom 2008), and they also are the only two of the
genus previously known from Colorado.
Gutierrezia elegans is known from five separate populations in and around Lone Mesa State Park, all
within a radius of about three miles. It occurs on San Juan National Forest and Bureau of Land Management
land adjacent to the state park. Among its known populations, we estimate that G. elegans is represented by
a total of over 4000 plants. Plants of the new species at the type locality occur as well-separated individuals
on very sparsely vegetated flats (slopes ca. 296) and sides of shallow washes. They grow along cracks in the
bare shale and in thin gravelly soil over the shale. Where the shale is not directly exposed, the new species
is easily seen to grow in the “crack-lines” of the underlying rock. Gutierrezia elegans is among the more
abundant species in this habitat. Pinus ponderosa occurs on nearby slopes.
At other localities, the soil is deeper and more sandy and Gutierrezia elegans occurs less commonly
among other species in a more densely vegetated community dominated by Artemisia nova. Gutierrezia elegans
is most abundant at the interface of the barren shale and surrounding sagebrush dominated areas, forming
a rim around the barren patches. Also abundant here are Artemisia ludoviciana, Heterotheca villosa, Achillea
lanulosa, Eriogonum racemosum, Astragalus haydenianus and various grasses, with Pinus ponderosa on sur-
rounding slopes. This site is at 7800 feet elevation and has slopes of 6-8%.
The rare and recently discovered Physaria pulvinata (O'Kane & Reve O grows with Gutierrezia
elegans on the shale ps within Lone Mesa State Park. The t ilar in their many-branched
caudex and caespitose, mound-forming habit. Physaria pulvinata i is known from similar shale barrens
in the Miramonte State Wildlife Area of San Miguel Co., about 23 miles north of the Lone Mesa State Park.
In addition to these two species, other endemics have been described from Mancos Shale habitats closely
clustered in the Four Corners area: Abronia bolackii Atwood, Welsh, & Heil, Proatriplex pleiantha (W. Weber)
Stutz & Chu, Sclerocactus mesae-verdae (Boissevain & Davidson) L.D. Benson, and Xanthisma paradoxum
(Turner & Hartman) Nesom & Turner (New Mexico Rare Plant Technical Council 1999; Nesom & Turner
ACKNOWLEDGMENTS
We are grateful to Scot Elder, Superintendent of Lone Mesa State Park, for his cooperation and support and
to John Strother and Ron Hartman for their editorial comments.
REFERENCES
Hei, K.D., S. O'Kane, AND L. Reeves (eds.). 2008 (in press). Flora of the Four Corners Region: Vascular Plants of the San
Juan River Drainage— Arizona, Colorado, New Mexico, Utah. Missouri Botanical Garden Press, St. Louis.
Lane, M.A. 1985. Taxonomy of Gutierrezia (Compositae: Astereae) in North America. Syst. Bot. 10:7-28.
Nesom, G.L. 2006. Gutierrezia (Asteraceae: Astereae). In: Flora of North America Editorial Committee, eds. Flora of
North America North of Mexico, Vol. 20. Oxford University Press, New York.
Nesom, G.L. 2008 (in press). Gutierrezia. In: Heil, K.D. S. O'Kane, and L. Reeves, eds. Flora of the Four Corners Region:
Vascular plants of the San Juan River Drainage — Arizona, Colorado, New Mexico, Utah. Missouri Botanical
Garden Press, St. Louis.
Nesom, G.L. AND B.L. Turner. 2007. Taxonomic review of the Xanthisma spinulosum complex (Asteraceae: Astereae).
Phytologia 89:371-389.
New Mexico Rare PLANT TECHNICAL COUNCIL. 1999. New Mexico Rare Plants. Albuquerque, NM: New Mexico Rare Plants
Home Page. http://nmrareplants.unm.edu (Latest update: 19 April 2007).
O'Kane, S.L. AND J.L. ReveaL. 2006. Physaria pulvinata (Brassicaceae), a new species from southwestern Colorado.
Brittonia 58:74-77.
SCHNEIDER, A. 2008. Southwest Colorado wildfl ,ferns, &trees. <www.swcoloradowildflowers.com> Accessed
September 2008.
JUNCUS DIGITATUS (JUNCACEAE),
A NEW ANNUAL RUSH FROM SHASTA COUNTY, CALIFORNIA, U.S.A.
Carol W. SAN Peter F. Zika
h Stre erbarium, Box 355325
EE California 958 > 15, U.S.A. vr of Washington
witham@ncal.ne Seattle, Washington 98195-5325, U.S.A.
Zikap@comcast.net
ABSTRACT
Juncus nis is an ad deseupedt bons two pu in the foothills of the Cascade Range in Shasta County, California. It is
separable ar fruits with an acuminate apex, longer capsules 11-17 mm, and fruits
J E
that are 3-4 times as oe as the tepals. A key to J ti i 1 for western North America.
RESUMEN
Juncus digitatus es una planta anual O de dos e is pue i la o de las Cascadas, en el panies de
cápsulas
Shasta, California. Es separable de J
más largas 11-17 mm; y tépalos 25-30% la lofgitud de los frutos. Se presenta una clave para Juncus sección Cabrio para la región
occidental de América del Norte.
Key Wonps: Juncus digitatus, Juncus section Caespitosi, California, Shasta County, vernal pools
Juncus section Caespitosi Cout. is defined as a group of 16 delicate annuals primarily from South Africa and
western North America (Kirschner et al. 2002a), alternatively included as annuals within subg. Graminifolii
(Engelm.) Buchenau (e.g., Brooks & Clemants 2000). Eleven members of section Caespitosi are indigenous
to California, including four endemics: Juncus triformis Engelm., J. leiospermus FJ. Herm. var. leiospermus, J.
leiospermus var. ahartii Ertter, and J. luciensis Ertter (Ertter 1986; Swab 1993). An additional endemic is
described here, first collected in 1991 by Dean W. Taylor.
— o C. Witham & Zika, sp. nov. (Figs. 1-3). Te: U.S.A. CaLtrornIa. Shasta Co.: vernal pool about 10 air km
E of Shingletown, 2 Jun 1993, Carol W. Witham 481 (notoryee: JEPS; isotypes: CAS, MO, OSC, RSA ).
A Junco triformi capsulis linearibus acuminatis 11-17 mm longis differt.
Plants annual, 4—10.5 cm tall. Leaves essentially basal, sheaths 2-15 mm long, with broad scarious frag-
ile margins tapering to the blade without auricles; blades filiform, 5-20 mm long, with acicular apex
0.2-0.5 mm long. Stems (culms) unbranched, 3-7 cm tall, 0.2-0.3 mm thick. Bracts often clustered like
an involucre, 2-8, ovate or narrowly ovate, acute, scarious or dark purple towards base, 1-1.8 mm long.
Pedicels 0.5-1.5 mm long. Flowers terminal, in clusters of (1-)2-8, trimerous. Tepals uniformly 6, nar-
rowly lanceolate, 3.5-4.4 mm long, 0.6-0.7 mm wide, the inner (petals) usually 0.4—0.7 mm longer than
the outer (sepals); midvein green with red border, 0.1-0.3 mm wide, often with scattered low papillae at
40x, the tips dark purple; with narrow pale margins 0.1-0.2 mm wide to the apex; tepal margins scarious;
apex acuminate to attenuate, erect to slightly recurved at tip. Stamens 3, 2-2.5 mm long; filaments 0.6-0.8
mm long; anthers often somewhat sagittate-based, 1.1—2.1 mm long, ca. 2 times the length of the filaments.
Style 1-1.5 mm long, elongating to 2.7 mm in fruit; stigmas (2)3, 1.6-5 mm long, reddish, exserted 1.5-3
mm beyond the tepals at anthesis. Capsules linear, often slightly curved, (2)3-valved, c. 3-4 times as long
as the tepals, (7-)11.3-17 mm long, (0.4-)0.7-1 mm wide, brown to red or dark red to dark brown, similar
in color to the tepals, the apex acuminate and gradually tapered into base of the persisting style. Seeds 9-20
per row and ca. 10-50 per capsule, ellipsoid to ovoid or narrowly ovoid, 0.5-0.65 mm long, 0.35-0.4 mm
wide, shiny, dark brown, at 10x with 10-12 strong longitudinal striations and faint cross-striations, base
blunt or with dark broad stipe 0.05 mm long, apex pale, short apiculate. Chromosome number unknown.
J. Bot. Res. Inst. Texas 2(2): 775 — 781. 2008
f Texas 2(2)
776
Fila 392021
1£27,T1 VII LIA 2227 1
4900.0 £,
Is. C. Anthers, fresh and old. D.
"
J
E. Mature capsule F. Seed, witt I
Ed
Fic. 1. Juncus digitatus. A. Habit. B. Fl
& Witham.)
Mel 4 7; A A fi fi Calif i 777
Etymology.—We have selected digitatus, a name that suggests the finger-like aspect of the unusual fruit
clusters.
Paratyees: U.S.A. CALIFORNIA. Shasta Co.: Shingle Creek along Highway 44, 29 May 1991; D.W. Taylor 11835 (JEPS, mixed sheet
with J. triformis); same site, 20 May 1992, C.W. Witham 443 (JEPS); same site, 27 May 1992; D.W. Taylor 12657 & C.W. Witham (JEPS);
same site, 2 Jun 1993; C.W. Witham 480 (CHSC, DAV, GH, JEPS, NY); same site, 3 Jun 1993, D.W. Taylor 13561 (UC); same site, 18 May
2007, C.W. Witham 1258 (DAV), same site, 4 Jun 2007, P. F. Zika 23015 & C.W. Witham (CAN, DAV, UC, US, WTU); Hamp Creek, 25 May
1993, D.W. Taylor 13498 (JEPS, UC); same site, 18 May 2007, C.W. Witham 1259 (DAV); same site, 4 Jun 2007, P. F. Zika 23021 & C.W.
Witham (DAV, GH, MO, NY, PRA, RSA, UC, WTU).
DISTRIBUTION AND ECOLOGY
Juncus digitatus is known only from two populations in Shasta County, from the foothills of the Cascade Range
at the northern end of the Sacramento Valley. Populations are in full sun, in the vernally damp ground of
seeps, vernal pools, and swales on gentle slopes over volcanic bedrock, at elevations of 660—790 m. Its com-
mon associates include Allium amplectens Torr., Centaurium venustum (A. Gray) B.L. Rob., Cyperus squarrosus
L., Deschampsia danthonioides (Trin.) Munro, Epilobium pallidum (Eastw.) Hoch & P.H. Raven, Eryngium ar-
ticulatum Hook., Isoetes nuttallii A.Braun ex Englem., Juncus bufonius L., J. acuminatus Michx., Mimulus gut-
tatus Fisch. ex DC., Muhlenbergia richardsonis (Trin.) Rydb., Navarretia intertexta (Benth.) Hook., Odontosto-
mum hartwegii Torr., Polygonum bolanderi Brewer, and Trichostema laxum A. Gray. Surrounding forest is
dominated by Pinus ponderosa Douglas ex Lawson & C.Lawson, P. sabiniana Dougl. ex D.Don, and Quercus
douglasii Hook. & Arn.
One population of Juncus digitatus grows within a few meters of plants of J. triformis, although the two
species vary in their flowering period and microsite preference. In the same vernal pool complex J. triformis
prefers slightly drier positions and is in fruit when J. digitatus is in flower in mid-May. Juncus digitatus fruits
in late May and early June.
RELATIONSHIPS
Juncus digitatus is most closely related to J. triformis, and the two have similar outcrossing pollination biol-
ogy, with anthers much larger than the filaments, spreading tepals, and exserted stigmas when in bloom.
Their habit, habitat, tepals, foliage and seeds are also similar; both species are in Juncus section Caespitosi
and endemic to California. Juncus triformis capsules are 1-3 mm long, including the beak. Juncus digitatus is
unique in the genus with its Lotus-like clusters of linear capsules 11-17 mm long. No other Juncus regu-
larly have capsules more than 10-11 mm long, among the 315 species covered in the comprehensive cata-
logues of Kirschner (2002a, 2002b), Balslev (1996), or Brooks and Clemants (2000). Juncus triformis tepals
generally exceed (rarely equal) the length of the capsule; in J. digitatus the mature capsule is about 3-4 times
as long as the tepals. Juncus digitatus capsules are linear, often curved, and gradually tapered to the persistent
style on the apex; in J. triformis the capsule is generally ovoid to ellipsoid, and the apex is usually abruptly
tapered to the persisting style base.
Our field work found that J. triformis is absent at one of the two populations of J. digitatus. This is sig-
nificant, demonstrating that the two species retain their morphological differences in the same habitat, or
when growing isolated from each other (Wagner & Wagner 1983). Intermediates are not known. This min-
gling in some sites without intermediates, and growing isolated elsewhere, parallels the situation among a
number of other western North American species of Juncus section Caespitosi reported in Ertter (1986) and
supports our argument that J. digitatus should be treated at the rank of species.
Ertter (1986) points out that Juncus triformis is unusually variable in several morphological features,
including “coloration, shape, and relative proportions of inner tepals, outer tepals, capsules, and seeds.”
Hermann (1948) was unaware of this extensive and continuous morphological gradation, and split from
Juncus triformis high elevation plants with relatively slender culms and large seeds, which he called J. me-
gaspermus F.J. Herm. After reviewing the variation in specimens of J. triformis from across California, using
the collections at CAS, CHCS, DS, GH, JEPS, OSC, POM, RSA, UC, UCR, and WTU, we agree with Ertter
778 J lof the Botanical R h institute of Texas 2(2)
A. WE n T)
Fic. 2. Photo of Juncus digitatus habit.
(1986) and Kirschner (20022) that the type of J. megaspermus is not a significant variant and belongs in the
synonymy of J. triformis. Nor have we found any other name in the synonymy of J. triformis that would apply
to and have priority over J. digitatus.
A key to annual Juncus with simple stems and terminal flowers in North America is provided to aid in
identification of the species. Mature capsules and ripe seeds are essential.
KEY TO JUNCUS SECTION CAESPITOSI IN NORTH AMERICA
. ts prominent, erect, appearing to be a continuation of the stem, much longer than
Sn eae outer tepals long-acuminate, much longer than acute to blunt inner tepals; Eurasian
introduction J. capitatus Weigel
Lower inflorescence bracts inconspicuous, not erect, much shorter than the inflorescence; outer tepals
similar in shape and length to inner tepals, sometimes inner tepals longer than outer tepals; native to west-
ern North America
2. Anthers 0.9-3.2 mm long, longer than filaments, styles 0.9-4 mm long (flowers outcrossing).
—
Witham and Zika, À new species of Juncus from California
Fic. 3. Photo of Juncus digitatus flower and immature capsules.
ve : Lafeha Bataniral D h Institute of Texas 2(2)
3. Capsules 11-17 mm long, about 3-4 times as long as tepals J. digitatus C. Witham & Zika
3. Capsules 1-5 mm long, shorter than or about as long as tepals.
4. Tepals acuminate; E a striated (at 15x); inner tepals colored to apex; inner tepal
margins narrowly sc J. triformis Engelm.
4. Tepais acute; ie e anginal striated; inner tepals not colored to apex, both inner and
outer tepal m | Jl J. leiospermus FJ. Herm
5. Flowers ius solitary a 2) pers stem a Jeiospenmus var. ahartii Ertter
5. Flowers usually 2-4 (rarely 7) per s var. leiospermus
2. ibas de 15- 0. 5 (0. 8) mm long, shorter "n ETE styles 0-0.5 mm long ETT self- pollinating).
6. lly striated (at 15x); flowers 1-2 (rarely 7) per stem; bracts 2, acute to acuminate.
7. pue usually 4.
8. Mature capsule usually subglobose, shorter and sometimes paler than tepal tips; seeds 0.5-0.8
m long; 2-3 seeds per locule, 4-9 seeds per capsule J. capillaris FJ. Herm.
8. Mature o elliptic to oblong, usually equaling or longer than similarly colored tepals; seeds
0.4-0.5 mm long; 4-7 seeds per locule, 10-30 seeds per capsule J. tiehmii Ertter
7. bs ace
9. Tepals usually longer and darker than the as 4-9 seeds per capsule; longitudinal ridges on
seeds often faint even at 30x; seeds 0.5-0.8 mm long J. capillaris FJ. Herm.
9. Tepals usually shorter and the same color as P capsule; 40-50 seeds per capsule; longitudinal
ridges on seeds weil-developed at 2e Sos 0.3-0. 2 mm long.
10. Stems usually with 2-3 (rarely 1 or Is ulti ly dark reddish;
0.4-0.6 mm ong) J. kelloggii Engelm.
10 C H ¡Wa | J | Ie | | n rA A
. JUICIO Usually Y I IOVVI ic
(tepal midveins Rd od distally); seeds 0.3- 04 mm lon ng J. luciensis Ertter
6. dA not longitudinally striated; flowers solitary; inflorescence bracts absent or 1-2, truncate to
acu
ue | + } t ct,
thickened below flower J. hemiendytus
FJ. Herm. var. abjectus (FJ. ERE Ertter
. Inflorescence bracts 1 = ED O not thickened below flower.
12. Inflorescence ry, apex truncate, sheathing stem; stamens 2-3 J. uncialis Greene
12. Inflorescence bracts 1—2, apex acute to blunt, not sheathing uin stamens 2-4.
13. Tepals usually 6, shiny, incurved and longer than capsule; inflorescence bracts 2 J. bryoides
FJ. Herm
13. Nus uen dull, erect or slightly outcurved and lly shorter than capsule; inflores-
ence bracts 1-2 J.t y . hemiendytus
CONSERVATION CONCERNS
Juncus digitatus joins a list of narrow endemics restricted to Shasta County, including Ageratina shastensis
(DM. Taylor & Stebbins) R.M. King & H. Robins., Ivesia longibracteata Ertter, Neviusia cliftonii Shevock, Ert-
ter & DW. Taylor, and Puccinellia howellii J.1. Davis. Like Juncus digitatus, several Shasta County endemics
are rare and have conservation issues. Due to the small populations and the restricted geographic extent of
Juncus digitatus, we have not provided exact locality data. The lead author has invested many field seasons
searching for this species, without adding more localities. Certainly more study and inventory is needed.
Nonetheless J. digitatus seems an appropriate candidate for attention from the conservation community, as
a narrow endemic on private lands in one county, in a wetland habitat that has been largely converted into
agriculture or rangeland. About half the available habitat at the Hamp Creek population has been lost in
recent years to a piped diversion of spring water.
ACKNOWLEDGMENTS
We would like to thank Barbara Ertter for prompting us to investigate this plant. We are grateful to Lowell
Ahart, Ed Alverson, Jan Kirschner, Frank Lang, Jim Nelson, Julie Nelson, and Dean W. Taylor for assistance,
to Ken Chambers for the Latin diagnosis, to Amanda Jorgenson for the Spanish abstract, to Linda Vorobik
for the artwork, and to the curators of the cited herbaria for loans or access to specimens. Steve Clemants,
Barbara Ertter and Jan Kirschner kindly reviewed and improved our manuscript. We thank the California
nel 17: ] A : Fl £i Calif i 781
Native Plant Society and its Sacramento Valley and Shasta Chapters, and the Lawrence R. Heckard Fund of
the Jepson Herbarium for grants which covered some of our expenses.
REFERENCES
BaLsLev, H. 1996. Juncaceae. Fl. Neotropica Monogr. 68:1-168.
Brooks, R. E. AND S.E. CLemanTs, 2000. Juncus. In: Flora of North America Editorial Committee, eds. Flora of North
America North of Mexico, Vol. 22. Oxford University Press, New York, NY. Pp. 211—255.
ERTTER B. 1986. The Juncus triformis complex. Mem. New York Bot. Gard. 39:1-89.
HERMANN, FJ. 1948. The Juncus triformis group in North America. Leafl. W. Bot. 5:109-120.
KIRSCHNER, J. (ed.). 2002a. Juncaceae 2: Juncus subg. Juncus, Species Plantarum: flora of the world Part 7. Australian
Biological Resources Study, Canberra, Australia.
KIRSCHNER, J. (ed). 2002b. Juncaceae 3: Juncus subg. Agathryon, Species Plantarum: flora of the world Part 8. Aus-
tralian Biological Resources Study, Canberra, Australia.
Swag, J.C. 1993. Juncaceae, Rush Family. In: J.C. Hickman, ed. The Jepson manual: higher plants of California.
University of California Press, Berkeley, CA. Pp. 1157-1165.
Waener, W.H. AND F.S. WAGNER. 1983. Genus communities as a systematic tool in the study of New World Botry-
chium (Ophioglossaceae). Taxon 32:51-63.
782 J lof the Botanical R h Institute of Texas 2(2)
BOOK REVIEW
SYLVAIN BEAUSEJOUR, with foreword & English translation by Paut Martin Brown. 2008 (expected). Orchidees
Indigenes du Quebec/Labrador (Native Orchids of Quebec/Labrador). d das Native, j
Boul. Manseau, Joliette, Quebec J6E 3E7 Canada. (Orders: } //orchid index.htm
info@nativeorchidsamerica.com, 450-755-2560; also ble from Orchid Books, B
com). $49.99 (CAD), $42.33 (USD), 176 pp., 380 illus., 9 3/4" x 11 3/4".
?
e p in I g to the wild orchids of Canada nde pod == been d duin and a
1 1 NT f, 11 A Ing 1 1
En i
Lahes. After many y Í pai l Syl I ] lwl i i bly tl I ifull l
th hids of north orth Arei Neither a aed guide nora Botanical treatise, ee niei de Quebec/Labrador (Native
Se of once + isa ak for the ey Į gray g in many years. Each species is treated
hnical inf i d howing th 1 distribution in tl i Habitat
T o o e
zi A 1 1 1 1 1 4 M Py 1 1 1
is imponen. Bun his dedication to finding all of un reds is bordering on the natia. I have bu privileged to o help him i in
locating some of the ele man of
t o
provided the Englis! Vh larger RE Tr was a pe ial xperien e that I had I t Bad Be sure
c "E
f£, 2 n F +1 1 1
y q OLLE DOOR
Derhid i» queso /T 4l 4 AT.
Orchid sof ipii de had a PRAE history of publication as the first
printing in Noveulber 2007 was so poorly bound that S books, and then after many months of
negotiations and broken promises, has finally bons ihe o And they p ] UE large itum heavy papers, and brilliant
colors all cene to this is volume. Althoug biased in favor of the book, 1 nie recommend it to all native os
enthusiasts Į ill sell out soon. Can we now look f i
by Sylvain on th hids of Ontario? Paul Martin Brown, 10896 SW 90th Terrace, Ocala, Florida 34481, U.S.A., er com.
J. Bot. Res. Inst. Texas 2(2): 782. 2008
TWO NEW SUBSPECIES OF SIDALCEA HICKMANII (MALVACEAE) IN CALIFORNIA
Steven R. Hill
iir hired PA S ~
6 S. Oak Stre
T Illinois 6 m U.S.A.
srhill@inhs.uiuc.edu
ABSTRACT
Revised treatments for Sidalcea for the oe Second ndi of the i Manual d the Flora M iid America have led to the
ern Cie As with the other four described subspecies, ihe two new taxa are found in very small isolated populations in upland
areas of open chaparral vegetation, the former from Napa County, and the latter from Lake County. They may represent relict popula-
tions. Unlike most other populations of the ee the two new taxa are not necessarily associated with serpentine. The two new
subspecies, and the others, may be vul grazing, foraging and human-caused changes in their environments.
Key Wonps: California, Sidalcea hick ii, Malvaceae, North America, rare plants
RESUMEN
1 1. €: 1.41 1 1 Asani dal lr A 7 la El f NT, Ih A E 1 n J 1A 1 E E -
e AME J
taxa adicionales. Se describen Sidalcea hicl ii subsp. napensi bsp. pillsburiensi lel de California. Tal como
1 1 : ] Lj 1 J 41 1 ^i = 1
pa i ¿E 1
A 1 qs 1 H del dadn de NT X 1 da nel dada de ] ake P 1 h]
© Y i por r F
relictas. Al trari ] ía del blaci de la especie, los d A i ig
serpentinas. Las B nuevas e. v las otras, pueden ser vul lr y bi ópi sus medios
INTRODUCTION
Sidalcea hickmanii Greene has long been known to exist in only a few widely scattered isolated populations,
generally on serpentine (Roush 1931; Hitchcock 1957), and the survival of these populations appears to be
dependent on occasional fires. Until recently, the species was thought to be a California endemic, but the
discovery of a population in southern Oregon, yet to be fully studied, has shown that this is not the case
(Halse, pers. comm.). This distribution can be interpreted as relictual for a species formerly more widely
distributed that has become separated into several isolated parts, each part subsequently diverging through
inbreeding. Work by Andreasen and Baldwin (2001) suggests that the species is not only basally divergent
within the genus but that it ‘evolved significantly more slowly than most of the other perennials’. Previ-
ously, four subspecies of S. hickmanii have been described; namely, the typical subspecies, subsp. hickmanii,
restricted to the Santa Lucia Mountains in Monterey County; subsp. anomala C.Hitchc., restricted to the
region of Cuesta Pass in San Luis Obispo County; subsp. parishii (Robinson) C.Hitchc., known from a few
sites in San Bernardino and Santa Barbara cos.; and subsp. viridis C.Hitchc., restricted to Big Carson Ridge
in Marin County. Reports of the species in Sonoma County have not yet been substantiated and several
coastal specimens examined from that county that had been reported as this plant were actually S. malvi-
flora. Specimens of unusual individuals of S. hickmanii at three sites have been brought to my attention by
their discoverers since the publication of the most recent treatment of the genus in the Jepson Manual (Hill
1993). An examination of the specimens and images of these plants led me to the conclusion that two new
Californian subspecies of this distinctive species needed description.
Sidalcea hickmanii Greene subsp. napensis S.R. Hill, subsp. nov. (Fig. 1) Tw U.S.A. Cairormia. Napa Co.: W of
Leoma Lakes at base of Mt. George, elevation 1380 ft; Mt. George quad., TON, R3W, NE/4 of NE/4 Sec. 29; chaparral, on brushy
J. Bot. Res. Inst. Texas 2(2): 783 — 791. 2008
784 Journal of the Botanical R h Institute of Texas 2(2)
EPSON HERSORIUM
diversity of California
E
+ IMAGED Minim C0.
N.N
y
1,61, ja, 18, 14.
ie oe er a a ae a
1% Re
pppn 1 NAME.: aeron hickmanii ssp. viridis
Pant Y: Malvace
COM NAME: SED TEhacketftson
ln
Ü SITE: west of Leo ma Lakes at base of Mt.George
y ELEVATION: 1380
UNTY QUAD: Mt. Geor
Ti £N. Ri Sit SEC: 29 NEGOfNES
COMMUNITY: Chaparral
HABE iy on brushy gradual slope, washed by
shee noff:openings ROCK: uri pra
Bree de biennial or perennial? -3 dm
tallscoroiia pale pink
COLLECTOR: Jake Puygt COL NUMBER: 2959
REC NO: 2434 DATE: May 1$, 1992
Fic. 1. Holot f Sidalcea hick iG bsp. napensis S.R. Hill, subsp. nov., Jake Ruygt 2959, 15 May 1992 (JEPS 110725).
¿Y
Hill, New subspecies of Sidalcea hickmanii from Californi a
gradual slope washed by sheet runoff, openings; rhyolite, biennial or perennial?, 2-3 dm tall; corolla pale pink, 15 May 1992, J.
Ruygt 2959 (HOLOTYPE: JEPS 1107251).
Herba p i di t , radice palari elongato. Caules pe erecti, + 20 ee cm, dis rubelli, en
noa : HERES A B 1: I Els I í. 10-25
mm longis, flexis in summis 2.5-3 mm; laminis latitudine ica quam tods, 10— Nc 20) mm ies x 15-25(34) mm latis,
3-5(-7)-lobatis palmatim prope basim, lobis dentatis profundi aia ovatis, 5-6 mm longis x 2-3 mm x 3—5-nervatis, ciliatis;
faka haci : 1 Inf] ; we -—
is 2-bracteatis, |
ribus, bracteolis 3 persistentibus, 5-6 mm longis x 0.7-1.0 mm latis, n 7-10 mm longis x 11-12 mm a a dense,
mes stellatis, lobis ciliatis; petalis roseis pallidis 9—11(-15) mm longis x 6-9 mm latis. Fructus schizocarpium, 6—7-mericarpiis 2 mm
2.0-2.5 mm longis x 1.5-1.8 mm latis, parietibus lateralibus laevibus, dorsis marginibusque 2—5-corrugatis-sinuatis, stria me-
diano in dorso; cuspide (mucrone) 0.
Perennial herb from a small erect caudex, generally with an elongated taproot or rootstock. Stems few, erect
(to slightly ascending), + 20—30(-40) cm tall, often tinted reddish (maroon), pubescent with scattered
tufted stellate hairs and a few simple hairs. Leaves few (« 6 per stem), pubescent with gen 6-rayed stellate
hairs; mid-stem leaves with petioles 10-36 mm long (to 6 cm on lower leaves), bent in the uppermost
2.5-3 mm (this portion pulvinus-like); blades wider than long, 10-17(-20) mm long x 15—25(-34) mm wide,
palmately 3—5(-7)-lobed nearly to the base, the cuneate lobes deeply few-toothed; stipules ovate, persistent,
5-6 mm long x 2-3 mm wide, 3-5-nerved, ciliate; a few leaves at the base occasionally deeply crenate.
Inflorescence a spiciform raceme, flowers scattered below, overlapping above, pedicels ca. 2 mm long, sub-
tended by 2 bracts similar to the stipules but more pubescent, 5.5-7 mm long; bractlets 3, persistent, 5-6
mm long x 0.7-1.0 mm wide; calyx 7-10 mm long x 11-12 mm wide, densely pubescent with stellate hairs,
lobes ciliate; petals pale pink, 9-11-15) mm long x 6-9 mm wide. Fruit a schizocarp with 6—7 mericarps
2 mm tall x 2.0-2.5 mm long x 1.5-1.8 mm thick, with smooth lateral walls, the back and margins with
2-5 wavy corrugations, single median line on back, cusp (mucro) 0.
The paratypes I have cited below include all other collections I have examined, most of which were
collected at the same site. A single robust specimen with 8 main flowering stems and a caudex 3 cm wide
was collected in 1942 at a second site west of St. Helena, but the current status of this population, as well
as other specific details, are unknown. On 7 May the lowermost flowers in the spikes had just opened.
Habitat information for the known extant population at the base of Mt. George suggests S. hickmanii subsp.
napensis is a plant of open mixed and chamise chaparral in rocky rhyolitic soils, at an elevation of 421
(417-427) m (1380 ft; 1370-1400 ft). The total number of plants is incompletely known, but the number
was approximated at 30 individuals (Ruygt, pers. comm.). When discovered on 29 April 1992, the first
flowers were opening, and flowering continued until at least 2 June 1992, at which time some fruits were
mature. Some of the plants discovered may have been browsed, and so the stems may exceed 3 dm in height
(Ruygt, pers. comm.). No subsequent observations have been made. This population is on private land, the
Wild Horse Valley Ranch, which may be developed as a vineyard (Ruygt, pers. comm.).
Etymology.—This new subspecies was brought to my attention in 1992 by Jake Ruygt who discovered
the plant while he was conducting botanical surveys as a subcontractor, and also as part of his ongoing
research on the flora of Napa County, and its epithet has been chosen to reflect its known distribution.
The plants were growing in a sparsely vegetated area, described as Chamise and Mixed Chaparral
(Ruygt, pers. comm.), with bare soil and abundant coarse gravel at a site where water puddles ephemerally
following rain. The immediate associates were Mimulus bolanderi A.Gray, Mimulus rattanii A.Gray, Navarretia
heterodoxa (Greene) Greene, and Phacelia suaveolens Greene. The typical local components in this plant com-
munity were Adenostoma fasciculatum Hook. & Arn., Arctostaphylos canescens Eastw., A. glandulosa Eastw., A.
stanfordiana C.Parry, Ceanothus purpureus Jepson, Pickeringia montana Nutt., Quercus wislizenii var. frutescens
Engelm., and Rhamnus californica Eschsch. The specimen matched best Sidalcea hichmanii subsp. viridis C.
Hitchc. and this was my first tentative identification in September 1992, but it obviously differed from that
786 J | of the Botanical R h institute of Texas 2(2)
in its deeply lobed leaves. Similar leaves are found on one other subspecies, S. hickmanii subsp. anomala,
known only from the vicinity of Cuesta Pass in San Luis Obispo Co., but the new subspecies differs from
that by its smaller bracts that are considerably shorter than the calyx, the fewer stems per plant, the essen-
tially unbranched inflorescence, and its rather different habitat and range.
The single, and first, specimen of the new subspecies was collected west of St. Helena in 1942 by Milo
S. Baker near the summit of Spring Mountain Road, apparently not far from the Sonoma County line, but
precise location details were not recorded on the label. This general area lies within an elevation of 427-609
m (1400-2000 ft). Jake Ruygt brought this specimen to my attention as well and it is most certainly the
same new subspecies.
PARATYPES. U.S.A. California. Napa Co.: divide between Santa Rosa and Napa Valley a f St. Helena town, near summit of
Spring Mt. Road, 7 id 1942, M.S. Baker 10193 e S. Baker ied 009782 at Ncc E sid of Mount oe ca. 4 mi E of nape,
elev. ca. 1380 ft; Ch I; lightl following rain
associated with Mimulus rattanii and Phacelia suaveolens, 2 Jin 1992, J Ruygt 3019 (VT!); W of Leoma Ge at = of Mt. George, eleva-
tion 1380 ft; Mt. PA o T6N, R3W, NE/4 of NE/4 Sec. 29; chaparral, on rocky flat, roadway; Rhyolite, biennial or perennial?,
1.5-2 dm tall; lla pink, 29 Apr 1992, J. Ruygt 2911 (Ruygt pers. coll.!).
Sidalcea hickmanii Greene subsp. pillsburiensis S.R. Hill, subsp. nov. (Figs. 2-4). Te: U.S.A. Caurornia. Lake
Co.: [vicinity of Lake Pillsbury] in tractor trail just under 2 mi S of Big Squaw Valley, T18N, R9W, SW/4 of SW/4 Sect. 8, in Mt.
Diablo Meridian and Base; GPS: NAD 27, 10S 0508308 E 4363785 N ra n 4375 N. Lat., -122.9375 W. ndis |, on slopes of
ephemeral d lj y ght r Arcto staphy-
los patula Ad J ! haris pilularis, Q berberi lif lia, Desch psi l , 12 Jul 2005, D.W. Isle 1124 with
Tara Athan (morore: JEPS 1130560.
Herba pus pue e pipe erectum puvum, Caules npe erecti, + a ida cm, viridulis, Pa: pilis stellatis
—20 mm longis;
fous latitudine plus quam iansiisdine lenfisbellatis. 8-15 mm Deis x 12-22 mm litis 7-nervatis palma inlobatis, crenatis
profundis a + 12-crenis; DRE owe +3 mm a x 1.3 mm latis, 1-nervatis, pilis similaribus atque in foliis.
Inflorescentia floril p prope basin 2- men vel ad i bracteis soli-
tariis. 1 . 1: 2i 11 : 1 E “1 1 + ut] 1 E 1 lin 2 ah]
ja] 1 4 c Ly O
mm latis, brevioribus bi 1 p 4—5 mm longis x x 6 mm latis, lis d ili llatis minuti il li
ES eo y LE [om
roseis pallidis 8-10 mm longis x 4 mm latis. F ] pium (4-)6—7-mericarpiis 1.5-2 mm altis x 2.0—2.3 mm longis x 1.0-1.3
mm latis, parietibus ] i gl il I 2—5- -corrugatis-sinuatis, ]i ind ide C ) 0.
Dwarf perennial herb from a small erect caudex. Stems several, erect, + (5—)10-30(-40) cm tall, greenish,
pubescent with appressed moderately dense stellate hairs. Leaves few (« 6 per stem), pubescent with tufted
stellate hairs; mid-stem leaves with petioles 6.5-20 mm long; blades wider than long, broadly fan-shaped,
8-15 mm long x 12-22 mm wide, palmately 7-veined, unlobed, rather deeply crenate, + 12-crenations;
stipules wide-lanceolate, + 3 mm long x 1.3 mm wide, 1-nerved, with hairs similar to those on the leaves.
Inflorescence with solitary axillary flowers or a short raceme, flowers scattered, those towards the base
2-bracted or with single bracts on those towards the apex, the single bracts oblong, cupped, much shorter
than the calyx; bractlets 3, oblong, persistent, 3 mm long x 0.8-1.0 mm wide, half as long as the calyx;
calyces 4-5 mm long x + 6 mm wide, densely puberulent, with all hairs minute, stellate; petals pale pink
(appearing nearly white in bright sunlight), 8-10 mm long x 4 mm wide. Fruit a schizocarp with (4—)6-7
mericarps 1.5-2 mm tall x 2.0-2.3 mm long x 1.0-1.3 mm thick, with smooth lateral walls, the back and
margins with 2-5 wavy corrugations, single median line on back, cusp (mucro) 0.
I have seen only the single collection, but the plant has also been documented rather well with digital
images. Habitat information and the images suggest that S. hickmanii subsp. pillsburiensis is a plant of open
chaparral on non-serpentine rocky soils of the Franciscan Formation, at an elevation of 701 m (2300 ft).
Only a single population has been documented and the total number of plants is unknown, though 24 were
large enough to be counted and protected from browsing animals within wire mesh enclosures. Additional
seedlings were found beneath Ceanothus shrubs in the Spring of 2006 (T. Athan, pers. comm.). When dis-
RUE LS
Memes National Forest - North Coast Ranges
pper Main Ecl River Watershed
| Sidaices Rickman E. Greene virulis Hatche.
| det. by Dr. Steven R. Hull
MEHBOGENO County.
Lug Tena, j bsc miles south of Bay Squaw Valles, Lake
county, California. TIEN, ROW, SW 18 of SW 1/4 Section 8
B. GPS: NAD 2* 108 9598308 E 4365745 N, on sloges oF
j enh ple S of Pinos attenuata, 2
“Zieh pank f Aisi Ceanothus foticsus, Arctostaphy! p
| Adenoscoma tesciculatum, cdo platea QUBE, DEEL
David W. isle 1124 07/12/2005
vits Tara Atao
' Studies in the Malvaceae
Sidolcea hickmuni
subsp. pillshuriensis 8. epi iet toy,
iv
Det: Ste
& ——.
Mendocino National Forest - North Coast Ranges
Upper Mzin Eel River Watershed
| Sidalcea apes E. Greene viridis Hitche.
| dct. by Dr. Steven R. Hall
MEENIBOCINO County, CA
Ta ‘Tractor Trail, just under 2 miles sou south of Big Squaw Valley, Lake
i County, C. W, SW 3/4 of SW 1-4 Section 8,
MDMB. GPS: NAD 27 10S 0508308 E 4363785 N, on siupes of
ephemeral ij, ©
Bacchaas pem QUBE, DFFL
David W. PH 1124 07 2/2005
with Tara
i *
d babes. Cn!
> *
x Studies in the Malvaceae
$ ë y Sidalcea hickmanii Greene
subsp. pillsburiensis S.R.Bill, soasp. nov.
. . pain
» e
* —
£ Hawes
i
epe Male Eel River bes
ic, F " die Hitch,
| det De Ser Steven P. im
MENDOCINO County, C.
icem Cats just E el soles south of Big Squaw Valley. Lake
or poc TIEN, ROW, SW iid of SW 1/4 Sectu 8,
NAD 27 105 0508308 E 4363 ird N rorat
m chaparral ad; DS of Pirus
sinus
i Laght pink fls. Assoc: Ceanothus fona, ados M
1 Adenostome Jasciculatum. Bacokans pilularis, QUBE, DEEL
Davii W. Isle 1124 07/12/2005
with Tera Athan j
Fic. 2. Holot ECHAS ra» Tm J
yl p. pillsburiensis S.R. Hill, subsp. nov., D.W. Isle 1124 with Tara Athan, 12 Jul 2005 (JEPS 113056).
Ingel Dag H ID
788 J h Institute of Texas 2(2)
covered on 7 July 2003, the plants were in flower. On 12 July 2005 plants were still in flower, and a few ripe
fruits were also found, and it continued to flower until at least 27 August 2005. The site was visited several
additional times in late 2005 and early 2006, but no information is available on more recent visits. This
single population is on Mendocino National Forest land (under federal ownership).
Etymology.—The subspecies epithet has been chosen to reflect its known distribution close to the
margin of, and upslope from, Lake Pillsbury.
This new subspecies was brought to my attention by National Forest botanist David Isle who, along
with fellow botanist David Toren, discovered the plant during surveys of a tractor fire line near Lake Pills-
bury in 2003 (“Tractor Line, Big Squaw Valley to 2001 Pillsbury Fire”). The plants were growing in a
sparsely vegetated area with bare soil and gravel on both slopes of an ephemeral drainage. In this habitat
the water flow would be limited to winter rain events, but no areas of pools were evident. The plant asso-
ciation (D. Isle, pers. comm.) best fits the description of uM oak—chamise chaparral, with dominants
that included Quercus berberidifolia Liebm. and Adenost iculatum, as well as the associates Arctostaphy-
los patula Greene, Arnica sp., Baccharis pilularis DC., Camissoña hirtella (Greene) Raven, Ceanothus foliosus
C.Parry, Cryptantha sp., Deschampsia elongata (Hook.) Benth., Epilobium minutum Lindl. ex Lehm., Galium
nuttallii A.Gray, Lotus denticulatus (E.Drew) Greene, Madia minima (A.Gray) Keck, Mimulus moschatus Doug.
ex Lindl., Nemacladus capillaris Greene, Pedicularis densiflora Benth. ex Hook., Penstemon heterophyllus Lindl.,
Polygala californica Nutt., and Potentilla cf. glandulosa Lindl. It adjoined a knobcone pine / chamise associa-
tion, dominated by Pinus attenuata Lemmon and Adenostoma fasciculatum. The images and sample sent to
me in 2005 matched best Sidalcea hickmanii subsp. viridis C.Hitchc. and this was my first tentative identifi-
cation, but it appeared to differ from that in its dwarfed habit, open inflorescence, and shorter bracts and
bracteoles. The crenate, truncate leaves can be found on that subspecies and on the typical subspecies as
well, but these were remarkably small on this new subspecies in comparison. Another difference noted was
in the pubescence of the upper stem; in this new subspecies the hairs are stellate but appressed, whereas on
subsp. viridis the hairs are distinctly tufted. The flowers appeared closer to bright white than pink when
fresh (D.Isle, pers. comm.).
Sidalcea hickmanii subsp. pillsburiensis has been collected twice. The first collection was made by David
Toren and David Isle on 7 July 2003, D. Toren 9318. This was mailed to me on November 1, 2005, but,
sadly, I have no record that this specimen ever arrived and there was no duplicate. The second collection
was made by David Isle, with Tara Athan, on 12 July 2005, and this specimen was sent separately and ar-
rived safely. This specimen consisted of 2 stem fragments and several mature fruits, from 3 separate plants
at the single site, and these have been deposited in the Jepson Herbarium (JEPS) to serve as the holotype.
Photographs of the plants were made on August 10 and 27, 2005, by David Isle (Fig. 4) and members of the
Sanhedrin Chapter of the California Native Plant Society, whose volunteer bers also enclosed 24 of the
previously located plants within wire-mesh cages to protect them from animal (deer) foraging—a problem
that had already been reported at the site. All of the living stems were quite short and very possibly animal-
browsed, but the associated dried stems from the previous season were 30—40 cm tall and probably not
browsed (D. Isle, pers. comm.). The chapter volunteers have shown an interest in monitoring these plants
and to look for any additional populations. The population appears to be vulnerable to heavy deer foraging,
and individuals within the tractor fire line may be damaged by its maintenance.
With the finding of these two new subspecies, there are now six named subspecies of Sidalcea hickmanii
in California, all of whose populations are small, isolated, and vulnerable. In order to better facilitate their
identification, the following key to these subspecies has been provided below.
It should be noted here that flower size in Sidalcea often varies by its sexuality—and the genus has
often been characterized by the presence of bisexual flowers as well as pistillate flowers in ombina
tions in the species and on individuals. It is easily observed that pistillate flowers (with aborted anthers) are
Hill, New subspecies of Sidalcea hickmanii from Californi 789
e $4
»
^
s
a *
*
k H5»
Fic. 3. n a of the plant material from the holotype sheet shown in Fig. 2 of Sidalcea hickmanii G bsp. pillsburiensis S.R. Hill
(JEPS113056).
Fic. 4. Sidalcea hick ii | pillsburiensis S.R. Hill subsp nov. PI I ht y D.W. Isle, 10 Aug 2005.
3
790 I falo Detani-nl D h Institut f Texas 2(2)
generally much smaller than bisexual flowers within a species, having petals even half the length of those
in bisexual flowers. The two newly described subspecies of S. hickmanii are represented by very few collec-
tions, and so the sample size is small. In all of these collections the flowers are bisexual and, therefore, no
size variation based on sexuality has been seen. However, I have been unable to find any such size variation
in the flowers of any of the subspecies of S. hickmanii to date (nor am I aware of any mentioned in the lit-
erature for this species). I am not yet willing to generalize that all S. hickmanii flowers are bisexual and show
no variation in size correlated with their sexuality, but I have, as yet, not observed such variation in the
species.
The following key uses the acronyms for bioregional distribution as found in the Jepson Manual (Hick-
man 1993)
KEY TO CALIFORNIA SUBSPECIES OF SIDALCEA HICKMANII
1. Most leaves deeply lobed + tc base.
2. Bracts > (8210-12 mm, += calyx; SCoRO: c San Luis Obispo Co. subsp. anomala
2. Bracts 5.5-7 mm, < calyx; NCoRI: Napa Co. subsp. napensis, subsp. nov.
1. Leaves not lobed, or deeply crenate to shallowly lobed much less than 1/4 length.
3. Bracts lanceolate, generally 6-10 mm x 2.5-4 mm wide, + « calyx; bractlets + = or « calyx; leaves above
uniobed or very shallowly lobed < 1/4 length; SCoRO, w WTR: Santa Barbara Co; SnBr subsp. parishii
3. Bracts linear, oo or oblong, 2-7 mm x 0.5-2 mm wide, << calyx; bractlets « calyx; leaves
ba always un
. Plant oy 4-8 dm, grayish-canescent; calyx densely stellate-puberulent, n hairs much
longer; bractlets 2-7 mm iong; largest leaves deeply cordate, closely crenate, often » 3 wide;
inflorescence + dense; ds Ms hickmanii
4. Plantgenerally « 4 dm, greenish, less hairy; calyx stellate-puberulent but generally not densely so, hairs
uniform in iength; bractlets 2-5 mm long; larger Ec truncate to wide-cuneate, coarsely crenate,
ped « 25 mm wide, wider in robust plant lly more open
. Bracts ofupper flowers dee flat or cupped; leafblades. 1-25 cm x2.5-4cm Aide stem above
with tufted hairs generally 0.5-1.2 mm long; stems generally > 30 cm long, flowers > 10, inflorescence
t spicate in age; calyx 4-7 mm long; n CCo subsp. viridis
Bracts of upper flowers 1, cupped; leaf blades 0.6-1.2 cm long x 0.7-2.1 cm wide; stem above with
appressed hairs generally 0.2-0.5 mm; stems generally « 30 cm long, flowers < 10, inflorescence not
spicate; calyx 4-5.5 mm long, NCoRH: n Lake Co. subsp. pillsburiensis, subsp. nov.
Ui
ACKNOWLEDGMENTS
I thank David W. Isle, Jake Ruygt, David Toren, and Tara Athan for not only bringing these plants to my
attention, but for their astute observations, plant and site images, and stimulating correspondence concern-
ing these taxa in recent years. I also thank Linda M. Prince, of Rancho Santa Ana, for generously sharing
images of the other subspecies as well as data from her molecular studies within this group. 1 would espe-
cially like to thank Bruce G. Baldwin and the Lawrence R. Heckard Endowment Fund of the Jepson Her-
barium for the funding to visit the UC / JEPS and CAS herbaria to study herbarium specimens in support
of my work on several Malvaceae genera for the upcoming revision of the Jepson Manual: Higher Plants of
California. The UC / JEPS and CAS staff were especially hospitable during my visit and very generous with
their time. Special thanks go to Rich Whitkus of Sonoma State University for the loan of the important
Sidalcea specimens collected by Baker from the herbarium (NCC), and to Richard Halse for his generous
sharing of expertise and specimens of the genus. A continuing contract with the Illinois Department of
Transportation is gratefully acknowledged for financial support towards research and publication of these
studies.
Hill, New subspecies of Sidalcea hickmanii from Californi "
REFERENCES
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checker mallows (Sidalcea, Malvaceae): evidence from 185-265 rDNA internal and external transcribed spac-
ers. Molec. Biol. Evol. 18(6):936-944.
Hickman, J.C. (ed.). 1993. The Jepson manual: higher plants of California. University of California Press, Berkeley.
Hu, S.R. 1993. Sidalcea. In: J.C. Hickman, ed. The Jepson manual: higher plants of California. University of Cali-
fornia Press, Berkeley. Pp. 755-760.
HitcHcock, C.L. 1957. A study of the perennial species of Sidalcea. Part |. Taxonomy. Univ. Wash. Publ. Biol.
18:1-79.
RousH, E.M.F. 1931. A monograph of the genus Sidalcea. Ann. Missouri Bot. Gard. 18:117-244.
: 15£sha Botanical D h Institute of Texas 2(2)
792
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and Adjacent Countries (within the borders of the former USSR). (ISBN 978-3-906166-61-2,
hbk.). Koeltz Scientific Books, Koenigstein, Germany. (Orders: www.koeltz.com or www.orchidbooks.
com). $212.00, 690 pp., color distribution maps, photographs, and watercolor plates.
E
a) TU fD. : e a T 1 1 [E t TS J 11 f +1
E ) p
Wide*tn he fonnd in A n ree ee pl 4 n m Wan ret USSR. The authors
studied orchids for more than 3 30 tun in different climati pe ola i n of the Cola Peninsula, temperate zones in Central
Russia, Central Ukraine, Balti ast territories, Medi Crimea and Caucasus, continental climate in Siberia
and Central Asia, and ic climate in the Far East of Russia. The first several chapt ist of detailed nds Bs the various
d ic Review, S cology and
authors and are entitled History and Contemporary T
Phytocoenology, Morphology, Ontogenesis, Rhythm of Seasonal Development and Reproduction, Mycoriüss. and hun Protection.
The bulk of A ped is eevee to the na Mud and they are very complete and easy to understand. Throughout the book
g O pa n in i bou dt A brief glossary contains not only the technical
] l Anglicized Russi hat help i Inow know that a Zapovendik is a highly protected area
under g l that prohibi i iviti ‘nature pese
A ] has this i i years and, judging data th 2004. Several genera and
ies also found i rth America are included, m. d found especially in Als: It is helpful to have more
ROPA on ie species. xd p Ld hubs Opripedium guttatum, C. cali Ced aristata, bici M ee
Liparis loeselii, Listera cordata, 1 g p. oligantha), Mal phyll th if P.
dilatata, P. chorisiana, and P. tipuloides. S lof native North A I his regi ] includ
pipacti helleborine, E. palustris, Listera ovata, and Z t tica. One of the most interesting treatments is in the
fth Lysiell i d ab d Tulotis. The latter ld include tł t North American
Many scientific ies rigi e written in a languaee other than English often md gam in EM dh but no so with this
book. The Englis! ] g p 1. Over 200 color Į p d by numerous
excellent l d a by 18 line d ing d 93 di i ps. Itis} I he book has i y significant
ue No RT mi is a i dd the es o or showing th ] cae gi ften mentioned in de book. For
r TICCD 341 L: 1 v TRAC g. If only one page
were ‘alae toa one outline map showing - S dan of D countries it w ds UM the book mud more able Fortunately
m. The names ne a idis of Dod countries
J
are slightly diffi f I d in the book, but easily dons. Then map can be printed
gary P
" A i 1 a hid fehl
ES 1/5 fp
reference. I hig
region.— Paul Martin Brown, 10896 S1 W 90th Terrace, Ocala, Florida 34481, U.S.A., naorchid@aol.com.
J. Bot. Res. Inst. Texas 2(2): 792. 2008
IPOMOEA TEHUANTEPECENSIS (CONVOLVULACEAE):
A NEW SPECIES FROM THE ISTHMUS OF TEHUANTEPEC, MEXICO
Leticia Torres-Colín, Rafael Torres-Colín,
and M P. Ramírez de Anda J. Andrew McDonald
tituto de Biología The University of Texas Pan American
Uem ii Autónoma de México Department of Biology
Departamento de Botánica 1201 West University Drive
Apartado Postal 70-367, Del. Coyoacán Edinburg, Texas 78541-2999, U.S.A.
04510 México, MÉXICO, D.F
ABSTRACT
A new and narrowly endemic species from the Tehuantepec District of Oaxaca, fpomaes tenane recenzis L. oi: pon R. Torres
Colín, M.P. Ramírez & J.A. Mi A is AME m e DER The new sp g p g y-seede d
lianas (1. subg. Eriospermum Verdc.) i I if 11 iti 1 reproducti Ur
moea ser. Mirandinae D.F. Austin). I } 1 is sl phological trai ith I l Matuda) D. Austin and
I. conzattii Greenm. i id regi f th Mexico but differs by gl } lami ith oł l 1 leaf! pleiocasial
cymes, glal licels less tl ] lak Is from 4-4.5 mm long and 23 muc m iin corollas ome to 3.0 cm
o
truncate
long. In contrast, T conzattii and I. concolora ei aa laminas with EA or truncate
i llas fi 3.0-4.5 cm nae
pubescent pedicels from 7-15 mm long, sericeous sepals from 5-8 mm lon
RESUMEN
Se describe eil I 1 is L. Torres-Colín, R. Torres- Colín, M.P. Ramírez & J.A. McDonald,
endémica del Distrito de Ao Oaxaca. d La nueva aon se ubica en un ENS de lianas con RON comosas (1. sube
Eriospermum Verdc.), | I j p y I mirandinae
D.F. Austin). I } laci 1 fold lora (M la) D Fans el ii
de las alee se distingue Sor sus hojas glabras con bases TUM a sibus inflorescencias en pleiocasios, pedicelos glabros de
menos de 6 mm de largo, sépalos glabros, 4-4.5 mm largo y 2-3 mm ancho y corollas glabras hasta 3 cm e ee conzattii y I.
concolora presentan hojas pubescentes con bases cordadas o truncado-obtusas, monocasios, pedicelos t
mm largo, sépalos sericeos, 5-8 mm largo, y corolas pilosas o sericeas de 3.0—4.5 cm larg
Recent floristic investigations in the Tehuantepec District of Oaxaca, Mexico, have uncovered ten species of
Ipomoea (Torres-Colín 1989; Torres et al. 1997), one of which is new to science. The new species belongs to
Ipomoea subg. Eriospermum Verdc. on the basis of lignescent stems and comose seeds and is accommodated
comfortably within Ipomoea ser. Mirandinae D.F. Austin s. str. (Austin 1977, 1979; = I. *microsticta complex
sensu McPherson 1979 or I. ser. Eriospermum s. lat., Austin & Huáman 1996) on account of its deciduous
leaves and relatively small (4.0—4.5 mm long), subequal sepals. These and a combination of ornithophilous
floral features, including red, hypocrateriform corollas, and exserted reproductive structures suggest close
relations with Ipomoea concolora (Matuda) D.F. Austin and I. conzattii Greenm., distinctions between which
are presented herewith.
Ipomoea Pu ar L. Torres-Colín, R. Torres-Colín, M.P. Ramírez & J.A. McDonald, SP. nov. (Figs.
1A—G). Te: ME
XICO. Oaxaca: Distrito as a no e to Arroyo de Las Minas, to the W of Rancho Limón,
24 km to P W Tehuantepec, 6?18'N, 95°27'W), 600 m, 23 Jan is Rafael Torres
C. 11255 & Cipriano Martínez R. TEE MEXU; tsorvees: ENCB, MEXU, MO, OAX, PAUH).
Species haec ab I. conzattii Greenm. et I. concolora (Matuda) D. Austin differt | labris basibus lami btusis vel subcordatis
1 lahri
pleiocymis pedunculis glabris pedicellis glabris maxima 6 mm longis sepalis labo 4.0-4.5 mm longis 2-3 mm |
maxima 3 cm longis
Twining lianas; Stems cylindrical, to 5 mm in diameter, bark yellow-brown, striate, glabrous, rarely squa-
mose, lenticels abundant. Leaves deciduous before anthesis; blades ovate to broadly ovate, (3.8-)5.5-11.0
J. Bot. Res. Inst. Texas 2(2): 793 — 797. 2008
794 I af tha Botanical D h Institute of Texas 2(2)
Jl ft J f. 411 190597) B
Fig. 1. | f. te} ic AV 2 £ : +h O Se | g 7 .C p
las : ‘th £l {frit £ And : ge J lla D gynoecium E Mat la f4 J Dafnal T. 11255) F Ventral surface
Tank 4 £ Adah: J Ia fF I
F
of mature seed. G. Persi 1892).
cm long, (2.5—3.5—8.5 cm wide, upper surface dark green, lower surface light green, chartaceous, glabrous
on both surfaces, apex acute, base obtuse or rarely subcordate, margin entire; petiole basally canaliculate,
distally terete, striate, (2.0-)3.0-8.7 cm long, 1-2 mm wide, glabrous. Inflorescence a pleiocasium 4.0—6.0
cm long, flowers 15-45, primary peduncle 1-3 mm long, glabrous, secondary peduncles 4-6 mm long,
bracteoles deciduous; pedicels 4-6 mm long, erect, glabrous, sparingly dilated at the apex, striate; sepals
Torres-Colín et al., A ies of | from Mexi 795
imbricate, reddish, ovate to broadly ovate, 4.0—4.5 mm long, 2-3 mm wide, chartaceous, glabrous, the
margins entire, apex obtuse, rarely acute, mucronate; corolla hypocrateriform, red, 2.5—3.0 cm long, tube
2.5-2.8 cm long, 5-6 mm in diameter, limb irregularly 5-lobate; segments 3—6 mm long; stamens 5, sub-
equal, exserted, filaments basally pubescent, anthers yellow; style exserted, glabrous, ca. 3 cm long, stigma
capitate. Fruit a capsule, ellipsoid, 9-13 mm long, 6-7 mm wide, 4-valvate, brown, chartaceous, dehiscent,
glabrous; seeds 4, brown, ellipsoid, three angled, 7 mm long, ca. 4.6 mm wide, the lateral margins comose,
hairs cream, ca. 7 mm long.
Habitat.—Ipomoea tehuantepecensis thrives in short tropical deciduous forest on dry gradients between
200—750 meters. The vines prefer steep slopes with thin soil mantles and often grow in association with
Bursera schlechtendalii Engl., Gossypium aridum (Rose & Standl.) Skovst., and Juliania adstringens Schltdl.
Distribution. —As presently known, I. tehuantepecensis is a narrow endemic of Oaxaca, Mexico, known
only from the District of Tehuantepec in the Municipalities of Santiago Laollaga and Santo Domingo Tehu-
antepec (Fig. 2).
Phenology.—Ipomoea tehuantepecensis flowers and fruits following the rainy season, from November to
March.
Etymology.—The species epithet refers to the Isthmus of Tehuantepec, in which the new taxon is ap-
parently restricted.
Specimens examined: MÉXICO. Oaxaca: Distrito de Tehuantepec, Municipio Santiago Laollaga, recorrido a las faldas del Cerro Indio
Dormido, al S de Laollaga, Álvaro Campos V. 4116 (MEXU, PAUH). Municipio Santo Domingo Tehuantepec, Cerro Guiengola, 11 km al NO
de Tehuantepec, Álvaro Campos V. 3831 y Cipriano Martínez R.(MEXU, PAUH); Álvaro Campos V. 4242 (MEXU, PAUH); Cerro Guiengola,
11 km al NE de Tehuantepec, Abisaí García M. 2802, L. Torres C. y L. Cortés A. (MEXU); subida a las ruinas del Cerro inde Leticia
Torres C. 243, R. Torres C. y C. Martínez R. (MEXU); Cerro Guiengola, ladera S entrando por Paso Alicia, Leticia Torres C. 306, R. Torres
C. y C. Martínez R. (MEXU ns ruinas del Cerro Guiengola, al N de nro Leticia Torres C. 336, R. Torres C., Pedro gai L.
y d ds Martínez R. (MEXU, PAUH); ladera S del Cerro Guiengola, de cal, Leticia Torres C. 774, R. Torres C., L. Cortés A.
Cipriano Martínez R. (MEXU, m Cerro Pene at the ruins, M Stafford 8 y RJ. NM (MEXU); Cerro Guiengola, al N de
d. Rafael Torres C.4892 y PAU
DISCUSSION
Ir tehuantey is and its cl latives, I. concolora and I.conzattii, belong to an assemblage of deciduous
lianas that extends from southern Mexico t thern South America, with a secondary center of diversity in
the Caribbean (Austin 1977). Ipomoea tehuantepecencis is distinguished by glabrous laminas with an obtuse
or subcordate leaf base, while the blades of I. conzattii are strigose and elliptical with obtuse bases. Laminas
of I. concolora are tomentose on the undersurface and basally cordate. Whereas I. tehuantepecensis produces
compound cymes with secondary peduncles from 4—6 mm long, I. conzattii and I. concolora exhibit simple
cymes with secondary peduncles that range from 2-4(-5) mm long. Glabrous pedicels of I. tehuantepecencis
are relatively short, ranging from 4—6 mm long, contrasting with those of I. conzattii and 1. concolora, which
are pubescent and 10-15 mm or 7-12 mm long (respectively).
Like many other Ipomoea species, I. tehuantepecencis is readily diagnosed by distinctive sepals, these
being reddish, glabrous, and relatively small (4-4.5 mm long, 2.3 mm wide). In contrast, sepals of I. conzattii
are greenish, sericeous, 5-7 mm long, and 4-6 mm wide, and those of I. concolora are reddish, sericeous,
7-8 mm long, and 3.5-4 mm wide. The brightly colored, glabrous corollas of I. tehuantepecensis are 2.5-3.0
cm long, the tube less than 2.8 mm in diameter at base and 5.0—6.0 mm from the middle to the throat,
while corollas of I. conzattii and I. concolora are pilose and sericeous (respectively), measuring from 3.0—4.5
cm long. These and other less discrete distinctions are summarized in Table 1
ACKNOWLEDGMENTS
The authors thank Cipirano Martinez Ramirez and Alvaro Campos Villanueva for collecting ample plant
materials during botanical explorations of the Isthumus of Tehuantepec, Oaxaca.
ID L Eze
796 f Texas 2(2)
O |. tehuantepecensis
a Bi / concolora
; @ |. conzattii
~~ 25°N
Mexico. Note that /
L
Tag. 1. Comp phology of IE lora, |. conzattii, and |
Ipomoea conzattii Ipomoea concolora Ipomoea tehuantepecensis
Blades Elliptic, base truncate to obtuse, Broadly ovate, base cordate, Ovate to broadly ovate, base
Inflorescence
Pedicels
Sepals
Corolla
both surfaces strigose
Monochasium, secondary
peduncles 3-5 mm long,
tomentose
10-15 mm long, tomentose
5.0-7.0 mm long, 4-6 mm wide,
greenish, sericeous, apex acute
to roundish, not mucronate
3.0-4.5 cm long, scarcely pilose,
tube 2.5-3.5 cm lon
0.5-0.7 cm wide, eimi Segments
1.0-1.5 cm lon
upper surface glabrous, lower
surface tomentose
Monochasium, secondary
peduncles 2-4 mm long,
sericeous
7-12 mm long, sericeous
7.0-8.0 mm long, 3.5-4.0 mm
wide,reddish, sericeous, apex
acute, not mucronate
3.3-4.0 cm long, sericeous, tube
3.0-3.6 cm long, 0.5-0.6 cm
wide, limb segments
0.3-0.4 cm long
obtuse to subcordate, glabrous
on both surfaces
Pleiocasium, secondary
peduncles 4-6 mm long,
glabrous
4-6 mm long, glabrous
4.0-4.5 mm long, 2-3 mm
wide, reddish, glabrous, apex
obtuse, mucronate
2.5-3.0 cm long, glabrous, tube
2.5-2.8 cm long, 0.5-0.6 cm
wide, limb segments 0.3-0.6
cm lon
Torres-Colín et al., A pecies of Ij from Mexi 797
REFERENCES
Austin, D.F. 1977. Realignment of the species placed in Exogonium (Convolvulaceae). Ann. Missouri Bot. Gard.
64:330-339.
Austin, D.F. 1979. An infrageneric classification for ipomoea (Convolvlulaceae). Taxon 28:359-361.
Austin, D.F. AND Z. Huaman. 1996. A synopsis of Ipomoea (Convolvulaceae) in the Americas. Taxon 45:3-38.
McPherson, G.D. 1979. Studies in the genus lpomoea (Convolvulaceae). Ph.D. Dissertation, University of Michi-
Torres, C.M.L. 1989. Estudio florístico y descripción de la vegetación del Cerro Guiengola en el Istmo de Tehuan-
tepec, Oaxaca. Tesis de Licenciatura, Escuela Nacional de Estudios Profesionales Iztacala, Universidad Nacional
Autónoma de México. México D.F.
Torres, C.R., L. Torres, P. DÁVILA, AND J.L. VILLASEÑOR. 1997. Listados florísticos de México. XVI. Flora del Distrito de
Tehuantepec, Oaxaca. Instituto de Biología. Universidad Nacional Autónoma de México.
B i laf tha Botanical D h Institute of Texas 2(2)
BOOK REVIEW
WILHELM BARTHLOTT, STEFAN POREMBSKI, RUDIGER SEINE, AND INGE THEISEN. 2007. The Curious World of Carnivo-
rous Plants, a Comprehensive Guide to their Biology and Cultivation. (ISBN 978-0881927924,
hbk.). Timber Press Inc., 133 S.W. Second Avenue, Suite 450, Portland, Oregon 97204-3527, U.S.A.
(Orders: www.timberpress.com, mail@timberpress.com, 800-327-5680, 503-227-3070 fax). $39.95,
224 pp., 158 color and black/white illustrations, 2 maps, 7 1/2" x 10 1/4”.
Contents:
1. Preface 16. Dioncophyllaceae: Hookleaf Family
2. The Flowers of Evil: An Introduction 17. Droseraceae: Sundew Family
3. Hal 18. Drosophyll Portug Sundew Family
4. iiS and Diversity 19. Lentibulariaceae: Bladderwort Family
5. Attracting and Trapping Mechanisms 20. Nepenthaceae: Asian Pitcher Plant Family
6. Digestion of Prey and Nutrient Uptake 21. Roridulaceae: Bug Plant Family
7. Prey 22. Sarraceniaceae: American Pitcher Plant Family
8. Guests at the Carnivores’ Table: The Commensals 23. Animal-Trapping Liverworts
9. Noncarnivorous Animal-Trapping Plants 24. Animal Trapping Fungi
10. The Evolution of Carnivory 25. The Carnivorous Plants of the World: A Comprehensive Listing
11. Growing Carnivorous Plants
12. Nature Conservation . Gloss
13. Bromeliaceae: Pineapple Family (Bromeliads) 27. Bibliography
14. Byblidaceae: Rainbow Plant Family
15. Cephalotaceae: Australian Pitcher Pl Famil 29. Carnivorous Plant Societies and Sources
] pl harbor deep darl ts, described as "against the
Sundews and waterwheels and corkscrews! Oh, my! TI
order of nature as didus by God" pas Carl Linnaeus in late 1700s. Set; apart Bom: bue plants € devious Reni use oe
tr
OME 1 + [o]
pu ine. Pd The gory behind thi i I t prise the topic of this book, initially lens by the
language in 2004, then re-released i pdated and revised English-language edition i
In chapters 2-4, the authors identify the open, low-nutrient, and moist sites that cl teri i plant habi d tl
bd of peus nan between 20%N and Ae latitude. Chapters 5-10 di the g lized y by which tl lant
1 tactile, A ^t. y $ ^ s 1 fadl
8 y prng
snapping, oe or suction), “digestion: d tyg Íp Belief i pletely darl f tl pl is dispelled hapt
f 1 1 1 r : 1 el
8 and 9 r
as a nutrient source. Carnivorous p t uniq facilitat f animal dan and ihe animal-killing capabilities of ‘ normal"
pus Gomever unintentional the out f their herbivory def ics) ied macussed: Chapters 11 and 12 comprise the
f, ; 1 ç Sd 4: E
any ca i Į g what to look for when picking out
Sind I ] maintain them, to the species best suited to begininere
ral n da 13 Eu take the reader deeper into tl jor groups of i pl li ing 1 dicot
AT it See | E T RA PES f 4l + -l x 1 E
g , Test y groups of p y associate
with carnivory J he li df i (the £ I did i i tested, as their prey are protozoans).
These chapters include preliminary discussions of each family, what it is related to, and what portion of its members are carnivores.
Also included are sections dd each Pe where OSO nOs "e PRO Lal features, trapping mechanism, digestion and
om life- a onto carnivores (Byblis
prey, selected species,
é o e 4
P | bs 1 1 1 : 1 - fel O Ea ip 1 (T1
gigantea: B or during certain
(Pinguicul daa Carnivory across habi i is noted i in epiphytes o and pos E
T£ 1 11
E E Eu D
$ 4 f. 1 L eu f£», Hs fl 31 = 1 mi fel
are related
the er of iN EGO appar, THER had pus Ri ered 62 ye TUS earlier by Carilo Golgi.
d botanical } ledge; it i i dibl
, DACKEU
This
semblage of facts, fom the ecology and physiology E carnivorous peus to the p of od names dnd us uses, to in pure and
hands
hand inis DOOR recom
ended ro Eon ni any interest whatsoever in cultivation of E and unusual plants.—Tiana E Franklin, Botanical Rudi
Institute of Texas, 500 East 4th Street, Fort Worth, Texas 76102-4025, U.S.A.
J. Bot. Res. Inst, Texas 2(2): 798. 2008
ILLICIUM GUAJAIBONENSE, ELEVATED TO SPECIES RANK AND COMPARED
WITH THE SUBSPECIES OF ILLICIUM CUBENSE (ILLICIACEAE)
Walter S. Judd J. Richard Abbott
Department of Botany iic of icd Herbarium
214 Bartram Hall 9 Dickinson Hall
ok x 118526 Florida n of oe History
Bs of Horid University of Florid
mE n 3261 a USA, Gainesville, Florida 32611, U.S.A.
wjudd@botany.ufledu jrabbottabotany.ufl.edu
Ashley Morris
Department of Biological Sciences
University of South Alabama
Mobile, Alabama 36688, U.S.A.
abmorrisgusouthal.edu
ABSTRACT
lies h guajail has] llected on the slopes of Pan de Guajaibón, Pinar del Rio, Cuba. The smooth epidermal
cells of the petioles un f tl plant d their red flowers with usually ol i tepals, which are recurved-
spreading, supr gnition of thi ity disti pecies, Illicium guajaib ! and anew ue is pov The
“224 : 1 led 1 Handed A t d d 1 ]
AUNT ly o r r Ev r r oO e A 44 E r
cubense.
RESUMEN
Illicium cubense subsp. guajaibonense ha sido re-encontrada en las vertientes pe Pan de ón Pinar del Río, ee Las ae
epidérmicas lisas de los peciolos y de los tallos de estas pene) junto a j p
lingulados, los cuales son recurvados-extendidos, de esta entidad una E distinta, Mlicium gua-
jaibonense, Pent lo qu prop inb cen Se provee una descripción ampliada para la esp I ida y se
compara con las p i geografi i de I. cubense
Key Wonops: Illicium, Illiciaceae, Cuba, Pinar del Rio
In the course of fieldwork in the province of Pinar del Rio in western Cuba the second author collected a
distinctive species of Illicium sect. Cymbostemon (see Morris et al. 2007) on the slopes of Pan de Guajaibón,
on the western edge of the Sierra del Rosario. Subsequent study of these specimens (along with herbarium
material of Illicium cubense A.C. Smith occurring in both western and eastern Cuba, as well as I. ekmanii A.C.
Smith, I. floridanum J. Ellis, I. hottense Guerrero, Judd & A.B. Morris, and I. parviflorum Michx. ex Vent., all
of which belong to sect. Cymbostemon) have led to the realization that these Pan de Guajaibón plants, which
had been described as a subspecies of Illicium cubense (Imkhanitskaya 1993; i.e., I. cubense subsp. guajaibo-
nense), are better understood as a distinct species. Thus, we propose here the new combination Illicium
guajaibonense for this distinctive Pinar del Rio entity.
Illicium cubense, a Cuban endemic, was considered by Imkhanitskaya (1993) to be widely distributed
on the island, and she grouped its rather variable populations into five subspecies. Our observations (based
on 29 collections, excluding duplicates) suggest that these four subspecies, in addition to I. cubense subsp.
guajaibonense, have consistent morphological differences (among many other polymorphic features) that
correlate with their geographical distributions, and they are thus easily diagnosed. As discussed in more
detail below, we found the epidermal features of the petiole and young stem to be especially useful in diag-
nosing these entities (although distinctive floral features also occur in subsp. guajaibonense). Of these sub-
species, I. cubense subsp. rangelense Imkhanitskaya, like subsp. guajaibonense, is restricted to western Cuba,
J. Bot. Res. Inst. Texas 2(2): 799 — 806. 2008
" i lof tha Botanical D h Institute of Texas 2(2)
where it is restricted to the Sierra del Rosario, near Rangel. Illicium cubense subsp. cubense, subsp. bissei
Imkhanitskaya, and subsp. guantanamense Imkhanitskaya are endemic to the mountains of eastern Cuba,
with subsp. cubense occurring in the Sierra Maestra, including the Gran Piedra range (southern "Oriente"),
subsp. bissei occurring in the Sierra de Nipe, Sierra de Cristal, Cuchillas de Moa, Sierra Mina de Iberia,
Cuchillas de Toa, and Cuchillas de Baracoa (all in northern “Oriente”), and subsp. guantanamense occurring
in the Sierra de Purial, Sierra de Imías, and the Meseta del Guaso (prov. Guantánamo).
Taxonomic reconsideration of the status of these remaining subspecies of Illicium cubense must await
field investigations and the study of additional specimens (especially of material in Cuban institutions) of
the taxonomically difficult eastern Cuban entities (subsp. bissei, subsp. cubense, and subsp. guantanamense)
as well as the western Cuban taxon, I. cubense subsp. rangelense. The latter is especially poorly collected, and
field studies of all of these entities would also be of value in working out the systematics of these plants (and
such studies are currently underway; Alejandro Palmarola B., personal communication). Thus, we restrict
our current focus to I. guajaibonense, since for this Pinar del Rio endemic we have adequate material to
understand its pattern of variation. Our additional collections (and especially the field observations of J.R.A.)
allow 1. guajaibonense to be described in more detail than previously has been possible, and it is now read-
ily distinguished from the various entities provisionally retained within : e We thus provide a full
description and a new combination below, along with a discussion of morphological features that distinguish
this new species from I. cubense. Finally, we note that our preliminary studies suggest that all of the subspe-
cies of I. cubense may be better treated at the species level, and these subspecies are briefly compared.
In the following description we have based measurements on leaves that are presumably fully ex-
panded, avoiding leaves at the apex of shoots if they are smaller than nearby leaves; we note that young
leaves often dry a different color than adult leaves. The measurements of both vegetative and floral parts are
based on dried herbarium material.
ids a (Imkhanitskaya) Judd & Abbott, comb. et stat. nov. (Fig. 1). illicium cubense A.C. Smith
guajaibonense ll nitskaya, Bot. di 78:2. 1993. Tre: CUBA. Prov. Pinar DEL Rio: northern slopes of Pan de Guajaibón,
16 a 1953, Bro. Alain & J. Acuña 2972 : HAC GH).
Shrub to ca. 2 m tall; bark grayish brown, slightly fissured; young stems brownish to grayish, 1.6-2.7 mm
in diameter near the lowest leaves, + terete to obscurely angled, with decurrent ridge below some nodes, the
surface smooth (although lightly to heavily wrinkled on drying) with rectangular epidermal cells that are
neither papillate nor bullate. Petiole (0.7-)1.1-1.3 mm in diameter near middle, slightly wider and swollen
at point of attachment to stem, narrowing to midrib, but difficult to delimit because blade often slightly
decurrent onto petiole, drying dark (blackish), 4-10 mm long, excluding apical portion with obvious
laminal flange, although there are often slight ridges on either side of the petiole adaxially, smooth, at least
when fresh, wrinkled on drying, the epidermal cells neither bullate nor papillate, sometimes arranged in
transverse rows; leaf blade thinly coriaceous, somewhat crisply succulent when fresh, and not particularly
tough or fibrous, fragmenting readily when dry, light green abaxially when fresh but reddish brown when
dried, dark green adaxially when fresh, more or less shiny (due to waxy cuticle) but dull grayish brown with
pale greenish tinge when dried, generally x elliptic to elliptic-obovate, occasionally ovate (the widest point
mostly at or above the middle, sometimes parallel sided for about 10 mm near mid-leaf), 3.4-9.5 cm long,
1.4—4.2 cm wide, the base acute to cuneate, forming a 25—45? angle, often with a small flange of decurrent
tissue along the upper petiole, the apex mostly + acute, sometimes obtuse and bluntly rounded, forming a
30—45(-70)? angle, rarely with a small emarginate indentation; margin minutely revolute, either uniform
throughout or the revolute portion slightly wider at base and narrowing toward apex, sometimes the extreme
apex with margin plane; midvein shallowly sunken above, becoming more or less flush with the blade near
the apex, when dried often inconspicuous for the apical 4-15 mm, slightly darker than blade, strongly raised
below, very rounded near base, becoming flattened and less distinct towards apex, although usually clearly
visible all the way to the tip, concolorous with blade to much darker, with epidermal cells nearly isodiamet-
ric near base, ranging to rectangular and then very elongate towards mid-leaf and apex, the dary veins
Judd et al., Illicium guajaibonense, comb. nov. 801
En
F- 3
1 - ' j
Fic. 1 Hipi es J £n lar LA faha Ahh 19007
, Y d
]
obscure (not visible on many leaves), but at least a few leaves with 2-5 visible y veins per side (but
generally only a small portion of the proximal part of the vein visible), loosely looping together near the
margin (i.e., brochidodromous) and with irregularly developed and only poorly visible tertiary veins (but
these only seen with transmitted light), with spicy sweet, non-anise fragrance when crushed. Flowers axil-
lary, but inflorescences distally clustered on branches and thus appearing terminal, with 1—4 inflores-
cences per stem, although apparently only one at a time with open flowers, the flowers 1—3 per cluster;
subtending bracts (in tight cluster at base of pedicel) (3—)4—5, rounded ovate, sometimes broadly so, ca. 0.5
x 0.8 mm, the margin ciliate to nearly entire, persistent through anthesis, but perhaps deciduous in fruit.
Pedicels slender, ca. 0.7 mm wide near middle, sometimes nearly the same width from base to apex, but
usually widest at base and narrowing to apex, 6-11 mm long at anthesis, nearly straight to recurved,
flexuous; with 0—3 bracteoles, usually 1 or 2 of these near pedicel base and 1 near apex, broadly rounded-
ovate, but curved around pedicel and sometimes appearing + triangular in side view, 0.6—1 x 1-1.2 mm,
the margin ciliate. Perianth segments (tepals) 12-16, the 3-5 outermost tepals similar to large bracteoles,
greenish (when fresh), broadly rounded-ovate to somewhat bluntly triangular, 1.2-1.8 mm long, 1.7-2.5
mm wide, with margin ciliate, the next 5-8 tepals much larger, 4.5—7.6 mm long, 2.5-4.1 mm wide, red
(or the outermost ones red and mottled with green or pale brown), non-papillose, obovate to lingulate to
sometimes slightly unguiculate (i.e., somewhat clawed at base, with narrow, pinched-in portion above
extreme base), recurved-spreading, weakly concave to convex adaxially, with margin ciliate (outermost) to
entire (innermost), and 2 or 3 of this set often cupped near the upper margins, the innermost 3—5 tepals
grading into the stamens in size (and 1 or 2 may even be stamen-like, with non-functional thecae), same
I Lafthna D * al D hindi
80 0 | | | | | | | | | / Jeurmal oft t f Texas 2(2)
color and shape as larger tepals, although none cupped or ciliate margined; generally the outermost green-
ish tepals persist after the reddish, petaloid tepals and stamens xd fallen, the surface of the tepals mostly
smooth except for drying-related wrinkles, with glandular idiol t as lighter punctations. Stamens
5-7, uniseriate, except for 1 or 2 that are sometimes intermediate E tepals, 3.33.7 mm long, 1-1.7 mm
wide, carnose (i.e., thicker and fleshier than the tepals when fresh, although not apparently so when dried),
red, the filaments laminar, the anthers poorly developed, with thecae introrse, sub-immersed in the con-
nective tissue, ca. 0.8 mm long, narrowing to a truncate apex. Carpels 7-8 in a single, tight, ring-like whorl,
2.5-3 mm long, the ovary laterally compressed, curving inwards above; style 0.8-1.2 mm long, erect to
curved-spreading. Fruits and seeds not known.
Distribution and ecology.—lllicium guajaibonense is endemic to Pinar del Rio, Cuba, where it has been
collected on the slopes of Pan de Guajaibón (at the western edge of the Sierra del Rosario) at 500—600 m
elevation, and at Loma Espanol, San Cristobal. At the Pan de Guajaibón locality the species occurs on baux-
itic soil in mesophyllous evergreen forest with Adiantum spp., Blechnum fragile (Liebm.) CV. Morton & Lel-
linger, Buchenavia tetraphylla (Aublet) R.A. Howard, Brysonima sp., Calophyllum antillanum Britton, Cithar-
exylum sp., Cyathea arborea (L.) Sm., Cyrilla sp., Daphnopsis sp., Dendropanax arboreus (L.) Decne. € Planch.,
Lagetta wrightiana Krug & Urb., Matayba apetala Radlk., Mecranium haemanthum Triana ex Cogn., Miconia
prasina (Sw.) DC., Ocotea leucoxylon (Sw.) Laness., Odontosoria sp., Ouratea elliptica (A. Rich.) M. Gómez,
Pachyanthus tetramerus Urb. Sr Ekman, Polystichum wrightii (Baker) C. Christensen ex Maxon, Pseudolmedia
sp., Sagraea capillaris DC., Selaginella spp., Talipariti elatum (Sw.) Fryxell, Ternstroemia sp., and Trichomanes
scandens L
Phenology.—Flowering material has been collected in e and July.
Additional coilections. CUBA. Pinar del Rio Prov.: Pan de Guajaibón, f Si del Rosario,
old military barracks a little below the summit, 500—600 m, between 22°47'17'N, 8372255. 8"W and 22?47'23.6"N, 83?21'59. 5"W (WGS
84 map datum), 8 May 2004 (fl) Abbott 18885, with Bécquer (FLAS, HAJB, NY, S, US); Loma Español, San Cristobal, 6 Jul, 1860-1864
(£D) Wright 1844 (GH).
DISCUSSION
Illicium guajaibonense belongs in the I. parviflorum complex, ae Illicium sect. Cymbostemon (as recircum-
scribed by Morris et al. 2007) because its stamens h ly thickened fil d sub-immersed
anther thecae. We note that the monophyly of this clade is s supported by molecular data (Morris et al.
2007). Vegetatively, it can be distinguished from the various entities retained within I. cubense subsp. cubense,
subsp. bissei, and subsp. rangelense by the smooth epidermis of its petioles (vs. microbullate, micropapillate,
or tuberculate-papillate; Fig. 2), smooth epidermis of its young stems (vs. micropapillate to tuberculate-
papillate, rarely microbullate or nearly smooth), and only shortly and apically flanged petioles (vs. petioles
with flange of lamina tissue extending to near base); see Table 1 for a more detailed comparison of these
taxa. The flowers of Illicium guajaibonense have red, obovate to lingulate inner tepals, which are recurved-
spreading and adaxially weakly concave to convex, and stamens narrowed to a truncate apex (Fig. 1), while
those of I. cubense subsp. cubense and subsp. bissei (Fig. 3) have yellowish green, suborbicular to elliptic or
ovate, adaxially concave, non-recurved tepals, and stamens abruptly narrowed to a differentiated, obtuse-
rounded apex. The stamens of I. guajaibonense are red while those of I. cubense subsp. bissei are peach-orange
with the apical portion purple-tinged; stamen color has not been recorded in subsp. cubense. The flowers of
I. cubense subsp. rangelense are poorly known, but the tepals of this taxon are broadly ovate to elongate-el-
liptic, oblong, or obovate, and also somewhat spreading, and the stamens are narrowed to a truncate apex.
Thus, the flowers of subsp. rangelense may be more similar to those of I. guajaibonense than to the populations
of I. cubense from eastern Cuba; unfortunately, the color of the tepals and stamens is unknown in subsp.
/curvature.
rangelense, and observations of living flowers are required in order to accurately record tepal sl
Illicium guajaibonense can be distinguished from the poorly known I. cubense subsp. guantanamense by
its red (vs. yellowish green) flowers; we have not compared the vegetative characters of I. guajaibonense with
I. cubense subsp. guantanamense as we have not seen herbarium material of this rarely collected taxon.
Judd etal., Illicium guajaibonense, comb. nov.
803
^x^
T rJ
Fic. 2. V.
ARE L AN]
18885, Fl AS) B.! cubense subsp
F 7
ji nanillata:
4536, GH). D. J. cubense subsp. i
P4
f
Bro. Clemente 6477, GH). C. I. cubense subsp.
san) [|
Bro. Alain 536, GH). Black bar — 1 mm.
Y papillate,
Fic. 3. Mici bissej fi
faha (ALL
Sierra de Nipe, Prov. de Holguín,
IP" 44 Dat . [|
f Texas 2(2)
804 Jour Mut wi
TABLE 1 MA Kalaaiceal sl 1 lien pes ha fl " £ Hl IH a ie Bo ] iA I3cb at lah]
material).
Taxon |. cubense I. cubense |. guajaibonense |. cubense subsp.
subsp. bissei subsp. cubense rangelense
Petiole flange to near base
Petiole texture microbullate
(Fig
2C)
micropapiilate (rarely
Stem texture
microbullate near
nodes)
Young stem (dried micro-ridges
condition
Stem epidermal isodiametric
cells
Leaf margin revolute (blade
metimes also
cupped)
Secondary veins obscure (mostly not
abaxially visible except on
some specimens
with wide shade
leaves)
Flower ccior;
Inner tepal shape yellowish green;
sub-orbicular to,
ovate or elliptic
to near base
micropapillate
(Fig. 2B)
LETS ft.
LE AL
micropapill
nearly smooth)
micro-ridges
isodiametric to
elongate
revolute (blade flat)
obscure (mostly
not visible)
yellowish green;
sub-orbicular to ovate
or elliptic, and
non-recurved
only short, apical
flange
smooth
(Fig. 2A)
smooth
wrinkled (smooth
when fresh)
isodiametric to
rectangular
revolute (blade flat)
generally visible,
especially on
largest leaves
red; obovate to
lingulate, and
recurved-spreading
to near base
tuberculate-papillate
)
(Fig. 2D
tuberculate- papillate
(some narrow,
sharp-pointed)
micro-ridges
elongate
revolute (blade
mostly flat, sometimes
slightly cupped)
obscure to sometimes
visible
color not recorded;
ovate to elongate-
elliptic, oblong, or
ovate, and
and non-recurved
spreading
Illicium cubense primarily occurs in the mountains of eastern Cuba, but subsp. rangelense, like I. guajai-
bonense, is endemic to western Cuba. Illicium cubense subsp. rangelense is phenetically distinctive due to its
tuberculate-papillate stems and petioles, and on some plants the papillae are even narrowed and sharply
pointed apically, looking like minute hairs (Fig. 2D). Additionally, the flowers of subsp. rangelense, as noted
above, are quite different from those of the eastern populations of I. cubense. Thus, I. cubense subsp. rangel-
ense, like I. guajaibonense, is easily distinguished from the other Illicium populations of Cuba. Were subsp.
rangelense better known, we also would have proposed raising it to specific rank, as both seem to be amply
distinct from each other, as well as from the populations of I. cubense from eastern Cuba. Unfortunately, it
has been poorly collected, and (as noted above) the flower color is unknown. The status of subsp. rangelense
is currently being investigated by A. Palmarola B. (as part of his revisionary study of Illicium in Cuba).
Illicium guajaibonense can be distinguished from Illicium parviflorum (of peninsular Florida; see Guer-
rero 1997) by its nearly unflanged (vs. + flanged) petioles, leaves with a spicy, sweet (vs. anise) fragrance,
differently shaped, red (vs. pale yellow) inner tepals, and 7-8 (vs. 10-13) carpels. It can be differentiated
easily from I. floridanum and I. mexicanum (both continental in distribution) because these two species have
more numerous and much more elongated tepals, as well as more numerous carpels. Finally, it is also dif-
ferentiated from the two Hispaniolan species, I. ekmanii and I. hottense (Guerrero et al. 2004) by its nearly
Judd et al., Illicium guajaibonense, comb. nov. 805
unflanged (vs. + flanged) petioles, and differently shaped, red tepals (vs. pale yellowish to pink or red tinged
in I. ekmanii; yellowish, tinged basally with salmon-color or yellowish with peach margins in I. hottense). In
addition I. hottense has outer tepals that are strongly papillate, i.e., the entire surface of the tepals is densely
papillate. The perianth segments of I. ekmanii lack papillae or are only very slightly papillate, i.e., with sparse
papillae in the central region of the tepals and not extending to margins. The tepals of I. guajaibonense are
consistently non-papillate. The petioles of I. ekmanii are omen or with sparse low papillae; those of I. hot-
11
tense are smooth, with a few scattered bumps, or rarely
Phylogenetic analyses using ITS, trnL, trnL-F, and trnG sequence sde (Morris et al. 2007) support the
monophyly of the New World species of Illicium, i.e., section Cymbostemon, as re-circumscribed. Within this
clade, I. floridanum and I. mexicanum are sister to a clade containing (among sampled taxa) I. parviflorum, I.
hottense, I. ekmanii, I. cubense subsp. bissei, and I. guajaibonense, i.e., the I. parviflorum complex. IB two
clades, in addition to being well-supported by molecular characters aio posterior probabilities of 1.00
and 0.98, respectively), reflect very different floral morphologies (Imkhanitskaya 1993; Guerrero 1997),
although we note that lingulate tepals have evolved independently in I. guajaibonense and in the common
ancestor of the I. floridanum + I. mexicana clade. Within the I. parviflorum complex, molecular resolution is
lacking due to short branch lengths, with each species exhibiting one to six pop. DNA characters.
This pattern is consistent with the differences among closely related species ved within the Old World
clade (Morris et al. 2007), indicating that such differences provide additional support for species recogni-
tion. Furthermore, short branch lengths within the New World clade, relative to long branch lengths between
the New and Old World clades, indicate recent speciation within this group, and suggest the possibility of
a history of extinctions within Illicium.
Red flowers are undoubtedly autapomorphic for Illicium guajaibonense, and this morphological feature,
along with the molecular autapomorphies indicated in Morris et al. (2007), indicate that it probably is,
therefore, a cladospecies, satisfying the requirement of the apomorphy species concept (Donoghue 1985;
Mishler & Theirot 2000; Judd et al. 2008). It is evident that I. guajaibonense also satisfies the morphological-
phenetic (Judd 2007) and diagnostic (Wheeler & Platnick 2000) species concepts.
Although Illicium guajaibonense and I. cubense (with its various subspecies) are probably each other's
closest relatives, as evidenced by the reduced number of carpels (Guerrero 1997), we believe that it is prob-
able that I. guajaibonense represents a distinct evolutionary lineage (see de Queiroz 2007), which would be
obscured by considering these Pan de Guajaibón plants within a broadly circumscribed I. cubense. The rank
of subspecies has been used in various ways, but could be interpreted to imply that the boundaries between
I. guajaibonense and the various entities within I. cubense are not entirely fixed. However, as we have shown,
this is not the case for I. guajaibonense, which is amply and consistently distinct from the various entities
within I. cubensis in both vegetative and reproductive characters. It is our view that the rank of subspecies
should be used for geographical or ecological variants with demonstrable intergradation.
Illicium guajaibonense, which may now be limited to a small forested region on the Pan de Guajaibón,
should be considered as “critically endangered" according to the guidelines of the IUCN red data book
categories (Lucas & Synge 1978). The very small population should be protected from development.
Pending a detailed monographic study of New World Illicium, the following preliminary key (with
treatment of I. guantanamense, for which we have not seen specimens, based on characters reported by
Imkhanitskaya, 1993) is provided as an aid to the identification of the species and subspecies of Illicium
occurring in Cuba.
1. Petioles and stems conspicuously tuberculate-papillate (i.e, with elongate, rounded to sharp projections
from the epidermal cells; Fig. 2D; generally tightly packed but with scattered flat, smooth spots where the
epidermal cells can be seen), with some papillae narrow and hair-like; plants of western Cuba (mountains
near Rangel) I, cubense subsp. rangelense
. Petioles and stems microbullate, micropapillate, or smooth (apart from wrinkling related to drying), but
never tuberculate-papillose
—
" ! Lafého Botanical D h Institute of Texas 2(2)
2. Flowers with inner tepals bright red, obovate to lingulate, and recurved-spreading; petioles and young
stems smooth (apart from wrinkling related to drying; Fig. 2A); plants of western Cuba (Pan de Guajaibón)
I. guajaibonense
2.H ith i Is yell | , suborbicular to elliptic or ovate, and non-recurved; petioles and
young stems microbullate, micropapillate, or smooth (apart from wrinkling related to drying); plants of
eastern Cuba
3. Petioles + smooth; planis. of pror AO I. cubense subsp. guantanamense
3. Petio! icrobull tern Cuba, prov. Granma, Holguín, Santiago de Cuba
andn Guantánar (Cuchillas de Baracoa
4. Petioles microbullate (i.e swollen ends of near li | cell | ti
rows of low, rounded, regular, tightly packed, E protuberances; Fig. 20; young stems micro-
papillate (sometimes microbullate near nodes); petiole flange usually to very base; Edu usually
somewhat abaxially curved (to more or less flat in shade); plants of northern "Oriente l. cubense
subsp. bissei
jed jecti fi | id | cells, with tiny, flat, ED
lata ft
, Small,
J
regions between the papillae; Fig. 2
flange usually ends 1-2 mm above petiole base (usually with raised t lines continuing to base);
leaves always + flat; plants of southern "Oriente" I, cubense subsp. cubense
ACKNOWLEDGMENTS
We thank Norris Williams, Keeper of the University of Florida Herbarium (FLAS) in the Florida Museum of
Natural History, and Kent Perkins, Collections Manager, for assistance in processing specimens. We also
thank Emily Wood, senior collections associate of the collections of the Harvard University Herbaria for the
loan of their Cuban Illicium. We thank Eldis Bécquer G. for his assistance to J.R.A. during his field work in
Cuba and Alejandro Palmarola B. for comments on an early version of this paper. We thank Steven Manchester
for his assistance in the production of Fig. 2 and Lorena Endara for assistance with the Spanish abstract.
REFERENCES
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alternative. Bryologist 88:172-181.
Guerrero, A. 1997. A revision of the species of Iilicium subsection Parviflora (Illiciaceae). M.S. thesis. University of
Florida, Gainesville.
Guerrero, A., W.S. Jupp, AND A.B. Morris. 2004. A new species of Illicium subsection Parviflora (Illiciaceae) from the
Massif de la Hotte, Haiti. Brittonia 56:346-352.
IMKHANITSKAYA, N. 1993. The genus Illicium (Illiciaceae) in the flora of the Antilles. Bot. Zurn. (St. Petersburg) 78:
1-15.
Juno, WS. 2007. Revision of Miconia sect. Chaenopleura (Miconieae, Melastomataceae) in the Greater Antilles.
Syst. Bot. Monogr. 81:1-235
Jupp, W.S., C.S. Campsett, E.A. KeLLOGG, PF. Stevens, AND M.J. DonocHue. 2008. Plant systematics: a phylogenetic
approach, 3 ed. Sinauer Assoc., Sunderland, Massachusetts.
Lucas, G. AND H. Synce (compiLers). 1978. The IUCN plant red data book. International Union for Conservation of
Nature and Natural Resources, Morges, Switzerland
Mister, B.D. AND E.C. Teror. 2000. The phylogenetic species concept (sensu Mishler and Theirot): monophyly,
apomorphy, and phylogenetic species concepts. In: Q.D. Wheeler and R. Meier, eds. Species concepts and
phylogenetic theory: a debate. Columbia University Press, New York. Pp. 44—54.
Morris, A.B., C.D. Bat, J.W. CLAvroN, W.S. Jupp, D.E. Sorms, anp PS. SoLts. 2007. Phylogeny and divergence time esti-
mation in /llicium with implications for New World biogeography. Syst. Bot. 32:236-249
WHEELER, Q.D. AND N.I. PLatnick. 2000. The phylogenetic species concept (sensu Wheeler and Platnick). In: Q.D.
Wheeler and R. Meier, eds. Species concepts and phylogenetic theory: a debate. Columbia University Press,
New York. Pp. 55-69,
NEOCALYPTROCALYX MORII (CAPPARACEAB),
A NEW SPECIES FROM CENTRAL FRENCH GUIANA
Xavier Cornejo Hugh H. Iltis
The New York Botanical Garden Department of Botany
200^ St. and Kazimiroff Ave. University of Wisconsin
Bronx, New York 10458-5126, U.S.A. 430 Lincoln Drive
jj bg.org jog gmail. Madison, Wisconsin 53706, U.S.A.
ABSTRACT
We deserihe N 1 ii, a distinctive localized end 1 idered to be a local sporadic variant of Capparis
leprieurii, cd to Fer Guiana. The new species is characterized 55 alternate leaves, eight stamens and ovoid, yellowish ovaries
when dried
d ET T.
E
Key Wonps: Capparaceae, endemic, French Guiana, Ni
RÉSUMÉ
I 11 éce N 1 ] ii, endémique de la Guy francaise, est décrite. re plante était auparavant considérée
Į yp
comme une variante locale di d is leprieurii. Ell érise p ées, huit étamines, et des ovaires
E L FE
ovoides et jaunátres à l'état sec.
Neocalyptrocalyx Hutch. (1967) is a neotropical genus of Capparaceae that comprises seven species, ranging
from the Amazonian lowlands of Brazil and Ecuador to the Guianas and eastern Venezuela. The genus was
recently resurrected and its taxonomical limits clarified (Cornejo & Iltis 2008). While studying the species
of Neocalyptrocalyx, we became aware that several collections from French Guiana, provisionally identified
and cited as Capparis leprieurii Briq. (Iltis et al. 1996: 378; Iltis & Mori in Mori et al. [eds.], 2002, Fig. 66,
pl. 38:e; also used by Iltis as a New Year's card, widely distributed in 1997), consistently differ by having
flowers with 8(to 9) stamens, ovoid rather than oblongoid ovaries, and alternate leaves, representing a new
species herewith described.
Dee morii Cornejo & Iltis, sp. nov. (Fig. 1). Tre: FRENCH GUIANA: Saúl and vicinity, Sentier a
st E of Eaux Claires, ca. 500 m from entrance to trail, ers moist forest, 3°37'N 53°12'W, ca. 300 m, 23 Sep 1994 (fl
Mori, C. Gracie & C. Zinder 24021 (nororveg: NY; isotypes: CAY,
NL 1 Vell.) H AT loni] ; + 1) Iltis affinis ] :1 1; ff, fAuieal ie ersrminibus
MODE 8cc 9) et iis ovati a ex (in BMC:
Tree to 45 m tall, trunk cylindric to base but not buttressed, sometimes angled, to 75 cm dbh, the outer
bark smooth and light brown to light grayish-brown, the inner bark cream or yellow, slash yellow, emitting
fetid aroma when cut; lepidote-stellate throughout. Leaves al ; petioles 1—1.7(—2.3) cm; mature blades
elliptic to oblong-elliptic or narrowly-obovate, 10-23 x 4-9(-11) cm, the base inconspicuously subcordate
to broadly obtuse, the margin entire to repand, the apex acuminate, coriaceous (in vivo), chartaceous (in
sicco), covered by lepidote-stellate hairs on both surfaces, glabrescent; and 9 to 13 main lateral veins on
each side of the midrib. Inflorescences very short corymbose racemes on peduncles to 2 cm long, some-
times apparently subfasciculate due to the absence of a peduncle, terminal, axillary and ramiflorous; floral
bracts linear, 1-2.5 mm, very soon deciduous; flower buds subspherical to widely elliptic nearly to anthesis;
pedicels 10-27 mm. Flowers nocturnal, the outer sepals widely ovate to suborbicular, 7-15 x 7-9 mm,
green to gray-green, entirely covering the flowerbud until anthesis, becoming widely divergent to reflexed
at anthesis, the inner pair of sepals ligulate-oblanceolate, ca. 6-10 x 2 mm; petals obovate, 12-20 x 10-12
mm, white, with cuneate base, wavy margin and rounded at the apex; stamens 8(—9), with filaments 13-25
mm, cream to creamy-white (in vivo), inserted in a single row, with hyaline uniseriate trichomes at the base,
the anthers ca. 2 mm; gynophore 13-25 mm, cream to creamy-white (in vivo), the ovary ovoid, 23 x 2-3
J. Bot. Res. Inst, Texas 2(2): 807 — 810. 2008
808 j lof the Botanical R h Institute of Texas 2(2)
Fic. 1. Neocaly l —À » P. Detail of abaxial surface of nearly fully expanded leaf (right) and enl ,
of stellate l pi lote trict (left) C. Inf in bud. D. M | ] | [ I [ lled d | t. le. E Aj I f fl
i c" zi I cu 4 laf! mad; MARRECI APA DRE an Mon nf Ic G
ad ino itu d Ud sión oy Label. (rom
Iltis & Mori in Mori et al. [eds.] 2002: 187, fig. 66. TI | number of lified by the fi tl
f. j A es : M lunt I ii, H £ n Le... 809
mm, yellowish, the gynophore and ovary are densely covered with whitish-stellate trichomes, the stigma
capitate, sessile, green, turning red at post-anthesis. Fruits unknown.
Additional specimens examined: FRENCH GUIANA. St. Elie: Piste de St. Elie, interfluve Sinnamary/Counamama, 5°20'N 53°0'W,
arbre IV-578, 5 Oct 1992 (fi), Sabatier & Prévost 4084 (CAY [not seen], WIS [2]; Station de la Piste de St. Elie (ECEREX), Parcelle PSEIV
(km 16,3), 5°17'N 53°3'W, 6 Nov 2003 (fl), Prévost & Sabatier 4791 (CAY [not seen], NY). Saúl: Monts La Fumée, Antenne Norte, 9 Oct.
1982 (fl), Prance & Boeke 28085 (CAY [not seen], NY); voucher for ecological study: Tree 81, 3°37'N 53°12'W, 200—400 m, 15 Sep 1982
(£D, Boom & Mori 1697 (MO [not seen], NY); voucher for ecological study: Tree 173, 3°37'N 53°12'W, 200—400 m, 24 Sep 1982 (st), Boom
& Mori 1813(NY); voucher for ecological study: Tree 227, 3°37'N 53°12'W, 200-400 m, 25 Sep 1982 (st), Boom & Mori 1841(NY); voucher
for ecological study: Tree 262, 3°37'N 53°12'W, 200—400 m, 7 Oct 1982 (fl), Boom & Mori 1897 (NY); voucher for ecological study: Tree
271, 3°37'N 53"12'W, 200—400 m, 8 Oct 1982 (fl), Boom & Mori 1904 (CAY [not seen], NY); voucher for ecological study: Tree 274, 3°37'N
53°12'W, 200—400 m, 8 Oct 1982 (st), Boom & Mori 1909 (NY); voucher for ecological study: Tree 280, 3°37'N 53°12'W, 200-400 m,
8 Oct 1982 (st), Boom & Mori 1914 (NY); voucher for ecological study: Tree 284, 3°37'N 53°12'W, 200—400 m, 8 Oct 1982 (st), Boom &
Mori 1917 (NY); voucher for ecological study: Tree 416, 3°37'N 53°12'W, 200—400 m, 14 Oct 1982 (st), Boom & Mori 2062(NY). Vicinity
of Faux Claires, on the Sentier Botanique, on ad pure of ie pipe to the pa ca. n m, 12 s 1991 n d & Gracie
22096 (CAY [not seen], NY, WIS). Saül and Claires, | a
tree 460-3, 3°37'N 53°12'W, 250-350 m, 12 Sep 1994 (st), Mori et P 23897 (NY.
In Neocalyptrocalyx the number of stamens is one of the helpful characters to differentiate species. Thus,
Neocalyptrocalyx nectareus, N. leprieurii and N. grandipetala have flowers with 16 to 18 stamens, N. eichleri-
anus has flowers with 30 to 37 stamens, these four species have the stamens inserted in two rows. However,
Neocalyptrocalyx morii has 8(to 9) stamens inserted in a single row. Neocalyptrocalyx maroniensis has flowers
with 10 to 20 stamens inserted in one to two rows, but the flowers of the latter species are distinctively
smaller. It is interesting to note that eight stamens is one of the characters for several well-defined species
among the neotropical Capparaceae with variously stellate indumenta (e.g. Capparicordis crotonoides [Kunth]
Iltis & Cornejo, Quadrella lundellii [Standl.] Iltis & Cornejo, Q. morenoi Cornejo & Iltis, and Q. steyermarhii
[Standl.] Iltis & Cornejo). A second character to reinforce the recognition of this species is that among the
neotropical species of Capparaceae, the shape and color of the ovaries in dried material do not show con-
spicuous variability. Neocalyptrocalyx nectareus and N. leprieurii have ovaries oblong-cylindrical and brown
(when dried), but the ovaries of N. morii are ovoid and yellowish (when dried). In addition, Neocalyptrocalyx
morii has alternate leaves, while N. nectareus and N. leprieurii usually have spiral leaves. Several of the studied
collections cda Been previously identified as a local variant of Neocalyptrocalyx leprieurii, but none of the
studied sp liate floral features between N. leprieurii and N. morii. These morphologi-
cal differences correlated with the uniform geographical distribution of the studied collections lead us to
recognize Neocalyptrocalyx morii as a distinct new species.
Etymology.—We are pleased to name this new species in honor of Scott A. Mori, the distinguished
specialist on neotropical Lecythidaceae, who collected the type (see details in Iltis et al. 1996: 378), and the
main editor of Guide to the vascular plants of central French Guiana.
Habitat and Distribution. Common but scattered in well-drained tropical moist forests of Saul and St.
Elie, French Guiana.
Phenology.—This nocturnal and presumably bat-pollinated species has been collected with flowers from
September to November.
ACKNOWLEDGMENTS
The authors thank C. Feuillet (US) and an anonymous reviewer for criticizing the manuscript. Bobbi Angell
and Scott Mori (NY) kindly provided the permission to reproduce the plate published in this article. Cony
Decock (MUCL) translated the abstract to French.
REFERENCES
CORNEJO, X. AND H.H. Iuris. 2008. Two new genera of Capparaceae: Sarcotoxicum, M pparis stat. nov., and the
reinstatement of Neocalyptrocalyx. Harvard Pap. Bot. 13:103-116.
laf + D H ID hi FM £f T,
810
HUTCHINSON, J. 1967. The genera of flowering plants (Angiospermae) Dicotyledones. Vol. 2. Clarendon Press,
Oxford.
luis, H.H., LJ. Cumaná, R.E. DELGADO, AND G.C. AYMARD. 1996. Studies in the Capparidaceae XVIII. A new giant-fruited
Capparis (C. muco) from Eastern Venezuela. Novon 6:375-384.
ILris, H.H ano S.A. Mori. 2002. Capparaceae (Caper family). In: S.A. Mori, G. Cremers, C. Gracie, J-J. de Granville,
S V. Heald, and J.D. Mitchell, eds. Guide to the vascular plants of central French Guiana. Part 2. Dicotyledons.
Mem. New York Bot. Gard. 76(2):186-188.
A NEW SPECIES OF LASIOLAENA
(ASTERACEAE: EUPATORIEAE: GYPTIDINAE) FROM BAHIA, BRAZIL
Nádia Roque Silvana C. Ferreira
Universidade Federal da Bahia Universidade Estadual de Feira de Santana
Departamento de Botánica Departamento de Ciéncias Biológicas
Av. Ademar de Barros, s/n, Ondina Herbário. Km 03, BR 116. 44.031-460
40.171-970, Salvador, Bahia, BRAZIL Feira de Santana, Bahia, BRAZIL
nroque@ufba.br
Harold Robinson
U.S. National Herbarium, NMNH
Smithsonian Institution, MRC 166
Washington, DC. 20013-7012, U.S.A.
ABSTRACT
Lasiolaena i d f the Bahia state and previously included six described species. The most a O of Lasio-
laena are the pel arranged le odd the tomentum on the stems, 1 d invol , and the flattened b the pappus. The
A fl Tala + f. 11th 1 1 d T LR |
new p Į
1 L 1 CS | E 1 1 11 th t 1 ] f f,
y p g small amina 4- 9 x 2-4 mm), d
phyllaries are lanate; th ical densel p de acd e Te is oe 5y coarsely o e qs
and pin ish I ially towards apex. The new sp i B i
ogy is presented.
Key Wonps: Asteraceae, Compositae, Brazil, Lasiolaena, Eupatorieae
RESUMEN
T 1 4 A 5 p tadas Pak ] Jj : T A
o L E. E
der 1 1 1 P | f, lad lt + dol tall 1 1 1 A 141 J
r 4 r r
T EA DEN ES d intr : f^ ; f] 1 pl PEN do tad 1 A
LUSH ia ICIIC Ulla MOL pd VESCLALIVA bM
del género la as hojas de Lasiolaena lycl triangul pequeñas (lámina de 4-9 x 2-4 mm), rev: PR y sésiles: los
ee e y DE reaS son Eas el ptáculo cónico es densamente piloso yt nomen. el E PANES está formado por
J 1 7 l äni
rosado, est Į I I E FSP a
información relativa a su habitat y fenclogia.
INTRODUCTION
Gyptidinae s.l. constitutes a complex and problematic subtribe and is, without doubt, polyphyletic (Hind &
Robinson 2007). Basically it is characterized for the weakly distant phyllaries, persistent, the leaves opposite
to spirally inserted, the capitula almost always with ten or more florets, and the pappus usually of many
capillary or sometimes plumose bristles (King & Robinson 1972).
The greatest ion of genera is in Brazil, especially in Northeast Brazil (Hind & Robinson 2007).
Of the 29 genera and ca. 135 species in the subtribe, 24 genera have representatives in the Brazil, 20 are
distributed in the State of Bahia, and nine genera are restricted to that state (King & Robinson 1987; Hind
19992, 2000).
The genus Lasiolaena was established by King and Robinson (1972) in part of morphological and ana-
tomical study that divided the artificial traditional concept of Eupatorium L. in 80 distinct genera. The most
distinctive features of Lasiolaena are the spirally arranged leaves, the tomentum on the stems, leaves and
involucres for which the genus is named, and the flattened bristles of the pappus. The conical receptacle of
the genus was originally thought to indicate relationship to Barrosoa R.M. King & H. Rob. and Conocliniopsis
P
R.M. King & H. Rob., but this character occurs widely among various groups of genera in the Gyptidinae
J. Bot. Res. Inst. Texas 2(2): 811 — 815. 2008
a 1 1 A£4hA Da iral n h Institut f Texas 2(2)
(King & Robinson 1972, 1987). Later, King and Robinson (1987) and Hind (19992, 2000) discussed the
closest relationships of Lasiolaena with Agrianthus Mart. ex DC., Catolesia DJ.N. Hind, Stylotrichium Mattf.
and Bahianthus R.M. King & H. Rob, all of which have densely spirally inserted leaves, but lack the flattened
bristles of the pappus.
Although Lasiolaena is a small genus endemic to ne State of Bahia and where most of six species were
described in the last 40 years, not all the species ar juately di ished in the keys provided by Hind
(1999b), and Oliveira (2006). The Dac used as diagnostic, iude the density of trichomes in the
corolla lobes, lamina, cypselae, and the length of thep t adequately tested. Consequently,
many newer specimens of Lasiolaena in herbaria are Wusdefeemined
Recently, we have collected a new species of Lasiolaena which has completely different vegetative and
floral morphology from all the known species. Here we describe the new species and include comments
about its habitat and phenology.
MATERIALS AND METHODS
The data presented are based on literature revisions and an analysis of Lasiolaena collections and types
available in herbaria ALCB, CEPEC, HRB, MBM, R, RB, SP, SPF, UEFS and US. Morphological studies using
Olympus SZH10 stereomicroscope were carried out on dried material.
TAXONOMIC TREATMENT
Lasiolaena lychnophorioides Roque, Ferreira & H. Rob., sp. nov. (Fig. 1-2). Tre: BRAZIL: Bara: Municipio de
Mucugé, Serra do Gobira, 13°5'S, 41°22'W, 16 Sep 2006, A.A. Conceição, D. Cardoso & E.R. Souza 1878 (notorvee: HUEFS!; isoryres:
ALCB!, K)
Ah il LoT A 1 ; Fal; . 1 m HI 1 a 1 IH! 1 2 foliis CO-
riaceis, discaloribus riangularibus laminis 6— ON PHA gi lutis, ilil I ] ico, dense piloso, setis pappi
Aiffort
lanceolatis, barbellatis p
Shrub 0.6 to 1.5m tall. Stems branched, pseudoverticillate, usually densely leafy in distal parts, densely
white-lanate. Leaves alternate, spirally arranged, coriaceous, discolorous, lamina 4—9 x 2-4 mm, triangular,
apex acute, margins revolute, entire, base truncate to cordate, sessile, adaxial surface dark-green, glabrous
since younger, white-lanate abaxially and cryptically glandular-punctate, glandular trichome multicellular,
5—T-seriate, venation obscure beneath. Inflorescence terminal, corymbiform, distinctly exceeding upper
stem leaves, 3-9 short-pedunculate capitula, ra 2-5(-7) mm long, densely lanate, 1—3-bracteolate,
bracteoles 3—4 x 0.3 mm, linear, apex acute, ab irface lanate; capitula ; involucre
campanulate, 7-10 mm tall x 6.5—8.5 mm diam., phyllaries 24-36, 4-6 x 1-1. 5 mm, oblong: -janeeo late:
apex acute, 3—4-seriate, distant, subequal, abaxial surface lanate; receptacle conical, epaleaceous, densely
pilose. Florets 30-36 per capitulum, bisexual, ca. 7 mm long; corolla pinkish, tube 3.5-4.0 mm long x 1
mm diam., cylindrical, moderately glandular-punctate; lobes 0.5—0.8 x ca. 0.4 mm, triangular, acute, sparsely
glandular-punctate, inner surface of lobes mamillose; an anther-appendages conspicuous, ca. 0.3 mm
long, oblong-ovate, rounded; bases of anther tl ther-collar flattened and thickened; style ca.
5 mm long, style-base lacking basal node, glabrous; style-arms 2.5 mm long, slightly clavate, conspicuously
short-papillose. Cypselae prismatic, 2-5 mm long, 5-ribbed, body setuliferous, setulae or long ‘twin-hairs’,
scarcely forked at apices, apices acute; carpopodium annuliform, obscurely procurrent on to base of ribs;
pappus-bristles 20-22, uniseriate, subequal, persistent, 2-3.5 mm long, lanceolate, laciniate at the base,
apices barbellate, acuminate into a bristle-like point, m especially towards tip.
Additi ined. Braz. Bahia: Muni de M id M do Gobira, 1400 m, 11 Dec 2003, N. Roque
s.n. (ALCB 64605, 64770); Morro do Gobira, 13°5'38"S, 41°22! 31"W, 04 Aug 2004, E.L. Borba, A.C.S. Pereira, P.L. Ribeiro & O.A. Oliveira
1805 (HUEFS, K); Morro do Gobira, 13°5'38"S 41°22'31"W, 04 Aug 2004, E.L. Borba, A.C.S. Pereira, P.L. Ribeiro & O.A. Oliveira 1813
(HUEFS, K); Serra do Gobira, plató, 13°5'57"S, 41°22'34"W, 15 Aug 2005, E.C.Oliveria & A.K.A. Santos 106 (ALCB, IBGE, HUEFS, K);
subida Sul para a Serra do Gobira, 08 Sep 2007, S.C.Ferreira, A.L.A. Córtes, M. Ibrahim & S.N. Monteiro 343 (HUEFS)
Distribution.—evidently endemic to the Mun. Mucugé, Chapada Diamantina, Bahia.
Habitat.—sandy soils in campos rupestres, often in scrubby hillside vegetation.
Roaue et al
q ,
Aa
PRU
S S
EN
VN
DESC
NE
Fic. 1.1 lychnophorioides (Borb |, 1813). A. Habit. B. D ly pil ptacle. C. Bracteole. D. Outer phyllary. E. Median phyllary. F. Inner
phyllary. G. Floret. H. Mamillose lobe. I. Conical and epal le ( | ). J. Barbellate. i i | K. Style
Anther witt led appendage. M. Cypsela and pappus. N. Carpopodi 1 ‘twin-hairs: 0. "Twin-hairs' in detail.
Journal of the Botanical Research Institute of Texas 2(2)
q”
Fic. 2. Lasiolaena lychnophorioides. A. Shrub amongst rocks in campos rupestres vegetation. B. Inflorescence corymbiform on tips of leafy branches and
leaves spirally arranged. C. Florets in details (phyllaries dense white tomentum). Pictures from Silvana C. Ferreira.
Roque et al., A ies of Lasiol fi Brazil 815
r
Phenology.—Flowering time: August-December.
Etymology.—The species is named for its similarity in habit to some Lychnophora species from Bahia.
DISCUSSION
Lasiol lychnophorioides has spirally bd d 1 leaves, a character very common in a number of gencia o the
Gyptidinae. S ] t i (stems, abaxial leaf face and involucre),
inflorescence, conical reade with trichomes (except L. carvalhoi) and pappus-bristles barbellate, acute
to acuminate, have reinforced the generic position adopted by the authors. On the other hand, it has a set
of distinctive morphological characters when compared with others species of Lasiolaena.
The leaves are coriaceous, discolorous, triangular, revolute and sessile. Revolute leaves are unknown
in Lasiolaena and indeed in the Gyptidinae. This specific feature is very common in some species of Lychno-
phora (Vernonieae) from Bahia, as Lychnophora phylicifolia, L. blanchetii, L. regis and L. triflora, revealing how
interesting are the convergent processes inside this family. Moreover, the stems, abaxial leaf surfaces and
phyllaries of Lasiolaena lychnophorioides are white-lanate, and not tomentose or puberulous, as commonly
observed in other species of the genus. The conical receptacle is densely pilose in the new species, what is
unknown in Lasiolaena and rare in Gyptidinae. Finally, the pappus is formed by lanceolate, coarsely barbel-
late bristles, which are pinkish, especially towards their tips.
Lasiolaena is one of the most distinctive genera of the Gyptidinae in the campos rupestres of Bahia, and
it occurs in small, geographically restricted populations. It is a prime candidate for studies on population
genetics and morphology that may be useful in measuring the level of variability in the species and help
establish conservation strategies
A phylogenetic study based on molecular data including 24 genera of this subtribe is under progress at
University of Feira de Santana, Brazil. Its main goal is to provide the first assessment of phylogenetic rela-
tionships among the Gyptidinae genera. Data from these results will help determine if Gyptidinae includes
at least three distinct clades, each representing a separate subtribe, as suggested by Hind and Robinson
(2007).
ACKNOWLEDGMENTS
We are grateful to Cássio Van den Berg for the Latin diagnoses and Paulo Takeo Sano for comments. We
also thank Natanael Nascimento and Daniela Guimarães for the illustration plate. Special thanks go to two
reviewers for useful suggestions and critical reading of the manuscript.
REFERENCES
HIND, D.J.N. 1999a. A new genus, Semiria (Compositae: Eupatorieae: Gyptidinae), and a discussion of its affinities
within the Gyptidinae of Bahia, Brazil. Kew Bull. 54:425—432.
Hino, DJ.N. 1999b [2000]. A new species of Lasiolaena (Compositae: Eupatorieae: Gyptidinae), and a synopsis
of the genus. Kew Bull. 54:915-925.
Hinn, DJ.N. 2000. A new genus, Catolesia (Compositae: Eupatorieae), and a discussion of its affinities within the
subtribe Gyptidinae of Bahia, Brazil. Kew Bull. 55:947—948.
Hind, D.J.N. AND H. Rosinson. 2007 [2006]. Eupatorieae. In: Kadereit, JW. & C. Jeffrey, eds. The families and genera
of vascular plants, Vol. 8. Flowering plants - Eudicots - Asterales. Part of series by Kubitzki, K., ed. Kubitzki's
authoritative encyclopedia of vascular plants. Springer-Verlag, Berlin, Heidelberg, New York. Pp. 510—574.
King, R.M. AND H. Robinson. 1972. Studies in the Eupatorieae (Asteraceae). XCVI. A new genus, Lasiolaena. Phyto-
logia 24:185-186.
KiNG, R.M. ano H. Rosinson. 1987. The genera of The Eupatorieae (Asteraceae). Monogr. Syst. Bot, Missouri Bot.
Gard. 22.
Ouveira, E.C. 2006. A familia Compositae no Municipio de Mucugé, Chapada Diamantina, Bahia. Dissertation.
Universidade Estadual de Feira de Santana, Brazil. Unpublished.
J lof the Botanical R h Institute of Texas 2(2)
816
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Store, 11030 East Boulevard, Cleveland, Ohio 44106, www.cbgarden.org, 216-707-2800). $39.95, 160
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Street, Fort Worth, Texas 76102- 4025, US. a
J. Bot. Res. Inst. Texas 2(2): 816, 2008
FOLIA TAXONOMICA 9. NEW SPECIES OF PASSIFLORA SUBG. DECALOBA
(PASSIFLORACEAE) FROM NORTHERN SOUTH AMERICA
Christian Feuillet John M. MacDougal
Department of Botany, MRC-166 Division of Arts and Sciences
Smithsonian Institution Harris-Stowe State University
PO. Box 37012 3026 Laclede Avenue
Washington, DC 20013-7012, U.S.A. St. Louis, Missouri 63103, U.S.A.
feuillec@si.edu threebrane@sigmaxi.net
ABSTRACT
Two new species of Passifl l Decalol d ibed f 1 South America: P. angusta and P. rufa eu status He
hia casas Hoehne is discussed. The d peci ferred to Passifl I Auriculata J.M. MacD
fpl J d lami ies, linear petals, att t l filaments, and s ridged seeds.
A key t th p 4 ru I E E I TJ ]
RÉSUMÉ
D è Illes de Passifl Decaloba du Nord de l'Améri | du Sud. P. angusta et P rufa, sont décrites. Le statut
] d Passifl ti Auriculata I.M. MacD g L&E
de P. P. ppal Hoehne est disce I
au vu des opercules plissés, des E pétiolaires et t laminaires, des petales ingas, des filaments de la
graines striées transversalement. |
Our work toward the treatment of the Passifloraceae for Flora of the Guianas (CF) and the study of subgenus
Decaloba (DC.) Rchb. (JMM) brought attention to collections of Passiflora L. that proved to represent new
species related to Passiflora auriculata Kunth, a tropical American species of subg. Decaloba. Many collections
from the Guianas previously referred to P. auriculata represent P. rufa sp. nov., a species actually more com-
mon than P. auriculata in French Guiana. Other collections from Brazil and the Guianas belong in two
other species, P. angusta sp. nov. from the Guayana shield and P. cryptopetala Hoehne from Amazonian
Brazil. The three species and P. auriculata share several characters with P. fanchonae Feuillet (French Guiana),
P. ferruginea Mast. (Peru), and P. jatunsachensis Schwerdtfeger (Ecuador) and are put in the same species
group. Killip (1938) placed P. auriculata and P. ferruginea in series [Auriculatae] (invalid: no Latin descrip-
tion) at the base of his subgenus [Plectostemma] (invalid: includes the type of subg. Decaloba).
We have proposed supersection Auriculata J.M. MacDougal & Feuillet (Feuillet & MacDougal 1999,
2003; MacDougal & Feuillet 2004) for those species. Passiflora supersect. Auriculata belongs to subgenus
Decaloba. The tip of growing stems is not cernuous. The laminar nectaries are scattered on the leaf blade
and are not marginal or submarginal. Two paired petiolar nectaries are present and are conspicuously en-
larged and elaborated laterally into flaps or ear-like appendages of the petiole, with the glandular area facing
downward. The flowers have reduced linear petals, attenuate coronal filaments, and a plicate operculum.
The seeds are transversely grooved.
The combination of laminar leaf nectaries position, a plicate operculum, and grooved seeds clearly
places supersection Auriculata in Passiflora subg. Decaloba. In this subgenus, supersect. Auriculata is basal
among the advanced groups that have lost the rpoC1 intron (Hansen et al. 2006). It still has petiolar nec-
taries like supersect. Cieca (Medik.) JM. MacDougal & Feuillet or supersect. Bryonioides (Harms) J.M.
MacDougal & Feuillet, yet is advanced in having nonmarginal laminar nectaries and grooved seeds like
supersect. Decaloba (DC.) J.M. MacDougal & Feuillet.
An interesting natural history note (L. Gilbert, per. comm.): Gilbert (University of Texas at Austin) has
"grown P. rufa since 1990 or so, prior to" the present “recognition of its status.” He has “also grown P. jat-
unsachensis for at least that long," as well as "P.auriculata from Brazil, Panama, and Costa Rica. Quite often,
J. Bot. Res. Inst. Texas 2(2): 817 — 824. 2008
818 Journal of the Botanical R h Institute of Texas 2(2)
large plants of all these clade members will mass flower in the same few-day window even if in different
greenhouses. It is as if there are ancestral clock genes shared by all of them that are independent of latitude
of the source populations." One of us (JMM) observed the same phenomenon in the greenhouses at MO in
the 1990s and that stimulated our interest in this group. It is possible that the plants are responding simi-
larly to an environmental trigger instead of a clock. It would be interesting to check if P. fanchonae, another
mass flowering species of this group in cultivation, follows the same pattern and synchronicity.
KEY TO THE SPECIES OF SUPERSECTION AURICULATA
1. Petiolar nectaries abaxial on D ped ene with and proximal to the leaf blade (Guianas) P. fanchonae
1. Petio ies al parate from the lea
2. Be blades E or " with 1 ae oe beneath (wide Neotropical distribution)
P. auriculata
2: i blades pubescent beneath.
Mic at least 3 times as long as wide sparingly hairy above, shortly hairy beneath (Guianas,
Eos uela) > angusta
3. Leaf blades usually much less than twice as long as wide or abundantly and obviously hairy.
4. Leaf blades! ly ovate or sub-orbicular, 3-8 x 3.5-7.5 cm, shortly and densely hyaline puberulent
on both sides (Amazonian Brazil) P. cryptopetala
4. Leaf blades broadly ovate to narrow ovate, 8-25 x 5-15 cm, indumentum colored and obvious,
hirsute above and thick velvety beneat
5. Vegetative parts with yellow trichomes; leaf blades elliptic, usually more than twice as long as
wide (Napo, Ecuador) P. jatunsachensis
5. Vegetative parts with ruf: icl ; leaf blades ovate to broadly ovate, usually once to twice
as long as wide
6. Stem angular; bracts setaceous, 1-2 mm long; petals greenish (San Martin, Peru) P.ferruginea
6. Stem terete; bracts narrowly triangular, 2.5 mm long; petals white (Guianas, NE Amazonian
Brazil) P. rufa
Passiflora angusta Feuillet Sr J.M. MacDougal, sp. nov. (Fig. 1 B, E, H; 2 A-C). Tre: GUYANA. CUruNI-MAZARUNI:
Utshe River Falls, 5°44'N, 61°08'W, 900-975 m, 24 May 1990, fl., T. McDowell €» D. Gopaul 2834 (HoLorvrE: BRG; isorvees: AAU, B,
COL, E MO, NY, TEX, U, US).
areas pee ad A Passi lore iain d aca = liis speciebus folii mm oblongis, laminis 3plo longioribus quam
; ab P. auriculata | tibus di
Herb ES xum tendrils, growing on shrubs. Stems subterete, weakly subangular, drying somewhat
striate and 1 t glaucous, with short pubescence, apex o straight. Tendrils weak,
[e]
short-pubescent. Stipules persistent, narrow-triangular, slightly m 0.7-1 mm long, apex acute, margin
entire, pubescent. Petiole 5-20 mm long, cylindrical to obscurely canaliculate, pubescent, 2 glands under
paired appendages, each appendage lateral, about 1/3 from the base of the petiole, saucer-shape, oval, with
a central gland, ring glabrous, about as wide as the gland, ca. 1.7 x 1.1 mm; blade chartaceous to coriaceous,
rigid, unlobed, narrow-oblong to narrow elliptic, or lanceolate to rarely obscurely 3-lobed and narrowly
ovate, (327-10 x 1-3(-3.5) cm, apex acute to slightly obtuse, base obtuse or rounded to slightly cordate,
margin entire, recurved, adaxially shiny, with scattered, short, stiff hair on the veins, abaxially dull, with
short, stiff hair, with (122—6(-10) laminar glands, not visible adaxially on dry material, round, with a swol-
len circular rim, venation pinnate, raised on both surfaces, usually the proximal vein on each side longer
but not reaching the margin, 3—5 main veins on each side. Inflorescences 2-flowered, axillary; peduncle
lacking; bract near the 2 bracteoles, similar to them; pedicels, 10-15 mm long, joint 1-1.5 mm below the
flower, shortly and densely pubescent; bract and bracteoles scattered in the basal 1/3 of the pedicel, persis-
tent, triangular to triangular lanceolate, 1-1.4 mm long, apex acute, margin entire, shortly and densely
pubescent. Flowers about 1.5 cm in diameter, yellowish green or yellow-white or green or green-white,
fragrant (Maguire 43838); hypanthium saucer-shape, short-pubescent; sepals green outside, white to yellow-
ish green inside, narrow triangular, 1.3-1.5 x 0.3-0.4 cm at base, apex acute to blunt in the same flower,
margin entire, short-pubescent outside, densely pubescent in the basal 1/3 inside spreading; petals white,
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Fic. 1. Passiflora rufa: A. Flowering stem; D. Leaf blade; G. Petiole; J. Seed; K. fruit. Passiflora iind : bed pee ot H. Seed. Passiflora
cryptopetala: G kear blader E, Petiole; Seed: pi D. bl fholotype: US); B. Maguire
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linear, 4 to % as long as the sepals, very thin, apex aristate, margin entire, glabrous on both surfaces, spread-
ing; corona with 2 rows of filaments, outer row 1-1.5 cm long, widened and flattened at base, curly and
attenuate near apex, purplish at base, light green or greenish yellow above, glabrous, inner row 2-5 mm
long, capitate, glabrous; operculum membranous, plicate, din. nectar ring not raised, densely and
minutely papillose; limen annular, not raised, loosely papill nical at base, then terete,
ca. 0.5 cm long, glabrous; stamen at the base of the ovary, Dicen menta narrow, slightly flattened, 0.7-0.8
cm long, glabrous, anthers dorsifixed, oblong, 3 x 0.7 mm; ovary globose, ca. 2 x 1.5 mm, densely covered
with short thick hair, styles pale yellow, ca. 5 mm long, glabrous, stigmas swollen, facing outward, ca. 1
mm in diameter. Fruits globose, about 1.2-1.5 cm in diameter, green or purplish black, short appressed-
pubescent, indehiscent; seeds black, ovate, axis slightly curved, flattened, about 3.5 x 3 mm, with 6-10
main transversal ridges.
The flowers are described on labels as being yellowish green (Henkel 1034), yellow-white (McDowell
3081), green (Maguire 43838), greenish (Steyermark 117272), or green-white (McDowell 2744, 2834; Kral 72111).
The corona is said to be purple at base and green distally (Henhel 1034), olive-green at base (Maguire 43838),
lavender (Steyermark 117816), or purple (McDowell 2834). Most collectors have noted the fruits to be green,
these sometimes with mature-looking black seeds, but the fruits apparently continue to mature until they
are blackish (Gentry 10570) or blackish purple (Benítez 5200).
Ecology and distribution.—Passiflora angusta is an herbaceous climber that crawls on shrubs and small
trees in savannahs and savannah-forest ecotones. It is known from Venezuela (Bolívar), western Guyana,
and Brazil (Roraima), between (270—460 and 1300 m.
Etymology.—The epithet angusta, Latin for *narrow;" refers to the narrow leaf blades compared to those
of P. auriculata.
PARATYPES. VENEZUELA. Bolívar: Mun. Gran Sabana, desde Santa Elena en el Km 284, 900 m, 8 May 1995, fr., C. Benítez & W. D'Arcy
O); Mun. Rau'l Leoni, Pierdra Pintada, area de Pequenna Minería Aza Karón, 270 m, 06?19'11"N, 63?28'00"W, 23 May 1992,
W. Diaz 843 (MO); Mun. Gran Sabana, 4.2 km NE de Uriman, 400 m, 05?24'N, 62°42'W, Mar 1986, A. Fernández 2456 (MO); Río Apon-
guao 2, near bridge over river, km 151-152 S of El Dorado, 1200 m, 16 Mar 1974, A. Gentry et al. 10532 (MO); 133 km 5 of El Dorado,
1300 m, 16 Mar 1974, fr., A. Gentry et al. 10570 (MO); Dist. Roscio, 3 km S of El Paují, 1050 m, 4?30'N, 61°35'W, 19 Oct 1985, B. Holst
& R.L. Liesner 2356 (MO); 7 km E of Kavanayen, 1100 m, 16 Dec 1984, fl., R. Kral 72111 (MO); Río Las Ahailas, 17 km E of El Pauji,
4?30'N, 61°30'W, 850 m, 30 Oct 1985, fl., R.L. Liesner 19182 (MO), 1 Nov 1985, fl., R.L. Liesner 19273 (MO), 1 Nov 1985, fr., R.L. Liesner
19277 (MO); 0-3 km W of El Polo, NW of El Pauji, 4°30'N, 61°40'W, 650—800 m, R.L. Liesner 19588 (MO); 0—6 km SE of El Pauji, 4°30'N,
61°35'W, 800—900 m, 9 Nov 1985, R.L. Liesner dii: an 0-2 km W of El Mcd 4?30'N, 61°35'W, 850-900 m, 10 Nov 1985, fl., R.L.
Liesner 19823 (MO); Dist. Piar, Río Aparamán, afflu Rí iray-merú rapids, 1.5 km S of SW corner O
500 m, 05°54'N, 62°15'W, R.L. Liesner & B. Holst 20203 (MO); Dist. Piar, Rio Acanán, EDELCA Campo Carrao, 5 km NW of NW co
of Amaruay-tepui, 5°57'N, 62°17'W, 470 m, 3 May 1986, R.L. Liesner & B. Holst 20540 (MO); 10 km of Karaurin Tepui, 5°19'N, ND
900—1000 m, 24 Apr 1988, fr., R.L. Liesner 23757 WARME 5 km S of San Ignacio de Yuruani, 4°59'N, 61°10'W, 850 m, 9 May 1988 fl., R.L.
Liesner 24402 (MO, US); Cumbre d Guaiquinima, entre el afluente del rio Carapo (1 km rio arriba del Salto $ I i) y los
peñascos al S del rio, 730—900 m, 05?44'4"N, 63°41'8"W, 24 May 1978, fl., J. Steyermark et al. 117272 (MO); Quebrada El Cajón, Puente
Luis Raúl Vásquez Z., 26.5 km al E de Icabarú, 750 m, 04°25'N, 61°32'W, 18 Dec 1978, fl., J. Steyermark et al. 117816 (MO), Distr. Roscio,
7.5 km NE de Santa Elena de Uairén, 880 m, 4°40'N, 61°04'W, 3 Dec 1982, J. Steyermark & R.L. Liesner 127570 (MO). GUYANA.
Cuyuni-Mazaruni: Imbaimadai Savannahs, near mouth of the Partang River, 460 m, 14 Jun 1960, fL, B. Maguire, S.S. Tillett & C.L.
Tillett 43838 (NY); Top of Warama Tipu, small Mt. across Karamang River, N to Paruima village, 5°50'N, 61°04'W, 530—610 m, 20 May
990, fr., T. McDowell & D. Gopaul 2690 (BRG, NY, US); Utshe camp, 0.3 km N of Utshe River, 5%5'N, 61°08'W, 975 m, T. McDowell &
D. Gopaul 2744 (MO); 7 km N of Paruima village, near Mt. Waleliwatipu, 5°54'N, 61°02'W, 980-1060 m, 30 May 1990, fr., T. McDowell
& D. Hughs 2982 (BRG, US); Macaw creek to Holitipu Camps, hillside of Kartabo Mt., 6°00'N, 61°05'W, 580-1100 m, 4 Jun 1990, fl. &
fr., T. McDowell & D. Gapaul 3081 (BRG, US). Potaro-Siparuni: Pakaraima Mts., south rim of summit ridge, Cipo Mtn., 2-4 km from
Cipo Creek, 4°54'N, 60°05'W, 1250 m, 28 Jan 1993, fL, T.W. Henkel, M. Chin & W. Ryan 1034 (BRG, U, US); Pakaraima Mts., headwaters
of the Mazaruni R., W of Imbaimadai, 5°43'N, 60°20'W, 21 Jun 1986, fr., J.J. Pipoly 7944 (BRG, CAY, NY, US). BRAZIL. Roraima: 1300
m, Dec 1909, fl., E. Ule 8665 (B?, K, MG, U, US).
Passiflora a Hoehne, Comm. Linh. Telegr. Matto Grosso Ann. 5: Bot. pt. 5:76; pl. 112. 1915.
(Fi , L; -F). Lecrore: BRAZIL: near the trijunction point between Amazonas, Pará & Mato Grosso, near San
Manuel, Feb 1912, fl., Hoehne Comissão Rondon 5178 (noroTyer: R; photograph in Hoehne 1915).
UTR
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talhe D Hu " tnla T CI fi A p I +). E Data: el : L +h leat, [| d tha A ] E finarrulum
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7 r
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unfolded (partial).—A-C. McDowell & Gopaul 3081 (US); D—F. Gentry 13058 (MO); G-I. Feuillet 4711
E
(holotype: US)
|| | af +b D H In KI Pa pde
822 Journal of f Texas 2(2)
Herbaceous to suffruticose climber with tendrils. Stems terete, minutely puberulent, apex of growing stem
straight, striate. Tendrils few. Stipules scale-like, triangular 0.2-0.3 mm long, pubescent. Leaves alternate;
petiole 1.5-2 cm long, pubescent, 2 glands under paired appendages, each appendage lateral between
middle and ca. 1/4 from the base of the petiole; blade wide-ovate or sub-orbicular, 3-8 x 3.5-7.5 cm, apex
obtuse or rounded or abruptly and shortly acuminate, base slightly cordate or round, margin entire or with
2 obscure teeth or mucros, shortly and densely puberulent on both sides, with 1-3 glands between the
midrib and the main lateral vein on each side, occasionally between the lateral vein and the margin. Inflo-
rescences 2-flowered, axillary; peduncle lacking; bract near the 2 bracteoles, similar to them; pedicels erect,
recurved when bearing the fruit, about as long as the petiole, pubescent, joint 1.5—2 mm below the flower;
bracteoles near the base of the peduncle, triangular, ca. 1 mm long, pubescent. Bow white, 2.53 cm in
diameter; hypanthium saucer-shaped, outside minutely puberulent, yellow ide; sepals lanceolate,
0.7-1 cm long, white adaxially, outside minutely puberulent; petals linear, oe and um narrower than
the sepals; corona in 2 rows, filaments of the outside row spreading, purplish at base, ligulate, filiform at
apex, those of the inner row oblique or erect, filiform, capitate; operculum membranous, plicate, erect
somewhat curved inward in apical half; androgynophore 3-4 mm long, glabrous; stamen anthers linear-
oblong, slightly purplish at margin; ovary ovoid, pubescent. Fruit globose, ca. 2 cm in diameter, with short
indumentum; seeds ovate, axis slightly curved, flattened, ca. 3 x 2 mm, with 5-6 main transversal ridges.
Killip (1938) offered a broad concept of P. auriculata Kunth and placed P. cryptopetala Hoehne in its
synonymy. We consider the plant to deserve recognition at the rank of species. The original description of
P. cryptopetala was illustrated by a photograph of a herbarium specimen. The above description was drawn
from Hoehne's original description and the specimens cited below. The dried fruits of Rodrigues & J. Chagas
4482 are dark colored, suggesting they might have been dark purple or black when ripe like in most species
in this group.
Ecology and distribution.—Amazonian Brazil, on river banks and non-flooded moist forests.
Vernacular name.—Maracujá bravo (Brazilian Portuguese: Chagas s.n. and Rodrigues & Chagas 4482).
Additional specimens. BRAZIL. m. Fazenda Dimona, Ms km N ou Manaus, 2°19'S, 60*05'W, 50-125 m, 15 Feb 1989, M. Pa-
checo, E. "Palheta & S. de S 202 (INPA, NY); ho, 8 May 1962, fr., W. Rodrigues & J. Chagas 4482
(INPA); Manaus, estrada da Forquilha, marg. mer da Cacheira Alta, 13 Aug 1956, J. Mr s.n. UAA wey soe Manaus-Itaco-
atiara Rd, km 84, 3 Dec 1974, fl., A. Gentry 13058 (INPA, MO). Pará & Mato Grosso: nea Amazonas,
Pará & Mato Grosso, near San Manuel, Feb 1912, fl. & fr., Hoehne Comissáo Rondon 5176 (R), 2 pen omissa Rondon 5177 (R). Pará:
Alto Tapajós, Vila Nova, perto da Cachoeira do Chacoráo, 21 Jan 1952, fr., J.M. Pires 3986 (US).
Passiflora rufa Feuillet Sr JM. MacDougal, sp. nov. (Fig. 1 A, D, G, J, K; 2 G-D. Tre: FRENCH GUIANA.
Montacnes DE Kaw: between Roura and Camp Caiman, ca. 10 km from Roura, 12 Mar 1988, fl., C. Feuillet 4711 (HOLOTYPE: US;
IsoryPES: CAY, P U).
Species haec ad Passifl Auricul i P. fe i laminis foli i ra
ERI
bh alii Jal rn O r, 1 i Ai
Herbaceous to woody climber with tendrils, to 15-20 m high, mostly rufous-tomentose. Stems terete, stri-
ate when dry, green to rufous, tomentose, apex of growing stem straight. Tendrils strong, rufous, tomentose.
Stipules usually persistent, narrow triangular, 5-7 mm long, acute, margin entire, rufous-tomentose. Peti-
ole reddish brown, canaliculate, 1.5—3.5 cm long, tomentose, 2 glands under paired appendages, each ap-
pendage lateral, about 1/3-1/2 from the base of the petiole, 1.5-2 x 2-3 mm, saucer-shape, rufous, oval,
with a central gland, greenish black; blade becoming chartaceous, ovate, 9-25 x 5-15 cm, apex acute, base
cordate, obscurely 3—5-lobed or margin entire, with a few laminar glands on each side in the sector defined
by the midrib and the 2 basal veins, round, with a raised rim, ca. 1 mm in diameter, adaxially dark green,
stiff hirsute (usually appressed when dry), abaxially pale green, rufous-tomentose, venation 3(—5)-palmate,
rufous beneath, raised on both surfaces, camptodromous, 1-3 main veins on each side, secondary venation
scalariform. Inflorescences 2-flowered, axillary; peduncle lacking; bract near the 2 bracteoles, similar to
them; pedicels 2, spreading, 1.5 cm long, rufous, rufous-tomentose, joint 1 mm below the flower; bract and
E Ilag IRA n | M . £ n. ag £ al e sh A H 823
2 bracteoles scattered in the basal 1/5, persistent, narrow triangular, 2.5 mm long, apex acute, margin entire,
rufous tomentose. Flowers facing upward, yellowish white, fragrant; hypanthium saucer-shaped, ca. 0.8 cm
in diameter, yellowish green to rufous green and rufous pubescent outside, yellowish green inside; sepals
triangular, 1.5-2 x 0.6—0.9 cm long, apex rounded with a subapical short awn, yellowish green to rufous
green and rufous pubescent outside, whitish and glabrous inside, spreading; petals linear, 7-8 x ca. 1 mm,
thin, apex acute, whitish transparent when dry, glabrous, spreading; corona with 2 rows of filaments,
outer row 1.5—2 cm long, laterally flattened, spreading, green at base and green spotted with violet brown
in the middle, the apical 1/3 filiform, yellow, contorted, inner row capitate, ca. 3 mm long, pale yellow-green,
white at apex; operculum plicate, erect, ca. 2 mm tall, whitish; nectary ring slightly swollen, pale yellowish
green; limen not raised, whitish; androgynophore terete, 0.6—0.7 cm long, greenish to green spotted with
violet; stamen just below the ovary, filaments flat, narrow, ca. 0.6 cm long, greenish white, anthers dorsifixed,
elliptic, 4 x 1.5 mm, yellowish; ovary globose, 2-2.2 x 1.8-2 mm, pale green, densely pubescent, 0.6 cm
long, greenish white, glabrous, stigmas ca. 1 mm in diameter, pale yellow. Fruits pendent, globose, 1.5—2
cm in diameter, black, hirsute, trichomes white, pulp white, indehiscent; seeds ovate, axis slightly curved,
flattened, about 3 x 2.5 mm, with 5-6 main transversal ridges, black.
The new species resembles P. ferruginea from Peru by its leaf size, shape, and rufous pubescence. Pas-
siflora rufa has 2—6 lateral veins reaching the margin, forming small teeth (Fig.1 A & D). In P. ferruginea the
main veins reaching the margin form a hardened mucro, an uncommon character in the genus.
Passiflora auriculata does not have the rufous pul of P. rufa and is mostly glabrous or glabrescent
with microscopic trichomes. In French Guiana, the ripe fruits of P. auriculata are yellow to orange, those of
P. rufa are dark violet to black, and those of P. fanchonae cherry to dark red.
Distribution and ecology.—Passiflora rufa has been e on des Guiana shield and the Amazonian
basin. The species of supersection Auriculata in French Guian a colonize diff: niches: Passiflora rufa grows
on slopes or well drained areas, on lateritic soil or forest humus, at the edge of forest, P. fanchonae has been
collected near temporary pools or drainage ditches in forested areas, on red soil or forest humus, and P.
auriculata is found at the edge of permanent pools or creeks, generally in open vegetation, mostly on sand.
Etymology.—The new species is named rufa for the rufous trichomes that cover the vegetative parts of
the plant.
Paratypes. BRAZIL. A M E da do Paredáo, 8 Apr 1943, fl., A. Ducke 1213 (MG leaf f ); Pará: Munic. Paragominas
17 km S of Ligação along Belém — Brasilia Highway, near km marker 1509, 250 m, 2 Mar 1980, fr., T. Plowman, G. Davidse, N.A. Rosa,
C.S. Rosário & M.R. dos Santos 9440 (MG). GUYANA. Barima-Waini: Aranka R. head, settlement W e Baramita, 7°25'N, 60°32'W, 107
m, 9 Apr 1991, fr., T. McDowell 4327 (BRG, US). SURINAM. Sipali Itany, 31 fi r. Koulé-Koulé, 25 Jul 1985,
st., C. Feuillet 2803 (CAY, US). FRENCH GUIANA: ona riv. basin, Road Cayenne — Régina, km 50, 11 Jun 1984, fl., C. Feuillet
1402 (CAY, P); km 48, 4 May 1979, fr., M.F. Prévost 575 (CAYO), US); Route de Bélizon, 11 km off road Cayenne — Régina, 52°20'W,
4?25'N, 19 Feb 1988, st., C. Feuillet 4577 (US); Route de Bélizon, 17 km off road Cayenne — Régina, 52°20'W, 4°25'N, 1 Jun 1988, C.
Feuillet 9857 (CAY, US); Station des Nouragues, 27 Feb 1987, seedlings, M.F. Prévost 2216 (CAY); Comté riv. basin, between Route Na-
tionale 2 and Cacao, 52°25'W, 4°33'N, 8 Jul 2005, fl., M.F. Prévost 4876 (CAY, K, MO, P, US), Montagne Cacao, 17 Jan 1983, fl., C. Feuil-
let 545 (CAY, P); 52°27'W, 4°33'N, 1 Jun 1988, fr., C. Feuillet 9848 (CAY, US); 3-4 km from Cacao, N slope, 52°30'W, 4°35'N, 25 Mar
1985, fr., L.E. Shog & C. Feuillet 5683 (CAY). Montagne de Kaw: between Roura and Camp Caiman, 15 Jun 1979, fr., G. Cremers 5780
(CAY(3), US); near Camp Caiman, ca. 22 km from Roura, 23 Nov 1980, fl., G. Cremers 6905 (CAY, US); 9 Jan 1981, fl., C. Feuillet 178
(CAY); western slope, 27 Dec 1980, fl. & fr., G. Cremers 6944 (CAYO), P, US); 17 Apr 1985, fr., C. Feuillet 2098 (CAY, P); 18 dud 1985, st.,
C. Feuillet 2105 (CAY, P); 8 km from Roura, 52°9'12"W, 4°33'N, 26 Nov 1985, fr., C. Feuillet 2884 (CAY, MO, P, US) € dlings, 2884A
(CAY); summital plateau, 28 Feb 1988, C. Feuillet 4672 (CAY); Oyapock riv. basin, near Cr. Patawa, near aan Oyapock, 17
Apr 1981, fr., P. Grenand & M.F. Prévost 2047 (CAY, US); 16 May 1983, fr., M.F. Prévost 1378 (CAY(2))
ACKNOWLEDGMENTS
We would like to thank L.E. Gilbert and P.M. Jergensen for reviewing the manuscript. We are indebted to
Jorge Fontella Pereira of the Museu Nacional/UFR] for providing digital photographs of the three specimens
collected by Hoehne and preserved at R. We are very grateful to Cathy Pasquale-Johnson for the fine illus-
trations. We would like to thank the curators of the herbaria CAY, INPA, MG, NY, P, U for lending the
» : Lafého Datanizal D h Institute of Texas 2(2)
material in their care. This paper is published as No. 85, in the Smithsonian's Biological Diversity of the
Guiana Shield Program publication series.
REFERENCES
FEuiLLET, C. AND J.M. MacDoucat. 1999. Infrageneric classification of Passiflora. In: XVI International Botanical Con-
gress, St Louis, MO.
FEUILLE, C. AND J.M. MACDOUGAL. 2003 [May 2004]. A new infrageneric classification of Passiflora L. (Passifloraceae).
Passiflora 13(2):34-38.
Hansen, A.K., L.E. GiLeenr, B.B. Simpson, S.R. Downie, A.C. Cervi, AND R.K. Jansen. 2006. Phylogenetic relationships and
chromosome number evolution in Passiflora. Syst. Bot. 31:138-150.
Kur, E.P. 1938. The American species of Passifloraceae. Publ. Field Mus. Nat. Hist, Bot. Ser. 19:1-613.
Linnaeus, C. 1753. Passiflora. In: Species Plantarum. Salvius, Stockholm. 2:955—960.
MacDoucaL, 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, OR. Pp. 27-31.
FOLIA TAXONOMICA 10. NEW SPECIES OF NAUTILOCALYX
(GESNERIACEAE: EPISCIEAE) FROM THE VENEZUELAN GUAYANA
Christian Feuillet
Department of Botany, MRC-166
Smithsonian Institution
PO. Box 37012
Washington, D.C. 20013-7012, U.S.A.
i
feuillecasi.edu
ABSTRACT
S i f Nautilocal 1 ibed f he Y | N il N. orinocensis. and N. ruber from
1
r 4: kd
the state of Amazonas; N. paujiensis, N. pusillus, N. roseus, and N. vestitus from the state of Bolívar.
RÉSUMÉ
Sept espèces de Nautilocalyx sont décrites de Guyane Vénézuélienne; Nautilocalyx crenatus, N. orinocensis et N. ruber de l'état
d'Amazonas; N. paujiensis, N. pusillus, N. roseus et N. vestitus de l'état de Bolívar.
The llections made in the last 50 years in Bolívar and Amazonas states of Venezuela uncovered
several new species. In the Gesneriaceae, the new species have a clear affinity for the mountains, the bases of
the tepuis, and the headwaters of nearby rivers. Of course, these locations reflect the nature of the collecting
efforts that took place during the last decades. Since the middle of the 20th century, there was a resurgence
of interest for the Gesneriaceae of the Guiana Shield (Leeuwenberg 1958; Feuillet & Steyermark 1999; Skog
& Feuillet 2008). At the middle of the 20th century, most species ly included in Nautilocalyx Linden
ex Hanst. were included in Episcia Mast. along with many species now placed in Paradrymonia Hanst. Nau-
tilocalyx was greatly augmented with the redefinition of Episcia (Wiehler 1973, 1978). As a group of 57 spe-
cies currently accepted (including the six described below), it is different from Fpiscia by the lack of stolons
(Wiehler 1973) and seems closer to Chrysothemis Decne. than to Episcia as shown by recent DNA analyses
which included one species of Nautilocalyx and of Paradrymonia (Smith 2000; Zimmer et al. 2002; Clark
& Zimmer 2003). All but Smith (2000) placed the Chrysothemis / Nautilocalyx clade close to Paradrymonia.
Clark et al. (2006) sequenced 155 species of the tribe Episcieae, including seven species of Nautilocalyx and
13 of Paradrymonia. Their analysis showed Chrysothemis nested in Nautilocalyx, itself nested in Paradrymonia.
Both Nautilocalyx and Paradrymonia are in great need of morphological and molecular studies to identify
monophyletic groups and of revisions to reflect the resulting clades.
1.—New pink- or red-flowered species of Nautilocalyx
Nautilocalyx crenatus Feuillet, sp. nov. (Fig. 1). Tw: VENEZUELA. Amazonas: Cerro Yapacana, alrededores del campamento
a lo largo del rio en las faldas en la parte suroeste, 3°45'N, 66°45'W, 825 m, 4 May 1970 (fl), J.A. Steyermark & G. Bunting 103068
(HOLOTYPE: US; isotypes: NY, VEN).
Nautilocalyci cataractarum affinis. Planta parviore; sepalis lanceolatis, integris; corollae rubrae, tubo bis longiore; glande non lobata;
ovario bis parviore differt.
Saxicolous herb; leaves clustered at apex of short stem; stem densely long-hairy in young parts. Leaves oppo-
site, equal in a pair: petiole lacking to 1 cm long, densely villous, trichomes 3.5—4.5 mm long; blade elliptic to
oblanceolate, 2.5-8 x 1.2-2.7 cm, base attenuate to decurrent, apex obtuse to slightly acute, margin crenate
to mostly bi-crenate, laxly ciliate, adaxially rugose, deep green or bronze, and laxly pilose with trichomes
1-1.5 mm long, abaxially lavender or purplish, densely antrorsely pilose on main veins, less so on veinlets,
each side of the midrib 6—7 strongly ascending main lateral veins anastomosing 3-5 mm from the margin.
J. Bot. Res. Inst. Texas 2(2): 825 —836. 2008
826 Journal of the Botanical R h Institute of Texas 2(2)
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Fic. 1. Nautilocaly tus, photograph byC Feuill
Inflorescence a reduced pair-flowered cyme, axillary, appearing fasciculate, with several flowers, pedicels
1.5-1.7 cm long densely appressed-villous. Flowers: sepals free, oblanceolate to oblong, slightly unequal,
the larger 4-6 x 2 mm, the smaller 3-4 x 1-1.5 mm, margin 0—3-crenate on each side near apex, densely
villous outside, sparsely pilose near apex within; corolla red, posture oblique to nearly transverse in the
calyx, spur narrow-oblong, 5 x 1-1.5 mm, glabrous, tube 2.3-3 cm long, 0.6 cm wide at throat, appressed-
villous outside, lobes suborbicular, 7 x 7 mm, sparsely appressed-pilose outside, glabrous within; stamens
£M PETI I £ then WM I r 827
Feuillet, N i
7 b J b
with anthers suborbicular-oblong, 1.2 x 0.9 mm; nectary gland dorsal, ovate, 1 mm long, apex obtuse to
rounded, glabrous; ovary long-ovoid, 2 x 1 mm, densely sericeous, style glabrous. Fruit not seen.
Distribution.—Nautilocalyx crenatus is known from Cerro Yapacana (Amazonas, Venezuela) where it
grows in crevices and on wet rocks, 800-1200 m. It was flowering in May.
Nautilocalyx crenatus resembles N. cataractarum Wiehler, from the state of Bolívar, in habitat, habit, and
leaf texture. It differs from N. cataractarum by an overall smaller plant habit and has flowers with sepals entire
at margin, a truly red corolla with a longer and nearly straight tube, an unlobed dorsal nectary gland, and
an ovary half the size. Nautilocalyx crenatus was Nautilocalyx *sp. F" in Feuillet and Steyermark (1999).
Etymology.—The epithet crenatus refers to the crenate or bi-crenate margin of the leaf blades.
Paratype: VENEZUELA. Amazonas: Cerro Yapacana, en la cumbre, 3°45'N, 66°45'W, 1000—1200 m, 5—7 May 1970 (£D, J.A. Steyermark
& G. Bunting 103180 (US, VEN).
Nautilocalyx paujiensis Feuillet, sp. nov. (Fig. 2). Ter: VENEZUELA. Bortvar: 17 km E of El Pauji by road and 64 km
W of Santa Elena by road, 4 km N of highway, Río Las Ahallas, ca. 4930'N, 61°30'W, 850 m, 28 Oct 1985 (fl), R.L. Liesner 19044
(HOLOTYPE: US; sorvetes: MO, VEN)
XI] quoc do eqno pp 1 :1
r r r
oblanceolatis 11 bol i-hirsuto differt
> g ;
Terrestrial herb, decumbent, 25-45 cm tall; stem hirsute, trichomes 3-4 mm long, red, internodes 5-12
cm long. Leaves opposite and decussate, equal in a pair: petiole 2-5 cm long, hirsute, trichomes red; blade
elliptic to oblanceolate, 7-13 x 3.5—5.5 cm, base asymmetrical, obtuse to acute, eet une at ud base,
i 1 :
apex obtuse, and W idely acuminate, Margin crenate to bicrenate
and teeth rarely so, adaxially apese -pubescent, surface flat, abaxially pubescent-hirsute on veins, on
both sides trichomes of dried specimens hyaline, 7—10 main veins on each side of the midrib. Inflorescence a
reduced pair-flowered cyme, axillary, appearing fasciculate, with 3—10 flowers; pedicels 1-4 cm long, hirsute,
trichomes red. Flowers: sepals free, oblong to ovate, 6-8 x 2-3 mm, with few teeth toward apex, hirsute
on both sides, trichomes red; corolla posture slightly oblique in the calyx, red, hirsute outside, trichomes
red, 3-4 mm long, basal gibbosity rounded, 2.5 mm long, 2-2.5 mm wide, tube 2-2.5 cm long, wider 2/3
from base, somewhat narrowed at the throat, lobes suborbicular, 5 x 5 mm, margin entire, glabrous within,
outside hirsute at base, androecium and gynoecium not seen (flowers glued). Young fruit ovoid, nearly as
long as the reddish calyx, hirsute.
Distribution.—Nautilocalyx paujiensis is known only from the type specimen collected 17 km east of El
Paují (Boltvar, Venezuela) were it was growing on mossy rocks and cliffs at 850 m elevation. It was collected
with a few flowers and young fruits in October
Nautilocalyx paujiensis resembles N. kohlerioides (Leeuwenb.) Wiehler, from the basin of the Oyapock
river in Amapá (Brazil) and French Guiana. It differs from N. kohlerioides by the red indumentum, the leaf
blades obtuse to acute at the base, the fasciculate flowers with oblanceolate sepals half the size and by the
hirsute indumentum on the corolla tube. Nautilocalyx paujiensis was Nautilocalyx *sp. D" in Feuillet and
Steyermark (1999).
Etymology.—The new species is named for the community of El Paují near which the species was dis-
covered.
iaa pusillus Feuillet, sp. nov. (Fig. 3). Ter: VENEZUELA. Bottvar: near El Paují, Río Cabass, waterfall, 4°30'N,
61°35'W, 800—900 m, 3 Nov 1985 (fl), R.L. Liesner 19429 (uororvpe: US; rsorvees: MO, VEN
Nautilocalyci } ichi affi lamina fl ioris, corollae non gibbosa, glande non lobata, ovario ter breviore differt.
t E
Saxicolous herb; whole plant hirsute or appressed-pubescent except the stamens and the style. Leaves op-
posite, equal or unequal in a pair: petiole lacking or to 3 cm long, densely strigose; blade linear-oblanceolate
to narrowly oblanceolate, 4.5—15 x 0.7-3.5 cm, base decurrent, apex acute to acuminate, margin minutely
serrate to bi-serrate on the apical 3/4, adaxially gray appressed-pubescent, abaxially hirsute on veins, with
5—6 main lateral veins on each side of the midrib, strongly ascending, ending near the margin but not clearly
en š Laftha Rataniral D h Institut f Texas 2(2)
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Feuillet, N peci f Nautilocalyx f he Vi lan G 829
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Fic. 3. Nautilocalyx pusillus, photograph by C. Feuillet of Liesner 19429 (holotype: US)
anastomosing. Inflorescence a reduced pair-flowered cyme, axillary, appearing fasciculate, 1-2-flowered,
pedicels 13-20 mm long, densely villous with purplish red spreading-ascending, 1.5 mm long trichomes.
Flowers: calyx with lobes free or nearly so, connate for 0-2 mm, lobes subequal, narrowly to linear lanceo-
late, 4-5 x 0.7-1 mm, acute at apex, densely hirsutulous outside, glabrous except near apex within; corolla
posture straight in the calyx, scarlet, not gibbous at base, tube 1.5—2 cm long, densely hirsute outside except
in the calyx, glabrous within, lobes unequal, upper 2 rounded, 2 x 2.5 mm, lower 3 ovate—oblong, 4 x 2.5
mm, stamens 4: filaments 8 mm long, inserted on the corolla 6 mm from the base, glabrous, anthers coher-
ent in 2 pairs, sagittate, 1.2 x 1 mm; gland oblong-lanceolate, 1.2 mm long, sulcate in upper half; ovary
ovoid, 1.5 x 1 mm, sericeous, style 9 mm long, glabrous, stigma bilobed, lobes short, eventually growing
to 1 mm long. Fruit not seen.
Distribution.—Nautilocalyx pusillus grows on boulders or in bluff crevices, 500—900 m, near El Paují
and Icabarú (Bolívar, Venezuela). It was flowering October to December.
Nautilocalyx pusillus resembles N. porphyrotrichus (Leeuwenb.) Wiehler, from Guyana and Venezuela.
It differs from N. porphyrotrichus by the smaller size of the leaves and the flowers, especially its ovary 1/3
the size, and by the lack of a basal gibbosity (“spur”). Nautilocalyx pusillus resembles also young N. arenarius
L.E. Skog & Steyerm., but this latter species has cup-shaped calyces with short lobes and white corollas
twice the size. Nautilocalyx pusillus was Nautilocalyx “sp. G” in Feuillet and Steyermark (1999).
Etymology.—The epithet pusillus refers to the small size of the plant.
" : lof tha Botanical D h Institute of Texas 2(2)
PARATYPES: VENEZUELA. Bolivar: 4 km of El Pauji, Río Chaberú, 750—900 m, 4°30'N, 61°36'W, 12 Nov 1985 (fl), R.L. Liesner 19904
(US); 5 km S of El E be M Río ra a of Río Icabarú, 4?23'N, 61°38'W, 520 m, 23 Oct 1985 (fD, R.L. Liesner & B.K.
Holst 18978 (MO, US, Y jón Luís Raúl Vasquez Z., 26.5 km al este de Icabarú, 4?25'N, 61°32'W,
750 m, 18 Dec 1978 (f), J. E a V RS nu & G. C. K. Dunsterville & E. Dunsterville 117834 (SEL, VEN).
fA t " 1 1 1 fA 4 dd
Nautilocalyx roseus Feuillet, sp. nov. (Fig. 4). Tw: VENEZUELA. Botivar: N sid pui, along Q
Honda, Mar 1969 (fD), G.C.K. Dunsterville & E. Dunsterville s.n. (GoLorvee: VEN)
Nautilocalici cataractarum affinis; internodo longiore, pedicellis longioribus, calycis lobis angustioribus, corollae tubo breviore, cylin-
draceo, antheris discretis, stylo breviore differt.
Terrestrial herb, decumbent; stems at least 10 cm long, densely appressed-pubescent, internodes 4—5 cm
long. Leaves opposite, decussate, equal or subequal in a pair: petiole densely villous, 1.5-2.5 cm long; blade
elliptic-oblanceolate or broadly oblanceolate, 10.5-11.5 x 3.8—4.5 cm, base cuneate, apes acute, mac
densely ] t at base, adaxially densely appressed villous, abaxially appressed
on the veins, 6-7 main veins on each side of the midrib, ascending, anastomosing near the margin. Inflo-
rescence a reduced pair-flowered cyme, axillary, appearing fasciculate, up to 12-flowered; pedicels slender,
15-20 mm long, densely villosulous, trichomes 1-1.5 mm long. Flowers small: calyx fused at base, cup 1
mm long, 3 mm wide, lobes linear-lanceolate, 5 x 0.7-0.8 mm, apex acute with a sclerified, cylindrical, blunt
tip, densely villosulous outside, trichomes 0.7-1.5 mm long, villosulous inside in apical half, glabrous in
basal half; corolla tubular, tube “pale gray rose”, 12 mm long, 3.5 mm wide for most of the length, slightly
dorsally gibbous making the base rounded and moving the insertion on the receptacle in a ventral position,
glabrous ventrally in the basal 4 mm, otherwise villous, lobes suborbicular, “deep rose", the two dorsal ones
4.5 x 3 mm, rounded, glabrous on both sides, the three others rounded, 3 x 3 mm, without pubescent in
basal part, otherwise glabrous, within glabrous; stamens four, included, inserted 3 mm from the base of the
corolla, filaments base lanceolate, 2 x 0.8 mm, sparsely ciliolate, apex filamentose, 5-6 mm long, spirally
twisted, glabrous, anthers not coherent, bilobed in basal half, 1 x 0.8 mm; nectary gland dorsal, about 1 x
0.5 mm, bilobed, glabrous; ovary narrowly ovoid, 2 x 1-1.3 mm, acute, densely sericeous, style glabrous,
6.5 mm long, stigma bilobed. Fruits not seen.
Distribution.—Known only from the type collection from Quebrada Honda, north of Auyan Tepui,
about 10 km west of Angel Falls in Parque Nacional Canaima. It was in bloom in March.
Nautilocalyx roseus was annotated by H. Wiehler as Nautilocalyx sp. aff. melittifolius (L.) Wiehler and
later by J.A. Steyermark as an elongated-stemmed form of the rosulate N. cataractarum Wiehler. In spite of
some similarities and of the geographical proximity of the type localities, N. roseus can be separated from
N. cataractarum (type from the base of Angel Falls) by a series of characters. The leaf margin of N. roseus is
serrate versus crenate and the pedicels are 15-20 vs. 5-10 mm long. The calyx lobes are fused into a cup
at base vs. free and their width is 0.7-0.8 vs. 1.2 mm. The corolla tube is cylindrical vs. ventricose and the
lobes are 3 mm wide and unequal (2 dorsal longer) vs. 5-7 mm wide and equal (or the 2 dorsal slightly
shorter). In the stamens, the anthers are free vs. coherent in a group of four and the filaments are spirally
twisted vs. not twisted. Finally the style is 6.5 vs. 9-10 mm long. Nautilocalyx melittifolius from the Lesser
Antilles is clearly different with pedunculate inflorescences, calyx lobes 3-7 mm wide, and ovary 4-5 mm
long.
Etymology.—Named roseus for the color “rose” (according to the specimen-label) of the corolla.
aca is ruber Feuillet, Sp. nov. (Fi Ig. 5). Tree: VENEZUELA. AMAZONAS: Dept. Atures, 4 km of Río Coro-Coro, W of
a de Yutajé, 9 km NW of settlement of Yutajé, along stream on plateau north of unnamed 1760 m peak, 5°41'N, 66°10'W,
1050-1300 m, 7 Mar 1987 (fl), R.L. Liesner & B.K. Holst 21728 (HoLoTYPE: US; isoTYPE: MO, VEN).
Nautilocal ffinis; 1 g oblanceolata, parviore, petiol lto | palis ob! latis, intus glabris, corollae
Terrestrial or saxicolous herb; stem pubescent to densely villous with 1-1.5 mm long spreading trichomes.
Leaves opposite, unequal in a pair: petiole lacking or to 3 cm long, densely villous; blade oblanceolate,
6-17 x (1.5-)2-5.5 cm, base appearing decurrent but mostly long narrowly attenuate and asymmetrically
Feuillet, N ies of Nautilocalvx f the V lan G 831
Fic. 4. Nautil ly , photograph byC Feuillet of D ille & D ill (holotype: VEN)
rounded at very base, apex rounded to acute, margin ciliate and wide crenate to bi-crenate, adaxially bullate,
silvery around the midrib, with 2-3 mm long trichomes on top of the bullae, otherwise glabrous, abaxially
conspicuously reticulate with areolae 2-4 mm diam., villosulous on the main vein, each side of the midrib
with 8-13 main lateral veins anastomosing 3-5 mm from the margin. Inflorescence a reduced pair-flowered
cyme, axillary, appearing fasciculate, pedicels slender, appressed-villous, 1.5-2 cm long. Flowers: sepals
subequal, free, oblanceolate, 6-8 x 1.5-2 mm, sparsely to densely pilosulous outside, with 1—few teeth near
apex and ciliate at margin, glabrous inside; corolla red to orange-red, tube 2.5-3.5 cm long, 3 mm wide
at base, 6 mm at throat, sparsely puberulent outside, glabrous within except near throat where sparsely
papillate, lobes 7 x 5 mm, the ventral one up to 9 x 6 mm, short appressed-pubescent outside; stamens:
filaments broadened at base where minutely ciliate, otherwise glabrous, anthers broadly oblong, cordate at
base, rounded at apex, 1.2 x 0.9 mm; nectary gland dorsal, 0.8—0.9 mm long and wide, obscurely 3 lobed at
apex, sulcate near base; ovary ovoid, 3.5 x 1.5 mm, short sericeous, style glabrous, 2 cm long. Fruit (ripe?)
ovoid, acute at apex, about 6 x 4 mm, pilosulous at apex.
Distribution. —Nautilocalyx ruber has been collected on the Serranía de Yutajé and Cerro Sipapo
(Amazonas, Venezuela) on stream banks or mossy rocks, 300-1500 m. It was flowering in December and
February-March, and fruiting in December.
The leaves of Nautilocalyx ruber are distinctive from other congeners. Their texture is reminiscent of N.
maguirei L.E. Skog & Steyerm. and of N. pemphidius L.E. Skog, but the blad eolate, |
832 Journal of the Botanical R h Institute of Texas 2(2)
Fic. & Nantilacah n photograph hv( Fauillet of 17 o Uni 21708 (I I US, H TI g h 3 2 cm for scale)
nearly decurrent to the petiole, but asymmetrically rounded at the narrow base. It differs from the white-
flowered N. pemphidius by its much larger and orange-red corollas, and from the red-flowered N. maguirei by
the shape and smaller size of the leaf blades, by the much shorter petioles, by the sepals oblanceolate and
glabrous adaxially, and by the shorter corolla tube. Nautilocalyx ruber was Nautilocalyx "sp. E" in Feuillet
and Steyermark (1999).
Etymology.—Named ruber for the red color of the corolla.
=
PanaTYPEs: VENEZUELA. Amazonas: Dept. Atures, Rio Coro-Coro, stream and slopes E of river, W of Serranía de Yutajé, 6 km N of
settlement of Yutajé, 5°41'N, 66°07'W, 320 m, 21 Feb 1987 (f1), R.L. Liesner & B.K. Holst 21300 (MO, US, VEN); Serranía de Yutajé, Caño
Yutajé, 1500 m, 21 Feb 1953 (fl), B. Maguire & C. K. Maguire 35368 (NY, US); Cerro Sipapo, between Phelps Camp and North Savanna,
1400 m, 17 Dec 1948 (fl & fr), B. Maguire & L. Politi 27746 (NY, US, US).
Although the color of the corolla is not considered here as a proof of phylogenetic relationship, for practical
reasons it might be useful to present the following key.
KEY TO THE PINK- OR RED-FLOWERED NAUTILOCALYX
FROM THE VENEZUELAN GUAYANA
1. Leaf blade ovate or obovate N. chimantensis L.E. Skog & Steyerm.
1. Leaf blade elliptic or oblanceolate.
2. Leaf blade surface flat
3. Leaf blades 12-30 cm long; calyx lobes about 10 times as long as wide______ N. fasciculatus L.E. Skog & Steyerm.
3. np B 6-12 Pu Ung; sav eee ot 3-7 ue as long as wide.
m appresse p anceolate, forming a short cup at the base N. roseus
4. Stem dite calyx lobes cblong to ovate, free N. paujiensis Feuillet
PA i blade bullat
5. Leaf blade da
6. Corolla tube up to 2.5 cm long N. maguirei L.F. Skog & Steyerm.
Feuillet, M . £M PETI [| £. L A [| P al 833
6. Corolla tube more than 2.5 cm long N. ruber Feuillet
5. Leaf blade elliptic.
7. Petiole up to 1.5 cm long.
8. Corolla tube about 1.5 cm long N t Wiehler
8. Corolla tube about 2 cm long N. crenatus Feuillet
7. Petiole more than 2 cm lon
9. Calyx lobes entire; corolla tube 2-3 cm long N. pusillus Feuillet
9. Calyx lobes dentate; corolla tube 3-5 cm long N. porphyrotrichus (Leeuwenb) Wiehler
2.—New white-flowered species of Nautilocalyx
Nautilocalyx orinocensis Feuillet, sp. nov. (Fig. 6). Tire: VENEZUELA. Amazonas: Upper Orinoco river, Sierra Guaharibo,
near Raudal de los Guaharibos, slopes of “Mt. Rimbaud,” light growth near the top, 30 Jul 1951 (fl), L. Croizat 429 (HoLorveE: NY;
photograph MO, negative ++ 05102
AT, 4) hrs] f£: fl “1 Tes “1,
Ò E E
lo bis longiore, corollae tubo ter longiore differt.
Terrestrial or saxicolous (?) herb; shortly pubescent throughout; internodes very short, leaves remaining
only on the young growth. Leaves opposite, equal in a pair: petiole 4—6 cm long, densely villous; blade
obovate, 7-9 x 3-5 cm, base narrowed and then rounded, slightly gi d asymmetric, apex broadly
rounded to obtuse, margin obscurely crenate, adaxially densely and minutely appressed-pubescent, abaxi-
ally densely appressed pubescent on main veins, loosely so on the tertiary veins and glabrescent between
the veins, 4—6 main veins on each side of the midrib. Inflorescence a reduced pair-flowered cyme, axillary,
appearing fasciculate, 2-4-flowered; pedicels 8-10 mm long, densely villous. Flowers: sepals narrowly
lanceolate, 11 x 1.5 mm, densely villous on both sides, ciliate at margin; corolla white, spur narrow, 2 x
1 mm, tube more than 3 cm long, densely villous outside (only corolla damaged, partly hidden by leaves,
glued on the herbarium sheet), glabrous within with papillae in apical part (in bud), lobes glabrous on both
sides (in bud); stamens recoiled in the basal third of the tube; nectary gland dorsal, ligulate-oblong, 1 x 0.6
mm; ovary narrowly ovoid, 2 mm long, densely hirsutulous, style more than 1.5 cm long. Fruit subglobose,
5x5 mm, densely pubescent.
Distribution.—Nautilocalyx orinocensis was discovered near the top of a slope on “Mt. Rimbaud,” Sierra
Guaharibo (Amazonas, Venezuela), in “light growth,” near the Upper Orinoco. It was flowering and fruiting
in July.
Nautilocalyx orinocensis resembles bryogeton (Leeuwenb.) Wiehler from Guyana. It differs from N. bry-
ogeton by much larger flowers: sepals 11 x 1.5 mm (versus 4-6 x 1-1.5 mm), corolla tube, more than 3 cm
long (versus 1.2 cm), ovary narrowly ovoid, 2 mm long (versus globose, 1 mm long). The two species are
similar in their vegetative parts. Nautilocalyx orinocensis was Nautilocalyx “sp. B" in Feuillet and Steyermark
1
Etymology.—The new species is named orinocensis after the Orinoco river.
Nautilocalyx vestitus Feuillet, sp. nov. (Fig. 7). Ter: VENEZUELA. BoLtvar: C del Cerro Guaiquinima, a lo largo del
affluente del Río Carapo, 1 km río arriba del Salto Szczerbanari, 5°44'N, 63°41'W, 730-750 m, 23-24 May 1978 (1D, J.A. Steyermark,
P Berry, G. C. K Dunsterville & E. Dunsterville 117244 (HoLorvrE: VEN
li 1 lesd hi lami 1 l lanceolata. corolla
r r ES E r o T
Planta tota
alba.
Saxicolous herb; stems covered with long brown trichomes. Leaves opposite, decussate, equal or subequal in
a pair: petiole 1—2.5 cm long, covered with long brown trichomes; blade obovate, 3-7 x 2-3.5 cm, covered
with long white trichomes, base acute to cuneate, apex rounded to obtuse, margin serrate-crenate, adaxially
densely pilose except on main veins, abaxially densely pilose, main veins 6—8 on each side of the midrib.
Inflorescence a reduced pair-flowered cyme, axillary, appearing fasciculate, 1—3-flowered; pedicels 4-5 mm
long, covered partly by the indumentum of the stem, covered with short trichomes. Flowers: sepals linear-
lanceolate, 6—7 x 1-2 mm, pale green, covered with long white trichomes, apex curved outward in bud;
corolla white in young bud (according to the specimen-label). Fruit not seen.
834
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Fic. 6. Nautil ly 7 js, photograph by C Feuillet of (roi;
429 (holotype: NY)
Distribution.—Nautilocalyx vestitus was found on the Cerro Guaiquinima (Bolívar) in Venezuela, on
shaded bluffs of narrow rocky quebrada along a small stream, 730-750 m. It was in flower bud in May.
Nautilocalyx vestitus is
jue among the Nautilocalyx species by its vegetative characters. It shares with
Episcia duidae Feuillet, E. rubra Feuillet, and an undescribed Paradrymonia, from the state of Amazonas, a long
and dense indumentum usually characteristic of plants able to survive exposed to strong winds. It will be
Feuillet, M : £ Maur I £ "T 1 p.
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836
necessary to study an open flower to assess the relationships of N. vestitus with other species of Nautilocalyx
Nautilocalyx vestitus was Nautilocalyx “sp. A” in Feuillet and Steyermark (1999).
Etymology.—The epithet vestitus refers to the dense, brownish or white indumentum that covers the
stems and leaves of the plant.
KEY TO THE WHITE- OR LAVENDER-BLUE-FLOWERED NAUTILOCALYX
ROM THE VENEZUELAN GUAYANA
. Leaf blade ovate or obovate.
2. Leaf blade ovate
2. Leaf blade obovate.
3. Lower surface of leaves with tertiary veins hidden by long hairs N. vestitus Feuillet
3. Lower surface of leaves with tertiary veins jous.
4. Leaf blade narrowly rounded; corolla w
4. Leaf blade widely rounded; corolla tube bim lobes lavender blue N. cordatus (Gleason) L.E. Skog
1. Leaf blade elliptic or oblanceolate.
5. Leaf blade surface bullate.
6. Leaf blade oblanceolate, margin large-crenate N. pens L.E. Skog
6. Leaf blade elliptic, margin short-crenate-serrate N. resioide ) Wiehler
5. Leaf blade surface flat
7. Leaf blade not decurrent on the petiole N. arenarius L.E. Skog & Steyerm.
7. Leaf blade decurrent on the petiole.
8. Leaves and bracts reddish
8. Leaves and bracts green or whitish
N. adenosiphon (Leeuwenb. Wiehler
N. orinocensis Feuillet
N. punctatus Wiehler
N. pallidus Sprague
ACKNOWLEDGMENTS
I would like to thank J.L. Clark and L.E. Skog for their careful reviews. I thank the curators of the herbaria
MO, NY, and VEN for lending the material in their care. This paper is published as No. 140 in the Smith-
sonian's Biological Diversity of the Guiana Shield Program publication series.
REFERENCES
CLARK, J.L., PS. HERENDEEN, L.E. SKOG, AND E.A. Zimmer. 2006. Phylogenetic relationships and generic boundaries in the
Episcieae (Gesneriaceae) inferred from nuclear, chloroplast, and Rp ga “a ás 54:313-336.
| for the evolu-
£ AH f
CLARK, J.L. and E.A. Zimmer 2003 A preliminary pl phylogeny OI AHODIECLU
tion of flower resupination. Syst. Bot. 28:365-375.
FEUILLET, C. AND J.A. STEYERMARK. 1999. Gesneriaceae. In: Steyermark, J.A., P. E. Berry, K. Yatskievych, and B.K. Holst, eds.,
Flora of the Venezuelan Guayana, vol. 5. Missouri Botanical Garden Press, St. Louis. Pp. 542-573.
LeEUWENBERG, A.J.M. 1958. The Gesneriaceae of Guiana. Acta Bot. Neerl. 7:291-444.
Skog, L.E. anp C. FeuiLLer. 2008. Gesneriaceae. In: M. J. Jansen-Jacobs, ed. Flora of the Guianas, Ser A: 26:1-136,
Royal Bot. Gard., Kew.
Smr, J.F. 2000. Phylogenetic resolution within the tribe Episcieae (Gesneriaceae): Congruence of ITS and NDHF
sequences from parsimony and maximum-likelihood analyses. Amer. J. Bot. 8/:883-897.
Wiener, H. 1973. Seven transfers from Episcia species in cultivation (Gesneriaceae). Phytologia 27:307-308.
Wiener, H. 1978. The genera Episcia, Alsobia, Nautilocalyx, and Paradrymonia (Gesneriaceae). Selbyana 5:1 1-60.
Zimmer, E.A., E.H. ROALSON, L.E. SkoG, J.K. BOGGAN, AND A. lonurm. 2002. Phylogenetic relationships in the Gesneri-
oideae (Gesneriaceae) based on nrDNA ITS and cpDNA trnL-F and trnE-T spacer region sequences. Amer. J.
Bot. 89:296-311.
FOLIA TAXONOMICA 11.
CONSPECTUS OF VARRONIA (CORDIACEAE: BORAGINALES)
IN THE GUIANA SHIELD WITH THREE NEW COMBINATIONS
Christian Feuillet
Department of Botany, MRC-166
Smithsonian Institution
PO Box 37012
Washington, D.C. 20013-7012, U.S.A.
feuillec@si.edu
ABSTRACT
J; ^N ; 1 r1 f. ` Chisld
The conspectus lists the 17 taxa of e Minus Varronia P. Browne (C
The synonymy and three new bi ded: Varronia bullata suba humilis (Jacq.) Feuillet, V. cremersii (Feuille)
Feuillet, and V. marioniae (Feuillet) Feuillet. Short comments and a key are provided to help identify the specie
RÉSUMÉ
Le conspectus liste les 17 taxa présents dans la flore du Bouclier MUR ou ican Su genre récemment rétabli Varronia P. Browne
(Cordiaceae). La synony i binai t bsp. humilis (Jacq.) Feuillet, V. cremersii
(Feuillet) Feuillet et V. marioniae (Feuillet) Feuillet. De bref: i lé fi is Į l
On the base of strong morphological and molecular arguments, Miller and Gottschling (2007) resurrected
Varronia P. Browne for the shrubby species of Cordia L. with spicate or condensed inflorescences. Varronia
species are easily recognized from Cordia by their spicate or condensed inflorescences and multi-stemmed
habit. They also differ by their leaves usually serrate with a craspedodromous venation and their porate
pollen grains.
Following most authors and especially Johnston (1935, 1936), all modern regional treatments included
Varronia in Cordia (Gaviria 1987; Estrada 1995; Miller et al. 1997). Borhidi et al. (1988) had made many
new names in Varronia for Cordia species and Miller (2007) published several new combinations for Central
America and the Venezuelan Guayana. The Boraginaceae (Feuillet et al. 2007) in the Checklist of the Plants
of the Guiana Shield was at the press when Miller and Gottschling (2007) was published. To provide an
update to the checklist and during the preparation of the treatment for Flora of the Guianas, a conspectus of
Varronia in the flora of the Guiana Shield with three new combinations should help the users and students
of the vegetation of northern South America.
The geographical area covered by this paper includes the following administrative or political entities:
the three Venezuelan states situated east of the Orinoco River (Amazonas, Bolívar, Delta Amacuro) and the
three Guianas (Guyana, Surinam, French Guiana).
1. Varronia bullata L., Syst. Nat., ed. 10, 916. 1759. Cordia bullata (L.) Roem. & Schult., Syst. Veg. 4:462. 1819. Tyre:
MAICA: P Browne s.n. (HOLOTYPE: LINN (Savage Catalog number 255.2); microfiche).
Cordia a , Prodr. 9:498. 1845. (non C. asperrima Spreng.). Varronia asperrima (DC.) Fresen, Bull. Soc. Bot. Genève, sér. 2
24: 155. 1933. Tree: JAMAICA: Bertero s.n. (G-DC, microfiche).
Distribution.—Present from Mexico and the West Indies to Peru and Venezuela. Miller et al. (1997) cite
Guyana for subsp. humilis, but 1 have not seen a Guyanan specimen for this taxon. In our area, only subsp.
humilis (next) is present.
la. Varronia bullata subsp. humilis (Jacq.) Feuillet, comb. nov. Basionya: Varronia humilis Jacq., Enum. Syst. Pl. 14.
1760. Cordia humilis Jacq.) G. Don, Gen. Hist. 4:383. 1838. Cordia globosa var. humilis Jacq.) I.M. Johnst., J. Arnold Arbor. 30:98.
J. Bot. Res. Inst. Texas 2(2): 837 — 842. 2008
" : lof *ha Rataniral D h Institute of Texas 2(2)
1949. lia glol bsp. humilis (Jacq.) Borhidi, Bot. Kózlem. 58:171. 1971 lia bull bsp. humilis Jacq.) Gaviria, Mitt. Bot.
Staatssamml. jMüteberi 23:189. 1987. 1 ia glol bsp. humilis Jacq.) Borhidi, Acta Bot. Hung. 34:385. 1988. Tyre: “Browne,
Civ. nat. hist. Jamaica 1:172, t. 13, f. 1” (ICONOTYPE).
Varronia a Desv., J. Bot. o 1:274. 1808. Cordia dasycephala (Desv.) Kunth, Nov. Gen. Sp. (quarto ed.) 3:76. 1818. Tyre:
VEN )
p Idt & Bo
B-
Humbolat © I
Varronia rai Enum. Syst. PL 14 1760. Cordia globosa Jacq.) Kunth, Nov. Gen. Sp. (quarto ed.) 3:76. 1818. Tyre: "Caribaearum
maritimis” (no original material seen).
Distribution.—In our area: Delta Amacuro, Bolívar. Also: Mexico, Central America, West Indies, central and
eastern parts of N Venezuela.
Varronia bullata subsp. humilis has terminal capitate MR with the fertile part 1.5-2 cm in
diameter. The corollas are 7-10 mm long with five lobes deeply d mucronate. Differing from it,
V. macrocephala Desv. has stellate trichomes and apically rounded corel lobes, and V. bonplandii Desv. from
north central Venezuela and V. sangrinaria (Gaviria) J.S. Mill. from western Venezuela have both terminal
and axillary inflorescences and unlobed or very shallowly lobed corollas.
2. Varronia cremersii (Feuillet) Feuillet, comb. nov. B Feuillet, Novon 13:435. 2003. Tyre: FRENCH
GUIANA: G. Cremers & J.-J. de Granville 13934 (HoLoTYPE: US; isoTYPES:: CAY, NY, P U).
Distribution.—In our area: known only from the type collection in southern French Guiana, on a rocky
outcrop. Endemic.
Varronia cremersii has capitate terminal and axillary inflorescences 1-1.2 cm in diameter, and the styles
first branching is near the middle and the second divisions halfway between the first and the tips. It is easily
separated from V. steyermarkii from Venezuela by the shape of the leaves (cf. key below). It differs from the
Brazilian (Bahia) V. harleyi (Taroda) J.S. Mill. by its shorter petioles, longer and narrower leaf blades that
are cuneate at base and acute to long acute at apex, and its shorter inflorescence peduncles and somewhat
smaller corollas (Feuillet 2003).
3. Varronia curassavica Jacq., Enum. Syst. Pl. 14. 1760. Cordia curassavica (Jacq.) Roem. & Schult., Syst. Veg. 4:460.
1819. Lithocardium curassavicum (Jacq.) Kuntze, Revis. Gen. Pl. 2:977. 1891. Tree: DUTCH ANTILLES: Curacao, E.P Killip & L.
Smith 1067 (weoTYPE: NY; ISONEOTYPE: US).
Lantana bullata L., Sp. Pl. 627. 1753. (non Varronia bullata L.) Montjolya bull L.) Friesen, Bull. Soc. Bot. Genéve, sér. 2, 24:142. 1933.
TYPE: Plukenet, Alamg. 393, tab. 221, fig. 3. 1696, "Salvia barbadensibus dicta, spica florum compactiori.”
Varronia martinicensis sensu J.B. Aublet, L.C. Richard, Lam., non Jacq. 1760.
Varronia macrostachya Jacq., Enum. Syst. Pl. 14. 1760 bua Ruiz & Bion Cordia macrostachya (Jacq.) Roem. & Schult., Syst. Veg. 4:461.
1819, “macrostachia” (non Spreng. 1825). Li (Jacq.) Kuntze, Revis. Gen. Pl. 2:977. 1891. Tree: COLOMBIA:
Cartagena (fide Johnston, 1935).
Varronia guianensis Desv., J. Bot. (Desvaux) 1:270. 1809. Cordia gujanensis (Desv.) Roem. et Schult., Syst. Veg. 4:460. 1819. Montjolya
guianensis (Desv.) Friesen, Bull. Soc. Bot. Genève, sér. 2, 24:142. 1933. Tyre: FRENCH GUIANA: Ex herb. Richard s.n. (HOLOTYPE: P;
ISOTYPE: P)
Cordia divaricata Kunth, Nov. Gen. Sp. 3:74. 1818. Lithocardium divaricatum (Kunth) Kuntze, Revis. Gen. Pl. 2:977. 1891. Varronia
divaricata (Kunth) Borhidi, Acta Bot. Hung. 34(3-4):391. 1988. Tree: VENEZUELA: Humboldt & Bonpland 72 (HOLOTYPE: P; ISOTYPE
B-W, photo. F US)
Cordia graveolens Kunth, Nov. Gen. Sp. 3:74. 1818. Cordia cylindrostachya var. graveolens (Kunth) Griseb., Fl. Brit. WI. 480. 1861. Var-
ronia graveolens (Kunth) Borhidi, Acta Bot. Hung. 34(3-4):391. 1988. Tre: VENEZUELA: Humboldt & Bonpland 1074 (uoLotver: P;
ISOTYPE: B-W)
Cordia canescens Kunth, Nov. Gen. Sp. 3:75. 1818. Lith di Kunth) K , Revis. Gen. Pl. 2:976.1891. Varronia canescens
(Kunth) Borhidi, Acta Bot. Hung. 34(3-4):390. 1988 (non Andersson 1853). Tree: COLOMBIA: Humboldt & Bonpland s.n. (HOLOTYPE:
P; rsorvpe: B-W, photo. E
Cordia spicata Willd. ex Roem. & Schult., Syst. Veg. 4:799. 1819. Tepe: VENEZUELA: Humboldt & B B P
Cordia rugosa Willd. ex Roem. & Schult., Syst. Veg. 4:801. 1819. Type: VENEZUELA: Humboldt € england n. (HOLOTYPE: B- Ww: Die to "
Cordia interrupta DC., Prodr. 9:491. 1845. Cordia cylindrostachya var. interrupta (DC.) Griseb., Fl. Brit. WI. 480. 1861. Lithocardium cylin-
drostachyum var. interruptum (DC.) Kuntze, Revis. Gen. Pl. 2:438. 1891. GUYANA: Hostmann 323 (Lecrorvre: G-DC). The specimen
cited in the text by A.P. de ene "Perrottet" 212, from French Guiana, is a mixture (fide Jobssion 1935). In an infra note, Alph.
iss.”
andolle said “et circa Surinam legit Hostmann ! 323 in
mu. verbenacea DC., Prodr. 9:491. 1845. Tree: BRAZIL: Gaudichaud 532 (uoLorvee: G-DC; isorvees: NY, DL)
Feuillet, Conspectus of Varronia in the Guiana Shield 839
C is DC., Prodr. 9:492. 1845. Lit) di if (DC.) Kuntze, Revis. Gen. Pl. 2:976. 1891. Varronia cuneiformis (DC.)
Borhidi, Acta Bot. Hung. 34(3-4):391. 1988. Tyre: VENEZUELA. Vargas(?18) 90 (G-DC)
Cordia rs DC., Prodr. 9:492. 1845. Lithocardium oxyphyllum O. Kuntze, Revis. Gen. Pl. 2:977. 1891. Tree: GUYANA: Parker s.n.
Cordia us DC., Prodr. 9:492. 1845. Lithocardium salicinum (DC.) Kuntze, Revis. Gen. Pl. 2:977. 1891. Tree: BRAZIL: Vauthier 204
SS TYPE: G- pes ISOTYPE: red ).
burgk, Fauna Fl. Brit. Guiana 960. 1848 (non (Ruiz & Pav.) Roem. & Schult. 1819).
Varronia cylindrostachya sensu Graham, Ann. Carnegie Mus. 22:240. 1934 (non Ruíz & Pav. 1799).
Distribution.—In our area: Delta Amacuro, Bolívar, northern Amazonas, Guyana, Surinam, French Guiana;
especially common near beaches. Also: Mexico, Central America, West Indies, South America except Chile
and Uruguay, mostly in dry vegetation below 2000 m.
Varronia curassavica is characterized by leaf blades that have, at least in our area, adaxially smooth or with
trichomes short erect or more often reduced to their wide conical base and abaxially with curved trichomes
limited to the veins, and spicate inflorescences that are terminal or opposite to the leaves or internodal, but
not axillary.
4. verona oh aaa pis , J. Bot. (Desvaux) 1:273. 1808. Cordia grandiflora (Desv.) Kunth, Nov. Gen. Sp. 3:77. 1818.
Desv) Kuntze, Revis. Gen. Pl. 2:977. 1891. Tyre: VENEZUELA. Humboldt & Bonpland 805 (HoLoTYPE: P;
ISOTYPES: B-W, p)
[Cordia rufa] Kl. in Schomburgk, Reis. Br. Guiana 960. 1848, nom. nud.
Distribution.—In our area: Delta Amacuro, northern Bolívar, northwestern Amazonas, Guyana. Also: Co-
lombia, Venezuela, northern Brazil.
Varronia grandiflora has terminal capitate inflorescences with unlobed corollas more than 3 cm long.
EH : ; Ln
5. Varronia marioniae (Feuillet) Feuillet, comb. nov. B , Novon 13:436. 2003. Tree: GUYANA,
Rupununi District, Dadanawa, 120 m elev., 2°49'N, 59°31'W, 6 June 1995, fl. & fr., MJ. Jansen-Jacobs, B.J.H. ter Welle, C. Gustafsson
& V James 3931 (noLoTYPE: US; isorvPESs: BRG, GB, MO, U).
Distribution —In our area: SW Guyana. Endemic.
Varronia marioniae has characteristic obovate leaf blades attenuate at base and round at apex, spicate
inflorescences terminal or axillary, and emarginate corolla lobes.
6. Varronia polycephala Lam. Tabl. Encycl. 1:418. 1791. Cordia polycephala (Lam.) LM. Johnst., J. Arnold Arbor. 16:33.
1935. IconorrrE: Plukenet, Phytogr. tab. 328, fig. 5. 1691. (see J.E. Sanchez, Fl. Colombia 14:45. 1995)
Lantana corymbosa L., Sp. Pl. 2:628. 1753. Tree: JAMAIC
Cordia corymbosa Willd. ex Roem. & Schult., Syst. Veg. 4:801. 1819. Varronia macrostachya Jacq., Enum. Syst. Pl. 14. 1760. Cordia mac-
rostachya (Jacq.) Roem. & Schult., Syst. Veg. 4:461. 1819, *macrostachia." Lithocardium macrostachyum (Jacq.) Kuntze, Revis. Gen.
1891. Tyre: VENEZUELA: Camara, Humboldt & Bonpland (B-W)
Varronia martinicensis sensu J.B. Aublet, L.C. Richard, non Jacq. 1760. Tere: VENEZUELA: Humboldt & Bonpland s.n. (noLorvre: B-W).
Cordia mariquitensis Kunth, in Humb., Bonpl. & Kunth, Nov. Gen. Sp. 3:75. 1818. Lithocardium mariquitense (Kunth) Kuntze, Revis.
Gen. Pl. 2:977. 1891. Ulmarronia mariquitensis (Kunth) Friesen, Bull Soc. Bot. Genéve, sér. 2, 24:181. 1933. Varronia mariquitensis
(Kunth) Borhidi, Acta Bot. Hung. 34:389. 1988. Tre: COLOMBIA: Tolima (P)
Varronia monosperma Jacq., Pl. Hort. Schoenbr. 1:18, t. 39. 1797. Cordia monosperma (Jacq.) Roem. & Schult., Syst. Veg. 4:463. 1819.
Varronia corymbosa Desv., J. Bot. (Desvaux) 1:275. 1809, illeg. (renaming of V monosperma Jacq. 1797), non L. ex Desf. 1804, nom.
nud. Cordia pé lii (Desv.) G. Don, ion Hist. punt ue non Willd. ex Roem. & Schult. 1819. Iconoryre: VENEZUELA: “Ex
J
Caracas. In cal
J que |
Distribution —In our area: eastern Bolívar, Guyana, Surinam. Miller et al. (1997) cite French Guiana for
Cordia polycephala, but I have not seen a specimen of this taxon from this French region. Also: West Indies,
Colombia, Venezuela, northern and coastal Brazil.
Varronia polycephala has inflorescences adnate to the petiole, branched, usually bearing a few reduced
leaves and with at extremities capitate-looking condensed cymes, and corollas 5-7 mm long, unlobed.
Contrary to the capitate inflorescence species, the calyx lobes are not acuminate and without tips projecting
freely from closed bud.
- . laf tha Botanical D h Institut f Texas 2(2)
7. Varronia polystachya (Kunth) Borhidi, Acta Bot. Hung. 34:393. 1988. Cordia polystachya Kunth, Nov. Gen. Sp. 3:7
[quarto]. 1818. Lithocardium polystachyum (Kunth) Kuntze Revis. Gen. Pl. 2:977. 1891. Tree: VENEZUELA: a
146 (HOLOTYPE: P; iso
Cordia canescens Willd. ex Roem. & Schult., Syst. Veg. 4:797. 1819, non Kunth 1818. Tree: VENEZUELA: Prope Maypure, Humboldt &
Bonpland s.n. (B-W)
Pin ae our area: Ronson pu To be expected in neighboring Colombia.
Vat are terminal and axillary, adnate or not to the petiole,
petiole up to 5 mm long and 5 be corollas. It differs from V. schomburgkii, which has inflorescences always
adnate to the petioles, by the young branches lacking curly trichomes, petioles 10-12 mm long and conical
to campanulate calyx, and from V. stenostachya which has petiole longer than 8 mm and unlobed corollas.
8. Varronia roraimae (I.M. Johnst.) J.S. Mill., Novon 17:374. 2007. Cordi imae I.M. Johnst. Fieldiana, Bot. 28:511.
1953. Tyre: VENEZUELA: Bolívar, Mt. Roraima, J. Steyermark 59003 (HoLorvrE: F; isotypes: GH, US, VEN)
aid E OUI area: por cue
land axillary, adnate to the petioles, the fertile part
is short and narrowly aed to narrowly elliptic, subglobose when young, and unlobed corollas. Contrary
to V. tomentosa, it has calyx lobes that are not acuminate.
9. Varronia schomburgkii (DC.) Borhidi, Acta Bot. Hung. 34:393. 1988. Cordia schomburgkii A. DC., Prodr. 9:490.
1845. Lithocardium schomburgkii (DC.) Kuntze Revis. Gen. Pl. 2:977. 1891. Tree: GUYANA: Rob. Schomburgk ser. 1, 406 (HOLOTYPE:
G-DC; sorres: BM, K, L, BU
Cordia aubletii sensu Pulle, Schomburgk non DC. 1
[Cordia lucida] Splitg. ex Pulle, Enum. Vasc. Pl. ae 397. 1906, nom. nud. Based on Splitgerber 206 (L).
Cordia patens var. polycephala sensu Miquel, non (as form) Cham. & Schltdl. 1
Cordia tobagensis Urban, Repert. Spec. Nov. Regni Veg. 16:39. 1919. Varronia FM (Urb.) Borhidi Acta Bot. Hung. 34:393. 1988.
TYPE: ee Broadway 4235 (HoLotyre: B n.v.; isoTYPE: GH).
Cordia tobagensis Urban var. broadwayi Urban, Repert. Spec. Nov. Regni Veg. 16:40. 1919. Tree: TOBAGO: Broadway 3072 (HoLoryrE: B
n.V.; ISOTYPE: GH).
Varronia guianensis sensu Graham.
Distribution.—In our area: Bolivar, Guyana, Surinam, French Guiana. Also: Trinidad-Tobago, northern
Brazil.
Va homl ii has terminal and axillary, spicate infl dnate to the petioles. The leaves
are erin strigose add somewhat shiny. The inflorescences are up to 25 cm long, including the fertile
and loosely flowered part 5-15 cm long. In flowers, the calyces are enlarged and cup-shaped above a short
tube or seldom from the narrow base, they are covered with small resinous granules, the lobes are strigose
outside. ena) NE me apex. Like most spicate inflorescence species, it has calyx lobes that are not
d lack g freely from closed buds. In contrast, V. spinescens (L.) Borhidi from western
Venezuela has aie ai scattered resinous granules that are strigose but at the very base, the trichomes
on the calyx tube are sometimes hirsute.
10. Varronia stenostachya (Gaviria) J.S. Mill., Novon 17:375. 2007. Cordia stenostachya Killip ex Gaviria, Mitt. Bot.
Staatssamml. München 23:243, fig. 1987. Tree: VENEZUELA: R. Spruce 3642 (HOLOTYPE: K; ISOTYPES: K, W)
Distribution.—In our area: Amazonas (Raudal de Maipures), Guyana (Mt. Shiriri). Endemic of granitic
outcrops.
TZ,
tenostachya has terminal and axillary, spicate infl adnate to the petiole that have
a thinner peduncle and fertile part than other species in our area. The petioles are 0.7-2.5 cm long and the
corollas are unlobed.
11. Varronia steyermarkii (Gaviria) J.S. Mill., Novon 17:375. 2007. Cordi kii Gaviria, Mitt. Bot. Staatssamml
München 23:200. 1987. Type: VENEZUELA: J. Saer d'Heguert 480 (HOLOTYPE: M; isorvees: E, VEN).
Distribution.—In our area: Bolívar. Also: Northern Venezuela.
Feuillet, Conspectus of Varronia in the Guiana Shield 841
Varronia steyermarkii has terminal capitate inflorescences and leaves with linear-elliptic blades and
petioles less than 3 mm long. The style is branching near the apex with short branches.
12. Varronia tomentosa Lam., Tabl. Encycl. 1(v. 2):419. 1792. Cordia tomentosa (Lam.) Roem. & Schult., Syst. Veg., ed.
is [R Sr Schult ] 4:459. 1819. Lithocardium tomentosum (Lam.) Kuntze Revis. Gen. Pl. 2:977. 1891. Montjolya tomentosa
Friesen Bull. Soc. Bot. Genève ser. 2, 24:183. 1933. Tyre: FRENCH GUIANA: s.loc., from herb. Jussieu (P-Lamarck)
Varronia martinicensis sensu Aublet, Hist. Pl. Guiane 1:232. 1775 (non Jacq. 1760).
Cordia aubletii DC., Prodr. 9:490. 1845. Lithocardium aubletii (DC.) Kuntze, Revis. Gen. Pl. 2:976. 1891. Varronia aubletii (DC.) Borhidi
ta Bot. Hung. 34(3-4):389. 1988. Tre: FRENCH GUIANA: Perrottet 211 (G-DC)
Distribution.—In our area: Surinam, French Guiana. Endemic.
Varronia tomentosa has spicate inflorescences that are terminal or axillary and adnate to the petioles.
The leaves are adaxially dull, scabrous with stiff ascending bristles. The inflorescences are up to 15 cm long,
including the fertile and densely flowered part 1.5—4 cm long and often club-shaped. The calyx tubes are
densely hairy or tomentose with intermixed resinous granules. Contrary to the other species with spicate
inflorescences in our area, even V. roraimae with short fertile parts, the calyx lobes of V. tomentosa are nar-
rowly triangular and acuminate and with acumen projecting freely from closed bud.
EXCLUDED SPECIES
Varronia cylindrostachya Ruíz & Pav., Fl. Peruv. 2:23. 1799. Cordia cylindrostachya (Ruíz & Pav.) Roem. & Schult., Syst. Veg.
59. 1819. Tyre: Fl. Peruv. 2: pl 147a “macrostachya.” 1799.
Distribution.—From Venezuela to Peru. Specimens identified as C. cylindrostachya in our area represent V.
schomburgkii.
Varronia ferruginea Lam., Tab. Encycl. 1:418. 1791. te: Ex horto (HoLotvee: P-LA). Cordia ferruginea (Lam.) Roem. & Schult.,
Syst. Veg. 4:458. 1819. = Varronia spinescens (L.) Borhidi, Acta Bot. Hung. 34:393. 1988. (cf. below).
Specimens identified as C. ferruginea in our area represent V. schomburgkii.
Varronia multispicata (Cham.) Borhidi, Acta Bot. Hung. 34(3-4):392. 1988. Cordia multispicata Cham., Linnaea 4:490. 1829.
Lithocardium multispicatum (Cham.) Kuntze, Rev. Gen. 2:977. 1891. Tyre: BRAZIL: Sellow 46 (HOLOTYPE: B, extant?; IsOTYPE: US)
Distribution.—Coastal Brazil. Does not reach the Guianas.
Varronia spinescens (L.) Borhidi, Acta Bot. Hung. 34:393. 1988. Cordia spinescens L., Mant. Pl. 2:206. 1771. Lecroryre: LINN
(Savage catalog N? 253.2) (cf. Miller, 1988).
DEDE m S Mexico to Bolivia and western ea
conical calyx at anthesis. S ifi
d in our area represent V. schomburgkii.
Varronia C is not present either in the Venezuelan states neighboring Venezuelan Guayana.
KEY TO THE VARRONIA SPECIES OF THE GUIANA SHIELD
Infl Ira. J pa |
: p cymose very condensed.
2. Flower heads « 5 mm diam, calyx lobes not acuminate V. polycephala
2. bcn heads » 5 mm diam, calyx lobes acuminate, acumen projecting freely from closed bud.
. Peduncles » 5 cm long; corollae » 2 cm lon n. V. grandiflora
A Peduncles mostly < 4 cm long; coro llae m long.
4. Petiole at least 5 mm long; calyx lobes nad abruptly long acuminate V. bullata subsp. humilis
4. Petiole up to 5 mm lon
5. Leaves ovate, seldom elliptic: peduncle 0.7-2 cm long; corolla lobes broadly elliptic, rounded at
5mm V. cremersii
5. Leaves linear to LOS. peduncle 1-4 cm long; corolla lobes triangular, acute at apex,
5-2x1.5-2m V. steyermarkii
. Inflorescences spic
6. Inflorescences ind or opposite to the ae or intemosan put never Ho ur V. curassavica
6. Inflorescences axillary or terminal and axilla to the petiole at
842 h Institute of Texas 2(2)
V. tomentosa
7. Calyx lobes acuminate, acumen projecting freely from closed buds
7. Calyx lobes not acuminate.
B. be^ J |
+ +h Tila h
t ERE Be mye rir Sie
9. Leaf bl laxially dull; infl ly fl D lla crenate (not 5-lobed) V. roraimae
. Leaf blad laxially shiny; infl | ly fl | (with irregular gaps between groups
of flowers).
- Calyx eap shaped with (seldom without) a short tubular base; corolla with 5 teeth ___ V. schomburgkii
. Calyx wide conical ao corolla without 5 teet V. polystachya
8. | luncl ite to the petiole base.
V. stenostachya
11. Corolla not lobed
11. Corolla 5-lobes.
12. Leaf blades elliptic, acute at apex
12. Leaf blades obovate, widely rounded at apex
V. polystachya
. marioniae
ACKNOWLEDGMENTS
I would like to thank the curators of the herbaria CAY, NY, P, and U for lending the material in their care. This
is number 141 in the Smithsonian's Biological Diversity of the Guiana Shield Program publication series.
REFERENCES
Bonuipt, A., E. GONDAR, AND Z. Orosz-KovAcs. 1988. The reconsideration of the genus Cordia. Acta Bot. Hung.
34:375-423.
ESTRADA, J. 1995. Cordia subgénero Varronia (Boraginaceae). Flora de Colombia 14:1-176.
FeuiLLET, C. 2003. Three new species of Cordia (Boraginaceae) from the Guianas. Novon 13:433-437.
FeuiLLer, C., J. GAVIRIA, R. Gómez, J.S. MiLLER, AND G. Ropricuez. 2007. Boraginaceae. In: V. Funk, T. Hollowell, P. Berry, C.
Kelloff, and S.N. Alexander. Checklist of the plants of the Guiana Shield (Venezuela: Amazonas, Bolívar, Delta
Amacuro; Guyana, Surinam, French Guiana). Contr. U.S. Nat. Herb. 55:224-226
Gaviria, J. 1987. Die Gattung Cordia in Venezuela. Mitt. Bot. Staatssamml. München 23:1-279.
JOHNSTON, I.M. 1935. Studies in the Boraginaceae X. The Boraginaceae of northeastern South America. J. Arnold
Arbor. 16:1—64.
JOHNSTON, I.M. 1935. Boraginaceae. In: A.A. Pulle, Flora of Suriname 4(1):306-333.
Miter, J.S. 2007. New Boraginales from tropical America 5: New names and typifications for neotropical species
of Cordia and Varronia. Novon 17:372-375
MiLLER J.S., J. Gaviria, R. GÓMEZ, AND G. RobríGUEZ. 1997. Boraginaceae. In: PE. Berry, B.K. Holst, and K. Yatskievych, eds.,
Flora of the Venezuelan Guayana 3:527-547.
Mier, J.S. AnD M. GorrscHuNG. 2007. Generic classification in the Cordiaceae (Boraginales): resurrection of the
genus Varronia P. Br. Taxon 56:163-169.
A NEW SPECIES OF THE FERN GENUS DORYOPTERIS (PTERIDACEAE)
FROM SOUTHEASTERN BRAZIL
Alexandre Salino
Universidade Federal de Minas Gerais
Departamento de Botánica - Instituto de Ciéncias Biológicas
Caixa Postal 486, 30123-970, Belo Horizonte, MG, BRASIL
salinogicb.ufmg.br
ABSTRACT
inhonl is Salino, known only from Minas Gerais, Brazil, is described and illustrated. TI
to Section Doryopteris due to the stipes with one vascular roni at ys bus ud Aedo venation. eee Jequitinhonhensis is is
trilobata J. Prado (Bahia, Brazil) and Di ply 3-lobed lamina
dit its du fronds, the lanceolate lobes with an acute apex, the ad much larger than the A lobes. Dorcpteris jequitinhonhensis
differs from both species by sterile leaves with margin sno not uu Mena crenate, and peels puberulous: is Rhone
d
1 f. 1 1
up to 0.1 mm long, Man A cells. From D. cordifoli y fronds g
buds; from the D. tril ttod 1 d in] OOE DEET
4 I >
Key Worps: Pteridaceae, Doryopteris, Minas Gerais, Brazil
RESUMO
n os e | 1 Cali 4 bla A 4 A Pay, $
yop Jeq i I Minas Gerais, Brasil é I I
Secáo D A 1 A 1 faz E T 2 3
mais proxima de D. "m Prado (Bahia - Brasil) e D. cordifolia eni Diels SUA Bs limas fértil trilebada: lobos lan-
lobo apical maior que os lobos basais. D 1 difere pela ee iid com dd
in
A tilagi 1 t d ] berulent tricomas até m 1 d i t
udi ate de D. ee pela auséncia de en na base da lamina estéril, e de D. trilobata pelo caule ereto ou decumbente, e
E
PALAVRAS CHAVE: Pteridaceae, Doryopteris, Minas Gerais, Brazil
INTRODUCTION
Doryopteris is a genus of about 30 species mostly of the southeastern Brazilian Highlands, with five endemic
to Madagascar and one pantropical (Mickel & Smith 2004). Besides an early revision by Tryon (1942) and the
surveys of Brade (1965), Tryon (1962) and Sehnem (1972), nothing more on the taxonomy of the Brazilian
species of the Doryopteris has been published, except a new endemic species from the Cadeia do Espinhaco
by Prado (1993), and a new species from the Atlantic forest (Yesilyurt 2007).
TAXONOMIC TREATMENT
Doryoptexs AT Salino, sp. nov. (Fig. 1). Ter: BRAZIL. Minas Gerais: Salto da Divisa, fazenda Santana,
enha (afluente do rio E MM 134 m alt., 16? 03'41.5"S, 40°03'23.5"W, 04 Mar 2004, A. Salino, J.A.
Lombardi R.C. Mota, T.E. Almeida & PL. V CB)
Pl Rhi t Id b Frondes di 1 1 f ilis 3.3—5.9 x 0.9—1.95 cm; frons
fertilis 9-10 x 0.75—0.82 cm; petiolus semi-cylindricus, sul in facie adaxiali; lami ilis i blonga vel obovata, ad basin
cordata vel sagitata, margine non pL A crenata, rare Integra) n fertilis iin vel elobi. Doryopteri cordifoliae (Baker)
milis in] 1 :
Diels et D. trilobatae J. Prado sim
tibus ad O
o O 16 E rF
o J o
" tal ] t A sal; ls
decumbenti, et p
Plants terrestrial. Rhizome erect to decumbent; scales linear to linear-lanceolate, with acute to acuminate
apex, 1-1.3 mm long, with sclerified dark brown band, margin hyaline. Fronds dimorphic, erect; sterile
frond 3.3-5.9 x 0.9-3 cm, fertile frond 9-10 x 1.5-3 cm; petiole semi-terete, adaxially sulcate, 17-28 x
J Rot. Res Inet Texas 200 243 — 846. 2008
844 J | of the Botanical R h Institute of Texas 2(2)
tem
tion, respectively.
, TES y
I L . J Ec
2
ile frond. C. Venati f the fertile frond. D. Rhizome scale. E. Sterile frond. F. Sterile frond. G. Porti
3
Salino, A ies of D teris f Brazil 845
0.38-0.58 mm in the sterile fronds, and 65—70 x 0.6—0.7 mm in fertile fronds, dark brown with one cylindri
cal vascular bundle at the base, basal scales (sometimes into the apical part) similar to those of the rhizome,
but with or without sclerified dark brown central band, puberulous, trichomes capitate, up to 3-celled, and
up to 0.1 mm long, light brown on adaxial surface, and with ciliform trichomes, 0.7-1.5 mm long on lateral
surfaces; lamina of the sterile frond entire, oblong or obovate with deeply cordate base or, sagitate, apex
obtuse, rounded or acute, margin hyaline, not sclerified, usually crenate, rarely entire; hydathodes present
on the upper surface; lamina of the fertile frond hastate, deeply 3-lobed, lobes entire, lanceolate, apex acute,
margin revolute, apical lobe larger than basal lobes, basal lobes forming a right angle to the apical lobe,
lamina of the both types of frond membranaceous to papyraceous, glabrous on both surfaces or glabrescent
on adaxial surface, trichomes ciliform, light brown, venation completely areolate on both types of fronds
without enclosed veinlets, and usually without free veins near the margin. Sorus marginal in vascular
comissure, continuous, including the sinus region, absent at the apex of the lobes; indusium formed by the
revolute margin; spores trilete, tetrahedral-globose, castaneous, sparsely cristate.
Etymology.—The specific epithet refers to the occurrence of the species only in the forest along the
Jequitinhonha River in Brazil.
Doryopteris jequitinhonhensis ncs: in section eo a due to the pe with one vascular bundle
at base and areolate venation. Doryopter in) is apparently closely related to D. trilobata J. Prado
(from Bahia, Brazil) and D. cordifolia dake Diels (from Madagascar) mainly in the deeply 3-lobed form
of the lamina of its fertile fronds, the lanceolate lobes with an acute apex, the apical much larger than the
basal lobes, and in the cordiform base of the sterile lamina. Doryopteris jequitinhonhensis differs from both
species by sterile leaves with margin hyaline, not sclerified, usually crenate, and petiole puberulous, with
capitates trichomes up to 0.1 mm long. From the D. cordifolia differs by fronds with margins not sclerified
and absence of proliferous buds; from the D. trilobata differs by rhizome erect to decumbent and petiole
sulcate adaxially.
In the general aspect (habit and shape of the sterile frond) D. jequitinhonhensis resembles D. rufa Brade,
but this species has fronds that are coriaceus with free venation, and belongs to the section Lytoneuron. An
identification key is presented to distinguish D. jequitinhonhensis from other similar Brazilian species.
Distribution.—Known only from the type collection from a population with about 30 individuals grow-
ing only along stream banks inside the deciduous forest in lowlands (100—170 m). Doryopteris trilobata and
D. rufa grow in open rocky habitats in the highlands (1200-1500 m).
KEY TO SPECIES DORYOPTERIS OF SOUTHEASTERN BRAZIL
SIMILAR TO DORYOPTERIS JEQUITINHONHENSIS
1. Venation of both steril | fertile fronds fi | DD filiform D. rufa
1. Venation of both sterile and fertile | i | te to acuminate.
2. Rhizome erect to decumbens ubi SeIn terere ip be puberuous with capitate trichomes
up to 0.1 mm; fronds thr margins not sclerified, y crenate jequitinhonhensis
2. Rhizome short ie terete without capitate tri ^ mes up to 0.1 mm; fronds coriaceous wit
margins sclerified and entire D. trilobata
ACKNOWLEDGMENTS
The author thanks Tarciso Filgueiras and Marcos E.G. Sobral for the Latin diagnosis, Conselho Nacional
de Desenvolvimento Científico e Tecnológico (CNPq) for a research fellowship grant (302594/2005-1),
and Miryan Morato Duarte for preparing the illustration. I thank Blanca Léon and an anonymous reviewer
for their critical comments on the manuscript. This work originated from the research project Inventário
Biológico nos Vales dos rios Jequitinhonha e Mucuri nos estados da Bahia e Minas Gerais, Brasil, and was
supported by the Ministério do Meio Ambiente (MMA-Brasil).
1 e£ the Datanizal D h Institute of Texas 2(2)
846
REFERENCES
BRADE, A.C. 1965. Contribuição para o conheci das espécies brasileiras do género Doryopteris (Polypodia-
ceae). Arch. Jar. Bot. Rio de Janeiro 18:9-72.
MickeL, J. AND A.R. SmitH. 2004. The pteridophytes of Mexico. Mem. New York Bot. Gard. 88:1—1054.
PRADO, J. 1993. A new species of Doryopteris (Section Doryopteris-Pteridaceae) from Brazil. Kew Bull. 48:13-16.
SEHNEM, A. 1972. Pteridáceas. In: R. Reitz, ed. Flora Ilustrada Catarinense. Herbário Barbosa Rodrigues. Itajaí, Santa
Catarina, Brasil. Pp. 1—244.
Tryon, R.M. 1942. A revision of the genus Doryopteris. Contr. Gray Herb. 143:1—79.
Tryon, R.M. 1962. The fern genus Doryopteris in Santa Catarina and Rio Grande do Sul, Brazil. Sellowia 14:51-59.
YesıLYuRT, J.C. 2007. Doryopteris majestosa (Pteridaceae), a new species from South America. Amer. Fern J.
2-219
NEW RECORDS OF POA (POACEAE) AND POA PFISTERI:
A NEW SPECIES ENDEMIC TO CHILE
Robert J. Soreng and Paul M. Peterson
Department of Bot
ids Museum of o History
mithsonian dr
TEM DC 20013-7012, U.S.A.
sorengr@si.edu, A
ABSTRACT
The distributi f ] ly endemic speci rd in Chile and Argentina are discussed. Poa hachadoensis var. pilosa and P.
mendocina are newly reported for Chile. Poa pfisteri is d I hile. A key to Nicoraepoa and Poa species
(except species of Poa sect. Dioicopoa) that grow in Chile from Coquiml ] ] is provided
RESUMEN
Se discuten las distribuci ingidas d i j démicas d w en Chile y iaa Tanto P. Rue var. pilosa
como Poa esa se citan nuevamente nes no Pos pfisteri se d una nueva esp de pid Se lie una
de Poa sect. Dioi ) n Chile ad
The genus Poa L. is highly developed in Argentina and Chile with 58 and 39 species, respectively, accepted by
Soreng et al. (2003, on-line version updated Jan 2008) in the Catalogue of New World Grasses (http://mobot.
mobot.org/W3T/Search/nwgc.html). Most of the Chilean species are shared with mE n two species
of P. sect. Dioicopoa E. Desv. (P. cumingii Trin. and P. paposana Phil.), and one new sp ribed here, are
thought to be endemic to Chile whereas 20 species are considered to be endemic to Argentina (Soreng etal.
2003, updated Jan 2008). Other than the Soreng et al. (2003) index, Poa has not been sy lly treated
in Chile since the early accounts by Desvaux (1854), Steudel (1854), and by R.A. Philippi who named 36 new
species between 1858 and 1891 (see Muñoz Pizarro 1960). Regional floras for Patagonia (Nicora 1978) and
Tierra del Fuego (Moore 1983) include species of Poa that occur in the southern third of Chile; these include
keys, descriptions, and illustrations. Negritto and Anton (2000) revised the genus for the five northwestern
provinces of Argentina, and Giussani (2000) revised Argentina's taxa of P. sect. Dioicopoa. These treatments
leave a major gap in our knowledge for the occurrence of Poa in Chile, especially for the northern two-thirds
of the country. Marticorena and Quezada (1985), with assistance of Oscar Matthei, pared down the number
of accepted species of Poa in Chile to 65 (no synonyms were listed). As Marticorena and Quezada (1985)
suggested, the nomenclature and taxonomy of many large genera in Chile was in chaos, and their listing of
taxa in these genera was only provisional. For example, of the 36 species of Poa named by R.A. Philippi only
three are accepted today (Soreng et al. 2003, updated on-line in Jan 2008; includes distribution by country,
synonymy, and classification with species placed in an infrageneric arrangement).
From November 2001 to February 2002 RJS (RJ. Soreng) visited Chile to collect and study the genus
Poa in the region. In CONC (Universidad de Concepción) and SGO (Museo Nacional de Historia Natural,
Santiago), the two main herbaria in Chile, herbarium specimens were found that appeared to represent seven
new taxa and another specimen was newly collected that potentially seemed to be new to science. RJS was
able to recollect all but one of them in Chile or on subsequent trips with PMP (P.M. Peterson) to Argentina
(2003 and 2006). This paper describes the fate of these eight unknowns, five of which were recently identi-
fied as occurring in Chile (Soreng et al. 2003), and one that is described here as a new species endemic to
Chile.
Three of these unknowns, Poa planifolia Kuntze, P. subenervis var. spegazziniana Nicora, and P. pugioni-
J. Bot. Res. Inst. Texas 2(2): 847 — 859. 2008
848 ] Vaf tha Ratanical D h institute of Texas 2(2)
folia Speg., were identified more readily than the others by comparison with type material. There are no
keys to identify the first two species. Although Muñoz Pizarro (1941) considered the first to be a Chilean
species from the type locality of Paso Cruz, Marticorena and Quezada (1985) did not mention it for Chile,
presumably as it had not been verified as occurring on their side of the border. The second two were re-
ported as occurring in Chile for the first time by Soreng et al. (2003), with some newer stations for them
reported by Soreng and Gillespie (2007). Poa planifolia and P. subenervis var. spegazziniana were found in the
high arid Andean valleys east of Santiago, and P. pugionifolia (often misspelled as "pungionifolia") was found
in the Region of Magallanes, Province of Ültima Esperanza in the Sierra Baguales, along with P. subenervis
var. subenervis that in Chile is also only known from this interesting mountain range. Soreng and Gillespie
(2007) transferred the later two species to the new genus Nicoraepoa Soreng & LJ. Gillespie, as N. pugionifolia
(Speg.) Soreng & LJ. Gillespie and N. subenervis (Hack.) Soreng & LJ. Gillespie [with subsp. subenervis and
subsp. spegazziniana (Nicora) Soreng & LJ. Gillespie], along with four other Patagonian species previously
separated as Poa subgen. Andinae Nicora; now N. andina (Trin.) Soreng & LJ. Gillespie, N. chonotica (Phil.)
Soreng & LJ. Gillespie, N. erinacea (Speg.) Soreng & LJ. Gillespie, and N. robusta (Steud.) Soreng & LJ.
Gillespie.
Although also i AN from several locations in Argentina, two other unidentified specimens were at-
tributable to Ni j vis subsp. subenervis and Poa hachadoensis Nicora var. hachadoensis, previously
reported from single collections in Chile (Nicora 1977, 1978). These taxa were accepted by Marticorena and
Quezada (1985) as occurring in Chile, but no vouchers had yet been identified for these taxa at CONC or
SGO
Three remaining unidentified specimens were still potentially considered to be new species of Poa, and
these, along with new vouchers for P. hachadoensis var. hachadoensis are discussed below.
RESULTS AND DISCUSSION
Poa mendocina Nicora & F.A. Roig.—One specimen from the high Andes east of Santiago (Villagrán et
al. 8484) certainly seemed new, until in 2003 a similar herbarium specimen from 3100 m in the Province
of Mendoza, Argentina, named “Poa mendocina ined." was discovered at BAA. This isotype had been filed
in the general collections rather than with the types at BAA, and thus was buried in this large genus. Only
later was it realized that Poa mendocina was already published and illustrated by Nicora & Roig (1998). This
name had made it into the Catalogue of New World Grasses as a tentatively accepted taxon, endemic to
Argentina (Zuloaga et al. 2008), as it was then known only from the type and a paratype that had not been
seen by Soreng et al. (2003). On another collecting trip to Argentina in 2006 we recollected this species in
the same region of Mendoza as the type.
Poa mendocina is in a distinctive species of Poa, now known from four localities in Argentina and Chile.
However, Poa ber. is not included in a key, descriptive floristic account, or revision of the genus in
Chile. P. ina has bisexual spikelets, short anthers (ca. 1 mm long), and lemmas with soft hairs along
the lower portion of the keel and marginal nerves. Most of the lower florets have a sparse but distinct web
of hairs on the dorsal side of the callus. In the treatment of Poa in northwest Argentina (Negritto & Anton
2000) this species keys out to P. laetevirens R.E. Fr. However, P. mendocina has relatively longer glumes that
are subequal to their adjacent lemmas, and longer spikelets. In the Flora Patagónica Poa treatment (Nicora
1978) P. mendocina keys to either Poa scaberula Hook.f. or P. hachadoensis Nicora. Poa mendocina differs from
the former species by having much larger spikelets that are few in number, and from P. hachadoensis by
having a shorter stature, and short compact panicles. Poa mendocina superficially resembles P. subspicata (].
Presl) Kunth, a more northern species that occurs in Colombia, Venezuela, and Ecuador to the Departments
of Ancash and Junín, Peru (and one record from Bolivia, Province of La Paz—Solomon 15876, MO—that
requires verification). P locina differs from P. subspicata by having shorter panicles and by lacking hairs
between the lemma keel and marginal veins. Poa mendocina is likely very closely related to Poa hachadoensis
var. pilosa. However, P. mendocina is only 2-15 cm tall, with contracted, compact, 1-4.5 cm long panicles
S | Peterson, N ds and ies of Poa for Chile 849
that have steeply ascending branches to 1 cm long (versus 18-50 cm tall, with wide open, [4-]4.5-12 cm
long panicles with spreading branches 1.5-4 cm long). Nicora and Roig (1998) were uncertain whether the
species is rhi or not, but further collections reveal it is cespitose, although its stems may be buried
and elongated when rooted in thatch of wet meadows (vegas).
Known specimens of Poa mendocina from Argentina and Chile: ARGENTINA. Mendoza: Department of San Rafael, district of El
Sosneado, cerro Volcán Overo, [ca. 34°36'21"S, 70*04'48"W], 3100 m, en la última vega alta, sobre el aqua entre cojines de musgos,
frecuente, 10 Feb 1955, A. dne Leal 16894 (HoLoryre: MERL; isorYrEs: BAA!, SI); Department of Las Heras, Cerro Pelado, [ca. 32°47'S,
69°09'15"W], 3000 m, formand mpactas, 19 Dec 1954, A. Ruiz Leal 16456 (paratype: MERL); Department of Malargue, Rio
Salado headwaters, lago on DN flank of Cerro ie ca. 6 km northwest of Las Leñas, 35%06'44"S, 70°07'39.6"W, 3075 m,
wet meadow and rocky slopes, 6 Mar 2006, P.M. Peterson, RJ. Soreng, D.L. Salariato & A.M. Panizza 19202 (SI!, US). CHILE. "pa
Metropolitana de Santiago: Cajón del Maipo, Hito Paso Internacional Maipo, [ca. 34°11'31"S, 69%49'55"W], 3325 m, 17 Feb 1995, C.
Villagrán, R. Villa & F. Hinojosa 8484 (SGO)).
Poa hachadoensis Nicora.—A second set of specimens that RJS thought might represent two new spe-
cies are attributable to two varieties of Poa hachadoensis. Nicora (1977, 1978) reported P. hachadoensis var.
hachadoensis, from Chile, Región of Bio-Bío, Las Lajas, along with three specimens in Argentina, Province
of Neuquén, Departments of Picunches and Aluminé, and illustrated it. In 2002 RJ. & N. Soreng collected
this variety in the regions of Araucanía and Bio-Bío, in Chile, and in 2006 we collected this variety in the
same Departments of Neuquén, Argentina, as cited by Nicora (1977, 1978).
Nicora (1977, 1978) reported P. hachadoensis var. pilosa from Argentina, Province of Neuquén, Depart-
ments of Minas, Lácar, and Province of Chubut, Departments of Río Senguerr and Futaleufü. Three old
collections by F.W. Pennell from Chile, Región Libertador General Bernardo O'Higgins, from the vicinity of
Sewell, belong to P. hachadoensis var. pilosa. These represent the only known locations for the species in Chile.
In 2006 Peterson et al. made the first collection of this variety in the province of Mendoza, Argentina.
Poa hachadoensis vars. pilosa and hachadoensis have curious distributions. The range of P. hachadoensis
var. pilosa extends for 1300 km from 34? to 46? S latitude along the Andes. The range of P. hachadoensis var.
hachadoensis covers 220 km north to south in the middle of the range of P. hachadoensis var. pilosa (which
has three allopatric stations to the north and four to the south of the typical variety), where the latter vari-
ety appears to be absent. The two varieties have not been collected together. Both varieties have a distinct
dorsal tuft of wooly hairs on the callus. Poa hachadoensis var. pilosa has hairs along lemma keel and marginal
veins, whereas the typical variety has glabrous lemmas. After viewing a series of collections and finding no
other characteristics to satisfactorily divide them, we are not inclined to separate them at the species rank.
However, because they are geographically isolated, they appear to warrant subspecific status. Both varieties
occur in mossy p habitats in meadows in Aurucaria forests to lower alpine.
J A 11 $ f : J al) 1 r Chile: ARCENTINA N DS l 2
Poa hachadoensis var
km E of Paso Pino Hachado, 38°39'41.3"S, 70°53'19.8"W, 1800 m, open, wet o nd PUN d and Festis 4 Feb 2003,
P.M. Peterson, RJ. Soreng & N.F. Refulio-Rodriguez 17466 (SI!, US); Aluminé, 1 y 11 towards Lago AS
39°05'2.7"S, 71?19'41.3"W, 1050 m, meadow with Juncus, Carex, and 1 Escall hA
antarctica, 3 Feb 2003, P.M. Peterson, R.J. Soreng & N.F. Refulio-Rodriguez 17409 (SU, US); ihe: collections from Argentina are cited | by
Nicora de 1978). CHILE. [without other location] in “02/03/1939”, A. Burkart, 9510 (SI); Region VIII: Bio-Bio, La Laja, [ca. 37?27'S,
, A. Burkart 27449 (SI, BAA); Parque Nacional Laguna Laja, southeast slope of Volcán Antuco, Estero el Aguado o del Volcán,
~ ien a lone aii east of Los nee ca. 90 km, 37°27'43"S, 71?19'05"W, 1430 m, broad, nearly treeless, volcanic
valley. l bbly muck , 21 Jan 2002, RJ. Soreng & N.L. Soreng 7177 (US!, CONC?; Region IX: Araucanía,
west entrance to Barges Nacional Congxillio, east end of Laguna Captrén, in E between TR poa and DU Nevada, east of
Temuco ca. 70 km, dace 714149" W, as m, wet prm me meadov d by old Araucaria
araucana-Nothofag : I 22 Jan 2002, RJ. onis e NE, Soreng 7192 (US!, CONC).
Poa hachad i pil llection in Argenti d all hers for Chile. ARGENTINA. Mendoza: Dep f Malarg
Andes, Rio Salado headwaters, Valle de Las Leñas, ca. 36 km northwest of Las Leñas on Highway 222, northwest of Malargue 65 km,
35°05'41.7"S, 70°08'10.6"W, 2740 m, dnd wet niani m Poa, Festuca, and clic 5 Mar 2006, P. M. Peterson, R.J. Soreng, D.L.
Salariato, A.M. Panizza 19188, (SI!, US!) Nicora (1977, 1978). CHILE. Region VI: Libertador
General B do O'Higgins, Sewell ca. 34°04'S, 70°22'W, in 1925, F.W. Pennell 12312 (SGON, 12314 (SGO)), 12324 (US).
oo
One last collection turned up at CONC, SGO, and US that appears to actually be a new species. By the time
" : In€tha Bataniral D h Institute of Texas 2(2)
RJS realized this was new, it was too late in the growing season to effectively search for it, as it flowers in early
November. A cursory search was made for it around the crossing of Rio Renaico by Highway 5, southeast of
Mininco. However, the grasses were in poor shape by the time we passed through the area in late January.
The species is considered to be dioecious. The SGO specimen appears to have been pulled up as a clump, the
staminate and pistillate panicles are not attached to a single base and may be from different individuals. The
US sheet appeared to have two separate clumps, one of each sex (they have been separated and remounted
on the holotype). In any case, the specimens are quite different from the other dioecious species of Poa in
South America, all 22 members of which diu. to Poa sect. Dioicopoa E. Desv. Unlike Dioicopoa species, the
Pfister specimens] moderately congested, somewhat lax, slender panicles with slender branches
that are naked in the lows: 1/2, and the pistillate and staminate spikelets are undifferentiated in size, form,
floret number, and pubescence. The lemmas are glabrous and the calluses have a well developed web, as in
Poa hachadoensis var. hachadoensis, but the anthers are 2-2.5 mm long. The blades are densely hispidulous-
strigulose adaxially on and between the veins, as in Poa sect. Madropoa Soreng subsect. Madropoa Soreng
and subsect. Epiles Soreng (seven and five spp. respectively, in North America, Soreng 2007). The new spe-
cies is tentatively placed in Poa subsect. Madropoa where it most closely resembles Poa diaboli Soreng & DJ.
Keil, a rare species of the California Floristic Province (Soreng & Keil 2004). If Poa pfisteri is confirmed to
belong to one of the above Madropoa subsections it would be the only occurrence of any species of either of
these anywhere south of the state of Durango, Mexico. The infrageneric relationships of some or all native
gynomonoecious species of South American Poa may be with “Poa nervosa" complex of Poa sect. Madropoa
(eight spp. of North America; Soreng 2007; Soreng & Keil 2003), but this needs further study. Curiously,
there is only one species of Poa sect. Dioicopoa in North America (Soreng 2007), Poa arachnifera Torr., nearly
all species of which have tightly contracted panicles with crowded spikelets, and none have hispidulous-
strigulose adaxial leaf surfaces.
The new species is here named for the collector, Don Augusto Pfister, who collected in Chile from 1941
to 1945 (according to Harvard University Herbaria, Index of Botanists, Index Herbariorum - Collectors,
http://asaweb.huh.harvard.edu:8080/databases/botanist, index.html [accessed May 2008]), but from 1932
to 1964 (according to CONC herbarium records database; Clodomiro Marticorena, pers. comm.). There
are 5080 sheets collected by Pfister in CONC, with duplicates in SGO, and US (at least for grasses). Pfister
collected plants in all regions of Chile, with the majority coming from around the Termas de Chillán, in the
Region of Bio-Bío, where he frequently bathed to alleviate his rheumatoid arthritis (Clodomiro Marticorena,
pers. comm.).
Poa pfisteri Soreng, sp. nov. (Figs. la—n; 2). Tex: CHILE. Recion XIII: Bio-Bío, Province a Santa ed. dad m 1
Nov 1943, A. Pfister s.n. (HOLOTYPE (pistillate and staminate plants): US-215030 CONC-
6191!, SGO-73895).
A Poa diaboli Soreng & DJ. Keil plantis dioeciis? (versus sequente gynomonoeciis), surculo sterili tantum extravaginali (versus extrav-
aginali et intravaginali), vaginis conatis 1/5-1/2 (versus 2/5-7/10), laminis caulis 0.8-1.4 mm latis (versus 1.5-2.4 mm), conduplicatis
g p I | 1 ramis paniculis
2-3 pro nodo (versus 1—2), tereter vel levi gulatis ( Ide angulosis) is parvis liff iatis su[ lapud angul
1 ( ifeste longioril latis supra disti gulos), flosculis pl que 3 vel interdum 4 [versus 3-6(-7)],
hills: lalibus] i} f levi ld bri linfi 1 ibus) glumis primis 2-3 m m lo ongis [versus
(2-)2.7-3.8 mm], 1-nervis (versus 3-nervis), calli pili seins magis apud donum concentrata (verus duco) lemmatis superficiebus
as vel ee sin ir dde apicem ieee raro vel dense, i MM parsim scabri ), palearum carinis sparsim
), differt
Plants perennial; dioecious (?); short rhizomatous and stoloniferous, loosely tufted, tillers extravaginal.
Culms 45-50 cm tall, 0.5-0.8 mm diam, geniculate above, slender, bases erect, frequently branching above
the base, terete; nodes terete, 1 exerted in distal 1/2-1/3 of culm. Leaf sheaths, uppermost closed 1/5-1/2
their length, weakly keeled, moderately to densely scabrous distally, sometimes lightly strigulose distally,
basal sheaths becoming gray-brown, fibrous, bases scabrous, glabrous, distal sheaths 2-3 times longer than
their blades; collars slightly thickened; ligules (0.2-)0.5-2.5 mm long, uppermost 1.5-2.8 mm long; lower
851
Jy A pip 4y ^ e
aa E
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Zi ZA
A Em DG
ZY qj
A. Liaule, adaxial view. D. Leaf blade
adaxial surface. E. Staminate spikelet. F. Pistillate spikelet. G. Lower glume. H. Upper glume. I. Pistillate floret, lateral view. J. Staminate floret, lateral
ew. N. Lodicules
eral vi
; lat
view. K. Lemma. L. Pistillate flower.
852
[| ||
#UMITIG VE
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TYPUS:
Poa
US. INATIUSAL HEREARE M
Poa nema 35
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INSTITUTO DË
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pfrsten) Senes
a
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BOTANICA
¡VE RSIBACZ
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Nombre cienihico : fa
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Obserszecnes: 12).
Familia Crsminesg. .
-
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Fic. 2. Poa pfisteri Soreng, holotype, A. Pfister 6191 (US-2150300). The upp
F
M ] Peterson, N Is and ies of Poa for Chile 853
ligules abaxially densely scabrous, apices obtuse or truncate; upper ligules abaxially smooth or sparsely
scabrous, apices acute, erose; sterile shoot blades to 25 cm long, involute with tightly enrolled margins,
abaxially smooth or very sparsely scabrous, adaxially densely hispidulous-strigulose on and between the
veins, ligules of sterile shoots like those of the distal shoots of the culm; cauline blades 4-8 cm long, 0.4-0.7
mm wide in side view, involute with tightly inrolled margins, thin, soft, abaxially veins prominent, smooth
except for scabrous keel and margins, adaxially densely strigulose on and between veins, narrowly prow-
tipped, mid-culm blades to 31 cm, uppermost 2.7-5.3 cm long, the longest on the culm. Panicles 4-11 cm
long, lax, broadly lanceoloid, loosely contracted, sparse to loosely congested, with 10-90 spikelets, lower
axis smooth, proximal internodes 1.5-2.8 cm long, with 2 or 3 branches per node; branches, the longest
2-3.5 cm long, loosely ascending, lax, capillary, terete to slightly angled, sparsely to moderately scabrous
proximally, moderately to densely scabrous distally, with 3-9 spikelets these mostly loosely overlapping in
the upper 1/2, pedicels commonly ca. 1 mm long, mostly shorter than their spikelets. Spikelets 5-6 mm
long, to 3 times longer than wide, laterally compressed, staminate and pistillate spikelets undifferentiated;
purplish at anthesis, florets (2-)3-4; rachilla internodes 0.5-1.5 mm long, visible from the side, smooth,
glabrous; glumes, distinctly keeled, slightly unequal, keels smooth or sparsely scabrous, sometimes with
a few hooks on the sides near the apex, margins with few hooks; lower glumes 2-3 mm long, 0.3-0.4 mm
wide in side view, subulate, 1-veined; upper glumes 2.9-3.5 mm long, 0.4-0.5 mm long in side view, nar-
rowly lanceolate, 3-veined; calluses with a slightly diffuse dorsally concentrated tuft of soft, wooly hairs
about 1/3-3/4 the lemma in length; lemmas 3.8-4.5 mm long, 0.6-0.7 mm wide in side view, narrowly
lanceolate, distinctly keeled, glabrous throughout, lightly scabrous along keel upper 1/2, smooth or lightly
scabrous near the margins and apex, intermediate veins faint to moderately prominent, margins narrowly
scarious, edges smooth or with a few hooks, glabrous, apices narrowly acute; paleas slightly shorter than
lemmas, sparsely to moderately scabrous over the keels, in the distal 1/2, between keels smooth, glabrous;
, anthers 2-2.5 mm long or vestigial
1
lodicules ca. 0.6 mm long, ovate, with a small lateral lobe; st
in pistillate flowers (0.2-0.4 mm long); ovaries glabrous with two styles and stigmas, vestigial in staminate
flowers. Caryopses 2 mm long (immature), fusoid, ventrally sulcate, glabrous, adhering to the palea. Chro-
mosome number unknown.
Distribution and Habitat.—CHILE. Region VIII: Bio-Bio, Province of Santa Bárbara. Known only from
the type collection in the central valley of Chile. The one collection is presumably from near the northern
Región IX Araucanía border, from slopes along the Río Renaico and tributaries east of Mininco, roughly
from 37°47'S, 72°48'W, where elevations range between 200 and 300 m. If the species is still extant, it likely
occurs in the adjacent Región of Araucanía.
ANNOTATED KEY TO SPECIES OF NICORAEPOA AND POA (EXCEPT SECT. DIOICOPOA SPECIES)
IN CHILE REGIONS COQUIMBO TO MAGALLANES
Accepted taxa also occur in Argentina unless stated to be *endemic to Chile." Introduced taxa are marked*.
See Poa in Catalogue of New World Grasses for synonyms (Soreng et al. 2003, and on-line). Authors are not
given for taxa in the key if the taxon was already discussed above.
1. Ligules 6-23 mm long, apex margins conspicuously lacerated, glabrous; plants without rhizomes, form-
ing robust tussocks, (0.5—)1—2.5 m tall; panicles tightly contracted; florets slightly contorted, twisted,
glabrous; lemmas with a stiff cusp or stout mucro up to 3 mm long; plants from islands (or possibly
mainland coast of Straights of Magellan), bluffs by the coast on the farthest southern reaches of Tierra
del Fuego, Magallanes (P. subgen. Ochlopoa sect. Parodiochloa) P. flabellata (Lam. Raspail
. Ligules mostly 0.5-6 mm long, but if elongated, then apex not conspicuously lacerated; plants rhizoma-
tous, or if non-rhizomatous, then mostly much less than 1 m tall; panicles tightly contracted to loose
and open; florets not at all contorted and/or twisted, glabrous or pubescent; lemmas rarely cuspidate or
mucronate, or awned, but if awned then the awn slender (but see also P darwiniana); plants widespread.
2. Leaf blades adaxially with multiple ridges and valleys on either side of the midrib, ribs broader than
the valleys; ligule margins ciliolate to ciliate; callus hairs (when present) forming a crown surrounding
—
J | of the Botanical R h Institute of Texas 2(2)
854
the base of the lemma, hairs straight or slightly ee plants with stout rhizomes (Nicoraepoa).
3. Plants forming low spiny mats, culms to a cm tall; leaf blades bluish-white, tightly fold-
ed, rigid, apex sharply pointed and pungen e. in moist, sub-saline ground interior
Patagonia (Sierra Baguales), NE Última Espera TAM N. pugionifolia
3. Plants not forming low spiny mats, culms usually over 20 cm Em pru green or bluish-white, flat
or loosely folded, somewhat rigid to lax, apex n pointed, but g; plants of wet coastal
sub-saline to interior fresh water meadow and riparian habitats.
4. Plants with viviparous spikelets.
5. Inflorescences narrowly lanceolate, erect, branches erect; florets glabrous; plants of
Oy elevations, generally in sub-saline Ed wetlands, e N. robusta
id, somewhat nodding ax, ascending; florets usually with at
“least a few hairs on the callus; elie a of SW sub- d and low interior regions in fresh
water wetlands, Bio-Bio t gallanes, rarely further sout N. chonotica
4. Bs without viviparous spikelets.
Florets glabrous; plants si se oo ee near d coast, Madalena N. robusta
: F with at Il d ]
7. Ligules ciliate, cilia as long or longer than the membranous base; leaf blades soft, flat,
short, mostly 2—6 cm long; plants from mossy vegas above treeline N. subenervis
8. Panicles 5-15 cm long, (0.5-)1-5 cm wide, contracted or ni open; plants y
Andean Valparaíso and Región Metropolitana subenervis subsp. spegazziniana
8. Panicles 3-6.5(-10) cm long, ca.0.5(-1) cm wide, contracted; Sans of Patagonia (Sierra
Baguales), NE Última Esperanza, Magallan subenervis subsp. subenervis
7. n ciliolate, the cilia much Dena ie the membranous per E ades 1 firm, flat or
d, elongated, g g Git subalpine
wet mea
‘Anat, | J + J
4 on ] AG.
OWS.
9. das contracted, 1
| Hol tl fthe awn, glabrous and scabrous or weakly
E the keel; gl lI ig or longer than adjacent i. plants from
upper elevations ftóm. Bio- Bío to Los Ríos, innequent southward to northern Aisén N. andina
9. Panicles loosely contracted to open, 2— 15c cm wide; lemmas obtuse to nie entire or
mucronate, but apex without lateral lobes, glabrous and scabrous, or also pubescent on
the keel and sides; glumes usually shorter than adjacent lemmas; plants from low to mid
elevations from Bio-Bio to northern Magallanes, rarely further south N. chonotica
2. Leaf blades adaxially with two grooves only, one on either side of the midrib, ribs not prominent and
broader than the intercostal regions; ligule margins smooth or scabrous or at most ciliolate; callus hairs
absent or present, usually originating in dorsal tuft from below the keel of the lemma (sometimes with
secondary tufts below the marginal veins), hairs plicate or woolly (web), less often forming a crown origi-
nating around the base of the lemma and then woolly or with hairs straight to slightly sinuous; plants
with slender rhizomes, or without rhizomes (Poa).
10. Plants with viviparous spikelets; plants of Tierra del Fuego and southern Patagonia mountains, Los
Lagos to Man
11. Plants with bulbous shoot I ; plants of disturbed ground, from northern Magallanes *P, bulbosa
subsp. vivipara (Koeler) Arcang.
. Plants without bulbous shoot bases; plants widespread (Poa subgen. Poa sect. Dioicopoa)
P. alopecurus (Gaudich. ex w Kunth subsp. Pueden. E f) D.M. Moore & Dogg.
Aisén to southern Magallanes; P. ob t d.—plants of rocky slopes
from upper forest openings to above treeline, from southern Araucanía to northern Aisén.
10. PI itl ivi (or ra ith tl but then of Maule and more northern regions);
rely YIU T LITE te,
plants widespread.
12. Plants with unisexual flowers only; anthers in staminate plants 1.3—3(-4) mm long.
13. Panicles loosely flowered, loosely contracted, somewhat lax; branches slender, plainly ex-
osed, naked in the lower 1/2; spikelets mostly 5-6 mm long; pubescence of staminate
and pistillate florets undifferentiated, lemmas glabrous and calluses pubescent; callus with
dorsally concentrated crinkled hairs; leaf blades adaxially hispidulous-strigulose pubescent
costally and intercostally; plants of presumably of steep rocky slopes in sclerophyllous
iere io openings, in the central vale) in Bio-Bío ds al nía
| ), contr ice erect y
P. pfisteri
=
UJ
ù Panicles densely fl
head J: p | |
the spikelets
T + J f T 4^ UT
yg us or very sparse-
| L
Maas pa or th | | call glal hrougl leaf blad
xially. | | | II lly; callus glal pubescent
P picat or called hairs on a tight dorsal tuft, and often from tufts below the marginal
veins, or | (Poa subgen. Poa sect. Dioicopoa).
Species of this section are re notoriously difficult to key out, adis staminate plants commonly
have glabrous or nearly glabrous lemmas and can only be keyed vegetatively (See Giussani
2000). It is not possible for us to Wie a satisfactory id for mes until me Chilean ses are
better known. H I nthe
region aie aes in this key, along with | distribution: P lor (
ex Mirb.) Kunth subsp. alopecurus— plants of mesic step! |f pening Magallanes; P.
bonariensis (Lam) Kunth
from central Chile, but most of this material seen by RIS v was s redetermined as P denudata, P
lanuginosa, or P paposana. Several CONC specimens from low to middle elevations in ilia
Chile, originally determined as P bonariensis, need further study. The species is common i
the pampas of Argentina and it is expected as a waif in Chile; P. cumingii—plants of Ban
dunes between Snipe and d E Orán Sey endemic To cd P. denudata
Steud.—plants of low | bluff m the coast of
Bio-Bío to Los Lagos extending to Argentina; P gayana F. Desv.—plants oft ze openings in
middle to upper forested to shrubby mountain slopes of the interior central Andes of Chile,
orumbo p Bio- ES mame poso Meis This species is highly variable and perhaps
repr ciformis in the alpine and a rhizomatous species
of lower elevations with a webbed callus, iid P lanuginosa; P. gayana x holciformis—
plants intermediate between these species are frequent and occur on rocky slopes from
high to medium altitudes in the Andes, Coquimbo to Maule; P. holciformis J. Presi—plants
of open treeless slopes in the high Andes, Coquimbo to Maule, expected on high dry slopes
further south also; P. lanuginosa Poir.— the typical form with webbed calluses occurs from
low elevations in the Central Valley to middle elevation forested slopes, Metropolitana to
Aisén, and a form with glabrous calluses occurs in riparian meadows in Patagonian steppe,
Magallanes; P. bra Steud.—plants of rocky open forest and alpine volcanic slopes in
the Andes, Bio-Bío to Los Lagos; P. paposana— plants of rocky slopes in fog zones of coastal
ls Japan o southern Ano POER to EAE i spiciformis (Steud) Hau uman &
.iberi
plants of dry P D
(Phil) Giussani—plants aay joa steppe, Magallanes.
12. Plants with some or all ter, 0.2-3 mm long.
14. Anthers 0.2—-1(—1.2) mm lon
15. Glumes exceeding or equaling the lower floret, similar in form and length; plants of ripa-
P.a
=
ian meadows in oe steppe, from Magallanes (Dissanthelium s.) ^ P. atropidiformis Hack.
16. Lemmas giabrou P. atropidiformis var. atropidiformis
16. Lemmas uie P. atropidiformis var. patagonica (Parodi) Nicora
(This Van incu only Hom AISeruna, is EXPO in Chile)
15. Glumes shorterthan the IVECO similar or | gth; p p
pubescent Il à t inal fl ithi il f ill ilv of
(Poa subgen. Ochlopoa sect. Micrantherae).
18. Anthers a 0.5 mm Tong: pran hes anang, spikelets e along the
| ] | y lid and
Metropolitana P. infirma Kunth
18. Anthers 0.5—1.0 mm long; branches ascending to spreading, neis ea ae
arranged aonig the branches; ae Ser WIBESpredes ofal Re Eglo . annua L.
and scabrous, al ti scent nt medially: all fl ts bi |
19. Por glabrous; rapis. and cal densely scabrous throughout, glabrous,
ort stiff scabrous mucro (Poa subgen. and sect. uncertain,
possibly eel pan plants of rocky coastal cliffs and slopes of the bs
pene a ca Fuego, ena (Cabo de mol ae) d i Parodi
| | | sed JS
4 ivi
1 J
or pubescent, witl MUL Iul», plar Iho widespread.
856
titute of Texas 2(2)
20. Inflorescences congested, contracted, spikelets crowded; branches quite
abrous; lemmas pubescent between the veins; callus with a dorasl
tuft of crinkled hairs, sometimes with secondary tufts below the mar-
ginal veins; plants of disturbed ground in upper forested to subalpine
shrub zones, from Mexico to Tierra del Fuego, Arica & Parinacota to Maga-
lanes in Chile (Poa subgen. Poa sect. Dasypoa)
oe open or contracted, spikelets not crowded; branches smooth;
mas glabrous between the veins; callus with a dorsal tuft of crinkled
rairs or glabrous, secondary tufts absent; plants of more stable habitats,
iy Metropolitana to Araucanía (Poa subgen. Poa sect. Homalopoa s.l).
ulms 4—20 cm tall; inflorescences ao ca. 0.5 mm wide, peda
spreading in anthesis to 1 cm wide; plants of high wet meadows
slopes in the central Andes, from Metropolitana
P. scaberula
N
o
C
Es mendocino
21. Culms 15-60 cm tall; inflorescences open, 2-5 cm wide; plants of wet
mossy meadows and slopes in forest openings to low alpine regions,
from Liberator General O'Higgins to Araucanía P. hachadoensis
22. Lemmas glabrous; plants of Bio-Bío to Araucanía A
var. "Bachaduens
22. Lemmas pubescent; plants of Liberator General O'Higgins. PR hos
var. pilosa
14. Anthers 1.3-3(-4) mm long.
23. Florets glabrous; blades flat, apexes A AR plants of wet meadows in the
central Andes, from Valparaíso to Metropolita alas E, Desv.
This taxon possibly originated from past eer hybridization between P. sub-
gen. Poa sect. Acutifoliae; P. subgen. Poa sect. Dioicopoa; P planifolia x P. 7 a
. Florets pubescent; blades flat, folded, or involute, apexes tapered or naviculate; plants in
various habitats, widespread.
24. Blades 5-10 mm wide, prominently keeled, flat to tightly folded, apex abruptly
attenuated, acuminate, and pungent; florets pubescent; lemmas thin and
lustrous, usually sparsely short pubescent along the keel; callus with a dorsal
tuft des lly hairs (webbed); plants of wet subsaline to freshwater meadows,
springs, and stream-sides above tree line in the central Andes, from Valparaiso
to Meal A (Poa subgen. Poa sect. Acutifoliae)
Blades 1—5(—7) mm wide, not pre ntly keeled, flat to folded, apex obtuse to acute,
not pungent; florets glabrous or or dd lemmas dull (not reflective) or at least
not especially thin and lustrous, glabrous or pubescent; callus glabrous or pubescent;
plants bid, habitats and distributions.
25. Lemmas pubescent only on the keel, or also sparsely so on the lower 1/4 of the
marginal veins i li a POOR ER IAE -nerved, often sickle-
No
UJ
P. planifolia
N
>
of wooly hairs
(webbed); plants of disturbed ground in forested zones, known from Valparaíso
south to Magallanes (Poa [subgen. Ochlopoa x Stenopoa] sect. Pandemos)
*P. trivialis L. subsp. trivialis
. Lemmas pubescent on the keel and marginal veins and sometimes on inter-
mediate veins and sometimes between the veins, intermediate veins faint to
pronounced; first glumes (1)3-nerved, rarely sickle-shaped; ligules truncate to
acuminate; callus glabrous or iene with a web or crown of hairs; plants
widespread and of various distribution
26. Sheaths compressed-keeled; Ses open; plante e to mesic dis
thickets, and open forests, from Tierra del Fueg but
likely in Aisén also (Poa subgen. Poa sect. Poa) P. yaganica Speg.
ius aida fom Rio BE is known DA from the bin The oe
Ro
Un
3
species similarto P pratensis, with a mes 2 with more compressed pss
more rounded to acute li p boda gumer
> Sheat hs OL noticeably im vM UE |
ted or ope
27. Pars rhizomatous, spreading; upper culm sheaths ee more than
1/4—2/5 the length; lemmas glabrous between the veins; callus with a
Mm
ON
NA
=~
f Poa for Chile 857
well developed dorsal web of crinkled hairs (Poa subgen. Poa sect.
oa) P. pratensis L.
Poa pratensis subsp. alpigena (Lindm.) Hiitonen has lemmas with
glabrous or sparingly pubescent intermediate veins, narrowly ovate
spikelets, narrowly pyramidal inflorescences with mostly smooth
branches, and only extravaginal shoots. It is apparently indigenous in
the Magallanes Region as a disjunct taxon from northern hemisphere
boreal regions. Introduced *P pratensis s.l. is known from disturbed
ground from Coquimbo to Magallanes, but its races have not been
sorted out taxonomically in Chile. The lemmas are glabrous between
the keel and marginal veins, spikelets more ovate, inflorescences pyra-
n. with smooth or scabrous branches, and some shoots usually in-
vaginal.
a ron mne strictly tufted; upper culm iiu. closed
1/10-1,
callus glabrous or r with a crown of hairs or r with a pon s PIC
hairs.
28. Callus with a crown of hairs 0.2—2.0 mm long around the base of
the lemma (Poa subgen. Stenopoa sect. Secundae).
29. Lemmas weakly keeled, pubescent on and between the veins
at least at the base, pubescence of the keel and between veins
little differentiated; panicles secund, contracted to loosely
contracted, much longer than wide (except when somewhat
open in anthesis); plants of subalpine to lower alpine open
slopes, from Valparaíso to Magallanes P. secunda
Presl. subsp. secunda
29. Lemmas distinctly keeled, sometimes glabrous E the
veins, but if pubescent between the veins present, then hairs
alta) longer on the keel a between the ves, panicles
plants of subalpine open slopes, from Bio-Bio to Magallan
P. stenantha D. var. stenantha
28. Callus glabrous or with a dorsal web at the base of the keel of the
lemma.
30. Upper culm node in the lower 1/3 of the culm; upper culm leaf
least a fe ew tl eins near the
as oron the intermediate veins; callus guo eu or webbed.
. Spikelets (3.8—)4—5 times as long as wide (when closed);
lemmas weakly Ten poA on the keel and
marginal veins and between the veins at least near the
base, the hairs about the same length between the
veins and those on the keel and marginal veins; callus
glabrous; plants of subalpine to lower alpine open slopes,
from Valparaíso to Magallanes (Poa A Stenopoa
sect. Secundae) ecunda subsp. secunda
. Spikelets 2-3 times as long as wide ias closed);
lemma distinctly keeled, pubescent between the veins
at least near the base or not, sometimes pubescent
on the keel and marginal veins only, or on these and
the lateral veins only, the hairs shorter between the
veins than those on the keel and marginal veins; callus
glabrous or webbed; plants of subalpine open p es,
from P to Magallanes (Poa subgen. Sten
sec oa) P. glauca v hee glauca
30. Upper culm aust in the middle to upper 1/3 of the culm; upper
culm leaf blades subequal or longer than the upper sheath;
lemmas glabrous between the keel and marginai veins; cal-
w
858 J | of the Botanical R h Institute of Texas 2(2)
Steno 7
32. Ligul | lydeveloped
ey les than 1/2 the Ena la length uba stretched
a forest
margins, known from Magallanes *P, nemoralis |.
32. Ligules 1—4 mm long, apex obtuse to acute; web moder-
ately well developed, usually more than 1/2 the lemma in
bumps or sparsely hispidulous; plants usually of riparian
meadows, known from Magallanes *P, palustris L.
ACKNOWLEDGMENTS
Our thanks are extended to the faculty and staff of the Departamento de Botánica, Universidad de Concep-
ción, especially Oscar Matthei and Clodomiro Marticorena for assistance in logistics, and whose collections
database and knowledge of the geography and flora were invaluable; the Smithsonian Institution's Atherton
Seidell and Restricted Endowment Funds and the Department of Botany, National Museum of Natural History
for providing funding to collect in Chile and Argentina; Alice Tangerini for the illustration, Alan Touwaide
for help with the Latin diagnosis; curators of BAA, CONC, SGO [especially Mélica Muñoz-Schick], SI, and
US, for their assistance with collections and loans; Nancy Soreng, the first authors traveling companion
and co-collector, for four months in Chile, for her diligent herbarium assistance, and for her patience and
spirit of adventure.
REFERENCES
Desvaux, E. 1854. Fam. CXLVI. Gramineas. In: C. Gay, Flora Chilena. Fain & Thunot, Paris. Pp. 233-469.
Giussani, L.M. 2000. Phenetic similarity patterns of dioecious species of Poa from Argentina and neighbouring
countries. Ann. Missouri Bot. Gard. 87:203—233.
MARTICORENA, C. AND M. QUEZADA. 1985. Catálogo de la flora vascular de Chile. Gayana, Bot. 42:1—157.
Moore, D.M. 1983. Flora of Tierra del Fuego. Missouri Botanical Garden, St. Louis.
Munoz Pizarro, C. 1941. Indice bibliografico de | hilenas. Ministerio de Agricultura Dept. de Genetica
Fitotecnica. Santiago, Chile.
Muñoz Pizarro, C. 1960. Las especies de plantas descritas por R.A. Philippi en el siglo XIX. Ediciones de la Univi-
versidad de Chile, Santiago.
Necritto, M.A. AND A.M. Anton. 2000. Revision de las especies de Poa (Poaceae) del noroeste Argentino. Kurtziana
28:95—136.
Nicora, E.G. AND FA. Rois. 1998. Novedades en Poa (Gramineae) para la provincia de Mendoza (Argentina). Hi-
Ckenia 2:273-275.
Nicora, E.G. 1977. Gramineas Argentinas nuevas. Hickenia 1:99—107.
Nicora, E.G. 1978. Gramineae. Flora Patagónica 8:1—583. Colect. Cient. INTA, Buenos Aires.
SonENG, R.J. 2007. Poa L. In: M.E. Barkworth, K.M. Capels, S.L. Long, LK. Anderton, and M.B. Piep, eds., Magnolio-
phyta: Commelinideae (in part); Poaceae, part 1, Flora of North America.north of Mexico, volume 24. Oxford
University Press, New York. Pp. 486—601.
SorenG, RJ, L.M. Giussani, AND M. Necnrrro. 2003. Poa L. In: RJ. Soreng, PM. Peterson, G. Davidse, E.J. Judziewicz,
FO. Zuloaga, TS. Filgueiras, and O. Morrone, eds. Catalogue of New World grasses (Poaceae): IV. Subfamily
Pooideae. Contr. U.S. Natl. Herb. 48:505—580.
Soreng, RJ. AND LJ. GiLesPie, 2007. Nicoraepoa (Poaceae, Poeae), a new South American genus based on Poa
subgenus Andinae, and emendation of Poa section Parodiochloa of the sub-Antarctic islands. Ann. Missouri
Bot. Gard. 94:821—849.
SORENG, R.J. AnD DJ. Ke. 2003. Sequentially adjusted sex-ratios in gy im, and Poa diaboli (Poaceae), a
new species from California. Madroño 50:300—306.
gand ped Chile 859
STEUDEL, E.G.V. 1854. Synopsis plantarum glumacearum, vol. 1. Plantarum graminearum. J.B. Metzler, Stuttgart.
ZULOAGA, F.O., O. Morrone, AND M.J. BELGRANO. 2008. Catálogo de las plantas vasculares del Cono Sur de Sudamérica.
Monogr. Syst. Bot. Missouri Bot. Gard. In press.
860 | laf tha Ratanical R h Institute of Texas 2(2)
BOOK REVIEW
BILL AND JAN MOELLER. 2000 (2*1 printing). Lewis & Clark: A Photographic Journey. (ISBN 0-87842-405-9,
pbk.). Mountain Press Publishing Company, PO. Box 2399, Missoula, Montana 59806, U.S.A. (Orders:
www.mountain-press.com, info@mtnpress.com, 1-800-234-5308, 406-728-1900). $18.00, 107 pp., 77
color photographs, 1 map of Lewis and Clark route, 8 3/8” x 9”.
By the nineteenth y the newly f ] blic of the United States of America was eager to find a commercial a north
west passage to th Mann ipn n The impediment was the vast ies between the Rd River s the Rocky M e
1 |e2n AAA
new republic Fortunately P a b dd States, Periods was in the mood to sell having cold the French treasury for his military
operations. vas paid to the French, and the MR iud was now in das aw of the United s
more than doubling its size. In 1803 cu Thomas Jefferson would give |
of Discovery, but history would remember it as the Lewis & Clark expedition. With $2, 500 alcatel by Congress, Jeff t out th
directives of the expedition:
CH eset er m ission is to explore the Mi i River, and such principal st of it, as, by its course and com-
{+l D spe ri Lf. +1 +A + J H 11 H P"
waters oj ,may ojjer 1
the continent, for the purpose of commerce.”
His bad bidder 29-year-old Meriwether Lewis would be in command of tl diti d take ch ft ]
d fai determine soil i In addition, he would document e various Indian tribes
along on their habits, clothing, dwellings, Hai tae ie on religions 33-year-old William Clark would be his ander
Hanne did on pe western add r, Clark g E g and geography. He VEM ee Isdes/onguudes
o cu 1 is
commercial intercourse. Alone the way they would meet a French Canadi DE d his Shoshoni wife, b who would plas
a vital role in Indian relations with the various tribes.
wo years, four months, and 10 day p d P {8,000 miles. The original cost allotted by congress: $2,500.
The final cost was dol 722.25. an over budget ( hing ) and finding igable wat the Pacifi
as hoped, th dit ] h ion of ihe à i 1 many land Waterways
of the northwest estern. The ine i Poma a wealth of material o on the e inns iu E the a Lewis and Clark xe
P4
He and good fortune).
it like pili nud Clark poc. d Bill Dane J ond AE taken on Dholos apne talents, love of travel and the
ctor, long I iographies and journeys. Lewis &
h Lh:-T : qc 6 pe L 1 Le TET à efl 1 1 SY h
pnic }
in front mputer. It takes ! giu d RV and BRUN ihe paths that UE ane we made over 200 iai ago. E
this time, t] k is finding and tl g gg g I g g
of the expedition. The photograpl ing showing tl } pediti led it with tive d d wonderful
1 1 1 1 1 1 ] f, 1 : 1 £1 h Clark and Lewis T 1 fon] eh 1, is 1i 1 :
L c O pl Cc ja] 4
and seeing what Lewis and Clarl I for the first time:
May 13, 1805, Fripay, (The Missouri Breaks—in what would be Montana):
“The hills and river cliffs which we passed today exhibit a most romantic appearance, The bluffs of the river rise to
the oli of ta 200 to 300 feet and in most places nearly perpendicular, they are formed of remarkable white
sandstone... "—Lew
Sept. 14-15, 1805, through the Bitterroot Mountains, tl in they had experi d, elevation 7,000 ft
and higher, eventually so exhausted and | ud y they tingt f their h
“11 I t and ld i part was in my life...."—Clark
o only iege is n the reference map is far too eu to eae de n s text - beautiful E It would have
educational ] and events related to
X r O r [24
O Hn eaey, Linny/Designer, Illustrator, il: a0005835 Dai il
J. Bot. Res. Inst. Texas 2(2): 860. 2008
LEPTOCHLOA (POACEAE: CHLORIDOIDEAE) IN COLOMBIA
Neil Snow Paul M. Peterson
Herbarium Pacificum Department of Bot
ishop Museum National Museum of es History
1525 Bernice Street mithsonian dnd
Honolulu, HI 96817, U.S.A Tei DC 20013-7012, U.S.A.
neil. snow@bishopmuseum.org eu.
Diego Giraldo-Canas
Instituto de Ciencias um
Itad de Cienc
daa Nacional de a
Apartado 7495, Bogotá D.C, COLOMBIA
dagiraldoc@unal.edu.co
ABSTRACT
A taxonomic summary is presented for the indes species of Leptochloa. The classification, nomenclature, ied affinities, and
M KS s variation of the genu di d. Leptochloa is cca in Colombia by six species and one nontypical subspe-
cies C ndn peci , as well d peci HUS E distributions, and
Ei]
wr He ecological cs The basionym Chloris dubia 1
RESUMEN
S t di ^mi de | j l bi del género Leptochloa Se analizan di lati la clas-
ificación ] latura, las afinidad éri iació fológica de los caracteres. El género Tenia está representado
yl
en Saas RR seis ESSE y una subespecie. " a e claves pare reconocer las ores nds en Combia así como
la di
la iconografía, geog y g g y ecológicas. Se
lectotipilica el nombre Chloris dubia Kunth
Leptochloa P. Beauv., sensu lato (including Diplachne P. Beauv.) is a genus of approximately 32 annual or
perennial species (Snow 1997). The native range of the genus is pantropical into warmer temperate regions,
with several species being weedy and widely distributed. Although Parodi (1927), McNeill (1979), Phillips
(1982), Nicora and Rúgolo de Agrasar (1987), and Nicora (1995) recognized Diplachne as a separate genus,
we agree with McVaugh (1983), Clayton and Renvoize (1986), Nowack (1994), Renvoize (1998), and Snow
(1997, 19982) that the number of florets, the size of the spikelets, shape of the caryopsis, arrangement of
spikelets on the panicle branches (i.e., secund or not secund), and other morphological characters inter-
grade strongly and are unable to support generic separation (Snow 1997). Moreover, based on a numerical
analysis of morphological characters, Phillips (1982) found that species placed in Diplachne and Leptochloa
overlapped one another in a principal coordinates scatter plot. Although it is likely (next paragraph) that
some elements of Leptochloa may need to be segregated in the future with further analyses, in our opinion it
is unlikely that such a generic segregate would correspond closely to Diplachne as recognized by the authors
cited above. Thus at this time it seems more appropriate to recognize Leptochloa in its broad sense (Peterson
et al. 1997; Snow 1997).
The monophyly of Leptochloa sensu lato was not supported when tested cladistically with morphological
and anatomical data, depending on the number of outgroups included (Snow 1997). Moreover, its monophyly
has not been tested in breadth using molecular data, and the relationship of Leptochloa with other genera is
far from certain (Columbus et al. 2007; Peterson et al. 1997, 2007). For example, in a restriction fragment
analysis of cpDNA using a few representatives of New World genera (Duvall et al. 1994), Leptochloa [L. dubia
(Kunth) Nees] was a member of a basal clade, sharing a common ancestor with Scleropogon Phil., Dasyochloa
J, Bot. Res. Inst. Texas 2(2): 861 — 874. 2008
Lat +k D + al D h eee
862 j f Texas 2(2)
Willd. ex Rydb., Munroa Torr., and Erioneuron Nash. In a strict tree with bootstrap analysis, species
of Leptochloa did not form a particularly strong clade with the other genera, and instead shared a common
ancestor in a clade that included the Muhlenbergiinae, Munroinae, Scleropogon, Sporobolus R. Br., Eleusine
Gaertn., Tridens Roem. & Schult., Tripogon Roem. & Schult., and Eustachys Desv. (Duvall et al. 1994). In a
phylogenetic study of Chloridoideae based on matK sequences, Leptochloa (L. dubia) formed a clade with
Coelachyrum Hochst. & Nees, and Astrebla F. Muell. emerged as sister to these (Hilu & Alice 2001). Other
genera in the C, clade of Hilu and Alice (2001) included Brachyachne (Benth.,) Stapf, Chloris Sw., Cynodon
Rich., Dinebra Jacq., Eleusine, Enteropogon Nees, Eustachys Desv., Lepturus R. Br., Lintonia Stapf, Microchloa R.
Br., Oxychloris Lazarides, Tetrapogon Desf., and Trichloris E. Fourn. & Benth. In another phylogenetic study
of Chloridoideae based on analysis of combined trnL-F and ITS sequences, the placement of species of Lep-
tochloa [L. dubia, L. fusca (L.) Kunth, and L. panicea (Retz.) Ohwi] in separate clades suggests a polyphyletic
origin (Columbus et al. 2007). In a phylogenetic analysis of rps16 seq , Leptochloa dubia formed a clade
with Eleusine [F. coracana (L.) Gaertn.], whereas in a tree based on analysis of waxy sequences, Leptochloa
formed a clade with Eleusine coracana and Dactyloctenium aegyptium (L.) Willd. and D. radulans (R. Br.) P.
Beauv. (Ingram & Doyle 2007). In summary, the monophyly of Leptochloa has not been corroborated with
molecular data, and phylogenetic relationships of Leptochloa with other genera remain obscure (Columbus
et al. 2007, Peterson et al. 2007). For these reasons Peterson et al. (2007) treated the genus as incertae sedis
within the tribe Cynodonteae.
Leptochloa is represented by 17 species in the New World (13 of these native), six of which occur in
Colombia (Peñaloza-Jiménez et al. 2002; Peterson et al. ur. 20 de Our taxonomic treatment contains keys
ied d mies. Only
, illustrations,
synonyms used frequently in Colombian literature are given. For I id 2 see now 1997, Peterson et
al. (2001), and Soreng et al. (2008). This study is based on th COAH
CAUP, COL, HUA, MO, and US, including the type specimens of most species studied. The present study aims
to contribute to the knowledge of the Colombian grasses by collaborating with the Biodiversity Program and
Colombian Flora Inventories.
for determining the species, descriptions, distributions,
Leptochloa P Beauv., Ess. Agrostogr. 71. 1812. Ter: Leptochloa virgata (L.) P. Beauv.
Plants annual or perennial; caespitose, lacking stolons and rhizomes but sometimes geniculate and root-
ing at the lower nodes in mesic habitats. Flowering culms 30-150 cm tall, ascending to erect, glabrous to
sometimes pilose, arising from fibrous roots. Leaves cauline; sheaths longer or shorter than internodes with
smooth margins; ligules a membrane or a ciliate membrane; blades narrowly lanceolate to linear, flat, often
inrolled on drying, not pungent. Inflorescence a panicle of spicate primary branches exserted (or lower
branches occasionally inserted proximally) along an elongated main axis; main axis scabrous; primary
branches spaced or sometimes subdigitate, ascending to spreading from main axis, terminating in a spike-
let; glabrous or scabrous. Cleistogamous spikelets absent or present in axils of lower sheaths (in L. dubia or
rarely in L. fusca subsp. fascicularis). Disarticulation above glumes, lemma and palea falling as a unit; callus
glabrous to sparsely sericeous. Spikelets solitary, laterally compressed or terete, subsessile to shortly («2
mm) pedicellate, distant but often imbricate, often secund; glumes shorter than spikelets, unequal, smooth,
glabrous or slightly scabrous; lower glumes 1-veined; upper glumes shorter than lower lemma, 1-3 -veined,
unawned; rachilla pronounced between florets, glabrous; florets 2-13 per spikelet, sterile florets often pres-
ent above fertile florets (and uppermost floret often reduced to small, awnless rudiment); lemma 3-veined
(rarely 4—5-veined in L. dubia at base), membranous, smooth, glabrous or hairy along veins and sometimes
between veins, apex entire, emarginate or lobed, awnless, mucronate, or awned; paleas glabrous to sericeous,
unawned, membranous, margins not enfolding fruit, smooth; lodicules truncate; stamens 2 or 2 nare
yellow or reddish-purple; stigmas 2. Fruit with adnate pericarp (sometimes weakly so); caryopses dorsiven
trally compressed or laterally compressed (Snow 19982). Base chromosome number of x - 10.
Comments.—Leptochloa and Diplachne were established by Palisot de Beauvois (1812). Leptochloa was
based on three species, while Diplachne was based on D. fascicularis (Lam.) P. Beauv. [= L. fusca (L.) Kunth
Snow et al., Leptochloa in Colombia 863
subsp. fascicularis (Lam.) N. Snow]. Leptochloa was later lectotypified with L. virgata (L.) P. Beauv. by Nash
(1915), which was ineffective according to Index Nominum Genericorum Art. 10.5 example 6. In addition,
both genera were placed in the same *cohors," which suggested that Palisot de Beauvois regarded them as
more or less closely related. In his "Tabula Methodica" he mentioned that the lemmas of Diplachne were shortly
awned, whereas those of Leptochloa were said to be awnless. However, this feature varies considerably and
the lemma of Leptochloa can be entire, emarginate, lobed, awned, mucronate, or unawned. The micromor-
phological characters of the lemma vary little in Leptochloa (Snow 1996), and with few exceptions, cork cells
are present, silica cells are absent, chloridoid bicellular microhairs are present, prickles are present, and
macrohairs are present. However, the occurrence of papillae on long cells and short cells is more variable,
suggesting their presence or absence may be ibm A informative within the genus (Snow 1996).
Ecology and geographic distribution —T1 g 1 ly
to tropical regions. Its ecological distribution mostly includes semi-open vegetation such as grasslands. The
worldwide in warm- temperate
species most frequently grow in periodically disturbed or seasonally saturated areas such as riparian cor-
ridors, ephemeral pools, or artificial impoundments, although L. dubia generally prefers well drained soils.
All widespread annual species are considered weedy in agricultural situations (Snow 1997) and some have
become established on continents far beyond their native range (Snow & Simon 1999; Snow 2004).
KEY TO THE SPECIES OF LEPTOCHLOA IN COLOMBIA
1. Sheaths sparsely to moderately pilose, the hairs papillose-based L. panicea subsp. brachiata
1. Sheaths glabrous to sparsely hairy, the hairs usually not papillose-based.
2. Ligules membranous and neither ciliate nor erose (but sometimes lacerate), 5-8 mm long, apically
attenuate L. fusca
2. Ligules membranous and sometimes ciliate or erose apically, less than 5 mm long, apically obtuse or
truncate
3. Plants perennial; ligules 0.2-1.5 mm long.
4. Apex of lemma broad and usually notched; hidden A present in lower leaf sheaths;
florets widely divergent (40-90°) from a during anthes L. dubia
4. Apex of lemma acute, not notched but sometimes awned; hidden inflorescences absent in lower
leaf sheaths; florets on slightly diverging rum rachilla at anthesis (if at all) L. virgata
3. Plants annual; ligules 2.2-3.8 long.
Culms more or less round in cross section; panicle branches mostly somewhat stiff: lemmas 2.4—3
mm long; o 0.6—0.8 mn long 4. L. panicoides
Cul 19]
| H Jy +. “dad |
IJVVCI
to prominently arcuate: lemmas 2.1— 2. 4 mm long; anthers 0.2—0.4 mm long 5. L. scabra
1. Leptochloa dubia (Kunth) Nees, Syll. Pl. Nov. 1:4. 1824. (Fig. 1, A-C). Chloris dubia Kunth, Nov. = (quarto
ed.) 1:169. 1816. nud pud aid G. a Prim. Fl. Esseq. 74. 1818. Diplachne dubia (Kunth) Scribn., rrey Bot.
Club. 10:30. 1 Rabd k., Anales Mus. Nac. Buenos Aires 11:121. 1904. nm pi (Kunth)
Kuntze ex Stuck., Anales Mus. Nac. Buenos Aires 11:128. 1904. Tree: MÉXICO: E Humboldt & A. Bonpland 4172 (LecrorvrE, here
designated: P!; DUPLICATES OF LECTOTYPE: K, K-microfiche, US-865876 fragm. ex P!).
Perennials, mostly erect or infrequently decumbent or sprawling. Culms (5-)30-110 cm tall, 1.0-4.5 mm
wide, unbranched or only as tillers from very base, round or flattened below; nodes glabrous, internodes
3-11 cm long, herbaceous, solid (or occasionally hollow by virtue of aging). Leaf sheaths longer or shorter
than internodes, sparsely pilose, especially below, and occasionally pilose (sometimes densely so) near the
collar, the hairs occasionally from papillose bases, the margins glabrous or somewhat pilose; collars green
or tan; ligules mostly 1.0-1.5 mm long, membranous, truncate, ciliate apically; blades (2228-35 cm long,
2-8 mm wide, mostly linear or somewhat lanceolate, scabrous above at base or with a sparse covering of
pilose hairs, glabrous to minutely scabrous below, flat but drying involute, midvein mostly prominent or
sometimes not. Panicles of two types, the apical ones generally exserted at maturity and the lateral ones
completely hidden in lower leaf sheaths; apical panicles 10-45 cm long, (2-)3-25 cm wide; the branches
(2-)5-15, (1.5-)3-19 cm long, alternate or infrequently subdigitate, ascending to reflexed, often somewhat
lax, minutely scabrous, the branch axils pilose or merely scabrous. Spikelets 4.0-12.0 mm long, imbricate
2(2)
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Snow et al., Leptochloa in Colombia 865
to distant, mostly sessile; florets 4-13; callus glabrous or with a few short hairs; lower glumes o Ra A = 8
mm long, membranous, narrowly triangular or ovate, scab long midvein, ;
acute; upper glumes 3.3-6.0 mm long, membranous, ovate to ole scabrous on em (and some-
times with 1 or 2 additional veins at least basally) and papillate on edges, acute; lemmas 3.5-5.0 mm long,
membranous, ovate to obovate or widely obovate, light brown, bronze, light to very dark green, the lateral
veins usually prominent but sometimes not so, sericeous at least along lower portions of lateral veins and
sometimes on midvein and between veins, the apex usually bifid, broadly acute, obtuse, or truncate, awn-
less or mucronate; paleas subequal to lemma, lanceolate, membranous above but somewhat cartilaginous
near base, distinctly ciliate on edges, sometimes sericeous between veins, apex acute; stamens 3; anthers
(0.3-)1.0-1.6 mm long, yellow. Caryopses 1.5-2.3 mm long, 0.9-1.0 mm wide, narrowly elliptic, elliptic or
very widely obovate in hilar profile, depressed obovate in transverse section, with a broad, shallow groove
on hilar side, smooth, brown; pericarp weakly adnate to the endosperm.
Distribution and habitat. —Native in the USA from Arizona to Oklahoma and Texas, southern Florida,
common in México, sporadically in the Caribbean and in M ica through Bolivia, more common again
from Bolivia and Chile through Paraguay, Uruguay, and southern Brazil and much of Argentina; in a variety
of soil and vegetation types, but most frequently on well-drained slopes, 100-2600 m, but occasionally to
3150 m in Peruvian and Bolivian Andes.
d.C MBIA. Nariño: Mun. Imúes, corregimiento El Pedregal, Pilcuán, 2000 m, 11 Jan 1989, B. Ramirez 1259
(COL, PSO). Carretera Pasto-Túquerres, 1820-2600 m, 30 Nov 1962, C. Saravia & R. Jaramillo 1869 (COL). Río Guaitara valley near
junction of Panamericana and road to Táquerres, 1700 m, 1 Mar 1986, J. Wood 5326 (COL, K, US).
2. Leptochloa fusca (L.) Kunth, Révis. Gramin. 1:91. 1829. Festuca fusca L., Syst. Nat., ed. 10, 2: 876. 1759, nom. cons.
prop. Diplachne fusca (L.) P. Beauv. ex Stapf, Fl. Cap. 7:591. 1900, nom. illeg. hom. Diplachne fusca (L.) P. Beauv. ex Stuck., Anales
Mus. Nac. Buenos Aires 11:128. 1904, nom. illeg. hom. Diplachne fusca (L.) P. Beauv. ex Roem. & Schult. Uralepis fusca (L.) Steud.,
Syn. Pl. Glumac. 1:247. 1855. Tree: PALESTINE: F. Hasselquist s.n. (LecrorvrE: LINN 92.211, designated by Phillips, Fl. Trop. East
Afr. Gram. 2:281. 1974).
Annuals, ascending to erect, sometimes geniculate and rooting at lower nodes. Culms to 130 cm tall (when
erect); 1-8 mm wide, branched or unbranched; nodes glabrous; internodes (0.5-)3-26 cm long, hollow,
herbaceous or sometimes firm. Leaf sheaths longer or shorter than the internodes, glabrous on sides and
margins; collars mostly light brown or green; ligules 5-8 mm long, hyaline, apically attenuate but often
becoming lacerated due to mechanical damage; blades (3-)5-50 cm long, 2-6 mm wide, linear, flat gla-
brous to somewhat scabrous above and below. Panicles 15-150 cm long, mostly 2-30 cm wide, partially
included at base to completely exserted; the branches (1.5-)3-20 cm long, (325-35 per panicle, alternate,
sometimes reflexed or steeply erect but mostly somewhat ascending, stiff, minutely scabrous, the branch
axils glabrous. Spikelets 5-12(-14) mm long, short pedicellate, more or less secund, sometimes distant near
base of branches but overlapping near branch tips; florets (3-)6-12; callus glabrous or hairy; lower glumes
1.0-3.5(-4.9) mm long, membranous, lanceolate to ovate, usually scabrous on the midvein, broadly acute
to acute, awnless or infrequently shortly awned; upper glumes 1.8-5.5 mm long, elliptic, usually ovate or
widely ovate, sometimes obovate, scabrous on midvein, obtuse (rarely) or acute at apex, rarely short-awned;
lemmas 2.3-6.0 mm long, membranous, elliptic, ovate, or lanceolate, the lateral veins distinct and often
slightly excurrent, more or less sericeous on lateral veins and the midvein; apex truncate, obtuse, to acute
or acuminate and sometimes bifid, awnless, mucronate, or awned, the awn to 3.5 mm long; paleas subequal
or slightly exceeding lemma, more or less sericeous on veins; apex acute or obtuse; stamens 1, 2 or mostly
3; anthers 0.2-2.7 mm long. Caryopses 1.0-2.4 mm long, 0.7-1.2 mm wide, elliptic, ovate, or obovate in
hilar profile, transversely elliptic to depressed obovate in transverse section, hilar groove lacking, smooth
or sometimes slightly rugose, brown; pericarp weakly adnate to the endosperm.
Distribution —Widespread and common to abundant in warm-temperate and tropical areas, between
approximately 49°N and 40°S in the New World and 40°N and 42°S in Old World; mostly below 2000 m.
Comments.—Populations of the Leptochloa fusca complex often appear morphologically distinct in lo-
866 I 1 nfthn Dat ical D h Institut f Texas 2(2)
cal areas, and dozens of names have been applied to regional forms throughout the Americas, Africa, and
Australia. Since all morphological characters intergrade when considered globally most local variants do
lti istical studies
not merit taxonomic recognition (Snow 1997). However, herbarium work and
(Snow, unpubl.) based on eleven population samples (n = 30) from America, Africa, and Australia, support
the recognition of four subspecies. The subspecies, which are usually easy to distinguish, are: L. fusca subsp.
fusca, a polymorphic paleotropical taxon adventive in a few areas in the New World (cola 1995); L. fusca
subsp. muelleri (Benth.) N. Snow, known from much of the interior of Australia (Sharp and Simon 2002);
L. fusca subsp. uninervia, native to the neotropics but adventive elsewhere; and L. fusca subsp. fascicularis,
native to the temperate and tropical regions of the New World.
KEY TO SUBSPECIES OF LEPTOCHLOA FUSCA IN COLOMBIA
—
. Lowermost panicle branches often inserted in upper sheath; uppermost leaf blade length usu-
ally exceeding length of panicle; leaf sous sometimes mottled with anthocyanin parus lem
often smokey wis at maturity and Po i Pa in oe half a.L. fusca ies fascicularis
1. Lowermost | g lly [er leaf | lade | = g y NOL tex-
not, sometimes dark green or it bedus but usually not as a distinct spot 2b.L. Ricca an uninervia
2a. Leptochloa fusca subsp. fascicularis (Lam.) N. Snow, Novon 8:78. 1998b. (Fig. 2, A-C). Festuca fasci-
cularis Lam., Tabl. Encycl. 1:189. 1791. Diplachne fascicularis (Lam.) P. Beauv., Ess. Agrostogr. 81, 160, pl. 16, f. 9. 1812. Cynodon
fascicularis (Lam.) Raspail, Ann. Sci. Nat., Bot. 5:303. 1825. Tyre. South America. D. Richard s.n. (HoLOTYPE: Pl; isotypes: US-2875408!,
US-2875408A fragm. ex P!).
Annuals, ascending to erect. Culms (3.5-)15-130 cm tall, 1-5(-8) mm wide, often branching, round; nodes
glabrous; internodes 3-18 cm long, herbaceous, hollow. Leaf sheaths mostly longer than the internodes,
glabrous on sides and margins; collar light green or brown; ligules 5-7 mm long, hyaline; blades 3-45 cm
long, 2.5-7 mm wide, linear, usually sparsely scabrous above and below, flat but becoming involute when
dry, midvein prominent. Panicles (1.5-)10-72 cm long, (124-22 cm wide, generally partially included
below; the branches 3-35, (0.5-)3-22 cm long, alternate, erect, ascending, or occasionally divergent, rigid
or slightly flexuous, minutely scabrous, the axils glabrous. Spikelets 5-12 mm long, distant to mostly imbri-
cate, pedicels less than 1 mm long; florets 6-12; callus glabrous; lower glumes 2-3 mm long, membranous,
lanceolate or somewhat asymmetric, scabrous on midvein, acute to aristate and occasionally short-awned;
upper glumes 2.5-5.0 mm long, membranous, elliptic to ovate, scabrous on midvein, obtuse, aristate, or
short-awned; lemmas 2.5-5 mm long, membranous, lanceolate, light to dark green, light brown or smoky
white, the lateral veins pronounced and extending to edges often as small teeth, sparsely sericeous along
lateral veins and often midvein, glabrous between veins; apex acute to attenuate, often mucronate or with
awns to 3.5 long; paleas generally subequal to lemma, elliptic, sericeous along veins; apex acute to obtuse;
stamens 3; anthers 0.2-0.5 mm long, yellow. Caryopses 1-2 mm long, 0.8-1.0 mm wide, elliptic to obovate
in hilar profile, transversely elliptic in transverse section, hilar groove lacking, smooth, brown; pericarp
weakly adnate to the endosperm.
Distribution and habitat.—Massachusetts (USA) and southern Ontario (Canada) west to Washington
(USA) and south to about latitude 40° S in Argentina; open mesic areas, brackish marshes along the eastern
US seaboard, agricultural lands, ruderal sites along roads, and saline flats. Elevation from sea level to 1850 m.
Specimens examined. COLOMBIA. La Guajira: Costa del Caribe, 1 li ] Arid M ya El Pájaro, 29 Jan 1964,
A. Dugand 6667 (COL, US).
2b. Leptochloa fusca subsp. uninervia (J. Presl) N. Snow, Novon 8:79. 1998b. (Fig. 2, D-F). Megastachya
s Presl, cd Haenk. 1:283. 1830. Poa uninervia (J. Presl) Kunth, Enum. Pl. 1:344. 1833. Eragrostis uninervia (J. Presl)
Steud., Syn. Pl. Glumac. 1:278. 1854. Brizopyrum uninervium (J. Presl) E. Fourn., Mex. Pl. 2:121. 1886. Leptochloa uninervia g.
Presl) Pup ENDE: Contr, U.S. Natl. Herb. 18(7):383. 1917. Diplac ia (J. Presl) Parodi, Revista Centro Estud. Agron.
18:147. 1925. Tere: MÉXICO: Haenke 101 (uecrorree: PRI, designated by Snow, Novon 8:79. 1998b; DUPLICATES or LECTOTYPE: LE-
TRIN-2424.01!, US-78697 fragm.!, Wt).
867
Snow et al., Leptochloa in Colombia
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Annuals or occasionally weakly perennial in regions lacking frosts,
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2-5 mm wide, often branching, round;
Leaf sheaths longer or shorter than the internodes, often flattened below, glabrous on sides and margins;
ligules 5-8 mm long, hyaline; blades (2-)5—37 cm long, 2.0-5.5 mm wide, linear,
usually densely scabrous above, sparsely to densely scabrous below, flat but becoming involute when dry,
5
collars mostly tan or green;
868 J | of the Botanical R h Institute of Texas 2(2)
midvein prominent. Panicles 10-57 cm long, (0.5-)3-18 cm wide, usually exserted; the branches (3-)10-60,
(0.3-)2-11 cm long, branches mostly alternate, P to erect, stiff or sea DOXUOUS, mun
scabrous, th Spikelets 5-10 mm long, rarely distant to normally imbricate ( ghtly
so), pedicels mostly less than 1 mm long; florets (3-)6-10; callus sparsely sericeous; Iove glumes 1.0-2.6
mm long, membranous, narrowly triangular, lanceolate, or ovate, glabrous or scabrous on midvein, acute to
aristate or mucronate; upper glumes 1.8-2.8 mm long, membranous, obovate to widely ovate, glabrous or
scabrous on midvein, obtuse, acute, or rarely mucronate; lemmas 2.0-3.6 mm long, membranous, ovate or
elliptic, light brown to very dark green or somewhat plumbeous, lateral veins more or less prominent and
generally extending to edges, sparsely sericeous below on lateral veins and often midvein, glabrous between
veins; apex broadly acute, more commonly obtuse to truncate, sometimes bifid or mucronate; paleas subequal
or slightly longer than lemma, membranous, elliptic, sericeous along veins; apex obtuse; stamens 3; anthers
0.4-1.0 mm long, yellow. Caryopses 1.0-1.5 mm long, 0.7-0.8 mm wide, elliptic, ovate, or obovate in hilar
profile, transversely elliptic in transverse section, hilar groove lacking, smooth or slightly rugose, brown;
pericarp weakly adnate to endosperm.
Distribution and habitat.—In the New World mostly south of latitude 37*N, south to Argentina, occa-
sionally adventive in Old World (Snow and Simon 1999); open mesic areas, agricultural lands, saline flats,
mangrove swamps. Elevation is from sea level to 1200 m.
Specimens examined. COLOMBIA. Atlántico: Entre Palmar de Varela y Ponedera, bosques xerofíticos, 10 m, 27 Ago 1960, A. Dugand
5304 (COL, US). Magdalena: Ciénaga, alrededores de Aguacoca, 20 m, 21 Feb 1959, R. Romero-Castañeda 7268 (COL, US).
3. Leptochloa panicea subsp. brachiata (Steud.) N. Snow, Novon 8:79. 1998b. (Fig. 3, A—E). Leptochloa brachiata
Steud., Syn. Pl. Glumac., 209. 1854. Tyre: GUADALOUPE [French Republic]: Duchassaing s.n. (noLorves: P!; US fragm.!).
Fleusine filiformis Pers., Syn Pl. 1: 87. 1805. Leptochloa filiformis (Pers.) P. Beauv., Ess. Agrostogr.: 71, 163, 166. 1812. Tyre: SOUTH
AMERICA: Richard s.n. (Horory: PU.
Annuals, ascending to erect. Culms (10-)30-150 cm tall, mostly less than 5 mm wide, frequently branched;
nodes glabrous; internodes mostly 5-30 cm long, hollow, herbaceous or sometimes firm. Leaf sheaths lon-
ger or shorter than the internodes, sparsely to moderately pilose (hairs with papillose bases); collars often
magenta; ligules 1.0-3.2 mm long, membranous but erose apically; blades 6-25 cm long, 2-21 mm wide,
linear, flat, glabrous to sparsely pilose on both surfaces. Panicles 8-30 cm long, usually 5-35 cm wide,
occasionally partially included at base; the branches (1-)3-19 cm long, (3-)10-100 per panicle, alternate,
ascending (mostly) to somewhat reflexed apically (but otherwise firm), minutely scabrous, the branch axils
glabrous. Spikelets 2-4 mm long, short pedicellate, more or less secund, sometimes distant near base of
branches but overlapping near branch tips; florets 2-5(-6); callus glabrous; lower glumes 1.6-4 mm long,
membranous, lanceolate, usually scabrous on the midvein, acute, awnless; upper glumes 1.6-3.6 mm long,
lanceolate, awnless; lemmas 1.3-1.7 mm long, membranous, lanceolate, more or less sericeous on lateral
veins and the midvein; apex acute, awnless; paleas subequal to lemma; apex acute or obtuse; stamens 3;
anthers 0.2-0.3 mm long. Caryopses 0.9-1.2 mm long, 0.5-0.6 mm wide, more or less rounded in hilar
profile, widely depressed obovate to obdeltoid in transverse section, hilar groove narrow and shallow, smooth
or sometimes slightly rugose, brown to reddish; pericarp adnate to the endosperm.
Distribution and habitat.—Southern third of United States through Mesoamerica and the West Indies
to southern Brazil and Argentina; disturbed, mesic, and agricultural sites, to ca. 2800 m
Comments.— Leptochloa panicea is a polymorphic species occurring throughout — of the warm
temperate and tropical regions, and is comprised of three subspecies (Snow 1997, 19983). Snow and Davidse
(1993) reported that the name then in use for the widespread New World taxon, L. filiformis (Lam.) P. Beauv.,
was an illegitimate later homonym that should be replaced by L. mucronata (Michx.) Kunth. Although not
the first to do so, Nowack (1994, 1995) recently placed New World specimens as a subspecies of the oth-
erwise Old World species L. panicea, and made the new combination L. panicea subsp. mucronata. Included
in Nowack's concept of L. panicea subsp. mucronata were two taxa formerly recognized as L. filiformis vars.
filiformis and attenuata, which Snow (1997, 1998b) treated (respectively) as L. panicea subsp. brachiata (Steud.)
Snow et al., Leptochloa in Colombia 869
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Fic. 3. Leptochl pani I p. brachiata. A, Habit. B. Inflorescence. C. Porti f the leaf sheath witt ing of papill based hai . D. Spikelet.
E. Floret with rachilla (lateral view).
870 Journal of tani i f Texas 2(2)
N. Snow and L. panicea subsp. mucronata (Michx.) N. Snow. The much more widespread taxon, referred to
earlier as L. filiformis or L. mucronata (Snow & Davidse, 1993), was given the new name L. panicea subsp.
brachiata (Snow 1998b).
Drawing on field experience and examination of over 4,000 herbarium specimens, Snow (1997) con-
cluded that the phenetic variation is sufficiently consistent to warrant recognition of three subspecies. Only
L. panicea subsp. brachiata is present in Colombia.
Vernacular name.—Pasto paja-mona (Valle del e
Specimens examined. COLOMBIA. Antioquia: Mun. Necocli, 4 N lí-Arbolete 1 , 160 m, 29 Sep
1986 Ji yid et a 298 (COL, HUA, MO). Mun. Arboletes, en playa, 30 Sep 1986, J. Betancur et di 316 (COL, HUA, MO). La Pintada,
00 m, 20 Oct 1947, C. Blachman, J. Mejía & F. Barkley 17-C284 (COL). Mun. Bolívar,
carretera Bolivar-Caramanta, len 5 arriba de T margen izquierda del Río Cauca, Cabrera 647 (MO). Vicinity of Medellín, 30 Jul
1927, R. Toro 327 (US). Atlántico: Entre Palmar de Varela y Ponedera, hacienda El Paraíso, orillas del Río Magdalena, 10 m, 28 Jan/8
Feb 1950, A. Dugand 4435 (COL). Barranquilla, El Prado, 60 m, 30 Dec 1958, A. Dugand 5074 (COL). Cerca de Barranquilla, carretera
a Juanmina, 20 m, 10 Jul 1960, A. Dugand 5 (COL, US). Puerto Colombia, Sabanilla, 21 Nov 1912, A. H k 9932 Al sur de
Barranquilla, cerca de Martillo, 5-10 m, 29 Abr 1960, L. Mora 1344 (COL). Barranquilla, year 1926, B. Paul C-17 (US). Bolívar: Mun.
Turbaco, jardín botánico “Guillermo Piñeres,” 6 Ago 1980, J. Espina 499 (COL). Cartagena, 20 Nov 1912, A. Hitchcock 9919 (US). Mun.
San Martín de Loba, corregimiento La Ribona, reserva natural El Garcero, finca La Buenaventura, Ramirez & Villa 4664 (MO). Cauca:
Between Río Mayo and Galindez, 800 m, 26 Oct 1983, J. Wood 4060 (COL, US). Cundinamarca: 14 km WNW of Melgar, 330 m, 8
Jan 1974, G. Davidse, A. Gentry & F. Llanos 5573 (COL, MO). Poblado de Nariño, zonas ruderales, 340 m, 15 Feb 1986, J. Fernández &
R. Jaramillo 5230 (COL). Montañuela, San Bernardo, 1400 m, 19 Mar 1961, J. Idrobo & Dumont 4438 (COL). Mun. Tocaima, ca. 500 m,
Dec 1934, E. Pérez-Arbeláez 2419 (COL), 2424 (COL). Mun. Útica, en granja ide 467 m, 13 Oct Pam C. Saravia PA Dus
5 km of El Nilo, 16 Jun 1985, J. Wood 4927 (US). Huila: Mun. El Agrado, quebrada La Yaguilga,
circundantes, ca. 700 m, Sep 1986, J. Fernández & G. Morales 6839 (COL). Magdalena: Santa Marta, 1898- 1899, H. Suh 213 (US), 2163
(US). Santander: Bajo Magdalena, Isla de Venado, inter Billete Blanco et El Presidio, 7 May 1926, S. Juzepczuk 4770 (US). San Gil, 1250
m, year 1983, J. Wood 3885 (COL). Bajo Magdalena, Barrancabermeja, 6 Abr 1926, G. Woronow & S. Juzepczuk 4428 (US). Tolima: Mun.
Espinal, 500 m, Sep 1859, M. Lindig 1033-P33. Valle del Cauca: Río Cauca, alrededores de Palmira, 1000 m, 3 Dec 1947, R. Cardeñosa et
al. 17-C937 (COL, US), R. Murgueitio et al. 17-C929 (US). Hacienda El Trejo, entre El Cerrito y Palmira, 1050 m, 28 Dec 1938/5 Jan 1939,
H. Garcta-Barriga 6339 (COL, US). Mun. Palmira, Ingenio La Manuelita, 1000 m, 29 Jul 1966, J. Páez 23 (COL). Mun. Palmira, granja
agrícola experimental, 5 Jan 1957, A. Ramírez 14 (COL). Loboguerrero, 1050 m, 12-14 Nov 1962, C. Saravia 1483 (COL). Mun. Palmira,
1100 m, no date, A. Villamizar s.n. (COL: 82436). Unknown department: Río Frío, 23 Jun 1906, H. Pittier 1584 (US).
4. Leptochloa panicoides (J. Pres!) Hitchc., Amer. J. Bot. 21:137: 1934. (Fig. 2, G-I). Megastachya panicoides J.
Presl, Reliq. Haenk. 1(4-5):283. 1830. Poa vede Q. Presl) Kunth, Enum. Pl. 1:343. 1833. ide panicoides (J. Presl) Steud.,
. Glumac. 1:278. 1854. Diplachne panicoides (J. Presl) McNeill, Brittonia 31:402. 1979. Tree: MEXICO: Habitat in México
ad Gs oui verosimiliter inlutosis, Haenke s.n. (HOLOTYPE: PR; isorvees: IT: LE-TRIN-2121.02!, MO-2 Pin MO-2109567 line
drawing!, US-78688 fragm
Annuals, often geniculate below but generally erect. Culms (7-)45-110 cm tall, 1-7 mm wide, branch-
ing (sometimes profusely), round; nodes glabrous; internodes (1-)4-10 cm long, herbaceous, hollow.
Leaf sheaths generally longer than the corresponding internodes, somewhat flatted below, glabrous, the
margins sometimes very sparsely ciliate along the lower half; collars sometimes magenta; ligules 2.2-3.8
mm long, membranous, truncate, somewhat erose apically; blades 4-20 cm long, mostly 4-8 mm wide,
linear or the shorter ones lanceolate, glabrous to minutely scabrous above and below. Panicles (1-)20-35
cm long, (324-12 cm wide, generally exserted; the branches (2040-90, 2.5-7 cm long, alternate, mostly
ascending, stiff to weakly flexuous, minutely scabrous, the axils glabrous. Spikelets 4-5 mm long, generally
somewhat imbricate, pedicles 1-2 mm long; florets (4-)5 or 6(-7); callus glabrous; lower glumes 0.9-1.9
mm long, membranous, narrowly lanceolate to lanceolate or sometimes falcate, minutely scabrous on the
midvein, acute; upper glumes 1.8-2.3 mm long, membranous, ovate, minutely scabrous on the midvein,
acute to obtuse; lemmas 2.4-3.0 mm long, membranous, narrowly elliptic to ovate, tan, maroon, or green,
lateral veins pronounced, projecting abaxially, and usually bright green, sericeous (sometimes scarcely so)
at base of lateral veins and/or midvein; apex acute to broadly acute, awnless or mucronate; paleas subequal
to ca. 2/3 as long as the lemma, membranous, ovate to elliptic, glabrous throughout; apex obtuse; stamens
3; anthers 0.6—0.8 mm long, maroon or yellow. Caryopses 1.1-1.4 mm long, 0.7 mm wide, elliptic in hilar
Snow et al., Leptochloa in Colombia 871
profile, depressed obovate in transverse section, hilar groove lacking, smooth or somewhat rugose, brown;
pericarp tightly adnate to the endosperm.
Distribution and habitat. —From central Mississippi and Ohio River drainages in the United States south
through Mesoamerica to Brazil; MERC in mesic sites. Elevation: sea level to 500 meters.
Specimens examined. COLOMBIA. A Beira Rí la f. la Wandurraga, 28 Sep 1946, G. Black & R. Schultes
46-158 (COL); Río Loretoyacu, Oct 1946, G. Black & R. Schultes 46-160 (COL, GH, M 46-184 (COL, bois Pet Amazónico, Loretoyacu
River, ca. 100 m, Oct es R. uid Black 8503 (COL, GH, US). C El Yopal lables, finca
La Armonía, d , ca. 400 m, 3 Nov 2007, D. Giraldo-Cañas et al. 4156 (CAUP, COAH, COL).
5. Leptochloa scabra Nees, Agrostogr. Bras. 2: 435. 1829. (Fig. 1, D-F). Diplachne scabra (Nees) Nicora, Hickenia
2(19):91. 1993. Tyre: BRAZIL: Pará, in ripa fluminum Amazonum, Tagipuris, K.EP von Martius s.n. (HOLOTYPE: MI; isotypes: BAA-1514
fragm ex M, US-88699 fragm.ex MI).
Vigorous annuals mostly erect, rarely geniculate at the base. Culms (12-)20-125 cm tall, (0.6-)1-6.2 mm
wide, branching, keeled and compressed; nodes glabrous; internodes 6-9 cm long, herbaceous or hard, hol-
low. Leaf sheaths usually shorter than or equal to internodes, often flattened below, glabrous to scabrous, the
margins glabrous; collars magenta to dark maroon; ligules (0.5-)1.5-2.5 mm long, thinly membranous with
numerous hairs arising from base of the blade, truncate, fimbriate to erose apically; blades (4-)25-35(-50) cm
long, (328-16 mm wide, linear to lanceolate, scabrous above and below, flat, midvein prominent. Panicles
(10-)15-35 cm long, (2-)7-12 cm wide, usually partially included; the branches 50-150, 25-12 cm long,
alternate, ascending to erect, flexuous to prominently arcuate, minutely scabrous, the axils glabrous. Spikelets
3.0-4.5 mm long, mostly imbricate, short-pedicellate; florets 4-6; callus glabrous; lower glumes 0.8-1.6 mm
long, membranous, narrowly triangular, triangular, or trullate, minutely scabrous on the midvein, acute
to mucronate; upper glumes 1.1-2.1 mm long, membranous, ovate, glabrous or scabrous, obtuse, acute, or
mucronate; lemmas 2.1-2.4 mm long, membranous, lanceolate to narrowly ovate, light green becoming
tan, the lateral veins extending to near edge of lemma but less distinct apically, sparsely sericeous along
lower 1/2 of veins; apex acute, awnless or mucronate; paleas subequal to lemma, membranous, lanceolate,
sparsely sericeous along lower part of lateral veins, bifid apically; stamens 3; anthers 0.2-0.4 mm long,
yellow or light brown. Caryopses 0.8-1.3 mm long, 0.3-0.5 mm wide, elliptic to obovate in hilar profile,
depressed obovate in transverse section, hilar groove lacking, smooth to slightly rugulose; pericarp weakly
adnate to endosperm
Distribution and habitat.—Native distribution from the United States of America (Alabama and Louisi-
ana) to the West Indies, Mesoamerica to Paraguay and Brazil. Its introduced range includes Africa (Angola
and Mali) and Papua New Guinea; creek banks, disturbed alluvial areas, river margins, agricultural fields.
Elevation 0-750 m.
Comments.—Leptochloa scabra can be confused with L. panicoides. The former is distinguished by its
shorter and more tightly imbricate spikelets, the keeled lemmas, its distinctly flexuous to arcuate panicle
— and culms that are somewhat flattened near the base.
d. COLOMBIA. Amazonas: Leticia, 17 Ago 1946, G. Black & R. Schultes 46-2 (US). Rio mee acu, Oct 1946, G.
Black & R. Schultes 46-185 (COL, GH, US). Río Caquetá, J. Duivenvoorden et al. 71 (MO). Antioquia: Mun. Chigorodó, hacienda Monte-
cristo, 25 km después de Apartado, cultivo de chontaduro (Bactris gasipaes), 40 m, 2 Oct 1986, J. Betancur et Pi 345 (COL, HUA, MO).
Atlántico: al sur de Barranquilla, Río Magdalena, en la isla frente a Puerto Giraldo, 5-10 m, 29 Abr 1960, L. Mora 1455 (COL). Bolívar:
Río Sinú, 10-100 m, 11 Feb 1918, F. Pennell 4195 (US). Boca Esmeralda, on Río Sinú, 200-400 m, 1 Mar 1918, F. Pennell 4554 (GH, NY,
US). Cauca: Mun. Patía, a 2 km de Mojarras, granja Universidad de Nariño, 700 m, 23 Abr 1988, B. Ramírez 1127 (COL, PSO). Chocó:
alrededores de la confluencia de los Ríos Chintadó y Truandó, 25 Mar 1958, P. Bernal 52 (COL, MO). Río Truandó, gallery between the
and Río Salado, Duke 11100 (NY). eas iid 202 Ee Mis cuales Bacco e an 400 m, Dec 1976, C. pai
m con, 21 OM Guaviare: Mun. San José I Fuga
aguas arriba en bordes de Río, los d i 200 m, 17 Feb 1996, R. López 997 Cos COL). ne Nro.
2, margen derecha del Rio — 23 Feb oe P. Pinto & M Sastre 1012 (COL). Huila: 3 km W of Palermo towards Santa María,
500 m, 16 Mar 1985, J. Wood 4749 (CO d entre caño Calrín y 1 tera, del km 10 al 15, 20 Jun 1970,
N. de López 472 (COL) Región de Santa MS. en Ciénaga, El Arsenal, alreded de la ciudad, 17 Feb 1950, R. Romero-Castañeda 1979
(COL). Meta: Reserva Nacional La Macarena, margen izquierda del Río Guayabero, al pi ] frente a la confluencia
| af ob D ? al D hi PA
872 Jou f Texas 2(2)
del caño Losada, 300 m, 3 Mar 1959, R. Jaramillo & J. Hernández 2079 (COL). Margen derecha del Río Guayabero, raudal de La Macarena
(Angostura Nro. 1), 350 m, 24 Jan 1959, P. Pinto & Bischler 343 (COL, US). Margen izquierda del Río Guayabero, 10 km abajo del caño
Losada, 350 m, p 1959, P. Pinto Muse 368 ds n Sucre: Mun. Tolú, pn La Bora 17 km Tolú-Coveñas, golfo de
Morrosquillo 0—5 m, 20 Sep 1990, J. Betancur
& M. Berrto 1994 (COL, HUA, MO, US). Valle "m Cauca: Mun. NoD lagen La Carmelita, sección San Martin, Ramos et al. 2828
MO). Mun. Yumbo, Finca Río Grande, pasture around the houses, 1200 m, 13 Jun 1998, D. Stancik 793 (COL). Unknown department:
J.C. Mutis 5428 (US).
6. M Saar virgata (L.) P Beauv., Ess. Agrostogr. 71, 161, 166, pl. 15, f. 1. 1812. (Fig. 1, G & H). Cynosurus
L. Syst. Nat. (ed. 10):876. 1759. Festuca virgata (L.) Lam., Tabl. Encycl. 1:189. 1791. Fleusine MC Pers., Syn. Pl. 1:87.
1805. Vues virgata (L.) G. Mey., Prim. Fl. Esseq., 74. 1818. Cynodon virgatus (L.) Raspail, An: i. Nat., Bot. 5:302. 1825.
Tre: JAMAICA: P Browne s.n. (Lgcrorvee: LINN-91.18! designated by Hitchcock, Contr. U.S. Natl. rs Di 1908).
Perennials, ascending to erect or infrequently geniculate and rooting at nodes. Culms 30-100 cm tall, (1-)3-5
mm wide, branched, round or occasionally flat; nodes glabrous; internodes 4-35 cm long, herbaceous or
firm, solid. Leaf sheaths longer or shorter than internodes, round, glabrous, or shortly sericeous to sometimes
pilose, particularly near the collar, the margins similar; collar green or tan; ligules 0.2-1.0 mm long, firmly
membranous or cartilaginous, truncate, somewhat fimbriate or erose apically; blades (2.5-)10-44 cm long,
4-16 mm wide, lanceolate, glabrous, scabrous, or sparsely pilose from small papilla above and below, flat,
midvein prominent or reduced. Panicles (5212-60 cm long, (2-)3-20 cm wide, exserted; the branches 9-60,
(1.5-)3-18 cm long, alternate or rarely subdigitate, ascending to nearly erect, flexuous, scabrous, the axils
usually pilose. Spikelets (1.7-)3-4 mm long, mostly imbricate or somewhat distant at base of panicle branches,
sessile or nearly so; florets (224 or 5-6); callus glabrous, scabrous, to velutinous or pilose; lower glumes
(1.0-)1.7-2.9 mm long, membranous, lanceolate, minutely scabrous on midvein, acute, attenuate, or rarely
obtuse; upper glumes (1.4-)1.7-2.5(-3.8) mm long, membranous, ovate to lanceolate, minutely scabrous on
midvein, obtuse, acute, acuminate, or rarely mucronate; lemmas (1.5-)2.3-3.6 mm long, chartaceous, ovate,
light brown to maroon but apparently never deep green, lateral veins faint and not extending to the edges,
nearly glabrous to sericeous on veins, glabrous or sparsely sericeous on sides; apex acute or rarely obtuse,
awnless or awned (awns up to 11 mm long in other parts of South America); paleas subequal to lemma,
firmly membranous or chartaceous, lanceolate to narrowly elliptic, glabrous or sparsely sericeous between
veins; apex acute; stamens 2; anthers ca. 0.5 mm long, light purple. Caryopses (1.0-)1.3-1.8 mm long, ca.
0.5 mm wide, narrowly elliptic to ovate in hilar profile, rounded shallowly obdeltate to widely obovate in
transverse section, hilar groove lacking or fairly pronounced but shallow or relatively broad and shallow,
smooth, mostly light reddish brown; pericarp tightly adnate to the endosperm.
Distribution and habitat.—From southern Texas, southern Florida, and the West Indies through Me-
soamerica and South America (excluding Chile) and locally in Papua New Guinea and Papua Province,
Indonesia (formerly Iryan Jaya); occurring in numerous vegetation and soil types, but mostly in more mesic
climates. Elevation: sea level to 1300 m.
Vernacular names.—Painoo (Casanare), Rabo de zorro (Tolima).
Specimens examined. COLOMBIA. Antioquia: Vuelta de Acuña, Río Magdalena, 125-130 m, 14 Jan a F. Pennell 3803 (NY, US).
Mun. Santa Fe de Antioquia, finca Cotové, 580 m, 24 Sep 1986, R. Pohl 15594 (HUA, MO, US). Vicinity dellín, 30 Jul 1927, R. Toro
332 (US). Mun. Santa Fe de Antioquia, along road Santa Fe de Antioquia-Anzá, ca. 1 km W of crossing of Río Tonuzco at its confluence
ith Río Cauca, near Finca Cotové, 615 m, 19 Sep 1987, J. Zarucchi et al. 5486 (COL, MO, NY). Atlántico: Entre Juanmina y Cuatrobocas,
as roe bosques áridos, 80-100 m, 15-16 Jan 1946, A. Dugand & R. Jaramillo 4037 (COL, US). Entre Palmar de Varela y Ponedera,
orillas del Río Magdalena, 10 m, 8-10 Jan 1954, A. Dugand 4704 (COL, US). Bolívar: E the Caño aia N to
Los Piñones, Island of Mompós, Lands of Loba, Abr-May 1916, H. Curran 260 (US). Cartagena, 20 Nov 1912, A. Hitch 9923 (US).
Magdalena, boca de bosque, 50 m, 18 Dec 1969, J. Idrobo 6309 (COL). Soplaviento and vicinity, 5-10 m, 16 Nov 1926, E. Killip & A. Smith
14606 (GH, MO, NY, US). Cañabetal, Río Magdalena, 90-100 m, 15 Jan 1918, F. Pennell 3883 (GH, MO, NY, US). Casanare: Mochuelo
y Tsamani, margen derecha del bajo Casanare, 3 na 6 d iod. E dd 525 (COL). Chocó: Río leer gallery between the boom
and Rio Salado, Duke 11079 (NY). Cundi zonas ruderales, 340 m, 15 Feb 1986, J. mena a i io
5206 (COL). Mun. Caparrapt, hacienda Saldaña, 1280 m, 10-13 Jun 1939, H. Garcia-Barriga 7742 (COL). La Mesa
de La Mesa a San Javier, 950-1320 m, 2-20 Jan 1947, H. García-Barriga 12135 (COL). Montañuela, San Bernardo, 1400 m, 19 Mar 1961,
J. Idrobo & Dumont 4440 (COL). Cune, 1200 m, Jul 1860, M. Lindig 1121-P122 (COL). Nariño, ca. 500 m, Jul 1930, E. Pérez-Arbeláez 460
Snow et al., Leptochloa in Colombia 873
(COL). Nilo, Jul 1930, E. Pérez-Arbeláez 504 (COL). Near La Mesa on the road El Colegio, 1000 m, year 1983, R. Wood 3588 (COL). Huila:
Mun. Villavieja, Fortalecillas, km 2 vía a Villavieja, 450 m, B. Ramírez 13169 (CAUP, COL). Río Saldaña to Natagaima, Rusby & F. Pennell
240 (NY). Magdalena: Santa Marta, 1898-1899, H. Smith 188 (GH, US). Santa Marta, 1898-1899, H. Smith 2160 (US). Meta: Los Llanos,
Río Meta, El Porvenir, 16 Oct 1938, J. Cuatrecasas 3692 (COL, NY, US). Los Llanos, río Meta, isla de arenal en Vuelta Mala, 17 Oct 1938,
J. Cuatrecasas 3790 (US). Maren a e es adds 10 km pod del caño Lozada, 350 m, 30 Jan 1959, P. Pinto & Bischler 366
800 1, 15 Jan 1989, J. Ortiz 1316 (C OD.
San Andrés, Providencia y Santa Catalina: Isla de San dates. ca. 5 m, no date, R. Tol 38 (con. Santander Bajo alo Isla
de Venado, inter Billete Blanco et El Presidio, 7 May 1926, S. Juzepczuk 4744 (US), 4750 (US). 6 de Palyón, 800 m, 2 Ago 1966,
A. Robinson & C. BRI 3023-A (US). San Gil, 1250 m, year 1983, R. Wood 3868 (COL). Chicamocha Valley, Pescadero ie in
stable sand along f the Río Manco, 700 m, 3 Jul 1984, J. Med 4490 (COL). Tolima: Mun. Espinal, en Calicheros frente al
molino San Martín, no dave: R. Echeverry 1080 (COL). Mun. Espinal, 500 m, Dec 1859, M. Lindig 1107-P106 (COL). Between Guamo
Saldafia, 300 m, 11 Nov 1983, R. Wood 4096 (COL). Valle del Cauca: ek del Valle, entre Hi. y Cabuyal, 1000 m, 2 Jun d
J. Cuatrecasas 14495 (US). Plana del Valle, extremo norte, Cartago, en San Jerónimo, 980 m, 15 Nov 1946, J. Cuatrecasas 22841 (US). La
Manuelita, near Palmira, eastern side of Cauca Valley, 1100-1302 m, Dec 1905-Jan 1906, H. Pittier 832 (US). Hacienda Vidal, ca. 3 km
delante de Vijes, vía Vijes-Yotoco, Ramos & Ramos 2278 (MO). Corregimiento de Mulaló, en la vía Yumbo- ios Ramos & Ramos 3151 (MO).
Entre Loboguerrero y Yumbo, 1000-1100 m, 14-18 Nov 1962, C. Saravia 1620 (COL). Mun. Cali, Uni 1 Valle, Silverstone-Sopkin
1996 (MO). Unknown departamento: J.C. Mutis 5427 (US), 5429 (US).
EXCLUDED NAME
Tridens virens Nees, Fl. Bras. Enum. Pl. 2:476. 1829. Mord virens (ees) uM Enum. Pl. 1:319. 1833. d ned virens
(Nees) Parodi, Rev. Fac. Agron. Vetrin. 6:14. 1927. Tyre: BRAZI
Permambucana, ad Joazeiro et alibi." We do not know oe eke type is hae and have not seen it. Chase and Niles (1962)
indicate that the “type is Leptochloa pei [=L. fusca subsp. fascicularis] or affine."
ACKNOWLEDGMENTS
We wish to thank the curators at BAA, COAH, CAUP, COL, HUA, LE, MO, SI, and US for their valuable help;
the Smithsonian Institution and Universidad Nacional de Colombia for supporting research visits by NS
and DGC to the United States National Herbarium; Utah State University for permission to use illustrations
that appeared in Flora of North America, Volume 25, prepared by Linda A. Vorobik and Karen Klitz; Victor
L. Finot and Stephan L. Hatch for providing helpful comments on the manuscript; and Alice Tangerini for
preparing all electronic plates. This article will contribute to projects “monograph of Leptochloa (Poaceae:
Chloridoideae)” of NS and PMP (in prep.), and "Estudios sistemáticos en gramíneas de Colombia: Parte II.,"
of the Universidad Nacional de Colombia (Bogotá) of DGC.
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WORLD GRAssES. (UPDATED UOUSLY). HTTP://MOBOT.MOBOT.ORG/W 3 T/SEARCH/NWGCHTML
ERAGROSTIS (POACEAE: CHLORIDOIDEAE: ERAGROSTIDEAE) IN COLOMBIA
Paul M. Peterson Diego Giraldo-Cañas
Department of Botan Instituto de Ciencias Naturales
National cl of Natural History Facultad de Ciencias
nian ho Universidad Nacional de Colombia
"P on pos 012, U.S.A. Apartado 7495, Bogotá D. C, COLOMBIA
DD n dagiraldoc@unal.edu.co
ABSTRACT
A ; : Ir 1 1 Ki : IE ;e Tl E "n Jin Canl] hian leer DK
A Tes 1 ` E. ; 1 ie DAT F niori 17. Lc 1
o r
and E rufescens Schrad. ex Schult
time in ALE F ford ]
jej [o]
descriptions, illustrations hical distributi ] holoeical and l
Y e e r T de O D
cal observations are Aud for ate native "a dventi ies of E is in Colombia. E i la (Kunth) Steud El Foe pana
Kunth) and E. unioloides (Retz.) Nees ex Steud. (= Pi ioloides Retz.) luded f he Colombian Fl i ll
as E. d are pue HRS i are eeu E to s non d a s une ex Steud. A La eae
) Mat
1
RESUMEN
di 5mi lel i lombi del gé E tis. Este gé 1 Colombia por 25
del 1 E. H 1 2e DAI E €— fT we
by E. rufescens S Schrad. ex Schult. se citan por
olombia. S 1 | l i l
de r r r X + F
lasd ipci leé , la iconografía,
r
especies y una subespecie,
í la fl
mbia, así
E D m 1 1 : o
Las o E. patula a Pour a Poa patula Kun) y E. , unioloides Qui. ) Nees ex Steud A Poa o Retz. ) se excluyen de la
cod o r
E A (A. Rich.) Hochst. ex Steud. Por otra parte, E. unioloides ha sido citad 1 áni j l
E
itl
para la Flora de Antioquia, p Imente el i de cl te a Poidium j ii (Hack) Matthei.
r E y o
Eragrostis is a large genus of approximately 350 species occurring in tropical, subtropical and warm temper-
ate regions throughout the world (Clayton & Renvoize 1986; Watson & Dallwitz 2008; Peterson et al. 1995,
1997; Lazarides 1997; Veldkamp 2002; Ingram & Doyle 2007, Peterson & Sánchez Vega 2007). There are
111 species of Eragrostis recorded in North, Central, and South America, and 67 native in South America
(Peterson & Boechat 2001; Peterson 2003; Peterson et al. 2001, 2007). The genus is characterized by having
many-flowered spikelets where the disarticulation of the lemma and palea occurs separately, lemmas that
are usually 3-nerved and unawned, longitudinally bowed-out paleas with ciliolate keels, paniculate inflores-
cences, and leaves with ciliate ligules (Peterson et al. 1997). Most species of Eragrostis occupy open habitats
with poor soils, and many occur in ruderal sites (Clayton & Renvoize 1986; Van den Borre & Watson 1994),
and their distribution exhibits wide altitudinal gradients and a high variation in humidity conditions, from
the sea level to 3600(-4000) m, and from pluvial environments to xeric habitats, respectively.
Colombia is placed in the Neotropical Region and encompasses 1,141,748 km?. The geomorphologi-
cal and environmental complexity of Colombia is tremendous where there is an altitudinal gradient from
sea level to over 5800 m, and climates can vary from wet and humid to xeric and dry. The country can be
divided into three major regions: the Andes with three Cordilleras (Occidental, Central, and Oriental), the
Pacific Region, and the Caribbean territories. In Colombia there are a multitude of habitats for plants to oc-
cupy and this has lead to its high biodiversity, second only to Brazil. The Poaceae are a diverse family that
occupies a myriad of terrestrial habitats in Colombia. According to our inventories, the Poaceae is represented
in Colombia by 811 species [a single genus (Agrostopoa Davidse, Soreng, and P.M. Peterson; Davidse et al.
2008) and 40 species are endemic to Colombia] that are currently placed in 157 genera. The grass family is
the third largest family of vascular plants in Colombia behind the Orchidaceae and Asteraceae.
J. Bot. Res. Inst. Texas 2(2): 875 — 916. 2008
876 tani i f Texas 2(2)
In Colombia the Panicoideae (350 species/63 genera), Pooideae (148 species/36 genera), and Chlori-
doideae (98 species/25 genera) are the richest grass subfamilies. The largest genera are Paspalum (83 spe-
cies), Panicum (48), Festuca (33), Chusquea (25), Eragrostis (25), Calamagrostis (23), Axonopus (22), Digitaria
(19), Setaria (15), Aulonemia (12, Judziewicz, per. comm.), Lasiacis (12), Agrostis (11), Muhlenbergia (10), and
Sporobolus (10). The species within these subfamilies exhibit a specific distribution that is correlated with
the altitude and/or with the rainfall patterns. Therefore, the Panicoideae are more diverse in the lowlands
and wet areas where precipitation is high, the Pooideae are best represented in the high elevation páramos,
and the Chloridoideae are best represented in dry areas at mid-levels of elevation.
Two centers of species diversity have been recognized in Colombia: the Llanos Orientales (Orinoco
Region) and the páramos Nolae Cañas 200r; ine Orinoco Region is donec inns C, e and
neotropical elements, whereas the p are l with C, grasses and I
genera. Despite different analyses, ER results indicate there is no clear pattern in regards to Ein in
these natural areas (Giraldo-Cañas 2007).
As a result of our studies, 11 new species and one new genus have been described, and 18 genera and
82 species have been recorded for the first time in Colombia (Giraldo-Cañas 2007). This serves to pe
J:
l country.
the tremendous variety and botanical interest present in Colombia, a floristically
The taxonomic knowledge of this family in Colombia is incomplete, and it is still possible to encounter novel
species and many new records for this country. There have been very few taxonomic studies investigating
Colombian grasses integrating the newest classifications based on molecular evidence. The data also indi-
cate a need to increase field exploration, primarily in the Llanos Orientales, Sierra Nevada de Santa Marta,
Nariño, and the Caribbean Region.
We report 25 species and one nontypical subspecies of Eragrostis for the Colombian Flora, three of the
species are cited for the first time for Colombia: Eragrostis mokensis Pilg., E. nigricans (Kunth) Steud., and E.
rufescens Schrad. ex Schult. Our taxonomic treatment contains a key for determining species, descriptions, dis-
tributions, specimens examined, illustrations, and synonymies. Only synonyms used frequently in Colombian
literature and those of South American origin are given. For further synonymy see Peterson et al. (2001), Peter-
son & Boechat (2001), and Soreng et al. 2008. This study is based on the examination of herbarium specimens
from COAH, CAUP, COL, HUA, MO, PSO, and US, including the type specimens of most species studied. The
present study contributes knowledge of the Colombian grasses in collaboration with the Biodiversity Program
and Colombian Flora Inventories.
TAXONOMIC TREATMENT
e Wolf, Gen. Pl. 23. 1776. Te: Eragrostis minor Host, Icon. Descr. Gram. Austriac. 4:15. 1809 (1ecrorvre, designated by
Ross, Ácta Bot. Neerl. 15:157. 1966).
Plants annual or perennial; usually synoecious, sometimes dioecious; caespitose, stoloniferous, or rhizoma-
tous; flowering culms (2-)5-150 cm tall, not woody, erect, decumbent, or geniculate, sometimes rooting at the
lower nodes, simple or branched; i d lid or hollow; bud initiation intravaginal, rarely extravaginal.
Leaf sheaths open, often with tufts of hairs at the apices, hairs 0.3-9 mm long; ligules usually membranous
and ciliolate or ciliate, cilia sometimes longer than the membranous DIESE Madam a hairs or membra-
nous and non-ciliate; blades flat, folded, or involute. Infl illary, simple
panicles, open to contracted or spike-like, terminal panicles usually a the upper leaves; pulvini in
the axils of the primary branches glabrous or hairy; branches not spike-like, not disarticulating. Spikelets
1-20 mm long, 0.5-4.5 mm wide, laterally compressed, with 2 to q florets; disarticulation below the fertile
florets, sometimes also below the glumes, acropetal with decid d lemmas but persistent paleas,
or basipetally with the glumes often persistent and the florets auly falling intact; glumes usually shorter
than the adjacent lemmas, usually 1(3)-nerved, not lobed, apices obtuse to acute, unawned; calluses glabrous
or sparsely pubescent; lemmas usually glabrous, obtuse to acute, 3(5)-nerved, usually keeled, unawned or
mucronate; paleas shorter than the lemmas, longitudinally bowed-out by the caryopses, 2-keeled, keels usually
Peterson and Giraldo-Cañas, Eragrostis in Colombia 877
short or long ciliate, intercostal region membranous or hyaline; stamens 2 or 3; ovaries glabrous; styles free
to the bases. Cleistogamous spikelets occasionally present, sometimes on the axillary panicles, sometimes
on the terminal panicles. Caryopses variously shaped; hilum short; embryo with an epiblast, scutellar tail,
and elongated mesocotyl internode (formula P+PF), endosperm hard. Chromosome base number of x = 10,
2n = 20, 40, 50, 60, 80, ae and dus 2, 4, 5, 6, 8, and 10 ploid (Watson & Dallwitz 2008).
I examined anatomically
Anatomical and biochemi ll species of Eragrostis that |
exhibit “Kranz” or C, leaf anatomy m alien Pilg. from South Africa, which exhibits C, leaf anatomy;
see Ellis (1984)], aud species have either chloridoid bicellular microhairs (with the broad, ies terminal
cell the same thickness as the basal cell) or panicoid bicellular microhairs (with a long, thin-walled terminal
cell) [Amarasinghe & Watson 1990]. Apparently, three C, biochemical types exist in Eragrostis: NAD-ME
(nicotinamide adenine dinucleotide co-factor malic enzyme), PCK (phosphoenolpyruvate carboxykinase),
and intermediates (Prendergast et al. 1986).
Taxonomic position.—The genus Eragrostis is placed in tribe Eragrostideae, which in the New World
includes three small subtribes (Cotteneinae, Uniolinae, and Eragrostidinae) that diverge as a clade at the
base of the chloridoids (Peterson et al. 2007). Character trends in the Eragrostideae include spikelets with
many florets, lemmas with 3 to 13-nerves, and many species adapted to xeric habitats. In the New World
the Eragrostidinae includes two native genera: Eragrostis with 111 species (86 native to New World); and
Steirachne Ekman with two species (P 2003; Peterson et al. 2007). TRE Eragrostidinae is characterized
by hairy or glabrous culm nodes, hairy or glabrous rachillas, entire that are awnless, mucronate,
or short-awned (only in the latter two genera), glabrous or scabrous mirae that are (1)3(5)-nerved, and
short basal microhair cells (15-75 pm) on the abaxial epidermis of the leaf blade. Morphologically, the two
species of Steirachne [S. parbat aD ) Renvoize and S. diandra Ekman] are hardly separable from Eragrostis
acutiflora, as all h te or subaristate, 3-nerved lemmas (Peterson et al. 2007). However,
the lemmas of the two species of Steirachne have more pronounced setulose lemmas and elongate caryopsis
(Renvoize 1984).
Monophyly of Eragrostis.—Based on nuclear and plastid DNA sequences, Ingram and Doyle (2003,
2004, 2007) tested the monophyly of Eragrostis and found that, with the inclusion of Acamptoclados Nash (E.
sessilispica Buckley), Diandrochloa De Winter, Neeragrostis Bush, and Pogonarthria Stapf, the genus is indeed
monophyletic. However, only 37 species of Eragrostis were included in their analysis, so any infrageneric
interpretations were beyond the scope of their work.
Infrageneric classification.—Based on spikelet disarticulation, Clayton (1974) and Clayton and Ren-
voize (1986) have arrived at a first approximation to natural groups and have presented a key to sections
Psilantha (K. Koch) Tzvelev, Eragrostis, Lappula Stapf, and Platystachya Benth. In most taxa native to the
Western Hemisphere, disarticulation of the spikelet is acropetal (florets first maturing at the base) and the
lemmas fall with the caryopses, leaving paleas attached to the rachilla. The common mode of disarticulation
for introduced species from Africa and Asia is basipetally, where the florets near the apex mature first.
Van den Borre and Watson (1994) investigated 53 species of Eragrostis and found that anatomical
characters, among others, support the recognition of two distinct groups: Eragrostis subgen. Eragrostis and
Eragrostis subgen. Caesiae Van den Borre & L. Watson. The most comprehensive attempt so far is Lazarides'
(1997) treatment of the Australian Eragrostis, in which he recognized six groups primarily based on spikelet
disarticulation. Lazarides (1997) correlates his classification with Van den Borre and Watson (1994), who
recognized subgenus Eragrostis and subgenus Psilantha, and with Amarasinghe and Watson (1990), who
investigated microhair morphaleey WE ed. de S (1998) informal treatment of Eragrostis for the
Flora of Zambesiaca is al pecies groups based on panicle, lemma,
and palea morphology in adios to spikelet disarticulation.
Based on caryopsis morphology, Boechat and Longhi-Wagner (2003) placed 49 of the 53 species of
Eragrostis that occur in Brazil into the following six groups: smooth-walled (six species), medianly reticulate
(14 species), roughly reticulate (10 species), finely late (nine species), alveolate (six species), and striate
878 Journal of tl Botanical R h Institut f Texas 2(2)
(four species). Overall morphological features led Peterson and Valdés Reyna (2005) to recognize the fol-
lowing four hypothesized lineages within 26 species of Eragrostis from northeastern Mexico: an Old World
group, the E. intermedia Hitchc. complex, the E. pectinacea complex, and the E. spectabilis (Pursh) Steud.-E.
secundiflora J. Presl group.
In addition, recent sy f Eragrostis from Argentina (Nicora 1998), Australia (Lazarides
1997), Bénin (Houinato et al. 2000), Bolivia (Renvoize 1998), Beal (Boechat & Longhi-Wagner 2000, 2001),
Congo (Kami 1993), Costa Rica (Pohl 1980), Cóte-d'Ivoire (Poilecot 1995), Ecuador (Peterson 2001), France
(Portal 2002), Guianas (Judziewicz 1991), Malesia (Veldkamp 2002), Mesoamerica (Davidse 1994), Mexico
(Beetle et al. 1991; Peterson & Valdés Reyna 2005), Niger (Poilecot 1999), Peru (Tovar 1993; Peterson &
Sánchez Vega 2007), the United States and Canada (Peterson 2003), Venezuela (Graterol et al. 1989), and
Zambesiaca (Cope 1998) have given us a good understanding of the species limits and their distribution.
However, there is no definitive treatment of the infrageneric classification of the entire genus.
Etymology.—The origin of the name is somewhat obscure. Nathaniel M. von Wolf (1776), who first
named Eragrostis, made no statement concerning the origin of its name. Clifford (1996) provides three pos-
sible derivations: from “eros” (love), and “Agrostis” (the Greek name for an indeterminate herb); from the
Greek “er” (early) and “agrostis” (wild), referring to the fact that some species of Eragrostis are early invaders
of arable land; or the Greek “eri-“ (a prefix meaning “very” or “much”, suggesting that the name means
many-flowered “Agrostis.” Watson and Dallwitz (2008) indicate that the derivation of Eragrostis is “from the
Greek ‘eros’ (love) or ‘era’ (earth) and 'agrostis' (a grass), a alluding to the characteristic, earthy (hu-
man) female aroma of the orina of many species.”
Ecology hic distribution.—The 25 species of Eragrostis in Colombia are widely distributed
among the coast ad icon coast), Andean mountains, llanos, forest regions, cultivated fields, and
city sidewalks. The annual species are more conspicuous along the sandy plain of the coast, the interior
valleys of the western slopes, and in the dry Andean valleys at lower elevations. Along the north coast, E.
tenella, E. ciliaris, E. cilianensis, E. hypnoides, and E. viscosa form part of the xerophytic vegetation.
The mountains species of Eragrostis are an integral component of natural, mid to high elevation eco-
systems. Eragrostis nigricans and E. pilosa are usually found growing along margins of cultivated fields while
many other species inhabit the edges of roads and trails. Eragrostis lurida, and E. pastoensis, are principally
found in the altiplano, and disappear in the páramos. In the mountainous region of central and southern
Colombia there is a greater number of perennial, native species of Eragrostis. Eragrostis hypnoides is the only
species commonly found along the coast and in the Amazonian forest in areas that are periodically flooded,
i.e., margins of rivers.
Caryopsis morphology.—The caryopsis contains many hological features that are important aids
in the identification of species and this information can be ue to infer hypothetical relationships among
the grasses (Boechat & Longhi-Wagner 2003; Colbry 1957; Peterson et al. 2007; Terrell & Peterson 1995). In
Eragrostis, grains can be terete, subterete, rectangular or trigonous in cross-section, and are sometimes com-
pressed either dorsally (on a plane with the embryo) or laterally (Peterson & Sánchez Vega 2007). The shape of
the grain can vary from spherical to much longer that broad (ellipsoid, obovoid, ovoid, rectangular-prismatic,
etc.). The embryo is located on the dorsal (abaxial) surface of the grain and the hilum is the tiny scare left
from the attachment of the funiculus found near the base on the ventral surface. The ventral (adaxial) surface
can be rounded, flattened or sometimes have a sulcus or groove running longitudinally along the body. The
surface of the grain can be smooth to variously sculptured and is often striate to reticulate. In the grasses
the surface or pericarp of the grain is almost always adnate, i.e., a true caryopsis; however, a few species of
Eragrostis (see E. japonica) can have loose pericarps that shed when the grain is moistened. Color of the grain
can vary from light brownish or whitish to reddish brown, and the grains can be translucent to opaque.
The most common caryopsis type found in Colombian Eragrostis is the rectangular-prismatic to ir-
regularly triangular, laterally-flattened, striate, reticulate-walled, and ventrally-grooved grain. This is found
predominately in species that occupy the altiplano such as E. lurida and E. pastoensis, as well as in E. lugens,
Peterson and Giraldo-Cañas, Eragrostis in Colombia 879
E. mexicana, E. nigricans, and E. tenuifolia (of African origins). Eragrostis pectinacea and E. pilosa are similar
to the last group, although these two species have grains that are striate only and are rectangular in cross-
section, so they are at least flattened ventrally but do not have a groove. Eragrostis maypurensis is unique in
possessing rhomboid reticulations without striations. Eragrostis tenella, E. cilianensis, E. ciliaris, E. curvula, E.
hypnoides, E. japonica, and E. maypurensis are all circular or elliptical to ovate/obovate in cross-section. Eragrostis
tenella, E. ciliaris, E. curvula, and E. japonica are striate or smooth with no evidence of reticulations. Eragrostis
ciliaris, E. japonica, E. pilosa, and E. curvula all have somewhat dorsally flattened caryopses. Eragrostis curvula
in addition to being strongly dorsally-flattened, can sometimes possess a shallow and broad ventral groove.
Economic importance.—The following Colombian species of Eragrostis are weedy around the World:
E. cilianensis, E. curvula, E. japonica, E. mexicana, E. pectinacea, E. pilosa, E. tenella, E. tenuifolia, and E. viscosa.
Eragrostis tef (Zucc.) Trotter and E. curvula are cultivated, and E. superba Peyr., a drought resistant species, is
used primarily for reseeding denuded land (Watson & Dallwitz 2008). Mi ef is a staple cereal in Ethiopia,
which is potentially of wide interest (Watson & Dallwitz 2008). Erag ili E. ciliaris, E. curvula, E. pas-
toensis, E pilosa and E.t if li 1 I Colombia (Wat m. 2008; pers obs.)
KEY TO THE SPECIES OF ERAGROSTIS IN COLOMBIA
p dbi moe or Dod without innovatio
. Sta els 0.1—1 mm long, mostly shorter than d spikelets.
3; PI ce Qiu ides the cilia 0.2—1 mm long; spikelets 1.8—3.3 mm long, 6-1 adi. lower
glumes 0.7—1.2 mm long; upper glume = .6 mm long; E uh 13 Mu m long E. ciliaris var. ciliaris
3. Paleas keels smooth to scabrous, tl ities less than 0.2 lets 6-15
10—43-flowered; lower glumes 1.4—2.5 mm long; upper glume 1.5— 26mm long; emna m
long 21. E. rufescens
2. Stamens 3; pedicels usually 1—4(—7) mm long, as M or longer than the spikelets.
4. is keels prominently ciliate, the cilia 0.2—1 m
. Plants viscid with particles of soil adhering to i sticky areas along the sheaths, culms, and blades;
v HN
spikelets (2—)2.5—5.5 mm long 25. E. viscosa
5. Plants not viscid and without particles of soil adhering to vegetative portions.
6. Paleas orbicular, winged 14. E. mokensis
6. Paleas obtuse, acute to truncate, not winged.
7. Spikelets (1—)1.5—2.2 mm long 23. E. tenella
7. Spikelets 5-20 mm long 4. E. cilianensis
4. Paleas smooth to scabrous, the vi less than 0.2 mm long
8. Plants mat-forming; panicles 1—3.5 cm long; erect portion of culms (2-)5-20 cm, tall, the per
portion prostrate and rooting at bu odes 7. E. hypnoides
8. Plants not forming mats; panicles 3—55 cm long; culms 15—130 cm tall, not prostrate or m at
the lower nodes
9. Ligules pennae glabrous 9. E. japonica
9. "E ciliate with a row or Hry wile d
with 1—3 the keels; spikelets 2-4 mm wide; disarticulation of
entire florets from Ber stónt rachilla 4. E. cilianensis
10. Lemmas without crateriform glands; spikelets 20 2.523) mm wide; disarticulation of the
e only, palea and rachilla usually persisten
1. Lemmas acuminate, the apices recurved, ae pulvini (axils of the up
pane) densely Pese. Me hairs up to 4 mm long 2. E. maypurensis
1 recurved, I pulvini II ? I
“to sparsely ciliate, the hairs if present ee uu 1.5 mm long.
12. Caryopses with a shallow p adaxial
13. Spikelets not end in Rude Sr e qe MC 15)-flowered, (4-)5—10(-11) mm
long; pedicels appressed to narrowly divergent, stiff 13. E. mexicana
13. Spikelets arranged in glomerules, 2-4(-5)-flowered, 2.6—3.8 mm long; an
spreading, divaricate and stout 5. E. nigricans
12. o without an adaxial groov
ps MM 5—1.5 n long, ar least 1/2 as long as the lowest lemmas; spikelets
.2—2.5 mm i or paired at the lowest 2 nodes; lem-
mas with moderately conspicuous lateral veins 17. E. pectinacea var. pectinacea
J lof the Botanical R h Institute of Texas 2(2)
880
14. Lowerglumes 0.3—0. se O, 8) m Em long, usually less than D aslongasthelowestlem-
mas; spikelets 0.6—1. y whorled at the lowest 2
Nees’ ans with i inconspicuous lateral veins 18. E. pilosa subsp. pilosa
1. Plants perennial, i , forming innovations at the basal nodes.
2. E. airoides
15. Spikelets 1.3—2 mm long with 1 or 2, rarely 3 florets
15. Spikeiets 2-23 mm long with 2 to many florets.
16. Stamens 3; caryopses usually with a shallow or deep adaxial groove.
17. Glumes not keeled, very unequal, the lower 0.2—0.6 mm long
17. Glumes keeled, equal to subequal, the lower 0.6-2.6 mm long.
18. Lemmas with lateral veins conspicuous.
24. E. tenuifolia
19. Caryopses 1—1.7 mm long; upper glumes 2-3 mm long 6. E. curvula
19. Cananea RU mmi g; Up; fee 1-2 mm cua
20. P y oranches not floriferous near! lary branches com
osed of loosely overlapping spikelets; pedicels 1.4-7 mm long 10. E. lugens
20. Panicles with the primary branches floriferous near base; secondary branches con-
densed into tightly glomerate lobes of spikelets; pedicels 0.1—1 mm long 11. E. lurida
subsp. lurida
18. Lemmas with n veins O U
21. Secondary | MI ] I i j 14 li Is 0.5 E 5)
m long 6. E. pastoensis
21. Secondary | b | ith spikelets diverging 50—90°from the main axis; Sate
2-)4-16 mm long.
22. Lemmas 1.2-1.8 mm long; blades densely hairy on adaxial surface; anthers 0.3—0.5
19. E. polytricha
mm long
22. Lemmas (1.6—)1.8—2.3 mm long; blades glab to sparsely hairy daxial :
nthers 0.5—0.8 mm long 8. E. intermedia
16. Stamens 2; cayopses Bn an eden goe
23. Le a apices a te tat ini with hairs
24. pin 24-5 mm Nes Ne: 1.7-4 mm long; lemmas 2— ón mm long 22. E. secundiflora
subsp. secundiflora
24. Spikelets 1-3 mm wide; glumes 1—2.8 mm long; lemmas 2—2.6 mm long.
25. Lemma apex recurved, yellowish-orange to greenish; spikelets 7-30 mm long; pedicels
0.3-1 mm long, appressed to the branches; upper glumes 1.5—2.8 mm long ___ 12. E. maypurensis
25. Lemma apex straight, not recurved, greenish to purplish; spikelets 5—7(—10) mm long;
pedicels 1—5 mm long, diverging 30—70° from the branch axis; upper glumes 1.4—2 mm
long 1. E. acutiflora
23. Lemma apices acute, ada slightly narrowed but never acuminate-attenuate; pulvini
glabrous, rarely with a airs.
26. Spikelets 2.4-5 mm dE glumes 1.7—4 mm long; lemmas 2-6 mm long 22. E. secundiflora
subsp. secundiflora
26. Spikelets 0.7—2(-2.2) mm wide; glumes 1—1.8(-2) mm long; lemmas 1.5-2.2 mm lon
27. Primary branches with spikelets congested or clustered near the base; spikelets 0.7—1.4
mm wide; anthers 0.2—0.3 mm long; caryopses flattened ventrall 20. E. prolifera
27. al branches naked near base, without a cluster of spikelets; spikelets 1.3—2(-2.2
m wide; anthers 0.4-0.6 mm long; caryopses rounded, not flattened ventrally 3. E. bahiensis
1. Eragrostis acutiflora (Kunth) Nees, Fl. Bras. E Em 2:501-502. 1829. (Fig. 1, A-C). Poa acutiflora Kunth,
Nov. Gen. Sp. 1:161. 1815 (1816). Tree: COLOMBIA fagdal E Humboldt & A. Bonpland 1603 (HoLoTYPE: Pl; ISOTYPES:
B-W, P, US-2891479 fragm. ex PT).
Caespitose perennials. Culms 20-65 cm tall, erect to geniculate spreading, glabrous and shiny below the
nodes. Leaf sheaths 1/3 to about as long as the internodes, glabrous, ciliate at the summit; ligules 0.1-0.3
mm long, ciliolate-membranous; blades 10-20(-25) cm long, 1.5-4.2 mm wide, flat, occasionally involute,
glabrous below, scaberulous above and sparingly ciliate near base. Panicles 8-32 cm long, 2-14 cm wide,
open to somewhat contracted; the ascending primary branches 1-10 cm long, somewhat densely flowered,
spreading 20-80? from the rachises; pulvini in axils of primary branches pilose; pedicels 1-5 mm long,
erect, spreading 30-70? from the branch axis, scaberulous. Spikelets 5-7(-10) mm long, (1-)1.2-1.8(-2.2)
881
Peterson and Giraldo-Cañas, Eragrostis in Colombia
NS
CIS
ESA A
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ee
Sy a
pe
4291, G). D.
AA IHynhon O CAL
J
4
(N.C. Fassett 25306). A Habit. B. Spikelet C. Caryopsis, dorsal view. El
:6
J
Fic. 1. El
Culm. E. Inflorescence. F. Spikelet. G. Caryopsis, dorsal view.
= 7 I ^f tha Ratanjral D h Institut f Texas 2(2)
mm wide, 7-14(-21)-flowered, narrowly elliptical, acute at both ends, strongly compressed, greenish straw-
colored towards the center to reddish-purple tinged near the margins; disarticulation entire floret above the
glumes or the lemmas falling individually leaving the paleas on the rachilla; glumes 1-2 mm long, narrowly
lanceolate with scaberulous keels; lower glumes 1-1.4 mm PA upper glumes 1.4 mm a Mad pou
1.4-2 mm long; lemmas 2-2.6 mm long, lanceolate to broadly ovate, chartaceous, keeled,
keel, lateral nerves evident; apex acuminate to attenuate or subaristate, scaberulous, often reddish- mu
paleas 1.5-2.1 mm long, bowed out below, hyaline, scaberulous along keels; apex acute; stamens 2, anthers
0.2-0.3 mm long, reddish-brown. Caryopses 0.5-0.9 mm long, obovoid to ellipsoid, finely longitudinally
striate, light brownish to reddish-brown. 2n - 40.
Distribution and habitat.—Eragrostis acutiflora ranges from México and the Caribbean through Central
America to South America where it has been reported from Bolivia, Brazil, Ecuador, French Guiana, Guyana,
Surinam, and Venezuela. It can be found growing in moist savannas, open disturbed areas, and sandy to
gravelly roadsides; 0-2000 m.
Specimens examined. COLOMBIA. Amazonas: Corregimiento departamental La Pedrera, río Caquetá, alrededores de la quebrada La
Tonina, 200—300 m, 18 Jul 1997, D. Giraldo-Cañas 2703 (COAH, COL). Antioquia: Puerto Berrío, ca. 125 m, 11 Jan 1931, W.A. Archer
1392 (US); Mutatá, 11 km Mutatá-Dabeiba, 250 m, J. Betancur et al. 378 (COL, HUA, MO); Andes, aeropuerto, 1200 m, 17 Sep 1986, R.W.
Pohl & J. Betancur 15521 (HUA, MO, US); Mun. San Carlos, río San Carlos, 920 m, 13 Oct 1981, C.I. Orozco 685 (COL). Arauca: Mun.
Arauca, inmediaciones de las instalaciones de la Universidad Nacional de Colombia, hacienda El Cairo, carretera Arauca-Tame, km 9,
200-300 m, 13 Jun d D. a Cañas Mn 3510 (COL). Caquetá: Mun. La Montañita, corregimiento El Santuario, vereda Las
Iglesias El Ceilán, 280 m, 27-30 Jun 2005, D. Giraldo-Cañas 3936 (COAH, COL); Mun. Florencia,
Barrio Villa Natalia, atas de pavimentos y andenes, 280 m, 27-30 Jun 2005, D. Giraldo-Cañas 3953 (COL); Mun. Florencia, carretera
Florencia-Neiva, río Hacha, a " un del primer puente, 280 m, 27-30 Jun 2005, D. Giraldo-Cañas 3956, 3958 ie COL). Casan-
are: Mun. El Yop b lables, entre el nuevo hospital y la Brigada del Ejército Nacional, 400 m, 3 Nov 2007, D.
Giraldo-Cañas 2 4155 (COL); Playones del río Upia, carretera a Villanueva, 300 m, 18 Mar 1986, J.L. dude s 5907 (COL).
Cauca: Río Patía, 1000 m, 2 May 1935, H. García-Barriga 4507 (COL). Chocó: Hoya del río Atrato, Beté, 50-60 m, 5 Apr 1982, E. Forero
8901 (COL). Córdoba: San Benito, 5 km carretera a San Juan, 35 m, 17 Jul 1973, J.M. Idrobo 6664 (COL). Cundihamurca Fusagasugá,
1680 m, 26 Mar 1961, J.M. Idrobo 4507 (COL); Entre Paratebueno y Maya, 19 Jun 1989, F.O. Zuloaga 4100 (COL, MO). Guaviare: Mun.
San José del Guaviare, trocha Nuevo Tolima, en cercanías del Batallón José Joaquín París, 250 m, Mar 1996, D. Giraldo-Cañas & R.
López 2574 ae COL, o) oe Mun. El a pnis La Yaguilda, 700 m, Sep 1986, J.L. Fernández Alonso 6840 (COL). La
dable, 4, Uribia rumbo a Maicao, 13 Feb 1963, C. Saravia 2229-A (COL, note: Saravia
2229-A at US is E. ciliaris), 19 Oct 1963, C. Saravia 2892 on Magdalena: Valle del río Cesar, cerca de Los Venados, 60 m, 30 Sep
1961, A. Dugand 5810 (COL). Meta: Mun. Villavicencio, carretera Villavicencio-Aeropuerto, piedemonte de la Cordillera Oriental, sitio
La Arenera, 2 km del puente sobre el río Guatiquía, 400 m, 10 Nov 2002, D. Giraldo-Cañas 3346 (COAH, COL); along Río Guatiquía,
near Villavicencio, ca. 500 m, 18-19 Mar 1939, E.P. Killip 34410 (US); Carimagua, NW of the research station, ca. 350 m, 19 Sep 1992,
S. A. Renvoize 5422 (COL, US). Santander: region about Landázuri, 70 km N of Vélez, ca. 700 m, 9 Jun 4, N. C. Fassett 25306 (COL).
Tolima: Ibagué, planta eléctrica de Mirolindo, 1200 m, 12 Mar 1965, R. Echeverry 1189 (COL). Valle del Cauca: between Uribe and
Sevilla, 1100 m, 2 Nov 1983, J. R.I. Wood 4083 (COL).
Guajira:
2. Eragrostis airoides Nee E pra eu Pl. 2: 509-510. 1829. (Fig. 2, A-C). Aira brasiliensis Raddi, Agrostogr.
Bras. 36. 1823. S Hack., Bull. Herb. Boissier, sér. 2, 4(3):278. 1904. à inu brasiliensis (Raddi) Herter,
Revista ade Bot. 6(5-6):145. 1940. Tyre: BRAZIL. Rio de Janeiro, G. Raddi s.n.(HOLOTYPE: PI; 1 ).
Airopsis Ven Griseb., Abh. Kónigl. Ges. Wiss. Göttingen 19:252. 1874. Tree: ARGENTINA. Tucumán: in monte Cuesta de Berico,
tz & G. Hieronymus 842 (uoLorvre: GOET, rsorvees: BA-38791!, US-76315 fragm.!).
ds inh Ekman, Ark. Bot. 11(4):42, t. 4, f. 1. 1912. Tyre: ARGENTINA. Misiones: Posadas, Bonpland Trans "Mitires Chico,” 20
n 1908, E.L. Ekman 714 (HoLoTYPE: S; isorvPES: BAA-1100 fragm.!, CORD!, SI!, US-602662!, US-77381 fragm.!).
Caespitose perennials with innovations, without rhizomes, not glandular. Culms 30-110 cm tall, erect, gla-
brous below the nodes. Leaf sheaths glabrous or pilose, hairs to 5 mm long; ligules 0.1-0.2 mm long; blades
8-22 mm long, (1-)2-4(-5) mm wide, flat to folded, glabrous abaxially, scabridulous adaxially. Panicles
18-70 cm long, 3-25 cm wide, diffuse, ovate; primary branches 4-20 cm, appressed or diverging 10—70?
from the rachises, naked basally; pulvini glabrous; pedicels 2.4-11 mm long, divergent. Spikelets 1.3-2
mm long, 0.8-1.8 mm wide, ovate to lanceolate, plumbeous, with 1 or 2, rarely 3 florets; disarticulation
acropetal, in the rachilla below the florets, glumes deciduous; rachilla prolonged above the terminal floret;
glumes lanceolate to ovate, membranous; lower glumes 0.8-1 mm long; upper glumes 1.1-1.4 mm long;
Peterson and Giraldo-Cañas, Eragrostis in Colombia
883
D. García 10, ©. A. Inflorescence. B. Spikelet. € Caryopsis, ventral view.
r.
Fig 2, E j iroides UJ.
D, E, H, 1). D. Habit. E. Inflorescence. F. Spikelet, G. Floret. H. Caryopsis, lateral view. I. Caryopsis, ve
ntral view.
884 j | of the Botanical R h Institute of Texas 2(2)
lemmas 0.8-1.2 mm long, ovate, membranous, plumbeous, keels and lateral veins inconspicuous, apices
obtuse; paleas 0.8-1.2 mm long, membranous, bases not projecting beyond the lemmas, apices obtuse; sta-
mens 3, anthers 0.3-0.5 mm long, purplish. Caryopses 0.4—0.5 mm long, ovoid, reticulate, reddish-brown.
2n = 36 (Davidse, pers. comm.).
Distribution and habitat.—Eragrostis airoides is a South American species known to occur in Argentina,
Bolivia, Brazil, Paraguay, Uruguay, Venezuela, the Caribbean, and introduced Brazos Co., Texas; it is found
in open grasslands, savannas, and cerrados (Boechat & Longhi-Wagner 2001).
Comments.—It is an enigmatic species, often treated as Sporobolus brasiliensis (Raddi) Hack, which it
resembles in its chromosome base number of x - 9 and caryopsis morphology, but its frequent possession of
- with more than one floret and its mode of spikelet disarticulation argue for its retention in Eragrostis.
MBIA. Cauca: Chisquio, ca. 1700 m, 4 Apr 1940, E. Asplund 10776 (US); Río Sucio to Río Piedras, W of
iocum 1500-1800 m, 3 bu 1922, F.W. Pennell & E.P. Killip 8190 (US); Loma de Bichiquí, near Toribío, Río Páez Basin, Tierradentro,
2000 m, Feb 1906, H. Pittier 1473 (US). Norte de Santander: Abrego, 1300 m, 25 Feb 1943, J.B. García & Cabrales 10 (COL). Valle del
Cauca: Pavas, Cordillera Occidental, 1500—1700 m, E.P. Killip 11636 (US). Unknown department: J.C. Mutis 5542 (US).
3. Eragrostis bahiensis Schrad. ex Schult., Mant. 2:318. 1824. (Fig. 1, D-G). Eragrostis pilosa var. bahiensis (Schrad.
ex Schult.) Kuntze, Revis. Gen. Pl. 3(2,2):353. 1898. Tree: BRAZIL: Maximilian Neowidensis s.n. (aoLotyre: LE).
Eragrostis expansa Link, Hort. Berol. 1:190. 1827. Tree: URUGUAY: Montevideo, E Sellow s.n. (sotyre: US-2850751 fragm.!).
Poa microstachya Link, Hort. Berol. 1:185. 1827. Eragrostis psammodes var. microstachya (Link) Doll, Fl. Bras. 2(3):153. 1878. Eragrostis
lios (Link) Link, Hort. Berol. 2:294. 1933. Tree: URUGUAY: Montevideo, E Sellow s.n. (HoLoTYPE: B; isorvees: BAA-989 fragm.
!, US-2850751!).
E firma Trin., Mém. Acad. Imp. Sci. Saint-Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 4,2(1):74. 1836. Tyre: BRAZIL.
Sáo Pauro: inter plantas a St. Paulo (noLorvPE: LE-TRIN-2342.011; isotyre: US-2891470 fragm.!).
Eragrostis blepharophylla Jedwabn., Bot. Arch. 5(3-4):197. 1924. BRAZIL: E Sellow 3688 (isorvee: US-2891454 fragm.!).
Eragrostis macra Jedwabn., Bot. Arch. 5(3-4):200. 1924. Tyre: BRAZIL: Nov 1887, A.EM. Glaziou 16624 (Lecrorvee: US-1280047! designated
echat & Longhi-Wagner, Iheringia, Bot. 55:147. 2001; isoLecroryrEs: C, US-2899241, US-1280048!
Eragrostis bahiensis forma riparia Burkart, Fl. Il. Entre Ríos 2:189. f.66. 1969. Tree: ARGENTINA. Entre Ríos: Concepción del Uruguay,
Isla Almirón Chico, Burkhart & Crespo 22874 (uotorvre: SI; isotype: BAA-1010)).
Caespitose perennials with innovations, without rhizomes, not glandular. Culms 25-95(-110) cm tall, erect,
glabrous. Leaf sheaths glabrous, summits hairy, hairs 13 mm; ligules 0.2-0.4 mm long; blades (8)12-40 cm
long, 2-5 mm wide, flat to involute, abaxial surfaces glabrous, adaxial surfaces scabridulous and glabrous
or long ciliate basally. Panicles 15-30(45) cm long, (48-17 cm wide, narrowly ovate, open to contracted;
primary branches 5-15 cm long, diverging 20—90? from the rachises, often capillary, usually naked ba-
sally; pulvini glabrous; pedicels 0.3-6 mm long, mostly appressed, scabridulous, always shorter than the
spikelets. Spikelets 6-15(18) mm long, 1.3-2(2.2) mm wide, narrowly lanceolate, plumbeous, occasionally
with a reddish-purple tinge, with 8-30(40) florets; disarticulation usually in the rachilla below the florets,
occasionally the lemmas falling separately, leaving the paleas on the rachilla; glumes lanceolate to ovate,
membranous to subhyaline, keeled; lower glumes 1-1.4 mm long; upper glumes 1.4—1.7 mm long; lemmas
1.5-2.2 mm, broadly ovate, leathery, scabridulous, lateral veins evident, apices acute; paleas 14-2.1 mm
long, hyaline, bases not projecting beyond the lemmas, keels scabridulous, apices acute to obtuse; stamens
2, anthers 0.4-0.6 mm long, reddish-purple. Caryopses 0.6-0.8 mm long, obovoid to ellipsoid, terete,
somewhat striate, reddish-brown. 2n = unknown.
Distribution and habitat.—Eragrostis bahiensis grows in sandy soils near river banks, lake shores, and
roadsides, at 0—1500 (-1850) m. Its range extends south from the Gulf Coast of the United States through
México to Peru, Bolivia, Paraguay, and Argentina.
d. COLOMBIA. Antioquia: Mun. Cocorná, en La Piñuela, carretera Medellín- "ORO intersección con De car-
uns San Francisco, 1000 m, 27 Feb 2005, D. Giraldo-Cañas 3870 (COL). Arauca: Mun. Árauca
de la Universidad Nacional de Colombia, ed - Cairo, carretera Arauca-Tame, km 9, 200—300 m, 13 Jun 2003, D. Giraldo-Cañas
3498 (COD). Cauca: Mun. Popayán iudad, Villa B Viento, 1850 m, 20 May 2001, B.R. Ramírez 14163 (CAUP, COL).
Chocó: Mun. Quibdó, en un pa del Barrio Medrano de la Universidad Tecnológica del Chocó, 90 m, 6 Nov
2005, D. Giraldo-Cañas 3974 (COL, HUA); Mun. Quibdó, carretera Quibdó-Guayabal, 23 Feb 1985, J. Espina 1474 (COL, MO); Hoya del
Peterson and Giraldo-Cañas, Eragrostis in Colombia 885
río San Juan, río Tamaná, afluente del San Juan, debajo de Santa Rosa, ca. 150 m, 10 Apr 1979, E. Forero et al. 4974, 4977 (COL, MO);
Quibdó Airport, 50 m, 24 Oct 1985, J.R.I. Wood 5117 (COL). Meta: Mun. Villavicencio, carretera Villavicencio-Aeropuerto, piedemonte
de la cordillera Oriental, sitio La Arenera, ca. 2 km del puente sobre el río Guatiquía, ca. 400 m, 10-14 Nov 2002, D. Giraldo-Cañas
3344 (COAH, COL, HUA); along road to Hac. Rubiales, E of Puerto Gaitán, 90 m, 10 Jun 1995, S. Legaard & C. Mayorga 17494 (COL);
Margen izquierda del río Guayabero, raudal de La Macarena, Angostura Nro. 1, 350 m, 20 Jan 1959, P. Pinto & Bischler 327 (COL).
Putumayo: Mun. Mocoa, alrededores de la población, 650 m, 20 Aug 1986, B.R. Ramírez 651 (COL, PSO). Valle del Cauca: Carretera
Dagua-Buenaventura, km 32, 540 m, 6 Feb 1961, J.M. Idrobo 4291 & Galeano (COL, US)
4. — cilianensis (All.) Vignolo ex Janch., Mitt. Naturwiss. Vereins Univ. Wien, n.s., 5:110. 1907. (Fig.
Poa pesti All; FL a 2:246. 1785. Eragrostis megastachya var. cilianensis (AIL) Asch. & Graebn., Syn. Mitteleur.
FL 2:371. 1900 ti i ) ET. Hubb., Philipp. J. Sci. 8(3):59-161. 1913. Erosion cilianense (AIL) Lunell, Amer. Midl.
Naturalist 4:221. 1937. Eragrostis multiflora var. cilianensis (All) Maire, Bull. Soc. Hist. Nat. Afrique N. 30: 369. 1939. Tyre: ITALY.
Ciliani, Bellardi s.n. (Lectotype: TO-8242 designated by E Vignolo, Malpighia 18:380. 1904; isorvess: BRI, K photo neg. 195711).
Caespitose annuals. Culms 15-45(-65) cm tall, erect or decumbent and prostrate, glabrous and shiny,
sometimes with crateriform glands below the nodes. Leaf sheaths 2/3 the length of the internodes above,
occasionally longer, glabrous, occasionally glandular, densely ciliate at the summit, the hairs often elon-
gate up to 5 mm long; ligules 0.4-0.8 mm long, ciliate; blades (125-20 cm x (1-)3-5(-10) mm, flat to
loosely involute, mostly glabrous below, scaberulous above occasionally with widely spaced elongate hairs,
sometimes glandular near margins. Panicles (3-)5-16(-20) x 2-8.5 cm, condensed to open, oblong to
ovate, the primary branches 0.4-5 cm long, ascending, appressed or diverging 20-80? from the rachises;
pulvini glabrous or ciliate; pedicels 0.2-3 mm long, erect, spreading to appressed. Spikelets 6-20 x 2-4
mm, 10-40-flowered, ovate-lanceolate, florets imbricate, plumbeous to greenish or hyaline; disarticulation
acropetal, between the florets from the base upwards, usually the entire floret, rachillas persistent; glumes
1.2-2.6 mm long, subequal, broadly ovate to aña sub- Pyae, membranous, keeled, usually with
crateriform glands along the keel, 1- or 3-nerved, f keel; lower glumes 1.2-2 mm
long, usually 1-nerved; upper glumes 1.2-2.6 mm long, often 3-nerved; lemmas 2-2.8 mm long, broadly
ovate, membranous, lateral nerves evident, strongly keeled, keels with 1-3 crateriform glands; apex obtuse
to acute; paleas 1.2-2.1 mm long, hyaline, keels scaberulous, sometimes ciliate, the cilia less than 0.3 mm
long; apex obtuse to acute; stamens 3, anthers 0.2-0.5 mm long, yellow. Caryopses 0.5-0.7 mm long, glo-
bose to broadly-short ellipsoid, striate and reticulate, circular to elliptical in cross-section, reddish-brown.
2n = 20, 40 (Bir & Sahni 1988).
Distribution and habitat. —Native to Europe; introduced in southern Canada, U.S.A, México, Central
America, Caribbean, Argentina, Bolivia, Brazil, Colombia, Ecuador, Paraguay, Peru, Uruguay, and Venezuela
(Peterson 2001; Peterson & Boechat 2001); a weed near cultivated fields, pastures, roadsides, and disturbed
dd 0-3000 m.
1. COLOMBIA. Bogotá D.C.: Bogotá, La Picota, g j i 1, Tunjuelo, 2650 m, 6 Apr 1941, H. García-Barriga
10519 (COL); border between Cesar, Norte de Santander, and des 20 kiló 1 de Abrego, Las Jurisdicciones, Cerro
de Oroque, 3700-3960 m, 19-21 May 1969, H. García-Barriga & R. Jaramillo-Mejía 19723 (COL, US). Cundinamarca: Tocaima, 400 m
no date, C. Saravia 4719 (COL). Huila: Río Cabrera, 2 km below confluence of Río Ambicá, 3 km WSW of Colombia, 1000 m, 15 Dec
1942, F.R. Fosberg 19318 (US); Garzón-Neiva, 400 m, no date, F.C. Lehmann 8742 (US). Nariño: Mun. Pasto, Panamericana, puente del
ee n : m, 7 Nov 1987, B.R. Ramirez 1037 (COL, PSO). Valle del Cauca: Río Calima, región del Chocó, margen derecha, lomas
de la Brea, 30-50 m, 18 May 1946, J. Cuatrecasas 21067 (COL, US); Cartago, Santa Ana de los "ed 950 m,19
Nov 1946, J. Cuatrecasas 23024 (COL, US); Yumbo, finca Río Grande, bosque El Ramón, 1600 m, 13 Jun 1998, D. Stancik 783 (COL)
5. Eragrostis ied (L) R. m Narr. Exped. Zaire 478. 1818. var. ciliaris (Fig. 3, F-J). Poa ciliaris L., Syst. Nat.
(ed. 10) 875. 17 L.) P Beauv., Ess. Agrostogr. 74, 167, 174. 1812. Cynodon ciliaris (L.) Raspail, Ann. Sci. Nat.,
Bot. 5:302. 1825. Eragrostis ciliaris (L.) Nees, Fl. Bras. Enum. Pl. 2:512-514. 1829. Tyee: JAMAICA: Browne s.n. (Lectotype: LINN
87.66! designated by Hüchcoce Cont U.S. Natl. Herb. 12:121. 1908).
Caespitose annuals. Culms (3-)9-75 cm tall, erect or geniculate in the lower portion, not rooting at the lower
nodes, glabrous. Leaf sheaths 1/2-3/4 as long as the internodes, hairy on the margins and at the apices, hairs
to 4 mm long; ligules 0.2-0.5 mm long; blades 1.8-12(-15) cm x 2-5 mm, usually flat, occasionally involute,
glabrous or ciliate basally. Panicles 1.717 x 0.2-1.5 cm, cylindrical, contracted, spike-like, branches forming
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Peterson and Giraldo-Cañas, Eragrostis in Colombia 887
glomerate lobes or sometimes more open, often interrupted in the lower portion; primary branches 0.4-4
cm, ascending, tightly appressed; pulvini usually glabrous, occasionally sparsely pilose; pedicels 0.1-1 mm
long, erect, shorter than the spikelets, glabrous. Spikelets 1.8-3.2 x 1-2 mm, 6-11-flowered, elliptical-ovate
to ovate-lanceolate, yellowish-brown, sometimes with a purple tinge, densely packed next to one another
forming glomerate lobes; disarticulation basipetal, glumes persistent; glumes 0.7-1.6 mm long, subequal,
ovate to lanceolate, keels scabridulous, veins commonly green, apices acute; lower glumes 0.7-1.2 mm
long; upper glumes 1-1.6 mm long; lemmas 0.8-1.3 mm long, elliptical-ovate to lanceolate, membranous,
keels scabridulous, lateral veins evident; apex obtuse to acute; paleas 0.8-1.3 mm long, membranous, keels
prominently ciliate, cilia 0.2-0.8 mm long, stiff and pectinate-thickened near base; apex obtuse to acute;
stamens 2, anthers 0.1-0.3 mm long, purplish. Caryopses 0.4-0.5 mm long, ovoid, striate, slightly dorsally-
flattened, elliptical in cross-section, reddish-brown. 2n - 20, 40 (Bir & Sahni 1988)
Distribution and habitat.—Apparently native to the paleotropics; introduced and naturalized in México,
U.S.A., Central America, Caribbean, Argentina, Bolivia, Colombia, Ecuador, Guianas, Paraguay, Peru, and
Venezuela (Nicora 1998; Peterson & Boechat 2001); growing along roadsides, on waste sites, in xerothermic
M NN saline Mm. and city sidewalks; 0-1600 m.
OLOMBIA. Antioquia: Mun. Medellín, predios de la Plaza de Botero, 1550 m, 4 Jan 2003, D. Giraldo-Cañas
3429 (COL); Medellín, Facultad de Agronomía, 1500 m, 10 Jul 1947, W.H. Hodge 6933 (US); Medellín, 1550 m, 22 Apr 1927, R.A. Toro
239 (US). Arauca: Mun. Arauca, inmediaciones de las instalaciones de la Universidad Nacional de Colombia, hacienda El Cairo, car-
retera Árauca-Tame, km 9, 200-300 m, 13 Jun 2003, D. Giraldo- bins e cn Atlántico: Entre PP Galapa, 80-100 m, 26
Nov 1960, A. Dugand 5380 (COL, US). Bolívar: along Mompós land of Mompós, Lands of Loba, Apr-May 1916,
H.M. Curran 245 (US); Cartagena, 20 Nov 1912, A.S. Hitchcock 9898 (US); Deron Island, mm Bay, 4 Nov 1926, E.P. Killip
E A.C. Smith 14122 (MO, E Cañabetal, río Magdalena, 90-100 m, 15 Jan 1918, FW. Pennell 3882 (MO, US); Cartagena, Caribbean
Coast, 0-1 m, 1-3 Oct 1922, F.W. Pennell 12007 (US). Casanare: Mun. El vu en áreas urbanas del centro de la ciudad, 400 m, 31 Oct
2007, D. Giraldo-Cañas 4133 (COAH, COL); Yopal, 400 m, year 1983, J. R.I. Wood 3815 (COL). Córdoba: Mun. Momil, finca El Paraíso,
en potreros, 39 m, 23 Feb 2005, D. Giraldo-Cafias 3853, 3855 (COL). MN Mun. dd hacienda La Guaira, río Pagüey, 350
m, 16 Oct 2004, D. Giraldo-Cañas 3781 (COL); Mun. Girardot, vía fé es, ribera del río La Magdalena, 250 m,
17 Oct 2004, D. Giraldo-Cañas 3787 (COL); Mun. Nariño, 340 m, 1 Mar 1986, J.L. Rm ud n 5514 (COL). Huila: Mun. Villavieja,
desierto La Tatacoa, 460 m, 28 May 2000, B.R. Ramírez 13193 (CAUP, COL). La Guajira: Mun. Uribia, RU Nazareth, playa
Nueva York, 0—5 m, 6 Jan 2005, J. Betancur et al. 11369 (COL); Maicao, 31 Mar 1962, C. Saravia 365 (COL, US); 2.6 km de Monevideo,
rumbo a Nazareth, 5 Abr 1962, C. Saravia & D. Johnson 467 (COL, US); Clausura Nopoipa, rumbo a Maicao, 4.5 s de Uribia, 13 Feb
1963, C. Saravia 2229 (COL, US), 2229-A (US, note: Saravia 2229-A at COL is E. acutiflora); Serranía La Macuira, entre el arroyo y la
duna Arehuara, 290 m, 4 Mar 1963, C. Saravia 2367 (COL, US); Serranía La Macuira, 12 Apr 1964, C. Saravia & M. E. de Saravia 3544
(COL, MO, US); Riohacha, € i ui G. Sinclair s.n. (US-865575). Magdalena: Río Frío, near sea level, 23 Jun 1906, H. Pittier 1583
(US); Isla de Salamanca Mahoma, 18 Nov 1966, M. Schnetter & R. Schnetter 112 (COL); Santa Marta, 1898-1901, H.H.
Smith 110 (COL, MO, US. Meta: Mun. Villavicencio, carretera Villavicencio-Aeropuerto, piedemonte de la cordillera Oriental, sitio La
Arenera, ca. 21 quía, ca. 400 m, 14—15 Dec 2002, D. Giraldo-Cafias 3386 (COL); Reserva Nacional de La
Macarena, margen aiad del río Guayabero, al pie de las mesetas con el caño Losada, 300 m, 5 Mar 1959, R. Jaramillo-Mejía 2108
(COL, MO, US). Nariño: cupa 45m, 10 ae ae P; us 1923 Ea San Andrés, Providencia y Santa Catalina: Isla de San
Andrés, 22-27 Apr 1948, G.R. Procto gan, 24 Dec 1988, J.H. Torres 3313-A (COL). Santander:
Granja agropecuaria Pedea al sur de Bucaramanga, 1000 m, 6 Oct 1966, A. Robinson 3167 (US); Prope Billete Blanco, 4 May 1926,
w & S. Juzepczuk 4671 (US); Bajo Magdalena, Isla de Venado, inter Billete Blanco et El Presidio, 7 May 1926, G. Woronow & S.
Juzepczuk 4748 (US). Tolima: Mun. Flandes, vía férrea entre Flandes y Girardot, ribera del río La Magdalena, 250 m, 17 Oct 2004, D.
Giraldo-Cañas 3792-A (COL); Piedras, 430 m, 29 Feb 1876, E. André 1868 (US). Valle del Cauca: Cisneros, río Dagua, 300—500 m, 5 May
1939, E.P. Killip 35597 (COL, M a Región Guayanesa, Mun. Puerto Carreño, laj
de Lajas y Cerro El Bita, rib , 40-100 m, 4-5 Jan 2004, D. Giraldo-Cañas & C. Parra 3621 (COAH, COD. Unknown
department: J.C. Mutis 5482 (US); Moritz s.n. (US-557611).
,entre Punta
6. Eragrostis curvula (Schrad.) Nees, Fl. Afr. Austral. Ill. 397. 1841. (Fig. 4, A-E). Poa curvula Schrad., Gott. Gel.
Anz. 3:2073. 1821. Tree: SOUTH AFRICA: Cape Province, Cape of Good Hope, Hess s.n. (HoLotyre: LE; isotype: LE-TRIN 2327.01!
lower middle specimen).
Caespitose perennials forming innovations at the base; culms (45-)60-150 cm tall, erect, glabrous or glan-
dular. Leaf sheaths 1/3-2/3 the length of the internodes, with scattered hairs, hairs to 9 mm long; ligules
0.6-1.3 mm long, ciliate; blades 12-50(-65) cm x 1-3 mm, flat to involute, abaxial surfaces glabrous, some-
times scaberulous, adaxial surfaces with scattered hairs basally, hairs to 7 mm long. Panicles 16-35(-40) x
888
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Eragrostis lugens (I. 5.
3218, J). F Habit. G. Inflorescence. H Spi kelet. |. F
Caryopsis, lateral vie
Peterson and Giraldo-Cañas, Eragrostis in Colombia 889
(48-24 cm, ovate to oblong. open; primary branches 3-14 cm long, diverging 10—80? from the rachises;
pulvini glabrous or not, the hairs up to 3 mm long; pedicels 0.5-5 mm long, appressed, flexible. Spikelets
4-8.2(-10) x 1.2-2 mm, 3-10-flowered, linear-lanceolate, plumbeous to yellowish; disarticulation irregular
to acropetal, proximal rachilla segments persistent; glumes 1.2-3 mm long, subequal, lanceolate, hyaline;
lower glume 1.2-2.6 mm long; upper glumes 2-3 mm long; lemmas 2-3.2 mm long, ovate, membranous,
lateral veins conspicuous; apex acute; paleas 2-3.2 mm long, hyaline to membranous; apex obtuse; stamens
3, anthers 0.8-1.3 mm long, reddish-brown. Caryopses 1-1.7 mm long, ellipsoid to obovoid, strongly
dorsally-flattened, ventral surface with a shallow, broad groove or ungrooved, smooth to striate, mostly
translucent, elliptical in cross-section, light brown, bases often greenish. 2n = 40, 50, 60, 80 (de Wit 1954;
Spies & Jonker 1987; de Wit 1954).
Distribution and habitat.— Native to southern Africa; introduced in the U.S.A, México, Costa Rica, Ar-
gentina, Bolivia, Brazil, Chile, Colombia, Peru, Uruguay, and Venezuela (Nicora 1998; Peterson & Boechat
2001); rocky slopes, margins of woods, roadsides, waste g d, and often used in reclamation; 10-2900 m.
Specimens examined. COLOMBIA. Valle del Cauca: Palmira, cultivada (a partir de semillas de la Argentina) en la Granja Agrícola,
1000 m, 19 Jul 1963, H. S. McKee 10489 (COL, US).
7. Eragrostis hypnoides (Lam.) Britton, Sterns & Poggenb., Prelim. Cat. 69. 1888. (Fig. 5, A-E). Basionw: Poa
hypnoides Lam., Tabl. Encycl. 1:185. 1791. WA hypnoides nio Beauv., Ess. Agrostogr. 74, 167, 175. 1812. Neeragrostis
hypnoides (Lam.) Bush, Trans. Acad. Sci. St. Louis 13:180. 1903. E les (1 ) Lunell, Amer. Midl. Naturalist 4:221.
Type: TROPICAL AMERICA: D. Richard s.n. (HoLoTYPE: P-LAM!; ISOTYPES: BAA- 1041!, NY fragm.ex P!, US-2850742 fragm. ex P!).
Annuals, stoloniferous, mat-forming, without innovations. Culms decumbent and rooting at the lower
nodes, erect portion (2-)5-12(-20) cm tall, often branched, glabrous or hairy on the lower internodes. Leaf
sheaths 1/3-1/2 as long as the internodes above, pilose on the margins, collars, and at the apices, hairs
0.1- S 6 mm Ms ligules 0. E 0.6 mm long, ciliate; blades 0.5-2.5 cm x 1-2 mm, flat to involute, abaxial
, adaxial pr 1 pubescent, hairs ca. 0.2 mm long. Panicles 1-3.5 x 0.7-2.5 cm,
terminal oF axillary ovate, open to somewhat congested; primary branches 0.1-0.5 cm long, appressed
to strongly divergent, glabrous; pulvini sparsely pilose or glabrous; pedicels 0.2-1 mm long, ciliate. Spike-
lets 4-13 x 1-1.5 mm, 12-35-flowered, linear-oblong, often arcuate, loosely imbricate, greenish-yellow to
purplish; disarticulation acropetal, paleas persistent; glumes 0.4-1.2 mm long, subequal, linear-lanceolate
to lanceolate, hyaline; lower glume 0.4-0.7 mm long; upper glume 0.8-1.2 mm long; lemmas 1.4-2 mm
long, ovate, strongly 3-nerved, the nerves greenish; apex acuminate; paleas 0.7-1.2 mm long, hyaline, keels
scaberulous; apex acute to obtuse; stamens 2, anthers 0.2-0.3 mm long, brownish. Caryopses 0.3-0.5 mm
long, ellipsoid, faintly striate and reticulate, laterally-flattened, somewhat translucent, elliptical in cross-
section, light brown. 2n = 20 (Davidse 1981).
Distribution and habitat —Native to the Americas found in North, Central, and South America (exclud-
ing Chile), and the Caribbean (Nicora 1998; Peterson & Boechat 2001); found along muddy or sandy shores
of lakes and rivers and moist disturbed sites; 0-1000 m.
Specimens examined. COLOMBIA. Amazonas: Loretoyacu, 1 Oct 1943, C.O. Grass! 10091 (US); Trapecio amazónico, Amazon River,
Leticia, 100 m, Sep 1946, R.E. De 8196 iius. b Trapecio Ámazónico, Loretoyacu River, 100 m, Oct 1946, R.E. Schultes & G.A.
Blach 8497-A (COL, US). Amazonas q A, Araracuara, sabana de la Angostura, 400 m, 21 Dec 1951, H. García-Barriga
& R.E. Schultes 14155 (COL, US). A Vsupésch ío Apaporis, Soratama, 14 Feb 1952, R.E. Schultes & I. Cabrera 16103 (COL, US).
Antioquia: Puerto Berrío, 125 m, 11 Jan 1931, W.A. Archer 1384 (US). Atlántico: Al sur de Barranquilla, río Magdalena, en la isla en
frente de Puerto Giraldo, 5-10 m, 29 Apr 1960, L.E. Mora 1465 (COL). Arauca: Mun. Arauquita, vereda La Osa, bloque Caño Limón,
plataforma petrolera Chipirón, 150 m, 31 Mar 2008, G.A. Silva 745 (COL). Bolivar: Soplaviento, and vicinity, 5-10 m, 16 Nov 1926,
E.P. Killip & A.C. Smith 14604 (COL, MO, US). Boyacá-Arauca: Los Llanos, Río Casanare about 15 km above Rondón, 200 m, 15 Mar
1939, O. Haught 2677 (US). Caquetá: Río Pepeya, Puerto Tokio, 250 m, 1 Mar 1976, J.M. Idrobo 8597 (COL). Casanare: Los Llanos,
along Meta, near Orocué, 140 m, 30 Mar dbi O. qeda! (COL, en US). Cauca: Cerca ho Puerto a ae 1000 m, 14 HUE 1968, :
Espinal 2380 (COL). Córdoba: Mun. Lori Estación Piscícola
del valle de los ríos Sinú y San Jorge), ca. 50 m, 22 Feb 2005, D Giraldo- Cañas et al. 3847-A cou HUA, pr a iS Nro.
2, margen derecha del río Guayabero, 200 m, 23 Feb 1969, P. Pinto 1010 (COL). Meta: M 10 km abajo
del caño Lozada, 350 m, 30 Jan 1959, P. Pinto 365 (COL, US). Santander: Río Masdsdine: Bs del río Sogamoso, 110- 120 m, 15 Jan
890
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Fic. 5. E (A. López M. & A Sagástegui A. 4023, D &E). A. Habit. B. Spikelet. C. Fl hed below, itl yopsi
D. Caryopsis, dorsal view. E. Caryopsis, lateral view. El tis tenella (S. Llatas Q. 1180, H & I). F. Cul ith Infl G Spikelet H Caryopsis, dorsal
view. I. Caryopsis, lateral view. Eragrostis viscosa (0. Haught 4464). H. Spikelet with sand grai hed to pedicel and | gl
Peterson and Giraldo-Cañas, Eragrostis in Colombia 891
1918, F.W. Pennell 3842 (US). Valle del Cauca: Buga, no date, M. A. Bonpland s.n. (COL-253787); NE of Cali, near Río Cauca, 1000 m, 3
Mar 1939, E.P. Killip & G. Varela 3470: (COL, US).
8. dis diam intermedia Hitchc., J. Wash. Acad. Sci. 23(10):450. 1933. (Fig. 2, D-D. Tee U.S.A. Texas: Bexar
. San Antonio, 3 Jul 1910, A.S. Hitchcock 5491 (HoLorvrE: US-1535749!; isotypes: US-908993!, US-15357501).
Caespitose perennials with innovations, not glandular. Culms (30-)40-90(-110) cm tall, erect, glabrous
below the nodes. Leaf sheaths overlapping, Y to about as long as the internodes below, sparsely pilose
on the margins, apices hairy, hairs to 8 mm long, not papillose-based; ligules 0.2-0.4 mm long; blades
(4-)10-20(30) cm long, 1-3 mm wide, flat or involute, abaxial surfaces glabrous, adaxial surfaces densely
hairy behind the ligules, elsewhere usually glabrous, occasionally sparsely hairy. Panicles 15-40 cm long,
(8.5-)15-30 cm wide, ovate, open; primary branches 4-25 cm long, diverging 20—90? from the rachises,
capillary; pulvini hairy or glabrous; pedicels 2-14 mm long, divergent. Spikelets 3-7 mm long, 1-1.8 mm
wide, narrowly lanceolate, olivaceous to purplish, with (3-)5-11 florets; disarticulation acropetal, paleas
persistent; glumes lanceolate to ovate, hyaline to membranous; lower glumes 1.1-1.7 mm long, narrower than
the upper glumes; upper glumes 1.3-2 mm long, apices acuminate to acute; lemmas 1.6-2.2 mm long, ovate,
membranous, hyaline near the margins, lateral veins inconspicuous, apices acute; paleas 1.4-2.1 mm long,
hyaline, narrower than the lemmas, apices obtuse to acute; stamens 3; anthers 0.5-0.8 mm long, purplish.
Caryopses 0.5-1.0 mm long, rectangular-prismatic, somewhat laterally compressed, with a well-developed
adaxial groove, striate, opaque, reddish-brown. 2n = ca. 54, 60, 72, ca. 74, 80, 100, 120.
Distribution and habitat —Eragrostis intermedia is an American native species, and grows in clay, sandy,
and rocky soils, often in disturbed sites; 0-2800 m. Its range extends from the United States through México
and Central America to South America.
Comments.—Eragrostis intermedia is similar to the more widespread E. lugens, but differs from that spe-
cies in having wider spikelets, longer lemmas, and caryopses with a prominent adaxial groove. A numeri-
cal taxonomic study of the Eragrostis intermedia complex was completed by Witherspoon (1975) where he
found much phenotypic overlap of individuals in his principal component and UPGMA cluster analyses
of E. intermedia with E. palmeri S. Watson, E. erosa Scribn. ex Beal, and E. hirta E. Fourn. Determination of
these species is often problematic and examination of this group, which additionally includes E. lugens and
E. hirsuta (Michx.) Nees, is needed to clarify species boundaries.
Specimens examined. COLOMBIA. Cundinamarca: Suesca-Nemocón, vereda Río Checua, hacienda Susatá, en cultivos de Acacia,
áreas secas, 2600-2800 m, 7 Jul 2000, J.L. Fernández et al. 18921-B (COL), 23 Aug 2000, J.L. Fernández et al. 19163 (COL); Mun. Suesca,
acienda Susatá, en matorral seco, 2650 m, 8 Sep 1999, Groenendijk & Rietman 1229 (COL), 26 Nov 1999, Groenendijk & Rietman 1277
(COL), 15 Nov 1999, Groenendijk & Rietman 1325 (COL), 2 Aug 2000, Groenendijk & Rietman 1534 (COL,
9. Eragrostis japonica (Thunb.) Trin., Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math. 1(4):405.
1830. (Fig. 6, A-D) japonica Thunb, Fl. Jap. 51. 1784. E i lla var. japonica (Thunb.) Roem. & Schult., Syst.
Veg. 2:576. 1817. Diandrochloa japonica (Thunb.) A.N. Henry, Bull. Bot . Surv. India 9:290. 1968. Roshevitzia japonica (Thunb.)
Tzvelev, Novosti Sist, Vys&. Rast. 7:50. 1970(1971). Tyee: JAPAN: Herb. Thundberg 2252 (uotorvee: UPS, microfiche IDC 1036!, K
photo!; isorvee: BRI fragm.
o
Sep
Poa glomerata Walter, Fl. Carol. 80. 1788. Eragrostis glomerata (Walter) L.H. Dewey, Contr. U.S. Natl. Herb. 2(3):543. 1894. een
abi (Walter) Schult., Mant. 2:327. 1824. Diandrochloa glomerata (Walter) Burkart, Bol. Soc. Argent. Bot. 12:287. 1
SOUTH CAROLINA: Walter s.n. (HOLOTYPE: BM!
Caespitose annuals. Culms 25-100(-115) cm tall, erect, sometimes geniculate below, branching from the
lower and middle nodes, glabrous and somewhat shiny below the nodes. Leaf sheaths 3/4 to 7/8 as long as the
internodes above, glabrous at the summit and along the upper margins; ligules 0.4-0.6 mm long, a delicate
membrane, glabrous; blades (4215-20(-25) cm x 1.5-6 mm, flat, glabrous below and scaberulous above,
sometimes auriculate near the base. Panicles 15-40 x 0.8-5 cm, lanceoloid, contracted, interrupted below,
the ascending, often appressed primary branches 2-10 cm long, spreading up to 30? from the rachises, the
branches scaberulous and shiny, floriferous near base; pulvini glabrous; pedicels 0.5-1.5 mm long, erect
and sinuous. Spikelets 2.2-3.8 x 0.8-1.3 mm, 4-12-flowered, oblong to narrowly lanceolate, yellowish-
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Peterson & N.F. Refulio-Rodriguez 13855, E-G
view. Eragrostis secundiflora (W.R. Anderson, M. Stieber & J.H. Kirkbride, Jr. 37109, K). |. Inflorescence. J. Spikelet. K. Floret.
Peterson and Giraldo-Cañas, Eragrostis in Colombia 893
brown to whitish and hyaline; disarticulation basipetal, the rachillas and glumes persistent; glumes 0.6-1
mm long, subequal, ovate to ovate-lanceolate, hyaline, faintly keeled, scaberulous along the keel; upper
glumes without a midnerve; lemmas 0.9-1.2 mm long, ovate, hyaline, the lateral nerves conspicuous below,
greenish, keeled, glabrous along the keel; apex acute; paleas 0.6-0.8 mm long, hyaline, scaberulous along
the keel near the apex; apex acute, often bifid; stamens 2, anthers 0.1-0.2 mm long, whitish to light brown.
Caryopses 0.3-0.4 mm long, obovoid, smooth or minutely irregularly striate, slightly dorsally-flattened,
pericarp often loose, elliptical in cross-section, reddish-brown. 2n - 20, 60 (Christopher & Abraham 1974;
Pohl & Davidse 1971 reported for E. glomerata).
Distribution and habitat.—Native in tropical regions of Asia; introduced in U.S.A., México, Central
America, Caribbean, Argentina, Bolivia, Brazil, Colombia, Ecuador, Guyana, Paraguay, Peru, Uruguay, and
Venezuela (Nicora 1998; Peterson & Boechat 2001); occurs in moist areas along rivers and streams usually
in d soils; 0-400 m.
; 1. COLOMBIA. A Río Caquetá, La Pedrera, 240 m, 1-4 Oct 1952, H. García-Barriga 14586 (COL). Atlántico:
Enl ha del río Magdal frente d illa, 6 m, 13 May 1960, L.F. Mora 1498 (COL). Casanare: Yopal, hacienda
La Milanesa 300 m, 10 Nov 1974, P. Carranza 11 (COL). je ells No data, 23 Sep 2000, C. Romero 108 (COL). Magdalena:
Santa Marta, 1898-1899, H.H. Smith 111 (COL, MO, US). Meta: Llanos, 190 m, 3 Jan 1974, G. Davidse & F. Llanos 5504 (COL, MO); Llano
de San Martín, 250 m, no date, H. Karsten s.n. (US-1126597); Llano de San Martín, 250 m, Jan 1856, J.J. Triana 352 (COL, US). Tolima:
Saldaña, vereda Baudá, vía Saldaña-Baudá, 360 m, 10 Jul 2002, A. Osorio 255 (COL). Vichada: Entrada del raudal San Borja, Ventanas,
00 m, 17 Mar 1971, P. Pinto & C. Sastre 1317 (COL, US).
10. Eragrostis lugens Nees, Fl. Bras. Enum. Pl. 2:505-506. 1829. (Fig. 4, F-J). Eragrostis pilosa var. lugens (Nees)
i Abh. ae di Wiss. pun 24:290. 1879. Tree: BRAZIL: E Sellow s.n. (LecroryrE: US-732957! designated by Davidse,
B, BAA-2932
Caespitose Į ials formi j t base. Culms 30-70 cm tall, erect, sometimes geniculate below,
glabrous and somewhat EC below the nodes. Leaf sheaths 1/2-2/3 as long as the internodes above, ciliate
at the summit and along the upper margins; ligules 0.2-0.3 mm long, ciliate; blades (5-)8-22 cm x 1-2(-2.2)
mm, involute, rarely flat, mostly glabrous above and below, scaberulous towards the apex and along mar-
gins, sometimes with scattered hairs along the margins, the hairs up to 7 mm long. Panicles 16-28 x 10-21
cm, open, ovate, the ascending primary branches 0.6-15 cm long, spreading up to 100? from the rachises,
the branches scaberulous, not floriferous near base; secondary branches composed of loosely overlapping
spikelets; pulvini ciliate, the hairs up to 7 mm long; pedicels 1.4-5(-7) mm long, erect, wiry, spreading.
Spikelets 2-4.5(-5) x 0.5-1.1 mm, 2-7-flowered, narrow lanceolate, light plumbeous to reddish-purple;
disarticulation acropetal, paleas and rachilla persistent; glumes 0.6-1.4 mm long, subequal, broadly ovate
to narrowly lanceolate, hyaline, keeled, scaberulous along the keel, sometimes reddish-purple; lower glume
0.6-1 mm long; upper glume 1.1-1.4 mm long, usually broader than the lower; lemmas 1.2-1.8 mm long,
broadly ovate, membranous, the distal margins hyaline, lateral nerves obscure to barely evident, keeled,
scaberulous along the keel near apex; apex acute; paleas 1.1-1.7 mm long, membranous to partially hyaline,
scaberulous along keels; apex obtuse; stamens 3, anthers 0.2-0.7 mm long, reddish-purple. Caryopses
0.5-0.6 mm long, obovoid to somewhat prism-shaped, terete to somewhat laterally flattened, with a weak
ventral groove, striate and reticulate, usually opaque, irregularly obovate to triangular in cross-section,
faintly reddish-brown to whitish. 2n = 40, 80 (Gould 1958).
Distribution and habitat —Native to the Americas with a broad distribution from U.S.A., Mexico, Cen-
tral America, Argentina, Bolivia, Brazil, Colombia, Ecuador, Peru, Uruguay, and Venezuela (Nicora 1998;
Peterson & Boechat 2001); sandy dunes, river banks, near cultivated fields, and open slopes; 0-2800 m
9910A
Specimens examined. COLOMBIA. Boyacá: along Río Soapaga, 12 km of Belén, 2460 m, 7 Nov 1944, F.R. Fosberg (US). Cauca:
Chisquio, finca Los Derrumbos, 1700 m, 3 Feb 1940, E. Asplund 10510 (US). El Ramal to Río Sucio, W of Popayán, 1600-1900 m, 3 Jul
1922, F.W. Pennell & E.P. Killip 8135 (US). Cundinamarca: Sabana de de entre Sibaté y San Miguel, 2750 m, 15 Aug 1939, J. Cua-
trecasas 6631 (COL, US). Huila: Cordillera Oriental, E of Neiva, 1500 m, 1-8 Aug 1917, H.H. Rusby & F.W. Pennell 1023 (US). Valle del
Cauca: Mozambique, N of La Cumbre, 16 Sep 1922, E.P. Killip 11277 m Carretera al mar, W of Cali, Cordillera lental, 1300 m,
13 Nov 1948, E.P. Killip & F.C. Lehmann 39795 (US). Pavas, Cordillera Occidental, 1500-1800 m, 12 May 1922, F.W. Pennell 5529 (US).
Unknown department: J.C. Mutis 5399 (US), 5401 (US
894 Journal of the Botanical R h Institute of Texas 2(2)
11. Eragrostis lurida J. Presl, Reliq. Haenk. 1(4-5):276. 1830. subsp. lurida (Fig. 7, A-J). Poa lurida (J. Presl)
Kunth, Enum. Pl. 1:342. 1833. Tyre: PERU: T. Haenke s.n. (moLorvrE: PR; isotypes: MO-21111461, PR, US-2941523 fragm. ex PRD).
Eragrostis contristata Nees & Meyen, Gramineae 31-32. 1841. Tyre: PERU: Laguna de Titicaca, Apr. 1831, Meyen s.n. (HOLOTYPE: B; ISOTYPES:
1 fragm. ex Bl, LE- TRIN-2326.01!, P!, US-2891461 fragm!).
Eragrostis bahiensis var. boliviensis Henrard, Meded. Rijks-Herb. 40:68. 1921. Tyre: BOLIVIA: La Paz: Larecaja, Sorata, 2900 m, Feb 1858,
G. Mandon 1332 (rotorves: L 908,88-303; isorvrrs: K!, L 908,88-342, Pl, US-31611271, US-256470!, US-911775 fragm. ex L!).
Caespitose perennials with innovations at base; culms (5-)15-75 cm tall, erect, sometimes slightly genicu-
late below, glabrous or occasionally with a tuft of hairs below the nodes, the hairs less than 1 mm long,
sometimes with an occasional elliptical orange gland. Leaf sheaths % to about as long as the internodes
above, densely white ciliate at the summit and along the margins, sometimes with scattered hairs on the
abaxial surface; ligules 0.5-0.9 mm long, ciliate; blades 1.5-16.5 cm x 1.5-3.5(-5) mm, flat to involute,
glabrous and sometimes shinny below and scaberulous above. Panicles 3-35 x 2-5(-7) cm, narrowly ovate,
contracted and condensed into tightly glomerate lobes, interrupted near base, spicate to narrowly ovate,
rachis glabrous, the ascending primary branches 0.6-5 cm long, densely-flowered, spreading 20-80? from
the rachises, floriferous near base; secondary branches condensed into tightly glomerate lobes of spikelets;
pulvini ciliate or glabrous; pedicels 0.1-1 mm long, ascending and appressed, wiry, scabrous. Spikelets
2.5-6 x 1-2.1 mm, 3-10-flowered, narrowly lanceolate to ovate, inflated to slightly compressed, dark to
light plumbeous, sometimes purple-tinged, rachilla often ciliate; disarticulation acropetal with glumes first,
then lemmas falling individually, paleas persistent on rachilla; glumes 1-2 mm long, subequal, broadly ovate
to lanceolate, membranous, margins hyaline, keeled, scaberulous along keel, sometimes 3-nerved; lower
glumes 1-1.5 mm long; upper glumes 1.2-2 mm long; lemmas 1.6-2.4 mm long, broadly ovate, membranous,
lateral nerves conspicuous, evident, keeled; apex acute, scaberulous, the minute prickle hairs appearing as
whitish dots under 10-20X; paleas 1.5-2.3 mm long, membranous to partially hyaline, scaberulous along
the keels; apex truncate to obtuse; stamens 3, anthers 0.4-0.6 mm long, reddish-purple. Caryopses 0.6-0.8
mm long, obovoid to ellipsoid, striate and reticulate, sometimes with a weak ventral groove, rectangular
with the lateral sides angled in cross-section, light reddish-brown. 2n = unknown.
Distribution and habitat.—Native to the central Andes from Bolivia, Colombia, Ecuador, and Peru
(Nicora 1998; Laegaard & Peterson 2001); dry rocky hillsides, slopes, sandy roadsides, and rocky alluvial
fans; 2000-3800 m.
Specimens examined. COLOMBIA. Cauca: Popayán, no date, F.C. Lehmann 6997 (US). Nariño: Cumbal, André 3524 (US). Unknown
department: J.C. Mutis 5517 (US).
12. Eragrostis maypurensis (Kunth) Steud., Syn. Pl. Glumac. 1:276. 1854. (Fig. 8, A-E). Poa maypurensis
Kunth, Nov. Gen. Sp. 1: 161, 162. 1815 (1816). Megastachya maypurensis (Kunth) Roem. & Schult., Syst. Veg. 2: 588. 1817. Tyre
VENEZUELA. Amazonas: Rio Orinoco, Apr, E Humboldt & A. Bonpland s.n. (oLotyre: Pl; isotypes: COL!, BAA-1053 fragm!, P!, US-
2850758 fragm. ex P-BONPL! & fragm. ex B-WILLD).
Caespitose annuals. Culms 25-45 cm tall, erect to ascending, often decumbent, many branched from the
base forming a rosette, mostly with 2-3(-6) nodes. Leaf sheaths Y-2/3 as long as the internodes above,
mostly glabrous, pilose at the summit and along upper margins; ligules ca. 0.5 mm long, ciliate; blades
6-12 cm x 2-4 mm, flat to loosely involute tow. the apex, scattered pilose near base and long margins to
pilose abaxially and adaxially, the hairs 1.5-5 mm long, papillose-based. Panicles 6-14 x (1.5-)3-7.5 cm,
open, narrowly ovate to oblong, more or less densely-flowered, primary branches 1.5-5 cm long, spreading,
solitary at a node, floriferous to base, branches diverging 0-70? from the rachises; pulvini pilose, the hairs
up to 4 mm long; pedicels 0-1.5 mm long, shorter than the spikelets, appressed, with a few scattered hairs.
Spikelets 8-15(-30) x 1.8-2.5(-3) mm, 12-35-flowered, narrowly lanceolate to ovate lanceolate, florets im-
bricate often appearing to be borne in fascicles, reddish-purple to yellowish; disarticulation acropetal with
the paleas and glumes persistent; glumes (1.5-)2-2.6 mm long, subequal, lanceolate-acuminate, strongly
to weakly keeled, scaberulous along the keel; lower glumes 2-2.6 mm long, usually longer than the upper;
upper glumes (1.5-)2-2.6 mm long; lemmas 1.8-3 mm long, ovate-acuminate, chartaceous, lateral nerves
895
Peterson and Giraldo-Cañas, Eragrostis in Colombia
Sats NEN
Se a
OU SS
E s Qm AS
Fis. 7. Eragrostis lurida (PM. Peterson, E.J. Judziewicz & R.M. King 9103 & PM. Peterson & E.J. Judziewicz 9327, A—H; P.M. Peterson & N.F. Refulio-Rodriguez
13993, | & J). A. Habit. B. Ligule. C. Inflorescence. D. Spikelet. E. Floret. F. Lemma, dorsal view. G. Palea, dorsal view. H. Palea enclosing the stamens,
pistil, and lodicules, ventral view. I. Caryopsis, dorsal view. J. Caryopsis, lateral view.
896 J lof the Botanical R h Institute of Texas 2(2)
Fic. 8. Eragrostis maypurensis (J.R. Swallen 3306, A—C; CM. Belshaw 3359, D & E). A. Habit. B. Spikelet. C. Floret. D. Caryopsis, dorsal view. E. Caryopsis,
lateral view. Eragrostis polytricha. F. Inflorescence. G. Spikelet, H. Floret.
Peterson and Giraldo-Cañas, Eragrostis in Colombia 897
and midnerve evident usually green; apex acuminate and recurved; paleas 1.5-2.2 mm long, shorter than
the lemma, hyaline; stamens 2, anthers 0.2-0.3 mm long, purplish. Caryopses 0.4-0.7 mm long, ovoid,
translucent, rhomboid reticulate without striations, obovate to circular in cross-section, reddish-brown. 2n
= 20 (Pohl & Davidse 1971).
Distribution and habitat.—Native to the Americas from México, Central America to South America in
Bolivia, Brazil, Colombia, Peru, and Venezuela (Nicora 1998; Laegaard & Peterson 2001); a fairly common
grass in the Amazonian region where it is found growing in savannas, sandy sites, along roadsides, and
disturbed areas; 100-1500 m.
Vernacular name.—"Colchón de pobre" (Casanare, J.M. Idrobo 5100).
Specimens examined. COLOMBIA. Amazonas: Araracuara, camino entre el Hospital y el Aeropuerto, ca. 200 m, 12 Sep 1977, L.E.
Aguirre-Galvis 1061 (COL); Río Caquetá, cerca del aeropuerto, 200-250 m, 18 Jul 1977, A. Fernández-Pérez 20061 (COL). Antioquia:
Bello, 1500 m, 29 n pus W.A. eid ds s E oe ue 19-20 Oct 1938, J. Cuatrecasas 3863 (COL, US); Mun.
El Yopal, sal 1,400 m, 3 Nov 2007, D. Giraldo-Cañas
4149, 4150, 4153 (COAH, COL); El Yopal, mM Mate psa 10 Oct 1962 ,J.M. arabe 5100 (COL); Mun. Paz d Ariporo, corregimiento
La Hermosa, finca Nicaragua, caño Pica Pico, 112 m, 25 Oct 2004, J.G. Ramírez 8705 (COAH, COL); Tauramena, quebrada Tauramena,
550 m, 30 Nov 1960, L. Uribe Uribe 3588 (COL). Cauca: along Río Palo at El Palo, 15 Dec 1943, E.L. Core 171 (US); Río Patía, 11 May
1935, H. Garcia-Barriga 4497 (COL, US). Guainía: Bassin de LAmazone, Río Negro a San Felipe, 200 m, 13-25 Nov 1952, H. Humbert
27487 (US); Puerto Colombia, opposite Venezuelan town of Maroa, and vicinity, ca. 280 m, 31 Oct 1952, R.E. Schultes 17927 (COL, US).
Guaviare: Mun. San José del Guaviare, trocha Nuevo Tolima, 11 Nov 1996, R. López & O. Rodríguez 1859 a COL). End Five a
N of Villavieja, upper basin of Río Magdalena, 400 m, 23 Jul 1950, S.G. Smith 1256 (COL, US). Magdalen
valle del río Magdalena, 100 m, 23 Dec 1948, J. Araque-Molina & F.A. Barkley 18- e nd e nue pon 1901, H.H. Smith 132
(COL, MO, US). Meta: Mun. Villavicencio, carretera Villavicencio-Aeropuert sitio La Arenera, ca
2 km del puente sobre el río Guatiquía, ca. 400 m, 14-15 Dec 2002, D. Giraldo- Cañas 3384 (COAH, COL, HUA); Río Meta, Cabuyaro, 235
m, 14 Oct 1938, J. Cuatrecasas 3612 (COL, US); Río Meta, La Ochovera, 28 Oct 1938, J. Cuatrecasas 4182 (COL, US); Villavicencio, Apiai,
500 m, 12 Nov 1938, J. Cuatrecasas 4730 (COL, US); Llanos Orientales, Puerto López, al SE de Ea alrededores de la laguna de
Yurimena, 175-200 m, 16 Sep 1958, R. Jaramillo-Mejía 1247 (COL, US); Carimagua, 200 m, 18 Sep 1992, S. A. Renvoize 5399 (COL, US);
San Juan de Arama, caño Curía, 440 m, 31 Dec 1986, D. Rivera 1265 (COL); Loma Linda, al sureste 5s San edi 300 m, 15 Sep 1966,
A. Robinson 3079 (US); Villavicencio, 450 m, Jan 1856, J.J. Triana 70 (US), 297 (US). Nariño: Carretera Past , alrededores del
río Juanambú, 1380 m, 1-5 Dec 1962, C. Saravia 1950 (COL). Santander: “Kilómetro 16," between Puerto una and Puerto Santos,
110-115 m, 29 Nov 1926, E.P. Killip & A.C. Smith 14825 (MO, US); Carretera Puerto Wilches-Sabana de Torres, km 46, 250 m, 16 Nov
1985, J.H. Torres 2816, 2817 eae oin no data, André 1924 (US); Melgar, 500-600 m, 4-5 Dec 1917, FW. P 112903 (US). Valle
del Cauca: Around Cali, Cauca Valley, 1000-1200 m, Dec 1905, H. Pittier 639 (US), 660 (US). Vichada: Región Guaya-
nesa, Mun. Puerto Carreño, afloramientos rocosos del tipo lajas, entre Punta de L Lajas y Cerro El Bita, ribera del río Orinoco, 40-100
m, 4-5 Jan 2004, D. Giraldo-Cañas & C. Parra 3648, 3655 (COAH, COL); Rio Orinoco, Puerto Carreño, 23-24 Oct 1938, J. Cuatrecasas
4072 (COL, US); ca. 10 km W of Las Gaviotas along road to Puerto Gaitán, 180 m, 30 Dec 1973, G. Davidse & F. Llanos 5367-A (COL,
MO, US); Gualandayas, ca. 100 km E of Gaviotas, 100 m, 31 Dec 1983, J. R.I. Wood 4213 (COL). Unknown department: F.C. Lehmann
778 (US), I. Linden 1554 (US), Moritz s.n. (US), J.C. Mutis 6094 (US), J.]. Triana 298 (US).
13. Eragrostis mexicana Morem ) Link, Fiort Berol. 1:190. 1827. (Fig. 9). Poa mexicana Hornem., Hort. Bot. ed
2:953. 1815. Tree: MÉXICO: cultivated fi ted in México, Sessé s.n. (Lectotype: MA! designated by
Vega, Ánn. Missouri Bot. Gard. 94:773. 2007).
Caespitose annuals. Culms 10-130 cm tall, erect, sometimes geniculate, glabrous, sometimes with a ring
of glandular depressions below the nodes. Leaf sheaths %-2/3 as long as the internodes, sometimes with
glandular pits, pilose near the apices and on the collars, hairs to a mm lone; papillose- d n 0.2-0.5
mm long, ciliate; blades 5-25 cm x 2-7(-9) mm, flat, abaxial g d
occasionally pubescent near the base. Panicles (5210-40 x (2-)4-18 cm, Ls than Y^ the height of the plani
ovate, rachises angled and channeled; primary branches 3-12(-15) cm long, solitary to whorled, appressed
or diverging to 80? from the rachises; secondary branches somewhat appressed; pulvini glabrous; pedicels
1-6(-7) mm long, almost appressed to narrowly divergent, stiff. Spikelets (425-10(-11) x 0.7-2.4 mm,
5-11(-15)-flowered, linear to linear-lanceolate or ovate to oblong, gray-green to purplish; disarticulation
acropetal; glumes 0.7-2.3 mm long, subequal, ovate to lanceolate, membranous; lemmas 1.2-2.4 mm long,
ovate, membranous, glabrous or with a few hairs, gray-green, lateral veins evident, often greenish; apex
acute; paleas 1-2.2 mm long, hyaline, keels scabrous; apex obtuse to truncate; stamens 3; anthers 0.2-0.5
898 ¡ lof tha Botanical R h Institute of Texas 2(2)
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D. Carvopsis
* F
Fic. 9. Eragrostis mexicana var. mexicana (I. Sánchez-Vega 4020, D). A. Inflorescence. B. Spikelet. C. Fl
lateral view. E ti j viresi I |
cens. E. Inflorescence. F. Spikelet G. Fl
mm long, purplish. Caryopses 0.5-0.8(-1) mm long, ovoid to rectangular-prismatic, laterally flattened, shal-
lowly to deeply grooved on the ventral surface, striate and reticulate, irregularly triangular in cross-section,
reddish-brown, distal 2/3 opaque. 2n = 60 (Pohl & Davidse 1971).
Distribution and habitat.—Eragrostis mexicana is native to the Americas and grows along roadsides, near
cultivated fields, city sidewalks, and in disturbed open areas, in lomas vegetation and slopes of the inter-
Andes of Colombia; 1000-2400 m. The species has been treated as two distinct subspecies by Koch and
Sánchez Vega (1985) and Peterson and Sánchez Vega (2007), both of which occur in Colombia.
Peterson and Giraldo-Cañas, Eragrostis in Colombia 899
KEY TO THE SUBSPECIES OF ERAGROSTIS MEXICANA
1. Spikelets ovate to oblong in outline, 1.5—2.4 mm wide; lower glumes 1.2—2.3 mm long; sum of the spikelet
width and lower glume length 2.7—4.7 mm; culms and sheaths sometimes with peg depressions
exicana subsp. mexicana
1. Spikelets linear to iinear-ianceolate, 0.7—1.4 wide; lower glumes 0.7—1.7 mm long; sum Red Spikelet width
and lower glume length 1.5—3.1 mm; culms and sheaths without glandular depressions 13b. E. mexicana
subsp. virescens
13a. Eragrostis mexicana (Hornem.) Link subsp. mexicana (Fig. 9, A-D).
E is limbata E. Fourn., Mexic. Pl. 2:116. 1886. Tree: MÉXICO: 1833, AJ.A. iue 4573 (LecroTYPE: P! designated by R. McVaugh,
EL Novo-Galic. 14:168. 1983 but specific herbarium not indicated, Bonpl housed at P; isotype: US-2941517
fragm!).
coe neomexicana niin exL.H. AMA nue U.S. Natl. Herb. 2(3):542. 1894. Tyre: U.S.A. New Mexico: Organ Mountains, 1881, G.
S.D. Koch and I. Sánchez Vega, Phytologia 58:379. 1985; isotypes: K!, US-822049!,
US-909912!).
Eragrostis alba J. Presl, Reliq. Haenk. 1(4—5):279. 1830. Poa alba (J. Presl) Kunth, Enum. Pl. 1:343. 1833. Tyre: PERÚ: T. Haenke s.n.
(HOLOTPE: PR; isotypes: MO-2111118!, US-2942409 fragm. ex P!).
Culms and sheaths sometimes with glandular depressions. Spikelets 1.5—2.4 mm wide, ovate to oblong;
lower a 1.2-2.3 mm long; sum of spikelet width and lower glume length 2.7-4.7 mm
d. COLOMBIA. Boyacá: Mun. Ráquira, cerca de los hornos de los artesanos, en inmedfiaciones del camino que
a al nd de La Candelaria, 2300 m, 19 Jul 2003, D. Giraldo-Cañas 3555, 3555-B (COL); Mun. Villa de Leyva, saliendo por el
Iguaque, en = afueras del casco urbano de Villa de Leyva, 2200 m, 18 Dec 2004, D. Giraldo-Cafias 3825, 3835,
3836 (COL): between Boavit Rio Chicamocha, 1500 m, 25 Jun 1984, i Wood is (COL). Cauca: Popayán, 1600-1800 m,
Feb 1887, F.C. Lehmann 4401 (US). Cundinamarca: Mun. Choachí, carretera CI le Choachí, km 2, 1600 m, 12 Nov 2001,
D. Giraldo-Cañas 3284 (COL). Nariño: C Past lrío] bú, 1380 m, 1-5 Dec 1962, C. Saravia 1954
(COL). Santander: 2 km al sur de Suratá, 1900 m, 4 Aug 1966, A. Robinson 3054 (US). Valle del Cauca: between Uribe and Sevilla,
1100 m, 2 Mar 1983, J. R.I. Wood 4082 (COL)
13b. Eragrostis mexicana subsp. virescens (J. Presl) S.D. Koch & Sánchez Vega, Phytologia 58(6):380. 1985.
(Fi ig. 9, E-G). Eragrostis vi esl, Reliq. Haenk. 1(4—5):276. 1830. Poa virescens (J. Presl) Kunth, Enum. PL 1:329. 1833.
Tree: CHILE: T. Haenke s.n. (HOLOTYPE: TR ISOTYPES: B!, BAA-1107 fragm. ex B!, LE-TRIN-2413.01!, US-2942410 fragm!).
Eragrostis leptantha Trin., Mém. Acad. inp. Sci. St. o Sér. 6, Sci. Math. 1(4):405. 1830. Poa leptantha (Trin.) Kunth, Enum.
. 1833. Tree: BRAZ I oji [Sao Paulo], 23 Apr, [Riedel s.n.] G.H. von Langsdorff s.n. (HoLotyPE: LE-
np 2361.01); sorres: BAA-1043 fragm.!, K, LE, MO-2111141!, NY!, P!, US-2941514 fragm.!).
Trin., Mém. Acad. Imp. Sci. Saint-Pétersl g, Sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 2(1):73. 1836. Eragrostis pilosa
ro inl
var. M a (Trin.) Hack., Anales Mus. Nac. Buenos Aires 11:133. 1904. Tree: BRAZIL: In cultis prope Rio de Janeiro, May-Jun
L. Riedel s.n. (HoLotyPE; LE-TRIN-2330.01!; isoLecroTYPES: LE, US- IU fragm. D.
Pega idis Phil., Verz. Antofagasta Pfl. 88. 1891. Tyre: CHILE. T. , 13 Mar 1885, R.A. Philippi 359 (HoLo-
E: SGO-PHI-359; BAA-1080 fragm.!, CORD!, $GO-37293; SGO-63537, SGO-62670, US-556538 fragm. ex SGO-PHIL-359!).
dobensis Jedwabn., Bot. Arch. 5(3-4):208. 1924. Tyre: ARGENTINA. Corposa: prope urbem, 21 Apr 1881, Galander 54b
paene BAA! designated by Boechat €: Longhi- Wagner, Iheringia, Bot. 55:164. 2001 US-2767407!).
Culms and sheaths without glandular depressions. Spikelets 0.7-1.4 mm wide, linear to linear-lanceolate;
lower glumes 0.7-1.7 mm long; sum d DRE NUS and lower glume length 1.53.1 mm.
Distribution and habitat.—T] to South America in Argentina, Bolivia, Brasil, Chile,
Colombia, Paraquay, Peru, and Ue introduced in Canada, México, and U.S.A. (Nicora 1998; Peterson
& Boechat 2001; Peterson & Sánchez Vega 2007).
Specimens examined. COLOMBIA. Antioquia: Mun. Sabaneta, parque central de la municipalidad, 1550 m, 3 Jan 2003, D. Giraldo-
Cañas 3428 (COL, HUA). Cauca: Mun. Popayán, sector norte de la ciudad, Villa del Viento, 1800 m, 7 Jan 2001, B.R. Ramirez 13911
(CAUP, COL). Quindío: Mun. Calarcá, entrada del jardín botánico del Quindío, 1500 m, 20-25 May 2005, D. Giraldo-Cañas 3893
(COL, HUA). Santander: Entre el cañón del Chicamocha y el valle de Rupala, 1000 m, 14. Jun 1962, C. Saravia 772 (COL). Tolima:
Cordillera Central andina, Mun. Ibagué, jardineras y andenes de la Plaza de Bolívar, 1200 m, 11-12 Jun 2005, D. Giraldo-Cañas 3903
(CAUP, COL, HUA)
900 Journal of tl Botanical R h Institut f Texas 2(2)
14. gni mokensis Pilg., Bot. Jahrb. Syst. 51(3-4):419. 1914. (Fig. 10, A-D). vee: EQUATORIAL GUINEA
NDO Po: Grasland (Bergweiden) von Moka im Südosten der Insel, 1200-1800 m, Nov 1911, J. Mildbraed 7102 (uoLorvre: B;
ISOTYPE: US-2941533!).
Eragrostis moritzii Jedwabn., Bot. Arch. 4:328. 1923. Tree: VENEZUELA. MeriDa: in der Alpinen Region, J.WK. Moritz 1570 (ISOTYPE:
US-2941535 fragm.!).
Caespitose annuals. Culms 10-30(-45) cm tall, erect or ascending, sometimes geniculate, branched at the
base and lower nodes, often with a small glandular ring below the glabrous nodes. Leaf sheaths overlapping
below to Y the length of the internodes above, sparsely pilose at the summit and along margins below, the
hairs up to 3 mm long and papillose-based; ligules 0.1-0.3 mm long; blades 3-8(-10) cm long, 2-4 mm
wide, flat, sparsely pilose with papillose-based hairs. Panicles 4-13 cm long, 2.5-7(-10) cm wide, narrowly,
ovate, open, the rachis angled; primary branches 2-7 cm long, ascending, stiff, spreading up to 90° from
the rachises; secondary branches widely spreading; pulvini glabrous, smooth; pedicels 0.5-2.5 mm long,
shorter than the spikelets. Spikelets 3-6.5(-7) mm long, 2-2.5 (-3) mm wide, (4-)10-20-flowered, broadly
ovate-lanceolate, the florets strongly imbricate, plumbeous to reddish-purple; disarticulation acropetal,
with the glumes first, then the lemmas and paleas falling as a unit; glumes 1-1.5 mm long, subequal, ovate,
membranous, keeled, scaberulous along the keel, apex acute; lemmas 1.2-1.6 mm long, 0.6-1.1 mm wide,
orbicular, chartaceous, lateral nerves absent or obscure, apex keeled and obtuse; paleas 1.1-1.4 mm long,
0.6-0.8 mm wide, orbicular, deciduous, winged, scaberulous, apex 3; anthers 0.3-0.5 mm
long, purplish. Caryopses 0.6-0.7 mm long, ovoid, striate and reticulate, flattened on the adaxial surface,
reddish-brown. 2n = unknown.
Distribution and habitat.—Native to western Africa (Clayton 1972), E. mokensis has been reported from
Brazil and Venezuela (Boechat & Longhi-Wagner 2001). Herein, we report it for the first time in Colombia.
This species occurs in rocky areas, along roadsides, and near cultivated fields and in waste areas often as-
sociated with Melinus mu icu P. Beauv. and cue ruderal plants; 1700- 2000 m.
omments.—E is very similar in habit, panicl and
and color to E. unio lees (Retz.) Nees ex Steud. However, E. mokensis can be separated from the latter by
having orbicular lemmas with obtuse apices (verses ovate lemmas with acute apices in E. unioloides), shorter
pedicels (0.5-2.5 mm long versus 2-10 mm long in E. unioloides), somewhat narrower spikelets [2-2.5(-3) mm
wide verses 2-3.4 mm wide], and florets with three stamens (two stamens reported in E. unioloides) [Clayton
72].
Specimen examined. COLOMBIA. Cauca: Highlands of Popayán, 1700-2000 m, 1889, F.C. Lehmann BT-635 (US).
ll spikel tshape
15. Eragrostis e (Kunth) Steud., oe Bot. ed. 2, 1:563. 1840. (Fig. 6, E-H). Poa nigricans Kunth, Nov.
Gen. Sp. 1:159. 18 ) & Schult., Syst. Veg. 2:586. 1817. Tree: ECUADOR: Chillo seca
Apr-May, E M SA. Bonpiand. 2291 (HOLOTYPE: P-Bonpl!; isorvees: BAA-1062 fragm.!, K photo!, LE-TRIN-2371.011, P!, US-
2891495 fragm. ex LE-TRIN! &!, fragm. ex PU).
i icm scabra Phil., Fl. Atacam. 55. 1860. Tree: CHILE. Atacama: prope Paposo, Dec 1851, R.A. Philippi 1051 (notorme: SGO-PHIL
TYPES: Bl, BAA-1089 fragm. ex B!, US 556539 fragm. ex SGO-PHIL 357! & photo!).
pol rahmeri Phil., Verz. Antofagasta Pfl. 88. 1891. Tree: CHILE. Tarapacá: Quebrada de Guaviña, 13 Mar 1885, R.A. Philippi s.n.
359 Uis LOTYPE: SGO-PHIL 359; isoryres: BAA-1080 fragm!, CORD!, SGO-37293, SGO-63537, SGO-62670, US-556538 fragm. ex
SGO-PHIL 3591).
Eragrostis de LN Bot. Arch. 5(3-4):205. 1924. Eragrostis nigricans var. tristis (Jedwabn.) Pilg., Notizbl. Bot. Gart. Paes
11:778. 1933. Tyre: BOLIVIA: ad pa ups 15 Feb 1910, Pflanz 359 (xecrorver: BAA-1101 fragm. ex BI, desig by Peterson
ez Vega, Ann. Missouri Bot. Gard. 94: 2007).
ONDE NM ci Bot. Arch. 5(3-4):202. 1924. Tyre: BOLIVIA: Prope La Paz, 1889, M. Bang 80 [collector erroneously cited as
Rusby in protologue] (HoLoTYrE: B; isorvees: Kl, US-8220651).
Caespitose annuals. Culms (10-)20-50(-80) cm tall, erect, sometimes geniculate below, mostly glabrous
and somewhat shiny below the nodes. Leaf sheaths 1/3-2/3 as long as the internodes, sparsely pilose at the
summit and along the upper margins, the hairs up to 2.5 mm long, j the margins glabrous; ligules
0.4-0.6 mm long, ciliate; blades 6.5-10 cm x 2-5 mm, flat, occasionally loosely involute near apex, glabrous
Peterson and Giraldo-Cañas, Eragrostis in Colombia 901
m
Y S e 3S EN
AKTENCERIN 2008 N E
G. t is (B.T. Lehmann 635). A. Habit. B. Ligule C. Spikelet. D. Floret. Eragrosti f (ER. Fosberg 22208). E Habit. F. ligule G
Spikelet. H. Floret.
m P l ^f tha Botanical D h Institut f Texas 2(2)
above and below, somewhat scaberulous near apex. Panicles (5-)7-24 x 2-7(-15) cm, oblong, somewhat
condensed, the spikelets arranged in glomerules that are widely spaced along the channeled rachis, primary
branches mostly 1-10 cm long, sinuous, ascending and spreading 20-90? from the rachises, solitary to
whorled below, scaberulous, secondary branches sinuous; pulvini glabrous to sparsely pilose, the hairs up
to 1.5 mm long; pedicels 0.4-2(-3) mm long, stiffly spreading, divaricate and stout, scaberulous. Spikelets
2.63.8-4.8) x 1-1.2 mm, 2 mm, 2-4(—5)-flowered, linear to narrowly lanceolate, grayish-green to purplish-
green; disarticulation acropetal, with the glumes first, then the lemmas falling, paleas persistent; glumes
1.0-1.4 mm long, subequal, ovate to lanceolate, membranous, keeled, scaberulous along the keel; lemmas
1.6-2.0 mm long, ovate, mou d eon glabrous or with a few scattered hairs, lateral nerves
metimes distinct; apex acute, ; paleas 0.9-1.4 mm long, hyaline, scaberulous along
the keels; apex truncate; stamens 3, anthers 0.3-0.5 mm long, purplish. Caryopses (0.6-)0.7-1.1 mm long,
ovoid, striate and reticulate, shallowly to deeply grooved on the ventral surface, translucent, irregularly
rectangular in cross-section, reddish-brown
Distribution and habitat —Native to the Andean mountains in Argentina, Bolivia, Chile, Ecuador, and
Peru (Nicora 1998; Peterson & Boechat 2001); occurs on rocky slopes, near cultivated fields, and disturbed
roadsides; 1000—4000 m. We report this aan for the first time in Colombia.
Specimens examined. COLOMBIA. $ Juan, 5 km arriba de California, 2733 m ,4Àug 1966, A. Robinson 3032 (US);
Carretera Bucaramanga—Cucuta, 1700 m, 29 Sep 1966, A. Robinson 3125 (US).
1 MR
16. Eragrostis C (Kunth) Trin., Mém. Acad. Imp. Sci. Saint-Pétersbourg, Sér. 6, Sci. Math., Seconde
t. Sci. Nat. 4(2):71. 1836. (Fig. 11 , A-J). Poa pastoensis Kunth, Nov. Gen. Sp. 1:160. 1815 (1816). Hs
(Kunth) Roem. & Schult., Syst. Veg. 2: 587. 1817. Tyee: COLOMBIA: Pasto, Mt. Arand, E Humboldt & A. Bonpland 2149 (HOLOTYPE:
P-Bonpl!; isotyres: Pl, P photo at Kl, US-2767397 fragm. ex P-Bonpl.!).
Poa montufari Kunth, Nov. Gen. Sp. 1:159. 1815 (1816). Eragrosti (Kunth) Steud., Nom. Bot. (ed. 2) 1:563. e id and
montufari (Kunth) Roem. & Schult., Syst. Veg. 2:586. 1817. Tire: ECI JADOR PicHincha: betw. Pi
Apr-May, E Humboldt & A. Bonpland s.n. (HoLorYPE: P-Bonpl.!; IsOTYPES: ae e P!, US-2891497! [fragm.] ex P-Bonpl.!).
Poa olmedoi Kunth, Nov. Gen. Sp. 1:159-160. 1815 (1816). Megastach | & Schult., Syst. Veg. 2:586. 1817. Eragrostis
olmedoi (Kunth) Steud., Nom. Bot. (ed. 2) 1:564. 1840. Tree: oe DURS E ade Ponpand sn (HOLOTYPE: P!).
Poa tenax Kunth, Nov. Gen. Sp. 1:160. 1815 (1816). E i om. Bot. (ed. 2) 1:564. 1840. Megastachya tenax
(Kunth) Roem. & Schult., iim ~ 2:587. 1817. Ter: ECUADOR: PE F Humboldt & A. ond. (HOLOTYPE: Pl; ISOTYPE:
US-2891490 fragm. ex P-Bo
Poa setifolia Benth., Pl. Hartw. 262. 1846. Eragrostis setifolia (Benth.) Steud., Syn. Pl. Glumac. 1:274. 1854, nom. illeg. hom. Tree: ECU-
ADOR: Quito, Hartweg 1452 (uoLorvee: Kl; isotypes: US-2826937 fragm. ex K!, US-2850739 fragm. ex CG! & fragm. ex P!, fragm.
!).
n lehmannii Pilg., Bot. Jahrb. Syst. 27(1—2):32. 1899. Tyre: ECUADOR: Ca. Baños ad flumen Tunguragua, 1800-2500 m, FC.
Lehmann 5283 [error for sane (isoryPe: US-2767406 fragm!).
Eragrostis virescens } lla Hack., Anales Mus. Nac. Buenos Aires 13:505. 1906. Type: ARGENTINA. Tucumán: Stuckert herb. arg.
aa ex "d Herb. arg. 3192€ (HOLOTYPE: W; isoTyPE: US-2942411 fragm.!).
E ienii Hack., Repert. Spec. Nov. Regni Veg. 6:157. 1908. Tree: BOLIVIA. Sup-Yuncas: Simpayas bei Yanacachi, O. Buchtien
428 (HOLOTYPE: W; isorvee: US-773880).
Caespitose perennials with innovations. Culms 30-90 cm tall, erect, sometimes geniculate below, glabrous
and somewhat shiny below the nodes. Leaf sheaths overlapping below, 3/4 as long as the internodes above,
ciliate at the summit and along the upper margins; ligules 0.2-0.5 mm long, ciliate; blades 5-40(-45) cm
x 1-4(-7) mm, involute or flat, glabrous to scaberulous below and scaberulous above, sometimes with
scattered hairs, the hairs up to 4 mm long. Panicles 8-45 x 1-27 cm, ovate, lanceolate to narrowly oblong,
contracted or open, rachis glabrous, the ascending primary branches 1-20 cm long, appressed to widely
spreading up to 80? from the rachises, the branches glabrous to scaberulous, not floriferous near base; sec-
ondary branches composed of loosely overlapping spikelets; pulvini ciliate or glabrous; pedicels 0.5-5 mm
long, erect, mostly appressed, sometimes with hairs. Spikelets 2-6 x 1-1.8 mm, 2-8-flowered, lanceolate
to oblong-ovate, inflated to slightly compressed, plumbeous, sometimes purple-tinged, rachilla sometimes
sparingly ciliate; disarticulation acropetal, with the glumes first then the lemmas falling individually, paleas
persistent; glumes 1-1.9 mm long, subequal, broadly ovate to lanceolate, membranous, sub-hyaline, keeled,
Peterson and Giraldo-Cañas, Eragrostis in Colombia 903
Ci“
Fic. 11. Eragrostis pastoensis (PM. Peterson, CR. Annable & M.E. Poston 8910 & PM. Peterson € E.J. Judziewicz 9268, A—H; P.M. Peterson & N.F. Refulio-
Rodriguez 13982, | & J). A. Habit. B. Ligule. C. Inflorescence. D. Spikelet. E. Floret. F. Lemma, dorsal view. G. Palea, stamen, and lodicule, lateral view.
H. Palea enclosing the stamens, pistil, and lodicules, ventral view. |. Caryopsis, dorsal view. J. Caryopsis, lateral view.
904 Journal of the Botanical R h Institute of Texas 2(2)
scaberulous along the keel; lower glume 1-1.4 mm long, narrow lanceolate; upper glume 1.2-1.9 mm long,
usually broader than the lower; lemmas 1.2-2.0(-2.1) mm long, ovate to broadly ovate, membranous, lateral
nerves obscure, keeled, especially towards the apex, scaberulous along the keel; apex acute; paleas 1-2 mm
long, membranous to partially hyaline, scaberulous along the keels; apex obtuse to truncate; stamens 3,
anthers 0.3-0.6 mm long, reddish-purple. Caryopses 0.4-0.9 mm long, obovoid to prism-shaped, striate
and reticulate, usually with a ventral groove, irregularly rectangular in cross-section, light reddish-brown
to translucent. 2n = 70 (Bowden & Senn 1962, as Eragrostis montufari).
Distribution and habitat.—Native to South America in Argentina, Bolivia, Brazil, Colombia, Ecuador,
Paraguay, Peru, Uruguay, and Venezuela (Nicora 1998; Peterson & Boechat 2001); occurs on dry rocky
hillsides, slopes, pastures, roadsides, barrancas and city sidewalks from 1400-3600 m.
ues un COLOMBIA. Bocorá D.C.: Bogotá, localidad de Usme, alrededores y riberas i río apto en imediaciones
del cas , 2800 m, 16 Sep 2001, D. Giraldo-Cañas 3235 (COL); Bogotá, en las g del Instituto
7 Ciencias Naturales, Campus de la Universidad Nacional de Colombia, 2600 m, 11 Dec 2003, D. Giraldo-Cañas 3616 (COL). Boyacá:
n. Villa de Leyva, camino entre la hacienda “Torcoroma de Arriba” y el cañón de Las Clusias, 2500 m, 18 Jul 2003, D. Giraldo-Cañas
Es 3539 (CORN M Tunja, campus de la Universidad ipie sa de Colombia, 2700 m, 12 Nov 2003, D. Giraldo-Cañas
3608 (COL). C Mun. Nemocón, la vereda Susatá y Nemocón, 2700 m, 4 Nov 2002, D. Giraldo-
Cañas 3320 (COL); Guasca, 1 Aug 1919, B. Ariste-Joseph A-368 (US); region of Bogotá, no date, B. Ariste-Joseph s.n. (US-1040145); Sabana
de Bogotá, entre Sibaté y San Miguel, 2750 m, 15 Aug 1939, J. sree 6640 (COL, US), Salto del Tequendama, 2500 m, 8 Mar 1939,
E.P. Killip 34006 (US), 34016 (US); Hoya del río Checua Loma, 250 m SE de San José, arenas del Cacho, 2640 m, 8 Dec 1966, Schrimpff
117 (COL); Andes de Bogotá, 2700 m, Oct 1856, J.J. Triana s.n. (US-1865282). Nariño: Mun. Sapuyes, El Espino, 3200 m, 15 May 1964,
L.E. Mora 2984-B (COL, PSO); Mun. Piedrancha, arriba de El Guabo, 2300 m, 15 May 1964, L.E. Mora 3005 (COL, PSO); Mun. Pasto,
corregimiento Chachagút, 2000 m, 1 Jun 1989, B.R. Ramírez 1579 (COL, PSO). Norte de Santander: pe 23 Mar 1935, WA
Archer 3232 (US); highway between Pamplona and Málaga, 24 Mar 1935, W.A. Archer 3241 (US); vici , 2300-2400 m, 27
Feb 1927, E.P. Killip & A.C. Smith 19782 (US). Santander: vicinity of California, 2000 m, 11-27 Jan 16 1927, E.P. Killip & A.C. Smith 16846
(US). Unknown department: J.C. Mutis 5518 (US), J.J. Triana 292 (US).
17. Eragrostis pectinacea (Michx.) Nees, Fl. Afr. Austral. Ill. 406. 1841. var. pectinacea (Fig. 12, A-E).
Poa pectinacea Michx., Fl. Bor.-Amer. 1:69. 1803. Tyre: U.S.A. Irumors: Michaux s.n. (HOLOTYPE: P-MICH!; isorvee: US-2851264 fragm.
ex P).
Eragrostis diffusa Buckley, Proc. Acad. Nat. Sci. nein 14:97. 1862. cds purshii var. diffusa (Buckley) Vasey, Contr. U.S. Natl.
Herb. 1 La 59.1 d Tere: U. a Texas: S.B. B H designated by Hitchcock, Man. Grasses U.S. 849. 1935, without
bari US-9162 D
Caespitose annuals Wa ssi pits. Culms 10-80 cm = erect to geniculate or decumbent below,
glabrous. Leaf sheath ping below, 2-3/4 as long as tl bove, hirsute at the apices, hairs to
4mm long; ligules 0.2-0.5 mm long blades 2-20 cm x 1-4.5 mm, flat to involute, abaxial surfaces glabrous
and smooth, adaxial surf: idul Panicles 5-25 x 3-12(-15) cm, ovoid to pyramidal, usually open,
sometimes contracted, primary branches 0.6-8.5 cm long, appressed or slightly diverging to 20° from the
rachises, solitary or paired at the lowest 2 nodes; pulvini glabrous or sparsely hairy; pedicels 1-7 mm long,
flexible, appressed to widely divergent, sometimes capillary. Spikelets 3.5-11 x 1.2-2.5 mm, 6-22-flowered,
linear-oblong to narrowly lanceolate, plumbeous, yellowish brown, or dark reddish purple; disarticulation
acropetal, paleas persistent; glumes 0.5-1.7 mm long, subequal, subulate to ovate-lanceolate, hyaline; lower
glumes 0.5-1.5 mm long, at least 1/2 as long as the adjacent lemmas; upper glumes 1-1.7 mm long, usually
broader than the lower glumes; lemmas 1-2.2 mm long, ovate-lanceolate, hyaline to membranous, grayish
green proximally, reddish purple distally, lateral veins moderately conspicuous; apex acute; paleas 1-2 mm
long, hyaline to membranous, keels scabridulous; apex obtuse; stamens 3; anthers 0.2-0.4 mm long, pur-
plish. Caryopses 0.5-1.1 mm long, rectangular-prismatic, slightly laterally flattened, striate and reticulate,
rectangular with nearly equal sides in cross-section, brownish. 2n = 60 (Koch 1974; Davidse 1981).
Distribution and habitat.—Native to the Americas; found in North America, Central America, the Ca-
ribbean, and most of South America (not known from Chile); it grows in disturbed sites such as roadsides,
railroad embankments, city sidewalks, gardens, and cultivated fields; 50-2600 m (Peterson & Boechat 2001).
Vernacular name.—“Hierba canto" (Huila, Giraldo-Cañas 3921).
905
Peterson and Giraldo-Cañas, Eragrostis in Colombia
V
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Fic. 12. Eragrostis pecti (PM. Peterson & N.F. Refulio-Rodriguez 13981, D & E). A. Habit. B. Spikelet. C. Fl p
Caryopsis dorsal view. E Caryopsis, lateral view E is pilosa (S. Llatos 1107, ) &K). F.P f cul ith alandul G. Culm. H. Inflorescence
l. Spikelet. J. Caryopsis, dorsal view. K. Caryopsis, lateral view.
3
" 2 lafehkn Datanizal D h Institut f Texas 2(2)
d. COLOMBIA. Antioquia: Mun. Sabaneta, pur central de la municipalidad, 1550 m, 3 Jan 2003, D. Giraldo-
oe 3427 (COL); Valle de Aburrá, Mun. Itagúí, Į jue [ pal del Barrio Simón Bolívar, a e iiie D. Giraldo-Cañas 3819
(COL); Valle de Aburrá, Mun. Envigado d 1 del “Almacenes Éxito”
1550 m, 4 Dec 2004, D. Giraldo- Cañas 3820, 3822, 3823 (COL, US); Medellín, 1500 m, 11 Jun 1930, W.A. Arder 102 (US); Mun. Medellín,
predios de la Universidad de Antioquia, 1450 m, 25 Sep 1986, R.W. Pohl 15601 (HUA, MO, US). Bogotá D. C.: Bogotá, unidad deportiva
El Salitre, área peatonal, Calle 63 con Avenida 68, 2600 m, 26 Jul 2005, D. Giraldo- Eu d E HUA, va Bogotá, La Picota, 2600
m, Feb 1934, E. Pérez-Arbeláez 4755 (COL, US). Boyacá: Mun. Santa María, 1, 900 m, 27 Sep 2007,
D. Giraldo-Canas 4121 (COL); Soatá, dry eastern slopes, 1800 m, 17 Sep 1938, J. d ped (COL, vs; 2 a wes o towards
Tipacoque, 1800 m, 3 Nov 1985, J.R.i. Wood 5132 (COL). Caquetá: Mun. Florenci
27-30 Jun 2005, D. Giraldo-Cañas 3962 (COAH, COL, US). Casanare: Mun. El ed en áreas urbanas del centro de la ciudad, 400 m,
= Oct s D. uaa © Cañas 4132, 4135 (COL). Cauca: Mun. Popayán, predios internos y jardineras abandonadas de la Facultad de
del Cauca, 1750 m, 7-11 Nov 2004, D. Giraldo-Cañas 3802 (COL). Chocó: Mun. Quibdó, en inmediaciones
de la penitenciaría municipal, calle 26 con carrera 10, 90 m, 6 Nov 2005, D. Giraldo-Cañas 3969, 3970 (COL, HUA). Cundinamarca:
Entre d aa i m, Oct idi H. de -Barriga 2417 (COL, US); Provincia de Río Negro, Mun. Pacho, carretera Pacho-La
Capilla, l 1, 1600 m, 13 Jan 2004, D. Giraldo-Cañas 3724, 3727, 3732 (COL); Mun. Nilo, hacienda
La Cidia río Pagüey, 350 m, 16 Oct 2004, D. Giraldo- Cañas 3775, ie OE Tocaima, Dec 1932, E. Pérez-Arbeláez 2418 (COL, US).
Huila: Mun. San Agustín, San Agust principal del parque arqueológico, 1700 m, 1-5 Jul
2005, D. Giraldo-Cañas 3919 EN (COD; Man San ree n, atau poa de San Agustin, andenes del Centro Ada del
parque arqueológico, 1700 m, 1-5 Jul 2005, D. Giraldo-Cañas 3921 (COL); Mun. Timaná, en p de E andenes del casco urbano,
1500 m, 1-5 Jul 2005, D. Giraldo-Cañas 3926 (COL). Meta: Mun. Villavicencio, Los Centauros, 400
m, 10 Nov 2002, D. Giraldo-Cañas 3341 (COL, US). Ano Mun. Quimbaya, eae principal scere Dav inen 1500 m, 20-25
May 2005, D. Giraldo-Cañas 3889 (COL, HUA). S fun. Barbosa, aceras, cerca de la Plaza Principal,
1700 m, 20 Jul 2003, D. miel Cañas 3563 (COL). Tolima: Cordillera e andina MR Ibagué, jardi andenes de la Plaza de
Bolívar, 1200 m, 11-12 Jun 5, D. Giraldo-Cañas 3910 (COL, HUA, US). V: del C Estación Piedras, entre Obando y Cartago,
980 m, 5 Feb 1961, J.M. ps 4249 (COL).
18. o E E ; P Beauv. bus ds A 71:162, 175. 1812. Hap m (Fig. 12, F-K). Poa pilosa
8.1753. T ajus, locustis, p oribus” in Scheuchzer, Agrostographia:
193, t. 4, 3. 1719 as dand by Du Puy et al., Fl. nm 472. 1993). dE ITALY, 9-10 Aug 1902, A. Kneucher,
Gram. Exsicc. XII, 344, (errrvre: B! designated by H. Scholz in Cafferty et al., Taxon 49:256. 2000; US-557051!)].
Caespitose annuals. Culms 8-45(-70) cm tall, erect or geniculate spreading below, glabrous or occasionally
with a few glandular pits. Leaf sheaths overlapping below, about 1/2 to as long as the internodes above, cili-
ate at the summit and collar or glabrous, the hairs up to 3 mm long; ligules 0.1-0.5 mm long, ciliate; blades
2-15(-20) cm x 1-2.5(-4) mm, flat, scaberulous above and glabrous below with a few hairs near the base.
Panicles 4-20(-28) x 2-15(-18) cm, ellipsoid to ovoid, open, diffuse, the ascending, capillary often droop-
ing, primary branches 1-10 cm long, spreading 10-80?(-110?) from the rachises, usually whorled on the
lowest two nodes, glabrous to scaberulous; pulvini glabrous to occasionally sparsely ciliate; pedicels 0.8—10
cm long, erect to flexuous, appressed to spreading, scaberulous. Spikelets (2-)3.5-6(-10) x 0.6-1.3(-1.8)
mm, (3-) 5- to 17-flowered, linear-oblong to narrowly ovate, plumbeous; disarticulation acropetal with the
glumes first then the lemmas and paleas falling individually, paleas easily deciduous; glumes 0.3-1.2 mm
long, narrowly ovate to lanceolate, hyaline, keeled, scaberulous along the keel; lower glume 0.3-0.6(-0.8)
mm long; upper glume 0.7-1.2(-1.4) mm long, usually broader than the lower; lemmas 1.2-1.8 mm long,
ovate-lanceolate, hyaline to membranous, grayish green dis and reddish purple near the apex, keeled,
scaberulous along the keel near apex, lateral ; apex acute; paleas 1-1.6 mm long, hyaline
to membranous, scaberulous along the keels; apex obtuse: stamens 3, anthers 0.2-0.3 mm long, purplish.
Caryopses 0.5-1 mm long, obovoid to prism-shaped, dorsally flattened, smooth to striate, rectangular in
cross-section, light brown. 2n = 20, 36, 40 (Bir € Sahni 1988).
Distribution and habitat.—Native in Europe, naturalized in North, Central, and South America (excluding
Surinam); occurs in disturbed habitats and along forest margins in sandy or gravelly sites and city sidewalks;
0-1700 m (Peterson & Boechat 2001).
Vernacular name.—"Maleza del arroz" (Tolima, F.A. Montealegre 11).
d paa Antioquia: Medellín, 1500 m, 2 Jul 1930, W.A. Archer 317 (US); Mun. Urrao, carretera a La
10 1 l , 1650 m, 10 Sep 1986, R.W. Pohl & J. Betancur 15478 (HUA, MO, US); vicinity of Medellín, n
r
carnación,
Peterson and Giraldo-Cañas, Eragrostis in Colombia 907
1927, R.A. Toro 264 (US). Arauca: Mun. Arauca, i diaci del i de la Universidad Nacional de Colombia, hacienda
E] Cairo, carretera Arauca-Tame, km 9, 200-300 m, 13 Jun 2003, D. Giraldo-Cafias 3493 (COL). Bolívar: along the Mompós-Juana
Sánchez trail, Island of Mompós, lands of Loba, Apr-May 1916, H.M. Curran 244 (US), 246 (US). Casanare: Mun. El Yopal, en áreas
urbanas del centro de la ciudad, 400 m, 31 Oct 2007, D. Giraldo-Cañas 4131, 4138 (COAH, COL). Cundinamarca: Mun. Nilo, hacienda
La Guaira, río Pagüey, 350 m, 16 Oct 2004, D. Giraldo-Cañas 3772 (COL); between Anolaima and Cachipay, 17 Apr 1935, W.A. Archer
3328 (US). La Guajira: Pájaro, 10 m, 28 Nov 1959, J. Cuatrecasas & R. Romero-Castafieda 25474 (COL, US); Clausura Napaipa, rumbo a
Maicao, 4.5 km de Uribia, 13 Feb 1963, C. Saravia 2224 (COL, US). a ae meus ira 1898-1899, H.H. Smith 2150 (COL, MO,
US). Meta: Mun. Villavicencio, carretera Villavicencio-Aeropuerto, pi illera Oriental, sitio La Arenera, ca. 2 km del
puente sobre el río Guatiquía, ca. 400 m, 10 Nov 2002, D. Giraldo- Cañas 3342-A (COL), 14-15 Dec 2002, D. Giraldo-Cañas 3388 (COAH,
COL, HUA). Nariño: Mun. Pasto, Panamericana, puente del Juanambú, 1000 m, 20 May 1989, B.R. Ramírez 1489 (COL, PSO). Quindío:
Mun. Armenia, Avenida Bolívar, Parque de La Vida, 1500 m, 29 Jul 2007, D. Giraldo-Cañas & J.C. Ospina 4095 (COL, HUA). Santander:
acs Apropecuaria Piedecuesta, al sur de Bucaramanga, 1000 m, 6 Oct 1966, A. Robinson 3165 (US); Mesa de los Santos, al sur de
ucaramanga, 1700 m, 30 Dec 1966, A. Robinson 3207 (US). Tolima: Ibagué, vereda Chucuni, finca Reinoso, 1100 m, 2 Jul 2006, F.A.
qui 11 (COL). Valle del Cauca: Cartago, Santa Ana de los Caballeros, 950 m, 19 Nov 1946, J. Cuatrecasas 23023 (US), 23037
(COL, US); between Uribe and Sevilla, 1100 m, 2 Nov 1983, J. R.I. Wood 4084 (COL).
19. cn polytricha Nees, Fl. Bras. Enum. Pl. 2:507-508. 1829. (Fig. 8, F-H). Poa e (Nees) Kunth,
m. Pl. 1:331. 1833. Tire: BRAZIL: Sellow s.n. (HoLoTYPE: B; isotypes: BAA-2668 fragm. ex B!, US-77386 fr.
Eragrostis lugens var. villosa Doll, Fl. Bras. 2(3):140. 1878. Type: BRAZIL. Minas Gerais: Caldas, 28 Mar 1868, Exp. Regnell III 1405 (LECTOTYPE:
S designated by Boechat & Longhi-Wagner, Iheringia, Bot. 55:117. 2 USD.
Peg fran Hitche., Amer. J. Bot. 2:308. 1915. Eragrosti florid Hitcl laa Ann. Missouri Bot. Gard.
64:328. 1977. Tyre: U.S.A. FLoriDa: Hillsborough Co., near Tampa, Mar, Curtiss 3494 (HoLoTYPE: US-726520! : BR,EI E
M, nee NY!, PH, TAES, TENN, US-748356!, US-823092!).
llen, Fieldiana, Bot. 28(1):18. 1951. Type: VENEZUELA. BoLtfvar: Gran Sab between Kun and Uaduara-parú, in
pue ab Río Kukenán, S of Mount Roraima, 1065-1220 m, 1 Oct 1944, J.A. oum 59062 (notorvee: US-1911657!; ¡SOTYPES:
E VEN
Caespitose perennials with innovations, without rhizomes, not glandular. Culms 30-62 cm tall, erect,
glabrous and shiny below the nodes. Leaf sheaths sometimes densely pilose dorsally and on the collars,
margins and apices hairy, hairs to 5 mm; ligules 0.2-0.4 mm long; blades 5-20(—33) cm long, 1-3.5 mm
wide, involute to flat, both surfaces with scattered hairs, adaxial surfaces densely hairy behind the ligules,
hairs to 7 mm long. Panicles 15-25 cm long, 5-27 cm wide, ovate, open; primary branches 0.6-15 cm long,
diverging up to 90? from the rachises, capillary, naked basally; pulvini hairy, hairs to 8 mm long; pedicels
1.4-10(-16) mm long, divergent. Spikelets (2.5-)3—5 mm long, 1.1-1.6 mm wide, 4—9-flowered, narrowly
lanceolate to linear-oblong, plumbeous; disarticulation acropetal, paleas persistent; glumes broadly ovate
to narrowly lanceolate, hyaline to membranous; lower glumes 1.1-1.6 mm long; upper glumes 1.2-1.8 mm
long; lemmas 1.2-1.8 mm long, broadly ovate, membranous throughout, lateral veins inconspicuous, apices
acute; paleas 1.1-1.7 mm long, membranous to hyaline, narrower than the lemmas, apices obtuse; stamens
3, anthers 0.3-0.5 mm long, reddish-purple. Caryopses 0.5-0.8 mm long, obovoid to somewhat prism-
shaped, laterally compressed, with a well-developed adaxial groove, finely striate, opaque to translucent,
reddish-brown. 2n = 60, 80
Distribution and habitat —Eragrostis polytricha grows in sandy and rocky areas, usually in open savannas;
0—2650 m. Eragrostis polytricha ranges from México through Central America to Argentina, Bolivia, Brazil,
Chile, Colombia, Guyana, Paraguay, Uruguay, and Venezuela (Boechat & Longhi-Wagner 2001).
Specimens examined. COLOMBIA. Cundinamarca: Hoya del río Checua Loma, 250 m SE de San José, arenas del Cacho, 2640 m, 8
Dec 1966, Schrimpff 122 (COL).
20. Eragrostis prolifera (Sw.) Steud., Syn. Pl. Glumac. 1:278. 1854. (Fig. 13, A-D). Poa prolifera Sw., Prodr. 27.
1788. Tree: GUADELOUPE: Swartz s.n. (HOLOTYPE: S; ISOTYPE: K!)
Eragrostis salzmannii Steud., Syn. Pl. Glumac. 1:277. 1854. Tyre: BRAZIL. Banta: P Salzmann s.n. (isotypes: K-photo!, US-9119011, US-
27674141)
Caespitose perennials with innovations, without rhizomes, not glandular. Culms 85-130(150) cm tall, stiffly
erect, glabrous below the nodes. Leaf sheaths glabrous or hairy at the apices, hairs to 4 mm long; ligules
908 J lof the Botanical R h Institute of Texas 2(2)
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Fic. 13. Eragrostis prolifera. A. Habit. B. Culm. C. Spikelet. D. Floret. Eragrostis tenuifolia (PM. Peterson & 0. Tovar 14049, E—G; P. Nuñez € V.&E. Bengoa
8731, H—J). E. Habit. F. Spikelet. G. Floret. H. Caryopsis, dorsal view. I. Caryopsis, ventral view. J. Caryopsis, lateral view.
Peterson and Giraldo-Cañas, Eragrostis in Colombia 909
0.1-0.2 mm long; blades 25—50 cm long, 1.5-6 mm wide, flat to involute, glabrous abaxially, scabridulous
adaxially, sometimes also with a few scattered hairs near the base. Panicles (10)20—50(60) cm long, 2-8(10)
cm wide, narrowly ovate, contracted to open; primary branches mostly 2-14 cm long, appressed or diverg-
ing up to 50°(-90°) from the rachises, spikelets congested near the base of the branches; pulvini glabrous;
pedicels 0.3-2.4 mm long, appressed, always shorter than the spikelets. Spikelets 3.2-10(12) mm long,
0.7-1.4 mm wide, (5)8-25-flowered, linear-lanceolate, stramineous to plumbeous, sometimes with a reddish
tinge; disarticulation acropetal, glumes first, then the lemmas, paleas persistent; glumes subequal, ovate to
lanceolate, hyaline; lower glumes 1-1.5 mm long; upper glumes 1.1-1.6 mm long; lemmas 1.1-1.82) mm
long, ovate, membranous, apices acute; paleas 0.8—1.7 mm long, hyaline, narrower than the lemmas, apices
obtuse to truncate; stamens 2, anthers 0.2-0.3 mm long, purplish. Caryopses 0.6-0.9 mm long, ovoid,
flattened ventrally, finely striate, reddish-brown. 2n = 40.
Distribution and habitat. —Eragrostis prolifera grows on beaches, in brackish water, and along roadsides,
at elevations below 20 m. Its range extends southward from U.S.A. through México and Central America
to Brazil, Colombia, and Venezuela (Boechat & Longhi-Wagner 2001). Boechat and Longhi-Wagner (2001)
reported E. prolifera as occurring in Peru but no specimens were seen by PMP in a treatment of Eragrostis in
Peru (Peterson & Sanchez Vega 2007).
Specimens examined. COLOMBIA. Atlantico: Puerto Colombia, a ae Nov 1912, A.S. Hitchcock 9931 (US). Bolivar: Cartagena
and vicinity, Dec 1928, B. Elias 649 (COL, US); Cartagena, Mamonal, lecl 0—20 m, 16 Nov 1990, R. Álvarez 12 (COL)
Boca Grande, near Cartagena, 0—5 m, 3 Nov 1926, E.P. Killip & A.C. Smith 14097 (COL, US), 14103 (MO, US); Cartagena, sand beach, 20
Nov 1912, A.S. Hitchcock 9900 (US), 9902 (US); Cartagena, 0—2 m, 1-3 Oct 1922, F.W. Pennell 12013 (US); Costa Caribe, Alrededores de
Cartagena, 5 m, year 1942, H. Apolinar 392 (COL); Islas del Rosario, 2 m, 14-17 Oct 1963, P. Pinto 685, 687, 701 (COL); Cartagena, playas
de Marbella, 0—5 m, 3 Feb 1962, C. Saravia & D. Johnson 31, 66 (COL). San Andrés, Providencia y Santa Catalina: Santa Catalina
Island, near the village, 0-5 m, 5 Feb 1985, J.R.I. Wood 4729 (COL). Sucre: Mun. Tolú, arroyo Amansaguapos, glofo de Morrosquillo,
Manglares, 0—5 m, 18 Sep 1990, J. Betancur & M. Berrío 1970 (COL, HUA, MO).
21. Eragrostis rufescens Schrad. ex Schult. var. rufescens, Mant. 2:319. 1824. (Fig. 10, E-H). den
inconstans var. rufescens (Schrad. ex Schult.) Nees, Fl. Bras. Enum. Pl. 2(1):495. 1829. Tree: BRAZIL: Maximilian Neowidens s
(HOLOTYPE: LE; isorvee: BAA-1087!).
Eragrostis polyneura Jedwabn., Bot. Arch. 5(3-4):205. 1924. Tree: BRAZIL: A.EM. Glaziou 18559 (soryre: US-79710 fragm.!).
Eragrostis acicularis Trin., Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math. 1(4):406. 1830. Tree: BRAZIL. PARANA: in arenosis ad
ripas fluv., Rio Jacaré, Dec, Riedel 1140 (moLorvrE: LE-TRIN-2299.01!; isorYPES: K, LE, P, SI, US-28914421, US-2765424 fragm.!).
Bi mn Salzm. ex Steud., Syn. Pl. Glumac. 1:277. 1854. Tyre: BRAZIL. Banta: in maritimis, 1838, T. Salzmann s.n. (HOLOTYPE:
-2572.015; isotypes: K, LE-TRIN-2572.02!, MO, US-2767409!, US-911819!, US-1614948)).
mh s. . Moore, Trans. Linn. Soc. London, Bot. 4:511. 1895. Tre: BRAZIL. BrasiLia: Hab. in cacumine montium Serra de
da prope Santa Anna de Chapada, Matto Grosso Exped, Spencer Le M. Moore, Botanist 131 (notorvee: BM).
Caespitose annuals with innovations, not glandular. Culms 5-45(-70)-35 cm tall, erect or decumbent,
often geniculate near base, glabrous below the nodes. Leaf sheaths shorter than the internodes, hairy at the
apices, hairs to 4 mm long; ligules 0.1-0.3 mm long; blades 6-20 cm long, (1-)23.5(-4) mm wide, flat to
involute, glabrous abaxially, scabridulous adaxially and with scattered hairs, the hairs 2-4 mm long and
more numerous near base. Panicles (3-)5-19 cm long, 1-6.5(-8.5) cm wide, ovate to narrowly oblong, con-
tracted to open; primary branches mostly 2-8 cm long, appressed or diverging up to 50? from the rachises;
pulvini hairy to glabrous; pedicels 1-2 mm long, appressed, always shorter than the spikelets. Spikelets
(526-15(-21) mm long, 1.5-2.2-3) mm wide, 10-43-flowered, linear-lanceolate, imbricate, stramineous
to plumbeous to reddish-purple tinged; disarticulation acropetal, glumes first, then the lemmas, paleas
persistent; glumes 1.4-2.6 mm long, subequal, narrowly lanceolate, membranous, 1-keeled, 1-nerved; lower
glumes 1.4-2.5 mm long; upper glumes 1.5-2.6 mm long; lemmas 1.6-2.3 mm long, ovate-acuminate,
chartaceous to coriaceous, lateral nerves evident, apices acuminate; paleas (0.8—)1.2-1.5 mm long, hyaline,
narrower than the lemmas, apices obtuse to truncate; stamens 2, anthers 0.2-0.3 mm long, reddish-brown
to dark-purplish. Caryopses 0.4-0.6 mm long, ovoid, smooth to finely striate, light reddish-brown with a
dark mark near base of the embryo. 2n = 60 (Davidse 1994).
Distribution and habitat.—Eragrostis rufescens var. rufescens is an American species and is distributed
910 J lof the Botanical R h Institute of Texas 2(2)
from Mesoamerica to Bolivia, Brazil, Paraguay, and Venezuela (Boechat & Longhi-Wagner 2001). This spe-
cies grows in savanna grasslands in open, sandy areas along rivers and in flats that are periodically flooded
by water, along roadsides, city sidewalksm, and cultivated fields; 1500-2500 m. This is the first report of
E. rufescens for the Colombian flora.
Comments.—Morphologically, E. rufescens is very similar to E. pectinacea. Nevertheless, E. rufescens has
longer glumes (1.4-2.6 mm long versus 0.5-1.7 mm long in E. pectinacea), only two stamens (versus three
in E. pectinacea), and ovoid caryopses (versus rectangular-prismatic in E. pectinacea).
Specimens examined. COLOMBIA. Antioquia: noa 27 Apr 1927, R.A. Toro 243 (US). Boyacá: Cordillera Oriental, along Río
Soapaga, 12 km E of Belén, 2460 m, 7 Nov 1944, F.R. Fosberg 22208 US).
22. Eragrostis secundiflora J. Presl, Reliq. Haenk. 1 (4-5):276. 1830. subsp. secundiflora (Fig. 6, I-K).
Poa secundiflora (J. Pres!) Kunth, Enum. Pl. 1:342. 1833. Tyre: MÉXICO: T. Haenke s.n. (HoLoTveE: PR; isotypes: photo K!, LE, MO-
1237641, US-79720 fragm. ex PR!)
Eragrostis compacta a ex ~ Syn. Pl. Glumac. 1:275. 1854. Tyre: BRAZIL. Banta: P Salzmann s.n. (noLoTYPE: Pl; sotyres: K!, MO!
1749!, US-9117
Eragrostis yucatana L.H. Harv. Bull. Torrey Bot. Club 81(5):406. 1954. Tyre: MEXICO. Yucatan: near Progreso, 11-15 Aug 1932, J.R.
Swallen 2933 (HoLoTYPE: US-15371941
Caespitose perennials with innovations, not glandular. Culms 30-75 cm tall, erect, glabrous below. Leaf
sheaths overlapping below, Ya as long as the internodes above, mostly glabrous, hairy at the apices, hairs to
4 mm long; ligules 0.2-0.3 mm long; blades 10-25(-40) cm long, 1-5 mm wide, involute, glabrous abaxi-
ally, scabridulous adaxially, sometimes also sparsely pilose. Panicles (3-)5-30 cm long, 1-15 cm wide, from
narrowly oblong, glomerate, and interrupted below to ovate and open; primary branches 0.5-12(-16) cm,
appressed or diverging up to 40? from the rachises, stiff; pulvini glabrous or sparsely hairy; pedicels 0—1(-3)
mm, appressed, flattened. Spikelets 6-16(-23) cm long, 2.4-5 mm wide, ovate to linear-elliptic, flattened,
stramineous, with reddish-purple margins or completely reddish-purple, with 10-45 florets; disarticulation
basipetal, florets falling intact and before the glumes; glumes ovate-lanceolate to lanceolate, membranous;
lower glumes 1.7-3 mm long; upper glumes 2.2-4 mm long, apices acuminate; lemmas 2-6 mm long, ovate,
membranous to leathery, apices usually acuminate or attenuate, sometimes acute; paleas 1.5-3 mm long,
membranous to leathery, narrower than the lemmas, apices obtuse, sometimes bifid; stamens 2, anthers
0.2-0.5 mm long, brownish. Caryopses 0.8-1.3 mm long, ellipsoid, somewhat laterally flattened, smooth,
reddish-brown. 2n - 40.
Distribution and habitat.—Eragrostis secundiflora subsp. secundiflora occurs throughout México, and in
South America it is found in Bolivia, Brazil, Guyana, Peru, and Venezuela. This species grows in sandy soils,
dunes, grasslands, beaches, and roadsides; 0-1700 m
Al ls Ev. RH d rid fl FP |
Specimens examined. COLOMBIA. Casanare:
ca. 200 m, 12 Jul 1963, J. Blydenstein & C. Saravia 1365 (COD. Meta: Al sur del río G hacienda Gibraltar, en terraza al ca.
200 m, 9 Sep , J. Blydenstein 1590 (COL). Santander: Mesa de los Santos, 1500 m, 11-15 Bes 1926, E.P. Killip & A.C. Smith 15241 (US);
Mesa de los Santos, al sur de Bucaramanga, 1700 m, 30 Dec 1966, A. Robinson 3188 (US).
23. Eragrostis tenella (L.) K Beauv. ex Roem. & Schult., Syst. Veg. 2: e 1817. (Fig. 5, F-D. Poa tenella L.,
: 69. 1753. M lla (L.) Bojer, Hortus Maurit. 369. 1837. Tyre: INDIA: A Herb. Linn. 87.33-
LINN! designated by velero: Blumea 47:164. 2002, IDC microfiche US!).
Poa amabilis L., Sp. Pl. 68. 1753. Eragrostis amabilis (L.) ions = Arn., Bot. Beechey Voy. 251. 1838, hom. illeg. Megastachya amabilis (L.)
P. Beauv., Ess. Agrostogr. 74, 167, 173. 1812. Cynod is (L.) Raspail, Ann. Sci. Nat., Bot. 5:302. 1825. Tyre: SRI LANKA: Herb.
Hermann 2: 59, no. 46 [Lecrorypz: BM! n designated i sess in Cafferty et al., Taxon 49:254. 2000, IDC microfiche US
Poa plumosa Retz., Observ. Bot. 4:20. 1786. E ) Link, Hort. Berol. 1:192. 1827. Eragrostis tenella var. peus (Retz.)
Stapf, Fl. Brit. India 7(22):315. 1897 yes Erüeiostis amabilis var. plumosa din ) E.G. Camus & A. Camus, Fl. Indo-Chine 7: 557.
1923. Tyre: INDIA: E. Tranquebaria, König s.n. (HOLOTYPE: LD; isorvee: K fragm
Eragrostis Ms var. patens Chapm. ex Beal, Grasses N. Amer. 2:479. 1896. Tyre: U.S.A. Georcia. Wayne Co.: Doctortown and Jesup,
Sep-Oct 1880, A.H. Curtiss 3493 (HoLorYrE: MSC; isotypes: NY!, US-748296!, US-821866!, US-909340!).
Caespitose annuals. Culms 5-40(-50) cm tall, erect to spreading, mostly glabrous and occasionally with
Peterson and Giraldo-Cañas, Eragrostis in Colombia 911
oblong glandular areas below the nodes. Leaf sheaths 1/2 as long as the internodes, ciliate at the summit,
collar, and along margins of the upper portion, the hairs to 4 mm long, stiff; ligules 0.2-0.3 mm long, ciliate;
blades 2-8 cm x 2-4 mm, flat to involute, glabrous below and scaberulous above, occasionally with a few
scattered papillose-based hairs below. Panicles 4-15 x 1-7 cm, open, narrowly ovate, primary branches 0.5-4
cm long, sometimes with irregular glandular areas below the branch bases, branches diverging 20-100? from
the rachises; pulvini ciliate or glabrous; pedicels 0.8-4(-7) mm long, mostly pendant, drooping to erect.
Spikelets (1-)1.5-2.2 x 0.9-1.2 mm, 4-8-flowered, ovate to oblong, reddish-purple to greenish; disarticu-
lation between the florets with a portion of the rachilla; glumes 0.4-1.1 mm long, unequal, ovate, hyaline,
keeled, the nerve commonly green, scaberulous along the keel; lower glumes 0.4-0.7 mm long; upper glumes
0.7-1.1 mm long; lemmas 0.7-1.1 mm long, ovate to broadly oblong, membranous, lateral nerves evident,
usually greenish, strongly keeled, scaberulous along keel; apex truncate to obtuse; paleas 0.6-1.1 mm long,
hyaline, keels ciliate, the cilia 0.3-0.5 mm long; apex obtuse to truncate; stamens 3, anthers ca. 0.2 mm
long, purplish. Caryopses 0.3-0.5 mm long, ellipsoid, faintly striate, elliptical to circular in cross-section,
translucent, light brown. 2n = 20 (Baquar & Saeed 1969)
Distribution and habitat.—Native in India; introduced in U.S.A., Mexico, Central America, Caribbean,
Brazil, Bolivia, Ecuador, Guianas, Paraguay, Peru, and Venezuela (Nicora 1998; Peterson & Boechat 2001);
occurs in open areas near cultivated fields, dry forests, city sidewalks, and along roadsides; 0-1600(-2700)
m.
Vernacular names.—"Grama ilusión” (Atlántico, Dugand 5866), “Paja ilusión" (Norte de Santander, Car-
vajalino & Díaz 43).
Comments.—Based on unpublished research by Otto Stapf, Bor (1960) pointed out that Eragrostis amabilis
(L) Wight & Arn. is the same species as E. tenella. Since Hooker (1896:315, cited by Koch 1978) was the first
to unite the two, and he used the latter name, E. tenella is the correct name for this species (Koch 1978).
Specimens examined. COLOMBIA. Antioquia: Mun. Medellín, predios internos del Jardín Botánico Joaquín Antonio Uribe, 1550 m,
2 Jan 2002, D. Giraldo-Cafias 3293 (COL, HUA); Valle de Aburrá, Mun. Envigado, creciendo en las grietas de andenes y pavimentos del
estacionamiento del supermercado “Almacenes Éxito," 1550 m, 4 Dec 2004, D. Giraldo-Cañas uae COD: iran o m, 1 des
1930, W.A. Archer 101 (US), Oct 1945, A. Fernández s.n. (US-2765421). Arauca: Mun. Arauca, i i la
Universidad Nacional de Colombia, hacienda El Cairo, carretera Arauca-Tame, km 9, 200-300 m, 13 Jun 2003, D. Giraldo-Cañas 3500,
3507 (COL). Atlántico: Barranquilla, campus de la Universidad del Norte, carretera Barranquilla-Puerto Colombia, km 5, 5-10 m, 11
Oct 2007, D. Giraldo-Cañas 4123, 4127 (COL); Puerto Colombia, 15 Mar 1949, A. Dugand 4308 (COL, US), 21 Nov 1912, A.S. Hitchcock
9935 (US); Barranquilla, barrio Altos del Prado, 50-70 m, 1 Nov 1961, A. Dugand 5866 (COL). Bolívar: Cartagena, near shore, 20 Nov
1912, A.S. Hitchcock 9897 (US). Caquetá: Mun. Florencia, Barrio Villa Natalia, grietas de pavimentos y andenes, ca. 280 m, 27-30 Jun
2005, D. Giraldo-Cañas et al. 3952 (COL). Casanare: Mun. El Yopal, en áreas urbanas del centro de la ciudad, 400 m, 31 Oct 2007, D.
Giraldo- "nin 4129, 4134 (COL). Córdoba: Mun. Santa Cruz de Lorica, centro de Lorica, 150 m, 24 Feb 2005, D. Giraldo- Caras 3857
o Mun. San A carretera San Pelayo-Lorica, en huerta casera de fríjol, 150 m, 26 Feb 2005, D. Giraldo-Cañas 3868 (COL).
C o, hacienda La Guaira, río Pagüey, 300 m, 16 Oct 2004, D. Giraldo-Cañas 3769 (COL); Mun. Girardot, vía férrea
de la ribera del río La Magdalena, 250 m, 17 Oct 2004, D. Giraldo-Cañas 3785 Add Magdalena: Santa
Marta, T 1898— 1901, H.H. Smith 2746 (COL, MO, US). Meta: Mun. Villavicencio, en área urbana, call Los Centauros,
400 m, 10 Nov igne D. Ed Cañas 3340 (COAH, COL). Nariño: Mun. Ipiales, Las Lajas, 2700 m, 7 Aug 1939, H García-Barriga
7841-A (COL). Nort Cúcuta y El Rosario, Feb 1941, Carvajalino & Díaz 43 (COL). Quindío: Mun. Quimbaya, parque
principal, en jardineras, 1500 m, 20-25 May 2005, D. Giraldo-Cañas 3891 (CAUP, COL, HUA). Santander: Bucaramanga, 1100 m, 4
Aug 1944, N. C. Fassett nue ma San Gil, 1250 m, year 1983, J.R.I. Wood 3884 (COL). Tolima: Mun. Flandes, vía férrea entre Flandes
y Girardot La Magdalena, 250 m, 17 Oct 2004, D. Giraldo-Cañas 3791 (COL); Cordillerra Central andina, Mun
Ibagué, jardineras y andenes de la Plaza de Bolívar, ca. 1200 m, 11-12 Jun 2005, D. Giraldo-Cañas et al. 3914 (COL). Valle del Cauca:
Plana del Valle, extremo N, Cartago, malezas en E 980 m, 15 Nov 1946, J. Cuatrecasas zs es US) Vichada: Región
Guayanesa, Mun. Puerto Carreño, afl de Lajas y el Cerro El Bita, ribera d rinoco, 40-100 m, 4-5
Jan 2004, D. Giraldo-Cañas & C. Parra 3625 (COL).
24. Eragrostis tenuifolia (A. Rich.) Hochst. ex Steud., Syn. Pl. Glumac. 1:268. 1854. (Fig. 13, E-J). Poa tenuifolia
A. Rich., Tent. Fl. Abyss. 2:425. 1850 (1851). Tyre: ETHIOPIA: in locis incultis Vallium prope Adoam, 18 Sep 1837, G.H.W. Schimper
92 (LECTOTYPE: P! designated by S. Phillips, Fl. Ethiopia 7:122. 1995; isotypes: GOET-5814, K!, L, WI, WAG, US-1127147 fragm.!).
Caespitose perennials. Culms 25-75 cm tall, erect to geniculate spreading below, glabrous, 2-3 nodes per
culm. Leaf sheaths overlapping below, 3/4 to as long as the internodes above, glabrous, pilose along the
" 1 Lattha Rataniral D h Institut f Texas 2(2)
margins and at the summit, the hairs up to 2 mm long; ligules 0.2-0.3 mm long, ciliate; blades 3.5-20(-30)
cm x 1-3 mm, flat to folded or loosely involute, glabrous to scaberulous above and glabrous below. Panicles
5-20 x 3-10(-12) cm, open, ovate to narrowly pyramidal, with a well developed peduncle up to 18 cm long,
the relatively few-flowered primary branches 0.5-6 cm long, spreading 40-90? from the rachises, solitary,
scaberulous; pulvini ciliate, often reddish, the hairs up to 2 mm long; pedicels 2-15 mm long, erect, capillary
and stiff, rct e Spikelets 6-12(-14) x 1.5-2.2 mm, 6- to 14-flowered, linear, plumbeous; disarticula-
tion acropetal with the gl first then the | falling, paleas mostly persistent; glumes 0.3-1 mm long,
very unequal, hyaline, fi keeled, glabrous; lower alone 0.3-0.6 mm long, subulate to linear-lanceolate;
upper glumes 0.5-1 mm long, lanceolate, usually broader than the lower; lemmas 1.5-2.0 mm long, ovate,
membranous, plumbeous, obscurely keeled, lateral nerves obscure, scaberulous near the apex; apex acute
to obtuse; paleas 1.3-1.8 mm long, membranous, plumbeous, scaberulous along the keels; apex obtuse;
stamens 3, anthers 0.2-0.4 mm long, red to white in age. Caryopses 0.6-1.1 mm long, ovoid, strongly later-
ally flattened, and curved on the adaxial side, striate and reticulate, with a deep ventral groove, narrowly
triangular in cross-section, brownish. 2n = 40 (Pohl & Davidse 1971; Morton 1993).
Distribution and habitat.—Native in North Africa and Asia, naturalized and rapidly spreading in the
New World tropics from México, Guatemala, El Salvador, Honduras, Costa Rica to Argentina, Bolivia, Chile,
Ecuador, Peru, and Venezuela; along roadsides, city sidewalks, soccer fields, and disturbed open areas;
100-3400 m.
Comments.—A population of Eragrostis tenuifolia in the Cordillera Oriental (Choachí, Cundinamarca:
Giraldo-Cañas 3288) has stolons and no other individuals of this species are known to have these struc-
tures.
Specimens examined. COLOMBIA. Antioquia: Mun. Itagüí, barrio Simón Bolívar, carrera 50 con calle 64, 1550 m, 2 Jan 2003, D.
Giraldo-Cañas 3424, 3426 (COL); Mun. Jardín, sitio La sedis addo wean Riosucio, km 5, 1900 m, 5 Jan 2003, D. Giraldo-Cañas
3432 (COL); Mun. Envigado, loma de El Escobero, 2 arretera Las Palmas, 1850 m, 26 Dec 2003, D.
Giraldo-Cañas 3619 (COL); Mun. Andes, aeropuerto, 1200 m, 17 Sep 1986, R.W. Pohl BE Betancur 15523 (COL, HUA, MO, US); Mun.
un vereda Las disse 18 "A oeste n Jardín, 1600 m, 19 Sep 1986, R.W. Pohl & J. Betancur 15550 (COL, HUA, MO, US). Boy-
á: Mun. males, 2500 m, 22 Jun 2003, D. Giraldo-Cañas 3526 (COL); Mun. Villa de Leyva, camino
entre la hacienda “Torcoroma de Arr n y a cañón de Las Clusias, 2500 m, 18 Jul 2003, D. Giraldo-Cañas 3553 (COL); Mun. Tunja,
y
,
campus de la n ersidad a r TE e uu ea d m, 12 Hox ae D. ded Cañas 3609 (COL). Cauca: Mun.
nid del Cauca, 1750 m, 7-11 Nov 2004,
D. Giraldo- Cañas 3810 (CAUP, COL); Popayán, airport, 1750 m, 2 Nov 1955, A. Saa 59 d Mun. Coconuco, 2500 m, Jun 1948, S.
Yepes-Agredo 351 (COL, US). Cundinamarca: Mun. and carretera Choacl hoachí, km 2, ca. 1600 m, 12 Nov 2001,
D. Giraldo-Cañas 3288 (COL); Mun. Soacha, autor ] Sibaté, km 17, en frente de 1 presa Pisos Al 2700 m, 3 Nov 2002,
D. Giraldo-Cañas 3312 (CAUP, COL); Mun. Nemocón S N hacienda Susatá, 2700 m, 4 Nov 2002,
D. Giraldo-Cañas 3317 (COL, HUA); Provincia de Río Negro, Mun. Pacho, carretera Pacho: La Capilla, alrededores del estadio municipal
de fútbol, 1600 m, 13 Jan 2004, D. nid de 3733 (COL); Mun. Suesca, hacienda Susatá, 2650 m, 10 Jul 2004, J. Groenendijk 1813
(COL). Huila: Mun. Pitalito, en griet del casco urbano, 1550 m, 1—5 Jul 2005, D. Giraldo-Cañas et al. 3930 (COL, HUA).
Nariño: Empatí, 1500-2000 m, 30 Apr 1939, A.H.G. Alston 8297 (US); 5 km S of Tangua, 14 Jan 196 T H.B. di uio 2 Pd
retera Pasto-Túquerres, 2600 m, 30 Nov 1962, C. Saravia 1812 (COL). Quindío: Mun. Calarcá, d
1500 m, 20-25 May 2005, D. Giraldo-Cañas 3894 (COL). Santander: Mun. Vélez, inmediaciones de la Feria de Ganado, 1700 m, 21
Jul 2003, D. Giraldo-Cañas 3566 (COL). Tolima: Cordillerra Central andina, Mun. Ibagué, jardineras y andenes de la Plaza de Bolívar,
ca. 1200 m, 11-12 Jun 2005, D. Giraldo-Cañas et al. 3904, 3908 (COL). Valle del Cauca: Calima Valley, NW of Restrepo, 1500 m, 6 Jul
1962, A.R. Bridgeman 50 (US); Palmira, Plana del Valle, predios de la Facultad de Agronomía de Palmira, 1000 m, 15 Jan 1964, López
Filgueiras 8498 (US); Mun. Yumbo, finca Río Grande, 1180 m, 12 Jun 1998, D. Stancik 797 (COL). Vichada: Caño Urimica, 250 m, 19
c 1971, I. Cabrera 1681 (COL)
25. Eragrostis viscosa (Retz.) Trin., Mem. Acad. Imp. Sci. St. Petersbourg Hist. Acad. 1:397 1830. (Fig. 5,
H). Poa viscosa Retz., Obs. Bot. 4:20 (1786). Tyre: INDIA: Malabar, König s.n. (moLorvrE: LD; isorvre: BM!)
Tufted annuals. Culms 13-50 cm tall, erect, sometimes spreading or geniculate at the lower nodes, viscid
below the nodes with a complete or partial ring of yellow glandular areas below the nodes. Leaf sheaths 1/2
the length of the internodes to overlapping, viscid, ciliate at the summit, collar, and along margins of the
upper portion, the hairs up to 4 mm long, stiff; ligules 0.2-0.5 mm long; blades 4-15 cm long, 2-4 mm wide,
Peterson and Giraldo-Cañas, Eragrostis in Colombia 913
flat, mostly glabrous with scattered papillose-based hairs, the hairs up to 4 mm long, the upper surface often
viscid and covered with adherent soil particles. Panicle 6-22 cm long, 2-9 cm wide, open, cylindrical to
ovate-lanceolate; ascending branches 0.5-6.5 cm long, spreading 20-90? from the rachises, viscid; pulvini
in the axils of the primary branches sparsely ciliate or glabrous; pedicels 0.5-5 mm long, erect. Spikelets
(2-)2.5-5.5 mm long, 1.2-2.2 mm wide, 5-9-flowered, ovate, compressed, reddish-purple to greenish-yellow,
rachilla viscid; disarticulation between the florets with a portion of the rachilla; glumes 0.8-1.5 mm long,
ovate to broadly ovate, hyaline to sub-hyaline, keeled, the nerve commonly green, scaberulous along the
keel; lower glumes 0.8-1.3 mm long; upper glumes 1-1.5 mm long; lemmas 1.1-1.8 mm long, ovate the
broadly oblong, membranous, lateral nerves evident, sometimes greenish, keeled, scaberul long the keel;
apex truncate to obtuse; paleas 1.1-1.8 mm long, bowed out, hyaline, uem ue along the MES the
hairs 0.3-0.6 mm long; apex obtuse to truncate; stamens 3, anthers 0.2-0.4 mm long, purplish. Caryopses
0.4-0.5 mm long, ellipsoid, translucent, light brown. Chromosome number 2n = 40, 60.
Distribution and habitat.—This species is distributed from México to Northern South America (Ecuador,
and Venezuela); and is found in disturbed areas near cultivated fields, roadsides, and along river banks;
0-2000 m
1. COLOMBIA. La Guaji Riohacha, 30 m, 30 Nov ei O. Haught d nn US); d de Uribia rumbo a
Mun. 29 Mar 1962, C. Saravia & D Jinn 328 (COL, US). Magd io Cesar, h región El Callao,
hacienda Santa Marta, 200 m, 29-30 Oct 1959, J. Cuatrecasas 6] R. Romero- Coen 24962 (COL, US); Valle del río Cesar, entre Vallito
y Mata de Indio, 60 m, 12 Feb 1961, A. Dugand 5595 (COL). Norte de Santander: Los Estoraques, La Playa, 14 Jan 1974, MJ. Balick 132
(COL); Carretera Cúcuta-Pamplona, La Esperanza, 1400 m, 19 Nov 1949, H. García-Barriga 13259 (COL). Valle del Cauca: Cordillera
Occidental, Yanaconas al Silencio, montaña La Victoria, 1700-2000 m, 5-10 Dec 1962, H. Garcia-Barriga 17610 (COL).
EXCLUDED SPECIES
Eragrostis uad (Kunth) Steud. (= Poa patula Kunth). This species is commonly referred to the Colombian
n local treatments, but the Colombian specimens belong to Ei iin d (pers. obs.). The
recent use of this name in Bolivia (Renvoize 1998), and Ecuador (Jorgensen & León-Yánez 1999),
places this as a synonym of E. tenuifolia. However, after studying the type 2 T ud fragment at US, we
are sure that this taxon is not a synonym of E. tenuifolia. Therefore, we are not including this name in
this treatment.
Eragrostis rd (Retz. 1 nen ex Bu is a unioloides Retz.). This species has been cited for the Antioquia's
Flora (www.mobot sed specimen R. Callejas & A. Echeverri 11494 (MOD), but this col-
ae belongs to aly passus mu ) Matthei.
ACKNOWLEDGMENTS
We wish to thank the curators at BAA, COAH, CAUP, COL, HUA, LE, MO, PSO, SI, US, and USM for their
valuable help; the Smithsonian Institution and Universidad Nacional de Colombia for supporting a research
visit of DGC to the United States National Herbarium; Utah State University for permission to use illustra-
tions that appeared in Flora of North America, Volume 25, prepared by Linda A. Vorobik and Cindy Roché;
Alice R. Tangerini for preparing illustrations and for preparing all electronic plates; and Victor L. Finot and
Emmet J. Judziewicz for reviewing the manuscript. This article contributes to "Estudios sistemáticos en
gramíneas de Colombia: Parte II.,” of the Universidad Nacional de Colombia (Bogotá) by DGC.
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PETERSON, PM., J.T. COLUMBUS, AND S.J. PENNINGTON. 2007. Classification and biogeography of New World grasses:
Chloridoideae. Aliso 23:580—594.
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Ann. Missouri Bot. Gard. 94:745—790.
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grasses (Poaceae): Il. subfamily Chloridoideae. Contr. U.S. Natl. Herb. 41:1—255.
PETERSON, PM. AND J. VaLDEs Reyna. 2005. Eragrostis (Poaceae: Chloridoideae: Eragrostideae: Eragrostidinae) from
northeastern México. Sida 21(3):1365—1420.
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(Poaceae: Chloridoideae). Sica 16:529—544.
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Kónigsberg.
BOUTELOUA (POACEAE: CHLORIDOIDEAE: CYNODONTEAE:
BOUTELOUINAE) DEL NORESTE DE MÉXICO
Yolanda Herrera Arrieta Paul M. Peterson
Instituto Politécnico Nacional Department of Botany
CIIDIR Unidad Durango-COFAA National Museum of Natural History
Durango 34220, MÉXICO mithsonian Institution
yherreraQipn.mx TA De 20013-7012, U.S.A.
ind cdi
Jesus Valdés-Reyna
Departamento de Botánica
Univerisidad Autonoma Agraria “Antonio Narro”
uenavista, Saltillo
Coahuila 25315, MÉXICO
jvaldes@uaaan.mx
RESUMEN
Se presenta el estudio t 5mico de Boutel ] te d pns pcd. Nuevo León y Tamaulipas). En el área de estudio
se reconocen en total 25 a nativas y tres vatibdades not se ilustra Bouteloua tamaulipensis G.J. Pierce ex
D. Pacheco y Columbus, sp. nov. Los nombres, aid s de akuri riha E. Fourn., Bouteloua burkii Scribn. ex S. ich eu
drummondii E. Fourn. ua Poi Torr. y les Mull Hal., son lectotipificados. S
determinar las especies, así como ee epus mapas de disitibución, especimen minados y sinonimia Ed: cada Ee de
Bouteloua en el noreste de Méxic
ABSTRACT
A i f Bouteloua f ] México (Coahuila, Nuevo León, and Tamaulipas) is given. A total of 25 native spe-
1 tl ieti ized i i rea. Bouteloua i A n Pierce ex D. Pacheco & Columbus,
sp. nov., is desorbed and illustrated. me names, Atheropogon atus E. Fourn., Bout ibn. ex S. Watson, Chondrosium
drummondii E. Fourn., asu pican Torr., and Chondrsium — Müll Hal., are Ioue A cd i ole
the species, descriptions, distril ps, Sp , and synony I d for all sp
eastern México.
Coahuila, Nuevo León y Tamaulipas o la región noreste de México cubre un área de 291,955 km? que cor-
responde al 15 % del área total de México (Fig. 1). La localización y nombre de cada Municipio en Coahuila,
Nuevo León y Tamaulipas se presenta en las Figuras 2, 3 y 4. Esta pu n pauco de dos pours
naturales conocidas como Desierto Chihuahuense y Desierto T;
como centro de origen y diversificación de especies de plantas de las zonas áridas y semi-áridas (Dávila-
Aranda y col. 2004; Dávila y col. 2006; Espejo Serna y col. 2000; Peterson y col. 2001, 2007). Como parte
de una revisión en curso, de la flora de gramíneas del noreste de México, para examinar la taxonomía y
distribución de las especies de la subtribu Boutelouinae, fue iniciada en apoyo a las industrias agrícola y
ganadera. En el presente estudio se tratan 25 especies y tres variedades no típicas.
La Subtribu Boutelouinae consta de un solo género, Bouteloua, con 57 especies (Columbus 1999; Co-
lumbus y col. 1998, 2000, 2007) con centro de origen en el suroeste de Estados Unidos y México, e incluye
la gramínea dominante de los pastizales americanos B. gracilis (Kunth) Lag. ex Griffiths (navajita azul, zacate
navajita).
La subtribu se teriza por presentar una inf] ia de I-muchas
gadas distribuidas a lo largo del eje principal, los ejes de las ramificaciones persisten o caen enteros (Allred
1982), espiguillas bisexuales o unisexuales (plantas monoicas o dioicas), o estériles, solitarias, pareadas, o
iG
J. Bot. Res. Inst. Texas 2(2): 917 — 981. 2008
- : laf tha Ratanical D h Institut f Texas 2(2)
Fic. 1. Localización de Coahuila, Nuevo León y Tamaulipas en México.
en tripletes, algunas veces la segunda gluma biseriada, aristada o sin arista, lemas 3-nervadas, aristadas o
sin arista, número básico de cromosomas x = 10, y con preferencia por los hábitats xéricos.
Como se ha entendido corrientemente para la Subfamilia Chloridoideae del Nuevo Mundo, las tribus
Cynodonteae y Zoysieae son hermanas, uniéndose con este clade los miembros de la tribu Eragrostideae
(Columbus y col. 2007; Peterson y col. 2007). Todas las tribus del árbol se apoyan en datos de secuenciación
del ADN de cloroplastos (región matK, trnL-F) y genoma nuclear (ITS) (Hilu y Alice 2000, 2001; Columbus
y col. 2007). Las relaciones entre las diez subtribus (Boutelouinae es una de ellas) dentro de Cynodonteae,
así como los 25 géneros no colocados, son inciertas dependiendo de estudios adicionales. En este punto
los agrostólogos no tienen idea clara de cómo son las relaciones entre estas diez subtribus Cynodonteae
(Columbus y col. 2007; Peterson y col. 2007). Sin embargo, hay un buen sustento molecular para mantener
la tribu Cynodonteae y uno morfológico para todas las diez subtribus.
Otros tratamientos taxonómicos importantes sobre Bouteloua que se consultaron para esta revisión
incluyen Griffiths (1912), Gould (1979), Reeder y Reeder (1980), McVaugh (1983), Beetle (1987), Herrera
Arrieta y Rzedowski (1990), Pohl (1994), Pacheco (2002), Wipff (2003) y Herrera Arrieta y col. (2004).
El siguiente tratamiento taxonómico contiene una clave para determinar las especies, descripciones,
mapas de distribución, especimenes examinados y sinonimias. Este estudio está basado en el examen de
especimenes de los herbarios: ANSM, ARIZ, B, BA, BAA, BM, CIDIIR, ENCB, GH, HINT (herbario personal
de George S. Hinton), IBUG, INEGI, K, LE, MEXU, MICH, MO, MSC, NMC, NMSU, NY, P, RSA, SI, SLPM,
TAES, TEX, UAT, US, UT, UTC, VT y W; incluyendo los especimenes tipo de la mayoría de las especies
estudiadas.
Bouteloua Lag., Varied. Ci. 2(4, 21):134. 1805. Tire: Bouteloua racemosa Lag.
Plantas anuales o perennes, cespitosas, estoloníferas o rizomatosas. Lígula membranácea, ciliada. Láminas
Harrara Arriata etal., B tal dal ta Aa Máws 919
Fic. 2. Municipios en Coahuila: Abasolo-1, Acuña-2, Allende-3, Arteaga-4, Candela-5, Castaños-6, Cuatrociénegas-7, Escobedo-8, Francisco |. Madero-9,
Frontera-10, General Cepeda-11, Guerrero-12, Hidalgo-13, Jiménez-14, Juárez-15, Lamadrid-16, Matamoros-17, Monclava-18, Morelos-19, Múzquiz-20,
Nadadores-21, Nava-22, Ocampo-23, Parras-24, Piedras Negras-25, Progreso-26, Ramos Arizpe-27, Sabinas-28, Sacramento-29, Saltillo-30, San
Buenaventura-31, San Juan de Sabinas-32, San Pedro-33, Sierra Mojada-34, Torreón-35, Viesca-36, Villa Unión-37 y Zaragoza-38.
foliares lineares, aplanadas o conduplicadas. Inflorescencia del subgénero Chondrosum: una a varias rami-
ficaciones primarias espigadas racemosas unilaterales; raquis de la ramificación persistente; espiguillas
numerosas, sésiles, pectinadas. Inflorescencia del subgénero Bouteloua: un racimo de 1 a numerosas rami-
ficaciones primarias espigadas cortas, unilaterales; ramificaciones desarticulándose como una unidad y las
espiguillas a veces también desarticulándose arriba de las glumas. Ramificaciones primarias espigadas por
lo general con 3 a numerosas espiguillas, insertas en 2 hileras a lo largo del lado inferior del raquis. Espi-
guillas comprimidas lateralmente, con un flósculo perfecto y 1 a 2 flósculos rudimentarios, estaminados o
920 Journal of the Botanical R h Institute of Texas 2(2)
Fic. 3. Municipios en Nuevo León: Abasolo-1, Agualeguas-2, Aldamas Los-3, Allende-4, Anáhuac-5, Apodaca-6, Aramberri-7, Bustamante-8, Cadereyta
Jiménez-9, El Carmen-10, Cerralvo-11, Ciénega de Flores-12, China-13, Doctor Arroyo-14, Doctor Coss-15, D González-16, Galeana-17, García-18,
San Pedro Garza García-19, General Bravo-20, Iturbide-30, General Escobedo-21, General Terán-22, General Treviño-23, General Zaragoza-24, General
Zuazua-25, Guadalupe-26, Los Herreras-27, Higueras-28, Hualahuises-29, Juárez-31, Lampazos de Naranjo-32, Linares-33, Marín-34, Melchor Ocam-
0-35, Mier y Noriega-36, Mina-37, Montemorelos-38, Monterrey-39, Parás-40, Pesquería-41, Los Ramones-42, Rayones-43, Sabinas Hidalgo-44,
Salinas Victoria-45, San Nicolás de los Garza-46, Hidalgo-47, Santa Catarina-48, Santiago-49, Vallecillo-50 y Villaldama-51.
Mí
estériles, modificados u ornamentados de forma variable. Glumas desiguales a subiguales, más cortas que
la lema, 1-nervada, la primera más corta que la segunda. Lema del flósculo fértil cartilaginoso, 3-nervada,
las nervaduras a menudo excurrentes como aristas. Pálea 2-carinada, el ápice 2-dentado o 2-mucronado.
Lema del flósculo rudimentario generalmente reducida, con aristas alargadas. Lodículas 2, estambres 3,
estilos 2. Fruto una cariopsis; embrión Y a 2/3 partes la largo de la cariopsis; hilo punteado. En el área de
estudio están presentes 25 especies.
Ha Arriata af 3l Bantal AAT Anda Mul
rrara al., 921
Fic. 4. Municipios en Tamaulipas: Abasolo-1, Aldama-2, Altamira-3, Antiguo Morelos-4, Burgos-5, Bustamante-6, Camargo-7, Casas-8, Ciudad Madero-9,
Cruillas-10, Gómez Farías-11, González-12, Güemez-13, Guerrero-14, Gustavo Díaz Ordaz-15, Hidalgo-16, Jaumave-17, Jiménez-18, Llera-19, Mainero-20,
El Mante-21, Matamoros-22, Méndez-23, Mier-24, Miguel Alemán-25, Miquihuana-26, Nuevo Laredo-27, Nuevo Morelos-28, Ocampo-29, Padilla-30,
Palmillas-31, Reynosa-32, Río Bravo-33, San Carlos-34, San Fernando-35, San Nicolás-36, Soto la Marina-37, Tampico-38, Tula-39, Valle Hermoso-40,
Victoria-41, Villagrán-42 y Xicoténcati-43.
922 || I af tl Das H In
pci AE 5 ill lesarticul | iguillas f
Ha]
espiguillas masculinas 2-floras (Buchloe).
6. B. dactyloides
. Plantas con flores perfectas, al menos las principales.
2. Espiguillas 3 en cada espiga; la o central corto estipitada, 3-flosculares, primer flósculo herma-
cum).
b laterales masculinos (Cathes
3. nudos de los
e largo; anteras 1.5 mm o más largas
estolones pce de 15 cm o más largos; ramas de la inflorescencia mayores
7. B. erecta
3. da de los estolones menores de 15 cm; ramas de la inflorescencia de 6 mm o más cortas
anteras usualmente Mes de l ae mm
B. tamaulipensis
2. Espiguillas (132 a ga, de dif ¿con 1 a 2(-3) rudimentos,
tados por una aa
. Espigas izas en la madurez, portando 1 a 12 espiguillas; las espiguillas permanecen
espigas.
5. Ramas de la inflorescencia (espigas), la mayoría, con una sola espiguilla.
6. Espigas pediceladas, con un pedicelo conspicuo de 2 a 3 m
veces represen-
insertas a las
m de largo; plantas con estolones
aéreos bien desarrollados; arista central del flósculo rudimentario (3.524 a 4.5 mm de
largo
15. B. pedicellata
24. B. uniflora
: : | , menor de 0.5 mm de largo; plantas sin estolones
aéreos ado arista central del fló dm Idimentario 2 a 3.5 mm de largo
5. Ramas de la pigas) con (1-2 a 12 i onus
7. Espigas (4-)9 a 80 por Do ius o péndulas, si son menos que 15, entonces las
espigas miden menos de 1 cm
8. Espigas 9 a 25(-35), de 5 a 10 mm de Sid portando (1-2 a 6 espiguillas; arista central E.
flósculo rudimentario 2.5 a 5 mm de lar
5. B. warnockii
8. Espigas (4-)30 a 80, de 1 a 3 cm de largo, comers 2 a 7 espiguillas; arista central
rudimentario 5 a 10 mm
del OR
7. Espigas 1 a 16 por inflorescencia, ro lent tentes, de 0.7 a4cmd
5. B. MODE
a sel ad eanes mee) mes de 1 cm.
9.
am
E ga
uv
,aristada
con 3 aristas iguales a subiguales; espiguillas aplicadas al raquis de la espiga, ampliamente
separadas una de otra
1. B. aristidoides
9. Plantas perennes; flósculo rudimentario oa desarrollado, eee o lanis
cui
19. B. rigidiseta
aplicadas o extendidas desde el raquis de la espiga u
10. Espigas portando (2 FA a5 Espada sn nin ise
10. Us portando 5a al te glab
11. Lá r , larg Duc dc d
sate -pubescentes en la eee adaxial
aC
2. B. johnstonii
. Láminas foliares ele ades. 2 us en la mitad basal, la mnan aSa llega a ser invo-
luta cuando seca, el ápice pina, glabras,
base papilosa en una o ambas S
12. Epes 3a 8, ae 7 a 10 mm de largo; raquis de las espigas y segunda gluma
4. B. chondrosioides
12. Fepigas 5 a 12, de 15 a 50(-70) mm de largo; raquis de las espigas y segunda
gluma og a escabrosas.
13. + rim hian necarralladas , fue
de $ a 21 mm n de largo; espigiillas 9a 13 mm nde largo
, con entrenudos
16. B. radicosa
espiguillas 6 a 9 mm ie id
2mm
4. Fe DE portando (8-)20 a 60(-1 illas; | iguill desarticul
18. B. repens
Id L T
a nforecenci una sola espiga unilateral.
lantas claramente anuales
"s e perennes.
21. B. simplex
aquis de la espiga prolongada 2 a 10 mm más allá del punto de inserción de la espi-
guilla distal, terminando en una aguja semirígida; segunda gluma con pelos de base
papilosa, al menos en algunas espiguillas
11. B. hirsuta
ieta et al., B tal ESI ta da Mv: 923
16. D cri ; : E l z H4 Ant PEE ns Fore SN | ; Em . |
gl glabra o pub te pero sin pelos de base papilosa.
| s | T J
17. Una sola espiga juis d piga y seg gl glal b ;
aqu claramente srqueado 20. B. scorpioides
e dal l | Hi hf. id
2 T T +
a ass piloso- papilosos; aquis recto, no a 10. B. gracilis
14. Inflorescenci pigas unilat
18. a o (al menos de ides Seales hispida o hirsuta con pelos de base
papilos
19, au de la espiga extendiéndose más adelante del punto de inserción de la Mia.
terminal 1. B. hirsuta
19, ale de la espiga sin extenderse más adelante del punto de inserción de la eem
terminal
20. Lema 3.5 a 4 mm de largo; us 3a8,de 1 a3 cm de largo; plantas cespitosas o
estoloníferas, pero nunca rizoma 4. B. parryi
20. Lema 4 a 5.5 mm de largo; Bs i -)2(-4), de (1.593 a 4.5 cm de largo; plantas
dEliérarmente edi cud 10. B. gracilis
18. Segunda gluma g'abra | | d e papilosa.
21. Entrenud del mo: " menos s los ias lanoso- ema
22. Primera gluma 3 a 4 mm de largo, segunda gluma glabra; lema de la flor perfecta de
6 a 7 mm de largo, claramente glabra 8. B. eriopoda
22. Primera giuma 2 a 3 mm de raro segunda ana Mss a scs eum ite; lema
de la flor perfecta 5 a 6 mm | |
barbada en la base 9. B. eriostachya
1 Entr, A rlal | | lah
" SENER f pol
iclo de vida corto 2. B. barbata
N
v2
23. Plantas claramente perennes.
24. e con (1-)2(-4) espigas.
ulmos conspicuamente ramificados en la mitad inferior, barbados en las
axilas de las ramas, con 4 a 5 nudos; segunda gluma 3 a 3.2 mm de largo, con
un mucrón de 0.8 a 1 mm de largo; base de la planta firme y relativamente
leñosa 17. B. ramosa
25. Culmos nunca ramificados, con 1 a 2(-3) nudos, glabros; segunda gluma 5 a
6.5 mm de largo, apiculada, no mucronada; base de la planta firme pero
nunca bios 10. B. gracilis
24. Inflorescen n 3 a 30 espigas
26. Plantas de base fuenemente dlls a dii d
27. e de
uA. de 10 : 30(-40) cm " de alto; espigas con 8 a 24(-32) espiguil-
glabra 23. B. trifida
28. Culos de 20 a 270 cm de alto; espigas con 40 a 100 espiguillas; se-
unda gluma esparcido pilosa a híspida . B. gracilis
i Espiguillas de 3 a 4.5 mm de largo; plantas de base ene
rizomatos
29. Segunda gluma 3 a 3.5 mm de largo; lema de la flor perfecta
5 mm de largo, e Ree nes ra-
evil p | flós culo
nao 3. B. chasei
29, n. gluma 2 a 2.5 mm de largo; lema de la flor perfecta
mm de largo, esparcidamente pubescente; raquilla
sin un cuu de pelos en la base del flósculo rudimentario
13. B. karwinskii
26. Plantas de base nunca rizomatosa.
30. Lema de la flor perfecta glabra a esparcido estrigosa, con 3 aristas gen-
eralmente de 4 a 5(-6) mm de largo 23. B. trifida
30. Lema de la flor perfecta pubescente en la base, con 3 aristas de 1 a 3 mm
largo
10. B. gracilis
924 J | of the Botanical R h Institute of Texas 2(2)
1. S aristidoides (Kunth) Griseb. var. aristidoides, Fl. Brit. WI. 537. 1864. (Fig. 5). Dinebra aris-
tidoides Kunth, Nov. Gen. Sp. (quarto ed.) 1:171. 1816. Atheropogon aristidioides (Kunth) Roem. & Schult., Syst. in 2:415. 1817.
Futriana as (Kunth) Trin., Gram. Unifl. Sesquifl. 242. 1824. Triathera aristidoides (Kunth) Nash, Fl. S.E. U.S. 137. 1903.
Tipo: MÉXICO: Crescit in asperis frigidis convallis Tolucensis (cercano Nevado Toluca), Sep, E Humboldt & A. ane 67 (HOLOTIPO:
P; isctiro: US fragm.!
Bouteloua ciliata Griseb., Abh. Kónigl. Ges. Wiss. Sonica 24:302. 1879. Tipo: ARGENTINA. Sata: cerca del Pasaje del Río Juramento,
21 Feb 1873, PG. Lorentz & G. Hieronymus 35 -A865569 fragm. ex GOET!).
Dinebra hirsuta J. Presl., Reliq. Haenk. 1(4-5):292. 1830 Eutriana hirsuta (J. a Kunth, Enum. Pl. 1:280. 1833. Tiro: PERÚ: Haenke
n. (SoLoTiPo: PR; isoriPo: US fragm.!).
Plantas anuales de vida corta, amacolladas. Culmos de 6 a 50 cm de alto, delgados, frágiles, simples, los
laterales curveado-erectos desde una base decumbente. Follaje muy escaso; vainas más cortas que los entre-
nudos; lígula de 0.2 a 0.5 mm de largo, un anillo de pelos cortos; láminas foliares de 2 a 4 cm de largo y 1 a
2 mm de ancho, delgadas, aplanadas o conduplicadas, superficie adaxial a menudo con escasa pubescencia
de pelos largos y duros en la base, que en ocasiones se extiende hacia el ápice. Inflorescencia de 2.5 a 10
cm de largo; ramificaciones primarias espigadas 6 a 16, de la 2(3.5) cm de largo, erectas o ascendentes,
caedizas, espaciadas, abiertas, portando 2 a 5-10) espiguillas; raquis aplanado, densamente pubescente
por lo menos en la base, ápice curvado, extendiéndose (2-)6 a 10 mm más adelante de la inserción de la
espiguilla terminal. Espiguillas de 6 a 8 mm de largo, permanecen insertas a las espigas, las inferiores de las
ramas aplicadas al raquis, sin flósculo rudimentario, con la lema sin arista a corto aristada; las superiores
de las ramas con un flósculo rudimentario reducido a una columna 3-aristada, de 2 a 6 mm de largo; glu-
mas muy desiguales, angostas, agudas o acuminadas, primera gluma de 2 a 2.5 mm de largo, aristiforme,
escabroso-puescente, segunda gluma de 6 a 7 mm de largo, ancha, esparcido pubescente principalmente
sobre los márgenes, a menudo abierta formando un amplio ángulo con la flor; lema de la flor perfecta 6 a
7 mm de largo, con aristas cortas de 0.4 a 0.6 mm de largo, escaso a abundantemente pubescente sobre la
nervadura media; pálea de 6 a 7 mm de largo, glabra. Flósculo rudimentario 1, lema de 8 a 10 mm de largo,
fuertemente 3-aristada; aristas de 6 a 8 mm de largo, aplanadas, escabrosas. Anteras de 3 a 4 mm de largo.
Cariopsis de 2 a 2.5 mm de largo, color amarillo cambiando a café en la madurez, lanceolado-elíptica. 2n =
40. Florece probablemente durante la pas
omentarios.—Se le reconocen dos variedades B. aristidoides var. aristidoides y B. aristidoides var. arizonica,
esta última muy poco común.
Distribución.—Sw de Estados Unidos a México; Sudamérica en: Argentina, Bolivia, Brasil, Paraguay y
Perú.
Hábitat.—Pastizales deteriorados, matorrales xerófilos, a orilla de cultivos y en lugares pedregosos y
secos; 245 a 1900 m.
Ejemplares examinados. MÉXICO. Coahuila: Coahuila, E. Palmer 1353 (US); Municipio de Escobedo, 33.5 km al SE del ejido Paso
de San Antonio, Rancho el Carrizal, 2 Oct 1972, 1150 m, F. Chiang s.n., T. Wendt & M.C. Johnston (MEXU); Municipio de Matamoros,
10 km carretera Matamoros-Saltillo, 5 Sep 1988, J. Espinosa 88 (ANSM, TAES); Municipio de Muzquiz, Sierra del Carmen, Ejido San
Francisco, 5 Sep 2005, 1424 m, P.M. Peterson 18817, J. Valdés-Reyna & C. Sifuentes (US); Municipio de Ocampo, Along the Río Grande,
1 km W of Boquillas de Carmen, 6 Sep 2005, 560 m, P.M. Peterson 18870, J. nia Reyna & C. Sifuentes (US); Municipio de Parras, El
Mesteño, Ejido 4 de Marzo, 29 Jul 1994, E.A. Roing 94 (ANSM); A de S , El eee carretera A E. Lyonnet
3762 (MEXU); 10 km al NW de San Fernando y 2 km al N de Loma de Piojo, H. García s.n. (SLP Pedro, 5.5 km al
de Nueva Delicias, vertiente E de la Sierra de las Delicias, al N de San Pedro de las uds Y Herrera LEUR M. de la Cerda & O.
Rosales 1489 (CIIDIR, HUAA); Municipio de Torreón, 3 km de Torreón, Fracc. La Florida, 1120 m, B. Rodriguez-C 4,9 (ANSM, COCA,
IEB); Sierra de Jimulco, proximidades al ejido La Trinidad, 25 Aug 1988, 1900 m, J.A. Villarreal 4414, M.A. Carranza & A. Rodríguez
(ANSM, MEXU); 13 Oct 1989, E. Palmer 513 (MEXU); Municipio de Viesca, 5 km antes de Viesca, 25 Aug 1988, 1500 m, J.A. Villarreal
4456, M.A. Carranza 6» A. Rodríguez (ANSM, CIIDIR). Tamaulipas: Municipio de Casas, Camino al Ejido 1* de Abril, 28 Jul 1983, 245
m, M.H. Cervera 153 (MEXU
2. icem barbata Lag. var. barbata, Varied. Ci. 2(4):141. 1805. (Fig. 6). Actinochloa barbata (Lag.) Roem. &
., Syst. Veg. 2:420. 1817. E (Lag.) Kunth, Revis. Gramin. 1:96. 1829. Chondrosum barbatum (Lag.) Clayton, Kew
apa en 417. 1982. Tiro: MÉXICO: Nee s.n. UNAM MA; isoripo: US-865567 fragm. ex MAI).
Harra
925
1 Ial em p Y EN
Fic.5. Distrib
Chondrostum polystachyum Benth., Bot. Voy. Sulphur 56. 1844. Bouteloua polystachya (Benth.) Torr., Pacif. Railr. Rep. 5(2):366, t. 10. 1857.
: ur: Bahía Magdalena, G.W Barclay s.n. (HoLomipo: K; isorieos: BM, US-A865566 fragm. ex K!).
Tiro: MEXICO. Baja CALIFORNIA S
drosi i ioides Müll Hal., Bot. Zeitung (Berlin) 14:347. 1856. Tiro: MÉXICO. BAJA CALIFORNIA: Patria, G.W Barclay s.n. (Lec-
)
TOTYPE, designado aquí: US-883457 ex K!, un espécimen con etiquetado "lower California.”).
teloua pumila Buckley, Proc. Acad. Nat. Sci. Philadelphia 14:93. 1862. Tio: U.S.A. New Mexico & Texas: C. Wright 754 (Lectotipo: PH,
designado por Griffiths, Contr. U.S. Natl. Herb. 14(3) 383. 1912; isorEcrOTIPOS: GH, US-81706)).
Chondrosium exile E. Fourn., Mexic. Pl. 2:137. 1886. Tiro: MÉXICO: J.L Berlandier 842 (HoLoripo: P; 1sotipos: US-A865678 fragm. ex
P!
1, W).
Chondrosium microstachyum E. Fourn., Mexic. Pl. 2:138. 1886. Bouteloua microstachya (E. Foum.) L.H. Dewey, Contr. U.S. Natl. Herb.
2(3):531. 1894. Tipo: MÉXICO: Guadelupe, 1865-1866, M. Bourgeau 667 (HoLomiro: P; isorros: LE, MO, US-74908!, US-87194 fragm.!),
Bouteloua arenosa Vasey, U.S.D.A. Div. Bot. Bull. 12(1): t. 34. 1890. Bouteloua barbata var. arenosa (Vasey) Beetle, Phytologia 54:1. 1983.
Tiro: MÉXICO. Sonora: Guaymas on the Gulf of California, 1887, E. Palmer 189 (oronro: US-817161).
Bouteloua micrantha Scribn. & Merr., Circ. Div. Agrostol. U.S.D.A. 32:8. 1901. 32:8. 1901. Tiro: U.S.A. Arizona: Pima Co.: Fort Lowell,
Sep 1900, D. Griffiths 1556 (noLoripo: US-81724)).
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Fic 6. Dictril y la D L b har
Plantas anuales, de ciclo corto, de tamaño y aspecto variable, según hábitat y grado de desarrollo. Culmos
de 5 a 30(50) cm de alto, separados y decumbentes o amacollados y erguidos en vegetación densa, nunca
desarrollando estolones; entrenudos sólidos, glabros. Hojas persistentes, caulinares y basales, de textura
suave; vaina con el margen membranoso-hialino de 1 a 5 cm de largo; lígula de 0.6 a 1 mm de largo, un
anillo de pelos cortos y largos, hialinos; láminas foliares de 1 a 9 cm de largo y 1 a 3 mm de ancho, aplana-
das, rectas; escabrosas en la superficie adaxial, con pubescencia de base papilosa a los lados de la nervadura
principal y en ocasiones en las ner d laterales; la superficie abaxial escabrosa, a veces con pelos hirsutos
de base papilosa; ápice acuminado. Inflorescencia con (227(-11) ramificaciones primarias espigadas por
culmo; ramificaciones 1 a 2.5 cm de largo y 1.5 a 5 mm de ancho, persistentes, unilaterales, purpúreas,
corto pediceladas, curvadas, portando 20 a 3045) espiguillas; raquis aplanado, escabroso. Espiguillas de
1 a 2.5 mm de largo, articulación por arriba de las glumas; glumas desiguales ovadolanceoladas, aquilladas,
marginadas irregularmente, prolongándose en un mucrón corto, primera gluma de 0.7 a 1.5 mm de largo,
Harrara Arriat 4 I, B tal dal ta da Mw 927
color verde pálido a hialina, segunda gluma de 1.5 a 2.5 mm de largo, glabra, púrpura marginada; lema
de 1.7 a 4 mm de largo, ligeramente pilosa en la base, con 3 aristas de 1.5 a 2 mm de largo, escabrosas, y
4 dientes intercalados con las aristas, color verde pálido; pálea de 1.5 a 4 mm de largo, color verde pálido,
con 2 aristas escabrosas, porciones laterales pilosas en toda su superficie. Flósculo rudimentario 1, de 1.5
a 4 mm de largo incluyendo las aristas, con 3 aristas purpüreas, escabrosas, divergentes, de 0.8 a 4 mm de
largo, y 3 lóbulos de 0.5 a 1 mm de largo, color verde claro con los márgenes púrpura; callo de 0.7 a 1 mm
de largo, con un anillo piloso. Anteras de 1 a 1.2 mm de largo, color amarillo-rojizo. Cariopsis 0.5 a 1.3 mm
de largo, angulosa-triangular. 2n = 20, 40. Florece de agosto a diciembre.
Distribución. —Amplia en Estados Unidos y México, encontrada también en Argentina.
Hábitat. —Orilla de caminos, bordos, arroyos; áreas de cultivo, suelos arenosos, gravosos y arcillosos;
matorrales desértico micrófilo, espinoso, subespinoso, y sarco-crasicaule; pie de monte; 650 a 1900 m.
Ejemplares examinados. MÉXICO. Coahuila: Municipio de Arteaga, Arroyo Canoas, 28 Jul 1993, 1600 m, F. Alcalá 22-A (MEXU);
Arteaga, 18 Sep 1991, 1600 m, Brigada Coahuila 229 (MEXU); Municipio de Cuatrociénegas, 2 millas NO de Cuatrociénegas, carretera
a Ocampo, 22 Sep 1990, 830 m, P. M. Peterson 9997, C.R. Annable y J. Valdés-Reyna (ANSM, US); 28 mi SO de Cuatrociénegas, 23 Sep
1990, 750 m, P.M. Peterson 10003, C.R. Annable & J. Valdés-Reyna (ANSM, US); km 11 E de Cuatrociénegas, 24 Sep 1955, 650 m, E.
997, L. García (ANSM, pound Sierra de la Madera, Cañón Charreteras, rancho Charreteras, 15 Sep 1993, 1750 m, J.A. Villarreal 7359,
M.A. Carranza & R. Rodrig ANSM); Sierra de San ae areas cercanas a la Poza de La Becerra, 25 Sep 1988, 950 m, A. Zarate 2
(ANSM); Municipio de G l Cepeda, 38 | y 40, 21 Jun 1964, 1400 m, G. Mick 84, K. Roe (TAES); Paila junction
on Mex 40, Parras, J.M. Flooh & J.D. Spears 75 (CHAPA); 41 mi iO of Saltillo, 5 Sep 1965, 1400 m, F.W. Gould 11536 (TAES, US); km 45
carr. 40 Saltillo-Torreón, en la desviación a Gral. Cepeda, Y. Herrera Arrieta & A. aa 1477 nupt dii 10 km 2 W de Saltillo,
carr. Saltillo-Gral. Cepeda, J. Valdés-Reyna, L.E. Rodriguez & R. Vazquez 15712 lava, 3 mi W de Monclava,
11 May 1979, desert grassland, 600 m, F.W. Gould 15565 & R.B. Shaw (CHAPA); Municipis de Maia Sierra del Carmen, Ejido San
Francisco, 5 Sep 2005, 1430 m, P.M. Peterson 18831, J. Valdés-Reyna & C. Sifuentes (US); 7.2 mi SE of eee on ee lowers Cuesta
de malena and Melchor Muzquiz, 01 Sep 2006, 1010 m, P.M. Peterson 19850, S. Lara-Contreras (US); Muni ipi a El Jardín,
slopes above Canyon del Diablo, 2 Sep 2006, 1418 m, P.M. Peterson 19882, S. Lara-Contreras (US); Municipio de Fide Rancho el
Barranquito, aprox. 50 km de Ocampo rumbo a Sierra Mojada, 2 Oct 1990, 1200 m, M.A. Carranza 574, FJ. Carranza (ANSM); Rancho
Las Gallinas, aprox. 43 km de Ocampo rumbo a Sierra Mojada, 2 Oct 1990, 1300 m, M.A. Carranza 565, F.]. Carranza (ANSM); Sierra de
la Madera, Rancho Laguna de la Leche, aprox. 62 km de oe rumbo a Sierra Mojada, 2 Oct 1990, 1000 m, M.A. Carranza 624, FJ.
Carranza (ANSM, TAES); Municipio de P. ,2kmalOd , 10 Sep 1960, F. Medellín 987 (CHAPA, MEXU); El Mesteño, Ejido 4
de Marzo, 28 Jul 1976, 1290 m, FA Roing 99 (ANSM); km 82 carretera Torreón-Saltillo, 23 Sep 1955, 1000 m, E. Hernández-X. 1997, V.
Mathus (ANSM, CHAPA, US); Sierra de Parras en el cañón de Gustrola, 10 Oct 1980, 1850 m, A. Rodríguez 253, M.A. Carranza (ANSM);
Parras, W.A. Archer 3405 (US); Municipio de Ramos Arizpe, Cañada el Diente, Sierra de la Paila, 19 Sep 1989, 1300 m, J.A. Villarreal
5193, M.A. Carranza (ANSM); La Sauceda, 17 Dic 1965, 1030 m, G. Cano 65-6 (TAES); Rancho Santo Domingo, 5 Nov 1982, 1680 m,
F. Coss-V (ANSM); Municipio de Saltillo, 26 km O de Saltillo, carretera Saltillo-General Cepeda, ejido La Noria, 21 Jul 1983, 1780 m,
J. Valdés-Reyna 1571, L.E. Rodríguez (ANSM); About 3 mi E of San José, 18 Aug 1941, I.M. Johnston 8216 (MEXU); 1898, E. Palmer 400,
514 (MEXU); Municipio San Pedro, 5.5 km al NW de Nueva Delicias, vertiente E de la Sierra de las Delicias, al N de San Pedro de las
Colonias, Y. Herrera Arrieta, M. de la Cerda & O. Rosales 1488 (CIIDIR, HUA A); San Pedro, Sierra de San Salvador, ladera, carretera San
Pedro-Cuatrociénegas, 12 Enel974, 900 m, J.S. Marroquín s.n. (ANSM); Municipio de Torreón, Plains near Jimulco, 24 Sep 1902, 1524
m, C.G. Pringle 11216 (MEXU, TAES); Sierra de Jimulco, proximidades al ejido La Trinidad, 25 Au ug 1988, 1900 m, J.A. Villarreal 4413,
M.A. Carranza & A. Rodríguez (ANSM); Colonia Estrella, Torreón, B. Rodríguez s.n. (IEB, SLPM); L.M. Villarreal 4105 (IBUG); 5 km al
NE de las Lilas, M.A. Carranza & L. García 7848 (IBUG); S of Torreon canyon between Jimulco and Jin o P.M. Peterson 8470, n
Valdés-Reyna (US). Nuevo León: Municipio de Allende, 153.2 km NW of Muzquiz on Hwy 53 quilla del Carmen and 8
NW of jct of road to Encantada, 12 Sep 1991, 1250 m, P.M. Peterson 10605, C.R. Annable (US); Municipio de Gal , El Salero, 19 E
2001, 1840 m, P.M. Peterson 15818, J. Valdés-Reyna (US); 19 km SE of El Salero, 19 Sep 2003, 1707 m, PM. Peterson 17837, J. Valdés-Reyna
& H.S. Hinton (US). Tamaulipas: Municipio de Bustamante, Rumbo a la Cabecera de Bustamante, 4 Sep 1981, 1600 m, J.G. Galván 31
(MEXU); Municipio de Miguel Alemán, Brecha de Pemex, 25 Oct 1991, 95 m, C.R. I ópez 202 (MEXU); Municipio de Tula, 40 km al S
de Tula, 25 Aug 1983, I. Nufiez 16 (MEXU)
3. Bouteloua chasei Swallen, po po Soc. Wash. 56:81. 1943. (Fig. 7). Chondrosum chasei (Swallen) Clayton, Kew
Bull. 37:418. 1982. Tipo: MÉXICO. N Leon: Galeana, Mesa, 5400 ft, 30 Jul 1939, A. Chase 7673 14 (nuovormo: US-1763078!).
Plantas perennes, amacolladas, rizomatosas. Culmos 15 a 30 (50) cm de alto, desde una base firme, nudosa;
entrenudos glabros. Hojas basales; vai ás cortas que los entrenudos, con pubescencia diminuta cerca de
los márgenes, barbados a los lados de la lígula; lígula de 3 mm de largo, un anillo de pelos; láminas foliares
L af sh D 1 ID hil bitit, £T,
928 J t Texas 2(2)
Fie 7 Distribución de B 1 n , l ta da M
de 3 a 10 cm de largo y 2 a 3 mm de ancho, cerrado-involutas, angostas y fuertes, arqueadas, pungentes,
esencialmente glabras. Inflorescencia de 2 a 4(-8) cm de largo, con 2 a 3(-5) ramificaciones primarias
espigadas; ramificaciones de 1.5 a 2(-2.5) cm de largo, erectas o algo curvadas, persistentes, portando 18 a
25(-30) espiguillas; raquis glabro, en ocasiones con pelos cortos sobre el margen. Espiguillas de 3.5 a 4.5
mm de largo, articulación por arriba de las glumas; glumas desiguales, la primera de 1.5 a 2 mm de largo,
la segunda de 3 a 3.5 mm de largo, mucronadas, pubescentes hacia el ápice y el nervio central; lema de la
flor perfecta de 4 a 4.5 mm de largo, densamente plateado-pubescente, pelos aplicados, de 0.6 a 0.8 mm de
largo, diminuto aristada desde un ápice bífido; arista de 2 a 3 mm de largo; pálea 3 mm de largo, pubescente
hacia un ápice 2-dentado. Flósculos rudimentarios 2, con un mechón denso de pelos en la base de ambos
flósculos rudimentarios, el primero de 3 a 4 mm de largo incluyendo la arista, el segundo de 1 a 1.5 mm
de largo, truncado, sin aristas. Anteras de 1 a 1.5 mm de largo. Cariopsis de 1.8 a 2.2 mm de largo, color
amarillo, lanceolado-elíptica. 2n = 40. Florece de julio a diciembre.
Harrara Arriata at I, B tal dal ta da Más 929
Distribución. —Especie mexicana de: Coahuila, Durango, Nuevo Leon, San Luis Potosí y Zacatecas.
Habitat.—Es un componente estricto de suelos yesosos y secos, asociado a plantas arbustivas; 1219 a
2150 m.
A E examinados. MÉXICO. Coahuila: Municipio de Ramos Arizpe, Rancho Santo Domingo, 10 Oct 1982, 1680 m, F. Coss-V.
s.n. (ANSM); Municipio de Saltillo, 28.2 mi S de Saltillo, carretera 54 a Concepción del Oro, Zac., 24 Sep 1990, 1860 m, P.M. Peterson
10030, C.R. Mik 6J. Valdés-Reyna (ANSM, US); 33 mi S de Saltillo, carretera 54, 17.7 mi E de La Ventura, 24 Sep 1990, 1720 m, P.M.
Peterson 10041, C.R. Annable & J. Valdés-Reyna (ANSM, US); 53 km S of Saltill , along hwy 54, S of Pop del Oro, ar 26 Aug
981, 1850 m, S.L. Hatch 4504, C.W. Morden & J. Valdés-Reyna (ANSM); Along road to C f Saltillo, 5 Sep
1963, 1890 m, J.R. Reeder 3621, C.G. Reeder. (ENCB, MEXU); Buenavista, 7 km S de Saltillo, carretera 54 Saltillo- Concepción del Oro,
Zacatecas, 9 Aug 1974, 1775 m, J.F. Cano 111 (ANSM); 7 kmal S de la Cd. de Saltillo, F. del Río s.n. (ENCB); 7 km S P5 Saltillo, carretera
54 Saltillo- Concepción del Oro, Zac., Nov 1973, 1776 m, M.A. González, s.n.(ANSM); 7 km S de Saltillo, t Saltillo-Concepción
del Oro, Zac., 28 Jul 1974, 1775 m, R.E. Rodríguez s.n. (ANSM); 40 km al SW de Saltillo rumbo a Zacatecas, en el km 295 de la carr. 54,
Y. Herrera Arrieta & A. Cortés 1457 (CIIDIR, HUAA); La Ventura, aprox. 80 km 5 de Saltillo, 10 Oct 1989, 1800 m, J. Valdés-Reyna 1989,
J.A. Villarreal & M.A. Carranza (ANSM). Nuevo León: Municipio de Aramberri, 7.5 km E of Puentes on dirt road, 19 Sep 2003, 1775
m, P.M. Peterson 17848, J. Valdés-Reyna & H.S. Hinton (US); Municipio de Doctor Arroyo, 50 km al S de San José de las Raices, sobre la
carretera a Matehuala, 14 Oct 1974, 1800 m, J. Rzedowski, 32274 (MEXU); Dr. Arroyo, Nov 1980, 1750 m, J.A. Ochoa 1273 (MEXU);
Municipio de Galeana, 2 km S del Salero, 1 km E de carretera 57 Matehuala- a 31 Jul 1983, 2050 m, J. Valdés-Reyna 1615, L.E.
Rodríguez (ANSM, ENCB); J. Valdés-Reyna & J.S. Marroquín s.n. (IBUG); Near San Rafael (Hwy 57), 19 Sep 2001, 1900 m, P.M. Peterson
15813, J. Valdés-Reyna (US); 5.6 mi E of jct of Hwys 57 and 58 towards Linares, 17 Oct 1995, 1950 m, P.M. Peterson 13294, M.B. Knowles
(US); La Chona, 19 Sep 2003, 1850 m, P.M. Peterson 17824, J. Valdés-Reyna & H.S. ini (US); 19 km SE of El Salero, 19 Sep 2003, 1707
m, P.M. Peterson 17840, J. Valdés-Reyna & H.S. Hinton (US); 20 km S of San Rob g hwy 57, 18 Aug 1983, 1820 m, S.L. Hatch 4941,
J. Valdés-Reyna & J. Kessler (ANSM); 5 km SE of intersection at Galeana along hwy 58, 54 is SE of San Roberto, 27 Aug 1981, 1350 m,
5.L. Hatch 4558, C.W. Morden & J. Valdés-Reyna (ANSM); About 1 Mi S of Galeana, 4 Sep 1991, 1524 m, K.W. Allred 5504, J.T. Columbus
& J. Valdés-Reyna (ANSM); About 3 Mi S of Galeana, 20 Jul 1958, D.S. Correll 19851, LM. Johnston (MEXU); Alrededor de Galeana, Sep
1980, J.A. Ochoa 1259 (MEXU); Ejido El Tokio, 22 Jul 1986, 1865 m, M.L. Avalos s.n. (ANSM, UAT); 19 Nov 1981, 1900 m, O.L. Briones
470, J. Treviño (ANSM); 29 May 1993, 1890 m, J.L. Mora 343 (UAT); 4 mi al SE de Galeana, J.F. Reeder & C.G. Reeder 3653 (ENCB); 3966
(US); Región de Galeana, E. Hernández & V. ania N-2042 See : mi E N de San sun JR. nd & C.G. ded dE a
Galeana, Nov 1980, 1650 m, J.A. Ochoa 998 (MEXI
26 Aug 1987, 1219 m, D. Bogler 127, T. Atkins (MEXU); La Becerra, 8 Sep 1989, 1950 m, G.B. Hinton 19644 (MEXU); Salinas del Refugio,
A. Rodriguez (ANSM); Santa Rita, 5 km S de Galeana, 11 Dic 1991, 2300 m, J.A. Villarreal 6318, M.A. Carranza (ANSM); 5 km al E de
Galeana, I Cabral 1083-b (IEB); Iturbide, Bosque-escuela, Universidad Autónoma de Nuevo León, 12 km E de Iturbide, 18 Nov 1987,
1500 m, I. Cabral 1083 (ANSM).
4. Bouteloua chondrosioides a Benth. ex S. Watson, Proc. Amer. oe a 179. 1883. (Fig. 8).
Dinebra chondrosioides Kunth, N n. Sp. (quarto ed.) 1:173, t. 53. 1816. At & Schult., Syst.
Veg. 2:416. 1817. Tiro: MECO. Michoacán: Sep, E Humboldt & A. E al otro: P; sonos: LE, US fragm.!).
Bouteloua havardii Vasey ex S. Watson, Proc. Amer. Acad. Arts 18:179. 1883. Tiro: U.S.A. Texas: Limpio Mts., V Havard 53 (notorio:
-817231)
Plantas perennes, cespitosas. Culmos de 20 a 50 cm de alto, numerosos, erectos, simples; entrenudos 2 a
5, glabros; nudos glabros. Vainas glabras; lígula 0.2 a 0.4 mm de largo, un anillo ciliado; láminas foliares
de 3 a 10(-18) cm de largo y 2 a 3 mm de ancho, aplanadas, glabras o piloso-papilosas. Inflorescencia de
3 a 7 cm de largo, con 3 a 8 ramificaciones primarias espigadas erectas o ascendentes, caedizas en la ma-
durez; ramificaciones de 7 a 10 mm de largo, ascendentes a adpresas, portando 8 a 12 espiguillas; raquis
de las ramificaciones fuertemente híspido, prolongado 3 a 5 mm más allá de la ramificación terminal, las
espiguillas congestas, traslapándose densamente. Espiguillas de 6 a 11 mm de largo, permanecen insertas
a las espigas; glumas con las quillas híspidas, los pelos largos con tintes purpúreos, primera gluma 2.5 a
3.5 mm de largo, angosta, segunda gluma 5 a 7 mm de largo, híspida en el dorso, ancha; lema de la flor
perfecta de 5 a 6.5 mm de largo, híspida en 3/4 partes de su superficie, especialmente sobre las nervaduras,
con 3 aristas iguales, de 2 a 3 mm de largo; pálea de igual tamaño que la lema o un poco mayor, híspida
sobre los nervios principales hacia el ápice. Anteras de 2.8 a 3.5 mm de longitud, amarillas; estípite glabro;
Flósculo rudimentario 1, reducido a 3 aristas de 2 a 7 mm de longitud, híspidas, la central frecuentemente
con alas membranáceas. Cariopsis de 2 a 2.5 mm de longitud, color amarillo, oblongo-elipsoide. 2n = 20,
40. Florece de agosto a diciembre.
930
IR h Institute of Texas 2(2)
i"
tn da MA
Fic. 8. Di
Distribución.—S de Estados Unidos a todo el O de México.
Hábitat.—Pastizal y matorrales en suelos calcáreos; 50 a 1600 m.
Ejemplares examinados. MÉXICO. Coahuila: Municipio de Piedras Negras, calcareous mesas near Díaz, 20 Abr 1900, 244 m, C.G.
Pringle 9018 (MEXU); Municipio de Saltillo, Buenavista 7 km S de Saltillo 54 Saltillo- Concepción del Oro, Zac., Nov 1975, 1600
m, A. Aznar s.n. (ANSM); 7 km $ de Saltillo, carretera 54 Saltillo-Concepción del Oro, Zac., Sep 1975, 1770 m, R. Palomo s.n. (ANSM).
)
Tamaulipas: Municipio de Jiménez, Ejido Sor Juana Inés de la Cruz, 29 Sep 1993, 150 m, R.A. Carranco 401 (MEXU).
5. Bouteloua curtipendula (Michx.) Torr., Explor. Red River Louisiana 300. 1853. Chloris curtipendula Michx., Fl.
Bor.-Amer. 1:59. 1803. Dinebra curtipendula (Michx.) P. Beauv., Ess. Agrostogr. 98, 158, 160. 1812. Eutriana curtipendula (Michx.)
Trin., Fund. Agrost. 161. 1820. Cynod ipendula (Michx.) Raspail, Ann. Sci. Nat. (Paris) 5:303. 1825. Andropogon curtipendulum
encl. Bot. ed. 2 1:90. 1840, nom. inval. Atheropogon curtipendulus (Michx.) E. Fourn., Mexic. Pl.
(Michx.) Spreng. ex Steud., Nom
2:138. 1886. Tiro: U.S.A. Iruors: 1795, A. Michaux s.n. (HoLOTIPO: P; isoTiro: US fragm.!).
Plantas perennes, por lo general densamente cespitosas; rizomas ocasionalmente presentes. Culmos de 30
u A HP S tal., B tal Aal ERE | M ÁAwi 931
a 100 cm de alto, erectos, simples; estolones ocasionalmente presentes; entrenudos 2 a 3, glabros, nudos
glabros. Vainas con pelos esparcidos de 1 mm de largo; lígula de 0.5 a 1 mm de largo, un anillo denso de
pelos; láminas foliares 5 a 20-35) cm de largo, y 3 a 7 mm de ancho, aplanadas, por lo general esparcido
piloso-papilosas, principalmente en la base y en los márgenes. Inflorescencia de 9 a 20(-30) cm de largo,
con (4-)30 a 80 ramificaciones primarias espigadas de 1 a 3 cm de largo, inclinadas, caedizas en la madu-
rez, portando 2 a 7 espiguillas; espiguillas de 6 a 8 mm de largo, permanecen insertas a las ramificaciones;
glumas desiguales, verdosas con tintes violáceos, escabrosas en la quilla, primera gluma de 3 a 4 mm de
largo, angosta, glabra, segunda gluma de 5.5 a 8 mm de largo, más ancha, aguda hacia el ápice; lema de la
flor perfecta 5 a 7 mm de largo, glabra a escabroso-pubescente, 3-nervada, las nervaduras extendidas como
mucrones hasta de 1 mm de largo. Anteras 3 a 4 mm de largo, color anaranjado, ocasionalmente amarillas.
Flósculo rudimentario 1, lema de 1.8 a 2 mm de largo, con 3 aristas, la central de 5(-10) mm de largo, las
laterales más cortas. Cariopsis de 2 a 2.3(-4) mm de largo, cilíndrica, color amarillo. 2n = 20, 40, 41-64
(Gould, 1979), apomícticas en su mayoría.
E —Gould did reconoce tres variedades. La variedad típica es de amplia distribución en
EUA; en Méxi | jemplares de Coah. y Qro, sin embargo, las variedades caespitosa
y tenuis son comunes y de Ss TET en México.
CLAVE PARA LA DETERMINACIÓN DE LAS VARIEDADES DE B. CURTIPENDULA
1. fole densamente cespitosas, sin rizomas; culmos erectos, sin formar estolones; láminas foliares 3 a 5 mm
ancho, gruesas y rígidas, sin pubescencia de base papilosa 5a. B. curtipendula var. caespitosa
1. arn ligeramente cespitosas, con rizomas o estolones o culmos decumbentes rastreros; láminas foliares
varios; pubescencia de uoce papllesa a veces present
2.
rígidos, fuertes, loma tol láminas foli le (3.535 a 7(-8)
mm bos ancho, color verde- aclote En | pubescencia de base infl a larga con 40 a 70
, anteras típicamente ias O ojo: kot ME 5b. B. curtipendula
var. curtipendula
15 .
2. Ri | + + | ral frAryil J } f, J rM
| | | l SU
foliares de 2 a 4 mm de ancho, color verde-amarillentas, glabras,
f i : 1 r x AT :
a . .
A
a F HY ; Uy Ila
ranja 5c. B. curtipendula
var. tenuis
5a. Bouteloua curtipendula var. caespitosa Gould & Kapadia, Brittonia 16:203, f. 43. 1964. (Fig. 9). Tro:
U.S.A. Arizona: Cochise Co. 3 mi E of Bisbee, 1725 m, 10 Aug 1962, EW Gould 10021 (HoLotiro: TAES; isoripos: ARIZ, DS, GH,
NY, TEX, UC, US-2473990!).
ro
Bouteloua racemosa Lag., Varied. Ci. 2(4): 141. 1805. Ad (Lag.) Roem. & Schult., Syst. Veg. 2:414. 1817. Tipo: MÉXICO:
MA)
Atheropogon aranana E. Fourn., Mexic. Pl. 2:139. 1886. Atheropogon acuminatus E. Fourn. ex Hemsl., Biol. Cent.-Amer., Bot. 3:564.
1885, nom. nud. Bouteloua acuminata (E. Fourn.) Griffiths, Contr. U.S. Natl. Herb. 14(3):406, f. 55. 1912. Tro: MÉXICO: Mirador,
Nov 1941, EM. Liebmann 583 (Lecroriro, designado aquí: P!; isoLecroriro: US-734843!).
Plantas densamente cespitosas, sin rizomas. Culmos 0.5 a 1 m de alto, rígidos, erectos, de base nudosa,
dura pero sin formar estolones. Láminas foliares de largo variable, 3 a 5 mm de ancho, gruesas y rígidas,
sin pubescencia de base papilosa. Inflorescencia de tamaño variable, con 30 a 80 id EE primarias
espigadas caedizas, portando 2 a 7 espiguillas. Espiguillas permanecen insertas a | , de color
cobrizo, café-amarillento a pajizo o verde, con diversas manchas púrpura. Anteras amarillas o anaranjadas,
ocasionalmente rojas o purpúreas. Cariopsis 2 a 2.3 mm de largo, color amarillo, oblongo-elipsoide. 2n =
20, 40, con aneuploide 2n = 42. Florece de junio a diciembre.
Distribución. —Amplia en casi todo México; Sudamérica en: Argentina, Bolivia, Ecuador, Perú, Uruguay
y Venezuela.
Hábitat. —Se le encuentra en una gran diversidad de condiciones que va desde suelos arenosos, rocosos,
arcillosos bien drenados de pastizales, matorrales y bosques de encino y pino; 115 a 2260 m
Ejemplares examinados. México. Coahuila: Municipio de Acuña, Serranías del Burro, Rancho El Bonito, cañón el Toro, Acuña, D.H.
932 J | of the Botanical R h Institute of Texas 2(2)
ES
f^. if
Riskind 2170 (ANSM); Municipio de Arteaga, Sierra Hermosa, en la Sierra de Zapali hacia el O rumbo al Carión del Carbón, 8 Aug
1979, 2260 m, R. López s.n. (ANSM); Municipio de Castaños, Camino al Ejido San Francisco, 8 Jul 1993, 900 m, E. Pérez 20 (MEXU); S
end of Puerto San Lázaro (Cuesta La Muralla), 0.5 Mi N of San Lázaro along route 57, 2 Oct 1960, 1250 m, M.C. Johnston 5882 (TAES);
Sierra de la Gavia, Cañón la Gavia, 6 km S de la casa camino al cañón, 20 Sep 1989, 1480 m, P.S. Hoge 320, M.E. Barkworth & M.A.
Carranza (ANSM); Al pie de la Sierra La Gavia, km 115 carr. 57 Saltillo-Monclova, Y. Herrera Arrieta, M. de la Cerda & O. Rosales 1487
(CIIDIR, HUAA); 25 mi SW de Sabinas, Monclova, F. Wynd & C.H. Mueller 217 (US); Municipio de Cuatrociénegas, Puerto del Norte,
Cuatrociénegas, L.H. Harvey 1208 (US); Sierra Las Cautivas, 31.6 mi W of Ejido Guayabas and 5.4 mi W of Dulces Nombres, 25 Sep 2007,
1833 m, P.M. Peterson 21071, J.M. Saarela (US); Municipio de Francisco I. Madero, Old Candelilla camp, canyon in NO f Si d
Delicias, about 3 km S of Puerta de Sardinas, 9 Aug 1973, 1200 m, M.C. Johnston, F. Chiang, T.L. Wendt, J. Henrickson & D. Morofka
MEXU); Municipio de G l Cepeda, Ejido La Rosa, carretera 40 Saltillo-Torreón, aprox. 20 km NE de General Cepeda, 5 Jul 1986,
1500 m, S. Vásquez 96, A. García (ANSM); 11 km W of Saltillo, G. Mick & K. Roe 19 (ENCB); Municipio de Ji , 6miS of Cd. Acuña,
24 May 1965, EW. Gould 11338 (TAES); Municipio de Morelos, Entre Nava y Morelos, J. Crutchfield, J.C. Marshall & M.C. Johnston 5884
(US); Municipio de Müzquiz, Cerca de Muzquiz, L.H. Harvey 1178 (US); 99.6 mi N of Melchor Muzqui uesta de Malena on Hwy 25
ds Bequillas del Carmen, 01 Sep 2006, 1574 m, P.M. Peterson 19831, S. Lara- Contreras (US); Madera del Carmen, 16.3 mi NE of Los
Uayr, Arriats at al., n ta] "al tn da y 933
Pilares on road towards nico Dos, 21 T dud 2235 m, P.M. d od did J.M. Saarela, S. Lara- Contreras & J. Reyna (US); 22 km
SE of La Cuesta del Pl q q y y from Villa Acuña, 7 Jun 1972, 1000 m, F. Chiang
7539-A, T.L. Wendt & M.C. Johnston (MEXU); 8 km SE de Palau, camino a Barroteras, 24 May 1977, 422 m, J. Valdés-Reyna 937 (ANSM);
La Cuesta del Plomo on lite Müzquiz- lla highway, 14 Sep 1972, 1750 m, F. Chiang 9205, T.L. Wendt & M.C. Johnston (MEXU);
os de Nava, Sierra El Jardín, Canyon del Diablo, 8 mi from jct of road towards Boquillas, 2 Sep 2006, 1336 m, P.M. Peterson
19864, S. Lara-Contreras (US); Sierra Ei Jardin, slopes above Canyon del Diablo, 2 Sep 2006, 1418 m, P.M. Peterson 19889, S. Lara-Con-
treras oe ae El ord. 3 ~ 2006, din m, P. M. Peterson 19915, S. I idiot (US); Municipio de Ocampo, 16 km by winding
] ation de San Vicente, 17 Sep 1972, 1050 m, F. Chiang
9366-G, T.L. Wendt & M. C. Johnston (MEXU), Sierra El Pino, 9.2 km SW of Rancho El Cimarron along the easter slopes, 14 Sep 1991,
1500 m, P.M. Peterson 10628, & C.R. Annable (US); Sierra Madera del Carmen, Villa Ocampo, J. Passini & M.F. Robert 5312 (ENCB); Ma-
dera del Carmen, 9.7 mi NE of Los pilares, 21 Sep 2007, 1902 m, P.M. Peterson 20943, J.M. Saarela, S. Lara- Contreras & J. Reyna (US);
Rancho experimental La Rueda, 87 km NE de Ocampo, brecha Ocampo-Boquillas del Carmen, 2 Jul 1991, 1350 m, M. Vásquez s.n.
(ANSM); D. Ibarra s.n. (CIIDIR); Y. Herrera Arrieta & A. Cortés 1456 (CIIDIR, HUAA); Sierra de la Madera, Rancho Laguna de la Leche,
aprox. 62 km de Ocampo rumbo a Sierra Mojada, 2 Oct 1990, 1000 m, M.A. Carranza 614, FJ. Carranza (ANSM, CIIDIR); Municipio de
Parras, km 30 carretera General Cepeda-Parras, 12 Sep 1986, 1500 m, J. Espinoza 156 (ANSM, TAES); Municipio de Ramos Arizpe, 12
km E of Ramos Arizpe on the Monterrey highway, 24 May 1973, 1250 m, M.C. Johnston 11214-A, T.L. Wendt & F. Chiang (MEXU); km
30, carr. 40 Saltillo-Monterrey, Ramos Arizpe, Y. Herrera Arrieta & A. Cortés 1474 (CIIDIR, HUAA); San Juan de las Bonitas, 19 Jul 1993,
1450 m, F. Alcalá 13 (MEXU); 20 Jul 1993, 1450 m, E. Pérez 29 (MEXU); Sierra de la Paila, Ejido el Cedral por el camino El Carmen, 4
Oct 1989, 1300 m, J.A. Miei 5333, M.A. Carranza & A. Rodríguez E one pi p M.F. Robert 4539, J. Passini (ANSM);
M.F. Robert 4601, J. Passini (ENC Robert 9118, J. Passini (UAG); 9 km S Saltillo on Mex 54 and 2.1 km W
on road to Microondas, just W of Enn Carneros, 18 Oct 1989, 2250 m, P) M. Peterson Be M.A. Carranza (US); 5 mi W of Chapul-
tepec on cutoff road between Hwy 54 & 57, 23 mi S of Saltillo, 16 ct 1995, 2410 m, P.M. Peterson 13269, M.B. Knowles (US); Buenavista,
7 km S de Saltillo, carretera 54 Saltillo-Concepción del Oro, Zac., 28 Jun 1972, 1750 m, R. Almeida 1580, M. Torres (ANSM); Rancho
experimental Los Angeles, 48 km S de Saltillo, carretera 54 paca Concepción del Oro, Zac., 4 Nov 1962, 1800 m, FW. Gould 10302
(TAES); 48 km S de Saltillo, carretera 54 Saltillo-C pci l Oro, Zac., 14 Nov 1992, 1800 m, J.A. Villarreal 6715, M.A. Carranza &
D.E. Lozano (ANSM); Sierra de Zapalinamé, Canón de San liso 5 km S de Saltillo, 25 Aug 1981, 1950 m, S.L. Hatch 4476, C.W.
Morden & J. Valdés-Reyna (ANSM, TAES); Cañón de San Lorenzo, 8 km S de Saltillo, 6 Oct 1979, 1950 m, R. López 6 (ANSM); 2 km al N
de Cañada Blanca, km17 carr. 57 Saltillo-Monclova, Y. Herrera Arrieta, M. de la Cerda €» O. Rosales 1482 (CIIDIR, HUAA); 22.5 mi SW of
Saltillo, FW. Gould 8693 (ENCB); Ls ein Torreón, Sierra de Jimulco, N-facing cliffs, NO-facing notches, 27 Jun 1972, 1600 m,
M.C. Johnston 11485, T.L. Wendt & F. Chiang (MEXU); E of Torreón, 5.1 km SE of Estacion La Cuchilla on Mex 40 to Saltillo, 20 Oct 1989,
1100 m, P.M. P di 8446, J. a ma J.A. Villarreal & P.A. Fryxell (US). Nuevo León: Municipio de Cadereyta Jiménez, En los
alred de Cadereyta, 22 Mar 1993, 320 m, N. Bazaldua 36 (MEXU); Municipio de Carmen, Meseta de Chipinque, San
Pedro Garza a 5 Jun 1992, M.M. Castillo 7 (MEXU); Municipio de Dr. A 12 mi N of Matehuala, % mi N of San Luis Potosi
line, 16 Sep 1963, 1800 m, F.W. Gould 10753 (TAES); Ejido Lagunita y R r. Arroyo, M.E. Pérez 120 (CIIDIR); Municipio
de Galeana, 1 mi E of San Marcos, 13 mi SE of Galeana, 13 Sep 1963, 1850 m, FW. pus 10726 (TAES); 1.5 km E of El Barrosito, 1.5
m O of Puerto Prieto, in S part of Sierra la Tomita, 19 Jun 1972, 2100 m, F. Chiang 8014, T.L. Wendt & M.C. Johnston (MEXU); 7 km of
hwy 57, on the ib to El ae near Cerro de la Conformidad, 19 Jun 1972, 2150 m, F. Chiang 8029a, T.L. Wendt & M.C. Johnston
(MEXU); km San Roberto, 18 Nov 1987, 1900 m, I. Cabral 1084 (ANSM, IEB); Sierra Madre Oriental, about 15
mi SO of Galeana, 13 Jul 1934, C.H. Mueller 1087, M.T. Mueller (MEXU); Galeana, V.H. Chase 7632 (US); SW de Galeana, C.H. Mueller &
H. Mueller 497 (US); Municipio de Iturbide, 1 mi O of Iturbide, 30 Sep 1976, 1570 m, J. Brunken 203, C. Perino (TAES); 11 mi E of Itur-
bide, Mex. 60, 30 Sep 1976, 700 m, J. Brunhen 198, C. Perino (MEXU, nid Municipio de Juárez, Ra. San Marcos, km 7 camino a
San Mateo, carretera Villa de Juárez- Cadereyta, 1 Jun 1993, J 28 XU); Municipio de I , Aprox. 5 km O del
Rancho Pascualito, 16 Feb 1980, 440 m, O.L. Briones 107 (ANSM); Municipio de Li ,Los Pinos, 12 Oct 1980, 1300 m, J.J. Ortiz s.n.
(ANSM); Rancho San José de los Hoyos, km 15 carretera Linares- iturbide 1 Abr 1993, J. Garza 20 (MEXU); 11 mi E de Iturbide, carr.
Linares-Galeana, M.C. Johnston 5841 (US); M , Facultad es e Universidad Autónoma de Nuevo León, km
Marín, 22 Oct 1992, M.M. Castillo 25 (MEXU); Municipio de Mont indad del Cerro del Hp ain uad F. D MCA. 14497
(US); km 105 carr. pi erralvo, N.I Bendech 205 (CIIDIR); sic de Sabinas Hidalgo, Carrete abinas Hidal
go, km 82, 24 Mar 1993, 380 m 1 inas Victoria, Sierra de dac 29 Ene 1993, M.M. Castillo
42 (MEXU); Municip ti novevo, 11 Sep 1986, 1150 m, 1. Cabral 515 (ANSM). Tamaulipas: Municipio de Busta-
mante, 10 km al NE de la une 5 Aug 15 1985, 1800 m, P. Hiriart 846, R. Poen & V. Juárez (MEXU); 2 km NO de Felipe Angeles
mbo a Bustamante, 9 Nov 1984, 1600 m, R. Díaz 180 (UAT); N Al 4 km al NO de González, 11 Jun 1992, 115 m,
" G. Galván 272 (MEXU); 5 km al N de González, Mar 1991, 110 m, J. G. Galván 259 (MEXU); Municipio de Jaumave, 7 km al N de Mag-
daleno Aguilar (Santiaguillo), 18 Sep 1976, 1850 m, F. González 9801, A. Castellanos & P. Zavaleta (MEXU); Sierra Madre rumbo a Jau-
mave, R. Carranco 089 (IEB); Municipio de Llera, La Herradura, 7 Oct 1980, 380 m, C. Pantoja 1 (MEXU); Municipio de Miquihuana, 2
km al SO de Miquihuana, 13 Aug 1989, 1600 m, ue Mora zm die UAT); Ejido Valle d Miquihuana, R. Carranco 062 (IEB);
3 mi al N de Miquihuana, L.R. Stanford 2465 illas, San Pedro, 6 km al NO de Municipio Palmillas, 13 Aug 1984,
1600 m, P. Hiriart 227, F.V. Juárez, M. González, A. McDonall, M. Ortega y R. Díaz (MEXU); ud. de San Carlos, Cerro del Diente, J.A.
Barrientos 080 (IEB); Municipio de Tula, 11 Mi S of Palmillas on the road to Tula, 18 Sep 1960, M.C. Johnston 5628, J. Crutchfield (MEXU,
934 Journal of the Botanical R h Institute of Texas 2(2)
US); La Tapona, 21 Oct 1982, 1900 m, R.A. Carranco 159 (MEXU); Municipio de Victoria, Altas Cumbres, 18 Oct 1982, 1240 m, J.G.
Galván 55 (MEXU); 11 mi SW de Victoria, L.R. Stanford 1995 (US); Municipio de Villagrán, 1 Mi E of Ejido de San Lázaro, 11 Oct 1959,
m, M.C. Johnston 4290, J. Graham (MEXU); On highway 85, between Linares and Ciudad Victoria, 111 km NO of Ciudad Victoria,
e 1977, 415 m, T.B. Croat 39199 (MEXU).
5b. Bouteloua curtipendula var. curtipendula (Fig. 10).
Atheropogon apludioides Muhl. ex Willd., Sp. Pl. 4:937. 1806. Tiro: NORTH AMÉRICA: (HoLotiro: ?; isoriro: US-1535780 fragm.!).
Eutriana affinis Hook. f., Trans. Linn. Soc. London 20:174. 1847. Atheropogon affinis (Hook. f.) E. Fourn., Mexic. Pl. 2:141. 1886. Tiro:
U.S.A. Missourt: St. Louis, Schweinitz s.n. (sitio: K); Texas: Drummond s.n. (siNriPo: US-1063204 fragm.!).
Bouteloua curtipendula var. aristosa A. Gray, Manual (ed. 2) 553. 1856. Bouteloua racemosa var. aristosa (A. Gray) S. Watson & J.M. Coult.,
Manual (ed. 6) 656.1890. Tiro: U.S.A. Iuuwors: Geyer s.n. (HoLoTIPO: GH!)
Plantas ligeramente cespitosas, con rizomas presentes, delgados o fuertes. Culmos erectos, rígidos, fuertes,
solitarios o en pequeños grupos. Láminas foliares de (3.535 a 7(-8) mm de ancho, aplanadas, color verde-
eos con eas de base papilosa. Inflorescencia tipicamente larga con 40 a 70 ramificaciones
, reflejas, sobre un eje fuerte, portando 3 a 7 espiguillas. Espiguillas permanecen
aoe a las A glumas y lema típicamente color púrpura o teñidas de púrpura. Anteras color
rojo o rojo-anaranjado, muy esporadicamente anaranjadas, amarillas o purpúreas. 2n = 40, se le conocen
varios casos de aneuploidía con 41 a 64 cromosomas. Florece de junio a diciembre.
Distribución.—Desde el S de Canadá, Planicies del centro de Estados Unidos al Sur hasta México en:
Coahuila, Chihuahua, Durango, Nuevo León, San Luis Potosí y Tamaulipas.
Hábitat.—Especie característica de pastizales, matorrales y bosques de encino y pino sobre suelos
arcillosos y bien drenados; 60 a 2500 m.
Ejemplares examinados. MÉXICO. Coahuila: M de Acuña, 1936, (MEXU); Municipio de Allende, 13 Mi SO of Ciudad Allende
toward Sabinas, 7 Oct 1959, 488 m, M.C. Johnston 4176, J. Graham (MEXU); Municipio de Area Arroyo Canoas, 28 Aug 1993, 1600
m, F. Alcalá 23 (MEXU); Municipio de Castaños, La Muralla km 135 carretera Saltillo-Monclova, 15 Nov 1974, 750 m, H.M. Garza s.n.
(MEXU); Municipio de Castaños, Paso de San Lázaro, Sierra La Gavia, 37.6 Mi S de Monclova, carretera 57, 22 Sep 1990, 1250 m, P.M.
Peterson 9982, C.R. Annable & J. Valdés-Reyna (ANSM, US); Sierra de la Gavia, Cañón de la Gavia, aprox. 6 km S de la casa camino al
cañón, 14 Aug 1986, 1480 m, J. Valdés-Reyna 1712, M.A. Carranza & D. Frame (ANSM, IEB); Municipio de Cuatrociénegas, Vecindad de
Aguaje del Pajarito, Cañón al W de Sierra de la Fragua, 2-3 kmN de Puerto Colorado, 1-3 Sep 1941, I.M. ee 8792 (US); Cañón del
Agua (N- Rs apium of on rocky hilltop along road 0.1 mi N of Ranchito, 26 Aug 1977, 1420 m, J. Valdés- bi diis T.L. ada
(ANSM); Cerros entr g Ocampo- -Cuatrociénegas 8 Jul 1987, 2500 m, A Rodríguez 939, J J. López & J.L
4
Sierra de San Marcos, cañón Grande, Ejido Estanque de Norias, aprox. 43 km O de la carretera 57 rumbo a Ejido Reforma, 1 Oct 1993,
0 m, M.A. pem 1709, d pe Encina, J. Fierro us R. es sities Cañón del agua, Cuatrociénegas, J. Valdés-Reyna & T. Wendt
VR-1009 (ANSM); Municip Sierra d in Cañón del Chilpitín, 5 Mi up (E) from mouth of canyon, near El Chil-
pitín, at juncti f mai yon, 7 Sep 1976, 1060 m, T.L. Wendt 1686, D.H. Rishind (MEXU); Municipio de Müzquiz, El Sauz, entron-
que carretera 57, 32 km S de Sabinas, 5 Sep 1990, 420 m, R. Vásquez 217 (ANSM); Palm Canyon, 19 Sep 1936, E.G. Marsh Jr. 980 (MEXU);
Sierra del Carmen, Ejido San Francisco, 5 Sep 2005, 1430 m, P. M. Peterson 18840, J. Valdés-Reyna & C. Sifuentes (US); Sierra rin del
Carmén, 17 Sep 1989, 1850 m, E.A. Estrada 1839 (ANSM); Sierra del Carmén, Cañón de la Carbonera, 12 Aug 1975, 1900 m, M.F. Robert
5633, J. Passini (ANSM); Canyon del Centinela, on Hacienda Piedra Blanca, 10 Jul 1936, 2200 m, L.F. Wynd 642, C.H. Muller (ANSM);
Mesa del Cinco, 13 Aug 1975, 2480 m, M.F. Robert 5528, J. Passini (ANSM); Sierra Maderas del Carmen, 17 Sep 1989, 1800 m, E.A. Es-
trada s.n. (ANSM), Sierra Madre Oriental, Cuesta la Encantada, 16 Aug 1975, 1400 m, M.F. Robert 5661-A, J. Passini (ANSM); Municipio
de O eor 5 d of Santa Elena. Vicinity of Santa Elena Mines, cal s Cruces, 16 Jul 1941,
(U); Si El Pino, 33.5 km W of indi El UMEN 14 Sep 1991, 1500 m, P.M. Peterson 10680, 10681 & C.R.
Annable (US); Cuesta de Zozoya, aprox. 38 km de O Mojada, 3 Oct 1990, 1600 m, M.A. Carranza 677, FJ. Carran-
a (ANSM); Ocampo, Aug 1972, 1240 m, R. Vásquez y Noreh (ANSM); Rancho experimental La Rueda, 87 km NE de Ocampo, brecha
Ocampo- as del Cármen, 31 Jue 1979, Pond m, M.T. Ruíz, M.A. Carranza & M. Vásquez (ANSM); D. i (ANSM); Sierra de la
Madera, 25 Jul 1973, J.S. A del Cármen Campo 1, 12 Aug 1975, 1750 m, M.F. Robert 5555, J. Passini (ANSM);
Sierra del Pino, Cañón de la Vaca 14 Aug 1975, 1450 m, M.F. Robert 5224, J. Passini (ANSM); Sierra La Encantada, rancho Puerto del
Aire, 8 Sep 1990, 1250 m, M.A. Carranza 729, J. Valdés-Reyna, P.A. Fryxell, R. Vásquez & O. Meza (ANSM, UAT); Sierra Madre Orientale,
San José de las Piedras, 15 Aug 1975, 1200 m, M.F. Robert 5661, J. Passini (ANSM); Municipio de Parras, 9 km S of Parras on Sierras
Negras, 3 Jul 1941, 2400 m. L.R. Stanford 195, K.L. Retherford & R.D. Northcraft (MEXU); Municipio de Ramos Arizpe, Sierra de la Paila,
Ejido el Cedral por el camino El Cármen, 26 Aug 1992, 1300 m, J. Valdés-Reyna 2171 (ANSM); Municipio de Sabinas, 35 mi NW del
Rancho Margaritas, 100 mi NW de Sabinas, F.W. Gould 10681 (US); 26 Jul 1973, 330 m, 2567 (ANSM); 85.4 km NW of sd ue on iid
53 towards Boquilla del Carmen, vía La Rosita, 12 Sep 1901, 810 m, P.M. Peterson 10574, C.R. Annable (US); M p km
E de Saltillo, carretera Saltillo-Torreón, 6 Nov 1972, 1400 m, R. Almeida 1631, 1640 (ANSM); A 2 km por la desviación Estación i.
L
m
B
sa da RA kere 935
E eden
ES
tn An RAK
Fic. 10 nie sz I^ Dor.
do, carretera Saltillo-Torreón, 19 Oct 1986, 1620 m, E. García s.n. (ANSM); About 4 Mi O of Saltillo, 12 Aug 1950, J.R. Reader 1656, C.G.
Reeder & L.N. Goodding (MEXU, TAES); Above the road on Diamonte, 14 Mi S of Saltillo, 2 Jul 1947, F.A. Barkley 7193, G.L. Webster &
C.M. Rowell Jr. (MEXU); Area de ipal, Buenavista, 21 Sep 1973, 1756 m, R. Almeida 2600 (MEXU); Cañón de San Lorenzo
rumbo a los Aguajes, camino a los Burros, 17 Aug 1979, 1900 m, R. López s.n. (ANSM); El Refugio, S de Saltillo, Jul 1942, 1600 m, E.
Lyonnet 3551 (MEXU); La Encantada a 17 km de Saltillo tera a Zacatecas, 9 Jun 1972, 1880 m, R. Almeida 1342, B.J. García s.n.
(ANSM); Lomas las Tetillas, 19 Jul 1993, 1700 m, J.A. García 19 (MEXU); R peri tal Los Angeles, 48 km S de Saltillo, carre-
tera 54 Saltillo-Concepción del Oro, Zac., 30 Jun 1978, 2000 m, J.S. Sierra s.n. (ANSM); Rancho Los Angeles, km 30 carretera Saltillo-
C pción del Oro, Zac., 31 Oct 1978, F. Cárdenas, M.A. Bernal (MEXU); Rumbo al Cañón El Cuatro, 19 Jul 1993, 1940 m, P. Moya 421
(MEXU); Saltillo, 1898, E. Palmer 407 (MEXU); Sierra de Catana, 40 km SO de Saltillo, 29 Jul 1992, 2700 m, J.A. Villarreal 6572, M.A.
Carranza., J. Valdés-Reyna, M. Vásquez y D.E. Lozano (ANSM); Sierra de Zapalinamé, 5 km S de Saltillo, in San Lorenzo canyon, 25 Aug
1981, 2000 m, S.L. Hatch 4493, C.W. Morden & J. Valdés-Reyna (ANSM, ENCB); Sierra de Zapalinamé. Lomas de Lourdes, 3 Oct 1974,
1800 m, V.M. Valdés s.n. (ANSM); Municipio de San Buenaventura, Sierra La Encantada, rancho Carrizalejo, entrada S al rancho Puerto
del Aire, 8 Sep 1990, 1200 m, M.A. Carranza 771, J. Valdés-Reyna, P.A. Fryxell, R. Vásquez & O. Meza (ANSM); Municipio de Buenavista,
29 Oct 1974, 1700 m, G. I. Flores s.n. (MEXU); 7 km S de Saltillo 54 Saltillo- Concepción del Oro, Zacatecas, 15 Oct 1974, 1750
e]
a A laf tha Botanical D h Institute of Texas 2(2)
m, Á. Rodríguez 104 (ANSM); 7 km S de Saltillo tera 54 Saltillo-C ión del Oro, Zac., 28 Aug 1982, 1756 m, J.A. Villarreal 1692
(ANSM); 7 km S de a carretera 54 cared Concepción del O Tae, 13 Aug 1980, 1850 m, J.A. Villarreal 862, R. López (ANSM).
Nuevo Leon: ón de Bust te, 18 Jun 1993, 600 m, P. Jáuregui 68 (MEXU); Grutas de Bustaman-
te, 18 Jun 1993, 1300 m, P. Jauregui 77 ous Municipio de Dr. Bree E js km W of Dr. Arroyo and 37 km E of Matehuala on ee
3, 13 Oct 1991, 1950 m, P.M. Peterson 11138, C.R. Annable (US); M l ,3 km S of El Salero, 19 Sep 2003, 1869
Peterson 17836, J. Valdés-Reyna & H.S. Hinton (US); 25 km E of San Robério along highway 58, 27 Aug 1981, 1740 m, S.L. ms nee
C.W. Morden & J. Valdés-Reyna (ANSM, TAES). Tamaulipas: Municipio de Aldama, 31 May 1980, R. Sandoval 9 (MEXU); Municipio de
Hidalgo, Ejido Nicolás Bravo, 29 e 1981, J. Cantú 41 (MEXU); Municipio de Llera, About 30 Mi S of Ciudad Victoria on highway 85,
toward Ciudad Mante, 28 Sep 1976, 450 m, F.W. Gould 14817 (ANSM, ENCB); Municipio de Victoria, Altas Cumbres, 19 Oct 1982, 1220
m, J.G. Galván 60 (MEXU). — am Allende, ini. highway 85, 2 Mi S of Allende. Just S of river and bridge, 13 Oct 1980, J.A.
Wussow 258, G. Landry (MEXU); ^ de G 1 Mi S of Saltillo on México 57, 1 Oct 1960, M.C. Johnston 5863*, J. Bio
(MEXU); 25 km E of San Roberto along TM 58, 27 ud 1981, 1740 m, S.L. Hatch 4570, C.W. Morden & J. Valdés-Reyna (ANSM); 2
W of Dieciocho de Marzo up road towards Cerro Potosí, 18 Oct 1995, 2130 m, P.M. Peterson 13322, M.B. Knowles (US); En los iS
res del poblado Santa Clara de González, 16 Jul 1995, 1940 m, N. Bazaldua 112 (MEXU); Hacienda de San José de Raices, 31 Jul 1935,
C.H. Mueller 2309b (MEXU); La Becerra, 12 Sep 1989, 2100 m, G.B. Hinton 19681 (MEXU); Municipio de General Zaragoza, Peña Neva-
da, 16 Nov 1977, H. Brailovsky (MEXU); 42 km SW of Hwy 85 on road towards Dulces Nombres, 23 Sep 2001, 1145 m, P.M. Peterson
15907, J. Valdés-Reyna (US). Municipio de Guadalupe, La Pastora, near La Ciudad de los Niños, Guadalupe, 15 Aug 1983, 500 m, S.L.
Hatch 4837, J. Valdés-Reyna & J. Kessler (ANSM); Municipio de Iturbide, 5 Mi above (O of) Iturbide on the me highway, 30
Sep 1960, M.C. Johnston 5849 (MEXU); Cañada Marrubial, Santa Rosa, 7 Jul 1985, p m, B.A. i ino 505 (MEXU); Municipio de Li-
nares, Ejido Rancho Viejo, 9 Oct 1980, 850 m, J.J. Ortíz s.n. (ANSM); Localizad galar, 1 Linares-
San Roberto, km 12, 2 Jun 1983, 440 m, M.M. Castillo 100 (MEXU); Municipio de M y, Sierra Mad ins, 29 Jul 1933, C.H.
Mueller 422, M.T. Mueller (MEXU); Santiago, 22 km O de Villa de Santiago, 2 Aug 1976, 1800 m, V. Valdés-Reyna 108 (ANSM). Tamau-
lipas: Municipio de Bustamante, 15 km al S de Miquihuana, 23 Aug 1984, P. Hiriart 300, V. Juárez & R. Molczadzki (MEXU); Cerro Ti-
naja. Vicinity of San José, 14 Jul 1930, 869 m. H.H. Bartlet 10324 (MEXU, US); Municipio de González, 14 mi E of Mante toward Gonzá-
lez, 10 Dic 1959, 183 m, M.C. Johnston 4917 (MEXU); Municipio de Gómez, Aserradero Revilla, 4 Nov 1982, 1700 m, G. Villegas 539
(MEXU); Municipio de Jaumave, Altas Cumbres km 160 carretera Victoria-Jaumave, 29 Jun 1985, 950 m, M. Martínez & Díaz de Salas
731 (UAT); S de Alta Cumbres, 10 Sep 1984, 1200 m, A. McDonald 853 (UAT); Municipio de Hidalgo, Sierra La Lagunita, 6.4 mi SE of
Aramberri on road towards Agua Fría, 19 Sep 2002, 1397 m, P.M. Peterson 16678, J. Valdés-Reyna & M. Sosa (US); Municipio de Miquihua-
na, Sierra de Los Lobos, Colonia Agrícola la Pena, 26 Nov 1981, 1800 m, J.A. Barrientos s.n. (MEXU); Municipio de San Carlos, Cerro de
San Carlos, 21 Aug 1980, 1000 m, G. Villegas 567 (MEXU); E La Gavia, 3 Sep 1981, 300 m, J.F. Iribe 73 COCA); Municipio de Soto la
Marina, 3 mi N of Soto la Marina on the road to Jiménez, 11 Dic 1959, 183 m, M.C. Johnston 4973, J. Crutchfield (MEXU); Ejido Verde
Chico, 23 Oct 1986, 60 m, J.F. Iribe 327 (MEXU); Municipio de Tula, 2 km al N de 2 veneer cerca de las Antonias, 12 Au
1300 m, F. González 4601, R.M. López & R. Dirzo (MEXU); 30 km al SO de Tula d Luis Potosí y Tamaulipas),
9 Aug 1972, 2000 m, i es 4439, R.M. López & R. Dirzo (MEXU); 5 km al O del Ejido El je 3 Jul 1985, 1500 m, P. Hiriart
759, V. Juárez & R J}; Ejido La Presita en el km 66 carretera Tula-Victoria, 18 Jul 1986, 1600 m, M. Mart & Díaz de
Salas 1221 (MEXU, UAT); Municipio de Victoria, 11 mi O of Victoria toward Jaumave, 29 Sep 1951, 914 m, M.C. Johnston 4141D (MEXU);
Los San Pedros, 18 Sep 1980, R.A. Carranco 615 (MEXU); On Jaumave road about 13 mi SO of Ciudad Victoria; near summits of Sierra
Madre, 13 May 1949, 1000 m, R. McVaugh 10500 (MEXU); 5 km O de Palmillas carretera Victoria-San Luis Potosí, 13 Jul 1984, 1580 m,
R. Díaz 8 (UAT); Municipio de Ciudad Victoria, 25 May 1988, 320 m, J. Delgado s.n. (UAT); Municipio de Villagrán, 3 mi N of Villagrán
on the Victoria-Linares highway, 12 Nov 1959, 488 m, M.C. Johnston 4672% J. Graham (MEXU).
5c. podus curtipendula var. tenuis Gould & Kapadia, Brittonia 16:201.1964. (Fig. 11). Tro: MÉXICO.
TECAS: 10 mi NW of Somrerete (68 mi SE of Durango), 30 Oct 1959, EW. Gould 9000c (notorio: TAES; isoripos: TEX, UC,
US- —
Plantas ligeramente cespitosas, rizomas generalmente ausentes; culmos delgados, fragiles y decumbentes
formando estolones bien desarrollados o culmos estoloniferos. Laminas foliares 8 a 15 cm de largo y 2 a
4 mm de ancho, aplanadas, de en la base y largamente apiculadas, color verde-amarillentas, las basales
enroscadas cuando secas. Inflorescencia con 4 a 9 ramificaciones primarias espigadas espaciadas en el
raquis; espiguillas color café a cobrizo, con tintes purpüreos. Anteras amarillas o anaranjadas. Cariopsis de
2 a 2.2 mm de largo, color amarillo, oblongo-elipsoide. Plantas sólo diploides 2n = 20 y aneuploide 2n = 42
cromosomas. Florece de julio a noviembre. Mapa 5c.
Distribución.—Especie mexicana encontrada en casi todo México.
Hábitat.—Pastizales abiertos sobre suelos fértiles; 710 a 1750 m.
Ejemplares examinados. MÉXICO. Coahuila: Municipio de General Cepeda, Ejido La Rosa, carr. 40, Saltillo-Torreón, aprox. 20 km
al NE de Gral. Cepeda, S. Vázquez & A. García 96 (ANSM, SLPM). Nuevo León: Municipio de Montemorelos, Rancho Sta. Ana, 2 kmal
E de Estación Huerta, aprox. 37 km SE de Montemorelos, FW. Gould 12800, 12860 (US); Dulce Nombre, F.G. Meyer & D.J. Rogers 2888
] Aal ta fa MAG 937
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aes
Fis. 11 DRiertelb ry In D
pio de Sabi Hidalgo, P M lique, 7 Jul 1940, I.K. Langman 2447 (US). Tamaulipas: 11 mi S of Palmillas camino a
Tula, M.C. Johnston & C. McMillan 6088 (US); M.C. Johnston & J. Crutchfield 5628, 5722%, 6074 (US); F. Martínez & G. Borja F-2262, F-2330,
F-2365 (US); J.R. Swallen 1685 (US); Hacienda Buenavista, 21 Jun 1919, E.O. Wootom s.n. (US); Municipio de Victoria, Altas Cumbres, 7
Oct 1976, 710 m, G. Villegas 17 (MEXU).
6. Bouteloua dactyloides (Nutt.) Columbus, Aliso 18(1):63. 1999. (Fig. 12). Sesleria dactyloides Nutt., Gen. N. Amer.
Pl. 1:65. 1818. Calant! dactyloides (Nutt.) Kunth ex Hook., Hooker's J. Bot. Kew Gard. Misc. 8:18. 1856. Buchloe dactyloides (Nutt)
Engelm., Trans. Acad. Sci. St. Louis 1:432, pl. 12 & 14, £ 1-17. 1859. Casiostega dactyloides (Nutt.) E. Fourn., Bull. Soc. Roy. Bot.
Belgique 15:470. 1876. Bulbilis dactyloides (Nutt.) Raf. ex Kuntze, Revis. Gen. Pl. 2:763. 1891. Tiro: U.S.A.: Plains of the Missouri
River, T. Nuttall s.n. (noLoriPo: BM; setro: PH?)
(US; M icipi
Planta baja, estolonífera, dioica u ocasionalmente monoica, formadora de césped. Culmos de 5 a 15 cm
de longitud, estolones con entrenudos de 1 a 4 cm de longitud. Vainas estriadas, glabras o pilosas en los
márgenes superiores y en la garganta. Lígula ciliada, de 0.8 a 1 mm de longitud. Láminas foliares de 6.5 a
10 cm de longitud, de 1 a 1.5(—2) mm de ancho, planas, enrolladas, escabrosas en ambas superficies. Inflo-
938 Journal of the Botanical R h institute of Texas 2(2)
Fic. 12 Mietribiirióm ra 2. L r} nd I ta da Y!
rescencia estaminada exserta, con 1 a 2(-4) ramificaciones primarias espigadas, de 1 a 1.5 cm de longitud.
Espiguillas de 4 a 5 mm de longitud. Flores casi transparentes. Glumas subiguales de 1.5 a 4 mm de longitud,
uninervadas; la segunda en ocasiones corto-aristada. Lema de 3 a 4.8 mm de longitud, con el ápice agudo
y mucronado. Pálea del mismo tamaño que la lema. Anteras color naranja-rojizo o salmón. Inflorescencia
femenina incluida en las 2 vainas superiores, que son cortas y ensanchadas. Ramificaciones primarias es-
pigadas 2, de 6 a 7 mm de longitud, de 3 a 4 mm de ancho. Espiguillas color crema, pubescentes o glabras,
con dientes o lóbulos en ocasiones aristados, en algunas espiguillas aristas ausentes; flores dorsalmente
comprimidas. Segunda gluma cartilaginosa, con nervios poco notorios. Lema de 4 a 5 mm de longitud,
levemente coriácea, lustrosa. Pálea un poco más pequeña angosta hacia el ápice, bidentada. Colectada con
flores en todos los meses del año.
Distribución. —Amplia, conocida del SW de Canadá; planicies del W de EUA; N y Centro de México.
Hábitat.—Pastizales, matorrales y bosques, en la orilla de caminos; 244 a 2700 m.
Harrara Árriata et al., B tal dal tn dan Mo: 939
Ejemplares examinados. MEXICO. Coahuila: Municipio d Arteaga, Ejido La Escondida, 21 Jun 1994, 2460 m, J.A. García 48 (MEXU);
21 Jun 1994, 2460 m, E. Pérez 56 (MEXU); Poblado La Ciruela, 22 Jul 1993, 2600 m, P.A. Lobato 22 (MEXU); Puerto Los Tejocotes en la
Sierra de Zapalinamé, 21 Jun 1979, 2260 m, R. López s.n. (ANSM); Sierra de los Lirios, Jul 1942, 2600 m, E. Lyonnet 3693 (MEXU); Sierra
Hermosa, en la Sierra de Zapalinamé hacia el O rumbo al Cañón del Carbón, 8 Aug 1979, 2270 m, R. López s.n. (ANSM); Municipio de
Muzquiz, Sierra del Carmen at Cuesta de Malena, 6 Sep 2005, 1630 m, P.M. Peterson 18863, J. Valdés-Reyna & C. Sifuentes (US); Municipio
de Nava, Level plains of the Río Grande Valley, just N of Río Escondido, 18 km SSO of Piedras Negras on México No. 57, 12 Sep 1978,
300 m, H.H. Iltis 21 (MEXU); Municipio de Parras, Arroyo seco en el arroyo El Tunal, 10 Aug 1984, 2000 m, A. Rodríguez 1437, M.A
s (ANSM); Municipio de Ramos Arizpe, La Sauceda, 5 Aug 1976, 1030 m, 5. podia s.n. — Sierra de la Paila, Ejido E
ral por el camino El Carmen, 26 Aug 1992, 1300 m, J. Valdés-Reyna 2163 (AN illo, Sierra El Astillero, 1.8 mi
SW mu Santa Rosa and 5.4 mi SW of San dies at trail naa db water pun. 21 Oct 2007, 2550 m, P.M. Peterson 21441, J.M. Saarela
& D. Stancik (US); 12 km N of the Coahu ] 54 between Saltillo-Concepción del Oro, Zacatecas, 26
Aug 1981, 1900 m, S.L. Hatch 4514, C.W. Morden & J. Maud RON ANS: 3kmS de la Ciudad de Saltillo, fracc. Parque de la Cañada,
16 Jul 1993, 1700 m, J. Valdés- pa M.A.C lad Autónoma Agraria Antonio Narro, orillas de la
carretera Saltillo-C l Oro, Zac., 6 Sep 1972, 1750 m, P.E. García s.n. (ANSM); 50 km S de Saltillo, carretera 54 a Concepción
del Oro, Zacatecas, tias El colere; >un Vir 2000m 2 A. Kaa 2060 J Valdés-Reyna & M.A. Carranza (ANSM); At Fraile,
59 km S of Saltillo. In playa valleys hills, 10 Jul 1941, 1967 m, L.R. Stanford 271,288, K.L.
Retherford & R.D. Northcraft (MEXU); Cañón de San dod. ubi demere 1 km E del pozo número 2, 22 Jul 1979, 1700 m, R.
López, J. Valdés-Reyna (ANSM); Los Ramones, 20 Jul 1993, 1550 m, E. Pérez 27 (MEXU); Rancho Los Angeles, km 30 carretera Saltillo-
Concepción del Oro, Zacatecas, 31 Oct 1978, 1960 m (MEXU); Rumbo le ipe E] Cuatro, 19 Jul 1995, 1940 m, P. Moya 422 (MEXU);
Saltillo, Apr 1898, E. Palmer 7 (MEXU); Sierra de Catana, 40 km SO , 29 Jul 1992, 2700 m, J.A. Villarreal 6570, M.A. Carranza,
J. Valdés-Reyna, M. Vásquez y D.E. Lozano (ANSM); Buenavista, 1 Oct 1991, 1870 m, Brigada Coahuila 86, 90 (MEXU); 7 km S de Saltillo,
carretera 54 Saltillo- Concepción del Oro, Zac., 12 Jul 1974, 1775 m, J.F. Cano 106 (ANSM). Nuevo León: Municipio de Anáhuac, Dis-
trito de Riego 04, Don Martín, 10 Abr 1985, 170 m, P.M. De la Garza (ANSM); Municipio de China, Rancho El Reparo, 25 Oct 1983,
150 m, J.F. Iribe 200 (MEXU); Municipio de Doctor Arroyo, Ejido La Escondida, 5 km N carretera 102 Doctor Arroyo-Galeana, 17 Jun
1992, 1740 m eee ipii 6522 M.A. nee oe Municipio de Galeana, Ejido 18 de Marzo, Nov 1980, 2000 m, J.A. Ochoa 982
(MEXU); E lred l d lat ález, 16 Jul 1993, 1940 m, N. Bazaldua 105 (MEXU); Hacienda Pablillo, 24
Aug 1936, M. oe 208 (MEXU); Sierra Madre Oriental; Pablillo about 15 mi SO of Galeana, 18 Jul 1934, 2438 m, C.H. Mueller 1086,
M.T. Mueller (MEXU), El Salero, 19 Sep 2001, 1840 m, P.M. Peterson 15821, J. Valdés-Reyna (US); Municipio de Lampazos de Naranjo,
Carretera a Lampazos cerca del rancho Santa beds 29 Sep 1980, 370 m, O.L. Briones 356 (ANSM); Municipio de Marin, A orillas del
canal Marín, carretera Marín-Higueras a 3 km de la cabecera municipal de Marín, 1 Jul 1993, P. Jáuregui 102 (MEXU); Municipio de
Salinas Victoria, La Soledad Salinas Victoria, ee 1981, 500 m, SA A. Ochoa 1134 (MEXU). Tamaulipas: Municipio de Burgos, Rancho
Labra, 20 Oct 1983, 150 m, C.R López 112 (MEXU); Munici[ te, Ejido La Cardona, 20 Oct 1982, 2050 m, G. Villegas 560
(MEXU); scan En 11 miN of Santander Jimé 1 San F highway, 25 Nov 1959, 244 m, M.C. Johnston 4829,
J. Graham (MEXU); ^ San Fernando, Ejido Palo Solo, 10 Feb 1994, 10 m, J. F. Iribe 396 (MEXU); Rancho El Canelo, 30 May
1980, 40 m, J.F. Iribe 2 (MEXU); Municipio de Soto la Marina, Ejido Charcos, 31 Oct 1984, R.A. Carranco 203 (MEXU); Municipio de
Tula, Joya 1, 28 Oct 1982, 1050 m, G. Villegas 549 (MEXU).
7. Bouteloua erecta (Vasey & Hack.) es a 18:63. 1999. (Fig. 13). Cathestecum erectum Vasey & Hack.,
Bull. Torrey Bot. Club 11:37, t. 45. 1884. Tro: U.S.A s: Bone Spring (Tornillo Cr.), between El Paso and Presidio, Aug 1883, V
Havard 2 (ectotiro: US-884020! designad GJ. Pierce en 1979 en la herb., validado en Contr. U.S. Natl. Herb. 41:38. 2001).
FE
Plantas perennes. Culmos de 10 a 30 cm de alto, erectos o algo geniculados, simples o ramificados, es-
toloníferos, estolones delgados y correosos, entrenudos alargados y arqueados, nudos pubescentes. Vainas
basales aglomeradas y pilosas en las garganta, las superiores distantes y glabras; lígula de 0.2 a 0.3 mm de
largo, ciliada; láminas foliares de 2.5 a 6 cm de largo y 1 a 1.2 mm de ancho, aplanadas a veces involutas,
generalmente basales, glabras a escasamente pilosas. Inflorescencia homomorfa, compuesta por 4 a 8 rami-
ficaciones primarias espigadas, de 6 a 8 mm de largo, generalmente color verde claro, dimorfas, una forma
compuesta de espiguillas estaminadas y la otra con la flor inferior pistilada o hermafrodita, la flor superior
y Te de n espiguillas laterales estaminadas; puna cars en una s (monomorfas) o
ill a misma infl das de 3 a 4 mm de largo; primera
tin
Fd la2mm di a segunda gluma de 2a 3 mm ds me lema de 3a 4 mm de largo, irregularmente
lobuladas, sin aristas. Espiguillas pistiladas de 4.5 a 5 mm de largo, glumas 3 a 3.5 mm de largo, similares,
vilosas, lema de 4 a 5 mm de largo, glabras, con una hendidura en la parte media hasta 1/3 de su longitud,
con aristas híspidas emergiendo entre los lóbulos y apenas excediéndolos. Flores estaminadas laterales y
superior similares a la pistilada pero d r porte, las aristas glabras y un poco más largas que los lóbulos.
Florece aparentemente de junio a noviembre.
, Lattha Rataniral D h Institute of Texas 2(2)
X S.l
SANA
CN A
Distribución.—Desde el S de Estados Unidos a México en: Chihuahua y Coahuila.
Hábitat. —En matorral desértico micrófilo y bosque de encino; 190 a 1125 m.
Ejemplares examinados. MÉXICO. Coahuila: Municipio de Ocampo, Along the Río Grande, 1 km W of Boquillas de Carmen, 6 Sep
2005, 560 m, P.M. Peterson 18872, J. Valdés-Reyna & C. Sifuentes (US); 33.5 km S de Boquill quiz higl I
Santa Rosa turnoff, 14 Sep 1972, 960 m, F. Chiang 9237, T.L. Wendt & M.C. Johnston (MEXU, TEX); Hills 5 km by winding road S Rancho
El Carrizal Grande, 2 Oct 1972, 1125 m, F. Chiang 9646, T.L. Wendt & M.C. Johnston (MEXU, TEX).
$ Af 1
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y NOL 141 ITOM Ue
8. Bouteloua eriopoda (Torr.) Torr., Pacif. Railr. Rep. 4(5):155. 1857. (Fig. 14). Chondrosum eriopodum Torr., Not.
Milit. Reconn. 153. 1848. Tiro: U.S.A. New Mexico: Las Cruces, Wooton 458 (neotipo: US-A865571 fragm.!, designated by Gould,
Ann. Missouri Bot. Gard. 66:397.1979).
Bouteloua brevifolia Buckley, Proc. Acad. Nat. Sci. Philadelphia 14:93. 1862. Tipo: U.S.A. Texas: May-Oct 1849, C.H. Wright 748 (LectorIPpO:
PH designado por Gould, Ann. Missouri Bot. Gard. 66:397. 1979; isoLecroripo: US-817151).
AT hy
a y
MEA I ov^
MON
Fic. 14 Dierxib az I Da; ioj F} J Ar M
Plantas perennes. Culmos de 25 a 60 cm de alto, tipicamente decumbentes, estoloníferos; entrenudos de los
culmos y estolones lanoso-pubescentes, al menos en parte. Vainas glabras, con márgenes hialinos, más cortos
que los entrenudos; lígula de 0.2 a 0.5 mm de largo, membranácea, ciliada; láminas foliares de 2.5 a 10 cm
de largo y 0.5 a 2 mm de ancho, aplanadas o involutas, con algunos pelos esparcidos principalmente en los
márgenes, de 2.5 a 3 mm de largo, con bases papilosas. Inflorescencia de 5 a 10(-18) cm de largo, con 3 a 8
ramificaciones primarias espigadas persistentes, delgadas, esparcidas; ramificaciones de 2 a 5 cm de largo,
portando 9 a 20 espiguillas; base del raquis de las ramas lanoso-pubescente. Espiguillas de 6 a 7 mm de largo,
barbadas en la base, articulación por arriba de las glumas; glumas desiguales, agudas o acuminadas, primera
gluma de 3 a 4 mm de largo, segunda gluma de 6 a 7 mm de largo, claramente glabra, con el nervio central
prolongado en una arista de 1 mm de largo, emergiendo de entre 2 lóbulos agudos; lema de la flor perfecta
de 6 a 7 mm de largo, barbada en la base, pubescente en el margen, estrechándose hacia el ápice, terminando
" : lafthe Rataniral D h Institute of Texas 2(2)
en una arista fuerte de 1.5 a 3 mm de largo, las aristas laterales muy reducidas; pálea de 6 a 7 mm de largo,
glabra. Anteras de 2 mm de largo, color amarillo; Flósculo rudimentario 1, lema del flósculo rudimentario
de 6 a 7 mm de largo, reducida a 3 aristas aplanadas, de 4 mm de largo, barbadas en la base. Cariopsis de
2.4 a 3 mm de largo, estrechamente elíptica. 2n = 20 (Reeder, 1967). Florece de agosto a octubre.
Distribución.— Centro y SO de Estados Unidos a México en: Coahuila, Chihuahua, Nuevo León, San
Luis Potosí, Sonora, Tamaulipas y Zacatecas.
Hábitat. — Presente en pastizales y matorrales sobre mesetas arenosas secas y sitios rocosos, asociada a
arbustos; 1250 a 1900 m.
Ejemplares examinados. MÉXICO. Coahuila: Municipio de Cuatrociénegas, 19 km al SE de Tanque Nuevo, Cuatrociénegas, H. García s.n.
(CHAPA, ENCB, SLPM); 8 km al N de Cuatrociénegas, camino a Ocampo, J. Marroquín 2672 (ANSM); Municipio de Muzquiz, Sierra del
Carmen, Ejido San Francisco, 5 Sep 2005, 1430 m, P.M. Peterson 18828, J. Valdés-Reyna & C. Sifuentes (US); Municipio de Nava, Sierra El
Jardín, 7.1 mi SW of Rancho La Mula, 4 Sep 2006, 1418 m, P.M. Peterson 19963, S. Lara- diit (US); Municipio de Ocampo, O base of
Picacho del Fuste, NE from Tanque Vaionetta, N facing montain-side, 23 Aug 1941, I.M. J 8420 (MEXU); Rancho prete La
Rueda, 87 km NE de Ocampo, brecha Ocampo-Boquillas del Carmen, Oct 1976, 1280 m, R. Vásquez & J.F. Sifuentes s.n. (ANSM, ENCB,
MER Renee ae La pupa 87 km NE de Ocampo, brecha Ocampo-Boquillas del Carmen, 2 Jul 1991, n m, M. Mei di
s.n. (A la Rueda, 8 Nov 1977, 1320 m, R. Vásquez s.n. (MEXU); Si La Encantada,
8 Sep 1990, 1250 m, M.A. Carranza 730, J. Valdés-Reyna, P.A. Fryxell, R. Vásquez & O. Meza (ANSM); Si La Encantada, rancho Puerto
del Aire, 8 Sep 1990, 1250 m, M.A. Carranza 768, J. Valdés-Reyna, P.A. Fryxell, R. Vásquez & O. Meza (ANSM, MEXU, UAT); Municipio de
Saltillo, 26 km O de Saltillo, carretera Saltillo-General Cepeda, ejido La Noria, 21 Jul 1983, 1780 m, J. Valdés-Reyna 1569, L.E. Rodríguez
& R. Vásquez (ANSM); Municipio de Sierra Mojada, San Antonio de los Alamos, E base of the Volcanic Sierra de San Antonio, 20 DUE
1941, I.M. Johnston 8251, 8257 (MEXU); Sierra de las Cruces, Sierra Mojada, E.G. Marsh 896 (US). Nuevo León: Municipio de Galeana
km 3 al E de Galeana, carr., E. Hernández & V. Mathus 1963 (ANSM). Tamaulipas: Municipio de Hidalgo, Estación B Cruz, 15 Nov
1976, 220 m, G. Villegas 34, A. Bolaños (MEXU).
9. Bouteloua eriostachya (Swallen) unn pios Torrey Bot. Club 94:7. 1967. (Fig. 15). Bouteloua eriopoda var.
eriostachya Swallen, Proc. Biol. Soc. Wash. 56: 943. Chondrosum eriostachyum (Swallen) Clayton, Kew Bull. 37:418. 1982. Tiro:
MÉXICO. Coanutta: cerca de Aguaje de VE 13 Sep 1941, I.M. Johnston 8718 (notorio: US-154692!; isoripos: GH, L
Plantas perennes, cespitosas. Culmos de 40 a 60 cm de alto, erectos, fuertes, con base nudosa; entrenudos
varios, los inferiores densamente lanosos. Lígula de 0.2 a 0.6 mm de largo, un anillo densamente ciliado;
láminas foliares de 4 a 10 mm de largo y 0.5 a 2 mm de ancho, aplanadas, involutas cuando secas, abun-
dantemente híspidas en la superficie adaxial. Inflorescencia de 10 a 18 cm de largo, con 3 a 8 ramificaciones
primarias espigadas esparcidas, persistentes; espigas de 2 a 5 cm de largo, portando 8 a 12 espiguillas; la
rama y el raquis densamente blanco-lanoso principalmente en la base. Espiguillas de 6 a 7 mm de largo,
sobre un estípite largo, articulación por arriba de las glumas; glumas desiguales, primera gluma de 2 a 3
mm de largo, glabra a escabroso-pubescente hacia el ápice, aguda a acuminada, segunda gluma de 6 a 7
mm de largo, vilosa a lanoso-pubescente, con un mucrón de 1 a 2 mm de largo; lema de la flor perfecta 5 a
6 mm de largo, esparcidamente pubescente sobre los nervios y barbada en la base, con 3 aristas, la central
más larga, de 1.5 a 3 mm de largo, las laterales en ocasiones reducidas; pálea de 5 a 6 mm de largo. Flósculo
rudimentario 1, lema del flósculo rudimentario con 3 aristas de 4 a 8 mm de largo, fuertes, escabrosas,
barbadas en la base. Anteras de 2 a 2.2 mm de largo. Cariopsis de 2.3 a 3 mm de largo, color amarillo,
lanceolado-elíptica. 2n = 60 (Reeder, 1967). Florece de septiembre a octubre.
Distribución. —México, endémica de Coahuila.
Hábitat.—En sitios rocosos y secos con matorral espinoso y planicies; 800 a 1200 m.
. MÉXICO: Coahuila. Municipio de Cuatrociénegas, 19 mi SW of Cuatrociénegas, 6 Sep 1965, 1300 m, J.R.
Render & C.G. icd 4522 [^ Carretera Cuatrociénegas - San Pedro de las Colonias, 17 Sep 1996, 800 m, J.T. Columbus 2843, M.
Porter (ANSM po, Sierra de las Cruces, E foothills 8 mi N of Santa Elena Mines, Johnston & Muller 1021 (MEXU);
Municipio de San Pedi, 5.5 5 km al NW de Nueva Delicias, vertiente E de la i de las Delicias, al N de San Pedro de las Colonias, Y.
Herrera Arrieta 1491, M. de la Cerda & O. Rosales (CIIDIR, HUAA); N Si Mojada, San Antonio de los Alamos, E base of the
Sierra San Antonio, Johnston 8257 (MEXU).
10. Bouteloua gracilis (Kunth) Lag. ex Griffiths, Contr. U.S. Natl. Herb. 14:375. 1912. (Fig. 16). Chondrosum
gracile Kunth, Nov. Gen. Sp. (quarto ed.) 1:176, pl. 58. 1816. Actinochloa gracilis (Kunth) Willd. ex Roem. & Schult., Syst. Veg. 2:418.
Harrara Arriat +5) D tal Aal aa da Mw
943
xh
SUA
Por
ox
Fic. 15 Rice ez la D "m AP 1 I ta da M
1817. Eutriana gracilis (Kunth) Trin., Gram. Unifl. Sesquifl. 240. 1824. Atheropogon gracilis (Kunth) Spreng., Syst. Veg. 1:293. 1824.
Tiro: MÉXICO. Guanajuato: La Bufa, Sep, E Humboldt & A. Bonpland s.n. (HoLoriro: B-W1628; isoriro: US-A865679 fragm.!).
AH
pog, lig tach N Gen. N. Amer. Pl. 1:78. 1818. Eutri lig } Nutt.) Kunth, Revis. Gramin. 1:96. 1829. Chondrosum
oligostachyum (Nutt.) Torr., Explor. Red River Louisiana 300. 1853. Bouteloua oligostachya (Nutt.) Torr. ex A. Gray, Manual (ed. 2)
553. 1856. Tiro: U.S.A.: Plains of the upper Missouri, T. Nuttall s.n. (notorio: BM; isoripo: PH!)
Chondrosum gracile var. polystachyum Nees, Linnaea 19(6):692. 1847. Tro: MEXICO. Aschenborn 153 (HoLormPo: ?; isoriro: US-883615
m.!)
Bouteloua stricta Vasey, Bull. Torrey Bot. Club15(2):49. 1888. Bouteloua gracilis var. stricta (Vasey) Hitchc., J. Wash. Acad. Sci. 23(10):454.
Po: U.S.A. Texas: western Texas, 1887, G.C. Nealley s.n. (notorio: US-81710!).
Bouteloua oligostachya var. major Vasey ex L.H. Dewey, Contr. U.S. Natl. Herb. 2(3):531. 1894. Bouteloua gracili
Dewey) Beetle, Phytologia 52(1):11. 1982. Tiro: U.S.A. Arizona: 1882, J.G. Lemmon 427 (BoLoripo: US-883
Bouteloua oligostachya var. pallida Scribn. ex Beal, Grass. N. Amer. 2:418. 1896. Bouteloua gracili pallid ex Beal) B. Boivin,
Naturaliste Canad. 94(4):521. 1967. Tiro: MÉXICO. CHIHUAHUA: 23 Aug 1885, C.G. Pringle 407 (notorio: US-817089).
var. major (Vasey ex L.H.
2010).
1
944 Journal of tl ical | Texas 2(2)
Fic. 16 Mets or Ja D L On I ta da AA
Plantas perennes, amacolladas. Culmos de 20 a 70 cm de alto; nudos 1 a 2 (3), glabros, ligeramente geni-
culados en ocasiones, nunca ramificados arriba de la base; entrenudos glabros. Vainas glabras, más cortas
que los entrenudos; ligula de 0.3 a 0.5 mm de largo, membranácea, ciliada; láminas foliares de 5 a 1530)
cm de largo y 1 a 2.5 mm de ancho, membranáceas. Inflorescencia de 2.5 a 57) cm de largo, con (1) 2 (4)
ramificaciones primarias espigadas persistentes; ramificaciones densas de (1.533 a 4.5 cm de largo y 4 a 7
mm de ancho, lineares, conspicuamente pectinadas, portando 40 a 100 espiguillas, distribuidas hasta el
ápice del raquis; raquis piloso o híspido, aplanado, recto, no arqueado. Espiguillas de 5 a 6.5 mm de largo,
color verde cambiando a púrpura obscuro en la madurez, articulación por arriba de las glumas; glumas
desiguales, primera gluma de 2 a 3 mm de largo, glabra, segunda gluma de 5 a 6.5 mm de largo, apiculada,
no-mucronada, usualmente esparcido pilosa o híspida, a menudo piloso-papilosa, pelos de 2 a 3 mm de largo,
en ocasiones no-pilosa pero con pubescencia corta y densa en los extremos, ocasionalmente con tubérclos
purpúreos en la madurez; lema de la flor perfecta de 4 a 5.5 mm de largo, con 3 aristas de 1 a 3 mm de
La vo Arriat: tal.,B tal eal ta fa Mv: 945
largo, barbada en la base y pilosa en las nervaduras, al menos la central, los pelos de 0.5 a 0.8 mm de largo;
pálea de 4 a 5.5 mm de largo, glabra. Flósculos rudimentarios 1 a 2, lema de la primera de 3.5 a 5.5 mm de
largo incluyendo las aristas, 3-aristada, con un mechón de pelos en la base del estípite; el segundo flósculo
cuando presente en forma de escama, de 2.5 a 3 mm de largo y ca.0.5 mm de ancho. Anteras de 2.2 a 2.5
mm de largo, color amarillo. Cariopsis de 2 a 2.2 mm de largo, color amarillo a amarillo-verdosa, angulosa
triangular. 2n = 20, 40, 60. Florece de julio a noviembre.
Distribución. —Ampliamente distribuída en Canadá, Estados Unidos y México.
Hábitat. —Principal elemento de los pastizales naturales de SW de EUA y NW de México. Se le reconoce
su excelente calidad forrajera; 1100 a 2600 m.
Ejemplares examinados. MÉXICO. es Ubu M de aa 2 du N of El E on = winding road to La Huerfanita, 13 Sep
1972, 1100 m, F. Chiang 9191, T.L. Wendt & M.C. Jol Aug 1936, E.G. Marsh Jr. 570, 693 (MEXU);
Municipio de Arteaga, Al N de la cañada el bd aprox. 2 a NO del Ejido Sierra Hermosa, 19 Oct 1979, 2270 m, R. López s.n.
(ANSM); Predio El Cristal, 21 Jun 1994, 2100 m, J.A. García 53 (MEXU); papse de Cuatrociénegas, Cañón de la Carretera, Sierra de
la Madera, along the road up the open main canyon, 16 Sep 1941, 12 LM. Sid 9159 gine ed Me a Francisco I.
Madero, Valle de Buenavista, a broad high montane valley enclosed by the Si cati
ta, to pe S de las Margaritas (to E); NO of Valle del Sobaco, 30 Aug 1975, 1575 m, m Wendt 1387- " E.J. Lott (MEXU); Municipio
de General Cepeda, km13 al W de Saltillo, E. Hernandez & V. Mathus N-1665 as E. He & V. dn: ie (ANSM); 20 km O de
Saltillo, carretera Saltillo-Torreón, 6 Nov 1972, 1400 m, R. Almeida 1639 (ANSM), y 40 toward Torreón,
25 Aug 1983, 1700 m, S.L. Hatch 5037, J. Valdés-Reyna (ANSM); Municipio de ME 99.6 mi N of Melchor Muzquiz at Cuesta de
malena on Hwy 25 towards Bequillas del Carmen, 01 Sep 2006, 1574 m, P.M. Peterson 19829, S. Lara- Contreras (US); 20 mi NW de
Hacienda la Baba, Múzquiz, F. Wynd & C.H. Mueller 435 (MEXU, US); La Cuesta del Plomo on the Múzquiz-Boquillas highway, 14 Sep
1972, 1750 m, F. Chiang 9203, T.L. Wendt & M.C. Johnston (MEXU); Sierra del Carmen, Ejido San Francisco, 5 Sep 2005, 1430 m, P.M.
Peterson 18837, J. Valdés-Reyna & C. Sifuentes (US); Sierra del Carmén, Campo 1, 12 Aug 1975, 1800 m, M.F. Robert 5311,5553, J. Passini
(ANSM); Sierra Maderas del Carmen, 17 Sep 1989, 1800 m, E.A. Estrada 1837 (ANSM); Madera del Carmen, 11.2 mi NE of Los Pilares,
21 Sep 2007, 1953 m, P. M. Peterson 20950, J.M. Saarela, S. Lara- Contreras & J. Reyna (US); Madera del Carmen, 16.3 mi NE of Los Pilares
on road towards Campo Dos, 21 Sep 2007, 2235 m, P.M. Peterson 20993, J.M. Saarela, S. Lara- Contreras & id Reyna (US); Ven ils de
Nava, Sierra El Jardin, Canyon del pini 8 mi from jet of road towards Boquillas, 2 Sep 2006, 1336 m, P.M. Peterson 19865, S. 1
Contreras (US), Si El Jardín, sl del Diablo, 2 Sep 2006, 1418 m, P.M. Peterson 19883, S. Lara-Contreras (US); Sierra
El Jardín, 4 Sep 2006, 1480 m, PM. Peterson 19945, S. Lara-Contreras (US); Sierra El Jardín, 8 mi SW of Rancho La Mula, 4 Sep 2006,
1394 m, P.M. Peterson 19958, S. Lara- Contreras (US); Municipio de Ocampo, Madera del Carmen, 9.7 mi NE 2 Los e: 21 Sep 2007,
1902 m, P.M. Peterson 20936, J.M. Saarela, S. Lara- Contreras — Reyna (US); 2 km O of Santa Elena, foothill las Cruces,
18 Jul 1941, R.M. Stewart 831 (MEXUJ; Sierra El Pino, 9.2 km SW of Rancho El Cimarron along the easter slopes, 14 Sep 1991, 1500 m,
P.M. Peterson 10626, & C.R. Annable (US); Sierra El Pino, 33.5 kmW of Rancho El Cimarron, 14 Sep 1991, 1500 m, P.M. Peterson 10691,
& C.R. Annable (US); Base of hills in palma-belt, near mouth of Cañón de la Cruz, about 20 km S of Ocampo, 16 Sep 1941, I.M. Johnston
9177 (MEXU); Rancho La Rueda, 92 km al NW de Villa Ocampo, M. Vázquez s.n. (SLPM); Between Palos Blancos and San Pedro, road
from Ocampo over the mountains to Puertecito (via cuesta de Zozaya), 19 Sep 1941, LM. or 9274 (MEXU); Mesa Grande, high
mesas 40 km NO of Hacienda de La Encantada, 14 Sep 1941, R.M. Stewart 1632 ( e cacho del Fuste, NE from Tanque
Vaionetta, N facing montain-side, 23 Aug 1941, I.M. Johnston 8424 (MEXU); Ran imental La Rueda, 87 km NE de Ocampo,
brecha Ocampo-Boquillas del Carmen, Sep 1976, 1530 m, R. Vásquez s.n. (ANSM); Rancho experimental La Rueda, 87 km NE de Ocam-
po, brecha Ocampo-Boquillas del Carmen, 1 Jul 1991, 1350 m, M. Vásquez s.n. (ANSM); Rancho Santa Teresa de la Rueda, Sep 1976,
1530 m, R. Vásquez (MEXU); Municipio de Parras, Cañón de Gustrola, 16 km al SE de Parras de la Fuente, 30 Ene 1981, 2000 m, A.
Rodríguez 185 (ANSM); J. Valdés-Reyna s.n. (SLPM); A. Orta & M. Carranza 13304 (UAG); Rancho el Tunal, 25 km ESE de Parras de la
Fuente, 5 Jul 1983, 2050 m, A. ls 1179, MA. peat ee M de Parras, Sierra de Parras en el arroyo La Tinaja, 13
Sep 1980, 1850 m, A. Rodríguez 254, M. N Arizpe, La Sauceda, 5 Aug 1976, 1030 m, J.S. Marroquín
s.n. (ANSM); San Juan de las Bonitas, 19 Jul 1993, 1450 m, E Alcalá 15 (MEXU); S e los Landeros, 19 Jul 1993, 1450 m, E. Pérez
21 (MEXU); Sierra de la Paila (Lado Norte) Cañada Becerros, 13 Oct 1989, 1700 ne Villarreal 5433, M.A. Carranza & L. Arce (ANSM);
Ejido el Cedral por el camino El Carmen, 4 Oct 1989, 1300 m, J.A. Villarreal 5332, M.A. Carranza & A. Rodríguez (ANSM); eae de
Saltillo, On highway 54 (road to Concepción del Oro), 6 km S de Saltillo, reforestation area Zapalinamé, near Universidad Auton
Agraria Antonio Narro, 18 Sep 1988, 1800 m, M.E. Barkworth 5112, P.S. Hoge, J. Valdés-Reyna-Reyna & M. A. Carranza P. (ANSM); 29 n
S of Saltillo on Mex 54 and 2.1 km W on road to Microondas, just W of Estacion; 18 Oct 1989, 2250 m, P.M. Peterson 8414, M.A. Carran-
za (US); La Angostura, Saltillo. M. Tórres 1589 (ANSM); 29 km al S de Saltillo, carr. Saltillo-Zacatecas, R. Almeida 1630 (ANSM); Sierra
Zapalinamé, Loma de Lourdes, R. Palomo s.n. (ANSM); Area del Rastro, 7 km S de Saltillo, 3 Aug 1973, 1800 Banda 2599 (ANSM);
Saltillo, 0.3 km E of Hwy t d to San Lorenzo, 10 Sep 1990, 1720 m, P.M. Peterson 10553, C.R. irem (US); Buena-
vista, 7 km S de Saltillo, carretera 54 Saltillo- Rui del Oro, Zac., 23 Nov 1975, 1600 m, A. Aznar s.n. (ANSM); 20 May 1986, 1785
m, E. García s.n. (ANSM, ENCB, IBUG, IEB); 8 Aug 1974, 1800 m, A. Rodríguez 131 SM Cañas e en la Sierra Zapalina-
mé, 3 km NO de Cuauhtémoc, 28 Sep 1979, 2 m, R. López s.n. (ANSM); I é, SE de Saltillo,
946 I Lafeha Dataniral D h Institut f Texas 2(2)
26 Aug 1986, 1800 m, J. Espinosa 28 (ANSM); La Encantada, 19 Sep 1991, 1860 m, Bri, huila 217, 220, 221, 231, 234, 235 (MEXU);
Paso de adu alS lu Saltillo, 14 Nov 1974, 1100 m, J.S. Marroquín 3059 (ANSM); Rancho experimental Los Angeles, 48 km S de
Saltillo, Itillo-Co ción del Oro, Zac., 15 Jul 1981, 1800 m, M.A. Carranza, M. T. Ruíz de León & M. Vásquez R. (ANSM);
25 Jul 1972, 1800 m, P.E. García s.n. E 26 Aug 1981, 1900 m, S.L. Hatch 4536, C.W. Morden & J. Valdés-Reyna (ANSM); 1 Jul 1974,
1725 m, R.E. Rodríguez 108 (ANSM); 1 Sep 1978, 2000 m, J.S. Sierra s.n. (ANSM); Jul 1976, 2200 m, R. Vásquez s.n. (ANSM); 14 Sep 1992,
J.A. Villarreal 6716, M.A. Carranza & D.E. Lozano (ANSM); Rastro de Saltillo, 7 km S de Saltillo, 7 Jul 1972, 1780 m, R. Almeida 1383
ae 5 mi W of E ad on ud road between Hwy 54 & 57, 23 mi S of Saltillo, 16 ct 1995, 2410 m, P.M. Peterson 13270, M.B.
apalina Lomas de Lourdes, 9 Sep 1975, 1770 m, R. Palomo s.n. (ANSM). Nuevo León: Municipio de Aram-
berri, 16 km NE of Sandi d to La Ascension, 19 Sep 2001, 2000 m, P.M. Peterson 15828, J. Valdés-Reyna (US); 7.5 ln E of Puentes
on dirt road, 19 Sep 2003, 1775 m, P.M. Peterson 17849, J. Valdés-Reyna & H.S. Hinton (US); Municipio de Bustamante, 10 km al NE de
la Verdolaga, 5 Jul 1985, 1800 m, P. Hiriart 849, R. Molczadzki & V. Juárez (MEXU); Municipio de Dr. Arroyo, 9.4 mi W of San Antonio
de Peña Nevada & 0.4 mi E of jet of Hwy 2t 22 Sep 2002, 1807 m, P.M. Peterson 16796, J. Valdés-Reyna & M. Sosa (US); 30 km ENE
of Dr. Arroyo, ENE of San Antonio de Pena M O paves of ndi Peña Nevada, 30 Jul 1989, 1950 m, G. Lane 4731 (ANSM); Doctor
Arroyo, Nov 1980, 1700 m, J.A. Ochoa 1189 (MEXU); Municip Gal , 12.3 mi S of Milagro, 18 Sep 2003, 2040 m, P.M. Peterson
17808, 17811, J. Valdés-Reyna & H.S. Hinton (US); 5.1 mi S of Milagro, 18 Sep 2003, 2000 m, P.M. Peterson 17818, J. Valdés-Reyna & H.S.
Hinton (US); Arroyo Hondo, Hacienda San José de Raices, 1 Aug 1935, C.H. Mueller 2310 (MEXU); 6.5 mi S of border of Coahuila and
Nuevo León on Hwy 57 towards Matehuala, 16 Oct 1995, 1920 m, P.M. Peterson 13288, M.B. Knowles (US); El Salero, 19 Sep 2001, 1840
m, P.M. Peterson 158823, J. Valdés-Reyna (US); Hacienda Pablillo, 5 Aug 1936, M. Taylor 56 (MEXU); Navidad, carretera 57, km 85 al N
de Saltillo, 23 Sep 1978, 1650 m, F. Cárdenas, M.A. Bernal pato. Rancho e 17 Oct 1989, 1950 m, G.B. Hinton 19861 (MEXU);
Valle de Santa Rita, Galeana, I. Siller 10858 (UAG). Tamaulipas: Munici , Camino a la Joya de San Francisco, 1950 m,
A. Brito (MEXU); Municipio de Jaumave, 1 km N del ejido San ern 10 Nov 1989, 680 m, A. Mora 1320 (UAT); Municipio de Mi-
quihuana, 4 kmN del ejido estanque de los Walle, 29 May 1986, 1560 m, L. Herndndez 1841 (UAT), 3 Sep 1981, 2460 m, R.A. Carranco
78 (MEXU); 4 kmO of Miquihuana, 4 Aug 1941, 2600 m, L.R. Stanford 647, K.L. Retherford & R.D. Northcraft (MEXU); 6 km al O de
Miquihuana, 6 Jul 1985, 2130 m, P. Hiriart 873, V. Juárez, R. Molczadzki, J.L. López, F. Ojeda & J.A. Gutierrez (MEXU); 9 km al O de Mi-
quihuana, 20 Sep 1984, 1560 m, P. Hiriart 338, V. Juárez (UAT); El Cargadero, 15 km O del estanque de los Walle, 25 Oct 1986, 1560
m, L. Hernández 2055 (UAT); La Peña, 8 Dic 1988, 2000 m, M. Cisneros 198 (MEXU); Rumbo al Aserradero, 4 Sep 1981, 1900 m, R.A.
Carranco 72 (MEXU); San José del Llano, 29 Nov 1984, J.F. Iribe 231 (MEXU); Valle Hermoso, 13 Aug 1977, 2000 m, G. Villegas 84, M.A.
Bolaños (MEXU); Municipio de Palmillas, 13.5 km al NO de Palmillas, 7 Jul 1985, 1600 m, P. Hiriart 904, V. Juárez & R. Molczadzki
(MEXU); 101.6 kmSWof Ciudad Victoria on Mex 101 towards San Luis Potosí, 9 ct 1989, 1720 m, P.M. Peterson 8314 & R.M. King (US);
Municipio de Tula, 30 km al N de Tula, 25 Aug 1983, I. Nuñez 30 (MEXU); 5 km al O del Ejido El Salitrillo, 3 Jul 1983, 1500 m, P. Hiriart
761, V. Juárez, R. Molczadzhi, J.L. López, F. Ojeda & J.A. Gutterrez (MEXU); La Tapona, 21 Oct 1982, 1900 m, M.H. Cervera 134, 134-B
(MEXU)
11. poe ewe E Varied. Ci. 2(4):141. 1805. Actinochloa hi (Lag.) Roem. & Schult., Syst. Veg. 2:419. 1817.
et. Hort. Brit. 1:455. 1826. Tro: MÉXICO: culti lin? 1 Lagasca s.n (LecrotipO: MA-
[2]
245716 desi 1 Griffiths, Contr. U.S. Natl. Herb. 14:372, 1912; 1isoLecroripos: BM, M-7433 LE-TRIN 2125.6 fragm. ex M!)
O I
Planta perenne, de vida corta o larga, ocasionalmente estoloníferas. Culmos de 15 a 75 cm de alto, mod-
eradamente ramificados o sin ramificar, con 3 a 6 nudos glabros a densamente híspidos con pelos de base
papilosa, erectos o decumbentes en la base. Hojas basales y caulinares; vainas glabras a papiloso-hirsutas,
más cortas que los entrenudos; ligula corto ciliada de 0.1 a 0.5 mm de largo; láminas foliares de 5 a 1830)
cm de largo y 1 a 2 mm de ancho, aplanadas o subinvolutas, casi glabras a hirsuto-pubescentes en la su-
perficie adaxial, con pelos papilosos en el margen y parte basal de la hoja. Inflorescencia de 10 a 40 cm de
largo; ramificaciones es epee E a 5, de 1.5 a 4 cm de largo y 6 a 9 mm de ancho, pectinadas,
persistentes, portando (10—)20 a 50 espig , subsésiles, con el raquis conspicuamente aplanado, proyec-
tándose 5 a 10 mm más allá de la inserción de la espiguilla distal, terminando en una aguja semirígida.
Espiguillas de 4 a 6.5(—7) mm de largo, color verde o púrpura oscuro, articulación por arriba de las glumas;
glumas desiguales, lanceoladas, terminando en una arista de 1 a 3 mm de largo, primera gluma de 1.5 a
2.5(3) mm de largo, angosta, hispida a lo largo de la nervadura media, segunda gluma de 3 a 6(-6.5) mm
de largo, papiloso-hirsuta a lo largo de la nervadura media, las papilas negras, los pelos hasta de 2 mm de
largo; lema de la flor perfecta de 4 a 6 mm de largo, 3-dentada, cada diente termina en una pequeña arista
de 1 mm de largo, conspicuamente pilosa a lo largo de los márgenes y de la nervadura central; pálea de
3.5 a 4.2 mm de largo, ovada, obtusa, pubescente, sin nervaduras visibles. Anteras de 2 a 2.5(-3) mm de
largo, color amarillo a crema. Flósculos rudimentarios 2 a 3, el primero con la lema de 2 a 5 mm de largo,
2-lobulada y 3-aristada, aristas híspidas, de 3 a 4—6) mm de largo, los otros flósculos reducidos a brácteas
Hu Ayviat tal., B tal dal tada Mw: 947
diminutas, a veces hasta de 0.5 mm de largo; raquilla de 1 a 2 mm de ancho. Cariopsis de 0.7 a 2 mm de
largo, color amarillo, anguloso-triangular. 2n = 20, 60.
Bouteloua hirsuta Lag. var. hirsuta (Fig. 17). Chondrosum hirtum Kunth, Nov. Gen. Sp. (quarto ed.) 1:176, t. 59. 1816.
Atheropogon hirtus (Kunth) Spreng., Syst. Veg. 1:293. 1825. Bouteloua hirta (Kunth) Scribn., Contr. U.S. Natl. Herb. 2(3):531. 1894.
Tiro: MÉXICO: E Humboldt & A. Bonpland s.n. (notorio: P; isorro: B-W-1626, M, LE-TRIN-2126.1!, US-A865681 fragm.!).
Ho n Engelm., Amer. J. Sci. 46:104. 1843. Bouteloua pappilosa (Engelm.) Torr., Explor. Red River Louisiana 300. 1853.
um papillosum (Engelm.) Torr., Explor. Red River Louisiana 300. 1853. Tiro: U.S.A. liuwors. Cass Co.: 1842, Geyer s.n.
pe MOD.
Chondrosum aschenbornianum Nees, Linnaea 19(6):692. 1847. Bouteloua a (Nees) Griseb. ex E. Fourn., Mexic. Pl. 2:137.
1886. Tiro: MÉXICO: A. Aschenborn 331 (HoLoriro: B: sorro: US-865680 fragm
Chondrosium foeneum Torr., Not. Milit. Reconn.153, pl. 12. 1848. Bouteloua foenea (Torr) Torr. ex S. Watson, Cat. Pl. 18. 1874. Tiro:
. New Mexico: Cooks Springs, 9 Oct 1846, WH. Emory s.n.(Lectoripo, designado aqui: NY-73939!
Chond lii E. Fourn. Mexic. Pl. 2:137. 1886. Tiro: U.S.A. Texas: Drummond 323 (LECTOTIPO, etude aquí: P!; ISOLECTOTIPOS:
LE, US-A865677 fragm. ex P!, US-2182917 fragm.!, US-1063203 fragm. ex E!
Bouteloua hirsuta var. palmeri Vasey ex Beal, Grass. N. Amer. 2:417. 1896. Tipo: MÉXICO: cultivated from seed, 1886, E. Palmer s.n.
(uoLoriPo: US!).
Culmos 15 a 40 cm de alto, Hone en la base, ] ificad iba de la base, con 4 a 6 nudos;
Hai
glabras o papiloso-hirsutas, pilosas en la garganta;
láminas foliares de5a 186 cm de largo y 1a2 mm de ancho, aplanadas a subinvolutas. Inflorescencias de 10
a 30 cm de largo; ramificaciones primarias espigadas 1 a 4, de 1.5 a 3 cm de largo, persistentes, portando
20 a 40 espiguillas, subsésiles; con el raquis aplanado, proyectándose hacia adelante de la espiga terminal.
Anteras de 2 a 2.5 mm de largo. Cariopsis de 0.7 a 1.52) mm de largo. Colectada con flor todo el año.
Distribución. —Ampliamente distribuida en Estados Unidos y México.
Hábitat.—Se le encuentra en áreas planas con pastizales abiertos y secos, con matorrales y/o bosque de
encino y en suelos bien drenados, rocosos; 50 a 2300 m.
Ejemplares examinados. MEXICO. Coahuila: Municipio de Acuña, Del Carmen mountains, 9 Ene 1936, E.G. Marsh Jr. 887 (MEXU);
Serranías del Burro, rancho El Bonito, cañón Los Ojitos, 19 Sep 1977, 1600 m, D.H. Riskind 2181 (ANSM); Municipio de Allende, 135.4
km NW of Muzquiz on Hwy 53 towards Boquilla del Carmen, 12 Sep 1991, 1380 m, P.M. Peterson 10578, C.R. Annable (US); Municipio
de Arteaga, Al N de la cañada el Columpio, aprox. 2 km NO del Ejido Sierra Hermosa, 19 Oct 1979, 2260 m, R. López s.n. (ANSM); S.L.
Hatch, C.W. Morden & J. Valdés- Sd 4535 ds La o carretera ana MAE Sierra de Arteaga, 7 Oct 1973, 2020 m,
R. García s.n. (ANSM); Sierra Herm del Carbón, 8 Aug 1979, 2300 m, R. López
s.n. (ANSM); inc E de Castaños, La Muralla, Si de la Gavia, carretera 57, 7 Oct 1989, 1400 m, A. Rodríguez 1301, M. Martínez &
J.M. Sosa (ANSM); 15 mi S of Castaños, J.R. Reeder & C.G. Reeeder 3951 (US); Municipio de Cuatrociénegas, Sierra de la Madera, vicinity
of ‘La Cueva’ in Corte Blanco fork of Charretera Canyon, 11 Sep 1941, 1615 m, I.M. Johnston 9058 (MEXU); Sierra Las Cautivas, 23.4
mi W of Ejido Guayabas along road towards Dulces Nombres, 25 2 2007, Vide m, P.M. Peterson 21063, J.M. Saarela ud Municipio
de Monclova, Sierra de la Gloria, in Cañón del Chilpitín , near El Chilpitín T
Sep 1976, 1060 m, T.L. Wendt 1685, D.H. Rishind MEXU) Municipio d de Múzquiz, Seia del Carmen O m Francisco: 5 Sep 2005,
, P.M. Peterson 18836, J. Valdés-Reyna & C ] Carmen, fl N 17 Sep 1992,
1200 m, J.A. Villarreal 6927, M.A. Carranza & R. Vae (ANSM); Madera del carmen, 7.2 mi NW of Pilares, 7 Sep 2005, 1700 m, P.M.
Peterson 18875, J. Valdés-Reyna (US); Sierra La Encantada, 140 km al N de Múzquiz, entronque Boquillas del Carmén brecha y túnel,
Flourita de México Unidad Minera, 6 km al SO pasando el túnel, 7 Sep 1990, 1500 m, M.A. Carranza 709, J. Valdés-Reyna, P.A. Fryxell
& R. Vásquez (ANSM); Municipio de Nava, Sierra El Jardín, Canyon del Diablo, 9 mi from jct of road towards Boquillas, 2 Sep 2006,
1418 m, P.M. Peterson 19880, S. Lara-Contreras (US); Si El Jardín, slopes above Canyon del Diablo, 2 Sep 2006, 1418 m, P.M. Peterson
19907, S. Lara-Contreras (US); Sierra El Jardín, 8 mi SW of Rancho La Mula, 4 Sep 2006, 1394 m, PM Beier DR 19959. S. Lara- Contreras
(US); Municipio de Ocampo, O base of Picacho del Fuste, NE from Tanque Vaionetta, N facing montain-side, 23 Aug 1941, I.M. Johnston
8425 (MEXU); Ocampo, 1 Aug 1972, 1260 m, R. Vasquez, Noriega (ANSM); Sierra El Pino, 33.5 km W of Rancho El Cimarron, 14 Sep
1991, 1500 m, P.M. Peterson 10652, & C.R. Annable (US); Sierra El Pino, 39.5 km W of Rancho El Cimarron, 14 Sep 1991, 1500 m, P.M.
Peterson 106 78, & C.R. Annable ee Madera ps Carmen, 9.7 mi NE of Los aa 21 Sep 2007, 1902 m, P.M. Peterson 20935, J.M. Saarela,
Lara-Contreras & J. Reyna (US); N p , Sierra de P de Gustrola, 1 Sep 1988, 2050 m, A, Rodríguez 566,
P. Antonio (ANSM); Municipio de Ramos Arizpe, Sierra de la Paila (Lado Norte) canada Becerros, 13 Oct 1989, 1700 m, J.A. Villarreal
5450, M.A. Carranza & L. Arce (ANSM); Sierra de la Paila, Ejido el Cedral ] El Carmen, 4 Oct 1989, 1300 m, J.A. Villarreal
5336, M.A. Carranza & A. Rodríguez (ANSM); Municipio de Sabinas, 85.4 km NW of ue on Hwy 53 towards Boquilla del Carmen,
vía La Rosita, 12 Sep 1991, 810 m, P.M. Peterson 10569, C.R. Annable (US); Municipi Saltillo, Hacienda Buena Vista al SE de la Uni-
versidad Autónoma Agraria Antonio Narro, 11 Sep 1981, 1600 m, G. Villaseñor s s.n. (MEXUJ; on carretera 54, 6 km S de Saltillo, area
1 laf tha Botanical D h Institute of Texas 2(2)
948
Fic. 17. Distribución de B
de Reforestación de Zapalinamé, near Universidad Autónoma Agraria Antonio Narro, 18 Sep 1988, 1900 m, M.E. Barkworth 5110, P.S.
Hoge, J. Valdés-Reyna & M.A. Carranza (ANSM); Rancho experimental Los Angeles, 48 km S de Saltillo, carretera 54 Saltillo- Concepción
del Oro, Zac., 26 Aug 1981, 1800 m, S.L. Hatch 4535, C.W. Morden & J. Valdés-Reyna (ANSM); 18 Oct 1989, 2140 m, P.M. Peterson 8428,
M.A. Carranza (US); 2 Sep 1978, J.S. Sierra s.n. (ANSM, MEXU); 14 Sep 1992, J.A. Villarreal 6714, M.A. Carranza & D.E. Lozano (ANSM);
Municipio de San Buenaventura, Sierra La Encantada, rancho Carrizalejo, entrada S al rancho Puerto del Aire, 8 Sep 1990, 1200 m,
M.A. Carranza, J. Valdés-Reyna, P.A. Fryxell, R. Vásquez & O. Meza (ANSM). Nuevo León: Municipio de Cadereyta Jiménez, 4 km NE de
Cadereyta Jiménez, 20 Abr 1973, 330 m, J.S. M quin 2502 (ANSM); en los alreded del poblado de Cadereyta, 22 Mar 1993, 320 m,
N. Bazaldua 35 (MEXU); Municipio de Doctor Arroyo, 21 km al N de Matehuala, San Luis Potosí, 19 Sep 1984, 1800 m, P. Hiriart 315, V.
Juárez & R. Molczadzki (UAT); 70 mi N of Matehual Hwy 61 towards Linares and 1.5 mi N of La Rosita, 19 Oct 1995, 1940 m, P.M.
Peterson 13361, M.B. Knowles (US); 9.4 mi W of San Antonio de Peña Nevada & 0.4 mi E of jet of Hwy 2 to Or., 22 Sep 2002, 1807 m,
P.M. Peterson 16795, J. Valdés-Reyna & M. Sosa (US); Municipio de Galeana, 2 mi W of Dieciocho de M p ] erro Potosí,
18 Oct 1995, 2130 m, P.M. Peterson 13324, M.B. Knowles (US); 25 km E of San Roberto along highway 58, 27 Aug 19
Hatch 4575, C.W. Morden & J. Valdés-Reyna (ANSM); km 636 carretera Galeana-San Roberto, I. Cabral 1087 (IEB), Municiy
Zaragoza, Camino a Tinajas, 5 Nov 1982, 1530 m, A. Brito 107 (MEXU); Municipio de Iturbide, 1 km NE de Cadereyta, 14 Oct 1971,
81, 1740 m, S.L.
imio de G 1
Li avy, A Bt eN l., B tal dal To PU Y PA 949
320 m, J. Valdés-Reyna 3 (ANSM); Bosque-escuela, Universidad Autónoma de Nuevo E 12 km E is aba 18 Oct 1987, 1500 m, I.
ds 1093 (ANSM); Ejido Santa Rosa, 29 Aug 1989, 1550 m, E.A. Estrada 1677 (ANS! , Monterrey, T. Tateoka
US). Tamaulipas: Municipio de Aldama, Playa Rancho Nuevo, 18 Aug 1985, 60 m, R. Diaz 28 (ANSM, UAT); Santa Rosa, 29 Aug
"n J.L. Ramos s.n. (MEXU); Municipio de Altamira, Rancho La Fortuna, 9 Oct 1980, 50 m, R. Sandoval 661 (MEXU); Municipio de
Burgos, Rancho La Granja, 18 Oct 1983, 200 m, J.F. Iribe 206 (MEXU); Municipio - Bustamante, Cerro Tinaja. Vicinity of San José,
14 Jul 1930, 731 m, H.H. Bartlett 10342 (MEXU); Municipio de Gómez, km 26, Rancho el Soldado, 28 P 1993, 150 m, O.M. Galván
603 (MEXU); Municipio de Hidalgo, Ejido Nicolás Bravo, 29 Oct 1989, J. Cantú 40 Re EXU); Municipi Mai , km 5 hacia Fl
Ejido Camarones, 25 Jul 1986, 750 m, M.H. Cervera 352 (MEXU); La Oveja (Venustiano Carranza), 19 Nov 1986, ii m, M.H. Cervera
398 (MEXU); Municipio de Matamoros, Dunas del Mezquite, 11 Jul 1981, J. Cantü 311 (MEXU); Muni El Angelito,
Colonia La Peña, 24 Nov 1981, 660 m, J.F. Iribe s.n. (MEXU); Municipio de Palmillas, 12 km al NO Palmillas, m 1976, 1700 m, F.
González 10013, P. Zavaleta eund Municipic de San Carlos, Piedra Iman, 2 km ESE de San José, Sierra de San Carlos, 22 Nov 1984,
m, O.L. Briones 1213 (ANSM); M de San Nicolás, Rancho La Trinidad, Oct 1983, 650 m, J.F. Iribe 201 (MEXU); Municipio
Soto la Marina, Ejido 5 de i 18 Sep 1981, 140 m, J.G. Galván 7 (MEXU); Ejido Benito Juárez, 3 Oct 1986, J.F. Iribe 323 (MEXU);
Ejido Los Arroyos, 22 Oct 1983, A. Brito 153 en Rancho San Isidro, 22 Oct 1986, 30 m F. dd d CERU oe de Tula,
22 km NO de Tula, límite entre | pios de Tula y Bustamante, 9 Feb 1984, 1400 m (UAT); 5-6 km al S del
Capultn, 10 Aug 1972, 1800 m, F. González 4454, R.M. López & R. Dirzo (MEXU); Municipio ^ Villagrán, 3 mi S of N I line on
the Villagrán-Linares highway, 12 Nov 1959, 400 m, J. Graham 4665, M.C. Johnston (MEXU, UAT); 1 mi al E de Morales y 5 mi al W de
Loreto, M.C. Johnston 5545-B (US).
12. Bouteloua johnstonii Swallen, Proc. Biol. Soc. Wash. 56:79. 1943. (Fig. 18). Tiro: MÉXICO. Coanuma: S de
anada Oscuro, near Tanque La Luz, 27 Sep 1942, I.M. Johnston 8491 (HoLoripo: NA-154624; isoripos: GH, US-18177901).
Plantas perennes, amacolladas, macollos de 10 a 20 cm de diámetro. Culmos de 10 a 40 cm de alto, den-
samente ramificados, de base dura, rizomatosa. Vainas glabras, con margenes hialinos; lígula un diminuto
anillo de pelos de 0.5 mm de largo; láminas foliares de 5 a 12 cm de largo y 1 a 2 mm de ancho, rígidas,
estrechas, curveadas en la madurez, involutas en toda su largo, con el ápice rígido como espina, abundante
lanoso-pubescente en la superficie adaxial, y en ambas superficies hacia la base, los pelos largos hasta de 4
mm de largo, cambiando a glabras con la madurez. Inflorescencia largamente exserta; ramificaciones pri-
marias espigadas 6 a 12, de 8 a 15 mm de largo y 5 a 10 mm de ancho, erectas o ascendentes, caedizas en la
madurez, portando 7 a 11 espiguillas; raquis aplanado, ligeramente escabroso, algo sinuoso. Espiguillas de
5 a 7 mm de largo, permanecen insertas a las ramificaciones; glumas desiguales, color párpura, delgadas,
agudas a acuminadas o cortamente aristadas, primera gluma de 3.5 a 4 mm de largo, angosta, largamente
acuminada, ligeramente escabrosa o corto pubescente en la mitad inferior, segunda gluma de 5 a 7 mm de
largo, esparcidamente hirsuta, con un mucrón corto; lema de la flor perfecta de 5 a 6 mm de largo, glabra con
algunos pelos adpresos hacia la base, redondeada en el dorso, los nervios se prolongan en aristas fuertes, las
aristas laterales cortas, escabrosas, la central más larga, de 2 a 3 mm de largo, emergiéndo entre dos lóbulos
membranáceos; pálea de 4 a 4.2 mm de largo, glabra. Anteras de 2 a 2.2 mm de largo, color amarillo claro,
con los filamentos pubescentes. Flósculo rudimentario 1 (2), el primero de 6 a 7 mm de largo, formado por
una columna glabra con 3 aristas aplanadas, fuertes, escabrosas, alternas con 4 aristas pequeñas, de 2.5 a
3 mm de largo, muy delgadas, escabrosas; segundo flósculo cuando presente muy reducido. Cariopsis de 2
a 2.2 mm de largo, color amarillo, oblongo-elipsoide. Florece de agosto a octubre.
Distribución. —Especie endémica mexicana conocida de Coahuila y Zacatecas.
Hábitat.—Planta escasa, presente en suelos yesosos; 1200 a 1600 m.
LU dp MEC Coala; Municipio de Francisco I. Madero, S part of Si de los O h 9.5 km E of
Pi Aug 1973, 1200 m, M.C. Johnston 12114 F. Chiang, T.L. Wendt &J. Henrickson (ANSM);
Municipio de G peda, Sierra de la Paila al O en el Ejido Nueva Imagen, 5 Oct 1985, 1400 m, A. id Ded iade E
Rancho. Parritas, W de Coahuila, base de la montafia, margen E del Valle de Acatita, R.M. S p
Sierra de la Paila, Valle de Parreños, 31 Oct 1987, 1600 m, J.A. Villarreal 4173, J.L. Elizondo & JJ. López (ANSM, CIIDIB.
13. oce karwinskii (E. Fourn.) Griffiths, Contr. U.S. Natl. Herb. 14(3):394, f. 47, t. 76. 1912. (Fig.
. Chondrosium karwinskii E. Fourn., Mexic. Pl. 2:137. 1886. Tiro: MÉXICO: Cañón de las Minas et Victoria, Karwinsky 1479
dins P; isoripos: LE, US-A865682 fragm.!).
Plantas perennes. Culmos de 20 a 55 cm de alto, delgados, erectos, ramificados desde una base dura y
fuertemente rizomatosa; entrenudos glabros. Vainas más cortas que los entrenudos, glabras, ciliadas solo
950 j lof the Botanical R h Institute of Texas 2(2)
Fic, 18. Distribución de B
en el margen, con algunos pelos largos en la garganta; lígula un anillo ciliado de 0.3 mm de largo; láminas
foliares de 3 a 9 cm de largo y 1 a 2 mm de ancho, aplanadas. escabrosas, esparcidamente pilosas en la
superficie adaxial, curvas. Inflorescencia con 3 a 6 espigas persistentes; ramificaciones primarias espigadas
de 1 a 1.8 cm de largo, espaciadas y casi erectas, portando 10 a 15 espiguillas. Espiguillas 3 a 3.5(-4) mm
de largo, articulación por arriba de las glumas; glumas desiguales, glabras a esparcidamente pubescentes
sobre la quilla, primera gluma de 1.5 a 2 mm de largo, estrecha acuminda, segunda gluma de 2 a 2.5 mm de
largo, más ancha que la primera, aguda; lema de la flor perfecta de 2.5 a 3.2 mm de largo, esparcidamente
aplicado-pubescente sobre las nervaduras y la base, las nervaduras laterales se transforman en 2 aristas
fuertes, de 1 a 1.5 mm de largo, no sobresalen del ápice agudo de la lema. Flósculos rudimentarios (1) 2,
lema del primer flósculo de 1 mm de largo, ancha, con las tres nervaduras prolongadas en aristas de 1.5
mm de largo; segundo flósculo de 1 mm de largo, incluyendo las 3 aristas escabrosas, raquilla glabra en la
Harrara Arriata at al , n *al lal ta da Mávi 951
GA
ia
eee
Fic. 19. nier HA il. D trl, L O T I ta Ada KAZ
base del flósculo rudimentario. Cariopsis de 0.7 a 1 mm de largo, color amarillo claro a café en la madurez,
estrechamente elíptica. 2n = 20. Florece de junio a noviembre.
Distribución.—México en: Coahuila, Nuevo León, San Luis Potosí y Zacatecas.
Hábitat. —Suelos salinos y yesosos, en pastizales y les de Larrea al NE de México; 1000 a 1650 m.
hal M `
astaños, Rancho de S Teresa, al S de Castaños (MEXU); Municipio de
Ejemplares examinados. MÉXICO. C pio d
General Cepeda, Ejido La Rosa, carretera 40 Saltillo-Torreón, aprox. 20 km NE de General Cepeda, 5 Jul 1986, 1230 m, S. Vásquez 84,
A. García (ANSM); km 45 carr. 40 Saltillo-Torreón, en la desviación a Gral. Cepeda, Y. Herrera Arrieta, M. de la Cerda & O. Rosales 1478
(CIIDIR, HUAA); Municipio de Ocampo, 1-2 mi O of Matrimonio Viejo about mogote at b f slightl
Sep 1941, I.M. Johnston 9370 (MEXU, US); Laguna de La Leche, 22 Jun 1978, 1000 m, J. Valdés-Reyna 1334, H. Garza & L.E. Rodríguez
(ANSM); S of Laguna de La Leche, 30 Aug 1941, I.M. Johnston 8618 (MEXU); Laguna de la Vaca, Ocampo, J. Valdés-Reyna s.n. (ANSM);
Municipio de Saltillo, 26 mi NNE de C ión del Oro, J.R. Reeder 4000 (US). Nuevo León. Municipio de Dr. Arroyo, Ejido Lagunita
y Ranchos Nuevos, M.E. Pérez 108 (CIIDIR); Municipio de Galeana, Just E of San Roberto jct. along hwy. 57, J.R. Reeder & C.G. Reeder
y saline and gypseons slope, 23
952 l Lafttha Prataniral D h Institut f Texas 2(2)
7060 a IEB, sn E flats, 3 mi S of Galeana, D.S. Bade 19851 S G.A. González 674 (SLPM); G.A. Gómez 715 (MEXU,
SLPM); J. B, SLPM); J.R. Reeder & C.G. Reeder 2926 (MEXU); J.M. Peña 674 (UAG); J. Rzedowski 8281, 11460 (ENCB);
Navidad, carretera 57, km 85 al N de Saltillo, 23 Sep 1978, 1650 m, F. Cárdenas & M.A. Bernal (MEXU).
14. Bouteloua parryi (E. Fourn.) Griffiths, Contr. U. S. Nan, Herb. 146): 381, f. 37, pl. 73. 1912. Chondrosium
parryi E. Fourn., Mexic. Pl. 2:150. 1886. Tiro: MÉXICO Luis Potosí, 1878, C.C. Parry & E. Palmer 943 14
(HOLoTIPO: BR?; isoTiros: US-81783!, US-883835!).
Plantas anuales o perennes de vida corta, cespitosas. Culmos de 10 a 60 cm de alto, erectos o a veces ge-
niculados, generalmente con estolones. Vainas más cortas que los entrenudos, los márgenes hialinos, con
pubescencia larga, esparcida, de base papilosa, más abundante cerca del collar; lígula ciliada de 0.5 a 0.8
mm de largo; láminas foliares de 1 a 15 cm de largo y 0.5 a 3(-4) mm de ancho, las superiores más cortas,
glabras a piloso-papilosas en ambas superficies, los pelos llegan a medir hasta 3.5 mm de largo, principal-
mente en los márgenes hacia la base. Inflorescencia de 6 a 30 cm de largo, con 3 a 8 ramificaciones primarias
espigadas persistentes; ramificaciones de 1 a 3 cm de largo, portando 25 a 65 espiguillas sobre un raquis
plano, piloso-papiloso; raquis sin extenderse más allá del punto de inserción de la espiguilla terminal.
Espiguillas de 3 a 4(-6) mm de largo, incluyendo las aristas, articulación por arriba de las glumas; glumas
desiguales, primera gluma de 1 a 1.8 mm de largo, hialina, glabra, aristada, segunda gluma de 3 a 3.5(—5)
mm de largo, corto aristada, la arista hasta de 1 mm de largo, emergiendo de un ápice bífido, piloso-papilosa
sobre la quilla; lema de la flor perfecta de 3.5 a 4 mm de largo, pilosa en la base, 3-aristada, aristas de 1 a
3 mm de largo, alternando con 3 lóbulos; pálea del mismo tamaño que la lema, dentada, 2-aristada, aristas
pequeñas, ligeramente pubescente en los márgenes. Anteras de 2 a 3 mm de largo, color crema a amarillo
claro. Flósculos rudimentarios 2, lema del primer flósculo de 2 a 3 mm de largo, barbada en la base, 3-aris-
tada, aristas de 2 a 3 mm de iargo, con lóbulos obtusos; lema del segundo flósculo sin aristas, reducido a
una escama. Cariopsis 1.8 a 2.5 mm de largo, anguloso-triangular.
Comentarios.—Especie con dos variedades reconocidas, sólo la una variedad en Coahuila.
Bouteloua parryi var. parryi (E. Fourn.) Griffiths. (Fig. 20).
Bouteloua polystachya var vestita S. Watson, Proc. Amer. Acad. Arts 18:177. 1883. Bouteloua vestita (S. Watson) Scribn. ex L.H. Dewey,
ontz. U.S. Natl. Herb. 2(3):531. 1894. Tio: MÉXICO: Sierra Madre S of Saltillo, E. Palmer 1357 (HoiortPo: GH).
Plantas anuales. Culmos de 20 a 60 cm de alto, erectos o abiertos basalmente, no estoloníferos. Láminas
foliares de 3 a 15 cm de largo y 2 a 3(-4) mm de ancho, piloso-papilosas. Inflorescencia de 15 a 30 cm de
largo, con 4 a 8 ramificaciones primarias espigadas, de 2 a 3 cm de largo, con 40 a 65 espiguillas. Espiguillas
de 3.5 a 4(-6) mm de largo. Florece de agosto a diciembre.
Distribución.—Desde el Sw de Estados Unidos a México en: Coahuila, Colima, Chihuahua, Durango,
Jalisco, San Luis Potosí, Sinaloa, Sonora y Zacatecas.
Hábitat.—Se le encuentra en laderas arenosas, mesetas y colinas rocosas, y a orilla de carretera.
Ejemplares examinados. MÉXICO. Coahuila: Municipio de Saltillo, Sierra S of Saltillo, E. Palmer 1357 (US).
E
15. Bouteloua pedicellata Swallen, N. Amer. Fl. 17:627. 1939. (Fig. 21). Tiro: MÉXICO. PuesLa: Chalchicomula,
9000 ft, 19 Aug 1910, A.S. Hitchcock 6302 (notorio: US: 8839501; isorieo: UST).
Plantas bajas perennes. one de ae a 25 cm du aa CrsanTo tando uo od y espiralado-aéreos,
hasta de 45 cm de largo. H inci dos, escabrosas, con
escasa pubescencia larga, disperse: ligula de 0.5 a0. 8 mm de largo, un anillo ciliado denso; Mas foliares
de 2 a 5 cm de largo y 0.4 a 0.7 mm de ancho, involutas, algunas aplanadas cerca a la inflorescencia, color
verde-grisáceas, escabosas en ambas superficies, con algunos pelos largos dispersos en la superficie adaxial,
O ae Hacia la id cerca ac G "s Inflorecencia 8 a 12(-15) cm de largo, largamente exserta,
con 1022 ifi di la madurez, divergentes o reflejas; ramificaciones
de 8 a 9(-10) mm de DS la mayoría spartans una sola espiguilla péndula, conspicuamente pedicelada,
pedicelo de 2 a 3 mm de largo; raquis aplanado, con márgenes ciliados. Espiguilla de 8 a 9 mm de largo,
permanece inserta a la espiga; glumas anchas, firmes, sin aristas, escabroso-pubescentes en las quillas, con
Fig. 20. Distribución de B.
tintes purpüreos, primera gluma de 4 a 5 mm de largo, con ápices obtusos, finamente ciliados hasta los
márgenes, segunda gluma de 8 a 9 mm de largo, aguda a acuminada, con la quilla escabroso-pubescente,
márgenes angostamente hialinos; lema de 7 a 8 mm de largo, glabra, con los márgenes hialinos, sin arista;
pálea de 6 a 7 mm longitud, angosta, ligeramente escabrosa. Anteras de 2.5 a 3 mm de largo, color amarillo
claro. Flósculo rudimentario 1, reducido a tres aristas escabrosas, diminutas, las laterales de 2 mm de largo
y la central más ancha y larga, de (3.54 a 4.5 mm de largo, no exserta de la espiguilla. Cariopsis de 2.4 a
3 mm de largo, color pajizo, oblongo-elipsoide. Florece de julio a septiembre.
Distribución.—México en: Guanajuato, Puebla, Tlaxcala y Veracruz; ahora encontrada en Nuevo
Leon:
Hábitat.—Sobre laderas rocosas, principalmente en pastizales; 1950-2500 m de altitud.
Ejemplares examinados. MÉXICO: Nuevo León: Municipio de Dr. Arroyo, 8.8 km W of Dr. Arroyo and 37 km E of Matehuala on Hwy
3,13 Oct 1991, 1950 m, P.M. Peterson 11140, C.R. Annable (US)
954 I l ^f tha Botanical R h Institut f Texas 2(2)
Fic 21. Distrihució lIa D L At all nt I ta dn Mv!
16. Bouteloua radicosa (E. Fourn.) Griffiths, Contr. U.S. Natl. Herb. 14:411. 1912. (Fig. 22). Atheropogon radicosus
Fourn., Mexic. Pl. 2:140. 1886. Bouteloua} id radicosa (E. Fourn.) Vasey ex L.H. Dewey, Contr. U.S. Natl. Herb. 2(3):533.
1894. Tiro: MÉXICO: near México City, 8 Jun 1865-18 Jun 1866, M. Bourgeau 450 ( P designado p ffiths, Contr. U.S.
Natl. Herb. 14:411, 1912; isotscroriPOs: BR, F GH, LE, MPU, US-81791!, US-87200 fragm. ex P!, US-883896 fragm.!).
Dinebra t ides Kunth, Nov. Gen. Sp (quarto ed.) 1:172, t. 51. 1816. Atheropogont les (Kunth) R & Schult., Syst. Veg. 2:415.
1817. Eutriana bromoides (Kunth) Trin., Fund. Agrost. 161, 162. 1820. Eutrianal ides (Kunth) E h, Revis. Gramin. 1:95. 1829
Tiro: MÉXICO. GUANAJUATO: "Crescit i peratis scopulosis aridis i
& A. Bonpland s.n. (notorio: P; sorro: US-865871 fragm. ex P y foto!).
Guanaxuato et Cubilete Mexicanorum," Sep, E Humboldt
Plantas perennes. Culmos de 40 a 80 cm de alto, erectos, base conspicuamente rizomatosa, rizomas muy
engrosados y escamosos; entrenudos de 1 a 2 mm de largo en la base, típicamente cubiertos por vainas firmes.
Hojas generalmente basales; vainas glabras, más cortas que los entrenudos; lígula de 0.5 a 1 mm de largo, en
El
forma de un anillo de pelos; láminas de 6 a 1030) cm de largo y 1.5 a 2.5 mm de ancho, aplanadas,
Nj
pa
A td
( $ 8 ra
Fic. 22. Dictril HA la Drontal ge 1 tn Adan TI
ciliadas en los márgenes, con pelos de base papilosa, de 2 a 4 mm de largo, a menudo esparcidamente pilosas
en ambas superficies. Inflorescencia de 6 a 15 cm de largo y 5 a 8 mm de ancho, con 5 a 12 ramificaciones
primarias espigadas, erectas o ascendentes, oblongas en el contorno general, desprendiendose íntegras en
la madurez; ramificaciones de 1.5 a 3 cm de largo, portando 6 a 12 espiguillas, por lo general barbadas en
la base, insertas sobre las % partes de un raquis glabro, así el raquis se prolonga 4 a 8 mm delante de la
espiguilla terminal. Espiguillas 9 a 13 mm de largo, incluyendo las aristas, permanecen insertas a la espiga;
glumas desiguales, 1-nervadas, con tintes purpúreos, primera gluma de 5 a 6 mm de largo, ancha, aguda
a acuminada, glabra o con escasa pubescencia en el raquis, segunda gluma de 6 a 7 mm de largo, glabra a
escabrosa en la quilla, acuminada; lema de la flor perfecta de 7 a 9 mm de largo, sin aristas o con 3 aristas
cortas, la central hasta de 2 mm de largo, glabra a escabrosa hacia el ápice; pálea ligeramente mas pequeña
que la lema, sin nervios notorios, glabra o con escasa pubescencia a lo largo de la parte central. Anteras de 3
a 4 mm de largo, rojizas. Flósculo rudimentario 1, lema del flósculo de 8 a 12 mm de largo, incluyendo las 3
956 j | of the Botanical R h Institute of Texas 2(2)
aristas fuertes, aplanadas, con tonos purpúreos; pálea angosta, casi tan larga como la lema. Cariopsis de 1.3 a
1.6 mm de largo, color amarillo, ovoide a estrechamente elíptica. 2n = 20,60. Florece de julio a diciembre.
Distribución.—Sw de Estados Unidos a casi todo México.
Habitat —En laderas secas rocosas, pastizales y matorrales xerófilos; 210 a 2000 m.
Ejemplares examinados. MÉXICO. Coahuila: Municipio de Castaños, 1 km N de cuesta La Muralla y restaurant la Muralla, 24 Sep
1977, 1150 m, J. Valdés-Reyna 1308, L.E. Rodríguez (ANSM); S end of Puerto San Lazaro (Cuesta La Muralla), 0.5 mi N of San Lázaro
along route 57, 28 Jul 1977, 1250 m, J. Valdés-Reyna 1102, T.L. Wendt (ANSM); S end of Puerto San Lázaro (Cuesta La Muralla), 0.5 mi
N of San Lá long route 57, 28 Jul 1977, 1250 m, T.L. Wendt & J. Valdés-Reyna (CHAPA, MEXUJ; Sierra de la Gavia, Canón de la
Gavia, aproximadamente 6 km S de la casa camino al cañón, 14 Aug 1986, 1480 m, J. Valdés-Reyna 1709, M.A. Carranza & D. Frame
(ANSM); Sierra de la Gavia, Cañón la Gavia, 6 km S de la casa camino al cañón, 20 Sep 1989, 1480 m, P.S. Hoge 306, M.E. Barkworth
& M.A. Carranza ee o og o poids al Casco del rape ON de Menchaca, 30 Aug 1980, 1180 m, R.
Vásquez 19, LE.R San Marcos, frent de Cuatrociénegas, 24 Jun 1989, 800 m, A.
Rodríguez 1221, M. MEUS & jJ. m (ANSM); Sierra de San Marcos, cañón Granda, Ejido Estanque de Norias, aprox. 43 km O de la
carretera 57 rumbo a Ejido Reforma, 1 Sep 1993, 1550 m, M.A. Carranza 1708, J.A. Encina, J. Fierro & R. Rodríguez (ANSM); Municipio
de General Cepeda, 20 km sobre la carretera de General Cepeda-Parras, 12 Sep 1986, 1250 m, J. Espinosa s.n. (ANSM); Municipio de
Müzquiz, Rancho La Peña, Sierra de Santa Rosa, 5 Dic 1981, 2000 m, R. Salgado s.n. (ANSM); Sierra Santa Rosa, 11 Ju 1976, 1800 m,
R. Vásquez s.n. (ANSM). Nuevo León: Municipio de Cadereyta Jiménez, Carretera a Reynosa, 2 kmO de la Refinería Cadereyta, 26 Sep
1979, 320 m, J.A. Villarreal, J. Valdés-Reyna (ANSM); Municipio de Cerralvo, 5.2 mi S of e on m towrds Ejido La Encantada,
21 Sep 2002, 1500 m, P.M. Peterson 16738, J. Valdés-Reyna & M. Sosa (US); Municipio d , In desert scrub 17 mi E of Monterrey,
1 Jul 1947, F.A. Barkley 7178, G.L. Webster & C.M. Rowell Jr. (MEXU); Municipio de G | Zaragoza, 4 km S of Zaragoza at jct of road
to Cerro Viejo-Tepehuanes, 20 Sep 2001, 1630 m, P.M. Peterson ERA J. Valdés. ue (US); Municipio de Iturbide, Canada Marrubial,
a Rosa, Hu 1985, 1320 m, B.A. Treviño 500 SM); Ejid 29 Aug 1989, 1550 m, E.A. Pta o o
E del Cerro Colorada; 17 Mar 1981, 500 m, 0. L. Briones 4
Los iuit cañón de las Ead 8 Sep 1982, 700 m, T.S.P. s.n. (ANSM); 8 mi de Linares sobre E carr. à Glens J Crutchfield 6 M. Es
Johnston 5835c (US); Municipio de Sabinas Hidalgo, Sabinas mus 29 Jun 1933, C.H. Mueller 419, M.T. Mueller (MEXU); Municipio de
Salinas Victoria, La Soledad Salinas Victoria, May 1981, 500 m, J.A. Ochoa 1130 (MEXU). Tamaulipas: Municipio Abasolo, 13 mi E de
Abasolo, desviación a la carr. San T. Jimenez, J. Crutchfield & M.C. Johnston 6140* (US); Municipio de Hidalgo, Ejido Nicolás Bravo, 30 Oct
1981, J.A. Barrientos 103 (MEXU); La Aurora, 17 Aug 1983, 280 m, J.F. Iribe 195 (MEXU); Municipio de Llera, Estación San Francisco,
Sep 1985, C.R. López 126 (MEXU); La Gloria, 14 Nov 1993, 210 m, J.E. López 152 dida pe de Miquihuana, 18 km al SO de
Miquihuana, 15 Aug 1972, 1700 m, F. González 4754, M. López & R. Dirzo (MEXU); ^ pio de San Carlos, Piedra Imán, 2 kmESE de
San José, Sierra de San Carlos, 22 Nov 1984, 790 m, O.L. Briones 1215 (ANSM); Municipio de Soto La Marina, 35 km de Victoria, sobre
la carr. a Casas y Soto la Marina, F. Martínez 2305 (US); Municipio de Tula, Camino a 5an José de las Flores, 23 Oct 1986, 1280 m, J.E.
López 10 (MEXU); Municipio de Victoria, Colonia La Estrella, Ciudad Victoria, 20 Sep 1985, J.G. Galván 85 (MEXU).
AMNTCORAN
17. Bouteloua ramosa Scribn. ex Vasey, U.S.D.A. Div. Bot. Bull. 12(1): t. 44. 1890. (Fig. 23). Bouteloua oligos-
tachya var. ramosa (Scribn. ex Vasey) Scribn. ex Beal, Grass. N. Amer. 2:418. 1896. Tipo: U.S.A. Texas. Presidio Co.: Chenate Mts.,
1888, G.C. Nealley 797 (Lecroriro: US-8832441, designado por J. Reeder, Taxon 35:153. 1986
Plantas perennes, con bases duras, nudosas y subrizomatosas. Culmos de 25 a 60 cm de alto, delgados, ge-
niculados, correosos, conspicuamente ramificados en la mitad inferior, barbados en las axilas de las ramas;
nudos 4 a 5, entrenudos glabros. Vainas más cortas que los entrenudos; lígula de 0.1 a 0.2 mm de largo, un
collar de pelos diminutos; láminas foliares de 2 a 5(-7) cm de largo y 1 a 2 mm de ancho, aplanadas con el
ápice involuto, membranáceas, nunca dísticas, acuminadas, pubescentes en la superficie adaxial y glabras
en la superficie abaxial. Inflorescencia de 5 a 8 cm de largo, con (1) 2 (4) ramificaciones primarias espigadas
persistentes; ramificaciones densas, de 1 a 2.53.5) cm de largo y 4 a 5 mm de ancho, pectinadas, portando
25 a 45(—60) espiguillas; raquis pubescente, casi glabro con el tiempo, terminando en una espiguilla a menudo
reducida y en forma de aguja. Espiguillas 4 a 5 mm de largo, articulación por arriba de las glumas; raquilla
con mechones de pelos de 1 mm de largo en la base de la flor perfecta y de 2 mm en la primera flor reducida;
glumas desiguales, agudo-acuminadas, glabras o esparcidamente pilosas en la base, primera gluma de 2 a
2.5 mm de largo, con una arista de 0.5 mm de largo, segunda gluma de 3 a 3.2 mm de largo, con un mucrón
de 0.8 a 1 mm de largo, escabrosa; lema de 4 a 5 mm de largo, densamente pubescente, excepto en el ápice,
con 3 aristas gruesas, de 1 a 1.2 mm de largo las laterales y de 1.5 a 2 mm de largo la central, escabrosas, de
color púrpura; pálea de 2.5 a 4-5) mm de largo, glabra. Flósculos rudimentarios 1 o 2, primer flósculo con
tres aristas fuertes, de 3 a 5 mm de largo, incluyendo las aristas, híspida; segundo flósculo cuando presente
Harrara Arriata at al., n tal dal tn da Mu 957
Fic. 23 Dictrib 145 dn D. L I ta da MM
en forma de escama diminuta. Anteras de 1 a 1.5 mm de largo. Cariopsis obovada, de 2 a 2.5 mm de largo,
color amarillo claro, lanceolado-elíptica. 2n = 40. Florece de julio a diciembre.
: e" se MA
D 4
Oomentarins
VITEL PELE LU».
de B. breviseta por diferentes
y
autores, entre ellos Gould (1979); sin embargo, revisando los tipos de ambas esp t grandes
diferencias, la principal es de tipo vegetativo. Mientras que B. ramosa presenta | l ificados, delgados
y correosos, hasta de 70 cm de alto; B. breviseta posee culmos no ramificados, no son delgados y correosos,
por lo general son cortos, de 20 a 2530) cm de alto, rígidos, decumbentes y con láminas foliares cortas,
duras y dísticas. B. breviseta solo fué encontrada en una localidad de México en Chihuahua, aparentemente
su distribución se restringe a Texas en EUA y Chihuahua en México.
Distribuición.—Desde Texas al N y Centro de México.
Hábitat.—Se le encuentra en laderas pedregosas y secas, a lo largo de arenas yesosas y calcáreas; 650
a 2700 m.
958 I | -£4h, Dasanizal D h Institut f Texas 2(2)
Ejemplares examinados. MÉXICO. Coahuila: Municipio de Castaños, 5 mi S of Castaños, 3 Sep 1960, 884 m, J.R. Reeder 3276, C.G.
Reeder & T.R. Soderstrom (TAES, Pn 2 ae a N de Cañada Blanca, km 17 carr. 57 Saltillo-Monclova, Y. Herrera Arrieta, M. de la Cerda &
O. Rosales 1481 (CIIDIR, HUAA); Al y La Gavia, km 117 carr. 57 Saltillo-Monclova, a un lado del € I
Y. Herrera Arrieta, M. de la Cerda & O. Rosales 1485 (CIIDIR, HUAA); Municipio de Cuatrociénegas, 19 mi SW of NEN 6 Sep
1965, 1300 m, J.R. Reeder & C.G. Reeder 4523 (US), Puerto del Norte, Cuatrociénegas, L.H. Harvey 1207 Mah 3 mi N of Cuatrociénegas,
28 Oct 1964, 823 m, J.R. Reeder 3940, C.G. Reeder (MEXU, TAES); Aprox. 8 km NO de Cuatrociénegas, Ocampo, 12 Ene 1974,
740 m, J.S. Marroquín 2672 (ANSM); Bolsón de Cuatrociénegas, 21 km S de Cuatrociénegas, carretera 30 Cuatrociénegas-San Pedro
de las Colonias, a orillas de la Sierra de San Marcos, 17 Sep 1993, 850 m, J.A. Villarreal 7255, M.A. Carranza & R. Rodríguez (ANSM);
Ladera baja de la Sierra de San Marcos, frente a las dunas de Cuatrociénegas, 24 Jun 1989, 950 m, A. Rodríguez 1111, M. Martínez &
J.M. Sosa (ANSM); 24 Jun 1989, 800 m, A. Rodríguez 1220, N. Moreno & J.J. López (ANSM); Laderas de la Sierra de San Marcos 24 mi SO
Cuatrociénegas, 23 Sep 1990, 750 m, P.M. Peterson 10012, C.R. Annable & J. Valdés-Reyna i US); Sierra de San Salvador, carretera
San Pedro-Cuatrociénegas, 12 Ene 1974, 1350 m, J.S. Marroquín s.n. (ANSM); Tip of Si San Marcos, NE-facing slope, 8 Jul 1968,
DJ. Pinkava 5110, E. Lehto & D. Keil (MEXU); cs de General Cepeda, Carretera a Paila, llanura, 24 Sep 1973, 1140 m, Clase de
Botánica (ANSM); Near La Rosa, O of Saltillo, 9 Oct 1964, 1350 m, J.R. Reeder & J. Morrow s.n. (TAES); Ejido La Rosa, carretera 40
Saltillo-Torreón, aprox. 20 km NE de General Cepeda, 5 Jul 1986, 1500 m, S. Vásquez 100, A. García (ANSM); Sierra de la Paila al O en
el Ejido Nueva Imagen, 5 Oct 1985, 1400 m, A. Rodríguez 1461 (ANSM, CIIDIR); km 45 carr. 40 Saltillo-Torreón, en la desviación a Gral.
UM Y. Herrera Arrieta, M. de la Cerda & O. Rosales 1479 (CIIDIR, n Due ae Puerto = MM ears i km, across the
Ils at Tanque Jerico, 22 Aug 1941, I.M. Jol M
orenzo
DIDda Valle J E
ri 7200 (US); Monclova, C.G. Pringle 414 (US); Municipio d , 7.2 mi SE of Jaboncill d ds Cuesta de malena
Melchor Muzquiz, 01 Sep 2006, 1010 m, P.M. Peterson 19854, 5. Pa Contreras (US): icipio de O , Laguna de La Leche,
22 nus 178, 1000 m, J. Va Mes 2 1333, H. Garza & L.E. Rodríguez (ANSM); Rancho OH HM, La Rueda. 87 km NE de Ocampo,
brecha Ocampo-Boquillas armen, Sep 1976, 1580 m, R. Vásquez 76 & E. Lp eee ih 1976, des R. ns (ANSM);
2 Jul 1991, 1350 m, M. Vasquez s.n. (ANSM); ewes Las Gallinas, ap km d Mojada, 2 Oct
1990, 1300 m, M.A. Carranza 569, FJ de la Madera, Rancho Laguna de la 1 UM imad
Ocampo rumbo a Sierra Mojada, 2 Oct 1990, 1000 m, M. A. Carranza 615, F.J. Carranza (ANSM); Municipio de Parras, Ejido 7 de Enero,
28 Aug 1982, 1450 m, L.E. Rodríguez s.n. (ANSM); La Peña, Ejido 4 de Marzo, 6 Aug 1976, 1170 m, E.A. Roing 129 (ANSM); Municipio de
Parras, 11 Sep 1993, 1310 m, G.B. Hinton 23189 (ANSM); Sierra de Parras, 29 Sep 1978, 1850 m, J. Valdés-Reyna 1354, M.A. Capó (ANSM);
Sierra de Parras aprox. 2 km SO del Ojo de Agua, 15 Sep 1980, 1800 m, A. Rodríguez 255%, M. Martínez (ANSM); Sierra de Parras, aprox.
6 km SO del Ojo de Agua, 30 Ene 1981, 1750 m, Á. Rodríguez 184 (ANSM); Municipio de Piedras Negras, 26 mi N de Saltillo, camino a
Piedras Negras, M.C. Johnston 5870* (US); Municipio de Ramos Arizpe, km 30, carr. 40 Saltillo-Monterrey, en Ramos Arizpe, Y. Herrera
Arrieta & A. Cortés 1454 (CIIDIR, HUAA); 15 km N de Saltillo, carretera 57 Saltillo-Monclova, 22 Aug 1984, 1200 m, J.A. Villarreal 2654,
M.A. aa 6 - Webbe (ANSM, CHAPA); isi s PIE Sierra de la pun. i ads bad 1300 m, J.A. Villarreal 5189, M.A dd
e Norias, 14 Sep 1973, 1200 m s Sabanier, aprox
20 Oct 1989, 1500 m, P.M. Petron: 8441, J. Valdés-Reyna & A. Rodriguez (ANSM); 23 Sep 1988, A. Rodriguez 1916, J. Valdés-Reyna & M.
Martínez (ANSM); 19 Oct 1989, 1410 m, P.M. Peterson 8441, J. Valdés-Reyna & P.A. Fryxell (US); San Juan de las Bonitas, 19 Jul 1993,
450 m, E. Bde 23 o zie in la Paila, campo experimental de La Sauceda, 19 Oct 1972, 1650 m, J.S. Marroquín 2334 (ANSM),
M c bo a Cuatrociénegas, 30 Sep 1989, 650 m, A. Rodríguez 1230, N. Moreno & J.J. López
(ANSM): 10 km de S bo a Cuatrociénegas, 30 Sep 1989, 650 m, A. Rodríguez 1231, N. Moreno & J.J. López (ANSM, CHAPA);
Municipio de Saltillo, Saltillo, E. dd 404 oim 12 = O de dpud carretera M camino a La estación de Microondas Vega, 9 Jul 1983,
2000 m, J. Valdés-Reyna 1533, L.E 6miO y 40 toward Torreón, 25 Aug 1983, 1680 m, S.L.
Hatch 5042, J. Valdés-Reyna Hn on del Pueblo, O de la Ciudad de Saltillo, 11 Sep 1990, 1650 m, J. Valdés-Reyna 2052, M. Martínez
& M.A. Carranza (ANSM, CHAPA); Buenavista, Saltillo, A. Rodríguez 131 (ANSM); Municipio de San Pedro About 121 li
Las Delicias on the E face of a ple Sierra, 25 Sep 1972, 1000 m, M.C. Johnston 9530, F. Chiang & T.L. Wendt (ANSM); 5.5 km al NW de
Nueva Delicias, vertiente E de la Sierra de las Delicias, al N de San Pedro de las Colonias, Y. Herrera Arrieta, M. de la Cerda & O. Rosales
1490 (CIIDIR, HUAA); Al 5W de s oen con Durango-Chihuahua), D. López 16 (CHAPA); Municipio de Sierra Mojada, San
Antonio de los Alamos, E base of the V. rra de San Antonio, 20 Aug 1941, I.M. Johnston 8249 (MEXU); Municipio de Torreón,
Cañón Picardias, 2 km antes de Juan Eugenio, Torreón, J.A. Villarreal, M.A. Carranza, A. Orta & J. Elizondo 2276 (CIIDIR); 11.7 km E of
Flor de Jimulco at an abandoned mine, 21 Oct 1989, 1730 m, P.M. Peterson 8456, J. Valdés-Reyna, J.A. Villarreal & P.A. Fryxell (US); 10
mi S de Jimulco, F. Shreve 9414 (US); Sierra de Jimulco, proximidades al ejido La Trinidad, 25 Aug 1988, 1900 m, J.A. Villarreal 4411,
M.A. Carranza & A. Rodriguez (ANSM, CIDIR, IEB); Municipio de Torreón, Sierra El Rosario, carr 49, 30 km al SE de Lerdo, Dgo., J.A.
Villarreal 5803,M.A. Carranza (ANSM); Municipio de Zaragoza, 16.9 km NE of San Miguel on road to Boquillas, 13 Sep 1991, 1180 m,
M. Peterson 10612, C.R. Annable (US). Nuevo León. Municipio de Galeana, 8 km al entronque de San Roberto, rumbo a Galeana, Y.
Herrera Arrieta & A. Cortés 1449 sae rods Municipio de Monterrey, Carr. Villa de García a Grutas, W de Monterrey, T. Tateoka
1087 (US). Tamaulipas: } Ejido El Aserradero, 4 Sep 1988, 2700 m, R.A. Carranco 81 (MEXU
18. Bouteloua repens (Kunth) Scribn., Bull. Div. Agrostol., U.S.D.A. 4:9. 1897. (Fig. 24). Dinebra repens Kunth,
Nov. Gen. Sp. (quarto ed.) 1:172. t. 52. 1816. Atheropogon repens (Kunth) Roem. & Schult., Syst. Veg. 2:416. 1817. Eutriana repens
Larra
ieta et al., B tal dal dn e E 959
Fic. 24. Distribución da R F}
(Kunth) Trin., Gram.Unifl. Sesquifl. 241.1824. Tiro: MÉXICO. Guerrero: Acapulco, “Crecit ad litora Ocwani Pacifici prope,” E
Humboldt & A. Bonpland s.n. (ectotiro: P, designado por McVaugh, Fl. Novo-Galiciana 14:91. 1983 en nombre solo).
ega Trin., Gram. Unifl. Sesquifl. 242.1824
AT
Heterosteca juncifolia Desv., Nouv. Bull. Sci. Soc. Philom. Paris 2:188.1810. Eutriana heterost
f A J: Hot + E ioi: TS +] E n 1 E E 5 Á +L Syret
Veg. 1:294. 1825. Eutriana juncifolia (Desv.) Kunth, Revis. Gramin.1:95. 1829. Dinebra juncifolia (Desv.) Steud., Nomencl. Bot. (ed.
mboldtiana Griseb., Mem. Amer. Acad. Arts, n.s. 8:532. 1862 (basado encendido Heterosteca juncifolia
Desv, 1 juncifolia Lag.). Bouteloua} ga (Trin.) Griffiths, Contr. U.S. Natl. Herb. 14(3):414, f. 59. 1912.
CUBA: La Habana, “Habitat in Antillis," A. Bonpland 132B (HoLoripo: P-DESV; isoripo: US-2830954 fragm.!).
Bouteloua bromoides Lag., Gen. Sp. Pi. 5.1816. Actinochloa bromoides (Lag.) Roem. & Schult., Syst. Veg. 2:420. 1817. Eutriana bromoides
(Lag.) Trin., Gram. Unifl. 241. 1824. Tiro: MÉXICO. Guerrero: Acapulco (noLoriPo: MA; isoripo: US-865568 fragm.!).
Atheropogon filiformis E. Fourn., Mexic. Pl. 2:140. 1886. Bouteloua filiformis (E. Fourn.) Griffiths, Contr. U.S. Natl. Herb. 14(3):413,
12. Tiro: MÉXICO: “Hacienda de la Naranja,” Karwinski 991b : LE, designado por McVaugh, Fl.-Galiciana 14:91. 1983
en nombre solo).
2) 1:510. 1840. Bouteloua hu
I A R 1
960 J | of the Botanical R h Institute of Texas 2(2)
Heterosteca rhadina Nash, Bull. Torrey Bot. Club 30:386. 1903. Tipo: PUERTO RICO: Near Ponce, 18 Nov 1902, A.A. Heller 6057 (uo-
ormo: NY-71113}; isoripo: US-426292!).
mi pubescens Pilg., Verh. Bot. Vereins Prov.Brandenburg 51:193. 1909. Tiro: GUATEMALA. Derro. HUEHUETENANGO: Malactan, 13
1896, E. Seler 3224 (isoripo: US-81707 fragm.!).
Plantas perennes, amacolladas. Culmos de 20 a 60 cm de alto, erectos o esparcidamente ramificados en el
tercio inferior a abierto-decumbentes, generalmente débiles, partiendo de una base firme y nudosa; entre-
nudos y nudos glabros a escasamente hirsutos. Vainas glabras, diminutamente hirsutas o con algunos pelos
dispersos; lígula de 0.2 a 0.5 mm de largo, un anillo de cilios blancos; láminas foliares de 4 a 12 cm de largo
y de 2 a 3(-5) mm de ancho, lineares, aplanadas, glabras o escasamente hirsutas, con pelos de base papilosa
de 2 a 4 mm de largo, cerca de la base y en el margen. Inflorescencia de 5 a 10 (14) cm de largo, largamente
exserta, solitaria, terminal, con 7 a 10 ramificaciones primarias espigadas caedizas en la madurez; espigas
de 1.5 a 2.5 cm de largo, ascendentes a patentes, espaciadas a lo largo del eje, pero apiñadas hacia el ápice
de la inflorescencia, cuneadas en la base, portando 6 a 10(-12) espiguillas, densamente imbricadas en la
mitad inferior del raquis. Espiguillas de 6 a 9 mm de largo, congestas, permanecen insertas; glumas anchas,
agudas, de color púrpura, primera gluma de 4 a 7 mm de largo, hispida en la quilla, con el ápice mucronado
a ligeramente bifido, segunda gluma de 6 a 9 mm de largo, más ancha, glabra, con el ápice mucronado; lema
de la flor perfecta de 5 a 9 mm de largo, glabra, rara vez barbada en la base, con 3 aristas cortas, hasta de
1 mm de largo, la central más fuerte y larga, ligeramente escabrosa; pálea de 4.5 a 8 mm de largo, glabra a
finamente estrigosa hacia el ápice, las nervaduras proyectándose en 2 pequeñas aristas del mismo tamaño
que las de la lema o un poco mayor. Anteras de 3.5 a 5.5 mm de largo, color anaranjado, amarillas o rojizas.
Flósculo rudimentario 1, bien desarrollado, estaminado; lema del flósculo de 5.5 a 7 mm de largo, incluyendo
las 3 aristas, aristas de 2 a 10 mm de largo, la central fuerte y escabrosa, las laterales más cortas, la mitad
del tamaño o un poco más largas que el cuerpo de la lema. Cariopsis de 2.4 a 3 mm de largo, ovoide a es-
trechamente elíptica. 2n = 20, 40, 60. Florece de agosto a noviembre.
Distribución.—S de Estados Unidos hasta todo México; Antillas: Cuba, Haití, Puerto Rico y República
Dominicana.
Habitat —En terrenos abiertos de pastizales, planicies o laderas sobre suelos arenosos o pedregosos,
en bosques de encino, de pino, oe y matorrales xerófilos; 50 a 1600 m.
1 p Jo M. L 30 Aug
$ o
Ejemplares examinados. MÉXICO. Coahuila: M ipio d Cuatrociénegas Entrada al Casco del R
1980, 1600 m, R. Vasquez 19%, L.E. Rodríguez (ANSM); Majo iS apace Entre el Coyote y Palo Blanco, Oct 1976, F. Gonzdlez 6309,
R.M. López & R. Hernández (MEXU); Rancho Santa Barbara, 28 Nov 1989, 190 m, J.A. Garcia s.n. (UAT); Municipio de Saltillo, N of
Saltillo on Mex 57, N of Lazaro Pass, and 0.5 km S of La Muralla RE 13 Oct 1989, 1150 m, P.M. Peterson 8379 & J. Valdés-Reyna
(US). Nuevo León: Municipio de Allende, Río Ramos, 1 km S de Allende, carretera 85, 29 Oct 1992, 405 m, J.A. Villarreal 6789, M.
Vásquez, D.E. Lozano & J. Valdés-Reyna (ANSM); Río Ramos, 4 km 5 de Allende, carretera 85, 24 Oct 1986, 460 m, J. Espinosa s.n. (ANSM);
Rio Ramos, 1 km al S de Allende, J.A. Villarreal, M.Vázquez, D.E. Lozano & J. Valdés-Reyna s.n. (CHAPA); 9 mi N of Allende on Mex 85, J.
Brunken & C. Perino 180 (CHAPA); Entre Montemorelos y Allende, FW. Gould 9822 (US); Municipio de Anáhuac, La Gloria, Jul 1978,
180 m, G. Villegas 241, Flores (MEXU); Municipio de Bustamante, Grutas de Bustamante, 18 Jun 1993, 1300 m, P. Jauregui 76 (MEXU);
d de Cadereyta Jiménez, Carretera a Bones a, 2 km O de la Refinería Cadereyta, 26 Sep 1979, 320 m, J.A. Villarreal & J. Valdés-
Reyna (MEXU); Municipio de Cerralvo, Puert trada del Rancho Tinajas a 5 km de la carretera Doctor González-Cerralvo, 14 Feb
1987, 540 m, P.A. Garcia d (MEXU); Municipio de Doctor Gone Carretera Doctor Gonza lez-Cerralvo, km 52, 19 May 1993, 400
m, N. AN ad a 80 (MEXU); de Galeana, El Saucit Galeana Linares, Nov 1980, 600 m, J.A. Ochoa 1268 (MEXU);
vo, General Bravo, Dic "cn 150 m, J.A. Ochoa 1161 Sache n mi E de i Bravo sobre E carr. a ue
J. Crutchfield & M.C. sn 6056 (US dd Falda de la Loma I
1Jul 1 P.J MEXU): Municipio de I EjidoI eu eones, alt. 250, 18 May 1980, J.J. Ortiz s.n. (ANSM);
Localizad I ho El o BE en ine carretera es San Roberto, km 12, 2 Jun 1993, 440 m, M.M. Castillo 80 (MEXU);
Municipio de Los Ramones, A la carretera Los Ramones, 14 Abr 1982, 210 m, J.A. Villarreal 1517 (ANSM); Ejido el Car-
rizo, 20 Oct 1976, 210 m, D. Dávila s.n. (ANSM); Los Ramones, 4 Abr 1985, 210 m, J.A. Villarreal 2877 (ANSM); Municipio de Marín,
Facultad de Agronomía, Universidad Autónoma de Nuevo León, km 17, Marín, 22 Oct 1992, M.M. Castillo 20 (MEXU); Municipio de
Monterrey, Aug 1911, 540 m, F. Abbon 6245, G. Arsene (MEXU); Near Monterrey, 26 Jun 1888, C.G. Pringle s.n.(MEXU); Valley of Mon-
terrey, 6 Jun 1889, C.G. Pringle 2547 S MIU B.C. m 1905 (US); 20 km ai NW de eim o S de Mentes J.M.
Weaver 588 (US); Near 934 km post, S g , 17 Jul 1955, H.M. Chute 280 Q pio de Muzquiz,
Larrara Arriata etal., B tal Anal ta daB 961
a La Pefia, Sierra ide nad Rosa, ind R. bx ad (CIIDIR, SLPM); Municipi Sabinas Hidalgo, Fracción del Río Sabinas,
4 Mar 1993, N. Bazaldua 41-A (MEXU); S de Sabinis Hidalgo, C.H. Mueller
419 (US); Municipio de Saltill ludin A. Ruiz 35 (ANS M); Buenavista, R. Palomo 357 (ANSM); Municipio de Santiago, Blanquillo,
22 Jun 1994, 370 m, G.B. Hinton 24439 (MEXU); Xeric O wall of Cañón de Potrero Redondo, 4 Jul 1935, C.H. Mueller 2108 (MEXU).
Tamaulipas: Municipio de Aldama, 20 km N of Aldama on the road to Soto La Marina, at the Hacienda La Concepción, 20 Sep 1956,
50 m, F. Martínez 1792, G. Borja (MEXU); Camino a la Barra del Tordo, 4 Mar 1987, P. Moya 205 (MEXU); Camino al Rancho El Olmo,
11 Jun 1992, 260 m, J.E. López 96 (MEXU); Ejido San Felipe, 11 Jun 1992, 230 m, J.F. Iribe 349 (MEXU); km 10 a Barra del Tordo, 24
Sep 1980, 160 m, J. Cantú 2 (MEXU); Poblado Piedras Negras, 13 Sep 1988, 180 m, J.G. Galván 236 (MEXU); Pozas del ejido E] Naci-
miento, 5.5 km SO de Aldama, 31 Aug 1984, 260 m, L. Hernández 1170 (UAT); Rancho La Concepción, 11 Jun 1981, 75 m, G. Villegas
457 (MEXU); Rancho La Granja, 3 Mar 1987, 180 m, P. Moya 204 (MEXU); Sierra de Tamaulipas, Region of Rancho Las Yucas, 40 km
NNO of GE near Paso Lajas (Ejido Las Yucas), 10 Oct 1957, R.L. Dressler 2326 (MEXU); Municipio de Altamira, 15 mi from Tam-
pico on the Mante highway, 27 Sep 1959, M.C. Johnston 4082, J. Graham (MEXU); Ejido Santa Juana, 23 Oct 1980, 90 m, J. Cantú 8
(MEXU); in de Antiguo Morelos, Ejido Guadalupe Mainero, 25 Sep 1981, 700, J.F. ide 100 MERE Santa Fé, 19 Aug 1981, 130
m, J.F. Iribe 58 (MEXU); J.F. Iribe 795 (1EB); Santa Teresa,19 Aug 1981, J.F. Iribe 66 (MEXU); Municipi de Casas, 35 km from Victoria
on the road to Casas and Soto la Marina, 3 Oct 1956, 280 m, F. Martínez F-2350, G. ndn L. E 64 km from Soto la Marina on the
(old, wading) aes to aed and Victoria, A Oct 1956, 350 m, F. Ae F-2361, G. Borja (MEXU); Rancho La Trinidad, 18 Sep 1984,
xj 1 Ejido El Nacimiento, 27 May 1990, 100m E. Crespo 304 (ANSM);
La Poza Azul, to del río Frí Te de la Biosfera Fl Cielo, 25 Aug 1993, 300 m, J.L. Mora 429 (UAT) ica de González,
24.5 km al E de Xicotencatl, 11 Jun 1992, 170 m, J.G. Galván 271 (MEXU); 5 km al N de ález, Mar 1991, 110 m, G. Bores 140 (MEXU);
Ejido Josefa Ortíz de Domínguez, Sep 1983, 380 m, M.H. Cervera 191 (MEXU); Sep 1983, C.R. López 91 (MEXU); Orilla Magiscatzin, 3
Mar 1987, 180 m, J.G. Galván 201 (MEXU); Rancho La Peñita, 21 Sep 1982, 90 m, J.L. Ramos 83 (MEXU); Municipio de Gómez, La
Herradura, 7 Sep 1983, 400 m, R.A. Carranco 196 (MEXU); La Minita, 18 Aug 1983, 370 m, J.L. Ramos s.n. (MEXU); Municipio de Hi-
dalgo, Sierra La Lagunita, 6.4 mi SE of Aramberri on road towards Agua Fría, 19 Sep 2002, 1397 m, P.M. Peterson 16674, J. Valdés-Reyna
& M. Sosa (US);15 km al SE de Hidalgo rumbo al Barretal, Mar 1969, 230 m, F. González 2337 (MEXU); 18 km al SE de Hidalgo, G. Vil-
legas 72 (MEXU); Estación Adelaida, 1 Oct 1986, 200 m, J.F. Iribe 312 (MEXU); Estación La Cruz, 220 m, G. Villegas 7, López (MEXU);
Marroquín, 15 Aug 1980, 360 m, M.H. Cervera 15 (MEXU), ipii 15 Aug 1980, 360 m, M.H. Cervera 540 (MEXU); Oyama, 21 Oct
m 220 m, G. Miis 27 (MEXU); San Francisco, 21 Feb 1985, 280 m, R. Díaz 288 (UAT); Municipio de Jaumave, Altas Cumbres
aumave 29 jun 1985, 950 m, M. Martí 742 (UAT): EjidoI a Unión, 9 Oct 1985, J.F. Iribe 241 (MEXU);
Municipio de Jiménez, 1 mi from Santander Jiménez on the road to Padilla, 20 Oct 1959, M.C. Johnston 4395, J. Graham (MEXU); Car-
retera Padilla-Jiménez, 23 Nov 1981, 210 m, J.L. Ramos 44 (MEXU); Municipio de Llera, About 30 mi S of Ciudad Victoria on highway
85, toward Ciudad Mante, 28 3P 1976, 460 m, — oe isi bad CHAPA); Ejido Portes Gil, 8 Sep 1981, 180 m, J.A. Barrientos
93 (MEXU); L.H. Harvey & J. V Victoria-Llera, 23 Sep 1986, P. Moya151 (COCA, IEB); Municipio
El Mante, Between Cd. Mante and Tasnoio: 10 mi s. village of AE Trinidad, A.A. Beetle M-1220, M. May, Q. Skinner, A. Ibarra, J.
Bradley & F. i d iMd EL a ditches on south side of Cd. Mante, A.A. Beetle M-1187 (CHAPA); 10 mi al S de Cd. Mante, F.W.
Gould 9552 (US); ^ , Ejido Boreal Real, 25 Jul 1986, 750 m, J.G. Galván 118 (MEXU); km 5 hacia El Ejido Camarones,
25 Jul 1986, 750 m, 1G. Galván 127, 128 (MEXU); La Oveja (Venustiano Carranza), 19 Nov 1986, 1800 m, M. Cisneros 103 (MEXU);
Municipio de Méndez, San Tomás, 18 Sep 1980, 110 m, R.A. Carranco 17 (MEXU); Santo Tomás, 110 m, R.A. Carranco 630 (MEXU);
Municipio de Miquihuana, El Angelito, Colonia La Peña, 24 Nov 1981, 660 m JA A. Barrientos 1267 (MEXU); Municipio de La Tamauli-
E Vicinity of the San Miguel, 26 Jul 1930, B.H. Harris Sia (MEXU); Munici los, Sierra de Tamalave, 3 Aug 1981,
m, J.F. Iribe 69 (MEXU); Municipio de San Carlos, 10 lat Morales, 26 Jun 1994, M. Martinez & Diaz
E s 2695 (MEXU); Ejido el Corralejo, 21 Aug 1980, 320 m, J.F. Iribe 7 (MEXU); Ejido el Corralejo, Aug 1980, 270 m, G. Villegas 420
(MEXU); Ejido La Gavia, 3 Sep 1981, 300 m, J.A. Barrientos 73 (MEXU); Rancho Loma Atravesada, 25 Jun 1992, 380 m, J.F. Iribe 375
(MEXU); Sierra de San Carlos. La Morita, 26 Jun 1930, 533 m, B.H. Harris 1933 (MEXU); Municipio de San Nicolás, Entre San José del
Alto (Las Adjuntas) y el Palmar, 1 Mar 1970, 700 m, F. González 2781, V.M. Toledo & E. Martinez (MEXU); Rancho La Trinidad, 19 Oct
1983, 650 m, J.F. Iribe 202 (MEXU); Municipio de Soto la Marina, Ejido Benito Juárez, 3 Oct 1986, J.F. Iribe 322 (MEXU); Entre Lavade-
ros y la Piedra, 18 Sep 1981, 140 m, A. Brito 9 (MEXU); Tramo San José de las Rusias-Ejido 5 de Mayo, 2 Oct 1986, 80 m, J.G. Galván
156 (MEXU); Municipio de Tula, Ejido La Laguna, 6 Nov 1986, 1150 m, P. Moya 166 (MEXU); Municipio de Victoria, Camino al Ejido
La Reforma, 23 Oct 1986, 700 m, G. Bores 118 (MEXU); Camino al Molino, 3 Sep 1986, 500 m, R.A. Carranco 247 (MEXU); 18 Nov 1986,
1000 m, J.E. López de la Cruz 27 (MEXU); Cañon del Novillo. Río San Marcos, 23 Jun 1989, 440 m, A. Mora 857 (UAT); Municipio de
Victoria, Ejido Vicente Guerrero, 6 Sep 1985, 800 m, M.H. Cervera 276 (MEXU); Ex-Hacienda de Nogales, Aug 1986, 340 m, P. Moya
116, 117 (MEXU); Rancho El Abra, 18 Sep 1985, J.G. Galván 82 (MEXU); Municipio de Villagrán, 3 mi S of Nuevo León line on the
Villagrán-Linares highway, 12 Nov 1959, 457 m, M.C. Johnston 4660, J. Graham (MEXU); Ejido Candelario Reyes, 15 Oct 1985, 500 m,
J.F. Iribe 250 (MEXU); Ejido Guadalupe, 28 Jun 1983, 360 m, J.F. Iribe 170 (MEXU); Ejido Guadalupe, 22 Oct 1985, 380 m, J.F. Iribe 255
(MEXU); Garza Valdéz, 19 Nov 1986, 290 m, J.G. Galván 192 (MEXU); La Huasteca, 24 Jul 1983, 450 m, M. H. Cervera 172 (MEXU); La
Huasteca, 24 Aug 1983, 450 m, J.F. Iribe 174 Gee On ee 85, uio oe sae Ciudad Vicóna 111 km NO of Ciudad
Victoria, 24 Jun 1977, 415 m, T.B. Croat 39 AE? j Zapata, 4 Feb 1981, J.F. Iribe 51 (MEXU);
Jul 1979, A. Bolaños 7 (MEXU).
962 | Vof the Botanical R h Institute of Texas 2(2)
19. Bouteloua rigidiseta (Steud.) Hitchc., J. Wash. Acad. Sci. 23(10):453. 1933. (Fig. 25). Aegopogon rigidisetus
Steud., m Pl. Glumac. 1:146. 1854. Tiro: U.S.A. Texas: 1856, Drummond 374 (HoLoriPo: P; isorrros: US-879824 fragm. ex Pl,
exE
-106 Of
S ragm
Bouteloua texana S. Watson, Proc. Amer. Acad. Arts 18:196. 1883. Polyodon texanus (S. Watson) Nash, Fl. S.E. U.S. 138, 1327. 1903.
Tiro: U.S.A. Texas: San Antonio de Bejar, Berlandier 1535 (Lecroriro: GH, designado por Griffiths, Contr. U.S. Natl. Herb. 14:404.
1912).
Plantas perennes, amacolladas. Culmos de 15 a 4050) cm de alto, delgados, débiles, erectos, muy espat-
cidamente ramificados en la base. Hojas principalmente basales; vainas glabras a ligeramente escabrosas;
ligula de 0.1 a 0.5 mm de largo, un anillo corto de cilios; láminas foliares de 4 a 10-17) cm de largo y 1 a 2
mm de ancho, generalmente aplanadas, en ocasiones los ápices involutos, terminando en un ápice muy fino,
glabras a finamente papiloso-pilosas, pelos de 1 a 1.5 mm de largo. Inflorescencia de 3 a 6 cm de largo (solo
donde se insertan las ramificaciones), con 5 a 14 ramificaciones primarias espigadas caedizas, articulación
de las espigas es 0.5 mm por arriba de la base, dejando un pequeño trozo poe BODIE el eje principal;
ramificaciones de 4 a 7(-9) mm de largo, portando (293 a 5 espiguillas; raqui rígido, con
el ápice aristado, profundamente trifurcado a bifurcado, las aristas laterales antrorsas. Espiguillas de 7 a 10
de largo, incluyendo las aristas, flabeladas; glumas desiguales, primera gluma de 3 a 4 mm de largo, segunda
gluma de 6 a 8 mm de largo, diminuto plateado-pubescente en las nervaduras, lanceolada, acuminada, corto
aristada; lema de la flor perfecta de 4 a 5 mm de largo, sin incluir las aristas, esparcido-pubescente en las
nervaduras, terminando en 2 aristas laterales escabrosas, divergentes, de 2 a 3 mm de largo, y 1 central más
fuerte, de 3 a 4(-5) mm de largo; pálea de igual tamaño que el cuerpo de la lema, glabra, con el ápice bifido.
Anteras de 2.5 a 3 mm de largo, blanco-amarillentas. Flósculo rudimentario 12), el primero reducido a
una columna 3-aristada, las aristas de 5 a 10 mm de largo, aplanadas, fuertes, escabrosas, alternas con 2
lóbulos intermedios, de 3 a 4 mm de largo, corto aristados; segundo flósculo cuando presente, similar al
primero pero más pequeño. Cariopsis de 3.5 a 3.7 mm de largo, estrechamente ovobada, aplanada. 2n- 28,
35, 40 (para localidades de EUA). Ha sido colectado en diferentes meses a través del año.
Distribución.—SE de Estados Pii a o en Mine D anon A
Hábitat.—Pastizales y | arcillosos; 115 a 1400
O L ?
m.
dos. MEXICO: Tamaulipas: Municipio de González, 24 km al NO de González, 11 Jun nes 115 m, P. Pied
6 (MEXU): Municipio de Jiménez, Ejido Sor Juana Inés de la Cruz, 29 Sep 1993, 150 m, J.G. Galvan 338 (MEXU); ^
Marina, km 10 carretera a Soto la Marina-Abasolo, 10 Oct 1991, 60 m, O.M. Galván 503 (MEXU); Municipio de Vigra. El Madrofio,
17 Mar 1994, 1400 m, J.G. Galván 358 (MEXU); El Refugio, 17 Mar 1994, J.F. Iribe 429 (MEXU); Victoria-Altas Cumbres, 21 Jul 1993,
1200 m, J.A. Franco 139 (MEXU)
20. Bouteloua scorpioides Lag., Gen. Sp. Pl. 5. 1816. (Fig. 26). Actinochloa scorpioides (Lag.) Roem. & Schult., Syst.
Veg. 2:420. 1817. Atheropogon scorpiodes (Lag.) Spreng., Syst. Veg. 1:293. 1825. Chondrosum scorpioides (Lag.) Kunth, Revis. Gramin.
1:94. 1829. Tipo: MÉXICO: “Habitat in N. Hisp.,” Sessé s.n. (HoLoTIPO: MA?).
Plantas perennes, formando pequeños macollos o manchones. Culmos de 10 a 2030) cm de alto, delgados,
erectos, poco ramificados en la base, glabros a ligeramente puberulentos cerca de los nudos. Hojas princi-
palmente basales; vainas glabras, con márgenes hialino-pubescentes y un mechón de pelos en la garganta,
los pelos de 1.5 a 3 mm de largo; lígula de 0.5 a 1 mm de largo, ciliada; láminas foliares de 2.5 a 15 cm de
largo, involutas, curvadas o flexuosas, escabroso-pubescentes del lado adaxial. Inflorescencia corta, con un
eje de aproximadamente 1 mm de largo, portando una ramificación primaria espigada persistente; raquis
claramente arqueado, glabro a escabroso; ramificación de 2.5 a 4(-5) cm de largo, unilateral, inserta en
el ápice del culmo, en ocasiones presenta una bráctea estrecha hasta de 7 mm de largo, portando 35 a 90
espiguillas, distribuidas hasta el ápice del raquis. Espiguillas de 5 a 7 mm de largo, articulación por arriba
de las glumas; glumas subagudas, glabras a escabrosas, verdes o con tintes purpüreos, primera gluma de 2
a 3 mm de largo, angosta, segunda gluma de 4 a 7 mm de largo, glabra a escabrosa; lema de la flor perfecta
de 3.5 a 5.5 mm de largo, barbada en la base y densamente pilosa sobre los márgenes y nervaduras en la
mitad inferior, los nervios se prolongan en aristas fuertes de 1.5 a 2 mm de largo la central y 1 a 1.5 mm las
UU arr, Avriot t al., B tal dal ta da MA 963
AM
Cue v
1 dM
ES
Fic. 25 Dictrit .. la Dorrtal, > ne I ta da Mv
laterales. Anteras de 2.8 a 3 mm de largo, color anaranjado. Flósculos rudimentarios 2, el primero de 5.5 a 6
mm de largo, con 3 aristas de 1.5 a 4 mm de largo, con un mechón de pelos en la base; segundo flósculo muy
reducido. Cariopsis de 1.8 a 2.4 mm de largo, color amarillo a café, anguloso- triangular. 2n = 20. Florece
aparentemente de agosto a diciembre, aunque se le encuentra en otras épocas del año.
Distribución.—Especie mexicana distribuida del N al Centro del país.
Hábitat. —Laderas rocosas con pastizal y matorral xerófilo; 1750 a 2300 m.
Ejemplares inados. MÉXICO. Coahuila: Municipio de Monterrey, 25 mi SW de Monterrey, F.A. Warnock 14820 (US); Municipio
de Piedras Negras, Cerca de Díaz, F.A. Barkley 14820 (US). Nuevo León: Municipio de Aramberri, 16 km NE of Sandia on road to La
Ascension, 19 Sep 2001, 2000 m, P.M. Peterson 15829, J. Valdés-R (US); Municipio de D Arroyo, Doctor Arroyo, Nov 1980, 1750
m, J.A. Ochoa 1267 (MEXU); El Jarro, 16 km al O de Doctor Arroyo, en laderas calizas, Oct 1972, 1850 m, F. González 4956, M.A. Capo
& M. Sánchez (MEXU); 9.4 mi W of San Antonio de Peña Nevada & 0.4 mi E of jct of Hwy 2 to Or., 22 Sep 2002, 1807 m, P.M. Peterson
16794, J. Valdés-Reyna & M. Sosa (US); Municipio de Galeana, Región de Galeana, E. Hernández & V. Mathus N-2046 (CHAPA); 14 mi
al N de Matehuala, J.R. Reeder & C.G. Reeder 3661 (US); 8 km al ent jue de San Roberto, rumbo a Galeana, Y. Herrera Arrieta & A.
964
IR h Institute of Texas 2(2)
Fic. 26. Distril
Cortés 1448 (CIIDIR, HUAA); km 150, carr. 57 Matehuala-Saltillo, a 20 km de San Roberto, Y. Herrera Arrieta & A. Cortés 1450 (CIIDIR,
; km 162, carr. 57 Matehuala-Saltillo, Y. Herrera Arrieta & A. Cortés 1452 (CIIDIR, HUAA); 12 km E d t ue San Roberto
sobre la carretera 58 camino a Galeana, 5 Sep 1986, 2300 m, J. Valdés-Reyna 1719, R. Banda (ANSM, TEB); 12 km SE of
highway 58, 27 Aug 1981, 1845 m, S.L. Hatch 4579, C.W. Morden & J. Valdés-Reyna (ANSM); 12.3 mi S of Milagro, 18 Sep 2003, 2040 m,
P.M. Peterson 17809, J. Valdés-Reyna & H.S. Hinton (US); 2 km S del Salero, 1 km E de carretera 57 Matehuala-Saltillo, 3 Jul 1983, 2050 m,
J. Valdés-Reyna 1624, J.S. Marroquín (ANSM); Along highway 57, about 12 mi N of San Roberto, 4 Sep 1991, 1768 m, K.W. Allred 5512, J.
Valdés-Reyna & J.T. C i ANSM); Rancho Aguililla, 17 Sep 1989, 1950 m, G.B. Hinton 19740 (MEXU); Municipio de Tula, La Tapona,
21 Oct 1982, 1900 m, M.H. Cervera 134-A (MEXU).
21. Bouteloua simplex Lag., Varied. Ci. 4:141. 1805. (Fig. 27). Actinochloa simplex (Lag.) Roem. & Schult., Syst. Veg.
418. 1817. Chondrosum simplex (Lag.) Kunth, Revis. Gramin. 1:94. 1829. Tiro: PERU: Née s.n.?(HoLomiro: MA; 1soripos: M, LE-
t
San Roberto on
TRIN-2127.02!).
Chloris procumbens P. Durand, Quib. Chlor. Spec.16. 1808. Chondrosum procumbens (P. Durand) Desv., Nouv. Bull. Sci. Soc. Philom.
Paris 2:188. 1810. Chondrosum procumbens (P. Durand) Desv. ex P. Beauv., Ess. Agrostogr. 1:41, 158, t. 9, f. 7. 1812. Atheropogon
Arriat tsi D tol dal ta da kii
965
Fic 27 Distri cz da Darrfal O L ] ta da MZ
procumbens (P. Durand) J. Jacq., Ecl. Gram. Rar. 2:16, pl. 12. 1813. Actinochloa procumbens (P. Durand) Roem. & Schult., Syst. Veg.
2:417. 1817 don p I PD 1) Raspail, Ann. Sci. Nat. (Paris) 5:303. 1825. Bouteloua procumbens (P. Durand) Griffiths,
Contr. U.S. Natl. Herb. 14(3):364, f. 27. 1912. Tiro: PHILLIPINES: Cultivated at Madrid, probably from México or South America,
P Durand s.n. (nororiro: MPU-1469; isoripos: CIDIR ex MPU-1469 foto!, CIIDIR ex MPU-1468 foto!, L, MPU-1468, US-80856
fragm. ex MPU-1468)).
Bouteloua prostrata hort. ex Lag., Varied. Ci. 2(4):141. 1805. Actinochloa 1 hort. ex Lag.) Roem. & Schult., Syst. Veg. 2:419. 1817.
Chondrosum prostratum (Lag.) Sweet, Hort. Brit. 1:455. 1826. Tipo: MÉXICO: Cult. de la semilla recogida originalmente por L. Nee
en México, Lagasca s.n.(HoLoTIPO: MA; isoTipos: MA-7454, LE-TRIN 2126.31).
Chondrosum humile P. Beauv. ex Kunth, Nov. Gen. Sp. (quarto ed.) 1:175, t. 56. 1816. Actinochloa humilis (P. Beauv. ex Kunth) Willd. ex
Roem. & Schult., Syst. Veg. 2:417. 1817. Eutriana humilis (P. Beauv. ex Kunth) Trin., Gram. Unifl. Sesquifl. 239. 1824. Atheropogon
humilis (P. Beauv. ex Kunth) Spreng., Syst. Veg. 1:293. 1825. Bouteloua tenuis var. humilis (P. Beauv. ex Kunth) Griseb., Abh. Kónigl.
Ges. Wiss. Góttingen 24:303. 1879. Bouteloua humilis (P. Beauv. ex Kunth) Hieron., Bol. Acad. Nac. Ci. 4:495. 1882. Tiro: ECUA-
DOR: "Crescit in frigidi iccatis, argillosis propter 11 ga Quitensi i delupensi,” E Humboldt & A. Bonpland
s.n.(HOLOTIPO: P; IsoTIPO: US-A865683 fragm.!).
966 ] lof tha Botanical R h Institute of Texas 2(2)
hond PB Kunth, Nov. Gen. Sp. (quarto ed.) 1:176, t. 57. 1816. Actinochl (PB Kunth) Willd. ex Roem.
& Schult., PE d 2: e 1817. Eutri is (P. Beauv. ex Kunth) Trin., Gram. Unifl. Sesquifl. 240. 1824. Bouteloua tenuis (P.
Beat Abh. Konigl. Ges. Wiss. NM 19:259. 1874. Tipo: MÉXICO: "Novae Hispaniae prope México et San
in de las Cuevas,” E Humboldt & A. Bondpland s.n. (notorio: P).
s pusilla Vasey, Bull. Torrey Bot. Club 11:6. 1884. Tiro: U.S.A. New Mexico: Kingman, Jun 1881, G.R. Vasey s.n. (HoLoriPo: US-
81725).
Bouteloua ce Phil., Verz. Antofagasta Pfl. 85. 1891.—Tiro: CHILE: Tarapaca, E Philippi s.n. (noLoTIPo: SGO-PHIL-267; IsoTIPOS:
SG 9, SGO-63605, US-A865572 fragm. ex SGO-PHIL-267!).
m E Phil., Verz. Antofagasta Pfl. 85. 1891. Bouteloua simplex Lag. var. rahmeri (Phil.) Henrard, Meded. Rijks-Herb. 40:66.
921. Tipo: CHILE: Tarapacá, R.A. Philippi 268 (notorio: SGO-PHIL-268; isorieos: SGO-37548, SGO-63604; US-A865562 fragm.
Pen
Bouteloua simplex Lag. var. actinochloides Henrard, Med. Rijks-Herb 40:65. 1921. Tiro: BOLIVIA: Cochypata, 7 Nov 1857, 3000 m, G.
Mandon 908 (notorio: L-908.86-226; sorro: US-A865564 fragm.!).
Plantas anuales. Culmos de 7 a 23(-35) cm de alto, usualmente postrados en la mitad inferior, ascendentes
distalmente a ocasionalmente erectos, glabros. Vainas glabras, mucho más cortas que los entrenudos; lígula
de 0.2 a 0.4 mm de largo, membranácea, ciliada; láminas foliares de 2 a 7 cm de largo y 0.5 a 1.1 mm de
ancho, aplanadas en la base, involutas distalmente, glabras o escabroso-pubecentes en la superficie adaxial,
márgenes con algunos cilios arriba de la lígula. Inflorescencia una ramificación primaria espigada unilateral
solitaria, persistente, de 1 a 3 cm de largo, generalmente curveada, portando 20 a 4060) espiguillas pec-
tinadas. Espiguillas de 4 a 5 mm de largo, articulación por arriba de las glumas; raquilla con un mechón de
pelos debajo de la flor perfecta y del primer flósculo rudimentario; glumas glabras o escabrosas en la quilla,
agudas a acuminadas, primera gluma de 2 a 3 mm de largo, estrecha, casi transparente, segunda gluma de
3.5 a5 mm de us más ancha; lema de 3 a 5 mm de largo, densamente pubescente sobre los nervios, estos
se prolongan tas, fuertes, aplanadas y escabrosas, el nervio central lleva la arista más larga, hasta
de 2 mm de largo, que emerge entre 2 lóbulos membranáceos, de 1 mm de largo; pálea ancha, ligeramente
más pequeña que la lema. Anteras de 0.8 a 1 mm largo, color amarillo claro. Flósculo rudimentario 1-2),
el primero formado por una columna de 1 mm de largo y 3 aristas fuertes escabrosas, aplanadas, de 2 a 3
mm de largo; segundo flósculo cuando presente constituido por pequeñas escamas. Cariopsis de 0.7 a 1.2
mm de largo, color pajizo, anguloso-triangular. 2n = 20, 40. Florece de julio a diciembre.
Distribución. —Desde Estados Unidos a México en casi todos los Estados; Sudamérica: Argentina, Bolivia,
Chile, Ecuador y Perú.
Hábitat. —Planicies secas, laderas rocosas, lugares perturbados y matorrales bajos; 1800 a 2150 m.
Ejemplares examinados. MÉXICO. Coahuila: Municipio de Piedras Negras, E de Piedras Negras, M. Villegas 482 (IEB); Municipio de
Saltillo, Saltillo, E. Palmer 397 (US); 12 km N of the Coahuila-Zacatecas state line along highway 54 between Saltillo-Concepción del
Oro, Zacatecas, pee Ru p p m, = L. sns be C W. Morden & J. Valdés-Reyna (ANSM); Cañón de San Lorenzo, en la Sierra de
Zapalinamé, i idad Autónoma Agraria Antonio Narro, 6 Oct 1979, 2080 m, R. López s.n. (ANSM);
Rancho experimental Los RS 48 s S de Saltillo, carretera 54 Saltillo- -Concepción del Oro, Zac., 15 Oct 1982, 1800 m, F. Coss-V.
s.n. (ANSM); Rancho experimental Los Angeles, 48 km S de Saltillo, carretera an iid pias dea M Oro, Zac., 3 Sep 1978, 1800
m, J.S. Sierra s.n. (ANSM); San Lorenzo canyon, of highway 54 (road f Bod: metros pud first pump
station, 20 Sep 1988, 2000 m, M.E. Barkworth 5144, J. Valdés-Reyna & P.S. Hoge (ANSM); Si de Z ltillo, in San
Lorenzo canyon, 25 Aug 1981, 1800 m, S.L. Hatch 4483, C.W. Morden e. Valdes Reyna ( (ANSM); Municipio de ma Ventura, aprox. 80 km
S de Saltillo, 23 os ae 1800 m, J.A. Villarreal 3502, M.A. IBUG). Nuevo León: Municipi pde Dr. RUE 21 km al
N de Matehuala, P 312 (MEXU); Municipio de Gal 73 mi S de Saltillo FW. Gould 10112 (US); 6.5 Coahuila
de
and Nuevo León on Hwy 57 towards A 16 Oct 1995, 1920 m, P.M. Peterson 13281, M.B. Knowles (US); M
21 Sep 2001, 1930 m, P.M. Peterson 15866, J. Valdés-Reyna (US). Tamaulipas Municipi
de POARTE, Joya de Herrera, 20 Oct 1982, 2150 m, C.R. López 45 (MEXU).
22. Bouteloua tamaulipensis G.J. Pierce ex D. Pacheco & Columbus, sp. nov. (Figs. 28-31). Cathestecum
tamaulipense Beetle, Las Gramíneas de México 2:177. 1987, nom. nud. Tipo: MÉXICO. TamauLiras: Municipio de Jiménez: on mesa
ca. 7.5 road mi NNE of Santander Jiménez along Hwy 101, 24?1857"N, 98?26'16"W, elev. ca. 200 m, 22 Sep 1996, J.T. Columbus
2885 (notorio: RSA!; isoripos: ANSM!, MEXU!, MO!, US!).
Boutelouae erectae (Vasey & Hack.) Columbus affinis, sed diff. i insigni
antheris brevioribus.
967
Plantas perennes, cespitosas, estoloníferas, políga-
mo-monoicas. Ent dos del tol de 4a 14
cm de largo, arqueados ligeramente, de color verde
a púrpura, nudos densamente pilosos. Culmos de
2 a 22 cm de alto, rectos y erectos o geniculados y
ascendentes, con 0 a 2 nudos vegetativos visibles, a
menudo con una rama en el nudo superior, nudos
usualmente pubescentes. Hojas principalmente
basales; vainas con un margen escarioso, ciliadas,
los pelos más largos en el ápice o restringidos a él
(a los lados de la lígula), vaina de las hojas jóvenes
sobre estolones abaxialmente pilosos, la mayoría
cerca de la base; lígula compuesta de pelos de
cerca de 0.2 mm de largo, láminas de hasta 10 cm
de largo y 1 mm de ancho, aplanadas o involutas
cuando secas, por lo general glabras abaxialmen-
te, adaxialmente con pelos largos espaciados a lo
largo de la nervadura central, margen escabroso.
Inflorescencia incluida dentro del follaje o alta y
bien exserta, compuesta de 3 a 8 ramas primarias
Fic. 29. Infl ia de Bi ji li; is (4.T. Columbus 2885)
Barra de escala — 0.5 cm.
968 J | of the Botanical R h Institute of Texas 2(2)
dísticas, con una inflorescencia lateral a menudo originándose desde el nudo superior; raquis de 0.5 a 3 cm
de largo, aplanado, ligeramente sinuoso, glabro, el ápice cortamente bífido, entrenudos de 0.2 a 0.6 mm de
"n ramas de 4 (en inflorescencias cortas) a 6 (en inflorescencias largas) mm de largo, péndulas, cada una
conteniend tadas, con las primeras glumas afrontadas, púrpuras o con tintes purpüreos
(glumas, icis: y/o aude d eje cerca de 1 mm de largo, no prolongado mas allá de la espiguilla terminal,
desarticulándose cerca de la parte media en la madurez, puberulento debajo del punto de abscisión, densa-
mente piloso por arriba de él, las espiguillas caen en grupo. Espiguillas de una rama dimórficas: espiguilla
terminal (central) más larga (de 3.7 a 5.2 mm de largo), primera gluma más corta, flósculos 3 ó 4, primer
flósculo hermafrodita (inflorescencias altas) o pistilado (inflorescencias cortas; estaminodios usualmente
presentes), los flósculos restantes estaminados o neutros, espiguilla proximal (lateral) más corta (de 1.5 a
3.8 mm de largo), primera gluma más larga, flósculos 2 6 3, neutros (inflorescencias cortas) o los primeros
uno a dos flósculos estaminados (inflorescencias altas). Glumas variables en tamaño, forma e indumento;
primera gluma sin nervaduras, glabra (raramente pubescente en las espiguillas laterales), algunas veces 1- 6
2-lobadas, primera gluma de la espiguilla terminal de 0.4 a 0.6 mm de largo y típicamente flabelada, trun-
cada y sin arista, primera gluma de las espiguillas laterales de 0.6 a 1.5 mm de largo, típicamente lanceolada
y aristada en el ápice; segunda gluma de 1.3 a 3.5 mm de largo, por lo general ampliamente lanceolada,
1-nervada, pilosa (densamente pilosa en mitad inferior a lo largo de la nervadura central), ápice terminando
en una pequeña arista o acuminado, algunas veces ligeramente 2-lobado. Flósculo superior algunas veces
rudimentario, de menos de 1.5 mm de largo y sin arista. Lema 3-nervada, 2- ó 4-lobada, usualmente con
los ápices obtusos o agudos, 3-aristada desde las nervaduras, la arista central originándose entre dos lobos,
las aristas laterales iguales a o ligeramente más largas que la arista central; lema del primer flósculo de 1.6
(espiguillas laterales) a 3.5 (espiguilla terminal) mm de largo, 2-lobada cerca de 1/3 del largo del cuerpo
de la lema (algunas veces dos lóbulos laterales también presentes), por lo general esparcidamente pilosa a
través de la parte central del dorso por debajo de los lobos, los pelos algunas veces limitados a cerca del
margen, aristas de 0.6 a 1.3 mm de largo, iguales a o excediendo el cuerpo de la lema por 0.3 mm o menos,
escabrosas; lema de los flósculos superiores (excluyendo la flor rudimentaria, si está presente) de 1.1 (espi-
guillas laterales) a 3.7 (espiguilla terminal) mm de largo, 4-lobada cerca de 2/3 o más del largo del cuerpo
de la lema, el cuerpo glabro o con algunos pelos cerca de la base de los lobos laterales, aristas de 0.3 a 3.0
mm de largo, usualmente excediendo el cuerpo de la lema por 0.1 a 1.3 mm, pilosas sobre la mitad inferior
y escabrosas en la mitad superior o completamente escabrosas. Pálea de 0.7 a 3.5 mm de largo, más larga
en la primera flor de la espiguilla terminal, en el p uc: casi niu en longitud a (y algunas veces
era excediendo) la lema, prog relación a la lema,
usualm con dos o más lóbulos en el ápice, por lo mu ker deu vis sobre los 2/3 superiores
a lo largo de las dos nervaduras, las nervaduras usualmente extendiéndose en aristas cortas de hasta 0.6
mm de largo. Anteras usualmente de menos de 1.5 mm de largo, con tintes purpúreos; estigmas púrpura
obscuro. Cariopsis no observada. 2n - 20 (Pierce 1979; Pacheco 2002).
Comentarios.—Bouteloua tamaulipensis está más cercanamente relacionada a ejemplares robustos de
B. erecta, taxon que se distribuye en el Desierto Chihuahuense hacia el noroeste de la distribución de B.
tamaulipensis (Pierce 1979; Pacheco 2002; J.T. Columbus inéd.).
Distribución.—Endémica de Tamaulipas, México. Conocida sólo de un área limitada al noreste de
Santander Jiménez.
Hábitat. —En claros de matorral xerófilo con árboles bajos y en áreas de perturbadas que son resultado
de la remoción de la vegetación leñosa; suelo rocoso calcáreo; cerca de 200 m de altitud.
€ 1 n E EE a
Ejemplares adicionales examinados. MÉXICO. Tamaulipas: Municipio de Jiménez
Fernando, 19 Oct 1956, F. Martinez Martínez & G. Borja Luyando F-2451 (MICH, TEX, US); 11 mi N a Santander e iih e
Fernando, 800 ft, 25 Nov 1959, J. Graham & M.C. Johnston 4828 (MICH, TEX); Bis mi d i y t,
s 1959, M.C. rd 4896 ne TEX); 7 km from S ler Ji do, 23 Oct 1975, GJ. Pierce 2339 (RM); 11
Fernando, roadside, 23 Oct 1975, GJ. Pierce 2340 (RM); 18 km from Santander Jimenez toward
Larrara Arriata
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San Fernando, 23 Oct 1975, G.J. Pi 2341 (RM); 20 km f S J San F do, roadside, 23 Oct 1975, G.J. Pierce
2343 (RM); El Encinal, 2 Sep 1981, Villegas 468 (MEXU); El Encinal, 23 Nov 1981, Brito 056 (MEXU); 5 mi NNE of Santander Jiménez
just off Hwy 101 along road E to La Misión, 24?1712"N, 98?27720"W, ca. 200 m, 2 Nov 1992, J.T. Columbus 2426 (ANSM, MEXU, MO,
RSA, UC, US); Ejido Sor Juana Inés de la Cruz, 190 m, 28 Sep 1993, Galván 319 (MEXU); a 8 km por la carretera 101 al N del pobiado
de Santander Jiménez, M.H. Cervera 403 (MEXU); S ler Jiménez-San Fernando, 15 km al N de Santander Jiménez, aprox.
200 m de la entrada al Ejido El Encinal, 24%23.867'N, 98?23.300'O, 150 m, 12 Sep 1999, D. Pacheco & J.O. Zambrano 2552 (HERZU,
IZTA, MEXU); Santander Jiménez, Aldama cenotes, J. Valdés-Reyna 2402 (ANSM).
23. Bouteloua trifida Thurb., Proc. Amer. Acad. Arts 18:177. 1883. (Fig. 32). Chondrosum trifidum (Thurb.) Clayton,
Kew Bull. 37:418. 1982. Tiro: MÉXICO. Coauuma: Monclova, Aug 1880, E. Palmer 1355 (HoLoTIPo: GH; isoripos: NY, US-A865565!,
US-821003!, US-883967,! US-2474825 foto!).
Bouteloua burkii Scribn. ex S. Watson, Proc. Amer. Acad. Arts 18:179. 1883. Bouteloua trifida Thurb. var. burkii (Scribn. ex S. Watson)
Vasey ex LH. Dewey, Contr. U.S. Natl. Herb. 2(3):532. 1894. Treo: U.S.A. Texas: west Texas, upper Concho, sandy plains, Apr
Harrara Arriat tal., B tal Aal ta da T PAR- 971
1882, J. Reverchon s.n.-3440 Curtiss’ distribution of North American plants (Lecroriro, designado aquí: US-883346!; ISOLECTOTIPO:
US-820915
Chondrosum trinii E. Fourn., Mexic. Pl. 2:136. 1886 teloua t rn.) Griffiths, Contr. U.S. Natl. Herb. 14(3):387, f. 41, 42, pl.
74. 1912. Tiro: U.S.A. Texas. Webb Co.: Laredo, Berlandier e otro: P; isoripos: K, US-A865684!, US-204622!, US-10845181,
US-883356!).
Chondrosum virletii E. Fourn., Mexic. Pl. 2:136. 1886. Tiro: México. San Luts Potosi: Aug 1857, Virlet 1373 (aotomipo: P; isoripo: US-
A865685 fragm.!).
Plantas perennes, amacolladas, con base rizomatosa. Culmos de 10 a 30(-40) cm de alto, delgados, finamente
escabrosos, no-ramificados excepto en la base rizomatosa; entrenudos glabros. Vainas mucho más cortas que
los entrenudos, finamente escabrosas; lígula de 0.2 a 0.6 mm de largo, un anillo de cilios; láminas foliares
de 2 a 8 cm de largo y 0.5 a 1.2 mm de ancho, principalmente basales, aplanadas, conduplicadas o involu-
tas, cuando involutas con el ápice puntiagudo, escabroso-puberulentas principalmente hacia la base y en
la superficie adaxial, con algunos pelos dispersos sobre el margen, de 1.5 mm de largo. Inflorescencia de 1
a 5(-8) cm de largo, con 3 a 7 ramificaciones primarias espigadas persistentes, espaciadas 1.2 a 1.8 (2) cm
de distancia, sobre un raquis recto, nunca en zigzag; ramificaciones de 1 a 3(-4) cm de largo y 3a 5 mm de
ancho, portando 8 a 24(-32) espiguillas, color verde-amarillento o con tintes purpúreos. Espiguillas de 5.8
a9 mm de largo, articulación por arriba de las glumas; glumas desiguales, muy delgadas, casi transparentes,
agudas, acuminadas o mucronadas, con el ápice bífido, primera gluma 3 a 4 mm de largo, segunda gluma
de 5 a 5.5 mm de largo, lisa, glabra, a menudo color púrpura-rojizo; lema de la flor perfecta de 1.2 a 2.2 mm
de largo, glabra, con 3 aristas de 4.5 a 5-6) mm de largo, aplanadas, redondeadas y pubescentes hacia la
base, escabrosas hacia el ápice; pálea de 1.2 a 2.2 mm de largo, con el ápice obtuso. Anteras de 0.5 a 0.6 mm
largo, color blanco amarillentas. Flósculo rudimentario 1, reuducido a una columna corta, de 1.5 a 2.5 mm
de largo y 3 aristas de 3.5 a 6 mm largo. Cariopsis de 0.7 a 1 mm de largo, color amarillo, oblongo-elipsoide.
2n = 20. Florece de mayo a diciembre.
Distribución. —Centro y S de Estados Unidos al N de México.
Hábitat.—Se la encuentra sobre planicies secas, laderas con pastizal y bosque de encino-pino, sobre
diversos tipos de material; 60 a 2000 m
Ejemplares examinados. MÉXICO. Coahuila: 2 mi NW de la Frontera, I.M. Johnston 7171 (US); Municipio de Allende, 13 mi SO of
Ciudad Allende toward Sabinas, 7 Oct 1959, 488 m, M.C. o 4177, d bini (MEXU); Municipio de Arteaga, Arroyo Canoas, 28
Jul 1993, 1600 m, F. Alcaláá 22 (MEXU?; Municipio de Castaños Santa Teresa, al S de Castaños, 1936 (MEXU); Rancho de
Santa Teresa, al S de Castaños, 19 Jun 1936, 910 m, L.F. pos 205, C.H. Muller URN Municipio de Cuatrociénegas, 5 km al E de
Cuatrociénegas, 24 May 1977, 740 m, J. apis SCA Reyna 967 (ANSM); Entrada al Casco del Rancho Potrero de Menchaca, 30 Aug
1980, 1600 m, R. Vásquez 58, L.E. Rodrí, NSM); Sierra de la Madera, Cañón Charreteras, rancho Charreteras, 15 Sep 1993, 1750
m, J.A. Villarreal 7351, M.A. Carranza & H Rodríguez ee O p d 54.8 m N of Monclava on Mex 57 at jct of road
to Progreso, 15 Oct 1989, 480 m, P.M. Peterson 8379, J. Valdés-Reyna 1 Cepeda, Ejido La D carretera
40 Saltillo-Torreón, aprox. 20 km NE de General C Piin 5 Jul 1986, 1230 m, S. Vásquez 98, A. García (ANSM); Terren
entre Hacienda Rosa y Hacienda Lechuguilla Hipolito, 1936 (MEXU); Municipio de Hidalgo, Rancho Santa Barbara, 28 Nov 1989, 190
m, J.A. García (UAT); Municipio de Monclova, Puerto Santa Ana, 27 Jul 1973, 1000 m, J.S. Marroquín 2458 (ANSM); 22 mi N de Mon-
clova, J.R. Reeder & C.G. Reeder 5010 (US); 43 mi N of Monclova, F.W. Gould 6410 (ANSM); km 25 carr. Saltillo-Monclova, M. Tórres 1593
(ANSM); 2 km al N de Cañada Blanca, km 17 carr. 57 Saltillo-Monclova, Y. Herrera Arrieta, M. de la Cerda 6 O. Rosales nie 1483 (CIIDIR,
HUAA); Municipio de Müzquiz, 7.2 mi SE of Jaboncillos on road towards Cuesta de malena and Melchor Muzquiz 2006, 1010
m, P.M. Peterson 19847, S. Lara- Contreras (US); Rancho “Las Pilas," Müzquis, E. García s.n. nud ño L.M. ue 16853 (IBUG);
81 lau, camino a Barroteras, Müzquiz, J. Valdés-Reyna VR-939 (ASNM, IEB); ^ N. área de encinos, aprox. 10
km O de la cabecera municipal rumbo a Nava, 13 Jul 1988, 300 m, A. Rodríguez 1012, 1032, J. Valdés- -Reyna & M. Solís (ANSM); Level
plains of the Río Grande Valley, just N of Río Escondido, 18 kmSSO of Piedras Negras on México No. 57, 12 Sep 1978, 300 m, H.H. Iltis
10, A. Lasseigne (COCA, MEXU, IBUG); Valle de Río grande al N del Río Escondido, 18 km SSW de Piedras eG d H.H. Iltis 15222 A.
a (UAG); Municipio de Ocampo, Base of hills in palma-belt, near mouth of Cañón de la Cruz, about 16 Sep
1, LM. Johnston 9179 (MEXU); S m Laguna de La Leche, 30 Aug 1941, I.M. Johnston 8619 (MEXU); Sierra de la Madera, R25 Io La-
guna de la Leche, aprox. 62 km de bo a Sierra Mojada, 2 Oct 1990, 1000 m, M.A. Carranza 621, F J. Carranza (ANSM);
Municipio de Ramos Árizpe, 63 mi 5 of Monclova on highway 57, 9 Aug 1959, 396 m, T. R. Soderstrom 361 (MEXU, US); El Cedral, Si-
erra de la Paila, 16 Oct 1986, 1300 m, J.A. Villarreal 3628, M.A. Carranza & R. Rodríguez (ANSM); 2 Nov 1988, J.A. Villarreal 4812, M.A.
Carranza (ANSM, IEB); Estación La Sauceda, 20 Sep 1964, 1030 m, J.S. Marroquín (ANSM); Estación Paredón, 5 Sep 1987, 1400 m, A.
972 | Inf the Botanical R h Institute of Texas 2(2)
Rodríguez 905, M.A. Carranza & J.M. Sosa (ANSM); Ramos Arizpe, 21 Sep 1975, 1650 m, R. Palomo s.n. (ANSM); San Gregorio, carretera
Monterrey-Saltillo, 16 Sep 1983, 1350 m, J.A. Villarreal 2191 (ANSM, CIIDIR); San Juan de las Bonitas, 19 Jul 1993, 1450 m, E. Pérez 25
(MEXU); Sierra de la Paila, Ejido el Cedral por el ino El Carmen, 26 Aug 1992, 1300 m, J. Valdés-Reyna 2180 (ANSM); km10 camino
Saltillo-Gral. Cepeda, W de Saltillo, J. Espinoza 135 (ANSM); Ejido La Rosa, carr. 40, Saltillo-Torreón, aprox. 20 km al NE de Gral.
Cepeda, S. Vázquez & A. Garcia 98 (ANSM); km 30, carr. 40 Saltillo-Monterrey, Ramos Arizpe, Y. H Arrieta & A. Cortés 1455 (CIIDIR,
HUAA); Cerro del Pueblo, al W de la cd. de Saltillo, 16 Jul 1983, 1400 m, J. Valdés-Reyna VR-1847, M.Á. Carranza (ANSM, CHDIR); 11
kmW of Saltillo, G. Mick & K. Roe 17 (MEXU); km13 al W de Saltillo, E. Hernández & V. Mathus 2022 (CHAPA); 12 km O de Saltillo,
carretera 40 camino a La estación de Mi las Vega, 9 Jul 1983, 2000 m, J. Valdés-Reyna 1530, L.E. Rodríguez (ANSM); Municipio de
Sabinas, 85.4 km NW of Muzquiz on Hwy 53 towards Boquilla del Carmen, vía La Rosita, 12 Sep 1991, 810 m, P.M. Peterson 10570, C.R.
Annable (US); Ejido Agujita, Mar 1977, 350 m, J.F. Sifuentes s.n. (ANSM); Municipio de Saltillo, 28.2 mi S de Saltillo, carretera 54 a Con-
cepción del Oro, Zac., 24 Sep 1990, 1850 m, P.M. Peterson 10014, C.R. Annable & J. Valdés-Reyna (ANSM, US); Rancho experimental Los
Angeles, 48 kmS de Saltillo, carretera 54 Saltillo- Concepción del Oro, Zac., 30 Jul 1979, 1800 m, J.S. Marroquín (ANSM); Rancho Los
Angeles, 48 km al S de Saltillo, P.E. García s.n. (ANSM); N of Saltillo on Mex 57, N of Lázaro Pass, and 0.5 km S of La Muralla Restaurant,
13 Oct 1989, 1150 m, P.M. Peterson 8353 & J. Valdés-Reyna (US). N León: Municipio de Anáhuac, 34 mi 5 f Laredo, 14 Nov
7
zh
Darrara Aeviats at al ; D kal tal + R | |? PA 973
Nai. R.C. =D 5812, R.M. Tryon (MEXU); Municipio de Apodaca, Carretera Monterrey-Miguel Alemán, 28 May 1972, 430 m, R.
Almeida 2 AT); Municipio de Bustamante, En a eid de Bustamante, 18 Jun 1993, 600 m, P. Jauregui 67 (MEXU); Municipio de
Cadereyta Jiménez, C d 26 Sep 1979, 330 m, J.A. Villarreal (ANSM); En los alrededores
del poblado de Cadereyta, 22 Mar 1993, N. Bazaldua 18 (MEXU); Raco San Angel, km 7 camino a San Mateo, carretera Villa Juárez-
Cadereyta, 25 Mar 1993, M.M. Castillo 47 (MEXU); Municipio de Cerralvo, 8.5 mi E of highway 54 going to Rancho el 30, Sierra de
Picachos near Cerralvo, 16 Aug 1983, 700 m, J. Kessler 4859 (ANSM); Municipio de China, 1 km N del rancho La Osca, 23 Mar 1972,
160 m, J. Valdés-Reyna s.n. (ANSM); Presa La Ceja, carretera 40, 13 Sep 1992, 100 m, J.A. Villarreal 6843, M. Vásquez, D.E. Lozano & J.
Wand -Reyna (ANSM); Municipio de Doctor González, 1 km al SE del poblado de Doctor González, rumbo a los fresnos a la orilla del río
or González, 19 May 1993, 360 m, N. Bazaldua 68 (MEXU); Carretera Doctor Gonzalez-Cerralvo, km 52, 19 T 1993, 400 m, N.
a. 81 (MEXU); Municipio de Galeana, Cerca de Galeana, Oct 1980, 1650 m, J.A. Ochoa 1102 (MEXU); M 3km
from El Milagro and 10 km from Icamole on > road between, 5 Jul 1976, 720 m, M.C. Johnston 11615%, T.L. p & F. Chang
(MEXU); Límites de Coahuila-Nuevo I p a Saltillo, Nov 1980, 1000 m, J.A. Ochoa 1213 (MEXU); Muni
eral Zaragoza, 9 km S of Aramberri on road towards Zaragoza, 20 Sep 2001, 1400 m, P.M. Peterson 15836, J. Valdés- rin (US): Muni-
cipio de Higueras, Falda de la Loma Higueras frente a las instalaciones de la Unión Ganadera de Higueras, 1 Jul 1993, P. Jauregui 84
(MEXU); Municipio de a Carretera Villa de Juárez-San Mateo, km 5, 18 May 1993, M.M. Castillo 69, 76 (MEXU); San Roque, 11 Sep
1976, 500 m, LA. Jiménez s.n. (ANSM); Municipio de bake de Naranjo, Lomas El Tanque, 16 May 1982, 380 m, O.L. Briones 949
(ANSM); Rancho Santa Elena, carretera Lampazos-Horcones, 5 km de terracería al N, 18 Oct 1992, J. Garza 9 (MEXU); Lampazos, Dic
1980, J.A. Ochoa 1233 (MEXU); Municipio de Linares, 11 mi S of Linares toward Villagrán, 12 Nov 1959, 457 m, M.C. Johnston 4653
(MEXU); Ejido el Porvenir, 19 May 1980, 200 m, JJ. roi s.n. ip edidi p wen en el rancho El Nogalar, ubicado en la carretera
m12, 2 Jun 1993, 440 m, M.M La Escondida, km 8, 19 Jun 1992, P. Jauregui 3 (MEXU);
Linares-San R
Rancho San José de los Hoyos, km 15 carretera Linares-Iturbide, 1 Abr 1993 , J. Garza 22 (MEXU); Municipio de Los Ramones, 2 km S
del entronque a la carretera Monterrey-Reynosa, 26 Nov 1980, 210 m, J.A. Villarreal 1019 a Municipio de Marín, Facultad de
ON Univ a Autónoma de Nuevo León, km 17, Marín, 14 Jun 1992, M.M. Castillo 5 (MEXU); Facultad de Agronomía, Uni-
versidad A N León, km 17, Marín, 15 Nov 1992, P. Jauregui 31 (MEXU); Municipio de Mina, Predio 'El Venado, 12 Jun
1976, 600 m, G. Villegas 2, A. Bolaños (MEXU); Municipio de Monterrey, Sierra Madre mountains, 3 Aug 1933, C.H. Mueller 421, M.T.
Mueller (MEXU); Valley of Monterrey, 13 Jul 1889, C.G. Pringle 2671 (MEXU); Obispado, J. Roybal 63 (US); Carr. Villa de Garcia a Grutas,
W de Monterrey, T. Tateoka 1086 (US); I. Cabral 123 (ANSM); Aprox. 30 km de Monterrey a Laredo, 1 km al E de la carr., J. Ortíz s.n.
(CHAPA); Carretera a Trinidad China, Dic 1980, J.A. Ochoa 1226, 1227 (MEXUJ; Dry hill d , Monterrey, 5 Jul 1988, C.G. Prin-
gle 1974 (MEXU, US); E 30 km de Monterrey a Laredo, 1 km al E de la carr., J. Cantú s.n. oe Muncie de Pesqueria, Car-
retera M P 2 km antes del poblado, 19 Nov 1986, 350 m, P.A. García 1847 (MEXU); N pio de Sabi Hidalgo, 5
2
Sabinas Hidal ] M y, 16 Jul 1955, H.M. Chute 272 MERU Fracción del Río Sabinas, junto al
ativo Ojo de Agua de SER idalga: 24 Mar 1993, N. Bazaldua 46 (MEXU); Municipio de Salinas Victoria, Planos Rancho La Barrosa,
May 1981, 550 m, J.A. Ochoa 1170 (MEXU); Rancho La Soledad, May 1981, 500 m, J.A. Ochoa 1109 (MEXU); Sierra de Mamulique, 29
Ene 1993, M.M. Castillo 40 pA aic de Santa Catarina, 17 mi E of Monterrey, 1 Jul 1947, F.A. Barkley 7161, G.L. Webster &
C.M. Rowell Jr. (MEXU); Muni de Santiago, San Juan Bautista, 28 Nov 1982, 1700 m, I. Cabral 123 (ANSM); Villa de Santiago, Dic
1980, 550 m, J.A. Ochoa 1152 (MEXU); Municipio de Villaldama, Mountains E of state highway 34, 1.4 km (0.9 mi) N of jct to Busta-
mante, dirt road 15.7 km (9.8 mi) E of highway, 15 Mar 1983, 420 m, C.P. Cowan 3823, K.C. Nixon (ANSM). Tamaulipas: Municipio de
solo, 2 km antes de llegar al ejido Morelos, 24 Ene1985, 40 m, R. Díaz 269 (UAT); Ejido La Esperanza, 23 Sep 1982, 110 m, J.F. Iribe
147 (MEXU); La Cascada, 16 Oct 1982, R.A. Carranco 282 (MEXU); Municipio de Antiguo Morelos, Ejido El Sauz, 18 Aug 1981, 160 m,
R.A. Carranco 34 (COCA, IEB); Municipio de Burgos, Rancho Tanques, Aug 1980, 250 m, J.F. Iribe 23 (MEXU); Municipio de Busta-
mante, Ejido Calabacillas, 19 Mar 1987, 1200 m, M. Cisneros 116 (MEXU); Municipio de Camargo, Carretera la Rivereña, a orillas del
camino adyacente al rea agrícola, 24 Oct 1980, J.F. Iribe s.n. (MEXU); Municipio de Casas, 2 km antes de Villa de Casas, 4 Jun 1987, 310
m, C.R. López 143, 144 (MEXU); 35 km from Victoria on the road to Casas and Soto la Marina, 3 Oct 1956, 280 m, F. Martínez 2300,
2349, G. Borja no Camino a la Lajita,18 Mar 1994, J.G. Galván 400 (MEXU); Rumbo Ejido 1° de Abril, 28 Jul 1983, 260 m, J.L.
Ramos 147 (MEXU); N de Gomez Farías, La Poza Azul, nacimiento del río Frío. Reserva de la Biosfera El Cielo, 25 Aug 1993,
300 m, J.L. Mora 428, A. Ma & L. Corral (UAT); Municipio de González, 19 km from Mante on the Tampico highway, 18 Sep 1956, F.
Martinez 1710, G. Borja (MEXU); 24.5 km al E de Xicotencatl, 11 Jun 1992, 170 m, J.G. Galván 269 (MEXU); Ejido Gustavo A. Madero,
14 May 1981, 80 m, J.A. Barrientos 19 (MEXU); Ejido Josefa Ortíz de Dominguez, Sep 1983, 380 m, P. Moya 35, 152 (MEXU); km 56
González-Llera, 14 Oct 1987, 250 m, J.G. Galván 215 ae Se vee 3 Mar 1987, 180 m, J.G. Galvan 202 (MEXU); Rancho
La Peñita, 21 Sep 1982, 90 m, J.L. Ramos 78 (MEXU); ^ 5mez, Ejido Acatlán, 5 Oct 1983, 180 m, P. Moya 40 (MEXU); km
26, Rancho el Soldado, 28 Sep 1993, 150 m lván 606 (MEXU); Rancho El Melón, 8 Sep 1983, 150 m, M.H. Cervera 187 (MEXU);
Rancho El Melón, 5 Sep 1987, 150 m, pu Galván 212 (MEXU); Municipio de Hidalgo, md El n 5 Nov 1985, J.F. Iribe 257
(MEXU); Estación La Cruz, Hidalgo, D. Villegas 011 (IBUG); E de e aUa Po d 9 Oct 1982, 630 m, P.
Moya 9 (MEXU); Ejido José Maria Morelos, 7 Oct 1982, 690 m, J.F. Antonio km132 carretera Victoria-Jaumave,
25 Abr 1985, 1280 m, M. Martínez & Diaz de Salas 285 (MEXU, UAT); Ejido San F i de Zorrilla, 11 Jun 1979, 700 m, A. Bolafios
6, 499 (MEXU); Municipio de Jiménez, 2 km del Rancho El Rivereño, 4 Dic 1986, 210 m, M.H. Cervera 400 (MEXU); 8 km al N del Po-
blado Santander Jiménez, 5 Dic 1986, 200 m, M. Cisneros 111 (MEXU); Adelante de la parida, G. Villegas 999 (IEB); Municipio de Llera,
974 Journal of the Botanica! R h Institute of Texas 2(2)
Ejido lø de Abril, 27 May 1982, 230 m, R.A. Carranco 121, 122 (COCA, IEB), Ejido 1g de Abril, 19 Abr 1982, 250 m, J.F. Iribe 115 (MEXU);
Ejido Portes Gil, J.F Iribe 95 (MEXU); Estación Forlón, Sep 1985, C.R. López 124 (MEXU); Estación San Francisco, Sep 1985, J.L. Ramos
184 (MEXU); Llera-Río guayalejo, 1 Jun 1992, 280 m J E: e de la nee ^d o Ph of Cancer on highway 85 at km 660, 3
Oct 1965, 300 m, K. Roe 2401, E. Roe (MEXU); Munici , 10 km al SE del Rancho El Alteño,
ilez 585 (MEXU); Isla Las Gallinas, frente al po pesq La Capilla. Laguna Madre, 10 Sep 1994, 10 m, J.L. Mora
478 (1 IAD: El Soldado, Matamoros, A. Brito s.n. (IBUG); Muni de Méndez, San Tomás, 19 Sep 1980, 110 m, J.A. Barrientos 633 (COCA,
MEXU); Municipio de Miguel Alemán, Brecha de Bener alt. 95, 25 Oct 1991, C.R. López 200 (MEXU); Tamaulipas, 10 mi E of Linares,
8 Aug 1930, H.H. Bartlett 10844 (MEXU); Municipio de Nuevo Laredo, 20 km al O de Ciudad Guerrero, Oct 1973, F. González 6324, R
M. López & R. ilius (MEXU); Municipio de Padilla, Campo Turístico Lago Vista, 13 Feb 1986, P. Moya 89 (MEXUJ; Municipio de
San Carlos, 1.5 km SE de Guayacán, Sierra San Carlos, 7 Dic 1984, 590 m, O.L. Briones 1372 (ANSM); Ejido el Corralejo, 21 Aug 1980,
m, R. Sandoval s (MEXU); Ejido La Gavia, 3 Sep 1981, 300 m v ee m aes cave Puerto Rico, 3 T 1981, 200 m, J.A.
Barrientos 71 (MEXU); Rancho El Coyote, 19 Oct 1983, 300 m, J.F. I Miguel2 hez, Mar 1970,
420 m, F. González 2890, V.M. Toledo & E. Martínez (MEXU); Santa Teresa, 4 Oct 1981, 330 m, J.A. Barrientos 76 (MEXU); N de San
Fernándo, J.N. Rose 24336 (US); San Fernando, 10 mi from the San Fernando- Mee ione ad Dc on the E road to Rey-
nosa (junction of the dirt road to Mendez), 13 Nov 1959, 152 m, M.C. Johnston 4705 J. Carboneros, 5
Jun 1987, 80 m, R.A. Carranco 323 (MEXU); t Palo Solo, 10 Feb 1994, 15 m, J.F. Iribe 161, 394 (MEXU); El Charco del Nido, 42 km
SE de San Fernando, 24 Ene 1985, 400 m, R. Díaz 252 (UAT); 9 Feb 1994, 10 m, J.E. Lope d de ji pii 159 (MEXU); Municipio de San
Nicolás, Camino a San Nicolás, 4 Sep 1981, "os m, J.A. Barrientos 85 (COCA, IEB); Muni to la Marina, Ejido 5 de Mayo, 17
Sep 1981, 145 m, A. Brito 11 (MEXU); Ejido 5 de Mayo, 18 Sep 1981, 155 m, J.G. Galván 3 (MEXU); km 10, tramo Soto la Marina-Aba-
solo, 10 Oct 1991, 60 m, O.M. Galván 517 (MEXU); La Coma, 14 May 1985, 60 m, J.A. Franco 24 (MEXU); Rancho El 13, 21 Oct 1986,
85 m, J.A. Franco 82 (MEXU); Rancho El Saüz, 23 Oct 1983, A. Brito 167 (MEXU); Rancho Los Arroyos, 22 Oct 1986, 150 m, J.F. Iribe
325 (MEXU); Rancho Los Tripones, 16 Jul 1986, J.A. Franco 33 (MEXU); Rancho San Alfonso, 11 Feb 1981, J.A. Barrientos 765 (COCA,
TEB); Tramo San José de las Rusias-Ejido 5 de Mayo, 2 Oct 1986, 80 m, J.G. Galván 67, 158 (MEXU); Municipio de Tula, Camino a San
José de las Flores, 23 Oct 1986, 1280 m, P. Moya 175 (MEXU); Camino al Ejido Tanque Blanco, 6 Nov 1986, 1080 m, J.G. Galván 183,
191 (MEXU); Municipio de Victoria, 1 km from Victoria on the Matamoros highway, 2 Oct 1956, 350 m, F. Martínez F-2246, G. Borja
(MEXU, US); Camino al Ejido La Reforma, 23 Oct 1986, 850 m, J.E. López de la Cruz 14 (MEXU); Colonia La Estrella, Ciudad Victoria,
20 Sep 1985, 380 m, M. Cisneros 55 (MEXU); Ejido Guadalupe Victoria, 17 Mar 1994, 330 m, J.F. Iribe 407 (MEXU); Rancho El Abra, 18
Sep 1985, 330 m, M. Cisneros 44 (MEXU); San Juan y el Ranchito, 18 Mar 1994, 200 m, J.G. Galván 349 (MEXU); Santa Librada, 7 Sep
1983, 410 m, J.F. Iribe 189 (MEXU); Victoria, Sierra de la Colmena, 19 Oct 1982, J.F. Iribe 161 (MEXU); Villagrán, Ejido Guadalupe, 28
Jun 1983, 360 m, J.F. Iribe 169 (MEXU); Garza Valdéz, 19 Nov 1986, 290 m, M.H. Cervera 392 (MEXU).
24. Bouteloua uniflora Vasey, Bot. Gaz. 16:26. 1891. Tipo: U.S.A. Texas. Crockett Co.: 1890, G.C. Nealley 222 (notorio:
US-23079971; isoripo: GH).
Plantas perennes, no rizomatosas. Culmos 10 a 60 cm de alto, delgados, glabros a escabrosos, erectos,
simples, sin formar estolones. Vainas glabras a pubescentes; lígula de 0.5 a 1 mm largo, un anillo de pelos;
láminas foliares de 6 a 20530) cm de largo y 1 a 2 mm de ancho, aplanadas, algo involutas hacia el ápice,
involutas al secarse, escabrosas en ambas superficies, con algunos pelos de 2 a 4 mm de largo dispersos
sobre los márgenes, más abundantes hacia la base. Inflorescencia de 5 a 15 cm de largo, con 15 a 70 rami-
ficaciones primarias espigadas caedizas en la madurez; ramificaciones de 7 a 15 mm de largo, usualmente
péndulas, con pedicelos cortos, menores de 0.5 mm de largo o sentadas, portando 1 (2) espiguillas; raquis
triangular, hasta de 7 mm de largo. Espiguillas 7 a 9 mm de largo, permanecen insertas a las espigas, color
pajizo-purpüreas; glumas desiguales, anchas, redondeadas a agudas en el ápice, escabroso-pubescentes
sobre la nervadura central, delgadas, color verde-amarillento o con tintes purpúreos, primera gluma de 3 a
4 mm de largo, segunda gluma de 7 a 8 mm de largo; lema de 7 a 7.5 mm de largo, aguda, sin arista, glabra,
con márgenes hialinos y ápice finamente escabroso; pálea 6.8 a 7 mm de largo. Anteras de 2.5 a 3 mm de
largo, color amarillo limón, brillantes. Flósculo rudimentario ausente, o bien, representado por 1 a 3 aristas
diminutas, la arista central más larga, de 2 a 3.5 mm de largo. Cariopsis de 2.2 a 2.5 mm de largo, color
amarilla, ovoide a estrechamente elíptica.
CLAVE PARA LA DETERMINACIÓN DE LAS VARIEDADES DE B. UNIFLORA
l. C de6a 12 cm de piod t | | le 10 a 30(-45) cm de
| | follaje), con 15 a 30(-40) espigas 23a. B. uniflora
var. coahuilensis
I \
Larvars Arvinta at] " D tal Amt ^d RA Awi 975
Laminas foliares de 15 a 30 cm de largo, rectas a ligeramente curvas, esparcidas en el culmo; culmos de 40
a 60 cm de largo; inflorecencias no sobresalen el follaje, con 50 a 70 espigas 23b. B. uniflora
var. uniflora
=
24a. al uniflora Vasey var. coahuilensis Gould & Kapadia, Brittonia 16:191. 1964. (Fig. 33). Tro:
AHUILA: Rancho Los Angeles, 40 km SW de Saltillo, 4 Nov 1962, EW Gould 10300 (mooto: TAES; isorieos: DS, GH,
TEX, m Déc
Culmos de 10 a 30(-40) cm de alto. Vainas glabras a pubescentes; láminas foliares de 6 a 12 cm de largo,
concentradas hacia la base de la planta, fuertemente curvas conforme se van secando, con pelos esparcidos
hasta de 2 mm de largo en el margen cerca de la base. Inflorescencia escaposa, con 15 a 30-40) ramifica-
ciones primarias espigadas, portando generalmente 1 espiguilla, rara vez 2 (cerca de la base); raquis de 5
mm de largo. 2n=20. Florece de junio a noviembre.
Comentarios.—Se le encuentra creciendo con B. curtipendula var. caespitosa cuyas plantas poco desar-
rolladas se parece a B. uniflora var. coahuilensis, se distingue de la primera por sus anteras color naranja.
Distribución.—Especie mexicana del N y Centro de México.
Hábitat.—Laderas rocosas secas con matorrales o prec 220 a 2650 m.
huila: Muni de Art li 6, E of Saltillo, 2 Sep 2005, 2100 m, P.M. Peterson
Ejemplares examinados. MÉXICO. C
18778, J. Valdés-Reyna (US); Cerro de los Elotes, 1 Aug 1975, 220m, m Robert 4466, ]. P ini (ANSM); Si de Arteaga, El Tunal, carr.
Estatal 658, P.S. Hoge 268 (US); I.M. Johnston 7299 (US); Municipio de Castaños, Al pie de la Sierra La Gavia, km 117 carr. 57 Saltillo-
Monclova, a un lado del Campamento San Lazaro, Y. Herrera Arrieta, M. de la Cerda & O. Rosales 1484 (CIIDIR, HUAA); Municipio de
Nava, Sierra El Jardín, slopes above Canyon del Diablo, 2 Sep 2006, 1418 m, P.M. Peterson 19904, S. Lara-Contreras (US); Municipio de
Saltillo, 1 mi N of Carneros, 8 Oct 1959, 1829 m, M.C. Johnston 4192% J. Graham (MEXU); Along road to Rancho Los Angeles, about
2 km E of junction with road to Zacatecas (Concepción del Oro, Zacatecas), limestone hills, 5 Sep 1991, 1981 m, K.W. Allred 5524, J.
Valdés-Reyna & J.T. Columbus (ANSM); Area de reforestación de la Universidad Autónoma Agraria Antonio Narro, 3 Oct 1980, 1900 m,
J.A. Villarreal 884 (ANSM); Carneros, camino a la torre de microondas, 3 km O de la estación, 30 km S de Saltillo, 1400 m, J. Espinosa
(ANSMD); El Refugio, S de Saltillo, 1600 m, E. Lyonnet 3552 (MEXU); La Encantada a 17 km de Saltillo carr. a Zacatecas, win ao 1600
m, Grupo de Agrostología (ANSM); Rancho experimental Los Angeles, 48 km 5 de Saltillo, carretera 54 Sa
catecas, 26 Aug 1981, 1800 m, S.L. Hatch 4534 (ANSM); Area de Reforestación de la UAAAN Buenavista, A A. eee 884 T D.
Griffiths 8404 (US); Rancho “Los Angeles,” 48 km al S de Saltillo, S.L. Hatch, C.W. Morden & J. Valdés-Reyna 4534 (ANSM); FW. Gould &
Watson 10538 (US); La Encantada, km 9 carr. Saltillo- LR del Oro, Z. Morales, G. García & S. Inzunza s.n. (CHAPA); Carneros,
camino torre de microondas, 3 km al W de la estación, 30 | de Saltillo, J. Espinoza 94 (ANSM); Rancho experimental Los Angeles,
48 km S de Saltillo, carretera 54 Saltillo-C pción del Oro, Zac., 30 Sep 1978, 1800 m, J. Santos Sierra s.n. (ANSM); 16 Jul 1977, 2200
m, J. nd acd 1113 inii leas ES Villa ane Sierra Madera del Carmen, Villa Ocampo, J. Passini & M.F. Robert 5542A
(CHAPA y 1 kmal N de Matehuala, San Luis Potosí, 19 Sep 1984, P. Hiriart 312, 313, V. Juárez
R. Molczadzhi (MEXU); 30 a ENE of D A ENE of San Antonio de Peña Nevada, O base of Cerro Peña Nevada, 30 Jul 1983,
m, G.L. Neso 4733 (ANSM, MEXU); M pio de Galeana, Sierra El pinal Alto, 1.1 mi N of San Pablo, 10 Sep 2005, 2507 m, P.M
Peterson 18940, J. Valdés-Reyna (US); 12 km SE of San Roberto on highway 58, 27 Aug 1981, 1845 m, S.L. Hatch 4582, C.W. Morden & J.
Valdés-Reyna (ANSM); Rancho Aguililla, 26 Aug 1989, 1900 m, G.B. Hinton 19588 (MEXU); 8 km al entronque de San Roberto, rumbo
leana, Y. Herrera Arrieta & A. Cortés 1447 (CIIDIR, HUAA); km 150, carr. 57 Matehuala-Saltillo, a 20 km de San Roberto, Y. Herrera
Arrieta & A. Cortés 1451 (CIIDIR, HUAA); km 162, carr. 57 Matehuala-Saltillo, Y. Herrera Arrieta & A. Cortés 1453 (CIIDIR, HUAA); 4 mi
S de Galeana, R.L. Mc Gregor, uae Harms, A ar es R. del Rosario & R. Seagal 92 (US); 1 mi al E de San Marcos, 13 mi SW de Galeana,
F.W. Gould 10724, 10743, 10755 (US bide, Ejido Santa Rosa, Iturbide, A.E. Estrada 1670 (UAG). Tamaulipas: Municipio
de Miquihuana, Colonia Ágrícola La Peña, 26 Feb 1981, 700 m, J.F Iribe-Duarte 104 (MEXU); Ejido Valle Hermoso, cañón de la Sierra
a P 4 Sep 1981, 2100 m, R.A. Carranco 65 (MEXU); Rumbo al Aserradero, Sep 1981, 2650 m, R.A. Carranco 71 (MEXU);
Nicolás, Rancho La Trinidad, 19 Oct 1983, 650 m, J.F. Iribe 203 (MEXU
24b. Bouteloua uniflora var. uniflora Vasey. (Fig. 34).
Culmos de 40 a 60 cm alto, muy similar a la var. coahuilensis. Vainas glabras o casi glabras; láminas foliares
de 15 a 30 cm de largo, bien distribuidas en los culmos, no concentradas hacia la base de la planta, no se
curvan al secarse, con pelos de 3 a 4 mm de largo en el margen cerca de la base. Inflorescencia no escaposa,
con 50 a 70 ramificaciones primarias espigadas, muy cercanas entre sí, portando 1 o 2 espiguillas sobre un
raquis pubescente, hasta de 7 mm de largo. 2n = 20. Florece de agosto a septiembre.
Distribución.—Desde Texas hasta México en Coahuila, Nuevo León y Tamaulipas.
1 Ll A£4b ^ Dadaminal D h Institut. f Texas 2(2)
976
ape.
zo ^
&
Ce
STO
EA ce
Fic. 33. Distribución d
Hábitat.—Suelos rocosos, laderas secas o moderadamente húmedas, en pastizales y matorrales; 600 a
2000 m.
XT. i. Ja n :11 A 1 Carmen
411 de 13541 NAT ALKA
33.1 KM IN VV
MÉXICO. Coal jui y ]
); Municipio de Castaños, S end of Puerto San Lazaro (Cuesta La Muralla),
12 Sep 1991, 1380 m, P.M. Peterson 10588, C.R. Annable (US
0.5 mi N of San Lázaro along route 57, 28 Jul 1977, 1250 m, J. Valdés-Reyna 1107, T.L. Wendt (ANSM); Municipio de Acuña, Serranías del
Burro, 12 km O del rancho el Bonito, cañón el Toro, 18 Sep 1977, 1625 m, D.H. Riskind 2144 (ANSM); Serranías del Burro, 12 km O del
rancho el Bonito, cañón el Toro, 18 Sep 1977, 1625 m, J. Valdés-Reyna 1184, D.H. Riskind (ANSM); Municipi t ] d
El Tunal, carretera estatal 65, 17 Sep 1989, 2390 m, P.S. Hoge 268, M.E. Barkworth & J. Valdés-R
pie de la Sierra La Gavia, km 117 carr. 57 Saltillo-Monclova, a un lado del Campamento San Lazaro, Y. Herrera Arrieta, M. de la Cerda &
, ju [^E e
eyna (ANSM); Municipio de Castaños, Al
I
O. Rosales 1486 (CIIDIR, HUAA); Municipio de Cuatrociénegas, Cafión del Agua (N-draining), mouth of canyon, vicinity of Ranchito, 25
ul 1977, 1425 m, J. Valdés-Reyna 1006, T.L. Wendt (ANSM); Mounth of Cañada Los Posos (E-draining) in N foothill region of range, near
of Tanque La Boquilla, 2.7 mi by road O of rancho Cerro de la Madera, 26 Jul 1977, 1420 m, J. Valdés-Reyna 1049, T.L. Wendt (ANSM);
Larrars Arriata at 5l : D tal "lal ta da Mw 977
Lic 34 Dictril Fa la D L T] T.)
Sierra de la Madera, vicinity of “La Cueva’ in Corte Blanco fork of Charretera Canyon, 11 Sep 1941, 1615 m, I.M. Johnston 9060 (MEXU);
Municipio de Saltillo, 53 km S of Saltillo, along highway 54, S of Concepción del Oro, Zac., 26 Aug 1981, 1850 m, S.L. Hatch 4509, C.W.
Morden & J. Valdés-Reyna (ANSM); On highway 54 (road to Concepción del Oro), 6 km S of Saltillo, near Universidad Autónoma Agraria
Antonio Narro, 18 Sep 1988, 1850 m, M.E. Barkworth 5107, P.S. Hoge (ANSM); Saltillo, 0.3 km E of Hwy to Zacatecas up to road to San
Lorenzo, 10 Sep 1990, 1720 m, P.M. Peterson 10552, C.R. Annable (US); Municipio de Zaragoza, Rancho Santa Cruz, aproximadamente
81 km N de Müzqui la carretera Múzquiz-Boquillas del Carmen y 12 km de brecha NO bo a Santa Cruz, 23 Sep 1977, 600 m,
J. Valdés-Reyna 1286, L.E. Rodríguez (ANSM). Nuevo León: Municipio de Aramberri, 75.8 km N of Dr. Arroyo on Hwy 61 and 3.2 km
S of Ascensión (La Chona), 13 Oct 1991, 2100 m, P.M. Peterson 11145, C.R. Annable (US); Municipio de Doctor Arroyo, 21 km al N de
Matehuala, San Luis Potosí, 19 Sep 1984, P. Hiriart 316, V. Juárez y R. Molczadzki (MEXU); Camino a Agua Nueva, 23 Jun 1981, 2000
m, O.L. Briones 695 (ANSM); 9.4 mi W cf San Antonio de Peña Nevada, 22 Sep 2002, 1807 m, P.M. Peterson 16789, J. Valdés-Reyna & M.
Sosa (US); Municipio de Galeana, Alrededor de Galeana, Sep 1980, J.A. Ochoa-Guillemar 1261 (MEXU); Navidad, carretera 57, km 85 al
N de Saltillo, 23 Sep 1978, 1650 m, F. Cárdenas & M.A. Bernal (MEXU); 12.3 mi S of Milagro, 18 Sep 2003, 2040 m, P. M. Peterson 17806,
J. Valdés-Reyna & H.S. Hinton (US); Municipio de General Zaragoza, 4 km S of Zaragoza at jet of road to Cerro Viejo-Tepehuanes, 20 Sep
m , Iattha Bataniral D h Institut f Texas 2(2)
Crt A 1
2001, 1630 m, P.M. Peterson 15848, J. Valdés-Reyna (US). T: ipio de Hidalgo, Sierra La Lagunita, 6.4
on road towards Agua Fría, 19 Sep 2002, 1397 m, P.M. Peterson 16677, J. Valdés- -Reyna & M. Sosa (US); Municipio de Miquihuana, 15
km al SE de Miquihuana, 23 Aug 1984, P. Hiriart 284, V. Juárez & R. Molczadzki (MEXU); Municipio de Victoria, Altas Cumbres, 26 Oct
1980, 1100 m, J.F. Iribe s.n. (MEXU).
25. Bouteloua warnockii Gould & Kapadia, SouthW. Naturalist 7:176. 1962. (Fig. 35). Tiro: U.S.A. Texas.
Culberson Co.: 2 mi W of Kent, 18 Jul 1961, Gould & Kapadia 9533 (nororio: TAES; isoriros MICH, MO!, SMU, SRSC, TEX, UC,
US-25513671).
Plantas perennes, caespitosas, formando macollos de 4 a 10 cm de diámetro, algunos con rizomas cortos.
Culmos 20 a 35(—50) cm de alto, delgados, erectos, tiesos, sin ramificarse desde la base; nudos anteriores a
la inflorescencia con tintes purpúreos. Hojas generalmente basales, verde-azulosas, con frecuencia glaucas;
vainas estriadas, escabrosas con pelos tiesos, dispersos, de 1 a 1.5 mm de largo, obscureciendose con la
edad (cambiando a color café); con un margen hialino, angosto, que forma dos pequeñas aurículas mem-
branáceas, de 0.5 mm de largo, en la base de las láminas foliares; lígula de 0.5 a 1.5 mm de largo, un anillo
de pelos y algunos pelos más largos de base papilosa; láminas foliares de 5 a 1525) cm de largo y 1 a 2(3)
mm de ancho, involutas y arqueadas, muy escabrosas en la superficie adaxial, con algunos pelos largos en
el área cercana a la lígula y otros pocos de 5 mm de largo con bases papilosas en los márgenes, erectas y
nd emma en una rua muy ane Inflorescencia de 5 a 15 cm de largo, exserta, con 9 a 25(-35)
das, inclinadas, caedizas en la madurez; espigas de 5 a 10 mm de
largo, raquis alado, Em sob los margenes, portando (1-)2 a 6 espiguillas, apifiadas en el cuarto
o tercio basal. Espiguillas de 5 a 6.5 mm de largo, permanecen insertas en las espigas, de color verde o con
tintes purpüreos o cafés; glumas desiguales, agudas o acuminadas, primera gluma de 3 a 4 mm de largo,
angosta, pubescente en la nervadura central y hacia el ápice, segunda gluma de 4 a 5.5 mm de largo, más
ancha, acuminada, casi glabra a finamente pubescente en la nervadura central; lema de la flor perfecta 5 a
6.5 mm de largo, glabra, con el ápice trífido, las nervaduras laterales se prolongan en aristas cortas, de 0.5 a
1 mm de largo, gruesas, escabrosas; pálea glabra o con escasa pubescencia entre los nervios, de igual tamaño
que la lema o un poco menor. Anteras de 2 a 2.2 mm de largo, color púrpura. Flósculo rudimentario 1,
bien desarrollado, lema del flósculo de 5 a 6.5 mm de largo, con 3 aristas de 2.5 a 5 mm de largo, fuertes y
escabrosas, li arista central más larga. Cariopsis de 1.8 a 2.6 mm de largo, color amarillo,
O
ovoide a estrechamente MM Js 20, con plantas aneuploides en Estados Unidos desde 21-40 (Gould,
1979). Florece de junio a agosto.
Distribución.—Desde Texas hasta México en Coahuila.
Hábitat.—Laderas pedregosas, cerca de arroyo, sobre suelos calizos; 1250 m.
Ejemplares examinados. MÉXICO: Coahuila. Municipio de Castaños, South end of Puerto San Lázaro (Cuesta la Muralla), 0.5 mi N of
E Lázaro iind. a sl J. Valdés-Reyna VR-1107 (ANSM); Municipio de San Pedro, 5.5 km al NW de Nueva Delicias, vertiente E de la
li N de San Pedro de las Colonias, Y. Herrera Arrieta, M. de la Cerda & O. Rosales 1492 (CIIDIR, HUAA).
NOMBRES DUDOSOS Y EXCLUIDOS
Cynosurus secundus Pursh, Fl. Amer. Sept. 2:728. 1813. Dinebra secunda (Pursh) Roem. & Schult., Syst. Veg. 2:711. 1917.
Chloris secunda (Pursh) Eaton, Man. Bot. (ed. 5) 173. 1829. Tiro: U.S.A. Louisiana: upper, Bradbury s.n. No revisado el tipo.
Erucaria tetrastachya Cerv., Naturaleza (México City) 1:349. 1870. Tiro: MÉXICO: = Bouteloua barbata Lag.? No revisado
el tipo
Bouteloua hirta var. minor Vasey ex L.H. Dewey, Contr. U.S. Natl. Herb. 2(3):531.1894. Tro: U.S.A. Texas: Central
Erucaria lutescens Cerv., Naturaleza (México City) 1:349. 1870. Tiro: MÉXICO: No revisado el tipo.
AGRADECIMIENTOS
Agradecemos al Instituto Politécnico Nacional (CIIDIR, COFAA, SIP); a la Comisión Nacional para el
Conocimiento y Uso de la Biodiversidad (CONABIO-B061) y al Consejo Nacional de Ciencia y Tecnología
Fic. 35. Distribución de R. L m I ta fa M
(CONACyT-53142), por el financiamiento otorgado a Y. Herrera para la realización del presente estudio. Así
también agradecemos el financiamiento del Instituto Smithsoniano, a través de la Oficina de Entrenamientos
de Investigación y Servicios para Vistantes de Investigación, y del Fondo de Dotación Restringida, del Pro-
grama de Estudios Escolares, Oportunidades de Investigación, la Fundación Atherton Seidell, el Programa de
Inventarios e Investigaciones de Biodiversidad por el apoyo a P.M. Peterson. Reconocemos el financiamiento
para viajes de colecta que otorga National Geographic Society 8087-06 otorgado a P.M. Peterson. Nuestro
más sincero reconocimiento a Armando Cortés Ortiz por su colaboración desinteresada en la elaboración
de los mapas que se incluyen. La descripción de Bouteloua tamaulipensis, así como las tres ilustraciones del
taxon, han sido provistos por Darisol L. Pacheco y J. Travis Columbus; Anabel Duarte preparó la ilustración
(Fig. 30) y Gilberto Ocampo contribuyó con la revisión de la descripción. A los curadores de los herbarios
980 I Lafiha Dataniz4l D h Institut f Texas 2(2)
ANSM, ARIZ, B, BA, BAA, BM, CIIDIR, ENCB, GH, HERZU, HINT (herbario personal de George S. Hinton),
HUAA, IBUG, INEGI, IZTA, K, LE, MEXU, MICH, MO, MSC, NMC, NMSU, NY, P, RM, RSA, SI, SLPM,
TAES, TEX, UAT, US, UT, UTC, VT y W, se les agradece las facilidades y atenciones recibidas para hacer
posible la revisión de los ejemplares a su cargo.
REFERENCIAS
ALLRED, K.W. 1982. Describing the grass inflorescence. J. Range Managem. 35(5):672-675.
BeeTLE, A.A. 1987. Las gramineaes de México, Tomo II (incluye Bouteloua, Buchloe v Cathesticum). Cotecoca, S.A.RH.,
México, D.F. Pp.53-104, 155-159 y 172-179.
CoLumbus, J.T. 1999. An expanded circumscription of Bouteloua (Gramineae: Chloridoideae): new combinations
and names. Aliso 18:61-65.
CoLumBUs, J.T, M.S. Kinney, R. PANT Y M.E. Siqueiros DELGADO, 1998. Cladistic parsimony analysis of internal transcribed
spacer region (nrDNA) sequences of Bouteloua and relatives (Gramineae: Chloridoideae). Aliso 17:99-1 30.
CoLuMaUS, J.T., M.S. Kinney, M.E. Siqueiros DeLGADO Y J.M. Porter. 2000. Phylogenetics of Bouteloua and relatives
(Gramineae: Chloridoideae): cladistic parsimony analysis of internal transcribed spacer (nrDNA) and trnL-F
(cpDNA) sequences. En: S.W.L. Jacobs y J. Everett, eds., Grasses: Systematics and Evolution. CSIRO Publishing,
Collingwood, Victoria, Australia. Pp. 189-194,
CoLumus, J.T., R. CeRROs-TLATILPA, M.S. KINNEY, M.E. Siqueiros-DeLGADO, H.L. Bett, M.P. GrieritH Y N.F. REFULIO-RODRIGUEZ.
2007. Phylogenetics of Chloridoideae (Gramineae): A preliminary study based on nuclear ribosomal internal
transcribed spacer and chloroplast trnL-F sequences. Aliso 23:565-579,
DAVILA- ARANDA, PR. LIRA-SAADE Y J. VALDES-REYNA. 2004. Endemic species of the grasses in México: A phytogeographic
approach. Biodivers. & Conservation 13:1101-1121.
Dávila, P, M.T. M&fA-SAULEs, M. Gomez-SANCHEz, J. VALDÉS-REYNA, J.J, Ortiz, C. Morin, J. CASTREJÓN Y A. Ocampo. 2006. Ca-
tálogo de las gramíneas de México. Universidad Nacional Autónoma de México and Comisión Nacional para
el conocimiento & uso de la Biodiversidad, México, D.F.
ESPEJO SERNA, A., A.R. LÓPEz-FeRRAR Y J. VALDÉS-REvNA. 2000. Poaceae. In: A. Espejo Serna & A.R. López-Ferrari (eds.), Las
Monocotiledóneas Mexicanas: una Sinopsis Florística, Partes IX-XI. Consejo Nacional de la Flora de México,
A.C, Universidad Autónoma Metropolitana-Iztapalapa, y Comisión Nacional para el Conocimiento y Uso de
la Biodiversidad, México, D.F. P. 10.
Goulo, FW, 1979. The genus Bouteloua (Poaceae). Ann. Missouri Bot. Gard. 66:348-416.
GriFFITHS, D. 1912. The grama grasses: Bouteloua and related genera. Contr. U.S. Natl. Herb. 14(3):343-428.
HERRERA ARRIETA, Y., PM. PETERSON Y M. DE LA CERDA Lemus. 2004. Revisión de Bouteloua Lag. (Poaceae). Comision
Nacional para el Conocimiento y Uso de la Biodiversidad y Instituto Politecnico Nacional, Durango, México.
Pp. 1-187.
HERRERA ARRIETA, Y. v J. Rzepowski. 1990. Gramineae, In: Rzedowski, J. y G.C. Rzedowski, eds. Flora Fanerogámica del
Valle de México, Instituto de Ecología, Centro Regional del Bajío, Pátzcuaro, Michoacán. Pp. 34-173.
HiLu, KW. v L.A. Auct. 2000. Phylogenetic relationships in subfamily Chloridoideae (Poaceae) based on matk
sequences: A preliminary assessment. In: S. W.L. Jacobs and J. Everett, eds. Grasses: Systematics and Evolution.
CSIRO, Melbourne. Pp. 184—188.
Hitu, KW. v LA. Auce. 2001. A phylogeny of Chloridoideae (Poaceae) based on matK sequences. Syst. Bot.
26:386-405.
McVauaH, R. 1983. Flora Novo-Galiciana. A descripti tofthe vascular plants of western México. Gramineae.
14:1-436. University of Michigan Press, Ann Arbor.
Pacheco, D.L. 2002. Estudio Sistemático del género Cathestecum (Poaceae: Chloridoideae: Boutelouinae). Tesis
Doctor en Ciencias, Fac. de Est. Sup. Iztacala. UNAM, México D.F Pp. 1-353.
PETERSON, P.M, RJ. SORENG, G. Davipse, T.S. FILGUEIRAS, F.O. ZULOAGA v E.J. Juoziewicz. 2001. Catalogue of New World grasses
(Poaceae): Il. Subfamily Chloridoideae. Contr. U.S. Natl. Herb. 41:1—255.
Lu Arrieta et al., B tal Anl duc WA RE eee 981
Peterson, P.M., J.T. CotuMaUs v S.J. PENNINGTON. 2007. Classification and biogeography of New World grasses: Chlo-
ridoideae. Aliso 23:580—594.
Pierce, G.J. 1979. A biosystematic study of Cathestecum and Griffithsochloa (Gramineae). Ph.D. dissertation. Uni-
versity of Wyoming, Laramie. Pp. 1-244.
PoHL, RW. 1994. 100. Bouteloua Lagasca, nom. cons. In: Davidse, G, M. Sousa S. y A.O. Chater, eds. Flora Meso-
americana, Vol. 6 Alimataceae a Cyperaceae. Universidad Nacional Autónoma de México, Missouri Botanical
Garden y The Natural History Museum (London), México. Pp. 293-295.
Reever, J.R. v C.G. Reener. 1980. Systematics of Bouteloua breviseta and Bouteloua bromosa. Syst. Bot. 5:312-321,
Wierr, J.K. 2003. 17.46 Bouteloua Lag. In: M.E. Barkworth, K.M. Capels, S. Long, and M.B. Piep, eds. Magnoliophyta:
Commelinidae (in part): Poaceae, part 2. Flora of North America North of México, volume 25. Oxford University
Press, New York. Pp. 250-269.
982 I l ^f tha Bataniral D h Institut f Texas 2(2)
BOOK REVIEW
ADA V. SEGRE. 2006. The Gardens at San Lorenzo in Piacenza, 1656-1665. In two parts: (ISBN 13: 978-0-
88402-302-9, hbk.; ISBN 10: 0-88402-302-8, pbk.). Dunbarton Oaks Research Library and Collection,
Washington D.C., U.S.A. (Orders: www.doaks.org; ms odi at www.hup.harvard.edu, 800-405-
1619). $95.00. 107 pp., color photos, maps, and ill gl ith accompanying Manuscript
ee adi 12 1⁄2" x 9 1/2".
— e destum E aun m 1656-1665 ied by adilior Ada Seat
compla rE hes Ph Dub i teenth-century Italian fl lens. Comi fi ] ist f a five-hundred
year old Italian planting notebook manuscript, she realized it could be of pones in revealing ra practices of the third
quarter of h century Italy. Though too late to include in her thesis, she promised to ret tic garden journal
housed in Dumbarton Oaks's Rare Books Collection and give it the comprehensive research it deserved. She speni two years on just
— hod dada. pus ownership of the garden described in the Paar The evidence pointed to Margherita de' Medici
of th taly.
iacenza, cities located south
Tho} 11 1,1 Gey mc qr duca 1 1 : ; ftho 1442 A daci dm 11 1
represents Segre's study. Inserted in qe back of ihe study book are xr painstaking, computer-driv en ees reconstructions
of the flower gardens: 1656, the Nine Star Garden 1662, the Nine Star Garden 1664, and the St 1665
study book is the pee reproduction x the ae ai ee oe notebook, also in spa to capture the feel and TRE of
the original. Thi century ] fl gardens at San Lorenzo. Here you may detect for
yourself, and follow along with her investigations of these drawings add who might have made them. Both books are incased in a rich
oxblood buckram-covered slipcase with blind embossing on the
en reconstruction is supplemented by appendices ne 2 transcriptions and translations of the original pane
nd list of Plants at San Lorenzo, 1656-1665. Her xn in this nip isa ido di contribution to the fiel
I ill , and cultural history as vcl
E
iT Annseasmw
as to the general reader and book calcu Heagy, Dee Illustrator,
J. Bot. Res. Inst. Texas 2(2): 982. 2008
REVISION OF TOCOYENA (RUBIACEAE: GARDENIEAE) FROM
THE STATES OF GOIÁS AND TOCANTINS AND A NEW SPECIES
ENDEMIC TO WHITE-SAND AREAS IN THE BRAZILIAN CERRADO
Piero G. Delprete'
Federal University of Goiás, Campus II
Institute of Biological Sciences - ICB-1
e of General o
4001-970 Goiânia, Goiás, BR.
ABSTRACT
my x 1 Est 38 ME f, 1 ni A - Tocantins pe | po ERNE ful Jh h E 1
a E A
on the species of Tocoyena Aubl. (Rubiaceae, Gardenieae) occurring in this area, and are pins here. Also, some recent colo
of Tocoyena f hi d in th f Goiás and T: i y attention to the veg fth reas. Although
y aii 1 g f h PAR A fel A n x rl R 31 “campinas” or “campinaranas”),
very roy poe] Im aah ] iti a A fth Cerrado Biome A cl di 7 4 4 - I lanew
endemic species of Ti is described and illustrated, and a call for for tl tion of tl is] hed
A key to the species of Tocoyena encountered in the states of Goiás and Tocina is also included.
Key Wonps: Tocoyena, Gardenieae, Rubiaceae, Goiás, Tocantins, white sands, Cerrado, Brazil, conservation
RESUMO
fot A detalhad h^ ^ d amm
Y
Durante a preparacáo do tratado das Rubiaceae para a Flora de Goiás e Tocantins, foram
de amostras de herbário sobre ás espécies de Tocoyena Aubl. (Rubiaceae, Gardenieae
Também, algumas coletas recentes de Tocoyena provenientes de áreas pe areias UN nos Estados de Goiás e Tocantins atraíram a
areia branca na Bacia Amazónica (conhecidas
P
d à qui apresentadas
minha atenção sobre estas localidades. Embora existam muitos
no Brasil como “campinas” ou “campinaranas”), se sabe muito p b á de areias b tziti Bioma Cerrado.
4
Uma curta discussáo sob é apresentada, uma nova écie endémica de Tocoyena é descrita e ilustrada, e um apelo por
Y
d ao d A 6 i lançado Una chave para as espécies de Tocoyena eue nos Estados de
e r e uy
Goiás e Tocantins é também incluída.
PALAVRAS CHAVE: Tocoyena, Gardenieae, Rubiaceae, Goiás, Tocantins, areias brancas, Cerrado, Brazil, conservacáo
During the preparation of the Rubiaceae treatment for the Flora de Goiás e Tocantins, detailed field and her-
barium observations with taxonomic implications were made on the species of Tocoyena Aubl. (Rubiaceae,
Gardenieae) occurring in this area. Tocoyena is a genus of about 25 species with Neotropical distribution,
represented by subshrubs, shrubs and trees. The genus was divided into two groups by Schumann (1891)
based on the shape of the corolla lobes and the lobed portion of the flower buds: 1) Acutiflorae K. Schum.,
with corolla lobes and flower buds acute [with T. foetida Poepp. & Endl. and T. guianensis K. Schum.]; 2)
Obtusiflorae K. Schum., with corolla lobes and flower buds obtuse [with T. formosa (Cham. & Schltdl.) K.
Schum., T. brasiliensis Mart., T. sellowiana (Cham. & Schltdl.) K. Schum. and T. bullata Mart.]. The taxo-
nomic position of the genus has always been clear, as it has been placed in the tribe Gardenieae in all major
classifications (Hooker 1873; Schumann 1889, 1891; Bremekamp 1934; Verdcourt 1958; Robbrecht & Puff
1986; Robbrecht 1988, 1993). Persson (1993, 1995, 1996, 2000) with morphological, palynological, and
molecular studies confirmed its position in the Gardenieae.
Several floristic and taxonomic treatments are available for the South American species of Tocoyena.
Schumann (1889) revised the species present in Brazil (and those of contiguous countries) recognizing six
species. Steyermark (1965) presented a synopsis of Tocoyena for the Flora of the Guayana Highlands in which he
Cedex 5, FRANCE, pdel
la Dart 7 la Mával, AMAP TA-A51/PS?_ Rlvd de la Lironde 24398 M
F
J. Bot. Res. Inst. Texas 2(2): 983 — 993. 2008
984 I l of tho Botanical R h Institut f Texas 2(2)
recognized seven species, complemented by important taxonomic information. Taylor and Steyermark (2004)
recognized six species present in Venezuela, five of which present in the Venezuelan Guayana.
Prado (1987) revised the species of Tocoyena occurring in Brazil, and maintained the two groups pub-
lished by Schumann. The twelve species recognized in her revision are fully described, beautifully illustrated,
with full synonymy, and many specimens cited. She also proposed two varieties for T. formosa, based mostly
on vegetative characters. However, this study is a master dissertation, and the taxonomic novelties are not
effectively published (McNeill et al. 2006: Art. 30.5).
Silberbauer-Gottsberger et al. (1992) revised the group Obtusiflorae, which is mostly distributed in the
Brazilian Cerrado and Caatinga. The delimitation and synonymy of several species differed significantly from
those of Prado. These authors proposed three varieties of T. formosa, supported by a multivariate analysis
using mostly vegetative characters, resurrected T. brasiliensis as a separate species, and presented a key to
all the species and varieties recognized in the group Obtusiflorae. Two recent floristic of Tocoyena
are also available from two states of Southern Brazil, Santa Catarina (Delprete et al. 2005) and Sáo Paulo
(Prado & Kinoshita 2007), dealing with one and three species, respectively.
The flowers of Tocoyena give off a pleasant scent, resembling that of Gardenia J. Ellis (Gardenieae),
reaching its highest intensity at night, and are pollinated by several species of sphingid moths (Silberbauer-
Gottsberger 1972; pers. obs.). Most species of Tocoyena have great ornamental potential, and it is surprising
Pn ny have not been e into cua for this purpose. In addition, the fruits of all the species of
are an important food for birds (e.g., macaws, parakeets, parrots), mammals (e.g., monkeys,
nli e tamarins, agoutis, coatis, ic and peccaries) and small marsupials (e.g., opossums) in
most Neotropical habitats.
Recently, some collections of Tocoyena from white-sand areas in the states of Goiás and Tocantins at-
tracted my attention to the vegetation of these localities. Many studies are available about white sand areas of
the Amazon Basin, known in Brazil as “campinas” or “campinaranas,” where they are more evident because
of the dramatic difference between their savanic vegetation and the surrounding tall canopy forests typi-
cal of the lowland Amazon. Many classic studies on the phytosociology and floristic composition of these
Amazonian areas are available, as, for example, those of Ducke (1907, 1913), Ducke & Black (1953, 1954),
Gottsberger & Morawetz (1986), Anderson (1981), Prance (1996), Lisboa (1976), and Takeuchi (1960a,
1960b, 1960c, 1961, 1962), among many others. On the other hand, little is known about the vegetation of
quartzitic white-sand areas within the Cerrado Biome; this is probably due to the fact that the contrast with
the surrounding vegetation is not as striking as that of Amazonian white sands.
The floristic composition of white-sand areas in the Cerrado Biome is still largely unknown, as botanical
collections in these areas are sparse and have been made only by botanists randomly passing through these
areas. The most important factor influencing species composition in the Cerrado Biome is soil composition,
as it has been amply demonstrated by many interdisciplinary studies, as for example those of Cole (1986,
1992), Emmerich (1990), and Haridasan (2001). Asa pip the E vegetaon of these areas is made of
two main components: 1) plani Ie also present int g Cerrado sensu stricto, but with smaller
individuals than th gint with red soil (pers. obs.), and 2) species of plants
(and animals; e.g., Bodripües et al. 2007) deni to these areas. The new species of Tocoyena described
below is certainly one of those narrow endemics. Finally, these white-sand areas are currently in extreme
danger of being completely destroyed, as even some ecologists indicated to *encourage the construction of
industries and housing in infertile or less fertile soils, as the white sand areas in several region of tabular
interfluvium" (liberal translation from Ab'Saber 1990, p. 25), and these areas should be the subject of urgent
conservation programs.
SYSTEMATIC TREATMENT
Tocoyena formosa (Cham. & Schltdl.) K. Schum. in Mart., Fl. Bras. 6(6):347. 1889. (Fig. 1). Gardenia formosa
Cham. & Schltdl., Linnaea 4:200. 1829. Tyee: BRAZIL: Southern Brazil, without locality, s.d., Sellow s.n. (HoLoTyPE: B, destroyed,
photo-B at NY
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Tocoyena hirsuta Moric. ex DC., Prodr. 4:375. 1830. Tree: BRAZIL: without locality, s.d., Collector Unknown s.n. (HoLoTyPE: G-DC).
Tocoyena velutina Mart., Flora 24 II, Beibl. 6:81. 1841. Tyre: BRAZIL: Maro Grosso: Cuiabá, Aug [s.d.] (fl), Martius s.n. (HOLOTYPE: M)
€ microdon Mart., Flora 24 II, Beibl. 6:82. 1841. Tocoyena is (Cham. & Schltdl.) K. Schum. var. microdon (Mart.) Gottsb. &
Ehrend., Pl. Syst. Evol. 181:163. 1992. Tyre: BRAZIL: Marannáo: Rio Itapicurú, 1809 (fl), Martius 3297 (Hotorvpr: M).
Tocoyena mue K. Schum., Bot. Jahrb. Syst. 25, Beibl. 60:18. 1898. Tocoyena stipulacea K. Schum. ex Glaziou, Mém. Soc. Bot. France
43. 1909, nom. nud. Tree: BRAZIL: Gorás: Between Macacos and Chico Costa, 9 Oct 1893 (fl), Glaziou 21485 (HoLotvrE: B,
destroyed, photo web!; isotypes: BR, C n.v., K n.v. photo web!, R; photo-B at NY)
Tocoyena mollis K. Krause, Notizbl. Kónigl. Bot. Gart. Berlin 6:202. 1915. Tree: BRAZIL: Banta: Remanso, catinga, Dec 1906 (fl), Ule 7143
(HoLorwE: B, destroyed, photo web!, sory: L n.v.; photo-B at E NY).
Tocoyena surinamensis Bremek., Rec. Trav. Bot. Neerl. 33:710. 1936. Tyre: SURINAME: upper m River, savanna near Platte Berge,
5 Dec 1935 (fl), H.E. Rombouts 323 (HOLOTYPE: U; ISOTYPES: E n.v., K n.v. photo web!, MO, U, US).
NUM formosa (Cham. & Schltdl. He o var. 1 b. & Ehrend., Pl. Syst ‘Evol 181:162. 1992. Tyre: BRAZIL: Sao
: Mun. Botucatu, 18 km N, 1 M ]-Pi eze de Maio, 22°45'S, 48?25'W,
550 m, 1 Nov 1971, I. Gottsberger & G. Gottsberger 11-11171 cone UB; ISOTYPES: BOTU, Knv um web!, GI n.v.
Taxonomic observations: Silberbauer-Gottsberger et al. (1992) recognized three widely sympatric varieties
in Tocoyena formosa, according to the results of a multivariate analysis using vegetative characters and field
observations in pares of Cerrado in the state of Sáo Paulo ea Bra”: However, a ie of about 900
herbarium sy d personal field observations of thi its 1 that
the leaves vary widely in shape and vestiture, ranging from densely abese to Bn os (or exception-
ally glabrous and minutely bullate) M the same populations throughout Brazil, but the floral characters
remain constant throughout its geographic distribution. For the above reasons, T. formosa is here treated as a
plastic species with pol hi ive characters, with the same delimitation proposed by Silberbauer-
Gottsberger et al. (1992), but tou any subspecific taxonomic ranks recognized.
In the southern portion of the state of Goiás, and more specifically in the Parque Nacional das Emas
and the Serra dos Caiapós, several collections were made of individuals reduced to a basal rosette of leaves,
practically sessile to the ground or with an aerial stem of up to 3-5 cm long, and a xylopodium trailing
horizontally underground up to 70 cm in length (or maybe longer, as they were broken off by the collectors).
Aside from the extremely reduced habit, the leaves, flowers and fruits of these individuals are practically
identical to those with shrubby or arboreal habit (Figure 1). Because of this, these individuals are treated
here as a sessile form of T. formosa without any taxonomic recognition. The reason for this peculiar habit is
unknown, but, because all these individuals were collected in shrubby campos (“campos sujos") on sandy/
rocky soils, it is probable that the depauperate soil low in nutrients and the frequent fires influenced their
subterranean habit. The collections examined with sessile leaf rosettes are listed here: BRAZIL: Gorás: Mun.
Mineiros, Parque Nacional das Emas, 11 Oct 1994 (fl), H.D. Ferreira 2744 (NY, UFG), 12 Nov 1998 (fl), H.D.
Ferreira 3608 (UFG), 26 Feb 1995 (fr), H.D. Ferreira & O. Silva 3452 (NY, UFG), 11 Oct 1994 (fl), H.D. Ferreira
3900 (UFG), 12 Nov 1998 (fl), H.D. Ferreira 3988 (NY, UFG); Serra do Caiapó, ca 60 km S of Caiapônia, on
road to Jataí, 27 Oct 1964 (fD, Irwin & Soderstrom 7425 (NY, RB).
Distribution and ecology. —Widely distributed in tropical South America, from Suriname and throughout
Brazil and Paraguay. Subshrub, shrub or tree (exceptionally reduced to a sessile rosette with a subterranean
xylopodium; Figure 1); highly plastic and adapted to a wide variety of habitats. One of the most representative
species of Neotropical savannas (Brazilian Cerrado), mostly in shrubby savannas (Cerrado sensu stricto"),
but also encountered at gallery forest margins, in mesophytic forests (“Cerradáo” of Brazil), shrubby campos
(campos sujos”), and on rocky outcrops (“campos rupestres”). Sometimes also found in disturbed vegetation
or even as a dnd species in secondary vegetation in the Cerrado Biome.
Sel d 1: BRAZIL: Banta: Mun. Sáo Desidério, di 11 Roda Velha and Estiva, 725 m, 12°42'S, 45°48'W, 7
Nov 1997 (ED, D. Alvarenga et al. 1024 (NY); Mun. Formosa do Rio Preto, Projeto Ouro Verde, Rod. Anel de Soja, 710 m, 11°26'S, 46°4'W,
14 Nov 1995 (£D, B.M.T. Walter et al. 2927 (CEN, NY). Tocantins: Mun. Porto Nacional, 10 Oct 1993 (f1), M. Alves s.n. (HTO, NY); Mun.
Pium, Ilha do bananal, Parque Nacional do Araguaia, Rio Javaés, 9°59'S, 50°7'W, 27 Mar 1999 (fr), M.A. Alves et al. 4192 (IBGE); Mun.
Dianópolis, 570 m, 11°40'S, 46°40'W, 30 Sep 2003 (fD, T.B. Cavalcanti et al. 3385 (CEN); Mun. Porto Nacional, Área da FAB, 7 Aug 1993
(fl), J. Ferreira et al. 4 (HTO); Mun. Araguaína, ca 10 km S of Araguaina, 300 m, 16 Mar 1968 (fr), Irwin et al. 21237 (NY); Mun. Palmas,
Serra do Lageado, between Rio Taquarassu and Rio Taquarassusinho, 48°13'S, 10°17'W, 13 Apr 1994 (fp), LY. Lima & F.P.R. de Jesus 274
Delprete, Revision of Tocoyena 987
(HEPH, HTO), 275 (HEPH); Mun. Tupiratins, Córrego Almoço, 814'S, 48°9'W, 27 Oct 2000 (fD), S.F. Lolis et al. 71 (HTO); Mun. Caseara,
Parque Estadual do Cantão, 22 Jan 2000 (imm fr, P.E. cal & M. Richter 752 ide NY); road Taguatinga-Aurora do Norte, km 9, 6 Dec
1991 (fl), B.A.S. Pereira et al. 2005 (IBGE NY); Mun. P ] d from BR-153 to Itaporá, 12 km W of Presidente Kennedy,
Faz. Primavera, along Ribeiráo Feínho, ca 3°25'S, 48°37'W, 400—500 m, e Feb 1980 (fr), Plowman et al. 8259 (NY, UB); Transamazónica,
junction of Nazaré, 10 Sep 1973 (fl), Rizzo 9273 (NY, UB, UFG). Gorás: Mun. Niquelándia, dirt road 27 km from Colinas do Sul, near
Rio Tocantinzinho, 14°11'S, 48°6'W, 480 m, 6 May 1998 (imm fr), D. Alvarenga et al. 1164 (IBGE, NY); Mun. Nova Roma, road towards
laciara, Faz. Santa Clara, 485 m, 13%45'S, 46°51'W, 29 Feb 2000 (fr), D. Alvarenga et al. 1286 (CEN, IBGE, NY); Chapada dos Veadeiros,
13 km by rd S of Terezina, ca 1,000 m, 16 Mar 1973 (fr), Anderson et al. 7254 (NY, UB); Serra Geral do Paraná, 7 km by rd of São Joao da
Aliança, 1,100 m, 22 Mar 1973 (fr), Anderson et al. 7677 (NY, UB); Mun. Sao Domingos, near Água Quente, Faz. Sáo Domingo, 13°37'S,
+6%44'W, 27 Oct 2000 (1D), M.A. da Silva et al. 4558 (IBGE, NY); Mun. Campinacu, Córrego Laginha, 13°47'S, 48°14'W, 495 m, 9 Oct
1991 (1D, T.B. Cavalcanti et al. 901 (CEN, NY); Mun. Alto Paraiso, São Jorge, dirt road to Vale da Lua, 14%10'S, 47°46'W, 870 m, 19 Sep
1998 (£D), Delprete et al. 6703 (BR, CAY, GB, HTO, NY, RB, UB, UFG); Mun. Alto Paraíso, Parque Nacional E dos Veadeiros, Faz.
Murungú, 23 Sep 1998 (fT), Delprete et al. 6772 (BR, GB, HTO, NY, UB, UFG); Mun. Mineiros, P E road from Água
Ruim to Furna Grande, 18°8'S, 52°44'W, 680 m, 5 Oct 1998 (st), Delprete et al. 6817 (K, MO, NY, UB, UFG); Mun. Caiapónia, Serra dos
Caiapós, road Caiapónia-Montevidiu, 5-20 km de Caiapônia, 17°5'S, 51°44'W, 680—700 m, 12 Oct 1998 (fl, imm fr), Delprete et al. 6828
(GB, K, NY, NX, UB, UFG); Mun. Pirenópolis, Parque Estadual da Serra dos an e do Morro do Cabeludo, 15°48'S, 48°49'w,
1280 m, 26 Nov 2005 (ID, Delprete et al. 9323 (RB, UFG); Mun. Pirenópolis, F bra Rabicho, Córrego das Contenas, 15°44'S,
49°2'W, 8 Jul 2006 (fr), Delprete et al. 9850 (NY, UFG); Mun. Mossámendes, Serra Dourada, Reserva A LE Rizzo, near housing,
26 Oct 1997 (f1), Gomes-Klein et al. 3308 SR bos 3320 (NY, UFG); Serra dos Cristais, 17°S, 48°W, ca 2 km N of Cristalina, 1,250
m, 5 Nov 1965 CD, Irwin et al. 9905 (NY. Preto, 1695, 47°W, ca 3 km E of Cabeceiras, 1,000 m, 19 Nov 1965 (fr), Irwin
et al. 10566 (NY, UB); Mun. Pirenópolis. Sn dos Pireneus, 1695, 49°W, Dec 1965 (ID, Irwin et al. 10869 (MO, NY, RB, UB); Serra dos
Cristais, 17°S, 48°W, 10 km W of Cristalina, 1,200 m, 4 Mar 1966 (fr), Irwin et al. 13448 (NY, UB); Serra Geral de Goiás, 14°S, 46°W, Rio
da Prata, 6 km S of Posse, 800 m, 6 Apr 1966 (fn), Irwin et al. 14419 (NY, UB); Mun. Monte Alegre, Faz. Nica, 530 m, 13?8'S, 46°39'W, 30
Oct 2000 (fr), F.C.A. Oliveira et al. 1159 (CEN, IBGE, NY); Mun. Goiânia, rod. Goiánia-Guapó, Córrego Pindaiba, 3 ae 1968 (£D, Rizzo
& Barbosa 2075 (NY, UFG); Mun. Brazlândia, Fazenda Alegre, 2 Oct 1990 (fl), A.H. Salles 1758 (HEPH, IBGE, NY). DISTRITO FEDERAL
Reserva Ecológica do IBGE, área da M 26 pide 1977 d et al. 122 (IBGE), 23 Oct 1978 (fD, 681 (IBGE), 11 Oct 1982 (m.
7525 (IBGE, UB); ca 5 km ^ Pl , 1,200 m, 16 Oct 1965 (fD, Irwin et al. 9248 (NY); a
of Riberáo Bananal with Lagóa Paranoá, 975 m, 13 Jan 1966 (fr), Irwin et al. 11627 (NY, UB); perto do Riberáo Sobradinho, 15%44
47°44'W, 1,000 m, 28 Sep 1981 pd ddp 4467 (NY, d Faz. Vale Verde, perto do Córrego Acude, 15%39'S, 47°28'W, 1050 m, 26
Aug 1981 (fD, Kirkbride 4515 (NY ; Fazenda Água Li mpo Experimental da UnB, 15°58'S, 47°55'W, 26 Sep 1983 (fl), Kirkbride
& L.V. Ferreira 5397 (NY); Brasilia, Parque Ecológico Nous m Marx, 23 Oct 2003 (fl), J.R. Santos & G.A. Moreira 120 (CEN). Minas
Gerais: Mun. Paraopeba, 30 Nov 1965 (f), R. Goodland 251 (NY); ca 25 km W of Montes Claros, road to Água Boa, 1,000 m, 23 Men 1969
(fr), Irwin et al. 23719 (NY); Mun. Patrocínio, ca 4 km N of Patrocínio, 1,000 m, 31 Jan 1970 (fr), Irwin et al. 25735 (NY); Mun. Formoso,
Parque Nacional Grande Sertáo Vereda, Fazenda Mato Grande, C fato Grande, near waterfall, 770 m, 15?19'S, 45°59'W, 18 Oct
1997 (fl), R.C. Mendonça et al. 3200 (IBGE, NY); Mun. visa , Fazenda do Diamante, base of Serra do Anjico, 600 m, 15 Apr 1931 (fn),
Y. Mexias 5630 (NY); Estrada Trés Marias a Corinto, km 6 dáo, 30 Nov 1976 p, GJ. Shepherd et al. 3827 (NY). SAo PAULO:
Mun. Botucatú, 18 km N, 14 km E of Sáo Manuel-Piracicaba highway, y station Treze de Maio, 22°45'S, 48?25"W, 550 m,
I. Gottsberger & G. Gottsberger 2222-1 (BOTU nv., GI n.v., NY, UB).
Tocoyena brasiliensis Mart. , Flora 24 II, Beibl. 6:82. 1841. Tyre: BRAZIL: Rio DE Janerro: “Monte Telegraphi prope Sebas-
tianopolin,” June [s.d.], Luschnath in Martius Herb. Fl. Brasil. s.n. (HOLOTYPE: BR).
Taxonomic observations.—Tocoyena brasiliensis was treated as synonymous to T. sellowiana (Cham. & Schltdl.)
K. Schum. by Prado (1987). Silberbauer-Gottsberger et al. (1992) treated it as a distinct species because of
the "corolla tube entrance villous; leaves obovate, 15-17 x 6—7.5 cm (vs. “corolla tube entrance glabrous;
leaves oblong-lanceolate, smaller, mostly 7-9 x 2.5-3.5 cm" in T. sellowiana). Delprete (2005) also treated
T. sellowiana as a distinct species, occurring from Bahia to Rio de Janeiro and Santa Catarina. Prado and
Kinoshita (2007) treated them as distinct species, but also wrote that further studies might demonstrate
that in fact they are synonymous.
Tocoyena brasiliensis is also similar to T. formosa, from which it is easily distinguished by the leaf blades
completely glabrous or rarely with puberulent secondary veins POINT and plane (vs. variously pubescent or
rarely glabrous and microscopically bullate) and for the veg: d reproductive p ing brownish-
black when dry (vs. turning olive green to brown when dry).
Distribution and ecology.—Widely distributed in Brazil, from Amapá, Pará, Maranháo, Pernambuco,
Bahia, Mato Grosso, Tocantins, Goiás, Minas Gerais to Sáo Paulo. Shrub or small tree encountered in gallery
988 J I of the Botanical R h Institute of Texas 2(2)
forests sometimes seasonally inundated, and less frequently in mesophytic forests, shrubby savanna and
rocky outcrops with superficial water table.
Selected specimens examined: BRAZIL: Tocantins: Mun. Couto-Magalhaes, road Couto Magalháes-Colinas de Tocantins, 5 km from
Couto Magalháes, 8°19'29"S, 49°9'25"W, 200 m, 4 Nov 2005 (fl), Delprete et al. 9266 (CAY, HPL, HTO, NY, UFG); Mun. Goiatins, Aldeia
Rio Vermelho, 230 m, 8°2'S, 47°15'W, 16 Apr 2001 (fr), E.S.G. Guarino & J.P. Krahó 750 (CEN); 10 km S of Araguaína, 300 m, 16 Mar 1968
(fr), Irwin et al. 21237 (NY, UB); Mun. Lajeado, Faz. Ribeiráo dos Mares, Rio Tocantins, 0788531, 8915570 UTM, 22 Oct 1999 (ID, S.F.
Lolis et al. 318 (HTO); Mun. Porto Nacional, 10 km from Posto do Trevo, 18 Nov 1997 (£D, L. Silva et al. s.n. (HTO, UFG); Mun. Miracema
do Norte [now Miracema do Tocantins], Aldeia dos Índios Xerente, 20 km from Miracema do Norte, 200 m, 9°53'S, 48°36'W, 11 Apr
1988 (st), L.A. Skorupa & J.N. Silveira 431 (CEN); Mun. Palmas, Rio Tocantins, 9°59'39"S, 48°20'49"W, 9 Oct 2000 (fD, E.A. Soares et
al. 1013 (HTO); Mun. Lajeado, road Palmas-Lajeado, 9°57'S, 48°20'W, 10 Oct 2001 (£D, E.A. Soares et al. 1623 (HTO); Mun. Brejinho de
Nazaré, Faz. Santa Angelina, Reserva 1, 11%6'47"S, 48°41'20"W, 5 Nov 2001 (f1), E.A. Soares et al. 1696 (HTO). Go1ás: Mun. Niquelandia,
Faz. Limoeiro, 420 m, 25 Nov 1992 (fD, S.P. Cordovil et al. 172 (CEN); Mun. Mineiros, Parque Nacional das Emas, Rio Jacuba, 17°54'S,
52°52'W, 790 m, 2 Oct 1998 (£D, Delprete et al. 6802 (GB, K, NY, RB, UB, UFG); Mun. Mineiros, Parque Nacional das Emas, área 3, 21
Oct 1989 (fD, H.D. Ferreira 2093 (NY, UFG); Serra dos Pireneus, 16°S, 49°W, ca 12 km 5 of Corumbá de Goiás, 1,000 m, 1 Dec 1965
(£D), Irwin et al. 10869 (NY, UB); Mun. Alvorada do Norte, Fazenda Largomar, 550 m, 14°31'S, 46°34'W, 28 Nov 2003 (£D, A.C. Sevilha et
al. 3616 (CEN); Mun. Serranópolis, RPPN Pousada Ni Araras, 51°59'W, 1892615, 560 m, 25 Oct 2005 (fl), L.F. Souza et al. 1232 (UFG);
Mun. a Novas, eee 9, ae ae Rio Mr 17°42'S, 48°32'W, 720 m, 27 Oct 1993 (fl), R.F. Vieira et al. 1740 (CEN).
ili v 1978 (£D, Heringer 17172 (IBGE, UB); just outside of Reserva Biológica
Aguas Emendadas, 3 Sep 1995 (fI), C. Proença et al. 1500 uem Maro Grosso: Mun. Santa Terezina, Faz. Codeara, Serra da Viúva, 12
km S of Santa Terezina, 10°40'S, 50°35'W, 10 Oct 1985 (fl), C.A. Cid Ferreira et al. 6404 (NY); Mun. Brilhante, Fazenda Progresso, 28
Oct 1970 (£D, Hatschbach 25286 (NY). Maro Grosso po Sur: Mun. Trés Lagoas, Fazenda Dr. José Mendes, 12°30'S, 40°29'W, 20 Oct
1964 (fD, J.C. Gomes 2369 (NY)
Tocoyena viscidula Mart., Flora 24 II, Beibl. 6:80. 1841. Tre: BRAZIL: Banu: "in silvis Caatinga mediterraneis," s.d., Martius
s.n. (HOLOTYPE: M n.v.; photo-M at UEC n.v., UFMT).
Taxonomic observations.—Schumann (1889) treated Tocoyena viscidula Mart. as a synonym of T. formosa,
without any additional comments. Prado (1987) and Silberbauer-Gottsberger et al. (1992) recognized this
species as rare and distinct, mostly because of observations made on 20th century collections. Accordingly,
it is treated here as a distinct species, easily recognizable by its leaf blades densely lanate on both sides.
Distribution and ecology.—A species mostly collected in the State of Goiás and the Distrito Federal,
with just a few collections known from the states of Bahia (where the type is from) and Piauí. A small shrub
10-70(200) cm tall, encountered only in shrubby fields (“campos sujos”) and rocky outcrops (“campos
rupestres") within the Caatinga and Cerrado Biomes.
Conservation.—A rare species that needs to be added to the category of critically endangered taxa, as
the last collections known were made in 1980—81. The habitats of this species are in need of an urgent
conservation program.
Phenology.—Flowering specimens were collected during the rainy season, from August to November,
and the only fruiting collecting known was made in February.
Specimens examined: BRAZIL: Gorás: Mun. e Serra dos Cristais, 7 Oct 1981 (fl), Hatschbach 44060 (MBM, UB); Mun. São
si da n 21 km from Nossa Senhora d Gabriel de Goiás, 1,000 m, 14 Oct 1980 (fD, G. Martinelli et al. 7485 (UB). DISTRITO
DERA enda Agua Limpa, Campus da UnB, 47°56'S, 15°57°W, 16 Oct 1979 (ID, César 638 (UB); Brasilia, TV Record, 23 Oct 1978
i ns etal. 681 (UB); s Saia Velha, R-1, 10 Feb 1961 (fr), Heringer 8135/329 (UB); Brasília, D.N.E.R., 10 Oct 1962 (ID, Heringer
9046 (NY, UB); Brasilia, Saia Velha, 22 Nov 1964 (fl), Heringer 9964 (NY, RB, UB); Brasilia, Catetinho, 15 Nov 1973 (fl), Heringer 13005
(UB); Fazenda Água Limpa, Campus da UnB, Vargem Bonita, ca 18 km from Brasilia TV tower, 8 Oct 1976 (£D, Ratter et al. 3736 (UB).
mE arenicola Delprete, sp. nov. (Fig. 2). Tree: BRAZIL: Tocantins: Mun. Miracema do Tocantins, road towards Lajeado
-070), 3 km from Miracema do Tocantins, cerrado vegetation on white sand, 9°35'13"S, 48°24'32"W, 230 m, 3 Nov 2005 (fl, fr),
PG. DA H. Lorenzi, J.A. Lombardi & E.G. Goncalves 9249 (noLorveE: UFG; isotypes: CAY, HPL, MO, NY, RB, UB).
PE £ pa A Aft ££. tiri 0.3— L In lih XO ]5 ] 1 1 imi vel
1 1 1 *AT 1 430 fhilt J ] ]
arboris dedu ad 10m( i ylop
horizontale ortam ciet praeterea corollis 3. PR 6.5 cm longis (nec 8.5-14 cm longis) et laminis foli inoribus | distinct
] 1 dium; stems simple
Subshrubs to small shrubs 0.3-1 m tall, with 5-15 st iginating from a
or few-branched, terete or slightly laterally compressed, Lu ceni to oe bark papyraceous,
Delprete, Revision of Tocoyena
B
989
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Fic. 2
ic R Detail af al
Delprete et al. 9249, UFG).
990 j Vaf tho Botanical R h Institute of Texas 2(2)
pale brown to reddish-brown. Stipules narrowly triangular, oblong-triangular to lanceolate, 6-12 x 3-4
mm, acute to acuminate, glabrous or sparsely-pubescent outside, margins glabrous or ciliate, covered with
coleters at basal portion and glab distally inside, persistent. Leaves subsessile to short-petiolate; petioles
2-10 mm long, canaliculate, densely pubescent; blades elliptic, oblong-elliptic to obovate, 1.5—6.5(-10.5)
x 0.5-3(-5.5) cm, acute at base, obtuse to acute at apex, papyraceous to subcoriaceous, plane or micro-
bullate, sparsely to densely pubescent above, tomentose-pubescent below, olive green to dark brown above
and olive green below when dry; with 8-11 secondary veins each side. Inflorescences condensed-cymose,
3-11-florous; sessile to short-pedunculate, peduncles to 10 mm long; rachis antrorse-pubescent, hairs pale
brown; secondary branchlets 1-7 mm long; bracts deltoid to narrowly triangular, 0.5-3 mm long, glabrous
or sparsely pubescent, persistent. Flower buds pale green, lobed portion obtuse. Flowers sessile (terminal
ones) or subsessile (lateral ones). Hypanthium turbinate, 3.5—4.5 mm long, glabrous or sparsely pubescent.
Calyx cupular or short-tubular, 3-5 mm long, 5-lobed, glabrous or sparsely pubescent outside, with a few
colleters among the teeth inside; lobes triangular to lanceolate, 1.5-3.5 mm long, ciliate. Corolla hypo-
crateriform, 3.6—6.5 cm long, white or cream-white, turning yellow at final stage of anthesis, hirtellous to
erect-pubescent outside, glabrous inside, dark brown when dry; tube narrowly cylindrical, 3-6 cm long,
slightly ampliate near the mouth; lobes elliptic to oblong-ovate, 6-10 x 8—9 mm, slightly asymmetrical,
obtuse, puncticulate inside, margins ciliate. Stamens sessile to subsessile; anthers oblong, 4—5 mm long,
shortly-apiculate, base bilobed. Style exserted, 3.5-6.5 cm long, thickened-clavate distally, glabrous; style
branches narrowly obovate, 4-4.5 mm long, dorsally tricostate. Fruit globose or subglobose, 1.8-2 cm in
diam, smooth (not ribbed), green and puberulent when young, yellow and glabrescent when ripe, turning
black when dry.
Taxonomic observations.—Tocoyena arenicola Delprete is similar to T. viscidula in being a subshrub with
a basal xylopodium, from which it is easily distinguishable by the characters given in the key (see below).
However, because of the vestiture of the vegetative and reproductive parts it is most similar to T. formosa with
which it has sometimes been confused. Tocoyena arenicola differs from T. formosa because it is a subshrub
to small shrub 0.3-1 m tall, with 5-15 stems originating from a single xylopodium (vs. shrub or treelet to
tree (0.5—)3—10(-12) m tall, commonly with a main trunk up to 20 cm dbh or exceptionally reduced to a
single basal rosette of leaves and a horizontal xylopodium), with corollas 3.6—6.5 cm long (vs. 8.5-14 cm
long), and leaf blades 1.5—6.5(-10.5) x 0.5-3(5.5) cm (vs. (7-)9-20(-24) x (528-15 cm).
Distribution and ecology.—A species known from a few collections in the extreme north of the state of
Goiás (Mun. Posse) and the central-southern region of the state of Tocantins (Mun. Dianópolis and Miracema
do Tocantins). Subshrub to small shrub, heliophilous, with a short basal xylopodium, and found only in
shrubby fields ("campos sujos") on quartzitic white sands within the Cerrado Biome, sometimes in associa-
tion with Melocactus spp. (Cactaceae) and Syagrus acaulis (Drude ex Mart.) Becc. (Palmae). The vegetation
present in these white-sand areas contains species also common in the Cerrado Biome, but stunted and
much smaller than when growing in the surrounding savanna with regular red soil (pers. obs.). In addition,
it also contains a suite of species endemic to these areas, and Tocoyena arenicola is one of those.
Conservation.—Known by just a few collections, it is a rare species that must be placed in the category
of imminent threat of extinction. This is supported by the fact that white-sand areas in the Cerrado Biome
are primarily utilized for the construction of houses and buildings, as they are not suitable for agriculture
and cattle ranching. In general, these areas should be urgently conserved, as they support many rare and
endangered species of plants and animals.
Phenology.—Flowering and fruiting collections were realized from September to November.
Additional specimens examined: BRAZIL: Goiás: Mun. Posse, Nova Vista, campo cerrado, 8 Oct 1976 (fl), Hatschbach 39048 (MBM,
NY); Mun. Posse, di 1 irj cerrado recently burned, sandy soil, 820 m, 14°6'S, 46°20'W, 16 Oct 2001 (f1), R.C. Mendonca et al.
4450 (CEN, IBGE); Mun. Posse, road between Guarani and Posse, sandy soil, 830 m, 14°57'S, 46°22'W, 19 Oct 2001 (f1), R.C. Mendonca
et al. 4508 (CEN, IBGE). Tocantins: Mun. Dianópolis, op d quartzitic sand, 660 m, 11°37'S, 46°26'W, 28 Sep 2003 (f1), T.B.
Cavalcanti et al. 3233 (CEN); Mun. Miracema do Tocantins, road towards Lajeado (TO-070), 3 km from Miracema do Tocantins, cerrado
Delprete, Revision of Tocoyena 991
on white sand, 9°35'13"S, 48°24'32"W, 230 m, 3 Nov 2005 (fl, £D, Delprete et al. 9252 (CAY, HTO, NY, SP, UFG); Mun. Dianópolis, area
with quartzitic sand, 720 m, 11%36'S, 46°28'W, 25 Sep 2003 (fD, A.O. Scariot et al. 759 (CEN).
KEY TO THE NATIVE SPECIES OF TOCOYENA
IN THE STATES OF GOIÁS AND TOCANTINS, BRAZIL
1. Subshrub to small shrub 0.1—1(-2) m tall, with 3-15 stems originating from a basal xylopodium.
2. Leaf blades 7-17 x 6-11 cm, densely lanate on both sides; corollas 10-12.5 cm long (tube 85-11
cm long), pia (shrubby fields and rocky outcrops) T. viscidula
2. Leaf blades 1.5-7 x 0.7-3 an e PAP above, sparsely lanate below; corollas 3.6-6.5 cm
long (tube 3- E long), hirt quartzitic white-sand) T. arenicola
1. Shrub, treelet to tree (0.5-)3-10(- 12)n m tall, commonly wit a main oun up to 20 cm dbh (exceptionally
individuals reduced to a basal rosette of leaves and yloj T. formosa, in open fields
in the Parque Nacional das Emas and in ifie Serra dos Caiapós).
3. Leaf blades chartaceous to e Pans compere 9 4PIOUS or rarely with puberulent
Ven a dark brown to blaci T. brasiliensis
3. plne or minutely | ll bubo velutinous-tomentose, hispid
or rarely glabrous above, puc velutinous- -tomentose or rarely glabrous below, grayish, olive green
to pale brown when dry; corollas tomentose, puberulent or glabrescent externally T. formosa
ACKNOWLEDGMENTS
This research was realized during a fellowship for Visiting Scientist from the National Council for Scientific
and Technological Development (Conselho Nacional de Desenvolvimento Científico e Tecnológico - CNPq)
of the Brazilian Government (grant 309885/2003-5), at the Federal University of Goiás (UFG), with the
coordination of Vera Lúcia Gomes-Klein (UFG). Field work in the state of Tocantins was supported by Harry
Lorenzi (HPL), and accompanied by him, Julio Lombardi (HRCB) and Eduardo Goncalves (Catholic Univer-
sity of Brasília). My gratitude also goes to Lubbert Westra (WAG) for the Latin diagnosis and reviewing the
first draft of the manuscript. The directors and curators of the following herbaria are kindly acknowledged
for loan of material, sending digital images, and/or providing working space during my visits: BR, CEN, F,
G, G-DC, HTO, IBGE, K, MO, MBM, NY, NX, R, RB, U, UB, UFG, UFMT, and US. I am very grateful also to
Joseph Kirkbride (USDA) and Claes Persson (GB) for the helpful comments on the manuscript.
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994 Journal of the Botanical R h Institute of Texas 2(2)
BOOK REVIEW
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J. Bot. Res. Inst. Texas 2(2): 994. 2008
CINCO NUEVAS ESPECIES DE PILEA (URTICACEAE) DE COSTA RICA
Alexander Rodríguez Alexandre K. Monro
Instituto Nacional de Biodiversidad | of E
Apartado Postal 22-3100 The Natural History M
Santo Domingo de Heredia, COSTA RICA London, S I. 5BD, UNITED] AT
arodrig@inbio.ac.cr a.monro@nhm.ac.uk
RESUMEN
Ci i le Pilea, endémicas de Costa Rica il las: Pil If; AL Rodr. & A.K. Monro, P. gamboana
Al. Rodr. & A.K. Monro, P. ida Rodr. & A.K. Monro, P. 1 angl lata Al. Rodr., A.K. Monro & L. Acosta, P. moragana Al.
Rodr. & A.K. Monro, ad l género y su ubicación bajo las clasificaciones pe Weddell y Killip.
ABSTRACT
EB f Pilea, endemic to Costa Rica, are described and ill ]: Pilea alf. Al. Rodr. & A.K. Monro, P. gamboana
AL Rodr. GA K. Mon P. Pod Al oa & A.K. Monro, P. longil lata Al. Rodr., A.K. Monro &1 a P. pin Bae io
R ithin Weddell’
Rodr. & A.K. Monro. T Wed
Pilea Lindl. es un género con alrededor de 715 especies (Monro 2004) por el mundo, de las cuales cerca de
36 están presentes en Costa Rica. Es un género pantropical (con la excepción de Australia) con su centro de
diversificación filogenético y morfológico en Asia del Sur y con un número de especies en América del Sur
y Las Antillas Mayores (Monro 2006). En los Neotrópicos, Pilea se distingue de otros géneros de la familia
Urticaceae por la presencia de hoja tas y estípulas intrapeciolares, la ausencia de tricomas urticantes,
el hábito terrestre, epífito o Spio, d periantio de las flores indie con 3 lóbulos desiguales y libres
y por la presencia de apéndices subapicales sobre los tépalos masculinos (éstos a veces ausentes, e.g, en P.
longibracteolata Al. Rodr., A.K. Monro & L. Acosta); además las especies de Pilea son generalmente hierbas
con tallos suculentos que pueden parecer a especies de Begonia.
fa e Rica y Panamá este género ha sido poco estudiado taxonómicamente. Entre las principales
las de Standley (1937) en Flora of Costa Rica, Burger (1977) en Flora Costaricensis
y Killip (1960) en Flora of Panama. En la región Mesoamericana otros aportes importantes son la Flora of
Guatemala (Standley & Steyermark 1952), la sinopsis de Mesoamerica por Monro (2001) y la revisión por
Flora Mesoamericana (Davidse et al. 1994) hecho por Monro (in prep).
Nuevas exploraciones en Costa Rica han resultado en la descripción de las siguientes cinco especies
nuevas.
Pilea alfaroana Al. Rodr. & A.K. Monro, sp. nov. (Fig. 1). Tiro: COSTA RICA. San José: Pérez Zeledón, Parque Nacional
Chirripó, cuenca Térraba-Sierpe, sendero Llano Bonito, 09°27'24"N, 83°32'47°W, 2450 m, 4 May 1997, E. Alfaro, B. Gamboa, R.
Aguilar & A. Picado 1200 (HoLoriPo: INB).
EA 1; 2 T 2 ; 131 4; ££,
1 J
Species P. pubescenti Liebm. affinis,
Hierba erecta o procumbente, 0.25—0.35 m de altura, terrestre, dioica o monoica; tallos subteretes, lige-
ramente acostillados, suculentos, densamente pubescentes, los tricomas 0.75-1.00 mm, simples, erectos,
rectos o curvados, con esparcidas glándulas rojizas y sésiles, los cistolitos fusiformes. Estípulas 7-14 mm
de largo y 1.0—1.2 mm de ancho, obovadas u ovadas a ovado-orbiculares, subcordadas en la base, obtusas
en el ápice, varias veces nervadas a inconspicuamente nervadas, persistentes, adpresas, ascendentes o di-
varicadas, glabras; hojas del mismo nudo subiguales en tamaño; pecíolos 5-8 mm de largo, subiguales a
levemente desiguales, con una relación en tamaño de 1: 1.0-1.7, esparcidamente pubescentes, los tricomas
0.75-1.00 mm, simples, erectos o aplanados, rectos o curvados, con esparcidas glándulas rojizas y sési-
J. Bot. Res. Inst. Texas 2(2): 995 ~ 1007. 2008
996 1 | £s a Datania D h Institut f Texas 2(2)
Silvia Troyo 08
Fic. 1. A-E. P alfaroana Al. Rodr. & A.K. Monro. A. Hábito; B. Flor femenina; C. Flor masculina; D Estípulas: E. Ci
hojas. A, C, D, E, basados en J. Gi 1918 (INB); B, basado en D. $ ía 4072 (
les; lámina 7-90 mm de largo y 6-85 mm de ancho, ovada a ovado-orbicular; base obtusa a cortamente
cuneada; ápice agudo; margen gruesamente dentado; nervadura palmada o subpalmada, con 3 a 5 nervios
saliendo desde la base o hasta 4 mm sobre la misma; haz densamente pubescente, los tricomas 0.5-1.5 mm,
simples, aplanados, rectos o curvados, glabrescente con la edad, con esparcidas glándulas rojizas y sésiles,
los cistolitos fusiformes o en forma de “V,” secando verde a verde-grisáceo; envés esparcida a densamente
pubescente, los tricomas 1.00-1.75 mm, simples, erectos o aplanados, rectos o curvados, con esparcidas
glándulas rojizas y sésiles, los cistolitos fusiformes o en forma de “V,” verde-pálido al secar. Inflorescencias
unisexuales, axilares, 1 por axila, presentes en los nudos terminales, dimorfas; inflorescencias masculinas
15-40 mm de largo y 5-12 mm de ancho, cimas capitadas y simples o poco y cortamente ramificadas, el
pedúnculo 15-25 mm de largo, densamente pubescente, los tricomas 0.75—1.00 mm, simples, erectos, rectos
o curvados, con esparcidas glándulas rojizas y sésiles, basalmente con brácteas pedunculares 1.00-2.25 de
Dad ig JAA , M a Ja D: 997
la largo y 0. Ee 1.25 mm de ancho, ovadas, basalmente glabras, hialinas, generalmente difíciles de observar;
flores en glomérulos, subsésiles o con pedicelos hasta 1.7 mm de largo y glabres , bracteolas
pedicelares 0.75-2.00 mm de largo y 0.20—0.75 mm de ancho, lanceoladas a oblongo-lanceoladas, glabras,
hialinas; inflorescencias femeninas 10—40 mm de largo y 5-25 mm de ancho, cimas simples o varias veces
ramificadas, el pedúnculo 10-30 mm de largo, glabrescente o con esparcidos tricomas ca. 2.5 mm de largo,
simples, erectos, rectos o curvados; flores femeninas distribuidas a lo largo del raquis de la inflorescencia,
subsésiles o con pedicelos hasta 0.5 mm de largo y glabros, bracteolas pedicelares 0.5-0.6 mm de largo y
0.2—0.6 mm de ancho, lanceoladas, glabras, hialinas. Flores masculinas con periantio 0.75-1.50 mm de
largo, glabro o esparcidamente pubescente, los tricomas 0.3-0.5 mm, simples, erectos, rectos, 4-lobulado,
los lóbulos 0.5-1.1 mm de largo, lanceolados a obovados, incisos 1/4—3/4 de la longitud del periantio, los
apéndices subapicales 0.70—1.50 mm de largo; estambres 4, los filamentos 1. i Ms mm d Mcd i anteras
ca. 0.5 mm de largo y 0.35 mm de ancho; pistiloide muy pequeño y ap
lóbulos del periantio levemente desiguales, el dorsal ca. 0.7 mm de largo; MEM 3; estigma diminuto,
penicilado, erecto. Aquenios 0.75-1.00 mm de largo (incluyendo el estigma) y ca. 0.75 mm de ancho, ovoides,
subcompresos, simétricos, verdes a verde-castaños, con una línea submarginal puncticulada.
Fenología. —Flores y frutos entre abril, mayo y junio.
Distribución y Hábitats. —Endémica de Costa Rica. Se conoce de los bosques pluviales de la vertiente
Pacífica en la cordillera de Talamanca, en Copey de Dota, en Llano Bonito del Parque Nacional Chirripó y
en Tres Colinas de Potrero Grande, 1500-2450 m. Esta especie es conocida de pocas localidades y se ha
observado creciendo entre robledales, principalmente a orillas de quebradas donde podría formar parches
densos.
Discusión. —Pilea alfaroana, según la clasificación de Weddell (1869), pertenece al grupo Dentatae y
según la clasificación de Killip (1900) al grupo Molles. Se reconoce por ser una hierba terrestre, en general
procumbente, con partes veg densamente pubescentes, hojas isófilas, margen gruesamente dentado,
estípulas conspicuas, hialinas, persistentes, obovadas u ovadas a ovado-orbiculares, cordadas en la base,
obtusas en ápice, por sus flores masculinas con periantio generalmente pubescente, por sus frutos con una
línea submarginal puncticulada y por presentar una distribución muy localizada.
En apariencia vegetativa es similar a P. pubescens Liebm. considerando que ambas especies poseen un
hábito regularmente procumbente y terrestre, pubescencia densa y hojas isófilas, ovadas a ovado-orbicular
y dentadas. Se distinguen así:
1. Ns 2-5 mm de largo, haz de la hoja Pss los tricomas 2-3 mm, inflorescencias bisexuales,
0-120 mm, TO ue bu 200 msn Pilea pubescens
Nod 7-14m „l ¡ 0.5-1.5 mm, inflorescencias unisex-
uales, 15-40 mm, Meise: ente 1500-2450 msnm Pilea alfaroana
Etimología.— Esta especie es nombrada en honor a Evelio Alfaro, un excelente y animoso parataxónomo que
aportó la colección Tipo de esta especie y trabajó en el INBio entre los años de 1995 y 2003, donde hizo
una colección con más de 5500 especimenes botánicos, entre otros logros significativos. Lastimosamente
en la actualidad se mantiene alejado de la actividad botánica, se estableció en Estados Unidos procurando
mejorar su condición económica.
Material e inado. COSTA RICA. Puntarenas: Buenos Aires, Potrero Grande, Tres Colinas, 09°06'17"N, 83°03'54"W, 1500 m, 25
Abr 2006, D. Santamaría 4072 (INB). San José: Pérez Zeledón, Reserva Forestal Los Santos, cuenca del Savegre, San Gerardo de Dota,
sendero Los Robles, 09°32'50"N, 83°49'00"W, 2200-2350 m, 22 May 1197, J. González & M. Zumbado 1918 (INB); Dota, Copey, Área
no Protegida, San Gerardo, sendero La Quebrada, 09°23'10'N, 83?48'23"W, 2300 m, 5 Jun 2002, A. Quesada 1017 (CR); Rivas, Parque
Nacional Chirripó, sector Llano Bonito, alreded le la casa, bosques de robledales, 09°27'34'N, 83°32'47"W, 2450 m, 30 Ago 2000,
A. Rodríguez 6383 (INB).
Pilea gamboana Al. Rodr. & A.K. Monro, T nov. (Fig. 2). Tiro: COSTA RICA: San José: Pérez Zeledón, Cuenca Térraba-
Sierpe, Puesto Tres Colinas, colectado a orillas del río G bosque primario, 09°07'27"N 83°04'29"W, 1700 m, 17 Set 1996, B.
Gamboa & A. Picado 639 (notorio: INB; isoriPo: MOS
rl ah feu 1 1 A e D 1:3
Species P. pittieri Killip affinis, p p
998 Journal of the Botanical R h Institute of Texas 2(2)
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Fic. 2. A-D. P. gamboana Al. Rodr. & A.K. Monro. A. Hábito; B. Flor femenina; C Flor masculina; D. Estípulas. A, D basados en B. Gi
basado en G. Herrera 3509 (INB).
Dar íg JRA , AE H Aa Dil 999
Hierba erecta o procumbente, 0.3—0.5 m de altura, terrestre o epilítica, dioica o monoica; tallos subteretes,
glabros, con m aa glándulas rojizas y sésiles, los cistolitos fusiformes. Estípulas 3-4 mm de largo y
1.0-1.5 m ancho, oblongas a oblongo-obovadas, agudas a obtusas en el ápice, con nervadura inconspicua,
decíduas o persistentes en los nudos terminales, ascendentes, glabras; hojas del mismo nudo subiguales,
con una relación en tamaño de 1: 1-3 (incluyendo el pecíolo); pecíolos 4-100 mm de largo, subiguales o
desiguales, con una relación en tamaño de 1: 1.3-3.0, glabros, ocasionalmente con esparcidas glándulas
rojizas y sésiles; lámina 10-150 mm de largo y 4-75 mm de ancho, ovada o elíptica a ovado-elíptica; base
subpeltada a peltada; ápice acuminado; margen cortamente dentado o denticulado; nervadura palmada o
subpalmada, con 3 ó 5 nervios evidentes saliendo desde la base o hasta 2 mm sobre la misma; haz y envés
glabrescentes, los cistolitos fusiformes o en forma de “V” sobre ambas superficies, secando verde-grisáceos,
verde-pajizos o rojizos. Inflorescencias unisexuales, axilares, 1 6 2 por axila, subiguales; inflorescencias
masculinas 10-45 mm de largo y 10-30 mm de ancho, cimas paniculadas, el pedúnculo 10-25 mm de
largo, glabro, basalmente con brácteas pedunculares 0.75—1.50 mm de largo y ca. 1.0 mm de ancho, del-
toides, glabras; flores masculinas con pedicelos 0.30-0.75 mm de largo, glabros, bracteolas pedicelares
0.30—0.75 mm de largo y 0.25-0.40 mm de ancho, carentes de nervios evidentes, glabras; inflorescencias
femeninas 10-40 mm de largo y 5-40 mm de ancho, cimas varias veces ramificadas, el pedúnculo 4-20
mm de largo, glabro, basalmente con brácteas pedunculares 0.50-0.75 de la largo y 0.50—0.60 mm de an-
cho, deltoides; flores femeninas subsésiles o pedicelos hasta 1 mm de largo, glabros, bracteolas pedicelares
0.30—0.75 mm de largo y 0.25-0.40 mm de ancho, carentes de nervios evidentes, glabras. Flores masculinas
con periantio 1.00-1.25 mm de largo, glabro, 4-lobulado, los lóbulos 0.75—1.00 mm de largo, oblongos a
oblongo-lanceolados, incisos ca. 3/4 de la longitud del periantio, los apéndices subapicales 0.10—0.25 mm
de largo; estambres 4, los filamentos 1.00-1.25 mm de largo, las anteras 0.4—0.5 mm de largo y 0.3-0.4
mm de ancho; pistiloide muy reducido; flores femeninas con lóbulos del periantio levemente desiguales, el
dorsal 0.50—0.75 mm de largo, los laterales 0.3-0.4 mm de largo; estaminodios 3, ca. 0.12 mm de largo y
de ancho, petaloides; estigma inconspicuamente penicilado, erecto. Aquenios 0.75-1.00 mm de largo (in-
cluyendo el estigma) y 0.60—0.75 mm de ancho, ovoides a elipsoides, sul I , Simétricos a levemente
asimétricos, café-pajizos, lisos.
Fenología.—Poco conocida, hasta el momento flores y frutos sólamente observados en septiembre y
octubre.
Distribución y Habitats —Endémica de Costa Rica. Se conoce en los bosques pluviales de la vertiente
Pacífica en la Cordillera de Talamanca, 1450-1750 m. La especie se ha observado creciendo principalmente
a orilla de ríos y quebradas.
Discusión. —Pilea gamboana, según la clasificación de Weddell (1869), pertenece al grupo Dentatae y
según la clasificación de Killip (1960) al grupo Molles. Se reconoce por el hábito terrestre o epilítico, por sus
hojas con la base subpeltada, las del mismo nudo subiguales, con el margen levemente dentado, las estípulas
pequeñas, tempranamente deciduas, por las flores masculinas y femeninas en cimas paniculadas y por su
localizada distribución.
Vegetativamente es similar a Pilea pittieri Killip, una especie igualmente endémica de Costa Rica, debido
a que ambas son plantas glabrescentes, por el tamaño y forma de las estípulas, por sus hojas subiguales y
lámina con frecuencia ovada. Sin embargo, se puede distinguir por los carácteres siguientes:
1. Hojas basifijas, flores masculinas dispuestas en cimas capitadas, el periantio con apéndices subapicales
0.5-2.0 mm Pilea pittieri
D^ f
. Hojas subpeltadas, fl linas di t i iculadas, el periantio con apéndices subapica-
les 0.10-0.25 mm Pilea gamboana
Etimología.—Esta especie es nombrada en reconocimiento de Billen Gamboa, un excelente parataxónomo
del INBio que ha recolectado ampliamente en Costa Rica, principalmente en la cordillera de Talamanca.
Material examinado. COSTA RICA. Puntarenas: Buenos Aires, Reserva Indigena Ujarrás, oo a EDAM CUM pop
Dúrika, 09?16'30"N 83%14'40"W, 1500 m, 6 Set 1995, R. Aguilar et al. 4295 (CR, INB); Ujarrás yé, sig
1000 J I of the Botanical R h Institute of Texas 2(2)
que dan a Olan, 09°17' aaa oe 45"W, 1450 m, a e Po G. Herrera 3509 (INB, MO). San José: Pérez Zeledón, cuenca Térraba-
Sierpe, casi km 116 a 1 rretera, das, 09°28'40"N, 83°41'25"W, 1750 m, 7 Oct 1997, B. Hammel 21094
& H. Kennedy 21094 (INB, CR, MO, BM).
Pilea herrerae Al. Rodr. & A.K. Monro, iu nov. (Fig. 3m ud COSTA RICA: Limón: Cantón de Guápiles, Los Angeles, San
Miguel, siguiendo el camino al Volcán Tra j g do, 10°06'50"N, 83°49'30"W, 1100 m,
26 Feb 1990, G. Herrera 3851 Acces INB: IsoTIPO: CR).
Species P. costaricensi Donn. Sm. affinis, sed ab ea inflorescentiis capitatis, floribus staminalibus majoribus et lobulo dorsali florium
pistillatorum majore differt.
Hierba erecta o procumbente, 0.4—1.0 m de altura, terrestre, dioica o monoica; tallos subteretes, suculentos,
glabrescentes, con esparcidas glándulas rojizas y sésiles, los cistolitos escasos, punticulados o cortamente
fusiformes. Estípulas 1-2 mm de largo y 0.50-0.75 mm de ancho, lanceoladas, agudas en el ápice, la ner-
vadura inconspicua, persistentes, ascendentes, glabras; hojas del mismo nudo anisófilas, con una relación
en tamaño de 1: 1.6-23.0 (incluyendo el pecíolo); hojas menores en ocasiones ausentes, subsésiles o con
pecíolos hasta 3 mm de largo, glabros, la lámina 3-17 mm de largo y 5-10 mm de ancho, linear-lanceolada
o elíptica; hojas mayores con pecíolos 5-15 mm de largo, glabros, con esparcidas glándulas rojizas sésiles, la
lámina 30-190 mm de largo y 10-55 mm de ancho, lanceolada, elíptico-oblanceolada o falcado-lanceolada;
base obtusa, levemente oblicua o a menudo subpeltada; ápice acuminado; margen entero a raras veces den-
ticulado; nervadura 3-plinervada a subpalmada, con 3 nervios evidentes saliendo 2-4 mm sobre la base;
haz y envés glabrescentes, a menudo con esparcidas glándulas rojizas y sésiles, los cistolitos fusiformes oen
forma de "V secando gris, gris-oscuros o verde-grisáceos. Inflorescencias unisexuales, axilares, 1 6 2 por
axila, monomorfas a ligeramente dimorfas; inflorescencias masculinas 5-50 mm de largo y 5-12 mm de
ancho, cimas capitadas, raras ocasiones cimas 2 6 3 veces cortamente ramificadas, el pedúnculo 2-45 mm
de largo, glabro, en ocasiones con esparcidas glándulas rojizas y sésiles, basalmente con brácteas peduncu-
lares 0.3-0.5 mm de la largo y 0.3-0.5 mm de ancho, ovadas, glabras; flores masculinas subsésiles o con
pedicelos hasta 1.75 mm de largo, glabros, con esparcidas glándulas rojizas y sésiles, bracteolas pedicelares
0.5-1.0 mm de largo y 0.25—1.00 mm de ancho, lanceoladas, glabras; inflorescencias femeninas en plantas
monoicas ubicadas en los nudos inferiores, 10-30 mm de largo y 5-12 mm de ancho, el pedúnculo 5-26
mm de largo, glabro; flores femeninas subsésiles o pedicelos hasta 2 mm de largo, glabros, bracteolas pedi-
celares 0.3—0.5 mm de largo y 0.5—0.7 mm de ancho, lanceoladas, glabras. Flores masculi n periantio
1.50-2.25 mm de largo, glabro, 4-lobulado, los lóbulos 0.75-1.00 mm de largo, incisos ca. 3/4 de la longitud
del periantio, los apéndices subapicales 0.50—0.75 mm de largo; estambres 4, los filamentos 1.25-1.50 mm
de largo, las anteras ca. 1.25 mm de largo y 0.75 mm de ancho; pistiloide reducido; flores femeninas con
lóbulos del periantio levemente desiguales, el dorsal 1.5-3.0 mm de largo, los laterales 1.25-1.50 mm de
largo; estaminodios 3, ca. 0.75 mm de largo y 0.7 mm de ancho, petaloides; estigma diminuto, penicilado,
erecto. Aquenios 2.0-2.5 mm de largo (incluyendo el estigma) y 0.25-1.75 mm de ancho, ovados, subcom-
presos, simétricos, café, lisos.
Fenología.—Flores y frutos en enero, febrero, marzo y abril, además en agosto, septiembre y noviembre.
Distribución y Hábitats. —Endémica de Costa Rica. Se conoce de los bosques húmedos y pluviales de la
vertiente Caribe en la cordillera de Talamanca, 700-1700 m.
Discusión. —Pilea herrerae, según la clasificación de Weddell (1869), pertenece al grupo Heterophyllae
y según la clasificación de Killip (1960) al grupo Centradenoideae. Se reconoce por ser terrestre y con hojas
del mismo nudo conspicuamente anisófilas, el D generalmente entero, la nervadura 3-plinervada a
subpalmada, en ocasiones las hojas más larg; (difícil de notar en especímenes), por sus estípulas
diminutas y por presentar flores en cimas M no ramificadas.
Esta especie es muy similar a Pilea costaricensis Donn. Sm., la cual también presenta hojas del mismo
nudo anisófilas y el margen de la lámina entero a levemente dentado. Se puede distinguir por los carácteres
siguientes:
n. Jr Dna ,N E da Dal 1001
2
[2]
o
>
E:
m
2
[7]
Fic. 3. A-E. P. herrerae Al. Rodr. & A.K. Monro. A. Hábito; B. Flor femenina; C. Flor masculina; D. Estípulas; E. Cistolitos en el envés de las hojas. A, D, E,
basados en J, Bittner 1456 (INB); B, C, basados en A. Rodríguez 11066 (INB).
1. Infl li i i mificadas, 5-10 mm de largo, flores masculinas con un pe-
riantio de 1.00-1.25 mm, flores femeninas con s lóbulo dorsal 1.0-1.5 mm, aquenio 0.5-1.0 mm de largo,
penalo, de las hojas mayores 9-45 mm Pilea costaricensis
| i m de largo, flores A
con un periantio de 1.50- 225 mm, flores iii con el lóbulo dorsal 1.5-3.0 mm, aquenio 2.0-2.5 m
de largo, pecíolos de las hojas mayores 5-15 m Pics herrerae
El tipo era erróneamente identificado como P. purulensis por Monro en 1999.
Etimología.— Gerardo Herrera, un botánico costarricense contemporáneo, es, sin la menor duda, uno
de los mejores recolectores. Su esfuerzo en colecta de plantas ha concluido en mültiples novedades para la
ciencia, incluyendo ésta, por tal razón dedicarle esta especie es un reconocimiento a su valioso esfuerzo.
Material examinado. COSTA RICA. Cartago: Jiménez, Pejibaye, Refugio de Vida Silvestre La Marta, 09?47'15"N 83°43'30"W, 700—800
m, 29 Set 2004, R. Kriebel & A. Soto 4877 (INB); Selva, Reserva Biológica EI Copal, 09?47'10"N 83%45'06"W, 1100—1200 m, 30 Ene 2008,
1002 Journal of the Botanical R h Institute of Texas 2(2)
G. Wood €» D. Wood 8058 (INB); Turrialba, Parque Nacional Barbilla, cuenca del Matina, Sendero Barthon, Punta Fría, Sección Corina-
Cerro Tigre, 09°55'15"N 83°23'55"W, 1600 m, 6 Mar 2001, E. Mora et al. 1835 (CR). Limón: Talamanca, Camp by Kivut, trail between
Kivut and Alto Lari, 1-2 km up to Alto Lari, primary forest, 1-2 km up to Alto Lari, primary forest, 09°24'15"N 83705057 W, 1100-1200
m, 7 Mar 1992, J. Bittner 1456 (INB, MO); entre cerro Chimü y cerro Matama, 1200 m, 29 Abr 1995, L. D. Gómez & G. Herrera 23540
(CR); ae de Talamanca, N flank of Fila de Matama in h ío Boyei, 09%45'N, 83°19'W, 1200-1300 m, 18 Ago 1995,
M. Grayum 11085 (INB); Bratsi, Parque Internacional La Amistad, Sendero Transtalamanca, alrededores de quebrada Lari, tributario
del Coén, 09°25'40"N 83°11'59"W, 1600-1700 m, 6 Feb 2007, A. Rodríguez et al. 11066 (INB); Bratsi, Reserva Indígena Bribri, 1.7 km
southwest of Kivut, 6.1 km southwest of Alto Lari, in primary forest with heavy epiphyte density, 09?23'45"N 83°05'40"W, 1500-1530
m, 27 Feb 1992, H. Schmidt et al. 623 (INB); Valle de la Estrella, Fila Matama, cerca de 11 km SW del pueblo de Aguas Zarcas, bosque
maduro dominado por Tetragastris panamensis, Calatola costaricensis, Quaribaea, Sloanea y Billia rosea, 09°49'04"N 83°09'31"W, 800-90
m, 1 Nov 2007, D. Solano 4870 (INB)
Pilea longibracteolata Al. Rodr., A.K. Monro & L. Acosta, sp. nov. (Fig. 4). Tiro: COSTA RICA. Carraco: Paraíso,
Parque Nacional Tapanti-Macizo de la Muerte, cuenca del Reventazón, Sector del Túnel, 09°42'10"N, 83°47'20"W, 1700 m, 8 Mar
2000, L. Acosta & L.D. Vargas 3200 (notorieo: INB; isoripos: CR, MO)
c - 1 P derana A " + 1l E d:
r O o
Hierba erecta o subarbusto, 0.5—1.0 m de altura, terrestre o epipétrica, dioica o monoica; tallos subteretes
a levemente 5-acostillados, suculentos, glabrescentes a densamente ferrugíneo-pubescentes en las ramitas
terminales, los tricomas 0.40—1.75 mm, simples, multicelul , adpresos, torcidos, con esparcidas glándulas
rojizas y sésiles, los cistolitos ausentes o al menos no evidentes. Estípulas 6-15 mm de largo y 10-12 mm
de ancho, ovado-lanceoladas, sagitadas en la base, agudas y bífidas en el ápice, con nervadura inconspicua,
persistentes, ascendentes, glabrescentes, las del mismo nudo unidas basalmente y ee el tallo (visibles
1
en nudos terminales), en Dunes Heu. pod una partiéndose hasta la mitad | , formando
en cada lado 2 aparent idas; hojas del mismo nudo isófilas; pecíolos
0.3-13.0 cm de largo, sübieuales a depuis con una relación en tamaño de 1: 1.2-6.5, glabrescentes a
esparcida o densamente ferrugineo-pubescentes en ramitas terminales, los tricomas 0.50-1.75 mm, simples,
adpresos, torcidos, con esparcidas glándulas rojizas y sésiles; lámina 2.5-23.0 cm de largo y 0.5-14.0 cm de
ancho, ovada a ovado-elíptica; base obtusa o subcordada a cortamente cuneadas; ápice acuminado; margen
dentado; nervadura palmada o subpalmada, con 3 nervios evidentes saliendo desde la base o hasta 4 mm
sobre la misma; haz glabrescente, los cistolitos fusiformes o en forma de “V,” levemente bullado (no o ape-
nas distinguible en material seco), secando castaño, castafio-rojizo o castaño oscuro; envés esparcidamente
pubescente sobre los nervios principales, densamente pubescente en hojas nuevas, los tricomas 0.50-1.75
mm, simples, adpresos, torcidos, con esparcidas glándulas rojizas y sésiles, los cistolitos fusiformes. Inflo-
rescencias unisexuales, axilares, 1 6 2 por axila, dimorfas; inflorescencias masculinas 10-40 mm de largo y
10-30 mm de ancho, cimas unilaterales, poco o no ramificadas, con flores p te caedizas, subsésiles
o con pedúnculo hasta 14 mm de largo, glabro, basalmente con brácteas pedunculares 5-7 mm de largo y
1.0-1.5 mm de ancho, oblongas a oblongo-lanceoladas, esparcidamente pubescentes, los tricomas ca. 0.75
mm, simples, adpresos, torcidos; flores masculinas subsésiles o con pedicelo hasta 5 mm de largo, glabro,
bracteolas pedicelares conspicuas, 5-12 mm de largo y 2-7 mm de ancho, ovadas o lanceoladas a oblongas,
con 3 a 10 nervios paralelos, esparcidamente pubescentes, al menos en el lado externo, los tricomas 0.2-0.5
mm, simples, adpresos, torcidos; inflorescencias femeninas ubicadas en nudos inferiores o entremezcladas
con las masculinas, 10—70 mm de largo y 10—40 mm de ancho, cimas poco a varias veces ramificadas, el
pedúnculo 5-40 mm de largo, glabrescente, basalmente con brácteas pedunculares 2.5-3.0 mm de largo
y 0.75-1.00 mm de ancho, oblongas a oblongo-obovadas, glabrescentes; flores femeninas subsésiles o con
pedicelos hasta 0.5 mm de largo, glabro, bracteolas pedicelares 0.75-2.50 mm de largo y 0.2-1.3 mm de
ancho, lanceoladas a oblongo-lanceoladas, uninervada o con 3 nervios paralelos, glabras. Flores masculinas
con periantio 1.5-2.0 mm de largo, glabro, 4-lobulado, los lóbulos 1.0-1.5 mm de largo, oblongos a oblongo-
lanceolados, incisos ca. 3/4 de la longitud del perianto, los apéndices subapicales ausentes o si presentes no
más de 0.25 mm de largo; estambres 4, los filamentos 1.5-2.0 mm de largo, las anteras 0.60—0.75 mm de
largo y 0.5—6.0 mm de ancho; pistiloide muy pequeño y apenas distinguible; flores femeninas con lóbulos
del periantio levemente desiguales, el dorsal 0.75-1.25 mm de largo, los laterales 0.60-0.75 mm de largo;
Dados IRA „N 4 ln Dil 1003
O silvia Troyo 08
gl " m
Fic. 4. A-E. P longib lata Al. Rodr., A.K. Monro & L. Acosta. A. Hábito; B. Flor femenina;. C. Flor masculina; D Estipulas
infl ia f i | envés de las hojas. A, B, C, D, E, basados en A. Rodríguez 5877 (INB)
y Ly Ney Uy E,
E fict^lit
estaminodios 3, ca. 0.12 mm de largo y 0.35 mm de ancho, petaloides; estigma diminuto, penicilado, erecto.
Aquenios 0.80—1.25 mm de largo (incluyendo el estigma) y 0.7-0.8 mm de ancho, ovoides, subcompresos,
levemente asimétricos, café, lisos, con uno de los márgenes ligeramente engrosado.
Fenología.—Flores y frutos entre marzo y abril y entre noviembre y diciembre.
Distribución y Hábitats.—Endémica de Costa Rica. Se conoce en los bosques pluviales de la vertiente Ca-
ribe en la cordillera de Talamanca, en el Parque Nacional Tapantí y en los márgenes del río Coén, 1550-1700
m. Esta especie es conocida únicamente de dos localidades y se ha observado creciendo principalmente
sobre paredes rocosas y húmedas dentro del bosque.
1004 Journal of the Botanical R h Institute of Texas 2(2)
Discusión.— Pilea longibracteolata, según la clasificación de Weddell (1869), pertenece al grupo Dentatae
y segün la clasificación d Killip (1960) al grupo Molles. Se reconoce por presentar estípulas singulares y
conspicuas, por su lámina foliar levemente bullada y por la presencia de grandes y evidentes bractéolas en
sus inflorescencias masculinas; además la pubescencia ferrugínea (al menos en las ramitas terminales) es
característica.
Etimología.—El epíteto latino longibracteolata hace alusión a las conspicuas bracteolas pedicelares pre-
sentes en las inflorescencias masculinas.
Material examinado. COSTA RICA. C iso, P. ional Tapanti-Macizo de la Muert del R én, colectado
a orilla de la calle, desde el cruce del río Pino hasta el "e 09°42'10"N, 83°47'20"W, 1700 m, 8 Mar 2000, L. Acosta & V. Ramirez
578 (CR, INB, MO); alrededores del tunel, 9°43'00"N, 83°47'20"W, 1700 m, 5 Abr 2000, A. Rodríguez 5877 (INB). Limón: Talamanca,
Parque Nacional cordillera de Talamanca, 1 km aguas abajo unión quebrada Kirigú y rio Coén, 09°23'20"N, 83°12'50"W, 1550 m, 29
Mar 1999, G. Herrera 6097 (INB, CR).
Pilea moragana Al. Rodr. & A.K. Monro, sp. nov. (Fig. 5). Tiro: COSTA RICA. P Ifito, Reserva Forestal Golfo
Dulce, Península de Osa, Cerro Rincón, Puerto Jiménez, cabeceras de los ríos Tigre y Rincón, 08°30'50"N 83°28'20"W, 700 m, 7
May 1993, R. Aguilar 1881 (HoLoripo: INB; isoripo: CR).
Tint A aly : u E n1 an : 157 n diffaort
Species P. i affinis, p pi
Hierba erecta o procumbente, 0.1—0.2 m de altura, aparentemente terrestre, dioica o monoica; tallos sub-
teretes, pubescentes, los tricomas 0.2—0.3 mm, simples, más o menos adpresos, curvados, con inconspicuas
y esparcidas glándulas rojizas y sésiles, los cistolitos esparcidos fusiformes. Estípulas 1-3 mm de largo y
0.50-1.25 mm de ancho, ovadas u oblongo-ovadas a oblongo-obovadas, obtusas en el ápice, con nervadura
inconspicua, persistentes, ascendentes; hojas del mismo nudo subiguales, con una relación en tamaño de
1: 1.0-2.5 (incluyendo el pecíolo); pecíolos 1-16 mm de largo, subiguales o desiguales, con una relación en
tamaño de 1: 1.2-3.4, pubescentes, los tricomas 0.2-0.4 mm, simples, más o menos adpresos, curvados,
con inconspicuas y esparcidas glándulas rojizas sésiles; lámina 2-45 mm de largo y 1.5—16.0 mm de ancho,
lanceolada a elíptico-lanceolada; base cuneada a levemente oblicua; ápice agudo a cortamente acuminado;
margen dentado; nervadura subpalmada o 3-plinervada, con 3 nervios evidentes saliendo 1-2 mm sobre
la base; haz glabrescente a esparcidamente pubescente, los tricomas 0.30-0.75 mm, simples, aplanados,
curvados, con inconspicuas y esparcidas glándulas rojizas sésiles, los cistolitos escasos, fusiformes o en
forma de "V? en ocasiones ausentes, a menudo levemente bullado; envés glabrescente a esparcidamente
pubescente sobre los nervios principales, los tricomas 0.25-0.40 mm, simples, aplanados, curvados, con
glándulas inconspicuas, esparcidas, rojizas y sésiles, los cistolitos fusif en ee de “V,” en ocasiones
ausentes. Inflorescencias unisexuales, axilares, solitarias, lig nte dimorfas; i ias masculinas
5-25 mm de largo y 3-5 mm de ancho, cima capitadas, el pedúnculo 4-22 mm de largo, glabrescente,
basalmente con brácteas pedunculares 0.2-0.4 mm de largo y ca. 0.3 mm de ancho, deltoides, glabras;
flores masculinas subsésiles o con pedicelos hasta 0.75 mm de largo, glabro, bracteolas pedicelares 0.5—1.0
mm de largo y 0.4-0.5 mm de ancho, lanceoladas, con nervios inconspicuos, café-pajizas; inflorescencias
femeninas ca. 15 mm de largo y 3-5 mm de ancho, cimas cort te ramificadas y pareciendo capitadas, el
pedúnculo 3-20 mm de largo, glabro, basalmente con brácteas pedunculares 0.2—0.4 mm de largo y 0.2-0.4
mm de ancho, deltoides, glabras; flores f inas con pedicelos 0.75-1.00 mm de largo, glabros, bracteolas
pedicelares 0.4—0.6 mm de largo y 0.25—0.50 mm de ancho, lanceoladas, sin nervios evidentes, café-pajiza,.
Flores linas con periantio 0.5—1.0 mm de largo, glabro, 4-partido, los lóbulos 0.50—0.75 mm de largo,
lanceolados, incisos ca. 3/4 de la longitud del periantio, los apéndices subapicales 0.75—1.25 mm de largo;
estambres 4, los filamentos 0.75—1.0 mm de largo, las anteras ca. 0.5 mm de largo y 0.4 mm de ancho; pis-
tiloide muy reducido; flores femeninas con lóbulos del periantio levemente desiguales, el dorsal 0.50—0.75
mm de largo, los laterales 0.4-0.5 mm de largo; estaminodios 3, ca. 0.12 mm de largo y de ancho, petaloides;
estigma diminuto, penicilado, erecto. Aquenios 0.8—1.0 mm de largo (incluyendo el estigma) y 0.6-0.7 mm
de ancho, elipsoides, subcompresos, simétricos, café-pajizos, con una línea punticulada submarginal.
a
Fenología.—Flores y frutos en mayo.
Dade. IRA , N 1 Ja Dil 1005
O Silvia Troyo 08
Fic. 5. A-E. P. moragana Al. Rodr. & A.K. Monro. A. Hábito; B. Flor femenina; C. Flor masculina; D. Estípulas; E. Cistolitos y pubescencia en el envés de las
hojas. A, D, E, basados en R. Aguilar 1881 (INB); B, C, basados en M. Moraga 227 (INB).
Distribución y Hábitats.—Endémica de Costa Rica. Se conoce de los bosques húmedos de la vertiente
Pacífica en la Península de Osa, 150—700 m.
Discusión.—Pilea moragana, según la clasificación de Weddell (1869), pertenece al grupo Dentatae y
según la clasificación de Killip (1960) al grupo Molles. Se reconoce por ser una hierba pequeña con tallos
pubescentes, al menos en las partes terminales, las hojas subiguales y con el margen gruesamente dentado,
con frecuencia pubescentes y levememente bulladas en el lado adaxial, por sus estípulas reducidas y persis-
tentes, las inflorescencias solitarias, dimorfas, las masculinas en cimas capitadas, en tanto que las femeninas
I I nf +l D H FD [rn
1006 f Texas 2(2)
TABLA 1. Comparación diagnósti Ig fológicos de Pil inata, P. cornuto-cucullata, P. rug P moragana.
P. acuminata . cornuto- P. rugosissima P. moragana
cucullata
Tamaño de estípulas 3-20 mm 3-7 mm 2.5-9.0 mm 1-3 mm
Tamaño de pecíolos mayores 15-90 mm 5-45 mm 10-70 mm 1-16 mm
Tamaño de inflorescencias 30-100 mm 5-70 mm 25-50 mm 5-25 mm
masculinas
Tamaño de infl 15-100 mm 2.5-60.0 ram 5-20 mm Ca. 15 mm
femeninas
Forma de inflorescencias Paniculadas, Paniculadas o Capitadas Cortamente ramif-
femeninas raras veces cimas espigadas cadas, pareciendo
espigadas cimas capitadas
Tamaño de apéndice sub- 0.3-1.25 mm 0.2-1.3 mm 2.5-4.0 mm 0.75-1.25 mm
apicales del perianto
masculino
Aquenios con una línea Si No No SÍ
bmarginal punticulada
Distribución sobre el nivel 400-1900 m 2000-3400 m 1800-2500 m 150-700 m
del mar
son cimas cortamente ramificadas, por sus aquenios con una línea punticulada submarginal y finalmente
por su restringida distribución.
Por sus hojas a menudo lanceoladas, gruesamente dentadas, sus tallos en ocasiones pubescentes y sus
inflorescencias capitadas, al menos las masculinas, esta especie podría ser confundida con P. acuminata
Liebm., P. cornuto-cucullata Cufod. y P. rugosissima Killip (ver Tabla 1).
Esta especie se conoce de pocas recolecciones, todas provenientes de Península de Osa en Costa
Rica.
1 Corcovado, El Tigre, Cerro oe 08°27-30'N 83°33-38'W,
, Estación Los Patos
Material examinado. COSTA RICA. P
150—650 m, 22 May 1988, C. Kernan & P. Phillips 493 (CR); Golfito, Parque Nacional Corcovado,
cerca de la Estación en bosque primario, 08%34'09"N 83°30'23"W, 250 m, 27 May 1995, M. Moraga 227 (INB).
AGRADECIMIENTOS
Los autores desean Aia a la aa Association and Linnean Society por aportar financiamiento
para la preparación de | , A Silvia Troyo por la elaboración de los detallados dibujos, a Norman
Robson (BM) por su ayuda en la e acd de las diagnosis latinas y a la Darwin Initiative que a través
del proyecto Tools for the management of La Amistad Binational Park' contribuyó con el trabajo de campo
para la recolecta de material botánico de la especie Pilea herrerae.
REFERENCIAS
Burcer, W. 1977. Pilea. In: Flora Costaricensis. Fieldiana, Bot. 40:246-272.
Davipse, G., S.M. Sousa, AND A.O. Chater. 1994. Introducción general. Flora Mesoamericana, vol. 6, Alismataceae
a Cyperaceae. Universidad Nacional Autónoma de México, Missouri Botanical Garden, The Natural History
Museum (London), Mexico City
Kuup, E.P. 1960. Urticaceae. En: Woodson, Jr., Schery, W. & collaborator, eds. Flora of Panama. Ann. Missouri Bot.
Gard. 47:179-192.
Monro, A.K. 2001. Synopsis of Mesoamerican Pilea (Urticaceae), including eighteen typifications and a key to the
species. Bull. Nat. Hist. Mus. London, Bot. 31:9-25.
Monro, AK. 2004. Three new species and three new names in Pilea (Urticaceae) from New Guinea. Contributions
to the flora of Mt Jaya XV. Kew Bull. 59:573-579,
Rede, { Monro, N sac da Dii 1007
Monro, AK. 2006. The revision of species-rich genera: a phylogenetic framework for the strategic revision of Pilea
(Urticaceae) based on cpDNA, nrDNA and morphology. Amer. J. Bot. 93:426-441.
Poot, A. 2001. Urticaceae. En: Stevens, W.D., C. Ulloa, A. Pool, and O. Montiel, eds. Flora de Nicaragua, Tomo III.
Monogr. Syst. Bot. Missouri Bot. Gard. 85:2479-2494,
STANDLEY, PC. 1937. Pilea. In: Flora of Costa Rica, part |. Field. Mus. Nat. Hist. Chicago. USA. Pp. 392-398.
SrANDLEY, P.C. AND J.A. STEYERMARK. 1952. Pilea. In: Flora of Guatemala, part IIl. Fieldiana, Bot. 24:410-422.
Weopett, H.A. 1869. In: De Candolle, A., Prodromus, systematis naturalis regni vegetabilis. 16(1). Paris. Pp.
4—163
1008 Journal of the Botanical R h Institute of Texas 2(2)
BOOK REVIEW
RICHARD B. PRIMACK. 2008. A Primer of Conservation Biology, 4% Edition. (ISBN 978-0-87893-692-2, hbk.).
Sinauer Associated Inc., 23 Plumtree Road, Sunderland, Massachusetts 01375, U.S.A. (Orders:
sinauer.com, orders@sinauer.com, 413-549-1118 fax). $49.95, 293 pp., 123 illustrations, 9 1/4"x 7 1/4".
pene its En nde this concise version of the larger Essentials of Conservation Biology 4th Ed. delivers an expansive review of the
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The field of jan bt lin tt 305 ld he rapid decline of biodiversity. The first four chapters
explore the P MEM ini P and: threats to biodiversity. Chapter five di he finality of extincti lits imp he global
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T 1 E da 1 l.«L: 11 1 “11 1 1 f NA AT E E 11-11
State M N list Curriculum.—Gary L. Hinds, Chemist, Rio Brazos Master Naturalist, and BRI T volunteer, Fort
Worth, Texas, U.S.A.
J. Bot. Res. Inst. Texas 2(2): 1008. 2008
THE DIOSPYROS SALICIFOLIA COMPLEX (EBENACEAE)
IN MESOAMERICA
Mitchell C. Provance Ignacio García Ruiz
Instituto Politécnico Nacional
UCR Herbarium
mu of Botany and Plant Sciences po tro nano de a
University of California, Riverside
m o 92521-0124, U.S.A. Ep. Sierra 28, 595 10 Jiquilpan, Michoacán, MÉXICO
mitchprovance@yahoo.com igarciar@ipn.mx
ip i aues
CR Herbari
siii ke Botany i Plant Sciences
University of California, Riverside
Riverside, California 92521-0124, U.S.A.
andrew.sanders@ucr. edi
ABSTRACT
T Fl T»; Taf 1 Je 1 a 1 1 1 der E NDS Seven
The taxonom 1 i g
species are currently | in Mesoamerica Icensis K hich occurs from Panama to Jalisco and Veracruz, Mexico,
is separable into nine o PIS aids Standl, a Te id western Mexico Mun MEME to Oaxaca, is separable into six
subspecies j; ij ex Willd p xico, and may be of recent
hybrid origin Di isl lell he Yucatan Peni ] 1 | y i Í Mexi d Central America, is
separable into two subspecies Pup nem intricata m Gray) Standl. i ly known only from the Cape region of Baja California Sur,
Mexico. A d here 1 1 1 1 1 ET T a .
sometimes form at the edges of their Pages Panamanian MOM of putative D. "M Jacq., belong to this complex, and the
recently a D. enana B. va seems to as hii ns are bon for D. salicifolia, D. acapulcensis, D. aequoris and
and five new combinations are made. A key to the taxa is
D. yucatanensis, Į P
provided, along with distributi ps, ill i d pl f ty] d living material
RESUMEN
€ 1 1 1 Ty Te fal 1 1 Pil lási 4lisi lti i Se conocen
r 4 £ + 4
1 a * NA RS TW 1 TOME $ 1 3 m A dJa D A Tos y "T
je E E q J y veracruz, México, se
1 1 TY E ar A! 1 quls dox. bd Ead ^ PE d 1 Cty es separable
r SE
en seis subesse Diospyros ee Hub. & Bonpl ex Willd. es endémica de la región de A ] G , México, y puede
ser de reciente ads lia nO cid Lundell, de la Península de Yucatán y regiones Dr n México y América
Central Į Į js 1 A Gray) Standl ode Baja California
Sur, iron án ci i quí, la hibridación i l ] j veces se forman
t b i ] tr de su rango CaL fios del putativo D. inconstans cq, pertenece a este
complejo, y el recientemente descrito D. inis add Walln., t l portan descripci D. salicifolia, D. acapulcensis
1 : 1 a
D. ini on Tenn I I Se aporta
j pas de distribución, ilustraciones y fotos de los tipos, y de material vivo.
INTRODUCTION
As part of a revision of Diospyros for Mexico, a study of the D. salicifolia complex was conducted in order
to address taxonomic inconsistencies in the literature. We used a combination of conventional taxonomic
tatisti d extensive fieldwork in Mexico, in order to resolve uncertainty regarding
methods, multi
the proper o of names, assess taxonomic boundaries, emend species descriptions, and construct a
useable key for the taxa. Early workers described a number of 3-4-sepalous Diospyros in Mesoamerica that
seem to form a natural group that we refer to as the Diospyros salicifolia complex. Morphometrics, including
J. Bot. Res. Inst. Texas 2(2): 1009 — 1100. 2008
1010 J | of the Botanical R h Institute of Texas 2(2)
multivariate statistics, were used to explore continuous variation and the utility of quantitative vegetative
and reproductive characters in the circumscription of taxa throughout the distribution of the complex in
Mesoamerica.
Diospyros salicifolia complex: female flowers solitary, rarely in a 3-flowered cyme with two flowers
aborting, or a single female flower terminating an otherwise male inflorescence; calyx lobes 3—4, accrescent
(mostly widening, but lengthening markedly in D. yucatanensis subsp. spectabilis); female flowering calyx
mitriform, sometimes tearing or dehiscing at the sinus as the fruit and/or calyx enlarges; styles 3, varying
from a fully fused column to free to the base, minutely hairy, stigmas bifid, corolla tube sericeous with
3—4 glabrous zones around the base; fruits with three pairs of locules (these pairs difficult to see in mature
fruit, but conspicuous during dissection of immature ovaries), the members of a given pair divided by an
apparently complete septum; fruit wall indehiscenct to vaguely or (infrequently) conspicuously dehiscent;
flesh + clear and gelatinous, but becoming reddish, translucent and vitreous when dry; leaves usually evenly
distributed along the stem; hairs throughout the plant with fluid-filled lumens, the fluid clear in life, but
usually turning reddish upon drying (essentially non-capitate simple glandular hairs-some plants also with
occasional non-capitate multi-branched glandular hairs); seeds reddish, especially in life, distributed by
birds (White 1978, pers. obs.) and mammals, probably carnivores (pers. obs.).
There are two Mesoamerican Diospyros species complexes that could be confused with the salicifolia
complex.
Diospyros tetrasperma complex: Female flowers usually solitary; calyx lobes 4—5, not markedly ac-
crescent; female flowering calyx not mitriform; interior of the fruiting calyx often ridged and hairy in the
upper tube and basal lobe, sinus indehiscent/not tearing; styles 2 pairs or 1 with 5 branches, stigmas 4—5;
fruits 4—5 loculed, the locules all divided by a seemingly complete septum; the flesh of the fruit + gelatinous
(rarely fleshy), becoming reddish, translucent and vitreous when dry; leaves sometimes crowded at the ends
of stems on short shoots. Members include D. tetrasperma Sw., D. anisandra S.F. Blake, D. johnstoniana Standl.
& Steyerm., D. bumelioides Standl., and D. yatesiana Standl. ex Lundell.
Diospyros campechiana complex: Female inflorescence 1-4-flowered; calyx lobes 4-5, not markedly
accrescent; female flowers not mitriform; fruiting calyx weakly attached to the fruit, tending to persist on
the pedicel rather than the fruit, the tube spreading and lobes patent to reflexed; style 1 with 2 branches;
fruits 1-4-loculed; the amount of flesh present relatively meager by comparison to members of other com-
plexes, reddish, translucent and vitreous when dry (probably clear and gelatinous in life); the hairs minute,
and include dark red to nearly black, slightly flexuose to straight, appressed (or nearly so) hairs; seeds few,
loosely contained in the fruit of herbarium specimens; leaves medium to large, not crowded on the ends of
the shoots, the petioles sometimes with transverse fissures. Members include D. campechiana Lundell, D.
hartmanniana S. Knapp, and D. panamense S. Knapp.
Members of the salicifolia complex were formerly treated as members of the genus Maba J.R. Forst. &
G. Forst. (e.g., Hiern 1873) or Macreightia A. DC. (e.g., De Candolle 1844). Herbarium specimens often have
been identified using Standley's treatment of the group under the genus Maba (1924), but typically bear the
appropriate combination in Diospyros following Standley (1935). Interestingly, no specimens determined as
D. portus Standl. (= Maba rekoi Standl.) were seen during this study, although this little-known taxon ap-
peared in Standley's treatment. Specimens from the Yucatan Peninsula and northern Guatemala are usually
identified as D. yucatanensis or D. spectabilis following Lundell (1937, 1942).
Several workers, such as B. Wallnófer, C. Whitefoord and S. Knapp, and F. White, ascribe Mesoamerican
material of this complex (with the exception of the Baja California endemic D. intricata, and D. amanap and
D. aff. inconstans from Panama) to D. salicifolia Humb. & Bonpl. ex Willd., sometimes with the postscript
'sensu lato.' The different opinions concerning the taxonomic boundaries within this highly variable com-
plex, and the trends seen in the identification of material, are related to the application of three names, D.
acapulcensis Kunth, D. salicifolia Humb. & Bonpl. ex Willd., and D. albens Presl. These names are all based
Provance et al., Diospyros salicifolia complex 1011
on collections from the Acapulco region of Guerrero held in historical European collections at Paris (P),
Willdenow (B-W), and Vienna (W) respectively.
TYPE MATERIAL FROM GUERRERO, MEXICO
The name Diospyros salicifolia, published by Willdenow in 1806, is based on material (“Humboldt & Bonpland
s.n.”) collected in the Acapulco region of Guerrero, Mexico, and is the oldest name in the complex. Unfortu-
nately, the holotype at Berlin (B-W19250-01) consists only of a branchlet with some very immature leaves,
one disconnected mature leaf, and one disconnected fruit (Fig. 1). In preparation of Flora Veracruz, Pacheco
(1981) had doubts about the application of the name D. salicifolia because the type material is insufficient,
and thus used the name D. veraecrucis Standl. for plants growing on the east coast of Mexico. However,
she mentioned Frank White's suggestion that D. salicifolia is the correct name for the species in Veracruz.
Whitefoord and Knapp (1998-onward) commented that the type specimen of D. salicifolia is *depauperate;"
but concluded that material they attributed to D. salicifolia in southern Mexico and Central America was in
agreement with the holotype of D. salicifolia based on microfiche (“microficha BM! ex B-W"). Recently, the
Botanic Garden and Botanical Museum Berlin-Dahlem has made it possible to view on the Internet a high
quality image of the type specimen of D. salicifolia held in the Willdenow Herbarium (B-W), including the
informative abaxial surface of the mature leaf.
The name D. acapulcensis was published in 1818 by Kunth, who worked on the collections of Humboldt
and Bonpland in the herbarium at Paris between 1815 and 1828 (Hiepko 1987). The type (Fig. 2a—b) is a
specimen in the Paris Muséum National d'Histoire Naturelle (P), unfortunately bearing roughly the same
scanty label data as the type of D. salicifolia at Berlin, "Humboldt & Bonpland s.n." The locality indicated on the
preprinted label is “Amérique Equatoriale.” The determination of the specimen as “Diospyros acapulcensis” is
in the handwriting of Kunth. Because of the general similarity of these two species and the inadequacies of
the type specimens, this collection was considered an isotype of D. salicifolia by Wallnófer (by annotation c.
2002) and Whitefoord and Knapp (1998-onward). Wallnófer also annotated this specimen as the lectotype
of D. acapulcensis in 2002. As far as we can tell, this E was not formally published.
The confusion of D l d D. salicifolia is | g. In the earliest treatment of the genus to
include members of the sa licifol ia complex, Alphonse De dale (1844) included D. salicifolia in his concept
of Macreightia acapulcensis (Kunth) A. DC. with a question mark. But, in his monograph of the Ebenaceae,
Hiern (1873) stated that “the foliage is sufficiently different” between D. salicifolia (= Maba salicifolia Hiern)
and D. acapulcensis (= Maba acapulcensis Hiern) for them to be treated as separate species. However, it is
unclear to us what material he actually compared, since in reference to Maba salicifolia, he cites having seen
only *Humboldt and Bonpland" from "Equatorial America," which would seem to refer to the type collection
of D. acapulcensis in Paris. However, this citation seems dubious, since his description of D. salicifolia is more
consistent with the type at Berlin (D. salicifolia) than the type at Paris (D. acapulcensis), and lacks a descrip-
tion of the dissected e mounted on the Paris specimen. We are a dus by the absence of the
Humboldt and Bonpl from th tion of speci h. Hiern indicates
in the introduction to ks monograph that he examined the type material in the Willdenow collection at
Berlin. It would be astonishing if he had not made use of the type collection of D. salicifolia. In reference to
D. acapulcensis, Hiern cites a single specimen, with the locality data “Mexico, Acapulco, Bonpland,” which
is consistent with label data on a sheet at Paris (P-271690) identified as D. acapulcensis, but not currently
available to us. Standley (1924) treated D. salicifolia and D. acapulcensis (under the genus Maba) as separate
species. In his key he separated them based on differences in leaf shape at the apex and lamina vestiture.
He considered material from Acapulco referable to D. salicifolia, but did not expand on the distribution of
D. acapulcensis, other than to state that the type was from Acapulco. The status of these species was still not
—
completely clear.
More recently, D. salicifolia and D. acapulcensis have been treated as synonyms, apparently beginning
with a broad concept of D. salicifolia by F. White, but also based on the fact that the two type collections
1012
Mus. Bot Barcel.
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Provance et al., Diospyros salicifolia complex
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(P-307187), the lectotype of Diospyros acapulcensis Kunth.
icifolia Humb. & Bonpl. ex Willd., from Humboldt & Bonpland
s.n. (B-W-19250-01).
- , 1 aftho Rataniral D h Institute of Texas 2(2)
(B-W-19250-01 and P-307187) have been considered duplicates by some workers. However, the type
specimen of D. salicifolia (Humboldt & Bonpland s.n.) in the Willdenow collection (B-W-19250- 01) does not
represent the same species as the Type a D. kac pdt at the Paris Muséum National d'Histoire Naturelle
(P-307187.) Theset ly considered duplicates. The B-W specimen
(Figs. 1, 2c) represents D. salicifolia, a taxon that is evidently endemic to coastal regions around Acapulco,
Guerrero including Isla Roqueta in Acapulco Bay. This species has been collected at Acapulco on a number
of occasions in recent times, but nowhere else. The Paris specimen of Humboldt & Bonpland s.n. (Fig. 2a—b)
represents Diospyros acapulcensis (P-307187, designated below as the lectotype). This taxon occurs through
much of western Mexico, and while it does occur near Acapulco, in the state of Guerrero it is more common
inland in the municipalities of Chilpancingo, Achotla, and Iguala. The closest collections to Acapulco that
we have seen are from about 15 km away at 200 m elevation near Rio Papagayo (Fig. 3).
These two specimens also cannot be part of the same gathering because they are in different pheno-
logical stages: both specimens have fruit, but the Berlin specimen (D. salicifolia) is almost entirely without
leaves except for very immature ones at the shoot apices. It represents a collection from the beginning of
the growing season, from a plant that had a few old persisting leaves from the previous growing season.
The single mature leaf on the collection is elliptical with a base that is rounded to the petiole. The Paris
specimen (D. acapulcensis) represents a plant from a later phenological stage, as it still has numerous mature
leaves, and is without immature leaves at the apex. The laminas on the Paris specimen are all oblanceolate
to obovate, and have a cuneate to slightly rounded base. When the abaxial leaf surfaces of the specimens are
closely compared (Fig. 2a, c) additional differences are readily apparent. The lateral veins of the Paris speci-
men are thick and straight and diverge at an acute angle, while those in the specimen at Berlin are finer and
diverge at a greater angle, gradually curving toward the leaf apex. The Paris specimen probably came from
near Acapulco, but from a slightly more interior location than the Berlin specimen. It is unclear to us what
facts Kunth had that indicated the Paris collection was made in a littoral zone, which seems a more likely
habitat for D. salicifolia. This information may have come from one of the other Paris specimens (currently
unavailable to us), in which case, that specimen should be evaluated carefully to see if it is D. salicifolia. Other
possibilities include miscorrelation of field notes (unknown to us) attributable to D. salicifolia collections,
or use of the term "littoral" in a very broad sense. There are two other specimens at Paris identified as D.
acapulcensis that are currently unavailable, but which seem to have label data similar to the chosen (below)
lectotype: Bonpland s.n. (P-271689) from *Mexico," and Bonpland s.n. (P-271690) from *Acapulco, Mexico."
The specimen annotated by Wallnófer as lectotype (P-307187) is consistent with the original description of
D. acapulcensis. This description gives a rather complete account of seed anatomy, which is consistent with
the carefully mounted dissected seed on the specimen favored by Wallnófer. We believe that P-307187 was
important in the initial description of D. cas and below we formally designate it as the lectotype.
The other species involved in this lissue, Diospyros albens C. Presl, was described in 1835
from material collected by T. Haenke at Acapulco in about 1791 (although the collection is undated). Based
on a photograph of the type (Haenke s.n., W-31942) obtained from Field Museum, the specimen appears to
be a collection of D. salicifolia with male flowers and young leaves. We cannot separate this material from
other specimens of D. salicifolia with young leaves (e.g., W. Boege 452), and the two names are here treated
as synonyms. The name D. albens appears occasionally in the literature and sometimes on herbarium
specimens. For instance, Standley (1924) seems to have used this name in reference to D. aequoris subsp.
tehuantepecensis, a subspecies proposed here for the state of Oaxaca, and later in reference to the nominate
subsp. of Diospyros yucatanensis (Standley 1930).
METHODS
Taxonomic Methods.—All of the relevant literature, loaned Specimens. MADE. types, and images of types,
that could be obtaine valuated. Additionally, ibutable to the salicifolia com-
I
Provance et al., Diospyros salicifolia complex 1015
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plex were collected by the authors in the states of Baja California Sur, Sinaloa, Nayarit, Jalisco, Michoacan,
Guerrero, Campeche, and Oaxaca, Mexico and the Nicoya Peninsula, Costa Rica. We have studied about
900 specimens referable to this complex.
We generally agree with de Queiroz (1998, 2007) that species can be thought of as "separately evolv-
ing metapopulation lineages," where lineage refers to an “ancestor-descendant series,” and metapopulation
refers to "an inclusive population made up of several lineages" (de Queiroz 2007) and appreciate his eclectic
approach to acceptable lines of evidence that can be used in assessing lineage separation. However, we
remain uncomfortable with the practical implications of this general approach for the working systematist,
especially given the ambiguities of lineages composing metapopulations, the temporal segments of which
in turn compose the other lineages called species, and feel that the “general lineage concept" is not yet op-
erational.
Our approach is been based on examination of morphological variation observed in a sample of speci-
mens preserved in herbaria of the world over the past 200+ years, and in living material encountered (and
photographed) during field studies in Mexico and Costa Rica. Collections were sorted into groups based on
1016 Journal of the Botanical R h Institute of Texas 2(2)
morphological characters, and secondarily based on geography. We believe that useful taxonomic groups
should have unique and generally coherent geographical distributions. Groups identified in our preliminary
sorting were considered taxonomic hypotheses warranting continued investigation. Comparisons were
made between specimens assigned to each group, and between those specimens and type specimens. We
also examined specimens in light of type descriptions, existing floristic and monographic treatments, and
the annotations of previous workers. Published applicable to the identified groups were determined,
and the names having priority were determined for each group. Emended descriptions were constructed
for species based on studied herbarium specimens and field observations.
De Queiroz does not discuss the use of subspecies or other infraspecific ranks, though these have been
widely used by taxonomists, and even though such ranks appear quite appropriate for populations in his
zone of lineage separation (de Queiroz 2007, fig. 1) where only some of the properties of species have been
acquitted: This is the way that infraspecific ranks have traditionally been used by biologists. Here we use
the pecific rank, perhaps in a way similar to other recent workers (e.g., Mulcahy 2007), to communicate
our hypothesis of an incipient species within a metapopulation, although in some cases this may represent
formerly separate species in the process of merger due to secondary contact resulting from migration and/or
environmental change. We are reluctant to describe populations as full species when it appears that botani-
cal exploration between these populations has been inadequate and where the morphological differences
separating them are not great. Further exploration could reveal populations with mixtures of the defining
characteristics of the two initial pau ase it is certain that at least some of our subspecies
would be treated as full species under p inal taxa in a cladogram as species
(e.g., Mishler & Theriot 2000), we have taken a conse vane course in treating them as subspecies. Future
studies may reveal that some of our subspecies have diverged more than we can now see and that they are
best treated as full species. In any event, we feel it is most important to call attention to these entities. Their
taxonomic status can always be changed later, but only if the taxa are first described so that other biologists
know they exist.
In accordance with this approach, the material assigned to m eris was examined so that consistent
differences in morphology, correlated with contiguou uld be detected. If morphological
differences between subgroups within a species wore to jo predictive of geography, and the subgroup
distributions were allopatric or parapatric, then recognition at subspecific rank was strongly considered.
We consider subspecies to be geographic races marked by one or more morphological features, not random
variants that appear throughout the distribution of a species. There should be a g is for the morpho-
logical variation observed between subspecies, although it is understood that differences in environmental
distribution complicate efforts to tease genetic effects apart from environmental influences, especially in
baseline taxonomic studies that may have few representatives for certain taxa. In an effort to minimize the
influence of environmental factors on our subspecies delimitation, we conducted analyses of covariance
using elevation (a proxy for variation in the environment) as a cross effect on leaf and inflorescence traits
between subspecies. These analyses (to be presented in a later paper) suggest that quantitative variation
in many of these traits is likely to have a genetic component. Furthermore, they highlight an interaction
between genetic and environmental factors for some of these traits. In other words, some characters varied
inversely with elevation for different subspecies. For instance, the venation parameter leaf width : distance
between lateral veins, decreased with elevation for D. acapulcensis ssp. guanacastensis, while it increased with
elevation for D. acapulcensis ssp. rivensis.
We expect subspecies to hybridize with other subspecies, and if we found that two occupying the same
range and habitat did not hybridize with one another, we would propose that they be treated as separate spe-
cies. But, we do not necessarily expect reproductive barriers to be complete between plant species, although
we expect them to be greater than between subspecies.
We carefully studied the floral anatomy of each taxon we could, but descriptions of internal floral
structures are still sometimes based on relatively few observations, and the floral anatomy of all taxa needs
Provance et al., Diospyros salicifolia complex 1017
more study. In this treatment we use ‘lanceolate’ in the sense of Jackson (1916), being broadest near the
lower third of the leaf, not at mid-leaf in the sense of Stearn (2000).
For new combinations, at least one specimen is cited for each relatively small political unit of occur-
rence (e.g., municipality, sauna. Most ae examined is cited for new subspecies. Collections of
interspecific hybrids and sul ited in Appendix 1. Descriptions apply to herbarium
material except when indicated: or when obviously referring to fresh material. Comments on ecology and
ethnobotany are based on specimen label data, published field notes, vegetation maps, and personal ob-
servations. Map coordinates geo-referenced by the authors are given in brackets. Distribution maps were
constructed using The Generic Mapping Tools 4.1.4 (Wessel & Smith 2006) accessed through OMC (Weinelt
2006) and amended using Adobe Photoshop (Ver. CS 8.0, Adobe Systems Incorporated, San Jose, CA).
Photographs and illustrations are by the first author unless specified otherwise.
Statistics Methods.—The dataset used in the multivariate statistics consisted of 360 fruiting collections
of Mesoamerican Diospyros referable to the salicifolia complex (a list of accessions is available from the first
author). Leaf measurements were based on an average of the three largest leaves, except (rarely) when only
two acceptable leaves were present and the specimen was considered especially important for inclusion.
The statistical software used was JMP Statistical Discovery (Ver. 6.0, Macintosh Version, SAS Institute Inc.,
Cary, NC).
Prior to principal component analysis (PCA), strongly correlated characters were identified and elimi-
nated. Since exclusion values are somewhat arbitrary, two such values (R « 0.7, R « 0.8) were explored.
When a correlation was above the exclusion value, one character was retained with preference given to
leaf length or leaf width, and lengths over ratios. Leaf length and width were highly correlated. Instead of
choosing arbitrarily which of the fundamental lamina measurements should be excluded, we decided to
retain two character sets at each exclusion value (i.e., one set with width excluded, the other with length
excluded). PCA was used on each of the four sets to discern the relative separation of species groups in
multidimensional subspace, and to identify the quantitative variables explaining the most variation in the
data. PC scores were scaled to unit variance, thus grouping is discerned from ‘GH biplots. Where the first
three Ë returned eigenvalues greater than one (e.g., many components were needed to explain the
observed variation), a varimax factor rotation f l using all of the components having eigenvalues
of one or more in order to make the results ene
TAXONOMIC TREATMENT
The majority of South American Diospyros specimens were unavailable for this study, and consequently
the distribution of the salicifolia complex in South America could not be addressed. Whitefoord and Knapp
(1998-onward) reported “D. salicifolia” for Columbia, but they did not cite a specimen. We have not seen a
specimen of D. salicifolia sensu lato from South America, BN we do not doubt the presence of D. acapulcensis
in Columbia. The cds of available South American sy lso meant that the taxonomic status of D.
inconstans Jacq. in N ica could not be add d completely. All putative Mesoamerican D. inconstans
specimens we examined were collected in Darien Province, Panama. This material is uniform and we believe
it represents a single species perhaps related to, but nonetheless different from, those species represented
by the smattering of collections we received from South America and the Caribbean. In our opinion the
South American specimens determined by other workers as D. inconstans represent more than one taxon.
The leaves of the Panamanian material seem consistent with an image seen online (Tropicos.org 2008) of
a sterile collection from Columbia (Bertero s.n. [MO-1905992]) annotated by B. Wallnófer as an “ “isotype”
of: Diospyros berterii ... = D. inconstans s. lat. Jacq.” At any rate, the Panamanian collections cited here in Ap-
pendix 2 as D. aff. inconstans are clearly attributable to the salicifolia complex, most obviously because of the
male flowers in 2-3-flowered cymes, solitary female flowers, the trimerous calyx which is clearly mitriform
in the flowers of both sexes, and the overall appearance of the fruiting calyx and mature fruit.
Wallnófer (2005) recently described a new species, D. amanap B. Walln., based on one collection from
1018 J | of the Botanical R h Institute of Texas 2(2)
Columbia and another from Darien Province, Panama. Although we have not had the opportunity to compare
our material with these collections, we think it is unlikely that the D. aff. inconstans specimens cited here
represent D. amanap, as they are not in accord with the species description and because Wallnófer annotated
one of the specimens we studied as D. inconstans sensu lato. Unfortunately, Wallnófer did not provide an
illustration or photograph of D. amanap, or make any comparisons of D. amanap to other taxa. The species
is described as having a trimerous accrescent calyx and probably a 6-locular fruit (Wallnófer 2005), and is
therefore likely to belong in the salicifolia complex.
KEY TO THE MESOAMERICAN SPECIES OF THE DIOSPYROS SALICIFOLIA COMPLEX
(EXCLUDING D. AFF. INCONSTANS AND D. AMANAP
. Petioles « 2 mm is arana ie SOC) mm long, 5- x mm wide, ee rounded a. the base
JOL Ing),
cuneate, s rselyt to Moderately hairy Delov sed
d calyx small (calyx ERG + alobo width 12- 16 mm); eur mostly « 2. 5 m tall; Cape peus E Baja
California Sur, including Isla Cerralvo D. intricata
. Petioles > 2 mm long; lamina 18-155 mm long, 11-65 mm wide, shape variable, density of abaxial vestiture,
and shape of abaxial o tlie fruiting calyx larger (calyx length + lobe width usually (15-)17-25(-32)
mm; trees or shrubs, tall; not known from Baja California.
2. Lamina Ls to a 18-110 mm d nee ae Ve. oval or nearly circular,
the base rounded to subcordate, often coarsely wri
3. ud female calyx tube 1.2-1.8 mm long; lamina d 60- Mou NM length : width = 2.2742
1; base more narrowly rounded than in D. aequoris (length : width at 0.25 = 3.3-4. 8: 1); apex rounded
oe width at 0.75 = 3-4: 1), lamina not wrinkled above, Gaver to sparsely hairy when mature;
3° venation inconspicuous biu faintly raised or not apparent y known only from the
immediate vicinity of Acapulco, Guerrero, « 200 m D. salicifolia
Flowering female calyx tube 2.3-6 mm M (1.5 mm long in one subsp. from interior Oaxaca); lamina
8-81 mm long, length : width — 1.1—2.8 : 1; base rounded to subcordate (length : width at 0.25 —
1.5-4.3 : 1); apex rounded to acute i. : width at 0.75 = 1.3-3.5 : 1), lamina often wrinkled above,
moderately to densely pubescent below when mature; 3* venation usually prominent below and im-
pressed above; m in vee < 1200 m D. aequoris
2. Lamina membranaceous t eous, rarely subcoriaceous, 45-160 mm long, lanceolate to ovate, or
oblanceolate to a sometimes elliptic, oblong or oval, the base rounded, attenuate, or cuneate,
decurrent on the petiole, sometimes abruptly, not coarsely wrinkled above; Jalisco, Mexico to Panama.
4. ue pou ay tube 1 2- 1 di mm long; lower lamina surface glabrate to pun hairy when
hairy I pi p! raight, or nearly so,
without swollen rernanthsirl ferti pill pl i tly k nown only from the
mmediate vicinity of Acapulco, Guerrero, < 200 m D. salicifolia
4, "Hein female calyx tube 2.5-5.5 mm long (1.8-2 mm long in one Central American subsp); lower
Ru SEE add d densely hairy AER noue or if glabrate, then with swollen remnant hair
pread in Mesoamerica.
5. Lamina lanceolate Or ovate, sometimes Goes obovate, or elliptic, abaxial vestiture consisting
of Palle mita suelen bases, comicios a las papillose gppestanee to the lamina, especially in
pi y ys opaq ; 27-3? veins slightly
| Jal with a raised vein-l ;fruiti licels 2- 16mm long; fruiting calyx tube
2.5-5.5 mm long (tube : E length = 0.2-1.2 : 1); plants often tall and slender; Yucatan Penin-
sula, Chiapas, Tabasco, SE Veracruz, Mexico, Belize, Guatemala, and NW Honduras, <
270(-1400) D. yucatanensis
. Lamina oblanceolate to obovate, ti ti without conspicuous papillae,
or the papillae minute and diffuse (except in one Ms collected subsp. of D. aequoris from Hondu-
ras), n sone) apparatus OB m opaque Ane cone picuous; 2 uh veins a Usually Tuan do
—
Ww
Un
spodki length = 1. = 18: 1); plants not markedly tall and slender; Jalisco, Mexico to Panama, not
known from the Yucatan Peninsula, < 1500 m . acapulcensis
l. Diospyros acapulcensis - Kunth, ey Gen. m 3:254. 1818. (Fig. 2a—b). Macreightia acapulcensis (Kunth) A. DC.
bilis 8:221. 1844. Maba acapulcensis M Hiern, Trans. oe Philos. Soc. 12:128.
1873. Tree: MEXICO. GUERRERO: Abi Equatoriale” el on label], “Crescit i , prope Acapulco,”
“Fructificat Februario" [in protologue], Humboldt & Bonpland s.n. (Lectorvee, here designated: P-307187, digital image!).
Provance et al., Diospyros salicifolia complex 1019
Trees, arborescent shrubs or shrubs, (12)22-12(-25) m tall, sometimes multi-stemmed clones, facultatively
deciduous, mostly dioecious, rarely flowers of both sexes in the same inflorescence (the mating system in
need of more study); trunk up to 40 cm dbh, bark « smooth (not slick) to irregularly roughened, or patchy,
often lichen encrusted (at least in W Mexico), often in part irregularly scaly, shallowly fissured or checked,
especially at major branch forks, coloration uniform to patchy, usually grayish; wood yellowish to light
brown when dry, reportedly very hard, slash nearly white to light yellow, rapidly turning yellowish, sap
clear; mature stems terete to subterete, rarely angular, nearly smooth to roughened, fissured, half-netted,
verrucose, or knotted, + glabrous, the epidermis grayish, reddish, or brownish, lenticels often present,
minute, circular, and bump-like to large and elliptic, sometimes fissure-like and forming a reticulum; 2nd
year stems terete to angular, smooth to half-netted, the bark sometimes sloughing, often longitudinally
ridged, minutely fissured, bumpy, or verrucose, glabrous to sparsely hairy, rarely moderately to densely
hairy, often with remnants of desiccated hairs, these sometimes coated with exudate, the epidermis viscid,
often black gland dotted, occasionally mottled with reddish-brown or black, lenticels usually present; Ist
year stems quadrangular to subterete, smooth to verrucose, sometimes sulcate, sparsely to densely hairy,
the hairs straight, wavy, curly, kinked, distally curved or hooked, 0.2-2 mm long, spreading to appressed,
sometimes retrorse, hairs often reddish (the hairs throughout the plant are mostly simple glandular hairs,
translucent in life, but variously colored in herbarium specimens due to darkening of a fluid in the lumen),
clavate glandular hairs often present, deciduous, leaving the epidermis viscid, the epidermis green in life,
grayish, reddish, or brownish in dry material, sometimes minutely lenticellate, bud scales ovate, concave,
covered with appressed hairs. Leaves alternate, ME entire; pes URS 7(—9) mm long, sometimes
minutely winged, glabrous to densely hairy, clavat d present, these deciduous,
leaving the epidermis viscid, sometimes without HE glandular hairs and the epidermis dull, surface
smooth to rugose, sometimes glaucous, pruinose or scintillant, lower e convex, He surface dd
ly concave or shallowly canaliculate to ; lamina cl when
mature in one subspecies (shade and spring leaves often membranaceous), 45— 125(-160) mm long, (15-
)18-60(-65) mm wide, the length to width ratio (1.7-)2-3.6(-4.4) : 1, mostly oblanceolate to obovate,
sometimes elliptic, oblong or oval, usually slightly wider in the distal half, sometimes obtrullate, rarely
lanceolate or ovate, clavate glandular hairs often present, deciduous, leaving the epidermis viscid, the epi-
dermis color variable, but darker above, base cuneate to long or short attenuate, decurrent on the petiole,
margin flat to revolute, sometimes ciliate, apex acute to obtuse, acuminate or tapered to a blunt point,
rounded, or mucronulate; lower lamina surface sparsely to densely hairy, rarely glabrate, the hairs vari-
able, often deciduous, the epidermis often viscid, coloration sometimes mottled, the stomatal A clear
or off-white, opaque, and conspicuous, and then may be mistaken for scales; upper
to moderately hairy, the hairs often wavy and upright to ascending, usually deciduous, their bases o
thickened, annular seeping glands sometimes appearing where hairs have fallen, stellate hairs occasionally
present, often crystal-papillose (with aggregates of opaque off-white excrescences), more obviously so in
thinner leaves (rarely on the abaxial lamina surface), sometimes tented (with minute well spread-out waves
or peaks along veins of the lamina surface is by differential en in herbarium pun Venation
arcolanguid (Provance & Sanders 2005) to hidodromous, someti , 3(—5)-ribbed,
often darkened on the lower surface of young leaves; midrib Bua prominent Delos glabrate to densely
hairy, smooth to rugose, sometimes glaucous, pruinose or scintillant, slightly impressed to convexly-raised
above near the base, flush to slightly raised distally, usually impressed along the lamina-midrib seam, hairs
tending to be upright, clavate glandular hairs often present, deciduous, leaving the epidermis viscid; lat-
eral veins 5-12 on each side of the midrib, often the basal pair and/or the second from the base, diverging
from the midrib at a more acute angle and with a straighter course than the others, raised below, flush to
slightly raised above; 3? veins obscure to conspicuously raised below, usually flush or inconspicuously
raised above; 4?—5? veins obscure to barely raised. Laminar extrafloral nectaries nearly always present
on the lower lamina surface (very rarely on the upper surface), tending to be near the midrib in the proximal
1020 J lof the Botanical R h Institute of Texas 2(2)
half of the lamina, minutely rimmed, flat to concave, slightly raised to sunken in the lamina, elliptic, round
or amorphic, mostly 0.3-1.5 mm in longest dimension, green in life, drying greenish, reddish, brownish or
black. Male inflorescences solitary (2-)3-5(-7)-flowered cymes in leaf axils of young (usually Ist year)
stems, cymes of male flowers rarely terminated by a female flower, moderately to densely hairy, the hairs
minute, nearly straight to wavy or kinked, upright to ascending, tawny or dull orange to reddish or dark
brown, epidermis sometimes viscid, straw, yellowish, dull orange, light brown, golden brown, reddish-
brown, or dark brown; peduncles 1-6 mm long, often angular; pedicels 0.5-3 mm long, broadening
distally, bracts linear-elliptic to ovate, 1-2.5 mm long, often navicular. Male flowers 3-4-merous; male
flowering calyx narrowly campanulate to campanulate or infundibuliform, exterior hairs moderate to
dense, sometimes glabrous near the sinuses, oe hairs minute, otherwise as variable as those of the stems
(but not retrorse), clavate glandular hair idermis often viscid, tube (1.5-)2-4 mm long,
3(-4) mm wide, sometimes keeled from sinus to d base, i nd -reduplicate in the bud, lanceolate to
ovate, 1.5-6 mm long, 1.5-4 mm wide, apex acute, i le corolla in life white, cream,
yellow, or red (reportedly, perhaps in error) in life, the exterior hairs in dry material off-white, pinkish,
tawny, yellowish, orangish, reddish or brownish, interior glabrous; male corolla tube urceolate to nar-
rowly urceolate, narrowing distally, (2.8-)3-7 mm long, 2-4 mm wide, with three shield-shaped, 1—2.5
mm long, glabrous zones at the base (shielded), or sometimes the basal 1/4—1/2 of the tube completely
glabrous, otherwise sericeous with an undercoat of minute curly to eee hairs, minute hairs (including
occasional retrorse hairs) j present between th le corolla lobes
spreading in life, lanceolate, ovate, rhombic or elliptic, 2-3.5 mm long, 2 8 mm wide, hais dense, minute,
curly to wavy, with some straight intramarginal hairs, these sometimes retrorse, median sericeous, at least
basally. Stamens 1214); filaments adnate to the corolla at or near the base, or inserted on the receptacle,
or both, usually in tiers or groups of six stamens, sometimes geminate, 0.8-2 mm long; anthers lanceolate,
ovate or oblong, 1.2-3 mm long, apex acuminate, rostrate, aristate, or bluntly pointed, opening by longitu-
dinal slits. Pistillode minute, densely hairy, the hairs wavy to straight. Female inflorescences solitary
flowers in leaf axils of young (often Ist year) stems, rarely 3-flowered cymes; female flowering pedicels
with hairs and epidermis similar to male inflorescences. Female flowers 3—4(—5)-merous; female flower-
ing calyx mitriform, exterior moderately to densely hairy, the vestiture and epidermis the same as in male
flowering calyces, interior vestiture similar to interior fruiting calyx, tube cupulate to cylindrical, 1.8—5.5
mm long, 3.5-5 mm wide, sometimes very thick, especially basally, lobes valvate-reduplicate in the bud,
winged, broadly rounded to ovate, (2)3-8.5 mm long, 4-10 mm wide, apex vaguely acuminate or bluntly
acute, nerves sometimes conspicuous; female corolla range of coloration similar to that of male corollas;
female corolla tube narrowly to short urceolate or subcylindrical, narrowing distally, 2.7-7 mm long,
2.3-3.5 mm wide, exterior vestiture similar to male corolla tubes, interior glabrous; female corolla lobes
spreading in life, lanceolate to ovate or rhombic, 2-4 mm long, 1-2 mm wide, margins often glabrous and
partly involute, exterior vestiture the same as for male flowers, interior glabrous; ovary depressed- globose
to subglobose, turbinate, or obturbinate, 2-3 mm long, 1.5-4.5 mm wide, sometimes vaguely 3-lobed,
sparsely hairy to sericeous, hairs variously colored, longer hairs sometimes present at the base, rarely with
shield-shaped glabrous zones similar to those of the corolla tube; styles 3(-4), usually fused into a column
part of the way and spreading distally, 1-3 mm long, usually densely hairy; stigmas bifid, about 0.8 mm
long, exterior sometimes hairy. Fruiting pedicels terete to angular, often broader distally, 0.5-9 mm long,
sometimes fissured or half-netted, sometimes lenticellate, vestiture persisting or becoming glabrous, epi-
dermis often viscid, bracts (0—)1—2, alternate or opposite, lanceolate to broadly ovate or deltate, 1.2-2.5 mm
long, 0.92.5 mm wide, flat, concave or navicular. Fruiting calyx slightly accrescent, campanulate or in-
fundibuliform to explanate, the sinus sometimes tearing, exterior vestiture as described for flowering caly-
ces or glabrate, often with pale desiccated remnants of original vestiture, epidermis often conspicuously
viscid and darkened, tube (3—4—10 mm long, sometimes bulbous at the base, interior sparsely to densely
hairs, the hairs usually appressed to subappressed and distally directed, variously colored, hairy to the base
Provance et al., Diospyros salicifolia complex 1021
or glabrous in the lower 1/10—1/2, epidermis sometimes with dark blotches, lobes obtuse to rounded,
rarely acute, reflexed (basally, near the apex, or along the margins) to ascending, (2-23-7.5(-9) mm long,
7-15(-16) mm wide, apex usually abruptly acuminate to a blunt point, interior intramarginal band of hairs
variable in width, the hairs usually dense, minute, wavy, curly or kinked, upright, epidermis usually viscid,
hairs of the median usually dense, straight, appressed, and similar in color to those of the intramargin. Fruit
a subglobose to depressed-globose berry, 1.5-3.5 cm in diameter, consistently with three pairs of locules;
flesh gelatinous, sweet when ripe, with variable amounts of remnant astringency, becoming vitreous, red
and translucent upon drying; epidermis smooth to orange-peel textured, glabrous to sparsely and weakly
hairy when mature, a basal ring of sul l, fine, straight hai ti present (the ring sometimes
adhering to the base of the calyx tube), the apex sometimes umbilicate, usually densely and minutely hairy,
in life green when unripe and yellow, orange, or light brown when ripe, rarely red, in herbarium material
yellow, dark yellow, orange, golden brown, or brown. Seeds 6, wedge-shaped (resembling an orange seg-
ment with rounded edges), 11-16 mm long, 5.5-8 mm radial depth, 3.5-6 mm wide, brick red to dark
brown, texture rugulose-foveolate.
KEY TO THE SUBSPECIES OF DIOSPYROS ACAPULCENSIS IN MESOAMERICA
1. Fruiting pedicels 0.3-1 mm on! a obtrullate, 40- S01 mm pins at “4 length, juu ou abend
hairy below when young, glabra en mature, the cut sli
ri sligh LIGAN S
annular giandular papillae after het hairs fall; fruiting Ed Bed Bede Fas to sparsely une pos
12-13 mm wide, apices reflexed; pine forests of SW Honduras, 750-1120 m subsp. mejocotensis
1. Fruiting pedicels > 2 mm long; lamina rarely obtrullate, rarely over 45 mm wide at 3/4 length; e and
vestiture of lamina and ip. js variable; fruiting calyx lobes 7-16 mm wide, apices reflexed or not;
Jalisco, Mexico to Panam g Honduras
2. Fruiting calyx lobes 7- 7 5 mm long, base of eal tube cm mature lamina (e.g. in fruiting specimens)
membranaceous to subchartaceous, apex narrowly acute to narrowly acuminate, base narrow (length :
width at 0.25 c. 5 : 1), abaxial hairs straight, appressed, Ll directed; male diss terminated 2 female
flowers; anthers yeliow with red dots; Chiquimula vicinity, SE Guatemala, 400-1000 m subsp. chiquimulensis
2. Fruiting calyx lobes usually 3-6 mm long, base of calyx tube swollen or not; mature lamina chartaceous
(rarely membranaceous or subcoriaceous), apex rounded to acute, sometimes acuminate, but not nar-
rowly acuminate (except in subsp. dwyeri, and sometimes in subsp. veraecrucis), less narrowed basally
(except in subsp. dwyerí) (length : width at 0.25 usually < 5 : 1), abaxial hairs variable; inflorescences not
as described above; anthers not red dotted; Jalisco, Mexico to Panama, including Guatemala
3. Hairs of lower lamina surface straight, appressed, distally directed
4. Fruiting pedicels » 6 mm long; fruiting calyx lobes « 9.5 mm wide; leaves narrow, especially basally
(length : wiih at van > 22 : » bs oy 7m US subsp. dwyeri
4. F | 10 mm wide; lamina sometimes narrow,
but less narrow basally (iength : width at 0.25 usually « 5: 1), Mexico to Costa Rica
5. Lower lamina 3° venation not raised, men apparent due ^ ied uir look the span
ofthe four centermost lateral veli sual les 2- m long; lamina 45- 105(- 1 20)
mm long, width at 0.75 length 15- 29 ak fruiting calyx tube E 5.5-9 mm long, the lobes
9-15.5 mm wide; S Mexico (except the Yucatan Peninsula), W Guatemala and El Salvador, <
1240 m subsp. veraecrucis
. Lower lamina 3° venation prominently raised, the span of the 4 centermost lateral veins usually
> 40 mm; petioles 4-8 mm long; lamina 78-130(-150) mm long, width at 0.75 length (2124-40
mm; fruiting calyx tube 3.5-7 mm long, the lobes 6.5-13 mm wide; often near the Pacific Coast
of Nicaragua and NW Costa Rica, « 150 m (rarely to 600 m) subsp. rivensis
3. Hairs of lower lamina surface curly to straight, not appressed, not strictly distally directed.
6. Lower lamina reddish-brown, the vestiture (including the midrib) dense, the hairs curly or kinked,
golden-brown to reddish-brown; petioles rugose, glaucous-pruinose and mauve to dark purple;
fruits n to 351 mi n diameter; Selen forests, id WC Nicele@ue subsp. pedromorenoi
6. L vestiture color various, glabrate to T hairy,
the pus straight to wavy; petioles aor as above; fruits < 30 mm in diameter; W Mexico, Guatemala,
Honduras, Nicaragua, and Costa Rica (one subsp. rare in El Salvador).
7 Lamina often elliptic, length : width 2.1-3.8 : 1; adaxial lamina surface often laden with clusters
of opaque white crystals (crystal-papillose), and often with minute peaks and waves along veins
Un
£ 4L aiD l.l "n
1022 Journal of the Bot f Texas 2(2)
caused by differential drying (tented); hairs on the abaxial midrib surface appressed to subap-
pressed; coastal Niceragua and Costa Rica, « 150(-600) m subsp. rivensis
Lamina oblanceolate to obovate, rarely elliptic, d dis witi 1 J- I dn 2 I adaxial iu
surface crystal-papillose or not, i
spreading or retrorse; W Mexico, Guatemala, Honduras, Nicaragua, and Costa Rica (one subsp.
rare in El Salvador).
N
8. Hairs of mea e gold to reddish- "gold; Halle el ie ond n Sidi pur with
swollen bases, deciduou ]
pu] E e rad +
to Costa subsp. guanacastensis
. Hairs of inflorescence and leaves not as above; fruits not drying dark brown; te e
Mexico, Guatemala, Honduras, Nicaragua, and rarely El Salvador (subsp. nicaraguensis).
9. Fruiting calyx glabrous inside near the base, lobes usually < 10.5 mm wide; lamina (63-)82-
140 mm long, mostly (20-)29-50(-63) mm wide, 24-43(-47) mm wide at 0.75 length; SE
co
Guatemala, Honduras, Arai. rarely collected in El Salvador subsp. nicaraguensis
9. Fruiting calyx hairy inside , lobes rarely < 11 mm wide; lamina 50-82(-127) m
long, usually (18-)23- a —44) mm wide, < 29 mm wide at 0.75 length; W Mexico and W
Guatemala subsp. acapulcensis
la. Diospyros acapulcensis Kunth subsp. acapulcensis. (Figs. 2a—b, 3, 4, 5, 19c).
Trees to shrubs, 2-14 m tall; trunk to 30 cm dbh., bark light grayish-brown; stems glabrous to sparingly
hirsutulous, young stems puberulent and wavy-hairy. Petioles (2.5—)3.5—6(-7) mm long, glab to densely
hairy, the hairs wavy to straight, ascending to retrorse, sometimes glandular above. Lamina chartaceous,
50—105(-128) mm long, (18223-39(-44) mm wide, length to width ratio (1.822—2.8( 3.3) : 1, oblanceolate
to obovate, sometimes nearly elliptic, base cuneate to acutely rounded, sometimes attenuate, decurrent on
the petiole, margin flat to minutely revolute, apex acute to broadly acuminate, occasionally rounded; lower
lamina surface sparsely to densely hairy, the hairs straight to wavy, ascending to upright, not strictly api-
cally directed, clavate glandular hairs present, but deciduous; epidermis dull light brownish- green, the
stomatal apparatus translucent, sometimes opaque and conspicuous; upper lamina surface sparsely to
densely erect-villous, epidermis sometimes crystal-papillose. Venation brochidodromous to arcolanguid;
midrib pubescent or hirsute to wavy-hairy below, especially along the sides, the hairs sometimes ascend-
ing, tawny or orangish, finer above, sometimes retrorse basally; lateral veins 5-8 per side, prominent
below, flush to raised and apparent above; 3? veins barely raised below, flush or slightly raised above.
Male inflorescences 1—3(—5)-flowered cymes; peduncles 1-2 mm long; pedicels 1.5-2 mm long, bracts
oblong-lanceolate, 1-2 mm long, navicular. Male flowering calyx sericeous inside to the base to glabrous,
ascending wavy-tomentulose outside, epidermis viscid and dark, tube 2-4 mm long, 3-3.5 mm wide, lobes
acute, 2-4 mm long, 2-3 mm wide; male corolla tube 3.5-6.5 mm long, 2-3 mm wide, exterior basal 1/3
glabrous, otherwise tawny sericeous, bordering region scantily retrorse-puberulent; male corolla lobes
lanceolate to oblong-ovate, 2 mm long. Stamens 12; filaments adnate in two tiers to the corolla, the in-
ner basal, the outer tier just above, 1-1.8 mm long; anthers oblong-ovate, 2 mm long, rostrate. Female
inflorescences densely hairy, bracts opposite, 1.5-2 mm long, similar to male bracts. Female flowering
calyx short-mitriform, interior cream to tawny sericeous to the base, the intramarginal band wide, exterior
tomentulose, with scant slightly longer, wavy, ascending hairs, tube cupulate, thickened near the base,
4—5.5 mm long, 4.5-5 mm wide, lobes broadly rounded, (2-)4-8 mm long, 8-10 mm wide; female corolla
tube 5.5-6 mm long, 3 mm wide, exterior lower 1/2 glabrous, upper 1/2 sericeous, the bordering region
scantily retrorse-puberulent; female corolla lobes lanceolate to ovate, 2.5 mm long, 1.3 mm wide; ovary
depressed-globose, 2.5 mm long, 4.5 mm wide, minutely cream sericeous; styles 3, + 1.8 mm long, partly
fused, densely hairy; stigmas vaguely bifid, 0.8 mm long, minutely appressed hairy; staminodes (3-)6C-9),
adnate to the corolla from the base to mid-tube, the free portion 1.2-1.4 mm long, sometimes inserted
on the receptacle, the sterile anthers knife-shaped, 1 mm long, basally asymmetric, the filament flattish.
Fruiting pedicels stout, 1-4.5(-6) mm long. Fruiting calyx rotate to campanulate, often splitting at the
sinuses, exterior glabrate or with persisting desiccated hairs, epidermis viscid, dark brown, tube (56-10
Provance et al., Diospyros salicifolia complex 1023
/Mie shu th hapl : h^ h En vi L kall I
Fic. 4. Trunk of D. h is subsp. acapulcensis. Chorros del Varal, Jali
checked. l m
mm long, often bulbous near the pedicel, lobes obtuse, (224—7(-9) mm long, (9.5—-)11-15(-16) mm wide,
reflexed, margins sometimes reflexed. Fruit subglobose to depressed-globose, 1.5-2 cm long, 1.5-2.5 cm
in diameter, with slightly wavy to straight golden subappressed hairs at the base, these dense at the apex,
epidermis often glaucous-pruinose and/or scintillant, yellowish to golden brown. Seeds 11-14 mm long,
6—8 mm radial depth, 4—5 mm wide.
Diospyros acapulcensis subsp. acapulcensis is nearly endemic to western and southwestern Mexico, but is
unknown from Oaxaca (Fig. 23), and is known elsewhere only from a single collection from Guatemala. It
is an infrequent to occasional component of matorral, tropical deciduous forest, seasonal evergreen forest,
and oak forest, in canyons and on slopes and mesas below 1350 m. Like many taxa in the complex, it may
persist or be spared along roadsides and at the edges of cultivated land. It resembles D. a. subsp. veraecrucis,
but is differentiated by the abaxial lamina pubescence (see subsp. veraecrucis) and the shape of the lamina.
It also closely resembles D. a. subsp. nicaraguensis, but that differs in having a lamina that tends to be longer
and wider, narrower fruiting calyx lobes, and a calyx tube that is usually not sericeous inside to the base.
Representative specimens. GUATEMALA. Ro oe de los Cuchumatanes, 800-1200 m, 30 iuda 1942, J.A. Steyermark
51584 (F). MEXICO. CHIAPAS Mpio Chi p horreadero, 8 m, li Aug 1 972, D.E. B 874 MO, CHAPA).
1024 J lof the Botanical R h Institute of Texas 2(2)
Fic. 5. Diospyros acapulcensis subsp. LT A. Fruiting branchlets (M.C. Provance et al. 9398, VERI. B. Male flower Le » Hinton a al boda ia
C. Female immature flower (J. Steyermark, F). D-E. Based on E. Carranca aE, hin M (IEB). D
EET YA I £. £ L J HI T HN
Adaxial surface of male calyx (Kruse ick MO). G-H. Based on J y
Provance et al., Diospyros salicifolia complex 1025
rán: 6.5 km W of Tuxtla Gutiérrez, 600 m, 8 Oct 1971, D.E. Breedlove 20121 (MO). Mpio. Tonalá: Tonalá, 100 m, 6 May 1988, J.I.
in 13814 (XAL). Mpio. Tuxtla Gutiérrez: Tuxla Gutiérrez-El Sumidero, km 16, 990 m, 19 May 1982, J.I. Calzada 9223 (IEB, iid
Mpio. Tzimol: 15 km S of Comitan, 1200 m, 19 Nov 1980, D.E. Breedlove & F. Almeda 47619 (MO). Mpio. Venustiano Carranza: abov
Soyatitán, 3400 ft, 25 Oct 1967, A. Shilom Ton 3105 (CAS). Mpio. Villa Corzo: near El Brillante, 650 m, 9 Oct 1974, D.E. Breedlove 38411
(MO). Corma. Mpio. Comala: cerca de Campo Cuatro, 1200-1300 m, 5 May 1998, R. Cuevas 5858 (ZEA). Mpio. Tecoman: 1.5-2.5 km
de la empresa “Apasco,” 400-560 m, 6 Apr 1996, R. Cuevas & L. Guzmán 5133 (ZEA). México. Mpio. Temascaltepec: Palmar, 13 May
1934, G.B. Hinton et al. 6044 (MO). Guerrero. Mpio. Acapulco de Juarez: Río Papagayo, 3 km NW de Xalapa, 200 m, 17 Jan 1994, R.
Fernandez N. 4929 (IEB). Mpio. Chilpancingo: Salto Valadéz, 1350 m, 25 Jan 1970, H. Kruse 2692 (MO); al N de Ácahuizotla, camino a
Palo Blanco, {17°23'N, 99°27'W], 930-1050 m, 19 Jan 1994, E. Carranza & E. Perez C. 4743 (IEB). Mpio. Iguala: E de Tierra Colorada,
300 m, 8 Jul 1966, J. mm e ei (MSC). Mpio. Mochitlán: Agua de Obispo, barranca del Cuarenta, 945 m, 15 Jul 1967, H. Kruse
1636 (MO). Mpio. S le, 310 m, 31 Jan 1983, J.S. Miller & P. Tenorio L. 553 (MO). Mpio. Taxco: 7 mi N of “Centro”
sign at Iguala, 25 Jan 1971 Bde icon 6 L, Spetzman 5 50 ena. NA sud Mpio. Autlan de Navarro: Arroyo El Coajinque, 1000 m,
3 Oct 1998, R 587. A). Mpio. C Pesada-Rancho Los Patos, 395 m, 18 Nov 2000, R. Cuevas
6910 (ZEA). Mpio. El Grullo: A El Colomo, 1000 m, 7 Nov 1988, F. Santana M. & W. Anderson 4068 (ZEA). Mpio. Puerto Vallarta:
cerca Fracc. Colinas Campestres, 150 m, 26 Aug 1990, M, Cházaro B. & O. Montes 6405 (IEB, UCR). Mpio. San Sebastián del Oeste: 18
km WNW of San Sebastian, 410 m, 7 jan 1990, T.S. Cochrane et al. 12026 (IEB). Mpio. Santa Maria del Oro: Santa Maria del c 900
m, 19 Dec 2005, I. Garcia R. & M.C. Provance 7359 (CIMI, UCR). Mpio. Tolimán: 8—9 km adelante de Campo Cuatro, 1400 m, R. Cuevas
4039 (ZEA). Micnoacan. Mpio. Aquila: 3 km E de Aquila, 250 m, 29 Feb 1980, B. Guerrero C. 742 ia oe La Huacana: Limon
eee 690 m, 15 Jun 2002, X. Madrigal S. & S. Ontivaros A. 4803 (EBUM). M: I ): Chorros del Varal,
950—1000 m, 30 Jun 2005, I. Garcia et al. 7122 (CIMI, UCR); Puente de Iturria, dio 04, M. C. E et al. 9398 (UCR). MonrzLos.
Mpio. Tepalcingo de Hidalgo: 1 km SE de El Limón, 1310 m, 25 Feb 1991, J. Bonilla 1434 (IEB, UCR)
Ib. n e Kunth subsp. chiquimulensis M.C. Provance, I. García & A.C. Sanders, subsp.
v. (Fi: igs. 19a—b). Tee: GUATEMALA. CHIQUIMULA: "Frecuente a la entrada de Chiquimula,” [ca. 14?48'N, 89°33'W], “100
m” asd 400-1000 m], 20 May 1963, A. Molina R. & A.R. Molina 12473 (noLorvrE: F-1622586).
Arbo Ores a Tu 1 Y I j 1 m Ta a E £f, 1 LIE NES AA E j
rectis a AME ii i inatis et pistillatis, is pistillatis i i pi
"uo" "T " :
Trees 4—5 m tall; trunk unknown; stems glabrous to sparsely hairy, the young stems sparsely hairy, the
hairs apically curved to slightly wavy and subappressed, clavate glandular hairs present. Petioles minutely
winged, 4 mm long, glandular-viscid, appressed to subappressed-puberulent below, sparsely ascending-
puberulent above. Lamina membranaceous to subchartaceous, 80-103 mm long, 25-35 mm wide, length
to width ratio 3— i I; Ao base acute to uii, cuneate, m on the petiole, margin + flat,
1 1 1 pa]
ciliate, L acuminate; y to moderately appr essed-
pubescent, clavate dila Male is present, ad ek me epidermis hent Brown, the os
apparatus l andi
present, the epidermis dark low densely patsillate, somewhat verted. Veuation brotbidedroniouts midrib
subprominent below, with scant minute, appressed hairs, sparsely and minutely hairy above, sometimes with
deltoid scales, BORD MEC fae veins pes 12 per side, fine below, slightly raised above; 3-4? veins
darkened below, i 3—4(—7)-flowered cymes (terminated by a female
flower), densely nuc. hairy; peduncles 2.5-3.5 mm long; pedicels 1-2 mm long, bracts lance-ovate,
2-2.5 mm long, navicular. Male flowering calyx finely subappressed wavy-hairy outside, epidermis dark
brown, viscidulous, tube 3 mm long, 3 mm wide, often keeled from sinus to base, mostly glabrous inside,
lobes ovate, 3.5-4 mm long, 4 mm wide, slightly winged, acuminate to a point, glandular, mostly sericeous
inside; male corolla tube 2.8 mm long, 2 mm wide, thick-walled, exterior base with three glabrous shield-
shaped zones, upper 1/2 densely buff sericeous; male corolla lobes (immature) thick, lanceolate, 1.5 mm
long, 1 mm wide. Stamens 12; filaments adnate to the corolla towards the base, but at unequal levels,
1 mm long, sometimes geminate, connective red; anthers lanceolate to lance-acuminate, 1.5-2 mm long,
yellow with minute red dots. Female inflorescences solitary flowers (those seen somewhat immature), or
terminal flowers in otherwise male inflorescences, bracts lanceolate, up to 2.5 mm long, + keeled. Female
flowering calyx densely hairy outside, the hairs minute, slightly wavy, golden, tube 4 mm long, 3.5 mm
wide, very thick, especially near the base, glabrous inside near the base, otherwise densely straight-hairy,
1026
f Texas 2(2)
laf +k
Fic. 6. Diospy pulcensis K bsp. chiq lensis M.C. Provance, I. García & A.C. Sanders, subsp. nov. Based on the holoty; (A. Molina R. & A.R.
Molina 12473, F-1622586). A. F gl hlet. B. Mal lla lobe (abaxial side). C. S (lateral view). D. S (adaxial ). E. immature
male flower. F. I t I lla. G. Ad Isurf fi lefl i 3 ly H. Abaxial surf. fi f lefi i E
ariel li aye! female corolla. K. Femal be (adaxial side). L. Pistil
Provance et al., Diospyros salicifolia complex 1027
these horizontally oriented, reddish-gold, lobes 8.5 mm long, 7.5 mm o pur up band Pn)
wide, hairs golden, median hairs! ly and distally oriented, the ep
6-7 mm long, exterior lower 1/2 glabrous, upper 1/2 with three pee spaced sericeous areas, the hairs
of the intervening regions minute, horizontally oriented, upper and lower-halves separated by an undulate
ring of minute retrorse hairs, epidermis dark reddish-brown; female corolla lobes lanceolate, 2-2.3 mm
long, 1 mm wide; ovary subglobose, 2 mm long, 1.5 mm wide, minutely cream sericeous, with clusters
of longer hairs from the base, three shield-shaped glabrous zones also me SON the base; styles 3,
fused into a 1.5 mm long column, minutely hairy; stigmas unknown (d 3, lanceolate,
adnate to the corolla. Fruiting pedicels stout, + 3 mm long, densely hairy Fruiting calyx campanulate
to infundibuliform, similar outside to the flowering calyx, tube 5-6 mm long, bulbous at the base, lobes
obtuse, 7-7.5 mm long, 11 mm wide, reflexed near the apex. Fruit (probably immature) + 1.5 cm diameter,
moderately hairy, the hairs minute, subappressed, nearly straight, gold, epidermis glaucous-pruinose, light
golden brown. Seeds unknown.
This taxon is notable for male flowering cymes terminated by female flowers; its large, thin, leaves with
acuminate apices coming to distinct points; horizontal hairs on the inside of the female calyx and the outside
of the female corolla tube; and yellow anthers with red dots. Further study may reveal that this taxon war-
rants species rank, but we are following a conservative course in placing it at the subspecific level, because
we've only seen a single specimen. Of related taxa, it is most similar to D. a. subsp. veraecrucis. The single
collection was made near Chiquimula, Guatemala, at an elevation of *100 m" (Figs. 23, 24). The reported
elevation is likely a typographical error for 1000 m, as the lowest elevation in the area is 400 m. This species
is named for the Department of Chiquimula, Guatemala, where the type specimen was collected.
-
lc. Mus o Kunth subsp. dwyeri M.C. aan I. Garcia & A.C. Sanders, imd nov. (Figs.
, 21€). Tee: PAN , Farfan Beach, road h, [8°56'N, 79°35'W, 1-5m], 3 Aug 1967, J.D. Dwyer &
S.M.V Hayden 7525 o MO-2067635!; isotypes: MO-2129488!, E "o image, UC, COL, “Moldenke” Ens at LL]).
E Á 1 45 £f, I D ESSET 1 : +1 ates s x ,etlaminis
O 4 e
uec caps meu 3 : EE 1 E 1
foliorum amplioribas ai rationi lengitudod g pressis ap versus.
Shrubs to 4.5 m tall; trunk unknown; stems glabrous when mature, young stems sparsely upright to
subappressed-villous and densely covered in shorter upright to ascending, curved to kinked, yellowish to
golden-brown hairs, the epidermis glistening, brownish mottled. Petioles minutely winged, 4—5 mm long,
yellowish, hairy below, densely and conspicuously yellowish hirsutulous to pubescent above. Lamina
subchartaceous to chartaceous, 60—70(-80) mm long, 15-20 mm wide, length to width ratio 3.2-4 : 1,
oblanceolate, base cuneate to narrowly acute, decurrent on the petiole, margin flat to minutely revolute,
apex narrowly acuminate with a blunt point; lower lamina surface sparsely + appressed-pubescent, the
hairs 0.5-1.5 mm long, densest near the margin, clavate glandular hairs present, the epidermis tan to light
greenish-brown, the stomatal apparatus translucent and inconspicuous; upper lamina surface sparsely
puberulent, the epidermis brownish-green, even to vaguely tented, densely crystal-papillose. Venation
arcolanguid; midrib prominent below, subappressed to appressed puberulent, the hairs mostly » 1 mm,
yellowish, clavate glandular hairs present, on the upper surface flush to slightly raised, puberulent to short
wavy-hairy, the epidermis yellowish; lateral veins 6—8 per side, yellowish, prominent below, apparent
above; 3°-4° veins fine and barely raised below, sometimes darkened, inconspicuous above. Male inflo-
rescences and flowers unknown. Female inflorescences and flowers unknown, except style remnants
on immature fruit columnar, + 3 mm long, minutely hairy. Fruiting pedicels slender, 6-8 mm long, bracts
narrowly oblong, minute, concave to navicular. Fruiting calyx densely hairy outside, the hairs minute,
wavy, subappressed to ascending, reddish-gold, epidermis dark, tube half-globose, 4—6 mm long, sparsely
golden sericeous inside, lobes spreading, rounded, 4—5 mm long, 7-9 mm wide, acuminate to an obscure
point, interior mostly reddish-gold sericeous, golden tomentulose at the intramargin. Fruit (immature)
globose, the hairs straight to curved, 0.1-0.5 mm long, appressed to subappressed except basally where
straight, fine, x 1 mm long, epidermis dark reddish-brown. Seeds unknown.
oo B <
Cva : :
l^ ¿libros
TAN
xr UNE RSS y HERBARLA
ANTI
al atgel | ik.
E Bitte a 4:
by White (1927-2994)
"i 4 D0 3
DAC S fuco S uo c ro W.*-
IA
PLANTS OF PANAMA
Pistituted by the Misscwsd Branca? Gantin Horton, (MOS
e i d me i HIT esit k
Family: gree bena Dorr
Eberacea
Diuapyros sr.
qRoPicos Ce D
Exsiccalao Ne: T7525 961.
pes d, Dwyer e, MOCZ, UC, 0
R ee tas r^ =
Aided by e proat Pega Us
Fic. 7. Holotype of D. acapulcensis Kunth subsp. dwyeri M.C. Provance, 1.
2067635).
García & A.C. Sanders, subsp. nov. (J.D. Dwyer & 5.M.V. Hayden 7525, MO-
Provance et al., Diospyros salicifolia complex 1029
This subspecies seems endemic to Panama, the only collection seen being from Farfan Beach, near
the Pacific entrance to the Panama Canal (Fig. 24). Vegetation mapped in this area includes semideciduous
forest and periodically inundated tree savanna with a grassy herb layer (ANAM and CBMAP 2000). There
may also be patches of swamp forests in the vicinity. Unseen collections of Diospyros salicifolia’ cited for
Panama (White 1978, D'Arcy 1987) may represent this subspecies, and include a collection from Cativo
Swamp, Darien Province (Duke 11739a). This taxon is easily separated from D. a. subsp. veraecrucis by its
much longer fruiting pedicels, and from subsp. D. a. subsp. rivensis by its shorter, narrower lamina, that is
especially narrower relative to width at 3/4 length, and calyx tube that is narrower relative to width as well.
This subspecies is named in honor of the late John Dwyer, collector of the type specimen and renowned
plant taxonomist, ethnobotanist, professor at Saint Louis University, and research associate of the Missouri
Botanical Garden.
1d. Diospyros acapulcensis Kunth subsp. guanacastensis M.C. Provance, I. García & A.C. Sanders, subsp.
nov. (F igs. 8, 202p TYPE: COSTA RICA, Guanacaste: Parque Nacional Palo Verde, Sector Catalina, Camino y Sendero Botija
Saliendo al Camp terrizaje del Refugio Palo Verde, 10°21'N, 85°21'W, 20 m, 26 Nov 1991, U. Chavarría 427 (morore: MO-
57006321; : INB)
rbores a y Vi. AMLO ALMAS VCOLIILICIILU
aei cas deo ac minii.
Trees 2-10 m tall; trunk to 30 cm in diameter, whitish to grayish-white; stems glabrous or with ashy
remnants of hairs when mature, densely hairy when young, the hairs ascending to upright, bright reddish-
orange to brownish, the epidermis shiny and viscid. Petioles 4—7 mm long, rugose, viscid and shiny, brick
red to dark reddish-brown, densely pubescent below, the hairs bright, blond to reddish-gold, shorter above.
Lamina chartaceous, 45—85(-95) mm long, 20-35 mm wide, length to width ratio (1.82-2.8 : 1, oblan-
ceolate to obovate or oblong-ovate, sometimes nearly elliptic, base acute to obtuse, abruptly decurrent on
the petiole, margin flat to minutely revolute, sometimes ciliate, apex acute to obtuse; lower lamina surface
ascending-pubescent, the hairs golden to ferruginous, clavate glandular hairs often present, the epidermis
dark reddish-brown and sometimes black-mottled, the stomatal apparatus translucent and inconspicuous;
upper lamina surface densely wavy-hairy, the hairs deciduous, leaving seeping annular papillae, clavate
glandular hairs often present, deciduous, leaving the epidermis viscid, the epidermis dark, often tented,
crystal-papillose. Venation eucamptodromous, viscid, prominent below, often 3-ribbed from the base; midrib
rugulose below, densely hairy, the long ascending to spreading hairs sometimes sticking to the epidermis,
midrib impressed above and wavy-hairy; lateral veins 6—9 per side; 3? veins on lower surface similar to
the midrib below. Male inflorescences 3—5-flowered cymes, partly ascending-pubescent, partly tomen-
tulose; peduncles 1-2 mm long; pedicels 1-2 mm long, bracts lance-ovate, 1.5-2.5 mm long, navicular.
Male flowering calyx glabrous inside except for a very narrow intramarginal band on the lobes, exterior
partly wavy subappressed to ascending-pubescent, partly tomentulose, the epidermis viscid, dark brown,
tube 3.5 mm long, 3 mm wide, sometimes vaguely keeled from the sinus to the base, lobes acute, 2-2.5 mm
long, 2 mm wide; male corolla tube 6 mm long, 2 mm wide, sericeous except for 3 basal shield-shaped
glabrous zones, + 1.5 mm long, these minutely puberulent distally; male corolla lobes ovate to elliptic, 3
mm long, 1 mm wide. Stamens 12; filaments adnate to the base of the corolla, a few sometimes inserted on
the receptacle, 1.5-2 mm long, sometimes geminate, red; anthers lanceolate, 2 mm long, aristate. Female
inflorescences and flowers unknown except styles remnants on immature fruit columnar, hairy, 3 in
number. Fruiting pedicels stout, 0.5—2.5(-5) mm long, fissured to shallowly grooved, sparsely to densely
hairy, the hairs reddish-gold, bracts ovate to deltoid and sharply pointed, 1.2-1.6 mm long, flat to vaguely
concave. Fruiting calyx semi-globose when immature, explanate and torn at the sinuses when mature,
densely hairy outside, in part tomentulose, in partly villous, the hairs tawny to brilliant reddish-gold, +
deciduous, the epidermis dark brown, shiny and viscid, tube 6-10 mm long, without a bulbous base, the
upper 1/2 densely hairy inside, the lower 1/2 glabrate, lobes obtuse, 3.5-7.5 mm long, 9.5-13 mm wide,
barely acuminate to a point, usually reflexed, densely hairy inside with wide intramarginal band, the hairs
1030
+
Texas 2(2)
427,N MO- 5700632) AB ture fi
([ ). E. Mal de). F. Stamen. G. Ad
J
I
M.C. Provance, I. García & A.C. Sanders, subsp. nov. A-B. Based on the holotype (U. Chavarría
C e A M oe 15605 (BH). C. Male flower. D. Male corolla
Provance et al., Diospyros salicifolia complex 1031
tawny. Fruit 1.5—2(-2.3) cm in diameter, gold to reddish-brown ascending-velutinous until nearly mature,
then glabrate, glaucous-pruinose, sometimes minutely gland-dotted, reportedly green (when mature) to
orange-brown (presumably when fully ripe) in life, epidermis brownish, and sometimes quite dark, in
herbarium specimens. Seeds 14 mm long, 6 mm radial depth, 4 mm wide.
This subspecies is geographically isolated from both D. a. subsp. nicaraguensis and D. a. subsp. pedro-
morenoi in Nicaragua, being endemic to dry tropical forest near sea level to 800 m in NW and central Costa
Rica (Fig. 24). In NW Costa Rica it has been collected on the Nicoya Peninsula at Barra Honda National
Park, the Island of San Lucas, and the southern slopes of the Cordillera de Guanacaste at Rincón de la Vieja
National Park. Other sites include the vicinities of Bagaces, Cañas, and Palo Verde National Park. The NW
limit of this subspecies appears to be Santa Rosa National Park. Material intermediate with D. a. subsp.
rivensis has been collected on Cerro El Hacha at 1100 m. Collections from near sea level at the SE extreme
of the Nicoya Peninsula, and from the margins of the Candelaria River in central Costa Rica are difficult to
place, and may be intermediates with D. a. subsp. rivensis. Some speci from the Department of Guana-
caste have rather distinctive leaves and fruiting calyces (as seen in the type) that are very viscid, have a dark
reddish-brown epidermis, and a brilliant reddish-gold vestiture. Annular glands on both leaf surfaces are
derived from deciduous hairs that have left the basal portion of the fluid-filled trichome and/or the swollen
region of the epidermis at the base of the trichome. These are particularly conspicuous in collections from
Palo Verde National Park and San Lucas Island. This subspecies differs from D. a. subsp. nicaraguensis in its
longer fruiting calyx tube, shorter distance between superadjacent lateral veins, intense golden to reddish-
brown vestiture and dark fruits. Compared to D. a. subsp. pedromorenoi, this subspecies has smaller fruits,
shorter fruiting pedicels, a longer fruiting calyx tube, and different stem, inflorescence, and abaxial lamina
vestiture.
dose e specimens. COSTA RICA. ALajuELA: entre Bagaces y Los Cañas, 29 May 1932, A.M. Brenes 15605 (BH). GUANACASTE: P.N.
ón de la Vieja, 700—800 m, 16 Oct 1990, Gerardo Rivera 780 MO); P.N. Palo Verde, 10-150 m, 9 Jan 1992, U. ión 475 Pe
near ac 50 m, 26 Apr 1972, P.A. Opler 761 (MO); 15 km S of Liberia, 170 m, 4 Jul 1971, W.E. Harmon & J.A MO);
Hacienda La Pacifica, near Cañas, 19 Aug 1985, D.S. Seigler DS-12401 (MO, ILL), same location, 14 Aug 1986, D.S. Seigler DS-12782 a
ILL); Llanos de Cortes, 24 Jan 1984, T.D. di 6 in iiis 11420 (MO); E.N. Rincón de la Vieja, Est. d Pailas, 800 m, 3 Nov
1993, R. Espinoza 618 (MO); Rosa N.P., t de Costa Rica, 100 m, 25 Jan 1983 Garwoo d et 6 (MO); P.N. Santa
Rosa, Bahía Salinas a bius Cecilia, 300 m, 14 Jul 1994, R. Espinoza 1119 (MO); Cantón de Nicoya, P.N. Barra Honda, 300—400 m, 9
Jun 1993, Reyes 164 (MO); Est. Biol. Palo Verde, 20-90 m, 20 Dec 1992, Quigley 931 (LSU); Tempisque, Est. Biol. ue Vi Er 10-100
m, 12 Dec 1990, U. Chavarría 199 (MO); Ferri del Tempisque del lado de Cañas, 28 Aug 1985, Poveda et al. 4023 (VT). Puntarenas: Isla
San Lucas, 0 m, 29 Aug 1985, N. Zamora V. 1087 (MO, VT); Cantón de Puntarenas Monteverde, Río Guacimal, 600-800 m, 7 Aug 1992
W. Haber et al. 11326 (BM).
le. s TRECE dues subsp. mejocotensis M.C. Provance, I. García & A.C. Sanders, subsp.
v. (Figs. 9, 21a—b). te: HONDURAS. Leurma: Río pan. 9 km de Gracias, [14°32'N, 88°31'W], 1000 m, 7-9 Dec
P C. Nelson, M. oon GA. la 184 (HoLOTYPE: MO-4594757 TYPE: TEFH-3807!).
A vl TH 1 f J I ji 1 UE s DOE 1 2 : TI 1 11 pp : Sj praesertim ad 3/4 longitudinem,
PEE e A EH f, B ie Of PEN ;NI ENSE ere S eT 1
Trees 6 m tall; trunk unknown; stems glabrous when mature, the young stems densely hairy, the hairs
mostly 0.2-0.6 mm long, slightly wavy, ascending to upright, colorless to reddish-brown, the epidermis
viscid, dark reddish-brown. Petioles 4-6 mm long, rugose, the epidermis very viscid, golden-brown, hairy
below with wavy to straight glossy hairs, these subappressed. Lamina chartaceous, 80-119 mm long, 35-54
mm wide (40-50 mm at 3/4 length), length to width ratio 1.9-2.5 : 1, obovate to obtrullate, base acute,
decurrent on the petiole, margin flat, apex obtusely-acuminate; lower lamina surface densely hairy when
young, the hairs curly to slightly wavy, clear to reddish-brown, glabrate when mature, the bases of the hairs
slightly swollen, leaving glandular annular papillae upon loss of the hairs; upper lamina surface initially
densely ascending wavy-hairy, also with some clavate glandular hairs, but mature leaves glabrate. Venation
arcolanguid to eucamptodromous, the epidermis viscid, reddish-brown; midrib prominent below and at
the base on the upper surface, vestiture similar to petiole; lateral veins 7-8 per side, finely raised above,
1032 J lof the Botanical R h Institute of Texas 2(2)
Fic. 9. Di D) | fi is Kunth sub I mejocotensis M C. Provance, | García & A.C Sanders, subsp nov. A. Fruiti g
the holotype (C. Nelson et al. 184, M0-4594757). B-D. Based on $. Blackmore & M. Chorley 3903 (TEFH). B. Male flower. C. M l
portion). D. Stamen.
hlets, based on
Provance et al., Diospyros salicifolia complex 1033
impressed along the midrib-lamina seam; 3° venation barely raised above and below. Male inflorescences
3(-6)-flowered cymes, pubescence similar to young stems; peduncles 2.5-4 mm long; pedicels 1-2 mm
long, bracts 1.8 mm long, narrowly elliptic, navicular. Male flowering calyx glabrous inside, minutely
reddish-brown kinky-hairy outside, with scattered short crooked ascending hairs, the epidermis viscid,
dark reddish-brown, tube 3 mm long, 3 mm wide, lobes acute, 4 mm long, 3 mm wide, glabrate inside, with
only a few straight appressed hairs near the apex and a narrow tomentulose intramarginal band, similar
to the tube outside, but the hairs sparser, sometimes subappressed; male corolla tube 5 mm long, 2 mm
wide, tawny to golden brown sericeous outside, except for 3 glabrous regions at the base, these sometimes
continuing onto the petal lobe (the hairs apparently readily deciduous), the epidermis dark reddish-brown;
male corolla lobes rhombic, strongly involute on one or both sides, 2.8 mm long, 2.8 mm wide. Stamens
12; filaments adnate to the corolla at or near the base, 1.4 mm long, dark brown; anthers lanceolate,
1.7-2.0 mm long, rigidly pointed, golden brown. Female inflorescences and flowers unknown. Fruiting
pedicels stout, 0.3-1 mm long, bracts 0. Fruiting calyx campanulate, glabrate to sparsely hairy outside,
sometimes with desiccated hairs stuck to the viscid, horizontally wrinkled, very dark epidermis, tube 7-8
mm long, base not bulbous, minutely tawny sericeous almost completely to the base, lobes widely obtuse,
5-6 mm long, 12-13 mm wide, vaguely acuminate to a blunt point, reflexed, tawny sericeous inside ex-
cept for a broad tomentulose intramarginal band. Fruit globose to subglobose, 2 cm in diameter, densely
subappressed golden puberulent near the apex, otherwise glabrate, epidermis glaucous-pruinose, slightly
scintillant, mauve, orange-brown where the cuticular wax has been removed. Seeds 13 mm long, 6 mm
radial depth, 3.5 mm wide.
This taxon is similar to D. a. subsp. pedromorenoi, but has nearly sessile fruit and obtrullate leaves. The
fruiting calyces are similar to those of D. a. subsp. veraecrucis, but the leaves are differently shaped and have
kinky to curly, upright, reddish hairs abaxially in contrast to the straight, appressed, distally-directed hairs
of D. a. subsp. veraecrucis.
This subspecies has been collected twice in the Mejocote River Valley in southwestern Honduras,
just E and NE of the Cordillera de Celaque (Figs. 23, 24); once in pine forests at 1000 m, and again along
a tributary of the Mejocote River at about 750 m, + 20 km NNW of this site, from “rocky cliffs over large
river" (S. Blackmore & M. Chorley 3903). The known collections of this taxon come from places near Celaque
National Park, and so it should be sought within the park boundaries along arroyos and watercourses at
lower elevations.
Specimen ined. HONDURAS. Lempra: near Las Flores, on road Santa Rosa de Copán- Gracias [14°42'N, 88°36'W, 752 m], 13
May 1987, S. Blackmore & M. Chorley 3903 (TEFH, BM not seen).
1f. Diospyros acapulcensis Kunth PRA == (Standl.). M.C. Provance, I. García & A.C. Sanders,
comb. et stat. nov. (Figs. 10, 11, 18a-e). Bason: Maba nicaraguensis Standl. Contr. U.S. Natl. Herb. 20:193. 1919.
Diospyros nicaraguensis duct Standl. Publ. ec. Inst. Wash. 461: 80. 1935. Tree: NICARAGUA. Granapa: 16 Feb 1906, C.F
Baker 629 (HOLOTYPE: US-862725 as microfiche UCSB).
Maba nicaraguensis Standl. J. Wash. Acad. Sci. 17:526. 1927. nom. superf. Tree: NICARGUA: near Managua, 16 Ápr 1926, D. Chavez 206
(HOLOTYPE: US-1266111).
Trees and shrubs, 1-12(-20) m tall; trunk up to 30 cm in diameter, sometimes with multiple trunks,
bark smooth, grayish to reddish-brown; stems sometimes with scant persisting hairs when mature, young
stems pilose to villous, sometimes sublanate, the hairs blond to dull orange, sometimes cod partly retrorse
hirsutulous, with clavate glandular hairs often present, but deciduous, as a result th is viscid, often
mottled with black shiny regions. Petioles narrowly edged, (323.5—6.5(-8) mm TM epidermis rugose,
viscid, becoming glabrous, but initially densely hairy above and below, the hairs wavy to straight and ascend-
ing to retrorse, clavate glandular hairs often present. Lamina membranaceous in spring, later chartaceous,
50-120C-140) mm long, (20228—47(-65) mm wide, length to width ratio (1.7-21.8—2.7(—3.3) : 1, elliptic to
obovate-elliptic or oblanceolate-elliptic, base one) to acae an Ru on m Linc pn
tO
flat to minutely revolute, apex acute to rounded, f y
1034 | Iaftho Patanical D it
f Texas 2(2)
A]
ALT
i —— |
OGXFORD UNIVERSITY I
Nase allocated 10 Uma spo
by E White dress
AM. Buupgnell Apel 199
i)
erparie
PLANT ICAR AGLI
V. DAKEN TERNAS M al Pa
; : E vi
" x dà gl] Wi ;
Fic. 10. Di py j ji is Kunth sut I j 9 ic fi G d Dept of Granada Nicaragua A. Abaxial leaf surf. (CF. Baker 157, ARIZ). B-C.
(CF Baker 157, MO). B. Fruiting | ile | hlets. C. Detail of fruitina E hlet
densely hairy, the hairs straight to wavy, ascending to upright, yellowish to dull orange, clavate glandular hairs
often present, but deciduous, epidermis brownish-green, the stomatal apparatus translucent to opaque often
conspicuous; upper lamina surface villous, clavate glandular hairs often present, but deciduous, epidermis
dark grayish-green, sometimes crystal-papillose. Venation eucamptodromous; midrib prominent below,
hirsute, especially along the sides, some hairs slightly wavy, hairs curly to straight above, mostly upright
Provance et al., Diospyros salicifolia complex 1035
Fic. 11. Diospyros acapulcensis subsp. nicaraguensis from 13.5 km W of Juigalpa, Dept. of Chontales, Nicaragua (M. Nee 27612, MO).
but ascending to retrorse basally; lateral veins (5—)7—10(-12) per side, prominent below, raised above; 3?
veins prominent below, usually raised above; 4? veins barely raised below, sometimes darkened in young
leaves. Male inflorescences 1—3-flowered cymes, densely minutely wavy-hairy; peduncles (3-)4-6 mm
long; pedicels 1.5-2 mm long, bracts lanceolate, 1.5 mm long. Male flowering calyx densely tomentulose
outside, the epidermis viscid, dark brown; tube 2.5—4 mm long, 3-4.5 mm wide, glabrous inside, lobes acute,
3.5-6 mm long, 2-4 mm wide, bluntly pointed, sericeous inside except for a narrow intramarginal band;
male corolla tube 5-7 mm long, 2-3 mm wide, cream to pinkish sericeous outside, with 3-4 shield-shaped
glabrous basal regions, glabrous and sericeous areas sometimes separated by a few minute retrorse hairs;
male corolla lobes white, cream, yellow, or red (reportedly, perhaps in error), lanceolate, ovate or rhombic,
2-4 mm long, 1-2 mm wide. Stamens 12; filaments adnate to the corolla at or near the base of the tube,
sometimes inserted on the receptacle, 0.8—1.5 mm long, sometimes geminate; anthers lanceolate to oblong,
1.5-2 mm long, apiculate to rostrate. Female inflorescence densely hairy, bracts 1—2, opposite or alternate,
1.6-2.5 mm long, lanceolate to broadly ovate, acuminate, slightly concave to navicular. Female flowering
calyx sericeous inside, except glabrous basally and lobes with a broad tomentulose intramarginal band,
1036 J | of the Botanical R h Institute of Texas 2(2)
densely and minutely wavy-hairy outside, tube cupulate, thickened, especially basally, 1.8-2 mm long, 5 mm
wide, lobes obtuse, 5-6 mm long, 7.5-8 mm wide; female corolla tube 5.5 mm long, 3 mm wide, upper
1/2 sericeous outside, basal 1/2 with 3-4 shield-shaped glabrous areas, straight retrorse hairs sometimes
separating the glabrous and sericeous regions; female corolla lobes ovate, lanceolate or rhombic, 3 mm
long, 1.8-2 mm wide; ovary subglobose to depressed-globose, densely and minutely hairy; styles 3(—4),
+ 1.3 mm long, partly fused, hairy; stigmas + 0.8 mm long, minutely hairy; staminodes + 6, adnate to
base of corolla, filaments, 1.8 mm long, sterile anthers 1 mm long, lanceolate, asymmetric basally. Fruiting
pedicels stout, (0.521—5(-7) mm long, glabrate to densely hairy. Fruiting calyx subrotate to campanulate,
densely hairy outside, sometimes glabrate, tube (324.5-8(-9) mm long, base often bulbous, lobes obtuse,
3—6.5(-7.5) mm long, (7-)9-13(-16) mm wide, reflexed; fruit subglobose to depressed-globose, 1.5-2.5
cm in diameter, glaucous, scintillant, yellowish, orangish or light brown, living material reportedly orange
when ripe. Seeds 14 mm long, 7 mm radial depth, 5 mm wide.
This subspecies is found from near sea level to 1500 m in a wide range of habitats, including tropi-
cal deciduous forest, semideciduous forest, and pine-oak forest, in the cordilleras of southern Nicaragua,
Honduras, northeastern El Salvador, and eastern Guatemala (Figs. 23, 24). This taxon is most similar to D.
a. subsp. guanacastensis, which has a longer fruiting calyx tube, and a gold to reddish-brown vestiture and
meh a It EG End with D. a. subsp. pedromorenoi in several areas of west-central Nicaragua.
from Isla} bito, Lake Managua, are difficult to place, and are O regarded as
lisotypes by Wallnófer
Lm |
intermediate with D. a. subsp. rivensis. The duplicates of C.F. Baker 157 are
(by annotation of the MO specimen in May 1999) because they “have the same label data!” However, in
addition to differences in collection number, the holotype C.F. Baker 629 (US) label reads “about dry hills”
while the C.F. Baker 157 collections (MO, ARIZ) do not. Moreover the labels are in different handwriting.
Specimens of C.F. Baker 157 (MO, ARIZ) bear Pomona College labels, while the holotype has the U.S. Na-
tional Museum label. There seems to be insufficient reason to consider C.F. Baker 157 collections isotypes.
EL SALVADOR. M in: P. in, Rio Sapo, 25 Jan 2000, J. Monterrosa et al. 15 (MO). San MIGUEL: San Luis
ies 9 ea 1975, J.R. Martinez 315 (TEFH). GUATEMALA. CHIQUIMULA: Caracol Mountain, 1.5 mi N of Quezaltepeque, 1200-1400
m, 7 Nov 1939, J.A. Steyermark 31396 (F). Zacapa: above Teculután, 250—275 m, 7 Jan 1942, J.A. Steyermark 42132 (F). HONDURAS.
CnorurEca: Tatumbla, 25 km de la capital, 1500 m, 28 Sep 1982, V.M. Pineda s.n. (MO). Comayagua: Chichipates, C. Nelson et al. 6746
(TEFH). Cortés: orilla del Río Humuya, 40 km N Santa Cruz de Yojoa, 100 m, 1-30 Nov 1980, C. Nelson et al. 5752 (TEFH, MO). EL
Paraíso: near Casitas, 900 m, 4 Dec 1946, L.O. Williams & A. Molina R. 11074 (MO). Francisco Morazán: slopes on Cerro Majicarán,
900 m, 14 Jun 1947, A. Molina R. 108 (MO). Intipuca: Quebrada Los Naranjos, Aldea San Juan, Magdalena, 10 Jan 1975, J.R. Martinez
347 (TEFH). La Paz: between Galeras and Lizapa Grande, 1000 m, 24 Jun 1947, A. Molina R. 167 (MO). NICARAGUA. Boaco: Sierra El
Espino, 500-600 m, 11 Nov 1982, P. Moreno 18526 (MO). Carazo: La Boquita, 13 Jul 1982, R. Kral 69373 (MO). CHINANDEGA: E del
Volcán Casita, 23 Jul 1982, M. Araquistain 2982 (MO). CHONTALES: 13.5 km W of Juigalpa, 100 m, 28 Aug 1985, M. ? 2 (MO); 14
km SSW of Comalapa, 150 m, 28 Aug 1983, M. Nee 27619 (MO). EsrELt: vicinity of Guava, 20 km of Esteli, 900 m, 5 Nov 1968, A. Molina
R. 23121 (MO). GRANADA: Granada, 16 Feb 1903, C. F. Baker 157. Jinotega: “El Eden,” 7 km a la entrada del camino viejo a Jinotega,
840—860 m, 25 Oct 1983, S. Vega & W. Robleto 99 (MO, TEFH). Leon: Santa Rosa del Peñón, 500-600 m, 11 Sep 1980, P. Moreno 2440
(MO). Mapriz: Cerro Quisica, 800-1100 m, 23 Nov 1979, W.D. Stevens 16181 (MO). Managua: El Tamrindo, + 300 m, 1 Dec 1983, P.
Moreno 22524 (MO); entrada a la hacienda San Ramón, 18 Oct 1982, P. Moreno 17919 (MO). MasaYa: Laguna de Apoyo, 100-140 m, 20
Sep 1981, P. Moreno 11140 (MO). Maracatra: El Caao—Caserio Puertas Viejas, 460 m, 25 Aug 1983, J.S. Miller 1286 (MO). Nueva SEGOVIA:
6 km N of Ocotal, 700 m, 15 Jun 1977, D. Neill 2145 (MO).
1g. Diospyros eran Kunth subsp. pedromorenoi M.C. Provance, I. García & A.C. Sanders, subsp.
nov. (Fi igs c—d). Tyee: NICARAGUA. Estert: San as de po Valle San Juan de Tranquera, entre San Juan de Limay
y Pueblo m 15:15) P RM 400—450 m, 5 Sep 1980, P 86 (HOLOTYPE: MO-3307430!)
Arbores et frutices a seu saad ssp. e id inp sed ud pus longioriuus ae a purpureus
-pri lMinosis ]
F
auc
(usque ad 3.5 cm diam).
Trees, rarely shrubs, 2-12 m tall; trunk unknown; stems glabrous to densely hairy when mature, the hairs
wavy, ascending, dull orange to golden-brown, becoming desiccated and ashy, those of young stems sometimes
mixed with sparse longer, wavy, spreading to ascending hairs, with clavate glandular hairs often present, but
deciduous and leaving the brown to dark brown epidermis viscid. Petioles 4-7 mm long, densely tomentulose
Provance et al., Diospyros salicifolia complex
1037
Pid 12. Di Á is Kunth sul | pedromorenoi M C. Provance, l. García & A.C. Sanders, subsp nov. A. F gi hl ith | j
l itos, Dept. of Estelí, Ni (P Moreno 10887 MO). B. Fruiting the holotype troimVälie San Juan de Tranquera,
ar ats Nicaragua (P. Moreno 2386, vo- SD eas lon P. Moreno 21526 (MO) from Hacienda San Ant Dept. of Boaco, Nicaragua
Malei D. Male calvx. E. Ad | F Di g fi imal gi f mal lla tube. G. Stamen
1038 J l of the Botanical R h Institute of Texas 2(2)
or with short golden-brown wavy hairs, but soon glabrous, rugose, glaucous-pruinose, to dark purple,
reddish-brown and shiny where rubbed, clavate glandular hairs sometimes present on the upper surface.
Lamina chartaceous, 60-122 mm long, 25-60 mm wide, length to width ratio (1.8-)2-3(ú3.3) : 1, broadly
obovate, oblanceolate or elliptic, base short-attenuate, margin often slightly revolute, apex subacute to obtuse
or rounded, sometimes slightly acuminate or tapering abruptly; lower lamina surface densely hairy, the
hairs short and curly to wavy, golden to reddish-brown, clavate glandular hairs often present, deciduous,
leaving the reddish-brown epidermis viscid, the stomatal apparatus translucent and inconspicuous, wpper
lamina surface glabrate to ascending wavy-hairy, deciduous, clavate glandular hairs often present, but
deciduous, the epidermis viscid, dark greenish-brown, papillate or sometimes crystal-papillose,. Venation
eucamptodromous, usually 3-5-ribbed; midrib prominent below, conspicuously wide, hairs denser on the
midrib than on the lamina, lower midrib glaucous-pruinose, shiny, reddish-brown; lateral veins 6—9 per
side, very prominent to oe below, slightly raised above; 3° veins prominent below; 4—5? veins
often darkened below. Male i 3—5(-6)-flowered cymes, densely buff to dull orange with wavy
hairs; peduncles 4—6 mm long; pedicels 1-3 mm long, bracts lanceolate, 1-1.6 mm long, conduplicate. Male
flowering calyx densely clear to light gold minute wavy-hairy outside, with clavate glandular hairs often
present, deciduous, epidermis reddish-brown, tube 2.0-3.5 mm long, + 3 mm wide, glabrous inside, lobes
lance-ovate to ovate, 3.0—5.5 mm long, 2-3 mm wide, barely winged or not, acute to acuminate, glabrous to
sparsely hairy inside with a narrow tomentulose intramarginal band, sometimes with sparse clavate glan-
dular hairs present; male corolla tube 3.2—5.5 mm long, + 2.5 mm wide, white sericeous outside with 3
small basal shield-shaped glal l lla lobes oblong-ovate, 2.5-3 mm long, + 1.8 mm wide.
Stamens 12; filaments date to the Sorelle in two 6-stamen tiers, the inner basal, the outer just above, +
1 mm long, dark reddish-brown. Female inflorescence and flowers unknown except style remnants on
immature fruit fused into a hairy column, 3 in number. Fruiting pedicels stout, 2-4 mm long, densely
hairy, with clavate glandular hairs sometimes present, epidermis reddish-brown, bracts (022, opposite,
often immediately subtending the fruit, broadly ovate, 1.8-1.9 mm long, flat. Fruiting calyx campanulate
to infundibuliform, densely minutely kinky to curly-hairy outside, the hairs, orangish to reddish-brown,
clavate glandular hairs sometimes present, but deciduous, leaving the reddish-brown epidermis viscid,
tube 5-7(-8) mm long, glabrous inside basally, lobes obtuse to acute, 4.5-6 mm long, (9212-15 mm wide,
abruptly acuminate to a blunt point, margins often reflexed, golden to reddish sericeous inside except for a
wide tomentulose intramarginal band. Fruit up to 3.5 cm in diameter, glabrous to scantily hairy, glaucous-
pruinose, dull yellow, mauve or purplish, often reddish-brown where rubbed, yellow to orange in life. Seeds
14-16 mm long, 6-7 mm radial depth, 5-6 mm wide.
This subspecies is notable for leaves with a broad, viscid, reddish-brown abaxial midrib, dense with
curly to kinked reddish-brown hairs, rugose mauve to purple glaucous-pruinose petioles, and large fruit
that are second in size only to D. aff. inconstans for the complex in Mesoamerica. It seems to be endemic to
Nicaragua (Fig. 24), with most collections from gallery forests and rocky slopes of west-central Nicaragua. It
forms intermediates with D. a. subsp. nicaraguensis in several areas. The fruit is known locally as ‘chocollo,
“chochoyo' or 'chocoyito' and is eaten when mature. This subspecies is named in honor of Pedro P. Moreno,
the collector of the type, and an expert on the flora of Nicaragua.
Specimens examined. NICARAGUA. Boaco: Hacienda San Antonio, 200 m, 14 Jun 1983, P. Moreno 21526 (MO); 1 km N of San José
de Los Remates, 650 m, 27 Jul 1983, W.D. Stevens & P. Moreno 22297 (MO); CHONTALES: ca. 13.1 km N of Hwy 7 on road to Cuapa, ca.
155 m, 15 on 1978, Pipoly 2029 (MSC); same location and date, W.D. Stevens 6493 (MO); Hacienda Veracruz, 120—475 m, 17 Jul 1983,
O), same location, 4-6 Aug 1983, W.D. Stevens 22368 (MO); 8 km S of Cuapa, 200—375 m, 21 Sep 1983, M. Nee 28260
(MO. EsreLí: Salto de Estanzuela, 1000 m, 10 Sep 1980, M. Guzmán et al. 1162 (MO); Llano Gualilica, 280 m, 2 Sep 1980, P. Moreno
2091 (MO); Paso León, 800—820 m, 29 Mar 1983, P. Moreno 210634 (MO); Los Cerritos, 860—900 m, 13 Sep 1981, P. Moreno 10887 (MO);
Portal de Belén, 660—800 m, 1 Aug 1983, P. Moreno 21845 (MO); Cerro Mae Río e 800-850 m, 3 Jan 1982, P. Moreno
14119 (MO); Res. Nat. Miraflor, 600—850 m, 23 D 99, R. Rueda et al. 125 San Juan de Limay, 250-300 m, 3 Sep
1980, P.M 2236 (MO); Llano el Pozo, 1200-1300 m, 26 Dec 1982, P. Moreno 19345 (MO); “Hacienda La Grecia, 518 m, 2 Sep 1980,
P. Moreno 2173 (MO) & 2134 (MO); Las Cuevas, 1000-1100 m, 1 Feb 1983, P. Moreno 19918 (MO); along road to Yalí, 820-1100 m, 14
Nov 1979, W. Stevens & A. Grijalva 15508 (MO). Leon: Hacienda Las Lajas, 300—350 m, 4 Sep 1980, P. Moreno 2295 (MO); Malpaísillo,
Provance et al., Diospyros salicifolia complex 1039
faldas del Cerro Negro, 100—200 m, 8 May 1984, M. Castro 122 (MO); La Chibola, 28 Jun 1984, M. Castro 143 (MO); Llano San Lorenzo,
250—300 m, 4 Sep 1980, P. Moreno 2327 (MO). Managua: camino entre "Puertas aes alma ” + 100-120 m, ap 1983, M
Araquistain 3584 (MO). MaracaLra: 3 km al E de Puertas Viejas, 400—500 m, 10 Feb 1 5 & 20090 (N
"Puertas Viejas,” 500—600 m, 21 Sep 1983, M. Araquistain 3668 (MO, BH); ca 5 km al NW de la ciudad de Matagalpa, 22 Jul 1983, A.
Grijalva & F. Ortiz 2825 (MO).
1h. Diospyros acapulcensis Kunth subsp. rivensis M.C. Provance, I. García & A.C. Sanders, subsp. nov. (Figs.
, 14, 15, 17a-i, 21d, 22a—b). Tre: NICARAGUA. Rivas: Isle Ometepe, Volcán Concepción, camino de "Los Hatillos” a
“Altagracia,” 11°34'N, 85%35-39%W 133-150 m, 13 Aug 1984, W Robleto 1088 (HoLoTYPE: MO-35083541; isorver: TEFHI).
A +) f. 1 g 1 +1: J Aff, 1 fA 1 1 a 11; T iL DEM tp
E E Is in F t O is
el jT : E E 1 hat E B ENTE acras : 1; 1h;
r E y r r - Ir
uri? f. Dele lad rere
O I
secus venas impariter projectis, y g
Trees or shrubs 2-10 m tall; trunk unknown; stems moderately to densely velutinous when young, the
hairs tawny to reddish-brown, slightly wavy or curved, subappressed, sometimes also minutely hirtellous,
glabrous or only sparingly hairy when mature, the remnant hairs shorter, curved, and more upright, clav-
ate glandular hairs often present, the epidermis sometimes very dark. Petioles sometimes winged, much
broader than thick, 4-8 mm long, epidermis relatively smooth, yellowish to brownish, shiny and viscid,
lower surface glabrate to velutinous, the hairs appressed to subappressed, straight to slightly wavy, upper
surface somewhat flattened to slightly canaliculate, puberulent, the hairs appressed to upright. Lamina
chartaceous to subcoriaceous when mature, 70—130(-150) mm long, 25-50 mm wide, length to width ratio
(2.1-)2.5-3.5(-3.8) : 1, widely obovate to narrowly oblanceolate or elliptic, often crystal-papillose, base cu-
neate to acutely short-attenuate, decurrent on the petiole, margin slightly thickened to revolute, sometimes
darkened, apex slightly acuminate to rounded; lower lamina surface velutinous to subsericeous, the hairs
usually straight, but also curved, appressed and subappressed, cream to tan, clavate glandular hairs often
present, the epidermis chartreuse to olive or brownish-green, sometimes papillose, the stomatal apparatus
obscure; upper lamina surface often tented, sparsely to moderately hairy, the hairs straight or curved,
appressed to ascending, sometimes deciduous, especially near the midrib, sometimes leaving a seeping
ring-like gland, epidermis light greenish-brown to brownish-green, shiny. Venation arcolanguid, some-
times 3-ribbed; midrib prominent below, subsericeous, the hairs curved to nearly straight, subappressed to
upright, thickish, tawny to golden-brown, clavate glandular hairs often present, but deciduous and leaving
the yellowish to reddish-brown epidermis viscidulous, midrib slightly impressed to raised above, impressed
along the lamina-midrib seam, pubescent, the hairs wavy to curved, + upright; lateral veins (526—9(-12)
per side, prominent below, raised above; 3? veins slightly raised to prominent below, barely raised above.
Male inflorescences 3—5(—6)-flowered cymes, densely pubescent, the hairs kinky, curved, or wavy, cream,
epidermis brown; peduncles 4-6 mm long; pedicels 2-3 mm long, bracts ovate to oblong-ovate, or deltoid.
Male flowering calyx moderately to densely hairy outside, the hairs variously straight, wavy, or curly, cream
to tawny, the epidermis orangish-brown to reddish, sometimes quite dark, tube 2.5-4 mm long, 3-3.2 mm
wide, glabrous inside, lobes acute, 1.8-4 mm long, 1.5-3 mm wide, glabrous inside, except for a narrow
tomentulose intramarginal band, and some scattered appressed wavy hairs on the median; male corolla
tube 4—5 mm long, 2-4 mm wide, cream, tawny, yellowish or reddish-pink sericeous outside except for
3 basal shield-shaped glabrous zones; male corolla lobes reportedly cream to silvery and pearlescent in
life, lanceolate to ovate or oblong, + 3.5 mm long, 1-2 mm wide. Stamens 12-14; filaments partly adnate
to corolla base, partly inserted on the receptacle, 1.2-1.5 mm long, rarely geminate, dark reddish-brown;
anthers lanceolate to ovate or oblong-ovate, 1.1-2 mm long, yellowish, nearly cylindrical prior to opening,
apex long-acuminate to blunt. Female inflorescences and flowers unknown. Fruiting pedicels 0.5—9
mm long, minutely hairy, often coated with exudate, bracts lanceolate to ovate, 1-2 mm long, capes
f
Fruiting calyx spreading to crateriform (in the widespread form) or panul vaguely
(in the pale-leaved form), usually splitting below the sinuses, minutely sericeous to tomentulose outside,
clavate glandular hairs sometimes present, the epidermis viscid, sometimes dark brown, sometimes black
gland-dotted, tube 4-7.5 mm long, upper 1/2—3/4 sericeous inside, the lower 1/2-1/4 glabrate, sometimes
1040
I lL af el
f Texas 2(2)
No 3508354
Fic 13 Holotype ntn ic KE
ensis M.C. Provance, |. García €: A.C. Sanders, subsp
Isle Ometepe Dept
of Rivas, Nicaraqua (W. Robleto 1088, MO- saa
Provance et al., Diospyros salicifolia complex 1041
[9
14. Pale-leaved form of D. is Kunth sul sis M.C. Provance, |. García & A.C. Sanders, subsp. nov. fi l de Apoyo, Dept. of
Masaya, Nicaragua (P. Moreno & J. C Sandino 6185, MO- ipd
with scattered, thick, cylindrical glands, lobes obtuse, (2.5-)3.5-7 mm long, 9-15 mm wide, spreading to
strongly reflexed, the apex often folded to an acute point, cream to tan sericeous inside except for a tomen-
tulose intramarginal band. Fruit 2-2.6(3) cm in diameter, glabrate, sometimes minutely hairy, tawny hairs
present at the apex, epidermis sometimes minutely red to black gland-dotted, usually glaucous-pruinose,
sometimes scintillant, dull yellow to brownish-orange or mauve, reportedly yellow or orange when mature,
and orange to red when ripe. Seeds 12-13 mm long, 5.5-7 mm radial depth, 4-5 mm wide.
This subspecies is found near and along the west coast of Nicaragua and northwest coast of Costa
Rica, as well as on the islands of Zapatera and Ometepe in Lake Nicaragua, Nicaragua (Fig. 24). It is mainly
found between sea level and 150 m in tropical deciduous forest near shorelines and along riverbanks, and
sometimes in marshy areas. There are also some mid-elevation occurrences around 700 m associated with
volcanic slopes. Specimens from the area between Lake Managua and Lake Nicaragua represent a form
with pale abaxial leaf surfaces (Figs. 14, 15, 17a—e, 21d) found in dry tropical forest and secondary matorral
between 100 and 350 m elevation. These pale-leaved plants also vary subtly in some characteristics of stem
and leaf vestiture from collections of the subspecies made elsewhere, and seem to have a different fruiting
1042 Journal of the Botanical R h Institute of Texas 2(2)
2888
42 Me
TRY : 292.
| gat n :
4 aM X ue SẸ A aie
\ oF ae
Re a, Que
Me
Fic. 15. Pale-leaved form of D. Icensis subsp. rivensis from Costado S de I le Villa Fontana, Dept. of Managua, Nicaragua (4. Grijalva 775,
MO).
Provance et al., Diospyros salicifolia complex 1043
calyx shape. However, the forms are indistinguishable following quantitative comparison of their vegetative
and fruiting inflorescence characters. Currently, we consider plants from the area between Lake Managua
and Lake Nicaragua to represent an interesting ecological variant.
Specimens that represent intergrades with D. a. subsp. nicaraguensis occur on the Island of Momotom-
bito in Lake Managua, with D. a. subsp. veraecrucis on the volcanic Isla de la Tigre off the southern coast of
Honduras, and with D. a. subsp. guanacastensis in Costa Rica. A combination of characters distinguishes this
subspecies from others, including a tented lamina, straight to slightly curved, appressed to subappressed
hairs on the abaxial lamina and midrib surfaces, and clusters of opaque white excrescences on the lower
lamina surface. This subspecies i resembles D. a. subsp. veraecrucis, but the lamina of D. a. subsp.
rivensis is more elliptic, and the leaf apex more rounded. In D. a. subsp. rivensis, hairs on the lower lamina
and midrib surfaces are not as straight and strongly appressed as in D. a. subsp. veraecrucis. Also petioles of
this taxon tend to be longer and thinner, and the fruiting calyx lobes narrower than in D. a. subsp. verae-
crucis.
Diospyros a. subsp. rivensis tends to have an elliptic leaf. This accounts for significant differences in
several lamina shape parameters between this taxon and D. a. subsp. nicaraguensis and D. a. subsp. guanacas-
tensis. Lamina width is also significantly narrower in D. a. subsp. rivensis than in D. a. subsp. nicaraguensis.
Venation is significantly different between D. a. subsp. rivensis and D. a. subsp. guanacastensis.
This taxon has been collected in a number of protected areas, such as Santa Rosa National Park on
the Santa Elena Peninsula in northwestern Costa Rica, and in Nicaragua at Reserva Natural Lagunetas de
Mecatepe, Parque Nacional Volcán Masaya, Zapatera Archipelago National Park, and in Cosigúina Volcano
Natural Reserve. The fruit is described as pale orange to red and sweet in late February to May. The local
name 'chocoyito' appears on collections of the pale-leaved form from the area between Lake Managua and
Lake Nicaragua, and on collections from the Department of Chinandega, and the islands of Zapatera and
Ometepe on Lake Nicaragua. The subspecific epithet refers to the Department of Rivas, where the holotype
was collected.
Specimens examined (except pale-leaved form). COSTA RICA. GUANACASTE: P.N. Santa Rosa, Península de Santa Elena, 50-150 m, 18
Aug 1994, B. iie id C. Cano 19581 (K); P.N. Santa Rosa, and de Santa Elena, 50 m, 15 ido 1993, B uin Cano 85 (MO); P.N.
Santa Rosa, Pení El 3 Dec 1993, R. E P.N. Santa Rosa, Peni 30 m, 30 Jul 1996, J.F.
Morales & A. Soto 5567 (MO). PUNTARENAS: Cantón de (bio Punta Agujas, 0—5 m, 19 Sep 1996, J.F. Morales 5832 (K). NICARAGUA.
CHINANDEGA: Res. Nat. Cosigüina, Los Placeres, 0—600 m [+ 37 m], 27 Mar 2001, R. Rueda et al. 15977 (MO). GRANADA: faldas del Volcán
Mombacho, md la Laguna de Juan Tallo, 65 m, 23 Jul 1980, M. Guzmán et al. 549 (MO); Isla Zapatera, Ensenada y Finca Sonzapote,
40—60 m, 20 Jan 1982, J.C. Sandino 1910 (MO); Isla Zapatera, sobre el camino a Sonzapote, 40—55 m, 20 Jan 1982, J.C. Sandino 1877
(MO); Isla Zapatera, entre Finca Sonzapote y Ladera NE de El Cerro, 100—300 m, J.C. Sandino 2056 (MO); Isla Zapatera, Cerro El Llano,
ca. 100 m, 13 Aug 1982, A. ipii 804 ae hw de d punta al N de Finca La Habana, 30—40 m, 24 Jan 1982, J.C. Sandino 2105
(MO); Isla Zapatera, costada Maria, + 100 m, 22 Nov 1982, A. Grijalva 1892 (MO, TEFH). Managua: El Paraguay,
km 26, carr. a San Rafael del Sur, 700 m, 9 Jun 1983, P. ^ 10); Finca Armenia, 600 m, 2 Oct 1982, A. & M.V. Grijalva 1330
on Volcán de ee 100—200 m, 2 Jun 1981, P. Moreno & J. Henrich 8947 (MO); S slope and rim of Apoyeque, 220—350 m, 18 Nov
, W.D. Stevens & P. Moreno 18448 (MO); "La Amistad,” a 7 km de La Concha, carr. a San Rafael del Sur, 550—600 m, 11 Dec 1980,
P. Moreno 5261 ao Masaya: P.N. Volcán Masaya, lava flow N of Volcán Masaya, 250 m, 29 Nov 1977, D. Neill 3036 (MO); P.N. Volcán
Masaya, NW side of El Comalito, 285 m, 8 Feb 1978, J.J. Pipoly 1732 (MSC); same location and date, W.D. Stevens 6256 (MO). Rivas: “El
Pegón," al SE de Rivas, 50 m, 21 Jan 1981, P. Moreno 6109 (MO); convergence of Río La Pita and Río Escalante, 20 m, 3 Aug 1978, W.D.
Stevens 9692 (MO); Isla Ometepe, Volcán Maderas, + T m, 16 Jun 1984, W. Robleto 895 (MO); Isla Ometepe, Volcán Maderas, sitio “Las
Cuchillas,” 400 m, 14 Jun 1984, W. Robleto 829 (MO); Isla de Ometepe, playa de Finca Santa Cruz y el Istmo de Istián, 17 Jul 19
Sandino 965 (MO); Isla de Ometepe, N shore of isthmus, near Santa Cruz, 25 Feb 1978, J.J. Pipoly 2110 (MSC); same date and un
W.D. Stevens 6623 (MO); Isla c Volcán Concepción, 50-80 m, 28 Oct 1984, W. Robleto 1444 (MO, TEFH); Isla Ometepe, 40-55
m, 14 Sep 1983, M
Specimens of pale-leaved form of D. acapulcensis subsp. rivensis examined: NICARAGUA. GRANADA: Laguna de Ápoyo, 110-180 m, P.
Moreno 11174 (MO). Manacua: Costado S de Lomas de Villa Fontana, + 250 m, 6 Aug 1982, A. Grijalva 775 (MO, TEFH); Las Sierritas,
W de Lomas Santiago, 160 m, 10 Jun 1982, J.C. Sandino 3048 (MO). Masaya: Camino entre restaurante Tenampa y Piedra Quemada,
7 Apr 1982, J.C. Sandino 2540 (MO); P.N. Volcán Masaya, W shore of Laguna de Masaya, 135-145 m, 31 Mar 1980, W.D. Stevens et al.
16871 (MO), x 11°59'N, 87°07'W, 135—145 m, 31 Mar 1980, W.D. Stevens, M. Guzmán & D. Castro 16871 pass Laguna de Masaya al NW
de la ciudad, 11°58'N, 86°07'W, 100—150 m, 24 Oct 1980, P. Moreno 4019 (MO); lado NW de la Laguna d ya, 12°00'N, 86°08’W,
1044 tani tit Texas 2(2)
140 m, 23 Feb 1981, P. Moreno & A. Lopez 7171 (MO); N de Volcán Santiago, Piedra Quemada, 12°01'N, 86°09'W, [200 m], 5 Aug 1981,
J.C. Sandino 1165 (MO); “Piedra Quemada” lava flow, 1 km NW of Volcán Santiago, [12*01'N, 86°10'W], 350 m, 28 Oct 1976, D. Neill
1111 (MO); a orillas de la lla Mcd 11°58'N, 86°08'W, 160 m, 21 Jan 1981, P. Moreno 6140 (MO); Laguna de Apoyo costado Este
[probably West, since the d phic coordinates, and elevation imply the crater near the west coast], 11°56'N, 86°04'W,
ca. 100, 24 Oct 1980, P. Moreno 3948 (MO).
e
li. ipsis acapulcensis Kunth subsp. veraecrucis (Standl.) M.C. Provance, I. García & A.C. Sanders,
et stat. nov. (Figs . 16, 18f—k, 19d). Basionym: Maba verae-crucis Standl. Contr. U.S. Natl. Herb. 18:119. 1916.
MUS verae-crucis (Standl.) Standl. Publ. Caen Inst. M 461: 80. 1935. Tyre: ats VERACRUZ: Catemaco, 300 m, 26
Apr 1894, E.W. Nelson 429 (HoLoTYPE: US-569278; isotype: US-569277)
Maba purpusii Brandegee, Univ. Calif. Publ. Bot. ee 329. 1920. MEXICO. Veracruz: Barranca de Panoaya, [ca. 19°10'N, 96°24'W],
Sep 1919, C.A. P t 51 UC-2049 MO, internet image!)
¡OLOTYPE:
NM D
Trees and shrubs 2-25 m tall; trunk up to 40 cm in diameter; stems glabrous or with some persisting hairs,
the young stems puberulent, the hairs sometimes curved, longer wavy subappressed hairs often present
towards the shoot apex. Petioles minutely winged, (2-)3—6(—6.5) mm long, yellowish-green, glabrate below,
puberulent above. Lamina membranaceous in spring, becoming chartaceous, (47-)60-105-120) mm long
(shade leaves to 160 mm long), (15220-40(—55) mm wide, length to width ratio (22.5—3.5(-4.4) : 1, oblan-
ceolate, obrhombic or obovate, sometimes nearly elliptic, base acute and short attenuate to cuneate, decurrent
on the petiole, margin flat, apex acutely rounded to acuminate and bluntly pointed, usually mucronulate;
lower lamina surface sparsely to moderately hairy, the hairs straight, appressed, distally oriented, white
to gold, clavate glandular hairs often present, epidermis light olive to brownish-green, sometimes minutely
papillose or crystal-papillose, the stomatal apparatus translucent and inconspicuous; upper lamina sur-
face sparsely hairy, clavate glandular hairs often present, epidermis darker than the lower lamina surface,
papillate or sometimes crystal-papillose. Venation arcolanguid to eucamptodromous, yellowish-green to
orangish; midrib prominent below, hairs straight and appressed, glabrate to hirsutulous or ascending to
upright puberulent; lateral veins 6—10(-12) pers side, ud Ta to A below, inconspicuous to
finely raised above; 3 veins usually obscure, M 1-3-flowered cymes,
densely pubescent with minute pe Dale, Rennes 1-3 mm DAE. pedicels 0.5-1 mm long, bracts
lanceolate, 1.0-2.5 mm long. Mal ly hairy outside, the hairs slightly wavy, ascending
to subappressed, clavate glandular hairs often nd giving the golden to dark brown epidermis a viscid
appearance, tube 2-3 mm long, glabrous inside except immediately below the sinuses, lobes lance-ovate to
ovate, (121.5—3.0(3.5) mm long, 2-2.5 mm wide; male corolla tube 5-7 mm long, 3 mm wide, mostly
sericeous outside, the basal 1-2 mm glabrous, or with 3 basal shield-shaped glabrous zones, each 1-2 mm
long; male corolla lobes lanceolate to ovate, 2-3.8 mm long. Stamens 12; filaments mostly adnate to the
corolla, 0.8—1.5 mm long; anthers lanceolate, 2.5-3.0 mm long, often acuminate. Female inflorescences
solitary, rarely a long-peduncled 3-flowered cyme with aborted lateral flowers. Female flowering calyx
densely and minutely subappressed curly to wavy-hairy, clavate glandular hairs often present, tube subcy-
lindrical, 3-5 mm long, lobes broadly rounded, (2.5-)3-5(-6.5) mm long, 4-7(-7.5) mm wide, often with
a blunt point; female corolla tube cylindrical, thickish, 6-7 mm long, 3-4 mm wide, the upper 2/3-3/4
sericeous outside, lower 1/3—1/4 glabrous, border between the regions x undulate; female corolla lobes
ovate to oblong-ovate, sometimes, 3.5—4 mm long, 2 mm wide, the margins somewhat involute; ovary flask-
shaped to obturbinate and vaguely 3-lobed, 2.5-3 mm long, 2.5-3 mm wide, hairy; styles narrow, 2-2.5
mm long, + fused, spreading distally; stigmas bifid; staminodes (1—)4—6, resembling fertile stamens, or
pale, broad, and inserted on the receptacle. Fruiting pedicels stout, (0.5—1—4.5(-9) mm long, glabrous
to moderately hairy. Fruiting calyx sparsely hairy outside, the hairs slightly wavy to straight, becoming
pale and desiccated, epidermis shiny and viscid, often nearly black, tube cupulate, (5-)6-9 mm long, often
tearing at the sinuses, usually bulbous at the base, densely hairy inside nearly to the base, the hairs short
and thickish to longer and fine, epidermis often with US blotches, lobes obtuse to rounded, Q. 5-23-TC-T.5)
mm long, (8—)10-14(-15) mm wide, often abruptl te to a blunt point, reflexed, just near
4
Provance et al., Diospyros salicifolia complex 1045
Cat. No. 03-016
INCHES
S
Herharium - sony rity of California Rive ride
pelas Kuh ssp. raent (Sandi 2 8L C
Sm
Pigs eee iC ais & AL Sa
Kc"
Dict fe Mich Prov ate
INS ui Gud DE ECOLOGIA, AY. QAM
+ FLORA DE VERACRUS »
BS Stand) Stand
EBENAURAR
far Viba a Coba Arras, ath
ken do T a Viera M ejus,
ae racrux, Móxs ama X Non uu 14" Wy
Forest way m
Dew fer: do him snr US Veris,
Kudonda Usus; Cerca
Cal dE EE N No 235
OR Pe ¡HL
veraecrucis. A. Fruiti hlet from La Vil Veracruz (J.E. González H. 235, IEB). B. Detail of fruiting I hlet
Fis. 16
from Cerro de la Mesa, Veracruz (R. Acosta P. & N. Acosta B. 199, IEB).
1046 | 1 afthn Bataniral D h Institute of Texas 2(2)
Fic. 17. Flowers of Diospyros mm. EE subsp. rivensis d red form), based on J.C Sandino 3048 (MO) from Las Sierritas, Dept. Managua,
T i yx. C. Pistillode. D. Mal lla lobe (adaxial view). E. Stamen FL, subsp. rivensis,
J.F. Morales 5832 (K) from Punta A , Puntarenas, Costa Rica. F. S | view [left], adaxial view [ ). G. Male corolla. H. Male
Ila laha ladavial view) IL. Adavial : a ' |
Provance et al., Diospyros salicifolia complex 1047
"S lan Ir Candinn 4442(MI £ CdaVolránM I , Dept of Leon Nicaragua
hi a Dlospyres acapileensis in A-E. subsp. ni (MU)
Male corolla tube C Male flower, D. A lla lobe (adaxial view). E. Stamen. F-G. subsp. veraecrucis, hacéd on
pienses from Fraylesca, i Mexico. F. Pistil. G. Female corolla. H—K. subsp. veraecrucis, based on B.F. Hansen & M. Nee 7482 from Emiliano
Zapata, Veracruz, Mexico. H. Pistil. I. Female flowering calyx. J. Female corolla. K. Staminode.
Pm : Lafého Bataniral D h Institute of Texas 2(2)
Fic. 19. Abaxial leaf surfaces of D. acapulcensis (scale = 1 mm). A. subsp. chiquimulensis, includi idril tary (A. Molina R. & A.R.
Molina 12473, F). B. subsp. chiquimulensis, margin detail (A. Molina R. & A.R. Molina 12473, F). C. subsp. acapulcensis, including midrib (1.1. Calzada 9223,
XAL). D. subsp. veraecrucis, including midrib (R. Acosta P. 741, IEB)
1049
Provance et al., Diospyros salicifolia complex
mi
W
a AT
LN a -
Der.
] ,
Th
-P MP
AS
-
w— 4
La
). C subsp. pedromorenoi, i
p. ,
PSI
including midrib
le = 1 mm). A. subsp. guanacastensis (from the holotype). B. subsp. guanacastensis,
LEE
10887, MO). D. subsp. pedromorenoi
L/D Ag
1050
oe. "S
SOS $ AN ^
a
Fi 21 Ahavial leaf curt: £n 1
1mm). A. subsp; m
T! subsp dwyeri,
ee including Rd in th
e holotype). B. subsp. mejocotensis,
rivensis (pale-leaved form), including
midrib, from P Moreno &l. C Sandino 6 185 (MO).
Provance et al., Diospyros salicifolia complex 1051
the apex or along the margins, sericeous inside, the intramarginal band of moderate width. Fruit globose,
slightly compressed or umbilicate, 272.5 cm in diameter, scantily hairy except at apically where densely hairy,
epidermis often glaucous-pruinose and scintillant, yellow to light brown, sometimes light reddish-brown,
probably golden brown when ripe. Seeds 15-16 mm long, 6-7 mm radial depth, 5 mm wide.
This subspecies occurs below 1240 m in tropical deciduous forests, semideciduous forests, and pine
forests of El Salvador, Guatemala, Veracruz, Oaxaca, and Chiapas, Mexico (Figs. 23, 24). It has straight, ap-
pressed, strictly distally oriented abaxial lamina hairs, versus the straight to wavy, ascending to upright, not
strictly distally oriented hairs of D. a. subsp. acapulcensis. Furthermore, compared to D. a. subsp. acapulcensis,
the leaves of D. a. subsp. veraecrucis also tend to be slightly longer and narrower relative to length, especially
at quarter length. The fruiting inflorescences of some specimens from El Salvador have peduncles 9-13
mm long with aborted lateral flowers that resemble long fruiting pedicels (e.g., Monro 3576, M.A. Renderos
385)
There is some suspicion that this taxon should be recognized at the species level, and we considered
accepting Diospyros veraecrucis Standl. circumscribed so as to include three additional subspecies, all with
rather straight, strongly appressed to ascending hairs on the abaxial leaf surface and/or abaxial midrib (e.g.,
subsp. chiquimulensis, subsp. rivensis, subsp. dwyeri). While we have not seen intermediates between D. a.
subsp. veraecrucis and D. a. subsp. acapulcensis from regions where their distributions closely approach one
another in Chiapas, Mexico, we are also not aware of these taxa occurring together. In the southern part of
its range D. a. subsp. veraecrucis seems to intergrade with D. a. subsp. rivensis on an island off the coast of
southern Honduras. Collections of the complex are lacking from the region between this island and popula-
tions of D. a. subsp. veraecrucis in El Salvador. In turn, D. a. subsp. rivensis forms intermediates with taxa
that have curly or wavy hairs, or hairs not strongly ascending (e.g., D. a. subsp. nicaraguensis). Consequently,
we are following a conservative course in recognizing this taxon only at the subspecific level.
Representative specimens. GUATEMALA. EsquINtLa: Finca Sabana Grande, 550 m, 15 Sep 1997, M. Nee et al. 47265 (BM); 4 km S of
Palin, 7 Aug 1970, W.E. Harmon & J.D. Dwyer 2945 (UMO). EL SALVADOR. AnvAcHAPÁN: San Francisco Menéndez, El Corozo, Mari-
posario, 380 m, 21 Jan 2000, J.M. Rosales 88 (MO, BM); P.N. El Imposible, 14 Apr 1999, R. Villacorta & W. Beredsohn 2884 (MO, BM);
vicinity of Ahuachapán, 800-1000 m, 9-27 Jan 1922, P.C. Pai 20328 (MO). Cañanas: Cinquera, zona protegida, 380 m, 15 Apr
2002, R.A. Carballo et al. 206 (MO). La LiseRTAD: Cant d , 400 m, 27 Nov 1999, A.K. Monro et al. 3097 (MO, BM); Area pro-
tegida o Buenavista, 320 m, 12 os 2000, R.A. Carballo et al. s.n. (MO); 1 de Q lt , 620 m, Dec 1987, M. Calderón
JBL-00582 (N S illa del Lago de Ilopango, 442 m, 11 Dec 1997, M.A Renders 385 (BM). SANTA ÁNA: San José Ingenio,
P.N. aa eneljocotón, 14°25'N, 89°21'W, 1120 m, 24 Sep 2001, E. Ramos 90 (MO); San José Ingenio, P.N. Montecristo, la vuelta
de los pinos, 14°25'N, 89°21'W, 600 m, V.M. Martinez 240 (MO). Cooperativa Las Lajas, 1240 m, 28 Nov 2000, A.K. Monro et al. 3576
(MO, BM). SONSONATE: Finca Las Victorias, 890 m, 25 Nov 2000, A.K. Monro et al. 3552 (BM). MEXICO. Curapas. Mpio. Acapetahua:
La Encrucijada, 0 m, 26 Jan 1981, V. Rico-Gray & I. Espejel 329 (XAL). Mpio. Arriaga: Río Las Arenas, 15 km NW of Arriaga, 130 m,
27 Aug 1974, D.E. Breedlove 36833 (MO). Mpio. Cintalapa: 3-5 km N of Cintalapa, 900 m, 22 Dec 1972, E.E. Breedlove & R.F. Thorne
30519 (MO). Mpio. Mapastepec: Guadalupe Victoria, 180 m, 22 Nov 1977, J.I. Calzada et al. 3986 (XAL). Mpio. Ocozocoautla de
Espinosa: near Derna, 800-1000 m, 16 Dec 1972, D.E. Breedlove & R.F. Thorne 30360 (MO); 15 km WNW of Ocozocoautla, 800 m,
D.E. Breedlove 28994 (MO). Mpio. Siltepec: Fraylesca, 8 Mar 1945, E. Matuda 5228 (F). Mpio. Tapachula: Santa Clara, 100 m, 9 Aug
1984, E. Ventura & E. López 133 (XAL, MO). Oaxaca. Mpio. Barrio de la Soledad: 2 km S de Alm oloya, 10 Dec 1980, R. Cedillo T. & D.
Lorence 495 (ILL). Mpio. Matías Romero: Campo Nuevo, 28 Apr 1981, R. Cedillo T. 714 (MO). Mpio. San ue Chimalapa: 3 km al
S de San Miguel Chimalapa, 22 Nov 1983, R. Torres C. 4132 (MO, CHAPA). Mpio. San Miguel del Puerto: 1.6 km al W de Xadani, 500
m, 27 Oct 2001, A. Saynes V. et al. 2642 (TEX); Finca El Istmo, 790 m, 1 Jan 2001, A. Saynes V. et al. 2342 (TEX). Mpio. Santa María
Chimalapa: ca. 6 km en línea recta al NE de Benito Juárez, 1000-1100 m, 3 Sep 1986, S. Maya J. 3799 (UCR). Mpio. Santo Domingo
Tehuantepec: "El Mangito,” 11.1 km SW de Buenos Aires, 800 m, 25 Nov 1983, R. Torres C. 4180 (MO). Veracruz. Mpio. Actopan: Cerro
de la Mesa (Sierra Manuel Diaz), 250 m, 30 Dec 1985, R. Acosta P. - N. a B. 199 (IEB); Sierra Manuel Diaz, 150 m, 5 Jul 1985, R.
Acosta P. & F. Vazquez B. 741 (XAL, XALU, IEB). Mpio. Alto I Barrios: Laguna de San Agustin, 10 m, 25 Feb 1983,
C. Gutierrez B. 1304 (IEB). Mpio. Apazapan: 2-6 km SE of Emiliano Zapata, 250—400 m, 27 Jun 1980, B.F. Hansen & M. Nee 7503 (NO,
USF, MU, WIS); same location and date, B.F. Hansen & M. Nee 7482 (FLAS, MO, USF, UMO). Mpio. Atoyac: 5 km S de Caballo Blanco,
350 m, 22 Aug 1985, R. Acevedo R. & F. Vazquez 440 (IEB). Mpio. Axocuapan: Ejido Coetzalan, alt 500 m, 6 Nov 1982, L. Robles H. 358
(IEB). Mpio. Catemaco: Isla Agaltepec, lado NW, 24 Apr 1974, F. Ponce C. & C. Alvarez C. 296 (CHAPA). Mpio. Comapa: Barranca de
Panoaya, 450 m, 12 Dec 1985, M.E. Medina A. & M. Ortiz D. 827 (IEB, XAL). Mpio. Covarrubia: cerca de Los Mangos, 17 Jan 1972, R.
Hernandez M. 1351 (CHAPA). Mpio. Dos Ríos: Plan del Río, 250 m, 25 Jul 1974, F. Ventura A. 10390 (IEB). Mpio. Emiliano Zapata:
1052 J | of the Botanical R h Institute of Texas 2(2)
-
E
j
j
=
p
-
al
type). B. 8. Hammel
E £V 2T
Fic. 22. Abaxial leaf vestiture, including midrib, f D,
&C Cano 19581 (K).
Corral Falso, 850 m, 30 Jun 1979, L. Pacheco & J.I. Calzada 9 (IEB); same location, 17 Jan 1973, M. Cházaro B. & J. Dorantes 19 (XAL).
Mpio. La Antigua: Rincón del Pirata, 30 m, 19 Nov 1982, P. M. Casasola et al. BD-1168 (MO). Mpio. Medellín: Paso del Toro, 18 Jan
1990, P. Zamora C. 1929 (IEB). Mpio. Naolinco: cerca de Almolongas, 950 m, 25 Apr 1992, M. Cházaro B. et al. 6908 (IEB, MO). Mpio.
Paso de Ovejas: 2 km SW de Cantarranas, 200 m, 30 Jan 1985, G. Castillo C. & M.E. Medina A. 3551 (XAL, XALU, IEB). Mpio. Puente
Nacional: Chichicastle: 50 m, 10 Dec 1976, F. Ventura A. 13729 (IEB). Mpio. San Andrés Tuxla: Laguna Encantada, 300 m, 2 Jul
1982, M. Nee et al. 24761 (BH, USF, WIS). Mpio. Tamiahua: E de Laguna Salada, 5 m, 26 Jun 1972, J. Dorantes et al. 1076 (MO). Mpio.
Tonalá: NW of Puerto Arista, 3m, 19 Oct 1971, D.F. Breedlove & R.F. Thorne 20869 (CHAPA, MO). Mpio. Xalapa: San Antonio, 800 m,
4 Feb 1975, F. Ventura Á. 10911 (UMO).
2. Diospyros aequoris Standl., Publ. Carnegie Inst. Wash. 461:80. 1935. (Fig. 25). Maba d Standl. Contr.
U.S. Natl. Herb. 18:118. 1916. Tre: MEXICO. Sinatoa. [Mpio. Culiacán]: “Collected in the vicinity of Guadalupe” [as on label],
“dry coastal thickets” [protologue], [24°15'N, 107°19'W, 3 m], 18 Apr 1910, J.N. Rose, PC. Standley & PG. Russell 14709 (HOLOTYPE:
US-637592; isorvrEs: MO, internet image!, C!). Not Diospyros latifolia Gúrke, Bot. Jahrb. Syst. 26:63. 1899
Trees, shrubs, or multi-stemmed clones from root and crown sprouts, 1-8(-12) m tall, facultatively de-
ciduous, dioecious; trunk recorded up to 30 cm in diameter, bark nearly smooth (but not slick), irregu-
larly roughened, or irregularly scaly, sometimes fissured or checked, especially at major branch forks, col-
oration sometimes patchy, usually grayish; wood hard, cream to dark yellow, dried branch wood light
yellow to golden brown, slash nearly white to light yellow, usually yellowish after several minutes, sap clear;
mature stems terete, smooth or fissured to half-netted, sometimes somewhat checked, glabrous to scurfy,
epidermis often shiny or viscid, variously colored, but mostly beige, reddish-brown, grayish, or castaneous,
1053
Provance et al., Diospyros salicifolia complex
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x: ssp. rivensis (pale form) [PANAMA pray mu b. ^^ X ]
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AÑ M m
+ = ssp. guanacastensis E A « Gub .
= SSP. Ve 5 TOM i E
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10° C = ssp. cuna DAS
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= intermediate rivensis / guanacastensis
*- ssp. dwyeri
-90° -88° -86°
Fic. 24. Distribut fD l C tA
black gland-dotted, usually lenticellate; 2nd year stems subterete to terete, sometimes angular, striate to
half-netted, often glabrous, or the persisting vestiture desiccated and ashy or covered with exudate, some-
times hispidulous with scattered longer hairs, epidermis coloration variable, minute round cream lenticels
commonly present; Ist year stems angular to terete, smooth or striate to sulcate, densely hairy, the hairs
variously straight to curly, minute to about 2 mm long, ascending to spreading, sometimes twisted, clear or
white to reddish (the hairs throughout the plant are mostly simple glandular hairs, translucent in life, and
the color of hairs in herbarium specimens is due to darkening of a fluid in the lumen), clavate glandular
hairs often present, deciduous, leaving the orangish to dark brown (green in life) epidermis viscid. Leaves
alternate, simple, entire; petioles subterete to terete, sometimes broader and flattened or caniculate above,
1.5-6(-7) mm long, margin usually not winged, but sometimes distally so, densely hairy, the hairs as vari-
able as those described for lst year stems except sometimes subappressed, epidermis smooth to rugose,
coloration highly variable, often shiny and viscid; lamina chartaceous to subcoriaceous, 18—80(-100) mm
long, 12-40(-54) mm wide, length to width ratio 1.1-2.8(-3.2) : 1, usually elliptic, oblong, or oval, base
mostly rounded to very widely rounded to the petiole (length to width ratio at 1/4 lamina length 1.5-4.3 :
1), clavate glandular hairs often present, but deciduous, the coloration of the epidermis in dry material vari-
able, often darker above than below, margin flat to revolute, sometimes undulate, apex usually rounded,
often widely (leaf length to leaf width at 0.75 length 1.3-3.5 : D; lower lamina surface moderately to
densely hairy, the hairs straight, wavy, or curly, minute to 2 mm long, subappressed to upright, white to
Provance et al., Diospyros salicifolia complex 1055
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Fic. 25. Isotype of D. aequor Standl. from the vicinitv of Guadal pe, Sinaloa (J.N. Rose et al. 14709, C). A Fruiting calyx. B. Leafy and fruiting! hi
C. Abaxial leaf surface D. Adaxial leaf surface.
1056 J | of the Botanical R h Institute of Texas 2(2)
dark reddish-brown, vestiture often a mixture of hair types, sometimes deciduous, sometimes persistent
and forming patches of desiccated or exudate covered hairs, epidermis dull to shiny or viscid, the stomatal
apparatus often opaque, off-white, and conspicuous; upper lamina surface usually wrinkled, usually
moderately to densely hairy, the hairs as variable as described for the lower lamina surface except usually
tending to be erect, often deciduous, the base of the hair sometimes thickened or raised, and forming a
seeping annular gland when the rest of the hair falls, the epidermis often shiny and viscid, sometimes
crystal-papillose (Fig. 26). Venation arcolanguid to brochidodromous, sometimes eucamptodromous,
sometimes obscured by the vestiture; midrib prominent below, usually densely hairy, but often becoming
glabrous, impressed to flush above, sometimes slightly raised near the base, sparsely to densely hairy, the
hairs straight to wavy, ascending to upright, clavate glandular hairs sometimes present, sometimes furfura-
ceous; lateral veins (3-)4-9 on each side of the midrib, the course usually tending to arc towards the apex
of the leaf with minor inflections, the vestiture and epidermis usually similar to that of the midrib, promi-
nent below, stout to fine, impressed to barely raised above, usually impressed slightly along the lamina-vein
seam, sometimes obscure; 3° veins apparent to prominent below, fine to stout, impressed to obscure above.
Laminar extrafloral nectaries abaxial, mostly round to oblong, usually 0.3- M ib mm long, sometimes
minutely rimmed, more common in the proximal 1/2 and close to the midrib. N i
2-3-flowered cymes, or solitary flowers in leaf axils of young stems (usually new growth), densely hairy, the
hairs as variable as those described for the lower lamina surface; peduncles 0.53 mm long; pedicels
0.8—5 mm long, bracts deltoid to linear-lanceolate, minute to 2 mm long. Male flowers 3-4-merous, +
pendant; male flowering calyx narrowly urceolate to narrowly campanulate, sometimes slightly winged,
exterior densely hairy, the hairs minute to short, straight to curly, ascending, off-white to golden or tawny,
clavate glandular hairs sometimes present, deciduous, leaving the epidermis viscid, light green in life, tube
2-4 mm long, 3-4 mm wide, interior usually glabrous, sometimes with scattered to sparse hairs, lobes
valvate-reduplicate in the bud, acute, 3—6(-7) mm long, 2.5-5(25.5) mm wide, sometimes with a central
keel, exterior densely hairy, interior intramarginal band of hairs variable in width, comprised mostly of
minute, curly, off-white to tawny, golden, or reddish hairs, the median glabrous to sparsely hairy, the hairs
minute to short, straight to slightly wavy, appressed; male lla white to cream in life; male corolla tube
salverform to narrowly urceolate or cylindrical, sometimes narrowing distally, 4.5-6.5 mm long, 3-4 mm
wide, with three shield-shaped, 0.3-2 mm long, glabrous zones at the base, or sometimes the basal part of
the tube completely glabrous, otherwise sericeous with an undercoat of minute curly to straight hairs,
minute hairs sometimes present between the sericeous and glabrous zones; male corolla lobes spreading
in life, lanceolate to ovate, elliptic, oblong, or rhombic, 2-4 mm long, 0.8-2.5 mm wide, densely hairy, the
hairs minute, curly to wavy (and then upright), or straight and appressed, especially along the left side,
sometimes straight and nearly retrorse or going in random directions, especially along the right margin, the
median sometimes raised outside, sericeous, at least basally, margins sometimes involute, especially near
the base. Stamens 9—12; filaments adnate to the tube in 1-2 well-defined tiers or somewhat irregularly,
often with some (rarely all) inserted on the receptacle, sometimes geminate, rarely filaments adjoined later-
ally; anthers mostly lanceolate, sometimes oblong, 2-2.7 mm long, sometimes ‘eared’ (with a minute lobe
at the base of the anther on each side of the connective), opening by longitudinal slits. Pistillode usually
minute, densely hairy, the hairs straight to wavy, lustrous. Female inflorescences solitary in leaf axils of
young (often Ist year) stems, densely hairy, the hairs and epidermis similar to those seen for male inflores-
cences; female flowering pedicels with hairs and epidermis similar to male inflorescences. Female flow-
ers 3-4-merous; female flowering calyx mitriform, campanulate to broadly campanulate, exterior
densely hairy, the vestiture a variable mixture of short hairs and long hair types, the longer hairs sparse,
tube usually cupulate, sometimes half-globose, (1.5-)2.3-6 mm long, 3-7 mm wide, lobes valvate-reduplicate
in the bud, winged, usually broadly rounded to subacute, sometimes acute, 3-8 mm long, 4-8 mm wide,
abruptly acuminate or bluntly acute, major nerves sometimes conspicuous; female corolla white to cream
in life: female corolla tube urceolate or short urceolate to subcylindrical, usually narrowing distally, 4-7.5
Provance et al., Diospyros salicifolia complex 1057
mm long, (2.5-33-4 mm wide, exterior vestiture similar to that described for male corolla tubes, interior
glabrous; female corolla lobes spreading in life, ovate to rhombic, sometimes oblong, (2.5)3-4 mm long,
1.8-2.5 mm wide, often glabrous near the exterior margins, the median often broadly raised outside, often
partly involute, exterior j similar to that of male flowers, interior glabrous, sometimes with a longi-
tudinal depression through the center; ovary obturbinate, ovoid, globose, + depressed-globose or somewhat
conical, usually about 2 mm in diameter, glabrous to densely hairy, the hairs straight, white, blond, cream,
tan tawny, or reddish, sometimes dimorphic; styles 3, spreading or fused into a column, 1.8-2.5 mm long,
usually with minute appressed hairs on the exterior, interior sometimes with some minute wavy hairs;
stigmas vaguely bifid to conspicuously v-shaped, rarely trifid; staminodes 0—6, adnate to the base of the
corolla tube or inserted on the receptacle, narrowly lanceolate, rarely ovate, 2-4.5 mm long. Fruiting pedi-
cels terete to angular, sometimes stout, often broader distally, 2-8(-12) mm long, subglabrous to densely
hairy, the hairs usually wavy to curly, often covered with exudates, clavate glandular hairs sometimes pres-
ent, deciduous, leaving the light orange to dark reddish-brown epidermis viscid, bracts 1-2, opposite to
subopposite, linear to ovate, 1.5-4 mm long, concave to conduplicate. Fruiting calyx slightly accrescent,
cupulate to explanate, sometimes tearing at the sinus along a zone of dehiscence, exterior vestiture as de-
scribed for flowering calyces or glabrate, often with pale desiccated remnants of the original vestiture,
epidermis often conspicuously viscid and darkened, tube (323.5—6.5(-8.5) mm long, interior glabrous to
hairy in the upper 1/2-2/3, the hairs usually straight and appressed, lobes obtuse to rounded, overall simi-
lar to the calyx lobes of female flowers except wider, rarely acute, 3-8(-9) mm long, (6-)6.5-15 mm wide,
reflexed (basally, near the apex, or along the margins), interior intramarginal band of hairs variable in width,
the hairs similar to those described for the intramargin of the male calyx, the median usually hairy, the
hairs usually straight and appressed. Fruit a globose to subglobose or obovoid berry, 1.5-2.5 cm in diam-
eter, consistently with three pairs of locules, glabrous to sparsely hairy, sometimes moderately hairy at the
base, often densely hairy at the apex, the hairs slightly wavy to e a basal ring of subappressed, fine,
straight hai (the ring may adhere to the base of the fruiti lyx tube); flesh gelatinous,
[e] o P
drying vitreous, seddishi and translucent; epidermis smooth to orange-peeled or wrinkled, sometimes
separating from the fruit wall upon drying, sometimes glaucous, pruinose or scintillant, mature fruit light
grayish-yellow to golden brown, green in life, golden brown when ripe. Seeds shaped like an orange seg-
ment, texture rugulose-foveolate, 11-17 mm long, 6-9 mm radial depth, 5-7 mm wide.
KEY TO THE SUBSPECIES OF DIOSPYROS AEQUORIS IN MESOAMERICA
. Lamina orbicular to oblong or oue. length : width « 1.6 : 1, base subcordate to rounded, length : width
at 025 « 22:1, length : width a « 17:1, margin flat, hairs below fine, soft, glossy, white, sometimes
gold; stems atropurpureous; mS s tube 2 5 mm long; mostly trees; Apatzingan Valley, Michoacan,
Mexico subsp. martineziana
1. Lamina shape variable, but length : n usually > 1.6: 1, base not subcordate, length : width at 0.25 > 2.2:
1, length : width at 0.75 » 22 : 1, margin often revolute, hairs below not as above; stems light gray to atro-
purpureous; fruiting calyx tube < ba mm long; trees or shrubs; Sinaloa to Oaxaca, W Mexico.
2. Fruit epidermis weakly attached to the fruit wall, often separated in dry rice dull light yellow;
lamina rhombic, oval, elliptic or onions, the hairs oe aten to sidad ter ashy; uae calyx small
(calyx length + lobe width 15-20 mm), the lo! pex E regions
of Mexico end e rero subsp. balsensis
2. Fruit ached to the fruit wall, usually not separated in dry specimens, not dull light
al
yellow; lamina rarely rhombic, hairs below variously colored; E calyx small to large (calyx length +
lobe width 16-28 mm), the lobes reflexed or not; W Mexico
3. Lamina chartaceous, not coarsely wrinkled, the margins
leaving a seeping glandular area, 29 venation os 3? venation slightly raised below
jul not revolute, h t d
inconspicuous above; fruiting pedicels 2.5-4 mm long; coastal W Guerrero, Mexico ubsp. chutlensis
3. Lamina subcoriaceous ana usually chanel ee when mature, ih margins often pu not
markedly undulate, h , usually not leaving conspicuous seeping areas,
2° venation brochidodromous, 3° venation prominent be low, impressed above; fruiting pedicels
2-8(-12) mm long; fruiting calyx small to large; Oaxaca, Jalisco, Nayarit, Sinaloa, Mexico.
1058 J lof the Botanical R h Institute of Texas 2(2)
4. Fruiting pedicels mostly 5.5712 mm long; fruiting calyx large, the lobes 11-14 mm wide; lamina
usually » 65 mm long, petioles Mn > 4n mm Jano. known only from coastal Oaxaca
subsp. rekoi
4. Fruiting pedicels mostly 2-5.5 mm long; fruiting calyx small to large, the lobes 6-15 mm wide;
lamina usually « 65 mm long, petioles often « 4 mm long; interior Oaxaca, Sinaloa, Nayarit and
coastal Jalisco, Mexico
5. Lamina bright Sen to Dege or reddish- brow
m above and ol Ie Below cat
the midrib, Ii | sli ightly VV o straight,
subappressed hairs, venation ll very nea pear: to pr brick ae or reddish-
brown; plants of Sinaloa, Nayarit and coastal Jalisco, Mexico subsp. aequoris
5. Lamina greenish-gray to gray above and below, vestiture below, including midrib, of wavy to
straight hairs, venation below prominent, cream to magenta or speckled with both colors;
known only from interior Oaxaca subsp. tehuantepecensis
2a. Diospyros aequoris Standl. subsp. aequoris (Figs. 25, 26, 27, 28, 38b).
Trees or shrubs, 1-4(-9) m tall; trunk grayish-white or gray to reddish- t glabrous, furfuraceous
or with remnant patches of desiccated hairs, young stems with short, dense, seddish curly hairs, and sparse
long, straight, spreading to ascending hairs. Petioles cylindrical to semi-terete, 1.5-4 mm long, densely
hairy, the hairs curly, sometimes nearly straight, ascending, reddish-brown, epidermis dark brown, viscid.
Lamina subcoriaceous, rarely chartaceous, (20-)26-66(-77) mm long, 15-28-34) mm wide, length to
width ratio 1.7-2.7(3.2) : 1, elliptic to obovate, rarely oval, gray to reddish-brown, darker above than below,
base obtusely rounded, sometimes short-attenuate, margin revolute, apex obtusely to acutely rounded; lower
lamina surface with dense reddish-brown tomentum, with some longer slightly wavy to straight, ascend-
ing to subappressed hairs, the vestiture deciduous, or becoming ashy and covered with exudate, clavate
glandular hairs often present, the epidermis olive-green to brownish, the stomatal apparatus translucent to
opaque, usually e upper lamina surface wrinkled, subglabrous to curly-villosulous, the hairs
deciduous, clavate glandular hairs often present. Venation brochidodromous, very prominent below, mark-
edly impressed above; midrib subsericeous below, with sparse shorter curly hairs, the epidermis viscid,
reddish, densely hairy above, the hairs wavy, ascending, furfuraceous, viscid; lateral veins 5-7(-8) per
side, prominent below, subsericeous; 3° veins prominent below, wavy-hairy. Male inflorescences solitary
flowers, with dense minute cream to golden wavy hairs; pedicel 4-6 mm long, bracts minute, + deltoid.
Male flowering calyx urceolate to subcylindrical, with dense, ascending, cream to golden pubescence,
tube 2-4.2 mm long, 3-4 mm wide, several straight, 1.5-2 mm long hairs inside near the base, shorter hairs
scattered distally, lobes uno triangular, 3-5 mm long, 2.5-3 mm wide, with scant short vaguely wavy
appressed hairs inside, t in very narrow; mal lla tube 5.5 mm long, 3 mm wide,
exterior cream to yellowish or reddish sericeous, with 3 shield-shaped glabrous basal regions, each 1.5 mm
long; male corolla lobes lanceolate to rhombic, 2.5-4.5 mm long, 0.8-2.5 mm wide. Stamens 9-12; fila-
ments adnate to the corolla in two distinct tiers, or all inserted on the receptacle, 1-1.5 mm long; anthers
oblong-lanceolate, 2 mm long, sometimes ‘eared’ basally, vaguely granulate apically. Female inflorescences
densely and minutely hairy. Female flowering calyx campanulate, sometimes broadly, densely reddish
tomentulose outside, with some longer spreading hairs, epidermis reddish-brown, viscid, tube cupulate,
3-3.5 mm long, 4 mm wide, the upper 1/2 densely hairy inside, lower 1/2 glabrate, lobes obtuse, 6-8 mm
long, 5-6 (7) mm wide, bluntly pointed to acutely rounded; female corolla tube 6 mm long, 2.5-3.3 mm
wide, upper half white sericeous outside, lower half glabrous, drying dark red; female corolla lobes ovate
to oblong, 3.5 mm long, 2.5 mm wide; ovary obturbinate to ovoid, minutely white to tan sericeous; styles
3, fused most of their length; stigmas 3, conspicuously v-shaped; staminodes 0-3, inserted on the recep-
tacle, slender, 4.5 mm long, dark red. Fruiting pedicels 2-5(-8) mm long, densely tomentulose, clavate
glandular hairs often present, epidermis reddish-brown or darker, bracts linear, 1.5 mm long, conduplicate.
Fruiting calyx rotate to explanate, densely golden to reddish-brown tomentulose outside, with occasional
longer straight hairs, glabrate, or the hairs desiccating, becoming ashy, and sticking to the viscid epidermis,
Provance et al., Diospyros salicifolia complex 1059
4 LI
Nu Ure P e i Ei
$ A "y d. » ^ a
e WEL di
A
Y iB
m
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r
Fic, 26 Detail af adayial | A D]
UCR).
tube (346.59) mm long, the upper 1/2-2/3 hairy inside, lower 1/2-1/3 glabrous, lobes acute to slightly
obtuse, (4-)5—7.5-9) mm long, (68-13-15) mm wide, spreading, or sometimes reflexed near the apex,
along the margins, or both, nerves often conspicuous, hairy inside, the interior intramarginal band wide.
Fruit globose, 1.5-2.5 cm in diameter, densely ascending wavy-hairy basally and apically, glaucous-pruinose
and scintillant, yellow to dark golden brown, sometimes black gland-dotted. Seeds 12-13 mm long, 6-7
mm radial depth, 5 mm wide.
This subspecies occurs in semi-arid oak forests on rocky soils in the Barranca Region of the west Dur-
ango-Sinaloa border region, on steep, rocky volcanic slopes with dry tropical forest in the foothills around
Culiacan, on the coastal plains of Sinaloa in thornscrub and dry tropical forest or thornscrub transitioning
to mangrove forest, and on inland mesas with Krameria in the northern extremity of Nayarit. Several, ap-
parently disjunct, populations occur on and near the coast of Jalisco in the vicinity of the Bahia de Chamela
(Fig. 40). Only two specimens with male flowers have been seen. In both cases the inflorescences consisted
of solitary flowers in leaf axils of new growth, representing exceptions to the 3-flowered cymes typically
seen in the complex. This subspecies is similar to D. ae. subsp. tehuantepecensis from inland Oaxaca, being
separated by differences (see key) in leaf surface, abaxial leaf vestiture, and lamina color (at least when dry).
Furthermore, ratio of leaf length to avera ge distance between centermost lateral veins is significantly higher
in D. ae. nd aequoris than D. ae. subsp. tehuantepecensis.
R MÉXICO. Durango. Mpio. Tamazula de Victoria: Sierra Tres Picos, [25°3'N, 10791], 3000 ft,19 Dec 1939,
H: s. Gentry 5286 (NA, MO). JaLisco. Mpio. La Huerta: Rancho Cuixmala, 200—300 m, 5 Nov 1991, E J. Lott et al. 4159 (UCR). NAYARIT.
Mpio. Huajicori: Zonteco, 110 m, 30 Aug 2004, M.C. Provance et al. 9606, 9607 & 9608 (UCR). StNALoA. Mpio. ld boi ene-
gita, SE of Badiraguato, 2000 ft, 29 Mar 1940, H.S. Gentry 5946 (NA, MO, ARIZ). Mpio. Cosalá Vado Hondo: a 1
Cosalá, 590 m, 9 Nov 1985, A. Garcia V. 172 (CHAPA). Mpio. Culiacán: + 4 km al sur de Culiacán, 50 m, 1 Nov 1990, R. Vega À et al.
1060 j loftheB ical R h Institute of Texas 2(2)
Provance et al., Diospyros salicifolia complex 1061
a
a » e
an i Piei
ean Mar
nuni A ~ : My A
ci n — E E - " is. La bai
7 » - ps
T 28. ig eid bois subsp: aequoris, M male flower, pps on Provance aa 233, WR: C-G. based on £. Guizar N. 3233 AA. E Fruiting
mature fruit. G H. Mature fruit, b
5946, ASU.
3972 (CIIDIR). Mpio. El Rosario: Playa de Chametla, + 9 km S de Chametla, 10 m, 5 Nov 2004, J.M. Aguilar P. 1608 (UCR); El Carrizo,
+ 9 km al N de Cacalotán, 50 m, 6 Nov 2004, J.M. Aguilar P. 1612 (UCR). Mpio. Mazatlan: Isla de la Piedra, 23 Jul 1994, E. Guízar N.
3233 (CHAPA, IEB). Mpio. Mocorito: El Alamo, along road from Mocorito to Surutato, 1500 ft, 3 Mar 1971, D.E. Breedlove 19111 (MO).
Mpio. San Ignacio: ca. 2 km SSW of Vigas (Loma) and ca. 10 km WNW of Duranguito, 14 m, 26 Aug 2004, M.C. Provance & J.S. Ross-
Ibarra 9525 (UCR).
2b. Di
subsp. balsensis M.C. Provance, 1. García & A.C. Sanders, subsp. nov. (Figs.
29, 30, 37j-m, 38c). Ter: MEXICO. Michoacan. Mpio. Tuzantla: km 68 carr. B. Juarez-Tuzantla, [19°14'N, 100°30' 30"],
790 m, bosque tropical caducifolio, 16 Dec 2004. I. Garcia R., M.C. Provance 6 J.A. Machuca 6863 (HOLOTYPE: CIMII; isotype: UCR!,
and to be distributed).
Frutices vel paa a ae oe ssp. aequori similis sed differt laminis foli pl jue latioribus, praesertim ad 1/4 et 3/4
longitudinem, 1 i brevioribus, et fructib e p
Shrubs, sometimes arborescent, 1.5-3(-5) m tall; trunk smooth to somewhat roughened, bark forming
large patches, varying from light to dark gray; stems glabrous when mature, but the young stems moderately
1062 Journal of the Botanical R h Institute of Texas 2(2)
Fic. 29. The holotype of D. aeauoris Standl. subsp. balsensis M.C. Provance, |. García & A.C. Sanders, subsp. fi km 68 Benito Juarez Tuzantla
ma LIN | W J A [d 1 fal TEN | t.
(I. Garcia R. et al. 6863, CIMI). A. F ). D. Fruit detail
Provance et al., Diospyros salicifolia complex 1063
Fic. 30. Di is «ul bal jc f, km 68 Benito luarez—Tuzantla. A Sida vi fth n Tab id fruit B. A fruit with cinky
p)
A A
J L : r ‘k badh t J : P" TON I £.
1064 lof the B ical R h Institute of Texas 2(2)
ascending hairy, the hairs wavy, sometimes twisting, clavate glandular hairs also present. Petioles 2—3(—5)
mm long, cream to yellowish tomentulose, with sparse longer slightly wavy hairs, older leaves sometimes
glabrate, the epidermis viscid, densely hairy above, the hairs straight to wavy, ascending to upright, cream
to yellowish. Lamina subcoriaceous when mature, (30-)36-60(-72) mm long, 23-36(40) mm wide, the
length to width ratio 1.5-2(-2.6) : 1, rhombic, elliptic or oval, base rounded, sometimes attenuate, margin
slightly revolute, apex ay to aora y rounded; lower lamina surface densely cream to yellow villous-
velutinous, the ] d near the apex, becoming desiccated, pale and ashy, or covered
with droplets of exudate, clavate glandular hairs often present, the stomatal apparatus opaque, white, and
conspicuous; upper lamina surface wrinkled, the vestiture similar to but less dense than on the lower
lamina surface, epidermis shiny, pale light green or gray to dark gray in herbarium material, sometimes
crystal-papillose. Venation brochidodromous, the epidermis yellowish, prominent below, villous-velutinous,
sparser above; midrib flush to slightly impressed above, impressed along the midrib-lamina seam; lateral
veins (5-)7-9 per side, impressed above, or flush and apressa along! a ied midrib seam; 3? veins
prominent below, impressed above; 4^—5? veins slightly imp above 23-flowered
cymes; peduncles + 2 mm long; pedicels 0.8-2 mm long, densely wavy-hairy, bracts linear-lanceolate, + 1-2
mm long, conduplicate. Male flowering calyx narrowly campanulate to urceolate-campanulate, thinnish,
cream, tawny or golden tomentulose outside, clavate glandular hairs often present, the epidermis golden
brown, viscid, Ad 2.8.2 mm long, Biabsous inside, ii pud triangular, 3-3.3 mm long, 2.8-3.2 mm
wide, glal ide, except for danarrow intramarginal band; male corolla
tube 5, 5-6.5 mm lone 3 mm wide, golden sericeous outside, with 3 shield-shaped glabrous basal regions,
x 2 mm long, bordered distally by minute wavy hairs; male corolla lobes obovate to rhombic, 2.0-2.5 mm
long, 1.5 mm wide, margins sometimes involute. Stamens 12; filaments in two subgeminate tiers adnate
to the corolla near the base, up to 2 mm long, reddish- brown; anthers lanceolate to narrowly lanceolate,
up to 2.7 mm long. Female inflorescences densely pinkish-orange tomentulose. Female flowering calyx
campanulate, narrowly winged, tube cupulate, 2-4 mm long, 3 mm wide, the basal 1/2-1/5 glabrous inside,
upper 1/2-2/3 sericeous, the intramarginal band wide, pinkish-orange short wavy-hairy outside, with sparse
straight ascending hairs, lobes thickish, acute, 5-5.5 mm long, 4-4.5 mm wide, vaguely acuminate to a blunt
point; female corolla tube 6 mm long, 3.5 mm wide, pinkish-orange sericeous outside, with 3 small glabrous
shield-shaped basal regions; female corolla lobes ovate, 2.55.5 mm long, 1.8-1.9 mm wide; ovary ovoid,
2.5 mm long, 2 mm wide, hi lous to stiffly ascending hairy, the hairs light blond; styles 3, fused basally,
spreading; stigmas bifid to trifid, long, spreading; staminodes 5, narrowly lanceolate, up to 4.5 mm long,
adnate to the base of the corolla, rarely inserted on the receptacle. Fruiting pedicels stout, 2-4(-6) mm
long, sparsely to densely wavy-hairy, bracts 1-2, linear-lanceolate to ovate, 3-4 mm long, + conduplicate.
Fruiting calyx shallowly rotate to explanate, tube 4-6 mm long, hairy, lobes widely rounded, (3-)4-5.5
mm long, (829-11 mm wide, apex sometimes cuspidate, reflexed. Fruit globose to obovoid, 1.5-2.5 cm
in diameter, glabrous to sparsely ascending puberulent, epidermis tending to separate from the remainder
of the fruit upon drying, dull yellow to golden brown, in living material greenish-yellow to yellow when
mature, golden brown when fully ripe. Seeds 16-17 mm long, 9 mm radial depth, 5-7 mm wide.
This subspecies grows in dry tropical forest on slopes and mesas between 600 and 800 m in the Sierra
Madre Occidental near the west end of the Sierra Madre del Sur in eastern Michoacan, western Guerrero,
and the southwest part of the State of Mexico (Fig. 39). In this subspecies the epidermis of the fruit is weakly
attached, a character useful in separating it from similar taxa. It is distinguished from D. ae. subsp. marti-
neziana by thicker, wrinkled, frequently revolute lamina with a different shape. Lamina shape ratios (i.e.,
length : width, length : width at 1/4 length, length : width at 3/4 length) being significantly higher in D. ae.
subsp. balsensis. In addition to differences in abaxial vestiture, the lamina of D. ae. subsp. chutlensis is thin-
ner, less wrinkled, and has an undulate margin. The venation in D. ae. subsp. balsensis is more prominent
than in D. ae. subsp. chutlensis. This subspecies is similar to D. ae. subsp. aequoris, but has shorter fruiting
calyx lobes, and the lamina is usually wider.
Provance et al., Diospyros salicifolia complex 1065
Specimens examined: MEXICO. Guerrero. Mpio. Ajuchitlán del Progreso: Cuahuilote, 29 Jan 1934, G.B. Hinton et al. 5568 (DES, K).
Mpio. Coyuca de Catalán: Paso de Arena, 5 Jun 1934, G.B. Hinton et al. 6117 (NA, K). Mpio. Zirándaro de los Chávez: A 14 km al
SW del Tálamo, 710 m, 6 Sep 1982, J.C. Soto N. & G. Silva R. 4364 (XAL). Mexico. Mpio. Temascaltepec: Bejucos, 610 m, 22 May 1933,
G.B. Hinton 3959 (K); same location, 28 Nov 1934, G.B. Hinton 7058 (K, ARIZ, NA, US, MO). MICHOACAN. Mpio. Arteaga: Puente San
Salvador, 630 m, 14 Mar 2005, P 9919 (UCR, CIMD. Mpio. Huetamo: Arroyo Seco, I. Garcia R. et al. 6873 (CIMI, m Eréndira,
800 m, 5 Nov 1977, J. Rzedowski 35495 (AAU, IEB). Mpio. Tuzantla: ca. km 68 carr. Zitácuaro-Tuzantla, 789 m, 16 Dec 2004, I. Garcia
R. et al. 6867 (CIMI, UCR); 9 km al NE de Tuzantla, 750 m, 20 Jun 1983, J.C. Soto N. 5325 (MO); 10 km al NE de Tuzantla, 700 m, 12
Dec 1971, F. Chiang et al. 547 (WIS, NA, SD, XAL)
2c. Diospyros aequoris Humb. & Bonpl. ex Willd. subsp. chutlensis M.C. Provance, I. García & A.C. San-
ders, subsp. nov. (Figs. 31, 32, 33, 39a, c). Tee. MÉXICO. Guerrero. Mpio. La Unión: Paraje “El Palomar," trayecto
de terracería a Chutla, 10 m, 4 Dec 1993, E. Guízar N. & L. Pimentel B. 3067 (HoLorvee: IEB!; isorver: CHAPA).
A vh E : 1e; TW +14 1 AFA Faliic? lu Ail i
? , Vena-
us 3 oa -1 3 ES
tione abaxiali subtiliore, et
Trees and multi-stemmed shrubs, 1.5—7.6 m tall; trunk somewhat smooth to fissured and scaly, the scales
very thick, irregular, whitish-gray to grayish-beige; branches smooth to scaly or irregularly roughened;
stems glabrous, or with some persisting ind hairs in the second year, grayish-beige in life, the young stems
densely short wavy-hairy, with di ight hairs up to 2 mm long near the shoot apex,
the epidermis viscid, light dull E nee to reddish-brown. Petioles 2-5 mm long, rugose, margin not or barely
winged, dull orangish-red to gray magenta or light purplish-brown, rarely yellowish, densely hairy below,
the hairs wavy or curved distaliy to wavy, off-white to tawny, densely hairy off-white to tawny wavy-hairy on
the upper surface. Lamina flat or sometimes wavy in older leaves, but not typically wrinkled, chartaceous,
43-72 mm long, 18-34 mm wide, length to width ratio 1.5-2.5 : 1, elliptic to oblong-obovate, young leaves
sometimes orbicular, the epidermis p light green in life, base rounded to barely decurrent on the petiole,
margin undulate, at least in life, lightly revolute, apex obtusely rounded; lower lamina surface
glabrate to densely hairy, the hairs wavy or curved distally, « 1 mm long, upright, off-white to tawny, with
sparse subappressed straight hairs « 2 mm long, clavate glandular hairs often present, deciduous, leaving
the brownish epidermis viscid, the stomatal apparatus subopaque, conspicuous; upper lamina surface
with dense, tawny, wavy hairs, and occasional stellate hairs, the epidermis dark greenish-brown. Venation
+ arcolanguid (the major latera! veins fading terminally, forming indistinct loops with superadjacent lateral
veins), midrib prominent below, with more long straight hairs than the lamina surface, impressed basally
on the upper surface, sparsely upright wavy-hairy, clavate glandular hairs often present, the epidermis
dull yellow to light magenta; lateral veins 5—8 per side, irregularly spaced, angles of divergence from the
midrib irregular, fine but prominent below, pubescent, the epidermis light dull orange to purplish-brown,
barely raised above, the lamina-vein seams impressed; 3° veins slightly raised below, inconspicuous above.
Male inflorescences and flowers unknown. Female inflorescences unknown. Female flowering calyx
(based on one immature fruiting calyx) densely tomentulose outside, tube thick, 2.3 mm long, + 3 mm
wide, lobes 7 mm long, 8 mm wide; female corolla tube 4 mm long, 3.5 mm wide, light reddish-cream
sericeous inside, the hairs sparser in the basal 1/3, the epidermis dark reddish-brown; female corolla
lobes ovate, 3 mm long, 1.8 mm wide; Pistil unknown. Fruiting pedicels 2.5-5 mm long, densely curly
to wavy-hairy, bracts opposite, lanceolate, + 2 mm long, concave. Fruiting calyx cupulate to explanate,
outside minutely hairy, the hairs kinked to wavy, the epidermis shiny and viscid, dark reddish-brown, tube
3.5-6 mm long, dehiscent below the sinus for a short distance, the upper 1/2 becoming sparsely sericeous
inside, glabrous basally, lobes obtuse, 4—7 mm long, 6.5-12 mm wide, vaguely pointed, laxly reflexed, tawny
to reddish-brown sericeous inside with a broad intramarginal band. Fruit 2-2.3 cm in diameter, golden
brown, sparsely pubescent, the hairs up to 2 mm long, translucent to tawny. Seeds 11-13 mm long, 6-6.5
mm radial depth, 4.5-5 mm wide.
The subspecies is endemic to the coastal plains of western Guerrero (Fig. 40). It typically grows on
loamy or silty sand at the base of eroding slopes and on roadside embankments in dry tropical forest, and
along the edges of planted fields. Known localities do not seem to be protected, but there are collections
1066 Journal of the Botanical R h Institute of Texas 2(2)
Fic. 31. A. Di py ie Crandl subsp chutlensis M.C. Provance, |. García & A.C. Sanders, subsp nov..I | he hol f P. j “El Palomar,”
n freer
Chutla, Guerrero, Mexico (£ Guízar N. & L. Pimentel B. 3067, IEB). B. F | lla of sub I chutlensis, based on G.B. Hinton et
al. 14583 from Atoyac, Guerrero.
Provance et al., Diospyros salicifolia complex 1067
Fic. 32 leaves of D aeaunric cuben
1 t
1068 J | of the Botanical R h Institute of Texas 2(2)
Fic. 33. Trunk (c. 9 cm dia.) of D. aequoris subsp. chutlensis f l li T. Hwv 200 and Chutla de N Guerrero.
) ] ji ,
from Barbulillas, several hundred yards away from an ecological reserve in the vicinity of Ixtapa. Fonseca
and Lozada-Pérez's (1993) description of “Diospyros acapulcensis” as one of the four most common shrubs
in the halophytic vegetation zone along the coast southwest of Laguna de Coyuca may be in reference to
this taxon. Although we have not seen specimens from their studies, based on the habitat and elevation, we
suspect these reports represent D. ae. subsp. chutlensis or D. salicifolia. However, Koch & Fryxell 82139 (US!,
CHAPA), collected near this locality represents D. ae. subsp. chutlensis. Diospyros ae. subsp. martineziana is
separated by its smaller petioles, smaller lamina shape ratios, soft, fine, light colored, glossy abaxial lamina
hairs, and brochidodromous venation. This taxon is quantitatively distinguished from Diospyros salicifolia
by its shorter leaves with smaller lamina shape ratios (Table 1. characters 3, 7, 8). This subspecies is known
to occur near, but not at Acapulco.
Provance et al., Diospyros salicifolia complex 1069
Tagle 1. Twenty
Character Abbrev. Type
1. lamina length LL leaf size
2. lamina width LW leaf size
3. lamina length : lamina width LL:LW leaf shape
4. petiole length ptL leaf size
5. lamina width at 0.25 length LW1/4 leaf size
6. lamina width at 0.75 length LW3/4 leaf size
7. lamina length : width at 0.25 length LL:LW1/4 leaf shape
8. lamina e an at - 75 a LL:LW3/4 leaf shape
9. lamina | tween four centermost lateral veins LV venation pattern
10. lamina width at 0.25 [eris width at 0.7 LW1/4:LW3/4 leaf shape
11. lamina width at 0.25 : lamina width LW1/4:LW leaf shape
12. lamina width at 0.75 : lamina width LW3/41W leaf shape
13. fruiting pedicel length pdL infl. size
14. fruiting sepal width SW infl. size
15. fruiting se ngth SL infl. size
16. fruiting calyx tube length CL infl. size
17. fruiting sepal width : fruiting sepa! length SW:SL infl. shape
18. fruiting sepal width : fruiting calyx tube length SW: CL infl. shape
19. fruiting sepal width : fruiting pedicel length SW:pdL infl. shape
20. fruiting sepal length: fruiting calyx tube length SL: CL infl. shape
21. fruiting sepal length : fruiting pedicel length SL:pdL infl. shape
22. fruiting calyx tube length : fruiting pedicel length CL:pdL infl. shape
23. fruiting calyx size (fruiting calyx tube length + fruiting sepal CX infl. size
width + fruiting sepal width)
Representative specimens. MEXICO. Guerrero. Mpio. Acapulco de Juarez: km 5.5 carr. Acapulco-Pie de la Cuesta, 60—160 m, 21
, S.D. Koch & P.A. Fryxell 82139 (US, CHAPA); Punta Gorda, Laguna de Tres Palos, 10 m, 11 Oct 1988, N. Diego 4920 (IEB).
Mpio. Átoyac de Alvarez: Atoyac, [17°12'N, 10026! W], 25-300 m, 30 Aug 1939, G.B. Hinton et al. 14583 (UC, IJ, K). Mpio. Coyuca de
Benítez: between Acapulco and Ixtapa, just N of El Zapote, 16 Jan 1983, J.S. Miller et al. 461 (MO). Mpio. José Azueta: Punta Ixtapa,
al W de Ixtapa Zihuatanejo, 30 m, 2 Apr 1991, G. Castillo C. & P. Zamora C. 6276 (XAL); Ixtapa, Col. Barbulillas, 0.5 km N of Hotel La
Costa, 25 m, 10 Mar 2005, Provance 10020A (UCR, CIMI) and Provance 10020B (UCR, CIMI). Mpio. La Unión de Isidoro Montes de
Oca: Felicianos, road N from Hwy 200, 45 m, Provance 9981 (UCR, CIMI); road N from Hwy 200 going towards Chutla, 61 m, Provance
9923 (UCR, CIMD); Joluta, several km N of Hwy 200, 61 m, P 9930 (UCR, CIMI); vicinity of Coyuquila and La Salada, along Hwy
200, 25 m, Provance 9933 (UCR, CIMI); Japoncito, dirt 24 on N side of Hwy 200, 26 m, Provance 9932 (UCR, CIMI); 46 mi NW of
Zihuatanejo, 7 Mar 1979, C.D. Johnson 667-79 (MO)
2d. Diospyros aequoris Standl. subsp. martineziana Conde ex Leavenworth) M. E Provance, I. García &
A.C. Sander TS, comb. et stat. nov. (Figs. 34,3 d). B ex Leavenworth, Amer. Midl
Naturalist 36:184. 1946. Tree: MEXICO. Michoacan: Tancitaro Region, Aaa, above 2500 ft, 20 Aug 1941, WC. Leavenworth &
H. Hoogstraal 1735 (HoLotvee: F).
Trees, sometimes shrubs, 1.2-8 m tall; trunk up to 30 cm in diameter, bark somewhat smooth (not slick),
with scattered checked regions, sometimes irregularly patchy, light to dark gray; stems scurfy to glabrous
and very dark or castaneous, the young stems densely hairy, the hairs slightly wavy to curly, spreading to
ascending, colorless to white, clavate glandular hairs sometimes present. Petioles 2—3(—5) mm long, glabrous
to densely straight to slightly wavy subappressed hairy, pubescent above. Lamina chartaceous, orbicular,
oblong or obovate, 19-61(-65) mm long, 13-40 mm wide, sapling leaves 70-87 mm long, 40-54 mm
wide, length to width ratio 1.2-1.5 : 1, base rounded to subcordate, slightly decurrent on the petiole, margin
flat, apex rounded to obtuse, sometimes retuse, sometimes subacute; lower lamina surface moderately to
densely hairy, the hairs wavy to straight, subappressed to upright, soft, fine, glossy, white to gold, clavate
glandular hairs i present, the epidermis, yellowish to dark green; upper lamina surface sparsely
1070 Journal of the Botanical R h Institute of Texas 2(2)
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ds et ENT aequoris shep. ipium ips Based on M.C. Provance & |. García R. 9393 (UCR). A. St itl [ į male fl B
Male flower M ll E. Geminate st F- l. Based on M.C. p piss 9394 (UCR).
F. Mal | Lateral fusion of filaments. H. Adaxial surface of male flowering calyx. I. G ased onJ.C. Soto N,
& E.M. Martínez 4189.) J. Male corolla. K. Male corolla lobe. L. Pistillod | ith fil i ted on tl tad x à Based on /. García
R. &CG 1o 6989 (UCR). M. F I hlet. N Fruiting calices. 0 il le ( ited) of fl
|, based o
Provance & I. Garcia R. 93964 (UCR). P-S. Based on M.C. Provance 8 García R. 9392 (UCR). P, Pistil of flower type li. Q. Female flower. R. Female corolla.
S. Female corolla lobe.
Provance et al., Diospyros salicifolia complex 1071
to densely hairy, the hairs uniformly curved to wavy, ascending to upright, soft, fine, white to gold, the
epidermis olive to dark green, hair bases raised and nearby epidermal cells enlarged. Venation brochido-
dromous, yellow to chartreuse; midrib prominent below, narrow, with spreading, straight to slightly wavy
hairs above; lateral veins (3—4—6(—7) per side, fine but prominent below, connecting with superadjacent
laterals inward from the margin, sparsely hairy, barely impressed to barely raised above (+ obscure); 3? veins
very fine below, obscure above. Male inflorescences solitary (1-)3—4(-6)-flowered cymes, densely cream
to bd itn bn O Wiese to wavy-hairy; peduncles 0.5-3 mm long; pedicels 0.5-2 mm long. Male
lightly winged, moderately to densely white to yellowish velutinous-pubescent,
the hae Somede MM, with tomentulose undercoat, epidermis viscid tube 2.5-3.5 mm long,
3.8-4 mm wide, glabrous inside, lobes acute, (3.5—)5.5-6(-7) mm long, 3-)4—4.5(-5.5) mm wide, some-
times keeled, the nerves sometimes obvious, densely hairy inside, the intramarginal band width variable;
male corolla tube 4.5-6 mm long, 3-4 mm wide, sericeous outside, with 3 pale pink 0.5-0.8 mm long
shield-shaped glabrous basal regions; male corolla lobes lance-ovate to elliptic, 3-3.5 mm long, 1.2-2.5
mm wide. Stamens 10-12; filaments adnate to the base of the corolla or inserted on the receptacle, 2-2.5
mm long, narrowed distally, free, geminate or joined laterally, rarely in threes; anthers lance-ovate, 2-2.5
mm long, sometimes rostrate. Female inflorescence densely hairy, bracts narrowly triangular, 1.0-1.3
mm long. Female flowers apparently dimorphic (types I and II below, both types can be found in one
individual); female flowering calyx densely tomentulose outside, with some longer, straight to slightly
wavy, + ascending, light gold hairs, tube half-globose, 5-6 mm long, 7 mm wide, the upper 1/2 densely hairy
inside, lower 1/2 glabrate with a few reddish glandular hairs, lobes 7-8 mm long, 7-8 mm wide, acuminate,
the nerves prominent, densely hairy inside with a broad intramarginal band; female corolla tube 6 mm
long, 3.3 mm wide, with 3 shield-shaped 3.5 mm long glabrous basal regions: flower type I corolla tube
basal regions pinkish, + overlapping; flower type II corolla tube basal regions dark red, not or barely
overlapping; female corolla lobes rhombic to ovate, 4 mm long, 2 mm wide; flower type I ovary ovoid,
minutely cream sericeous; flower type II ovary shaped like a barrel, glabrous to scantily hairy, epidermis
golden; flower type I styles 3, free, arching outward; flower type II styles 3, in a fused column; stigmas
bifid. Fruiting pedicels 2-4 mm long, bracts slender, 1.8-2 mm long, about 0.3 mm wide, conduplicate.
Fruiting calyx shallowly rotate to cupuliform, tomentulose to wavy-hairy outside, much of the vestiture
including the longer hairs, deciduous in age, sometimes persisting on the conspicuously viscid, epidermis
tube 5—5.5 mm long, densely hairy inside, lobes spreading, obtuse to rounded, 3-8 mm long, 8.5-10.5 mm
wide, the margins usually revolute. Fruit globose, subglobose or obovoid, 2-2.3 cm in diameter, glabrous
or sparsely hairy, the hairs straight to slightly wavy, fine, ascending, white to golden, clavate glandular hairs
sometimes present, epidermis glaucous, pruinose, scintillant, tending to separate from the remainder of the
fruit upon drying, light grayish-yellow to golden brown, mature fruit light green in life. Seeds 13-14 mm
long, 6 mm radial depth, 5.5 mm wide
This subspecies grows in dry deciduous forest of the Apatzingan Valley, Michoacan, between 320 and
700 m (Fig. 40). Its occurrence has not been documented for any protected area. It is an occasional tree at
the type locality on the slopes of Acahuato, north of Apatzingan, Michoacan, where it is sometimes spared
adjacent to milpas to produce shade for workers (pers. obs).
Representative specimens: MÉXICO. Michoacan: Mpio. Apatzingan: Acahuato, 671 m, 2 Jul 2004, M.C. Provance & L Garcia R. 9392,
9393, 9395, 9396A, 9396B, 9394 (CIMI, UCR); Acahuato, 750 m, 17 Feb 2005, I. Garcia R. & C. Conrado 6989, 6983 (CIMI, UCR); above
Apatzingan, 2000 ft, 13 Aug 1941, W.C. Leavenworth & H. Hoogstraal 1502 (MO, ILL), same location, 1490 (MO, ILL), same location, 1494
(MO, ILL); Mt. Apatzingan, 2000 ft, 20 Aug 1941, W.C. Leavenworth & H. Hoogstraal 1728 (MO). Mpio. Huetamo: En Pataseo [Patasco],
6 km al E de Quetzería, 300 m, 18 Jul 1982, J.C. Soto N. & E.M. Martínez 4189 (XAL, CHAPA, MO, ILL). Mpio. La Huacana: Sierra
Las Cruces, ca. 6.5 km (by air) west-southwest of Los Ranchos, 600 m, 6 Dec 2003, Steinmann 3911 (UCR). Mpio Parácuaro: "Paso de
Yeguas” [Crucero de las Yeguas], 17 km a W de Cuatro Caminos, 250 m [+ 320 m], 18 Oct 1982, R. Torres C. 1541 (CAS
2e. Diospyros aequoris Standl. subsp. rekoi (Standl.) M.C. Provance, I. García & A.C. Sanders, comb. et
stat. nov. (Figs. 35, 38a). Bastonym: Maba rekoi Standl. Contr. U.S Natl. Herb. 20:193. 1919. Tee: MEXICO. Oaxaca. [Mpio.
1072 Journal of the Botanical R h Institute of Texas 2(2)
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d F Male corolla. G. Male corolla lobe :
J. Leaves (abaxial surface). B-C, G. Based on T.B. Salvato & M.C. Provance 247 (UCR). D, H-1. Based on 7.8. Salvato & M.C. Provance 246 (UCR). E, |. Based
on T.B. Salvato & M.C. Provance 293 (UCR). F. Based on T.B. Salvato & M.C. Provance 244 (UCR)
fit E F 31 J ne.
Provance et al., Diospyros salicifolia complex 1073
San Pedro Pochutla]: Puerto Angel, [15°40'N, 96°29'W, 1 m], 28 Sep 1917, B.P Reko 3429 (HT: US-842523). Diospyros portus Standl.
Publ. Carnegie Inst. Wash. 461(4):80. 1935.
Not Diospyros rekoi Standl., J. Wash. Acad. Sci.17: 527. 1927.
Shrubs, rarely trees, 1-4 12) m tall, sometimes with multiple stems from root and crown sprouts; trunk
somewhat smooth (not slick) to checked, the bark whitish with gray to gold patches; stems glabrous, some-
times hispidulous or with longer desiccated hairs in the 2nd year, young stems densely puberulent, with some
additional longer, spreading to ascending hairs. Petioles 3—)4—6(-7) mm long, glistening, dark, reddish,
densely minutely wavy-hairy on the lower surface, the hairs sparser and longer above. Lamina subcoria-
ceous, oblong or elliptic to obovate, (440-)50—90(-100) mm long, 16—35(-45) mm wide, the length to width
ratio 2.2—2.8 : 1, base broadly rounded to the petiole, margin revolute, apex broadly to obtusely rounded
(revolute lamina margins sometimes forming a false point); lower lamina surface moderately velutinous
to wavy-velutinous, the hairs ascending to upright, clavate glandular hairs often present, deciduous, leaving
jus pde» conspicuously viscid, the stomatal apparatus opaque and conspicuous to translucent; upper
wrinkled when mature, velutinous to wavy-velutinous. Venation brochidodromous; midrib
prominent below, ascending velutinous, impressed above, velutinous to wavy-velutinous, clavate glandular
hairs often present; lateral veins 4—8 per side, prominent below, impressed above; 3? veins prominent
below, impressed above; 4° veins sometimes darkened below. Male inflorescences 1-3-flowered cymes,
hirsutulous; peduncles 1-3 mm long; pedicels 0.3-2 mm long, bracts linear-lanceolate, 2-3 mm long,
conduplicate. Male flowering calyx densely puberulent, the epidermis viscid, tube 3-4 mm long, 3-3.5
mm wide, glabrate inside, some scattered minute hairs in the lower 1/2, lobes acute, (3.5-)5-6 mm long,
3-5 mm wide, glabrous to sparsely puberulent inside, the intramarginal band somewhat narrow; male co-
rolla tube 4.8—5.5 mm long, 3 mm wide, mostly sericeous outside, but basal 0.3-0.8 mm glabrous; male
corolla lobes oblong, ovate or rhombic, 3-4 mm long, 2 mm wide; Stamens (1112; filaments adnate to
the tube near the base, irregularly or vaguely in two tiers, or inserted on the receptacle, 2 mm long, some-
times subgeminate; anthers lanceolate to oblong, 2-2.4 mm long, sometimes minutely lobed basally on
each side of the connective (‘eared’), sometimes apiculate. Female inflorescences wavy-hirsutulous with
orangish ascending hairs, bracts linear-lanceolate, 2-3 mm long, conduplicate. Female flowering calyx
campanulate, straight to wavy-hairy, the hairs upright to ascending, orangish, tube cupulate, 2.5-3 mm
long, 5 mm wide, lobes obtusely rounded, 5.5-6 mm long, 6.5 mm wide, barely acuminate; female corolla
tube 5.5—7.5 mm long, 3.5-4 mm wide, exterior sericeous nearly to the base; female corolla lobes ovate
to rhombic, 3.3-4 mm long, 2.5 mm wide; ovary globose to somewhat conical, x 2 mm in diameter, usually
glabrous, sometimes with 3 rows of long hairs running from the base of the ovary to the base of the style
column; styles 3, column + 2.5 mm long; stigmas bifid, + 0.5 mm long, somewhat spreading; staminodes
3(-5), opposite to the petals, flat, the sterile anther narrowly lanceolate, 2 mm long, the filament 2 mm long,
adnate to the base of the corolla for 0.5-0.8 mm. Fruiting pedicels 4—8(-12) mm long, often glabrate.
Fruiting calyx rotate, glabrate to sparsely straight to wavy-hairy, the epidermis dark brown and conspicu-
ously viscid, tube 4.5—6.5 mm long, hairy inside to the base, lobes 5-8 mm long, 11-14 mm wide, sericeous
inside with a broad intramarginal band. Fruit 1.5—2.2 mm in diameter, epidermis reportedly yellow when
mature. Seeds unknown.
This subspecies i demic to the coastal region of Oaxaca (Fig. 40). It is common on sandy mesas with
littoral scrub near the beach, and on rocky seaside cliffs where it tolerates at least small amounts of salt-spray.
It also occurs a few kilometers inland on steep rocky slopes with dry tropical forest. Saplings are sometimes
very common along disturbed sandy roads near the beach. This subspecies forms intermediates with D. ae.
subsp. tehuantepecensis several kilometers east of Bahias de Huatulco, but can generally be distinguished
from it by its longer and wider lamina, longer petioles, larger calyx including wider fruiting sepals, longer
fruiting pedicels, and longer tube.
Representative specimens: MEXICO. Oaxaca. Mpio. San Pedro Mixtepec: Puerto Escondido, 22 Sep 1982, R. Cedillo T. & R. Torres C.
1813 (MO, XAL). Mpio. San Pedro Pochutla: Tahueca, 10-15 m, 3 Aug 2003, T.B. Salvato & M.C. Provance 244, 245, 246, 247 (UCR).
1074 J lof the Botanical R h Institute of Texas 2(2)
Mpio. S María H 1 erro San Antonio, Bahia de San Agustin, 50—60 ft, T.B. Salvato & M.C. Provance 293 (UCR). Mpio. Santa
María Tonameca: Playa Agua Blanca, 35 m, 4 Aug 2003, T.B. Salvato & M.C. Provance 363, 366 (UCR
2f. Diospyros aequoris Standl. subsp. tehuantepecensis M.C. RS E dius = A. Ç: E subsp.
nov. (Figs. 36, 37a, 39b). Tre: MEXICO. Oaxaca. Mpio. Tel 1 Cerro Guiengola,
16°22'55.4"W, 95°18'27.3"W, 300 m, 6 Aug 1994, J.L. Panero & J.I. T 4490 roe IEB-48849!; isotypes CAS!, TEXI).
ne f Ty I 1 1 1 EP A Aste 1 £A g 1 I g +1 11/4 et 3/4
longitudinem, infi ti vel magentea vel colori! bal ]
Trees or shrubs, 2—5 m tall; trunk unknown; stems glabrous to pulverulent or scurfy, the young stems
pilosulose to villosulous, with some longer slightly wavy ascending reddish-orange hai the shoot apex.
Petioles 2-45) mm long, hairs similar to young stems, rugose below and sometimes SOS epidermis
magenta to yellowish or cream, sometimes variously striated. Lamina cl , elliptic
or oblong, sometimes obovate, rarely oblanceolate, 20—56(-67) mm long, 12-24-27) mm wide, length to
width ratio (1.6—)2.1—2.6 : 1, base rounded to the petiole, margin sometimes revolute near the base, apex
obtusely rounded, sometimes acutely; lower lamina surface hairs moderate to dense, straight to wavy,
ascending to subappressed, clavate glandular hairs often present, the epidermis usually light gray, the sto-
matal apparatus opaque, white, and conspicuous; upper lamina surface usually wrinkled, villous, clavate
glandular hairs often present, the epidermis densely papillose. Venation brochidodromous to arcolanguid,
below off-white to gray pilosulose to villosulous, the epidermis cream to magenta or speckled with both
colors; midrib prominent below, impressed above except basally where slightly raised, densely villous, the
epidermis yellow orange; lateral veins 5—7 per side, prominent below, slightly impressed above; 3? veins
raised below, slightly impressed above. Male inflorescences 3-flowered cymes; pedicels 1-2 mm long,
densely pubescent. Male flowering calyx densely pubescent to wavy-hairy outside, hairs ascending, off-
white, tube urceolate, 4 mm long, 3.5-4 mm wide, weakly winged, glabrous inside, lobes triangular, 4—5
mm long, 3-3.2 mm wide, ascending to slightly spreading, glabrous inside with a narrow intramarginal
band; male corolla tube 5.5-6 mm long, 3 mm wide, the upper 1/2 sericeous outside, lower 1/2 with 3
shield-shaped glabrous basal areas, + 1.8 mm long, bordered distally by minute curly ascending hairs; male
corolla lobes rhombic, 3 mm long, 1.8-2 mm wide, sometimes involute basally. Stamens 12; filaments
adnate to the base of the corolla in a single tier, rarely inserted on the DRM 2 mm long; anther? lan-
ceolate, 2.3 mm long. Female inflorescences densely pubescent. Femal ly minutely sericeous
outside, with sparse longer, wavy, ascending to spreading hairs, tube shallow-eupulate: 1.5 mm long, 4mm
wide, glabrate inside, lobes obtusely rounded, 6-7 mm long, 7-7.5 mm wide, densely hairy inside with a
broad intramarginal band; female corolla tube 4.5 mm long, 3 mm wide, similar to male corolla tube,
the glabrous basal regions 3 mm long, 2.2 mm wide; female corolla lobes rhombic, 3.5 mm long, 2 mm
wide; ovary depressed-globose, 2 mm long, 3 mm wide, cream sericeous; styles 3, upright, not fused, 1.8
mm long, minutely sericeous abaxially, white tomentulose adaxially; stigmas 3, bifid; staminodes 3, the
filaments adnate to the corolla base, 1.5-2 mm long, the sterile anther narrowly lanceolate, 1.5 mm long.
Fruiting pedicels 2.5—-6.5(-8) mm long, densely ascending-pubescent. Fruiting calyx explanate, densely
puberulent outside, hairs cream to orange; tube 3—)4—5(-7) mm long, lobes broadly obtuse, (3.5-)4-8 mm
long, (7-)8.5-12.5 mm wide, spreading to reflexed. Fruit globose, 1.5-2.5 cm in diameter, glabrous to
sparsely puberulent, the hairs minute, straight, appressed to ascending, epidermis amber to dark yellow,
green in living material when mature. Seeds 11-12.5 mm long, 8 mm radial depth, 5-6 mm wide.
This subspecies occurs in dry tropical forests, between 50 and 1100 m, in the southwest part of the
Isthmus of Tehuantepec, Oaxaca (Fig. 40). It has been described as frequent at three sites with limestone
substrates but has also been reported from sandy soils and disturbed sites. The occurrence of this taxon on
Cerro Guiengola is significant because dry tropical forests of this region are considered high in quality, and
have consequently been recommended for formal conservation following a study by Gordon et al. 2004).
The species forms hybrids with D. ae. subsp. rekoi in the vicinity of Tehuantepec and in coastal regions near
Salina Cruz, and in the Municipio of Santiago Astata.
Provance et al., Diospyros salicifolia complex
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Fic. 36. Holotype of D. a came Misi cis C Provance, 7 García & A.C. Sanders, subsp. nov. from Mixtequilla, Oaxaca (J.L. Panero
1075
1076 J lof the Botanical R h Institute of Texas 2(2)
Fic. 37. Diospyros aequoris Standl. A-1. subsp. tehuantepecensis. A—D. Based on J.L. Panero & J.1. Calzada 449 (CAS). A. Styles and stigmas. B. Female
flower. C. F. l lla tul D.F | lla lobe (al ial vi ) E-i ased on J. R fi ki 33065 (MOU) E Male flower. F. Adaxial f; f |
calyx. G. Male corolla tube (adaxial view). H. Stamen. 1. Male corolla.lobe. J-M. subsp. balsensis, based on G.B. Hinton 3959 (K) from Bejucos, Mexico.
J. Male calyx K. Adaxial surf. f male fl i ly L. Male corolla. M. Stamen
El
Provance et al., Diospyros salicifolia complex 1077
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Fic. 38. Abaxial leaf vestiture D. aequoris, including midrib (scale — 1 ILA subsp rekoi (T. Salvato & M.C. Provance 487, UCR). B. subsp. aequoris (M.C.
Provance et al. 9607, UCR). C. subsp. balsensis (M.C. Provance 9919, UCR). D. subsp. martineziana (M.C. Provance & I. Garcia R. 9396B, UCR).
1078 J | of the Botanical R h Institute of Texas 2(2)
i ap eni specimens. MEXICO. Oaxaca. Mpio. Guevea de Humboldt: Cerro de Picacho, Jul 1914, C.A. Purpus 7444, 7176 (MO).
io. Ixtepec: Cerro Blanco por la carretera Juchitán—Ixtaltepec, 200 m, 15 Feb 1982, J.I. Calzada & J. Elizondo 8508 (X AL). Mpio.
Magdalena e er 8 km al SW de La Pajima, 285—330 m, 29 Aug 1994, G. Flores-Franco & J.I. Calzada 3621 (NLU, ARIZ). Mpio.
Magdalena Tlacotepec: 4 km del ojo de agua de Tlacotepec hacia la Ollaga, 260 m, 26 Sep 1984, R. Torres C. 6277 (MO). Mpio. San
Pedro Huamelula: Playa Grande, 2.1 km ab E, 50 m, 31 Jul 2000, S. Salas M. 3213 (IEB). Mpio. Santa Maria Jalapa del Marqués:
Rancho Ricardo, al N. de Buenos Aires baS 12 Sep 1985, R. Torres C. & C. Martínez 7312 (MO). Mpio. Santiago
Lachiguirí: Aproximadamente 10 km al NE de " cortina de la Presa Benito Juárez, camino de terraceria a Santiago Lachiguirí, 100 m,
24 Aug 1994, G. Flores-Franco 3556 & J.I. Calzada (ARIZ). Mpio. Santo Domingo Ingenio: 10 km al NE de La Ventosa, 2 Dec 1980, R
Cedillo T. & D. Lorence 503 (ILL). Mpio. Tehuantepec: 22 km WNW de Tehuantepec, sobre la carr. a Oaxaca, 200 m, 22 qe 1975, I
alia ies (MOU); 5 km al SW de Tehuantepec, 40 m, 28 Jun 1982, R. Torres C. & R. Cedillo T. 684 (CHAPA,
; O de Tehuantepec, 400 m, 5 Feb 1988, C. Martinez R. 1288 (IEB); Piedra de Cal, 5 km al NO de Jeanine 50 m, 18
Feb pene P. Tenorio L. & R. Torres C. 5480 (MO, CHAPA); "Rancho Lizet,” 6.3 km al SW de la entrada a Buenos Aires, 190 m, 26 Mar
1984, R. Torres C. & C. Martínez R. 4787 (MO, CHAPA); Cerro Guiengola, 1110 m, 27 Nov 1983, R. Torres C. 4194 (CHAPA); Cerro Pozo
Zorillo, entrando por Las Pilas, 8 Feb 1988, C. Martinez R. 1297 (IEB); 20 km al O de Tehuantepec, entrando por Las Pilas, 5 Jan 1988,
C. Martínez R. 1158 (MO)
3. Diospyros salicifolia Humb. & Bonpl. ex Willd. Sp. Pl. Editio quarta 4:1112. 1806 (Figs. 1, 2c, 39d,
41, 42, 43). Maba salicifolia (Humb. & Bonpl. ex Willd.) Hiern, Trans. Cambridge Philos. Soc. 12:129. 1873. Tre: MEXICO.
GUERRERO: near Acapulco, no date, Humboldt & Bonpland s.n. (GoLoTveE: B-W-19250-01, digital images!).
Diospyros albens C. Presl, Reliq. Haenk. 2:62. 1835. Macreightia albens (C. Presl) A. DC., Prodr. (DC.) 8:221. 1844. Ebenus albens (C.
Presl) Kuntze, Revis. Gen. Pl. 2:408. 1891. fer. MEXICO. GUERRERO: “Habitat ad portum Acapulco Mexicanorum,” date unknown,
T. Haenke s.n. (HOLOTYPE: W-31942, as photo F9).
Trees, 1.5-6 m tall, facultatively deciduous; trunk unknown; wood unknown; mature stems terete, half-
netted, glabrate sometimes black gland-dotted, epidermis tan; 2nd year stems similar to mature stems; 1st
year stems apadranp niar becoming terete, moderately Biy the hairs usually minute, thick, appressed,
fluid-filled (the hairs throughout the plant are mostly si lular hairs, likely translucent in life, and the
color of hairs in herbarium specimens is due to denia B. a fluid in the lumen), associated with glandular
punctae, rarely some longer wavy subappressed hairs present near the tip of the growing shoot, lenticels
narrowly elliptic, few in number, bud scales deeply concave, triangular, 1.5-2 mm long, moderately to densely
hairy, the hairs strongly appressed, similar to those on the young stems, glandular punctate, sometimes
with minute clavate glandular hairs. Leaves alternate, simple, entire, closely spaced on new growth; peti-
oles subterete, at least distally, 2—)3—5 mm long, sparsely to moderately hairy above and below, the hairs
wavy, appressed to ascending, yellowish; lamina chartaceous, narrowly elliptic to oblong, oblanceolate,
sometimes obovate, 60—80(-110) mm long, 22-32(-35) mm wide, the length to width ratio (2.3-)2.5-3.2
: 1, somewhat revolute, at least along the lower 1/2 of the margins, apex broadly to acutely rounded, base
rounded to the petiole, very rarely cuneate; lower lamina surface moderately hairy in immature leaves,
glabrate to sparsely hairy when mature, the hairs 0.2-0.9 mm long, appressed to subappressed, straight,
or nearly so, more frequent in the basal 1/4 of the leaf, sparsely to moderately clavate glandular hairy, the
glandular hairs amber colored, the epidermis gray in herbarium material, sometimes viscid, the stomatal
apparatus often opaque, white, and conspicuous; upper lamina surface glabrous to sparsely pubescent
with weak hairs, the epidermis weakly papillate, brown to ed Bray and ida ya in herbarium
material, epidermal cells conspicuous with age. Venation brocl guid; midrib narrow
and prominent on the lower surface, usually < 1 mm wide, sparsely to moderately hairy, the hairs 0.3-0.9
mm long, slightly wey to nearly straight, subappressed; the midrib raised aes on the upper surface,
convex, tally, becoming flush with the lamina, glabrate to sp y pubescent, rarely moderately
pubescent, the TER + wavy, the epidermis light yellow above and below; lateral veins 6-9 on each side
of the midrib, diverging from midrib between 45-90", rounding towards the apex distally, intersecondary
veins inconspicuous; 3? veins usually faint below, sometimes slightly darkened and more apparent, faintly
raised or not apparent on the upper surface; 4? veins sometimes slightly darkened and apparent below.
Laminar extrafloral nectaries 0.2-0.6(-0.8) mm in dimension, circular to elliptic. Male inflorescences
solitary (1-)23-flowered cymes in leaf axils of young stems (our description of the male inflorescence and
Provance et al., Diospyros salicifolia complex 1079
"At =
ICHELUTR T AGE
PETENS E
E d.
5 - T RM E
Fic. 39. Abaxial leaf i includi g idrib (scale = 1 ). A. D. aequoris subsp. tlensis (f the hol tyr ). B. D. aequoris subsp. tehuantepecensis
(ftom the holotype). C. D. aequoris subsp. chutlensis (M.C. Provance 9930, UCR). D. young leaf, D. salicifolia (W. Boege 452, FR).
lof the Botanical R h Institut f Texas 2(2)
1080
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TERENI 2008 Apr 19 01:09:23 OMC- Martin Weinek
£n.
is Standley and D. salicifolia Humb. & Bonpl. ex Willd
Fic. 40. Distributi
flowers is approximate, and lacks detail because the only male specimen we have seen is a photo of the type
of D. albens); peduncles 2 mm long; pedicels 1-1.7 mm long. Male flowers 3-merous; male flowering
calyx narrowly campanulate, tube 2 mm long, 2 mm wide, lobes valvate-reduplicate in the bud, narrowly
triangular, 3-5.5 mm long, 2.2 mm wide; male corolla tube urceolate, 3 mm long, 2 mm wide, sericeous;
male corolla lobes 2.5 mm long. Stamens of unknown number; filaments at least sometimes adnate
to the corolla near the base; anthers lance-ovate, 1.5-2 mm long, opening by longitudinal slits. Female
inflorescences solitary flowers in leaf axils of young stems; female flowering pedicels 3-5 mm long,
vestiture similar to that of young stems. Female flowers 3-4-merous; female flowering calyx mitriform,
exterior moderately hairy, the hairs very minute, subappressed, light yellow to amber, apparently borne
from glandular punctae, these most obvious near the sepal margins, tube cupulate, 1.2-1.8 mm long, 3 mm
wide, interior unknown, lobes valvate-reduplicate in the bud, ovate to widely ovate, winged, 6.5 mm long,
6.5 mm wide, faintly 3-6-veined; female corolla unknown; ovary unknown; styles (based on remnants
on fruits) 3, fused into a column with a dark basal band, densely wavy-hairy, the hairs cream to amber;
stigmas unknown. Fruiting pedicels 2-5 mm long, bracts 2, + opposite, deltoid, 0.3 mm long. Fruiting
calyx slightly accrescent, explanate to rotate, the exterior glabrate to sparsely hairy, the hairs slightly wavy
to straight, subappressed; epidermis glandular punctate, glands often black, tube 3-4.5(-6) mm long, not
bulbous at the base, interior lower 1/2 glabrous, otherwise moderately white hairy, the hairs long, fine, glossy,
Provance et al., Diospyros salicifolia complex 1081
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straight and appressed, lobes spreading, broadly ovate to oblong-ovate, 4-6(—6.5) mm long, 8-10 mm wide,
sometimes acuminate, often retlexed along the margins, with a wide interior intramarginal band of hairs, the
hairs dense, minute, curly, the median similar to the upper 1/2 of the fruiting calyx tube. Fruit a « globose
berry, 2-2.5 cm in diameter, with three pairs of locules; flesh probably gelatinous when fresh, vitreous, red
se J | of the Botanical R h Institute of Texas 2(2)
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and translucent upon drying; epidermis smooth to orange-peel textured, sparsely to moderately hairy near
apex and base of the fruit, otherwise glabrous or scantily appressed hairy, the epidermis yellow to dark yel-
low when mature, often pruinose. Seeds 5—6, wedge-shaped (resembling an orange segment with rounded
edges), 15 mm long, 8 mm radial depth, 6 mm wide, reddish-brown, texture granular-foveolate.
Diospyros salicifolia is endemic to dry tropical forest and littoral scrub between 1 and 100 m, in the vi-
Provance et al., Diospyros salicifolia complex 1083
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cinity of Acapulco and Isla Roqueta (Isla Grifo) at the mouth of Acapulco Bay, Guerrero (Fig. 40). Protected
forests exist in the vicinity of Acapulco, but it is unknown if the species occurs in these areas. Because of
its restricted range, apparent association with littoral scrub, and proximity of a rapidly developing tourism
center, the species should be considered threatened. Local names associated with this species include “ta-
malototzin” and “sapotillo.” The fruiting calyx of this species is quite small and similar to D. aequoris subsp
chutlensis. However, the leaves of D. salicifolia are longer and have a greater lamina shape ratio. The foliage
of collections of D. acapulcensis subsp. veraecrucis from coastal regions of Chiapas sometimes resembles D.
salicifolia, but fruiting calyces have the longer tube typical of subsp. veraecrucis. Material from this region
should be evaluated more thoroughly when more collections are available.
Representative specimens. MEXICO. iron M BAR: de Juarez: Isla Roqueta, I.K. Langman 3305 (NA); portum Acapulco,
Haenke 31942 (W photo); C | 6°46'12"N, 99°47'24"W, 1-3 m, (The label incorrectly places Copacabana
in "Puebla/Tlaxcala." The dated is suspect also.), 15 Jul 1966, W. ne 452 (FR); Viveros El Huayacán, La Poza, 22 May 1979, W.L. Forment
790 Und 2 a Ww eni Puerto Marqués, 19 Jan 1981, W.L. Forment 1388 (CAS, JBSD, MO); Puerto Marqués, 28 Jan 1981, W.L. Forment
vicinity, Oct 1894—Mar 1895, E. Palmer 372 (MO); Revolcadero, 27 Jan 1981, W.L. Forment 1317 (SD, XAL).
1338 (X
1084 J lof the Botanical R h Institute of Texas 2(2)
4. Diospyros yucatanensis Lundell. Publ. Carnegie Inst. of Wash. 478:217. 1937. Tres: GUATEMALA. Peren. San
Andres [as on label], "collected on the steep north bank of Lake Petén near San Andres" [protologue], 4 May 1933, Lundell 3237
(HOLOTYPE: MICH; 1soTYPE: F-685309, as image!, LL).
wee or DUE 2—15(-28) m tall, Hentam ie deciduous, dioecious; trunk often relatively tall and slender,
; wood unknown; mat , half-netted to finely reticulated, glabrous, epider-
mis dark reddish-brown to grayish-brown, sometimes lenticellate, sometimes black gland dotted; 2nd year
stems subterete to terete, nearly smooth, minutely ridged or sulcate, glabrous to densely hairy, the hairs when
present often pale and desiccated; 1st year stems angular to subterete, sometimes rugulose, moderately to
densely hairy, the hairs short, wavy to curly, sometimes longer and nearly straight, orangish to reddish-brown
(the hairs throughout the plant are mostly simple glandular hairs, likely translucent in life, and the color of
hairs in herbarium specimens is due to darkening of a Haig in HE En ee glandular hairs sometimes
present, deciduous, leaving the epidermis viscid, j , reddish-brown.
Leaves alternate, simple, entire; petioles (2-)3.5-8(-8.5) mm long, so ee winged, oben only distally,
glabrous to densely hairy, epidermis usually rugose, often viscid, g 1s, sometimes pruinose,
lat 1 lamina hartaceous,
sometimes scintillant, golden to very dark brown, barely
(52-)59-106(-123) mm long, 19-41-48) mm wide, length to width ratio (1.8-)2. 13. 5(—4) : 1, cea
or ovate, sometimes oblanceolate, obovate, or elliptic, at 3/4 length mostly 12-30(-35) mm wide, base acute
and short attenuate to broadly rounded, usually somewhat decurrent on the petiole, margins flat to slightly
revolute, apex acuminate to acute, sometimes rounded; lower lamina surface glabrate to densely hairy,
curly to slightly curved, sometimes straight, cream to tawny to bright orange-red, hair bases usually swollen
and + papillose, epidermis grayish, sometimes viscid, the stomatal apparatus usually opaque and conspicu-
ous; upper lamina surface glabrous to moderately hairy, the hairs wavy, upright, epidermis sometimes
shiny, sometimes crystal-papillose. Venation brochidodromous to arcolanguid, the epidermis yellowish to
reddish-brown and sometimes viscid below; midrib prominent below, glabrous to densely hairy, the hairs
short, curly, with some longer, straight, subappressed, bright reddish-brown hairs; lateral veins 5-8 on
each side of the midrib, their courses mostly arching, thin but + prominent below, often with long, straight,
bright reddish-brown hairs, fine and barely raised to flush above, impressed along the lamina-vein seam;
3? veins subprominent below, though usually somewhat obscured by tHe vestiture, usually obscure above,
sometimes slightly impressed; 4°-5° veins obscure. Laminar ies usually on basal half of
leaf, often near midrib, sometimes on intersecondary veins, mostly elliptic, 0.3-1 mm long, concave. Male
inflorescences 1—5-flowered cymes in leaf axils on young stems, densely hairy, the hairs minute, curly to
wavy, reddish; epidermis viscid and dark brown; peduncles 2-6 mm long; pedicels 1-3 mm long, bracts
ovate or deltoid to linear lanceolate, 1-3 mm long. Male flowers 3—4-merous; male flowering calyx nar-
rowly campanulate, exterior moderately to densely hairy, the hairs straight to curly, golden to reddish, the
epidermis reddish-brown, tube 3-5.5 mm long, 3.5-5.5 mm wide, interior entirely glabrous, or glabrous in
the lower half and densely hairy above, the hairs straight, appressed, lobes valvate-reduplicate in the bud,
oblong-ovate to lance-ovate or triangular, 2-4 mm long, 2.8-3.5 mm wide, acute to subacute, sometimes
barely acuminate, interior intramarginal band narrow, moderately to densely hairy, the hairs minute and
curly, the median hairs moderate to dense, the i porre or d so, appressed to subappressed; male
corolla white, yellowish or pinkish; mal rical, narrowing distally, 6—10 mm long, 3-4
mm wide, exterior with 3 small shield-shaped glabrous ee at base, or with a narrow glabrous basal band,
sometimes with minute wavy hairs just distal to glabrous areas, otherwise tawny to reddish sericeous with
a dense undercoat of minute curly hairs; male corolla lobes oblong-ovate, 3-5 mm long, 1-2.3 mm wide,
exterior densely hairy, the hairs curly to straight, subappressed to ascending, reddish-gold to brownish, the
epidermis reddish to dark reddish-brown. Stamens 10—14; filaments adnate to the corolla at or near the
base, in two well-defined tiers, or with a few stamens inserted on the receptacle, sometimes a few geminate,
(1.522-3.5 mm long; anthers lanceolate to ovate or oblong-ovate, 2.3-4.5 mm long, lower anthers usually
shorter, apex sometimes rostrate or cuspidate, sometimes thickened or granular near the apex; pistillode
Provance et al., Diospyros salicifolia complex 1085
minute, hairs dense, wavy or straight. fendi icut solitary flowers in leaf axils of young stems,
the vestiture similar to male infl g pedicels with hairs and epidermis similar to
£. 1
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mitriform, y winged,
male inflorescences. Female flowers 3-4C5)-me ‘ous
tube 2.5—3.3 mm long, 4—5 mm wide, lobes valvate-reduplicate in the bud, bluntly acute to slightly rounded,
3.3-6 mm long, 3.5-5.5 mm wide; corolla tube urceolate, narrowing distally, 3-5 mm long, 2.5-3 mm
wide, exterior reddish sericeous except for some glabrous parts near the base; corolla lobes lance-ovate to
ovate, 3.5-4.5 mm ORB, l. NA 5mm ue JOE ovoid, sericeous, the hairs reddish and fluid-filled; styles
3, columnart Fruiting pedicels 2-14(-16) mm
long, Bsoadening Pede ee to pis hairy, the hairs shor. wavy, subappressed to appressed, often
pale and desiccated on mature fruit, epidermis reddish to brownish, bracts not seen. Fruiting calyx rotate
to explanate, exterior hairs similar to fruiting pedicels, tube 3-6(—7.5) mm long, interior sparsely to densely
hairy, the hairs very short, fine, straight and appressed, but glabrous in the basal 1/8—1/3, lobes accrescent,
broadly rounded to oblong, 3-10(-12) mm long, (7-)9-14(-15) mm wide, spreading to reflexed, the margin
sometimes conspicuously undulate, tip rounded, tapered or barely acuminate, ti ing narrowed
to a point due to folding of the distal margin, intramarginal band very wide, sparsely to i dead hairy,
the hairs minute, curly, erect to ascending, amber; the epidermis puncticulate viscid; median sparsely to
moderately hairy, the hairs similar to those of inner calyx tube, present only near the base of lobe. Fruit a
globose to depressed-globose berry, 1.5—2.5 cm in diameter, sometimes umbilicate, often partly dehiscent,
consistently with three pairs of locules, immature fruit sparsely reddish-brown hairy, mature fruit glabrate
except near apex where hairs may be dense, and at base where a ring of subappressed, fine, straight hairs
is sometimes present (the ring sometimes adhering to the base of the fruiting calyx tube); flesh gelatinous;
drying vitreous, reddish and translucent; epidermis texture + orange-peel p» often pruinose, scintillant,
sometimes glaucous, golden brown to reddish-brown, sometimes yellow. S d lil ge seg ,
texture rugulose-foveolate, 11-14 mm long, 6-7 mm radial depth, 4-5 mm EN
The range of this species includes the Yucatan Peninsula, the highlands of northern Chiapas, Tabasco,
southeast Veracruz, and northeast Guatemala (Fig. 49). The southern limit of its range seems to be near
Lake Izabal, Guatemala. Collections from *El Rancho" on the southern slopes of the Sierra de las Minas in
Guatemala are difficult to place, and are thought to represent hybrids between the nominate subspecies
of D. yucatanensis and Diospyros acapulcensis subsp. nicaraguensis. The species seems to be associated with
thin rocky soils throughout its range. Lundell (1937) described two species in the salicifolia complex for
the Yucatan Peninsula: Diospyros yucatanensis and D. spectabilis. We recognize only one species, Diospyros
yucatanensis, separated from other members of the complex based on differences in leaf shape and several
inflorescence size and shape parameters. Diospyros yucatanensis, even when sterile, is easily separable from
D. aequoris and D. acapulcensis. Its leaves are frequently lanceolate to lance-ovate, a feature not seen else-
where. On the abaxial surface of the leaf the hair bases are usually swollen and slightly viscid. This gives a
characteristic papillose appearance to older leaves that have lost their hair shafts. Characteristics of some
floral features also seem be of taxonomic significance, such as the longer corolla tube, differences in the
distribution of the external vestiture on the corolla tube, and a trend toward fourteen stamens rather than
twelve. However, it would be desirable to evaluate the constancy of these characters further when there is
access to larger samples of flowers. We have reduced D. spectabilis to subspecific status under D. yucatanensis
because it shares the aforementioned leaf characters and trends in floral morphology, and because interme-
diates between the taxa are common. Still, D. y. subsp. spectabilis warrants subspecific recognition because
throughout most of its range it can be differentiated from the nominate subspecies by its longer pedicels,
longer and distinctively shaped fruiting sepals, and generally narrower leaves. The ranges of the two taxa
are more or less parapatric.
KEY TO THE SUBSPECIES OF DIOSPYROS YUCATANENSIS IN MESOAMERICA
1. Fruiting pedicels 2-6(-8) mm long; calyx lobes 3-6(-8) mm long; lamina wide, tá in the ei haff,
usually 19-30 mm wide at 3/4 length; usually « 350 m, but occasionally to 140 subsp. yucatanensis
1086 Journal of the Botanical R h Institute of Texas 2(2)
1. Fruiting pedicels mostly 6-16 mm long; calyx lobes 4.5-10(-12) mm long; lamina narrow, espe-
cially in the upper-half, usually 12-23 mm wide 3/4 length; « 250 m subsp. spectabilis
4a. Diospyros yucatanensis Lundell subsp. spectabilis (Lundell) M.C. Provance, I. García & A.C. Sanders,
comb. et stat. nov. (Figs. 44, 45, ua 48 a-b). Basionym: Diospyros spectabilis Lundell, Publ. Carnegie Inst. Wash
478:218. 1937. Tee: MEXICO. CAMPECHE: Tuxpeña, “secon f Tuxpeña” [protologue], [18°27'N, 90*5"W, 213 m],
11 Oct 1931, C.L. Lundell 807 (4oLorvee: MICH; isorvpes: Kl, iL, MO, 2 as digital image!, WIS!
Trees or shrubs, 2-10(-14) m tall; trunk up to 13 cm in diameter; mature stems glabrous, rarely with
remnant hairs in the 2nd year, the young stems glabrate to sparsely villosulous, some hairs subappressed,
the epidermis viscid, pruinose, scintillant. Petioles (2—)3.5—8(-8.5) mm long, sometimes with a thin wing,
rugose, golden to dark reddish-brown, sometimes prui and scintillant, viscid, glabrous to sparsely pu-
bescent or wavy-hairy below, the hairs appressed to subappressed, ascending to subappressed villosulous
above. Lamina lanceolate or ovate to elliptic, sometimes oblanceolate or obovate, (52259-96(-123) mm
long, 19-38(-48) mm wide, length to width ratio (1.822.1-3.5(-4) : 1, lamina at 0.75 length 12-24(35)
mm wide, base acute, sometimes obtuse, short attenuate and abruptly decurrent on the petiole, margins
slightly revolute, apex acuminate or acute, sometimes rounded; lower lamina surface glabrate to tomen-
tulose, the cream, tawny, or reddish hairs readily deciduous, the hair bases swollen, giving a dense papil-
lose appearance to te oo pde to Cor) iaa the stomatal apparatus dense, usually opaque and
ly villosulous, epidermis viscid and shiny. Venation
prechidoaronous to arcolanguid; midrib below proce to densely reddish-brown hairy; lateral veins
finely raised Hoos ees Ps along the lamina-vein seam above; 3? veins barely raised below,
slightly imp 1-3-flowered cymes, tomentulose and sparsely subappressed
pubescent; peduncles 4-6 mm long; pedicels 2-2.5 mm long, bracts ovate to deltoid, 1-1.3 mm long, +
concave, readily deciduous. Male flowering calyx moderately to densely reddish-gold puberulent outside,
the hairs minute, + straight, not overlapping, tube 3-4 mm long, 3.5—4 mm wide, glabrous inside, or with
scattered hairs below the sinuses, lobes oblong-ovate to lance-ovate, 2-4 mm long, 3.5 mm wide, acute or
barely acuminate, minutely sericeous inside with a narrow intramarginal band; male corolla tube 6—7.5 mm
long, 3.2 mm wide, sericeous and either glabrous for 0.5-0.8 mm basally, or with 3 minute shield-shaped
glabrous basal regions; male corolla lobes lanceolate to lance-ovate, 3.5—5 mm long, 1-1.8 mm wide, the
hairs reddish-gold. Stamens 10—14; filaments adnate to the corolla « basally in two well-defined tiers,
2-2.5 mm long, red; anthers lanceolate to lance-ovate, 2.3-3 mm long. Female inflorescences 7-9 mm
long, bracts unknown. Female flowering calyx narrowly winged, densely reddish subappressed wavy-
hairy outside, sometimes with some straight ascending hairs, vestiture inside unknown tube 3.3 mm long,
4 mm wide, lobes acute, 3.3 mm long, 3.5 mm wide; corolla tube 4-5 mm long, + 3 mm wide, reddish
sericeous outside, otherwise unknown; corolla lobes lance-ovate, 3.5-4.5 mm long, 1.5 mm wide. Pistil
unknown except style remnants on immature fruit 3 in number. Fruiting pedicels 3—)5—14(-16) mm
long, glabrate to moderately hairy, viscid. Fruiting calyx tube 3-6(-7.5) mm long, upper 2/3 sparsely to
moderately hairy inside, lower 1/3 glabrous, lobes rapidly and markedly accrescent, broadly rounded to
oblong, (3.5)4-10(-12) mm long, (8-)9-14(-15) mm wide, rounded to barely acuminate to a blunt point,
spreading to reflexed, margins reflexed at the sinus and along the distal margin conferring the appearance
of a narrowly acute to acuminate point, sparsely to moderately hairy inside with a wide intramarginal band.
Fruit globose to depressed-globose, sometimes umbilicate, immature fruit sparsely reddish-brown hairy,
mature fruit glabrous except moderately hairy at the apex.
This northern sub rows on thin, rocky, cal in decid d subdecid forests,
and secondary growth a or these vegetation types. It occurs below 250 m elevation throughout the
Yucatan Peninsula, Mexico, (Fig. 49), in the states of Campeche, Quintana Roo, and Yucatan. lt also has
a limited range in northeast Guatemala and Belize. This taxon is separable from the nominate subspecies
by its longer fruiting pedicels, longer and slightly wider fruiting sepals, and slightly shorter and narrower
lamina. There seems to be a difference in lamina pubescence, but it is subtle, and of limited use since lamina
Provance et al., Diospyros salicifolia complex 1087
f LS
Ñ ie T
Vira NC
|
|
Fic. 44 Isotype of D. tabili iell (D. is Lundell sul tabilis (Lundell) M.C. Provance, I. García & A.C Sanders, comb. nov.), from
Tuxpeña, Campeche, Mexico, CL. Lundell 807 (WIS).
1088 I Infl Dak . aiD LI EE fTexas 2(2)
Fic. 45. Diospy yucat is sub bilis f M i, Yucatan, Mexico, C. Chan 6864 (CIQR)
Provance et al., Diospyros salicifolia complex 1089
Fic. 46. Di pyros yucat is subs t is f l P Itza, Guatemala (B. Wallnöfer & F.M. Tut-Tesucun 5941, MO).
J | of the Botanical R h Institute of Texas 2(2)
1090
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3 mm ——
Fi. 47. ipn yucatanensis Lundell. ink oR n from Tikal s P (c k Lundell 16257, IJ). A. Pistil. B. d sti C
Female co lla lok lyx. G-L. mie uatanens fom: Son sidro Tabasco
Mexico (E. Matuda 3378, NA). G. Male flower. H. Adaxi l surf. f male fl i lyx. | Male oroli tube ), K. Male
corolla lobe (abaxial side). L. Stamen. Me 0. subsp. y is from Río Dul Tzabal G k 39427 ' E). M. Stamen. N. Male corolla
Male flower. P-S. subsp Roo, Mexico (C. Gallardo et al. 2220, MO). P. Ad f f mal g ly Q. Male corolla
PE
tube. R. Male flower. S. Stamen.
1091
Provance et al., Diospyros salicifolia complex
VAS
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e
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1 mm). A. subsp. spectabilis (J.D. Dwyer 12747). B. subsp. spectabilis (Schultz
1530, UCR). C. subsp. yucatanensis (Cabrera 14830, MO). D. subsp. yucatanensis (C.L. Lundell 6807, ARIZ).
é
including midrib, of D
1092 J l of the Botanical R h Institute of Texas 2(2)
22° |
D. yucatanensis i p- TE I aN i
Q = ssp. yucatanensis | m e eec eh
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INTE 2008 Apr 20 16:45:08 | OMC- Martin Weinott
Fic. 49. Distribution of Diospyros yucatanensis.
vestiture tends to be deciduous in both, though the vestiture of this taxon seems to be even less persistent
than in the nominate subspecies.
Representative specimens. BELIZE. Corozal: 5 km NW of Chan Chen, 10 m, 16 Nov 1989, M.J. Balick 2189 (BRH). ORANGE Wark: from
mi 50, Northern Hwy. to new river toll bridge, 7 Jun 1974, J.D. Dwyer 12747 (UMO, WIS, IJ, BRH, MO). Stan Creek: All Pines, 0 m, 12
May 1976, G.R. Proctor 36205, 36207 (I. GUATEMALA. Peren: P.N. Río Azul, 0.5 km antes de llegar al Cedro Bosque, 20 Feb 2000,
M. Peña-Chocarro et al. 859 (BM); Dos Lagunas, 9 km E of village, 25 Nov 1960, E. Contreras 1651 (IJ, F). MEXICO. CAMPECHE. Mpio.
Calakmul: central chiclera Buenfil (Calakmul), 24 Feb 1994, J.M. Kantún 305 (UCAM). Mpio. Campeche: 1 km S de Samulá, 40 m, 1
May 2003, C. Gutiérrez B. 7761 (UCAM). Mpio. ce e carr. und ro Ba a Pital, 5 m, 29 Jan 1986, C.
Chan 6102 (CIQR). Mpio. Hecelchacán: carr. 5 km E de Rejón, 3 Feb 2002, G.G. Cabrera M. 12
(UCAM). Mpio. Hopelchén: Rancho El Carmen, near m 33-35 km S of Xmaben, 200 m, 13 Mar 1990, A.C. Sanders et al. 9752
(SD, UCR). Mpio. ae entre San Pedro y Santa Rita, 28 Oct 1997, P. Zamora C. & H. Uc C. 5890 (UCAM). QUINTANA Roo. Mpio.
Benito Juarez: Cancun, 25 Feb 2005, H. Kennedy & F. Ganders 5944 (UCR). Mpio. Cozumel: Isla Cozumel, 1 km NW del Faro de la
Punta Celerain, 22 Nov 1985, E. Cabrera et al. 9804 (MO). Mpio. Felipe Carrillo Puerto: 7 km E of Carrillo Puerto, 2 Nov 1984, D.
Neill et al. 5766 (MO). Mpio. Lazaro Cardenas: Yalahau Region, El Eden Reserve, ca. 30 mi NW of Cancún, 6 m, 26 Jun 2000, G.P.
Schultz et al. 1513 (MO, UCR), and about the same location 12 Jul 2000, G.P. Schultz et al. 1530 (UCR). Mpio. Othon P. Blanco: 500 m
S del Cenote en Bacalar, 15 May 1999, R. Durán et al. 3271A (MO). Mpio Solidaridad: X'cacel a X'cacelito, ca. 13 km N de Tulum, 14
Provance et al., Diospyros salicifolia complex 1093
Jun 1998, C. Gallardo et al. 2220 (MO). Yucatan. Mpio. Chemax: 3 km W de Chemax, 16 Jul 1985, E. & H. Cabrera 8868 (MO). Mpio.
Conkal: camino Conkal a Yaxkukul, 5 m, 4 Nov 1997, F. Tun, J.A. González-Iturbe 335 (SEL). Mpio. Halacho: Rancho El Naranjo, 12
m, 8 Sep 1988, L.M. Ortega T. & E. Mena P. 787 (UCR). Mpio Maxcanú: Chunchucmil, 5 km a la redonda, 11m, 4 Aug 1986, C. Chan
6864 (CIQR). Mpio. Muna: km 10 carr. Muna a Opichén, 50 m, 24 Sep 1984, C. Chan 3991 (UO). Mpio Oxkutzcab: zona arq. de Sayil,
25 Dec 1985, E. & H, Cabrera 10368 (CIQR, MO). Mpio. Tekax: Tekax, 18 Nov 1992, F. May 775 (CIQR). Mpio. Tixcacalcupul: S de
Xchak Hua, 13 Nov 1991, M. Méndez et al. 398 (MO). Mpio. Tizimín: 6 km W de Tizimín rumbo a Sucilá, 22 Sep 1996, R. Durán et
al. 2619 (MO). Mpio. Tunkas: carr. rumbo Quintana Roo pueblo, 23 m, 19 Jan 1982, E. Ucan et al. 1840 (NO). Mpio. Tzucacab: hills
above Tzucacab, 16 Feb 1983, D.A. White 271 (NO, JBSD). Mpio. Valladolid: Xuilub, 22 m, 24 Jun 1989, M.C. Sanchez 342 (UAGC).
Mpio. Yaxcaba: Tixcacaltuyub, 24 m, 29 Sep 1981, C. Vargas & P. Sima 540 (NO); S.E. Kancabconot [Kancabdzonot], [27 m], May 1917,
G.F. Gaumer 23862 (C, MO).
4b. Diospyros yucatanensis Lundell subsp. yucatanensis (Figs. 46, 47a—o, 48c-d).
Trees or shrubs, 3-15(-28) m tall; trunk up to e cm in diameter; stems usually glabrous, vestiture some-
times persisting into the 2nd year, the y densely orangish to reddish-brown tomentose
or wavy-hairy, sometimes with some ón o hairs, clavate glandular hairs sometimes present,
deciduous, leaving the dark reddish-brown epidermis viscid and shiny. Petioles (3.5—)4—6.5(-8) mm long,
+ rugose, golden or reddish-brown to very dark brown, viscid or glaucous, densely ascending pubescent
below, densely short wavy-hairy above. Lamina oblong ovate to oblanceolate, sometimes elliptic to obo-
vate, rarely lanceolate, (65270—106(-117) mm long, (20225-41(-47) mm wide, the length to width ratio
(1.8-)2.2-3(-3.4): 1, lamina at 0.75 length usually 19-30 mm wide, yellow-green in life (Lundell 1937), base
rounded to the petiole or abruptly short-decurrent on the petiole, margins slightly revolute, apex acuminate
or acute, sometimes rounded; lower lamina surface densely bright orange-red to tawny tomentulose, the
hairs sometimes deciduous, the a: bases Pen swollen, the nen reddish-brown to gray, the stomatal
lish wavy-hairy, the epidermis
apparatus dense, usually or
sometimes crystal-papillose. Venatici brochidedromaus to arcolanguid; midrib densely bright reddish-
brown tomentulose, with some longer, straight, sul 1 hairs; lateral veins mostly prominent below,
often bright reddish-brown velutinous, flush to finely raised above; 3° veins + raised below and obscured
by the vestiture, slightly impressed or obscure. Male inflorescences 1—5-flowered cymes, tomentulose to
wavy-hairy, the epidermis us dark EID dis 2-5 mm long; pedicels 1-3 mm long, bracts linear
lanceolate, 1-3 mm long. Male fl y golden to reddish tomentulose to short wavy-hairy
outside, with some longer hairs, tube 634. 2 5.5 mm long, 4.5—5.5 mm wide, the upper 1/2 sericeous,
lower 1/2 glabrous, lobes oblong-ovate to triangular, 3.5 mm long, 2.83.5 mm wide, subacute to barely
acuminate, sericeous inside with a somewhat narrow tomentulose intramarginal band; male corolla tube
(6.5-)9-10 mm long, 3-4 mm wide, sericeous except for 3 shield-shaped glabrous basal regions, 0.5-1.5
mm long, and some very minute wavy hairs just distal to these; male corolla lobes oblong-ovate, 3-5 mm
long, 1.5-2.3 mm wide, reddish to brownish sericeous outside. Stamens 12-14; filaments adnate + at the
base of the corolla, a few stamens sometimes inserted on the receptacle, sometimes geminate, (1.5—)2-3.5
mm long; anthers lanceolate to ovate or oblong-ovate, 3-4.5 mm long, sometimes rostrate or cuspidate, apex
thickened or granular. Female inflorescences 1-3 mm one pes ae (not seen by us), reported by
Lundell (1937) to be oblanceolate and 3mm long. Femal densely hairy, tube 2.5 mm long,
5 mm wide, sericeous except near the base where glabrous, lobes cue to obtuse, 6 mm long, 5.5 mm
wide, slightly rounded or tapering to a blunt point, sericeous inside with a wide tomentulose intramarginal
band; corolla tube 3 mm long, 2.5 mm wide, the upper 2/3 reddish sericeous outside, lower 1/3 with 3
overlapping shield-shaped glabrous regions; corolla lobes ovate, 4 mm long, 3 mm wide; ovary ovoid,
reddish sericeous; styles 3, spreading; stigmas bifid; staminodes 3, adnate to the corolla tube near the
base, anther-like. Fruiting pedicels 2-68) mm long, densely hairy. Fruiting calyx tube (33.5-5.5(=7)
mm long, sericeous except near the base, lobes widely oblong, 3-6(-8) mm long, (7-)9-12(-13) mm wide,
rounded or obtusely tapered to a blunt point, spreading, sericeous except near the base, the hairs of the
intramarginal band sparser than in the flowering calyx; epidermis puncticulate. Fruit globose, immature
fruit sparsely tawny to reddish-brown hairy, mature fruit + glabrous, but densely hairy near the apex.
1094 Journa | ofthe Botanical R h Institute of Texas 2(2)
The two collections Lundell used to describe male flowers were based on collections from Yucatan by
Gaumer; one from Kancabdzonot, the other without a specific locality. The specimen from Kancabdzonot
is representative of D. y. subsp. spectabilis, while the collection lacking a precise locality is inadequate for
determination to the subspecific level. The nominate subspecies of D. yucatanensis has a more southern
distribution than D. y. subsp. spectabilis (Fig. 49). It occurs in the eastern extreme of coastal Veracruz,
Tabasco, parts of Campeche, the northern highlands of Chiapas, Mexico, the mountainous parts of Belize,
and through a large part of northern Guatemala. We have seen three collections that are biogeographically
difficult to explain at this time, two from northern Quintana Roo, including one from the southern part of
Cozumel, and another from near Popolna, and a collection from near Sayil in the state of Yucatan. It grows on
thin or rocky soils in evergreen forest, deciduous forests, and secondary forest derived from these vegetation
types, usually between sea level and 360 m, but at elevations up to 700 m in Belize, and 1400 m in Chiapas,
Mexico. It is sometimes associated with cliffs. The distribution of this taxon apparently overlaps with both
D. acapulcensis subsp. veraecrucis and D. a. subsp. acapulcensis in northwest Chiapas. We have seen single
collections of hybrid material between D. yucatanensis and each of these subspecies from this region.
p specimens. BELIZE. Cavo: brecha al W del “Rancho Indio Suizo,” 8 mar 1985, E. Cabrera et al. 7719 (CIQR). ToLEDO: ca.
Medina Bank, 800-1200 ft, 23-27 Apr 1976, G.R. Proctor 35991 (MO); Mountain Pine Ridge, San Agustin, Jul-Aug 1936,
C.L. Lundell 6807 (ARIZ). GUATEMALA. Baja Veraraz: El Mago, 26 Oct 1968, E. Contreras 8015 (XAL, F). IzaBAL: Bay of Santo Tomás,
0-2 m, 13 Apr 1940, J.A. Steyermark 39366 (F); Puerto Mendez, bordering Sarstun River, 5 km, 9 Oct 1969, E. Contreras 9293 (MO); Río
Dulce, N side, between Livingston and 6 mi up river, 1-25 m, 14 Apr 1940, J.A. Steyermark 39427 (E). Peren: Lake Petén, 3 May 1933,
C.L. Lundell 3194 (F); Umgebung des Westufers des Lago Peten Itza, La Providencia” (= “La Trinidad”), 170 m, 17 Aug 1993, B. Wallnófer
& F.M. Tut-Tesucun 5941 (MO); San Miguel, 14 Dec 1967, E. Contreras 7316 (F, I); Tikal N.P., top of Temple Two, 8 Jul 1959, C.L. Lundell
16257 (IJ, F). MEXICO. Campecue. Mpio. Carmen: cerca de la Est. del Carmen, del ICMyL, UNAM, ca. 9 km E de Cd. Carmen, 23
Nov 1987, E. & H. Cabrera 14830 (MO, IEB); Colonia Nueva Coahuila, 10 m, 15 Jan 1982, C. Chan 1118 (XAL). Cmiaras. Mpio. Las
Margaritas: W side of Laguna Miramar, 350 m, 11 Feb 1973, D.E. Breedlove 33386 (CHAPA). Mpio. Ocosingo: 7 km E de Fl Carmen
Pataté, 900 m, 10 Apr 1991, E. Martínez S., C.H. Ramos & C. Eufrosio 24581 (XAL). Mpio. Ocozocoautla: Res. del Ocote, 380 m, 6 May
1983, J.I. Kod 10003 or Male: pe La Montaña, desviación a Sn. Juan Bautista, 20 m, 5 Jul 1999, C. Gutiérrez B. & J. Balam
6607 (UCAM). Mpio. S Fernando and Moravillas, 840—940 m, 15 Feb 1987, T.B. Croat & D.P. Hannon 65041
(MO). deni. Roo. Mpio. Cozumel: parte S de la isla, 4 m, 27 Aug 1985, C. Chan 5619 (CIQR). Mpio. Lazaro Cardenas: 2 km E
de Popolna, 12 Nov 1980, E. Cabrera & L. Cortez 256 (MO, CIQR). Mpio. Solidaridad: 12 km S del Ejido Laguna OM, 7 Aug 1980, O.
Téllez & E. Cabrera 3058 (MO). TABASCO. idt na ada. 9 Dec di A. nodo et al. 186 ae MO, ARIZ, XAL); San Isidro,
7-11 Jun 1939, E. Matuda 3378 (K, NA, CAS, MO). Mp Z v 1986, E. & H. Cabrera
12471 (MO). Mpio. Huimanguillo: aen 0 m, 2 Dec 1983, F. Ventura A. 20848 a XAL, 1B) ins Tenosique: Tenosique,
14-16 Jun 1939, E. Matuda 3401 (K, NA, MO, CAS). Veracruz. Mpio. Pajapan: | del ón, 0-5 m, 28 Mar 1980, L. Gutierrez 91
(XALU). Yucatan. Mpio. Oxkutzcab: 4 km W de Sayil, 20 m, 25 Oct 1984, A. Espejo et al. 1235 d
Diospyros yucatanensis Lundell var. e Lundell, Contr. Univ. Michigan Herb. 7:45. 1942.
Tere: MEXICO. Yucatan: Chichen Itzá off Kaua road in advanced deciduous forest [as on label], “along Kaua road, east of Chichen
Itzá, in advanced d forest” eco [20°40'W, 88°33'N, 29 m], Jun 11 E pons [specimen], 1938, C.L.
Lundell & Amelia A. Lundell 7509 (uotorvee: MICH; isorvegs: WTU!, LL, digital image!).
Lundell described this taxon based on its long male eun Leid The small amount of material we
in leaf and infl ubspecies, while the male flowering
have seen sl ilariti
peduncles (e.g., C. Chan 5110) seem rather long for each subspecies. We suspect this name is synonymous
with D. y. subsp. spectabilis, but would like to see material with fruit from Chichen Itzá before taking a
MM on the status of this taxon.
Addi examined: Mpio. Izamal: K'axek, rumbo a Tunkas, 20 m, 17 Apr 1985, E. Ucan et al. 3812 (CIQR). Mpio. Tinum:
Tinum, alrededor del cenote Xholak, en Chichen Itzá, 25 m, 25 May 1985, C. Chan 5110 (CIQR); Chichen Itzá, in second growth, C.L.
Lundell & A.A. Lundell 7434 (CAS, WIS).
MULTIVARIATE ANALYSES RESULTS AND DISCUSSION
A total of 23 leaf and fruiting inflorescence cha were used in the analyses (Table 1). From these, four
character subsets were derived from the matrix, with lamina length and lamina width treated as priority
characters, and R « 0.7 and R « 0.8 used as exclusion values (Table 2). A summary of the resulting biplots
Provance et al., Diospyros salicifolia complex
TasLe 2. Subsets of cf
1095
Subset 1. Lamina length priority, R « 0.7
Subset 3. Lamina width priority, R « 0.7
lamina length
lamina length : lamina width
lamina width at 0.75
lamina width at 0.25 : lamina width
lamina width at 0.75 : lamina width
lamina length : distance between four centermost veins
calyx tube length
lamina width
lamina length : lamina width
petiole lengt
lamina width at 0.25 : lamina width
lamina width at 0.75 : lamina width
lamina length : distance between four centermost veins
pedicel length
sepal width
epal length
calyx tube length
Subset 2. Lamina length priority, R « 0.8
Subset 4. Lamina width priority, R « 0.8
lamina length
lamina length : lamina width
petiole lengt
lamina width at 0.25
lamina width at 0.75
lamina length : lamina width at 0.25
lamina width at 0.25 : lamina width at 0.75
lamina width at 0.25 : lamina width
lamina width at 0.75 : lamina width
lamina length : distance between four centermost veins
lamina width
lamina length : lamina width
petiole lengt
lamina length : lamina width at 0.25
lamina width at 0.25 : lamina Hn at 0.75
lamina width at 0.25 : lamina
lamina width at 0.75 : lamina
lamina length : distance E four centermost veins
pedicel length
sepal width
pedicel length Sepal length
calyx tube length
sepal width : calyx tube length
calyx tube length
sepal width : calyx tube length
for PCA of the subsets is provided in Table 3. A biplot for subset 2 (leaf length given priority, R « 0.8)
is presented for demonstration purposes (Fig. 50). Overall, biplots supported the four species groups
identified by conventional taxonomic methods. In biplots of PC1 vs. PC2, Diospyros acapulcensis, D.
yucatanensis and D. aequoris each tended to form distinct clusters that overlapped to varying degrees
in the area where the PC-axes converged. Diospyros salicifolia collections typically formed a subcluster
in this region of overlap. Diospyros intricata formed a cluster most closely associated with D. aequoris,
but effectively separated from other taxa along the PC2-axis. The putative interspecific hybrid usu-
ally grouped with collections of D. acapulcensis. Separation of the taxa into groups along the PC3-
axis was usually poor, although D. intricata separated completely from the other taxa in subset 4.
The intermediate position of D. salicifolia in the biplots suggests a hybrid origin for D. salicifolia. 1f this
is the case, then based on geography, the likely parent taxa are D. aequoris, perhaps a form similar to subsp.
fürs a western ae. and the nominate subspecies of D. up lina we as no direct
y more similar
occurs between these taxa. I
to D. aequoris ron chutlensis than the nominate form of D. acapulcensis. However, leaf a of D.
salicifolia, such as lamina size, shape, and venation, usually group clearly with one putative parent or the
other, and often represent the extreme measurements for that character among the three taxa. In general,
specimens of D. salicifolia are rather consistent with respect to qualitative and quantitative characters. At
any rate, it seems most unlikely that specimens of D. salicifolia represent F1 hybrids of the possible parent
taxa, since neither of these species occurs with D. salicifolia, nor are the parent taxa currently believed to be
E
F
1096 J | of the Botanical R hi te of Texas 2(2)
Taste 3. Summary of biplots resulting from PCA of subsets 1—4.
Axes Separation
Subset 1.
PCI D. GCODMICEISI T is s from D. yucatanensis and D. aequoris with minor overlap.
PC2 D. intricat arated from all other taxa. D. aequoris is RN from D. yucatanensis with
slight eo: and is docs to separate from D. aca nsis.
Subset2. PCI D. id is nearly completely Paoa from D. E Diospyros aequoris is
S rated fr acapulcensis mall amounts of over rlap.
PC2 D. aequoris is ted I from D. acapulcensis with small amounts of overlap, and separated from
D. yucatanensis with minor overlap. D. intricata is effectively separated from all other taxa.
Subset 3.
PCI D. acapulcensis and D. yucatanensis are separated id minor overlap.
PC2 D. | from D | D. acapulcensis, but with some overlap.
D. intricata is s effectively separated from all iuis taxa.
Subset 4.
PCI D. acapulcensis and D. i completely separate
PC2 D from D. yucatanensis bl it with some SO up and tends to separate from
D. acapulcensis but With considerable overlap.
PC3 D. i | p y from other taxa.
As pem oia s Poln f thybrid origin for D. salicifoli ld not be add 1 sufficiently by
noh I g ; : f. g i his] yl } eee
c
ellite markers, ex — cross iizati i t
1 1 j f TTA 1; ff : 1
The D. intricata cluster was most oan with the D. ia cluster. Diospyros intricata is generally
thought to be endemic to the Cape Region of Baja California Sur and Isla Cerralvo in the Gulf of California
(We have not yet been able to verify material recently reported from an isolated upland locality on Isla
Tiburón [Wilder et al. 2008]). Consequently, it is separated from all other taxa of the salicifolia complex by
the Sea of Cortez. The range of D. intricata is more closely approached by D. aequoris (in Sinaloa) than D.
acapulcensis (in Jalisco). We have not seen any collections that appear to be intermediate between D. aequoris
and D. intricata.
The characters with the highest loadings for the rotated principal components (RPC) are reported in
Table 4. High loadings in the rotated factor patterns were most often seen in inflorescence size variables,
although leaf size characters had the highest loadings for subset 2. While inflorescences size characters
proved useful in explaining the variation in the data set, they were not much used during conventional
taxon delimitation (perhaps some in the case of D. yucatanensis). Leaf size and shape cl were useful
in explaining variation in the data as well, while inflorescence shape characters were relatively less impor-
tant. Conversely, some characters that were important during conventional taxon delimitation (ex hair
base morphology, leaf thickness, prominence of venation, etc.) were not considered during PCA. It should
be remembered that interpretation of the rotated factor patterns does have a subjective element. Still, it
appears significant that PCA found groups consistent with those found during conventional taxonomic
procedures, even though the RPC highlighted a somewhat different set of characters. It thus provides a
somewhat independent confirmation of the existence of the identified groups.
ACKNOWLEDGMENTS
We would like to express our appreciation to UCMEXUS and the UCR Center for Conservation Biology,
CIIDIR-IPN, Jiquilpan, Michoacan, and Dr. Giles Waines (UCR), for their support of taxonomic work
on Mesoamerican persimmons. We would also like to thank Paulo T. Sano and an anonymous reviewer
for their many useful comments on the manuscript. Access to microscopes was kindly made possible by
Martha L. Orozco-Cardenas at the UCR Plant Transformation Research Center. We thank Guy Nesom for
Provance et al., Diospyros salicifolia complex 1097
Taste 4. Characters with highest loadings for RPC of subsets 1—4.
RPC 1 RPC2 RPC3 RPC4 RPC5
Subset 1.
pdL LL SW LL:LW
SL 1W3/4 CL
Subset 2.
LL 1W1/4:1W3/4 pdL LL:LW1/4 SW
ptL LW1/4:LW SL CL
LW1/4 LW3/4:LW SW:CL
LW3/4
Subset 3.
pdL SW LW LL:LW
SL CL ptL LLY
Subset 4.
pal LW1/4:LW3/4 LL:LW SW LW
SL LW1/4:LW LL:LW 1/4 CL ptL
SW:CL LW3/4:LW
providing Latin diagnoses, Ed Plummer for technical support, Jose Aguilar Patiño (UAS), Francisco Santana-
Michel (ZEA), Antonio Machuca (CREG), Teresa B. Salvato (UCR), C. Conrado, Jeffrey Ross-Ibarra, and Randy
Richmeier for field assistance. We are grateful to Jeffrey Ross-Ibarra for his assistance with the statistical
investigations. Much appreciated specimen loans and images were provided by: AAU, ARIZ, B, BH, BM,
BRH, C, CAS, CHAPA, CIIDIR, CIMI, CIQR, DES, E, EBUM, F, FLAS, FR, GB, HUAA, IEB, JJ, ILL, JBSD,
K, LSU, MAD, MSCH, MO, MOU, NA, NLU, NO, P, SD, SEL, TEFH, TEX, UAGC, UC, UCAM, UCR, UCSB:
UMO, US, USF, UTEP, XAL, XALU, VT, WIS, WTU, ZEA.
APPENDIX 1
Rey bspecific intermediates and interspecific hybrid specimens:
Di isi with D. acapulcensis subsp. rivensis. COSTA RICA. Se
Cerro El Hacha, PN. Guanacaste, 1100 m, 15 Sep 1991, R.E. Espinoza 120 (MO, BM). San José: Rio si BRA
900 m, 7 Jan 1970, W.C. Burger & R.L. Liesner 7180 (MO). Puntarenas: Nicoya Peninsula, Curt, 2-75 m, 31 Aug 19 Rt p
Sanders et al. Wee (UCR); Canton de o n eun Rig Canarana; 700 m, 10 Aug 1993, J.F. inis 1576 (MO).
Di is. NICARAGUA. I S NE
de | la Isla de Momotombito, 200 m, 21 Oct 1979, M. Araquistain 371 (MO); Sector W de la Isla Momotombito, 150-200
m, 27 Jan 1980, M, Araquistain & P. Moreno 1061 (MO). Manacua: Campus of Escuela Americana, 200 m, 20 Oct 1976,
E Neill 1088 (MO):
Ia (C
l 1 i with subsp C Corpus, ca. 100
m, 0,5 Sep 1982, WD. Stevens 21802 (MO); EsteLí: Llano El Pozo, 18 km E de | Estelí, 1260-1300 m, 23 May 1983, P. Moreno
21380 (MO). Leon: Volcán Momotombo, 200 m, 29 Jan 1980, M. Araquistain & P Moreno 1082 (MO). MatacaLra: Valle
PueblaMidor 500 ı m, 2 c 1984, F Moreno 22958 (M O).
Di with D. acapulcensis subsp. veraecrucis. HONDURAS. Vare: Amapala,
Isla de la Tigre, fad W de ED Vanno pn m, 4 Apr 1981, C. Nelson & H. Martínez C. 7743 (mo, TEFH);
Di ith subsp. martineziana. MÉXICO. ^
NW de San Jeronimo, carretera Huetamo- Churumuco, um m, 20 Mar as da Soto N. et al. 2687 (MO, XAL, ILL).
odis aequoris subsp. rekoi dui with subsp O. Oaxaca: Mpio. Salinas Cruz: El Coyol,
W de Salina Cruz, 17 Mar 1 LP Ramamoorthy et al 2014 E te o Astata: Playa La Colorada,
1.8 E al W, 40 m, 30 Jul 2000, EH Er 3163 (IEB); Mpio. Tehuantepec: Cerro Lieza, 2 km NW de Tehuantepec, 1
_ Nov 1996, Cipriano! M if (MO).
ol |
F
Icatanensis. GUATEMALA. Peten: Río
Pasion, ‘Laguna San Juan Acul, 11 Feb 1964, C.L. Lundell 17946 (F). MEXICO. PE Mpio. pacis Entre basia
y Venustiano Carranza, 9 Aug 1996, P Zamora C. & D. Méndez D. 5348 (UCAM); Mpio. E A
8 m, 16 Nov 2003, C. Gutiérrez B. 8036 (UCAM); Mpio. Hopelchén: Xcochkax, 30 m, 9 Mar 1987, JN. Labat 1918 (IEB).
1098 Journal of the Botanical R h Institute of Texas 2(2)
o
o
o
B PC2
y
o T " x
*
x
o o o i x
o d d x x
% o
o x
o o ? "
o xo x
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o x
g o B ] P x x 2 x *
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oo e. Xx * o. 7 xy. * x
o 9o o o o s ig i x
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9o P o0 o x D ic x
e? o 2 s oF x * x x x
o 9o o NE o9 cu x yx PCT
oP y © e 2 LM" * x x Xem
o ES o 8 e x x
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o o x e^ x *x di Mu
o eS o? x e x
SP $ e 9
o 0% S e *% x
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i o Πo e A ES
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e e
o oo oo è . š E *
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9 *
o " e e
ee +
e
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s
$
+
? a O Diospyros acapulcensis
.
. 0 D.aequoris
e e
A D. intricata
A
. © D. salicifolia
X D. yucatanensis
. ^
Y D.acapulcensis x D. yucatanensis
Fic. 50. Biplot of PC1 vs. PC2 for subset 2 (leaf length given priority, R « 0.8).
Provance et al., Diospyros salicifolia complex 1099
Diospyros yucatanensis subsp. yucatanensis x D. acapulcensis subsp. acapulcensis. Mpio. Chiapa de Corzo: Río
Gr rijalva, 10 mn W of Chapa de Cae 500 m, iis May 1972, s Breedlove 25167 (MO).
ecrucis. MEXICO. CHIAPAS: Mpio Ocozocoautla
de Espinosa: 3 km N of Ocozocoautla along road to Mal Paso, 1 E 1973, D.E. Breedlove 32870 (CHAPA).
Diospyros yucatanensis subsp. yucatanensis x D. acapulcensis subsp. nicaraguensis. GUATEMALA. BAJA VERAPAZ:
Opp. El Rancho, 2300 ft, 12 Jan 1908, WA. Kellerman 7990 (F), and same location, 2500 ft, 5 Jan 1908, WA. Kellerman
7640 (F [2 sheets]).
APPENDIX 2
Specimens examined of Diospyros aff. i t PANAMA. Dann: El Real, 1-3 km S. of town, near sea level, 7 Jan 1975,
A. Gentry 13458 (MO, TEFH); El Real, 1 Mar 1972, A. Gentry 4508 (MO); Río Tuira, vicinity of El Real, 14 Jun 1959, WL. Stern et al.
455 (MO); same location, 1 Jul 1959, WL. Stern et al. 763 (MO); El Real, near airport, 13 Jul 1971, T.B. Croat & Porter 15467 (TEFH);
El Real, Quebrada Trapiche, [without date], J.A. Duke & N. Bristan 324 (MO).
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A TAXONOMIC REVISION OF CRATAEGUS SERIES LACRIMATAE (ROSACEAE)
J.B. Phipps and K.A. Dvorsky
Department of Biology
The University of Western Ontario
London, Ontario N6A 5B7, CANADA
johipps@uwo.ca
ABSTRACT
This pue reviews Ms ser. Lacrimatae (R J; a dd Re of ser. Flavae, sens. auctt. ENG 1500 qe were
1 ] United States flora. A
HEN with differentiation from allied series. Taehtybwo distinct and several subsidiary species are treated, broadly blowing spe-
d by Beadle, who had recognized 54 species. One i^ the E treated, C. lancei, is new to science. À key is
ndis to all taxa treated. All names np are id Im Mone: vided, 20 of the species are provided with
the fi ane in ps are — Most, if not = m es 1 of
line illustrations, mostly for t ti I
1 fh 1 f o4 da
Crataegus habitats. The series is marked by normally pendulous branching, EUN yellow to orange aU small leaves and a B degree
of glandularity.
, southeastern United States, xeromorphy
Key Wonps: Crataegus, ser. Lacrimatae, C. lancei sp. nov., Rosaceae
RESUMEN
E t ícul isa Crat I (R 1 d del Flavae, sens. auctt. S d
o + eo o
a een Bus el estudio de est p te, do abund de la flora del sureste de los rd Unidos. Se da una
1 : i la Aft, ; 1 dT :
iJ L P y val las subsi-
dias, siguiendo en] les el pto de especi blecido por Beadle, que ! ido 54 especies. Urade] peci
tratadas, C. lancei, es nueva para la ciencia. S t ] los | d Se tipifi | 1 se
aportan descripciones detalladas de las especies, ilustraciones de 20 Bas las e la me por Pa imera vez y los pri I
de deae l7ent as La inp oría, si no todas, de] p i ]
fueso aleuna d 1 litate d " Jon 1
e e I o E
1 J441 f "n A A 11 1 $ IS A A 1 dil m p:
I ya, hojas pequeñas y 8 &
INTRODUCTION
This paper continues an ongoing revision of denies Ji Crataegus, n poorly or imperfectly known
ries Aestivales , Parvifoli iae, Pulcher-
series, in the southeastern United States. The p
rimae, Bracteatae and Triflorae, and Apricae (Phipps (19882, b), Phipps and Dvorsky (2006, 2007), Phipps,
O'Kennon and Dvorsky (2006), Phipps, Lance and Dvorsky (2006)).
Series Lacrimatae is most similar to ser. Apricae but is part of what American authors during much
of the last century have included in Flavae, a problem which is discussed in Phipps and Dvorsky (2007).
The two series Lacrimatae and Apricae are differentiated as follows: whereas all but two dwarf species of
ser. Lacrimatae have more or less pendulous terminal branches, at least at maturity, and all have distinctly
zig-zag young twigs, 20 stamens and white to cream anthers, members of series Apricae lack conspicuously
pendant branches, have unbent or only slightly bent nodes, most commonly 10 stamens and often have pink
to purple anthers. In addition, ser. Lacrimatae species usually have less venous leaves, their veins spreading
at a narrower angle. It would also appear to be the case that a good many members of ser. Apricae are less
xeromorpic than any in ser. Lacrimatae. In herbaria confusion of smaller-leaved species of ser. Lacrimatae
with C. uniflora (ser. Parvifoliae) is not uncommon, but more or less straight, non-pendulous twigs are defini-
tive for ser. Parvifoliae. In addition, the number of styles and nutlets is usually greater and the calyx lobes
are normally much larger and much more deeply dissected in the latter series. Also see discussion under
ser. Parvifoliae (Phipps & Dvorsky 20063).
J, Bot. Res. Inst. Texas 2(2): 1101 — 1162. 2008
1102 J l of the Botanical R h Institute of Texas 2(2)
There are two rather well defined groups within ser. Lacrimatae which are here described as subseries.
These differ primarily in leaf shape, size, and texture, in twig thickness, and in fruit size, shape, and color.
The first subseries, Tenues (spp. 1-8a), consists of more slender plants, with shorter and finer thorns, gener-
ally smaller and thinner leaves, often smaller flowers, smaller fruit and generally thinner twigs. The second
subseries, Robustae (spp. 9-22), consists of sturdier plants, with stouter and generally longer thorns, larger
leaves, often larger flowers, usually larger fruit and generally thicker twigs. Many of the species of both
these groups are densely white-hairy, typical xeromorphs, though a few are glabrous. Nearly all have yellow
(sometimes red-cheeked) to orange or ruddy fruit.
No species names correctly attributable to ser. Lacrimatae appeared before 1900 when Beadle com-
menced his studies in this large group and Ashe described a few species. However, it is to Beadle that credit
must be given for being the first to observe and comprehensively record their variation. Beadle eventually
(1903) recognized 54 species in nine groups that might equate to subseries. Nearly all later floristic workers
have shown a reluctance to get to grips with Beadle’s demonstration of this large amount of variation, and
drastically reduced Beadle’s total of species without discussion, thus, none of these treatments represents a
critical re-examination of the group. Murrill (1942), however, in a local study of Florida hawthorns, added
several species which either seem to represent rare, local forms, or can be synonymized, and none of them
is upheld here. W.W. Ashe also made quite numerous collections, particularly in North Carolina, but de-
scribed relatively few taxa in this group, all of which are either synonymized here or untypifiable. In spite
of the attention of these three authors, one of the species treated here, C. lancei, had escaped notice (or else
perhaps equates to an untypifiable Ashe name), and is thus new to science.
In the view presented here, Beadle did name too many species, which often differ from one another by
only slight differences in fruit shape or tomentum. However, the opposite extreme of attempting to lump all
or most of the variation into a very few species, if done with an eye to objectivity rather than procrustean
systematics, is a very difficult exercise. Nevertheless, that is what important floras of the region since Beadle’s
time have habitually done, for example, Tidestrom (1933), Radford et al. (1968) and Wunderlin and Hansen
(2003), who typically used some or all of C. flava (Aiton 1789), C. lacrimata (Small, 1901), C. floridana (Sarg.
1902) and C. michauxii (Pers. 1806). These four species names will now be considered.
Crataegus flava has understandably found favor since it is a a old name and became the ype of the
long tablished ies Flavae (Loudon) Rehder. Nevertheless, careful e nation of i shows
it to be a particularly distinct member of series Flavae as previously delimited (Phipps 19882). ENT it proves
so different from the remaining Flavae that it is now transferred to ser. Intricatae (see discussion in Phipps
& Dvorsky 2007). Faxon's illustration of C. flava (something like C. condigna Beadle or C. colonica Beadle is
shown) in Sargent's Silva vol. IV, of 1890, was no doubt influential as the source of the misapplication of C.
flava in a generic sense for ser. Lacrimatae species. Although Sargent (1902) later issued a corrected descrip-
tion and illustration of C. flava no attention appeared to be given to it. The 1902 correction is of a plant far
more like true C. flava though it differs in stamen number. Crataegus lacrimata, on the other hand, is one of
the most typical, yet distinct and easily recognized members of ser. Lacrimatae and it is difficult to see why
its interpretation should be widened. In any case, there are earlier names available from the series. Sargent's
1902 account of C. floridana has no illustration and explains that the 1890 ‘flava’ illustration should apply
to it. This would appear to have resolved the confusion, at least from Sargent's perspective. However, unac-
countably, it did not and instead it merely provided a rather imprecisely illustrated C. floridana for the ranks
of later floristic accounts. Crataegus florid ly understood, was nonetheless another poor choice for
one-to-few species accounts of the groups since while there is no question that it is a typical member of ser.
Lacrimatae, it is not only a less clear-cut species than C. lacrimata but as many as 11 species accepted here
all have earlier names, and therefore the name lacks appropriate priority if combined with any of them.
Particular probl d the much used name C. michauxii which many floras, starting with Beadle
(1903), have treated as a member of the southeastern United States flora, where it is routinely construed to
be a member of ser. Lacrimatae (under different group names) more or less as understood here. Persoon's
name C. michauxii was a new one for C. glandulosa of A. Michaux (1803), itself homonymous to C. glandulosa
AL: in. L D D» Efrat H I j + 1103
of Moench (1785) and of Georgi (1776) which are Eurasian. Persoon’s protologue gives: “Habitat in Carol.
Super.”; “spinosis...stipulis, calycibus imaque fol. parte glandulosis, fol. subrotund. breviter lobatis subpu-
bescentibus; fl. subsessilis,” which is of little diagnostic help. Moreover, the name C. michauxii is rejected
for use in this paper because the type specimen at P of Michaux's C. glandulosa is simply a piece from a
vigorously growing extension shoot of an unidentifiable species. Any attempt to effect an accurate modern
identification from such defective material seems impossible unless at some future date it were to be done
by molecular methodology. One thus has to wonder about Beadle's rationale in resuscitating the name C.
michauxii when his species concepts were narrow. Of course, Beadle's (op. cit.) usage itself is probably at least
approximately interpretable (but I have seen no material named this in Beadle's hand). Subsequent authors
simply did not investigate the nomenclatural problem, preferring their own interpretation of Beadle's usage.
A practicable solution might appear to be to epitypify C. glandulosa Michx. on excellent modern reproductive
material that is a presumed best match for Beadle's usage of C. michauxii. However, the later applications
of this name are too inconsistent or vague to make such an exercise worthwhile. Thus, those authors who
include all or most Floridian members of series Lacrimatae and Apricae under C. michauxii ignore the type
issue. However, if ser. Lacrimatae were to be treated as a single species, the earliest available unquestionably
suitable names become C. alabamensis Beadle and C. senta Beadle, both described in 1900.
This study is based on the examination of well over 1,400 herbarium specimens of ser. Lacrimatae
including borrowings from 35 herbaria and fairly large collections of duplicates received from R. Lance.
It has greatly helped this study to have made my own (JBP) considerable collections of ser. Lacrimatae, in
many cases with matching fruit and flower specimens. Such fieldwork also enables one to observe habit
and color differences that are not evident in herbarium specimens as well as, of course, observing many
times more specimens than actually collected. This level of sampling greatly facilitated identifying repeat-
ing patterns of variation, sometimes with surprising ease. The repeating patterns are matched against type
material, nearly always Beadle's, which is generally of high quality. There were relatively few good quality
specimens that are intermediate between accepted taxa. Where "annotations" are referred to under several
species discussions this alludes to JBP annotations made prior to final decisions on the correct name and
these provisional names were applied because the size and complexity of the project meant that it became
inappropriate to retain loans until final decisions could be made.
Some of the characteristics found useful in the taxonomy of ser. Lacrimatae include thorn frequency
and size, leaf general shape and size, tomentum of pedicel, hypanthium and fruit, flower and fruit size,
margination of leaf (manner and extent to which lobed and toothed). A number of species possess long-
petiolate leaves that can flutter in the wind or droop (become "floppy") in calm air. These characteristics are
so helpful when learning taxa that they are used routinely in our keys and descriptions even though they
will not be evident in herbarium specimens. Also, there is some indication that the form of extension shoot
leaves in this series, usually passed over in Crataegus taxonomy due to its frequently extreme plasticity as
llas shortage of identifiable herbarium examples, may prove of taxonomic value when better documented.
For example, C. florens Beadle and C. dolosa Beadle (a form of C. integra (Nash) Beadle) apparently consis-
tently possess more or less circular, lobeless extension shoot leaves. As with all modern specialist Crataegus
taxonomy, short-shoot leaf-shape has been found particularly valuable and we have attempted to articulate
this more precisely than Beadle did, both in words and with the numerous line illustrations.
Beadle's keys give somewhat imprecise descriptions of the leaf shapes which are often so diagnostic in
Crataegus, and this makes positive identification from the keys a challenge, to put it mildly. However, loans
of critical types, principally from A, plus easy access to the digitized type collections at US and NY (where
the type specimens of many Beadle names are unfortunately attributed to W.W. Eggleston rather than the
real collector) greatly helped in making critical determinations. However, names due to Ashe have often
presented more or less insuperable typification difficulties of the kind previously discussed in Phipps et al.
(20062). Fortunately, it is possible to confidently match all the here-accepted names in Lacrimatae against
type material without needing to apply any names of Ashe or Murrill.
1104 j | of the Botanical R h institute of Texas 2(2)
It became clear early in this study that Beadle was the only previous worker who had made any sense
of the great variation in ser. Lacrimatae. The 22 species recognized here nevertheless represent a substantial
reduction from the 54 species found in Beadle's final account (1903) without being procrustean. As in all
the first author's taxonomies that use a similar approach, the complete lack of or at least, low frequency of,
intermediates is a critical consideration in coming to decisions on taxon delimitation. The highly dissected
variation patterns encountered are thus at pesent most easily treated as rather narrow species some of which
may eventually prove to equate to apomictic microspecies, although several other species turn out to be
quite variable, e.g., C. crocea and C. condigna. However, nothing is yet known about the breeding systems
or chromosome numbers of ser. Lacrimatae, the count for C. agrestina Beadle in Talent et al. (2005) actually
representing a ser. Pulcherrimae individual with correct usage of this name so such valuable interpretive
information cannot be used.
Series Lacrimatae has a similar geographical range to ser. Apricae, but is far more abundant (Fig. 1), with
1458 records in our database compared to 274 for the former series. This far greater abundance of material
made most taxonomic decisions easier than for the treatment of ser. Apricae (Phipps & Dvorsky 2007), the
predecessor to this paper.
This paper provides the first revision of the species included by Beadle (1902, 1903) in his subgroups
Michauxianae, Integrae, Dentaiae, Attritae, Sentae, Anisophyllae, Colonicae, Recurvae and Lepidae plus elements
of Visendae. (Crataegus fruticosa Sarg. 1902, repeatedly collected around the time of its description and not
seen since, is perhaps a hybrid between a member of ser. Lacrimatae m ser. Aestivales and is excluded.)
Most illustrations are firsts and Crataegus meridiana is the only widely ies not illustrated, both
because it is somewhat obscure to characterize and because of lack of ied. doen presumptively typical
material. Most maps are also firsts and are presented on a county frequency basis. Most of the illustrations
and all the maps are firsts. Complete descriptions are provided for all main species. More obscure forms are
treated as subordinate taxa, usually without illustration or map. Often these have abbreviated descriptions.
All taxa and synonyms used are typified. Synonymy, however, is somewhat incomplete (as it is for most
North American series), here largely due to problems with Ashe names. As in other recent publications in
this series, Ken Dvorsky is responsible for creating the maps. An appendix of cited specimens is provided.
TAXONOMIC PART
Series Lacrimatae J.B. Phipps, Taxon 37:113. 1988. Tree species: Crataegus lacrimata Small, Torreya 1:97. 1901.
Shrubs, sometimes very small, or small trees; bark on trunks black or dark gray and highly corrugated (pre-
sumably fire-resistant), unlike that of most other Crataegus series; ultimate branches pendulous, particularly
in the very slender-twigged species, also zi the nodes, often purple-brown at 1 year; thorns mostly 1—5
cm long, none to numerous, x straight, uuu. slender, very dark at 1 year. Leaves deciduous, petioles very
short to long (10-40% length blade); blades small (1-4 cm long), ovate to oblong in general shape, usually
shallowly lobed; margins entire to toothed, very glandular; lateral veins 1—5/side, exiting mainly or wholly
in the apical part of the blade; tomentose, at least when young in most species. Inflorescences 1-4(-7)-flow-
ered; the branches in many species densely appressed white-pubescent, bearing small, deciduous, linear,
membranous, gland-margined bracteoles with generally high persistence through anthesis. Flowers 10—20
mm diam.; hypanthium often tomentose; calyx-lobes + narrow-triangular, entire to glandular-dentate; petals
+ circular or rarely elliptic, white; stamens 20, anthers usually white to cream; styles 3—5. Fruit 6-12 mm
diam., pyriform to orbicular, yellow through orange and copper to red, one face often colored most brightly,
smooth or with scattered hairs; calyx-lobes usually reflexed; nutlets 3—5, dorsally grooved, sides plane. Most
species ripen their fruit in August, before sympatric hawthorns.
Series Lacrimatae constitutes a characteristic and frequently abundant element of the sand plains,
scrubby areas and pinelands of the southeastern United States coastal plain and piedmont from Louisiana
east of the Mississippi, south to central Florida thence north on both sides of the Appalachians and extreme
southeastern Virginia. Treated here are 22 usually fairly narrowly defined species and six more obscure
1105
Tonig ani
diry
| ‘Ore eens >
RAN
á i A VE d
MEO
LN
24
gms 25 -80
IN 61 - 100
800 Miles
iii o
y Mumber of Records
Fic. 1. County level distribution map of collated species.
entities indicated by adding a subscript to the number of a similar species. This compares to the 54 species
recognized by Beadle (1902, 1903)
LII (leaf incision index or IFI in Latin) refers to the degree to which lobed leaves are dissected and
ranges from 0% (without lobes) to 100% (cut to the midvein). Different sinuses may vary in LII and the value
given refers to a typical value for the deepest sinus.
KEY TO SERIES LACRIMATAE
ile Lial4m Aft. | | i H las i} ho hal J]
HI
ACNACI
f movement
in slight wind, else dwarf shrubs 0.5-1 m tall at maturity; twigs slender, often only 1.5 mm thick at about 5 cm
m the tip; thorns fine, usually 1-4 cm long; blades usually 1-4 cm long, HELM (eIDopinig or fluttering
many species; fruit usually small, 6-8 mm diam., often yellow r. Tenues).
2 me blades 1—2(-3) cm long, as broad as long to pes longer than broad (1-1.4: 1.
. Dense twiggy shrubs seldom reaching 2 m tall, not bna lacrimate except sometimes in larger
specimens; often flowering at 0.3 m tall; leaves gl emi-evergreen; blades « 2 cm long, + isodia-
metric to ta oo TUNES HOF lobed to obscurely lobed across the i end, Ai margins
| leaf-marginal teeth; eaves very
much like short- shoot leaves but larger and shallowly koa
a rrimean (subse
callo CU lc
1. C. lepida
Uu
. Larger shrubs or small trees 2-7 m tall at maturity, n ; flower
ing at less than 1 m tall; ees not agony deciduous in fall; blades1-2(- 3) cm long, nearly always
broadestin apical a , USI jally d 1-2 but al
to + +, į | y per LE - 1 |. J +!
{7
TOC ay
sion par
eeply and somewhat irregularly lob
tip, thei large obtuse distal end Bang quite
d ed
2. Leaf-blades 2-5 cm long, significantly longer than broad (1.4—2:1).
4. Inflorescence branches and hypanthium + glabrous.
2. C. egens
1106
AV Ul idi Yi
5. Leaf- dad 1-1.75 cm long, teeth + regular; petals 5-7 mm long; ripe fruit yellow with a red
chee
8. Leaf-blades proportionately broa
spathulate, 1.5-2.0:1; tips Rien to flattish, at most the end very flat triangular or Mie
t
e.
9. Leaves proportionately longer, + cuneate-spathulate 5. C. crocea
9. Leaves proportionately shorter, broad rhombobovate to + broad ellipti
7. Leaf-blades narrowly rhombic in general shape, margins nearly entire
f Texas 2(2)
3. C. lacrimata
5. Leaf- im 2.25- 4.00 cm long: teeth larger and more irregular; petals 7-9 mm long; ripe ip deep
or dull orange-copper 3a. C. sp. cf. C. lacrimata
4. ibo. branches and hypanthium pubescent to tomentose.
. Leaf margins entire to subentire, or, if toothed, teeth very fi d ly restricted to terminal
part of leaf.
7. Leaf-blades oblong-obovate to very narrowly elliptic in general shape, margins entire to
ii serrato-crenulate across the tips, sometimes with occasional small sharp teeth on the
es.
d Leaf-blades narrow elliptic to narrow o i B 3:1; ¡Ups i 5. e crocea (typical form)
(villaris form)
(incana form)
8a. C.sp
6. Leaf du distinctly, though often shortly, lobed and/or toothed.
10. didis wiggy 5 shrubs seldom reaching 2 m tall, not notably lacrimate except sometimes in
lar mens; often flowering at 0.3 m tall; leaf-blades cuneate to narrow elliptic, often
o parallel: sided margins sharply e i ot euh < a cm meno.
10. Larger shrubs or small trees 2-7 m tall at matur
flowering at less than 1m al leaf blades 1 5-4c cm long, more re obtrullate i in general shape, +
lobed apically, g t piae e
11. Leaf isl ded j | |, well-defined or sometimes
obscure or obtuse; petioles S long (25-5096 ae of blades thin, capable of
bun. in wind; thorns few or none, + slender, ca. 2 cm lon
eaf apices obtuse to flattish in general shape; c obscure or lacking young or
| dentate
more noticeable at maturity when small, subacut g y crenate-dentate
4. C. munda
5a. C. vicana
12. Leaf aie rounded to triangular in general shape, with well-defined subterminal
+/- acute lobes
13. BS usually 1 -3 cm long, +/- stiff to fluttering in wing; | les 5-10 mm long.
14. LII on broader leaves < 20%; leaves at anthesis larger than fl tensi
shoot leaves unlobed or lobed much less than LII 80%
14. Lil on broader short-shoot leaves ca. 2096; leaves at anthesis about size of
flowers; larger extension-shoot leaves deeply lobed (LII to 8096)
6. C. quaesita
13. Blades often reaching 3-4 cm, though normally some shorter, larger able to flut-
6a. C.
sp. aff. resima
ter in a wind; petioles 10-20 mm long 7. C. floridana
1 1. I J Load g | LL L | | |! L | +
(< 25% length of! lade); blades stiff and fl ing in wind; very thorny, tl t stout
3-5 cm long 8. C. invicta
1. Habit more robust and stocky though ultimate twigs usually x pendulous in mature specimens; twigs
thicker, at least 1.5 mm and often 2-3 mm wide at about 5 cm from the tip; thorns, if present, normally rela-
tively stout, 2-5 cm long; blades often rather thick and stiff, commonly 2.5-4.0 cm long; fruit usually larger,
8-12 mm diam., often cooper cuore at maturity (subser. Robustae).
15. Leaves in most specimens without lobes, but when present, these subterminal, few and obscure and
very short-triangular or merely undulate, margins toothed or not.
16. Leaf marginal teeth, if any, merely denticulate.
7. Thorns generally absent; most leaf-blades 2.5-4 cm long; flowers 15-20 mm diam; fruit 10-12
mm diam.
18. Leaf-blades 2-4 cm long, many at least twice as long as b
tip obtuse to cuspidate, sides evenly tapered from | just below broadest point t to cien
with petiole; petiole 10-15 mm long; leaves floppy' in na
16. C. lassa
18. Leaf-blades often 2.0-3.5 cm long, 1.2-1.8 x as long a xe oad, usually proportionately
broader than above (but occasionally proportionately narrower), general shape somewhat
Lacrimat 1107
variable, + obovate, broad rhombobovate to + broad elliptic, or even suborbiculate; tip
varied in form, acute to subcuspidate, alternatively truncate-obtuse; and sometimes ob-
scurely undulately lobed around broadest point; petiole 3-8 mm long, leaves stiff 11. C. integra
17. Thorns long, numerous; leaf-blades 2-2.5 cm long; flowers 12-15 mm diam; fruit 6-9 m
diam
5 C. colonica
16. Margins with well-defined teeth.
19. Hypanthium and pedicels tomentose.
20. Margins with fine regular teeth; leaves lacking lobes.
21. Leaf-blades broad-elliptic to narrow obovate or obovate in general outline, obtuse to
somewhat cuspidate across the tip; inflorescences 2-7-flowered; flowers 14-25 mm
diam.; thorns absent or scarce, if present, 20-30 mm long.
22. Leaf-blades narrow obovate to broad elliptic in general shape; apex rounded to
broadly acute; flowers 13-20 mm diam
22. Leaf- hens broad- ids to cuneate, + flattened at apex or slightly NY
9. C. condigna
flow ps 25 mm is C. alabamensis
21. Leaf-blad hombi tl Il
is tip; c 1-3-flowered; flowers ca. 12 mm diam; thorns abundant, 35-50
m lon 13. C. pexa
20. on uns coarse, irregular teeth, some leaves with one shallow lobe per side 15b. C. pulla
ent.
19. E nd and pedices n to Tu -pubesc
ith 2- "ed of lateral veins; twigs nidum"
| nutlets 2-3 . C. teres
eaf FT argit al
inflorescence h h I | } |
23. EX iid sd code and/or sharp, dentate; ids with 1-4 pairs of lateral veins; many
pubescent or subglabrous; styles and
nutlets 3—5.
24. pM quite large, 2.0-4.5 cm long; pairs of lateral veins 2 —4; inflorescence
branches thinly to moderately pilose-pubescen 15
24. iden oe 1.5-3.5 cm long; pairs of lateral veins 1-2; inflorescence branches
subglabro 15a. C. attrita
gla
15. At least 40% ice blades Fev lobed, but sometimes
25. Blades 2-5 cm long at maturity; most leaves + narrow in dn genetal shape, many >1.75:1, elliptic to
. C. florens
Il ft also toothed.
narrow-obovate or narrow-obtrullat
hu Mate leaf-blades 30-50 mm oe 2 Il sul inal lobes per side; flowers 17-20
wide 17. C. lanata
26. es d blades 15-30 mm ong, uu lones well- o and + sharp (except sometimes in
Cla where mat -16 mm wide
27. Loles fedey absent) 0-1 per side, small, subacute teeth crenate to crenate- as
lateral veins 1—3 per side; inflorescences 3-7-flowered 0. C. lancei
27. Lobes (present on all leaves) 1-2 d side, and Mi well-defined, sharp; lateral veins =
per side; inflorescences 2-4-flowered 21. C. senta
25. ien 1.5-4 cm long at maturity; Munda or more leaves broader in general shape, mainly ca.
1.6:1.
E Bades usually broadest towards the tip, + broad-obtrullate to obtrullate-rhombic or obovate
d shape, rather longer than wide, lobes (0—)1—-3 per side subterminal, distinct; anthers
2 po stiff, often persistently lanate; lobes 1-2 per side, subterminal to lateral,
8. C. furtiva
generally rather obscure
29. Leaf-blades stiff or floppy, persistently lanate or not; at least one prominent subterminal
lobe per side.
30. Inflorescences 2-4-fl i: ll ining f [ ,lobes 1-2 per
side, sharp at E when young teeth sharp not.
31. Leaf-blad teing bes 1-2 per side; marginal teeth
anal oa seule 19. C. meridiana
31 broad-obovate i | shape; lobes, usually 2 per side;
to broad y I!
sica teeth very sharp 2. C. dispar
30. | » 37 A Hg Int tl ] Ily1( 0) per side, broad but
23. C. exilis
acute; marginal teeth very small, sharp
1108 j l of the Botanical R h Institute of Texas 2(2)
28. Blades broadest in the middle, rhombic in general shape, a little longer than wide, with one
distinct + acute lobe per side; anthers pink to purple [C. egregia (ser. Apricae)]
Subseries Tenues J.B. Phipps, subser. nov. (spp. 1-8). TYPE SPECIES: C. recurva Beadle.
Habitus vulgo aliquantum tenuis cum ramulis ultimis pees gracilibus pendulisque et capacibus motus in vento exiguo, duobus
speciebus fruticosis nanis 0.5-2.0 ltis i li graciles, saepe solum 1.5 mm crassi ca. 5 cm ab apice; spinae
graciles, vulgo 1-4 cm longae; laminae vulgo 1-4 cm ne AN tenues, demissae vel tremaefactae in vento levissimo in
ibus speciebus; fructus vulgo parvi 6-8 mm diametro, pyriformes vel late-ellipsoidei vel interdum suglobosi, saepe flavi cum
carmesinis genis.
Habit usually rather slender with ultimate branches particularly fine and pendulous and capable of move-
ment in slight wind except in two dwarf shrubby species 0.5-2.0 m tall at maturity; twigs slender, often
only 1.5 mm thick at about 5 cm from the tip; thorns fine, usually 1-4 cm long; blades usually 1-4 cm long,
rather thin, drooping or fluttering in many species; fruit usually small, 6-8 mm diam., pyriform to broad
ellipsoid, occasionally subglobose, often yellow with a crimson cheek.
1. Crataegus lepida Beadle, Biltmore Bot. Stud. 1:36. 1901. (Fig. 2). Tre: U.S.A. Georcia. Ware Co.: Waycross, 24 Aug
1901, C.D. Beadle 3041 (Lectotype, selected here, US).
Shrubs, often less than 1 m tall but sometimes reaching 2 m, dense and twiggy, not notably lacrimate except
sometimes in larger specimens, capable of flowering at 0.3 m tall; bark of trunk scaly, dark grey; young
twigs A repe pubescent, 1-year old twigs brown, zig-zag at nodes; thorns 0.6—2 cm long, fine, straight.
Leaves late deci wintergreen; petioles 25-40% length of leaves, pubescent, glandular; blades 0.6-1.5
cm long, broadly obovate tox isodiametric in general shape; tip very obtuse or with a slight apiculus, base
abruptly contracted into the winged upper portion of the petiole; sometimes with extremely shallow lobes
across apical third; margins minutely glandular-crenate to sometimes with conspicuous distal glandular-
scalloped teeth distally; venation craspedodromous, 1-2 main lateral veins per side; surfaces very slightly
1
appressed-pilose above when young, more particularly so on the veins, + pubescent below, especially on the
veins; coriaceous and often somewhat glossy; stipules herbaceous, falcate, gland-margined. Inflorescences
1-3-flowered; branches appressed-pubescent, bearing deciduous, linear, membranous, gland-margined
bracteoles. Flowers ca. 12 mm diam.; hypanthium externally pubescent; calyx-lobes abaxially pubescent,
triangular, margins + entire; petals + circular, white; stamens 20, anthers ivory; styles 3-4. Fruit 8-15 mm
diam., slightly longer than broad, + pubescent, yellow-green to yellow or orange when ripe, black gland-
dotted; calyx-lobes recurved or broken off, nutlets ca. 3, dorsally grooved, sides plane.
Common name.—Dwarf Hawthorn.
Distribution (Fig. 3).—This is a scarce species occurring in the southeastern United States in scattered
localities from central Florida to Georgia and South Carolina where it is found in dwarf scrub. Remarkably,
a sterile specimen, Thomas 57507, from Louisiana, appears to be this species.
In central Florida plants flower and begin their extension growth very late, in May, long after growth
of other nearby plants has started, however, further north, anthesis about the second week of April seems
more normal for this species. The former are the most wintergreen populations. Particularly small-leaved
specimens with numerous, small, sharp marginal teeth may represent C. garrettii of Murrill.
This is a dwarf semi-evergreen shrub, very attractive, with dark, glossy, often suborbiculate leaves that
are very striking in the field. Larger specimens might be confused with C. egens.
2. Crataegus egens Beadle, Biltmore Bot. Stud. 1:85. 1902. (Fig. 4). Twe: U.S.A. Fronma. Liberty Co.: Bristol, 1 Apr
1901, T.G. Harbison 4037 (LectorvrE, selected I GH; IsOTYPES: A, US).
? = Crataegus cirrata Beadle, Biltmore Bot. Stud. 1:101. 1902. Tipe: U.S.A. ALABAMA. Russell Co.: Girard, no date, C.D. Beadle 4122 (1ec-
TOTYPE, selected here: U
Comment.—Beadle 4123, also Gn Girard, was collected 9 Apr 1910.
Shrubs or small trees to 5 m tall, slender, with pendulous branches, seldom flowering at less than 1 m
tall; extending shoots white-canescent; 1-year old purple-brown, older darker or grayer; unarmed or with
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1110 J | of the Botanical R h Institute of Texas 2(2)
i Y DIAL:
[s DAT Neb i
rel > ee |
H
acm USE A A e Number of Records
O za — 9-14
ee AMETE ae NS 15 - 24
;
A
í
r N
E
400 800 Miles M
Georgia)
' gia)
thorns 1-2 cm long, fine, straight or slightly recurved, blackish to deep gray. Leaves deciduous; petioles
6-18 mm long, very slender below the wing margin, pubescent, gland-margined; blades 1-2(-3) cm long,
short-obtrullate to broadly obovate in general shape, broadest in the apical half, cuneate to slightly rounded
at the base and tapering into winged petioles, subacute at the apex, usually with 1—2 short, generally sub-
acute lobes per side across the apical end; margins obscurely crenate to evidently crenotoserrate in distal
half, glandular; venation craspedodromous, main veins 1—2 per side, exiting at or beyond at the widest part;
when young thinly hairy, soon glabrate except on the veins below. Inflorescences 1-2-flowered; branches
densely appressed white-pubescent, bearing a few deciduous, linear, membranous, gland-margined bracteoles.
Flowers 12-16 mm diam.; hypanthium white-tomentose; calyx-lobes 2-3 mm long, pubescent externally,
gland-margined; petals + circular, white; stamens 20, anthers ivory; styles usually 3. Fruit 8-10 mm diam.,
subglobose, + glabrous, deep dull yellow or same with red cheek; calyx-lobes appressed, sometimes late
circumscissile; nutlets ca. 3, dorsally sulcate, sides smooth.
Distribution (Fig. 5).—This species is found in the southeastern United States where it is locally com-
mon in Georgia and north central Florida, outliers occurring in Clark and Henry Cos., Alabama and Jasper
Co., South Carolina. There used to be a fine specimen, fully 6 m tall, on the University of Florida campus
at Gainesville in the 1980s.
The drooping, long-petiolate leaves are very small and short-obtrullate to broadly obovate in shape.
In general the proportions are broader than 1.4:1. There is some variation in leaf lobing from quite sharp
to rather blunt and even obscure. An additional characteristic of this species is the way in which extension
shoot leaves may flare outwards towards the tip, their obtuse and large terminal end being quite deeply
and somewhat irregularly lobed. Specimens of C. egens of lower stature (1—2 m tall) may be distinguished
from C. lepida by that species’ nearly or completely unlobed, coriaceous and shiny leaves. Specimens with
2
J
=]
a
"
3
-à
-—
—
-
1 cm. S. Laurie-Bourque del.
4
J
1 M.L. Fernald & B. Lona 6604 (US). fruiting Scale harc —
CERCANO
ZADDINCIN
Fic. 4, Li
1112 j | of the Botanical R h Institute of Texas 2(2)
Fic. 5. C
leaves of the above general shape but more elongated may be referred to C. cirrata and are thus somewhat
intermediate with C. floridana.
3. jioa lacrimata Small, Torreya 1:97. 1901. (Fig. 6). See also Pl. 27 in Phipps et al. (2003). ire: U.S.A.
oripa. Okaloosa Co.: Crestview, 8 Apr 1899, C.D. Beadle 17 (Lectorvre, selected here: US).
Shrubs to 5(-8) m tall; twigs zig-zag at the nodes; growing shoots pubescent, 1 to 2-year old shoots gray or
purple-gray; thorns 1.5-3.0 cm long, straight, purple-gray. Leaves decid ; petioles 2-6 mm long, slender,
pubescent; blades 1-2 cm long, oblong, + narrowly obtrullate or occasionally elliptic, the tip truncate to obtuse
to occasionally cuspidately subacute, subentire to toothed in distal half or very occasionally very slightly
lobed; veins 1-2 per side, exiting in the distal half of the leaf; surfaces glabrous. Inflorescences 1-3-flowered;
pedicels glabrous, bearing + caducous , linear, I eglandular to more usually few-gland-margined
bracteoles; flowering 8 April ‘91 in the Florida panhandle. Flowers ca. 15 mm wide; ea externally
glabrous; calyx-lobes 2-3 mm long, triangular, abaxially glabrous, margins + entire; ular, white;
stamens 20, anthers cream, styles 3-4. Fruit ca. 8 mm wide, + orbicular, glabrous, dos yellow blushed
red, or red; calyx remnants patent-reflexed; nutlets dorsally grooved, sides plane.
Common name.—Weeping Hawthorn.
Distribution (Fig. 7).—This species is found over the Florida panhandle and adjacent Alabama and ex-
tends through Georgia into South Carolina. It is extremely abundant between Pensacola and Panama City,
and locally common elsewhere in pinewoods and open scrub.
Crataegus lacrimata occurs as an attractive, + upright, usually single-stemmed shrub with slender +
weeping branches and small, narrow, glabrous, bright green leaves and is easily recognized among ser.
Lacrimatae from all except the species 3a. This and the next are the only members of ser. Lacrimatae to
commonly possess eglandular bracteoles.
1113
1114 J l of the Botanical R h Institute of Texas 2(2)
Number of Records
0
BN 61-71
3a. Crataegus sp. cf. lacrimata Small, Torreya 1:97. 1901.
Shrubs 2-4 m tall; twigs of the current year glabrous, twigs zig-zag at nodes; l-year old twigs deep dull
purple; thorns mostly 1-4 cm long, few to numerous, straight. Leaves deciduous; petioles 2-6 mm long,
pubescent, glandular; blades 2-4 cm long, narrowly oblong to narrowly oblanceolate, unlobed; distal third
markedly toothed with gland-dotted teeth; base narrowly cuneate; venation craspedodromous with about 2
veins per side diverging at a narrow angle from the midrib and exiting in the distal quarter or third; mostly
glabrous except for tufts of hairs in the axils of the veins below. Inflorescences 1-3(-4)-flowered; branches
glabrous, bearing a few deciduous, linear, membranous, + eglandular, bracteoles. Flowers ca. 18 mm diam.;
hypanthium externally glabrous; calyx-lobes narrow-triangular, glabrous, margins gland-toothed; petals
+ circular, white; stamens 20, anther cream; styles 3. Fruit 7-15 mm diam., + orbicular, glabrous, usually
deep or dull orange-copper when ripe; calyx remnants variably present; nutlets ca. 3, dorsally grooved,
laterally smooth.
Distribution.—This entity occurs in the southeastern USA from the Florida panhandle to Georgia and
South Carolina. It is found in open scrubland, principally on sand or sand-plains where it is locally com-
mon.
It is very similar to C. lacrimata but differs in its conspicuously larger leaves, flowers and fruit.
4. Crataegus munda Beadle, Biltmore Bot. Stud. 1:38. 1901. (Fig. 2). Tre: U.S.A. Soura Carora. Lexington Co.:
Batesburg, 31 Jul 1900, C.D. Beadle 2821 (1ecrorves, selected here: US).
? = Crataegus geniculata Ashe J. Elisha Mitchell Sci. Soc. 17:16. 1900. Comment. henti ial of this taxon can be located but
the protologue suggests something very like C. munda in whose main range it lies.
Stiff, compact thorny shrubs 0.5-2 m tall, or occasionally (? misidentified) much taller; twigs zig-zag, those
of the season densely appressed-pubescent, green, l-year old twigs purple brown-gray; thorns 1-3 cm
um: I Dvorsky, Revision of Crat iac Tanna’ 1115
i w tf, i AT *
Hj LH K ae
Vas
Number of Records
‘0
400 PES ,... B00 Miles
long, slender, + straight, purple-brown at 2 years. Leaves deciduous; petioles 0-5 mm long, winged above,
pubescent only when young, glandular; blades 8-18 mm long, narrowly obovate to cuneate to oblanceolate,
with + straight sides, tapered to the base; tip typically blunt, and sometimes obscurely 3-lobed, more rarely
acute; margins finely serrate-crenate in distal half, the teeth glandular; venation craspedodromous with 1-2
pairs of lateral veins, these exiting above the broadest part of the leaf; thinly pubescent above and below
even at maturity, + densely pubescent below near the vein axils; dark green when young, generally shiny
and thin-chartaceous at maturity; stipules thin, approximately obtriangular. Inflorescences 1-2(-3)-flow-
ered; branches + thinly appressed-pubescent, bearing deciduous, linear, membranous, gland-bordered
bracteoles. Flowers ca. 12-15 mm diam.; penin REAY pubescent cud. pe triangular,
1
gland-toothed, abaxially pubescent mainly; p , white; stamens 20, y or rarely light
purple (C. visendiformis); styles 3-5. Fruits ca. 7-8 mm diam., subglobose, subglabrous, red; calyx remnants
appressed; nutlets 3-5, dorsally sulcate, laterally smooth.
Distribution (Fig. 8). —This is one of the most widespread species in the series. It occurs from North
Carolina to central Florida, southeastern Alabama and South Carolina with scarce outliers occurring in
Mississippi (Jones, Forest Cos.) and Virginia (Nansemand Co.). It is found mainly on sandy soils, in open
brushy places or among pine.
Crataegus munda has a stiff compact habit, like C. lepida, that generally lacks the pendulous twigs so
generally typical of the series. Herbarium specimens of C. munda somewhat resemble C. lacrimata, but,
instead, have pubescent pedicels and shorter petioles, are more likely to have pointed leaves and have more
pronounced marginal teeth. Similarities may also exist to narrower forms of C. recurva, which, however, has
more generally pendant, + entire leaves, usually much longer petioles and more nearly tomentose inflores-
cences and hypanthia. Crataegus munda is one of the few species that regularly flowers at under 1 m tall and
resembles C. lepida in this respect. It has been recorded with attractive pink flowers east of Geneva, Georgia.
1116 I 1 nf ak Dat * ^al D E 14. f Texas 2(2)
Fic. 9. Line drawing of Crataeg from J.B. Phipps 5132 (UWO) and R.K. Godfrey 79497 (FSU), flowering and J.B. Phipps 6762 (UWO), fruiting. Scale
bars — 1 cm. S. Laurie-Bourque del.
ipt y, Revisi g : ] 1117
A similar form perhaps keying out to C. geniculata Ashe, with generally larger, more sharply toothed
leaves, 3-4 veins per side, larger flowers with less hairy pedicels and subpyriform fruit has been found by
R. Lance in northeastern Georgia.
5. Crataegus crocea Beadle, Biltmore Bot. Stud. 1:113. 1902. (Fig. 9). Tex: U.S.A. Froupa. Marion Co.: Citra, 21 Aug
901, C.D. Beadle 44452 (LecrorvrE, selected here: US).
Crataegus curva Beadle, Biltmore Bot. Stud. 1:109. 1902. Tyre: U.S.A. FLoriDa. Duval Co.: Jacksonville, 15 Aug 1901, C.D. Beadle 40822
(LecrorrrE, selected here: US). U.S.A. FLorma. Duval Co.: Jacksonville, 30 Mar 1901, C.D. Beadle 4082, fragment (Epytvre, selected
e, NY).
Crataegus frugalis Beadle, Biltmore Bot. Stud. 1:100. 1902. Tyre: U.S.A. Georcla. Dougherty Co.: along river above Albany, 3 Apr 1901,
e 4092 (LECTOTYPE, here selected, US).
Crataegus recurva Beadle, Biltmore Bot. Stud. 1:106. 1902. Tyre: U.S.A. Flora. Marion Co.: Ocala, Mar 1901, C.D. Beadle 40072 (LEc-
, here selected, A)
Crataegus villaris Beadle, Biltmore Bot. Stud. 1:108. 1902. Type: U.S.A. FLorna. Marion Co.: Citra, 27 Mar 1901, C.D. Beadle 4042 (1sc-
TOTYPE, selected here: US).
? = Crataegus adusta Beadle, Biltmore Bot. Stud. 1:110. 1902. Type: U.S.A. Forma. Alachua Co.: Gainesville, 1901, C.D. Beadle 4065
(Lectotype, selected here: US)
? = Crataegus incana Beadle, Biltmore Bot. Stud. 1:113. 1902. Tyre: U.S.A. FLORIDA. Liberty Co.: Bristol, 24 Aug 1901, T.G. Harbison 4918
(LECTOTYPE, selected here: US). U.S.A. FLorIDA. Liberty Co.: Bristol, 29 Mar 1901, T.G. Harbison 4020, fragment (eptrvpz, selected here,
Shrubs or small trees 2—6 m tall; habit quite lacrimate; twigs of the season densely appressed white-pubes-
cent, l-year old gray-brown to purple-brown; older gray; thorns 1-4 cm long, none to plentiful, straight, +
fine, purple-brown to gray. Leaves deciduous; petioles 2-6 mm long, densely pubescent, glandular at least
initially; blades 1-2.5 cm long, narrowly oblong to narrowly oblanceolate or narrow-elliptic or more com-
monly spatulate in general shape, generally tapered rather fast into a cuneate base, the tip subacute, obtuse
or truncate, + entire or occasionally obscurely lobed; margins gland-dotted when young, occasionally rather
obscurely crenate-serrate at the tip; venation craspedodromous with 1-4 narrowly diverging veins per side
exiting beyond the widest part of the leaf; conspicuously pubescent along the veins below, slightly so above;
often rather densely pubescent at first on the adaxial and slightly pubescent on the abaxial surface, later
glabrescent. Inflorescences 1—3-flowered; branches densely appressed- white- nuca bearing deciduous,
linear, membranous, gland-margined bracteoles. Flowers 16-18 mm diam.; hypanthiu ll
white-pubescent; calyx-lobes ca. 4 mm long, narrow-triangular, gland-margined, abaxially thinly beto
petals elliptic or + circular, white; stamens 20, anthers cream, styles usually 3. Fruit ca. 8 mm diam., sub-
globose, glabrescent to somewhat pubescent, yellow to copper-orange or reddish-orange when-ripe; calyx
remnants reflexed when present, sometimes circumscissile; nutlets 3, dorsally grooved, laterally smooth.
Distribution. —Crataegus crocea ranges from southwestern Alabama to central Florida and through
Georgia and South Carolina just into North Carolina (Fig. 10) where it is locally common on sand plains
and in open scrubby places. It is particularly abundant in Alachua and Marion Cos., Florida.
Crataegus crocea is somewhat variable and will normally be found annotated as C.recurva in my an-
notations. The form selected as typical actually represents one pole of variation. However, it is the most
distinctive compared to other taxa in ser. Lacrimatae and is locally common at least in northern Florida. It
has narrow-elliptic leaves perfectly smooth in outline, acute at the apex, the distal margins sometimes finely
denticulate. Its median leaf shape is similar to the narrowest leaf in the flowering inset. Other forms with
similar leaves may be referred to C. frugalis and, if with larger leaves at maturity sometimes slightly lobed
distally, to C. recurva. Crataegus curva has a somewhat similar leaf form with broader and shorter leaves and
shorter petioles. Commonly, however, leaves of specimens here referred to C. crocea are wider than the type
and somewhat spatulate, as illustrated in Fig. 9 - these represent C. villaris (which is keyed out). All the
foregoing forms intergrade. Forms such as C. adusta which have somewhat similar shaped leaves to those
of C. villaris ee larger i in au parts than typical C. crocea) and often with strongly crenulate to obscurely
lobed leaf ap iate with C. condigna. Finally, other specimens with shorter and broader leaves,
1118 Journal of the Botanical R h Institute of Texas 2(2)
Number of Records
0
N
E
400 0 400 800 Miles
Fic. 10. C level distril f G
rather like those of C. dapsilis (ser. Robustae), but which retain the typically small fruit of subser. Tenues are
referred to C. incana, a form characterized by a particularly dense tomentum when young.
There should usually be no difficulty separating C. crocea from C. munda of this subseries and from the
small-leaved C. condigna of subser. Robustae, both of which have prominently toothed leaf margins, nor from
C. lacrimata with its glabrous inflorescences. Larger-leaved forms of C. cro cea with spatulate-cuneate blades
and obscurely crenate-serrate apices may resemble the C. illudens form of C. lassa from Selma, Alabama.
Crataegus crocea, like many in this subseries, can form an attractive, delicate shrub which should be very
suitable for xeriscaping.
5a. Crataegus vicana Beadle, Biltmore Bot. Stud. 1:104. 1902. Tre: U.S.A. Frormpa. Lake Co.: Tavares, 17 Aug 1901,
C.D. Beadle 4010? (Lecrorvez, selected here: US). U.S.A. Frona. Lake Co.: Tavares, 21 Mar 1901, C.D. Beadle 4010, fragment (EPYTYPE,
selected here, NY.
1 1 1 lar] LAK th t 1 1 1 11 iul n
).
ment.—the epitype well di
o
m laxa Beadle Biltmore Bot. Stud. 1:103. 1902. Tyee: U.S.A. Arasama. Russell Co.: Girard, 8 Apr 1901, C.D. Beadle 4117, fragment
(LECTOTYPE, here selected, NY)
?= Cato a tada p Bot. Stud. 1: pus ie Tire: U.S.A. oe a ao near oe 20 ene fe C.D. Beadle
4001?
equate to C. vicana. U.S.A. FLoripa. Marion Co.: near Duda 21 Mar 1901, C.D. Beadle 4001 (eprtvee, selected iere NY). Comment.—
This is a cl ] 1ditional ial i ded but, even so, the epitype is only a fragment.
Crataegus vicana is somewhat i diate between C. crocea and C. quaesita and consists of forms with long-
petiolate leaf-blades 2.5-4 cm long (i.e., generally much longer than in C. crocea) and length-width ratios of
1.7-1.9:1 that are strongly crenate, especially young, but barely lobed with 0—1 very small subterminal lobes
per side. It somewhat resembles in general shape the broader-leaved C. crocea types with blunt or cuspidate
PL: in I Davies £ frat H [| H 1 1119
drawi f: Crat ita f J.B. Phipps 6492 (UWO), fl i d of C. floridana from J.B. Phipps 6791, flowering and R. Lance
PP \ y Pf E
s.n. (UWO), fruiting. Scale bars = 1 cm. S. Laurie-Bourque del.
narrow-obovate leaf-blades. The flowers are ca. 15 mm diam. and the fruit is smallish and pyriform-ellipsoid
6-8 mm diam, yellow or yellowish orange sometimes splotched with red (Beadle).
Distribution.— Crataegus vicana is a quite common entity of north-central Florida and Georgia with a
single Alabama record.
1120 J | of the Botanical R h Institute of Texas 2(2)
It may be compared to C. crocea (adusta form) but usually has a more triangular to obtuse, rather that
cuspidate, leaf apex that, in addition, is more irregular, at least at maturity. Also, there are similarities to
the slender-leaved forms of the polymorphic C. condigna.
6. Crataegus quaesita Beadle, pe Bot. Stud. 1:33. 1901. (Fig. 11). Tw: U.S.A. Fiona. Gadsden Co.: River
Junction, 3 Apr 1900, C.D. Beadle 2080 selected here: US)
Crataegus viaria Beadle, Biltmore Bot. Stud. 1:101. 1902. Tree: U.S.A. FLORIDA. Duval Co.: Jacksonville, 29 Mar 1901, C.D. Beadle 4065
LECTOTYPE, selected here: U
Crataegus resima Beadle, manor Bot. Stud. 1:100. 1902. Tre: U.S.A. GEORGIA. mA Co.: in no nd - D. Beadle 4089? (Lec-
TOTYPE, selected here: NY 435966; isotypes: NY, US). Comment.—The NY specimen YT it al
a flowering stem on the same sheet.
Shrubs or small trees 2-5 m tall; habit lacrimate; twigs of the season + densely appressed white-pubescent,
l-year old gray-brown to pue ows older gray; thorns 1-2 cm long, usually present, straight, + fine,
purple-brown to gray. Leaves decid ; petioles 2-6 mm long, densely pubescent, glandular at least initially;
blades 1.53.0 cm long, narrowly broad-cuneate to obtrullate in general shape, generally tapered rather fast
into a cuneate base, the tip acute to subacute; 1—2 small sharp lobes per side terminally, occasionally rather
obscurely crenate-serrate at the tip; margins gland-dotted when young; venation craspedodromous with
1-4 narrowly diverging veins per side exiting beyond the widest part of the leaf; conspicuously pubescent
along the veins below, slightly so above; often rather densely pubescent at first on the adaxial and slightly
pubescent on the abaxial surface, later glabrescent, somewhat stiff or floppy at maturity. Inflorescences
1-3C-4-flowered; branches densely appressed- white-pubescent, bearing deciduous, linear, membranous,
gland-margined bracteoles. Flowers 10-13 mm diam.; hypanthium externally white-pubescent; calyx-lobes
ca. 4 mm long, narrow-triangular, gland-margined, abaxially thinly pubescent; petals elliptic or + circular,
white; stamens 20, anthers cream (?purple, see comment); styles usually 3-4. Fruit ca. 8-10 mm diam.,
subglobose, glabrescent to pubescent, mainly reddish-orange to red when-ripe; calyx remnants not always
present, reflexed when so; nutlets 3-4(5), dorsally grooved, laterally smooth.
Distribution (Fig. 12).—Northeastern Florida and Georgia, sandy soils.
Beadle's protologue for C. quaesita claims the anthers are purple, but if so, this would be nearly the only
instance in ser. Lacrimatae, which series it fits in all other respects. The leaves of this species are somewhat
like a very elongated C. egens, especially in its viaria form. Crataegus resima and C. viaria are good matches
for C. quaesita, although the viaria form has less sharp lobes than the type form. Crataegus meridiana (subser.
Robustae) is rather similar but more robust and with larger flowers.
6a. Crataegus sp. aff. resima Beadle, Biltmore Bot. Stud. 1:100. 1902. (Fig. 14).
This is a very local species with some similarities to C. quaesita. The mature, expanded leaves, 2-3 cm
long, are however, much more sharply lobed and very sharp-tipped. Moreover, at anthesis these leaves are
particularly small, only ca. 1 cm. long and thus very little expanded whereas in C. quaesita the leaves are
nearly full size then. The flowers are 10-12 mm diam., the plant is very thorny, and the extension shoot
leaves are 3 cm wide, + isodiametric in general shape, with deep, narrow sinuses presenting a marshallii-
like appearance.
Distribution.—Most specimens are from near Jacksonville, Florida but several are from Georgia.
7. Crataegus floridana Sarg., Bot. Gaz. 33:124. Feb pia (Fig. 1D. Tr: U.S.A. Fora. Duval Co.: Momie 18 Jul
ear, A.H. Curtiss 8 (LECTOTYPE, selected e A). Commen 1
ii) another sheet from A, also labeled "TYPE" and ive l ial, is A.H. Curtiss s.n., Mar 25 1900, same
imd this ne id Lond the ae ones with much n more sharply pointed Su probably uum au plant and perhaps
r o A y
Crat isophylla Beadle, Biltmore Bot. Stud. 1:99. Apr 1902. Tyre: U.S.A. FLorIDA. Duval Co.: Jacksonville, 30 Mar 1901, C.D. Beadle
4067 o selected here: NY; isotype: US).
Crataegus versuta Beadle, Biltmore Bot. Stud. 1:112. a 1902. Tyre: g = A. GeorciA. Dougherty Co.: Albany, 25 Aug 1901, C.D. Beadle
1? (Lectotyee, selected here: US). Comment. lly lobed form.
nL: in 1 n oe ff feat H [| H qe 1121
Fic. 12 f. yl las TRE I £ fynt.
Shrub to 6 m tall; bark of trunk dark brown or rimose; branches drooping; ultimate twigs zig-zag, dark
purple-brown at 2 yrs; thorns ñne, 2 cm long; dark purple-brown at 2 yrs. Leaves deciduous; petioles slen-
der, 1-2 cm long, blades 1.5— 4.0 cm long, spatulate to obtrullate in general outline, somewhat obscurely to
+ distinctly and subacutely lobed in the apical region; margins crenate to serrate in apical region, margins
smooth proximally, strongly gland-dotted young; venation craspedodromous, main lateral veins 2-3/side;
+ pubescent on both surfaces, persistent. Inflorescences 2-4-flowered; branches canescent-tomentose,
bearing deciduous, linear, membranous, gland-margined bracteoles. Flowers 12-15 mm wide; hypanthium
tomentose; calyx-lobes narrowly triangular, abaxially tomentose; margins + entire, petals 5, + circular, white;
stamens 20, anthers cream; styles 3-5. Fruit ellipsoid to subspherical, 6-8 mm diam., subglabrous or with
some residual pubescence, orange; nutlets 3—5, dorsally grooved.
Distribution (Fig. 13).—Crataegus floridana is found in northern Florida, particularly around Jacksonville,
and extends into Georgia with single records from Kershaw Co., South Carolina and Brunswick Co., North
Carolina. It occurs on dry ground.
The delicate, long-petiolate leaves of Crataegus floridana may often be observed fluttering in the wind.
Typical material of this species has long-petiolate leaves much longer than wide that are flared above the
petiole and with shortish but somewhat acute lobes across the distal end. There is a gradation on the one
hand to leaf forms with similar proportions but obscure or blunt lobes and on the other to forms with
rather deep (LII 20—2596) and sharp lobes where the length:width ratio is as low as about 1.25. Crataegus
floridana normally has the largest leaves of the Tenues group but has all the group characteristics of very
short thorns, thin leaves, fine, very pendulous twigs, etc. The lectotype of C. floridana, with its trilobed
leaves, is an exact match for the illustration of fruiting material of the latter species (Sargent, Silva, 1890,
t.189). Sources of plentiful later confusion were that Sargent (1890) mistakenly placed C. floridana under C.
flava (see explanation under C. floridana in vol. 13 of the same work (Sargent 1902)) and that the illustration
1122 J lof the Botanical R h Institute of Texas 2(2)
peer yo Beek? ^ A
VH s ET a
A PCs ar
i Ya, Wa Lt
LC VTL
Sy (
Number of Records
¿0
í
Idi 4-8
5 == 9-14
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cited in Sargent (1890) clearly shows a different species, perhaps C. crocea. Crataegus anisophylla is a name
that may be applied to the more markedly lobed forms. Much of the material of C. floridana was annotated
C. anisophylla by me before the type of C. floridana was located. Also note comments under C. meridiana.
8. Crataegus invicta Beadle, Biltmore Bot. Stud. 1:37 1901. Tre: U.S.A. Georcia. Wayne Co.: Doctortown, no date, fruiting
specimen, no collector or number (i&crorvre, selected here: US 96933 1).
Small shrub 1—2.5 m tall, slender; branches flexuous at the nodes, extending shoots dense-canescent, rap-
idly becoming dark below the indumentum, purple-brown at 1-year old; usually very thorny, thorns 3-5
cm long, fine, straight, dark purple-brown at lyr, becoming gray. Leaves deciduous; petioles 1-3 mm long,
pubescent, gland-margined; blades 1.5—2 cm long, obovate-cuneate in general shape, shallowly lobed in the
apical part, obscurely so in narrower or smaller leaves; margins obscurely crenate or crenato-serrate, teeth
glandular; venation craspedodromous, veins 13 per side, exiting at or beyond at the widest part; surfaces in
late spring to early summer moderately densely covered with a crisped pubescence; very glossy (probably dark
green) adaxially at maturity. Inflorescences 1-2(3)-flowered; branches very dense white-pubescent, bear-
ing a few small, semi-persistent, linear, gland-margined bracteoles. Flowers 12-15 mm diam.; hypanthium
externally + densely pubescent; calyx-lobes narrow-triangular, abaxially thinly, margins glandular-serrate;
petals + circular, white; stamens 20, anther colour unknown; styles 3—5. Fruit 6-8 mm diam. by mid-June,
+ pyriform, fully ripe fruit not recorded.
Distribution.—Dry sandy soils, southeastern Georgia and adjacent South Carolina, scarce and local.
Crataegus invicta is a distinctive species capable of flowering at under 1.5 m tall, as in the type collec-
tion. It is somewhat similar to the cirrata form of C. egens but differs in its relatively shorter petioles, typi-
cally much more thorny habit, and blunter and more shiny, apparently oriaceous, foliage in the little
material seen.
ies Lacrimat 1123
m
Fic. 14. Line drawing of Crataegus sp. aff. C. resima from Curtiss 811 (NY), floweri j fruiting. Scale bars = 1 cm. S. Laurie-Bourque del.
1124 J | of the Botanical R h Institute of Texas 2(2)
8a. Crataegus sp.
This is a typical subser. Tenues shrub with strongly lacrimate branches, short, fine thorns and narrowly
thombobovate leaf-blades, i.e., with the lateral vertices nearer the distal end and entire margins; the long-
cuneate bases taper into the pedicels which are 50-75% the length of the blades; flower diameter is 12-16
mm; styles 2-4; fruit pyriform, 7-12 mm diam., orange, usually flushed red.
This is a local form with a very distinctive leaf shape that does not appear to match any named entity,
is probably not described and would key out next to C. crocea. It is known from northern Florida is very
distinctive.
Subseries Robustae J.B. diio subser. nov. d 9-21). Tyee Species: C. furtiva Beadle.
Hah: 1 A id 1415 : Ts : e2 -3 mm
lati ca. 5 cm D doce: spinae, si adsunt, 2—5 cm longae, vulgo a le: e saepe 2.5—4 cm longae, et RUE in vento
levissimo solum in C. lanata et C. lassa; fructus vulgo grandiores, 8-12 mm diam., saepe cuprini in maturitate.
Habit relatively robust and stocky though ultimate twigs usually + pendulous in mature specimens, twigs
thicker, often 2-3 mm wide at about 5 cm from the tip; thorns, if present, normally relatively stout, 2-5 cm
long; blades commonly 2.5-4.0 cm long, often rather thick and stiff, fluttering in a light wind only in C.
lanata and C. lassa; fruit usually larger, 8-12 mm diam., often copper-colored at maturity.
Members of this subseries also characteristically have thicker and often larger leaves than in subser.
Tenues although this is not true of C. condigna and some forms of C. integra.
9. Crataegus condigna Beadle, Biltmore Bot. Stud. 1:35 1901. (Fig. 15). Tre: U.S.A. Flora. Gadsden Co.: River
Junction, 5 Sep 1899, C.D. Beadle 1095 (LecrorvrE, selected here: US).
ien ravenelii pui Bot. Gaz. ie 122. Feb 1902. Tyre: U.S.A. Georcia: Augusta, sand hills, 1901, A. Cuthbert 411 (ectorvee, here
designated, from Aiken, SC, mentioned by Sargent and which might have included preferable
e Dune have not us located. The oops without precise date, is in full flower.
Crataegus arguta Beadle, Biltmore Bot. Stud. 1:102. Apr 1902. Tyre: U.S.A. ALagama. Russell Co.: Girard, 9 Apr 1901, C.D. Beadle 4123
(LecrorTYPE, selected here: US).
= Crataegus audens Beadle, Biltmore Bot. Stud. 1:114. Apr 1902. Tyre: U.S.A. FLORIDA. Gadsden Co.: Chattahoochee, 30 Aug 1901, T.G.
Harbison 4963 (Lectory, selected here: US). U.S.A. FLoriDa. Gadsden Co.: Chattahoochee, 8 Apr 1901, T.G. Harbison 4097 (EPYTYPE,
selected here, US).
Crataegus clara Beadle, Biltmore Bot. Stud. 1:95. Apr 1902. Type: U.S.A. FLorIDA. Liberty Co.: Bristol, 4 Apr 1901, T.G. Harbison 6033
ndi a paie NY).
Biltmore Bot. Stud. 1:93. Apr 1902. Tere: U.S.A. FLORIDA Alachua Co.: Gainesville, 22 Aug 1901, C.D. Beadle
Jle 4060) t a different species
1
40602 OA aae here: US). Comment.—A flowering syntype at NY (Beadle 4060) app p
perhaps C. illudens.
Crataegus insidiosa Beadle, Biltmore Bot. Stud. 1: du d 1902. Tree: U.S.A. RAE Dale Co.: Ozark, 11 Apr 1901, T.G. Harbison 4115
(LecrorvrE, selected here: NY). Comments.—The is w W.W. Eggleston and seems indistinguishable from
LOM Ic totype i gly
?= pon rava Beadle, Biltmore Bot. Stud. 1:91. Apr 1902. Tyre: U.S.A. Floripa. Leon Co.: Tallahassee, no date, T.G. Harbison 4064
(LecrorTrYPE, selected here: US).
C. rimosa a Biltmore Bot. Ta 1:107 Apr 1902. TYPE: U.S.A. FLoripa. Marion Co.: Citra, 21 Aug 1901, C.D. Beadle 4043? (LECTOTYPE,
selected here: US 981163). U.S.A. FLonipA. Marion Co.: Citra, 27 Mar 1901, C.D. Beadle 4043 ( , selected here: NY)
? = Crataegus alachuaniformis Murrill, Castanea 7:21. 1942. U.S.A. Forma. Alachua Co.: Sugarfoot, Aug 27 1940, WA. Murrill 34276
(Lectotype: FLAS; isorYrE: US).
? = C. praeformosa Murrill, Castanea 7:25. 1942. U.S.A. FLorpa. Alachua Co.: 2 mi S of Warren's Cave, 10 Sep 1940, WA. Murrill 34618
(Lectotype: FLAS; IsoTYPE: US)
Intricate shrub, 1-2.5 m tall or large shrub or small tree to 3-6 m tall (-10 m in the type); branchlets
flexuous; extending twigs densely appressed-pubescent; 1-year old purple-brown; older dark gray; usually
thorny, 2-year old thorns (2) 3-4 cm long, + straight, rather fine, blackish gray. Leaves deciduous; petioles
5-8 mm long, ca. 20-35% length of blade, dense-pubescent, glandular, blades usually 1.5-3.0 cm long,
obovate-spatulate in general outline; base tapered into petiole, apex often cuspidate, subacute to acute;
unlobed or with the merest hint of lobes across the distal portion; margins evenly and distinctly dentate to
Phipr { Dvorsky, Revision of Crat ae ee 1125
Fic. 15. Line drawing of Crataegus condigna from J.B. Phipps 6578 (UWO), flowering and R.K. Godfrey 80642 (UWO), fruiting. Scale bars = 1 cm. S.
Laurie-Bourque del.
1126 | af el Dat « om! D h Inrtitt f Texas 2(2)
Number of Records
0
11-3
crenate-serrate, the teeth gland-tipped; venation craspedodromous, 3—4 veins per side exiting in the distal
part of the leaf; at first pubescent above, below scabrous-hairy on the surface and tomentose on the main
veins; ln d Apu green young, coriaceous and somewhat shiny at maturity. Inflorescences 2-4-flowered;
appressed-pubescent, bearing deciduous, linear, membranous, gland-margined bracteoles.
Flowers 13— 163 mm diam.; hypanthium externally white-canescent, less densely above; calyx-lobes 3 mm
long, narrow-triangular, margins gland-toothed, pubescent abaxially; petals + circular, white; stamens 20,
anthers cream; styles 3-5. Fruit 9-12 mm high, subglobose to sometimes ellipsoid, orange-red or orange at
maturity, sometimes with e nid pea yates reflexed; mn 3—5, dorsally grooved, sides plane.
Distribution (Fig. 16).— al f south-central Alabama, North and
South Carolina, as well as "i coria "m northern Florida. It is found in open areas and thin woodland,
usually on sandy soil where it can be quite common.
This is an abundant and variable species, characterized by unlobed but distinctly toothed leaf margins.
The extreme variation in stature suggests the existence of at least two forms but in the herbarium they
cannot be distinguished. Forms with smaller leaves and more delicate branching, e.g., typical C. condigna,
represent something of a link to subser. Tenues, and a few might just key out to to a broad-leaved form of C.
crocea. On the other hand, forms of C. condigna well-developed leaves may resemble narrower versions of
those of C. florens (but with smaller leaf marginal teeth) and, like the latter, C.condigna may also have circular
extension-shoot leaves. However, the indumentum of both inflorescence and leaves is quite different from
C. florens. The typical form is small-leaved, C. compitalis represents a particularly large-leaved form, while
C. clara is a common and average form.
10. Crataegus alabamensis Beadle, Bot. Gaz. 30:342. 1900. (Fig. 17). ‘tye: U.S.A. ALasama. Montgomery Co.: Mont-
gomery, 16 Apr 1900, C.D. Beadle 2168 (Lecrorvre, selected here: A).
Phipy IR ky, R "un £r g : n » x 1127
Fig 17 Te J. s. £f. I ^E * d.
Bourque del.
C Anderson 4133 (FLAS), flowering and R. Kral 31417 (VDB), fruiting. Scale bars = 1 cm. S. Laurie-
1128 Journal of the Botanical R h institute of Texas 2(2)
? = Crataegus adunca Beadle, Biltmore Bot. Stud. 1:87. 1902. Tyre: U.S.A. FLorIDA. Leon Co.: Tallahassee, 28 Aug 1901, T.G. Harbison
4941 (LecrorrrE, selected h
? = Crataegus dapsilis Beadle, SN Bot. Stud. 1:89. 1902. Tyre: U.S.A. FLORIDA. Lake Co.: Lane Park, near Tavares, 17 Aug 1901, C.D.
Beadle 4836 (Lectotype, selected here: US)
Shrubs or small trees to 5 or 6 m tall; ultimate branches somewhat pendulous, extending twigs densely
tomentose, l-year old twigs dark gray, older gray; thorns 2-3 cm long, generally sparse or even lacking,
slender, + recurved, dark blackish grey at 1 year. Leaves deciduous; petioles slender, 20-30% length of
blade or even shorter, pubescent, glandular; blades 2-3 cm long at anthesis, a little larger at maturity, broad
oblong-cuneate in general shape with tip flattened to slightly cuspidate; sides lacking lobes, tapered regularly;
margins regularly and finely crenato-serrate, this i nearly or completely deappeannë towards the
base; venation craspedodromous with 3-5 lateral veins per side; when young surfaces subglabrous above,
puc ony on the veins below, shiny above at maturity. Inflorescences 3-6-flowered; branches tomentose,
, linear, | , gland-margined bracteoles. Flowers 20-25 mm diam.; hypanthium
ER tomentose; calyx-lobes ron triangular, abaxially appressed white-pubescent in the centre
of their laminas, edges subglabrous, margins with gland-tipped teeth; petals + circular, white; stamens 20,
anthers yellow; styles 3-5. Fruit 10-15 mm diam., subglobose to pyriform, glabrate, red at maturity, often
very glaucous on drying; calyx-lobes spreading; nutlets 3-5, dorsally grooved, sides plane.
Distribution.—Generally scarce, C. alabamensis occurs in south central Alabama and northern Florida
near Tallahassee. It is found in open woods. The type location is recorded as being on on clay soil near
Montgomery, Alabama, a somewhat unusual habitat for this series.
Crataegus alabamensis may be recognized by its beautifully formed leaves with conspicuously, if finely,
crenato-serrate margins and, in the type, unusually broad leaves for this general shape (none narrower than
1.6:1). It may be confused with larger forms of the variable C.condigna, but see the key. Crataegus adunca has,
unusually for this series, violet anthers. Crataegus dapsilis is a small-leaved form rather like broad-leaved
forms of C. crocea.
11. Crataegus integra (Nash) Beadle, Biltmore Bot. Stud. 1:87. 1902. (Fig. 18). Crataegus flava Aiton var.
integra Nash, Bull. Torrey Bot. Clu b 22:150 1895. Te: U.S.A. FLorma. Lake Co.: vicinity of Eustis, 1-15 Jul 1894,
G.V. Nash 1142 (LecrorrrE, selected here: A; isoryPE: US).
? tans Beadle. Biltmore Bot. Stud. 1:88. 1902. Tyre: U.S.A. Mississipi. Lowndes Co.: Columbus, 14 Sep 1901, T.G. Harbison.
066 - 4187? (Lectotype, selected here: US). U.S.A. Mississtrrt. Lowndes Co.: Columbus, 25 Apr 1901, TG. Harbison 4187 (EPYTYPE,
ee here,
Crataegus dolosa B Beadle, Biltmore Bot. Stud. 1:90. 1902. Tye: U.S.A. ALapaMa. Henry Co.: Abbeville, 9 Sep 1901, TG. Harbison 5036
(Lectoryre, selected here: US).
irn ns Beadle, 1:80. 1902. Tire: U.S.A. ArABAMA. Russell Co.: Girard, 26 Aug 1901, C.D. Beadle 4868 ( , selected here:
US).
Shrub or small tree 2-5 m tall; bark of trunk ashy gray, rough or scaly; ultimate branches + pendulous; twigs
at 1 yr old + flexuous, purple-brown or blackish, older dark gray; extending twigs densely white-canescent,
usually thornless, or rare thorns 2-3 cm long, these slightly recurved, blackish or purple-brown at 1 yr old.
Leaves deciduous; petioles short, usually < 25% length of blade, densely pubescent, blades white-hairy at
anthesis, particularly on the veins below; 1.5-4 cm long, obovate-cuneate to narrow-obovate in general
shape, the tip generally + acute, essentially entire else with somewhat obscure, wavy lobes; sides gradually
tapered into the winged upper part of the petiole; margins glandular-denticulate to subentire, + pubescent
above and below when young, especially on the veins below, glabrescent; venation craspedodromous, 2-4(5)
lateral veins per side on all but small leaves; + coriaceous. Inflorescences 2-4-flowered; branches tomentose,
bearing deciduous, oblong-linear, membranous, gland-bordered bracteoles; flowering mid-March (Florida)
to mid-April. Flowers 15-20 mm wide; hypanthium externally tomentose; calyx-lobes 4 mm long, narrow-
triangular, abaxially + tomentose, margins finely toothed, teeth glandular; petals + circular, white; stamens
20, anthers cream or ivory; styles 3-5. Fruit typically (@-)10-12 mm diam., subglobose, yellow blushed red
to orange-red, pubescent; calyx-lobes spreading; nutlets 3—5, dorsally grooved, sides plane.
y, Revisi g i 1129
Mit.
| i fG j fi JB Phi 7 6650 (UWO) flowering and A lance 2195 (l MO) fruitina Scale hare =1m S. Laurie-Bourque
1130 J I of the Botanical R h Institute of Texas 2(2)
MES
UNAS
i
400 0 40 .— à 1 1 1 800 Miles
Fic. 19. € laual detek t £r m
4? F ES I
Distribution (Fig. 19).—Occurring in the southeastern United States, concentrated in the northern half
of peninsular Florida and extending to Alabama, southern Georgia and South Carolina, Crataegus integra is
one of the more common members of series Lacrimatae in its main area of distribution. It may be found in
sand plains, dry scrubby places, thin woodland, etc.
Crataegus integra is a fairly variable species in leaf shape and is notable for its almost perfectly entire
leaf-blades, the margins of which often have numerous, barely detectible leaf-teeth. In well-presented speci-
mens this species frequently displays a considerable variety of leaf size even on one short shoot. It has more
veiny leaves than most species of ser. Lacrimatae and for this reason might key out to ser. Apricae. However,
all other characteristics place it squarely with ser. Lacrimatae. One of the most different entities in the C.
integra complex is C. sodalis (= C. dolosa). This entity has clearly denticulate margins and rather large sub-
orbiculate extension shoot leaves. Crataegus integra differs from the illudens form of C. lassa in its generally
rather smaller leaves and lacks the distinctive cuneate-cuspidate shape of the latter species which are also
sometimes obscurely subterminally lobed. Crataegus integra is particularly common in the northern half
of peninsular Florida where C. lassa is rare. Crataegus constans, with less copious inflorescence tomentum,
longer petioles, and proportionately larger and thinner leaves is perhaps a shade form of C. integra.
12. Crataegus colonica Beadle, Biltmore Bot. Stud. 1:104. 1902. (Fig. 20). Tree: U.S.A. Souta Carora. Beaufort Co.:
Bluffton, Jun i JH ME s.n. ae ae iid id
Comment.—B tl hi i Bluffton in 1882. There are three potential
types at A: a flowering specimen Crec" d ma T se a vere specimen (“rec'd 18 Jul 1899”) and a ue pups eo
all by Mellichamp from the lity The neotype has attached to it an Arnold A
labeled “SL 897. C. d Beadle a trees. ond which implies, but d I Mellichamr 1882 specimen is a
lectotype. T} } i itl ived Jun specimen, so i i I
specimen. 1 decided that it ld be best HM y 1882 Mellichamp speci f he typ ity even
Dh: IDuarelky Ravician nf (rat ies Lacrimat 1131
Fic. 20. Line drawi f: C colonica ( } fi LB. Ph ipp 6511 (UWO), flowering and J.B. Phi & 6765 (UNO) fruiting; and C. pexa (at bottom)
from J.B. Phipps 6502 & 6576 (UWO), fl i 1 Bil Herb. s.n. (US), fruiting. Scale bars — 1 cm. S. Laurie-Bourque del.
1132 J | of the Botanical R h Institute of Texas 2(2)
i = ii Records
1-
IT 4-8
EL 9-14
SS 15 - 24
zum 25 - 60
N
E
400 0 400 800 Miles S
Fis. 21. C y level distributi: I f C. colonica and C. pexa.
though it is sterile and requires an epitype. U.S.A. Sours CAROLINA. Beaufort Co.: Bluffton, ‘received’ 18 Jul 1899, J.H. Mellichamp s.n.
ment.—A l i is ch f it
EPYTYPE, selected here, A). Com:
Shrubs 1.53 m tall; bark of trunk ashy gray, rough or scaly; ultimate branches + pendulous; twigs at 1 yr
old + flexuous, purple-brown or blackish, older dark gray; extending twigs densely white-canescent; usu-
ally very thorny, thorns 2-4.5 cm long, straight or slightly recurved, blackish or purple-brown at 1 yr old.
Leaves deciduous; petioles short, ca. 1596 length of blade, densely pubescent; blades white-hairy at anthesis,
particularly on the veins below; 1.5-2.5(23) cm long, obovate in general shape, sides gradually tapered into
the winged upper part of the petiole; tip acute and + cuspidate to + obtuse, essentially entire to subentire,
though distal margins somewhat denticulate; white-tomentose above and below when young, especially
on the veins below, glabrescent adaxially; venation craspedodromous, 2—4(—5) lateral veins per side on
all but small leaves; + coriaceous. Inflorescences 2—4-flowered; branches tomentose, bearing deciduous,
oblong-linear, membranous, gland-bordered bracteoles; flowering early to mid-April. Flowers 12-15 mm
wide; hypanthium externally tomentose; calyx-lobes 4 mm long, narrow-triangular, abaxially + tomentose,
margins finely toothed, teeth glandular; petals + circular, white; stamens 20, anthers cream or ivory; styles
3—5. Fruit typically 6-9 mm diam., pyriform or subglobose, green turning red or blushed red in mid-late
August, pubescent; calyx-lobes spreading; nutlets 3-5, dorsally grooved, sides plane.
Distribution (Fig. 21).—This species occurs somewhat locally in the southeastern United States, espe-
cially the Carolinas, being particularly abundant in south-central North Carolina. It is found on sand hills,
in dry sandy woodlands, sandy roadsides, etc.
Crataegus colonica is a rather distinct from most other species of this series on account of its thorniness
combined with a rather distinctive leaf-shape reminiscent of forms of C. crus-galli together with its small, +
DL: in 1 Dawie: ff rat H | H f 1133
pyriform fruit. However, whether the next entity, which has a similar distribution and thorniness, should
be included, widening the phenetic width of C. colonica, is not yet clear.
13. Crataegus pexa Beadle, Biltmore Bot. Stud. 1:116. 1902. (Fig. 20). Tyee: U.S.A. NorTH CaroLia. Rowan Co.: near
Salisbury, 28 Apr 1897, Biltmore Herbarium 300b (Lectorvre, selected here: US 981163).
“Small shrub," on only specimen with indication of stature, differing from C. colonica as follows: branchlets
very flexuous; extending twigs densely white-tomentose, 1-year old twigs dark gray; older gray; thorns
generally numerous, at 2 years 3—5(-7) cm long, slightly to strongly recurved, purple-black at 1 year, older
blackish, slender. Leaves deciduous; petioles 3-8 mm long, 10-20% length of blade, tomentose and some-
what glandular young; blades 1.5—3 cm long, obovate to obovate-rhombic in general outline, often narrowly
so; apex acute to obtuse, not or obscurely lobed; margins strongly crenate-serrate for most of their length;
venation craspedodromous, 2-3 veins per side; white-tomentose both sides young, indumentum thinner
at maturity; somewhat coriaceous and dark at maturity. Inflorescences 1-3-flowered; branches tomentose,
bearing deciduous, oblong-linear, membranous, gland-margined bracteoles. Flowers 12 mm diam.; hypan-
thium externally dense-tomentose; calyx-lobes + linear, margins gland-toothed, tomentose abaxially; petals
+ circular, white; stamens 20, anther colour not recorded; styles ca. 4. Fruit 8-10 mm diam., subglobose (a
little taller than broad), ?orange at maturity, tomentose; calyx-lobes reflexed; nutlets ca. 4, dorsally grooved,
sides plane.
Distribution (Fig. 21).—This entity is widely distributed in North Carolina and there are a few records
from South Carolina and Georgia. It is found in dry woodlands, etc.
Crataegus pexa resembles C. colonica in its very thorny nature, most vegetative characteristics and gen-
eral distribution. However, unlike the latter, its leaf margins are normally conspicuously crenate-serrate.
Particularly thorny taxa are unusual in ser. Lacrimatae and as this species has such a similar distribution
to the very thorny C. colonica more work on possible conspecificity is needed. Compared to C. condigna,
plant stature is probably generally lower, the leaf blades are relatively broader, the thorns longer and always
abundant and the twigs are particularly zig-zag. Also the leaf margination is different, see figures. Uncertain
of the correct name at the time, C. pexa has been annotated as C. *acantho-dentata."
14. Crataegus teres Beadle, Biltmore Bot. Stud. 1:43. 1901. Tre: U.S.A. ALabama. Montgomery Co.: Montgomery, 15 Apr
1900, C.D. Beadle 2169 , selected here: US; isotvre: NY).
Yao: 1 E 1 1 fo x «TIC A
5 types at Vo alla
a) 1 kl E
omments. p yI p g
NY I bl inal h tl he | 1 uniformly blunt leaf-tips.
o 4 ES
Shrubs or small trees, 3-6 m tall; with only slightly flexuous twigs; extending shoots subglabrous; 1-year
old tawny to reddish-brown; 2-year old dark gray; 2 year old thorns 23 cm long, + fine to fairly stout, +
straight, shiny blackish-gray. Leaves deciduous; petioles 0.3-0.7 cm long, somewhat pilose young, very
glandular; blades 2-3.5 cm long, narrowly obovate in general shape, apex overall flat-triangular to rounded
or + truncate, often minutely cuspidate; + evenly tapered proximally; lobeless; margins finely dentate, the
teeth gland-tipped; venation craspedodromous, (2-)3-4 veins per side; sparsely hairy on the veins adaxi-
ally, glabrous abaxially; thin, shiny and dark green above at maturity, when slightly coriaceous; extension
shoot leaves broad-elliptic to suborbiculate, larger, often slightly lobed, strong-toothed. Inflorescences
2-6-flowered; branches subglabrous, bearing deciduous, linear, membranous, gland-bordered bracte-
oles. Flowers ca.16 mm diam.; hypanthium glabrous; calyx-lobes 4-5 mm long, narrow-triangular, with
glandular-serrate margins; petals + circular, white; stamens 20, anthers pale yellow; styles 2-3. Fruit 8-10
mm diam., te llipsoid, red; calyx-lobes 7 mm long, spreading, thinly hirsute adaxially, narrow-triangular,
gland-margined; nutlets 2-3.
Distribution.-—Pine woods and sandy fields, south-central Alabama, scarce.
Crataegus teres and the following two species, C. florens and C. attrita, appear to form a natural group
and may be difficult to distinguish. Characters that unite them are the + lobeless leaves with + sharply
denticulate margins that are subcoriaceous and bright to deep green at maturity, subglabrous vegetative
1134 1 Lafelo Dotaniz4lD h Institut f Texas 2(2)
Fic. 22. Line drawing of Crataegus florens from J.B. Phipps 7685 (UWO), flowering and J.B. Phipps 6768 (UWO), fruiting. Scale bars = 1 cm. S. Laurie-
Bourque del.
DL: In L D n Ef $ + || H $ 1135
parts, and inflorescence branches that lack the dense tomentum characteristic of many members of series
Lacrimatae and which instead, range from subglabrous to pilose. Crataegus teres has perhaps the most indi-
vidual characteristics of the three, manifesting less of a zig-zag twig, having the smoothest leaf tips, more
venous leaves (usually 3—4 per side), subglabrous to pilose inflorescences and only 2-3 styles and nutlets,
an ensemble of characters not unlike what might be found in ser. Crus-galli. It is the least common of the
three.
15. Crataegus florens Beadle, Biltmore Bot. Stud. 1:94. E = 22). Tyre: U.S.A. Mississippi. Lowndes Co.: Columbus,
Apr 25 1901, T.G. polit 4176, ee oe selected n re:
Comments.—Thoug ing tl l identification. As with other
O © E rr
NY fn 31 1 i F 1 1 a : a 1 XXE WAJVA TOY
YV. vy. Eggleston but vv
to (hase of the original and a true, collector.
Shrubs or small trees, 3-6 m tall; with flexuous branchlets; extending shoots thinly to more densely
spreading-pubescent; 1-year old tawny to reddish-brown; 2-year old dark gray; 2 year old thorns 2-4.5 cm
long, + fine to fairly stout, straight to slightly decurved, shiny blackish- gray. Leaves deciduous; petioles
0.5-1.5 cm long, somewhat pilose young, Pana ae 2—4.5 cm long, narrowly obovate to obovate in
general shape, apex often + cuspidate, else obt te at the tip, fairly rapidly tapered into the petiole;
usually without trace of lobes, sometimes with 1-2 obscure lobes in the distal part; margins + dentate, the
teeth large at least at anthesis, gland-tipped; venation craspedodromous, 2—4(1-2 in type) veins per side;
sparsely hairy on the veins above and below, elsewhere glabrous; shiny and dark green above at maturity,
thin, but slightly coriaceous; extension shoot leaves broad-elliptic to + orbicular, large, often slightly lobed,
strong-toothed. Inflorescences 3—6-flowered; branches thinly spreading-pubescent, bearing deciduous,
linear, membranous, gland-bordered bracteoles. Flowers 15-18@Q0-25, Beadle) mm diam.; hypanthium
externally pubescent; calyx-lobes 4-5 mm long, narrow-triangular, with glandular-serrate margins; petals
t circular, white; stamens 20, anthers pale yellow; styles 3(—5, Beadle). Fruit 8-15 mm diam., + globose,
orange-red to red; calyx-lobes 7 mm long, spreading, thinly hirsute adaxially, narrow-triangular, gland-
margined; nutlets 3(—5, Beadle).
Distribution (Fig. 23).— Crataegus florens occurs mainly from central Mississippi to the Carolinas and
Florida and is quite common in central Alabama. However, 1 have seen one unambiguous specimen from
West Feliciana Parish, Louisiana (this is still east of the Mississippi, however). This species is found in brushy
thickets on sand-plains, often in more mesic habitats than other members of ser. Lacrimatae.
Crataegus florens is differentiated from C. teres by the des given in the key and also discussed
under the latter species. Here we expand on the protol to include all forms of smaller
flower dimensions and with larger, more Suena dende jeas, thus including those with 3-4 veins per
side but in other respects still quite different from C. teres. With this broadened circumscription, C. florens
is a fairly common species. Differences from C. attrita are discussed under that species.
Crataegus florens is somewhat early flowering as interpreted here, before most other sympatric Lacri-
matae, and much earlier than the type. The flowers are also much smaller than the protologue dimensions
and as the veins are fewer than in the type it may finally prove that typical florens is very rare and therefore
much of the material here attributed may eventually have to be transferred to a broadened C. attrita. But at
the moment these matters must be left in abeyance. Crataegus florens as interpreted here is a striking plant
with zig-zag twigs and fine glossy deep green leaves, superficially resembling members of ser. Crus-galli.
15a. Crataegus attrita Beadle, Biltmore Bot. Stud. 1:98. 1902. Tre: U.S.A. ALagama. Dale Co.: Ozark, 11 Apr 1901, T.G.
Harbison 4116 (Lectotype, selected here: US).
Shrubs or small trees 2-5 m tall; trunk bark rough or rimose; branches “recurved” (? = drooping); one year
old twigs dull gray; extending twigs glabrous, dark greenish—red; thorns ca. 2 cm long, + straight, dark
grayish-black at one year. Leaves deciduous; petioles 3096 length of blade, slender, very short pubescent in
the sulcus, with a few sessile marginal glands; blades 2-3.5 cm long, cuneiform in general shape, tapered to
P l nf tha Rataniral D h Institut f Texas 2(2)
1136
m
Es
< RA
Sha, EAS See CUA ED
|^.
E s [7
S
RM cde
jx
7 a of Records
409 0 400 800 Miles
Fic, 23. C
the base, the apex rounded in general outline; margins dentate, the teeth gland-tipped, occasionally with
a shallowly subterminal lobe on each side; venation semi-camptodromous with 13 lateral veins per side
originating in the proximal part of the leaf and ending in the widest (distal) third of the blade; surfaces
glabrous except for some tomentum along the proximal end of the midvein on both sides and tufts of hair
in the main axils abaxially; thin. Inflorescences usually 2-3-flowered; branches glabrous or extremely
sparsely pilose, bearing deciduous, membranous, linear, gland-margined bracteoles. Flowers 18-22 mm
wide; hypanthium glabrous to very sparsely hairy; calyx-lobes 4—5 mm long, x ligulate from a broad base,
+ entire with some marginal glands, abaxially glabrous; stamens ca. 20, anthers cream; styles 3-5. Fruit
10-14 mm diam., subglobose, glabrous, at maturity yellow splashed red; calyx-lobes neither seen (JBP) nor
recorded (Beadle); nutlets 3—5.
Distribution.—This is a rare species known only from southeastern Mississippi (Lowndes Co.), south-
eastern Alabama (Dale and Greene Cos.) and adjacent Florida (Washington Co.). Another specimen only
differing in its densely pubescent inflorescence is known from South Carolina's 'Haw Ridge.
Cratcegus attrita is essentially similar to C. florens and C. teres and is generally intermediate between
the two. Its dense tufts of hair in the abaxial vein axils are reminiscent of C. aestivalis.
15b. Crataegus pulla Beadle, Biltmore Bot. Stud. 1:96. 1901. Ter: U.S.A. Mississipri. Lowndes Co.: Columbus, 25 Apr
1901, T.G. Harbison 4181 (LecrorvrE, selected here: NY)
Comment.—there is a good fruiting syntype at US.
Crataegus pulla is a little known species apart from excellent type material and the good protologue. It is
a shrub 4-5 m high, with pendulous branches, + obovate leaves which are at first x tomentose, and then
variably somewhat glabrescent, notable for its coarsely and rather irregularly toothed leaves often with a
single, small, somewhat obscure lobe on each side. The flowers are borne on 2—5-flowered inflorescences
Phipps and Dvorsky, Revision of Crat ies Lacrimat 1137
Fic 2A li " ; £f rnt I £ ID Phipy 6481(UWO), flowering and J.B. Phipt
6702 (UWO), fruiting. Scale bars = 1 cm. S. Laurie-Bourque
1138 tani i f Texas 2(
and are 14-17 mm wide. The fruits are subglobose to ellipsoid, 9-12 mm wide, yellowish or orange-yellow
flushed red.
Distribution.—Crataegus pulla was described from sandy flats along the Tombigbee River, Mississippi.
16. Crataegus lassa Beadle, Biltmore Bot. Stud. 1:29. 1901. (Fig. 24). Tee: U.S.A. ALasama. Dale Co.: Selma, 5 Aug
1899, C.D. Beadle 871 (Lecrorrre, selected here: NY)
Crataegus illudens Beadle, Biltmore Bot. Stud. 1:111. 1902. Tyee: U.S.A. FLorma. Marion Co.: Citra, 28 Mar 1901, C.D. Beadle 4055, frag-
ment (LECTOTYPE, selected here:
Crataegus panda Beadle, Biltmore Bot. Stud. 1:89. 1902. Tree: U.S.A. FLORIDA. Leon Co.: Tallahassee, Sep 1901, T.G. Harbison 4051 (Lec-
TOTYPE, selected here: A; isorvre: NY)
Shrub or small tree 2-5 m tall; bark of trunk ashy gray, rough or scaly; ultimate branches + pendulous; twigs
at 1 yr old + flexuous, purple-brown or blackish, older dark gray; extending twigs densely white-canescent;
often thornless, or with rare thorns 3-4.5 cm long, straight or slightly recurved, blackish or purple-brown
at 1 yr old. Leaves deciduous; petioles 0.5-1.5 cm long, usually 25-33% length of blade, densely pubescent,
gland-margined; blades 2-4 cm long at maturity, obovate-cuneate to narrow obovate in general shape, the
tip often strikingly cuspidate, essentially entire; sides gradually tapered into the winged upper part of the
petiole; margins glandular, denticulate to subentire; into the winged upper part of the petiole; margins
glandular, denticulate to subentire; venation craspedodromous, 2—4(—5) lateral veins per side on all but
small leaves; + white-pubescent above and below when young, especially on the veins below, glabrescent;
+ coriaceous. Inflorescences 3—5-flowered; branches tomentose, bearing deciduous, linear, membranous,
gland-bordered bracteoles; flowering early to mid-April. Flowers 15-20 mm wide; hypanthium externally
tomentose; calyx-lobes 6-8 mm long, narrow-triangular, abaxially + tomentose, especially at base and in
centre, margins finely toothed, teeth glandular; petals + circular, white; stamens 20, anthers cream or ivory;
styles 3-5. Fruit typically (8210-12 mm diam., subglobose, yellow blushed red to orange-red, pubescent;
calyx-lobes spreading-recurved or lost; nutlets 3-5, dorsally grooved, sides plane.
Distribution (Fig. 25).—Crataegus lassa occurs in the southeastern United States from central Alabama
to South Carolina and northern Florida and is particularly abundant around its type locality of Selma. It
inhabits sand plains, dry scrubby places, thin woodland etc.
Crataegus lassa is a handsome plant with its normally large foliage and big, floppy leaves. The type
material has rather shorter leaves than most specimens seen commonly near to Selma today and is notable
for having a few small terminal lobes. Crataegus illudens is barely different but has somewhat longer leaves
and leaf tips more often quite entire. Crataegus panda is a form with a slightly shorter and relatively broader
leaf, minutely crenate on the margins. There are also a few intermediates with C. integra.
17. oco de lanata Beadle, Biltmore Bot. Stud. 1:86. 1902. (Fig. 26). Tyre: U.S.A. GEORGIA. Gwennett Co.: near banks
of Yellow R. below McGuire's Landing, 21 Jun 1899, C.D. Beadle 587 (Lecroryre, selected here: US).
? = Crataegus amica Beadle, Biltmore Bot. Stud. 1:97. 1902. Type: U.S.A. Fioripa. Marion Co.: Ocala, 20 Aug 1901, C.D. Beadle 4004?
(LecToTYPE, selected here: NY). U.S.A. FLoriDa. Marion Co.: Ocala, 21 Mar 1901, C.D. Beadle 4004 (Lectortvee, selected here: US).
Crataegus ? inops Beadle, Biltmore Bot. Stud. 1:96. 1902. Tyre: U.S.A. ALañaMa. Dale Co.: Ozark, 1 Apr 1901, T.G. Harbison 4113 (Lecto-
TYPE, selected here: US)
Shrubs to 4 or 6 m, broad-spreading when mature, branchlets often pendant; expanding twigs densely
appressed white-pubescent; l-year old brown to purple-brown, glabrous; thorns 1-4 cm long, + straight,
none to plentiful, slender, purple-brown. Leaves deciduous; petioles 3-6(-10) mm long, pubescent when
young, strongly glandular; blades 3-5 cm long, narrowly to broadly obtrullate in general shape, or sometimes
more obovate; tip acute to subacute, the base usually gradually tapered to the petiole; with 1-2(-3) short,
but well-defined lobes per side across the terminal end; margins subentire to obscurely crenate-serrate,
glandular; venation craspedromous with 2-3 strong lateral veins exiting above the broadest part of the leaf;
pubescent above and below when young and at maturity. Inflorescences 1-4-flowered; branches densely
white-canescent, bearing deciduous, linear, | , gland-margined bracteoles. Flowers about 17-20
Lacrimat 1139
4°
E A
Number of Records
—>0
N
t E
400 0 400 800 Miles
ea z = : S
Fic. 25. C level distri p of Crataegus |
mm diam.; hypanthium externally densely canescent; calyx-lobes 5 mm pun narrow triangular, abaxially
; styles 4—5. Fruit up to
pubescent, mates paud toothed; petals + circular, white; stamens 20, y
9 mm broad, subglob hat pyriform, generally orange or ruddy to crimson, somewhat pubescent;
calyx-lobes xilesed: nutlets 4—5, dorsally sulcate, laterally smooth.
Distribution (Fig. 27).—This species occurs in the southeastern USA from Alabama to the Carolinas and
northern Florida and is generally common. However, there is one record from Chesterfield Co., Virginia. It
is found in pine barrens and open scrubby places, most commonly on sand.
Crataegus lanata is one of the most abundant taxa in series Lacrimatae and is noted for its very large
(for this series), usually floppy, shallowly-lobed, kite-shaped leaves, somewhat white pubescent at all times.
The leaves are typically clearly but shortly lobed (see illustration) and this is the common taxon that has
been annotated C. inops. The fruit of C. lanata ripens to an attractive copper to red. Though the leaves are
typically of quite a different shape from the illudens form of C. lassa (also illustrated), some intermediates
with C. lassa, sens. str., do exist. Crataegus lanata also intergrades with a long-leaved lassa type with a few
very short subterminal lobes and these intermediates have mostly been annotated C. ?inops.Other variants
with broader, more obovate leaves have been annotated as "cf. inops" and these may be the same as C. amica.
These intermediates have as a rule been annotated to the taxon with which they were most similar
18. Crataegus furtiva Beadle, Biltmore Bot. Stud. 1:81. 1902. (Fig. 28). Tyee: U.S.A. Georcia. Dougherty Co.: Albany,
24 Aug 1901, C.D. Beadle 4865 (Lectotype, selected here: NY)
Shrubs, 3-4 m tall or small trees to 6 m; extending twigs densely appressed white-pubescent, 1-year old twigs
purple-brown; older deep gray; usually unarmed but sometimes with thorns, about 2-3 cm long, straight,
these same color as twigs of the same age. Leaves deciduous; petioles 3-6 mm long (unwinged part), widen-
1140 Journal of the Botanical R h Institute of Texas 2(2)
Fic 26 li A n £ f. L n £. I D Dhi 40 19707 (hath INIA fawaving ans JD Dhi E77?) (WO). fruiting. Scale harc = 1
rT A Li J Li H Y
cm. S. Laurie-Bourque del.
est { Dvorsky, Revision of Crat sae lacrimal 1141
ing above into a broad wing, pubescent, glandular; blades 1.5-2.5 cm long, broadly obtrullate or obdeltate
to obtrullate in general shape; base tapered very rapidly into the winged upper part of the petiole; often
shallowly and bluntly 3-lobed across the top, occasionally only sharply cuspidate, alternatively with more
acute tip in the narrower-leaved forms; margins finely and obscurely crenate, the teeth gland-tipped; vena-
tion craspedodromous with 2-3 lateral veins per side, these diverging at a narrow angle from the mid-vein;
surfaces densely tomentose when young, variably glabrescent later. Inflorescences 2—4-flowered; branches
decid , + linear, 1 , gland-margined bracteoles; anthesis
densely tomentose-canescent, | g
mid-April in central Georgia. Flowers 14-16 mm diam.; hypanthium externally densely white appressed-
pubescent; calyx-lobes ca. 5 mm long, narrowly triangular, abaxially appressed white-pubescent, margins
with gland-tipped teeth; petals + circular, white; stamens 20, anthers cream; styles 3. Fruit 8-15 mm diam.,
broad-ellipsoid to + orbicular, glabrous to thinly pilose, orange to red; calyx-lobes reflexed; nutlets ca. 3,
dorsally grooved, laterally smcoth.
Distribution (Fig. 29).—This species is concentrated in Georgia and South Carolina with a few addi-
tional records from North Carolina, Alabama and Florida. It is found in open brushy places, most usually
on sandy soils, often among thin pine.
Crataegus furtiva, as it is usually seen, has among the smallest leaves of the thick-twigged group and is
one of the most densely hairy of all series Lacrimatae when young. Its usually broad, obtrullate leaves (as
illustrated) are quite distinctive. It represents what was at first held to be C. lanata. However, the narrower
leaved form, which is also quite common, is the type form. In spite of some variability in leaf shape and
fruit size, shape and residual indumentum, this species is generally easily recognized.
19. Crataeg idi Beadle, Biltmore Bot. Stud. 1:115. 1902. Tir: U.S.A. Arapama. Dale Co.: Ozark, 11 Apr 1901,
T.G. Harbison 4114 (LectoTYPE, selected here: US; sory: fragment at NY).
1142 J l of the Botanical R h Institute of Texas 2(2)
Fic. 28. Line d
ing of Q furtiva from J.B. Phipps 6555 and 7688 (both UWO), flowering and J.B. Phipps 6674 (UWO), fruiting. Scale bars = 1
cm. S. Laurie-Bourque del.
Shrubs or small trees 3-7 m tall; branchlets zig-zag at nodes; elongating twigs appressed-pubescent, 1-year
old gray-brown to blackish gray; unarmed or with occasional thorns 2-3.5 cm long, + straight, slender,
purple-brown at 1 year, becoming gray. Leaves deciduous; petioles 3-9 mm long, persistently pubescent,
glandular; blades 2-4 cm long at maturity, 1/3 smaller at anthesis, narrowly to broadly cuneate to obtrul-
late in general shape, shallowly and quite sharply 1-3 lobed subterminally at anthesis, a bit more bluntly at
maturity; ends rounded to subacute in general shape; margins crenate-serrate, glandular; venation craspe-
dodromous, with 2-3 lateral veins per side; pubescent on both surfaces, dense-tomentose abaxially on veins,
often becoming floppy. Inflorescences 2-4-flowered; branches appressed-canescent, bearing deciduous,
linear, membranous, gland-margined bracteoles. Flowers 18-20 mm diam.; hypanthium externally densely
Dh: In L Darte’ ffuant : I : 4 1143
Number of Records
0
400 0 400 800 Miles
canescent; calyx-lobes 4-6 mm long, narrow-triangular, abaxially somewhat pubescent, margins gland-
toothed; petals + circular, white; stamens 20, anthers ivory; styles 2-4(—5,Beadle). Fruit 9-12 mm diam.,
subglobose, somewhat pubescent, orange-yellow to ruddy; calyx-lobes on a distinct collar, reflexed; nutlets
2—4C-5, Beadle), dorsally sulcate, sides plane.
Distribution.—This species occurs in the southeastern United States where it is known from Georgia,
South Carolina and north central Florida. There are also a few records from Alabama. It is found in open
brushy places and woodland.
Crataegus meridiana is a rather poorly understood complex of long-petiolate, terminally lobed forms
but more shortly or obscurely so than in C. senta. The general form of the leaf is not unlike that of that of
C. quaesita although extension shoot leaves are more likely to be + isodiametric in C. meridiana. Crataegus
meridiana is quite thorny and the thorns tend to be slender. The somewhat similar C. lassa has rather larger
leaves with blunt or obscure lobes and sl floppy leaves with C. meridiana. Some more delicate specimens
of this species might key out via subser. Tenues to C. floridana. See also comments under C. quaesita.
20. Crataegus lancer E Phipps, T nov. Fig. 30). Tyee: U.S.A. NortH Cagouna. Buncombe Co.: NW of Weaverville, New
Stock Road at Al Sep 2008, R. Lance 3408 (notorvee: UWO; isotypes: A, GAM, UNC, US).
dia — Crataegus irá may prove to be ue same as C. eei Ashe E Hs VAR e Soc. 17:17. 1900) but Ashes
E [] "ug
Frutices vel arbores parvae, 5-8 m alti; li extensi d pubescentes vel i ini brunnei vel atri, veteriores
atrocinerei; spinae nullae vel numerosae, 3-4 cm longae, purpureo-brunneae vel queman atrae in anno secundo. Folia decidua;
petioli 3-7 cm longi, pubescentes, glandulosi; laminae 2-3 cm longae, anguste-obovatae vel obtrullatae vel interdum oblanceolatae
in forma UN contractae in basi, lateriores "Dope peus um acuti vel id pus UMS cum puer 3) o c Pe
enis per lat tus,
renato
crenato-serratae;
imo canescentes vel floccosae solum noe xiédism venam, rss in maturitate, P. anto pilorom male in
ube , lineares, membranaceas
pr
principalibus axillis. Inflorescentiae 3-7
1144 tani i f Texas 2(
glandulo-marginatas bracteolas. Flores 10-12 mm diam.; hypanthi 3 b te, lobi calycis 3— A triangulares,
tis; petala + circularia, alba; stamina 20 theri ] i 3—4. Fructus 10 mm
lati + orbiculares, saepe e vel n lobi eats patento-reflexi; pyrenae 3—4, dorsaliter sulcatae, lstetibis planis.
Shrubs or small trees 5-8 m tall; elongating twigs dense-pubescent to tomentose; 1-year old purple-brown
to blackish, older dark gray; thorns lacking to numerous, 3-4 cm long, purple-brown to blackish in second
year. Leaves deciduous; petioles 3-7 mm long, pubescent, glandular; blades 2-3 cm long, narrowly obovate
to obtrullate or occasionally oblanceolate in general shape, + contracted at the base, broadest near the apex;
apex pointed to obtuse; end shortly 1-2(3) lobed per side; margins glandular, crenato-serrate; venation
craspedodromous, 1-3 veins per side, terminating beyond the widest part of the leaf; surfaces canescent to
wooly hairy only near the midvein at first, tending to subglabrate at maturity except for tufts of hair in the
main axils abaxially. Inflorescences 3—7-flowered; branches pubescent to densely pubescent, bearing de-
ciduous, linear, membranous, gland-margined bracteoles. Flowers 10-12 mm diam.; hypanthium externally
densely pubescent; calyx-lobes 3-4 mm long, triangular, + pubescent abaxially, margins gland-toothed;
petals + circular, white; stamens 20, anthers cream; styles 3-4. Fruit 10 mm long, + orbicular, often copper
to deep red, + pubescent; calyx-lobes patento-reflexed; nutlets 3-4, dorsally sulcate; sides plane.
Etymology.—I am pleased to honor Ron Lance, collector of the type, in the name of this species. Ron
is one of the very few serious modern collectors of Crataegus in the southeastern United States.
Distribution.—This is a fairly common member of the thick-twigged group found from Alabama to both
Carolinas as well as northern Florida and occurs in open scrub and woodland.
This is one of the narrow-leaved members of the thick-twigged group and when the narrower type of
leaf, which is broadest about the middle, is plentiful it makes one of the most distinctive species in ser. Lac-
rimatae. However, leaf shape varies towards + elliptic-oblong, unlobed, crenate-margined on the one hand
to narrow-obovate (rarely narrow-ellipt-rhombic) with single well-defined but short, subterminal, tooth-like
lobes each side on the other. Leaves on extension shoots only are likely to resemble those of C. senta when
they are narrowly obtrullate and sharply lobed. Other striking features are inflorescences more floriferous
than usual for the series and often dark orange to deep red, cherry-like fruit. Crataegus lancei may prove to
be the same as C. yadkinensis but Ashe's protologue is insufficient for certainty and no authentic material
of that species has been discovered. Annotations are mainly under C. ?cullasagensis, an error deriving from
its type locality and Ashe's rather obscure leaf description, the error being corrected when the type of that
species was discovered to belong to ser. Pulcherrimae or perhaps Intricatae.
21. Crataegus senta Beadle, Bot. Gaz. 30:341. 1900. (Fig. 30). Tyre: U.S.A. Nort Carona. Buncombe Co.: Biltmore,
abandoned fields, 11 May 1899, Biltmore Herb. C-18 (ectorvre, selected here: US; IsOTYPE: A).
Shrubs or small trees to 6 m tall; 1-year old wood purplish-brown under an exfoliating wax, older twigs deep
dull gray; thorns absent or numerous, if present 1.5—5 cm long, straight to slightly recurved, purplish-brown.
Leaves deciduous; petioles 0.5-1 cm long, 3096—4096 length of blade, roughly pubescent, black-glandular;
blades ca. 3 cm long, oblong-spatulate to cuneate in general shape; base narrow cuneate, a acute, generally
cuspidate; sharply lobed with 1-2 acute lobes per side in the distal portion; y crenate-serrate
almost to base, very black-glandular at maturity; venation craspedodromous with (2-)3- AS) lateral veins
per side; dull green and sparsely hairy above at first, paler below and pubescent along the main veins and
in the axils. Leaves on extension shoots larger, relatively broader and more deeply lobed. Inflorescences
3-7-flowered; branches dense short-canescent, bearing deciduous, linear, membranous, gland-margined
bracteoles; anthesis early May near Asheville, North Carolina. Flowers 15-20 mm diam.; hypanthium ex-
ternally tomentose; calyx-lobes narrow triangular, abaxially + pubescent, margins glandular-serrate; petals
+ circular, white; stamens 20, anther color not recorded; styles 3-5. Infructescences with 13 fruits; fruit 1
cm diam., subglobose, deep red, punctate, surface mostly glabrous; calyx-lobes patent to reflexed, narrow
triangular, margins t entire; PEN 3—5, dorsally grooved, sides smooth.
Distribution (Fig. 31). T tal very similar distribution to C. lancei andisa quite widespread
species known to me am wesen North Carolina, South Carolina (where it is most common), northern
DL: in L Dres £ f unt H I H n 1145
Fic. 30. Line drawings of: C. Jancei (top) from R. Lance s.n., 3 May 1999 (UWO), flowering and R. Lance 2197 & 98-51 (UWO), fruiting; and C. senta (bottom)
from J.B. Phipps 6486 & 6554 (UWO), fl I R.K. Godfrey 82692 (UWO), fruiting. Scale bars = 1 cm. S. Laurie-Bourque del.
mas 1 Lafeka Rataniral D h Institut f Texas 2(2)
Number of Records
Fic. 31. County level distribution map of C. senta.
Florida and Georgia. However, there is also a disjunct record from West Feliciana Parish, Louisiana (not
mapped). It inhabits dry pine woods, open scrub and sand plains like other members of the series.
This taxon most resembles C. dispar but its leaves have a narrower form and are not so lanate. Smaller
leaves of C. senta have fewer veins (only 1—2 per side). Crataegus senta is very handsome in fruit.
22. Crataegus dispar Beadle, Biltmore Bot. Stud. 1:28. 1901. (Fig. 32). Tv: U.S.A. South Carolina: Aiken Co.: Aiken,
)
27 Jul 1900, C.D. Beadle 2800 ( selected here: US).
à Edi
Shrubs or small trees to 5 m; thorns none or several, 3-4 cm long, straight, purple-brown at 2-year old;
extending shoots densely appressed-canescent; at 2-3 years, dark gray or gray-brown. Leaves deciduous;
petioles 5-15 mm long, densely appressed- pubescent when young, glandular; blades 2.3-3.5 cm long, +
obovate-spatulate in general outline, acuminate at the tip and at the base gradually tapered to the petiole;
generally with about two very acute subterminal lobes per side, the sinuses with LII 15-20%; margins
strikingly toothed, the teeth gland-tipped; venation craspedodromous, 2-3 veins/side; + glabrate above and
below except near the veins. Inflorescences 2-4-flowered; branches densely appressed-pubescent, bearing
deciduous, linear, membranous, gland-margined bracteoles. Flowers ca. 13-16 mm diam.; hypanthium
densely pubescent; calyx-lobes 3 mm long, narrow, pubescent abaxially, margins gland-toothed; petals +
circular, white; stamens 20, anthers ivory; styles 4—5. Fruit 8-12 mm diam., orbicular, nearly glabrous to
slightly pubescent, bright orange-red or reddish; calyx-lobes reflexed; nutlets 4-5, dorsally ridged, laterally
smooth.
Distribution (Fig. 33).—This is a somewhat scarce but distinct plant of the southeastern United States
that is concentrated in South Carolina but has scattered records from Georgia, Florida and Alabama. In
South Carolina I have seen it growing in extremely dry conditions.
Crataegus dispar is one of the most easily recognized members of this series, even identifiable in speci-
Phipy "—————— frat iec Lacrimat 1147
Fi. 32. Line drawing of Crataegus dispar from J.B. Phipps 6505 plus pl f (UWO), flowering, J.B. Phipps 6685 (UWO), fruiting and J.B. Phipps
6501 (UWO), sterile, Scale bars = 1 cm. 5. Laurie-Bourque del.
1148 J lof the Botanical R h Institute of Texas 2(2)
Number of Records
^ 40
j1-3
Id 4-8
== 9-14
—
sass
400 0 400 800 Miles
mens lacking reproductive parts but otherwise adequate and sometimes from elongation shoots alone. Of
note are the broad extension-shoot leaves which are often deeply incised into widely spreading, sharp seg-
ments lobed nearly to the sinuses, rather as in C. marshallii. The interesting foliage of this plant, added to
its attractive flowers and fruit, make it potentially a beautiful ornamental for the dry landscape.
IMPERFECTLY KNOWN TAXA
Crataegus exilis is mentioned here because it has fairly distinctive type material but little else is known about
it. The other two are only known from Ashe's protologues and would need typification to validate them.
However, authentic material has not been located.
23. Crataegus exilis Beadle, Biltmore Bot. Stud. 1:76. 1902. Tee: U.S.A. Georcia. Dougherty Co.: Albany, 25 Aug 1901,
C.D. Beadle 40932 (Lecrorvez, selected here: US).
Shrubs 2—4 m tall; bark of trunk rough; ultimate branches + spreading (1); twigs at 1 yr old + flexuous, color
not recorded; extending twigs not recorded; thorns 1-1.5 cm long, straight, color not recorded. Leaves de-
ciduous; petioles short, ca. 25-30% length of blade, pubescent, at least young; blades 1.5—2.5(-3) cm long,
+ cuneate-obtrullate and very sharply lobed and toothed at anthesis, broad obovate to broad rhombelliptic
in general shape, sides tapered into the winged upper part of the petiole; tip subacute; 0—1 blunt distal lobes
per side, LII « 1096; margins with distal half crenatoserrate; pubescent at anthesis, particularly on the veins
below, glabrous at maturity; venation craspedodromous, ca. 3 lateral veins per side; firm to subcoriaceous
at maturity. Inflorescences 3-7-flowered; branches pilose-pubescent, bearing deciduous, oblong-linear,
membranous, gland-bordered bracteoles; flowering early to mid-April. Flowers ca.15 mm wide; hypanthium
externally pilose-pubescent; calyx-lobes 3-4 mm long, narrow-triangular, margins subentire to finely
nL: IR L Davie: f fg» H || H 1149
toothed, teeth glandular; petals + circular, white; stamens 20, anthers cream or ivory; styles 3-5. Fruit 5-7
mm diam., globose, red; calyx-lobes usually erose; nutlets 3—5, dorsally grooved, sides plane.
Distribution —On sandy soils and along streambanks in south-central Georgia.
The appearance of C. exilis is very distinctive but unfortunately it is known only from single flowering
and fruiting syntypes. It somewhat resembles a larger C. calva (ser. Apricae) but has a merely pilose inflo-
rescence, slightly different leaf shape and somewhat different margination.
Known only from protologues:
Crataegus cuthbertii Ashe, J. Elisha Mitchell Sci. Soc. 17:15. 1900.
Crataegus pearsonii Ashe, J. Elisha Mitchell Sci. Soc. 17:10. 1900.
APPENDIX OF CITED SPECIMENS
Arrangement alphabetical by species, state and y (number following county | ber of records for county).
DAS alabamensis Beadle
ALABAMA. Barbour Co.: (1) Eufaula NWR at old hwy. 165, 6 Apr 1993, D. Drennan 82 (UWO). Dallas Co.: (4) Selma, 11 Apr
1912, A. a (DOV); 8-10 mi E of Selma, 3 Oct 1998, J.B. Phipps and R. Lance 7798 (UWO); Selma, 19 Apr 1920, T.G. Har-
bison 15733 (NCU); near Selma, T.G. Harbison s.n. (NCC). Montgomery Co.: (4) Montgomery, 9 Aug 1899, Biltmore Herb. 899
(A); Montgomery, 16 Aug 1900, Biltmore Herb. 2168 (A, GH)); Montgomery, 15 Apr 1900, Biltmore Herb. s.n. (A). FLORIDA. Leon
Co.: (7) near Tallahassee, 12 Apr 1931, EJ. Palmer, 38560 (A); N side of Tallahassee, 27 March 1976, L.C. Anderson 4133 (FLAS,
FSU, GH); SW of Tallahassee, 24 May 1983, R.K. Godfrey 80656 (UWO); Along Sharer Rd, S of l-10, 27 March 1983, L. Anderson
4133 (UWO); 7 mi NW of Tallahassee, WA. Murrill s.n. (FLAS). GEORGIA. Baker Co.: (1) Limesink Lake, 22 Aug 1977, A. Kral s.n.
(UWO). Richmond Co.: (1) Augusta, 19 Jul 1900, A. Cuthbert 4 (DOV). SOUTH CAROLINA. Aiken Co.: (1) no locality, 31 May
1882, H.R. Ravenel 23 (MO). Kershaw Co.: (1) 0.1 mi E of Richland, 18 Jul 1958, J.A. Duke 1487 (NCU).
E attrita Beadle
BAMA. Greene Co.: (2) Smith Lake area, 14 Apr 1969, J.L. Thomas 1975 (ALU); Smith Lake, 18 May 1967, H. Naugle s.n.
um Coffee Co.: (2) SE of Ella, 8 Jun 1968, R. Kral 31417 (VBB, VDB). FLORIDA. Washington Co.: (2) 1 mi W of Millers Ferry, 3
Apr 1958, R.K. Godfrey 56374 (DUKE, USF). MISSISSIPPI. Lowndes Co.: (1) Chitlin Corners area, 15 mi S of Columbus, W side
of Tombigoee River, 16 Apr 1973, WC. Holmes s.n. (NATC). SOUTH CAROLINA. Chesterfield Co.: (1) Haw Ridge, 23 Apr 1933,
T.G. Harbison s.n. (NCU)
Crataegus colonica Beadle
NORTH CAROLINA. Bladen Co.: (3) W shore of White Lake, 10 Jun 1948, W.B. Fox s.n. (NCSC); On Big Turnbull Creek, 23 Aug
1953, L. Melvin s.n. (NCU); 1 mi W of White Lake, 1 May 1969, R.L. Wilbur 10747 (DUKE). Cumberland Co.: (6) Route 87, 5 mi
SE of 1-95, 4 Jul 1981, J.B. Phipps 511 (UWO); 700m S of Mileport 36 on 1-95, S of Fayetteville, 18 Aug 1993, J.B. Phipps 6766
(UWO); 400m S of Mileport 36 on 1-95, S of Fayetteville, 18 Aug 1993, J.B. Phipps 6765 (UWO); Exit 40 of l-95 and US 301, SW
corner, 18 Aug 1993, J.B. Phipps 6764 (UWO); Exit 40 of l-95 and US 301, SW corner, 6 Apr 1991, 48. Phipps 6511 (UWO); 15 mi
S of Fayetteville, 16 Sep 1936, E. Wherry s.n. (^). Robeson Co.: (13) 3.5 mi SSW of St. Paul's, 20 Sep me c ee (AUA,
CM, 2*DHL, KY, LAF, LYN, NC, 2*USCH, WVA, WILLI); 1-95 N of exit 33, 10 Aug 1993, J.B. Phipps 6671 (UW! Co.: (2)
1.1 mi NNW of Waycross, 28 Jun 1857, J. Haesloop 30133 (NCU); no locality, 28 Apr 1933, T.G. Harbison s.n. seat Wilson Co.:
(1) 24 mi SSW of Black Creek, 21 Jun 1958, A.E. Radford 35726 (NCU). SOUTH CAROLINA. Beaufort Co.: (1) Bluffton, 1882, G.
Engelman 22359 (MO). Lexington Co.: (2) Route 302, SW of Edmund, 10 Apr 1991, J.B. Phipps 6502 (UWO); US 321, 2 km S of
l-26, W side road, 20 Aug 1992, J.B. Phipps 6576 (UWO).
Crataegus condigna Beadle [excluding Alabama records identified as C. insidiosa]
FLORIDA. Alachua Co.: (5) 7 mi N of Gainesville, 8 Aug 1972, S.T. Anderson 57 (FLAS); Gainesville, 27 Aug 1940, WA. Murrill s.n.
(A); E of Gainesville, 2 Apr 1940, WA. Murrill s.n. (FLAS); Gainesville, 17 Aug 1940, WA. Murrill s.n. (FLAS); Gainesville, 2 Jun 1940,
W.A. Murrill s.n. (FLAS). Columbia Co.: (1) SE of Ft. White, 30 Mar 1956, R. Kral 2188 (FSU). Gadsden Co.: (9) Aspalaga, no date,
Biltmore Herb. 297 (NCU); River Junction, 8 Sep 1897, A.H. Curtiss 5982 (FLAS, CM, GA, GH, MO, NCU, NY); Aspalaga, 9 Sep 1950,
H. Kurz s.n. (NCU). Jackson Co.: (1) 3.3 mi N of Marianna-Cottondale, 16 Sep 1950, H. Kurz s.n. (NCU). Leon Co.: (1) Tallahassee,
no date, 7.G. Harbison 5461 (NCU). Liberty Co.: (1) no locality, 24 Mar 1951, H. Kurz s.n. (FSU). Marion Co.: (1) Gainesville, 30
Jun 1939, W.A. Murrill s.n. (GA). Suwannee Co.: (2) 1 mi S of O'Brien, 3 Apr 1940, WA. Murrill & W.B. Tisdale s.n. (FLAS, GA); 7 mi
W of Live Oak, 21 Aug 1939, Tisdale & W.A. Murrill s.n. (FLAS). Wakulla Co.: (2) St. Marks, 12 Apr 1920, T.G. Harbison 15699 (NCU);
SW of jct. of US 98 and Co. Rd. 365, 21 May 1983, R.K. Godfrey 80642 (UWO). Washington Co.: (1) 5 mi SE of Vernon, 31 may
1985, L.C. Anderson 8192 (FSU). GEORGIA. Chatham Co.: (1) no locality, 21 Apr 1959, EO. Mellinger s.n. (GH). Columbia Co.:
(1) E of Harlem on 278, 18 Apr 1992, J.B. Phipps 6557 (UWO). Dougherty Co.: (1) Albany, 18 Sep 1909, WW. Eggleston 5123
(MO). Richmond Co.: (6) Augusta, 16 Apr no year, C.S. Sargent s.n. (A); Augusta, 8 Apr 1919, T.G. Harbison 41 (A); Augusta, 8 Apr
1150 i of the Botanical R h Institute of Texas 2(2)
1919, 7.G. Harbison 14847 (NCU); Augusta, no date, T.G. Harbison 6057 (NCU); Augusta, 11 Apr no year, 7.G. Harbison s.n. (NCU);
Augusta, 30 Sep 1900, A. Cuthbert s.n. (FLAS). Screven Co.: (1) 1 mi N of Statesville, 4 Sep 1968, A. Lazor, R.K. Godfrey and J. Lazor
1067 (FSU). NORTH CAROLINA. Carteret Co.: (2) Lennox Point, E of Beaufort, 12 Sep 1947, H.T. Scofield, L.A. Whitford and W.B.
Fox 1167 (NCSC); Lennox Point, 16 Jul 1965, M. Creech s.n. (DUKE). Craven Co.: (2) New Bern, 17 Apr 1918, 7.G. Harbison 14065
(NCU); 2.8 mi E of Ft. Barnwell, 9 Jun 1958, A.E. Radford 37668 (NCU). Duplin Co.: (1) 2.6 mi NE of Magnolia, 27 Apr 1957, H.E.
Ahles 24052 & G. Ramseur (NCU). Florence Co.: (1) E of Clausen, 24 May 1957, CR. Bell 7529 (USF). Lenoir Co.: (2) 2 mi W of La
Grange, 22 Jun 1957, A.E Radford 25681 (NCU); 2 mi W of La Grange, 6 May 1957, A.E. Radford 22117 (NCU). McDowell Co.: (1)
1 mi SE of jct. NC 264 and US 221, 1 Sep 1956, H.E. Ahles 17770 (NCU). Moore Co.: (1) 2 mi E of Carthage, 25 Apr 1948, WR. Fox
& R.K. Godfrey 1382 (NCSC). New Hanover Co.: (1) 4 mi W of Wilmington on road to Elizabethtown, 26 Apr 1936, W.C. Coker s.n
(NCU). Polk Co.: (1) Columbus, 20 Aug 1921, D.C. Peattie 1329 (NCU). Sampson Co.: (1) 3.8 mi E of Falcon, 5 May 1957, H.E. Ahles
& H. Laing 24548 (NCU). Wayne Co.: (1) 4 mi W of Pikeville, 29 Apr 1956, J. Rose 56 (NCU). SOUTH CAROLINA. Aiken Co.: (5) no
locality, Apr 1890, H.W. Ravenel 3 (A); near Windsor, 14 Apr 1932, EJ. Palmer 39877 (A); no locality, 21 Apr 1880, H.W. Ravenel s.n.
(A); no locality, 12 Sep 1909, WW. Eggleston 5048 (MO); no locality, 31 Mar 1952, W.R. Kelley & WT. Batson s.n. (USCH). Allendale
Co.: (1) SC 3 at 8.1 km S of US 301, 19 Apr 1992, J.B. Phipps 6565 (UWO). Bamberg Co.: (1) SW of Govan, 6 Apr 1957, H.E. Ahles
& J.G. Haesloop 22207 (NCU). Barnwell Co.: (1) no locality, 31 Mar 1952, WR. Kelley & W.T. Batson s.n. (USCH). Berkeley Co.: (1)
West side of county, 7 Apr 1957, (NCU) H.E. Ahles & J.G. Haesloop 22390 (NCU). Calhoun Co.: (2) no locality, no date, H.B. Mooney
& T.H. Stork s.n. (USCH); 2 mi N of St Matthews, 31 Mar 1957, H.E. Ahles & J.G. Haesloop 21751 (NCU). Cherokee Co.: (1) 0.4 mi N
of Blacksburg, 0.5 m W SW of jct. US 29 & Co. Rd. 52, 18 Sep 1957, H.E. Ahles 34198 (NCU). Chester Co.: (1) 3 mi E of Leeds, 20
Apr 1957, OM. Freeman 57143 (NCU). Chesterfield Co.: (4) Sugar Loaf Mountain, 13 May 1932, B.E. Smith s.n. (NCU); Jct. US 1
and Co. Fd. 20, S of Cheraw, 16 May 1976, J.C. Solomon 1951 (CM); SW of Cheraw, 16 May 1976, D.E. Boufford 18306 (MO); Haw
Ridge, near McBee, 3 Apr 1935, V. Matthews & B.E. Smith s.n. (USCH). Dillon Co.: (1) 2.6 mi NW of Smithboro, 18 Apr 1957, G.
Ramseur & H.E. Ahles 23300 (NCU). Florence Co.: (2) E of Clausen, 24 May 1957, CR. Bell 7529 (NCU, USF). Hampton Co.: (1) SC
3 at 25.2 km S of US 301, 19 Apr 1992, J.B. Phipps 6563 (UWO). Lexington Co.: (3) no locality, Mar 1963, WR. Kelley & WT. Batson
s.n. (USCH); US 32, abt. 2 km S of 1-26, 20 Apr 1992, J.B. Phipps 6577 (UWO); US 321 0.5 km S of US 30, Allen Valley Rd., 20 Apr
1992, J.B. Phipps 6578 (UWO). Marlboro Co.: (1) 4 mi S of NC state line, 4 Jun 1958, J.A. Duke 892 (NCU). Orangeburg Co.: (3)
US 178, 5.6 mi NW of Orangeburg, 30 Mar 1957, H.E. Ahles & .G. Haesloop 21561 (FLAS, NCU); W of Copeland, 26 Sep 1957, H.E.
Ahles & J.G. Haesloop 35116 (NCU). Richland Co.: (1) near Columbia, 19 May 1934, EJ. Palmer 42411 (A).
Crataegus crocea Beadle
ALABAMA. Dallas Co.: (1) Rte. 140, Mt. Carmel Church, 8-10 mi E of Selma, 3 Oct 1998, J.B. Phipps & R. Lance 7806 (UWO). Pike
Co.: (4) 5 mi SE of Troy, 29 Sep 1979, S McDaniel and C Duncan 22940 (ALU, FSU, GA, IBE). FLORIDA. Alachua Co.: (36) W of
Gainesville, 22 Mar 19??, WW. Ashe s.n. (FSU); near Gainesville, N of Sevenville, 01 Aug 1940, WA. Murrill s.n. (FLAS, GA); Jct. of
Fla 26 and Millhopper Rd., W side of Gainesville, 22 Mar 1968, WG D'Arcy 2370 (FLAS, LAF, MO, NY, USF); W side of Payne's
Prairie near Gainesville, 19 Jul 1940, WA. Murrill s.n. (FLAS); Gainesville, 31 Jul 1940, WA. Murrill s.n. (3*FLAS); Gainesville, 07 Apr
1931, EJ. Palmer 38429 (MO); Gainesville, 21 Aug 1940, WA. Murrill s.n. (FLAS); Gainesville, 02 Jul 1939, WA. Murrill s.n. (2*A);
Gainesville, 29 Jul 1939, WA. Murrill s.n. (FLAS); Gainesville, 02 May 1941, WA. Murrill s.n. (GA); Gainesville, 21 Jul 1940,
Murrill s.n. (FLAS); Gainesville, 26 Feb 1982, J.B. Phipps 5133 (UWO); Gainesville, 26 Feb 1982, J.B. Phipps 5131 (UWO); Gainesville,
26 Feb 1982, J.B. Phipps 5130 (UWO); Gainesville, 26 Feb 1982, J.B. Phipps 5132 (UWO); Payne's Prairie Nature Reserve, 17 Aug
1993, J.B. Phipps 6767 (UWO); Payne's Prairie Nature Reserve, 17 Aug 1993, J.B. Phipps 6759 (UWO); Payne’s Prairie Nature Reserve,
12 Mar 1993, J.B. Phipps 6653 (UWO); Payne's Prairie Nature Reserve, 12 Mar 1993, J.B. Phipps 6661 (UWO); Paynes Prairie Nature
Reserve, 12 Mar 1993, J.B. Phipps 6657 (UWO); Payne's Prairie Nature Reserve, 12 Mar 1993, J.B. Phipps 6656 (UWO); Payne's
Prairie Nature Reserve, 12 Mar 1993, J.B. Phipps 6654 (UWO); Payne's Prairie Nature Reserve, 12 Mar 1993, J.B. Phipps 6763 (UWO);
Payne's Prairie Nature Reserve, 12 Mar 1993, J.B. Phipps 6658 (UWO); Payne's Prairie Nature Reserve, 12 Mar 1993, J.B. Phipps
6659 (UWO); Kanapaha Rd., 0.5 mi from end, 12 Mar 1993, J.B. Phipps 6670 (UWO); S.R. 346. SE of Archer, 12 Mar 1993, J.B. Phipps
6665 (UWO); Rte. 24, SW of Kanapaha and 0.4 km SW of Rte. 127, 12 Mar 1993, J.B. Phipps 6669 (UWO). Citrus Co.: (8) Rosehill
1 m N of Hernando Co, 21 Apr 1959, G.R. Cooley and RJ. Eaton 6460 (FSU, GH, USF); 1 mi E of Holder, 26 Mar 1965, J. Beckner
661 (FLAS); Citrus Springs, 10 Mar 1993, J.B. Phipps 6636 (UWO); E side of US 41, abt. 3 mi N of Hernando at old bend in road,
10 Mar 1993, J.B. Phipps 6638 (UWO); E side of US 41, abt. 3 mi N of Hernando at old bend in road, 10 Mar 1993, J.B. Phipps 6639
(UWO); W side of US 41, S of Citrus Springs, 10 Mar 1993, J.B. Phipps 6637 (UWO). Clay Co.: (12) Magnolia Springs, 09 Apr 1920,
T.G. Harbison 5674 (A); Green Cove Springs, 29 Jun ae ee Ue iste: Magnolia Springs, 09 Apr 1920, T.G. Harbison
5677 (A); Hibernia, March 1869, W.M. Canby s.n. (GH) 09 Apr 1920, T.G. Harbison 5680 (A); Green Cove Springs,
26 Jun 1940, E.D. Murrill s.n. (A, MO); Green EUR Springs, 01 Apr 1941, ia and WA. Murrill s.n. (GA); Magnolia Springs, 09
Apr 1920, T.G. Harbison 15670 (A); S.R. 16 on N side, 200 m W of side road south to Clay Co. Fairground, 11 Mar 1993, J.B. Phipps
6643 (UWO); S.R. 16 on N side, 200 m W of side road south to Clay Co. Fairground, 11 Mar 1993, J.B. Phipps 6641 (UWO); Penney
farms and Green Cove Springs, 11 Mar 1993, J.B. Phipps 6642 (UWO). Dade Co.: (1) SW 189 Ave and Mowry St, 18 Mar 1969,
GN. Avery 609 (FLAS). Dixie Co.: (1) Old Town, 30 Mar 1920, 7.G. Harbison 5603 (A). Gadsden Co.: (3) no locality, 06 Sep 1940,
WA. Murrill and W.B. Tisdale s.n. (FLAS); Open pineland, 0.75 mi W of Rosedale, 15 Sep 1983, R.K. Godfrey 80955 (UWO); 1.4 mi
from Dellwood, 25 Apr 1983, R.K. Godfrey 80559 (UWO). Gilchrist Co.: (3) Between Newberry and Trenton, 11 Mar 1975, R.K.
AL: nm ” n os £ fyat H I H t 1151
Godfrey 74173 (2*FSU, MO). Hardee Co.: (1) 10 mi W of Bowling Green, 22 Jul 1977, E. Kuczynski s.n. (USF). Hernando Co.: (3)
Chinsegut Hill, 18 Mar 1958, J. Monachino and G.R. Cooley 5644 (USF); Brookesville along Crystal River Rd, 12 Mar 1936, H.H.
Hume s.n. (FLAS); In pine-oakwoocs SW Florida Water Management District lands, 8 mi S of Brooksville, no locality, 15 Mar
1977, T.F. Rochow s.n. (USF). Hillsborough Co.: (12) Tampa, USF campus, 02 Aug 1961, O. Lakela 24519 (FLAS, FSU, GH); Tampa,
USF campus, 11 Mar 1961, O. Lakela 23807 (FLAS, FSU, GH}; Tallahassee, 04 Apr 1923, T.G. Harbison 6073 (A); USF Ecology area,
16 Mar 1983, D. Richardson 901 (USF); USF Campus, 14 Apr 1963, Students s.n. (USF); E side of USF campus Tampa, 25 Mar 1961,
O. Lakela 23925 (USF); no locality, Jun 1971, O. Lakela 32160 (USF); Tampa, campus of South Florida University, 8 Mar1993 J.B.
Phipps 6639 (UWO). Jackson Co.: (4) Dellwood, 28 Jun 1972, R.K. Godfrey 71431 (FLAS, FSU); Marianna, 15 Mar 1937, L.E Arnold
and E. West s.n. (FLAS); Chattahoochee, 03 Apr 1900, CS. Sargent s.n. (DOV). Kershaw Co.: (1) US 1, 8 km SW of lights at
Bethura, 6 Apr 1991, J.B. Phipps 6505 (UWO). Lafayette Co.: (2) 4 mi W of Mayo, 21 Aug 1939, WA. Murrill and W.B. Tisdale s.n.
(2*FLAS). Lake Co.: (2) Tavares, 22 Mar 1901, Biltmore Herb. 4020 (A); Tavares, 15 May 1900, A.H. Curtiss 6617 (GH). Leon Co.: (7)
Along Sharer Rd. N side of Tallahassee, 27 Mar 1976, L.C. Anderson 4133 (FLAS); Tallahassee, 06 Apr 1920, T.G. Harbison 5464 (A);
near Tallahassee, 09 Apr 1929, EJ. Palmer 35213 (2*A); W end of W Campus FSU, 09 Mar 1950, H Kurz s.n. (FSU); Tallahassee, 27
Mar 1977, R.K. Godfrey 75772 (FLAS, FSU). Liberty Co.: (3) Tallahassee, 12 Apr 1972, P Elliot 224 (FSU); Torreya State Park, 04 Jul
1972, R.K. Godfrey 71510 (FLAS); Torreya State Park, 22 Mar 1982, R.K. Godfrey 79452 (FSU). Live Oak Co.: (1) no locality, 13 Apr
1900, S.M. Tracy 6851 (MO). Madison Co.: (6) Ellaville, 21 Aug 1939, WA. Murrill and W.B. Tisdale s.n. (3*FLAS); near Ellaville, 05
Sep 1940, WA. Murrill and W.B. Tisdale s.n. (FLAS, GA); Ellaville, 21 Aug 1939, E.J. Palmer s.n. (A). Marion Co.: (22) N side of road
near Citra, 26 Jul 1940, WA. Murrill s.n. (FLAS); 8 mi S of Ocala, 23 Jun 1983, N.L. Mawhinney 234 (USF); Citra, 13 Jun 1940, WA
Murrill s.n. (FLAS); Cherryhill, 29 May 1942, J.R. Watson s.n. (FLAS); Citra, 26 Jul 1940, WA. Murrill s.n. a Citra, 10 Aug 1939,
W.G. Murrill and Hixon (sp?) s.n. (FLAS); 4 mi W of Ocala, 28 Mar 1958, G.R. Cooley and RJ. Eaton 5756 (GH, USF); E part of Ocala,
17 Jun 1940, WA. Murrill s.n. (FLAS); Ocala, 20 Aug 1901, Biltmore Herb. 134007 (GH); Ocala, 21 Mar 1901, Biltmore Herb. 134007
(GH); Ocala, 06 Apr 1929, EJ. Palmer 35171 (A); Dunnellon, 11 Aug 1940, WA. Murrill s.n. (FLAS); near Citra, 02 Apr 1941, WA.
Murrill s.n. (GA); Citra, 02 Apr 1941, WA. Murrill s.n. (FLAS); Citra, 21 Aug 1901, Biltmore Herb. 4038 (GH); Citra, 22 Mar 1901, Biltmore
Herb. 4038 (GH); Citra, 13 Jun 1940, Watson and WA. Murrill s.n. (FLAS); 1 mi NNE of Dunellon, 10 Apr 1993, J.B. Phipps 6633
(UWO); 1 mi NNE of Dunellon, 10 Apr 1993, J.B. Phipps 6635 (UWO); 1 mi NNW of Dunellon, W of US 41, S of humpback bridge,
10 Apr 1993, /.B. Phipps 6634 (UWO); 1 mi NNW of Dunellon, W of US 41 at the humpback bridge, 13 Apr 1993, J.B. Phipps 6644a
(UWO). Ocala Co.: (3) 3 mi 5 of Ocala, Aug 1939, WA. Murrill s.n. (A); no locality, August 1901, Biltmore Herb. R4007 (A); no local-
ity, Mar 1901, Biltmore Herb. R4007 (^). Orange Co.: (6) Apr 1923, G.K. Jennings and Q.. ?? s.n. (CM); Along US441, 2 mi SW of 2
Ellwood, 31 Mar 1977, L. Baltzell 9408 (FLAS); 1 mi W of US hwy. 441 and State hwy. 421, 07 Mar 1965, LL. Wiggins 19357 (FLAS);
mi S of Ocoee SE corner of Lake Apopka at NE corner of jct. of Fla 50 C-526 and C-439, 20 Apr 1981, R.P Wunderlin and J.
Beckner 8955 (USF); no locality, 16 Mar 1924, EW.H. 8965 (GH); Orlando 24 May 1929, W.C. Edwards s.n. (FLAS). Osceola Co.: (2)
Kissimmee Township 27, 01 Mar 1938, M.L. Singletory s.n. (DUKE); N side of Fla 532, 1.5 mi E of Interstate 4 NW of Loughman,
04 Jul 1977, A.G. Shuey 1942 (USF). Pasco Co.: (1) 2.5 mi E of hwy. 41 and 54, 25 Mar 1961, G.M. Riegler 23895 (USF). Polk Co.:
(3) W of Bartow, 08 Jun 1931, J.B. McFarlin 6391 (A, FLAS); 3 mi NNW of W Frostproof, 9 Apr 1993, J.B. Phipps 6630 (UWO). Su-
wanee Co.: (2) 7 mi W of Live Oak, 21 Aug 1939, WA. Murrill and WB. Tisdale s.n. (A); Hildreth, 10 Jul 1940, WA. Murrill s.n. (FLAS).
Wakulla Co.: (2) 1 mi S of Crawfordville by US Rte. 319, 25 Mar 1982, R.K. Godfrey 79460 (IBE); near Wakulla River, 10 Apr 1931,
EJ. Palmer 38509 (^). Walton Co.: (1) Sand Hills, Chapman s.n. (NY). ?? Co.: (1) no locality, no date, Chapman s.n. (NY). GEORGIA.
Bibb Co.: (1) 7 mi SE of Macon, E side of State Rd 87, 12 Apr 1940, WH. Duncan 1906 (GA). Calhoun Co.: (1) Albany, 18-19 Sep
1909, WW. d 5117 (MO). Charlton Co.: (1) near Trader's Hill, 05 Aug 1959, E Cypert s.n. (GA). Decatur Co.: (5) Woods
edge by hwy. 97 6 mi NE Bainbridge, 15 Jul 1976, RA Norris 3178 (NATC); 34 mi W of Brinson near Seminole Co. line, 31 Mar
1982, R.K. Godfrey 79498 (2*FSU); Mt Pleasant community by Ga Rte. 97, 10 mi N of Bainbridge, 31 Mar 1982, R.K. Godfrey 79497
(FSU, UWO). Lowndes Co.: (3) 4 mi NW of Hahira, 20 Jul 1965, WR. Faircloth 2476 (GA, MO, NCU). Pulaski Co.: (1) near Hawk-
insville, 26 Jun 1902, R.M. Harper 1374 (MO). Screven Co.: (1) 17.6 km N of Newington, 18 Apr 1992, J.B. Phipps 6562 (UWO).
Talbot Co.: (4) Rte. 90, 2 km ENE of Geneva, 9 Apr 1991, J.B. Phipps 6483 (UWO); Rte. 90, 2 km ENE of Geneva, 9 Apr 1991, J.B.
Phipps 6480 (UWO); Rte. 96, abt. 1 mi E of Junction City, 9 Apr 1991, J.B. Phipps 6493 (UWO); Rte. 96, 1 mi E of Junction City on
S side of id, 9 Apr 1991, J.B. Phipps 6490 cS Tatnall Co.: (2) 2 mi W of Reidsville, 03 dos a ^ mdp (GH, MO).
NORTH CAROLINA. Cumberland Co.: (2) NC53 and Co. Rd 2228, N of Jerome, 16 May 1 Solomon 5511 (CM, MO).
Johnston Co.: (1) 2 mi S of Smithfield, 03 P BA Radford and Stewart 152 (NCU). Sou QUA Beaufort Co.: (2)
Bluffton, J. Mellichamp s.n. (NY); Bluffton, 1876, J. Mellichamp s.n. (NY). Calhoun Co.: (1) 4.3 mi ENE of St. Matthews, 14 Apr
1968, S.W. Leonard 1264 (NCU). Darlington Co.: (1) Hartsville, 03 Sep 1909, WW. Eggleston 4932 (MO). Kershaw Co.: (1) near
Camden, 19 Apr 1932, EJ. Palmer 39956 (A). Marlboro Co.: (1) 12 mi N of Bennetsville near SC 38-166 jct., 30 Sep 1956, A.E.
Radford 18977 (NCU). Richland Co.: (1) 5335 Hardscrabble Rd, Blythewood, 08 Apr 1994, D.F Adcock 78 (USCH).
—
rier pid Beadle
ALABA towah Co.: (1) Hinds Road Gadsden, 19 Sep 1999, R. Lance s.n. (UWO). FLORIDA. Jackson Co.: (1) Calhoun-
join e i 5 mi S of I-10, 21 Jun 1982, R.K. Godfrey 79891 (UWO). GEORGIA. Burke Co.: (3) S of McBean and N of Crystal
Lake Road, 13 Aug 1993, J.B. Phipps 6706 (UWOJ; Route 56 N of intersection with 80 just N of Lake Crystal Road, 13 Aug 1993,
J.B. Phipps 6501 (UWO); S of McBean and N of Crystal Lake Road, 13 Aug 1993, J.B. Phipps 6708 (UWO). Clark Co.: (1) Carlton
1152 j | of the Botanical R h Institute of Texas 2(2)
Woods, 20 Apr 1923, Miller 3213 (GA). Richmond Co.: (3) Augusta, 31 Mar 1923, 7.G. Harbison s.n. (NCU); Augusta, 11 Apr
1916, WW. Ashe 19 (2*NCU). SOUTH CAROLINA. Aiken Co.: (8) no locality, 3 Apr 1882, H.W. Ravenel s.n. (GH); 14 Apr 1885,
CE. Flavian s.n. (3*N Y); SE of N Augusta, 12 Apr 1962, H.E. Ahles and PJ. Crutchfield 56348 (NCU); Route 39 exactly 10.7 mi WNW
of Wagener, 12 Aug 1993, J.B. Phipps 6685 (UWO); US 278 1.7 mi E of SC 19, 12 Aug 1993, J.B. Phipps 6686 (UWO); Route 39
exactly 10.7 mi NW of Wagener, 10 May 1991, J.B. Phipps 6505 (UWO). Chester Co.: (2) where SC 72 crosses Broad Road, 4
Nov 1981, J.B. Phipps 5127 (UWO); where SC 72 crosses Broad Road, 4 Nov 1981, J.B. Phipps 5128 (UWO). Edgefield Co.: (1)
9 mi SSW of Trenton, 12 May 1957, A.E. Radford 22501 (NCU). McCormick Co.: (1) Sumter National Forest SE of Clark, 16 Apr
1958, H.R. Totten 1613 (NCC).
Crataegus egens Beadle
ALABAMA. Clark Co.: (1) no locality, 18 Jul, C Mohr s.n. (MSC). Henry Co.: (1) Hwy. 95 S mm 46, 1 Mar 1993, J.B. NELSON Dren-
nen 74 (UWO). FLORIDA. Alachua Co.: (9) Gainesville, 21 Jul 1940, WA. Murrill s.n. (FLAS); Gainesville, 25 Jul 1940, WA. Murrill
and RJ. Wilmet s.n. (FLAS); S.R. 346 SE of Archer fenceline, 12 Mar 1993, J.B. Phipps 6664 (UWO); S.R. 346 SE of Archer fenceline,
12 Mar 1993, J.B. Phipps 6662 (UWO); S.R. 346 SE of Archer fenceline, 12 Mar 1993, J.B. Phipps 6663 (UWO); Gainesville, 26 Feb
1982, J.B. Phipps 5129 (UWO); Rte. 24, SW of Kanapaha and 0.4 km SW of SW127 St. [CH], 12 Mar 1993, J.B. Phipps 6668 (UWO);
Gainesville, University of Florida Campus, 12 Mar 1993, J.B. Phipps 6652 (UWO); University of Florida grounds, 24 Jun 1983, R.K.
Godfrey 80734 (FSU). Clay Co.: (23) Hibernia, Mar 1869, W.M. Canby s.n. (MO, NCU, 4*NY); Magnolia Springs, 9 Apr 1920, T.G.
Harbison 5673 (A); Magnolia Springs, 9 Apr 1920, T.G. Harbison 15664 (NCU); Magnolia Springs, 23 Sep 1923, T.G. Harbison 15673
(A, NCUJ; Magnolia D ; Apr ior 1G. flee AN (NCU); Magnolia Springs, 9 Apr 1920, T.G. Harbison 15775 (NCU);
Magnolia Springs, 23 Sep 1923, T.G. f ny ove Springs, 16 Aug 1993, J.B. Phipps
6750 (UWO); Between E qus and Green Cove: M 16 Aud 1993, J.B. Phipps 6751 (UWO); Between Penny Farms and
Green Cove Springs, 16 Aug 1993, J.B. Phipps 6755 (UWO); Between Penny Farms and Green Cove Springs, 16 Aug 1993, J.B.
rx 6754 (UWO); Between Penny Farms and Green Cove Springs, 16 Aug 1993, J.B. Phipps 6753 (UWO); Between Penny Farms
n Cove Springs, 16 Aug 1993, J.B. Phipps 6752 (UWO); Between Penny Farms and Green Cove Springs, 11 Mar 1993,
P Phipps 6646 (UWO); Between Penny Farms and Green Cove Springs, 11 Mar 1993, J.B. Phipps 6645 (UWO); Between Penny
Farms and Green Cove Springs, 11 Mar 1993, J.B. Phipps 6647 (UWO); Between Penny Farms and Green Cove Springs, 11 Mar
1993, i Phipps 6648 (UWO). Duval Co.: (3) Jacksonville, 1873, CE. Faxon s.n. (GH); Old fields near Jacksonville, 1 Aug 1894, A.H.
Curtiss 4552 (UWO); Old fields I I ma A.H. Curtiss 4552 (UWO). Gadsden Co.: (1) Aspalaga, 9 Sep 1950,
H. Kurz s.n. (FSU). Jackson Co.: ge, r 1964, S. McDaniel 4012 (IBE, FSU). Lafayette Co.: (4) Old Town,
20 Mar 1920, T.G. Harbison 5611 $ NCU); Old Town, 30 P a 0, 7.G. Harbison 5623 (A); Old Town, 30 Mar 1920, T.G. ipis
5610 (A). Lake Co.: (1) near Paisley, 25 Apr 1954, E. Pritchard 690 (FSU). Liberty Co.: (6) Bristol, 23 Aug 1901, Biltmore Hen
(A); Bristol, 21 Aug 1901, Biltmore Herb. 6755 (GH); Bristol, 1 Apr 1901, Biltmore Herb. 4037 (A); Bristol, 2 Apr 1902, Biltmore died
6023 (GH); Torreya Park, 24 Mar 1951, H. Kurz s.n. (FSU); Fla. Rte. 270 vicinity of Sweetwater community, 12 Apr 1982, R.K. Godfrey
79569 (UWO). Madison Co.: (4) Ellaville, 5 Sep 1940, WA. Murrill and W.B. Tisdale (FLAS); Ellaville, 3 Apr 1940, WA. Murrill and W.B.
Tisdale (FLAS); 13.9 mil E of Madison by US Rte. 90, 28 Apr 1982, R.K. Godfrey 79687 (UWO); 13.9 mi E of Madison, 28 Apr 1982,
R.K. Godfrey 79688 (UWO). Marion Co.: (2) Ocala, Apr 1900, M. Rodman s.n. (USF); Citrus, 10 Aug 1939, WA. Murrill and Hixon
Psp] s.n. (FLAS). St. John's Co.: (2) St. Augustine, Feb 1875, M.C. Reynolds s.n. (MO, NY). Suwanee Co.: (3) 1 mi E of Ellaville, 31
Mar 1957, R. Kral 4363 (FLAS, FSU, LAF). GEORGIA. Albany Co.: (1) no locality, 22 Jun 1901, H.C. Comtip s.n. (NCU). Camden
Co.: (1) 10 mi E of Folkston, 5 Apr 1941, WH. Duncan 3147 (GA). Decatur Co.: (1) 6 mi NE Bainbridge, 27 Aug 1948, R.F. Thorne
and W.C. Muenscher 8655 (GA). Early Co.: (1) no locality, 29 Jul 1946, WH. Duncan 6796 (A). Emmanuel Co.: (1) no locality, 30
Mar 1931, EJ. Palmer 38270 (A). Liberty Co.: (1) near Hinesville, 24 Apr 1944, W.C. Grimm s.n. (CM). Lowndes Co.: (4) no locality,
27 Mar 1980, K.D. Perkins and W. Judd 2596 (FLAS, FSU, WILLI, UWO). Randolph Co.: (1) N of Cuthbert, 28 Mar 1948, R.F. Thorne
and W.C. Muenscher 7688 (GA). Schley Co.: (1) Ken Brown Conservation Easement, 4 Jul 1993, J.B. NELSON Drennen 83 (UWO).
Screven Co.: (1) 6 mi NE of Penn ed Apr 1948, A. Cronquist 5004 (GH). Sumter Co.: M ) NW of Arnericus, 30 Jul 1901, R.
Harper 1143 (MO). Tabares :(3) E of Geneva, 13 Aug 1993, J.B. Phipps 6725 (UWO); 2 km ENE of Geneva, 13 Aug 1993,
J.B. Phipps 6727 (UWO); 2 km ENE of Geneva, 9 Apr 1991, J.B. Phipps 6485 (UWO). Tatnall Co.: B )2 a 3 "on
A. Cronquist 4949 (US). Worth Co.: (1) 3.7 mi NNW of Sylvester, 21 Aug 1966, WR. Faircloth 4030 (NCU). SOUTH LINA
Jasper Co.: (3) 1.2 mi NE of junction of Co. Route 462, 24 Sep 1969, S.W. Leonard and A.E. Radford 2778 (MO, NCUJ; : a e
Creek, E of S-219 and S-17, W of Grays, 8 May 1984, CA. Aulbach-Smith and J.B. Nelson 3080 (USCH)
NO
N
MAD florens Beadle
ALAB Barbour Co.: (1) no locality, 11 Aug 1927, KM. Wiegand and W.E. Manning 1356 (HUH). Covington Co.: (1) 2 mi
W of sts on US 84, 15 May 1970, R. Kral 38517 (FSU). Dallas Co.: (8) Rte. 140, Mt. Carmel Church, 8-10 mi E of Selma, 3
Oct 1998, J.B. Phipps and R. Lance 7803 (UWO); Rte. 140, 8-10 mi E of Selma near abandoned church, 10 Apr 1998, J.B. Phipps
7686 (UWO); Rte. 140, 8-10 mi E of Selma near abandoned church, 10 Apr 1998, J.B. Phipps 7685 (UWO); Rte. 140, 8-10 mi E of
Selma near abandoned church, 3 Oct 1998, J.B. Phipps and R. Lance 7801 (UWO); Rte. 140, 8-10 mi E of Selma near abandoned
church, 3 Oct 1998, J.B. Phipps and R. Lance 7802 (UWO); Rte. 140, 4.5 mi E of Selma, 10 Apr 1998, J.B. Phipps 7681 (2*UWO); S
outskirts of Selma, 15 Oct 1984, J.B. Phipps 5484 (UWO). Lowndes Co.: (1) NE of Whitehall Holy Ground Battlefield Memorial
Park, 10 Apr 1998, J.B. Phipps 7678 (UWO). Montgomery Co.: (1) Beziel Springs Church Rd. S of Alabama 24 near Jack Creek,
"m 1 Dwarcku Ravician nf (rat Sm 1153
1 Oct 1998, 8. Phipps and R. Lance 7785 (UWO). Sumter Co.: (1) Old Bluffport off S.R. 28 about 7 mi due E of Livingston along
wooded tracks oo qu mi from Tombigbee, 11 Apr 1998, J.B. Phipps 7696 (UWO). Lee Co.: (1) no locality, 1 Aug 1902,
T.G. } 5 (N A. Gadsden Co.: (1) Fla Rte. 269, 23 May 1982, R.K. Godfrey 79896 (UWO). Hamilton Co.: (1)
Withlacoochee e n pm 6, 1 Oct 1950, H. Kurz s.n. (FSU). eid e ( E de en 11 Sep 1950, H. Kurz s.n. (FSU).
Jefferson Co.: (1) Monticello, 26 Mar 1981, L.H. Lighthipe s.n. (NY). Le locality, 10 Jul 1940, WA. Murrill s.n. (FLAS).
GEORGIA. Baker Co.: (1) no locality, 22 Aug 1977, R. Kral 60870 TA e Co.: (1 ) jm hwy. 129 about 8.8 mi N of
Pulaski Co. line 0.8 mi S of Big Indian Creek Crossing, 10 Apr 2001, R. Lance 2126 (UWO). Liberty Co.: (1) About the Altamaha
River Swamp, 18 Jun 1895, J.K. Small s.n. (NY). Lincoln Co.: (2) no locality, 2 Sep 1940, WH. Duncan 2832 (2*NCSC). Lowndes
Co.: (1) Md inl Province, 28 Jun 1965, WR. Faircloth 1902 (MO). Macon Co.: (2) Just E of Oglethorpe on W side of Flint
River, 8 Oct 1 W.H. Duncan 1750 (2*NCU). Richmond Co.: (1) Augusta, 2 Aug 1902, A. Cuthbert s.n. (FLAS). Sumter Co.:
(2) no ane ra 1976, R.A. Norris 2863 (IBE, NATC). Taylor Co.: (1) Coastal Plain Province, 16 May 1974, WR. Faircloth 7530
(NCU). LOUISIANA. West Feliciana Parish: (1) 10 Aug 1916, R.S. Cocks 4022 (NO). MISSISSIPPI. Rankin Co.: (1) South end of
Ross Barnet Reservoir, 13 Apr 1984, J.B. Phipps 7709 (UWO). NORTH CAROLINA. Carteret Co.: (1) Beaufort, s.n. (UWO). Greene
Co.: (1) Sand ridge ca Y mi NE of Browntown, 22 Jun 1960, H.E. Ahles and J. Haesloop 53705 (NCU). Lenoir Co.: (1) Rte. 903
few mi S of La Grange, dirt driveway W side of road N of turkey farm, 18 Apr 1993, J.B. Phipps 6768 (UWO). Wayne Co.: (1) 2.5
mi W of Lenoir Co. line, 14 Jul 1949, W.B. Fox 2665 (NCSC). SOUTH CAROLINA. Aiken Co.: (2) 5 Aug 1898, H. Egert s.n. (2*MO).
Chesterfield Co.: (2) near McBee, 18 May 1934, EJ. Palmer 42382 (HUH); Haw Ridge near McBee, 17 Jul 1933, T.G. Harbison s.n.
(NCU). Clarendon Co.: (1) Coclough Pond, 15 Sep 1939, R.K. Godfrey and R.M. Tryon 1461 (NYBG). Kershaw Co.: (1) no locality,
3 May 1923, 3 May 1923, T.G. Harbison 7028 (HUH). Lexington Co.: (1) About 1 mi N o US 178 S of Swansea, 11 Sep 1939, J.B.
Phipps 6675 (UWO). Richland Co.: (1) no locality, 1 Jul 1908, WW. Ashe s.n. (NCU).
piel floridana Sarg.
FLORIDA. Alachua Co.: (3) Gainesville, 31 Jul 1940, WA. Murrill s.n. (FLAS); Rte. 24 SW of Kanapaha and 0.4 km SW of SW127
St, 12 pe 1993, J.B. Phipps 6666 (UWO); Paynes Prairie Nature Reserve, 17 Aug 1993, J.B. Phipps 6761 (UWO). Baker Co.: (2)
Lake City, 2 Apr 1923, T.G. Harbison 6062 (A); Lake City, 7 Apr E 0, T.G Harbison 5654 (A). Citrus Co.: (1) Pineola, 3 Apr 1930, FS.
M 7000 (GA). Clay Co.: (3) Magnolia Springs, 9 Apr 1920, T.G. Harbison 5662 (A, NCU); High Land, 2 Jul 1929, Hugh O'Neill
AS). Duval Co.: (10) Jacksonville, March no year, A.H. neis 811 (CM, 2*GA, MO); ped 25 Mar 1901, A.H. Curtiss
S.N. Jacksonville, 16 Jul no year, A.H. Curtiss s.n. (A); Jacksonville, 15 Aug 1901. Biltmore Herb. 4076 (GH); Jacksonville, 17 Mar
1893, A.H. Curtiss 4216 (MO); Jacksonville, 25 Mar 1902, Bayutou s.n. (FLAS); Mond ne Herb. 4067 (GH). Gadsden
Co.: (2) Aspalaga, 5 Apr 1964, 5. McDaniel 4100 (IBE, FSU). Gilchrist Co.: (1) W of Newberry, 11 Jul 1940, W.A. Murrill s.n. (FLAS).
Hamilton Co.: (2) 5.3 mi W of Jasper, 8 Jun 1950, H. Kurz s.n. (FSU); Floodplain eee 1 Oct 1950, H. Kurz s.n. (FSU). Hernando
Co.: (1) Chinsegut Hill Wildlife Refuge, 3 Apr 1958, G.R. Cooley and R. Eaton 5865 (FSU). Levy Co.: (1) near Chiefland, 7 Apr 1931,
J. Palmer 38435 (A). Liberty Co.: (1) Bristol, 7 Sep 1940, W.B Tisdale and WA. Murrill s.n. (FLAS). Madison Co.: (4) Ellaville, 3 Apr
40, WA. Murrill and W.B. Tisdale s.n. (FLAS); Cherrey Lake, 22 Aug 1940, WA. Murrill s.n. (FLAS); 3.2 mi W of Co. line, 8 Apr 1961
Barbara Moore 126 (FLAS); Ellaville, 21 Aug 1939, WA. Murill and W.B.Tisdale s.n. (FLAS). Marion Co.: (3) Summerfield, 11 Aug
1939, WA. Murrill s.n. (FLAS): S of Juniper Creek, 1 Sep 1941, W. H. Duncan 4158 (GA); 1 mi NNW of Dunellon W of US41 at the
humpbacked bridge at Chatmar, 15 Aug 1993, J.B. Phipps 6743 (UWO). Putnam Co.: (1) 1 mi N of Orange Springs, no date,
W.A. Murrill s.n. (FLAS). Suwanee Co.: (2) Luraville, 9 Aug 1946, West and Arnold s.n. (FLAS); Between US hwy. 90 and railroad,
23 Mar 1977, D.L. Martin 993A (FLAS). GEORGIA. Baker Co.: (1) SW of Newton, 20 Aug 1941, WH. Duncan 4116 (GA). Bibb
Co.: (1) 13 mi SW of Macon, 24 Oct 1967, R.L. Lane 1637 (NCSC), Brooks Co.: (1) 6 mi SW of Hyattville, 14 Jul 1965, R. Dean and
W. Faircloth 2181 (NCU). Chattahoochee Co.: (1) NE of Cusseta along Liberty Hill Rd., 20 Sep 1999, R. Lance s.n. (UWO). De
Kalb Co.: (2) Stone Mt. near Yellow River, 15 Aug 1899, A.H. Curtiss 6520 (A, GH). Dougherty Co.: (1) no locality, 19 Sep 1909,
WW. Eggleston 5119 (MO). Elbert Co.: (1) S of Bowman, 11 Apr 1977, R. and D. Coile 472 (GA). Jefferson Co.: (1) 0.2 mi NE of
Ogeechee River, 26 Jul 1967, J.R. Bozeman and J. F. Logue 10777 (NCU). Lee Co.: (2) McGuire's Mill, 2 Aug 1895, J.K. Small s.n. (A);
N of Md bridge over Muckalee Creek, 11 May 1940, W.H. Duncan 2347 (FLAS). Mitchell Co.: (2) 0.2 mi E of Flint River,
29 Jun 1966, J. Bozeman 5217 (NCU); no EN 23 Aug 1940, R. McVaugh 5248 (GA). Richmond Co.: (1) Augusta, 16 Apr 1923,
As heron 7006 (A). Sumter Co.: (4) 10 mi NE of Americus, 25 Sep 1939, WH. Duncan 1670 (GA, NCU); 10 mi NE of Americus,
Sep 1939, WH. Duncan 1671 (GA, NO ibo Co.: (8) Rte. 96 about 1 mi E of Junction City, 14 Aug 1993, J.B. Phipps 6714
ques "e 96 about 1 mi E of Junction City, 13 Aug 1993, J.B. Phipps 6715 (UWO); Rte. 96 about 1 mi E of Junction City, 13
Aug 1993, J.B. Phipps 6716 (UWO); Rte. 96 about 1 mi E of Junction City, 9 Aug 1993, J.B. Phipps 6492 (UWO); Rte. 96 about 1
mi E of Junction City, 9 Aug 1993, J.B. Phipps 6495 (UWO); Rte. 96 about 1 mi E of Junction City, 13 Aug 1993, J.B. Phipps 6717
(UWO); Rte. 96 about 1 mi E of Junction City, 13 Aug 1993, J.B. Phipps 6719 (UWO); 2 km ENE Geneva, 13 Aug 1993, J.B. Phipps
6723 (UWO). Tatnall Co.: (1) 2 mi W of Reidsville, 21 Aug 1940, WH. Duncan 2729 (GA). Terrell Co.: (1) 2 mi SE OF Parrot, 8 Oct
1939, W.H. Duncan 1756 (NCU). Toombs Co.: (1) Vidalia, Apr 1914, M. Huger s.n. (A). Webster. Co: (1) 1.3 mi W of Kinchafoonee
Creek, 27 Jul 1966, J.R. Bozeman 6728 (NCU). Wilcox Co.: (2) N of Abbeville, 7 Jui 2007, J.R. Bozeman 5518 (2*NCU). NORTH
CAROLINA. Brunswick Co.: (1) Smith Island, 29 Jul 1965, AW. Cooper 2856 (NCSC). SOUTH CAROLINA. Kershaw Co.: (1) 6
mi S of Cawdenon, 17 Jul 1933, TG. Harbison and H.R. Totten s.n. (NCU)
1154 Journal of the Botanical R h Institute of Texas 2(2)
Crataegus furtiva Beadle
LABAMA. Barbour Co.: (2) Eufaula NWR, 05 Aug 1994, D. Drennen 99 (UWO); Hwy. 431 S at Nat Guard Armoury 4 mi N of
Eufaula, 18 Aug 1993, D. Drennen 88 (UWO). Coffee Co.: (1) 1 mi S jct. AL 87 and US 84, 08 Jun 1968, R. Kral 31417 (UWO). Dallas
Co.: (5) 2 mi E of Selma, 11 Apr 1900, CS. Sargent s.n. (A); Selma, 12 Apr 1912, T.G. Harbison 10842 (NCU); Rte. 140 Mt Carmel
church 8-10 mi E of Selma, 03 Oct 1998, J.B. Phipps and R. Lance 7796 (UWO); 140 Mt Carmel Church 8-10 mi E o Selma, 03
Oct 1998, J.B. Phipps and R. Lance 7800 (UWO); Rte. 140 8-10 mi E of Selma, 03 Oct 1998, J.B. Phipps and R. Lance 7799 (UWO).
?? Co.: (1) no locality, 14 Apr 1902, T.G. Harbison 4114 (NCU). FLORIDA. Alachua Co.: (1) NE Alachua, 10 Aug 1993, J.B. Phipps
6756 (UWO). Lafayette Co.: (1) 6 mi W of Branford, 11 Jul 1940, WA. Murrill s.n. (FLAS). Lake Co.: (2) W side of Fla 19, 23 Mar
1981, D. Richardson, R.P. Wunderlin and B. Hausen 8134 (2*USF). Leon Co.: (2) W of Tallahassee, 17 Mar 1982, R.K. Godfrey 79429
(IBE); 1.5 mi W of Miccosukee, 07 Aug 1951, H. Kurz s.n. (FSU). Liberty Co.: (2) Bristol, 16 Mar 1890, GSS 18 (NCU); Torreya State
Park Entrance, 17 May 1983, R.K. Godfrey 80627 (FSU). Marion Co.: (1) Ocala, 11 Aug 1939, WA. Murrill s.n. (FLAS). Sumter Co.:
(1)3 mi Sof Coleman, 02 Apr 1985, B Hansen 10346 (USF). Suwanee Co.: (1) E of Suwanee River US 90, 28 Apr 1982, R.K. Godfrey
79693 (UWO). Wakulla Co.: (1) St Marks, 25 Sep 1913, T.G. Harbison 1209 (A). GEORGIA. Ben Hill Co.: (3) 1.5 mi S of Bowen
Mill along US-129, 30 Jul 1964, WAR. Faircloth 1433 (GA, MO, NCU). Bibb Co.: (3) 13 mi SW of Macon, 24 Oct 1967, R.L. Lane 1637
Q5 D Mill, 23 Apr 1898, EF Andrews s.n. (AUA); 10 Jun 1937, AM Laessle s.n. (GAM). Burke Co.: (3) River Rd at 18 km N of
, 19 Apr 1992, J.B. Phipps 6569 (UWO); River Rd at 18 km N of US 301, 19 Apr 1992, J.B. Phipps 6571 (UWO); River Rd at 18
n i. e US 301, 12 Aug 1993, J.B. Phipps 6701 (UWO). Clarke Co.: (2) Athens, 16 May 1925, J.H. Miller s.n. Mi Athens, 16 Apr
929, K.M. Drewry E7783 (GA). Colquitt Co.: (1) 5 mi N of Doerum, 25 Aug 1990, D. Drennen 24 (UWO). Decatur Co.: (4) 1 mi S
b Faceville, 28 May 1983, R.K. Godfrey 80661 (FSU); 4 mi N of Mt Pleasant Community, 31 Mar 1982, R.K. c 79496 (FSU); 1
mi S of Faceville, 28 May 1983, R.K. Godfrey 80662 (FSU); 34 mi W of Brinson, 31 Mar 1982, R.K. Godfrey 79498 (FSU). Elbert Co.:
(1) S of Bowman, 11 Apr 1977, N Coile 472 (NCSC). Heard Co.: (1) no locality, 29 May 1963, J.C. Ware s.n. (GA). Gwinnett Co.:
(1) Mr. McGuire's Mill banks of Yellow road, 02 August 1895, J.K. Small s.n. (A). Jefferson Co.: (1) 1.2 mi S of Co. line along GA
17,05 Aug 1966, J.R. Bozeman 7071 (NCU). Mitchell Co.: (1) 3 mi SW of Hopeful, 23 Aug 1940, R. McVaugh 5248 (A). is
Co.: (18) Augusta, 09 Apr 1919, T.G. Harbison 43 (A); Augusta, 12 Apr 1901, A. Cuthbert 402 (NCU); Augusta, 05 Apr 1914, CS.
Sargent 520, (2*A); Augusta, 31 Mar 1913, T.G. Harbison 1038 (A); Augusta, 02 Apr 1900, A. Cuthbert 4d (NCU); Augusta, a Apr
1923, WW. Ashe s.n. (NCU); Augusta, 15 Apr 1923, T.G. Harbison 7002 (A); Augusta, 09 Apr 1919, T.G. Harbison 51 (A); Augusta,
16 Apr 1923, T.G. Harbison 7003 (A); Augusta, 07 Jul 1900, A. Cuthbert 3 (DOV); Augusta, 31 Mar 1923, T.G. Harbison 6058 (A);
Augusta, 02 Apr 1913, T.G. Harbison 1042 (A); Augusta, 09 Apr 1919, T.G. Harbison 45 (A); no locality, 14 Aug 1958, J.A. Duke 2025
(NCU); 10 mi E of Bath Rd, 14 Aug 1958, J.A. Duke 2118 (NCU); Augusta, 04 Nov 1914, T.G. Harbison 11 (A); Summerville near
Augusta, 3 Oct 1914, T.G. Harbison 1041 (A). Meriwether Co.: (1) 5 mi W of Greenville, 30 Apr 1961, E.E. E Barclay
3414 (FSU). Schley Co.: (2) Ken Brown Conservation Easement 5 mi NW Ellaville, 01 Sep 1993, D. Drennen 91 (UWO); Ken Brown
Conservation Easement 5 mi NW Ellaville, 01 Sep 1993, D. Drennen 92 (UWO). Sumter Co.: (1) Americus, Apr 1897 2 W. Huger
s.n. (NY). Talbot Co.: (1) Rte. 96 1 mi E of Jct. City, 13 Aug 1993, J.B. Phipps 6713 (UWO). Tatnail Co.: (1) 2 mi W of Reidsville, 03
Apr 1948, A. Cronquist 4949 (GA). Thomas Co.: (1) Thomasville, Mar 1890, Canby s.n. (DOV). Troup Co.: (1) West Point, 12 Jun
1899, s.n. (NCU). NORTH CAROLINA. Bladen Co.: (1) Along big turnbull creek, 25 Aug 1993, M. Lionel s.n. (NCC). Cumberland
Co.: (1) Steadman (across river), 20 Apr 1933, T.G. Harbison and H.R. Totten s.n. (NCU). ?? Co.: (2) no locality, no date, WW. Ashe
s.n. (NCU); no locality, May 1938, L. Hunt s.n. (NCU). SOUTH CAROLINA. Aiken Co.: (7) near Windsor, 15 Apr 1932, EJ. Palmer
39876 (A); 3 mi N of Aiken, 03 Apr 1939, R.T. Clausen 3769 (A); Graniteville, 23 Apr 1899, H. Eggert s.n. (MO); near Windsor, 15
Apr 1932, EJ. Palmer 39857 (A); 1.1 mi SW of Saluda-Aiken line, 11 Apr 1962, H.E. Ahles and PJ. Crutchfield 56159 (NCU); 12-15
Sep 1909, WW. Eggleston 5064 (CM); US 278 ca 7 km E of jct. Ma SC 19, 03 Apr 1992, J.B. Phipps 6574 (UWO). Allendale Co.:
(4) 1 mi E of Martin, 11 Sep 1956, CR. Bell 5183 (GH, NCU, NY); no locality, 06 Apr 1953, WA. Kelley and WT. Batson s.n. (USCH).
Anderson Co.: (3) 2.5 mi SE of Fair Play, 31 May 1956, A.E. Radford 11992 (2*GA, NCU). Barnwell Co.: (2) Bowl State Park, 17
Oct 1956, C.L. Porter 343 (USCH, UWO). Calhoun Co.: (1) 4.3 mi ENE of St Matthews, 14 Apr 1968, S.W. Leonard 1264 (NCU).
Chesterfield Co.: (1) US 1 and Co. 20 S of Cheraw, 16 May 1976, J. Solomon 1951 (MO). Clarendon Co.: (1) no locality, 30 Jul
1982, JF. Logue 8L5635 (USCH). Darlington Co.: (2) Hartsville, 03 Jun 1941, D.E. Smith 955 (NCU); 10 May1920, A. Rehder 965
(A). Edgefield Co.: (1) 4.5 mi S of Trenton 01 Jun 1957, A.E. Radford 26442 (NCU). Kershaw Co.: (4) 6 mi SW of Camden, 14
Jul 1933, T.G. Harbison s.n. (2*NCU); SW of Camden, 15 Jul 1933, 7G. Harbison s.n. (NCU); 6 mi SW of Camden, 17 Jul 1933, 7.G.
Harbison s.n. (NCU). Lexington Co.: (2) Bethel church, 13 Apr 1974, D.E. Boufford 12808 (MO); 2 mi E of Edmund, 25 Apr 1980,
J.B. Nelson and WT. Batson 1527 (FSU). Richland Co.: (2) E of Old Hartsville Guard, 05 Apr1991, J.B. Nelson 10242 (USCH); 13.4
i W of Camden, 14 Aug 1958, H.E. Ahles and J.A. Duke s.n. (NCU). Sumter Co.: (1) 5 mi WNW of Pinewood, 19 Apr 1957, A.E.
Radford 20984 (NCU). ?? Co.: (1) Sandhills “A; 11 Jul 1952, WA. Kelley 419 (USCH).
S purus (Nash) Beadle
ALABAMA. Barbour Co.: (1) Lake Pt. State Park, 5 Jun 1992, DJ. Drennan 00044 (UWO). Calhoun Co.: (1) Anniston Natural
History MET 30 Aug 1998, J.B. Phipps aid E und 7772 (UWO). Dallas Co.: (1) Rte. 140, 8-10 mi E of Selma, 10 Apr
1998, J.B Phipps 7683 (UWO). Macon Co.: (1) 7 NE of Tuskegee, 20 Jul 1967, Ross C. Clark 16456 (NCU). Pike Co.: (1) S of
Brunridge, 17 Oct 1927, WW. Ashe &n. a. Co.: (1) Jct. Kellytown Rd and Valley Rd, 23 Sep 1999, J.B Phipps and
nL: In I Dawe? £ frat H I H $ 1155
R.J. OKennon 8071 (UWO). FLORIDA. Alachua Co.: (12) Hawthorn, 27 Feb 1933, JW. Kea s.n. (FLAS); Gainesville, 7 Apr 1931,
EJ. Palmer 38416 (A); Gainesville, 25 Jul 1940, Wilmot and W.A. Murrill s.n. (FLAS, GA); Hawthorn, 4 Aug 1940, Watson and WA.
Murrill s.n. (FLAS); Gainesville, 6 Mar 1952, G.R. Cooley 1443 (USF); Gainesville, 25 Sep 1940, WA. Murrill s.n. (GA); Gainesville, 31
Mar 1940, WA. Murrill s.n. (FLAS); Gainesville, 15 Jul 1936, Survey Pasture s.n. (FLAS); Gainesville, 17 Aug 1940, WA. Murrill s.n.
(FLAS); Gainesville, 12 Apr 1937, WA. Murrill s.n. (FLAS); Paynes Prairie Nature Reserve, Chacala Trail, 21 Mar 1993, J.B Phipps
6660 (UWO). Baker Co.: (2) near Macclenny, 24 May 1941, WA. Murrill s.n. (GA); E side of Trail Bridge, 5 Jun 1936, H.H. Hume s.n.
(FLAS). Citrus Co.: (4) 2 mi N of Pineola, 11 Mar 1956, R. Kral 2069 (FLAS, FSU); Rose Hill, 21 Apr 1959, G.R. Cooley 6460 (GH); 4 mi
W of Hernando on SR486 at Anthony Ave, 15 Aug 1993, - B en ee lie hd Co.: (2) Green Core Springs, 26 Jun 1940,
WA. Murrill s.n. (NCU); Green Core Springs, 26 Jun 1940, J W.A. Murrill s.n. (NCU). Columbia Co.: (2) Lake City, 30
Jul 1927, E. Manning 1350 (GH); 6 mi E of Branford, 1 Oct i H. Kurz s.n. uM De Soto Co.: (2) Old Pine Level Community
Centre, 28 Aug 1976, A.G. Shuey 170: (2*USF). Duval Co.: (1) Jacksonville, 17 Mar 1893, A.H. Curtiss 4216 (NY). Escambia Co.: (1)
University of Fla. Grounds, 24 Jun 1983, R.K. eid 80734 (FSU). Gadsden Co.: (3) 6.8 mi E of Chattahoochee, 23 Apr 1967, S.
McDaniel 8757 (FLAS); 12 mi W of Quincy, 6 Sep 1940, W.B. Tisdale and WA. Murrill s.n. (FLAS); 14 mi W of Quincy, 16 Sep 1940,
W.B. Tisdale and WA. Murrill s.n. (FLAS). sepu e (1) 6 mi N of Brooksville, 30 Jul 1961, O. Lakela 24508 (USF). Highlands
Co.: (4) Sebring, J.B. McFarlin s.n. (FLAS); Sebring, 9 Mar 1948, R. Garrett s.n. (GA); Sebring, 24 Jul 1949, R. Garrett s.n. (FLAS); NE
of Sebring, 1 Aug 1948, R. Garrett and L.E. Arnold 20 (FLAS). Hillsborough Co.: (5) 0.8 mi from Polk Co. line, 26 Mar 1961, O.
Lakela 23944 (FLAS, USF); Hillsborough River State Park, 3 May 1974, S.M. Kreps s.n. (USF); no locality, 3 Aug 1971, O. Lakela 32161
(USF); Old Field, 13 Mar 1904, A. Fredhold 6335 (GH). Jackson Co.: (3) Cottondale, 5 Apr 1923, T.G. Harbison 6077 (A, NCU); N of
Sneads, 10 Jun 1982, R.K. Godfrey 79855 (UWO). Lake Co.: (10) no locality, 2 Apr 1978, K. and J. Daubenmire 126 (USF); Eustis,
1-15 Jul 1894, G. Nash 1142 (DOV, GH, MO); no locality, 28 Feb 1944, FWH 17911 (GH); no locality, 13 Mar 1981, RP Wunderlin
8880 (USF); Mt. Plymouth. 26 Nov 1960, B. Moore s.n. (FLAS); Leesburg, 17 Aug 1939, Rude and W.A. Murrill s.n. (FLAS); Eustis, 5
Aug 1901, Biltmore Herb. 4022 (A); no locality, 14 May 1929, WW. Ashe s.n. (NCU). Leon Co.: (2) E of Tallahassee, 23 Aug 1951,
H. Kurz s.n. (FSU); 7 mi NW of Tallahassee, ot a nie: Liberty Co.: (2) Tallahassee, 6 Oct 1920, T.G. Harbison 5711 (A);
Tallahassee, 27 Aug 1902, Biltmore Herb. 6043 (GH). Madi o.: (3) Cherry Lake, 22 Aug 1940, WA. Murrill s.n. (FLAS); Cherry
Hill, 29 May 1942, WA. Murrill s.n. (FLAS); EE 5 Sep ón W.B. Tisdale and W.A. Murrill s.n. (FLAS). Marion Co.: (3) Sum-
merfield, 11 Aug 1939, WA. Murrill s.n. (FLAS); o Springs, 7 Jun 1984, R.F Thorne 57941 (USF); 1 mi S of Reddick, 15
Sep 1940, WA. Murrill s.n. (GA). Orange Co.: (8) 1. W of McDonald, 24 Feb 1918, C. Barker 476 (3*A); 1.5 mi W of McDonald,
24 Feb 1918, C. Barker 475 (A); 2 mi £ posue zi 1918, C. Baker 476 (A); near oe 7 Jul d C Due JE
of Mt. Dora, 17 Aug 1939, WA. Murrill s.n. (FLAS); 1.5 mi N of Plymouth, 23 Apr 1982, B. Han 9 JSP).
Polk Co.: (9) Frostproof, 16 Mar 1949, R. Garrett s.n. (FLAS); Haines City, 17 Mar Ts J eae 4511 (A); 15 mi N of Lakeland,
28 Mar 1964, R. Wall 34 (USF); Lake Hamilton, 26 Sep 1962, H.S. Conrad s.n. (FLAS); no ere no date 1919, Grace K. Jennings
6346 (CM); no locality, Dec 1919, Grace K, Jennings s.n. (UWO); Lake Hamilton, 17 Mar 1931, EJ. Palmer 4447 (A); Haines City,
4 Ale 1 c J. McFarlin 4510 (A); 3 rni NNW of West Frostproof, N of Xeriscape Nursery, 15 Aug 1993, J.B Phipps 6748 (UWO).
o.: (9) near Edgar, 10 Nov 1940, Watson and WA. Murrill s.n. (FLAS); 5 mi S of Melrose, 29 Mar 1942, WA. Murrill s.n.
ae near Bn Lake, 10 Nov 1940, WA. Murrill s.n. (GA); Johnson, 9 Jun 1940, WA. Murrill s.n. (FLAS); 0.8 mi W of Putnam
Hall, 11 Mar 1993, J.B. Phipps 6650 (UWO); 2 mi N of SR100 on SR309, 11 Mar 1993, J.B. Phipps 6649 (UWO); 3.1 mi E of Melrose,
8 Jun 1982, 5. Humphrey 126 (FLAS); Cowpen Lake, 4 Aug 1940, W.A. Murrill s.n. (A); N side Putnam Hall, 11 Mar 1993, J.B Phipps
6651 (UWO). sumiar o (2) St. Catherine's Island, 25 Aug 1916, T.G. Harbison 12707 (NCU); 1.5 km NE of Center Hill, 10 Mar
1993, J.B Phipp NO). Suwanee Co.: (3) Branford, 21 Aug W.B. Tisdale and WA. Murrill s.n. (FLAS); Hildreth, 10 Jul
1940, WA. Murrill s.n. (A (A i Between Branford and Hildreth, 17 Mar 1 Exploration Party 1939 (FLAS). Wakulla Co.: (3) Tiger
Hammock, 13 Aug 1951, H. Kurz s.n. (FSU); SW of jct. US Rte. 198 Co. a 365. 21 May 1983, R.K. Godfrey 80641 (UWO); SW of jct.
US98 and Co. Rd.365, 21 May 1983, R.K Godfrey 80643 (UWO). ?? Co.: (3) no locality, Jun 1894, A.S. Hitchcock s.n. (MO); no date,
S.B. Buckley s.n. (MO); no date, A.W. Chapman s.n. (NY). GEORGIA. Decatur Co.: (1) 6 mi NE of Bainbridge, 15 Jul 1976, RA.
Norris 3178 (BE, Dodge Co.: (1) NE of Little Ocomulgee, 6 Jul 1966, J.R. B. 5412 (NCU). Houston Co.: (1) no locality, 1
Jun 1979, D. Dixon 1398 (NATC). Jefferson Co.: (2) 7 mi SW of Wrens, 20 Jun 1941, W. Duncan 3370 (GA, NCSC). Liberty Co.:
(6) 10 mi E of Taylor's Creek, ^ s ain W. md 2713 (A, DHL, 3*GA, NCSC). Lowndes Co.: (1) 5.5 mi NW of Valdosta, 3
May 1973, W. R. Faircloth 7254 o.: (1) no locality, 8 Jul 1966, J. R. B 5745 (NCU). Randolph Co.: (1)
Cuthbert, 29 Jul 1928, WW. aes (NCI ). dde d Co.: (3) Augusta, 31 Mar 1923, T.G. Daño 6059 (A); Augusta, 11 Apr,
T.G. Harbison s.n. (NCU); Augusta, 25 Jul 1900, Biltmore Herb. 2779 s ud Co: a Hwy. 82 W of Graves, 1 Jun 1992, DJ.
Drennan 00041 is: Mic e Co.: (1) On Altamaha River, 26 Jul 1941, 49 (GA). Webster Co.: (1) Negro Church,
no date, s.n. (NCU). SOUTH CAROLINA. Aiken Co.: (2) no locality, 1 e 1952, WR. Kelly 23 ET no locality, no date 1910,
W.W. Ravenel s.n. ia (1) 2 mi out of Barnwell, 15 Jul 1933, T.G. Harbison s.n. (NCU). Beaufort Co.: (3) no locality,
25 Aug 1916, I.G. Harbison s.n. (A); no locality, 31 May 1930, EJ. Palmer 5496 (A); Ladies Island, 30 Ms 1917, CF. Batchelder 4868
(GH). Calhoun Co.: (1) 4.3 mi ENE of St. Matthews, 3 Jul 1957, H.E. Ahles 30299 (NCU). Lexington Co.: (3) near Lexington, 19
Mod 1964, hi eie (USCH); a 2 mi NE of Lynches River, 24 Aug 1958, J.A. Duke 2258 (NCU); W side US321, 1.8 km S of 1-26,
Aug 1993, Dp WO). Richland Co.: (1) no locality, 29 Jun 1964, J.R. Russell 22 (USCH).
TT
1156 J | of the Botanical R h Institute of Texas 2(2)
Crataegus invicta Beagle.
GEORGIA. Appling C N side Big Satilla Creek, 9 Jun 1967, J.R. B 9242 (NCU). Berrien Co.: (1) 3.2 mi W of Nashville,
4 May 1968, WR Faircloth 5122 (GA). Brooks hs ix M. 5 mi NNW ILE i a 1965, WR. Faircloth 2005 (FSU, MO, NCC); 3
mi E of Quitman, 27 Jun 1966, J.R. Bozeman CU). Colquitt Co f Funston, 21 May 1973, T. McCarty s.n. (MO)
Early Co.: (2) Railroad Station Scaffold off US Rte. 84, 1 May 1982, RK Godfrey 79713 (FLAS, FSU). Lanier Co.: (4) no locality, 2
May 1956, R. Kral 2493 (FSU); 7.8 mi NNE of Lakeland, 50 yards S of Mud Creek, 18 Jul 1966, WAR. Faircloth 3579 (GA, MO, NCO).
Wayne Co.: (1) 1.3 mi N of Glynn Co. line E of Mt. Pleasant, 21 Jul 1966, J.R. Bozeman 6205 (NCC). SOUTH CAROLINA. Allendale
o.: (2) 0.1 mi NNE jct. Co. Rte. 49 and 26 on Co. Rte. 49, 13 May 1956, H.E. Ahles and CR. Bell 12522 (NCU, SC).
Fei e Small
ALABAM tauga Co.: (1) near Prattville, 10 Apr 1966, A.C. Koelling s.n. (AUA). Baldwin Co.: (2) no locality, 21 Mar 1888, E
Mohr s.n 200 Sandhills, 14 Oct 1923, WW Ashe s.n. (NCU). Dale Co.: (1) near W boundary of Dale Co, 15 Apr 1931, EJ.
Palmer 38645 (A). Dallas Co.: (2) Selma, 11 Apr 1912, T.G. Harbison 10831 (NCU); Selma, 10 Apr 1912, T.G Harbison s.n. (A). Pike
Co.: (4) 5 mi SE of Troy, 29 Sep 1979, S. McDaniel and C. Duncan 22940 (TENN); 5 mi ESE of Troy, 28 or 29 Sep 1966, 5. McDaniel
7805 (FSU, VDB); On hwy. 19, 4 mi E of hwy. 223, 11 Sep 1986, A.R. Diamond 2881 (AUA). Russell Co.: (2) 1 mi N jct. US 931 on
Ala 169, 22 Jul 1968, R. Kral 31806 (GA, UWO). FLORIDA. Alachua Co.: (3) W side of Gainesville, 22 Mar 1968, W.G. D'Arcy 2370
(UWO); Paynes Prairie, Gainesville, 10 Aug 1940, W.A. Murrill s.n. (FLAS); SW of Payne's Prairie, Gainesville, 21 Aug 1940, WA.
Murrill s.n. (FLAS). Bay Co.: (22) 2 mi W of Rte. 231 jct. with FL Rte. 20., N of Youngstown, 14 Mar 1982, R.K. Godfrey 79413 (FSU);
2 mi W of Rte. 231 jct. en FL Rte. 20 by Rte. 20, N of Youngstown, 14 Mar 1982, R.K. Godfrey 79412 (FSU); 2.5 mi W of jct. FL
Rte. 20 and US Rte. 231 by Rte. 20, 02 Apr 1982, R.K. Godfrey 79509 (10*FSU); 1 mi W of Calhoun-Bay line by FL Rte. 20, W of
Clarksville, 14 Mar 1982, R.K. Godfrey 19411 (FSU); 1 mi E of Econfina Creek by FL Rte. 20, 14 Mar 1982, R.K. Godfrey 19414 (FSU);
Street perpendicular to Pearl Ave. Panama City Beach, 06 Nov 1982, R.K. Godfrey 80187 (FSU, UWOJ; 1.8 mi W of jct. of FL Rte.
20 and US Rte. 231 by Rte. 20, 02 Apr 1982, R.K. Godfrey 79508 (UWO); 1.8 mi W of jct. of FL Rte. 20 and US Rte. 231 by Rte. 20,
02 Apr 1982, R.K. Godfrey 79507 (2*FSU, UWO); 8 mi N of jet. Fla. 20 along Fla. 231 2 mi S of Jackson Co. line, 11 Apr 1966, J.
Beckner and W. si 3 (FLAS); 0.1 mi W of jct. with Fla. 77 on Fla. 20, 01 May 1982, R. Wunderlin and J. Beckner 9325 (USF).
Calhoun Co.: (2) 16 mi W of Altha, 10 May 1964, S. McDaniel 4431 (FSU, IBE). Escambia Co.: (14) N North Pensacola, 12 Jul 1969,
R.K. Godfrey 68622 (FSU); Univ. W FL campus grounds, 24 Jun 1983, R.K. Godfrey 80734 PA no locality, 07 Apr hen ES:
Sargent 7665 (NY); near Pensacola, 14 Apr 1889, CS. Sargent s.n. (A); Pensacola, 07 Apr 1900, C.S. Sargent s.n. (A, GH ); Pensa-
cola, 14 Apr 1894, T.G. Harbison s.n. (A); In Pensacola, opposite Bayview Mem. Cemetary, e Mar bis dd 9980 (usn) Pa
mi W of eastern gate of reservation on Santa Rosa Island, 30 Mar 1967, C. and S. Chapman 0455 (FLAS). Hill h Co.: (3)
Northeast border of USF campus, 09 Sep 1960, J. Ray Jr. 10226 (USF); USF Campus, 11 Mar 1961, O. ps 23807 (USP); USF
ee 02 Ag 1961, O. Lakela 24519 (USF). Holmes Co.: (1) Choctawatchee River, W of Millers Crossroad by FL Rte. 2, 17 Mar
2, R.K. Godfrey 79423 (FSU). Levy Co.: (1) By US Rte. 19 between Gulf Hammock and Lebanon Station, 26 Mar 1983, R.K.
cd 80362 (FSU). Liberty Co.: (1) Torreya State Park, 04 Jul 1972, R.K. Godfrey 71510 (FSU). Marion Co.: (2) Dunnellon, 11
Aug 1940, WA. Murrill s.n. (FLAS); About 1 mi NNW of Dunnellon W of US 41, 15 Aug 1993, J.B. Phipps 6744 (UWO). Okaloosa
Co.: (70) Valparaiso, 09 Aug 1928, WW. Ashe s.n. (NCU); 4.8 km N of US Rte. 90, 27 May 1973, D.E. Boufford and H.E. Ahles 9484
(A); 1 mi. N of Deerland, 02 Aug 1954, ES. Ford 3886 (FLAS); 2 mi. NW of US 90 on FL Rte. 4, 20 Aug 1950, H Kurz s.n. (2*FSU);
Valparaiso, 23 Mar 1927, WW. Ashe s.n. (5*NCU); 3 mi N of Fort Walton, 02 Apr 1966, R. Kral 26149 (FSU, GA, TENN, UWO); near
Shoal River on US Rte. 90 and E of town of Crestview, 27 May 1973, D.E. Boufford and H.E. Ahles 9492 (A); N of Niceville along
FL 85, 19 Jun 1956, Mr. and Mrs. H.A. Davis 11604 (WVA); 2 mi W of Laurel Hill, 02 Aug 1961, R.K. Godfrey 61299 (FSU); 200 yds N
of Rd 220 ca. 8 mi NNE of Niceville, 24 Aug 1966, D.B Ward and C. Chapman 5987 (FLAS); 3.5 mi SW of Laurel Hill, 02 Aug 1961,
R.K. Godfrey 61316 (FSU); near jct. of Eglin Rd. 238 and Eglin Rd. 40, W of head of Rogue Creek, 22 Jul 1977, K.D. Perkins with J.B.
Nelson 340 (FLAS); 0.4 mi E of Shoal River on US 90, 21 May 1976, J.C. Solomon 5621 (FSU); along Rte. 90 about 1 mi E of Shoal
River, 27 Jul 1981, D.S. Correll and H.B 52224 (USF); Jct. of US Rte. 90 and Shoal River, then 3 mi N on dirt road NE of Crestview,
27 May no locality, no date 1973, H.E. Ahles with D.E. Boufford 77031 (CM); 17 mi NW of Ft Walton Beach, 10 May1967, D.B. Ward,
C. Chapman and R.R. Smith 6396 (FLAS, LAF, NCU); S of Shoal River on FL Rte. 87, 04 Jun 1951, H. Kurz s.n. (FSU); Milligan, 05 Apr
1949, S.C. Hood 1829 (FLAS); 1 mi N of Niceville Center by FL Rte. 85, 02 Apr 1982, R.K. Godfrey 79518 (FSU, UWO); N side of US
90 about 8 km E of Crestview 0.5 km W 4 lane hwy, 08 Apr 1991, J.B. Phipps 6476 (UWO); 2.5 mi E of Niceville by FL Rte. 20, 02
Apr 1982, R.K. Godfrey 79517 (FSU, UWO); rest area on 110 ca. 1 mi E intersection with hwy. 85 5 of Crestview, 23 May 1989, LE.
Brown 13628 (UWO); 1 mi N of Niceville Center by FL Rte. 85, 02 Apr 1982, R.K. Godfrey 79519 (FSU, UWO); 3.4 mi E of Holt by
US Rte. 90, 02 Apr 1982, R.K. Godfrey 79521 (FSU, UWO); near Deerland, 27 Apr 1956, R.K. Godfrey 54630 (DUKE, FLAS, FSU, NCU,
UWO); US 90, 6 mi E of Crestview, 14 Aug 1993, J.B. Phipps 6736 (UWO); US 90, 6 mi E of Crestview, 14 Aug 1993, J.B. Phipps
6739 (UWO); US 90, 6 mi E of Crestview, 14 Aug 1993, J.B. Phipps 6738 (UWO); US 90, 6 mi E of Crestview, 14 Aug 1993, J.B.
Phipps 6737 (UWO); US 90, 6 mi E of Crestview, 14 Aug 1993, J.B. Phipps 6740 (UWO); US 90, 6 mi E of Crestview, 14 Aug 1993,
J.B. Phipps 6735 (UWO); N side of US 90 about 8 km E of Crestview 0.5 km W of 4 lane hwy., 08 Apr 1991, J.B. Phipps 6475 (UWO);
Hwy. 4 just S of Baker and 2.9 km NW of Milligan at US 90, 14 Aug 1993, J.B. Phipps 6734 (UWO); Hwy. 4 just 5 of Baker and 2.9
km at Richmond Rd. and 2.9 km NW of Milligan at US 90, 04 Apr 1991, J.B. Phipps 6473 (UWO); Hwy. 4 just S of Baker and 2.9
km at Richmond Rd. and 2.9 km NW of Milligan at US 90, 04 Apr 1991, J.B. Phipps 6474 (UWO); Crestview rest stop on 110 E, 1
Jun 1989, DJ. Drennen 20 (UWO); 4.6 mi E of Holt by US Rte. 90, 02 Apr 1982, R.K. Godfrey 79520 (FSU, UWO); Milligan, 05 Apr
~
DL: In [| Das: £ft Hi I 1 $ 1157
1949, S.C. Hood x (FLAS); Rte. 85, 2 mi SW of Valparaiso, 04 Apr 1975, LJ. Uttal 10996 (NCU); 3 mi N of Ft. Walton, 02 Apr 1966,
R. Kral 26149 (IB , 28 Mar 1951, EL. Tyson 537 (FLAS); Rest area beside I-10 1 mi E of Shoal, 29 Mar 1975, L.L. Anderson
3922 (FSU); W side of Rte. 85 ts mi ie of bridge over Garnier Bayou, 15 Mar 1982, G. Wilhelm 9945 (USF); Valparaiso, 23 Mar, T.G.
Harbison s.n. (NCU); 1 mi NE of Ft. Walton, 11 Sep 1950, H. Kurz s.n. (FSU); 3 mi E of Shalimar and S of Fla 85, J. Beckner and W.
DArcy 1119 (FLAS); Crestview, 08 Apr 1899, CS. Sargent s.n. (A); Valparaiso, 27 Mar 1918, WW. Ashe s.n. (NCU); Valparaiso, 26 Mar
192, WW. Ashe s.n. (NCU); Tallahassee, 12 Jun 1950, H Kurz s.n. (FSU); W of Crestview, 20 Apr 1982, H.B. and DS. Correll 53779
(NCU). Santa Rosa Co.: (6) 17 mi E of Pensacola on US Rte. 98, 11 Aug 1950, H Kurz s.n. (FSU); Rte. 87, 8.5 km S of 110, E side of
road, 08 Apr 1991, J.B. Phipps 6472 (UWO); 1 mi W of Wynn Haven Beach by US 98, 04 Apr 1982, R.K. Godfrey 79562 (FSU, UWO);
FL 878.5 km 5 of 110 E side of road, 14 Aug 1993, J.B. Phipps 6733 (UWO); FL 87 8.5 km S of 110 E side of road, 14 Aug 1993, J.B.
Phipps 6732 (UWO). Volusia Co.: (2) Haw Creek, Seville, 16 Jul, A.H. Curtiss 6203 (2*A). Walton Co.: (14) 3 mi. S of Mossy Head
jet, 19 Jul 1966, C. Chapman and G. Crosby and R.R. Smith 1069 (FLAS); Along FL 285, 2.8 mi SW of jct. UC Rte. 90, 14 Sep 1970,
D.B. Ward and J. Hunter 7421 (FLAS); S of US 98 W of Grayton Beach, 13 Jul 1971, H.A. Davis 15687 (FLAS); FL 280, 4 mi S of Mossy
Head, 24 Aug 1966, D.B. Ward and C. Chapman 5998 (FLAS); 7 mi W of DeFuniak springs N of hwy. 90, 10 Aug 1957, ES. Ford
5289 (FLAS); along FL 285 ca. 3 mi S of jct. with US 90 at Mossy Head, 29 Jun 1966, J. Beckner, C. Chapman and R.R. Smith 1353
(FLAS); 1 mi E of Sandestin Beach by US Rte. 98, 04 Apr 1982, R.K. Godfrey 79563 (2*UWO); DeFuniak Springs, 14 Apr 1899, T.G.
Harbison s.n (A); DeFuniak Springs, 14 Apr 1931, EJ. Palmer 38619 (A, GH); near DeFuniak Springs, 14 Apr 1931, EJ. Palmer 38617
(A, MO); 6.5 mi W of Portland, 11 Aug 1966, C. Chapman 1258 (FLAS). Washington Co.: (3) on Orange Hill W of FL 273 and ca.
7 mi SE of Chipley, 16 Oct 1972, D.B. Ward and N.E. Lee and A. Gholson s.n.(FLAS); S of Chipley, West FL, 26 Mar 1928, WW. Ashe
s.n. (NCU); Holmes Valley, 26 Mar 1928, WW. Ashe s.n. (NCU). ?? Co.: (2) Nat. Forest ieu a pen d 26 Mar 1927,
WW. Ashe s.n. (NCU); Florida Nationa! Forest, 26 Mar 1927, WW. Ashe s.n. (NCU). GEORGIA o.: (1) Ft. Benning
Military Res., 16 mi SE of Columbus, 03 Apr 1980, R.R. Haynes 7665 (ALU). Cherokee Co.: (1 " ) Canton Rd 140, RR3, Box 1180, 19
Apr 1984, E.G. Meyer and PM. Mazzeo 20064 (UWO). Clarke Co.: (1) near River by Bobbin Mill, 08 Apr 1930, J.M. Reade E4634
(UWO). Lanier Co.: (1) 2.3 mi SSE of Lakeland, 26 Oct 1968, G. Lloyd, N. Gillis and W.R. Faircloth 5653 (GA). Lowndes Co.: (1) 10
mi WSW Valdosta, 30 Aug 1968, R.A. Norris 1051 (IBE). Madison Co.: (1) Danielsville 2 mi NE, 19 Aug 1927, KM. Wiegand and
WE. Manning 1357 (GH). Randolph Co.: (1) Cuthbert, 09 Sep 1928, T.G. Harbison s.n. (NCU). Talbot Co.: (4) Rte. 96, about 1 mi
E of Junction City, 13 Aug 1993, J.B. Phipps 6712 (UWO); Rte. 96, about 2 km ENE of Geneva, 09 Apr 1991, J.B. Phipps 6488
(2*UWOJ; 2 km ENE of Geneva, 13 Aug 1993, J.B. Phipps 6720 (UWO). SOUTH CAROLINA. Aiken Co.: (3) S.C. 391 11.2 mi S of
Batesburg, 18 Jul 1958, J.A. Duke 1512 (NCUJ; Woodlanders Nursery, 1128 Colleton Ave, 11 Apr 1984, KG. Meyer and PM. Mazzeo
19864 (UWO); 1 mi E of Wagener, 24 Dec 1941, WC. Coker s.n. (NCU). Lexington Co.: (1) Batesburg, 29 Apr 1911, EA. McGregor
609 (US). Orangeburg Co.: (1) 5 mi W of Orangeburg on US Rte. 301, 26 Sep 1957, H.E. Ahles 35077 (NCU).
—
~
—
MA picis Beadle
ALABAM arte On (2) Eufaula NWR at entrance of old hwy. 165, 06 na 1993, DJ. Drennen M ell Bird ad at
Baker Co des N y on old hwy. 165, 30 yrds E of fireline, 19 August 1993, DJ. D
20 Apr 1916, R.S. Cocks s.n. (NO); Sardis, 15 Apr 1918, R.S. Cocks s.n. (NO); Selma, 12 Apr 1912, T.G. hs 842 (HUH). Greene
Co.: (1) Entrance to Smith Lake, 14 Apr 1967, T.D Naugle T995 (ALU). Lee Co.: (1) Phoenix City, 28 Sep 1906, T.G. Harbison 10045
NCU). Russell Co.: (2) Phoenix City, 26 Sep 1905, T.G. Harbison 10044 (2*NCU). Tuscaloosa Co.: (1) Tuscaloosa, 05 Apr 1927,
WW. Ashe s.n. (NCC). FLORIDA. Citrus Co.: (11) 4.5 mi NE of Holder, 3 May 1981, L. Baltzell and D.W. Hall 11387 (2*FLA, FLAS,
WILLI); Withlacoochee State Forest, 27 May 1973, P Genelle and G. Fleming 1883 (GA, NCU, TENN, USF); Flora City, 19 Apr 1931,
EJ. Palmer 38393 (HUH); Pineola, 03 Aor 1930, ES. Blanton 7000 (2*HUH). Columbia Co.: (2) no locality, 21 Jul 1918, T.G. Harbison
s.n. (HUH); Lake City, 21 Jul 1918, 7.G. Harbison 14576 (NCU). Hernando Co.: (4) Chinesgut p Refuge, 03 Apr 1958, G.R.
Cooley and R.J. Eaton 5865 (USF); 0.7 mi S of Nobleton, 17 Jun 1976, L. ori and D.B. Ward 8620 (FLAS); 4 mi NE of Brooksville,
18 May 1965, J. Beckner and D.B. Ward 4694 (FLAS, WILLI). Jackson Co.: (2) 1.4 mi W of Sneads, 19 May 1966, R.K. Godfrey 67637
(FLAS, FSU). Lake Co.: (1) Vacant lot in Tavares, 15 May 2000, A.H. Curtiss 6617 pd Liberty Co.: (2) Telogia, 23 Jul 1940, Arnold
and West s.n. (FLAS); Rock Bluff, 06 Aug 1927, K.M. Wiegand and WE. Manning 1355 (HUH). Marion Co.: (1) Summerfield, 11 Aug
1939, WA. Murrill s.n. (FLAS). Sumter Co.: (1) Sumterville, 04 Apr 1946, Arnold and dis n. D Suwanee Co: (1) Hildreth,
10 Jul 1940, WA. Murrill s.n. (FLAS). Washington Co.: (1) 1 mi W of Miller's 03 Apr 1958, R.K. God 74 (FSU). Volusia
Co.: (1) Low woods bordering Haw Creek, 07 Jun 1900, A.H. Curtiss 6677 (GA). GEORGIA. pte Co.: (1) 5.3 km S of Co. line
on US 441, 01 Jun 1973, H.E. Ahles and D.E. Boufford 9693 (A). Chattooga Co.: ex Summerville near Augusta, 03 Sep 1914, TG.
Harbison 2=1041 (HUH). De Kalb Co.: (1) Bordering Yellow River near Stone Mt., 15 Aug 1899, A.H. Curtiss 6520 (MO). Gadsen
Co.: (1) 1 mi S of River Jct, 06 Sep 1940, WA. Murrill and W.B. Tisdale s.n. (FLAS). Gwinett Co.: (8) McGuire's Mills, 11 Sep 1902,
Biltmore Herb. 7105 (2*GH); On the Yellow River near McGuire's Mills, 17 May 1895, J.K. Small s.n. (NY); Yellow River, 11 Sep 1902,
C.D.Beadle 7105 (Gray, HUH); Rocky woods near Stone Mt, 15 Aug 1899, A.H. Curtiss 6519 (DOV, HUH); Rocky woods bordering
Yellow River near Stone Mt, 15 Aug 1899, A.H. Curtiss 6520 (MO). Hart Co.: (1) 5 mi SW of Hartwell, 20 May 1934, EJ. Palmer
42445 (HUH). Lee Co.: (1) 3 mi SE of Smithville, 11 May 1940, WH. Duncan 2306 (GAM). Lincoln Co.: (1) 3 mi W of Liston, 02
Sep 1940, WH. Duncan 2832 (DUKE). Long Co.: (1) 0.5 mi N of Beard's Bluff, 10 Apr 1963, J.R.Bozeman 2534 (NCU). Lowndes
Co.: (3) 10.3 mi S of Valdosta, 28 Jun 1965, WR. Faircloth 1902 (2*GA, NCU). Madison Co.: (1) 2 mi W of Comer on S fork of
the Broad River, 26 Apr 1942, W.K. Duncan 4781 (GA). Richmond Co.: (9) 1.5 mi E of GA-56, 04 Aug 1966, J. R. Bozeman 6952
(2*NCU); Summerville near Augusta, 3 Oct 1914, 7G. Harbison 1041 (A); Augusta, S.T. Olney and J. Metcalfe s.n. (HUH); Augusta,
as
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El n : A/D h Inet.
1158 Journal of tute of Texas 2(2)
S.T. Olney and J. Metcalfe 29 (NCSC); Augusta sandhills, 08 Jul 1900, A. Cuthbert 3 (NCU); Augusta, 14 Aug 1902, A. Cuthbert s.n.
(2*FLAS); Augusta, 09 Apr 1919, T.G. Harbison s.n. (HUH). Tatnall Co.: (1) NE of Altamaha river, 12 Jul 1966, J.R. Bozeman 5967
(NCU). Wilkinson Co.: (1) 5.3 km S of Baldwin Coline on US Rte. 441, 01 Jun 1973, D.E. Boufford and H.E. Ahles 9693 (HUH).
NORTH CAROLINA. Brunswick Co.: (1) N of Southport, Pineland, 24 Apr 1935, A.C. Matthews s.n. (DUKE). Cumberland Co.:
(1) near Stedman, 7.G. Harbison s.n. NED MEN Hanover Co.: (3) Carolina Beach, 14 May 1937, Mills and Shrink s.n. (NCSC); Mill
Pond, Wilmington, 28 Jun 1905, J.M. A (DUKE); Cape d 28 Jul 1949, WB. Fox 2820 (NCSC).
Sampson Co.: (1) near Stedman across the river, 28 Apr 1933, T.G Harbison and LR. Totten s.n. (NCU). ?? Co.: (1) Western North
Carolina, 23 May 1901, WW. Ashe 1949 (CM). SOUTH CAROLINA. Aiken Co.: (1) y 278 about 2 km E of jct. with bush 3 m tall,
29 Apr 1992, J.B. Phipps 6573 (UWO). Chesterfield Co.: (1) Haw Ridge near McBee, 17 Jun 1933, 7.6. Harbison and H.R. Totten
n. (NCU). Kershaw Co.: (6) Camden, 12 May 1902, T.G. Harbison 6124 (HUH); Hartsville, 07 Apr 1935, BE Smith 219 (USCH); no
pere 18 May 1934, EJ. Palmer 42393 (SC); near Camden, 19 Apr 1932, EJ. Palmer 39953 (HUH); 4 mi W of Bethune, 07 Sep
1939, R.K. Godfrey 8021 (GH, NY). Lancaster Co.: (1) 0.4 mi NE of little Lynches Creek, 06 Jun 1957, ie Haesloop 27422 (NCU).
Newberry Co.: (2) Billy Dreher Island, 18 Aug 1971, L.H. Buff 410 (FLAS, USCH). Oconee Co.: (2) 4 mi NW of Walhalla, 04 May
1956, H.E. Ahles and Bell 13923 (NCU); no locality, 30 Apr 1926, T.G. Harbison 8008 (HUH). Richland P (1) Between 6" Division
Rd. and Wildcat Rd, 27 May 1992, J.B. Nelson and B.S. Long 12594 (USCH). Sumter Co.: (1) no locality, 13 Jun 1993, J.B. Nelson
14225 (USCH). VIRGINIA. Chesterfield Co.: (1) Cambridge, 5 Aug 1845, R. Gray s.n. (GH).
Crataegus lancei J.B. Phipps
ALABAMA. Barbour Co.: (1) Eufaula NWR, 15 Oct 1993, D. Drennan 95 (UWO). Clayton Co.: (1) no locality, 20 Oct 1930,
EJ. Palmer 5658 (A). Greene Co.: (1) Smith Lake area, 14 Apr 1969, J. Thomas 1975 (NCU). FLORIDA. Alachua Co.: (1) W of
Gainesville, 20 Aug 1940, W.A. Murrill s.n. (GA). Clay Co.: (1) Hibernia, Mar 1869, W.M. Canby 1278 (CM). GEORGIA. Burke
Co.: (1) River Road, 18 km N of US 301, 12 Aug 1993, J.B. Phipps 6692 (UWO). Dougherty Co.: (1) Albany, 18-19 Sep 1909,
W.W. Eggleston 5118 (MO). Habersham Co.: (2) Tallulah and Taccoa Falls, 8 Aug 1893, J.K. amal (GH, E Richmond
Co.: (1) Augusta, sand bar ferry, no date, A Cuthbert s.n. (FLAS). NORTH CAROLINA 24 Apr
1935, A.C. Matthews s.n. (NCU). B Co.: (2) NW of Weaverville, 3 May 1999, R. Lance s.n (UWO); N Stock Road and
Alpine Meadows, 21 Oct 1998, R. Lance 98-51 (UWO). Mecklenburg eo (1 nod no ds W.W. Ashe s.n. (NCU). Moore
Co.: (1) Drowning Oak, 8 May 1975, J.F. Matthews, E. Edwards & T.P. H j M o.: (1) Franklin, 27 Apr no year,
T.G. Harbison 520 (A). SOUTH CAROLINA. Chesterfield Co.: (1) Haw Ridge, 22 pus 1933, T.G. sonde s.n. (NCU). Kershaw
Co.: (1) SE side of US 1, 8 km S of Bethune, 11 Aug 1993, J.B. Phipps 6672 (UWO). Oconee Co.: (1) Walhalla, 30 Apr 1926, T.G.
Harbison 16481 (NCU). Richland Co.: (1) near Ft. Jackson, 20 Jun 1958, J.A. Duke 1342 (NCU).
Crataegus lassa Beadle
ALABAMA. Barbour Co.: (6) Eufaula NWR, 6 Apr 1993, DJ. Drennan 81 (UWO); SR id 15 Apr 1992, DJ. Drennan 00042
(UWO); Eufaula NWR, 19 Aug 1993, l Drennan ud eie Eufaula NWR, 3 Aug D 00059 (UWO); Lake Point
State Park, 2 Jun 1989, DJ. D NWR, 15 Apr 1992, DJ. D E a Dallas Co.: (18) Selma,
no date, T.G. Harbison s.n. (NCU); near Selma, 19 Apr 1920, T.G. Harbison s.n. (NCU); Selma, 15 e. 1970, R. Kral 38586 (IBE, FSU,
WILLD; Selma, 11 Apr 1912, TG. Harbison 10836 (2*NCU); Selma, 11 Apr 1913, T.G. Harbison 836 (A); Selma, 19 Apr 1915, 7.G.
Harbison s.n. (A); Selma, 12 Apr 1912, TG. Harbison 843 (A); Selma, 9 Apr 1920, 1.G. Harbison ud (NCUJ; E side of 80 outskirts
of Selma, 6 Apr 1984, J.B. Phipps and T.C. Wells 5324 (2*UWO); E side of 80 outskirts of Selma, 6 Apr 1984, J.B. Phipps 5326 (UWO);
E side of 80 outskirts of Selma, 6 Apr 1984, J.B. Phipps 5327 (UWO); E side of 80 outskirts of Selma, 6 Apr 1984, J.B. Phipps 5329
(UWO); Outskirts of Selma, jct. US3O & AL 41, 7 Apr 1991, J.B. Phipps 6471 (UWO); Rte. 140 8-10 mi E of Selma, 10 Apr 1998,
J.B. Phipps 7684 (UWO). Macon Co.: (3) no locality, 18 Jul 1967, James Wright s.n. (AUA); Tuskegee National Forest, 11 Apr 1976,
AH. Sessler 016 (AUA, UWO). Pike Co.: (1) 5 mi SE of Troy, 28 Jun 1970, R. Kral 39798 (UWO). Russell Co.: (1) Rte. 431 SW of
Phoenix City 2.9 km SW of Russell Co. Rd 41,8 Apr 1991, J.B. Phipps 6479 (UWO). FLORIDA. Alachua Co.: (2) March, A.H. Curtiss
811 (GA); Paynes Prairie Nature Reserve, Chacala Trail, 17 Aug 1993, J.B. Phipps 6760 (UWO). Citrus Co.: (2) Citrus Springs, NW
corner Athens Dr. and Sherman Dr,15 Aug 1993, J.B. Phipps 6745 (UWO); E side of US 41, 3 mi N of Hernando, 15 Aug 1993, J.B.
Phipps 6746 (UWO). Gadsden Co.: (3) Aspalaga, 5 Apr 1964, Sidney McDaniel 4101 (IBE, FSU); 3 mi S of River Junction, 6 Aug
1927, WE. Manning and KM. Wiegand 1353 (GH). Leon Co.: (2) Tallahassee, Sep 1901, Biltmore Herb. 4947 (A); Tallahassee, Apr
1901, Biltmore Herb. 4051 (A). GEORGIA. Burke Co.: (1) River Rd. A, 18 km N of US 301, 19 Apr 1992, J.B. Phipps 6567 (UWO).
Columbia Co.: (3) US 278, 4.5 km N Camp Gordon, 13 Aug 1993, J.B. Phipps 6702 (UWO); US 278, 4.5 km N Camp Gordon, 13
Aug 1993, J.B, Phipps 6703 (UWO); US 278, 4.5 km N Camp Gordon, 13 Aug 1993, J.B. Phipps 6704 (UWO). Randolph Co.: (1)
Cuthbert, 29 Mar 1918, T.G. Harbison s.n. (A). Richmond Co.: (5) Augusta, 16 Apr 1923, T.G. Harbison 7005 (2*A); Augusta, Jul,
A. Cuthbert 107 (FLAS); Augusta, 9 Apr 1919, T.G. Harbison 48 (A); Spur 56 5 km in, 18 Apr 1992, J.B. Phipps 6558 (UWO). Talbot
Co.: (5) Rte. 96 2 km E of Geneva, 9 Apr 1991, J.B. Phipps 6484 (UWO); Rte. 96 2 km E of Geneva, 9 Apr 1991, J.B. Phipps 6486
(UWO): Rte. 96 1 mi E of Junction City, 9 Apr 1991, J.B. Phipps 6494 (UWO); Rte. 96 2 km E of Geneva, 9 Apr 1991, J.B. Phipps
6481 (2*UWO). Taylor Co.: (1) 3 mi $ of Junction City, 15 Apr 1978, Mary P Grant 078 (AUA). SOUTH CAROLINA. Aiken Co.:
(1) Hwy. 4, 7 km E of junction w. Rte. 394, 16 Apr 1991, J.B. Phipps 6513 (UWO). Calhoun Co.: (2) 4.3 mi ENE of St. Mathews, 14
Apr 1968, S.W. Leonard 1264 (NCU, TC). Lexington Co.: (2) US 178, 4 km E of jet. 5C113, 20 Apr 1992, J.B. Phipps 6579 (UWO);
Rte. 302 Clay Hill, 10 Apr 1991, J.B. Phipps 6503 (UWO).
"T { Dvorsky, Revision of Crat aciari 1159
Crataegus lepida Beadle
FLORIDA. Alachua Co.: (1) 1 mi W of Fla. ??, 04 Apr 1981, K.E. Woeste 3 (WILLI). Citrus Co.: (1) About 1.5 mi W of US 98, Aug
1995, 5. Riefler s.n. (UWO). Clay Co.: (1) Middleburg, 26 Jun 1940, WA. Murrill and Bratley s.n. (FLAS). Highlands Co.: (16) 4 mi
N Lake Placid, 28 Feb 1961, E.J. Palmer 10400 (USF); 3 mi N Lake Placid, 25 Apr 1960, H.A. Gleason and R.J. Eaton 9757 (2*USF);
4 mi N Lake Placid, 07 Oct 1960, J.D.Ray, O. Lakela and J. Patman 10400 (FLAS, FSU, USF); E of Sebring 2.8 mi from jct. with 17
at Sebring, 11 Sep 1987, J.D. Skean and WS. Judd and P Alcorn 2132 (UWO); Sebring, 23 Apr 1964, LJ. Brass 33210 (FLAS); US 17
jct. N of RR near jct. of Lake Letta Rd, 9 May 1982, J. Eckner 2574 (USF); N of Lake Placid, 29 Sep 1079, W.S. Judd with D.B. Ward
and B. Judd 2504 (FLAS, FSU); Lakemont, 5 Sep 1981, R.P Wunderlin and J. Beckner 9127 (USF); Josephine Creek, 5 Oct 1960, LJ.
nen 15319 (FLAS); Avon Park, 10 Apr 1948, R. Garrett 171 (FLAS); S of Polk Co. Line, 3 Jun 1961, O. Lakela 24226 (USF); Sebring,
ep 1934, J.K. Small and E. West s.n. (FLAS). Hillsborough Co.: (2) 1.4 mi SE of centre of Sebring, 09 Mar 1993, J.B. Phipps 6631
uc 1.4 mi SE of centre of Sebring, 15 Aug 1993, J.B. Phipps 6749 (UWO). Osceola Co.: (4) 5 mi NW of Loughman, 14 Oct
1960, J.D. Ray, O. Lakela and J. Putman 10456 (FLAS, 2*USF); Kissimmee, 9 Apr 1938, M.L. Singletory s.n. (DUKE). Polk Co.: (4) At
Greenlefe Resort, 21 Jun 1979, D. Richardson and B. Hansen 5752 (USF); 3 mi E of Lake Hamilton, 10 Aug 1963, H.S. Conard s.n.
(FLAS); Vicinity of Great Masterpiece Lake, Wales, 28 Jul 1962, O. Lakela 25203 (USF); Marion Creek, 9 Sep 1931, J.B. McFarlane
6643 (USF).GEORGIA. Liberty Co.: (2) 4 mi SW of Hinesville, 22 Jul 1927, KM. Wiegand and WE. Manning 1348 (GH); 10 mi N
A 18 Apr 1954, W. Duncan 17804 (GA). Pierce Co.: (1) Blackshear, 28 Jun 1901, A.H. Curtiss 6824 (DOV). Screven
o.: (2) Rte. 24, 12 mi S of US 301, 12 Aug 1993, J.B. Phipps 6691 (UWO); Rte. 24, 12 mi S of US 301, 12 Aug 1993, J.B. Phipps
a o Tatnall Co.: (1) no locality, 12 Jul 1966, J.R. Bozeman 6001 (NCU). ?? Co.: (1) no locality, no date, H Corey s.n. (NY).
LOUISIANA. Richland Parish: (1) S of start of woods E of cemetery and LA 133 and N of I-20 Sec 4 and 9, 11 Apr 1978, R.D.
Thomas and N. Dawson 57507 and 1087 (NCU). SOUTH CAROLINA. Colleton Co.: (1) 0.7 mi SE jct. Co. 172 and 28, 05 Sep
1956, H.E. Ahles and C.R. Bell 17918 (NCU). Lexington Co.: (1) 4 mi W of Lexington, 09 Apr 1985, J. Seto, E. Tuggle, R. Wayson and
C. Smith-Aulback s.n. (USCH)
~
MA munda Beadle
ALABAMA. Baldwin Co.: (5) Gateswood, 02 May 1903, S.M. Tracy 8698 (GH, MSC, NCU, 2*US). Conecuh Co.: (1) S of Evergreen,
15 at 1923, WW. Ashe s.n. (NCC). Covington Co.: (1) 19 mi N of Florala on US 331, 27 Apr 1969, R. Kral 34546 (GA). Cullman
Co.: (1) St. Bernard, 11 May 1934, W. Wolf s.n. (NO). De Kalb Co.: (1) E side of Rock Creek about Y? mi from mouth, 25 Sep
1948, R.M. Harper 4096 (ALU). Escambia Co.: (1) Co. Rd 18 0.5 mi N of jct. with US 31-29, 07 Apr 1967, R.L. Clark 10017 (NCU).
Etowah Co.: (2) Gadsdale, 29 Jul 1900, WR. Maxon and C.L. Pollard 345 (A, GA). Troy Co.: (1) no locality, 2 Jul 1880, C. Mohr s.n.
(ALU). ?? Co.: (1) St. Bernard, 01 May 1910, W. Wolf s.n. (ALU). FLORIDA. Alachua Co.: (3) near Gainesville, 06 May 1940, WA.
Murrill s.n. (GA); no locality, 15 Apr 1930, Loucks and West s.n. (FLAS); E side of Gainesville, 11 Feb 1937, West and Arnold s.n.
(FLAS). Brevard Co.: (1) Rockledge, 14 May 1897, J. Crawford s.n. (DUKE). Citrus Co.: (2) Crystal River, 24 Jun 1941, LE Arnold
and E. West s.n. (FLAS); Orlando, S Riefler Nursery, 28 Apr 1996, S. Riefler s.n. (UWO). Columbia Co.: (1) Lake City, 01 May 1893,
Layne s.n. (FLAS). Escambia Co.: (2) 5 mi W of Century, 08 Aug 1954, ES. Ford 42576 (FLAS); 3.5 mi W of Barnett Crossroads
and 1 mi W of Big Escambia Creek, 08 Jul 1981, G. Wilhelm 9212 (USF). Gadsden Co.: (2) Aspalaga, Apr no year, 300c (GH, US).
Highlands Co.: (1) N of Lake Placid near jct. of US Rte. 27 and Josephine Creek, 29 Sep 1979, B. Judd, D.B. Ward and WS Judd
2504 (UWO). Jackson Co.: (2) 1.6 mi N of Marianna, 28 May 1973, D. Boufford and H.E. Ahles 77051 (LSU); 1.5 mi N of Marianna,
27 May 1973, D. Boufford and H.E. Ahles 9511 (A). Leon Co.: (1) Tallahasse, Apr 1843, M. Rugel s.n. (NY). Liberty Co.: (2) By FL
Rte. 12, jct. with FL Rte. 171, N of Bristol, 22 Mar 1982, R.K. Godfrey 79444 (FSU); 11 Apr 1957, LJ. Uttal 5382 (LYN). Marion Co.:
(1) 3 mi S of Ocala, 11 Aug 1939, WA. Murrill s.n. (FLAS). Okaloosa Co.: (1) E of Rock Creek near Hurricane lake, 16 Jun 1980,
G. Wilhelm 7534 (USF). Suwanee Co.: (1) Hildreth, 21 Aug 1940, WA. Murrill s.n. ee Volusia Co.: (1) no locality, 01 Aug 1900,
A.H. Curtiss 6703 (NCU). GEORGIA. Barrow Co.: (1) 1.2 mi W of Windsor, 25 Jun 1941, WH. Duncan 3564a (GA). Berrien Co.: 3 :
Nashville, between Marion St and GA 125, 19 Apr 1981, G.F. Joye 423 (LSU). ds e (2) Rte. 56 S of McBean, 13 Aug 19
J.B. Phipps 6710a (UWO); N of intersection with 80, 09 Apr 1991, J.B. Phipps 6496 (UWO). Candler Co.: (2) 0.1 mi W of ee
Creek and 0.1 mi N of road that crosses creek, 22 Apr 1975, D.J. Drapalik and WW. Walker 69 (NCU); 11 mi WNW of Statesboro
and NW of Upper Lotts Creek Church, 11 Apr 1977, D.J.Drapalik and C. Thomas 39 (NCU). Charlton Co.: (1) near Folkston, 23
Mar 1940, J.H. Miller 20 (GA). Clarke Co.: (1) Along Oconee River, SE edge of university campus, 16 Apr 1955, M. Duncan s.n
(GA). Liberty Co.: (1) 11 mi E of Taylor Creek, 27 Apr 1940, WH. Duncan 2236 (GA).Rabun Co.: (2) Canyon at Tallulah Falls, 16
Aug 1893, J.K. Small s.n. (GH); Tallulah River Creek, Sep 1999, R. Lance s.n. (UWO). Richmond Co.: (10) near Augusta, 23 Apr
1900, C.S. Sargent s.n. (DOV); Augusta, 11 Apr 1916, WW. Ashe s.n. (NCU); Augusta, 27 Apr 1902, A. Cuthbert s.n. (FLAS); US 1,
10 mi E of Briar Creek, 16 May 1958, J.A. Duke 665 (NCU); US 1, E of Bath Rd, 14 Aug 1958, J.A. Duke 2051 (NCU); 2.9 mi E of jct.
US 1 and 88, 19 Jul 1958, J.A. Duke 1636 (NCU); Augusta, no date, S.T. Olney and J. Metcalf 221 (GH); Augusta, no date, S.T. Olney
and J. Metcalf 220 (NCSC); Augusta sandhills, 27 Apr 1902, A. Cuthbert s.n. (FLAS); Augusta, 9 Aug 1902, A. Cuthbert s.n. (FLAS).
Screven Co.: (2) 3.5 mi N jct. GA 17 on 301, 11 Jun 1961, H.E. Ahles 54343 (NCU); 17.6 km NE of Newington, 18 Apr 1992, J.B.
Phipps 6561 (UWO). Taylor Co.: (1) 3 mi E of Rupert, 12 mi S of Butler, 02 May 1942, WH. Duncan 5101 (GA). Talbot Co.: (2) 2
km ENE of Geneva, 09 Apr 1991, J.B. Phipps 6587 (UWO); 2 km ENE of Geneva, 13 Aug 1993, J.B. Phipps 6729 (UWO). Thomas
Co.: (1) Along roadside US 19 2 mi N of Thomasville, 26 Apr 1952, S.C. Hood 451 1 (FLAS). Turner Co.: (2) Along US 75 4.5 mi NE
of Ashburn, 03 Jun 1964, W.R. Faircloth 1054 (GA, NCU). Walton Co.: (1) S side of Ga 138, 35 mi W of Athens, 22 Apr 1981, G.F
Joye 545 (LSU). Washington Co.: (1) no locality, 30 Apr 1942, W.H. Duncan 4937 (GA). MISSISSIPPI. Forrest Co.: (3) E of Pep's
=
1160 J lof the Botanical R h Institute of Texas 2(2)
Point, 14 Sep 1969, K.E. Rogers 1404-A (GH, UWO); Just E of Pep's Point, 14 Sep 1969, K.E. Rogers 1404-B (NCC). Jones Co.: (1)
5.5 mi SW of Ellisville, 12 Jun 1960, J.G. Teer 190 (IBE). NORTH CAROLINA. Cumberland Co.: (2) 2.3 mi SW of Hope Mills, 11
Oct 1957, H.E Ahles 36604 (NCU); On Rte. 87 near jct. with NC 210, 07 Aug 1948, ET. Browne Jr. 294 (KY). Durham Co.: (1) Duke
Forest, 08 Apr 1951, WT. Batson 751 (NCU). Fayetteville Co.: (2) no locality, 21 Apr 1909, WW. Eggleston 4929 (CM); no locality,
26 Aug 1908, WW. o 4013 IMSS). Franklin Co.: (1) near r Rte. 56 about 5 5 miW of Castalia, 26 in 1963, RJ. Wilbur 6793
DUKE). Harnett Co.: (3 SW of St prings, 11 Jul 1968, S.W. Leonard, A Burnh 1745 (LAF, NCC, USCH).
Hoke Co.: (1) 0.4 mi SW e Ashley Heights, 11 May 1957, M. Murana E a 24974 (NCU). Lenoir Co.: (1) Few mi S of La
Grange, 16 Apr 1991, J.B. Phipps 6510 (UWO). McDowell Co.: 5 mi WSW of Dysortville, 1.6 mi ENE of US 221,01 Sep 1956,
H.E. Ahles and C.R. Bell 17794 (NCC). Moore Co.: (3) 4 mi SW of Pinebluff 01 n 1927, WE. Manning and K.M. Weigand 1344 (GH);
no locality, 20 Jul 1930, 11941 (DUKE); Wildlife Club, SR 1004 at Drowning Creek, 06 May 1976, T.P. Huntley and J.F. Matthews
CU). Richmond Co.: (1) 4 mi E of Hamlet, 1 Jul no year, KM. Wiegand and W.E. Manning 1345 (G). Rowan Co.: (1) Vicinity
of Faith Post Office, 14 Aug 1981, JK. Small and AA Heller s.n. (MSC). Rutherford Co.: (1) Midway between Rutherford and
Chimney Rock, 08 Aug 1932, W.C. Coker s.n. (NCC). Scotland Co.: (2) 7 mi S of Laurinburg on No 501, 05 Mar 1937, M.L. Loker
2 (NCUJ; Juniper Creek 1.3 mi NE of Silver Hill, 08 May 1957, H.E. Ahles 24779-A (NCU). SOUTH CAROLINA. Anderson Co.: (3)
no locality, 19 Apr 1912, J. Davis 7757 i des yard, 19 Apr 1912, J. Davis 7701 (GH, US). Berkeley Co.: (1) 1.8 mi E of jet. SC
Rte. 45 and Rte. 6 on SC Rte. 6, 07 Apr 1 J.G. Haesloop and H.E. Ahles 22403 (NCU). Chesterfield Co.: (1) Little Black Creek
near SC 265, 05 Jun 1956, A.E. Radford (NCU). Es ol Co.: (3) Co. Rte. 360 E of Co. Rte. 1040, 2 mi W of downtown
Hartsville, 13 May 1988, C. Horn and D. Rayner 2610 (USCH); N of Lauther's Lake, 03 Aug 1941, BE. Smith 952 (USCH); Hartsville
03 Aug 1940, B.E. Smith 952 (NCC). Kershaw Co.: (2) near US 601, 1.5 mi N of Camden, 04 Jun 1957, A.E. Radford 23733 (FSU,
NCU). Lancaster Co.: (2) near Kershaw, 26 Jul 1906, H.W, House 2605 (MO); 0.4 mi NNE of Little Lynches Creek on US 601 and
1 mi E on dirt road, 06 Jun 1957, J.G. Haesloop and H.E. Ahles 27425 (NCC). Lexington Co.: (1) 04 mi W of S. Lake Drive, 21 Sep
1987, S.R. Hill 18843 (USCH). Oconee Co.: (1 n no locality, 18 Oct e WW. Ashe sn. (NCU). Orangeburg Co.: (1) 4.7 mi WNW
jct. US 301-601 and SC 4 on SC 4, W of Orangeburg, 18 May 1957, J.G. Haesloop and H.E. Ahles 25272 (NCU). Richland Co.: (1)
Headwaters of Cobb's Pond within St. Lo Range Training Area "i 01 Jun 1992, a inne and J.G. Nelson 681 (USCH).
=
Y
—
Crataegus pexa Beadle
GEORGIA. Bryan Co.: (1) 1.9 mi NE of Pembroke, 30 Aug 1961, J.A. Boole Jr. 1215 (NCU). Richmond Co.: (1) Augusta, 25 Apr
1900, A. Cuthbert 11 (DOV). Tatnall Co.: (1) 3.7 mi SW of Glennville, 24 Aug 1965, M.G. Padgett 9 (GA). NORTH CAROLINA.
Craven Co.: (1) New Bern, 20 Apr 1919, T.G. Harbison s.n. (A). Cumberland Co.: (2) 1.4 mi SW of Fayetteville, 27 Jun 1957, J.
Haesloop and H.E. Ahles 29718 (NCU); 2 mi E of Stedman, Sep 1931, H.R. Totten s.n. (NCU). Harnett Co.: (7) Pineview, 3 Jun 1957,
H. Laing 1542 (NCU); 0.1 mi W of Johnsonville, 8 May 1957, H. Laing 1211 (2*NCU); Spout Springs, 18 Apr 1929, B.W. Wells s.n.
(NCSC); 0.1 mi S of Coates, 15 May 1957, H. Laing 1315 (NCU); 1.4 mi W of Angier, 24 Apr 1957, H. Laing 1035 (NCU); 18 mi S of
Sanford, 27 Jun 1958, J.A. Duke 1 145 (NCU). Jackson Co.: (1) Cherokee, 20 August 1891, H.C. Beardslee and C.A. Kofoid s.n. (GH).
Johnson Co.: (2) 2 mi S of Smithfield, 3 May 1940, Radford and Stewart s.n. (GA); 2 mi SE of Four Oaks, 29 Jul 1970, R.M. Downs
and E. Upchurch 11424 (NCSC). Lenoir Co.: (1) 1.5 mi W of Jenny Lind, 11 Oct 1958, CJ. Burk s.n. (NCU). Montgomery Co.: (1)
NE corner of Co, 31 May 1953, Lionel Melvin s.n. (NCU). Moore Co.: (5) 5 mi E of Carthage, 12 May 1958, J.A. Duke 506 (NCU);
E of Harnett Co. line, 25 Apr 1948, RJ. Campana and R.K. Godfrey s.n. (NCSC); Southern Pines, 19 Jul 1895, J.W. Blankinship s.n.
(GH); Southern Pines, 19 Jul 1895, J.W. Blankinship s.n. (GH); 0.7 mi N junction US 1 and US 15, 19 Jul 1958, J.A. Duke 1367 (NCU).
New Hanover Co.: (2) Wilmington, 15 Apr 1911, C S. Williamson s.n. (CM); Wilmington, 1867, W.M. Canby s.n. (NY). Rowan Co.:
(1) 2 mi NE of Spencer, 26 May 1956, A.E. Radford and H.E. Ahles 12900 (NCU). Sampson Co.: (2) 1.2 mi N of Salemburg, 22 Apr
1957, RL. Wilbur 5221 (DUKE, FSU). Wayne Co.: (2) 5.6 mi NW of Seven Springs, 21 Jun 1957, A.E. Radford 25539 (NCU); 0.2 mi
N of Neuse Road on N.C. Route 111, 22 Sep 1950, WB. Fox with S.O. Boyce and R.K. Godfrey 4312 (NCSC). Wilson Co.: (1) 1 mi
NW of Sims, 28 Jul 1958, A.E. PIG 37992 (NCU). SOUTH CAROLINA. Kershaw Co.: (1) 4 mi W of Bethune, 7 Sep 1939, R.K.
Godfrey 8021 (MO). Spart g Co.: (1)near Campobello, 20 Jun 1942, E.M. Walker 3489 (NCU).
Crataegus quaesita. Beadle
i ill Aar 1940, WA. Murrill s.n. (A); Gainesville, o WA. Murrill s.n. (A); Gainesville,
19 Jul 1979, K.D. Perkins à BJ Juda 657 us Clay Co.: (11) Magnolia Springs, 9 Apr 1920, T.G. Harbison 5667 (A); Magnolia
RAE 9 Apr 1920, T.G. Hart 5679 (A, NCUJ; a SpUnds, 2 1920, a G Harbison 5670A ); Magnolia Springs, 9 Apr
76 (A
20, T.G. Harbison 5 T ); Magnolia Springs, 9 Apr 1920, T.G lia Springs, 7 Apr 1920, 7.G. Harbison
ps (A); Magnolia Springs, 9 Apr 1920, T.G. Harbison um ); Magnolia Springs, 21 Mar 1981, G. Martin s.n. (MO); Hibernia,
Mar 1869, WM. Canby s.n. (GH, NY). Columbia Co.: (1) Lake City, 1 Apr 1920, T.G. Harbison 5661 (A). Duval Co.: (6 racial
25 Mar 1901, A.H. Curtiss s.n. (GH); Jacksonville, Mar no year, A.H. Curtiss s.n. Dub ad in fruit, no date, A.H. Curtiss s.
(NCU); Jacksonville, 1 Aug 1894, 2d d GA) Jagconvile 7 Apr 1885, CE. Faxon s.n. (NY); Jacksonville, 8 Apr 1885, CE
Faxon 1858 (A). Gadsden Co.: (1) B Mt. Pleasant, 14 Mar 1937, apio Party (FLAS). pn
Co.: (1) 4 mi W of Mayo, 6 Sep o > Murrill & W.B. Tisdale s.n. (FLAS). Lake Co.: (1) Ocklawanna R. and Umatilla, 13
1940, Watson & W.A. Murrill s.n. (FLAS). Leon Co.: (3) W of Tallahassee, 17 Mar 1982, R.K. Godiey à im Hail 79429 (FLAS); a
lahasee, 9 Mar 1914, C.S. Sargent s.n. (A); 5 mi W of Tallahasee, 11 Apr 1931, E J.Palmer 38517 (A). Liberty Co.: (2) Bristol, 6 Sep
1940, WA. Murrill & V/B. Tisdale s.n. (FLAS); Torreya State Park, 19 Jul 1938, WP Porter s.n. (FLAS). Madison Co.: (1) Eilaville, 21
=
DL: nm ky, R ee Efrat H [| 1 1161
TT
Aug 1940, WA. Murrill & W.B. Tisdale s.n. (FLAS). Marion Co.: (6) Ocala, 14 Apr 1940, WA. Murrill 8 Watson s.n. (FLAS); Ellavale, 21
Aug 1940, WA. Murrill & W.B. Tisdale s.n. (FLAS); near Ocala, 11 Aug 1939, WA. Murrill s.n. (FLAS); Rainbow Springs, 11 Aug 1940,
WA. Murrill s.n. (FLAS); 9 mi W of Weirsdale, 22 Mar 1970, L. Baltzell 1750 (FLAS); Ocala, 8 Mar 1883, J.D. Smith s.n. (GH). Suwan-
nee Co.: (3) 1 mi E of Ellaville, 31 Mar 1957, R. Kral 4363 (NCSC); 1 mi E of State Park, 31 Mar 1982, H.B. & D.S. Correll 53520 (NY);
Hildreth, 11 Jul 1940, W.A. Murrill s.n. (FLAS). GEORGIA. Bulloch Co.: (2) 4 mi SW of Statesboro, 27 Mar 1962, G.P DeWolf 1722
(GA, NCU). Floyd Co.: (1) Rome, 25 Apr 1901, Biltmore Herb. 4194 (A). Gwinnett Co.: (1) Anniston, 6 Apr 1900, CSS. Sargent s.n.
(A). Lowndes Co.: (1) no location, 27 Mar 1980, K.D. Perkins & BJ. Judd 126 (FLAS). Randolph Co.: (1) 3 mi NW of Cuthbert, 12
Apr 1913, T.G. Harbison 1364 (A). Tattnall Co.: (1) 1.9 mi NW of Reidsville, 11Jun 1961, H.E. Ahles 54272 (NCU). Taylor Co.: (1) 1
mi W of Howard, 22 Mar 1938, J.H. Pyron & R. McVaugh 2287 (GA). Washington Co.: (2) E of Cowpen Creek, 12 Oct 1941, WH.
Duncan 4331 (GA, NCSC). Webster Co.: (2) no location, 8 Oct 1939, WH. Duncan s.n. (GA, NCU). Wheeler Co.: (1) along Little
Ocmulgee River, 12 dl 1942, WH. Duncan 4715 (GA)
Crataegus sp. aff. re
FLORIDA. Duval e P Jacksonville, Mar, A.H. Curtiss 811 (GA, 2*GH, MO, 2*NY).
Crataegus vicana Beadle
ALABAMA. Barbour Co.: (1) Eufaula NWR, 6 Apr 1993, D.Drennan 80 (UWO). FLORIDA. Alachua Co.: (4) Gainesville, 28 Jun
1940, WA. Murrill s.n. (FLAS); near Micanopy, 6 Apr 1931, EJ. Palmer 38412 (A); o 7 Apr 1931, EJ. Palmer 38424 (A) Rte.
24, SW of Kanapaha and 4 km SW of SR 127, 12 Mar 1993, J.B. Phipps 6667 (UWO). Baker Co.: (3) near Glen Saint Mary, 1 Apr
1924, H.H. Hume s.n. (A); Lake City, 19 Mar 1892, PH. Rolfe s.n. (FLAS, ie he Co: (1 ee e Springs, ae 1940, WA.
Murrill s.n. (FLAS). Gadsden Co.: (1) Chattahoochee, 9 Mar 1897, coll (MO). Highlands Hammock
State Park, 5 Jul 1949, R. Garrett s.n. (FLAS). Lafayette Co.: (1 ) Old Town, 31 Mar 1919, T.G. usn 5627 (NCU). Leon Co.: (5)
4 mi W of Tallahassee, 7 Apr 1958, R.K. Godfrey 56422 (AUA, DUKE, FSU, LAF, USF). Levy Co.: (1) near Chiefland, 7 Apr 1931, EJ.
os (A). sen th (5) Bristol, Mar 1890, WM. Canby s.n. (DOV); Tallahassee, 4 Apr 1972, P Elliot 2 (FSU); Tallahassee, 6
, Tallahassee, 2 Apr 1920, T.G. Harbison 5647 (A); 15 mi W of Tallahassee, 2 Apr 1959, R.K.Godfrey
58290 E Madison Co.: e >) Ellaville 21 Aug 1939, WA. Murrill s.n. (FLAS); no locality, 31 Mar 1914, T.G. Harbison 1434 (A).
Marion Co.: (3) near Ocala, 6 Apr 1929, EJ. Palmer 35185 (A); 4 mi W of Ocala, 20 Mar 1958, G.R. Cooley & RL Eaton 5756 (FSU,
USF). Suwannee Co.: (2) 1 mi E of State Park, 31 Mar 1982, D.S. & H.B. Correll 53520 (USF); 5 mi S of Live Oak, 21 Auf 1939, WA.
Murrill & W.3. Tisdale s.n. (FLAS). GEORGIA. Berrien Co.: (1) 4.5 mi NE of Five Mile Creek, NE of Weber, 23 Jun 1966, JR. Bozeman
e ien Bibb Co.: (1) 7 mi Se of Macon, 12 Apr 1940, W. Duncan 1906 (GA). Bryan Co.: (1) E of Canoochee River, 14 Jul
043 (NCU). Decatur Co.: (3) 0.75 mi W of pisc near Seminole Co. line, 31 Mar 1982, R.K. Godfrey 79488
Ee is N of Mt. Pleasant Community, 31 Mar 1982, R.K. Godfrey 79496 (FSU); Rte. 97 and railroad, 1 mi S of Faceville, 28 May
1983, R.K. Godfrey 79488 (FSU). Floyd Co.: (1) Rome, 5 May 1899, dh Canby 28 (DOV). Lowndes Co.: (2) E of I-75 on N side of
GA 375, 27 Mar 1980, W.S. Judd & K.D. Perkins 2596 (LAF); 5.3 mi S of Valdosta, 21 May 1970, C. Scott 13 (MO
=
ACKNOWLEDGEMENTS
Susan Laurie-Bourque is thanked once more for her fine line illustrations, as is The National Sciences and
Engineering Research Council of Canada for continuing support to the first author. The authors are also
grateful to the curators of the following 32 herbaria for their patience during the long loan necessary to evalu-
ate very large number of specimens for this complex study: A, ALU, AUA, CM, DHL, DOV, DUKE, FLAS,
FSU, GA, GH, IBE, KY, LAF, LYN, MO, MSC, NATC, NCC, NCSC, NCU, NLU, NO, NY, TENN, UNA, US,
USCH, USF, VDB, WILLI, WVA. Significant cooperation and help in the field was supplied at various times
by Robert O’Kennon, Botanical Research Institute of Texas, Fort Worth, Texas and Ron Lance of Chimney
Rock Park, North Carolina. Antony Littlewood, Dept of Classical Studies, University of Western Ontario,
is thanked for checking the Latin descriptions. Various student volunteers assisted in the databasing of
specimens which provided the data for mapping.
REFERENCES
Arron, W. 1789. Crataegus flava. In: Hortus Kewensis 2:19.
BeaoLe, C.D. 1902. New species of thorns from the southeastern United States, Il. Biltmore Bot. Stud. 1:62. 1902.
BeapLe, C.D. 1903. Crataegus. In: J.K. Small, Flora of the southeastern United States, ed. 2. Author, New York.
GEoRG, J.G. 1775. Bemurkungen einer Reise im Russischen Reich im Jahre 1772. iii. Nachtrage: 273.
MicHAUx, A. 1803. Flora Boreali-Americana. 1:288.
MotNcH, C. 1785. Verzeichniss ausländischer Bäume und Stauden: 31.
MunniL, W.A. 1942. New Florida hawthorns. Castanea 7:19-30.
1162 J lof the B ical R h Institute of Texas 2(2)
Persoon, C.H. 1806. Crataegus michauxii. In: Synopsis Plantarum 2:38. Treutel and Wuerte, Paris and Cottam,
Tubingen.
Phipps, J.B. 1988a. Re-asessment of Crataegus flava Aiton and its nomenclatural implications for the Crataegus
serial name Flavae (Loud.) Rehd. and its sectional equivalent. Taxon 37:108-113.
Pus, J.B. 1988b. Crataegus (Maloideae, Rosaceae) of the southeastern United States, |. Introduction and series
Aestivales. J. Arnold Arbor. 69:401-431.
Phipps, J.B. AND K.A. Dvorsky. 2006. Crataegus series Parvifoliae and its putative hybrids in the southeastern United
States. Sida. 22:423-445.
Phipps, J.B., RJ. O'Kennon, AND K.A. Dvorsky. 2006. Review of Crataegus series Pulcherrimae (Rosaceae). Sida
22:973-1007.
Phipps, J.B., R. Lance, AND K.A. Dvorsky. 2006. Crataegus series Bracteatae and Triflorae (Rosaceae). Sida 22:1009-
1025.
Phipps, J.B., R.J. O'KENNON, AND R. LANCE. 2003. Hawthorns and medlars. Timber Press, Portland.
Raproro, A.E., H.E. AHLES, AND C. Rrrcuie BeLL. Manual of the flora of the Carolinas. University of North Carolina Press,
Chapel Hill.
SARGENT, C.S. 1890. Silva of North America. IV:113, t. 189.
SARGENT, C.S. 1902. Silva of North America. XIII, suppl.:155, t. 693.
TALENT, N. AND T.A. Dickinson. 2005. Polyploidy in Crataegus and Mespilus (R Maloideae) luti infer-
ences from flow cytometry of nuclear DNA amounts. Canad. J Bot. 83:1268- 1304.
TipEsTROM, |. 1933. Crataegus. In: J.K Small, Manual of the southeastern flora. Univ. North Carolina Press, Chapel
Hill.
WUNDERLIN, R.P. AND B. Hansen. 2003. Guide to the vascular plants of Florida, 2" ed. University Press of Florida,
Gainesville.
INDEX OF NAMES
d i ti | di ion in italics; a?" indicates synonymy is uncertain;
Accepted taxa
numbers represent ordinal sequence of species in text or page tb
Crataegus
?adunca Beadle, 10
adusta Beadle, 5 floridana Sargent, 15 munda Beadle, 4
alabamensis Sende 10 frugalis Beadle, 5 panda Beadle, 16
?amica Beadle, fruticosa Sargent, 1104 pearsonii Ashe, 1149
anisophylla e 7 furtiva Beadle, 18 pexa Beadle, 13
arguta Beadle, 9 garrettii Murrill, 1108 pulla Beadle, 15b
attrita Beadle, 15a ?geniculata Ashe, 4 quaesita Beadle, 6
?audens Beadle, 9 glandulosa Michx., 1102 ?rava Beadle, 9
cirrata Beadle, 2 i Moench, 1103 ravenelii Sargent, 9
clara Beadle, 9 i Georgi, 1103 recurva Beadle, 5
colonica Beadle, 12 illudens Beadle, 16 resima Beadle, 6
compitalis Beadle, 9 ?incana Beadle, 5 rimosa Beadle, 9
condigna Beadle, 9 ?inopina Beadle, 5a Robustae, subser. nov., 1124
?constans Beadle, 11 ?inops Beadle, me senta Beadle, 21
crocea Beadle, 5 insidiosa Beadle, sodalis Beadle, 11
cuthbertii Ashe, 1149 integra (Nash) M 11 Tenues, subser. nov., 1108
curva Beadle, 5 invicta Beadle, 8 teres Beadle, 14
?dapsilis Beadle, 10 lacrimata Small, 3 ?versuta Beadle, 7
dispar Beadle, 22 lanata Beadle, 17 viaria Beadle, 6
dolosa Beadle, 11 lancei J.B. Phipps, sp. nov., 20 vicana Beadle, 5a
egens Beadle, 2 lassa Beadle, 16 villaris Beadle, 5
exilis Beadle, 23 ?laxa Beadle, 6 ?yadkinensis Ashe, 20
flava Aiton, 1102 lepida Beadle, 1
flava var. integra Nash, 11 meridiana Beadle, 19
florens Beadle, 14 michauxii Persoon, 1102
DICERANDRA MODESTA (LAMIACEAE): RAISE IN RANK FOR A
DISJUNCT PERENNIAL IN A NEW COASTAL CLADE IN FLORIDA
Robin B. Huck
Florida eis of Natural History
sity of Florida
TEM ae 32611, USA.
mints@bellsouth.net
ABSTRACT
T 1 f. 1 1 J A 1 Ty 1 J D 1 1 RE, 1 h; iT
r I It
p to be a disj ligned with the Di ira species of the Átlantic Coastal Ridge of eastern Florida not with Dicerandra frutescens
1 ia] Te [; x 1 H Ts f£ * 11 : " x » 1h z 1
of the Lake g Florida p Į £j
distinguish Di 1 lesta f tl bers in the new coasta l clade.
Key Wonps: Dicerandra modesta; Lake Wales Ridge; Atlantic Coastal Ridge; southeastern U.S. endemics; pollination
RESUMEN
e y t
1 las especies de Dicerandra de la cadena tera del te de Florida y no con
F
1 A do]
t p F P 7 E
Dicerandra frutescens de la cadena de Lake Wales de la y interi lde Florida. Las dif i ] la, la orna-
C Ll TA PPM 1 ] +
1 Ps ey] 1
New molecular phylogenetic evidence (Oliveira et al. 2007) necessitates a review of the status of Dicerandra
frutescens Shinners subsp. modesta Huck, and, when supported by distinctive morphological and pollina-
tion characters, requires a raise in rank to species so that this new taxon can be represented correctly in the
upcoming volume of the Lamiaceae for the Flora of North America Project. This designation is in keeping
with the concept of species within the group.
pend modesta (Huck) Huck, comb. et stat. nov. Di f si modesta Huck, Novon
2001. Tre: U.S.A. FLoriDA. Polk Co: Black Road, E of Dundee, sandy nape 30 m, 9 Sep 1999, RB. Huck 5555 (HOLOTYPE:
e. ISOTYPES: MO!, USF!).
An endemic genus of the southeastern United States characterized by spurred anthers, Dicerandra has 4
annual species in D. subgenus Dicerandra and, now, 6 perennial species in D. subgenus Kralia, the latter
all rare and found on relict, island-like sandy ridges in Florida (Huck 1987, 1992, 2001, 2007; Huck &
Chambers 1997; Huck et al. 1989). With a pale yellow to pinkish-white corolla, a geniculate corolla tube
and smooth spurred anthers typical of adaptations to bee fly pollination, Dicerandra modesta was originally
recognized as a subspecies of D. frutescens, a taxon with a similar morphology and adaptation (Huck 1987,
2001; see Deyrup & Menges 1997). Further, this previous subspecies was thought to be yet another of the
many endemics of the Lake Wales Ridge (Huck et al. 1989; Christman & Judd 1990), the interior ridge of
central Florida. However, in the molecular phylogenetic study of Oliveira et al. (2007), both an analysis of
ITS data and a combined analysis of ITS and chloroplast data sets showed Dicerandra modesta was surpris-
ingly not allied with D. frutescens, but instead, was included in a well supported coastal clade with coastal
species D. thinicola H.A. Mill. and D. immaculata Lakela. Another tree based on chloroplast data supported
the distinctness of these perennial species (Oliveira et al. 2007). In contrast to Dicerandra modesta, these two
coastal taxa have pale purple to purplish-red to pink corollas, smoothly bent corolla tubes and ornamented
“brush” anthers spurs typical of bee pollinator adaptation.
Dicerandra thinicola and D. immaculata (and its variety D. immaculata var. savannarum Huck) grow in
isolated sand pine [Pinus clausa (Chapm. ex Engelm.) Vasey ex Sarg.] - oak scrub (Quercus geminata Small;
J. Bot. Res. Inst. Texas 2(21 1163 — 1164, 2008
" ; lef the Bataniral D h Institute of Texas 2(2)
Q. myrtifolia Willd.) plant communities near the ocean 60 miles from one another along the 300-mile-long
Atlantic Coastal Ridge of Florida, some 60 to 80 miles to the east of the type locality of D. modesta found in
a similar habitat on the parallel central ridge. Both Dicerandra modesta and D. thinicola are tetraploids and,
at least based on ITS data, are more closely related to each other than to hexaploid D. immaculata (Huck &
Chambers 1997; Oliveira et al. 2007).
Dicerandra modesta is a chamaephytic perennial with floral shoots arising from a branched woody base
and the inflorescence is a verticillaster with 2-3 flowers to a cyme, blooming successively in an exuberant
burst from late August through September. The specific name modesta meaning "shy" is derived from the
tendency of the pale yellow-colored corollas to blush to pink when the flowers are in full anthesis. The very
slender, geniculate tube of the corolla flares to a trumpet-shape, with the upper lobe an ovate-shaped stan-
dard, marked with trellised red-purplish markings and the lower tripartite lobe, separated from the upper
lobe by a deep sinus, is marked with spots of the same color. Four stamens are declinate along the lower
lobe of the corolla and the anthers have large pollen sacs divided by a glandular, widened connective. Each
pollen sac is subtended by a linear, glabrous or a few-haired spur terminating in a forked tip of 3—5 cells.
Dicerandra modesta is apparently reproductively isolated from the two coastal species by geography, pollina-
tor preference and, in the case of D. thinicola (which blooms in October and November), a displacement of
blooming time. A key to this small genus will be included in the treatment in the Flora of North America
Project.
ACKNOWLEDGMENTS
I thank D.L. Black for observations on the pollination of Dicerandra modesta. 1 thank H.L. Chambers and
an anonymous reviewer for their insightful comments and T. Lindler, and especially K. Perkins and N.H.
Williams (FLAS) for their continuing generosity and support.
REFERENCES
CHRISTMAN, S.P. AND W.S. Jupp. 1990. Notes on plants endemic to Florida scrub. Florida Sci. 53:52—73.
Devrup, M. AND E.S. Mences. 1997. Pollination ecology of the rare scrub mint Dicerandra frutescens (Lamiaceae).
Florida Sci. 60:143-157.
Huck, R.B. 1987. Systematics and Evolution of Dicerandra (Labiatae). Phanerog. Monogr. 19:1—343. J. Cramer,
Gebruder Borntraeger, Berlin, Germany.
Huck, R.B. 1992. Overview of pollination biology in the Lamiaceae. In: R.M. Harley & T. Reynolds, eds. Advances
in Labiate Science. Royal Botanic Gardens, Kew, UK. Pp. 167-181.
Huck, R.B. 2001. Two new infraspecific taxa in Florida Dicerandra (Labiatae). Novon 11:417—420.
Huck, R.B. 2007. Clarification of the type locality of Dicerandra linearifolia (Labiatae). Rhodora 109: 387—394.
Huck, R.B. AND H.L. CHamseRs. 1997. Polyploidy: a factor in the evolution of Dicerandra Benth. (Labiatae). Edinburgh
J. Bot. 54:217-229,
Huck, R.B., W.S. Jupp, W.M. WHrrEN, J.D. SKEAN, Jr, R.P. WUNDERLIN, AND K.R. DELANEY. 1989. A new Dicerandra (Labiatae) from
the Lake Wales Ridge of Florida with a cladistic analysis and discussion of endemism. Syst. Bot. 14:197—213.
Ouiveira, L.O., R.B. Huck, M.A. GrrzeNpANNER, W.S. Jupp, D.E. Sorts, AND PS. Souris. 2007. Molecular phylogeny, bioge-
ography, and systematics of Dicerandra (Lamiaceae), a genus endemic to the southeastern United States.
Amer. J. Bot. 94:1017—1027.
TWO NEW NAMES IN THE CARIBBEAN FLORA
Alain H. Liogier
Botanical Research Institute of Texas
500 E 4" Street
Fort Worth, Texas 76102-4025, U.S.A.
The Flora of Hispaniola has been published from 1990 to 2000, describing the plants known at that time
from the Island of Hispaniola. As happens for every Flora, many additions have been made to the existing
knowledge of the plants in the Island; furthermore, quite a number of taxonomical changes have been made
and need to be made known to the scientific community. I have been working on the project of revising the
published volumes; these volumes include the Dicotyledons, less the Cactaceae, that have been left for the
specialists in that family. In this process, I have found two illegitimate names, published by mistake, and
these names have to be re-named.
Salvia toaensis Alain, nom. nov. Salvia nigrescens Alain, Contr. Ocas. Mus. De La Salle 15:12. 1956; non Salvia nigrescens Bull.,
Retail List 121: 170. 1876.
Hedyotis ekmanii Alain, nom. nov. Hedyotis nigrescens (Urb. & Ekm.) Alain, Phytologia 54:103. 1983; non Hedyotis nigrescens
Merr., J. Arnold Arbor. 23:193. 1942.
ACKNOWLEDGMENTS
I thank Scott Zona for his comments on the paper.
J. Bot. Res. Inst. Texas 2(2): 1165. 2008
1166 J | of the Botanical R h Institute of Texas 2(2)
BOOK REVIEW
James K. AGEE. 2007. Steward’s Fork: A Sustainable Future for the Klamath Mountains. (ISBN-10:
0520251253, ISBN-13: 978-0520251250, hbk.). University of California Press, 2120 Berkeley Way,
Berkeley, California 94704-1012, U.S.A. (Orders: www.ucpress.edu, orders@cpfsinc.com, 800-777-
4726). $42.00, 306 pp., black/white photographs, line illustrations, 7 maps, 6" x 9".
The Klamath Mountains, and particularly the Trinity and Marble Mountains, of California are the area that cid discusses with
much love and insight. But his insight f value to those living beyond these mountains as well. Th the use of the
a c the continued use and care of the land, and the principles of how to responsibly manage th ystems can be
hs th ld.
The p chapters in the text cover the geologic development of the area, the flora and fauna, and the native peoples. À story of
Agee's efforts to retrace and best Alice Eastwood's collecting survey along Canyon Creek in 1901 is engaging and enlightening, par-
e so when he es that even ow a Steet in phan uh was iunii to cues in count. de n is his attempt and
fth
t the
indigenous peoples and their oa on bn ed stem, IA livestock grazing, and water issues. Final ais cover m views on
f,
I pror
bl hi ies of ti d pl ke tl 1f both real and attainable. He writes with consid-
erable sil and is pe to use i memories as both dd and al Regarding choices | Is that hild riding inner tubes on
the Stu kh landa | dunked or flipped over. But he retrieved
his tube and tried again. He had limited knowledge of what d t but adjusted hi long the way. We now have the
same challenge in choosing a course for the Klamath M i d for t} f y
St T's Forki Hill lwit 11 imag 1 maps. Concludi he bool index, appendices, and kabl
Ld E NE bibliography i ged by char d cites books, periodicals and web sites. À ho fi he bibliography
1 1 l-; 1 1 1 : SRI 1 1 ` : e 1271
of ALLY LLC YY
for any library or for anyone pt in the area. It will appeal to general readers as well as professionals and specialists.— Gary L.
Jennings, Botanical Research Institute of Texas, 500 East 4th Street, Fort Worth, Texas 76102-4025, U.S.A.
J. Bot. Res. Inst. Texas 2(2): 1166. 2008
TROCHANTHERA LEPIDOTA GEN. AND SP. NOV., A FOSSIL ANGIOSPERM
INFLORESCENCE IN DOMINICAN AMBER
George O. Poinar, Jr. Kenton L. Chambers
Department of Zoology Department of Botany and Plant Pathology
Oregon State University Oregon State University
Corvallis, Oregon 97331, U.S.A. Corvallis, Oregon 9733 i U.S.A
Alex E. Brown
629 Euclid Avenue
Berkeley, California 94708, U.S.A.
ABSTRACT
T. 1 1 1 oy 11 A 1, tan | 1 | “1 1 f£. T : y PES
& sp. nov., req y pitate male intiorescence at anthesis, is described from Tertiary Dominican
amber. The gens is oe by small, staminate fl isi 1 i fl 1 bracts ona nd recep-
tacle. Each fl of 3-4 stamens, with distally connivent filaments ad bilocular a showing longitudinal extrorse
o Toe perianth i is ea bya Het, appressed, truncate MNA Meu the alme. dun ne feminis ded
had
m belo puber ulent. The fo y,
A. and Moraceae.
Key Wonps: amber, Bal 1 , Dominican Republic, fossil flower, Greater Antilles, Moraceae, Tertiary
E
RESUMEN
RUD pidon pen & sp. nov., ROREM D una uu masculina capitada en antesis, es descrita del ámbar
£ m pee , "E
1 t o Is
fil : 1; 1 hil T
plano en un me esférico Cada is se compone de 3-4 estambres,
con dehiscencia 1 una vaina corta, truncada y apresada alrededor de la columna
de mnes Es resto de nd de callo e de » cabezuela no tiene hojas y es ligeramente puberulento. El fósil, aunque de
j las familias Bal y Moraceae.
INTRODUCTION
Dominican amber is well known as a preservative agent of both animal and plant remains. Flowers may
retain their structural details so clearly that they often can be assigned to extant genera (Poinar 1992) and
can be used in paleontological reconstructions (Poinar & Poinar 1999).
During an investigation of floral remains in Dominican amber, we discovered a LA niu
male inflorescence of uncertain affinity. Its morphology is compared with the families B
Moraceae, with which it shares many essential characteristics. The fossil's exceptional Sue dde allows
microscopic study of the flowers, which are here illustrated. Due to its unique combination of features, the
plant is described below as a new genus and species.
MATERIALS AND METHODS
The fossil originated from mines in the northern mountain range (Cordillera Septentrional) of the Dominican
Republic, between the cities of Puerto Plata and Santiago. Dating of Dominican amber is controversial, with
the youngest proposed age of 20—15 mya based on foraminifera (Iturralde-Vincent & MacPhee 1996) and the
oldest as 45—30 mya based on coccoliths (Cépek in Schlee 1999). Most of the amber is secondarily deposited
in turbiditic sandstones of the Upper Eocene to Lower Miocene Mamey Group (Draper et al. 1994).
The amber piece containing the fossil is rectangular in outline, measuring 15 mm in length, 10 mm in
width and 8 mm in depth. The holotype is deposited in the Poinar amber collection maintained at Oregon
State University.
J. Bot. Res. Inst. Texas 2(2): 1167 — 1173. 2008
1168 J | of the Botanical R h Institute of Texas 2(2)
DESCRIPTION
Trochanthera G.O. Poinar, K.L. Chambers & A.E. Brown, gen. nov. Tre Spears: Trochanthera lepidota G.O. Poinar, K.L.
Chambers & A.E. Brown, sp. nov.
Diagnosis.—Stem leafless and minutely puberulent, total length unknown. Head depressed-globose, com-
pletely covered with persistent glabrous, flat-topped, closely connivent bracts in + a hexagonal pattern.
Staminate flowers numerous, emerging over the entire head from between the edges of the bracts, each
consisting of a column of distally connivent filaments and 3—4 anthers; anthers connivent, not fused, each
with 2 ovate, unilocular thecae dehiscing extrorsely by longitudinal slits. Perianth represented by a short,
truncate, erose sheath enclosing base of emergent filament column. Pistillate flowers unknown, absent from
the described head or hidden beneath the bracts.
Trochanthera lepidota G.O. Poinar, K.L. Chambers & A.E. Brown, sp. nov. (Figs. 1-6). Tye: HISPANIOLA.
Dominican REPUBLIC: amber mine in the dede mountain a oa een e boudin amber miner s.n.
Corvallis,
(HOLOTYPE: catalogue number Sd-9-148A
Oregon 97331, U.S.A.).
Description.—Staminate flower head: width 7.3 mm, length 5.3 mm; attached stem piece 2.6 mm long,
1.2 mm thick; peltate bracts of inflorescence 408(371-468) um in diameter, anthers of emergent stamens
280(226-323) um long, 113(96—145) um wide; length of exposed filaments 0-193 um; perianth a short (up
to 60 um) sheath enclosing base of filament column; pollen grains spherical, 5-12 um in diameter, out of
position for detailed view of exine morphology.
Etymology.—Genus name from Greek “trochos” ball, and “antheros” flowery. Species name from Greek
“lepidotos” scaly.
DISCUSSION
We consider the fossil's distincti j linfl , consisting of a small, glo! tacl pletely
covered with flat-topped (presumably peltate) bracts, Between which ien numerous comple NA
flowers, to have its closest analogs in two modern angiosperm groups—family Balanophoraceae and family
Moraceae, tribe Dorstenieae (formerly Brosimeae). Both these groups have modern representatives in the
Caribbean region. We nido not been able to identify a modern genus to which the fossil can confidently be
assigned, although it sl istics with both families. The following discussion, where Trochanthera
is compared with each of these putative relatives, calls attention to the striking parallelisms in inflorescence
morphology between the two taxa, which otherwise differ markedly in habit—small holoparasitic herbs
versus tall tropical forest trees.
Balanophoraceae are a morphologically distinctive group of holoparasitic angiosperms occurring in
tropical and subtropical regions of the Old and New Worlds (Hooker 1856; Engler 1889; Harms 1955,
Nevling 1960; Kuijt 1969; Hansen 1976, 1980; Nickrent 1997 onwards; Stevens 2001 onwards; Gonzáles
2004). They are obligate subterranean parasites, usually forming a tuberous attachment to host roots from
which naked or bracteate inflorescence-bearing stems arise. By upward growth of the stem, the flowering
portion is exposed at the soil surface or elevated on a fleshy, usually upright stalk, the plants at this stage
often being described as “mushroom-like.” Inflorescences are variously shaped, forming a loose or compact
(tightly branched) raceme in some genera and a flat-topped or club-shaped to spherical head in others (Kuijt
1969, p. 125). The flowers are unisexual, the plants being dioecious or monoecious with the two sexes, in
the latter case, either mixed or in separate parts of the inflorescence (or on separate heads from the same
tuber). Pistillate flowers, in particular, are often minute and ly numerous, while the usually synangial
anthers of staminate flowers take a great diversity of forms (Kuijt 1969, pp. 128, 130). A 3—6-lobed peri-
anth is most often present in staminate flowers and may act as a protective covering over the anthers as the
young inflorescence pushes through the soil. A different kind protection for such inflorescences, occurring,
for example, in the New World genera Helosis and Corynea, is a complete covering of peltate bracts, fitting
and giving the capitulum a somewhat pineapple-like appearance
tightly edge-to-edge in a g
i
Poinar et al., Trochanthera lepidota gen. and sp. nov. 1169
Fics. 1-5, Trochanthera lepidota. 1. Entire flowering head. Scale bar = 1.4 mm. 2. Group of st ing edg
Sca e ar= 150 um 2 $. kh £L J lj d i tal, I L I n + L H E $l L +L 1: L Pas | E
(arrow). Scale bar = 180 pm. 4. Portion of head showing peltate | ( ) with antl ging | heir margins. Scale bar = 360 pm.
[4 Dall P^ LH 4 ah c
la | ER
J JU um.
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tions of peltate bracts (B). Scale bar = 130 um.
(Hooker 1856, pls. XIII, XV; Hansen 1980, fig. 11). The development and function of boss bracts is well
described by Hansen (1978). In the two genera mentioned above, the peltate bracts are decid following
emergence of the capitulum, and the small staminate flowers then become evident as they emerge from a
dense, mat-like covering of clavate hairs (Hooker 1856, pl. XIII). The tiny pistillate flowers remain embed-
ded in these hairs with only the styles and stigmas exposed.
Trochanthera differs from the condition described for Helosis and Corynea, among others (see Hansen
1978, 1980), in that a covering of tight-fitting bracts persists at anthesis, and the staminate flowers force
their way out between the connivent edges of the bracts (Figs. 1—4, 6). In the process, some flowers elon-
gate farther than others, and occasional anthers are broken and left behind, as it were. It seems remarkable
that the delicate tubular perianth (Figs. 3, 6) is also able to emerge, at least in some flowers, in the process
of the staminal column forcing itself between the bracts. A E system for the emergence of staminate
Balanophora wrightii, illustrated by
flowers from between protective bracts occurs in the
Nickrent (1997 onwards), but in it the male flowers are fewer on he head, they possess a broadly 3-lobed
perianth, and the anthers form a synandrium with transverse dehiscence, as in some other Balanophora
taxa. In his discussion of inflorescence structure in the related Asian Balanophora species B. harlandii and B.
wilderi, Hansen (1978) refers to the clavate bracts covering the pistillate portion of the heads as “spadicles,
although this usage may be inappropriate (R.K. Eberwein pers. comm.). These intergrade with the broader
sub-peltate type in the staminate part of the heads. We cannot be sure to what extent Hansen's term spadicles
applies to Trochanthera, and we prefer the term peltate bracts even though their 3-dimensional form cannot
be determined from the fossil.
The stamens of Balanophoraceae are extremely diverse, especially in the construction of the anthers.
Poinar et al., Trochantl lepidota gen. and sp. nov. 1171
Anthers similar to those of Trochanthera are found in the South African genus Mystropetalon; that is, they
are of an unspecialized, bithecal type, dehiscent extrorsely through two longitudinal slits. Mystropetalon
happens also to be characterized by the putatively plesiomorphic trait of male and female flowers each with
vestigial organs of the opposite sex (Nickrent et al. 2005; Kuijt 1969, p. 127), whereas pistillodes are not
present in male flowers of Trochanthera. Compared with Helosis and Corynea, and with the Asian Balanophora
taxa mentioned above (Hansen 1972), Trochanthera's anthers are notably larger than in the 2 New World
genera (see illustrations in Hooker 1856; Hansen 1980) and are extrorse rather than introrse (Figs. 1-4, 6).
The anthers are not fused into a synandrium that releases pollen through horseshoe-shaped or transverse
slits, pores, etc., as is common in Balanophora (Hansen 1972). The filaments of the fossil may be proximally
fused, as in these other genera, but are only closely connivent distally (Figs. 2-4, 6).
The short, tubular, sheathing, unlobed perianth of Trochanthera most resembles that of Corynea crassa
Hook. f. var. crassa (Hooker 1856, pl. XIII; Hansen 1980, fig. 11). On the other hand, a prominently 3-lobed
type is characteristic of staminate flowers in C. crassa var. sprucei and also of Helosis, while in Mystropetalon
and Balanophora species the perianth is well developed and 3-6-lobed. An unlobed, sheathing perianth is
present in staminate flowers of the Asian Rhopalocnemis (Harms 1935, fig. 163), but its anthers form a single
multichambered synangium.
The stalk or stem of Trochanthera is leafless, at least distally (Fig. 1). This is comparable to various genera
of Balanophoraceae (Hooker 1856; Kuijt 1969; Hansen 1980; Nickrent 1997 onwards). More or less bracteate
stems are also common in the family.
A remarkably similar form of staminate inflorescence to the above occurs in the Brosimum alliance of
Moraceae, tribe Dorstenieae (Berg 1989; Rohwer 1993; Datwyler & Weiblen 2004; Zerega et al. 2005; for-
merly Brosimeae of Engler 1889; Hutchinson 1967; Berg 1972). Congested, often globose flowering heads
are frequent in Moraceae, as for example the members of tribes Castilleae (Olmedieae) and Dorstenieae
(Brosimeae) described and illustrated by Berg (1972). Heads may be bisexual (one or more pistillate flow-
ers mixed with the staminate ones) or unisexual (staminate or pistillate only, Berg 1989, fig. 11.3). These
are borne on short, axillary peduncles, one to several per node (Woodson 1960; Berg 1972) and are in the
same size range as Trochanthera. Because of the covering of peltate bracts in Trochanthera, between which the
staminate flowers emerge, the most promising comparisons are with similarly bracteate Brosimum species.
Other genera having small, globose heads of male flowers, such as Coussapoa (Woodson 1960, now in fam-
ily Cecropiaceae; Rohwer 1993; Datwyler & Weiblen 2004), Helicostylis of tribe Dorstenieae (Olmedieae of
Berg 1972), or Poulsenia (Woodson 1960, assignable to tribe Castilleae per Datwyler & Weiblen 2004) lack
peltate bracts and have flowers with a 3—4-parted perianth and stamens either free and more numerous or
1-2 and often connate.
The Trochanthera inflorescence appears completely staminate, whereas species of Brosimum often, but
not always, have bisexual heads with a single, deeply embedded pistillate flower accompanying numerous
staminate ones. However, the style of this female flower may wither early, and the head may then appear
staminate. Woodson (1960, p. 130) quotes Standley as to the difficulty of finding “with ordinary herbarium
specimens...even one pistillate flower.” We could not find an exposed style in the fossil.
Flowers with 2—4 stamens occur in Brosimum species of section Brosimopsis (genus Brosimopsis of
Hutchinson 1967, p. 172), but a well developed, 4-lobed, pubescent perianth is present, and stamens are
clearly separate with slender filaments (Berg 1972, p. 178). In section Piratinera (Berg 1972, p. 181) a perianth
is absent in the staminate flowers and may be replaced by an enclosing bract (e.g., B. gaudichaudii, p. 185).
However, there is only 1 stamen, which has a broad anther connective unlike Trochanthera. Other Brosimum
species with a reduced or absent perianth and only 1 stamen are found in section Brosimum (Berg 1972, p.
169); they differ from Trochanthera in having either a peltate anther (B. alicastrum) or a tiny one only 0.2
mm in length and breadth (B. acutifolium). The peltate bracts of these and other Brosimum taxa differ from
Trochanthera in being circular in outline, rather than closely connivent and more or less hexagonal (Woodson
1960, pp 131-133; Berg 1972, figs. 62, 66, 71, 73). In Brosimum subgenus Ferolia (Berg 1972, p. 188) the
I n€ ob D A] h l "T £T.
1172 J t Texas 2(2)
stamens, if more than 1, are divergent with slender filaments. The sheathing perianth, if present, is minutely
ciliate apically. The peltate bracts are usually puberulent. The stamen, if 1, usually has the anther bent at
90°, with a thickened connective (all data from Berg 1972). In this comparison, therefore, the fossil cannot
be accommodated in the genus Brosimum, and its description as a separate taxon is supported.
Recognition of Trochanthera as a distinct genus will assist in future comparison of this unique fossil
with extant members of Balanophoraceae and Moraceae, both of which are diverse in floral morphology and
inflorescence structure. In Balanophoraceae, we prefer not to assign the plant to a variable modern genus
like Balanophora (Hansen 1972), since taxonomic circumscriptions may change in the future, especially
when DNA methods are applied to the family. A principal authority on the family, Bertel Hansen, was admit-
tedly conservative in his taxonomic approach (Friis 2006), and he merged 9 later synonyms into the single
genus Balanophora. Additionally, other generic unions have recently been proposed (Eberwein & Weber
2004), which strengthen Old and New World taxonomic affinities in the family. Further, when comparing
Trochanthera with representatives of Moraceae, it is again unclear that a close relationship exists between it
and particular modern genera, including Brosimum and allied taxa.
Molecular Sue of the Dus nd relationships of parasitic angiosperm families have advanced
rapidly in recent y Barkman et al. 2007). Based on such work, the Balanophoraceae have
been firmly scene’ to order EN (Nickrent & Franchina 1990; Nickrent 1997 onwards; Stevens 2001
onwards), but their exact position was uncertain until the work of Nickrent et al. (2005). Based on 4 loci
from the plastid, nuclear, and mitochondrial genomes, this study clearly placed 3 selected members of this
family within a clade containing 3 genera representative of Santalaceae, Olacaceae, and Loranthaceae.
Knowledge of the relationships of Moraceae with allied families, and of generic groups within the fam-
ily, has been clarified by recent molecular studies (Sytsma et al. 2002; Datwyler & Weiblen 2004; Zerega et
al. 2005). Changes in the familial and tribal assignments of certain genera mentioned above are based on
Datwyler and Weiblen (2004) and Zerega et al. (2005). The latter paper involved a combined phylogenetic
analysis of 26S and ndhF gene sequences and included proposals for phylogeography and divergence times
of the higher taxa. For Moraceae tribe Dorstenieae, a stem group age of 48 mybp was estimated, while for
Brosimum and its 2 closest relatives, the crown group age was placed at 28 mybp. Because the family place-
ment of Trochanthera is uncertain, and our analysis may not have exhausted all possibilities, we can only
point out that the fossil falls within an age of interest for presumed evolutionary diversification in both of
the families discussed above. M Qe aes parallelisms abound in the angiosperms, and in this case, a
similar form of infl in ifferent families prevents us from proposing a more exact assignment
for this distinctive Tertiary me
ACKNOWLEDGMENTS
The authors thank the herbarium curators of the New York Botanical Garden (NY) and Missouri Botanical
Garden (MO) for the loan of specimens of Balanophoraceae and Moraceae used in our study. We are grateful
to P.K. Endress, R.K. Eberwein, and an anonymous reviewer for their comments and suggestions. We also
thank J. Strother for providing us with samples of Brosimum.
REFERENCES
Barkman, T.J., J.R. McNEAL, S-H. Lim, G. Coat, H.B. Croom, N.D. Young, AND C.W. DerAMPHiLIS. 2007. Mitochondrial DNA
suggests at least 11 origins of parasitism in angiosperms and reveals genomic chimerism in parasitic plants.
BMC Evol. Biol. 7:248.
Bera, C.C. 1972. Olmedieae, Brosimeae (Moraceae). Fl. Neotropica 7:1-228.
Bera, C.C. 1989. Systematics and phylogeny of the Urticales. In: PR. Crane and S. Blackmore, eds. Evolution, sys-
tematics and fossil history of the Hamamelidae. Vol. 2. Clarendon Press, Oxford. Pp. 193-220.
DarwyrLer, S.L. AND G.D. Weeen. 2004. On the origin of the fig: phylogenetic relationships of Moraceae from ndhF
sequences. Amer. J. Bot. 91:767-777.
Poinar et al., Trochantt lepidota g and sp. nov. 1173
Draper, G., P. MANN, AND J.F. Lewis. 1994. Hispaniola. In: S. Donovan and T.A. Jackson, eds. Caribbean geology: an
introduction. The University bis n bus RE aude Mic Jamaica. Pp. 129-150.
EBERWEIN, R.K. AND A. Weer. 2004. £. L l | | gy and transfer to Helosis. Bot.
J. Linn. Soc. 146:513-517.
ENGLER, A. 1889. Balanophoraceae. In: A. Engler and K. Prantl, eds. Die Natürlichen Pflanzenfamilien III. 1. Verlag
Wilhelm Engelmann, Leipzig. Pp. 243-263.
ENGLER, A. 1894. Moraceae. In: A. Engler and K. Prantl, eds. Die Natürlichen Pflanzenfamilien III. 1. Verlag Wilhelm
Engelmann, Leipzig. Pp. 66-98.
Fais, |. 2006. Bertel Hansen (1932-2005). Taxon 55:1055-1056.
GowzALEz, F. 2004. Balanophoraceae. In: N. Smith, S.I. Mori, A. Henderson, D.W. Stevenson, and S.V. Heald, eds.
Flowering plants of the Neotropics. Princeton University Press, Princeton, NJ. Pp. 41-43
Hansen, B. 1972. The genus Balanophora J.R. and G. Forster. A taxonomic monograph. Dansk Bot. Arkiv. 28:
1-188.
Hansen, B. 1976. Balanophoraceae. Fl. Malesiana 7:783-805.
HANSEN, B. 1980. Balanophoraceae. Fl. ND 23: iu
Hansen, B. AND K. ENGELL. 1978. Inflores« leae, Lophophytoid | Scybalioideae (Balano-
phoraceae). Bot. Tidsskr. 72:177-187.
Harms, H. 1935. Balanophoraceae. In: A. Engler and H. Harms, eds. Die Natürlichen Pflanzenfamilien 16b. Verlag
Wilhelm Engelmann, Leipzig, pp. 296-339.
Hooke, J.D. 1856. On the structure and affinities of Balanophoreae. Trans. Linn. Soc. London 22:1-68.
Hutchinson, J. 1967. The genera of flowering plants (Angiospermae). Clarendon Press, Oxford.
ITURRALDE-VINCENT, M.A. AND R.D.E. Macphee. 1996. Age and paleogeographic origin of Dominican amber. Science
273:1850-1852.
Kuur, J. 1969. The biology of parasitic flowering plants. University of California Press, Berkeley.
Newune, L.i. 1960. Flora of Panama: Balanophoraceae. Ann. Missouri Bot. Gard. 47:303-308.
Nickrenr, D.I. 1997 onwards. The parasitic plant connection. <www.parasiticplants.siu.edu>
Nickrent, D.I. AND C.R. Franchina. 1990. Phylogenetic relationships of the Santalales and relatives. J. Molec. Evol.
31:294-301.
NickRENT, D.I., J.P. DER AND FE. ANDERSON. 2005. Discovery of the pl ynthetic relatives of the"Maltese mushroom"
Cynomorium. BMC Evol. Biol. 5:38
Ponar, Jr, G.O. 1992. Life in amber. Stanford University Press, Stanford.
Ponar, JR, G.O. AND R. Pomar. 1999. The amber forest. Princeton University Press. Princeton, NJ.
Rouwer, J.G. 1993. Moraceae. In: K. Kubitzki, J.G. Rohwer, and V. Bittrich, eds. The families and genera of vascular
plants. Springer-Verlag, Berlin. Pp. 438—453.
SchLee, D. 1999. Das Bernstein-Kabinett. Stuttgarter Beitr. Naturk,, Ser. C, 28.
Srevens, PF. 2001 onwards. Angiosperm Phylogeny Website. Version 8, June 2007 [and more or less continuously
updated since]. http:// bot.org/MOBOT/research/APweb
Sytsma, KJ., J. Morawerz, J.C. Pines, M. NepOKROEFF, E. Conti, M. ZiHRA, J.C. HALL, AND M.W. Chase. 2002. Urticalean rosids:
circumscription, rosid ancestry, and phylogenetics based on rbcL, trnl-F, and ndhF sequences. Amer. J. Bot.
89:1531-1546
Woobson, R.E. 1960. Flora of Panama. Moraceae. Ann. Missouri Bot. da LAO
ZEREGA, N_J.C., W.L. CLEMENT, S.L. DATWYLER, AND G.D. WElBLEN. 2005. Biogeography liverg ti in the mulberry
family (Moraceae). Molec. Phylogen. Evol. 37:402-416.
1 1 £41. Des sa) D h Inctitut f Texas 2(2)
1174
BOOK NOTICE
NEAL K. VAN ALFEN, GEORGE BRUENING, AND WILLIAM O. Dawson (eds). 2008. Annual Review of Phytopathol-
ogy: Volume 46. (ISBN 978-0-8243-1346-3, hbk; ISSN 0066-4286). Annual Reviews Inc., 4139 El
Camino Way, PO. Box 10139, Palo Alto, CA 94303-0139, U.S.A. (Orders: www.AnnualReviews.org,
science@annualreviews.org, 800-523-8635, 650-493-4400, 650-424-0910 fax). $84.00 indiv., 417
pp., 7 5/8" x 9 3/8".
Contents of Volume 46:
1. The Phenotypic Expression of a Genotype: Bringing Muddy Boots and Micropipettes Together—Roger Hull
2. The Origin of Ceratocystis fagacearum, the Oak Wilt Fungus—Jennifer Juzwik, Thomas C. Harrington, William L. MacDonald, and David
N. Appel
The pud M A o iden id iss s Most Familiar (Yet iid Known) Plant Pathogens— Dean A. Glawe
Plants as a E a Bacteria—Heather L. Tyler and Eric W. Triplett
TheO f Plant Pat! Agro-Ecosystems aa and Bruce A. McDonald
Melotto, William Underwood, and Sheng Yang He
illigan
Role of Somn in Plant Iia a and Foliar E. Diseases—Maeli M
Models of Fungicide Resistance Dynamics—Frank van den Bosch and Christopher A. Gi
pd in Fingal Physiology db Haas, Martin Eisendle, and B. Gillian Turgeon
rece Góhre and Silke Robatzek
i RUM dM
Microbial
10. Yeast asa Model Host to Explore Plant Virus-Host Interactions—Peter D. Nagy
11. pad in two Worlds: The Plant and Insect Lifestyles of Xylella Doce M Chatterjee, Rodrigo P. P. Almeida, and Steven
12. identification and Rational a of Novel Antimicrobial Peptides for Plant Protection—Jose F. Marcos, Alberto Muñoz, Enrique
rez-Payá, Santosh Misra dB pez-García
13. Direct and Indi t Rol f Viral S f RNA Silencing i in Pathogenesis—Jua nA. Díaz-Pendón and Shou- Wei i Ding
14. Insect Vector Interactions with Per enil Transmitted Viruses—Saskia A. Hogenhout, El-Desouky Ammar, Anna E. Whitfield, and
DAN
Melee: Redinbaug!
15. Plant V. I Materials and Their Use in Nanotechnology— Mark Young, Debbie Willits, Masahi Uchida, and Trevor
Douglas
16. Epidemiological Models for I i d Persist f Pathog Christopher A. Gilligan and Frank van den Bosch
J. Bot. Res. Inst. Texas 2(2): 1174. 2008
CRETACIFILIX FUNGIFORMIS GEN. AND SP. NOV., AN EUPOLYPOD FERN
(POLYPODIALES) IN EARLY CRETACEOUS BURMESE AMBER
George O. Poinar, Jr. Ron Buckley
iod of Zoology 9635 Sumpter Ridge
tate University Florence, Kentucky 41042-8355, U.S.A.
e ind 97331, U.S.A.
ABSTRACT
An eupolypod fern, Cretacifilix ee gen. & sp. nov. crm is described from four partial, fertile leaf blades in a piece
of Early Cretaceous Burmese amber. Di : oval, thick-walled sporangia with vertical, short annuli not extending
more eae half the See of the MR deeply we circular sori with brown, multicellular, round to reniform, per-
in d at the sinus and ruggose-tuberculate, oval, monolete spores. This discovery shows that eupolypod ferns and
angiosperms were evolving concurrentiy at the Burmese amber site in the Early Cretaceous.
Key Wonps: amber, Myanmar, Early Cretaceous, eupolypod fern
RESUMEN
; 1 J nido Kgentanifil:. f. TJ " - {Dal Aialac 2h E E ee fiel
Se describe un helecho eupoly; : g gen. & sp. no YI I I j ;
de una po = ámbar del Cretácico temprano de Birmania. 1 ract diagnósti D gi l pared gruesa con
:11 LE A A ] 1; < q A Pi 1 T E 1 f: J Tot J
AERE E O > E ,
i f , :
con indusios y s marrones, multicelulares, de redondos a reniformes, unidos en el seno, y esporas monoletas, ovales, rugoso-
Feta Hd he > 1 hel 1 1 JT 22 1 E 1 H : Ws ]
tuberculadas I polyp y gios[
Cretácico temprano en el lugar del ámbar birmano.
INTRODUCTION
Amber from Burma (Myanmar), which has been known for centuries, contains a variety of insect and plant
fossils (Poinar et al. 2006). A recent discovery revealed the remains of four fertile pinnules of an eupolypod
fern in a piece of Burmese amber. Fern remains are rare in amber and only three species have been described.
They are Alethopteris serrata Caspary (1881) and Pecopteris humboldtiana Góppert and Berendt (1845) from
Eocene Baitic amber (both of the above extinct genera had been established previously for Carboniferous
ferns) and Grammitis succinea Gómez (1982) in Miocene-Oligocene Dominican amber.
The present study describes the Burmese amber fern using characters associated with the sori, indusia,
sporangia and spores.
MATERIALS AND METHODS
The amber was derived from a mine first excavated in 2001, in the Hukawng Valley, southwest of Maing-
khwan in the state of Kachin (26920'N, 96°36’E) in Burma (Myanmar). This new amber site, known as the
Noije Bum 2001 Summit Site, was assigned to the Upper Albian of the Farly Cretaceous on the basis of
paleontological evidence (Cruickshank & Ko 2003). Nuclear magnetic resonance (NMR) spectra and the
presence of araucaroid wood fibers in amber samples from the Noije Bum 2001 Summit site indicate an
araucarian (probably Agathis) tree source for the amber (Poinar et al. 2007).
The single piece of amber containing the fern remains (Fig. 1) was roughly rectangular in shape,
measuring 12 mm long, 6 mm wide and 1 mm in depth. Observations, drawings, and photographs were
made with a Nikon SMZ-10 R stereoscopic microscope and Nikon Optiphot compound microscope (with
magnifications up to 700x). It is not possible to determine whether the blades were pinnate, bipinnate or
tripinnate. The lengths of the four fertile pinnules in the amber are: pinnule A, 3 mm; pinnule B, 4.9 mm;
pinnule C, 5.2 mm and pinnule D, 2.4 mm. Each pinnule contains one or more sori on their abaxial surfaces
(Figs. 1, 5).
J. Bot. Res. Inst. Texas 2(2): 1175 — 1182. 2008
1176 J lof the Botanical R h Institute of Texas 2(2)
DESCRIPTION
Cretacifilix G.O. Poinar & R. Buckley, gen. nov. Tre Species: Cretacifilix fungiformis G.O. Poinar & R. Buckley (Polypodiales).
Diagnosis.—Rhizome, stipe, rachis and costa unknown. Fronds of unknown length; pinnules lobed, mostly
glabrous with few scattered unicellular hairs on surface and margins; veins free, not reaching margins, not
ending in hydathodes; sori medial, orientated mostly in alternate positions in two rows, with mostly one,
but sometime two sori bone on single pinnule lobe, deeply embossed, resulting adaxial pustules at ends of
veins; sori covered by brown, thick, glabrous, multicellular, round to reniform, persistant indusia with stalk
attached by inner end of sinus with distal and lateral openings; some indusia bent backwards, revealing
ribbed stalks; sporangial stall posed of 1-3 rows of cells; sporangia oval, thick-walled, walls composed
of small, narrow, elongate cells; annulus vertical, short, broad, not extending more than half circumference
of sporangia, with less than 10 thick walled cells; spores small, oval, ruggose-tuberculate, monolete.
Cretacifilix fungiformis G.O. Poinar & R. Buckley, sp. nov. (Figs. 1-5). Tec: MYANMAR. Kacr: Noije Bum 2001
Summit Site in the Hukawng Valley, SW of Maingkhwan in the state of Kachin (26°20'N, 96°36'E), 2005-2007, collector unknown
(HoLoTYPE: catalogue # B-11, amber collection of Deniz Erin in Istanbul 34744, Turkey). This collection is available for study. Further
tal A
contact information can b m Ron Buckley at ronbuckley@fuse.net.
Description. —Four fertile pinnules: indusia (N = 22) opening outward, 416(360—504) pm in width, 233
um in length; indusial stalk 142(104-208) um wide, width of cap, 183(90-436) um; cells of indusia thick
walled, many filled with discrete, minute bacilliform bodies ranging from 3-4 um in length and 1-2 um in
width; stalk of indusia often ribbed (possibly from desiccation); sporangia oval-shaped; capsule walls com-
posed of thick walled, narrow, elongate cells; sporangia (N = 10), 158(140-189) um in length, 115(81-150)
um in width; annulus short, containing less than 10 thick-walled cells and extending no more than half
i f f sporangia; length annulus (N - 5), 5324-72) um; width annulus (N = 3), 5304— 72); spores
(N = 8) monolete, oval, 21(16-29) um long and 14(11-20) um wide, surface ruggose-tuberculate, perispore
not evident; hairs on margins and veins of pinnules range from 28-195 pm in length.
Etymology.— Cretacifilix is derived from the Latin Cretaceus for “chalky” indicating the Cretaceous
period and from the Latin “filix” for fern: fungiformis is derived from the Latin "fungi" for mushroom and
the Latin “forma” for shape in reference to the resemblance of the indusia to fruiting bodies of chanterelles
and some shelf-like mushrooms.
DISCUSSION
Since the vegetative structures and habit of Cretafilix are unknown, systematic placement is based on the
reproductive characters. The presence of a vertical annulus that partially encircles the spore case (Figs. 2,
5) places the fossil in the Eupolypods clade of the Polypodiales (Schneider et al. 2004; Smith et al. 2006).
It was not possible to place Cretacifilix in an extant family because of its unique combination of characters.
Surficial sori with reniform indusia are characters of the families Dryopteridaceae and Woodsiaceae (and
are also known from some members of the Thelypteridaceae, Tectari dI iopsid ) as define
by Smith et al. (2006). However members of the Dryopteridaceae and Woodsiaceae possess reniform, trilete
spores and the spores of Cretacifilix are oval and monolete. Also the indusia of these families are usually not
multicellular and thick as with Cretacifilix (Bower 1928; Sledge 1973).
Another character of Cretacifilix not found in the Dryopterid Woodsi ply embossed
sori (Plate 2). This charac ter closely resembles that found in the North American Pleopeltis polypodioides (L.)
E.G. Andrews & Windham of the Polypodiaceae (Cobb et al. 2005). This species, commonly known as the
resurrection fern, has the characteristic of curling up and appearing dead during periods of drought, but
then recovering when conditions are favorable. If embossed sori are associated with survival during periods
of aridity, it would suggest that Cretacifilix might have been subjected to similar climatic conditions. The
resurrection fern occurs on tree branches (Cobb et al. 2005), which is a likely habitat for Cretacifilix and
would explain how fronds of the fossil made contact with resin on the trunk of the araucaroid tree.
de] J ] 1
and Sp. nov (Polypodiales) 1177
| » N
e
" + i
v
Ar tht, E " ic AR n ah f PNG I ADD G loft
ight). Scale Bar = 1.2 mm. B. Pinnule B. Arrows
Fic. 1 g ber y
show brown, persistent indusia. Scale Bar — 0.6 mm.
2(2)
senda at T
ICAd»
1178
£ ic A Pustl
leurface &rala Bar — 1 mm BR. Inducium with ra-
A PELO ERO"
ar = 110 um C. Hairs (
F
d Scale B
y
Poi 1 Buckley, Cretacifilix fungiformi g and sp. nov. (Polypodiales) 1179
TOME TNT
wall. Arrow shows annulus. Scale Bar = 126 pm. C. L | vi porangi it | Vertical vi porangi
annulus. Scale Bar = 53 um.
Asian dryopterid species with similar sori are Arachniods standishii (Moore) Ohwi, Dryopteris lacera
(Thumb. O. Kuntze and Dry opteris chinensis (Bak.) Koidz. However, in the former two species, the veins
extend to the tips of the pinnules and the sori are either on the veins or in opposite rows, and in the latter
species, the annulus has 17 thick- walled cells and extends more than half way around the circumference
of the sporangium (Lee 1999).
Th 1 fextant l fi tend least two third d the ci f fthe
sporangial capsule (Moran 2004). The pu annuli, thick, narrow cell walls of the sporangial capsules and
small, oval spores TD to me unique features of Cretacifilix. The small bacilliform bodies in the indusial
cells could indicate a sy iation. No reports of similar bodies in the indusial cells of extant ferns
could be found.
The presence of an eupolypod fern in the Early Cretaceous supports other records of fossil Polypodiales
ferns from the same period (Deng 2002). Based on molecular studies, eupolypod subclades are thought
to have diversified in the Late Cretaceous (Schneider et al. 2004), but it is obvious that they were already
present in the Early Cretaceous and evolving concurrently with early angiosperms, several of which have
1180
I laf
SVULIIGI VI
+L
R h Institute of Texas 2(2)
elongate cells. A
4
g
£J pill.
= 37 um. B. Sporangium with a
n { Buckley, Cret DOE E SAE g and sp nov. (Polypodiales) T181
rH £ Ikad ineliici and tur^
Ei 5 ( T aT 2 [yy ic A Pi
sporanaia
»purailyta,
been recovered from the same Burmese amber fossil site (Poinar 2004; Poinar & Chambers 2005; Poinar et
al. 2007).
All evidence indicates that the Burmese amber forest was a tropical-subtropical rainforest with many
floral characteristics of the Waipoua kauri forest in Northern New Zealand. The Waipoua forest is the only
virgin Agathis forest in the world today and comprises a mixed dense ecosystem of flowering plants, ferns,
lycopods and moss on 90 sq km of land. Of the 241 plants that have been recorded, 178 species are seed
bearing and 63 are ferns or lycopods (McGregor 1948).
ACKNOWLEDGMENTS
The authors thank Alan Smith for assistance throughout this study, Kenton Chambers and Peter Bernhardt
for supplying pertinent literature and Roberta Poinar for comments on earlier versions of the manuscript.
REFERENCES
Bower, FD. 1928. The ferns (Filicales). Vol. IIl. The Leptosporangiate ferns. Cambridge University Press, London.
Caspary, R. 1881. Über neue fossile Pflanzen der Blauen Erde, des Bernsteins, des Schwarzharzes und des Braun-
karzes. Schrift. Physik.-Ókonom. Gesellsch. Kónigsberg 22:22-31.
Cose, B., E. FARNSWORTH, AND C. Lowe. 2005. A field Guide to ferns and their related families. 2" Ed. Houghton Mifflin
Co, Boston.
CRUICKSHANK, R.D. Ab K. Ko. 2003. Geology of an amber locality in the Hukawng Valley, northern Myanmar. J. Asian
Farth Sci. 21:441—455.
1182 Journal of the Botanical R h Instit f Texas 2(2)
DENG, S. 2002. Ecology of the Early Cretaceous ferns of Northeast China. Rev. Palaeobot. Palynol. 119:93-112.
Gómez, L.D.P. 1982. Grammitis succinea, the first New World fern found in amber. Amer. Fern J. 72:49-52.
GOPPERT, H.R. AND G.C. Berenot. 1845. Der Bernstein und die in ihm befinlichen Pflanzenreste des Vorwelt. In: Die
im Bernstein befindlichen organischen Reste der Vorwelt, Berendt, G. C. (Ed) 1:1-125.
Lee, T.B. 1999. Illustrated flora of Korea. Hyang-munsa, Seoul.
Mcerecor, WAR. 1948. The Waipoua Forest. Abel, Dykes Limited, Auckland.
Moran, R.C. 2004. A natural history of ferns. Timber Press, Portland.
Ponar, JR, G.O. 2004. Programinis burmitis gen. et sp. nov., and P laminatus sp. nov., Early Cretaceous grass-like
monocots in Burmese amber. Australian Syst. Bot. 17:497—504.
POINAR JR, G.O. AND K.L. CHAMBERS. 2005. Palaeoanthella huangii gen. and Sp. nov, an early Cretaceous flower (An-
giospermae) in Burmese amber. Sida 21:2087-2092.
Ponar, JR, G.O., R. BUCKLEY, AND A. Brown. 2006. The secrets of Burmese amber. Mid American Paleontol. Soc.
29:20-29.
POINAR, Jr, G.O., K.L. CHAMBERS, AND R. Buckiev. 2007. Eoépigynia burmensis gen. and sp. nov, an Early Cretaceous
eudicot flower (Angiospermae) in Burmese amber. J. Bot. Res. Inst Texas 1:91-96.
POINAR, JR., G., J.B. LAMBERT, AND Y. Wu. 2007. Araucarian source of fossiliferous Burmese amber: spectroscopic and
anatomical evidence. J. Bot. Res. Inst. Texas 1:449—455.
SCHNEIDER, H.E., K.M. ScHUETTPETA, R. PrYER, S. CRANFILL, S. MAGALLÓN, AND R. Lupia. 2004. Ferns diversified in the shadow
of angiosperms. Nature 428:553—557,
SLEDGE, W.A. 1973. The dryopteroid ferns of Ceylon. Bull. British Mus. (Nat. Hist) Bot. 5:1-43.
SMITH, A.R., KM. Pryer, E. ScHuerrPELZ, P. Korau, H. SCHNEIDER, AND PG. Wor. 2006. A classification for extant ferns.
Taxon 55:705-731.
AN EARLY CRETACEOUS ANGIOSPERM FOSSIL OF
POSSIBLE SIGNIFICANCE IN ROSID FLORAL DIVERSIFICATION
George O. Poinar, Jr. Kenton L. Chambers
Department of Zoology isis of Botany and inis Pathology
Oregon State University on State University
Corvallis, Oregon 97331, U.S.A mun Oregon 9733 i USA.
Ron Buckley
9635 Sumpter Ridge
Florence, Kentucky 41042-8355, U.S.A.
ABSTRACT
Lachnociona terriae gen. & sp. nov. is described from an Early Cretaceous flower preserved in Burmese amber. Lac)
resented by a small, hirsute, Rd. pistillate flower in a evenly. ls ncs pues 10 ra Dn and a a
5-carpellate pistil with 5 stout, erect, connivent or
stigmas. The gynoecium is na lanate- ind Tace d the filaments show thin pe of vestigial anther tissue at their tips. The
perianth is interpreted as a calyx, with th 2 p g ilies Brunelliaceae
and Cunoniaceae (order Oxalidales), and on selected les fi 2 oth id ord lus Saxifragal
RESUMEN
Se describe Lachnociona terriae gen. & sp. nov. d DU Lach-
r E
1 CA PA | : dam f
nociona está representada por una flor pequeña, hirsuta, f
filamentos estaminodiales, y un pistilo 5 lad 5 estilos f , erectos, conniventes o connatos, con los extremos muy asias
y recurvados, ia con estigmas ventrales decurrentes. Fl gi te lanoso-hirsuto. Tres de los filamentos
g perianto se i con la corola ausente. Se
discuten
E
1 hI 1 : r4 1 f. 215 n 17:
E
3 y Cunoniaceae E (iden Oxalidales), y en ejemplos selectos de
otros dos órdenes de rósidas y de Saxifragales.
INTRODUCTION
Burmese amber is first mentioned in works dating from AD 100, when an amber trade route was established
between that country and China. From AD 100 until 1936, the Burmese amber mines supplied amber for
jewelry and fossils. Social upheavals closed the mines for 63 years, but in 1999, extraction procedures be-
gan anew. The fossil was derived from a mine first excavated in 2001, in the Hukawng Valley, southwest of
Maingkhwan in the state of Kachin (26°20'N, 96°36'E) in Burma (Myanmar). This new amber site, known
as the Noije Bum 2001 Summit Site, was assigned to the Upper Albian of the Early Cretaceous on the basis
of paleontological evidence (Cruickshank & Ko 2003), placing the age at 97 to 110 mya. Nuclear magnetic
resonance (NMB) spectra and the presence of araucarioid wood fibers in amber samples from the Noije Bum
2001 Summit Site indicate an araucarian (possibly Agathis) tree source for the amber (Poinar et al. 2007).
Within the past few years, amber from this area has yielded a wealth of arthropod and plant remains
(Poinar et al. 2005). While insects dominate, some very i ing angiosperm fossils have been recovered
up to now, including flowers with possible affinities to the families Monimiaceae (Poinar & Chambers 2005)
and Cornaceae (Poinar et al. 2007), as well as two early bambusoid pau vena 2004). In the present
paper, we describe an additional flower from the same location, with simi il and extant members
of eurosid orders Oxalidales, Cucurbitales, Sapindales, and eudicot order Saxifragales.
MATERIALS AND METHODS
An entire, well-preserved female flower with staminodes (accession # ab-294) is embedded in a round piece
J. Bot. Res. Inst. Texas 2(2): 1183 — 1192. 2008
1184 J lof the Botanical R h Institute of Texas 2(2)
of amber with a diameter of 40 mm and a thickness of 5 mm. Examination and photographs were made with
a Nikon stereoscopic microscope SMZ-10 R at 80x and a Nikon Optiphot microscope at 800x.
DESCRIPTION
Lachnociona G.O.Poinar, K.L. Chambers & R. Buckley, gen. nov. Tere Srectes: Lachnociona terriae G.O. Poinar, K.L.
Chambers & R. Buckley, sp. nov.
Diagnosis.— Flowers small, pedicellate, unisexual, the type specimen functionally pistillate, 5-merous,
actinomorphic, apparently apetalous; sepals 5, regular, separate, ovate-lanceolate, recurved, ciliate, lightly
hirsutulous on both surfaces, venation obscure; staminodes 10, in 1 or 2 whorls, filaments conspicuous,
slender and tapering, the tip retrorsely hooked in some, the fully preserved ones all non-functional, anthers
absent or represented by thin vestigal tissue on 3 staminodes; gynoecium 5- carpellate, densely lanate-hirsute,
ovary superior or perhaps half-inferior, it and the disc (if present) concealed by the pubescence, styles con-
nivent or partly united, their tips sharply acute, recurved-divergent in the upper 1/4-1/3, the stigma probably
decurrent ventrally; staminate flower and fruit unknown.
Lachnociona trae G. O.Poinar, us B Chambers d R. Buckley sp. nov. (Figs. 1-4). Tre: MYANMAR (Burma):
Vall V (26° ind e m ons a 2001, unknown
vind miner s.n. Creme accession no. db: Bi pus amber, d ited in tl bul, Turkey, 34744).
I
ing Ron Buckley at ronbuckley@fuse.net.
Description. “Hower ic 5.6 mm; flower Puis 5 mm; sepal length 3.60 mm, width 1.8 mm; length of
trichomes on sepals 0.25-0.36 mm; staminodes exserted, most broken or chewed off; length of remaining
filaments 1.80-1.90 mm; length of vestigial anthers 0.11-0.88 mm; width of largest anther 0.20 mm; length
of gynoecial column 3.30 mm; length of divergent upper lobes of style 1.20 mm, length of subtending bract
on pedicel 1.10 mm; distance from base of bract to summit of hirsutulous pedicel 2.90 mm.
Etymology.—Genus name from the Greek “lachnos,” woolly hair, and “kion,” column, with reference to the
projecting stylar portion of the gynoecium. Species named in honor of the collector, Mrs. Terri Millinoff.
DISCUSSION
The salient features of the fossil, as described above, are its unisexuality, apparent apetaly, 10 staminodes,
densely pubescent, 5-carpellate gynoecium, stout stylar column, and acute, spreading stigmatic lobes. The
most reasonable interpretation of the ovary is that it is either superior or, judging from the expanded distal
region of the pedicel, half-inferior. The vestigial anther tissue on 3 of the filaments is noteworthy, suggesting
unisexuality by abortion, hence derivation from hermaphrodite ancestors. In framing a hypothesis for the
systematic position of Lachnociona, we have searched for present-day families which best fit the above group
of traits, if reasonable assumptions are made about features not clearly visible in the fossil. Because of the
thick covering of darkly pigmented hairs, microscopic study of this structure could not resolve the choice
between fused or connivent styles. As well, the recurved perianth parts with dark coloration at their base
prevented observation of any tiny, outer perianth parts beneath them, and we thus interpret the perianth
as a calyx, no E corolla ae ene Ol or oe
ly associated families Brunelliaceae and Cunoniaceae
(Oxalidales) (APG II 2003; Stevens 2001 onwards; Soltis et al. 2005), both groups occurring mainly in the
Southern Hemisphere with rep ives, in the latter case, in the Australian-Southeast Asian region. We
interpret the stout, densely hairy, darkly colored (in the fossil) columnar structure to be, all or in part, 5
connivent-appressed styles. The Late Cretaceous fossil Platydiscus (Schónenberger et al. 2001; Schónenberger
& von Balthazar 2006; ns zt 2000) was Ded in our analysis, because of its initial association with
Cunoniaceae. Also, Anisoy ll of order Cucurbitales, and Aphanopetalum (Aphanopeta-
laceae), of order As, came under consideration. Eurycoma (Simaroubaceae, order Sapindales) was
added for a comparison involving a different interpretation of the perianth of the fossil.
Table 1 arison of floral traits of Lachnociona with four living members of Brunelliaceae
and Cunoniaceae, together with mda. Anisophyllea, Aphanopetalum and Eurycoma. Brunelliaceae and
Poinar et al., Lachnociona terriae gen. and sp. nov 1185
-
2
BLE 1. C i f Lachnoc ith fossil and i (Platydiscus, Spiraeanthemum, Pullea, Vesselowskya), B y
phyllea), Aphanopetal (Aphanopetalum), and Si t (Eur) )
Lachnociona Platydiscus Spiraeanthemum Pullea
1. Pedicel present - +, absent - 0 T + + +,0
2. Sepals free - +, connate* — 0 + + ,0 +
3. Sepal number 5 4 4(5) 5(6)
4. Sepals spreading - +, upright — O t + 0 +
5. Petals absent - +, present - 0 + 0 + +
6. Flowers unisexual - +, bisexual — 0 + 0
7. Stamen number 10 8 8(10) 10(12)
8. Ovary superior - +, +, inferior — O +or0 0 + 0
9. Ovary Pr - +, syncarpous - 0 +or0 0 +" 0
10. Carpel nu 5 4 (2-3)4 or 5 2
11. Styles den +, separate — 0 +ord 0 0 0
12. Style tips acute - +, capitate — O + +° 0 +
13. Stigma decurrent - +, terminal — 0 +? +° 0 0
14. Gynoecium hairy - +, glabrous — 0 + + + +
15. Free styles connivent-+,spreading-O +? +° O(+) O(+)'
(genera continued) Vesselowskya Brunellia Anisophyllea Aphanopetalum Eurycoma
1 + + 0 + +
2 TS " + + +(small)
3 3(4) (4)5(6) (3)4(5) 4 5
4 ? +or0 0 0
5 +or 0° + 0 O(+, small) +(large)
6 + + (0) + 0 (0)
7 6(8) (8)10(12) (6)8(10) 8 5 large
8. + + 0 0 +
9. 0 + 0 0 +
10. 2 (3-4)5(6) (3)4(5) 4 5
11. 0 0 0 + Dor+
12. T + + + +
13, + + + 0 +
14. + + 0 0 +
15. 0 04 + - +
Footnotes: * t ing basal fusi pistil in 1/2 inferior ovaries; *— a Dey free above; * — laterally connate at base;
—style tips broken below ti tructed itl I sli ?); °° — spreading above in reconstruc-
tion, but connivent below in figs. 21, 34 in Schonenberger et al., 2001; 1 — connivent when is see s Diclison 1975, fig. 15h; ?— female
flowers have only sepals, Dickison 1989; 3 — separating in fruit.
References: 1970; Dickison 1975, 1989; Engler 1930a,b, 1931; Friis et al. 2006; Hermsen et al. 2006; Heywood 1978, p. 158;
Hoogland 1979; Hufford & Dickison 1992; Kubitzki 2004, 2007; Matthews et al. 2001; Matthews & Endress 2004; Nooteboom 1962;
Orozco Pardo 2002; Perry 1949; Schónenberger et al. 2001; Smith 1952, 1985; Takhtajan 1996.
Cunoniaceae, imes united into one family, have always been considered as closely related in traditional
theories of angiosperm phylogeny (e.g., Engler 1930a, b; Thorne 1992; Cronquist 1981). The fossil is similar
to Brunellia in being unisexual, apetalous, 10-stamened, and having a 5-carpelled, pubescent gynoecium
with acute style tips whose stigmatic surface is probably ventral and decurrent. Brunellia is apocarpous, the
carpels at most being immersed in the disc (Orozco Pardo 2002; Kubitzki 2004). Its separate styles may be
vertical in early flower (Cuatrecasas 1970; Kubitzki 2004), with recurved tips. In Cunoniaceae, the carpel
number is usually 2 and the stigmas, except for Vesselowskya, are terminal and often capitate (Bradford et
al. 2004). However, higher carpel numbers are known, e.g., in Pullea, Aistopetalum, Spiraeanthemum, and
: laf tha Botanical D h Institute of Texas 2(2)
1186
Fic. 1. Fl f Lachnocí jae in B ber. Holotype speci Scale bar = 1.43 mm.
Acsmithia, the latter two genera comprising a possible morphological link with Brunelliaceae (Orozco Pardo
2002; but see Bradford & Barnes, 2001, for DNA evid ing their inclusion in Cunoniaceae). Genera
of Cunoniaceae with unisexual flowers include Sviraeanilicmum: Pancheria, and Vesselowskya, and the latter
genus is unique in the family in showing a ventrally decurrent stigma like that of Brunellia (Dickison 1989,
fig. 4.3; Hufford & Dickison 1992) and putatively like that of Lachnociona. That the ovary of the fossil might
be half or more inferior (judging from the expanded apex of the floral pedicel) was mentioned above; in
this, it would resemble Schizomeria (Matthews & Endress 2002) and Pullea (Dickison 1975) and differ from
Spiraeanthemum and Vesselowskya. The styles may be erect and partially connivent in some Cunoniaceae (at
Poinar et al., Lachnociona terriae gen. and sp. nov. 1187
Fics. f Lachnociona terriae i
Note reflexed tip of filament. Scale bar = 123 pm.
least in early flower), as illustrated for Pullea and Acsmithia by Dickison (1975, fig. 15h, and 1989, fig. 4.2B,
respectively) and Hoogland (1979, fig. 1). Of the above-mentioned genera, petals are absent in Brunellia, Spi-
raeanthemum, Acsmithia, Pullea, and (in pistillate flowers) Vesselowskya (Dickison 1989). None of the modern
genera listed have styles pubescent nearly to the tip like Lachnociona.
Aphanopetalum (Aphanopetalaceae) is included in Table 1; it has a 4-carpelled gynoecium with half-
inferior ovary, and a stout stylar column with spreading, terminally-stigmatic tips (Dickison 1975, fig. 1a).
Dickison (op. cit.) shows a median longitudinal section of a flower nearing anthesis; its thick style of 4
fused carpels with spreading, bluntly pointed tips very much resembles the stout (but more acutely tipped)
gynoecium of Lachnociona. Significant differences are evident, however, in the androecium of 8 large, fertile,
nearly sessile anthers of Aphanopetalum, the 4-merous calyx, and the presence of 4 poorly developed pet-
als. The flowers of Aphanopetalum are glabrous, as well. This genus has an interesting taxonomic history,
having been placed originally in Cunoniaceae (Engler 1930b; Dickison 1975; Hufford & Dickison 1992),
but it was found by Bradford and Barnes (2001) to be better placed in or near Saxifragaceae, according to
their analysis of the TrnL-F intron of the chloroplast tRNA-Leu gene. Based on the morphological work of
i A chew T h Institute of Texas 2(2)
1188
Fic. 4. D n ££] Elack H . PD
Dickison et al. (1994) and the molecular phylogenetic studies by Fishbein et al. (2001) and Hermsen et al.
(2006), Aphanopetalum is now placed as a monotypic family of the order Saxifragales (Stevens 2001 onwards;
Soltis et al. 2005).
Platydiscus is a well-studied charcoalified Late Cretaceous fossil flower from sand and clay sediments
of the Kristianstad Basin in Scandia, Sweden (Schónenberger et al. 2001, see drawing of a reconstruction
in Friis et al. 2006). The perianth is 4-merous, with small, densely glandular petals and spreading sepals,
stamens are 8 and fertile, and the gynoecium of 4 carpels has a half-inferior ovary and connivent, erect styles.
The spreading stylar tips are reconstructed as acute with a ventral slit; the stigmatic region is not described
but may well have followed the slit as, e.g., in Brunellia and Vesselowskya (our suggestion, not the authors”.
The connivent styles, with broken off tips, are well illustrated by Schónenberger et al. (2001, figs 34 and
40). Features of the gynoecium of this fossil, as illustrated and reconstructed, are thus quite reminiscent of
Lachnociona as we visualize it. Differences between our fossil and Platydiscus are that the latter is 4-merous
throughout, bisexual, and petaliferous. Its age, given as Late Santonian—Early Campanian, is approximately
20 my younger than that of the fossil reported here from the Early Cretaceous.
The possible phylogenetic relationship of Platydiscus to Cunoniaceae and Anisophylleaceae, and that of
the latter two groups to each other, was discussed by Matthews et al. (2001), who pointed out the anomalous
position of the latter family in the order Cucurbitales (see also Tobe & Raven 1988). However, this ordinal
assignment is well borne out by recent molecular studies (Schwarzbach & Ricklefs 2000; Zhang et al. 2006).
Poinar et al., Lachnociona terriae gen. and sp. nov 1189
The flowers of Anisophyllea are similar in many ways to those of C i d Platydiscus, as cited in detail
by Matthews et al. (2001). Like Lachnociona, Anisophyllea is unisexual, with two whorls of stamens, separate
acute styles, and stigmas decurrent ventrally (Matthews et al. 2001; Matthews & Endress 2004). It differs
in being petaliferous and usually 4-merous throughout, and its ovary is fully inferior (Table 1). These and
numerous other traits of Anisophylleaceae are discussed by Matthews and Endress (2004), giving support
to its placement in Cucurbitales despite its divergence in floral form from other families of the order.
An alternative interpretation of the perianth of Lachnociona, namely that the conspicuous organs are
petals and that an inconspicuous calyx is out of sight beneath them, leads to comparison with a different
rosid group, the family Simaroubaceae of order Sapindales. We have selected the Southeast Asian genus
Eurycoma (Engler 1931, fig. 175; Nooteboom 1962, p. 206) for such a comparison (Table 1). In Eurycoma,
the conspicuously hairy flowers are unisexual, the 5-lobed superior ovary of 5 free carpels bearing 5 erect,
connate or connivent styles which are acute, recurved at the tip, and stigmatic ventrally. In fruit, the 5 carpels
separate and spread far apart, as occurs also in Brunellia. The androecium of pistillate flowers consists of 5
normal-sized stamens with abortive anthers, alternating with 5 scale-like, much modified staminodes. The
5 hairy petals are spreading-erect and have 5 small sepals at their base. Other than stamen number, there-
fore, and reflexed rather than erect petals, Lachnociona would resemble Eurycoma—if we hypothesize that a
small calyx is hidden below the “petals.” The fossil thus bears an ambiguous relationship to Simaroubaceae,
which we cannot resolve with certainty. In phylogenetic analyses, Simaroubaceae, sensu stricto, are placed
in the rosid II clade (Fernando et al. 1996; Gadek et al. 1996; APG II 2003; Stevens 2001 onwards; Soltis et
al. 2005; Schónenberger & von Balthazar 2006).
In summary, the case for postulating a relationship of Lachnociona to the clade containing Brunelliaceae
and Cunoniaceae rests on the similarity of the following critical floral features: unisexuality, 5-merous ac-
tinomorphic perianth, apetaly, abundant pubescence, presence of 10 staminodes in the pistillate flower, 5
uper a stout gynoecial column (which pu be) comprised of 5 connivent styles, and sharply acute stylar
ventrally decurrent sti A superior ovary or ovaries, if present, would be consistent
with these oe butifthe ovary is Ea as half-inferior, a similarity would exist to Pullea and a few
other Cunoniaceae (Bauera, f. Dickison, 1975; Ceratopetalum, f. Engler, 1930b; Codia, f. Hufford & Dickison,
1992) as well as to Aphanopetalum and the completely epigynous Anisophyllea. A decurrent “sutural” stigma
is characteristic of Brunellia (Kubitzki 2004) and Vesselowskya of Cunoniaceae (Dickison 1989; Hufford &
Dickison 1992).
Our suggestion of a connection with Brunelliaceae and Cunoniaceae cannot be proven, given the dif-
ficulty in observing certain critical features of the gynoecium of Lachnociona. However, we believe that the
totality of floral features favors placement of the fossil in the rosid clade of eudicots (e.g., Fernando et al.
1995; Soltis & Soltis 2004; Soltis et al. 2005; Schónenberger & von Balthazar 2006). Except for Saxifragales,
the four orders from which we have selected comparative ampie are in this large group of angiosperms.
According to Friis et al. 2006, p. 7), “(t)he presence of distincti lpate pollen grains in Late Barremian-
Early Aptian strata (about 120 Myr) shows that eudicot angiosperms were established in geographically
widespread regions early in the history of angiosperms.” Estimates for the ages of crown and stem groups
of basal eudicots were presented by Anderson et al. (2005) using rbcL sequences and multiple fossil refer-
ence points. Their inferred stem and crown group ages for rosids are 112 mybp and 108 mybp, respectively,
and for Saxifragales, the figures are 108 mybp and 102 mybp. In a different analysis, Crepet et al. (2004)
dated the minimum age of the “rosid I" clade as 94+ mybp. Both these estimates are younger than that of
Wilkstróm et al. (2001), which dates the stem group of the rosids at 125 mybp. As pointed out by Friis et
al. 2006, p. 3D, “(a) rapid diversification of eudicots, with the emergence of all lineages of basal eudicots
before the end of the Albian, has....been inferred, using molecular dating methods." In this context, the
eudicot flowers that have recently been described from Burmese amber of the Late Albian (Poinar 2004;
Poinar & Chambers 2005; Poinar et al. 2007), together with the present report, may help open a window
to particular features of floral evolution in this period of eudicot diversification.
1190 J l of the Botanical R h Institute of Texas 2(2)
The flower described here may have been insect-pollinated, as proposed earlier by Santiago-Blay et al.
(2005), and it could have been visited by Melittosphex burmensis, a small bee recently described from these
deposits (Poinar & Danforth 2006). The flower also shows evidence of herbivory, as there is obvious damag
to the filaments and anthers by chewing insects.
ACKNOWLEDGMENTS
The authors thank Roberta Poinar for reading earlier drafts. We are grateful for technical suggestions by 2
anonymous reviewers.
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A PHYLOGENY OF BEJARIA (ERICACEAE: ERICOIDEAE)
BASED ON MOLECULAR DATA
Catherine M. Bush and Kathleen A. Kron
Wake Forest University
1834 Wake Forest Road
Winston-Salem, North Carolina 27106, U.S.A.
bushcm3@wtu.edu
ABSTRACT
Phyl ic relationships of f the fif ized fB ined usi lecular (ndhF 5', matK; chloroplast
a vas; nn daté ipd PE were performed in PAUP dis i gene, combined e data ix total onus
7 J J
O Na Dod 0.2 TH lad 1 Pn holorini n 1 sh
W 1U
Bryanthus and
strong support. Bejaria racemosa, the ony species e North America, is tonis —— as sister to all other Bejaria eens
RESUMEN
Las relaciones suena de 10 de nu I idas de Bejaria f inad ilizando d lecul NES, matK,
1; J 1 A A 1 1 1 T h; A
y waxy). I I PAUP para cada gen, p yp
totales. I AID ` 1 Men 1 1 waxy. | PES A 1 1 1 3 1 4
4 4 E e Li r Jr
totales. E t álisis la tribu Bejarieae (Bejaria, Bryanthus y Ledoth ) filética. E dos | álisis, los miembros
muestreados de Bejaria filéti porte alto. Bejaria racemosa, la úni pecie de Norte América, f ] m
hermana de los otros miembros de Bejaria.
Bejaria Mutis ex. L. (Ericoideae: Ericaceae) contains fifteen species distributed in South America, Central
America and the Caribbean with B. racemosa occurring in the southeast United States (Clemants 1995).
The Ericaceae typically have a corolla that is fused basally and a five-merous corolla; therefore Bejaria was
thought to be the most ancestral within the family due to its free petals and higher merosity (Stevens 2007,
Camp 1941). This idea is not supported in recent studies (e.g., Kron et al. 2002) that show Enkianthus Lour.
(Enkianthoideae) as sister to all other taxa in the Ericaceae. Among the eight recognized subfamilies in
Ericaceae the Ericoideae clade contains five named groups (tribes): Bejarieae, Empetreae, Ericeae, Phyl-
lodoceae and Rhodoreae. Relationships among these five tribes are unresolved (Kron et al. 2002). Bejaria,
Bryanthus S.G. Gmel. and Ledothamnus Meisn. comprise Bejarieae (Kron et al. 2002). Ledothamnus is a genus
of nine species occurring in the Guyana Highlands and Bryanthus is monotypic, found only in Kamtchatka
and Japan (Mabberley 1997).
The most recent taxonomic revision of Bejaria was based on morphological characters and there are two
sections in the genus: sect. Racemosae and Bejaria (Clemants 1995). Fourteen of the species are in sect. Be-
jaria, while B. racemosa is the only member of sect. Racemosae, unique in its chartaceous leaves, the “stalked”
appearance of its inflorescence (i.e., there is a relatively long area at the base of the raceme that does not
contain leaves or flowers) and numerous distinct leaf anatomy characteristics (Clemants 1995).
After being discovered in 1767 in Colombia by Jose Celestino Mutis, Bejaria has been taxonomically
revised three times in its entirety (Fedtschenko & Basilevskaja 1928; Mansfeld & Sleumer 1935; Clemants
1995). Fedtschenko and Basilevskaja (1928) focused their revision on the characteristics of pubescence and
also inflorescence type. Mansfeld and Sleumer (1935) based their revision on plant i and corolla
size. Finally, Clemants (1995) based his monograph of the genus primarily on corolla shape, as opposed to
the traditional use of indumentum and inflorescence characters.
Four species of Bejaria—B. aestuans, B. resinosa, B. sprucei, and B. mathewsii—are widespread in distri-
bution, particularly B. aestuans which is found from Mexico to Bolivia (Clemants 1995). The other eleven
J. Bot. Res, Inst. Texas 2(2): 1193 — 1205. 2008
I Inha D -ai D MENTRE
1194 J al oft t fTexas 2(2)
species are considered to be endemic and several are in need of conservation due to habitat destruction and
spreading human habitation (Clemants 1995).
Bejaria is typically diagnosed by a combination of morphological features such as its seven-merous
flowers, free petals, capsular fruits, non-appendaged anthers, and viscin threads among the pollen tetrads
(Clemants 1995). Individually, many of these a are shared with other members of Ericaceae
(Stevens 1971). The only known potential phological synaj phy for the genus is tetracytic stomata
(Stevens 197 1).
An unusual characteristic of Bejaria is the presence of tubular corollas despite the fact that all species have
separate petals (Clemants 1995). There are four iM S Corolla ans among is 15 species of Bejaria
(Clemants 1995). The majority of species h llas (B. aestuans, B. cubensis,
B. imthurnii, B. neblinensis, B. racemosa, B. sema B. x Gan 1995). Several other species
exhibit tubular corollas (B. ledifolia, B. mathewsii, B. resinosa, B. sprucei and B. zamorae; Clemants 1995). Two
species (B. nana and B. tachirensis) have small globose corollas (Clemants 1995). Finally, B. infundibula has
a funnel-shaped corolla (Clemants 1995).
The variation seen in coroila shape within Bejaria may reflect different pollination modes and serve
as reproductive isolating mechanisms for the separate species, but very little is actually known about the
pollinators of the genus (Clemants 1995). For example, B. resinosa predominantly self-pollinates and only
occasionally outcrosses (Kraemer 2001). When a biotic pollinator is involved, however, the long, tubular
corollas of B. resinosa were thought to be pollinated by long-billed hummingbirds (Kraemer 2001). In a
pollination study by Kraemer (2001), B. resinosa was visited more often by bumble bees (Bombus funebris).
Bumble bees may pollinate B. resinosa more Pis d than hummingbirds, but the birds may play an oc-
casional, but significant role in long persal of pollen for the species (Kraemer 2001). In the study,
short-billed hummingbirds were common but visited the flowers rarely, most likely due to the inaccessibility
of nectar in the long tube (Kraemer 2001). Nectar robbers (flowerpiercers; Diglossa humeralis) were also a
common visitor to these flowers but did not assist in pollination (Melampy 1987; Kraemer 2001). Out of
the 15 species of hummingbirds at the site, only two were long-billed and both occurred infrequently in
the area (Kraemer 2001). These data appear to suggest that hummingbird pollination is indeed rare for the
long-tubed B. resinosa (Kraemer 2001).
Corollas that are spreading or campanulate are also visited by bees (Clemants 1995). For instance, B.
racemosa (spreading or rotate corolla shape), from the southeastern United States, was visited by the com-
mon honey bee Apis melifera (Clemants 1995). Bejaria aestuans, with a campanulate corolla, was visited by
bees (Clemants 1995). No documentation for the successful pollination of these species by the visitors is
available. Pollinators for the infundibular or globose corolla types are not known.
Ethnobotanically, species within the genus have been used on both continents independently as "fly
paper" due to the thick resin secreted from glandular hairs on both the flowers and leaves (Clemants 1995).
It has been shown that the resin on B. racemosa flowers is as strong as commercially produced fly adhesive
(Eisner & Aneshansley 1983). Species of Bejaria have also been used for woodworking, medicinal purposes,
and until late in the 19% century as a popular ornamental (Clemants 1995).
There has never been a published cladistic analysis of Bejaria. The goals of this study were to address:
(1) the monophyly of Bejarieae and Bejaria using molecular data (2.) the relationships within the genus;
particularly the relationship of B. racemosa to all other sampled species.
MATERIALS AND METHODS
Taxon sampling
Ten species (of the 15 recognized by Clemants 1995) of Bejaria were included in the molecular study. Bejaria
cubensis, B. nana, B. steyermarkii and B. tachirensis excluded because the chosen gene regions could not
be successfully amplified from available material. Bejaria neblinensis is known only from the type locality
(Cerro de la Neblina, Venezuela) and the very few specimens of this species were not available for loan or
for DNA extraction.
Bush and Kron, Phyl g f Bej ia (Eri ) 1195
For all analyses, 13 species representing other clades within Ericoideae were included (Table 1). In the
chloroplast anal (ndhF, matK and combined chloroplast analyses) ten species of Bejaria were included in
the ingroup. ior all analyses containing the nuclear waxy data set (waxy and total combined data analyses),
B. infundibula, B. mathewsii and B. sprucei were excluded from the ingroup due to large amounts of missing
data. Representative Ericoideae taxa were chosen based on previous work within the Ericaceae (Kron et al.
2002). All analyses were rooted with Cassiope mertensiana. Cassiope D. Don (Cassiopoideae) was found in
previous analyses to be sister to the Ericoideae (Kron et al. 2002). Voucher information for the molecular
study is presented in Table 1.
DNA extraction, amplification, and sequencing
Plant material originally dried for flavonoid analysis in 1983 was provided by Steven Clemants (Brooklyn
Botanic Gardens). These leaves were air-dried and no other means of preservation was used (i.e., silica gel,
refrigeration; S. Clemants, pers. comm.). DNA was extracted from these -20 year old leaves and other ma-
terial using an Epicentre Leaf Extraction kit (Epicentre Technologies, Madison, Wisconsin, U.S.A.) or by
using the CTAB protocol of Doyle and Doyle (1987). Less degraded DNA was obtained from the "air"-dried
leaves compared to similarly aged herbarium sheets. All extractions were cleaned with a Qiagen cleanup kit
mes hor di and Assay Technologies, Valencia, California, U.S.A).
loroplast genes, ndhF and matK, were amplified using standard polymerase chain reaction (PCR)
protocols and previously published primers (Olmstead et al. 1993; Steel & Vilgalys 1994). The ndhF gene
encodes a subunit of the nicotinamide dehydrogenase complex and is ca. 2223 base pairs long (Olmstead
& Sweere ids Alverson et al. 199 The e ob iE complex functions in direct hydrogen transfer
between th t 1 1) ide ring (Cook & Cleland 1981). The matK gene encodes a ribosomal
maturase and it is located in the intron of oak a gene that encodes the transfer RNA for lysine Johnson
& Soltis 1994). The matK gene is approximately 1500 base pairs long Johnson & Soltis 1994). The nuclear
gene included in this study is waxy, which encodes the granule-bound starch synthase (GBSSI; Miller et
al. 1999). This gene has 13 translated exons and is approximately 4500 base pairs long (Peralta & Spooner
2001). The gene functions in the synthesis of amylose (Mason-Gamer et al. 1998). For this study, the region
between exons 9 and 11 was amplified for a total of 628 base pairs (Miller et al. 1999; Peralta & Spooner
2001). Bejaria aestuans, B. imthurnii, B. racemosa and B. resinosa were cloned for the waxy gene and because no
multiple copies were found the remainder of the species were not cloned. All PCR products were gel isolated
and cleaned using Qiagen gel isolation kits (Qiagen Sample and Assay Technologies, Valencia, California,
U.S.A). Sequencing was performed on an ABI 3700 machine located at the DNA sequencing facility of Wake
Forest University. All sequences were edited using Sequencher 3.1.1 (Gene Codes Corporation, Ann Arbor,
MD and were aligned manually. GenBank accession numbers are listed in Table 1.
Phylogenetic analyses
Parsimony analyses were performed in PAUP* 4.0b2 (Swofford 1999). Characters were unordered and gaps
within the molecular data were treated as missing data. All characters were equally-weighted and only
those that were parsimony informative were included in the analyses. For ndhF, only the 5’ end of the gene
was used due to extensive missing data in several taxa in the 3’ end. The total number of base pairs for the
ndhF 5 gene region is 1318. The three gene regions (ndhF, matK, and waxy) were analyzed separately using
the heuristic search option with 1000 replicates, TBR branch swapping, and random step-wise addition.
Bootstrap analyses were used to estimate clade support and bootstrap scores (bt) were recorded (1000
replicates of TBR branch swapping with 100 replicates in the heuristic search; Felsenstein 1985). Conflict
between trees was assessed by inspecting the results from different analyses for medium to high bootstrap
scores (greater than 80%) that supported different relationships between the taxa. There was no strongly
supported conflict between the two chloroplast genes and the waxy gene region so all data were combined
for a total evidence analysis.
For the Bayesian analyses, each individual gene set was run in ModelTest 3.7 (Posada & Crandall
1998) to determine the best substitution type (Nst) and rate distribution model (rates) for each gene region.
1196 J lof the Botanical R h Institute of Texas 2(2)
Taste 1. Bejari | other species in the Tribe Bejari pled, sources of material, vouchers, and GenBank
Taxon Voucher ndhF matK waxy
Bejaria aestuans Mutis ex. L. Luteyn 14175 (NY) DQ002362 DQ002346 DQ000589
Bejaria imthurnii N.E. Br. Clemants & Delascio 2453b (NY) A DQ002363 DQ002347 DQ000590
Bejaria infundibula Clemants Clemants €: Fernandez 2007c (NY) DQ002364 DQ002348 N/A
Bejaria ledifolia Bonpl. Clemants 2483b (NY) DQ002365 DQ002349 DQ000592
Bejaria mathewsii Fielding Clemants 2046 (NY) DQ002366 DQ002350
&G
Bejaria racemosa Vent. Kron 2070 (NCU) DQ002367 DQ002351 DQ000594
Bejaria resinosa Mutis ex. L. Luteyn 14133 (NY) DQ002368 DQ002352 DQ000595
Bejaria sprucei Meisn. Clemants & Delascio 2479 (NY) DQ002369 DQ002353 N/A
Bejaria subsessilis Benth. Clemants 2278a (NY) DQ002370 DQ002354 DQ000596
Bejaria zamorae Clemants Bush 212 (LOJA) DQ002371 DQ002355 DQ000597
Bryanthus gmelinii D. Don Stevens DNA 192 (WFU) EU669881 AF440413 EU669884
Cassiope mertensiana (Bong.) s.n. (AA 75-83) DQ002372 U61346 DQ000598
G. Don
Ceratiola ericoides (Michx.) Kron 2069 (WFU) DQ002373 U61334 DQ000599
Elliottia bracteata (Maxim.) s.n. (RBGK 1979-5019) DQ002374 U61339 DQ000600
Hook. f.
Empetrum atropurpureum s.n. (RBGK 1988-433) DQ002375 U61355 DQ000601
Fernald 8: Wiegand
Kalmia angustifolia L. Kron DNA156 (WFU) DQ002376 DQ002356 DQ000602
Kalmiopsis leachiana Kron DNA177 (WFU) DQ002377 U61323 DQ000603
(Henderson) Rehder
Ledothamnus guyanensis Meisn. Picon & Williams 2910 (NY) EU669882 AF440419 FU669885
Phyllodoce caerulea Bab. s.n. (RBGK 1940-1013) DQ002378 DQ002357 DQ000604
Phyllodoce empetriformis D.Don s.n. (RBGK 1978-2120) DQ002379 DQ002358 DQ000605
Rhododendron grande Wight s.n. (RBGK 1969-8606) DQ002383 DQ002360 EU669886
Rhododendron hippophaeoides s.n. (RBGK 1932-1022) EU669883 U61353 EU669887
Balf.f. & W.W. Sm.
Rhodothamnus chamaedistus Rchb. s.n. (RBGK 1989-459) DQ002381 U61321 DQ000607
Three separate analyses were performed in MrBayes v.3.1.2 (Ronquist & Huelsenbeck 2003): waxy, com-
bined chloroplast and total combined data. The individual ndhF and matK analyses were not performed
using Bayesian because the results from each were predominantly unresolved in the parsimony analysis
and the combined chloroplast tree represents both data sets. For the multiple gene analyses, the genes were
partitioned in MrBayes and the appropriate models were applied for each gene region. Each analysis ran
for 1,000,000 generations and was sampled every 100 generations. The burn-in period was the first 2,500
sampled generations. The model used for each gene region is as follows: ndhF: HKY+G; matK: TVM+G and
waxy: HKY+G. Posterior probability values (,P) were calculated in MrBayes.
RESULTS
Analysis of chloroplast sequences
The aligned ndhF data set for 23 taxa was 1318 base pairs long. For this parsimony analysis, 1719 most par-
simonious trees were found (L = 531; C.I. = 0.59; R.I. = 0.72; data not shown). There were 217 informative
characters (16.5 96). The aligned matK data was comprised of 1541 characters and included the same taxa
as in the ndhF analysis. In the matK analysis, 198 most parsimonious trees were recovered (L = 470; CI. =
0.63; R.L. = 0.78; data not shown) and 216 characters were informative (14.0 96). Results of the separate ndhF
and matK parsimony analyses were similar in support and topology to the combined chloroplast analysis so
only the results of the latter will be discussed.
The combined chloroplast parsimony analysis resulted in 431 informative characters (2859 included
Bush and Kron, Phyl g f Bejaria (Eri ) 1197
td y
base pairs) and 9 most parsimonious trees (L = 1012; C.I. = 0.60; R.I. = 0.74). Bejaria is found to be mono-
phyletic in this analysis (100% bt; Fig. 1). Bejaria infundibula is sister to B. mathewsii (91% bt) and this clade
is in a polytomy with B. imthurnii and B. subsessilis (7396 bt; Fig. 1). The four taxa in this clade are placed in
a trichotomy with Bejaria aestuans and B. zamorae (6296 bt; Fig. 1). Sister to this clade is B. resinosa (10096 bt),
followed by B. sprucei (6596 bt) and B. ledifolia (5096 bt; Fig. 1). Finally, B. racemosa is sister to all other Bejaria
members sampled (Fig. 1). Outgroup relationships based on this analysis show Clade A (Phyllodoceae, Kron
et al. 2002) as sister to Bejaria, but without support (Fig. 1). Clade B (Empetreae + Rhodoreae, Kron et al.
2002) is placed sister to Clade A + Bejaria and Clade C is sister to the remaining sampled taxa (Fig. 1).
A monophyletic Bejaria was also recovered in the Bayesian combined chloroplast analysis (10096 ,P;
Fig. 2). In this phylogeny, B. imthurnii and B. subsessilis are sister to each other (5296 ,P) as are B. infundibula
and B. mathewsii (100% ,P) and the two clades are sister to each other (97% „P; Fig. 2). Bejaria aestuans and
B. zamorae are in a polytomy with the previous clade (88% ,P; Fig. 2). Sister to this larger clade is B. resinosa
(100% „P; Fig. 2). Bejaria ledifolia and B. sprucei are unresolved with respect to this clade (100% „P; Fig. 2).
Finally, B. racemosa is sister to all other Bejaria sampled (100% „P; Fig. 2). The relationship of Clade A to
Bejaria is the same in this analysis as in the parsimony results (Figs. 1, 2). However in this analysis Clade B
is sister to Clade C, although with low posterior probability (Fig. 2).
Analysis of waxy (nuclear) sequences
The aligned portion of waxy for 20 included taxa was 628 bases long (region between exons 9-11). The
parsimony analysis resulted in 450 trees with a length of 175 (C.I. = 0.71; R.I. = 0.83). There were 93 infor-
mative characters in this analysis (14.896). This analysis also supported a monophyletic Bejaria (10096 bt,
Fig. 3). It places B. aestuans, B. imthurnii, B. ledifolia, B. resinosa, B. subsessilis, and B. zamorae in a polytomy
(9496 bt) with B. racemosa sister to this clade (10096 bt; Fig. 3).
In the Bayesian analysis of the waxy data, Bejaria is monophyletic (10096 ,P; Fig. 4) but there is very
little resolution within the genus. Bejaria racemosa is sister to all other Bejaria members, which are in a
polytomy (99% „P; Fig. 4). Ledothamnus guyanensis is sister to Bejaria (89% „P; Fig. 4). Although placement
of the outgroup taxa varies between parsimony and Bayesian analyses neither of these results are strongly
supported.
Analysis of combined nuclear and chloroplast sequences:
The combined data set was 3487 aligned bases long. There were 513 informative characters (14.7%). The
parsimony analysis produced 6 trees 1142 steps long (C.I. = 0.62; R.I. = 0.73). In this combined analysis,
Bejaria is monophyletic with 10096 bootstrap support (Fig. 5). Bejaria imthurnii and B. subsessilis are sister
clades (9596 bt) inside a polytomy including B. aestuans and B. zamorae (8996 bt; Fig. 5). Sister to this poly-
tomy is B. resinosa (10096 bt; Fig. 5) and sister to this larger group is B. ledifolia (8596 bt; Fig. 5). Finally, B.
racemosa is sister to the remaining taxa sampled (Fig. 5). Among the outgroup taxa Clades B and C are placed
as sister with very low support (5696 bt) and sister to this clade is Clade A (Fig. 5).
In the Bayesian analysis of all combined data, Bejaria is monophyletic (10096 ,P; Fig. 6). The clade B.
imthurnii + B. subsessilis (100% ,P) is in a polytomy with B. aestuans and B. zamorae (100% ,P; Fig. 6). Sister
to this clade is B. resinosa (10096 ,P) followed by B. ledifolia (100% „P; Fig. 6). Finally, B. racemosa is sister to
all other Bejaria members sampled (Fig. 6). Similar to the parsimony analysis Clades B and C are placed as
more closely related to each other than to Clade A (Figs. 5, 6). However, the Bayesian analysis places Clade
A sister to Bejaria (Fig. 6).
DISCUSSION
Both parsimony and Bayesian total evidence analyses show Bejaria as monophyletic and B. racemosa as
sister to the remaining Bejaria sampled. Deeper nodes of the tree prevent conclusions about the closest rela-
tives to Bejaria as can be seen by the difference in placement of Clade A as sister to Bejaria in the Bayesian
results (Fig. 6) or as (Clade A (Clade B + Clade C )) in the parsimony results (Fig. 5). Clade C contains two
members of the Bejarieae as recognized by Kron et al., (2002). In their study they circumscribe Bejarieae
1198 J | of the Botanical R h Institute of Texas 2(2)
Bejaria aestuans
62 bejaria zamorae
88 Bejaria imthurnii
Bejaria subsessilis
100 97
91 Bejaria infundibula
*65 100 Bejaría mathewsii
50 Bejaria resinosa
100 ; Bejaria sprucei
Bejaria ledifolia
Bejaria racemosa
Kalmia angustifolia
Elliottia bracteata
Kalmiopsis leachiana
Phyllodoce caerulea
Phyllodoce empetriformis
Rhodothamnus chamaedistus
100 Ceratiola ericoides
14 | L- Empetrum atropurpureum
100 100 Rhododendron grande
Rhododendron hippophaeoides
97 m Bryanthus gmelinii
100 L— Ledothamnus guyanensis
Cassiope mertensiana
Fic 1 Strict £0 ; : (l 1012, C1— 0.60, RI 0.74) htainad in +h lysi Fs) hina il pl (5' ILE
VIZ, Xl — V.UU, Tu
r
mati. Vil + L L 7 L L Pi L L "T tet I 4 e ERO.
: A : D F F
see text). A sinal
Y
analyses. Finally
E ?
ID S oM
| Bay i lysi (I ior | l bility | ). Clades A, B and C
as containing Bejaria, Bryanthus and Ledothamnus and map a single (homoplasious) synapomorphy for the
tribe, that of separate petals. This character also occurs in other genera within the Ericaceae (i.e., Elliot-
tia, Ceratiola, Corema, Chimaphila, Orthilia and Pyrola; Kron et al. 2002) some of which are represented as
outgroups in this study.
Bejarieae is not monophyletic in any analysis in this study although the relationships outside of the
terminal groups are poorly supported. Of the Bejarieae, only Ledothamnus and Bryanthus are strongly sup-
ported as closely related in both the parsimony and Bayesian results of the total evidence and chloroplast
analyses (Figs. 1, 2, 5, 6). This clade has several potential morphological synapomorphies such as ericoid
leaves, adaxial calyx stomata, a style articulated with the ovary and the lack of endothecium (Kron et al.
Bush and Kron, Phylogeny of Bejaria (Ericaceae)
Bejaria aestuans
Bejaria zamorae
*52 Bejaria imthurnii
Bejaria subsessilis
Bejaria infundibula
Bejaria mathewsii
Bejaria resinosa
100
100 Bejaria ledifolia
Bejaria sprucei
Bejaria racemosa
Elliottia bracteata
Kalmia angustifolia
Kalmiopsis leachiana
Phyllodoce caerulea
Phyllodoce empetriformis
Ceratiola ericoides
Empetrum atropurpureum
Rhododendron grande
Bryanthus gmelinii
Ledothamnus guyanensis
Cassiope mertensiana
Rhodothamnus chamaedistus
Rhododendron hippophaeoides
1199
t numbers
Fic. 2 Si I £ E JD H loete aftha Dn ALE J "A
Ed F ?
1 tha k
" Eno. J L hal L3 L "dli PE habiles
An asterisk next
4
Finallv.
J
Bayesi lysis (Į ior probability sl ). Clades A, B and C
2002). Parsimony and Bayesian analyses of waxy data do show Ledothamnus and Bryanthus in different posi-
tions within the trees (Figs. 3, 4), but these are not well supported either by bootstrap or posterior probability
measures.
In his monograph of Bejaria, Clemants (1984) performed a morphological cladistic analysis of all 15
recognized species of Bejaria using 16 characters, eight of which were related to floral morphology. Variation
within Rhododendroideae (Kron 1997) was so significant that its use as an outgroup required designating
B. racemosa as a dean d obi id i urea = ah in the study (Clemants 1984). Only four characters
did not vary in th y outgroup (
; Clemants 1984). The cladogram produced from
1200 J | of the Botanical R h Institute of Texas 2(2)
Bejaria aestuans
Bejaria imthurnii
94 Bejaría subsessilis
99 Bejaria zamorae
100 Bejaria resinosa
Bejaria ledifolia
Bejaria racemosa
Flliottia bracteata
Kalmia angustifolia
100 88 Phyllodoce caerulea A
77 |99 Kalmiopsis leachiana
100| 73
Phyllodoce empetriformis
Rhodothamnus chamaedistus
100 Ceratiola ericoides
64 Empetrum atropurpureum
Bryanthus gmelinii B
2 100 Rhododendron grande
Rhododendron hippophaeoides
Ledothamnus guyanensis
Cassiope mertensiana
Fic, 3 Strict £13920 tr H . (L 193, Cl 20.74, RI 0.85) has J: L ly H Esl J Datania tE "nm la, B.
mathewsii and B. J
present k L hak L 4l tet I r EAD J L hal sha h E els A hake! A
F F F F F
. I L
E]
27
- Ls
" -— has kal hal
Finally, a singie number Delow
D H VL “al J H I.
Fr l x» I lei :
i Bayesi lys (posterior prot bility } ). Clades A, B and C
the four characters placed B. racemosa as sister to all other Bejaria species (Clemants 1984). In the cladogram
based on these four characters, a mucilaginous epidermis was a synapomorphy for all species in Central,
South America and Cuba (Clemants 1984). Bejaria racemosa had the autapomorphy of stomata on both leaf
surfaces (Clemants 1984). Flower shape was divided into three states: tube-shaped, cup-shaped and funnel-
shaped (Clemants 1984). Bejaria infundibula alone possessed a funnel-shaped corolla, B. nana and B. tachirensis
possessed cup-shaped corollas and the presence of tube-shaped corollas was a synapomorphy for the clade
containing B. ledifolia, B. mathewsii, B. resinosa, B. sprucei, and B. zamorae (Clemants 1984). Bejaria racemosa
and B. nana share the presence of an apical bracteole (compared to basal or medial) but this character has
possibly evolved twice within Bejaria (Clemants 1984). The analyses in the present study represent a much
smaller sample of Bejaria than Clemants (1984) due to problems with DNA extractions and PCR amplifica-
Bush and Kron, PI yl g TUNE
94.
99
100
*89
100
64
84
80
100
*5]
100 88
77 | 99
100 13
Bejaria aestuans
Bejaria imthurnii
Bejaria ledifolia
Bejaría resinosa
Bejaria subsessilis
Bejaria zamorae
Bejaria racemosa
Ledothamnus guyanensis
Ceratiola ericoides
Empetrum atropurpureum
Bryanthus Mii CN
Rhododendron m
Rhododendron hippophaeoides
Elliottia bracteata
Kalmia angustifolia
Kalmiopsis leachiana
Phyllodoce caerulea
Phyllodoce empetriformis
Rhodothamnus chamaedistus
Cassiope mertensiana
1201
L
dibula, B. mathewsii and B. j
probabilities. An asteri i tt |
L Lal
) Clades A, B and C
tion of old material as well as obtaining rare taxa. However our outgroup taxa include many members of
J
: 1
pl i 219 Uni y
compared to those of Clemants' (1984).
The revision of ue m e ERE (1995) euterca Ho previous revisions of the genus due to its focus
on corolla hape asa
f
10
(with the exception of Empetreae) so our results are useful when
However, the total evidence analyses in this
study (Figs. 5, 6) indicate tab a tubular rel such as iet found in B. ledifolia, B. resinosa, and B. zamorae
has likely evolved as least twice within Bejaria. The placement of clades with taxa that possess bell-shaped
or spreading corollas (B. aestuans, B. imthurnii, and B. subsessilis) intercalated within clades that represent
1202
100
95
100
dio
—
100
100
| | af aL Dat * 1n
Bejaria aestuans
Bejaria zamorae
Bejaria imthurnii
Bejaria subsessilis
Bejaria resinosa
Bejaria ledifolia
Bejaría racemosa
Bryanthus gmelinii
Ledothamnus guyanensis
Ceratiola ericoides
Empetrum atropurpureum
Rhododendron grande
Rhododendron hippophaeoides
Kalmia angustifolía
Elliottia bracteata
Kalmiopsis leachiana
Phyllodoce caerulea
Phyllodoce empetriformis
Rhodothamnus chamaedistus
Cassiope mertensiana
f Texas 2(2)
(L = 1142, Cl = 0.62, RI = 0.73) CANIT
+ L k h +
la, B. mathewsii and B. p
L Ļ
E DIQUCICA
sa hanth
Y
Finally.
J
5 OE
) Clades A, B and C
are labeled (see text).
taxa with tubular corollas indicates that corolla shape
within the genus.
J
e
may not be phylogenetically informative
Phylogenetic relationships among the species of Bejaria sampled in this study are identical in both the
Bayesian and parsimony total data analyses and are well supported. In our analyses a novel, strongly sup-
ported relationship between B. imthurnii and B. subsessilis was found (Figs. 5, 6), although Clemants' (1984)
study does not support this relationship. Instead, his results indicate that B. subsessilis is sister to B. cubensis
Bush and Kron, Phvl f Beiaria (Eri
Ll q - + *
mmm Bejaria aestuans
89 Bejaria zamorae
100 95 Bejaria imthurnii
100 Bejaria subsessilis
Bejaria resinosa
Bejaria ledifolia
Bejaria racemosa
Elliottia bracteata
Kalmiopsis leachiana
Phyllodoce caerulea
Phyllodoce empetriformis
Rhodothamnus chamaedistus
Kalmia angustifolia
Ceratiola ericoides
Empetrum atropurpureum
- Rhododendron grande
Rhododendron hippophaeoides
= Bryanthus gmelinii
100 — Ledothamnus guyanensis
Cassiope mertensiana
1203
AD
,
t, numbers above
al 4- 4- Mm
2
ondieatac that +h J I L FEAT L H ID H I Linally
Pr F F 4 7 , 4
ID A Y Lh
ly ; ( tari n bility f ). Clades A, B and C
due to the shared presence of a long floral bract (data not shown). Bejaria imthurnii and B. subsessilis do not
share geographic ranges: B. imthurnii is endemic to the summits of Cerro Roraima and Cerro Kukenán on
the border of Venezuela, Brazil and Guyana while B. subsessilis is endemic to Loja, Ecuador (Clemants 1995).
Bejaria imthurnii shares some diagnostic morphological characters with B. steyermarkii and B. neblinensis
(Clemants 1995). The three taxa share small, short-petioled leaves and campanulate corollas (Clemants
1995). Clemants (1995) noted that B. subsessilis possessed some characters that were intermediate between
- a Laféha Botanical D h Institute of Texas 2(2)
B. aestuans and B. resinosa (e.g., rachis similar to B. aestuans, subsessile leaves with obtuse leaf bases like B.
resinosa) suggesting that B. subsessilis possibly arose as a hybrid between B. aestuans and B. resinosa. In addi-
tion, some populations of B. subsessilis have distorted tubular corollas and two of the four pollen grains in
the tetrads are aborted (Clemants 1995).
The relationships of Bejaria aestuans and B. zamorae to the B. imthurnii + B. subsessilis clade (89% bt,
100% ,P respectively; Figs. 5, 6) are not resolved. While these species share several morphological traits,
a or not they inc relationship remains to be tested. Bejaria aestuans is the most geographically
widespread and phol li iable species in the genus: it is found from Mexico to Bolivia (Clemants
1995). cin zamora, conversely is aaa to the Rio Zamora valley in Ecuador (Clemants 1995). Bejaria
ters with both B. aestuans and B. sprucei (not sampled
in the total data analyses; Clemants 1995), Bejaria zamorae has distinct thin and long pedicels and a strongly
tubular corolla like B. sprucei. However, B. zamorae also exhibits slightly exserted stamens and a 7-merous
corolla similar to B. aestuans (Clemants 1995). Bejaria sprucei exhibits a disjunct geographic distribution
both north and south of the Ecuadorian volcanic region: one region includes the Guyana Highlands area
and the mountains near Táchira and Anzoátegui, Venezuela and the other geographic region for the species
is in northeastern Peru near San Martin (Clemants 1995).
Bejaria resinosa is sister to the clade containing B. aestuans, B. zamorae, B. imthurnii and B. subsessilis
(Figs. 5, 6). Bejaria resinosa shares tubular flowers and long calyx lobes with B. ledifolia and B. mathewsii (not
sampled in total evidence analyses) and these are considered indicators of relationship by Clemants (1995).
In general, our results agree that B. ledifolia and B. resinosa are S related cas 5, 6) but they are not
placed as sister in our analyses, indicating that such cl te leaf bases, short petioles,
and long calyx lobes may not be phylogenetically informative.
This study indicates that additional approaches to resolve these closely related species are desired. A
more complete sampling of the species within the genus, especially the rare and endemic taxa is important.
Additional morphological cladistic work that builds on Clemants' (1984, 1995) studies are likely to improve
our understanding of relationships within the genus and hyper-variable chloroplast or nuclear markers may
be able to differentiate among these taxa more clearly. Population-level studies of widespread taxa such as B.
aestuans and B. mathewsii which include samples from the geographic and morphological extremes of each
species could clarify the potentially complicated relationships within this genus.
ACKNOWLEDGMENTS
We thank Steven E. Clemants (t, 2 Nov 2008) for his guidance and review of this paper as well as the use
of his dried Bejaria materials. We also thank the anonymous reviewers for their helpful suggestions. We
would like to acknowledge the financial support from Wake Forest University and the National Science
Foundation (DEB 0433857).
REFERENCES
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CiEMANTS, S.E. 1984. Revision of the genus Befaria (Ericaceae). Unpublished Ph.D. thesis. New York Botanical
Garden.
CLEMANTS, S.E. 1995. Ericaceae subfamily Rhododendroideae. 6. Bejaria. In: J.L. Luteyn, ed. Ericaceae, Part II, The
superior-ovaried genera. Flora Neotropica Monograph 66:54-106. New York Botanical Garden, New York.
Cook, PF. ano W.W. Ci&L AND. 1981. Mechanistic deductions from isotope effects in multireactant enzyme mecha-
nisms. Biochemistry 20:1 790-1796.
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DovLe, J. AND J. Done. 1987. A rapid DNA isolation procedure for small quantities of fresh leaf tissue. Phytochem.
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EISNER, T. AND DJ. ANESHANSLEY. 1983. Adhesive strength of the insect-trapping glue of a plant (Befaria racemosa).
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FEDTSCHENKO, B.A. AND N.A. BASILEVSKAJA. 1928. Revision of the genus Bejaria Mutis. Bot. Gaz. (Crawfordsville)
85:299-322.
FELSENSTEIN, J. 1985. Confidence limits on phylogenies: an approach using the bootstrap. Evolution 39:783-791.
JOHNSON, L.A. AND D.E. Souris. 1994. matK DNA sequences and phylogenetic reconstruction in Saxifragaceae s. str
Syst. Bot. 19:143-156.
Kraemer, M. 2001. On the pollination of Bejaria resinosa Mutis ex Linne f. (Ericaceae), an ornithophilous Andean
paramo shrub. Flora 196:59-62.
Kron, K.A. 1997. Phylogenetic relationships of Rhododendroideae (Ericaceae). Amer. J. Bot. 84:973-980.
Kron, K.A., W.S. Jupp, PF. Stevens, D.M. Crayn, A.A. ANDERBERG, PA. GADEK, C.J. QUINN, AND J.L. LuTEYN. 2002. Phylogenetic
classification of Ericaceae: molecular and morphological evidence. Bot. Rev. 68:335-423.
MasgerLy, DJ. 1997. The plant-book: a portable dictionary of the vascular plants. Second Edition. Cambridge
University Press. Cambridge, UK.
MANSFELD, R. AND H. SLEUMER. 1935. Revision der Gattung Befaria Mutis. Notizbl. Bot. Gart. Berlin-Dahl-
em12:235-276.
MAson-GAmer, R.J, C.F. We, AND E.A. KeLLOGG. 1998. Granule-bound starch synthase: structure, function, and phy-
logenetic utility. Molec. Biol. Evol. 15:1658-1673.
Metampy, M.N. 1987. Flowering phenology, pollen flow and fruit production in the andean shrub Befaria resinosa.
Oecologia 73:293-300.
Mier, R.E., M.D. RAUsHER, AND P.S. Manos. 1999, Phylogenetic systematics of Ipomoea (Convolvulaceae) based on
ITS and waxy sequences. Syst. Bot. 24:209-227.
OLMSTEAD, R.G. AND J.A. Sweere. 1994. Combining data in phylogenetic systematics: an empirical approach using
three molecular data sets in the Solanaceae. Syst. Biol. 43:467-481.
OLMSTEAD, R.G., J.A. Sweere, AND KH. Wotre. 1993. Ninety extra nucleotides in the ndhF gene of tobacco chloroplast
DNA: A summary of revisions to the 1986 genome sequence. Pl. Molec. Biol. 22:1191-1193.
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STEELE, KP. AND R. VitaaLvs. 1994. Phylogenetic analyses of Polemoniaceae using nucleotide sequences of the
plastid gene matK. Syst. Bot. 19:126-142.
STEVENS, PF. 2007. Angiosperm Phylogeny Website. Version 8, June 2007.
STEVENS, PF. 1971. A classification of the Ericaceae: subfamilies and tribes. Bot. J. Linnaean Soc. 64:1—53.
SWOFFORD, D.L. 1999. PAUP* Phylogenetic analysis using parsimony (*and other methods). Version 4.01. Sunder-
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1206 J lof the Botanical R h Institute of Texas 2(2)
BOOK REVIEW
SEAN CARRINGTON. 2007. Wild Plants of Barbados. (ISBN-10: 1405094079, ISBN-13: 978-1405094078, pbk.).
MacMillan Caribbean Publishers, dan Towns Road, Oxford, OX4 3PP, United Kingdom. (Orders:
www.macmillan-caribbean.com, illan.com, +44 (0)1865-405843 fax). $10.69, 248 pp.,
color photographs, 8 1/4" x 5 1/2".
This fully revised and pa edition now ao Sa ae species previously unrecorded for bride This book aims to en-
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, over half with cee eee E nn ea s easy-to-follow Se iss accurate identification.
uel are dd d Family, identified the local e. Details are provided about the
tion, its habitat in Barbados l ink i lating to i
Bousticu] terms are explained and TER with labeled li glossary, both of which are useful for the begin-
ner. A O and i n is oa me use in a ae to ae Hay the v ple Ea br oie pin ue the basics of plant
y g yp page bibliography, index, and
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1993 had g520r pag I I
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I g I tion. This makes the current edition much easier to use.
Highly dedos public libraries ith an i in the flora of the Caribb d college and universities with an acti
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ds or Ls biology progra
os Carrington is Professor of Plant easel at the ao of the West Indies, Barbados, carrying out research on the
biochemistry f Caribbean flora. He is the author of Wild Plants of the Eastern Caribbean and lead
author of A-Z of Barbados Heide — Gary L. Jennings Botanical Research Institute of Texas, 500 East 4th Street, Fort Worth, Texas 76102-4025,
U.S.A.
J. Bot. Res. Inst. Texas 2(2): 1206. 2008
UNDERSTANDING OLIGACTIS (ASTERACEAE: LIABEAE):
THE TRUE IDENTITY OF O. SESSILIFLORA AND O. VOLUBILIS
Diego G. Gutiérrez
División Plantas Vasculares
Paseo del Bosque s.n., La Plata, Buenos Aires, B1900FWA, ARGENTINA
digutier@fcnym.unip.edu.ar
ABSTRACT
E (rooted Oligacti ins 13 species grouped 1 bs in hiopsis and Olig " Oligactis has f
species that occur mainly i lombia and fu ] entral America and Venezuela. New synonyms Oligactis garcia-barrigae
and O. valeri are proposed under the species O. volubilis and O. sessiliflora, respectively. The synonyms Liabum meridense, O. biattenuata,
and O. boyacensis are tranferred from O. volubilis to O. sessiliflora. O. volubilis is illustrated for the first time. In addition, a key to the spe-
d
cies of the subgenus Oligactis is provide
Key Wonrs: Asteraceae, Colombia, Compositae, Liabeae, Oligactis sessiliflora, O. volubilis
RESUMEN
El s oe P nar ue 13 especies agrupadas en dos subgéneros: ATE y oe El oa Oligactis
olombia, de | ly Venezuela. Se proponen a
Oligactis garcia- Barcode y O. valeri como sinónimos de las especies O. volubilis y O. sessiliflora j Se trafieren los sinónimos
iii meridense, O. ud y O. oce de O. a a O. sessiliflora. Se ilustra a O. volubilis por primera vez. Además, se provee
F
PALABRAS CLAVE: Asteraceae, Colombia, Compositae, Liabeae, Oligactis sessiliflora, O. volubilis
Oligactis (Kunth) Cass. contains 13 species confined to cloud forests from western Venezuela to northern
Peru and one species only that reaches forests of Panama and Costa Rica (Dillon & Sagástegui Alba 1994;
Funk et al. 1996). Oligactis is distinct among Liabean genera by its scandent or climbing habit, pinnatin-
erved leaves, capitula grouped in axillary or terminal capitulescences, and glandular cypselae. A modern
view of Oligactis was provided by SH M who illustrated the first evolutionary tree for Liabeae.
Phylogenetic analyses based on } d that Oligactis is closely related to Liabum
Adans. (Bremer 1994; Funk et al. 1996; Cuine 2004). on the other hand, in a preliminary phylogenetic
study conducted using sequences of the internal transcribed spacer (ITS) 1 and 2 and the 5.85 gene, Oli-
gactis was sister to and closely related to the natural group formed by Dillandia Funk & H. Rob. and Liabum
(Gutiérrez et al. 2007).
Originally, Oligactis was described by Kunth (1818) as a section of genus Andromachia Humb. & Bonpl.
(currently a synonym of Liabum) with three species: Andromachia nubigena Kunth, A. sessiliflora Kunth, and
A. volubilis Kunth. Years later, Bentham (1873) included the species of Oligactis in Liabum s.l. section Oli-
gactis. During a hundred years, this general point of view of Liabum was followed by different authors (i.e.
Hoffmann 1897; Blake 1927) with only Rydberg (1927) and Cabrera (1954) recognizing separate genera
with the general characters of Liabum.
In 1964, Aristeguieta recorded six species of Liabum s.l. from Venezuela, among them L. sessiliflorum
(Kunth) Less. and L. volubile (Kunth) Less. However, some features indicated by Aristeguieta in L. volubile
did not fit with its original description. Subsequent check-lists followed Aristeguieta (Badillo 1996; Badillo
2001).
In 1974 Robinson and Brettell resurrected Oligactis and created two subgenera: Andromachiopsis and
Oligactis. Also, they transferred several species originally described under the broad concept of Liabum to
J. Bot. Res. Inst. Texas 2(2): 1207 — 1213. 2008
laf 4 D eal D h lnrtitt
1208 J f Texas 2(2)
Oligactis, most of them to subgenus And hiopsis. According to those authors, subgenus Oligactis included
eight species: the three species described by Kunth (1818) under Andromachia section Oligactis, plus O. biat-
tenuata (Rusby) H. Rob. & Brettell, O. boyacensis (Cuatrec.) H. Rob. & Brettell, O. latifolia (Hieron.) H. Rob.
& Brettell, O. mikanioides (S.F. Blake) H. Rob. & Brettell, and O. valeri (Standl.) H. Rob. & Brettell. Then,
Robinson (1983) recognised six species under subgenus Oligactis since he excluded O. nubigena (Kunth) Cass.,
noticed O. biattenuata and O. POE as E and de the last species MUS O. garcia-barrigae
H. Rob. (Robinson 1980). He hypot! 1 the most sf element of the genus
Oligactis due to its axillary or terminal lira: six to 10 florets per capitulum, and papillose anther
apical appendages.
In spite of floristic treatments and check-lists from Costa Rica (Standley 1938), Ecuador (Robinson
1978; Robinson 1999), Peru (Dillon & Hensold 1993), and Venezuela (Aristeguieta 1964; Badillo 2002), a
revision of Oligactis has not been made. While performing a revision of the genus Liabum (Gutiérrez 2004),
species belonging to Oligactis subgenus Oligactis have been studied. The object of this work is to determinate
the real taxonomic identity and synonyms of O. sessiliflora (Kunth) DC. and O. volubilis (Kunth) Cass.
MATERIALS AND METHODS
TI hol land ic studies were based on specimens, photos and/or dipitali m kept at GH,
K, LP, NY, P, SL MO, US, and VEN. The data derived from the sp pr by information
from the literature. For microscopic examination, vegetative lucti
with a clearing process, stained with 296 safranin, and mounted on mune slides. Drawings were made
using an Olympus CH2 microscope with camera-lucida.
were rehydrated, treated
RESULTS AND DISCUSSION
The status of Oligactis volubilis
In the protologue, Kunth (1818) distinguished of Andromachia volubilis (currently O. volubilis) by its scandent
habit, lanceolate-linear leaves, and pedunculate capitula in terminal panicles with approximately six florets
per capitulum, and A. sessiliflora (currently O. sessiliflora) by its shrubby habit, oblong leaves (and ovate, ac-
cording to the original illustration), axillary and terminal panicles, and sessile and densely grouped capitula
with approximately eight florets per capitulum. In spite of several traits in common, both species can be
easily distinguished (Table 1).
Aristeguieta (1964) under the broad concept of Liabum first treated Lial idense Badillo as a synonym
of L. volubile based on the specimens Gehriger 427 (type of L. meridense) and Badillo 613. In his description,
he characterized L. volubile by its lanceolate leaves (2-4 cm wide), pedunculate capitula, and 10-15 florets
per capitulum. However, all the specimens studied by Aristeguieta seem to belong to O. sessiliflora because
none of the specimens analysed had narrowly elliptical or linear leaves, the most conspicuous trait of O.
volubilis. On the other dmi. he pu E e orum by its elliptical to lanceolate leaves (2.5-6 cm
wide), sessile to slightl itula, and ly 10 florets per capitulum. According to this
O
author both O. veluos m O. sessi "us would dud almost the same lamina width and amount of florets
per capitulum.
n 1980, Robinson described O. garcia-barrigae which was distinguished from O. volubilis (and O.
sessiliflora) by its narrow leaves, often closely spaced or clustered on lateral branches, and six florets per
capitulum (three ray and three disc florets). However, my analysis revealed that the type specimens of O.
garcia-barrigae are strikingly similar to the type of Oligactis volubilis (Fig. 1). The specimens of both taxa
share the following CO of oa narrowly a or lineal leaves, densely grouped capitula
six florets per capitulum (commonly three ray and three disc florets).
in axillary panicles, and apr y
Thus, O. garcia-barrigae is proposed as a synonym of O. volubilis.
On the other hand, Robinson (1983) included the Colombian species Oligactis biattenuata and O.
boyacensis described by Rusby and Cuatrecasas respectively, under O. volubilis and maintained the concept
Gutiérrez, Oligactis sessiliflora and 0. volubilis 1209
Tase 1. Characters in Oligactis sessiliflora and O. volubilis.
Character O. sessiliflora (Kunth) DC. O. volubilis (Kunth) Cass.
Leaf arrangement Decussate, not grouped on nodes Dec uped on nodes
Lamina shape Elliptical or ovate, sometimes narrowly een c or ee
Lamina width 2 cm to 6 cm
Lamina margin Subentire or serrulate
Leaf margin intrasection Pianate Revolute
Capitulescence Axillary and terminal paniculiform or Axillary and terminal paniculiform cymes
glomerate racemiform cymes
Involucre Sessile or slightly pedunculate Slightly pedunculate
Ray florets 3to6 3
Disc florets 7to9 3
of Aristeguieta regarding Liabum meridense as synonym of the last species (O. volubilis). However, both
Colombian's species (O. biattenuata and O. boyacensis) and L. meridense fit strickly with O. sessiliflora specially
regarding leaf shape, quantity of florets per capitulum, and ratio between ray and disc florets. Therefore, the
illustration of O. volubilis (Robinson 1983) actually represents O. sessiliflora. According to the findings above
described, L. meridense, O. biattenuata, and O. boyacensis are excluded from O. volubilis.
Oligactis volubilis (Kunth) Cass., in E Cuvier, Dict. sci. nat., ed. 2, 36:17. 1825. ~ 2). Andromachia volubilis
Kunth, in nia Bonpland & Kunth, Nov. aa sp., ed. Folio, 4: 80. 1818. Tree: Probably COLOMBIA. Department and locality
boldt & Bonpland s.n. (HoLotyre: P [digital image LP!]) ies cem below).
Diplostephium volubile (Kunth) Spreng., Syst. veg. 3 543. 1826. Liabum volubile (Kunth) Less., Linnaea 6:704. 1
Cgaetts garcia-barrigae H. Rob. Linn IS 46: a 1980. Syn. nov. Tyre: COLOMBIA. Cundinamarca-Meta: Cordillera Oriental, vertiente
lillera de Cerro Verde, 1900-3200 m, 26-28 Dec 1962, Garcia-Barriga 17689
, Fómeque,
(HoLoTyPE: COL; isotypes GH!, US [digital ee PI).
The Antisana (currently Antizana) mountain (Ecuador) was cited by Kunth (1818) with doubt as the original
locality of Andromachia volubilis. In addition, the holotype kept at P does not have any data of the locality
and there are no additional type materials of this species. Robinson (1999) understood the locality cited by
Kunth was a mistake and excluded Oligactis volubilis from Ecuador. Thus, Colombia is probably the country
where the type was collected by Humboldt and Bonpland, since the specimens assigned to O. volubilis from
Costa Rica (Standley 1938), Panama (Robinson 1999) and Venezuela (Aristeguieta 1966), actually belong
to O. sessiliflora. On the other hand, the synonym O. garcia-barrigae proposed in this work was collected in
central Colombia and its types currently represent the unique materials of O. volubilis with reliable data of
collection area.
The status of Oligactis sessiliflora
The analysis revealed that the type specimens of Liabum meridense, Oligactis biatenuatta, O. boyacensis (ex-
cluded in this work from O. volubilis), and O. valeri match Oligactis sessiliflora. The specimens of those taxa
share the following combination of characters: elliptical, ovate or sometimes narrowly ovate leaves, densely
grouped capitula in axillary and terminal panicles or glomerate racimous cymes, sessile or slightly pedun-
culate capitula, and eight to 15 florets per capitulum. Thus, L. meridense, O. biatenuatta, O. boyacensis, and
O. valeri are proposed as synonyms of O. sessiliflora.
Lm
Oligactis sessiliflora (Kunth) DC., Prodr. 5:96. 1836. Andromachia sessiliflora Kunth, in Humboldt, Bonpland & Kunth, Nov
gen. sp., ed. Folio, 4:80. 1818. Tyre: Probably COLOMBIA. D 1 locality i it in Regno Peruviano? Humbold
& Bonpland s.n. (HoLorvrE: P [photos LP!, SI!]; isotyre: B [bhoto FM 18129, SI!], probably destroyed). Oligactis apodocephala Cass., in
E Cuvier, Dict. sci. nat., ed. 2, 36:17. 1825. Diplostephium sessiliflorum (Kunth) Te Syst. veg. 3:543. 1826. Liabum sessiliflorum
(Kunth) Less., Linnaea 6:703. 1831. Oligactis sessiliflora (Kunth) H. Rob. & Brettell, Phytologia 28:58. 1974.
1210 J I of the Botanical R h Institute of Texas 2(2)
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Gutiérrez, Oligactis sessiliflora and O. volubilis 1211
Fic. 2. Oligactis volubilis. A. Habit. B. Capitulum. C. Involucral bract. D. Ray floret. E. Disc floret. F. Style of disc floret. G. Anther. H. Cypsela and pappus
(García-Barriga 17689, GH).
1212 J lof the Botanical R h Institute of Texas 2(2)
Liabum tovarense VM. Badillo, Bol. Soc. Venez. Ci. Nat. 10:314. 1946. Tyre: VENEZUELA. Aracua: Colonia Tovar, 1800-2000 m, Dec
En esed (noioivee: VEN [digital image LP!]; isorvrE: US [digital i pis p
Rusby, Descr. S. Amer pl.:159. 1920. Syn. nov. Tyre: COLO S Marta, Sierra del Líbano, 5500
ft, 19 Jan 1898/1899, Smith 2013 (HoLorvee: NY [digital image LP!]; tsotyees: K, MO, P [photos LP!, SI!], S, US [digital image LP!]).
Oligactis biattenuata (Rusby) H. Rob. & Brettell, Phytologia 28:57 197
Liabum boyacense Cuatrec., Revista Acad. Colomb. Ci. Exact. 6:61. 1944. a nov. Tre: COLOMBIA. Boyaca: Cordillera Oriental, entre
Moniquirá y Arcabuco, 2150 m, 25 Feb 1940, Pérez Arbeláez & Cuatrecasas 8164 (HOLOTYPE: COL; isotypes NY [digital image LP!]),
US [digital image LP!]). Oligactis boyacensis (Cuatrec.) H. Rob. & Brettell, Phytologia 28:57. 1974
d meridense V.M. Badillo, Bol. Soc. Venez. Ci.. Nat. 10:314. 1946. Syn. nov. Tyre: VENEZUELA. MtnrpA: Tabay, selva de la Isla y El
n, 2500-2700 m, 8 Sep 1930, Gehriger 427 (notorvee: VEN [digital image LP!]; isoryre Ms [digital image LP!]).
me Pa Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 18:1490. 1938. Syn. nov. Type: COSTA RICA. San José: Near Finca La Cima,
above Los Lotes, N of Copey, 2100-2400 m, 21/22 Dec 1925, Standley 42555 (noLoTYeE: F; isotype: US [digital image LP!]). Oligactis
valeri (Standl.) H. Rob. & Brettell, Phytologia 28:58. 1974.
Iconography.—Kunth 1818, tab. 338; Robinson 1983: 40, fig. 9 (sub nom. Oligactis volubilis).
Candolle (1836) recognized under Andromachia the three species of Kunth's section Oligactis. In the
synonymic list he assigned the COHIAIBAHOR Oligactis sessi Ets to dcn However, Cassini never made this
O la (Cassini 1825). Recently, Robinson and
=
combination because he created tł
Brettell (1974) made the combination Oligactis sessiliflora. Never ticles according to The International Code
of Botanical Nomenclature (ICBN), Art. 33.1, 2, and 4, the valid combination was made by Candolle.
The current four species of Oligactis subgenus xii can be distinguished as follows:
. Leaf lamina linear to narrowly elliptical, i ti lum usually 6-flowered O. volubilis
. Leaf lamina elliptical to ovate, margins subentire or serrulate; cabitülum of (8310-15 florets.
2. Leaf adaxial surface hirsute O. latifolia
2. a adaxial surface glabrous
. Sessile or subsessile capitula densely grouped in panicles or glomerate racemes O. sessiliflora
A Pedunculate capitula laxly grouped in panicles O. mikanioides
ACKNOWLEDGMENTS
Thanks are given to Gisela Sancho and the reviewers for helpful comments on this manuscript, and Luis
Hernández Chong for helping with type materials of VEN, and M. Alejandra Migoya for the illustrations.
Also, thanks are given to the herbaria for the use of specimens, information, and digital images of types.
Without such inter-institutional cooperation, this work would be impossible. This work was supported by
the Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET 1999-2004), Argentina.
REFERENCES
Aristecuteta, L. 1964. Liabum. In: T. Lasser, dir. Flora de Venezuela 10:744—751. Instituto Botánico, Dirección de
Recursos Renovables, Ministerios de Agricultura y Cría. Caracas.
Bapo, V.M. 1997. Los géneros de las Compositae (Asteraceae) de Venezuela. Ernstia 6:51—168.
BapiLLo, V.M. 2001. Lista actualizada de las especies de la familia Compuestas (Asteraceae) de Venezuela. Erns-
tia11:147-215.
BentHam, G. 1873. Ordo 88: Senecionidae. In: G. Bentham and J.D. Hooker, Gen. pl. 2:435-453. London.
Blare, S.F. 1927. New South American species of Liabum. J. Wash. Acad. Sci. 17:288-303.
Bremer, K. 1994, Asteraceae. Cladistics and classification. Timber Press, Portland.
CABRERA, AL. 1954. Compuestas sudamericanas nuevas o críticas. Il. Notas Mus. La Plata, Bot. 17:71-80.
CANDOLLE, A.P. 1836. Andromachia and Liabum. Prodr. 5:95—97. Paris (Treuttel & Wurtz).
Cassini, H. 1825. Oligacte. In: F. Cuvier, ed. Dict. sci. nat, ed. 2, 36:16-18. Paris (Reprinted 1975, RM King and H.W.
Dawson, eds. Cassini on Compositae, vol. 2. Oriole Editions. New York).
Ditton, M.O. AND N. HtNsoLD. 1993. Oligactis. In: L. Brako and J. Zarucchi, eds. Catalogue of the flowering plants
and gymnosperms of Peru. Monogr. Syst. Bot. Missouri Bot. Gard. 45:300-301.
DiLLON, M.O. AND A. SAGÁsTEGUI ALBA. 1994. Estudios en la tribu Liabeae (Asteraceae) en Perú: Il. Una nueva especie
de Oligactis precedente del norte del Perú y sur del Ecuador. Arnaldoa 2:25-30.
Gutiérrez, Oligactis sessiliflora and O. volubilis 1213
Funk, V.A., H. Rosinson, AND M. Ditton. 1996. Liabeae: Taxonomy, phylogeny and biogeography. In: DJ.N. Hind and
H.J. Beentje, eds. Compositae: Systematics. Proceedings of the International Compositae Conference, Kew,
1994. Pp. 545-567.
Guniérrez, D.G. 2004. Revisión sistemática y análisis cladístico del género Liabum Adans. (Asteraceae, Liabeae).
Unpublished D. Phil. Thesis. Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata,
Argentina.
Gutierrez, D.G., N. GARCIA-JACAS, AND A. SUSANNA. 2007. Preliminary phylogenetic affinities of Microliabum within
Liabeae (Asteraceae) inferred from ITS sequences. Darwiniana (suplemento) 45:87-88.
Hoffmann, O. 1894. Compositae: Senecioneae-Liabinae. In: A. Engler and K. Prantl, eds. Nat. Pflanzenfam.
4:284—285. Leipzig.
Kunth, C.S. 1818. In: F. W.H.A. von Humboldt, AJ.A. Bonpland, and C.S. Kunth, eds. Nov. gen. sp, ed. Folio, vol. 4.
Maze, Paris. Pp. 1-247 and tabs 301-412.
RoBINSON, H. 1978. Compositae-Liabeae. In: G. Harling and B. Sparre, eds. Flora of Ecuador 8:3-63.
Rosinson, H. 1980. Studies in the Liabeae (Asteraceae). XIV. New species of Liabum and Oligactis. Phytologia
45:99-103.
ROBINSON, H. 1983. A generic review of the tribe Liabeae (Asteraceae). Smithsonian Contr. Bot. 54:1—69.
Rosinson, H. 1999. Oligactis. In: PM. Jorgensen and S. León-Yánez, eds. Catalogue of the vascular plants of Ecuador.
Monogr. Syst. Bot. Missouri Bot. Gard. 75:300-301.
ROBINSON, H. AND R.D. BrerreLL. 1974. Studies in the Liabeae (Asteraceae). Il. Preliminary survey of the genera.
Phytologia 28:43-63.
RyosERG, P.A. 1927. (Carduales) Carduaceae, Liabeae, Neurolaeneae, Senecioneae (pars). N. Amer. Fl. 34:
289-360.
STANDLEY, P.C. 1938. Liabum. Flora of Costa Rica. Publ. Field Mus. Nat. Hist., Bot. Ser. 18:1489—1491.
1214 J | of the Botanical R h Institute of Texas 2(2)
BOOK REVIEW
JEFFREY GREENE (AUTHOR), WITH MARGARET BAMBERGER (ILLUSTRATOR). 2007. Water from Stone: the Story of Selah,
Barmberger Ranch Preserve. (ISBN 10: 1585445932, ISBN 13: 978-1585445936, hbk.). Texas A&M
University Press, Lewis Street, 4354 Texas A&M University, College Station, Texas 77843-4354, U.S.A.
(Orders: www.tamu.edu/upress, 800-826-8911). $24.95, 232 pp., illustrations, 6" x 9".
Growing up in rural Ohio uin d. ken RE and World War II, Bamberger is atan did age to usn waste, Kn food
productively, and admire the har with the land. His mother ve hi
a book that would set the course for his life: xen mas by Louis Bromfield, a visionary American advocate for land restoration.
Inspired by his new role model, uper ies ib n I ever pu RA I want to do |n n did."
AC Ta ff, then
5
1YULMOr JEL
to "o and EO of Church’ s Fried Chicken, t i ionall ized tionist. In fact, Greene tells two icum
"
related stories: the evolution of one man's Pss sense, Cappie proli incentives to land restoration and
the creation of a Texas Hill Country preserve y principles. It is one of the Texas Hill cm s
greatest conservation success stories.
Bamberger bought what he describes as the sorriest piece of land in Blanco County and entered upon a decades-long effort to
restore the ecological bal f 5,500 that had been ind destroyed by more than a century of misuse. oe his preserve
Selah, from the Old Testament term meaning “pause and reflect”, Bamberger dedicates himself and hi o protecting species
and educating school children, conservation das germen rend and everyone else who will listen to = central message,
delivered with s zeal: Wi th, and anyone can
avid and his wife, Margaret, have received many BEER and he has been nur in de New P ud in ee and on
} in doing so
[2]
CNN and es news. But until now, no one has fully told the story of how a man with
transformed himself.
Jeffrey Greene is also the author of ne memoir French Spirits and t llections of poetry. The book includes illustrations, a
bibliography and index. It would be a val publi llege library.— Gary L. Jennings, Botanical Research Institute of
Texas, 500 East 4th Street, Fort Worth, Texas 76102-4025, U.S. A.
J. Bot. Res. inst. Texas 2(2): 1214. 2008
SENECIO CYMBALARIOIDES, S. SUBNUDUS, AND S. SUBPELTATUS,
HISTORY OF A MUDDLE (ASTERACEAE: SENECIONEAE)
1 F. Mais
Nati IH Nederland. Leiden branch
Post box 9514, 2300 RA, Leiden, THE NETHERLANDS
veldkamp@nhn.leidenuniv.ni
ABSTRACT
1; fth m i S i balarioides, S. subnudus, and S.
An attempt is made to disentangle the history an
cu seais a: all three consisting. Bs d i poi iid The SERIE confusion was exacerbated by further
errors, ; fly sketched here. The South African S. subnudus DC.
is BOOD
RESUMEN
is m un intento E esclarecer n E y a nea de B5 combinaciones Senecio ia S. subnudus, y S.
las per M Ones y elucidaci de | gl ] ] ] las | juí. Se lectotipifica el fri c
subnudus DC
INTRODUCTION
Senecio L. is a cosmopolitan genus with 1500+ species. Morphological research in the past and in present
molecular analyses indicate that it is polyphyletic and that it should be broken up in numerous smaller
genera. Ás this is a nomenclatural exercise I here give what I think are the correct names in Senecio and its
split-offs whenever possible. According to the present insights Packera Á. & D. Lóve and Roldana La Llave
are "good" genera.
In his revision of the Compositae in the Great Prodromus De Candolle (1838) used the combination
Senecio subnudus twice, once for a species from South Africa (p. 405, # 381), and again for one from the
United States (p. 428, # 522). The first to choose between the two was Buek (1840), closely followed by
Steudel (1841). Unfortunately, the first maintained # 381 and renamed +4 522, while the second did it the
other way around.
Apparently, it is not generally known that Buek made an index to the Prodromus, and, moreover, most
are not aware that when De Candolle's treatment was based on specimen(s) in G-DC there are IDC micro-
fiches with photographs of the specimen(s), arranged according to volume, page, and species number.
Senecio subnudus DC. (+ 381) — Senecio oxyriifolius DC.
Senecio subnudus DC., Prodr. 6:405. 1837 [publ.1838], # 381, non p. 428, # 522. Senecio subpeltatus Steud.,
mencl. ed. 2, 2:565. 1841 (nom. superfl.). Tyre: AFRICA CAPENSIS. Distr. SrenLensosch: Ecklon 787.56.8 A‘ 1835 (LECTOTYPE,
designated here: uoLorvee: G-DC; IDC MICROFICHE 800).
= Senecio oxyriifolius DC., Prodr. 6:405. 1837 [publ. 1838], # 379 ('oxyriaefolius").
Harvey (1865) was the first to include the equally old Senecio peltatus DC., S. subnudus DC. (& 381), and S.
peltiformis DC. in S. oxyriifolius DC. (see Stearn 1955), and if this ae is accepted, pu latter is the correct
name in Senecio L. The species is part of a clade of South African and M ts of the 'Gynuroid'
group, and will have to be transferred to another or even a new genus (P. Pelser, pers. comm.).
J. Bot. Res. Inst. Texas 2(2): 1215 — 1218. 2008
1216 J lof the Botanical R h Institute of Texas 2(2)
Senecio subnudus DC. (+ 522), Senecio cymbalarioides H. Buek — Packera subnuda (DC.) Trock
& T.M. Barkley
Senecio E ss eee 6: oe 1837 [publ. 1838], ie non p. 405, # 381. Steud., Nomencl., ed. 2,
22 Bue
p. Synon. Cand. 2:vi. 1840, nom. nov. Senecio aureus L. var. subnudus A. Gray,
s d
Syn. a N. Amer. 1, 2:391. 1884. Senecio pauciflorus Pursh var. subnudus Man. Fl. Pl. Calif. 1154. 1925. Packera cymbalarioides
Weber & A. Love in WA. Weber, Phytologia 55:9. 1984, non WA. Weber and A. Lóve (1981). Packera buekii Trock & T.M.
Barkley, Sida 18:387. 1998, nom. superfl., see note. Packera subnuda Trock & T.M. Barkley, Sida 18:635. 1999. Type: AMERICAN
WEST COAST: Columbia River, Douglas A* 1830 (HoLotyPE: G-DC; IDC wicRoricHE 800), ‘as HON. Soc. Honric. Lonp.’ (This herbarium
was broken up in auctions between 1856 and 1859. I do not know bii the e went. There is a dol pd inK
1
“ex Herb. Peu ” with a predl Bees not rondes: asin ki DC I when material
l label, so a h itt had to! de)
to De C was only g
= Senecio cymbalarioides H. Buek. or Packera subnuda (DC.) Trock & T.M. Barkley.
The status of the rejected name S. subnudus DC. # 522 has been clarified by the addition of Note 3 to Art.
53.6 in the ICBN (2006):
Art. 53.6. When two or more homonyms have equal priority, the first of them that is adopted in an effectively published text (Art. 29,
30, 31) by an author who simultaneously rejects the other(s) is treated as having priority. Likewise, if an author in an effectively
as having priority.
Note 3. A1 ] f dor Art 52
should a SM to another n . be effected.
Ex. 20. Mimosa cineraria L. (1759), based on M. cinerea L. (Sp. Pl. 517, non 520. 1753; see Art. 53 Ex. 18), was transferred to Prosopis by
dus nen as P ane (L.) oe ONE the correct name in Prosopis would have been a combination based on M. cinerea
According to Trock and Barkley ne followed dby Trock (2006) S. subnudus DC. # 522 belongs to Packera
Á. & D. Lóve. Trock and Barkley (1998) had called it P. buekii which is superfluous as they should have used
the epithet subnuda under Note 3 above. They corrected this a year later (1999) to Packera subnuda (DC.)
Trock & T.M. Barkley.
Senecio subpeltatus Sch.-Bip. — Roldana subpeltata H. Rob. & Brettell
The second author to coin the combination Senecio subpeltatus was Schultz “Bipontinus” (1856) for a Mexi-
can species. This is a later homonym of the one by Steudel (1841). Curiously, no one seems to have picked
this up and proposed an alternative name in Senecio for it. When transferred to Roldana it is to be cited as
Roldana subpeltata H. Rob. & Brettell (Art. 58.1), without bracketed authorship, i.e., not as [Sch.-Bip.] H.
Rob. & Brettell.
In older Codes “square-bracketed authorship” was used to indicate that there was something “wrong”
with the basionym, e.g. based on a pre-Linnaean one, or on an invalid or illegitimate base. Unfortunately,
this useful application has quietly vanished after the Edinburgh Code (1966). The reference was still given
in the index of the Seattle Code (1972), but not in the text.
Senecio subpeltatus Sch.-Bip., in Seem., Bot. Voy. Herald 311. 1856, non Steud. (1841). Roldana subpeltata H.
Rob. & Brettell, Phytologia 27:424. 1974. Tre: MEXICO: Sierra Madre, Seemann 1984 (HoLorYrE: BM; isorrrE: K, P, F-photo, GH).
= Roldana subpeltata H. Rob. & Brettell
Dr. A.M. Funston (in sched.) regarded the K collection as the lectotype, but Seemann’s “study set” for the
Herald voyage is in BM (Stafleu & Cowan 1985: 474) and the holotype should be there.
Senecio cymbalarioides Nutt., non H. Buek
Senecio cymbalarioides Nutt., Trans. Amer. Phil. Soc. II, 7:412. 1841, non H. Buek (1840). Packera cymbalarioides
WA. Weber & A. Love, Phytologia 49:46. 1981. Tree: U.S.A. Orecon: Nuttall s.n. (HoLOTYPE: BM)
The situation has become more complicated and confusing because Weber and Lóve made the combina-
tion Pachera balarioides twice: once in 1981 based on Nuttall (1841), and again in 1984, based on Buek
P4
Veldkamp, Senecio cymbalarioides, S. nudus, and S. subpeltatus 1217
(1840). The latter one is therefore a later homonym of the first one, and not an isonym, as was implicitedly
suggested by Weber and Lóve (1984), who apparently thought that the same taxon was involved and that
merely changing the bracketed author would ‘correct’ their earlier combination.
In Senecio the correct name is Senecio streptanthifolius Greene according to Cronquist [(1994: 184),
with extensive synonymy]. If it is a Packera the name is Packera streptanthifolia (Greene) W.A. Weber &
Á. Lóve (1981: 48), see Trock (2006: 599) who did not accept the varieties distinguished by Bain (1988).
Senecio balarioides subsp. biensis Calder & R.L. Taylor
Senecio M rares (author not given; H. Buek, see below) subsp. moresbiensis Calder & R.L. Taylor,
d. J. Bot. 43:1399. 1965. Senecio tas Greene var. moresbiensis (Calder & R.L. Taylor) B. Boivin, Naturaliste
a 93:1051. 1966, nom. inval., see note moresbiensis (Calder & R.L. Taylor) B. Boivin, Phytologia
23 (1972) 108. Senecio moresbiensis (Calder & R.L. Taylor) G.W. i & Ruyle-Dougl., Canad. J. Bot. 56:1710. 1978. Packera
moresbiensis daga & R.L. Taylor) J.E Bain, Novon 9:457. 1999. Packera subnuda (DC.) Trock & T.M. Barkley var. moresbiens
(Calder & R.L. Taylor) Trock, Sida 21:289. 2004. Tyre: CANADA. BritisH CoLumBIA: Moresby Isl., Takakia lake, Calder, ie &
Taylor 23066 rbd DAO).
The authorship of the specific combination was not given by Calder & Taylor. In 1968 they cited Nuttall as
the author, but Taylor (1978) said that Buek had been intended. 1 suspect she fell in the same trap as Weber
& Lóve previously did.
Boivin made th bination Senecio streptanthifolius Greene var. moresbiensis B. Boivin, which is invalid,
as it lacked a full and direct reference to the sera Since in Senecio this seems to be the name required at
the varietal level, the necessary combination needs to be validated. As a species it is Senecio moresbiensis
iud in E L. in G.W. Douglas > Ruyle-Dougl.
name is P, bnuda (DC.) Trock & T.M. Barkley var. moresbiensis (Calder
S RL. pen Trock or Packera moresbiensis (Calder & R.L. Taylor) J.F. Bain. Trock (2004: 289) opted to
recognise the varietal status. That in Trock (2006) Pachera streptanthifolia is followed by Packera subnuda is
not indicative of any other relationship than an alphabetical one.
ACKNOWLEDGMENTS
I want to thank C. Flann (WAG) for pointing out the problem, P.B. Pelser (MU) for his comments on S.
subnudus DC. #381, D.G. Frodin (K) for the information on the 19th century herbarium of the Royal Horti-
cultural Society, DJ.N. e ES for finding a possible isotype of Senecio subnudus DC. in K, T.S. Quedensley
(TEX) for information on R bpeltata, and D.K. Trock (CAS-DS) and G. Nesom (BRIT) for their critical
reviews. The resumen was made by R. Camara Loret, Leiden.
REFERENCES
BAIN, J.F. 1988. Taxonomy of Senecio streptanthifolius. Rhodora 90:287.
BarkLEY, T.M. 1999, The segregates of Senecio, s.l., and Cacalia, s.l., in the flora of North America North of Mexico.
Sida 18:635, 668, 670.
Buek, H.W. 1840. Genera, species et synonyma candolleana alphabetico ordine disposita. Nauk, Berlin. 2:vi.
CALDER, J.A. AND R.L. TayLOR. 1968. Flora of the Queen Charlotte Islands. Part I. Systematics of the vascular plants.
Canad. Dept. Agric. Res. Branch Monogr. 4(1). Queens Printer, Ottawa. (n.v.).
Cronauist, A. 1994. In: Cronquist, A. et al, Intermountain flora. New York Botanical Garden, Bronx. 5:180, 184.
Dt CANDOLLE, A.P. 1837 [publ. 1838]. Prodromus systematis regni vegetabilis. Treuttel and Würtz, Lille. 6:405, 428.
Harvey, W.H. 1865. In: Harvey, W.H. and O.W. Sonder, Flora capensis. Hodges, Smith & Co., Dublin. 3:376-377.
NurraLL, T. 1841. Descriptions of new species and genera ... of the Compositae. Trans. Amer. Phil. Soc. Il, 7:412.
ScHuLtz, C. (“BIPONTINUS”). 1856. In Seeman, B. The botany of the voyage of H.M.S. Herald. Reeve & Co., London. P. 311.
STAFLEU, F.A. AND R.S. Cowan, 1985. Taxonomic literature, ed. 2. Bohn, Scheltema 8: Holkema, Utrecht, etc. 5:474.
STEARN, W.T. 1955. Buek and Steudel's conflicting names for species of Compositae described in de Candolle's
Prodromus. Candollea 15:67.
Journal of the Botanical R h Institute of Texas 2(2)
1218
STeuDEL, E.G . 1841. Nomenclator botanicus, ed. 2. Cotta, Stuttgart, Tubingen. 2:565
Tavlor, R.L. 1978. In: Taylor, R.L. and B. MacBryde. New taxa and nomenclatural changes with respect to vascular
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Trock, D.K. 2006. Packera. In: Flora of North America north of Mexico. Oxford University Press, New York, Oxford.
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TRock, D.K. AND T.M. Barkley. 1998. Seven new nomenclatural combinations and a new name in Packera. Sida
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flora- IV. Phytologia 55:9.
2C DNA CONTENT VALUES IN AMARANTHUS (AMARANTHACEAE)
Donald B. Pratt Shalini N. Jhangiani
Stephen F Austin State University 12031 Hedgegate Drive
artment of Biology Houston, Texas 77065, U.S.A.
PO. Box 13003 SFA Station
Nacogdoches, Texas 75962, U.S.A
prattdb@sfasu.edu
Robert J. Wiggers
Stephen F Austin State University
alt of Biology
PO. Box 13003 SFA Station
ee Texas 75962, U.S.A
ABSTRACT
Feulgen densitometry was used to measure 2C DNA content values from 15 accessions of 12 species of Amaranthus representing all
subgenera. Species showed a 3-fold variation in range from 0.89 pg (Amaranthus viridis) to 2.73 pg (Amaranthus tricolor). Values are
4 : A Pd 14 Á f DNA 7 di a
F
RESUMEN
Se usó la densitometría de Feulgen para medir los valores del contenido de 2C DNA de 15 accesiones de 12 especies de Amaranthus
que representan todos los subgéneros. Las especies mostraron una variación de 3-veces 89 pg (Amaranthus viridis) a 2.73
pg (Amaranthus tricolor). Se comparan los valores con informes previos y se discuten las implicaciones taxonómicas de los valores de
contenido de DNA
INTRODUCTION
Amaranthus L. (Amaranthaceae) is a genus of about 70 species of monoecious or dioecious annuals of world-
wide distribution. The genus is economically important, including the cultivated grain amaranths, potherbs,
and ornamentals as well as a suite of agricultural and range weeds. Traditionally the genus has been divided
into two subgenera, subg. Amaranthus (the monoecious amaranths) and Acnida (dioecious amaranths) based
on breeding system (Robertson 1981). The most recent classification system divides Amaranthus into three
subgenera: Amaranthus (with 12 species), Acnida (with 9 species), and Albersia (with 49 species) based on
breeding system and characteristics of ue fruit and flowers (Mosyakin & Robertson 1996).
Chromosome counts of the A la base count of x = 8 or x = 9 (Turner 1994). Amaranthus
itself appears to be paleopolyploid with counts of n = 16 or n = 17 (Grant 1959a; Grant 1959c; Greizerstein
& Poggio 1992; Greizerstein & Poggio 1994), and genomes behave as diploids during meiosis with the
exception of Amaranthus dubius, an allotetraploid with n = 32 between A. spinosus and an unknown species
(Grant 1959b; Greizerstein & Poggio 1992). Cytogenetic research d 1 that variation between n =16
and n -17 can occur within individuals of a single population, the extra chromosome in individuals with
counts of n = 17 is derived via primary trisomy from n = 16 (Pal et al. 1982; Griezerstein € Poggio 1994). In
addition to chromosome number, karyotypic work has demonstrated that Amaranthus chromosomes are of
small size (Grant 1959a; Grant 1959c; Griezerstein & Poggio 1994); possess a single NOR locus (Griezer-
stein & Poggio 1994); and demonstrate a wide variety of C banding patterns putatively due to differences
in heterochromatin content (Griezerstein & Poggio 1994).
Aside from chromosome counts, very little research has been directed towards analysis of DNA con-
tent in Amaranthus. To date, 2C DNA content in picograms has been recorded for 14 species in Amaranthus
(Griezerstein & Poggio 1994; Bennett et al. 2000; Jeschke et al. 2003; Rayburn et al. 2005). Taxonomic
J. Bot. Res. Inst. Texas 2(2): 1219 — 1223. 2008
1220 J | of the Botanical R h Institute of Texas 2(2)
sampling of the DNA content studies is highly skewed. Of the 14 reported counts, nine reports come from the
weedy and cultivated species of subgenus Amaranthus (which contains an estimated 12 species), two reports
from subg. Acnida (which contains nine species), and three reports from subg. Albersia (which contains 49
species). However, given the wide range of previously reported DNA content values, it would be prudent to
expand sampling of subg. Acnida and Albersia.
The objectives of this study were therefore twofold: 1) to obtain estimates of DNA content from a wide
variety of species of Amaranthus including representatives of all three subgenera using an image analysis
densitometry technique; and, 2) to explore possible taxonomic significance of DNA content variation.
MATERIALS AND METHODS
Seed from fifteen accessions and twelve species of Amaranthus were donated by the USDA North Central
Regional Plant Introduction States (NCRPIS). Known 2C DNA values for Amaranthus were obtained from
the database at Kew Gardens (data.kew.org/cvalues/), and Sorghum bicolor Pioneer cultivar 8695 was chosen
as the reference standard based on its 2C DNA content of 1.74 pg (Johnston et al. 1999) which compared
favorably to the range of values (0.96—2.46 pg) reported in the Kew database.
Amaranth seeds were stratified at 4°C for one month and plants grown in a greenhouse. Roots were
fixed in 3:1 ethanol: glacial acetic acid solution and stored in 7096 ethanol at 4°C. Fixed root material was
hydrolyzed in 5N HCl for 30 minutes and stained in freshly prepared and charcoal-filtered Fuelgen stain
for three hours, and destained for 15 minutes in distilled water. Fuelgen stained roots tips were softened
45 minutes in 296 w/v cellulysin enzyme in 0.001 M EDTA pH 5.6 solution, replaced with 4596 acetic acid,
squashed, cryofixed in liquid nitrogen, and dried overnight before adding coverslips.
Chromosome squashes were photographed on an Olympus BX50 Light Microscope using a Nikon
UR-E6 Coolpix MDC Lens attached to a Nikon Coolpix 5000 digital camera. Twenty to thirty images of the
stained nuclei per specimen were uploaded, analyzed using a wavelength of 560 nm (Hardie et al. 2002),
and integrated optical density (1.0.D.) values were calculated using Image Pro Plus 5.1 (Media Cybernetics,
Inc., Maryland).
IOD values for nuclei were pooled after discarding the top and bottom 596 of values if an ANOVA in-
dicated no difference between the means from different images (Hardie et al., 2002), and mean IOD values
were calculated from the pooled data. A coefficient of variation (COV) was calculated (mean / stdev.) and
the IOD data was considered usable only if the COV was less than 1096 (Vilhar et al. 2001; Hardie et al.
2002). Absolute DNA contents in picograms for all Amaranthus populations were determined comparing
the mean IOD values to the mean IOD for S. bicolor standard that was processed in the same stain batch as
[Amaranth 10D/ ]*1.74 pg
the Amaranthus spp using the following calculation: OS
RESULTS
Mean 2C DNA content values are reported in Table 1. Among the fifteen accessions sampled we found 3-fold
variation in 2C DNA contents, ranging from a low of 0.89 pg in A. viridis to a high of 2.73 in A. tricolor. The
two subspecies of A. graecizans had a 1.1-fold difference in values (1.33 and 1.46). The two accessions of
A. arenicola had 1.31 fold difference in content values (1.15 and 1.5). The two accessions of A. blitum had a
1.14-fold range in content values (1.06 and 1.21). Subgenera Acnida and Albersia showed a 1.14 and 1.07-fold
increase in DNA content values over values found in subgenus Amaranthus. Within subg. Albersia, section
Pyxidium showed a 1.8-fold increase in DNA values over sections Blitopsis and Pentamorion.
DISCUSSION
We report 2C DNA content values from 15 accessions of 12 species (Table 1), and provide reports for eight
species and two sections (Blitopsis and Pentamorion) that had previously been unreported in the literature
(Table 2), expanding the sampling from Amaranthus to 23 species. Our values compare favorably to previ-
ously reported values (Table 2).
Pratt et al., DNA content in Amaranthus
£ Lael
TABLE 1. Li
, species, and 2C DNA content values
1221
Taxon Location Accession 2C
Subgenus Amaranthus 1.23
A. caudatus Peru PI 490440 1.32
A. dubius Panama Ames 25792 1.23
A. powellii bouchonii Germany PI 572261 1.13
Subgenus Acnida 1.40
A. arenicota Kansas PI 607459 1.51
A. arenicola Kansas PI 599671 1.15
A. greggii PI 632240 1.54
Subgenus Albersia 1.32
Sect. Blitopsis 1.02
A. acutilobus Germany PI 633578 1.22
A. blitum India PI 608661 1.06
A. blitum ndia PI 610262 1.21
A. viridis Maldives PI 536439 0.89
Sect. Pyxidium 1.84
A. tricolor Pennsylvania PI 599683 2.73
A. graecizans silvestris India Ames 24671 1.46
A. graecizans thellungianus Mauritius PI 549157 1.33
Sect. Pentamorion 1.01
A. crassipes Cuba Ames 10339 0.97
A. standlevanus Argentina PI 61380 1.02
Values from subg. Amaranthus range from 1.04 pg in A. hybridus to 1.9 pg in A. spinosus with a composite
average (our values as well as all other reported values) of 1.29 pg (Table 2). Two of the reported values, A.
retroflexus 1.7 pg and A. spinosus 1.9 pg, are suspiciously high (1.31 and 1.4-fold higher than the average)
compared to other reported values for these species and for other member of subg. Amaranthus (Table 2).
If these two values are removed, the range of values for the subgenus decreases to 1.04-1.35 pg, with a
composite average of 1.23 pg which matches our values closely (Table 2).
Several DNA content values from subg. Amaranthus are noteworthy. The 1.23 pg DNA content for the
allotetraploid A. dubius is particularly intriguing (Table 1). The most likely 2C DNA content value of the
known parental species, the diploid A. spinosus, is 1.10 pg (Table 2). If these values for A. dubius and A.
spinosus are correct, then either the DNA content of A. dubius has undergone a major contraction following
the polyploidization event, or the unknown parent has a very small genome size. The reported values for A.
hybridus and A. quitensis (a South American form of A. hybridus) range from 1.04 to 1.30 pg, with an average
of 1.18 pg. This species is a wide-ranging, cosmopolitan weed that originated in the Americas. The range
in values may represent geographic or taxonomic patterns of distribution that should be better examined.
Reported values from subg. Acnida (the dioecious amaranths) represent a 1.62-fold range from 0.95
to 1.54 pg in DNA content (Table 2). This subgenus is closely related to species of Subg. Amaranthus as de-
termined by crossing data (Murray 1940; Trucco 2006). The large range of reported values may represent
considerable changes in DNA content following the evolution of dioecy. There is a 1.3 fold difference in
2C DNA content values for A. arenicola. The accessions were collected from the Arkansas River in Kansas
two years apart by different collectors. Amaranthus arenicola is a wild species endemic to the area, and has
received little study. These data indicate the possibility that the area may be a center of diversity for the
species and further research into the species is warranted. The reported DNA content value for A. palmeri
1222 Journal of the Botanical R h Institute of Texas 2(2)
Tag 2. DNA lues fr
OurValue Literature Our Value Literature
Subgenus Amaranthus Subgenus Albersia
A. caudatus 1.32 1.30! Sect. Blitopsis
1.35? A. acutilobus 1.22
1.265 A. blitum 1.06
A. cruentus 1,30! A. blitum 1.21
1.26? A. viridis 0.89
A. dubius 1.23 Average values 1.02 na
A. hybridus 1.30! Composite average 1.02
1.04?
A. hypochondriacus 1.30! Sect. Pyxidium
1.26? A. albus 1.184
A. powellii powellii 1.20' A. blitoides 1:151
1.148 A. tricolor 2.73 1.80
A. powellii bouchonii — 1.13 1.20* 246
A. quitensis 1.20! A. graecizans silvestris 1.46
A. retroflexus 1.70! A. graecizans thellungianus 1.33
1.233 Average values 1.84 1.65
A. spinosus 1.90! Composite average 1.73
1.10%
Average values 1.23 1.30 Sect. Pentamorion
Composite average 1.29 A. crassipes 0.97
A. standleyanus 1.02
Subgenus Acnida Average values 1.01 na
A. arenicola 1.51 Composite average 1.01
1.15
A. greggii 1.54 'Bennett, Bhandol, & Leitch 2000
A. palmeri 0.95? *Greizerstein & Poggio 1994
A. tuberculatus 1.34? ? Jeschke, Tranel, & Rayburn 2003
Average values 1.40 1.14 ^Rayburn et al. 2005
Composite average 1.30 *RGB Kew Gardens DNA C-Values Database
(0.95 pg, Table 2) is very low compared to other dioecious waterhemps, but is in-line with recorded values
for A. spinosus (1.10 pg, Table 2). These DNA content values support Wassom and Tranel's (2005) AFLP data,
which placed A. palmeri in a clade with A. spinosus rather than with other dioecious amaranths.
We provide DNA content data for six new species and eight accessions from subg. Albersia (Table 2).
Albersia is the largest subgenus of Amaranthus, although most of the species are poorly known (Mosyakin
and Robertson 1996). DNA content values in the subgenus ranged from the smallest value (A. viridis with
0.89 pg) to the largest (A. tricolor with 2.73 pg) in the study (Table 2) indicating that there is a great deal
of variation in DNA content values within the subgenus. Values within sections Blitopsis (1.02 pg) and
Pentamorion (1.01 pg) represent the lowest values in the study (Table 2). Values within section Pyxidium are
highly variable, ranging from 1.15 pg for A. albus and 2.73 pg for A. tricolor, a 2.4 fold range of DNA contents.
This is not surprising as Mosyakin and Robertson (1996) report that the section was morphologically and
phytogeographically “unsatisfactory,” and state that it could probably be further divided into subsections,
a conclusion that might be supported by DNA content values.
Accessions of three species (A. albus, A. blitum, and A. powellii powellii) were sampled, but DNA contents
could not be reliably quantified due to the presence of abnormally large, darkly stained nuclei that were
me n by side with nuclei of normal appearance. These large nuclei are most likely attributable to
yploidy, a condition that is frequent in Chenopodium and A t gs (B Kolano, pers.
comm., manuscript in preparation).
Pratt et al., DNA content in Amaranthus 1223
To date, differences in genome size have been used to identify hybrid individuals (Trucco et al. 2005;
Trucco et al. 2006). In addition to increasing reported DNA content values within Amaranthus by eight ad-
ditional species, we observed that patterns in DNA content might be taxonomically informative both within
and among species (Table 2). Additionally we note the contraction of the genome following polyploidization
in A. dubius and the presence of endopolyploidy in Amaranthus.
REFERENCES
BENNETT, M.D., P. BHANDOL, AND 1... Lerch. 2000. Nuclear DNA amounts in Angiosperms and their modern uses—807
new estimates. Ann. Bot. 86:859-909,
GRANT, W.F.1959a. Cytogenetic studies in Amaranthus |. Cytological aspects of sex determination in dioecious
species. Canad. J. Bot. 37:413-417.
Grant, W.F. 1959b. Cytogenetic studies in Amaranthus Il. Natural interspecific hybridization bet Amaranthus
dubius and A. spinosus. Canad. J. Bot. 37:1063-1070.
Grant, W.F. 1959c. Cytogenetic studies in Amaranthus Ill. Chromosome numbers and phylogenetic aspects.
Canad. J. Genet. Cytol. 1:313-328.
GRIEZERSTEIN, E.J. AND L. Pocaio. 1992, Estudios citogenéticos d is híbridos i pecíficos de Amaranthus (Ama-
ranthaceae). Darwiniana 31:159-165,
GRIEZERSTEIN, E.J. AND L. Poscio. 1994. Karyological studies in grain amaranths. Cytologia 59:25-30.
HARDIE, D.C., R. GREGORY, AND D.N. Heneenr. 2002. From pixels to picograms: A beginner's guide to genome quanti-
fication by feulgen image analysis densitometry. J. Histochem. Cytochem. 50:735-749.
Jeschke, M.R., PJ. TRANEL, AND A.L. RAYBURN. 2003. DNA content analysis of smooth pigweed (Amaranthus hybridus)
and tall waterhemp (A. tuberculatus): implications for hybrid detection. Weed Sci. 51:1-3.
JOHNSTON, J.S., M.D. Bennett, A.L. RAYBURN, D.W. GALBRAITH, AND H.J. Price. 1999. Reference standards for determination
of DNA content of plant nuclei. Amer. J. Bot. 86:609—613.
Mosyakin, S.L. AND K.R. ROBERTSON. 1996, New infrageneric taxa and combination in Amaranthus (Amaranthaceae).
Ann. Bot. Fenn. 33:275-281.
Murray, M.J. 1940. The genetics of sex determination in the family Amaranthaceae. Genetics 25:409-431.
PAL, M, R.M. PANDEY, AND T.N. KHOSHOO. 1982. Evolution and improvement of cultivated amaranths. IX. Cytogenetic
relationship of the two basic chromosome numbers. J. Heredity 73:353-356.
RAYBURN, A.L, R. McCLoskty, T.C. TATUM, G.A. BoLLERO, M.R. JESCHKE, AND PJ. TRaneL. 2005. Genome size analysis of weedy
Amaranthus species. Crop Sci. 45:2557—2562.
RosERTSON, K.R. 1981. The genera of Amaranthaceae in the southeastern United States. J. Arnold Arbor. 62:
Trucco, F, T. Tatum, A.L, RAYBURN, AND P.J. TRANEL. 2005. Fertility, segregation at a herbicide-resistance locus, and genome
structure in BC, hybrids from two important weedy Amaranthus species. Molec. Ecol. 14:2717-2728.
Trucco, F, T. TATUM, K.R. ROBERTSON, A.L. RAYBURN, AND PJ. TRANEL. 2006. Characterization of waterhemp (A. tuberculatus)
x smooth pigweed (A. hybridus) F, hybrids. Weed Technol. 20:14-22.
Turner, B.L. 1994, Chromosome numbers and their phyletic interpretation. In: H.D. Behnke and TJ. Mabry, eds.
Caryophyllales evolution and systematics. Springer-Verlag, New York, USA. Pp. 27-43.
VILHAR, B., J. GREILHUBER, J.D. Koce, E.M. Temsch, AND M. Demastia. 2001. Plant genome size measurement with DNA
image cytometery. Ann. Bot. 87:719-728.
WASSOM, J.J. AND P.J. TRANEL. 2005. Amplified fragment length polymorphism-based genetic relationships among
weedy Amaranthus species. J. Heredity 96:410-416.
1224 || l of tha Rat seal D en Peer mers
f Texas 2(2)
BOOK REVIEW
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J. Bot. Res. Inst. Texas 2(2): 1224. 2008
MINUARTIA MACRANTHA (ALSINOIDEAE: CARYOPHYLLACEAE):
MORPHOLOGICAL CIRCUMSCRIPTION, GEOGRAPHICAL RANGE,
AND PHYLOGENETIC AFFINITIES
Ronald L. Hartman Richard K. Rabeler
Rocky Mountain lia University of Michigan Herbarium
Department of Botany 3600 Varsity Drive
University of Wyoming Ann Arbor, Michigan 48108-2228, U.S.A.
Laramie, Wyoming 82071-3165, U.S.A. rabeler@umich.edu
rhartman@uwyo.edu
ABSTRACT
Minuartia macrantha is here defined to include M. filiorum. The traditional apena ipud dan the two iiia taxa
are discussed. Previously, M. macrantha was considered endemic to Colorado; now Minuartia
filiorum o M. mocinha) i is not E no to M. Maii uA to a recent floristic t treatment mich erme it as a variety of
the latter. A the M. rossii-stricta complex. This alliance includes the northern
Rocky Mountain M. austromontana, recently found sympatric us M. macrantha in northwest Wyoming. Minuartia stricta is discussed
briefly as it is often confused with M. macrantha in Colorado
RESUMEN
Se define aquí Minuartia macrantha po ou BRS M. lie Se ps e ae meres puce entre los E
r
supuestos taxa. Previamente, M.
e e o
Minuartia filiorum (= M tha) tá lacionad M. rubella, fl
1 iedad d alti Mi i } id l ña del lejo M. rossii-stricta. Esta
x L P5 El
1; : y ] ña de las M. ñas R M. austromontana 1 i impátri M. macrantha en el
noroeste de Wyoming. Minuartia stricta se di I yaq funde f M. macrantha en Colorado.
Our intent is to elaborate on the morphology of Micrantha macrantha s.l. (including M. filiorum), and the
degree of variability of diagnostic characters. Furthermore, the marked change in geographical range is
documented (see specimens cited and Fig. 1). Finally, the presumed phylogenetic affinities of M. macrantha
are discussed.
Bassett Maguire is recognized for his superb taxonomic treatments of Arenaria, including currently
recognized segregate genera (Eremogone, Honckenya, Minuartia, and Moehringia), in North America and his
collaboration with C. L. Hitchcock on Silene. Subsequent to Maguire's work, Stanley L. Welsh and associates
at Brigham Young University obtained numerous collections from Utah and vicinity that were determined
as M. filiorum. Correspondingly, floristic studies in Colorado and Wyoming (COLO, RM) contributed a
wealth of material, chiefly from these core southern Rocky Mountain states, that were determined as M.
macrantha. Unfortunately, there was relatively little communication between these two areas, although a
number of duplicates were exchanged. When we examined these collections during the preparation of the
treatment of Minuartia for the Flora of North America (Rabeler et al. 2005), we found the specimens filed as
M. macrantha and M. filiorum to be more difficult to segregate than the literature suggested.
MORPHOLOGY
Maguire (1946) described Minuartia filiorum and later (1958) distinguished it from M. macrantha (both
under Arenaria) on the basis of habit (annual or weakly perennial vs. obviously perennial with numerous
procumbent stems to 10 cm), cymes (1-5- vs. 3(-5)-flowered), flowers (not showy vs. large and showy),
sepal length and shape (3.5—4.8 mm, ovate-lanceolate vs. 4.5—5 mm, broadly lanceolate), and petal to sepal
ratios (shorter than to equaling sepals vs. conspicuous, exceeding them, to 8 mm long, respectively). While
J. Bot. Res. Inst. Texas 2(2): 1225 — 1231. 2008
126 | | | | | doumalof tanical h Institute of Texas 2(2)
Fic. 1. Maj ElL L n kyM . U.S.A., mmn
of M. stricta.
some populations may be differentiated using these features, many cannot be so recognized. For example,
the number of flowers per inflorescence, the length of the petals relative to the sepals, and sepal length are
more variable than portrayed. On the other hand, the mature sepals of both taxa are strongly ribbed (Fig.
2a) and the seeds are ially identical. We concur with W.A. Weber and R. Wittmann's annotation of the
th tha (Rydb.) W.A. Weber 1983) as inseparable from M. macrantha;
Alc
holotype of M. filiorum (as
our independent conclusion (Rabeler et al. 2005).
4
1227
irich 2910 BRY ) and M. t
(Elliott 11048a, RM). Note scale bars
macrantha.
1228 J | of the Botanical R h Institute of Texas 2(2)
Minuartia macrantha (Rydb.) House, Amer. Midl. Naturalist 7:132. 1921. Alsinopsis macrantha Rydb., Bull. Torrey
Bot. Club 31:407. 1904. Arenaria macrantha (Rydb.) A. Nelson, New Man. Bot. Centr. Rocky Mt. 186, 608. 1909, non Schischk.
Fl. URSS 6:522, 886. 1936. Alsinanthe macrantha (Rydb.) WA. Weber, Phytologia 51:369. 1982. Tree: U.S.A. Coronapc: La Plata
Co. [Montezuma Co.]: La Plata Mts., Little Kate Basin, common alpine form, 14 Jul 1898. C.F Baker, F S. Earle, & S.M. Tracy 678
(HoLorvrE: NY, photo!; rsorvee: NY, photo!; RM, 2!).
Arenaria filiorum Maguire, Bull. Torrey Bot. Club 73:326. 1946. Minuartia filiorum (Maguire) McNeill, Rhodora 82:498. 1980. Arenaria
rubeila (Wahlenb.) Sm. var. filiorum (Maguire) S. L. Welsh, Rhodora 95:393. 1993. Tree: U.S.A. Uran. Kane Co.: [Iron Co.]: N
Lake, [9030 ft], common, gravelly beaches, 13 Jul 1940. B. Maguire 19472 (morore: NY, pl GH, UTC, UC).
Common nanes.—House’s stitchwort, large-flower sandwort, beach sandwort.
Plants perennial, cespitose or mat-forming, or sometimes annual, glabrous throughout. Taproots
occasionally filiform, but usually somewhat thickened to moderately stout, often woody. Stems erect to
numerous and procumbent, green, 2-20 cm long, internodes of all stems 0.3-1(-2) times leaf length. Leaves
moderately to tightly overlapping (proximal cauline), variably spaced, progressively more so distally (distal
cauline), connate proximally, with loose, scarious sheath 0.3-0.8 mm long, blade straight to slightly out-
curved, green, flat to triquetrous distally, 1-3-veined, fleshy, midvein more prominent than 2 lateral veins,
subulate to linear, 5-10 mm long, 0.5-1.2 mm wide, flexuous, margins not thickened, scarious, smooth,
apex green, rounded, thickened and navicular, shiny; axillary leaves present among proximal cauline leaves.
Inflorescences terminal, 2—5(-9)-flowered, open to strict cymes or more often a solitary flower; bracts broadly
subulate, herbaceous or scarious-margined proximally. Pedicels 0.2-1.5 cm long. Flowers perigynous,
5-merous; hypanthium disc-shaped; sepals ovate to lanceolate (herbaceous portion lanceolate), 3.5—5.5 mm
long, to 6.5 mm in fruit, margins thinly scarious (0.1-0.2 mm wide), inrolled near apex, strongly 3-ribbed,
especially in fruit, rarely weakly so, ribs rounded, apex green or purple in part, blunt to sharply acute or
acuminate, not hooded but often incurved; petals oblong to obovate, (0.7—)1.2—1.5(-2) times sepal length,
apex rounded to blunt, entire (Fig. 2); stamens 10, those opposite the sepals with basal nectaries 0.3-0.4 mm
long, 0.4—0.5 mm wide, triangular, tapering distally, each with a shallow adaxial groove, collectively with
the brief hypanthium forming a nectar dish; styles 3, filiform, 1-3 mm long with stigmatic papill
or nearly so. Capsules on stipe ca. 0.2 mm long, broadly ovoid, 3-3.8 mm long, somewhat shorter than
to slightly exceeding the sepals, valves 3, recurved at apex. Seeds 8-13, black, suborbiculate with radicle
Men to rounded beak, somewhat M 0.7-1 mm us tuberculate; tubercles low, rounded.
ranitate
Collecti I d tated fili Maguire are indicated by an “m” followed by the
Be barum abbreviation and literature citation where appropriate (e.g., (m; | USES), (m; NY, UTC, cited Madrorio 8:262. 1946)). Up to
three collections are ic E county.
ZONA. C ino Co.: San Francisco Mts., 12,000 ft VUE 1938, Little 4664A (USFS); Kaibab Plateau, De
Motte e 8,700 T ers 1977, Reeves 5634 (ASU). COLORADO. Archulet tz Lake trail, 6 Sep 1998, Heil 12 (SC). Clear
Jul 1888, Eastwood s.n. (COLO). Conejos Co.: S of Blue Like ur ih a gol pd gai ni Rid (RM).
Dolores Co.: o: Navajo Ba Basin, 11,140—12,100 ft, 26 Jul 1994, Moore 3441 (RM) k, 10,200-11,800
ft, 27 Jul 1 33 (RM). Garfield Co.: neag oa Elk Creek, Flat Tops, 10,000-10,400 ft, 24 Jul 1990, H 25915 (RM);
eine Mesa 10,300-10,500 ft, 24 Jun 1990, I 38 (RM); Indian Camp Pass, 9,800—10,000 ft, 6 Aug 1991, Vanderhorst 3824
RM). Gunnison Co.: Horse Basin, 11,600 ft, 7 Aug 1950, ‘angen 70 (COLO); Comanche Creek, 12,000 ft, 14 Jul 1951, Langenheim
1361 SR oe Ridge area, 12,450 ft, 30 Jul 1989, Lehr 1262 (COLO); Double Top Mt., 11,500—11,900 ft, 14 Jul 1997, Taylor 3293
(RM). Hi ak area, 10,420—11,425 ft, 11 Jul 2002, Heil & Mietty 19182 (SJC). Lake Co.: Weston Pass, 12,000 ft, 14
Jul 1940, Gierisch 1298 (USFS). La Plata Co.: Silver Mesa near Lake Marie, 12,200 ft, 11 Jul 1993, Cramer 67 (COLO); Chicago Basin,
12,000 ft, 13 July 1934, Penland 1075 (COLO); Lime Mesa, 21 Aug 1982, Siplivinsky & Bech 4897 (COLO). Park Co: Weston Pass, 11,700
ft, 28 Jul 1985, Hartman 6357 (RM); Mt. Bross, 3,500—3,900 m, 25 Jul 1982, Weber et al. 2115 (COLO, RM); North Star Mt., 12,300 ft, 3
Jul 1954, Weber 8751 (COLO). Pitkin Co.: ridge E of Avalanche Ck, 12,000 ft, 14 Jul 1952, Langenheim 2106 (COLO). Rio Grande Co.
Wet Canyon, 2.6 mi NW of Elwood Pass, 10,500 ft, 24 Jul 1996, Heil 10367 (SJC). San Juan Co.: near Ironton, 21-31 July 1899 Curtis s.n.
(m; NY); Ruby Pass, 12,700 ft, 6 Jul 1996, Hogan 3023b (COLO); Stony Basin, 12,500 ft, 29 Jul 1983, Rottman 1753 (COLO). San Miguel
o.: near Trout Lake, 12,000 ft, 21 Aug 1924, Payson & Payson 4192 (RM). NEVADA. Clark Co.: Charleston Peak, 3,400 m, 8 Aug 1935,
Clokey 5460 (m; NY, UTC). NEW MEXICO. Bernalillo Co.: Kiwanis Point, Sandia Crest, 15 mi NE of Albuquerque, 10,000-10,200
ft, 28 Jul 1964, Baad 1035 (MICH), 23 Jun 1976, Wagner 2344 (MO), 22 Jul 1999, Sivinski 4997 (RM, UNM) (see Hartman and Sivinski
2006). UTAH. Beaver Co.: Tushar Mts., Dugway above Big Jim Flat, 3,050-3,150 m, 27 Jul 1991, Atwood 16176 oe Daggett Pa
“Gates of Birch Creek," 8 mi WSW of McKinnon, Wyoming, 9,600—9,700 ft, 30 Aug 1984, Tuhy 2176 (UTC)
[T prm A Dahal „Mi + +l (C ITI ) 1229
Fork Rock Creek, 14 mi N of Tabonia, 10,700 e o ae. pera & Jepson 15904 (BRY, RM); S rim of S Fork Rock Creek, TZN R8W
Sec. 24, 10,700 ft, 15 Aug 1995, Huber & 2910 Boulder Mt., 9,800 ft, 17 Aug 1995, Welsh & Atwood 26242 (BRY, COLO).
Garfield Co.: T31S R3E Sec. 36, 10 Sep 1942, Ellison 942- 13 (USFS); Clinetop Mesa, 10,300—10,500 ft, 24 Jun 1990, Hartman 25938
(RM); Boulde: Mt. nae Creek pi 10,600—10,700 ft, 6 Aug 1996, Huber 3415 (BRY); 2 mi N of Posey Lake, Aquarius Plateau,
10,000 ft, 29 Jun Maguire 20 : cited Madrono 8:262. Poe Iron Co.: 4 mi E Brian Head Peak, 11,000 e 23 Jun 1940,
pn pcc (m; NY, UTC): it of S 1 Left Hand Canyon, M Pl 3,100 m, 9 Aug 1986, N. 17 ; BRY). Juab
Mt. Nebo, 10,500-11,500 o 1980, Collins & Harper 864 (BRY). Kane Co.: Spruce Camp, Navajo Lake, 15 Jul 1940, ir
19541 (UTC). San P field Canyon, 10,928 ft, 8 Aug 1940, Maguire 19988 (m; NY, UTC, cited Madrorio 8:262. 1946);
T205 R4E Sec. 19, 10,000 ft, 22 Aug 1945, Ellison 4533 den USFS); Mora Mt., 11,000 ft, 24 Jun 1977, Lewis 4760 (BRY). Uinta Co.:
Dyer Mine, 5 Jul 1902, Goodding 1256 (RM). WYOMING. Sublette Co.: Palmer Peak, 10,600-11,400 ft, 5 Aug 1994, Hartman 40378
a oss d oes 10,400—11,180 ft, 2 Aug 1994, Hartman 49240 oe Teton Co.: Moose Basin divide, 9,140 ft, 22 Aug 2007, Scott
ide, 9,700 ft, 4 Aug 2004, Varga et al. s.n. (Grand Teton National Park).
GEOGRAPHIC RANGE, ECOLOGY, AND PHENOLOGY
Prior to Rabeler et al. (2005) and Hartman and Sivinski (2006), the published geographical range of M.
macrantha s.str. was Colorado while M. filiorium was credited to the southern portions of Colorado, Utah,
and Nevada (Maguire 1946, 1958). Based on recent collections and annotations, M. macrantha s.l. is now
known to occur throughout western Colorado, from northeast to southwest Utah, at one location in south-
ern Nevada, and most recently in restricted areas of central Arizona, central New Mexico, and northwest
ee SEE PN citations and Fig. 1).
lyf found on l t bstrat Iti gl few labels indicate volcanics.
Habitats vary from Festuca idahoensis Elmer meadows to parklands and other open areas often associated with
Picea engelmannii Parry ex Engelm., Abies bifolia A. Murray bis, Pseudotsuga menziesii (Mirb.) Franco, Populus
tremuloides Michx., and Pinus contorta Douglas ex Loudon var. latifolia Engelm. With increase in elevation,
it is found in krummholtz and alpine meadows, ridges, and scree and talus slopes. The elevational range is
2100-3700 m.
Flowering extends from early June into S ber; fruit tseen until late July to September.
Y
PHYLOGENETIC HYPOTHESES
The relationships between Minuartia (Arenaria) macrantha s.str., M. filiorum, M. rubella, and the M. rossii-
stricta complex (Wolf et al. 1979; McNeill 1962) have been in dispute.
Relationship of Minuartia macrantha to Minuartia rubella (sect. Tryphane)
In preparation for the flora of Utah, Welsh (1993) transferred Arenaria filiorum to a variety of A. rubella. He
distinguished var. filiorum from var. rubella by herbage (glabrous vs. typically glandular), sepal length (3.5-6.7
mm. vs. 2.8-4.5 mm), and seed length (0.7-1 mm. vs. 0.4-0.6 mm, respectively). Comparable character states
(Welsh et al. 2003) for A. macrantha are: herbage glabrous, sepals 4-5 mm long, seeds 0.7-1 mm long.
Under M. macrantha, Welsh et al. (1993) provided the following comment: “The plant, long obscured
within the mass of A. rubella in Utah collections, is still poorly known, and might represent nothing more
than an extreme phase of that species. It stands mainly on the feature of petals substantially surpassing the
sepals, which are longer than for A. rubella var. rubella, but not for A. rubella var. filiorum.
In contrast to Welsh's quotes, Minuartia macrantha s.l. and M. rubella are quite distinct; the two species
are easily distinguished in the field without magnification. The former is consistently glabrous while M.
rubella is invariably and prominently stipitate-glandular throughout the vegetative structure (see Rabeler et
al. 2005). Furthermore, the two species differ in leaf morphology (1-nerved, fleshy vs. 3-nerved, not fleshy)
and seed size (0.7—1 mm vs. 0.4-0.7 mm, respectively).
The relationship between M. macrantha s.l. and M. rubella, is distant at best (see below). Minuartia
macrantha is restricted to the western United States while the latter taxon is circumboreal extending south
into western North America. Maguire (1958) considered Arenaria macrantha s.str. and A. filiorum to be "the
southern complement of A. rossii” and did not include Minuartia (Arenaria) rubella in this complex. In fact,
1230 Journal of the Botanical R h Institute of Texas 2(2)
M. rubella traditionally has been placed in the highly polymorphic Minuartia section Tryphane (Fenzl) Hayek
(3-10 taxa), and is the sole member found in North America (McNeill 1962). The lectotype for the section
Tryphane is M. verna (L.) Hiern (chromosomal base x = 12, see below), with which Mi tia rubella has often
been confused.
Relationship of Minuartia macrantha with sects. Sclerophylla versus Alsinanthe
Two other relevant sections of Minuartia recognized by McNeill are Sclerophylla Mattf. (lectotype, M. michauxii
dus Mattf.) and Alsinanthe (Fenzl) Graebn. (lectotype, M. stricta (Sw.) Hiern). Section Sclerophylla is quite
f seven species. Two of these taxa were not seen (=?) by McNeill: ?Arenaria filiorum
and 2A, macrantha. Consequently, he did not make the combination Minuartia filiorum at that time. In our
opinion, neither of these taxa belong in this section, one characterized in part by “leaves rigid, recurved
and very thick median nerve.”
Weber and Wittmann (2000) recognize Tryphane rubella (Wahlenb.) Rchb. on one hand and Alsinanthe
macrantha and A. stricta (Sw.) Rchb. on the other. This generic realignment was predicated in part on differ-
ent chromosomal base numbers: x = 12 for Tryphane (Hartman 1971; Love and Love 1975a and x = 15 for
Alsinanthe (Love & Lóve 1975a). Although there are no known chromosome counts of M. macrantha, one with
a base number of x = 15 would be consistent with Maguire’s notion of M. macrantha being closely related to
M. rossii. Wolf et al. (1979) reported counts based on x = 15 for the three members of the M. rossii complex,
a base number also documented for M. stricta (Love & Love 1975b). Recent molecular investigations of the
Caryophyllaceae have included Minuartia (Nepokroeff et al. 2001, Fior et al. 2006) and both suggested that
Minuartia is polyphyletic. Nepokroeff et al. (2001 and pers. comm.) found M. rubella and M. rossii appearing
in unrelated clades; unfortunately, M. macrantha was not sequenced.
Distribution of the Minuartia macrantha
Previously, comments were made as to a possible affinity of Minuartia macrantha s.l. to the M. rossii complex.
Wolf et al. (1979) recognized three species in this alliance: M. rossii, M. elegans (Cham. and Schltdl.) Schis-
chk., and M. austromontana Wolf and Packer. The last was a substitute name for Arenaria rossii var. apetala
(Maguire non Fenzl; see Hartman and Nelson 1998, p. 57). The first two species are largely high Arctic in
distribution whereas M. austromontana extends south along the spine of the Rocky Mountains from central
Alberta, Canada, through Montana and west-central Wyoming, with a disjunct population in the Wallowa
Mountains, Oregon, U.S.A. (Cusick 2299, in part, GH!; Peck 22576, WILLU! at OSC). One geographic outlier
from the Uinta Mountains, Utah (Harrison & Harrison 10926, BRY!) cited by Wolf et al. (1979) represents a
specimen of M. macrantha. In Wyoming, four historical records (pre-1960, RM, USFS; Wolf et al. 1979) of
M. austromontana were known from the Beartooth Plateau, the Teton Range, and the northern Wind River
Range. Based on recent collections (1978-2007, RM), 41 additional sites have been documented from the
Absaroka, Gros Ventre, and Teton Ranges and the Big Horn Mountains. By contrast, the southern Rocky
Mountain M. macrantha is now documented from Wyoming; an apparent disjunct from the northeastern
Utah or northern Colorado. It is known from three sites (1994-2007, RM) from the Teton Range and the
Gros Ventre Mountains. Other state records include Arizona and New Mexico.
Both M. austromontana and M. macrantha are sympatric in the Gros Ventre Range of northwestern
Wyoming with no hint of hybridization. Their differences in gestalts and detailed morphology allow for
easy separation (see Rabeler et al. 2005; M. rossii, M. elegans, M. austromontana, and M. macrantha appear
near one another in the key to species of Minuartia, couplets 37 to 42).
Distribution of Minuartia stricta relative to Minuartia macrantha in Colorado
Minuartia stricta (Sw.) Hiern also occurs in Colorado and is disjunct in California as well from its normal
range that extends from Alaska, across northern Canada, to Greenland and Eurasia (Rabeler et al. 2005,
Nannfeldt 1954). It is here mentioned as it is frequently confused with M. macrantha. The key to species of
Minuartia (Rabeler et al. 2005) is terminated by M. stricta and M. dawsonensis (Britton) House (couplet 43).
Minuartia macrantha and M. stricta differ as follows: sepals 3.5-5 mm vs. (1.5-)2.53.2 mm long in flower,
eye I! Dahal „Mi "3 "n (C phyll ) 1231
to 5.5 mm vs. 4 mm long in fruit (Fig. 2); seeds 0.4—0.6 mm vs. 0.7-1 mm long, respectively; and subtle
leaf and habit differences. The documented distributions of M. macrantha and M. stricta in Colorado, where
they are known to co-occur, are shown on Fig. 1
Specimens of Minuartia stricta in the southern Rocky Mountains are listed below (based on holdings
at COLO and RM):
COLORADO. Boulder C I 11 l llins Pass, 11,300 ft, 18 Jul 1969, Brown 38 (COLO). Garfield Co.: E side of
Heart Lake, 10,600 ft, 26 Jun 1963, Miller 300 (COLO). Gunnison Co.: North Italian Mountain, 13,300 ft, 1 Aug 1949. Langenheim 455
(COLO). Hinsdale Co.: Mesa Seco, above Spring Pass, 12,400 ft, 31 Jul 1964, Weber 12167 (COLO). Huerfano Co.: 0.5 mi W of Iron
Nipple, 12,000 ft, 17 Aug 1999, Elliott 11048a (RM). Park Co.: Sheridan Mountain, above E. Mine, 13,790 ft, 12 Jul 1967, Weber
13303 (COLO); Hoosier Ridge, east of pass, 12,500 ft, 8 Jul 1951, Weber et al. 6551 (COLO). Summit Co.: Hoosier Ridge, 12,000 ft, 10
Jul 1959, Hultén & Weber 11054a (COLO).
ACKNOWLEDGMENTS
We are grateful to the following institutions for loans of specimens: ASU, BRY, COLO, GH, MICH, MO, NY,
OSC, SJC, UC, UNM, and UTC. We acknowledge Jennifer Ackerfeld for assistance with photomicroscopy;
Klara Varga and David Scott for access to recent collections from Grand Teton National Park.
REFERENCES
Fior S., PO. Karis, G. Casazza, L. MINUTO, AND F. Sata. 2006. Molecular phylogeny of the Caryophyllaceae (Caryophyl-
lales) inferred from chloroplast matK and nuclear rDNA ITS sequences. Amer. J. Bot. 93.399-411.
Hartman, R.L. 1971. Chromosome numbers in Caryophyllaceae from Wyoming and adjacent states. Bull. Torrey
Bot. Club 98:276-280.
HARTMAN, R.L. AND B.E. NeLson, 1998. Taxonomic novelties from North America north of Mexico: A 20-year vascular
plant diversity baseline. Monogr. Syst. Bot. Missouri Bot. Gard. 67:1—59.
HARTMAN, R.L. AND R. Sivinski. 2006. Plant distribution reports: Minuartia macrantha. New Mexico Bot. 35:3.
Love, A. AND D. Love. 1975a. Nomenclatural notes on Arctic plants. Bot. Not. 128:497-523.
Love, A. AND D. Love. 1975b. In: IOPB chromosome number reports XLIX. Taxon 24:501-516,
McNeLL J. 1962. Taxonomic studies in the Alsinoideae: |. Generic and infrageneric groups. Notes Roy. Bot. Gard.
Edinburgh 24:79-155
Macur, B. 1946. Studies in the Caryophyllaceae— ll. Arenaria nuttallii and Arenaria filiorum, section Alsine.
Madroño 8:258-263.
Macuire, B. 1958. Arenaria rossii and some of its relatives in America. Rhodora 60:44-53.
NANNFELOT, J.A. 1954, Some notes on Minuartia stricta (Sw.) Hiern and allied species. Nytt Mag. Bot. 3:159-170.
NEPOKROEFE, M. Er AL. 2001. Origin of the Hawaiian subfam. Alsinoideae and preliminary relationships in Caryophyl-
laceae inferred from matK and trn1. C-F sequence data. In: Botanical Society of America. 2001. Botany 2001
Abstracts. Columbus, OH.
RaBELER, R.K., R.L. HARTMAN, AND F.H. Utech. 2005. 24. Minuartia. in: Flora of North America Editorial Committee, eds.
Flora of North America North of Mexico. Oxford University Press, New York and Oxford. 5:116-136
WEBER, W.A. AND R.C. WitTMANN. 1992. Catalog of the Colorado flora: a Biodiversity Baseline. Univ. Press Colorado.
WEBER, W.A. AND R.C. WitTMANN. 2000. Catalog of the Colorado flora: a Biodiversity Baseline. Electronic ion revised
March 11, 2000 (http;//cumuseum.colorado.edu/Research/Botany/Databases/catalog.html).
WELSH, S.L. 1993. New taxa and new nomenclatural combinations in the Utah flora. Rhodora 95:392-421.
WeLsH, S.L., N.D. Arwoop, S. GoopaicH, AND L.C. Hiaains (eds.). 2003. A Utah flora. ed. 3. Brigham Young University,
Provo, Utah.
Warr, S.J., J.G. Packer, AND K.E. Denrorp. 1979. The taxonomy of Minuartia rossii (Caryophyllaceae). Canad. J. Bot.
57:1673-1686.
—
1232 Journal of the Botanical R h institute of Texas 2(2)
BOOK REVIEW
RONALD M. Lanner. 2007. The Bristlecone Book: A Natural History of the World's Oldest Trees. (ISBN
0-87842-538-1, pbk.). Mountain Press Publishing Company, PO. Box 2399, Missoula, Montana 59806,
U.S.A. (Orders: www.mountain-press.com, 800-234-5308). $12.00, 118 pp., color illustrations, 6" x 9".
Ronald M.Lanner is a native of Brooklyn, New York, whose teenage interest in natural history led him to n ne a DK He was the
editor of the Western Journal of Li foren for its first eleven yes Now retired, he no with his Il ponderosa
pines of Placerville, California, wh g ics. It was at tthe Institute where
he was employed early in his career, and he developed his lifel dmiration fi
his coniferous trees. His major research interests
1 I and the ecological and evolutionary
have included natural hybridization in dl effects of aging on trees, bud d
effects e mutualisms of birds and p
ost recent book is The e Book: A a ndi of idi a s Oldest Trees e api was pe ii pain
j| Caria (1999), Made For Each Other: A Symbiosis of Birds and Pines (1996), Autum n
990), Trees of the Great Basin: A Natural History (1984), and The Piñon Pine: A Natural and Cultural History (1981).
Professor Lanner has written ne T n history of the bri d foxtail pi f the western United Pese trees that
thrive in a harsh f years. Th PESehasl 1 1: oe)
right and | gs togetl he fi ptivating f 1 ] d foxtail pines, with a Special an on Great Basin
Tu ,th ld rft th
I have eal ow pes wonderful trees, eae arbon dating and * tree ring a curve”, but I
admit that I did 1 How old can really be? How
do they deal with soil erosion in the cold, windswent peaks of America's Mountain West What ae are ly selected to
prolong life?
In the fi I lescribes tl i Basin Bristl Pines, Rocky ee Bristlecone Pines, mu un
um!
Pines. In tl } , he di he pi d describes what they are made of. Hi
text. E Lin chapter is devoted to ie ii nd of n pne and the fourth chapter is aerated to various shapes and forms. He
g, forest fires, bark beetles, and White Pine Blister Rust. The
final chapter is devoted to a discussion of wnat ciales hese] trees unique: their great age, including recorded ages of living and dead
tre en the secret of longevi
e is an excellent pU a very helpful glossary, and an index. This book will make a good addition to any public or
E: pid —Gary L. Jennings, Botanical Research Institute of Texas, 500 East 4th Street, Fort Worth, Texas 76102-4025, U.S.A.
J. Bot. Res. Inst. Texas 2(2): 1232. 2008
CORRECT BASIONYM AND COMBINATION FOR
PAPPOSTIPA BARRANCAENSIS (POACEAE)
Konstantyn Romaschenko
M.G. Kholodny
Institute of Botany
National Academy of Sciences of Ukraine
01601 Kiev, UKRAINE
kromaschenko@gmail.com
In the recent paper (Romaschenko et al. 2008: 184), the incorrect basionym was cited for the combination
Pappostipa barrancaensis making the new combination invalid. The new combination is validated here with
the correct basionym.
Pappostipa barrancaensis (FA. Roig) Romaschenko, comb. nov. Basion: Stipa barrancaensis EA. Roig, Revista Fac.
Ci. Agrar. Univ. Nac. CO nain 46, t. 9, 26c. 1964 [1966]. Tree: ARGENTINA: Menpoza: Dpto. Malalhue, entre Ranquil Norte y
Coipo Lauquen, di acia el norte, 20 Dec 1960, A. Ruiz Leal 21547 (HoLoryrE: MEN).
REFERENCE
ROMASCHENKO, K., PM. PETERSON, R.J. SORENG, N. GARCIA-JACAS, O. FUTORNA, AND A. E 2008. Molecular phylogenetic
analysis of the American Stipeae (Poaceae) resolves J polyphyletic: evidence for a new genus,
Pappostipa. J. Bot. Res. Inst. Texas 2:165-192.
J. Bot. Res. Inst. Texas 2(2): 1233. 2008
1234 Journal of the Botanical R h Institute of Texas 2(2)
BOOK REVIEW
ANGELA Leiva, photographs by GonzaLo Canett1. 2007. Trees of Cuba. (ISBN-10: 1405029056, ISBN-13: 978-
1405029056, pbk.). MacMillan Caribbean Publishers, Between Towns Road, Oxford, OX4 3PP, United
Kingdom. (Orders: www.macmillan-caribbean.com, caribben@macmillan.com, +44 (0)1865-405843
fax). $10.70, 112 pp., color p 8 1/2" x 5 1/4".
This book has t ] f ] yut for anyone wishing to identify tl ly seen trees in Cuba, whether in
the city, beside the iUad. the beach or in other p fthe island. There are well 800 sp f growi he island, thus
this vol p y ple of ti 1 ] variety to be found in Cuba.
ing all of th t i d f the | known varieties, nearly one hundred species are covered in this
Ede pu species has a full page devoted to it Men at least one e color photograph. Details included are flowers, seeds, and bark.
y uses and g p I
This guide will be especially useful to the g l reader. A balanced ge of both nati d cultivated speci h it
usefulness as a per a It includes a bibliograpl lind f both d scientifi li ded for
anyone ith Cuba and the Caribbean region
Ein Leiva is a Professor of the University of Havana and is the Director of the n Botanical Ena A is in
| i fth Cuban plants Gonzalo C
publicite photography. He is a photo-reporter for the Cuban Ministry of Science, iaa and the Environment.—Gary L. Jennings,
Botanical Research Institute of Texas, 500 East 4th Street, Fort Worth, Texas 76102-4025, U.S.A.
J. Bot. Res. Inst. Texas 2(2): 1234. 2008
LOS ENCINOS (QUERCUS: FAGACEAE) DE COAHUILA, MEXICO
José A. Villarreal Q., Juan A. Encina D. y Miguel A. Carranza P.
Departamento de Botánica
bg AVE Anrnri Ant, i
Buenavista
25315 Saltillo, Coahuila, MEXICO
RESUMEN
E Jin da] b dus O ] do de Coahuila. La ri lafl d i incl 32 especies
I I e r L3
Se incluye 1 lave para separar las especies, d ipción de cad de ellas, datos de distribución, datos ecológicos e PES
ABSTRACT
A fel A. f 1 f 1 il s l Th A p TI 1 x 1 £1 = 1 J
: g I 32 species. À key to npe
cies, t ic d if ti 1 inf i g li Eg di il i d l g is included. Each E 11 illustrated.
INTRODUCCION
El género Quercus, al cual pertenecen los encinos y robles, es el más numeroso de la familia Fagaceae, sus
representantes son árboles y arbustos, siendo el componente principal de los bosques de encino y de pino-
encino, de las regiones montañosas de México. Los encinos presentan una gran diversidad taxonómica,
variación morfológica y distribución con un rango ecológico amplio. Para Méxi ti alrededor de 150
especies, con centros de diversidad en la regiones montañosas del centro y este del país (Nixon 1993b).
Los encinos usualmente presentan un habito de crecimiento arbustivo en regiones con climas xéricos
donde pueden llegar a formar matorrales densos, mientras que en condiciones mésicas, crecen especies
árboreas, que integran comunidades boscosas.
Coahuila es el tercer estado más grande de la república Mexicana, se ubica en la parte media de la
región septentrional del país. En su territorio se localiza la parte terminal de la Sierra Madre Oriental y su
continuación en una serie de montañas aisladas con distribución en el centro y norte del estado. Aunque
el clima árido es característico de Coahuila, en las zonas montañosas se presenta un clima templado y sub-
húmedo que permite el crecimiento de comunidades de encinos (Encina y Villarreal 2002).
La flora de encinos del estado está integrada por 32 especies, teniendo la mayor diversid
del noroeste y las del sureste. La riqueza más alta, con 16 especies, se concentra en la Sierra del Carmen.
Seis especies restringen su distribución al área del estado y regiones circunvecinas (Quercus carmenensis,
Q. coahuilensis, Q. gravesii, Q. hintoniorum, Q. invaginata y Q. saltillensis). Un análisis previo sobre la flora de
encinos en el estado de Coahuila llevado a cabo por Encina y Villarreal (2002) en el estado de Coahuila,
muestra que las sierras de la región centro están mas relacionadas con las del noroeste que con las del sur-
d en las sierras
este.
Los encinos son un grupo de plantas fáciles de colectar y ubicar como género por su hábito arbóreo
o arbustivo, hojas simples y alternas y el fruto tipo bellota. Sin embargo su taxonomía no es sencilla, entre
las dificultades que presenta están un número alto de especies, la variación morfológica dentro de la misma
especie, su variación ecológica y la hibridación, por lo que su manejo requiere de atención especial. Las
descripciones de las especies y la clave fueron elaborados con base en materiales de herbario y bibliografía.
Los datos de distribución general y local, de referencias bibliográficas y colectas, y la información ecológica
de datos tomados en campo. Los dibujos iuda MR con pese en Hia disponible de herbario. Este
trabajo presenta una revisión taxonómica d para Coahuila, descripciones,
datos ecológicos e ilustraciones de cada una de Be
J. Bot. Res. Inst. Texas 2(2): 1235 — 1278. 2008
1236 J | of the Botanical R h Institute of Texas 2(2)
MATERIALES Y METODOS
El presente trabajo es el resultado de la revision de informacion sobre el género poe para el estado de
Coahuila. Se colectó material botánico durante los años 2000 al 2006 en las y del estado para
enriquecer la colección de material de referencia y se consultó las colecciones de los herbarios ANSM, CFNL
y TEX. El criterio taxónomico que se siguió es en parte el de Muller (en prensa) y el de Nixon (1993a y b).
Las descripciones de las especies y la clave fueron elaborádos en base a materiales de herbario y bib-
liografía. Los datos de distribución general y local de referencias bibliográficas y colectas, y la información
ecológica de datos tomados en campo. Los dibujos son obra de Miguel Agustín Carranza Pérez, para su
elaboración generalmente se emplearon varios ejemplares del material disponible del herbario ANSM.
Se presenta una descripción de las especies, donde se menciona en primer término el nombre científico,
los sinónimos se escriben entre paréntesis seguido de la descripción de cada especie. Se incluye además su
distribución a nivel general y para el estado; así como información sobre los tipos de vegetación donde se
presentan, las especies asociadas, rangos altitudinales, e información general sobre su abundancia.
Generalidades del área de estudio
El estado de Coahuila se ubica en el norte de la republica Mexicana, entre los 24°32'13 y los 29°52'47” de
latitud norte, y entre los 99°50'30" y los 103°57'03" de longitud oeste (mapa).
Clima.—Por su ubicación el estado se caracteriza por presentar climas continentales secos, que van
desde los semicálidos, hasta templados en las partes altas. La precipitación es escasa, de 300 a 500 mm,
para todo el estado. La temperatura es extremosa, la humedad atmosférica baja y la evapotranspiración es
elevada. El clima es extremoso, con veranos muy calurosos e inviernos fríos. La parte nororiental tiene los
climas más húmedos y cálidos, el occidente es muy seco, mientras que el centro y sur tienen climas desde
secos y templados hasta templados subhúmedos en las cumbres de las sierras.
Geología y Suelos. —Los sedimentos marinos continentales del paleozoico al cuaternario forman en su
mayoría el territorio coahuilense. 1 calizas del mesozoico son las más típicas, los plegamientos tienen
dirección Este-Oeste en el sur y Noreste-Sureste en el centro y del estado. Existen afloramientos de rocas
ígneas del triásico al cuaternario. Rocas metamórficas paleozoicas afloran en áreas pequeñas y dispersas.
Los suelos que se presentan con mayor frecuencia son Xerosoles y Yermosoles, que son suelos localiza-
dos en zonas con climas áridos. Los Litosoles y Regosoles se presentan en áreas montañosas, son de origen
calcáreo y se mezclan con Rendzinas, con mayor contenido de materia orgánica (Anónimo 1983).
Fisiografía.—En el estado se reconocen las siguientes provincias fisiográficas (Villarreal et al. 1996):
1) Provincia de la Planicie Costera del Golfo que ocupa el noreste del estado. Esta formada por una
serie de llanuras y lomeríos bajos con declives suaves con altitudes que van de los 550 m a los 2200 m. La
vegetación que caracteriza a esta región es el Matorral Tamaulipeco.
2) Provincia de la Sierra Madre Oriental, formada por la terminación en el sureste del estado de la
cordillera de la Sierra Madre, proveniente del centro del país, continuándose hasta el noroeste del estado a
través de una serie de sierras aisladas. Las partes más altas alcanzan los 3400 m y la vegetación presente es
el Matorral Submontano en las partes bajas y exposiciones xéricas, y bosque en las partes mésicas.
3) Provincia del Desierto Chihuahuense, formada por valles, bolsones y lagunas ubicadas en el oeste
y centro de estado, en la región del Altiplano Mexicano. Es la región más árida y las altitudes van desde los
800 a los 1900 m. La vegetación característica son los diferentes tipos de Matorral Desértico.
Vegetación.—De on seis E de vegetación citádos para Coahuila por Villarreal y Valdés (1995), los en-
cinos están [ el Matorral Submontano y el Bosque de Montaña, y ocasionalmente
en el Matorral "Donau pido y la Vegetación Riparia.
Matorral Submontano.—Este tipo de vegetación ocupa cerca del cinco por ciento de la superficie del
estado. Esta formado por arbustos y árboles pequeños que crecen en laderas y cañones con condiciones
de alta humedad. Este tipo de comunidad puede presentarse en forma tanto dispersa como muy densa,
mezclándose con elementos del Matorral Desértico en las partes bajas de las sierras y extendiéndose hasta
Villarreal et al., Q
1237
104° 103 102° 101° 100°
30°) 30°
29°7 [29
28" F28g
27°] PIZ
26" r26°
25° r 25°
104° 103° 102° 101° 100°
Mapa | IP ER | dal J Jt ht y | g n
1. Sierra de Arteaga 9. Sierra de La Fragua 17. Sierra de Santa Rosa
2. Sierra de Zapalinamé 10. Sierra de La Gloria 18. Sierra Encantada
3. Sierra de La Concordia 11. Sierra Pájaros Azules 19. Sierra del Pino
4. Sierra de Parras 12. Sierra Delicias 20. Sierra Cruc
5. Sierra de Jimulco 13. Sierra de La Menchaca 21. Sierra del Cármen
6. Sierra de La Paila 14. Sierra de La Madera 22. Serranía del Burro
7. Sierra de La Gavia 15. Sierra del Fuste 23. Planicie Costera del Golfo
8. Sierra de San Marcos 16. Sierra Mojada
1238 Journal of the Botanical R h Institute of Texas 2(2)
el Bosque de Montaña en las porciones más altas. Es frecuente encontrar Quercus intricata, Q. invaginata,
Q. pringlei, Q. pungens, Cercocarpus montanus, C. ae vee m pti Ptelea trifoliata, Rhus
l Ungnadia speciosa,
virens, R. microphylla, Berberis trifoliata, Cali
Colubrina greggii, Mimosa biuncifera, Brahea iman Dassin an M y Peedi texana.
Bosque de Montafia.—Este tipo de vegetación presenta las siguientes comunidades:
1) Bosque de Encino. En valles intermontanos y cañones de las sierras, donde el clima es templado y
subhümedo. Usualmente sigue al Matorral Sul tano en altitudes de 1200 a los 1800 m, y ocupa menos
del 196 del estado. Esta dominado por árboles en poblaciones esparcidas o densas, muy homogéneas, donde
es frecuente Quercus gravesii, Q. grisea, Q. laceyi, Pinus cembroides, P. arizonica, Garrya ovata, Juniperus flaccida,
Arbutus xalapensis, Fraxinus cuspidata y Prunus serotina.
2) Bosque de Pino. Este tipo de comunidad dominada por especies del género Pinus, ocupa cerca del
296 de la superficie. Es frecuente en laderas y partes altas de sierras con climas templados y subhümedos. El
mas comün es el Bosque de Pino Pirionero, donde las especies de Pinus cembroides, P. remota y P. pinceana son
dominantes. Otras especies asociadas son Juniperus saltillensis, J. erythrocarpa, Quercus intricata, Q. striatula,
Q. pringlei, Rhus virens, Yucca carnerosana, Tillandsia recurvata y Bouteloua gracilis. Otras comunidades están
dominadas por Pinus arizonica en las sierras del centro y norte del estado, mientas que en la Sierra Madre
Oriental se presenta Pinus pseudostrobus, P. greggii, P. hartwegii, P strobiformis y P. teocote.
3) Bosque de Oyamel. Localizado en las partes altas de las sierras, en altitudes que van de los 2300 a
los 3000 m. Ocupa el 2% del área y está dominado por Pseudotsuga menziesii y Abies vejarii. Otras especies
frecuentes son Quercus saltillensis, Q. greggii, Arctostaphylos pungens, Pinus strobiformis, Cupressus arizónica y
Salix lasiolepis.
4) Vegetación Subalpina. Se presenta en las cimas de la Sierra de Arteaga, cubriendo menos del 1% del
estado. La comunidad está dominada por arbustos y herbá donde Pinus culminicola, P. hartwegii, Quercus
hintoniorum y Agave montana son frecuentes.
Matorral Tamaulipeco.—Se distribuye en planicies y lomeríos del noreste del estado, ocupa cerca del
20 96 del área, en lugares de los 240 a los 850 m. Esta dominado por Acacia rigidula, Leucophyllum frutescens
y Prosopis glandulosa. En lugares húmedos como cañadas u orilla de arroyos o donde el manto acuífero es
superficial, se presenta Quercus fusiformis en forma aislada o en comunidades densas llamadas localmente
“motas.”
Vegetación Riparia.—Restringida a los pocos ríos y arroyos que existen en el estado tributarios del
Río Bravo. Estas comunidades crecen en los márgenes de corrientes de agua, donde la humedad permite
su desarrollo. Las especies mas frecuentes son Taxodium mucronatum, Platanus glabrata, Salix nigra, Carya
illinoensis, Acacia farnesiana y ocasionalmente Quercus fusiformis.
TRATAMIENTO TAXONOMICO (FAGACEAE)
Árboles, algunas veces arbustos. He) as simples, a tenes, con venación pace, de enteras a lobuladas, gen-
eralmente con estipulas caedizas. F t pigas; perianto
de 4—7 lóbulos; flores masculinas con el mismo o doble número de estambres que las piezas del perianto,
aunque a veces pueden llegar a 40; flores f inas con un perianto de 4-8 lóbulos, pistilo tricarpelar, cada
una protegida por un involucro basal. Fruto una nuez o aquenio agrandado, en grupos de 13, protegidos
en la base o envueltos completamente por una cúpula de escamas o espinas.
Comprende 9 géneros y unas 800 especies, en su mayor parte originarias de zonas templadas y sub-
tropicales del Hemisferio Boreal (Nixon 1997). Proporcionan algunas de las maderas más importantes del
mundo, con múltiples aplicaciones, además de producir corcho, taninos, frutos comestibles y plantas de
interés ornamental.
Quercus L., Sp. Pl. ed. 2, 2:994. 1753. Esrece Tiro: Quercus robur L.
Arboles o arbustos monoicos; ramillas usualmente acanaladas; yemas cubiertas por escamas ciliadas; es-
típulas deciduas o raramente persistentes; hojas alternas, persistentes o deciduas, pecioladas, limbos con
Villarreal et al., Q (Fag ) de Coahuil 1239
bordes enteros, aserrados, dentados o lobulados. Flores masculinas en amentos colgantes, cada flor con un
perianto de 4—7 lóbulos y 6 a 12 estambres, con anteras y fil delgados; las femeninas solitarias
o en grupos pequeños, cada una en el interior de un involucro constituido por numerosas escamas, pistilo
de tres cárpelos que forman un ovario trilocular, cada lóculo con dos óvulos y tres estilos libres. Fruto una
bellota, conteniendo una nuez (aquenio agrandado), una semilla viable, los otros cinco óvulos restantes son
abortados y se adhieren a la semilla desarrollada, rodeada en la base por una cúpula formada por escamas
imbricadas, maduración anual o bienal.
El género está formado por unas 400 especies (Nixon 1997), en su mayoría originarias de las zonas
templadas del hemisferio norte, con algunas presentes en el sureste de Asia y solo una especie en el norte
de Sudamerica (Q. humboldtii en Colombia). En México son abundantes principalmente en las áreas mon-
tañosas, la mayor diversidad se presenta en el centro y sur del territorio, siendo el país con el mayor número
de especies. En Coahuila se estima que existen unas 32 especies ingidas a las áreas montañosas y unas
cuantas en las planicies bajas del noreste del estado.
Se les conoce con el nombre de encino, roble, ahuatl, y en ingles como oak. Al parecer en Coahuila se
usa en general el nombre de “encino” para reconocer a las especies de Quercus, a excepción de los encinos
arbustivos que se les conoce como “encinos chaparros” o “charrasquillos.” Los principales caracteres para
el reconocimiento de las especies están basados en las hojas. La forma, tamaño, borde, color, indumento y
venación, entre otras, son mi más LM en su diferenciación. npe especies, suelen ser Aen. en
la xs cl F1 T 1 J
3 1 morfológic a variar
dentro de la misma especie según sean Pues jóvenes o Aulos: o incluso dentro de ln misma planta
dependiendo de las partes donde se presenten. Otros carcteres importantes en la taxonomía son el tipo de
indumento en la hoja, que puede variar en densidad y tipo de pelos que lo forma. También la morfología de
la bellota es importante, como el tamaño de la cúpula y nuez, además de la forma de las escamas.
I. Quercus sect. Quercus.
Árboles o arbustos. Corteza usualmente cara escamosa, gris o blancosa. Hojas con margen entero, den-
tado o lobado, dientes mucronados. Flores estamina das con cáliz de 2-6 lóbulos, anteras apiculadas. Flores
pistiladas con cáliz adnado al ovario y estilo alargado. Fruto anual, cupula con escamas con base engrosada
y nuez glabra a tomentulosa.
Quercus arizonica Quercus laceyi Quercus pungens var. pungens
Quercus carmenensis Quercus laeta Quercus pungens var. vaseyana
Quercus fusiformis Quercus mohriana Quercus rugosa
Quercus gambelii Quercus muehlenbergii Quercus sinuata
Quercus grisea Quercus oblongifolia Quercus sinuata var. breviloba
Quercus greggii Quercus polymorpha Quercus tuberculata
Quercus intricata Ouercus potosina
Quercus invaginata Quercus pringlei
II. Quercus sect. Lobatae, Hort. Brit. 385. 1830.
Árboles o arbustos. Corteza lisa a fisurada, café a negruzca. Hojas con margen entero, dentado o lobado,
dientes y lóbulos usualmente terminados en aristas. Flores estaminadas con cáliz de 2-6 lóbulos, anteras
usualmente retusas. Flores pistiladas con cáliz libre del ovario y estilo linear a espatulado. Fruto bianual,
raramente anual, cupula con escamas con base plana, algunas veces engrosada y nuez tomentosa.
Quercus coahuilensis Quercus fulva Quercus hypoleucoides Quercus saltillensis
Quercus canbyi Quercus gravesii Quercus hypoxantha Quercus sideroxyla
Quercus emoryi Quercus hintoniorum Quercus mexicana
1240 J | of the Botanical R h Institute of Texas 2(2)
CLAVE DE LAS ESPECIES DE QUERCUS DEL ESTADO DE COAHUILA
i b | f | la superficie del haz; margen cartilaginoso, engrosado,
moderado o bod revoluto.
2. Margen de la hoja con dientes aristados, ial te en la mitad distal de la hoja, si ntero enton
ces con el aes akado pa ae cala glauco c [e amarilento, AN
3. Envés con indumento prendible.
4. Hojas amllaniente elípticas a alee ihe envés con tomento amarillo-cafezusco de pelos
eae ae y de po EM Quercus fulva
tomento denso amarillo-cremo-
soa a blanquecino de pelos fasciculados, sin p simples
dd
5. Margen con t ntes en la mita s con ápice redondeado a obtuso; bas
cordada a redondeada: peciolos 3-10 mm de largo Quercus MESES
5. Margen pud re) sab ioe acne en Me mitad distal; limbos con el ápice frecuentement
agudo; baser 5-15 mm lar Quercus odere:
rgo
3. Envés con indumento lanoso-tomentoso, denso, fácilmente desprendible, o sin lo anterior entonces
la epidermis con pelos glandulares.
eeu ae a ve formado por Mou: de dd ae restringidos a las axilas de la
iado ercus hintoniorum
6. Indumento del enu: formado de pels R que iem totalmente la pan pelos
glandulares ausentes.
imbos comúnmente con 1-3(-4) dientes por m bos en el ültimo tercio de la hoja; indu-
Que
mento amarillo-blanquecino de pelos con radios e OS ercus coahuilensis
7. iia con 4-6 dientes en cada lado, de unos: a n E del limbo; indumento e amaril-
lento de pelos fasciculados no isses ercus hypoxantha
2. Margen entero o con dientes cortos terminando en mucrón, en especial hacia la mitad distal de la hoja;
az comúnmente verde oscuro y lustroso.
8. Limbos menos de 5 cm de largo, si son má d | envés es bl te tomentoso;
margen fuertemente revoluto, venación no fienemente reticulada.
9. Margen con 3-5 dientes en EUM ~ raras veces entero; indumento lanoso y rizado, café amaril-
lento; limbos obovados a elíptic Quercus greggii
9. Margen coma ences raras nc con 3-5 dientes en cada lado; indumento corto, denso,
amarillo uercus intricata
8. Limbos de 4-15(- 20) cm de largo, si son más cortos entonces el envés no es notablemente tomen-
toso; a p d venación fuertemente reticula
10. Lim
ampliamen ovados a elípticos, fuertemente ¢ cóncavos hacia el envés; epidermis del
SING Eon pelos glandulares rojizos: uercus rugosa
lados, pl pidermis sin pelos glandulares.
. Hojas de 4a 8 cm largo con el apice agud btuso; pecíolos hasta 1 cm de largo___ Quercus arizonica
licias de Hasta l6 emi de larg | ápi gost te obtuso; pecíolos de 1.5 a 3 cm de
largo Quercus polymorpha
l. nin con las nervaduras no fuertemente impresas en la superficie del haz; margen ligeramente revoluto
plano.
e Margen lobulado, con la escotadura muy profunda, cada lóbulo terminando en una arista flexible de
le largo
13. Limbos de las hojas a u Bbovae0> le 6-8 en ius ancho, típicamente con tres dientes en la
parte apical de La hoja y con dos a tres | a lado Quercus gravesii
13. ee de dui hojas Mende eos de 2-4 cm de ancho, ápice acuminado y tres a cinco
| Quercus canbyi
12. Margen entero o lobulado, si presenta lóbulos entonces con la escotadura no profunda y terminando
en un mucrón o arísta de hasta 2 mm de largo
14. Ápice agudo, algunas veces obtuso, terminando en una arista fácilmente quebradiza; envés con
Ur MEE M oad "n B DM . :
o con dientes aristados.
. Margen siempre entero; hojas lll Ovadas a avons es con ones de
aspecto punteado, esparcido p estipitados,
con radios retorcidos y algo enredados Quercus mexicana
. Margen con dientes aristados, algunas veces entero, plano a cartilaginosamente engrosado;
=
un
—
un
Villarreal et al., Q (F } de Coahuil
g ) 1241
a]
axilas de la nervadura media.
16. Limbos gruesos y coriáceos, lanceolados; haz verde claro; envés con pelos amarillos fas-
ciculados hasta e ganto
16. Limbos delgados y flexibles, la bl | lados; !
glabro Quercus saltillensis
Ápice nae a redondeado, si es agudo terminando en un mucrón; envés con indumento
fácilmente desprendible; margen algunas veces entero, frecuentemente con dientes notorios,
aniciilad S +
p t
17. E A ^ P I
uercus emoryi
A
o EOS oe ie
18. dentado, con los di i li te lobulado,
old a ramillas café rojizas
19. |
J + IE ri | i | |
endurecidos muy ásperas al tacto; limbo con el margen crispado Quercus pungens
var. pungens
19. Haz muy lustroso, con indumento esparcido, ligeramente aes al tacto; limbo con
el eid casi plano -
us pungens var. vaseyana
18. Mar , irregulares, si ebd márgenes ondu-
ladamente erispados entonces, los lóbulos y el ápice del limbo son usualmente redon-
deados; ramillas café grisáceo
20. Limbos delgados y n obovados u oblanceolados; haz verde oscuro y lustoso
Quercus sinuata var. breviloba
20. Limbos gruesos y rígidos, oblongos a elípticos; haz verde claro, poco lustroso
Quercus fusiformis
17. Envé ind de pelos fasciculados, a veces esparcidos dando el aspecto de glabro, o
crn nroaminant
arg
DI ee Tende m tomento Ta
21.
a joi ia d veces entero.
22. Hojas de 2- 6(- 8) cm largo; limbos lanceolad ta obl
tomento flocoso o con parches de puis. pronto glabresce
23. Limbos verde grisáceo opaco en ambas superficies, pm o con indumento
persistente cerca de la base de las nervaduras; ápice y dientes no mucronados;
nervaduras del envés muy prominentes y las del haz con las venillas muy reticu-
ladas y esclerotizadas formando un retículo blanquecino evidente
a
Quercus
oblongifolia
23 Limbos conh rel d ] ^ | alabra o con ind + |
k vCIüc o verde , CQ I!
niilar nani IH + J A z z
1 Iud pl r
. | J
P IcCUulCuliaud»tiu
esclerotizados, s sin sofa un eeu PAra evidente Quercus pringlei
loi si menores, enton
J Mo! ml Li
[EN ri 1 J x Lk
22. |
i
+
g glabro o con indumento inconspicuo o
solo en las nervaduras principales.
24. Margen con lóbulos redondeados oblongos y llegando cerca de la nervadura
media a sinuado; ápice redondea
25. Hojas glaucas o verde arnarillentas, glabras al madurar; ramillas maduras y
pecíolos glabrecentes, café rojizos; envés comúnmente glabro Quercus laceyi
25. noes verde ¢ oe: con ae esparcidos, PERNES ramillas madur ras y
3 f I t
Quercus gambelii
24, Margen dentado o EIE lobulado, si es entero o crenado entonces con el
ápic btu
26. noja T E reris s laterales cen y anastomosados hacia el bee
epando; Puis rojizos cerca de la base;
came dela copula engrosadas En la bas Quercus tuberculata
26. Hoj | marg
margen repando, dentado o con lóbulos cortos; pecíolos no rojizos cerca de
la base; escamas de la cúpula casiplanas Qu
ercus muehlenbergii
1242 Journal of the Botanical R h Institute of Texas 2(2)
21. Envés con indumento, no glabro; margen entero o con pocos dientes toscos, nunca
lobul
27. + g ME ll H | + Hii + , J tal
tomentoso, cubriendo total tel | | ipales, algunas veces es espar-
cido o tomentuloso.
28. Li h lí +j | J] g + J T J ; AI H g el Ar
y terminand un mucrón.
29. Hojas 3-6(-8) cm de largo, ind to del 5s d gris-blanquecino, de
pelos o Quercus grisea
29. Hojas 1-2.5(-4) cm de largo, ind to del envés café-amarillento, de pelos
fasciclados pie: Quercus striatula
28. Li ;marg dientes, si ent t
las nervaduras las: Api dondeado, raras veces agudo.
30. Margen de la cúpula involuto; haz verde claro, opaco; limbos obovados algu-
nas veces oblongos, con dientes toscos crispados en la mitad distal de la
hoja Quercus invaginata
30. Margen de la cüpul luto; l | | lim
a elípticos, ent n dientes cortos Que ercus mohriana
27. Envé ] t ind to esparcido, cubriendo la epidermis o solo a través
de las beet d. raras veces denso
31. Hojas de 5-15 cm largo; limbos elípticos, oblongos u obovados Quercus laeta
31. Hojas comúnmente menos de 5 cm de "hs rara vez más grandes; limbos obo-
Mare O dl ago las oblongos
32 iio A `
. Y 7 f 1
sos en el envés; margen de la hoja con dientes terminados en mucrones
Quercus potosina
. Margen de la cúpula no involuto; limbos delgados y flexibles, sin puntos
H | EA ri ¡PE r T Aj + FAL mil
UJ
No
,
A
irregulares Quercus carmenensis
DESCRIPCIÓN DE LOS ENCINOS DE COAHUILA
Quercus arizonica Sarg., Gard. & Forest. 8:92. 1895. (Fig. 1). Tiro: E.U.A. Arizona: Fort Huachuca, C.S. Sargent s.n.
(HoLorIPo: GH).
Quercus sacame Trel., Mem. Natl. Acad. Sci. 20:89, pl. 142. 1924.
Quercus endemica C.H. Mull., Amer. Midl. Naturalist 18:846. 1937.
Árbol o d ce 8-12 m de alto; corteza gruesa, fisurada, gris claro; ramillas de 1.5-2.5 mm de grueso,
I
conp f ides, subagudas o redondas de 3 mm largo, café amarillento opaco; estípulas
persistentes (ilmsi o subuladas, de 5-8 mm de largo, pecu aa peces o subpersistentes,
gruesas y coriáceas; pecíolos de hasta 1 cm de largo, con ind idos; limbos
elípticos u oblongos hasta oblanceolados, de 4—8 cm de largo y e cm de ancho, ápice agido a obtuso, base
cordada o redondeada, margen cartilaginoso revoluto, entero o con di del ápice, terminando
en un mucrón, haz algo lustroso, con indumento diminuto y esparcido, de pelos fasciculados, envés opaco,
con pelos fasciculados estipitados, abundantes, con frecuencia glauco; nervaduras 9-11 en cada lado, rami-
ficadas y anastomosadas hacia el margen, impresas en el haz, muy prominentes y reticuladas en el envés.
Inflorescencias masculinas de hasta 3 cm de largo, raquis tomentoso, con flores abundantes. Inflorescencias
femeninas de hasta 2 cm de largo, con tomento corto, de 2—6 flores. Fruto anual, solitario o en pares, subsésil
o con pedúnculos cortos; cúpula hemisférica, de 8-14 mm de ancho y 5-7 mm de alto, márgenes gruesos,
escamas ovadas, base con tuberculo central, tomentosa, ápice redondea do, delgado, apresado, puberulento
y café, nuez ovoide de 8-12 mm de largo, café claro, glabras, incluida cerca de la mitad en la cúpula.
Distribución.—Se distribuye desde el suroeste de Arizona, Nuevo México, suroeste de Texas (Trans-
Pecos) a través de la Sierra Madre Occidental en el este de Sonora, Chihuahua, Durango y Baja California.
En Coahuila se encuentran poblaciones relicto en las sierras del Carmen y Las Cruces al noroeste del estado.
Es una especie característica del bosque de pino-encino y bosque de encino, asociándose principalmente
(F ) de Coahuil 1243
Villarreal et al ,Q
con especies de Quercus gravesii, Q. mohriana, Q. hypoleucoides, Q. 1 ek M sinuata var. breviloba, Q. rugosa, Q.
muehlenbergii, Juniperus flaccida, Rhus aromatica, Garrya ovata, Arl I Salvia regla, Pinus remota, P.
arizonica var. stormiae, Acer grandidentatum, Tilia mexicana, Prunus serotina y CM arizonica entre otras.
Crece en altitudes entre 2100-2600 m, principalmente sobre laderas medias a altas, con exposición norte y
noroeste, así como entre cañones, con suelos derivados de rocas ígneas o sedimentarias. Especie considerada
como rara para el área del estado.
Notas.—Se reconoce por sus hojas gruesas elípticas a oblongas, de más de 4 cm de largo, margen entero
o con pocos dientes mucronados, envés verde amarillento con venación prominente e indumento denso.
Escamas de la cápula con base tuberculada. Puede confundirse con Q. hypoleucoides por las caracteristicas
de la "
M Acuña: si del Carmen, western scarp, oak woodland forest, 2,500 m, 13 jul 1997, S. Wood, G. Harper y
D. Doan- Crider s.n. TURA El Mirador, 28°96'N y 102°57'W, 2,266 m, 17 jul 1997, S. Wood, G. Harper y D. Doan-Crider s.n. (ANSM).
Quercus puit E , Mem. Natl. Ácad. 20:188, pl. 379. 1924. (Fig. 2). Tiro: MÉXICO. Nuevo León: Monterrey,
C.G : B; isoripos: GH, M
Quercus karwinskii Trel., Mem. Natl. Acad. Sci. Whas. 20:188. 1924.
Quercus graciliramis C.H. Mull., J. Arnold Arbor. 17:177. 1936
Arbol de 8-15 m de alto; ramillas rojizas, glabras, con pocas lenticelas, de 2-3 mm de grueso; yemas café,
ovoides, acuminadas de 2-3 mm de largo; estípulas ciliadas, caedizas. Hojas deciduas, delgadas; pecíolos
glabros de 15-20 mm de largo; limbos lanceolados, atenuados hacia el ápice, de 5-10 cm de largo y 2-34)
cm de ancho; base generalmente cuneada, en algunas ocasiones redondeada; margen con tres a cinco pares
de lóbulos cortos, anchos en la base, aristados, haz glabro o con escasos pelos estrellados di tos y envés
con mechones de pelos fasciculados estipitados en las axilas de las venas y ocasionalmente a lo largo de la
nervadura central y la base del pecíolo; nervaduras 6-10 en cada lado. Inflorescencias masculinas de 23
cm de largo, pubescentes. Inflorescencias femeninas de 5-10 mm largo. Fruto sésil, en grupos de uno a
tres, sobre un pedúnculo de cerca de 4 mm; cúpula de 6-10 mm de diámetro y de 4—7 mm de alto; escamas
ovadas delgadas, con el ápice truncado, café rojizo y puberulentas; nuez ovoide, de 12-15 mm de largo,
incluida de un medio a un tercio en la cía
Distribución.—Su área de distrib lel tados de Nuevo León y Tamaulipas. En Coahuila
se presenta en la Sierra de Pájaros Azules, en él centro del estado, en los límites con Nuevo León. Crece en
bosque de encino asociada con Quercus invaginata, Q. laceyi, Q. gravesii, Q. polymorpha, Juglans microcarpa,
Fraxinus greggii, Chiococca alba, Colubrina greggii, Rhus virens, Pistacia texana y Ungnadia speciosa, en altitudes
entre 700-850 m, sobre laderas bajas y en el fondo de cañones húmedos, con exposición noreste. Especie
rara para el área del estado.
Notas.—Se reconoce por las hojas delgadas, casi glabras, verde oscuro, lanceoladas de menos de 10 cm
de largo, margen con 3-5 pares de lóbulos cortos y aristados, envés con pelos fasciculados en la nervaduras
principales. Escamas de la cúpula casi planas. Presenta semejanzas morfológicas con Quercus gravesii.
Material examinado: Mpio. Candela: sierra Pajaros Azules, rancho Santa María, limites con Lampazos N. L., 780 m, 14 jul 2002, JJ.
Medellín V. 243,341 (CFNL).
Quercus carmenensis C.H. Mull., Amer. Midl. Naturalist 18:847. 1937. (Fig. 3). Two: MÉXICO. Coamura: Mpio.
Acuña, sierra del Carmen, canyon oa EL. Wynd y C.H. Muller 639 (4oLoriPo: ILL; isoriros: GH, MO).
Quercus undulata Torr., Ann. Lyceum. Nat. Hist. N.Y. 2:248. 1828.
Arbusto rizomatoso de 0.5-2.0 m de alto, o árboles de hasta 12 m de alto; corteza gris clara, a cuadros o
surcada; ramillas de 1-1.5 mm de grosor, de color rojo, con lenticelas pálidas y prominentes y con pelos
multirradiados yemas casi redondas de 1-1.5 mm de largo, café claro; estípulas persistentes de 3-6 mm
de largo, en forma de arista. Hojas deciduas, delgadas y flexibles, con pecíolos de (225-10 mm de largo y
casi 1 mm de grosor, glabrescentes, rojos; limbos oblongos a obovados, de (223-5 cm de largo y 13 cm
de ancho, ápice obtuso a redondeado, base cuneada a redondeada, margen entero, ondulado, lobulado o
vous ] af tha Ratanical D h Institute of Texas 2(2)
iv] ]
Tom
Fic. 1. Quercus arizonica. A
ln £. : Im
variación en Fic. 2. Quercus canbyi. A, rama con hojas, amentos y bellota; B, hoja; C, pelo
i > 1 £. E J J binih, J n hallota: E "wt ^ i
J
tipitado: D, bellota: E
:
Fic. 3. Q ic A Los ; B, nri Ia E ja; C Fic. 4. Q nn n A, rama con hojas; B, | " I I H I
| li lados: D, bellota: E de la cúpul
pelo fasciculado. haz; G p ; I
Villarreal et al., Q (F ) de Coahuil 1245
toscamente dentado hacia el ápice; haz verde oscuro, con indumento de pelos multirradiados diminutos y
esparcidos, envés verde claro, verde amarillento o ligeramente amarillo rojizo, con indumento más promi-
nente; nervaduras 9-12 en cada lado, ramificadas y anastomosadas cerca del margen o terminando en un
mucrón en los dientes cuando estos ocurren. Inflorescencias masculinas de 2—3 cm de largo. Inflorescencias
femeninas de 1-5 cm de largo de 23 flores sobre pedúnculos tomentosos y delgados. Fruto anual, solitario
o en pares, subsésil o con un pedúnculo corto; cúpula (inmadura) con escamas agudas, café claro, canes-
centes; nuez no conocida.
Distribución.—Distribuida principal la Sierra del Carmen, en el noroeste de Coahuila, presente
en los bosques mixtos de pino-encino y bosque de encino. Se asocia generalmente con Cupressus arizonica,
Pseudotsuga menziesii, Pinus strobiformis, P. arizonica var. stormiae, Quercus sideroxyla, Q. hypoleucoides, Q.
rugosa, Q. gambelii, Garrya ovata, Salvia regla, Arbutus xalapensis, los pungens, Acer grandidentatum
y Juniperus deppeana. Se le encuentra en los 2200-2500 m de altitud; ee en la región de la Sierra del
Carmen, es poco frecuente en las montañas Chisos de Texas. Crece en valles intermontanos altos y en laderas
altas con exposición norte y noroeste, como arbusto rizomatoso, sobre suelos someros y en condiciones de
poca humedad, mientras que en cañones húmedos puede ser un árbol de hasta 12 m o más de alto. Con-
siderado como especie endémica del área de Megacoahuila (Villarreal y Encina 2005). Muy relacionado con
Q. undulada, posiblemente solo se trate de una variación de esta especie.
Notas.—Se reconoce por sus hojas delgadas, casi glabras, obovadas a oblongas de menos de 5 cm de
a margen entero, con pocos dientes a cortamente lobado, con pelos multirradiados esparcidos.
M lo: Mpio. Ocampo: El Jardín ición N de la si del Carmen, 2,200 m, 8 jun 1991, J.A. Pérez 1765 (ANSM); sierra
del Carmen, Moreno ER 28?98'N y 102°62'W, 2,165 m, 26 ago 1997, S. Wood, G. Harper y D. Doan-Crider s.n. (ANSM)
Quercus coahuilensis K. C. Nixon et C.H. Mull., Brittonia 45:150. 1993. (Fig. 4). Tiro: MÉXICO. Coanuma: sierra
del Pino, I. M. Johnston y C.H. Muller 567 (Hotoriwo: BH-CHM
Arbusto o raras veces árbol hasta de 8 m de alto, corteza obscura a lros; ramillas de 1-1.5 mm de grosor con
indumento gris o café rojizo, glabrescente; yemas ovoides, de 3-4 mm de largo, café rojizo, con puberulencia
blanquecina hacia el ápice; estípulas subuladas, de 5-6 mm de largo, caedizas. Hojas persistentes; pecíolos de
3—5(—7) mm de largo, con tomento gris persistente; limbos de (1.5—)3—4(-6) cm de largo, de (121.5—3 cm de
ancho, estrechamente elípticos a oblongos, o algunas veces obovados, ápice agudo a redondeado terminando
en arista corta, base redondeada o por lo común cordada, margen plano o un poco revoluto, con 1-34) pares
de dientes cortamente aristados, haz lustroso, verde amarillento o grisáceo, glabro o con indumento amaril-
lento persistente a lo largo de la nervadura media, con pelos diminutos, crispados, fasciculados dejando ver
puntos traslácidos (bajo el microscopio), envés con pelos fasciculados, con radios enredados desde la base,
amarillo-blanquecinos, glabrescente, epidermis con ampulas planas, la nervadura media frecuentemente
tornándose rosada con la edad; nervaduras 5-7 en cada lado, irregulares y desiguales, impresas en el haz,
prominentes en el envés. Inflorescencias masculinas cerca de 2 cm de largo. Inflorescencias femeninas 1
o raramente 2 sobre pedúnculos tomentosos de 1-4 mm de largo. Fruto bianual, solitario o raras veces en
pares, subsésil o sobre peauncutos de 3-4 mm de largo; cúpula hemisférica, de 9-11 mm de ancho, escamas
de la cúpula delgadas y floj apresadas, glabras y café claro, márgenes con cilios grisáceos, las puntas
erosas; nuez ovoides, café claro, con puberulencia densa, incluida la mitad en la cúpula.
Distribución. —Su distribución comprende la sierra del Diablo en el sureste de Chihuahua y en Coahuila
se distribuye en sierra del Pino, del Fuste, La Fragua, La Madera, San Marcos, Parras, Jimulco y La Con-
cordia. Forma parte del matorral submontano y del bosque de pino-encino, algunas veces en los bosques
de encino, asociándose con Pinus cembroides, P. remota, P. arizonica var. stormiae, Quercus pringlei, Q. greggii,
Q. intricata, Q. gravesii, Q. invaginata, Dasylirion cedrosanum, Rhus aromatica, R. virens, Arbutus xalapensis,
Arctostaphylos pungens, Nolina cespitifera, Fraxinus cuspidata, F. greggii, Prunus serotina, Acer grandidentatum,
Cercocarpus fothergilloides, C. montanus y Flourensia retinophylla. Crece entre los 1850-2800 m de altitud, en
laderas bajas con exposición norte y noreste en el matorral submontano, en laderas altas con exposición
sur en bosques de pino. En las sierras de la Madera y del Pino es más abundante, que en las sierras del sur
1246 J lof the Botanical R h Institute of Texas 2(2)
del estado. Se presenta como arbusto en el matorral submontano, y en el bosque de encino algunas veces
se presenta como árbol de hasta 10 m de alto. Considerado como endémico para el área de Megacoahuila
(Villarreal y Encina 2005).
Notas.—Se reconoce por sus cue coriáceas, ps a oblongas, menos de 6 cm de largo, usualmente
verde amarillentas, margen con dient tados, envés con vención prominente, es característico
la presencia de abundantes pelos fasciculados con radios enredados Escamas de la cúpula delgadas y lisas.
Relacionado con Quercus hintoniorum y Q. hypoxantha.
Material examinado: Mpio. Parras: sierra Playa Madero, cañón Pinalosa, = Santa Victoria, 25°09'N y 101°34'W, 2,300-2,600 m. 6
jun 1995, M.A. Carranza y J.A. Encina 153 (ANSM), M. A 2210 (Ah
Quercus emoryi Torr., WH. Emory, Not. Milit. Reconn. 151, pl 9. 1848. (Fig. 5). Two: E.U.A. Texas: Emory s.n.
(morotro: NY)
Quercus hastata Liebm., Overs. Danske Vidensk. Selsk. Forh. 1854.
Quercus balsequillana Trel., Mem. Natl. Acad. Sci. 20:123, pl. 220. 1924.
Quercus duraznillo Trel., Mem. Natl. Acad. Sci. 20:122, pl. 220. 1924.
Árbol de (5215-20 m de alto, corteza negra áspera y surcada; ramillas de 1-2.5 mm de grosor, café rojizo,
con tomento denso, amarillento; yemas de 1.5-3.0 mm de largo, café lustroso; estípulas caedizas, de 5-8
mm de largo. Hojas subpersistentes, gruesas y coriáceas; pecíolos de 5-8(-12) mm de largo, glabrescentes;
limbos lanceolados, elíptico-oblongos u ovados, de 3-6(-8) cm de largo y de 1—2.5(-3.5) cm de ancho,
verde claro a amarillento, ápice agudo, acuminado y mucronado o con arista corta, base truncada, cordada
o redondeada; margen plano o algo revoluto, entero, algo lobulado o con dientes aristados; hojas con haz
liso y brillante, glabro o con pelos fasciculados, envés menos brillante, con penachos prominentes de pelos
fasciculados estipitados a lo largo de la Henn HE o de cerca p pee o glabro; nervaduras de
5—8(-10) en cada lado, ramificadas. Infl los cortos, 1-2 flores
tomentosas. Fruto anual, solitario o en pares, sobre pedünculos de 1-2 mm de largo, cúpula de 10 mm de
diámetro, hemisférica, escamas de la cúpula apresadas, ovadas y delgadas, ápice glabro, café-castaño, base
con puberulencia grisácea, nuez oblonga, elíptica u ovada, de 10-15 mm de largo, de 5-9 mm de ancho,
café rojiza, ápice piloso, incluida cerca de la mitad en la cúpula.
Distribución.—Se distribuye desde Arizona, Nuevo México y las montañas del Trans-Pecos en Texas,
llegando a México a través de la Sierra Madre Occidental en los estados de Chihuahua y Durango. En
Coahuila se encuentra solo en la sierra del Carmen al noroeste del estado, forma parte de los bosques de
pino-encino y bosques de encino, iándose con especies de Quercus grisea, Q. laceyi, Q. gravesii, Q. mohri-
ana, Q. arizonica, Q. hypoleucoides, Arbutus xalapensis, Acer grandidentatum, Salvia regla, Crataegus greggiana,
Fraxinus cuspidata, Garrya ovata, Pinus remota, P. arizonica var. stormiae, Prunus mexicana, Juniperus flaccida,
J. scopulorum y Leucaena retusa. Su rango altitudinal es de 1500-2300 m, sobre laderas medias con suelos
derivados de rocas ígneas, con exposición norte, noroeste o en fondos de cañones boscosos con arroyos,
donde forma poblaciones más o menos densas. Por su distribución, se considera raro en Coahuila.
Notas.—Se reconoce por sus hojas gruesas, elípticas a oblongas, usualmente menos de 6 cm de largo,
verde amarillentas, con apice agudo cortamente aristado, casi glabras con pelos fasciculados en mechones
en la nervadura media. Escamas de la cúpula delgadas y lisas.
Material examinado: Mpios. Acuña y Ocampo: sierra Madera del Carmen, rancho El Secadero, cañón Poblano, 29°00'N y 102°20'W,
2,100—2,800 m, 20 ago 1994, M.A. Carranza y J.A. Encina 110 (ANSM).
Quercus fulva Liebm., ENS "Kongek pu. Vidensk. Selsk. Forh. Medlemmers Arbeider. 183. 1854. (Fig.
6). Tiro: MEXICO n B.C. Seemann 1973 (norotiro: C; rsorrro: GH).
Quercus rosei Liebm., Mem. Natl. Acad. Sci. 20:137, pl. 260. 1924.
Arbusto o árbol de (325—8(-10) m de alto; corteza surcada, gris obscura; ramillas de 2.5-5 mm de grosor,
con tomento denso amarillento; yemas ovoides a cortamente agudas, de 3-4 mm de largo, pubescentes;
estípulas lineares, escariosas. Hojas generalmente deciduas, rígidas, coriáceas y planas; pecíolos de 10-23
Villarreal et al., Q (F ) de Coahuil 1247
mm de largo, tomentosos; limb lípti obovados, suborbicul flabelados, de (6-)8-15 cm de largo,
de 4—8(-12) cm de ancho, ápice nda, redondeado, obtuso o subagudo, comúnmente aristado, base
cordada o redondeada, margen revoluto, cartilaginoso, entero o con 6-11 dientes cortos, cortamente aristados;
haz verde pálido poco lustroso, glabro excepto la nervadura media, las aos pue impresas,
] además de pelos
envés con tomento denso persistente, amarillo-cafesuzco, formado de p
vermiformes; nervaduras 6—10 en cada lado, ascendentes, ds Inflorescencias Hansen aa de 2-3
cm de largo, pubescentes. Inflorescencias femeninas con 1-3 flores, usualmente en pares, con pedánculos
de 2—5(-15) mm, largo, tomentosos. Fruto bianual, solitario, o en pares, sobre pedúnculos de de 10-15
mm, cúpula casi hemisférica, de 13-20 mm de ancho, de 6-10mm de largo, con el márgene ligeramente
involuto, escamas café con indumento pálido, un poco engrosadas en la base, ápice delgado, apresado; nuez
ampliamente ovoides o casi esféricas, de 9-12 mm de largo, de 8-11 mm de ancho, café claro, incluida
cerca de la mitad en la ce
Distribución. —Se dist I suroeste de México en los estados de Jalisco, Nayarit y Sinaloa, llegando
hasta el norte a través de la Sierra Madre Occidental a Chihuahua y Durango. En la Sierra Madre Oriental
se distribuye en Nuevo León y Coahuila. En Coahuila se han encontrado poblaciones relicto en la sierra de
Arteaga y sierra de Zapalinamé al sureste del estado. Crece en bosque de pino-encino y bosques de oyamel,
asociándose con Quercus hintoniorum, Q. greggii, Q. saltillensis, Pseudotsuga menziesii, Pinus greggii, P. teocote, P.
pseudostrobus, P. rudis, Prunus serotina, Arbutus xalapensis, Cupressus arizonica, Populus tremuloides, Ceanothus
buxifolius y Garrya ovata. Se desarrolla en altitudes de 2000-2800 m, en laderas medias con exposición
noroeste o en cañones húmedos donde es abundante, su abundancia disminuye en bosques más secos de
la exposición sureste. Es poco frecuente en los cañones de Los Lirios y Jamé en la Sierra de Arteaga. La
distribución restringida de esta especie la hace aparecer como rara en el área del estado.
Notas.—Se puede reconocer por sus hojas coriáceas, obovadas a suborbiculares de 6-15 cm de largo,
verde claro, ápice redondeado y margen con dientes aristados, envés con abundantes pelos simples y multir-
radiados. Bellotas casi esfericas, cúpula con margen involuto, las escamas de la cúpula con base ensanchada,
e n y ápice corto. Puede confundirse con Quercus potosina por la forma de las hojas.
M M Art ino Los Lirios-Laguna de Sanchez, ca. 25°19'N y 100°26'W, 2,200 m, 2 nov. 1994. J.A. Villarreal
y J.A. Encina 125 (ANSM); si B as Vigas, cañón de Jamé, rancho » Tu ca. Bá km jn NE a 25?26'N y 100°20'W, 2,400—2,600
m, 8 may 1995, M.A. mod Encina 144 (ANSM). Mpio. Salti El Chicharrón, vereda entre el ejido sierra
Hermosa y cañón de San Lorenzo, 25?20'N y 100°55'W, 2,532 m, 6 jul 2000, J.A. Ent R. ot 518 (ANSM).
Quercus fusiformis Small, Bull. Torrey Bot. Club 28:357. 1901. (Fig. 7). ~ virginiana Mill. var. fusiformis (Small)
Sarg., Bot. Gaz. 65:448. 1918. Tiro: E.U.A. Texas: Kerrville, H. Lacey s.n. (notorio: NY
Quercus oleoides Schltdl. et Cham. var. quaterna C.H. Mull., Contr. Texas Res. Found., Bot. Stud. 1:76, tab. 54, 55. 1951.
Arbusto o árbol rizomatoso de 5—10(-12) m de alto; corteza áspera gris o negra; ramillas de 1.5-2.5 mm
de grosor, con tomentulo amarillo-rojizo a gris; yemas subglobosas, café rojizas, puberulentas; estípulas de
20-25 mm de largo, subuladas, pubescentes, caedizas. Hojas persistentes, coriáceas; pecíolos de 2-8 mm
de largo con indumento denso; limbos oblongos, elípticos u obovados, de 2-7 cm de largo, de 0.5-3 cm
de ancho, ápice agudo a redondeado, cortamente mucronado, base cuneada a redondeada, margen entero
a irregularmente dentado, algo revoluto o casi plano, haz verde brillante y glabro o con indumento estrel-
lado esparcido, envés gris plateado, con abundantes pelos estrellados, cortos y adpresos, nervaduras 8-10
en cada lado, con algunas intermedios, muy ramificadas, inconspicuas en el haz, poco prominentes en el
envés. Inflorescencias masculinas de 3.5-4.5 cm de largo sobre pedúnculos delgados con pelos glandulares
esparcidos. Inflorescencias femeninas de 1.5-5.5 cm de largo con 2 a 3 o más flores. Fruto anual, solitario o
en grupos de 2 a 3, sobre pedúnculos de 1.5-8 mm de largo, glabros, cúpula de 7-12 mm de ancho, 7-10 mm
de alto; escamas ovadas engrosadas en la base, con indumento denso y grisaceo, ápice agudo, rojo oscuro,
puberulento; nuez ovoide, subfusiforme a fusiforme de 1.5-2.0 cm de largo, 1-1.3 cm de ancho, glabra y
café opaco, incluida un medio a un tercio en la cúpula.
Distribución.—Se distribuye desde Oklahoma y Texas, en México a lo largo de la Sierra Madre Oriental
1248 J | of the Botanical R h Institute of Texas 2(2)
Fic. 5 Quercus emoryi A I j y bellota; B iació I j ;C, Fic. 6. Quercus fulva. À Í j y j l
pelo fasciculado estipitado; D, bellota; E de la cüpul y por el haz; C, pelos ramificado y vermiforme; D, bellota; E, escama de
la cüpula.
y ( i /
VAL V
fusiformis. A hoi bellota: B, hoias: C, pelo Fic. 8. Quercus gambelii. A
ey 707; wr F 2 T (d 4
C pelo fasciculado: D, bellota: E de la cúpul
"1! , f
Fic
estrellado; D, bellota; E, escama de la cúpula
(F ) de Coahuil 1249
Villarreal et al ,0
en Nuevo León y Tamaulipas. En Coahuila es encontrado en la Planicie Costera del Golfo, al noreste del
estado, en las serranías del Pune la sierra de fa y La Encantada. Se le encuentra en el matorral
submontano como u p o bien com bolito en la transición con el matorral tamaulipeco,
así como en el TÉ de encino, asociándose con Dyospiros texana, Bumelia lanuginosa, Calia secundiflora,
Quercus invaginata, Q. mohriana, Q. sinuata var. breviloba, Ungnadia speciosa, Rhus virens, Condalia hooheri, C.
warnockii, Vauquelinia corymbosa y Juglans microcarpa. En el bosque de encino se asocia a Carya illinoenisis,
Celtis reticulata y Ulmus serotina. Se presenta hasta los 1900 m de altitud, siendo común entre los 600-1300
m. En la Planicie Costera, donde es más abundante, forma bosques densos en lugares húmedos, prefiere
suelos arenosos y profundos o terrenos calizos en laderas bajas, con exposiciones norte y noroeste. En los
márgenes de arroyos, ríos o lugares con humedad forma bosques de grandes árboles, su abundancia dis-
minuye hacia las sierras del sur del estado, es raro en el valle de Saltillo. En el municipio de Ramos Arizpe
se presentan árboles relicto a través de cauces de arroyos. En el noroeste de Nuevo León es frecuente en el
cauce de arroyos intermitentes desde 300 hasta 800 m de altitud, en suelos pedregosos y condiciones de
clima árido (Marroquín 1985). Aunque es comün en gran parte del estado su existencia se ve amenazada
por las actividades del hombre.
Notas.—Se reconoce por las hojas elípticas a obovadas usualmente de menos de 7 cm de largo. Es car-
acterístico el indumento de pelos estrellados escamosos y apresados, el envés grisáceo y las bellotas sobre
pedünculos de hasta 8 mm de largo, con nuez algo fusiforme, estriada y la cápula generalmente corta, con
escamas con base Ced
M I uevas, arroyo Las Vacas, 29°18'N y 100°58'W, 284 m, 12 sep 2002, J.A. Encina y O. Saucedo
1057 (ANSM). Mpio. Candela: sierra i Pisos Azules, campo Santa María, 4 km al SW de San Guillermo, 27°03'N y 100°53'W, 920 m,
11 nov 1997, M.A. Carranza et al. 2763 (ANSM), M.A. Carranza y D. Sanchez 2881 (ANSM). cerros cercanos a Candela, ca. 3 km al SE,
26? 50'N y 101°20'W, 1,900 m, A. Rodríguez y M. Martinez 1105 (ANSM). Mpio. Jiménez: ejido San Vicente, cerca del río San Diego, 29?
O6'N, y 100°55'W, 310 m, 13 sep 2002, J.A. Encina et al. 1086 (ANSM). Mpio. Muzquiz: río San Juan, al E del ejido Morelos, 28°00'N y
101%42'W, 484 m, 3 nov 2005, J.A. Encina e I. Ramirez 1586 (ANSM); sierra Santa Rosa, rancho El Silencio, 27°55'N y 101°50'W, 1,300
m, 2 may 2004, S. Lara iod R. VE 103 (ANSM); sierra Santa Rosa, al sur de Muzquiz, 27?49'N y 101°32'W, 760 m, 5 nov 2005, J.A.
Encina el R Mpio. Pied N 1 , ejido El Moral, 28°51'N y 100°42'W, 270 m, 12 ago 2001, M.A.
Carranza e I. Ramirez 3521 (ANSM). Mpio Ramos Arizpe: El Carmen, carr. 57, ca. 25 km al N de Saltillo, 25 oct 1994, J.A. Villarreal y
M.A. Carranza 7937 (ANSM). Mpio. Sal 1 l San Lorenzo, al SO de la ciudad, 25°23'N y 101°00'W, 1643 m, 8 jun 1996, J.A.
Encina 177 (ANSM). dens Villa Unión: camino de Nava a Villa Unión, rancho La Mota, 28°18'N y 100°44'W, 15 ago 1997, M. González
y A. Torres 2924 (ANSM). Mpio. Zaragoza: río San Antonio, ca. 12 km al N de Zaragoza (carr. 29), 28°32'N y 101%07'W, 240 m, 27 oct
1999, J.A. Villarreal y M.A. Carranza 8971 (ANSM); río San Rodrigo, ca. 25 km al N de Zaragoza, carr. 29 28°43'N y 100°55'W, 330 m,
9 sep 1995, M.A. Carranza 2544 (ANSM).
Quercus gambelii Nutt., J. Acad. Natl. Sci. Phila. 1(2):179. 1848. (Fig. 8). Quercus douglasii Hook. et Arn. var. gambelii
(Nutt.) A. DC., Prodr. (DC.) 16 Q,1):24. 1864. Q. undulata Torr. var. gambelii (Nutt.) Engelm., Trans. Acad. Sci. St. Louis 3:393.
1877. Tiro: E.U.A. New Mexico: Santa Fe, W. Gambel s.n. (isoro: GH).
Quercus utahensis Rydb., Bull. N.Y. Bot. Gard. 2:202. 1901.
Quercus novomexicana Rydb., Bull. N.Y. Bot. Gard. 2:208. 1901.
Quercus marshii C.H. Mull., Amer. Midl. Naturalist 18:848. 1937.
Quercus lesueuri C.H. Mull., Amer. Midl. Naturalist 27:476. 1942.
Arbusto o árbol de hasta 20 m de alto; ramillas de 1.5-2.5 mm de grosor, glabras o con pelos fasciculados;
yemas ovoides de 3-4 mm de largo, café claro con indumento esparcido; estípulas usualmente caedizas,
subuladas, de 4-6 mm de largo. Hojas deciduas, membranosas; pecíolos de 10-25 mm de largo, pubes-
centes o casi glabros; limbos ovados a obovados u oblongos a elípticos, de (428—12(-16) cm de largo, de
(2.52)4-6(-8) cm de ancho, ápice redondeado, base truncada a cuneada, margen con 4-6 lóbulos en cada
lado, oblongos, enteros o dentados, con el ápice redondeado a subagudo, las escotaduras de los lóbulos al-
canzando usualmente más de la mitad de distancia a la nervadura media; haz lustroso verde oscuro, glabro
o con pelos fasciculados cortos formando un indumento diminuto y esparcido, envés verde opaco algunas
d de pelos fasciculados de pocos radios; nervaduras 4—6 en cada lado, muy rami-
veces glauco, con i
ficadas y anastomosadas cerca de la nervadura media, un poco salientes en el haz, prominentes en el envés,
1250 Journal of the Botanical R h Institute of Texas 2(2)
Inflorescencias masculinas de 2-3 cm de largo, con tomento esparcido. Inflorescencias femeninas de 3-8 mm
de largo, con 13 flores en la parte terminal. Fruto anual, solitario o en pares, subsésil o sobre pedúnculos
hasta de 1 cm de largo; cúpula de 7-15 mm de ancho, de 5-7 mm de alto; escamas de la cúpula ovadas,
engrosado-tuberculadas, ápice corto, angosto, base con indumento gris; nuez ovoide a elípsoide, de 12-15
mm de largo, de 8-10 mm de ancho, café clara, casi glabra, incluida cerca de un medio en la cúpula.
Distribución. —Se distribuye en las montañas Rocallosas en los estados de Wyoming, Utah, Nevada,
Arizona, Colorado, Nuevo México y oeste de Texas, llega hasta la Sierra Madre Occidental al norte de Chi-
huahua y Sonora. En Coahuila se le encuentra en las si del Carmen y La Madera, en el norte y centro del
estado. Se localiza en bosques de encino y de pino-encino, asociándose con Quercus carmenensis, Q. arizonica,
is var. coahuilensis,
la
Q. greggii, Pseudotsuga menziesii, Pinus arizonica var. stormiae, P. strobiformis,
Populus tremuloides, Arctostaphylos pungens, Arbutus xalapensis, Acer grandidentatum y n stolonifera. Se
localiza entre 2500-3000 m de altura. En Coahuila presenta un alto grado de aislamiento, restringiéndose
para algunas de las cimas montañosas del centro y norte, crece cerca de laderas en exposición noroeste y
valles intermontanos altos. La especie es rara en el área del estado.
Notas.—Se reconoce fácilmente por sus hojas delgadas, verde obscuro, oblongas, elípticas u ovadas, con
4-6 pares de lóbulos, casi glabras. Escamas de la cúpula con ápice estrecho. Solo se le conoce de las sierras
del Carmen y la Madera.
Material examinado: Mpio. Acuña: sierra del Carmen, roadside collection between El Uno and Main Camp., 2,300 m, 25 ago 1997, S.
Wood, G. Harper y D. Doan-Crider s.n. (ANS
Quercus gravesii Sudw., Check List For. Trees U.S. 86. 1927. (Fig. 9).
Quercus texana Buckley var. chesosensis Sarg., Bot. Gaz. 65:423. 1918.
Quercus texana Buckley var. stellipila Sarg., Bot. Gaz. 65:424. 1918.
Quercus stellipila (Sarg.) Parks ex Cory, Rhodora 38:405. 1936.
Quercus chesosensis (Sarg.) C.H. Mull., Amer. Midl. Naturalist 18:850. 1937.
Quercus shumardii Buckley var. microcarpa (Torr.) Shinners, Field & Lab. 24:37. 1956.
Árbol de hasta 13 m de alto; corteza negra, áspera y surcada; ramillas de 1-2 mm de grosor, café rojizas
con pocas lenticelas, con indumento amarillento, glabrescentes; yemas agudas de 3-6 mm de largo, café
rojizo; estípulas caedizas, de 5-10 mm de largo. Hojas deciduas, delgadas y coriáceas; pecíolos hasta de
15 mm de largo, glabrescentes; limbos ovados u obovados, de (528-12(-14) cm de largo, de (4-26-8(-12)
cm de ancho, ápice agudo a obtuso, base ns a redondeada, margen con 23 lóbulos en cada lado con
sinuosidades profundas y abiertas, al n lamente la mitad a tres cuartos de distancia a la nervadura
media, los lóbulo dos y enteros con pocos dientes en la parte distal, los lóbulos terminales elongados,
dentados o ligeramente lobulados, terminados en aristas, haz verde claro, lustroso, glabro, con indumento
escaso de pelos fasciculados, envés verde claro o cobrizo, opaco, glabro o con algo de indumento fasciculado
diminuto; nervaduras una en cada lóbulo, salientes en ambas superficies. Inflorescencias masculinas de
6—7 cm de largo, con pelos fasciculados esparcidos. Inflorescencias femeninas de hasta 15 mm largo, 1-3
flores sobre pedánculos glabrescentes café rojizo. Fruto bianual, solitario o en pares, sobre pedúnculos de
4-12 mm de largo, cúpula de 1-1.5 cm de ancho, de 5-8 mm de alto, hemisférica; escamas de la cúpula
redondeadas, café claro, con indumento diminuto, denso, margen ciliado; nuez ovoide, de 12-15 mm de
largo y de 8-10 mm de ancho, café claro opaco, con puberulencia densa, glabrescente, incluida de un tercio
a un medio en la cúpula.
Distribución. —Se distribuye en las montañas del Trans-Pecos al oeste de Texas y la confluencia de los
ríos Bravo y Pecos, llega hasta Coahuila encontrándose en la mayoría de las montañas del norte, noroeste
y del centro del estado (sierra del Carmen, La Encantada, del Burro, La Madera, La Gloria y La Paila) y en
forma aislada se distribuye en la sierra de Zapalinamé, al sureste del estado. Se reportan híbridos con Q.
coahuilensis para la sierra de la Madera. El hibrido presenta hojas como Q. gravesii con lóbulos cortos a di-
entes cortos, aristados y envés ampulado (Muller, en prensa). Crece en el bosque de pino-encino y bosque
de encino asociándose con las siguientes especies: Quercus laceyi, Q. hypoleucoides, Q. grisea, Q. emoryi, Q.
(F ) de Coahuil 1251
Villarreal et al ,Q
invaginata, Q. mohriana, Q. muehlenbergii, Pinus arizonica var. stormiae, P. strobiformis, P. remota, Juniperus
flaccida, Cupressus arizonica, Prunus serotina, Acer grandidentatum, Arbutus xalapensis, Cercis canadensis,
Cornus stolonifera, Leucaena retusa, Fraxinus berlanderiana, en lugares con abundante humedad se encuentra
asociado con Ungnadia speciosa, Prunus serotina, Juglans microcarpa, J. mollis y Tilia mexicana.Se desarrolla
en elevaciones entre 1200-2500 m, sobre laderas medias y altas, con exposición norte y noroeste, en valles
intermontanos o al lado de arroyos en cañones húmedos, frecuentemente formando bosques densos. La
especie es considerada endémica para el área de Coahuila.
Notas.—Se reconoce fácilmente por sus hojas delgadas, verde claro, casi glabras, ovadas a obovadas,
usualmente de 8-12 cm de largo, con 23 pares de lóbulos terminados en aristas prominentes. Escamas de
la eee planas.Presenta semejanzas en morfología de la hoja con Q. canbyi.
Material Acuñ ias del Burro, rancho El Bonito, 29%01'N y 102°07'W, 1,625 m, 19 sep 1977, D. Riskind y C.H.
Muller 2201 (ANSM, TEX. Mpio. Candela: si zules, i ada Las Sabinillas, 27°02'N y 100°54'W, 1,300
997, M.A. Carranza et al. 2819 (ANSM). Mpio. Castaños sierra La Gavia, rancho La Gavia, 26?10'N y 101°18'W, 1,295 m,
10 ago ies M.A. Carranza y J.A. Encina 170 (ANSM). Mpio. Cuatro Cienegas: sierra San Marcos, cañón Grande, ejido Estanque de
Norias, 27°40'N y 102°50'W, 1,550 m, 1 oct 1993, M.A. Carranza y J.A. Encina 1683 (ANSM); sierra de La Madera, cañón Charreteras,
rancho Charreteras, 27?12'N y 102%42'W, 1,750 m, 15 sep 1993, J. A. Villarreal ai M.A. REA in nee Mpio. Muzquiz: sierra
Santa Rosa, ejido La Esperanza, 28°08'N y 102°00'W, 1,300 m, 8 sep 1990, R 1 Rosa (La Babia), rancho
La Babia, cañón Rincón de María, 28°27'N y 102°04'W, 1,450 m, 21 ago 1999, M.A. Carranza et al. 3077 (ANSM). Mpio. Ocampo:
del Pino, ejido Acebuches, cañón La Vaca, 28°15'N y 102°59'W, 1,850 m, 1 oct 2003, M.A. Carranza e I. Ramirez 3958 ra sierra
Maderas del Carmen, rancho El Secadero, 29%00'N y 102°20'W, 2,100 m, 20 ago 1994, M.A. Carranza y J.A. Encina 2033 (ANSM); Sierra
La Encantada, cuesta de Malena 28°35'N y 102°10'W, 21 ago 1994, M.A. Carranza y J.A. Encina 113 (ANSM). Mpio. Ramos Arizpe: sierra
de La Paila, ejido El Cedral, 26°02'N y 101°23'W, 1,300 m, 20 ago 1987, J.A. Villarreal y M.A. Carranza 3939 (ANSM). Mpio. Zaragoza:
rancho La Escondida, 28°53'N y 101°57'W, 1.200 m, 6 may 2003, M. Guerrero y O. Ballesteros s.n. (ANSM)
Quercus greggii (A. DC.) Trel., Contr. U.S. Natl. Herb. 23:185. 1922. (Fig. 10). Quercus reticulata Kunth var. greggii
A. DC., Prodr. (DC.) 16 (2.1):34. 1864. Quercus greggii (A. DC.) Trel. f. subglabra C.H. Mull. in I.M. Johnst., J. Arnold Arbor. 25:440.
1944. Tiro: MÉXICO. Coanuna: Arteaga, San Antonio de las Alazanas, J. Gregg 389 (HoLoriro: US). Q. loeseneri Trel., Mem. Natl.
Acad. Sci. 20:79, pl. 110. 1924.
Quercus derrumbaderoensis C.H. Mull., Amer. Midl. Naturalist 27:471. 1942.
Arbusto o arbolito de 0.5—5(-12) m de alto; corteza gris escamosa; ramillas de 1.5-3 mm de grosor, con
pelos fasciculados diminutos, densos, café amarillento; yemas subredondas, de 1 mm de diámetro; estípulas
lanceoladas, pubescentes de 4-5 mm de largo. Hojas persistentes, coriáceas; pecíolos de 3-6(-10) mm de
largo, pubescentes; limbos ovados, obovado-elípticos u oblongos de (1.5-)2-3(55) cm de largo, de (1-)1.5-4
cm de ancho, ápice obtuso a casi redondeado, base redondeada a cordada, margen entero a ondulado,
engrosado, revoluto, con 3—5 dientes mucronados o lóbulos poco profundos en la mitad distal de la hoja,
haz verde oscuro, algo lustroso con indumento en las nervaduras, envés densamente lanoso-tomentoso,
con pelos fasciculados y rizados, café amarillento y pelos glandulares; nervaduras prominentes, 5-6(-8)
por cada lado, ramificadas, impresas en el haz y elevadas en el envés. Inflorescencias masculinas de 3—4.5
cm de largo, con pelos fasciculados. Inflorescencias femeninas de 1—3.5 cm de largo con 2-4 flores. Fruto
anual, en grupos de 1-4, pedúnculo tomentoso, de 5-30 mm de largo; cúpula hemisférica de 10-15(-20)
mm de diámetro, escamas café rojizo, base engrosada y pubescente, ápice delgado, apresado, las escamas
del margen rebasando el borde; nuez ovoide-elíptica, de 10-17 mm de largo, glabra, incluida alrededor de
un medio o más en la cúpula.
Distribución. —Se distribuye desde el centro-este de México, en los estados de Hidalgo, Puebla y San
Luis Potosí, llega al noreste a través de la Sierra Madre Oriental hasta Nuevo León, en Coahuila se encuentra
en las montañas del centro, sur y sureste del estado, en las sierras de La Madera, de Parras, La Concordia,
de Zapalinamé y de Arteaga. Es un componente de los bosques de pino-encino, encino, oyamel y en la veg-
etación alpina-subalpina. Se asocia principalmente con Quercus hintoniorum, Q. saltillensis, Q. coahuilensis,
Pinus pseudostrobus, P. culminicola, P. arizonica var. stormiae, P. rudis, P. strobiformis, Cornus stolonifera, Garrya
ovata, Arctostaphylos pungens, Pseudotsuga menziesii, Abies durangensis var. coahuilensis, Acer grandidentatum,
Ceanothus buxifolius, Arbutus xalapensis y Prunus serotina. Se localiza entre los 2400-3400 m de altitud, en
1252 J | of the Botanical R h Institute of Texas 2(2)
cimas montañosas, laderas altas con exposición norte y noroeste, se encuentra además en cañones boscosos
hümedos y semihümedos o en terrenos semisecos, sus hábitos de crecimiento varían desde tipo arbustivo o
arbolito de 3-5 m, disminuyendo su altura con la altitud. Es una especie muy abundante en las montañas
del sureste del estado y frecuente en la Sierra de la Madera. La población de la sierra de la Madera reportada
como Q. greggii f. subglabra, es al parecer una introgresion de la especie con Q. pringlei (Muller, en prensa). En
bosques de oyamel es un arbusto rizomatoso que junto con otros encinos y arbustivas forman un matorral
denso y de difícil acceso, denominado chaparral por Marroquín (1976) y Valdés y Aguilar (1983). Especie
común en las áreas donde habita.
Notas.—Se caracteriza por las hojas coriáceas, ovadas a elípticas u oblongas, de menos de 5 cm de
largo, margen entero o con 3-6 pares de dientes mucronados, haz verde oscuro y envés con indumento
café-amarillento. Escamas de la cúpula con base engrosada. Puede confundirse con Quercus coahuilensis, Q.
hintoniorum y 2 b Li
M 1 i de Arteaga, El Coahuilón, 2,940 m, 14 may 2000, G.B. Hi 27538 (ANSM): Los Oyameles,
(=)
2,800 m, 6 mar 2003, E Estrada 15271^ (ANSM, CFNL); sierra Hermosa (sierra de Artega), cerro La Campana, 2 470 7 m, 27 jun 1979,
L. Arce s.n. (ANSM); La Siberia, 25?15'N y 100°29'W, 2,800 m, 6 sep 1994. J.A. Villarreal et al. 116 (ANSM); Las Vigas, cañón de Jamé,
rancho El Tirol, 25?26'N y 100°30'W, 2,450 m, 8 may 1995, M.A. Carranza y J.A. Encina 2164 (ANSM). Mpio. Cuatro Cienegas: sierra
de La Madera, cañón del Invierno (El Infiernillo), 27°04'N y 102°28'W, 2,750 m, 28 ago 1974, T. Wendt 634 (ANSM). Mpio. General
Cepeda: sierra La Concordia-Pilote de Fierro, ejido La Casita, 25°10'N y 101°25'W, 2,950 m, 24 oct 1991, M.A. Carranza y L. Garcia 1252
(ANSM). Mpio. Parras: sierra de Parras, arroyo Seco, 2,000 m, 12 abr 1981, A. Rodriguez y M.A. Carranza 283 (ANSM). Mpio. Saltillo:
sierra de Zapalinamé, El Penitente, 25°21'N y 100°54'W, 2,710 m, 4 may 2004, J.A. Encina et al. 1271 (ANSM); sierra La Concordia (sierra
Catana), 25°15'N y 101°12'W. 2,750 m, 29 ago 1992, J.A. Villarreal et al. 6556d (ANSM).
pie grisea Liebm., Overs. Kongel. Danske Vidensk. Selsk. Forh. Medlemmers Arbeider. 171. 1854. (Fig.
11). Tro: E.U.A. Texas: C. Wright 665 (HoLoripo: G-DC; isoripos: GH, MO, NY).
Arbusto o árbol de 3—7(-10) m de alto, corteza ania gris claro; ramillas de 1-2 mm de grosor, con indu-
mento gris amarillento, glabrescente; ovoides de 2 mm de largo, glabras, café rojizo; estípu-
las persistentes, subuladas o aA de 3-6 mm de BEBO. puces a deciduas o subpersistentes,
coriáceas; pecíolos de 3-10 mm de largo, tomentosos; limb a elípticos u ovados, de (1.5-)3-58)
cm de largo, de (0.5—)1.5—3(-4) cm de ancho, ápice agudo a chases mucronado, base cordada, redondeada
o cuneada, margen un poco revoluto, entero o con dientes cortos terminando en un mucrón, haz verde
opaco, glabro, con pelos multirradiados muy esparcidos y diminutos, envés verde grisáceo o blanquecino,
opaco, con pelos multirradiados densos; nervaduras 6 a 10 en cada lado, muy ramificadas y anastomosadas,
un poco salientes en el haz, prominentes en el envés. Inflorescencias masculinas de 2-4(—7) cm de largo,
flores vellosas. Inflorescencias femeninas de 0.5-4 cm de largo con 1-8 flores tomentulosas. Fruto anual,
solitario o en pares, subsésiles o en pedúnculos de hasta 2-3 cm de largo; cúpulas hemisféricas, de 8-15
mm de ancho, de 4-10 mm de alto, escamas de la cúpula ovadas a oblongas, base ligeramente engrosada a
tuberculada, con tomento canescente, ápice delgado, apresado, café rojizo con puberulencia esparcida, las
escamas del margen de la cúpula elongadas; nuez ovoide o elipsoide, de 12-18 mm de largo, de 8-12 mm
de ancho, café claro, glabra o con indumento fino, incluida cerca de la mitad en la cúpula.
Distribución.—Se distribuye desde Arizona, Nuevo México, oeste de Texas, noroeste de Nuevo León,
norte de Chihuahua, Durango, sur de Zacatecas y norte de Jalisco. En Coahuila se encuentra al norte y centro
del estado en las sierras del Carmen, Las Cruces, del Fuste, del Pino, La Encantada, La Paila, de Arteaga y
Zapalinamé, en matorral submontano, bosque de encino y de pino-encino. Se asocia con Pinus remota, P.
arizonica var. stormiae, Acer grandidentatum, Quercus emoryi, Q. mohriana, Q. gravesii, Q. intricata, Q. laceyi,
Juniperus flaccida, Crataegus greggiana, Ungnadia speciosa, Garrya ovata, Fraxinus cuspidata, F. berlanderiana,
Dasylirion cedrosanum, Cercocarpus montanus y Arbutus xalapensis. Crece entre los 1400-2350 m de altitud,
en macizos tafi con laderas bajas, colinas rocosas secas, valles o en fondos de cañones con suelos de
rocas ígneas o calizas. Es frecuente en exposiciones noroeste o suroeste de las montañas del norte y común
en la sierra de La Paila, así como en algunas localidades en las sierras del sureste. A orillas de arroyos gen-
eralmente forma colonias pequeñas de arbolitos o árboles de hasta 10 m y en colinas secas forma matorrales
Villarreal et al., Q (F ) de Coahuil 1253
Fic, 9. Quercus gravesii. A, hojas y bellotas; B, variación en hoja; jui m Quercus greggii. A, rome con bojas, amentos y bellotas; B,
C
J
lado: D, bellota: E de la cúpul ja; G p D, bellota; E, escama
de la cüpula.
h cm
n
ybe D
| 3 | I iculado: D, bellota: E, escama
; Vr] 7 Vy ; E,
Fic. T1. Quercus grisea. A, rama con hojas, amentos y bellotas; B, variación Fic. 12. Q hintoni A, rama con hojas,
n H I la: Je J n hallata: E A I 4 1 1 S LN
de la cúpula.
Ingel Dat . [|
1254 Journal UI R h institut f Texas 2(2)
densos de 13 m de alto. Especie común en las áreas donde habita.
Notas.—Se reconoce por las hojas coriáceas, elípticas a lanceoladas, usualmente de menos de 8 cm
de largo, ápice agudo y mucronado, margen generalmente entero co con pocos dientes mucronados, haz
verde opaco y envés grisáceo con indumento denso de pelos multirradiados. Escamas de la cúpula con base
engrosada. Se le puede confundir con Quercus mohriana.
Material inado: Mpio. Candela: si de Paj Azul S Maria, 27?01'N y 100°50'W, 3 jun 1997, P. Cruz 07 (ANSM).
Mpio. General Cepeda: sierra La Concordia, 4.5 A al NW del sido La Casita, 25°14'N y 101?25'W, 10 oct 1996, J.A. Encina y J. Zavala
259 (ANSM). Mpio. M del Carmen, Los Lirios, caf poral, 25°54'N y 102°30'W, 15 sep 1992, J.A. Villar-
real y M.A. Carranza 7042 (ANSM). Mpio. Ocampo: sierra del Pino, ejido dicebüches cañón La Vaca, 28?15'N y 102°59'W, 2 oct 2003,
M.A. Carranza e I. Ramirez 3957 (ANSM); sierra La Encantada, cuesta Malena, 28°54'N y 102°30'W, 29 mar 1992, M.A. Carranza y J.
Noriega 1515 (ANSM). Mpio. Parras: sierra Playa Madero, cañón Pinalosa, ejido Santa Victoria, 25°09'N y 101°24'W, 6 jun 1995, M.A.
Carranza y J.A. Encina 150 (ANSM). Mpio. Ramos Arizpe: sierra i La ien ejido El Cedral, valle Loma zem nba 101°35'W, 1
may 1991, J.A. Villarreal et al. 5825 (ANSM) Mpio. S Itill 6, cañón Boca Negra, del Cuatro, 25°21'N
y 100°55'W, 21 ago 2004, J.A. Encina et al. 1354 (ANSM); Mipo. DH rancho La Escondida, 28°55! N y 101°51'W, 6 may 2003, M.
Guerrero y O. Ballesteros s.n. (ANSM).
Quercus hintoniorum K. C. Nixon et C.H. Mull., Brittonia 45:147. 1993. (Fig. 12). Two: MÉXICO. Coanuma: Los
Lirios, R.M. Straw y M. Forman 1361 (HoLotIPO: US).
Arbusto rizomatoso o arbolito de hasta 5 m de alto, corteza gris oscura, lisa; ramillas café rojizas de 1-2
mm de grosor, con puberulencia esparcida, volviéndose glabras; yemas ovoides, cerca de 2 mm de largo,
café oscuro; estípulas lineares, cerca de 5 mm de largo. Hojas subpersistentes, pecíolos de 3-6 mm de largo,
glabros, rojizos; limbos elípticos u obovados, de (223-6 cm de largo, de 1-4 cm de ancho, ápice redondeado
o subagudo, base fuertemente cordada, raras veces da, margen plano o revoluto, sinuado, sublobulado,
raras veces entero, con (4-)5-8 dientes irregulares en cada lado, especialmente hacia el ápice de la hoja,
dientes con aristas de 1-2 mm largo, la hoja frecuentemente conduplicada arriba del pecíolo, haz verde os-
curo, moderadamente lustroso con pelos fasciculados esparcidos, glabrescente, envés con pelos glandulares
uniseriados y pelos fasciculados, estipitados y esparcidos, la superficie claramente visible, con pequeños
parches densos de pelos fasciculados y erectos restringidos a lo largo de las axilas de la nervadura media y
las nervaduras secundarias; nervaduras laterales 4—5 en cada lado, irregulares y desiguales, frecuentemente
ramificándose cerca del margen. Inflorescencias masculinas de 5-6 cm de largo. Inflorescencias femeninas
de 1 o 2 flores sobre pedúnculos tomentosos de 2-7 mm de largo. Fruto bianual, cúpula turbinada, escamas
de la cúpula con puberulencia amarillo-rojiza, nuez hemisférica a ovoide, incluida un tercio a un medio en
la cúpula.
Distribución.—Se conoce del sureste de Coahuila (Sierra de — y en Nuevo Leon (en los municipios
de Santiago, Galeana y Rayones). En Coahuila se le encuentra en localidades de la Sierra Madre Oriental como
La Siberia, Las Vigas, El Coahuilon, La Marta, San José de los Nuncios y en las proximidades de Los Lirios.
Es común en bosque de pino-encino y oyamel, así como en los limites de la vegetación alpina-subalpina. Se
asocia principalmente con Quercus greggii, Q. fulva, Q. sa Miet Bun setae Ors P greggii, a
menziesii, Abies vejarii, Populus tremuloides, Garrya ovata, C
y Prunus serotina. Crece entre los Rd p m de altitud, en laderas Medias y altas con O ios note y
blaci i ,enl Se presenta
noroeste, donde forma |
como arbusto PEN de 2-3 m de alto en cimas a laderas m y cañones húmedos. Especie
frecuente en las áreas donde habita, es considerada endémica para el área de Megacoahuila (Villarreal y
Encina 2005).
Notas.—Se puede reconocer por las hojas coriáceas, elípticas a obovadas, de menos de 6 cm de largo,
ápice redondeado a obtuso, margen conduplicado con 4-8 pares de dientes con aristas cortas y envés con
pelos glandul y pelos fasciculados en las nervaduras. Escamas de la cúpula casi planas. Relacionado con
Quercus greggii, Q. coahuilensis y Q. hypoxantha.
Material examinado: Mpio. Arteaga: sierra de Arteaga, La Siberia, 25°15'N y 100°29'W, 2,400 m, 7 sep 1995, J.A. Encina y M.A. Car-
ranza 176 (ANSM); sierra Las Vigas, cañón de Jamé, 25°26'N y 100°30'W, 2,450 m, 8 may 1995, M.A. Carranza y J.A. Encina 140 (ANSM).
Mpio. Ramos Arizpe: sierra San José de los Nuncios, 1,765 m, 19 jul 1991, G.B. Hinton 21097 (ANSM, TEX).Mpio. Saltillo: sierra de
Zapalinamé, cerro el Penitente, 25?21'N y 100°54'W. 2,710 m, 15 nov 2004, J.A. Encina et al. 1381 (ANSM
Villarreal et al., Q (F ) de Coahuil 1255
Quercus hypoleucoides A. Camus, Bull. Mus. Hist. Nat. Paris. Ser. 2(4):124. 1932. (Fig. 13).
Quercus confertifolia Torr., U.S. Mex. Bound. Surv. Rpt. 207. 1840.
Quercus hypoleuca Engelm., Trans Acad. Sci. St. Louis 3:384. 1876.
Árbol de 10 a 20 m de alto; corteza dura, hendida y negra; ramillas rojas, de 2-3 mm de grosor, lenticelas
poco evidentes, con indumento gris, glabrescentes; yemas ovoides, de 3 mm de largo y de 2 mm de ancho,
café rojizas; estipulas caedizas. Hojas persistentes, gruesas y coriáceas; pecíolos de 5—7(-15) mm de largo,
pubescentes, glabrescentes; limbos lanceolados, de 5-8-11) cm de largo, de 1.5-3.0 cm de ancho, ápice
agudo, terminando en arista, base redondeada a cuneada, margen fuertemente revoluto, entero o con po-
cos dientes terminados en aristas, haz verde oscuro o glauco, glabro o puberulento; envés con indumento
blanquecino o canescente denso, de pelos fasciculados sesiles o cortamente estipitados; nervaduras 10-12
en cada lado, con algunas intermedias que se desvanecen, ramificadas y no evidentemente anastomosadas
cerca del margen, algo impresas en el haz, salientes y nervadura media glabra en el envés. Inflorescencias
masculinas de 3—6 cm de largo, blanco pilosas. Inflorescencias femeninas de 5-10 mm de largo, 1-2 flores
en la parte terminal, pedúnculo con pelos fasciculados. Fruto anual o algunas veces bianual, solitario o en
pares, sobre pedúnculos de hasta 15 mm de largo; cúpula hemisférica, de 8-12 mm de diámetro y 7-10
mm de alto, con escamas obtusas delgadas y apresadas, tomentulosas a glabras, café pálido; nuez oblonga
a ovoide de 12-14 mm de largo y de 7-9 mm de ancho, puberulenta, glabrescentes, incluida cerca de un
tercio en la cúpula.
Distribución.—Se distribuye desde Arizona y Nuevo México, oeste de Texas hasta la Sierra Madre Oc-
cidental en Sonora y Chihuahua. En Coahuila solo se encuentra en la región de la sierra del Carmen en el
bosque de encino y bosque de pino-encino. Se asocia con Quercus gravesii, Q. laceyi, Q. arizonica, Q. muehlen-
bergii, Q. arizonica, Q. rugosa, Pinus strobiformis, P. remota, Pseudotsuga menziesii, Cupressus arizonica, Prunus
serotina, Juniperus deppeana, J. flaccida, Garrya ovata, Arbutus xalapensis, Salvia regla, Acer grandidentatum y
Tilia mexicana. Crece en elevaciones de 1700—2500 m sobre laderas altas y cafiones, con exposiciones norte
o noroeste, en cañones húmedos a través de arroyos (cañón del Centinela en la sierra del Carmen), como
árbol de hasta 20 m de alto. Especie considerada rara en el área de Coahuila.
Notas.—Se reconoce por las hojas gruesas, lanceoladas, de 5-10 cm de largo, con apice agudo y aristado,
margen entero o con pocos dientes aristados, haz verde y envés canescente, con indumento denso de pelos
fasciculados, venación prominente. Escamas de la cúpula planas. Relacionado con Quercus sideroxyla,
Material examinado: Mpio. Acuña: roadside on Maderas del Carmen, 2,300 m, 13 jul 1997, S. Wood, G. Harper y D. Doan-Crider s.n.
(ANSM), Mpio. Ocampo: sierra del Carmen, 1950 m, 17 sep 1989, A.E. Estrada 1817 (ANSM); sierra del Carmen, 28°58'N y 102°33'W,
2,200 m, 12 ago 1975, M.P. Robert y J. Passini 6583 (ANSM); El Jardín, exposición N de la sierra del Carmen, 2,200 m, 8 jun 1991, J.A.
Pérez 1766 (ANSM).
Quercus hypoxantha Trel., Mem. Natl. Acad. Sci. 20:170, pl. 339. 1924. (Fig. 14). Tiro: MÉXICO. Coamura:
mountains near Saltillo, C. G. Pringle 10227 (notorio: GH).
Quercus errans Trel. f. graciliramis C.H. Mull., J. Arnold Arbor. 17:169. 1936)
Arbusto de hasta 2 m de alto, corteza gris obscura, lisa; ramillas de 0.5-1 mm de grosor, con puberulencia
café-amarillenta, densa y persistente; yemas ovoides, cerca de 3 mm de largo, glabras, amarillo-rojizo;
estípulas de 2 mm de largo. Hojas subpersistentes; pecíolos de 3-6 mm de largo, glabros, rojizos; limbos
obovados a orbiculares, de (3—)4—5.5 cm de largo y (1. P a 5 cm de ancho, ápice redondeado a obtuso,
base fuertemente cordada, el limbo f lo, margen ondulado, revoluto, con 3-4(-6)
dientes aristados irregulares en cada lado especialmente d el ápice de la hoja, dientes irregularmente
orientados, con aristas de 1-2 mm largo, haz con tricomas fasciculados diminutos, esparcidos, envés con
indumento de pelos fasciculados estipitados, glabrescente; nervaduras 4—5(-6) en cada lado, irregulares
y desiguales, frecuentemente ramificadas cerca del margen. Inflorescencias masculinas 5-6 cm de largo.
Inflorescencias femeninas 1 o raramente 2 flores sobre pedúnculos tomentosos de 1-4 mm de largo. Fruto
bianual, solitario, sobre pedúnculos de 2-6 mm de largo; cúpula hemisférica, de 10-12 mm de ancho y 7-8
1256 J | of the Botanical R h Institute of Texas 2(2)
mm de alto, escamas de la cápula delgadas y poco apresadas, ápice redondeado, café claro, puberulentas;
nuez ovoide, de 14-18 mm de largo y 10 mm de ancho, café claro, con puberulencia esparcida.
Distribución.—Se distribuye en la Sierra Madre Oriental, desde San Luis Potosi, hasta Nuevo León. En
Coahuila solo se encuentra en el sureste del estado en las sierras de Arteaga y Zapalinamé. Crece en el ma-
torral submontano, así como en los límites con el bosque de pino y oyamel, asociándose con Quercus greggii,
Q. striatula, Purshia plicata, Garrya ovata, Arctostaphylos pungens, Pinus arizonica var. stormiae, Rhus virens y
Cercocarpus fothergilloides. En bosques de oyamel se asocia con Pseudotsuga menziesii, Cupressus arizonica y
Abies vejarii. Se localiza desde los 1900-2850 m de altitud, en laderas con exposiciones norte, noroeste o
noreste, generalmente sobre terrenos con roca madre expuesta, donde forma matorrales. Es comün en la
localidad del Penitente en la sierra de Zapalinamé. Especie considerada como rara para el estado.
Notas.—Se puede reconocer por las hojas coriáceas, obovadas a orbiculares, usualmente de menos de 5
cm de largo, ápice redondeado a obtuso, margen conduplicado con 4-6 pares de dientes con aristas cortas
y envés con indumento denso de pelos fasciculados. Escamas de la cápula planas. Relacionado con Quercus
greggii, Q. coahuilenses y Q. hintoniorum.
Material examinado: Mpio. Arteaga: predio Santa Anita, sierra ae Nuevo, A N de San Juan de Dolores, 25?26'N y 100°34'W,
2,800 m, 27 ago 2005, J.A. e 1410 (ANSM). Mpio. Saltil é, cañón de San Lorenzo, 25?23'N y 101°00'W,
1980-2060 m, 3 ago 1995, M.A. Carranza 2222 (ANSM) si de Zapali $ El] Penitente, exposición NW, 25?2]'N y 100°54'W, 2,710
m, 15 nov 2004, J.A. Encina et al. 1267, 1382, 1395 (ANSM).
Quercus intricata Trel., Contr. U.S. Natl. Herb. 13:185. 1922. (Fig. 15). Tiro: MÉXICO. Coanuna: Buenavista, J, Gregg
256 (HoLoriro: US; isoripo: GH).
Quercus microphylla Nee var. crispata A. DC., Prodr. 16(2):36. 1864.
Arbusto de 0.5-3 m de alto, gregario, intrincadamente ramificado; ramillas de 1-1.5 mm de grosor, con to-
mento corto, grisáceo-amarillento; yemas esféricas, de 1-1.5 mm de largo, café-rojizo; estipulas persistentes,
subuladas, de cerca de 2 mm de largo, pubescentes. Hojas persistentes, gruesas y duras; pecíolos de 2-4
mm de largo, tomentosos; limbos ovados a usualmente oblongos o elíptico-oblongos, de 1-2(—5) cm largo
y 0.5-1.5 cm de ancho, ápice obtuso, base cordada o cuneada, margen fuertemente revoluto y a menudo
crispado, entero o con pocos dientes, haz lustroso, con pelos fasciculados esparcidos; envés con indumento
denso y persistente, amarillo cremoso, de pelos fasciculados estipitados, glabrescente; nervaduras 6-9 en
cada lado, impresas en el haz, salientes en el envés. Inflorescencias masculinas de 2-3 cm de largo, con
raquis tomentoso y flores muy esparcidas. Inflorescencias femeninas de 3-10 mm de largo, 1-5 flores con el
pedúnculo tomentoso. Fruto anual, solitario o en pares, subsésil o en pedúnculos tomentosos de hasta 1.5
cm de largo, cúpula cerca de 1 cm de diámetro y 7-8 mm de alto, escamas ovadas, canescentes y engrosadas
en la base, con el ápice delgado y rojizo, apresado; nuez ovoide, de 9-12 mm de largo y 8-10 mm de ancho,
casi glabra, color café claro, incluida de un medio a un tercio en la cúpula.
Distribución. —Con distribución desde el oeste de Texas, noreste de Chihuahua, hasta el centro-sur
de Nuevo León, Zacatecas y noroeste de Durango. En Coahuila se distribuye en el noroeste, centro y sur,
en casi todas las sierras. Es parte importante en el matorral submontano y en la transición con el mator-
ral rosetófilo y bosque de pino piñonero. Se asocia con Pinus cembroides, P. remota, Vauquelinia corymbosa,
Quercus striatula, Q. invaginata, Q. pungens var. vaseyana, Q. mohriana, Q. pringlei, Q. coahuilensis, Cercocarpus
Du E flaccida, E HE qe texana, Arctostaphylos pungens, Dasylirion cedrosanum, Rhus virens,
Ptelea trij : Amelanchier denticulata, Fraxinus greggii, Calia secundiflora y Mimosa texana.
Se desarrolla en decies adea apareciendo desde los 1600 hasta 2700 m de altitud, sobre laderas
bajas con exposición norte, noreste, este y suroeste sobre terrenos pedregosos. Abunda en los márgenes de
arroyos en el matorral submontano y en el estrato arbustivo en bosques abiertos de pino piñonero. Especie
común en toda el área. Es conocido localmente como encino chaparro o charrasquillo.
Notas.—-Se reconoce fácilmente por ser un arbusto con hojas gruesas, oblongas a elípticas, usualmente
de 1-3 cm de largo, margen revoluto, entero o con dientes, haz verde amarillento y envés blanco-cremoso
con indumento de pelos fasciculados, venación evidente. Escamas de la cúpula con base engrosada.
Villarreal et al., 0 (F ) de Coahuil 1257
Fic. 13. Q hypol les. A, hojas y bellota; B, variaciónen Fic, 14. Q h ha. À, rama con hojas, amentos y bellota; B,
hoja; C, pelo fasciculado; D, bellota; E de la cúpul variación en hoja; C, pelo fasciculado estipitado; D, bellota; E, escama
de la cúpula.
- = i
PN A aras
Fic. 15. Quercus intricada. A, hoi beil ; B, vari id hoi Fic. 16. Qi j g A, j y
lad ;D bellota; E, de la cünul hojas; C lo fasciculad ;D bellota: E le la cü
: 1o£tha Bataniral D h institute of Texas 2(2)
1258
Material lo: Mpio. Acuña: si del Carmen, cañón del Diablo, 29°11'N y 102°47'W, 10 ago 1974, T. Wendt 551 (ANSM, TEX).
Mpio. Arteag de Art 25?25'N y 101°47'W, 1,800 m, 2 nov 1996, J.A. Encina et al. 121 (ANSM).
ü i La Gavia TRR a Gavia, 26°10'N y 101*18' W, 1,500 m, 10 ag0 1995, M.A. Carranza y J.A. Encina 173 (ANSM).
Mpi
pam Cuatro Cienegas: sierra de La Madera, cañón del Invierno, 27°04'N y 102°28'W, 2,725 m, 28 ago 1974, T. Wendt y E. Lott 636
(ANSM, TEX); sierra San Marcos, ejido de Norias, cañón Grande, 26?30'N y 101?40"W, 2,500 m, 19 oct 1993, M.A. Carranza et al. 1852
(ANSM). Mpio. General Cepeda: sierra La Paila, valle de Parreños, 25?28'N y 101°25'W, 1,600 m, 12 sep 1989, J.A. Villarreal y M.A.
Carranza 464 (ANSM); sierra L a Concordia, ejido El Tejocote, 25?14'N y 101°37'W, 2,130 m, 13 jul 2002, J.A. Encina et al. 1251 (ANSM).
Mpio. Ocampo: rancho La Cruz, 27?15'N y 102°40'W, 1,800 m, 11 sep 1991, M.A. Carranza y L. García 1189 (ANSM); sierra El Pino
ejido Acebuches, cañón La Vaca, 28°15'N y 102°59'W, 1,850 m, 2 oct 2004, M.A. Carranza e I. Ramirez 4017 (ANSM). Mpio Parras:
ejido Tanque Nuevo, 25?15'N y 101°44'W, 1,950 m, 11 jul 2002, J.A. Encina et al. 1244 (ANSM). Mpio. Ramos Arizpe: sierra San José
de los Nuncios, 1,630 m, 19 jul 1991, G.B. Hinton 21075 (ANSM); sierra La Paila, cañón Loma Prieta, 25?51'N y 101°30'W. 1,600 m, 23
oct 1985, J.A. Villarreal et al. 3118 (ANSM). Mpio. Saltillo: sierra La Concordia, Carneros, 18 nov 1974, J. Marroquín 3048 (ANSM), sierra
del Duraznillo, ejido Tinajuela, 25%04'N y 101°17'W, 14 ago 2004, J.A. Encina et al. 1348 (ANSM), sierra La Concordia, ejido San Blas,
cañón Troncones, 23?17'N y 101°18'W, 2 oct 1994, M.A. Carranza et al. 3157 (ANSM).
Quercus invaginata Trel., Mem. Natl. Acad. Sci. 20:87, pls. 137, 138. 1924. (Fig. 16). Tiro: MÉXICO. Ramos
Arizpe, sierra La Paila, C. A. Purpus 5029 (HoLorIPO: B; IsoTIPOS: G O).
Quercus invaginata Trel. f. purpusiana Trel., Mem.Natl. Acad. Sci. 20:87, pl. 138. 1924. Tiro: MÉXICO: Ramos Arizpe, sierra La Paila, C.
30 sorro: NY))
Arbusto o árbol de 1—3(-9) m de alto, con corteza gris escamosa o surcada; ramillas con indumento blan-
quecino, denso y persistente; yemas ovoides, de 1-3 mm de largo, café rojizo; estípulas setáceas y deciduas
de 2-3 mm de largo. Hojas coriáceas deciduas; pecíolos de (325-8 mm de largo, gris tomentoso; limbos
obovados a oblongos o elípticos, de (2-)3—5(-6) cm de largo y (1-)2-3.5 cm de ancho, ápice redondeado a
subagudo u obtuso, base cuneada a obtusa o redondeada, rara vez subcordada, margen ondulado, crispado,
con dientes toscos mucronados o raramente subentero; haz verde claro y opaco, con pelos fasciculados
esparcidos, envés con indumento denso blanco-grisáceo de pelos fasciculados; nervaduras 6-10 en cada
lado, anastomosadas y terminando en un mucrón, poco impresas en el haz, muy prominentes en el envés.
Inflorescencias as de 2-3 cm de largo. Inflorescencias femeninas de 5-10 mm de largo, con 1-3
10 , hasta de 3 mm de largo. Fruto anual, solitario o en grupos de 23, subsésil
flores en 1
o sobre un pedánculo de 5- 1017) mm de largo, cápula turbinada, de 10—20 mm de diámetro, al madurar
con los márgenes marcadamente involutos, escamas agudas con tomento grisaceo, base algo engrosada;
nuez elíptica a oblonga, de 10—15 mm de largo, café claro, glabra excepto en la parte apical, incluida cerca
de un tercio en la cúpula.
Distribución.—Se distribuye en el centro y noroeste de Coahuila, en las sierras de La Paila, La Gavia,
La Madera y del Pino, y en el centro-norte de Nuevo León, como un componente importante en matorral
submontano, en la ecotonía con el matorral rosetófilo, así como en bosques de pino-encino y de encino. Se
asocia con elementos de Pinus remota, P. arizonica var. stormiae, Quercus pringlei, Q. gravesii, Q. intricata, Q.
laceyi, Q. coahuilensis, Fraxinus greggii, Ungnadia speciosa, Vauquelinia corymbosa, Pistacia texana, Juniperus
flaccida, Dasylirion cedrosanum, Calia secundiflora, Gochnatia hypoleuca, Ptelea trifoliata, Cercocarpus montanus,
Acacia berlandieri y Colubrina greggii. Crece en altitudes de 1600—2200 m, presentándose disperso desde
los 1250 m, sobre laderas bajas con exposiciones norte, noroeste y en fondo de cañones, forma poblaciones
arbustivas densas en el matorral submontano, es frecuente en los bosques de encino y de pino-encino. En el
matorral rosetófilo se presenta en forma esparcida. Abunda en las sierras del centro del estado y disminuye
con el aumento de la altitud y la latitud. Marroquín (1985) la reporta para el noroeste de Nuevo Léon, fre-
cuente en el cauce de los arroyos, sobre laderas de exposición oeste entre los 600—1700 m de altitud. Especie
considerada como endémica para el área de Coahuila y comün en los lugares donde habita.
Notas.—Se reconoce por las hojas coriáceas, obovadas, oblongas a elípticas, usualmente de menos
de 5 cm de largo, ápice redondeado a obtuso, margenes ondulados a crispados, dentado-mucronados, haz
verde, glabro y envés con indumento de pelos fasciculados. Cracterísticamente la cápula presenta el margen
involuto y las escamas con la base ligeramente engrosada. En algunos casos pueden presentarse semejanzas
foliares con Q. grisea.
Villarreal et al., Q (F ) de Coahuil 1259
Material examinado: Mpio. panis sierra Pájaros Azules, campo Santa María, cañada La Sabanilla, 27°02'N y 100°54'W, 5 dic 1997,
M.A. Carranza y D. Sánchez 2882 (ANSM). Mpio. Castaños: sierra La Gavia, cañón La Gavia, 26?20'N y 101°15'W, 10 sep 1987, J.A.
ee y M. a Carranza 4037 TAN sierra La ae canón Chilpitin, 26°48'N y ae W, 7 Sp 1976. T. Wendt y D. Riskind 1684
(ANSM, TEX). Mpio. C il , cañón Grande, ejido E 27?40'N y 102°50'W, 1 oct 1993,
M.A. Carranza et al. 104 (ANSM), sierra La Madera, mouth of cañada Los Posos, 27?07' N y 102°27'W, 26 jul 1977, T. Wendt y J. Valdés
1052 (ANSM). Mpio. Monclova: sierra La Gloria, cañada La Tienda, 26°56'N y 101°25'W, 11 ago 1995, M.A. Carranza y J.A. Encina 2376
(ANSM). Mpio. Ocampo: sierra La Encantada, cuesta Malena, 28°54'N y 102°30'W, 29 mar 1992, M.A. Carranza et al. 1514 (ANSM).
Mpio. Ramos Arizpe: sierra La Paila, cañón El Diente, 26°09'N y 101°30'W, 5 oct 1988, J.A. Villarreal y M.A. Carranza 4666 (ANSM).
rs laceyi Small, Bull. c Bot. Club 28:358. 1901. (Fig. 17). Tiro: E.U.A. Texas. Kerr Co.: Lacey’ ranch near
errville, H. Lacey s.n. (HoLoTIPo: NY).
Quercus breviloba Sarg. f. laceyi (Small) Trel., Mem. Nat. Acad. Sci. 20:102. 1924. Quercus breviloba Sarg. ssp. laceyi (Small) A. Camus,
Chenes 2:280. 1939
Arbusto o árbol de 5-8(-10) m de alto; corteza gruesa, fisurada y gris; ramillas de 1-3 mm de grosor,
acanaladas, pubescentes, pronto glabrescentes, café rojizo; y ovoides a cónicas de 2-6 mm de largo, con
indumento esparcido; estípulas deciduas, subuladas, de 5 mm de largo. Hojas delgadas, deciduas, glaucas;
pecíolos de 3-10 mm de largo, delgados, glabros o con indumento esparcido; limbos oblongos, elípticos a
obovados, de 3—7(-10) cm de largo y 2.5—4(-6) cm de ancho, ápice redondeado u obtuso, base redondeada o
cuneada, rara vez cordada, margen cartilaginoso, irregularmente sinuado o con pocos lóbulos en cada lado,
raramente dentado o entero; haz glabro, o con pelos fasciculados escasos, verde azulado, grisáceo, ligeramente
brillante, envés con pelos fasciculados escasos, glabrescente, verde amarillento o glauco; nervaduras 7-9 en
cada lado, irregulares, ramificadas y anastomosadas, poco salientes en ambas superficies. Inflorescencias
masculinas de 3-6 cm de largo. Inflorescencias femeninas de 5 mm de largo, con 1-3 flores. Fruto anual,
solitario o en pares, subsésil o sobre pedúnculos de 1 cm de largo, cúpula de 15-18 mm de ancho, de 4-7
mm de alto, escamas de la cúpula ovadas, base engrosada, con poco tomento y ápice delgado poco apresado,
glabro, café rojizo; nuez cónica, ovoide o subcilíndrica de 1.5-2(3) cm de largo y 9-14 mm de ancho, glabra
excepto cerca del Due in claro, incluida menos de la mitad en la cúpula.
Distribución. el centro (Planicies Edwards) y oeste (Trans-Pecos) de Texas, en México a
través de la Sierra Madre Oñenal en Coahuila, Nuevo León y probablemente en Tamaulipas. En Coahuila se
encuentra li distribuido en la sierras del noroeste y centro, desde la sierra del Carmen, serranias
del Burro, det Pino, de San Marcos A la sierra La Paila. En las sierras de Zapalinamé y La Concordia, se
encuentran poblaciones aisladas. Es una especie propia del bosque de pino-encino, así como del bosque de
encino, asociándose con especies de Quercus gravesii, Q. invaginata, Q. mohriana, Q. grisea, Q. muehlenbergii,
Q. emoryi, Q. hypoleucoides, Rhus virens, Juniperus flaccida, Pinus arizonica var. stormiae, P. remota, Arbutus
xalapensis, Cercis canadensis, Prunus serotina, Dus mexicana, ol viscosa, Acer grandidentatum, Ungnadia
speciosa, Fraxinus berlanderiana, Pistacia t y] pa. Crece en altitudes entre
1000-2500 m, sobre laderas con exposición norte, noroeste u dete en fondo de cari Se presenta como
arbolito en laderas pedregosas con suelos someros, mientras que en cañones protegidos con humedad y a
orillas de arroyos es un árbol. Es abundante en las sierras del centro-norte del estado, raro en las sierras del
sur del estado. Especie común en el área donde habita.
Notas.—Se reconoce por las hojas delgadas, glabras, verde azulosas, oblongas a obovadas, de hasta 10
cm de largo, ápice obtuso a redondeado, margen sinuado o con lóbulos cortos y redondeados, nervación no
resaltada. Escamas de la cúpula con base engrosada.
Quercus laceyi ha sido reconocido y separado de Q. glaucoides, con el cual se ha confundido en algunos
tratamientos florísticos. Esta última especie se ha encontrado que restringe su distribución al centro y sur
de México (Nixon y Muller 1992).
Mat do: Mp ü ias del Burro, rancho El Bonito, 29*01'N y 102°07'W, 1,625 m, 19 sep 1977, D. Rishind y C.H.
Muller 2200 (ANSM, E Mpio. Candela: sierra de Pajaros Azules, campo Santa María, 27%01'N y 100°50'W, 1,800 m, 4 jun 1997, P.
Cruz 10 (ANSM). Mpio. Castaños: sierra La Gavia, rancho La Gavia, 26?10'N y pod. 1,295 m, 10 ago 1995, M.A. Carranza y J.A.
Encina 171 (ANSM). Mpio. Cuatro Cieneg San Marcos, ejid q Norias, cañón Grande, 27°40'N y 102°50'W, 1,550
1260 J | of the Botanical R h Institute of Texas 2(2)
m, l oct 1993, M.A. Carranza et al. 105 (ANSM); sierra La Madera, cañón La Hacienda, 27?05'N y 102°25'W, 1,600 m, 11 may 1973
(ANSM, TEX). Mpio. General Cepeda: sierra La Concordia (sierra El d as E Mimbre, ejido El Na 25°16'N y 101°
37W, 1,900 m, 6 jun 1995, M.A. Carranza y J.A. Encina 2194 (ANSM). M Moncl Castaños), sierra La Gloria, cañón
El Cono, 26?49'N y 101917'W, 1,220 m, 6 sep 1976, T. Wendt y D. Rishind 1628 (ANSM, TEX). Mpio. M iz: si S Rosa, rancho
El Silencio, 27°55'N y 101°50'W, 1,300 m, 2 may 2004, S. Lara y R. Pineda 100 (ANSM), sierra Santa os (La Babia), rancho La Babia,
cañón Rincón de María, 28°27'N y 102°04'W, 1,450 m, 17 sep 1999, J.A. Villarreal et al. 8767 (ANSM). Mpio. Ocampo: sierra del Pino,
po Acebuches, cañón La Vaca, dud y 102°59'W, 1 dnd m, sep 2003, M.A. Carranza e I. Ramirez 3982 uus sierra 2
102?
en, canyon La Fronteriza, I Alamo, about 0.8 mi N of rancho San Isidro, 28? 54'N y 32W,
1510 m, 12 aug 2004, D. Riskind, J Valdés y J. HEN in 23858, 23865 (ANSM, TEX). Mpio. Ramos Arizpe: sierra La E camino a El
Cedral, 25°59'N 101?28"W. 1,250 m, 8 oct 2005, J. A Villarreal e I. Ramirez 9291 (ANSM). Mpio. Saltillo: sierra de Zapalinamé, cañón
por el camino a El Cuatro, 25%21'N y 101°55'W. 2,100 m, 21 ago 2004, J.A. Encina et al. 1352 (ANSM). Mpio. Zaragoza: serranias del
Burro, rancho La Escondida, 28°56'N y 101°51'W, 1,200 m, 6 may 2003, M. Guerrero y O. Ballesteros s.n. (ANSM)
Quercus laeta Liebm., Overs. Kongel. Danske Vidensk. Selsk. Forh. Medlemmers Arbeider. 179. 1854. (Fig.
18). Tro: MÉXICO. Hidalgo: Casa Grande, Hartweg 419 (notorio: B).
Quercus centralis Trel., Mem. Natl. Acad. Sci. 20:61. 1924.
Quercus prinopsis Trel., Mem. Natl. Acad. Sci. 20:61, pl. 58. 1924.
Quercus obscura Trel., Mem. Natl. Acad. Sci. 20:71. 1924.
Quercus transmontana Trel., Mem. Natl. Acad. Sci. 20:71, pl. 89. 1924.
Quercus pallescens Trel., Mem. Natl. Acad. Sci. 20:89, pl. 141. 1924.
Quercus bipedalis Trel., Mem. Natl. Acad. Sci. 20:89, pl. 143. 1924
Quercus clivicola Trel. et C.H. Mull., Bull. Torrey Bot. Club 66:149. 1936.
Árbol de 5—8(-15) m de alto, raramente arbusto; corteza café rojiza o negra; ramillas de 1.5-3 mm de
grosor, acanaladas, con indumento amarillo y pelos glandulares, lenticelas abundantes; yemas ovoides, de
2-3 mm de largo; estípulas membranosas, pilosas, de 4-6 mm de largo, caedizas. Hojas coriáceas, deciduas,
verde amarillentas o rojizas; pecíolos de 2-5(-8) mm de largo, pubescentes; limbos elípticos a oblongos,
lanceolados u obovados, de 5—9(-15) cm de largo y 2-3(-6.5) cm de ancho; ápice obtuso, redondeado o
acuminado, base redondeada o cordada, margen engrosado y revoluto, entero o con 2-11 dientes mucrona-
dos; haz verde claro y lustroso, con pelos fasciculados diminutos y pelos glandulares simples; envés pálido
con indumento amarillo de pelos fasciculados, cortamente estipitados y pelos glandulares rojizos a lo largo
de la nervadura principal, epidermis ampulosa; nervaduras 7-12 en cada lado. Inflorescencias masculinas
de 4-6 cm de largo, pilosas y con pelos glandulares. Inflorescencias femeninas de 1-3 flores. Fruto anual,
solitario o en grupos hasta de tres, de 1-24) cm de largo, cúpula hemisférica, de 12-15 mm de diámetro,
escamas engrosadas y pul en la base, ápice estrecho, obtuso y apresado, nuez ovoide de 10-18 mm
de largo, incluida un medio o un tercio en la cúpula.
Distribución. —Se distribuye en la República Mexicana en los estados de Durango, Sinaloa, Zacatecas,
Aguascalientes, Nayarit, Jalisco, Michoacán, Guanajuato, Hidalgo, Distrito Federal, Puebla, San Luis Potosí
y Nuevo León. En Coahuila se le encuentra en el centro y sureste, en las sierras La Paila, La Concordia,
Zapalinamé y de Arteaga, en bosques húmedos de encino y bosque de pino-encino. Se asocia con Pinus
arizonica var. stormiae, Juniperus flaccida, Quercus saltillensis, Q. laceyi, Cupressus arizonica, Arbutus xalapensis,
Cercocarpus montanus, Garrya ovata y Crataegus baroussana. Crece en altitudes de 1400-2900 m, sobre laderas
bajas, con exposición norte, en cañones y valles i suelos profundos o bien en los márgenes
de arroyos, donde se presenta como un árbol, mientras en laderas con suelos someros es común encontrarlo
como un arbusto o arbolito. Es frecuente en las sierras de Zapalinamé, La Concordia y de Arteaga, raro en
la sierra La Paila. Especie común en los lugares donde habita.
Notas.—Se reconoce por las hojas coriáceas, verde amarillentas, elípticas a oblongas, de hasta 15 cm
de largo, ápice obtuso, margen entero o con 2-11 pares de dientes, haz casi glabro y envés con indumento
de pelos fasciculados, venación prominente. Escamas de la cápula con base engrosada, prolongada hacia el
ápice.
Material examinado: Mpio. Arteaga: sierra de Arteaga, Potero de Abrego, camino a Los Lirios, 25?20'N y 100°28'W, 2,300 m, 30 abr
1987, J.A. Villarreal y M.A. Carranza 3681 (ANSM). Mpio. General Cepeda: sierra La Concordia, Pilote de Fierro, ejido La Casita, 25°10'N
y 101°25'W, 2,950 m, 13 ago 1993, J.A. Encina et al. 101 (ANSM). Mpio. Ramos Arizpe: sierra La Paila, ejido El Cedral, 26°02'N y
Villarreal et al., Q (F ) de Coahuil 1261
rx vez
D z La?
Fic. 18. Quercus laeta. A
Fic. 17. Quercus laceyi A, j " j y bellotas; B, variaci ja;
C, pelo fasciculado; D, bellota; E de la cúpul C, pelo fasciculado; D, bellota; E, escama de la cúpulA, vista frontal y
lateral.
Fic. 19. Q icana. A, hojas y bellotas; B, variación en Fic. 20. Quercus mohriana. A, rama con hojas y bellota; B, variación en
hoja; Gp lo fasciculad di dados; D, bellota; E, escama de hoja; C, pelo fasciculado; D, bellota; E, del íl la, vista lateral
la cúpula. y frontal.
1262 J | of the Botanical R h Institute of Texas 2(2)
101°23'W, 1,400 m, 2 nov 1988, J.A. Villarreal y M.A. Carranza 4784 (ANSM). Mpio. Saltillo: si pali $ ón ]
a El Cuatro, 25%21'N y 100°56'W, 2,100 m, 21 ago 2004, J.A. Encina et al. 1350 (ANSM).
Quercus mexicana Humb. et Bonpl., Pl. Aequinoct. 2. 35, pl. 82. 1809. (Fig. 19). Tro: MÉXICO. GUANAJUATO:
Santa Rosa, A. Bonpland 4218 (HoLoTIpO: B).
Quercus rugulosa M. M leotti, Bull. Acad, Brux. 10:209. 1843.
Quercus pen C.H. Mull. pa Midl. Naturalist 27:477. 1942.
Árbol de 3-8(-20) m de alto; ramillas café oscuro a grisáceo glabrescentes; yemas ovoides, de 2-3 mm de
largo, café brillantes; estípulas caedizas, lineares, liguladas, de 4—5 mm de largo, densamente pubescentes.
Hojas deciduas, algo rígidas; pecíolos de 3-6(—10) mm de largo, pubescentes; limbos ovados, elípticos u
oblongos, de 3-6(-8) cm de largo y 1-2.5 cm de ancho, ápice agudo u obtuso, raras veces redondeado,
terminando en una arista corta y quebradiza, base cordada a redondeada, margen entero, cartilaginoso,
revoluto y algunas veces crispado; haz algo brillante y glabro, con indumento de pelos fasciculados a lo
largo del nervio medio; envés con Bueno denso de pelos fasciculados, con radios contortos, enredados,
aislados, formando punt bre la epidermis; nervaduras 7-10(-15) en cada lado, ramificadas
y anastomosadas, prominentes en el envés, no evidentes en el haz. Inflorescencias masculinas de 3-5 cm de
largo, raquis lanoso. E femeninas de 3-6 mm de largo, con 1-3 flores, pedúnculos de hasta 5
mm de largo, indum . Fruto bianual, solitario o en grupos de 2-3 sobre un pedúnculo de 1-5 mm
largo; cúpula EN A de 10-15 mm de ancho y 6—9 mm de alto, escamas delgadas, apresadas, glabras
o puberulentas, ápice obtuso o redondeado, base un poco engrosada; nuez ovoide, de 12-15 mm de largo y
10 mm de diámetro, café claro, puberulentas, incluida de un tercio a un medio en la cüpula.
Distribución.—Se distribuye en el centro y sureste de México en los estados de San Luis Potosí, Queré-
taro, Hidalgo, Puebla, México y Chiapas, hacia el noreste a través de la Sierra Madre Oriental en Tamaulipas
y Nuevo León. En Coahuila se encuentra solo en la sierra de Zapalinamé al sureste del estado, en bosques
de encino y pino-encino. Se asocia con componentes de Quercus saltillensis, Q. rugosa, Q. greggii, Fraxinus
cuspidata, Cercis canadensis, Salix lasiolepis, Juniperus flaccida, Cupressus arizonica, Arbutus xalapensis, Prunus
mexicana, Pinus cembroides y Bumelia lanuginosa. Crece entre los 1900—2750 m de altitud. Abundante en
el bosque de encino, en laderas bajas donde llega a formar un bosque denso. Común en el cañón de San
Lorenzo, en la sierra de Zapalinamé. La especie es considerada como rara en Coahuila.
Notas.—Se puede reconocer por las hojas elípticas a oblongas de hasta 8 cm de largo con ápice agudo a
obtuso, cotamente aristado, el margen entero, glabras. Es caracteristico el enves con venación prominente,
con mechones de pelos en las axilas de la nervadura principal con las secundarias, con pelos fasciculados
con radios contortos y algo enredados. Bellota casi esferica con escamas de la cúpula con base ligeramente
engrosada.
Material examinado: Mpio. Arteaga: sierra La Atravesada, 25?14'N y 100°54'W, 2,480 m, 2 nov 2003, E. Padilla 1167 (ANSM). Mpio.
Saltillo: sierra de Zapalinamé, cañón de Santa Rosa, rumbo a los Aguajes, 25?19'N y 100°57'W, 2,427 m, 11 feb 2002, J.A. Encina et al.
695, 721 (ANSM); sierra de Zapalinamé, cañón de San Lorenzo, 25?23'N y 101°00'W, 2,060 m, 3 ago 1995, M.A. Carranza y J.A. Encina
162 (ANSM), J.A. Encina y R. Cardenas 462 (ANSM).
Quercus mohriana e - N.Y. Bot. Gard. 2:219. 1901. (Fig. 20). Tro: E.U.A. Texas. Val Verde Co.: S.B. Buckley
s.n. (Lecroripo: TEX; ISOTIP
Arbusto rizomat árbol de 0.5-5 m de alto; ramillas de p mm de grosor, con indumento gri ; yemas
1-2 mm de Ens ovoides o redondeadas, café rojizo oscur ; estípul d , subuladas
a filiformes, de 3 mm de largo, pubescentes. Hojas dines o deciduas, coriáceas; pecíolos de 2-5 mm
de largo, pubescentes; limbos oblongos a elípticos, de (1.5—)3—5(-8) cm de largo y (1.0-)2-3(-3.5) cm de
ancho, ápice iis u ed veces agudo, base redondeada a cuneada, margen poco revoluto, crispado o
plano, entero u ond tado o denticulado, haz verde oscuro, lustroso, con pelos multirradiados
diminutos, esparcidos, envés con mime denso, grisáceo; nervaduras 8—9 en cada lado, ramificadas y
notoriamente anastomosadas, algo prominentes en el envés. Inflorescencias masculinas de 2-3 cm de largo,
con indumento esparcido. Inflorescencias femeninas de 2-8 mm de largo, 13 flores. Fruto anual, solitario o
Villarreal et al., Q (F ) de Coahuil 1263
en pares, subsésil o con pedúnculos de hasta 1 cm de largo, cúpula de 8-18 mm ancho y 5-12 mm de alto,
escamas de la cúpula triangular-ovadas hasta oblongas, base engrosada con indumento canescente, ápice
elongado y estrecho, rojizo, glabro y adpreso; nuez elipsoide u ovoide, de 8-15 mm de largo y 5-12 mm de
ancho, con indumento esparcido, café claro, incluida cerca de la mitad en la cúpula.
Distribución.—Se distribuye en Texas y en las sierras del norte de Coahuila como serranías del Burro,
sierra del Carmen, La Encantada, hasta el centro en las sierras de la Madera, San Marcos y La Paila. Forma
parte del matorral submontano, en la ecotonía con el matorral rosetófilo, en bosque de encino y bosque de
pino-encino. Generalmente se asocia con Quercus sinuata var. breviloba, Q. gravesii, Q. grisea, Q. intricata,
O. pungens var. vaseyana, Q. laceyi, Bumelia lanuginosa, Ptelea trifoliata, Rhus virens, R. aromatica, Vauquelinia
corymbosa, Pinus remota, Arbutus xalapensis, Ungnadia speciosa, Cercocarpus i , Leucaena retusa, Cercis
canadensis y Fraxinus greggii. También se le puede encontrar en el matorral rosetófilo, donde ocurre en forma
esparcida con elementos de Nolina cespitifera, Dasylirion cedrosanum, Yucca thompsoniana, Leucaena retusa,
Calia secundiflora y Acacia berlandieri. Sus rangos e van desde los 750-2200 m en el matorral
submontano del noroeste del estado. Se le encuentra en lizas y laderas bajas, en exposiciones norte,
noreste u oeste, en los cañones llega a ser un arbolito de 4—6 m. Frecuente en las sierra del Carmen y ser-
ranías del Burro, escaso en la sierra La Paila. Especie comün en las áreas donde habita. Se reportan algunas
introgresiones con Q. grisea para Texas (Muller, en prensa).
Notas.—Se reconoce por las hojas coriáceas, oblongas a elípticas, usualmente de menos de 8 cm de largo,
pm BO UR entero, ondulado o con pocos dientes, haz Jen oscuro, e y envés opaco, con
1
indumen lti li Escamas de la cüpula i base engrodasa.
[e]
Se puede contando con Quercus grisea o Q. sinuata.
Material examinado: Mpio. Acuña: sierra del Carmen, cañón El Diablo, mina El Popo, 29?12'N y 102°45'W, 1,600 m, 29 jul 1973,
M.C. Johnston et al. 11929 (ANSM); serranias a Burro, inp El Bonito, 29°01'N y 102°07'W, 1,625 m, 19 s 1977, D. une y C.H.
e (ANSM. TEX). Mpio. Cuat haca, 30 ago 1980, L. Rodríguez y R. Vasquez 39 (ANSM). Mpio.
Santa Rosa (La Babia), rancho Le a Babia, cañón Rincón de María, 28°28'N y 102°04'W, 1,450 m, 27 abr 1975, T. Wendt
y D. Riskind 943 (ANSM, TEX); sierra La Encantada, mina Las Cabras, 16 ago 1976, R. Vasquez s.n. (ANSM). Mpio. Zaragoza: serranias
del Burro, rancho Los Potros, 28°55'N y 101°45'W, 1,100 m, 14 jun 1987, D. Castillo 507 (ANSM).
Quercus muehlenbergii Engelm., Trans. Acad. Sci. St. Louis. 3:391. 1877. (Fig. 21).
Quercus prinus L. var. acuminata Michx., Hist. Chenes Amer. Sep.. 5:3. 1801.
Quercus acuminata (Michx.) Sarg., Card. & Forest i 8:55. 1895.
Quercus brayi Small, Bull. Torrey Bot. Club 28:3
Quercus sentenelensis C.H. Mull., Amer. Midl. CURE 18:849. 1937.
Árboles de hasta 30 m de alto; corteza café grisácea, delgada, escamas alargadas, angostas y frágiles; ra-
millas de 1.5—3(-4) mm de grosor, con indumento fino, esparcido o glabras, café; yemas de 3-3.5 mm de
largo,esféricas a ovoides, estípulas subuladas de 3-4 mm largo, con indumento E alan o pa Hojas
deciduas, delgadas y algo coriáceas; pecíolos de 1-3 cm de largo, glabros o con ind ; limbos
lanceolados, oblanceolados a obovados, de (5—)8-15(-21) cm de largo y (2.5—)4—8(-10) cm de ando ápice
agudo a acuminado, base cuneada o Funda margen repando, dentado o con lóbulos poco profundos,
redondeados o acuminados y terminand o en un mucrón, haz glabro y lustroso de color verde oscuro, envés
glauco, con pelos simples y fasciculados con pocos radios, pequeños, esparcidos; nervaduras 8-12 por cada
lado, con algunas intermedias, terminando en el margen, ramificadas, un poco elevadas en el haz y promi-
nentes en el envés. Inflorescencias masculinas en amentos de 4-6 cm de largo. Inflorescencias femeninas
de 13 de 0.5-2 cm de largo. Fruto anual, solitario o en pares, subsésil o con un pedúnculo corto, cúpula de
18-22 mm de ancho y 10-12 mm de alto, escamas triangulares a rombicas, poco engrosadas, con indumento
gris; nuez oblonga a ovoide, de 15-20 mm de largo y 10-13 mm de ancho, glabra excepto cerca del ápice,
café claro, incluida cerca de un cuarto a un tercio en la cúpula.
Distribución.—Se distribuye por todo el centro-este de los Estados Unidos desde Nuevo México, Texas,
Oklahoma, Arkansas, Louisiana, Wisconsin hasta Maine y Ontario en Canadá. En México se distribuye por
la Sierra Madre Oriental en los estados de Coahuila y Nuevo León, en Coahuila se encuentra en la sierra
1264 Journai of the Botanical R h Institute of Texas 2(2)
del Carmen, del Burro, Santa Rosa, La Madera, La Gloria y La Concordia, en bosques de encino, de pino
y de pino-encino. Se asocia con especies de Quercus gravesii, Q. laceyi, Q. hypoleucoides, Q. mohriana, Acer
grandidentatum, Pinus arizonica var. stormiae, Cupressus arizonica, Prunus serotina, Garrya ovata, Tilia mexicana,
Ulmus multinervosa, Ungnadia speciosa y Juniperus deppeana. En el sur del estado se asocia además con Quercus
laeta, Q. saltillensis, Pinus cembroides, Juniperus flaccida y Arbutus xalapensis. Su rango altitudinal oscila entre
1300—2800 m, especialmente en lugares protegidos, o en cafiones con abundante humedad, sobre laderas
altas con exposición noroeste, o bien confinado a los márgenes de arroyos en montafias boscosas, como árbol
de 8-15 m. Común en las sierras del Carmen y La Madera, raro en la sierra La Concordia. Especie común
en Coahuila, sin llegar a ser abundante.
otas.—Se reconoce fácilmente por las hojas delgadas, oblanceoladas y grandes, de hasta 21 cm de largo,
con ápice agudo, 8-12 pares de nervaduras laterales prominentes, margen lobulado a dentado, pecíolos de
1-3 cm de largo. Escamas de la cúpula triangulares a rombicos, con engrosamiento central.
Material iaa Mpio: ae sierra del Carmen, El Cinco, 28°97'N y 102°57'W, 2,286 m, 3 sep 1997, G. Harper, J. Medel y D.
Doan-Cider s.n. (ANSA i Burro, rancho El Bonito, cañón El Bonito, 29°01'N y 102°07'W, 1,650 m, 17 sep 1977, D. Rishind y
C.H. Muller 2098 (ANSM, TEX). Mpio. G La Concordia, 6 km al NE de La Victoria, 25?15'N y 101°20'W, 2,770 m,
29 oct 1987, J.A. Villarreal et us 4093 gi Mpio. pears sierra Santa Rosa, rancho El olen, 27°55'N y 101°50'W, 1,300 m
may 2004, S. Lara y R. Pineda 102 (ANSA Rosa (La Babia), rancho La Babia, cañón Rincón de María, 28°27'N y 102°04'W,
1,450 m, 17 sep 1999, LA. Villarreal et al. 8772 (ANSM).
Quercus oblongifolia Torr., Sitgreaves, Rep. Exped. Zuni & Colo. R. 173, pl. 19. 1853. (Fig. 22). Tiro: E.U.A.
ARIZONA: Woodhouse s.n. (HoLoTIPO: NY; rsoriPo: MO).
Árbol de 6-10-16) m de alto, troncos de hasta 30 cm de diámetro, corteza surcada gris o blanquecina;
ramillas de 1-1.5 mm de grosor, con pelos fasciculados, esparcidos, glabrescentes, café rojizo, con lenticelas;
yemas subesféricas a ovoides, de 1-3 mm de largo, café rojizas, glabras; estípulas de 3-4 mm de largo,
subuladas, persistentes. Hojas deciduas, subcoriáceas, subpersistentes; pecíolos de 22)3-4(-8) mm de
largo, puberulentos a glabrescentes, rojizos; limbos oblongos a elípticos, en ocasiones lanceolados u ovados,
de (2-23—6(-8) cm de largo y (0.5—)1-2(-2.5) cm de ancho, ápice redondeado a obtuso, base redondeada,
cuneada o cordada, margen engrosado, ligeramente revoluto, entero, ondulado o irregularmente dentado
hacia el ápice; haz verde grisáceo a ligeramente azulado, opaco y glabrescente, envés glauco, glabro, o con
diminutos pelos fasciculados esparcidos, usualmente restringido a la base del nervio medio; nervaduras
6—8(-10) en cada lado, prominentes en el envés, un poco salientes en el haz, en un retículo blanco y escle-
rosado. Inflorescencias masculinas de 3-4 cm de largo, raquis puberulento. Inflorescencias femeninas de
2-15 mm de largo con 1-5 flores, pedunculadas. Fruto anual solitario o en pares, subsésil o sobre pedúnculos
cortos; cúpula de 10-13 mm de ancho y 6-8(-13) mm de alto, escamas ovado-oblongas, engrosadas, con
indumento gris, ápice rojizo; nuez ovoide u oblonga de 12-17(-19) mm de largo y (72)10—12 mm de ancho,
café claro, glabra o con puberulencia cerca del ápice, incluida cerca de un tercio en la cúpula.
Distribución.—Se distribuye desde Arizona, oeste de Texas, Sonora, Baja California, Chihuahua y Du-
rango, donde es abundante. En Coahuila se encuentra en las sierras del centro y noroeste que comprende
las sierras de Las Cruces, La Madera, San Antonio y Hechiceros. Crece en comunidades del matorral sub-
montano, bosque de encino y de pino-encino. Se asocia con Quercus intricata, Q. pringlei, Q. grisea, Lindleya
los pungens, Pinus arizonica
mespiloides, Rhus virens, R. aromatica, Agave falcata, Arbutus xalapensis,
var. stormiae, P. remota, Garrya ovata, Ceanotl ii, Dasylirion cedrosanum y Nolina cespitifera. Con rangos
altitudinales de 1550-1900 m, se encuentra en tek fondo de cafiones con bosque de encino, mientras que
en el matorral submontano crece sobre laderas bajas de exposición noroeste y oeste, con suelos derivados
de rocas ígneas. Se reporta una población en sierra de Hechiceros con hojas con indumento denso, lo cual
parece ser una introgresion con Q. invaginata (Muller, en prensa). La especie es rara y poco abundante en
Coahuila.
Notas.—Se reconoce por las hojas subcoriáceas, verde grisáceas, oblongas a elípticas de hasta 8 cm de
largo, ápice redondeado a obtuso, mergen entero a ondulado, glabras o con pubescencia esparcida y diminuta
Villarreal et a! ,Q
(F ) la faahiil 1265
0.25 r
2 mm
2
ul
J
Fic. 21. Q hlenbergii. À hojas y bellotas; E: dein Fic. 22. Q blonaifol H bell B, variación en
en hojas; C, pelos fasciculado y simple; D, bellota; E d pula, hoja; C, pelo fasciculado; D, bellota; E, le la cúpul
vista mad y lateral.
TR. d
desi
DM
A
-— t
DE
Wie
Mz
C Y 1010mm
Fic. 23. Q l ly í ha. À, rama con hojas, tos y bellotas; B, Fic 24. Quercus potosina. A ias v bell B, variación en hoja:
iación en hoja; C, pelo fasciculado; D, bellota; E de la cúpul C pelo fasciculado; D, bellota: E del
1266 Journal of the Botanical R h Institute of Texas 2(2)
de pelos fasciculados, venación esclerosada. Bellotas con cápula corta y escamas con engrosamiento central.
Algunas muestras pueden presentar algunas semejanzas que lleven a la confusión con Q. laceyi.
Material examinado: Mpio. Cuatro Cienegas: cañón La Barrica, picacho El Pajarito, 27%01'N y 102°23'W, 2,260 m, 28 ago 1975, T.
Wendt y E. Lott 1344 (ANSM, TEX).
Quercus polymorpha Schltdl. et Cham., Linnaea 5:78. 1830. (Fig. 23). Tiro: MEXICO. Veracruz: Jalapa, hacienda de
Schiede 501B : HAL; isoripPo: GH)
Arbol de 10—15(-20) m de alto, con corteza gris, escamosa o surcada; ramillas de 2-3 mm de grosor, de
color café rojizas, con indumento amarillento, glabrescentes, con numerosas lenticelas; yemas ovoides, de
3-5(-10) mm de largo, de color café rojizo, con pelos cortos apresados a glabrescentes; estipulas subuladas
y pubescentes de 10-13 mm de largo, persisten cerca de las yemas terminales. Hojas deciduas, coriáceas;
pecíolos de (6—)15—25(-35) mm de largo, casi glabros, amarillentos, delgados y eng ] la base; limbos
oblongo-elípticos u ovados al lados u obovados de 5-10(-15) cm de largo y 3-6(-8) cm de ancho, ápice
angostamente obtuso, rara vez acuminado, base generalmente redondeada a cordada, margen ligeramente
cartilaginoso y revoluto, entero o p li mucronados, localizados en la mitad superior de la hoja; haz
verde oscuro y glabro, envés glauco ceroso, opaco, o con indumento gris amarillento de pelos fasciculados;
nervaduras 10-12(-14) a cada lado, muy ramificadas y anastomosadas cerca del borde, hundidas en el haz
y muy prominentes en el envés, formando un retículo. Inflorescencias masculinas de 6—8 cm de largo, vel-
losas. Inflorescencias femeninas de 5-10 mm de largo, con una o dos flores. Fruto anual solitario o por pares,
sobre pedúnculos de 5-30 mm de largo; cúpula hemisférica de 12-15(-20) mm de diámetro, con escamas
engrosadas en la base, agudas y apresadas, blanquecinas; nuez elipsoide de 14—20(-25) mm largo y 8-13
mm de ancho, café claro, incluida aproximadamente la mitad en la cúpula.
Distribución.—Se distribuye desde el suroeste de Texas, Nuevo León, Tamaulipas, Veracruz, San Luis
Potosí, Jalisco, Hidalgo, Oaxaca y Chiapas hasta Guatemala. En Coahuila se presenta en la sierra de Pájaros
Azules, en el centro del estado, en los límites con el estado de Nuevo León. Crece en bosque de encino
asociada con Q. invaginata, Q. laceyi, Q. gravesii, Q. canbyi, Juglans microcarpa, Fraxinus greggii, Chiococca
alba, Colubrina greggii, Rhus virens, Pistacia texana y Ungnadia speciosa, en altitudes entre 700—850 m, sobre
laderas bajas y en fondo de cañones húmedos, con exposición norte o noroeste. Especie rara para el área del
estado.
Notas.—Se puede reconocer por las hojas coriáceas, verde oscuro, cerosas, oblongo a ovadas, de 5-15
cm de largo, base redondeada a cordada, margen entero o con pocos dientes cortos, envés con nerviacion
reticulada resaltada. Bellotas de 15-30 mm largo, escamas de la cúpula triangulares, con tuberculo basal.
Material examinado: Mpio. Candela: sierra de Pajaros Azules, rancho Santa María, en los limites con Lampazos N. L. 875 m, 23 nov
2002, J.J. Medellin V. 433, 445 (CFNL).
NE potosina Trel., Mem. Natl. Acad. Sci. S 84, pl. 123. 1924. (Fig. 24). Tro: MÉXICO. San Luis P re
arry y E. Palmer 838 PM MO; isoripos: GH, M
ign potosina Trel. f. exilis Trel. Mem. Natl. Acad. Sci. 20:84, pl. 124. 1924.
Quercus potosina Trel. £. aperta Trel., Mem. Natl. Acad. Sci. 20:84, E 125. 1924.
is cordifolia Trel., Mem. Natl. Acad. Sci. 20:84, pl. 1
Quercus jaralensis Trel., Mem. Natl. Acad. Sci. 20:86, m 133. 1924.
Arbusto de 1-3 m o raras veces árbol de hasta 7 m de alto; corteza gris, escamosa; ramillas de 1-3 mm de
grosor, rojizas, con pelos fasciculados y pelos glandulares cortos; yemas ovoides, de 2-4 mm de largo, gri-
sáceas, pubescentes o glabrescentes; estípulas filiformes a subuladas, de 6—10 mm de largo, pilosas. Hojas
persistentes, gruesas y coriáceas; pecíolos de 2-3(-6) mm de largo, con indumento gris persistente, rojizos;
limbos obovados, elípticos, hasta suborbiculares, de (2-J3.5-5(-9) cm de largo y (1-)2-3.5(-6) cm de ancho,
ápice redondeado hasta agudo, base cordada a redondeada, margen algo revoluto con dientes toscos termi-
nando en mucrones, haz algo lustroso, con pelos fasciculados, envés glauco-ceroso, ad con O
1,41 1; 1
formado de pelos f estipitados en la base de la nervadura principal y
Villarreal et al., Q (F ) de Coahuil 1267
limbo, frecuentemente con puntos resi color ambar; nervaduras (6—)8(-10) en cada lado, prominentes
en el envés, la nervadura media y principales rojizas. Inflorescencias masculinas de 2-2.5 cm de largo, con
flores densas, raquis velloso. Inflorescencias femeninas de 2-12 mm de largo, 1-3(-4) flores. Fruto anual,
solitario o en grupos de 2-4, subsésil o sobre pedúnculos de hasta 8 mm de largo velloso; cúpula hemisférica
de 12-15 mm de ancho, con el margen involuto, las escamas poco engrosadas en la base, con el ápice agudo,
un poco apresadas, indumento de pelos cortos y apresados; nuez ovoide, de 12-15 mm de largo y 8-12 mm
de ancho, puberulenta o glabrescente, incluida cerca de la mitad en la cúpula.
Distribución.—Se distribuye desde Chihuahua, Durango, Zacatecas, San Luis Potosí, Aguascalientes,
Jalisco, Guanajuato, Querétaro e Hidalgo. En Coahuila se encuentra solamente en el extremo sureste del
estado en la sierra de Arteaga, tae en i cañón ee duae en is de pino-encino, asociándose con
Pinus cembroides, P. teocote, P. pseud Arbut I , Cercocarpus montanus,
Quercus saltillensis, Q. ruo y Q. laeta. Crece enun ang alitudisat de 2300-2650 m sobre laderas bajas
con exposición norte y sureste, en bosques abiertos de pino-encino, como arbusto o arbolito de hasta 5 m
de alto. Especie con categoria de rara en Coahuila.
Notas.—Se reconoce por las hojas gruesas obovadas a suborbiculares, usualmente menos de 6 cm de
largo, con ápice redondeado, margen con dientes mucronados, envés con pelos fasciculados, multirradiados
y puntos amarillos. Cúpulas con con margen involuto y escamas ligeramente engrosadas en la base. Se le
= confundir con Quercus fulva por la forma de las hojas.
Material inado: Mpio. Art i de Arteaga, rancho El Tirol, cañón de Jamé, 2,450 m, 24 jun 1994, J.A. Villarreal et al. 7719
(ANSM), sierra Las Vigas, cafion de Jamé, 25%26'N y 100°20'W, 2,500 m, 8 may 1995, M.A. Carranza y J.A. Encina 2162 (ANSM).
Quercus pringlei Seemen, Bot. s Syst. 29:96. 1900. (Fig. 25). Tipo: MÉXICO. Coanuma: Carneros Pass, C.G. Pringle
2382 (1ecrorIPO: B; isoripos: GH, T
Quercus filiformis C.H. Mull., Amer. Midl. Naturalist 27:473. 1942.
Arbusto o árbol pequeño de (0.50—1—4(-10) m de alto; corteza delgada, fisurada y gris; ramillas de 1-2 mm
de grosor, café oscuro a grisáceas, glabrescentes; yemas ovoides de 1-1.5 mm de largo, café oscuro, glabras;
estípulas filiformes, de 3—5 mm de largo, glabras. Hojas persi y coriáceas; pecíolos de 2-5 mm de largo,
indumento esparcido, rojizos; limbos lanceolados hasta ] lípticos u oblongos, de (122—3(-4.5)
r
cm de largo y (0.5-)1-2(3) cm de ancho, ápice agudo, obtuso o redondeado, mucronado, base redondeada,
cuneada o cordada, margen algo engrosado, revoluto o plano, entero, denticulado, a toscamente dentado,
dientes los; haz glabro, raramente con indumento diminuto fasciculado, esparcido o restringido al
nervio medio, envés ee azuloso, con indumento fasciculado esparcido, glabrescente; nervaduras 7-8(-12)
en cada lado, irregulares, ramificadas y anastomosadas cerca del margen, poco salientes. Inflorescencias
masculinas 1-2 cm largo, con indumento esparcido. Inflorescencias femeninas 1-2 flores, subsesiles. Fruto
anual solitario o en grupos de 2-4, subsésil o en pedúnculos de 1-10 mm de largo; cúpula hemisférica, de
8-10 mm de ancho, escamas con la base engrosada e indumento canescente, ápice café oscuro, delgado,
apresado; nuez elíptica de 8-10 mm de largo, glabra, café claro, levemente estriada, incluida de un tercio a
un medio en la cúpula.
Distribución. —Se distribuye en Zacatecas, San Luis Potosí, Nuevo León, Coahuila y Durango. En Coa-
huila se encuentra en el noroeste, centro-oeste del estado en las sierras del Pino, La Madera, hasta el sur y
sureste en las sierras de Parras y de Carneros en comunidades de matorral submontano, en bosque de pino
piñonero y bosques de encino, asociándose con Pinus cembroides, P. remota, Juniperus flaccida, J. erythro-
carpa, Calia secundiflora, Lindl iloides, Rhus virens, Mimosa biuncifera, Quercus striatula, Q. coahuilensis,
Agave falcata, Cercocarpus fothergilloides, Dasylirion cedrosanum y Eysenhardtia texana. Crece en altitudes de
1800-2800 m, donde forma matorrales en sitios con exposiciones norte, noreste y sur, sobre laderas bajas
y cimas con poca humedad, en suelos someros y pedregosos, prefiere crecer en lugares abiertos y soleados.
Abunda en las sierras del sur del estado y es común en las sierra La Madera y del Pino. Una población de
plantas de la sierra La Madera, con tallos postrados, alargados y hojas delgadas, fue descrito como Quercus
1268 J | of the Botanical R h Institute of Texas 2(2)
filiformis (Johnston 1944). Especie comün en las áreas donde habita. Es conocido localmente como: encino
chaparro o charrasquillo.
Notas.—Se reconoce por ser generalmente un arbusto bajo, de hojas coriáceas, elípticas a oblongas,
usualmente de menos de 3 cm de largo, glabras o casi glabras y venación inconspicua. Bellotas de menos de
1 cm de largo y escamas de la cúpula con base engrosada.
Material inado: M i La Marta, 25°12'N y 100°21'W, 2,920 m, 14 jul 1997, A. A. Moreno 022 (ANSM). Mpio. Cua-
REN cañón de la Barrica, 27°O1'N y 102°23'W, 2,300 m, 20 ago 1975, T. Wendt y E. Lott 1216 (ANSM, TEX).
Mpio. General Pid 30 km al W de dn dE 25°19'N y 101°44'W, 1,940 m, 4 sep 1992, J.A. Villarreal et al. 6537 (ANSM).
Mpio. Parras: sierra Playa Madero, cañón Pinalosa, ejido Santa Victoria, 25°09'N y 101°34'W, 2,400 m, 6 jun 1995, M.A. Carranza y
J.A. Encina 2203 (ANSM). Mpio. Saltillo: sierra Catana 25%15'N y 101°12'W, 2,800 m, 29 jul 1992, J.A. inet et al. 6471 (ANSM),
sierra Zapalinamé, cañón de San Lorenzo, 25°23'N y 101°00'W, 1990 m, M.A. Carranza y J.A. E 2223 (ANSM), sierra de Carneros,
25°07'N y 101°07'W, 2,100 m, 15 jun 1995, J.A. Villarreal y M.A. mana 8184 (ANSM).
Quercus depo Liebm. var. pungens, Overs. Kongel. Danske Vidensk. Selsk. Forh. Medlemmers Arbeider.
171. 1854 (Fig. 26). Quercus undulata Torr. var. pungens (Liebm.) Engelm., Trans. Acad.Sci. St. Louis 3:392. 1877. Tiro: E.U.A.
Texas: C. Wright 664 Gsotipo: MO; sintiro: NY).
Arbusto o árbol de 4—6 m de alto; corteza delgada, gris; ramillas de 1-2 mm de grosor, con indumento
amarillento; yemas esféricas u ovoides cerca de 2 mm de largo, café rojizo oscuro con indumento esparci-
do; estípulas subuladas, de 3-4 mm de largo, pubescentes. Hojas persistentes duras y tiesas; pecíolos de
2-3C10) mm de largo, con indumento amarillento denso; limbos elípticos a oblongos, de 2-4(-9) cm de
largo y 1.5—2.5(-4) cm de ancho, ápice agudo u obtuso, terminando en un mucrón, base redondeada o
raras veces cuneada, margen crispado, dentado o con lóbulos agudos, terminados en mucrón, ligeramente
revoluto, haz opaco o poco lustroso, con pelos estrellados, diminutos, esparcidos, envés canescente, de pelos
seríceos simples y pelos estrellados; nervaduras 5-8(-14) en cada lado, muy ramificadas, más evidentes en
el envés. Inflorescencias masculinas de 1-2 cm de largo. Inflorescencias femeninas con 13 flores. Fruto
anual, solitario o en pares, subsésil o sobre pedúnculos de 2-3 mm de largo; cúpula hemisférica, de 10-13
mm de ancho y 6-8 mm de alto; escamas de la cúpula ovadas o con frecuencia estrechas y oblongas, base
engrosada, con indumento grisáceo, ápice delgado y apresado, glabrescente o puberulento, café rojizo; nuez
ovoide a subcilíndrica, de 8-12 mm de largo y 6-8 mm de ancho, café claro, glabra, incluida una cuarta
parte a la mitad en la cúpula.
Distribución. —Se distribuye en Texas, Nuevo México, Arizona y Chihuahua. En Coahuila se encuentra
en las sierras del centro-oeste, suroeste y noroeste, como sierra Mojada, Las Cruces, El Pino, La Madera y
Jimulco, en el matorral submontano, en la ecotonía con matorral rosetófilo bosque encino y de pino-encino.
Se asocia generalmente con Pinus remota, P. johannis, Juniperus flaccida, Quercus intricata, Q. pringlei, Q. gravesii,
Ceanothus greggi, Lindleya mespiloides, Cercocarpus montanus, Garrya ovata, Rhus virens, Arctostaphylos pungens,
Acacia greggii, Dasylirion cedrosanum, Nolina cespitifera y Flourensia retinophylla. Se encuentra en altitudes de
1200-2400 m, sobre laderas bajas, con exposiciones norte, sur y sureste, con suelos superficiales, derivados
de rocas calizas o ígneas. En algunas comunidades puede formar un matorral denso de 1-2 m de alto. En
arroyos intermitentes presenta un crecimiento de arbolito o árbol de 4—6 m, formando colonias esparcidas.
Es menenie en la sierras n al oeste Se estado.
, duras, de cl rillentas, lustrosas, ásperas altacto,
elípticas a s e de menos de 4 cm de largo, margen con lóbulos crispados y mucronados
(pungentes), indumento de pelos estrellados diminutos y apresados. Bellotas de cerca de 1 cm de largo,
wi con escamas de la cúpula casi circulares, engrosadas y con ápice delgado.
l do: Mpio. Acuña: si del Carmen, Styrax canyon, 29°11'N y 102°45'W, 1,750 m, 22 sep 1973, T. Wendt y E. Lott 75
(ANSM, TEX). Mpio. Cuatro Cienegas: sierra de San Marcos y Pinos, 2,159 m, 14 jul 1999, R. Marroquin s.n. (ANSM). Mpio. Ocampo:
sierra La Encantada, cuesta Malena, 28°35'N y 102°10'W, 21 ago 1994, M.A. Carranza et al. 114 (ANSM).
Quercus pungens Liebm. var. vaseyana (Buckley). C.H. Mull., Contr. Texas Res. Found. Bot. Stud. 1:70.
1951. Quercus vaseyana Buckley, Bull. Torrey Bot. Club 10:91. 1883. recs undulata Torr. var. vaseyana o Rydb., Bull.
N.Y. Bot. Gard. 2:218, pl. 30. 1901. Tiro: E.U.A. Texas: Pecos river, S.B. Buckley s.n. (Lecroripo: NY, ISOLECTOTIPO:
(F ) de Coahuil 1269
Villarreal et al., Q
Quercus sillae Trel. Mem. Natl. Acad. Sci. 20:102, pl. 171. 1924.
Arbusto o arbolito de 2-3 m de alto; corteza delgada, gris, fisurada; ramillas de 1-1.5 mm de grosor, café
rojizo, con pelos estrellados, glabrescentes; yemas ovoides de 1-1.5 mm de largo, café rojizo oscuro o gris;
estípulas caducas, subuladas, 3 mm largo, con indumento esparcido. Hojas deciduas; pecíolos de 2-5 mm
de largo, con indumento estrellado; limbos coriáceos, lanceolados a oblongos, de 2.5-4(-6) cm de largo y
0.6-2 cm de ancho; ápice agudo o raras veces obtuso, base cuneada a redondeada, margen cartilaginoso y
engrosado, casi plano con 3-5 dientes toscos en cada lado o con lóbulos poco profundos, agudos u obtusos
y terminando en un mucrón o enteros, haz verde oscuro, lustroso, glabrescente o con tomento estrellado
esparcido, envés seríceo, con pelos diminutos estrellados apresados y pelos simples; nervaduras 3-4(-6)
en cada lado, ligeramente salientes en el haz, prominentes en el envés, las principales glabras y conspicuas
en contraste con el indumento del envés. Inflorescencias masculinas de 2.5-3 cm de largo, con tomentulo
corto. Inflorescencias femeninas de 3-5 mm de largo, 1-2 flores. Fruto anual, solitario o en grupos de tres,
subsésil o sobre pedúnculos de de 2-3 mm de largo; cúpula hemisférica, de 7-10 mm de ancho y 3-4 mm
de alto; escamas de la cúpula ovadas, base engrosada, con pelos cortos, ápice delgado, apresado, rojo oscuro,
glabro; nuez subcilíndrica, de 10-12 mm de largo y 5-8 mm de ancho, café claro, glabra, incluida de un
tercio a un quinto en la cüpula.
Distribución. —Se distribuye en Texas, Chihuahua, Coahuila y Nuevo León. En Coahuila se le localiza
en las sierras del norte, centro y sur, como La Encantada, La Madera, La Gloria, de Parras, en matorral sub-
montano, en la transición hacia el matorral rosetófilo y bosque de pino-encino, asociándose con Quercus
intricata, Q. invaginata, Q. striatula, Q. pungens var. pungens, Q. mohriana, Pinus cembroides, P. remota, Goch-
nathia hypoleuca, Dasylirion cedrosanum, Juniperus flaccida, Fraxinus greggii, Rhus virens, Lindleya mespiloides,
Cercocarpus montanus, Calia secundiflora, Acacia berlandieri y Ptelea trifoliata. Esta presente en altitudes de
1050-2500 m, en arroyos intermitentes, cañones o laderas bajas rocosas con exposición norte, sur y oeste.
Es comün que forme matorrales densos, en arroyos se presenta como arbolito o árbol esparcido de 7-8 m
alto. Es frecuente en las sierras del centro-norte y escaso en las sierras al sur del estado.
Notas.—Se reconoce por las hojas coriáceas, duras, verde claro a amarillentas, muy lustrosas, ligeramente
ásperas al tacto, elípticas a oblongas, usualmente de menos de 4 cm de largo, limbos casi planos, indumento
de pelos estrellados diminutos, apresados y esparcidos. Bellotas de cerca de 1 cm de largo, subsesiles, con
escamas de la E casi circulares, engrosadas y con ápice delgado.
Material lo: M del Carmen, San Ysidro ranch, 1,803 m, G. Harper y D. Doan-Crider s.n. (ANSM). Mpio. Cua-
tro Cienegas: sierra de La Fragua, valle Grande 26%48'N y 102?33'W, 1,700 m, 29 may 1999, C. Zermeño s.n. (ANSM). Mpio Parras:
sierra de Parras, arroyo Seco, 2,500 m, 11 jun1981, A. Rodríguez y M.A. Carranza 778 (ANSM). Mpio. Ramos Arizpe: sierra San José de
los Nuncios, 1,600 m, J.A. Villarreal y M.A. Carranza 7053 (ANSM). Mpio. San Pedro: sierra Las Margaritas, cañón Blanco, 26°32'N y
102°53'W, 1,290 m, 21 oct 2006, M.A. Carranza et al. 4603 (ANSM). Mpio. Torreón: sierra de Jimulco, 25?10'N y 103°15'W, 2,000 m,
26 sep 1995, J.A. Villarreal y M.A. Carranza 8329 (ANSM).
Quercus rugosa Née, Anales Ci. Nat. 3:275. 1801. (Fig. 27). Tiro: MÉXICO. Epo. De Mexico: Née s.n. (HoLoripo: MA).
Quercus reticulata Humb. et Bonpl., Pl. Aequin. 2:(35)40, pl. 86. 1809.
Quercus veliifera Trel., Mem. Natl. Acad. Sci. 20:73, pl. 90. 1924.
Quercus durangensis Trel., Mem. Natl. Acad. Sci. 20:73, pl. 91. 1924.
Quercus diversicolor Trel., Mem. Natl. Acad. Sci. 20:73, pl. 92-94. 1924.
Quercus rhodophlebia Trel., Mem. Natl. Acad. Sci. 20:74, pl. 95. 1924.
Quercus ariifolia Trel., Mem. Natl. Acad. Sci. 20:74, pl. 97. 1924.
Arbol de (5210—30 m de alto; tronco de hasta 1 m diámetro; corteza escamosa café claro a ; ramillas de
(1-)2-3(-4) mm de grosor, pubescentes a glabrescentes, con lenticelas color café; yemas ovoides, de 2-5 mm
de largo, café; estípulas persistentes, lineares u oblanceoladas de 5-8(-12) mm de largo, pubescentes. Hojas
deciduas, muy rugosas; pecíolos de 5—7(-10) mm de mace pube cce. a glabros, rojizos; limbos gruesos,
rígidos y coriáceos, obovados, elíptico-obovad , de 5-10(-20) cm de largo y 3-7(-10)
cm de ancho, ápice redondeado u obtuso a veces agudo, base redondeada o cordada, margen usualmente
revoluto, engrosado, sinuado o con dientes hacia el ápice de la hoja, los dientes terminan en un mucrón de
1270 J lof the Botanical R h Institute of Texas 2(2)
Fig. 25. Quercus pringlei. A hojas y bellotas; B, variación en hojas; ^ Fis. 26. Quercus pungens. A, rama con hojas y bellotas; B, variación en
hoja; C, pelo estrellado; D, bellota; E, de | " la, vista frontal
C, bellota; D, escama de la cüpula. ja; G p
y lateral.
is. À, rama con hojas, y bellotas; B,
haia f facririlado v facciculad ; D, bellota;
Fic. 27. Quercus rugosa. A, rama con hojas, amentos y bellotas; B, variación ^ Fic. 28. Q
hoia: € fasciculad di dad j D, bellota; E escama iació
Jur EN J
Ana er |
de la cúpula. E, escama de la cúpula.
Villarreal et al., Q (F ) de Coahuil 1271
1—2 mm, envés cóncavo a plano, haz verde oscuro y algo lustroso, glabro o con pelos glandulares y pelos
fasciculados a lo largo del nervio medio, envés con indumento formado por pelos fasciculados con radios
algo enredados y pelos glandulares vermiformes, amarillo rojizos, epidermis glauco-cerosa, marcadamente
ampulosa y papilosa; nervaduras 8-10(-12) en cada lado, ramificadas, impresas en el haz, muy prominentes
en el envés, formando un retículo. Inflorescencias masculinas de 3—5 cm de largo, con tomento fasciculado.
Inflorescencias femeninas de 5-10 cm de largo, con pedúnculo delgado pubescente, 15—20 flores. Fruto
anual, solitario o en grupos de 2-58) sobre pedúnculos de (1.5-)6-10 cm de largo; cúpula hemisférica
de 8-15 mm de ancho y 6-10 mm de alto; escamas de la cúpula con el ápice delgado, café y glabras, bases
engrosadas con tomento canescente; nuez ovoide a elíptica de 9-28 mm largo y 7-14 mm de ancho, café
claro y glabra, incluida cerca de la mitad en la cúpula.
Distribución.—Se distribuye desde Arizona, Nuevo México y Texas, Baja California, Chihuahua, Dur-
ango, San Luis Potosí, Aguascalientes, Colima, Jalisco, Michoacán, Guanajuato, Querétaro, Estado de México,
Hidalgo, Tlaxcala, Puebla, Veracruz, Distrito Federal, Morelos llega hacia el sur hasta Oaxaca, Chiapas y
Guatemala. En Coahuila se encuentra al norte del estado en la sierra del Carmen y al sureste en la sierra de
Zapalinamé, en bosques de encino y pino-encino. Se asocia con especies de Quercus carmenensis, Q. grisea,
Q. gravesii, Q. muehlenbergii, Q. arizonica, Q. mexicana, Q. saltillensis, Q. sideroxyla, Arbutus xalapensis, Pinus
arizonica var. stormiae, Pseudotsuga menziesii, Arctostaphylos pungens, Cercis canadensis, Acer grandidentatum,
Prunus serotina, Fraxinus cuspidata, Garrya glaberrima, Cupressus arizonica y Salix lasiolepis. Se le encuentra
en sitios con elevaciones generalmente de 2000—2700 m, en laderas altas con exposición norte y noroeste,
así como en cañones húmedos y márgenes de arroyos, forma bosques densos de árboles de hasta 10 m de
alto. Es frecuente en algunas localidades de la sierra del Carmen y en el cafión de San Lorenzo en la sierra
de Zapalinamé. La especie se considera rara en el área del estado.
Notas.—Se reconoce fácilmente por sus hojas coriáceas, gruesas, rugosas, verde oscuro y usualmente
concavas, obovadas a suborbiculares, generalmente menos de 10 cm de largo, ápice redondeado a obtuso,
margen con dientes cortos mucronados, envés con pelos glandulares y pelos fasciculados con radios riza-
dos. Bellotas de 1-3 cm de largo, en racimos largos (hasta 10 cm de largo), escamas de la cúpula con base
tuberculada.
M O lel Carmen, 28°58'N y 102°33'W, 2,200 m, 16 ago 1975, M.F. Robert y J. Passini 6602 (ANSM),
sierra del Carmen, El rand: 2 200, 8 jun 1991, J.A. Pérez 1767 (ANSM). Mpio. Saltillo: cañón de San Lorenzo, 25°20'N y 100°57'W,
2,292 m, 20 jul 2000, J.A. Encina 168, 497 (ANSM).
Quercus saltillensis Trel., Mem. Natl. sae Sci. 20:183, pls. 368, 369. 1924. (Fig. 28). Tiro: MÉXICO. Coanuma:
Carneros Pass, E. Palmer 1277 (n
Quercus carnerosana Trel., Mem. Natl. Acad. Sci. 20:183, pl. 369. 1924. Tipo: MÉXICO. Coanuma: Carneros Pass, C.G. Pringle 2802
(HOLOTIPO: AA).
Arbusto o arbolito 1-5 m n Pid con corteza obscura fisurada; ramillas de (131.5-2.5 mm de diámetro, con
indumento blanqu in fé-rojizos; y ovoides, de 3 mm largo y 2 mm de diámetro,
escamas ados. café das estipulas caedizas. Hojas deciduas y delgadas; pecíolos de 3-9 mm de largo,
glabros; limbos lanceolados a oblongo-lanceolados, de 2—4.5(-8) cm de largo y 0.8-2(3.0) cm de ancho,
ápice agudo a atenuado, aristado, base redondeada o cordada, margen algo cartilaginoso, plano, entero o
con dientes poco a muy evidentes, aristados; haz verde oscuro, lustroso, glabrescente, envés con indumento
blanquecino, denso, e S con pd fasciculados RUE y pelos con 2 fasciculos, en las axilas del
nervio medio; nervaduras 6-10 en cada lado el envés. Inflorescencias masculinas
3—5 cm de largo, con indumento esparcido, lóbulos del T fimbriados. Inflorescencias femeninas de 5-15
mm de largo, con 1-2 flores sobre pedúnculos puberulentos o glabros. Fruto anual, solitario o en pares,
sobre un pedúnculo de 3-7 mm de largo; cúpula hemisférica o turbinada, de 5-10 mm de alto y 7-12 mm
de ancho; escamas casi planas, con indumento amarillo-rojizo, ápice agudo, café; nuez ovoide, de 8-15 mm
de largo y 6-9 mm de ancho, café claro, casi glabra, incluida cerca de un tercio a la mitad en la cúpula.
Distribución.—Se distribuye en el sur y sureste de Coahuila en las sierras de Arteaga, La Concordia,
1272 J lof the Botanical R h Institute of Texas 2(2)
Parras, Carneros, Zapalinamé, se extiende a la sierra La Paila y hasta la sierra de Astillero al norte de Za-
catecas, al cerro El Águila en San Luis Potosí y a través de la Sierra Madre Oriental al centro-suroeste de
Nuevo León. Crece en bosques de pino-encino, de encino, y en la transición entre el bosque de pino-encino y
oyamel, asociándose con Pinus cembroides, P. pseudostrobus, Pseudotsuga menziesii, Cupressus arizonica, Quercus
hintoniorum, Q. mexicana, Q. greggii, Q. laeta, Q. striatula, Juniperus flaccida, J. erythrocarpa, Prunus serotina,
Ceanothus buxifolius, Juglans microcarpa y Arctostaphylos pungens, en exposiciones sur y sureste se asocia con
algunas especies de la familia Agavaceae. Se presenta entre 1900-2800 m, en laderas medias y bajas, con
exposiciones norte y noroeste. En lugares húmedos es un arbolito, mientras en los límites del bosque de
pino-encino y de oyamel se presenta como arbusto. Es abundante en las sierras de Arteaga, Zapalinamé y
La Concordia, mientras que en la sierras La Paila y de Parras, es escaso
jas delgadas, verde oscuro, lanceoladas a oblongas, usualmente de menos
de 5 cm de largo, apice generalmente agudo, margen entero entero o con dientes aristados, envés con pelos
fasciculados simples y pelos fasciculados dobles. Escamas de la cápula casi planas. Puede confundirse con
Quercus mexicana o Q. striatula por la forma de las hojas.
Material examinado: Mpio. Arteaga: Los Lirios-El Cercado, 2,410 m, 29 jul 1995, G.B. Hinton 25439 (ANSM), Las Vigas, cañón La Car-
bonera, 25?20'N y 100°39'W, 2,300 m, 5 jun 1987, J.A. Villarreal y M.A. Carranza 3742 ae sierra de Arteaga, cañón Jamé, 25°26'N
y 100°30'W, 2,600 m, 13 ago 1996, J.A. Villarreal y J. Zavala 8406 (ANSM). Mpio. G ] La Concordia, ejido La Casita,
25°14'N y 101°25'W, 2,200 m, 10 oct 1996, J.A. Encina y J. Zavala 178, 274 (ANSM). Mpio. Partie sierra de Parras, ejido El Capulín,
25°23'N y 102°13'W, 2,050 m, 6 may 1983, A. Rodríguez et al. 875 (ANSM). Mpio. Ramos Arizpe: sierra La Paila, La Sauceda, 14 nov
5, J. Marroquín 1063 (ANSM). Mpio. Saltillo: sierra de Zapalinamé, cañón Boca Negra, 25%21'N y 100°55'W, 2,270 m, J.A. Encina y
R. Cardenas 642 (ANSM), sierra Zapalinamé, lomas de Lourdes, 1,800 m, 19 may 1974, J. Marroquín 2759 (ANSM).
Notas.—Se reconoce por las]
Quercus sideroxyla Humb. et Bonpl., Pl. Aquinoct. 2:(34)39, pl. 85. 1809. (Fig. 29). Tiro: MÉXICO. Guanajuato:
Villalpando, A. Humboldt y A. Bonpland s.n. (HoLoTIpO: P).
uercus omissa A. DC., Prodr. (DC.) 16 (2.1):28. 1864.
Quercus epileuca Trel., Mem. Natl. Acad. Sci. 20:126, pl. 229. 1924.
Quercus incarnata Trel., Mem. Natl Acad. Sci. 20:127, pl. 231. 1924.
Árbol de 5—10(-20) m de alto, con troncos de 0.5-1.0 m de diámetro, corteza grisácea, gruesa y agrietada;
ramillas de 2-3 mm de grosor, gris oscuro, con puberulencia negra; yemas ovoides, de 2.5 mm de largo;
estípulas filiformes, caedizas. Hojas gruesas y rígidas; pecíolos de 3-6(-10) mm de largo, pubescentes,
con puberulencia negra; limbos oblanceolados a obovados o elíptico-oblongos de 2.5—4-6(-10) cm de
largo y (122-4(—5) cm de ancho, ápice agudo a obtuso, base redondeada o cordada, margen cartilaginoso,
engrosado y revoluto, denticulado o con dientes prominentes en la mitad distal, cortamente aristados, haz
opaco, verde amarillento, glabrescente, con pelos fasciculados, usualmente restringidos al nervio principal,
envés cubierto por indument o denso y persistente, blanco amarillento a café pálido, con pelos fasciculados,
con radios ondulados, epidermis con ampulas o papilas prominentes; nervaduras 5—7(—9) en cada lado,
ramificadas, alcanzando el diente o anastomosadas cerca del margen, algo impresas en el haz, prominentes
en el envés. Inflorescencias masculinas cerca de 4 cm de largo, con tomento fasciculado, con muchas flores.
Inflorescencias femeninas de 5-12 mm de largo, 1-2 flores. Fruto anual o bianual (flores y frutos contempo-
ráneos), solitario o en pares, sésiles o con pedúnculos de 1-10 mm de largo, cúpula hemisférica, de 7-13 mm
ancho y 4-7 mm de alto, escamas delgadas, obtusas, café claro y glabras o con indumento escaso, con una
costilla media tenue; nuez ovoide, café claro, de 10-13 mm de largo y 10 mm de ancho, con puberulencia
gris, glabrescente, incluida cerca de un tercio en la cúpula.
Distribución.—Se distribuye en la Sierra Madre Occidental en los estados de Chihuahua, Durango, Mi-
choacán, Zacatecas, Aguascalientes, Guanajuato, en la Sierra Madre Oriental desde Coahuila, Nuevo León
y San Luis Potosí. En Coahuila solamente se encuentra en la sierra del Carmen, en bosque de pino-encino,
asociado con Quercus rugosa, Q. carmenensis, Q. hypoleucoides, Cupressus arizonica, Arctostaphylos pungens,
Garrya ovata, Pinus strobiformis, Pinus greggii, Pseudotsuga menziesii. Crece entre 2300—2600 m de altitud,
en valles intermontanos altos y laderas altas con exposición norte o noroeste. En cañones, es frecuente se
desarrolle como árbol de hasta 10 m de alto. Especie rara en el estado.
(F ) de Coahuil 1273
Villarreal et al ,Q
Notas.—Se le puede reconocer por los pecíolos con puberulencia negra, las hojas gruesas, oblongas
a obovadas, generalmente de menos de 6 cm de largo, margen revoluto, dentado con aristas cortas, envés
T
amarillento con nervaduras prominentes y pelos fasciculados multirradi Escamas de la cápula con una
costilla tenue. Se puede confundir con Quercus hintoniorum o Q. hypoxantha por la forma de las hojas.
Material examinado: Mpio. Acuña: sierra del Carmen, campo Dos, 28°59'N y 102°36'W, 2,270 m, 9 ago 2004, D. Rishind et al. 2975
(ANSM).Mpio. Arteaga: sierra Rancho Nuevo, al E del ejido San Juan de Dolores, 25?25'N y 100?33'W, 2,660 m, 22 sep 2001, J.A.
Encina y S. Cruz 959 (ANSM).Mpio. Saltillo: sierra Zapalinamé, cañón de San Lorenzo, 25%20'N y 100°55'W, 2,532 m, 6 jul 2000, J.A.
Encina y R. Cardenas 455 (ANSM
portan lo que al parecer son híbridos entre Quercus sideroxyla y Q. mexicana: Mpio. Arteaga: sierra de Artega, cañón La
Carbonera, 25*16'N y 100°35'W, 2,150 m, 21 jun 1995, J.A. Encina et al.161 (ANSM), San Francisco Tarcuy, 2,285 m, 29 oct 1994, G.B.
Hinton 24926 (ANSM)
Quercus sinuata Walter, Fl. Carol. 235. 1788. (Fig. 30).
Quercus durandii Buckley, Proc. Acad. Nat. Sci. Phila. 12:445. 1860.
Se distribuye en el sur de Estados Unidos desde el sur de Carolina hasta el centro y oeste de Texas, llegando
al noreste de México a los estados de Coahuila, Nuevo León y Tamaulipas.
Quercus sinuata Walter var. breviloba (Torr.) C.H. Mull. in I.M. Johnst., J. Arnold Arbor. 25:439. 1944.
Quercus obtusifolia D. Don var. breviloba Torr., U. S. Mex. Bound. Surv, Rpt., 2:206. 1859. Quercus breviloba (Torr.) Sarg., Gard. &
Forest 8:93. 1895. Quercus durandii Buckley. var. breviloba (Torr.) Palmer, Amer. Midl. Naturalist 33:516. 1945
Quercus annulata Buckley, Proc. Acad. Sci. Phila. 445. 1860.
Quercus san sabeana Buckley, in Young, E Tex. 507. 1873.
Arbusto, raras veces árbol de hasta 5 m de alto; corteza gris, delgada y frágil; ramillas de 2-3 mm de grosor,
café grisáceo, glabras o con pelos esparcidos; yemas de 2-3 mm de largo, ovoides, glabras o con indumento
esparcido, café-rojizas; estípulas caedizas, de 3—5 mm de largo, filiformes y liguladas, pubescentes. Hojas
deciduas, delgadas y flexibles; pecíolos de 2-3 mm de largo, d o c D obovados a
oblanceolados u oblongos, de 2.5)3-6(-8) cm de largo y 2-4 cm do, base cuneada
u obtusa, raras veces atenuada, margen plano, entero a irregularmente denade y one lobulado,
haz glabro y lustroso, o con pelos esparcidos, envés opaco, canescente, con indumento de pelos fasciculados
diminutos, apresados, y pelos simples en la nervadura media; nervaduras 7-11 en cada lado, frecuentemente
con intermedias, ramificadas y anastomosadas hacia el margen, algo salientes en el haz, muy prominentes
en el envés. Inflorescencias masculinas de hasta 8 cm de largo, con flores laxas. Inflorescencias femeninas
de 3-7 mm de largo, 1-3 flores, con indumento denso. Fruto anual, solitario o en pares, subsésil o sobre
pedúnculos pubescentes de hasta 7 mm de largo; cúpula de 10-12 mm de ancho y 6-8 mm de alto; escamas
de la cúpula ovadas, obtusas, bases engrosadas y pubescentes, el ápice delgado y puberulento, café rojizo;
nuez ovoide a elipsoide de 7-15 mm de largo y 7-10 mm de ancho, glabra, café claro, incluida cerca de un
cuarto en la cúpula.
Distribución.—Se distribuye en Oklahoma, Texas, Coahuila, Tamaulipas y el centro de Nuevo León.
En Coahuila se encuentra en las sierras del Carmen, Santa Rosa (sierra de San Manuel) y La Gavia (sierra
del Oso), forma parte del matorral submontano, bosque de encino y bosque de pino-encino. Se asocia gen-
eralmente con Quercus invaginata, Q. morhiana, Q. arizonica, Q. laceyi, Q. gravesii, Rhus virens, R. aromatica,
Bumelia lanuginosa, Colubrina greggii, Juniperus flaccida, Pinus remota, Leucaena glauca y Ungnadia speciosa.
Se presenta entre 1200-1600 m de altitud, en laderas bajas con exposición norte y noreste, en cañones o
laderas con rocas calizas. Frecuente en las sierras del norte, su abundancia disminuye hacia las sierras del
centro del estado. Especie poco abundante en el área del estado.
Notas.—Se reconoce por las hojas delgadas, lustrosas y casi glabras, oblongas a obovadas, generalmente
de menos de 6 cm de largo, ápice obtuso a redondeado y base cuneada, margen entero, dentado a cortamente
lobulado, envés con pelos fasciculados, diminutos y escasos. Bellota ovoide (corta y ancha) con escamas
de la cúpula con base engrosada. Se le puede confundir con Quercus grisea o Q. mohriana e incluso con Q.
laceyi, por la forma de las hojas.
1274 J | of the Botanical R h Institute of Texas 2(2)
Mat M Santa Rosa, hacienda La Babia, cañón Rincón de Maria, 28°28'N y 102°02'W, 1,200 m, 27
abr 1975, T. Wendt y D. Rishind 959 (ANSM, TEX).
Quercus striatula Trel., Mem. Natl. Acad. Sci. 20:93, pl. 151. 1924. (Fig. 31). Two: MÉXICO. Duranco: R. Endlich
s.n. (HOLOTIPO: B).
Quercus cordifolia Trel., Mem. Natl. Acad. Sci. 20:84, pl. 125. 1924. Tiro: MÉXICO. Coanuna: Carneros Pass, E. Palmer 1278 (isotipo:
Quercus microphylla Neé, Anal. Cienc. Nat. 3:264. 1801.
Quercus striatula Trel. f. otinapensis Trel., Mem. Natl. Acad. Sci. 20:94, pl. 152. 1924.
Arbusto o arbolito de (0.3—) 0.5-2.5 m de alto, con crecimiento colonial rizomatoso; ramillas de 1-2 de mm
grosor, amarillo rojizo, glabrescentes; yemas ovoides, de 1-1.5 mm de largo, café rojizo, con las escamas
ciliadas; estípulas de 1-5 mm de largo, filiformes o subuladas, caedizas. Hojas deciduas, planas, un poco
coriáceas; pecíolos de 1.5—3 mm de largo, pubescentes; limbos oblongos hasta ovados o lanceolados de
(13234) cm de largo y (0.5-)1-2 cm de ancho, ápice agudo a obtuso, mucronado, base redondeada a
cordada, margen entero o con pocos dientes toscos y mucronados, en ocasiones ondulado, haz opaco a poco
lustroso, verde a verde grisáceo con pelos fasciculados esparcidos, envés con indumento gris-amarillento,
denso, de pelos fasciculados estipitados; nervaduras de 6—8(-10) en cada lado. Inflorescencias masculinas
cerca de 2 cm de largo, raquis pubescente. Inflorescencias a de (4—-)10—20(—-40) mm de largo, con
1-28) flores. Fruto anual, solitario o en pares, subsésil o sol los de 4-8(-12) mm de largo, cúpula
hemisféricas de 8-12 mm de ancho y 5—7 mm de alto, las escamas engrosadas en la base con indumento
denso, ápice delgado y glabro, de color café rojizo, flojamente apresadas; nuez ovoide o subesferica, de 8—12
mm de largo y 7-10 mm de ancho, glabra, café claro, incluida de un tercio a un cuarto en la cúpula.
Distribución. —Se distribuye en San Luis Potosí, Zacatecas, Durango y Nuevo León. En Coahuila se en-
cuentra en el sur y sureste del estado, en las sierras de Jimulco, Parras, La Concordia, Carneros, Zapalinamé,
Arteaga, llega hasta el centro en la sierra La Paila, en comunidades del matorral submontano, bosque de
pino y en bosque de pino piñonero. Generalmente se asocia con Quercus pringlei, Q. intricata, Q. saltillensis,
Rhus virens, Dasylirion cedrosanum, Purshia plicata, Juniperus saltillensis, Calia secundiflora, Lindleya mespiloides,
Agave scabra, Pinus cembroides y Mimosa biuncifera. Se desarrolla en altitudes entre 1800-2700 m, como un
arbusto o subarbusto rizomatoso, formando parte de los matorrales en las laderas de colinas y montañas,
en las exposiciones noreste, este y oeste, con suelos someros. Crece en el matorral submontano y el bosque
de pino piñonero en las sierras ubicadas en el sur del estado, ocasionalmente se le encuentra a orillas del
bosque de pino, de pino-encino y de oyamel, es escaso en la sierra La Paila.
Notas.—Se puede reconocer las hojas delgadas, verde grisáceas, lanceoladas a oblongas, menos de 4
cm de largo, ápice agudo a obtuso y base cordada a redondeada, margen entero o con dientes mucronados,
envés amarillento con indumento denso de pelos fasciculados. Escama de la cápula con base engrosada.
i confundir con Quercus saltillensis por la forma de las hojas.
M ] : Mpio. Arteaga: sierra de Arteaga, cañón de Jamé, 25°26'N y 100°30'W, 2,600 m, 13 ago 1996, J.A. Villarreal y J.
Zavala 8404 (ANSM), La Siberia, 25°15) N y 100°29'W, 2,400 m, 21 jun 1995, J.A. Encina et al. 156 (ANSM). Mpio. Castaños: sierra La
Gavia, rancho la Gavia, 26°10'N y 101°18'W, 1,500 m, 10 ago 1995, M.A. Carranza y J. Encina 173 (ANSM). Mpio. General Cepeda:
sierra La Concordia, ejido La Casita, 25°13'N y 101°25'W, 2,200 m, 27 sep 1996, J.A. Encina y J. Zavala 183 (ANSM). Mpio. Saltillo:
rancho Los Angeles, 25°06'N y 101°06'W, 2,100 m, 11 nov 1994, J.A. Encina 130 (ANSM), sierra de Zapalinamé, El Penitente, 25°21'N
y 100° 55W, 2,445 m, 15 sep 2005, J.A. Encina et al.1494 (ANSM). Mpio. Torreón: sierra de Jimulco, camino hacia la cima, 25°10'N y
103°15'W, 2,000 m, 26 sep 1995, J.A. Villarreal y M.A. Carranza 8324 (ANSM)
us t lata Liebm., Overs. Kongel. Danske Vidensk. Selsk. Forh. Medlemmers buius 181. 1854.
(Fig. 32). Tro: MÉXICO. Smaroa?: between Mazatlán and Durango, B.C. Seemann 1970 (notorio: K; sorro
Quercus idonea Goldman, Contr. U.S. Natl. Herb. 16:321. 1916.
Quercus standleyi Trel., Contr. U.S. Natl. Herb. 23:181. 1922.
Quercus monterreyensis Trel. et C.H. Mull., Bull. Torrey Bot. Club 63:151. 1936.
Árbol de hasta 10-12 m de alto, con tronco de hasta 30 cm de diámetro; corteza delgada, gris, surcada;
ramillas de 1.5—3 mm de grosor, café o café rojizas, con lenticelas prominentes, pubescentes a glabrescentes;
Villarreal et al., Q (F ) de Coahuil 1275
MAI big.
B, variación en Fic. 30. Quercus sinuata. A, rama con hojas; B, hoja; C, pelo fasciculado;
D, bellota.
J 7
I
-g
<.
=
ay
aj
S
a
D.
5
=
=
"(1
4
ba
Tem!
B. hoia por haz
ja |
P | 1 ^ I
>
Fic. 31. Quercus striatula. A, rama con hojas y bellota; B, variación dela Fic. 32. Q berculata. A, hojas y
oja; C | fasciculad D, bellota; E, de | fj | y envés: C, pelo fasciculad ipitado; D, bellota, E,
vista frontal y lateral.
yemas ovoides, obtusas o agudas, de 1-2(3) mm de largo, glabras; estípulas subuladas de 4-6 mm de largo,
pubescentes, persistentes solo en las yemas terminales. Hojas deciduas, coriáceas; peciolos de 4-10(-20)
mm de largo, rojizos cerca de la base, glabros o pubescentes; limbos oblongos, de elípticos a ovados u obo-
vados, de (28-12-24) cm de largo y (1.524-6(-12) cm de ancho, ápice obtuso a agudo, base cordada
a redondeada, raras veces cuneada, margen ligeramente cartilaginoso, engrosado, entero o dentado en la
[| | af 41 D A ID h lartik t
1276 f Texas 2(2)
porción distal de la hoja; haz verde lustroso, finamente reticulado, con el margen ondulado, glabro o con
pelos fasciculados esparcidos, envés verde con tonos café amarillento, glabro o con indumento fasciculado
estipitado, usualmente restringido a las nervaduras; nervaduras 8-10 en cada lado, un poco impresas en
el haz, formando un retículo prominente en el envés, ramificadas y anastomosadas hacia el margen. In-
florescencias masculinas de 5-8 cm de largo. Inflorescencias femeninas de hasta 3 cm de largo con 2-3
flores sobre pedúnculos cortos. Fruto anual, solitario o en pares, subsésil o con pedúnculos cerca de 5 mm
de largo, cúpula de 12-15(-20) mm ancho y 10-12 mm de alto, escamas engrosadas (tuberculadas) y con
puberulencia canescente en la base, ápice café, delgado y glabro; nuez ovoide de 15-20(-25) mm de largo,
con pelos diminutos, de pelos simples cerca del ápice o glabrescente.
Distribución.—Se distribuye en el noroeste de la República Mexicana, en los estados de Sonora, Chi-
huahua, Sinaloa y Durango, así como en Aguascalientes, Michoacán y Guanajuato llega hasta el este a la
Sierra Madre Oriental en Nuevo León. En Coahuila se encuentra en el centro-este del estado solamente en
la sierra La Gavia, particularmente en el cañón La Gavia y en la sierra Pajaros Azules, en bosque de encino,
se asocia con Quercus invaginata, Q. laceyi, Q. gravesii, Juglans mollis, J. microcarpa, Fraxinus greggii, Colubrina
greggi, Rhus virens, Pistacia texana y Ungnadia speciosa. Generalmente crece en altitudes entre 1400-2200
m, sobre laderas bajas y en fondo de cañones húmedos, con exposición norte o noroeste. Especie rara para
el área del estado.
Notas.—Se reconoce por las hojas coriáceas, verde oscruro, brillantes, casi glabras, oblongas a obova-
das, usualmente de menos de 12 cm de largo, margen entero a dentado, envés con pelos fasciculados en la
nervadura media. Bellotas de 1.5-2.5 cm de largo, con escamas de la Da con base tuberculada.
, 795 m, 24 nov 2002,
M :A] nado: M C le] : Pai A zi hoS M los lí Lampazos N.I
J
J.J. Medellin V. 489 (CEND.
GLOSARIO
Basado en Font Quer (1979) y Moreno (1984).
s +
ESTÍPULAS dea laminares que se forman en número
cada lado de la base folia
ACUMINADO C
ulo menor or de 45 grados
en án
AreLPADO Con E largos que forman una mata densa
e fieltr
AMENTO Inforscenci espigada, decidua, generalmente
peq y unisexuales.
S. Con I id pul vejigas.
ANASTOMOSADO Caracterizado por tener dos o más nervios
uelven a unir.
Arista Estructura setosa, elongada y fl
e una vena desnuda cubierta por tejido corneo.
Bettota Aquenio agrandado o nuez, derivado de un
ovario infero, rodeada por un involucro seco llamado
ible, consistiendo
`
MAI h J J
CAEDIZO
CANESCENTE (cia) indumento dus se vuelve blanco o
o-grisáceo con el tiem
Castaño De color café-rojizo
Citano Con pelos marginales.
CONDUPLICADO Doblado a lo largo del eje central o nervio
dondeados.
Crispapo Con los márgenes divididos, levantados a in-
OS
" . laz al: I £, ri
copa, iodeando el fruto o la flor femenina en encinos.
EscLERosApo Endurecido, como de piedra.
FILIFORME E forma alargada y del on
FimBrIADO Franjeado o con cilios gruesos y densos.
GLasRO Sin Dui tipo de indumento lampifio.
hus ti
De colar verde claro, con matiz ente
Il
VET.
azulado.
Impreso Nervaduras marcadas en la superficie del haz.
InDuMENTO Conjunto de pelos, glándulas, escamas, etc., que
Era el A ni | | +
2 r
F|
Cii Iad P
INvo.uro Con | á superficie
superior o ventr al.
LaNoso Con pelos largos, suaves y entrecruzados seme-
jantes a la lana.
LIGULADO
parte abrazadora.
NOME
T £, J ce I I!
p
llamados lóbulos
equeña, que se proyecta a partir de la
rvadura, cubierta por tejido corneo
Osovapo De forma ovada, pero con el ápice más amplio
Ovapo Con la hase más ampia que el Spices.
OBLANCEOLADO D
ancho que la base):
PaPiLoso C
forma más o > menos cónica) en la superficie.
lA ] COEM AT J
Villarreal et al ,Q
PELOS GLANDULARES Glándula epidérmica peliforme. Pelo con
una estructura globosa secretora en el ápice, puede
ser unicelular o multicelular, peltado, pedúnculado
ésil.
Pins Con pelos largos y suaves.
PuBERULENTO Con pelos simples muy finos, cortos y escasos.
Diminutamente pubescente.
Pusescente Con pelos simples y delgados, finos y rectos.
PuLveRULENTO Con una cubierta de polvillo diminuto.
Repanpo Escasamente sinuado
RevoLuTo Con | árg
inferior o dorsal.
Seríceo Con pelos largos y sedosos, generalmente ad-
presos
I I Ļ | Er
SINUADO C
1277
SuBULADO Estrechado hacia el ápice hasta rematar en
punta fina.
Tomento Conjunto de pelos largos y muy entrecruzados
ue cubren totalmente la superficie
TowENTULOSO Escasa o cortamente ‘erento diminuta-
m ntoso
TRICOMA ee del tejido epidérmico, a menudo en
de pelo.
TunBINADO En fi d
y ancho en el ápice.
VgLurINO Cubierto por pelos densos, largos, suaves y rectos,
como el terciopelo
VeLLoso Con pelos largos y suaves curvados pero no
4i + h lak
afelpados.
Vermirorme Alargado, como en forma de gusano.
I
mi T F EN +
central de la estructura; especialmente del margen.
AGRADECIMIENTOS
Agradecemos a Ross A. McCauley y Susana Valencia Avalos la revisión y sus comentarios al manucrito
original. A la Universidad Autónoma Agraria Antonio Narro su apoyo a traves del proyecto 0205 2503. A
Erika Reyna la elaboración del mapa.
REFERENCIAS
ANÓNIMO. 1983. Síntesis geográfica de Coahuila, México. Instituto Nacional de Estadística, Geografía e Informática.
Secretaría de Programación y Presupuesto.
BANDA, R. 1974. Contribución al conocimiento de los encinos del estado de Nuevo León, México. Tesis. Fac. de
Ciencias Biológicas. UANL. Monterrey, N.L, México
CorreLL, D.S. AND M.C. JoHNsTON. 1970. The manual of the vascular plants of Texas. Texas Research Found. Renner,
Texas.
ENCINA, J.A. v J.A. VILLARREAL. 2002. Distribución y aspectos ecológicos del género Quercus (Fagaceae), en el estado
de Coahuila, México. Polibotánica 13:1-23.
ESPINOSA, J. 1981. Fagaceae. In: Rzedowski, J. y G.C. de Rzedowski, eds. Flora fanerogámica del Valle de México.
Vol. 1 C.E.C.S.A. México, D.F. Pp. 104-114,
FONT Quer, P. 1979. Diccionario de Botánica. Ed. Labor. Barcelona, España.
GonzáLez, R. 1992. Sinonimia de los encinos mexicanos. Memorias. Ill Seminario Nacional sobre utilización de
encinos. Rep. Ci. Fac. Ci. Forest., UANL. 15(2):876-892.
Marroquín, J.S. 1976. Vegetación y florística del nordeste de México, I. Aspectos sinecologicos en Coahuila. Rev.
Soc. Mex. de Hist, Nat. 36:69-101
MARROQUÍN, R.A. 1985. El género Quercus L. al noroeste del estado de Nuevo León. Tesis. Fac. de Ciencias Biológicas
UANL, Monterrey, N.L. México.
Martínez, M. 1981. Los encinos de México. An. Inst. Biol. Méx. No. 8, 2a. Ed. Comisión Forestal, Michoacán,
México.
MCVAUGH, R. 1974. Fagaceae. In: Flora Novo-Galiciana. Contr. Univ. Mich. Herb. 12(3):1-93.
Moreno, N.P. 1984. Glosario botánico ilustrado. C.E.C.S.A. México, D.F. 1a. edición.
Mutter, C.H. 1940. Oaks of the Trans-Pecos, Texas. Amer. Midl. Naturalist. 24:703-728
Muiter, C.H. 1951. The oaks of Texas. Contr. Texas Res. Found. Renner, Texas. 1:21-323.
MuLLer, C.H. (In press). Chapter 29: Fagaceae. In: Henrickson, J. and M.C. Johnston, eds. Flora of the Chihuahuan
Desert Region. Chihuahuan Desert institute, Alpine, TX. Pp. ca. 30.
Nixon, K. AND C.H. MULLER, 1992. The taxonomic resurrection of Quercus laceyi Small (Fagaceae). Sida 15:57-69.
NIXON, K. AND C.H. Mutter. 1993. The Quercus hypoxantha complex (Fagaceae) in northeastern Mexico. Brittonia
45:146-153.
1 lof tha Botanical R h Institut f Texas 2(2)
1278
Nixon, K.C.1993a. Infrageneric classification of Quercus (Fagaceae) and typification of sectional names. Ann. Sci.
Forest. 50:25-34.
Nixon, K.C. 1993b. The genus Quercus in Mexico. In: T.P. Ramamoorthy, R. Bye, A. Lot and J. Fa, eds. Biological
diversity of Mexico: origins and distribution. Oxford University Press. New York. Pp. 447-458.
Nixon, K.C. 1997. Fagaceae. In: Flora of North America and north of Mexico. NY and Oxford. 3:1-590.
Pinkava, D.J. 1984. Vegetation and flora of the Bolson of Cuatrocienegas Region, Coahuila, Mexico: IV summary,
Endemism and corrected catalogue. J. Arizona-Nevada Acad. Sci. 19:23-47.
Roprlcuez, G., A., J.A. VILLARREAL Y J, VALDES R. 1994. Vegetación y flora de la Sierra de Parras, Coahuila. Agraria
10(2):79-109,
VALDEZ, T, V. Y M.L. Acuitar. 1983. El género Quercus en las unidades fisonómico-florísticas del Mpio. de Santiago,
N.L. México. INIF-SARH. México. Bol.Tec. No. 98.
VALENCIA, S. 2004. Diversidad del género Quercus (Fagaceae) en México. Bol. Soc. Bot. Mex. 75:33-53.
Vazquez, M.L. 2006. Trichome morphology in selected Mexican red oak species (Quercus section Loabatae). Sida
22:1091-1110.
VILLARREAL, J.A., VALDÉS, J. Y J. L. ViLLaseñor. 1996. Corología de las Asteraceas de Coahuila, México. Acta Bot. Mex.
VILLARREAL, J.A. Y J. VALDÉS R. 1993. Vegetación de Coahuila. Manejo de Pastizales 6(1-2):9-18.
VILLARREAL, J.A. 1994, Vegetación y flora de la Sierra de La Paila, Coahuila, México. Sida 16:109-138.
VILLARREAL, J.A. 2001. Listados floristicos de México. XXIII Flora de Coahuila. Instituto de Biología. Universidad
Nacional Autónoma de México.
VILLARREAL, J.Á. y J.A. Encina D. 2005. Plantas vasculares endémicas de Coahuila y algunas áreas adyacentes,
México. Acta Bot. Méx. 70:1—46.
THOMAS WALTER TYPIFICATION PROJECT, VI:
NEOTYPES FOR AN ADDITIONAL 18 WALTER NAMES
Daniel B. Ward
dE of Botany
sity of Florida
NE s 32611, USA.
ABSTRACT
Flora Caroliniana was oa ae uh Walter " 1788. B fth ly date of this fl any of its species were new to sci-
nce. and t] in the A ican Soutl un: Walter kept no herbarium, recent
T
efforts nave chosen later OE as re cds types. Her ini } ftl dd d, with selection of
P y p g p
18 Thomas Walter names.
rr Es JF E
RESUMEN
Flora Caroliniana fue PE por qme bud en 1788. Penge a la fecha ind: de esta flora, muchas de estas especies fueron
nuevas a la ciencia, y l por Y g I el Sureste de América. Aunque i no id un
herbario, esf i I gido colecci 5 : 1 1 A 1 :
ion d i iad 18 nombres adicionales de Thomas Walter.
In the 1780s, Thomas Walter (1740-1789) owned and operated a rice plantation on the Santee River, Berkeley
County, South Carolina. His interest in the plants of the area led him to compile a simple flora in which
he gave them descriptions and Latin names. This book, Flora Caroliniana (1788), is the first treatment of
American plants employing the binomial nomenclature and sexual classification system of Linnaeus. Of
even greater importance, the early date of this publication has given many of his names priority over those
of later authors.
This paper largely completes typification of the plant names published by Walter in his pioneer Flora.
By the selection of type specimens to represent these names, they become fixed in their meaning, bringing
stability to the nomenclature of this portion of the sala flora.
Walter kept no herbarium (Ward 20072). In the ab thentic original materials, the International
Code of Botanical Nomenclature (McNeill et al. 2006) permits selection of another specimen—a neotype—
to represent the missing type.
The Thomas Walter Typification Project (Ward 2006a, 2007b, 2007c, 2007d, 2008) and ancillary
studies (Ward 2006b, 2007e, 2007f, 2007g) have addressed the typification of many of the names assigned
to American plants by Walter. These studies also treat aspects of the relationship between Walter and the
Scottish horticulturist John Fraser. Fraser's herbarium (the Fraser/Walter herbarium, BM), gathered in the
Carolinas and Georgia in 1787, was briefly seen by Walter, but is largely irrelevant to typification of Walter's
names. Selection of collections made by other persons is the only pathway to stability of the great majority
of Walter's names.
In previous numbers of the Project (2007d, 2008), the Harvard University Herbaria (GH) were the
source of selected specimens. Here, for those species not adequately represented in the holdings of that
institution, the herbaria of the University of South Carolina (USCH) and Duke University (DUKE) have
provided specimens of quality appropriate for selection as neotypes.
With inclusion of the species treated here, essentially all of Walter’ that are in common use have
now been assigned types, as have an appreciable number of Walter's names usually referred to synonymy.
Still other of his names that have been treated as synonyms, or are unidentified as to their modern mean-
ing, remain unaddressed. A subsequent number of the Project, now in preparation, will provide an index
to typification or other disposition of all Thomas Walter names.
I. Rat. Res. Inst. Texas 2(2)- 1379 — 1283. 2008
1280 j | of the Botanical R h Institute of Texas 2(2)
An additional 18 of Walter's species are here assigned neotypes. Specimen citations, if not otherwise
attributed, are to the Fraser/Walter herbarium (BM) as described elsewhere (Ward 2006a). The typifications
are presented in the format employed previously, in alphabetical sequence, using the names given them by
Thomas Walter.
LYPIFICALIONS
WALTER'S NAME: Amsonia Tabernaemontana Walter (p. 98)
MODERN NAME: Amsonia tabernaemontana Walt.
Typical Amsonia tabernaemontana, with ovate leaves, is rare in eastern SC, frequent westward. The narrow-
leaved var. salicifolia (Pursh) Woodson is frequent on the SC coastal plain. Walter described his plant with
leaves "ovatis." Spm. 5-C, labeled “Amsonia Tabernaemontana” by Fraser, has the broader leaves of the typical
variety; there is no evidence it was seen by Walter. Though the ovate-leaved form may not have been the
variation most familiar to Walter, to preserve accepted usage (Woodson 1928: 406) a wide-leaved speci-
men, Smith 1114, 3 June 1941 - USCH, from Witherspoon Island, Darlington County, South Carolina, is here
selected as NEOTYPE for Amsoria tabernaemontana Walt. It has been suggested that Walter's name must have
been taken from Tabernaemontana amsonia L. (1762) and that Walter's type must therefore be the same as
Linnaeus (J.S. Pringle, pers. comm., Dec 2005). In such case, Walter's name would be superfluous and thus
illegitimate. Selection of a Walter neotype leaves the suggestion in the realm of conjecture.
WALTER'S NAME: Anemone caroliniana Walter (p. 157)
MODERN NAME: Ánemone caroliniana Walt.
Very rare in SC (3 counties, all inland), rare also in central GA. This distant range indicates the species
was probably brought to Walter by John Fraser. There is no specimen in the Fraser/Walter herbarium (BM).
Pyron 2109, 19 Mar 1938 - DUKE (annot. Carl S. Keener 1980), from 3 mi. SW of Irwinville, Irwin County,
Georgia, is here selected as NEOTYPE for Anemone caroliniana Walt.
WALTER'S NAME: Arethusa spicata Walter (p. 222)
MODERN NAME: Hexalectris spicata (Walt.) Barnhart
Infrequent throughout. Blake (1915: 136) has stated Walter's diagnosis is "quite distinctive" of this spe-
cies. There is no specimen. Evett s.n., 9 July 2000 - USCH (annot. John Nelson 2000), from Rucker Road,
Chapin, Richland County, South Carolina, is here selected as NEOTYPE for Arethusa spicata Walt., basionym
of Hexalectris spicata (Walt.) Barnhart.
WALTER'S NAME: Athanasia paniculata Walter (p. 201)
MODERN NAME: Verbesina walteri Shinners [- Ridan paniculata (Walt.) Small]
Verbesina walteri is rare in SC, known only from three counties. Walter's epithet is not transferrable to Verbe-
sina (not V. paniculata Poir., 1808). Shinners (1964) formed a new name, basing it upon Walter's name and
description. No specimen has been identified in the Fraser/Walter herbarium (BM). The species is infrequent
through much of its range, west to Texas. Available material, though from far beyond the reach of either
Walter or Fraser, is believed representative of plants Walter might have known. Thieret 24848, 27 Sep 1966
- DUKE (annot. James R. Coleman 1969), from 8 mi. SE of Denham Springs, Livingston Parish, Louisiana,
is here selected as neoryre for Athanasia paniculata Walt., the basis for Verbesina walteri Shinners.
WALTER'S NAME: Frasera caroliniensis Walter (p. 88)
MODERN NAME: Swertia caroliniensis (Walt) Kuntze
Very rare (3 counties in SC.: Abbeville, Greenwood, Laurens, all on piedmont); undoubtedly a Fraser dis-
covery. A supposed specimen (48-B), labeled “Frasera Caroliniensis" by Fraser, has since been identified as
Dodecatheon meadia (Britten 1921: 70). Horn 6581, 8 May 1993 - USCH, from Sumter National Forest, ca.
11 mi. NE of Clinton, Laurens County, South Carolina, is here selected as NEOTYPE for Frasera caroliniensis
Walt., basionym of Swertia caroliniensis (Walt.) Kuntze.
Ward, Thomas Walter ay pification VI Iditi l I ty I 1281
WALTER'S NAME: Hydrolea quadrivalvis Walter (p. 110)
MODERN NAME: Hydrolea quadrivalvis Walt.
Frequent in eastern SC. No specimen has been identified. Horn 4246, 30 July 1990 - USCH, from just N of
Laurel Hill Plantation, 2 mi. SE of Goose Creek, Berkeley County, South Carolina, is here selected as NEOTYPE
for Hydrolea quadrivalvis Walt.
WALTER'S NAME: Hypericum tubulosum Walter (p. 191)
MODERN NAME: Triadenum tubulosum (Walt.) Gleason
Very rare in SC Q counties, both inland from Walter's Berkeley Co.); rare in central GA; not known in
NC. But Walter's description seems unambiguous, clearly contrasted with T. virginicum and T. walteri. This
probably is a Fraser discovery, perhaps obtained on his trip to the Flint River, west-central GA, in spring
of 1787. There is no specimen. Allison 10271, 16 Oct 1997 - DUKE, from NE corner of Ft. Benning Military
Reservation, 17.5 mi. NE of Cusseta, Chattahoochee County, Georgia, is here selected as NEOTYPE of Hypericum
tubulosum Walt., basionym of Triadenum tubulosum (Walt.) Gleason.
WALTER'S NAME: Iris hexagona Walter (p. 66)
MODERN NAME: Iris hexagona Walt.
Very rare in SC (known only in Charleston Co.). There is no specimen. Hutto s.n., 4 June 1993 - USCH,
from branch of Wambah Creek, Hampton Plantation State Park, Charleston County, South Carolina, is here
selected as NEOTYPE for Iris hexagona Walt.
WALTER'S NAME: Laurus melissaefolia Walter (p. 134)
MODERN NAME: Lindera melissifolia (Walt.) Bloom
Rare; in SC known only in 3 counties (Berkeley, Colleton, Darlington). Spm. 63-C was named “Laurus melis-
saefolium" by Walter and later annotated "Benzoin melissaefolium" by A. J. Kosterman. Its leaves are smaller than
other specimens of that species. Spm. 63-H is similar, but was annotated by Kosterman as “Benzoin odoriferum,”
a synonym of L. benzoin (L.) Blume, which it could well be. An independent specimen of certain identity,
Williamson 1550, 23 July 1961 - USCH, from 1 mi. NE of Honey Hill, Darlington County, South Carolina, is
here selected as NEOTYPE for Laurus melissaefolia Walt., basionym of Lindera melissifolia (Walt.) Bloom.
WALTER'S NAME: Melanthium hybridum Walter (p. 125)
MODERN NAME: Melanthium hybridum Walt. [= Veratrum latifolium (Desr.) Zomlefer]
Rare in SC, frequent in western NC; likely a Fraser discovery. No specimen has been identified. Nelson 8431,
12 Sept 1989 (2 sheets) - USCH (annot. D. E. Kennemore), from W of Marietta, Greenville County, South
Carolina, is here selected as NEOTYPE for Melanthium hybridum Walt.
WALTER'S NAME: Melanthium racemosum ? Walter (p. 126)
MODERN NAME: Tofieldia racemosa (Walt.) BSP.
Frequent in eastern SC. Walter was the first to form the combination Melanthium racemosum. [Others: M.
racemosum Michx., 1803 (= M. virginicum), and M. racemosum Roth, 1821 (= Iphigenia indica).] His expression
of doubt does not invalidate his name (Art. 34.1). No specimen has been identified. Pittman & Darr s.n., 1
July 1999 - USCH, from Shirley tract, Lynchburg-Savanna area, near SC 327, Lee County, South Carolina,
is here selected as NEOTYPE for Melanthium racemosum Walt., basionym of Tofieldia racemosa (Walt.) BSP.
WALTER'S NAME: Ophioglossum crotalophoroides Walter (p. 256)
MODERN NAME: Ophioglossum crotalophoroides Walt.
Very rare in SC (two counties, one of which is Berkeley). There is no specimen. Douglas s.n., 12 March
1991 - USCH, from along Simons Road, 1/2 mi. SW of SC 68, just W of Yemassee, Hampton County, South
Carolina, is here selected as neoTYPE for Ophioglossum crotalophoroides Walt.
1282 ical h Institute of Texas 2(2)
WALTER'S NAME: Pharnaceum maritimum Walter (p. 117)
MODERN NAME: Sesuvium maritimum (Walt.) BSP.
Infrequent along SC shore. No specimen. Nelson 5850, 28 July 1987 - USCH, from South Island, Yawkey
Wildlife Center, Georgetown County, South Carolina, is here selected as NEOTPE for Pharnaceum maritimum
Walt., basionym of Sesuvium maritimum (Walt.) BSP.
WALTER'S NAME: Rajania ovata Walter (p. 247)
MODERN NAME: Brunnichia ovata (Walt.) Shinners [- Brunnichia cirrhosa Banks ex Gaert.]
Rare in the Carolinas, but known from three coastal SC counties (Charleston, Georgetown, Jasper). Shinners
(1967) accepted Walter's description as applying to the plant also known as Brunnichia cirrhosa. Spm. 91-A,
marked by Walter as “Rajania (monoica)" and noted by Fraser to be the "ovata" of the Flora, is a worthless
scrap. Aulbach-Smith 3144, 15 July 1984 - USCH, from near Forks Lake Bluff, ca. 5 mi. NW of Hardeeville,
Jasper County, South Carolina is here selected as NzoTYPE for Rajania ovata Walt., basionym of Brunnichia
ovata (Walt.) Shinners.
WALTER'S NAME: Silphium scabrum Walter (p. 217)
MODERN NAME: Silphium scabrum Walt. [- Silphium dentatum Ell.]
Common throughout. Silphium asteriscus L. is a more scabrous plant, but is essentially absent from SC. Guy
Nesom (pers. comm., Aug 2007) has identified Walter's name as S. dentatum. Spm. 98-C was labeled “Silphium”
by Walter and “Scabrum” by Fraser; it is a fragment without meaningful characters. Nelson 5571, 9 June 1987
Q sheets) - USCH (annot. J. A. Clevinger 1999), from West Springs, Union County, South Carolina, is here
selected as NEOTYPE for Silphium scabrum Walt.
WALTER'S NAME: Sophora lanceolata Walter (p. 135)
MODERN NAME: Baptisia lanceolata (Walt.) Ell.
Rare in SC sandhills; perhaps a Fraser discovery. There is no specimen. Stanford 16, 4 April 1993 - USCH,
from Compartment 84, Savannah River site, Barnwell County, South Carolina, is here selected as NEOTYPE
for Sophora lanceolata Walt., basionym of Baptisia lanceolata (Walt.) Ell.
WALTER'S NAME: Thymbra caroliniana Walter (p. 162)
MODERN NAME: Macbridea caroliniana (Walt.) Blake
Infrequent in eastern SC. Blake (1915: 132) noted Walter's description to be "definitive." No specimen has
been identified. Horn 2362, 28 Aug 1987 - USCH, from Shaws Creek, 4 mi. N of Aiken, Aiken County,
South Carolina, is here selected as Neotyre for Thymbra caroliniana Walt., basionym of Macbridea caroliniana
(Walt.) Blake.
WALTER'S NAME: Utricularia fibrosa Walter (p. 64)
MODERN NAME: Utricularia fibrosa Walt. [- Utricularia striata LeConte ex Torr.]
Rare on SC coastal plain, frequent in piedmont. There is no specimen. Utricularia fibrosa Walt. was left in
limbo by Taylor (1989: 586) who determined Walter's plant was either U. biflora Lam. (1791) or U. striata
LeConte ex Torr. (1819), but then rejected both. Nelson 12474, 18 May 1992 - USCH, from Training Area
25D, Fort Jackson, Richland County, South Carolina, a plant identifiable also as U. striata, is here selected
as NEOTYPE for Utricularia fibrosa Walt.
ACKNOWLEDGMENTS
The specimens listed here, not available from our other cooperating herbarium, were made available by
John B. Nelson (USCH) and Robert L. Wilbur (DUKE). Their cooperation in locating adequate materials of
these less well-known species is greatly appreciated. As before, the Spanish abstract has been prepared by
Christine M. Housel (ABT). And as always, Charles E. Jarvis (BM), by virtue of his long-ongoing efforts to
typify the names of Carl Linnaeas, is godfather to the entire Thomas Walter Typification Project.
ale Arntiam Wie additinnal ptyr 1283
REFERENCES
BLaxe, S.F. 1915. Some neglected names in Walter's Flora Caroliniana. Rhodora 17:129-137.
BnrrrN, J. 1921. Thomas Walter (1740?-88) and his grass. J. Bot. 59:69—74.
McNeitt, J, F.R. Barrie, V. DEMOULIN, D.L. HAwKsworTH, K. MARHOLD, D.H. NicoLson, J. PRADO, P.C. SitvA, J.E. Scoc, J.H. WIERSEMA,
and N.J. Turano (eds.). 2006. International code of botanical nomenclature (Vienna Code). Gantner Verlag;
Ruggell, Liechtenstein.
SHINNERS, L.H. 1964. Notes [re Verbesina walteri]. Sida 1:253.
SHINNERS, L.H. 1967. Brunnichia ovata (Walter) Shinners, comb. nov. (Polygonaceae). Sida 3:115.
TAYLOR P. 1989. The genus Utricularia—a taxonomic monograph. Kew Bull., Addit. Ser. 14:1—724.
WarrtR, T. 1788. Flora Caroliniana. London.
Warp, D.B. 2006a. Thomas Walter Typification Project, |: Observations on the John Fraser folio. Sida 22:
1111-1118.
Warp, D.B. 2006b. Silene catesbaei, rather than Silene polypetala, the correct name of the endangered Fringed
Catchfly. Castanea 71:324—329.
Warp, D.B. 2007a. The Thomas Walter herbarium is not the herbarium of Thomas Walter. Taxon 56:917-926.
Warp, D.B. 2007b. Thomas Walter Typification Project, Il: The known Walter types. J. Bot. Res. Inst. Texas 1:
407-423.
Warp, D.B. 2007c. Thomas Walter Typification Project, Ill: Lectotypes and neotypes for 20 Walter names, as rec-
ognized in the Fraser/Walter herbarium. J. Bot. Res. Inst. Texas 1:425-430,
Warp, DB. 2007d. Thomas Walter Typification Project, IV: Neotypes and epitypes for 43 Walter names, of genera
A through C. J. Bot. Res. Inst. Texas 1:1091-2007,
Warp, D.B. 2007e. What in the world did Thomas Walter mean by Xxxxx yyyyy? Part one: The complete unknowns.
Phytologia 89:228-235.
Warp, D.B. 2007f. What in the world did Thomas Walter mean by Xxxxx yyyyy? Part two: The quite doubtfuls.
Phytologia 89:300-314.
Warp, D.B. 2007g. Quercus sinuata Walter—the hybrid of Q. falcata and Q. phellos—rediscovered and neotypi-
fied. Castanea 72:177-181.
Warp, D.B. 2008. Thomas Walter Typification Project, V: Neotypes and epitypes for 63 Walter names, of genera D
through Z. J. Bot. Res. Inst. Texas 2:475-486.
Woopson, R.E. 1928. Studies in the Apocynaceae. Ill: A monograph of the genus Amsonia. Ann. Missouri Bot.
Gard. 15:379-427.
ZIMMERMAN, J.H. 1958, A monograph of Veratrum. Ph.D. diss., Univ. of Wisconsin, Madison.
" I ^f tha Botanical D h Institut f Texas 2(2)
1284
BOOK REVIEW
Jackie M. Poore, WILLIAM R. Carr, DANA M. PRICE, AND JASON R. SINGHURST. 2007. Rare Plants of Texas: A Field
Guide. (ISBN-13: 978-1585445578, ISBN-10: 1585445576, pbk.). Texas A&M University Press, Lewis
Street, 4354 Texas A&M University, College Station, Texas 77843-4354, U.S.A. (Orders: www.tamu.edu/
upress, 800-826-8911). $35.00, 656 pp., color photographs and maps, drawings, 5 3/4" x 9 1/4".
Texas Parks and Wildlife Department botanists, Jackie M. Poole and David H. Riskind, published their first report in 1987 about Texas'
deitas or a pas T Mu. included only 21 Eus The authors began work on the new book in 2000. Over 200
d, imperiled, Rare Plants of Texas. Because most of these plants are
too rare to be mentioned, much less ds in standard field guides, dn a handful of botanists have known what these plants look
like in their natural oap
Complete with photographs, line drawings, and county Js oe ae sie. E KE listed candidate and species-of-
concern plants in Tas Individual itat flowering time, federal
and state status, similar species, and published ref Th I | id 1 hapters on the state's vegetation
E F F r o
f Texas; federal, state, and ot] king methods; ti ive pl hods;
a Bim guideline
his E ia isa E de den ids at the rare evils a i The ee will be asp En interest for
i
>
J
43 f 1; J> 1 1
the excellent Rare Plants of Texas wil ] And it i
personnel, non- a entities, cm concutit natural resource d naturalists, academics, and M
] | listed, candidate, or o of concern plants of Texas.
fi ] there is a section on the various natural (vegetation) regions of Texas. This
sectio into detail about the land ma ME various types of plants typically found in the 11 regions, illustrated by a colored map
of the sara regions of Texas. It is noted pod that rare ds are still being found today in Texas as much of the state has not been
p > 1 ES fil E 1 1 J 1 E J E 1 1 1 b MET
4 I a r
ily human-related are followed by the section on natural regions. This is followed by sections on “Management and Restoration of Rare
Plants” and “Nomenclature and Species eh A SUN oh the official “Wildlife Diversity Reporting Form” issued by the Texas Parks
Wildlife D , is also included, along with instructions for its use
a Treatments is the core section a F book. Each i plan is depicted in a drawing, a color picture, aud a location on a
c on name, family name, EE status Spam m itlives, and a
each plant
and
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detailed dd Itiple col d ink drawing v erall, 232 species of plants are illustrated
with 247 color photographs, 234 color distribünon maps, sd 215 superbly rendered line drawings by artist Linny Heagy, who has been
2000
associated
Following the species treatments is a short section on ie was dnd ne din book and why. A very detailed glossary of
terms, a 79-pag pag ud
Up until the last few decad latively little ] | ftl 's pl pecies in dang decline, thus making
conservation action decisions difficult. Now every Texan has the opportunity to take the fi p by gnizing ly what we are
in ee of losi
1 i 1 1 hes lla f, J 1 1 14 1 f 1 1 11 "m 1
g
layman interested in idu field. Those with a deep i in tl ic will iate it tl For ihe readership it was de ned for,
it works well and provides a glimpse of t! y rare plants found across Fa great state of Bn The decision to use a flex binding,
enabling readers to easily scan both pages devoted to each species without fear ling, is heartily applauded by this
reviewer.
ts of Ti i j ibuti 1 ledge ab Texas plants. The detailed inf i l I I
"Rare Plan
A 1 1 Mel 1
excellent line drawings
plants in the state,” said a M. Digg Jr. Ph D., an ae Ones DIG of n “Further, anyone generally interested in
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Xd5 DOLANY
tne history of pl ion in tt "— Gary L. Jennings, Botanical Research Institute of Texas, 500 East 4th Street, Fort
of Ti
Non Texas 76102-4025, US. Á.
J. Bot. Res. Inst. Texas 2(2): 1284. 2008
CARYOPHYLLACEAE IN NORTH AMERICA: CHANGE OVER THE LAST 110 YEARS
Richard K. Rabeler Ronald L. Hartman
Mis of joa Herbarium Rocky Mountain Herbarium
Varsity Drive Department of Botany
Ann TS Michi Huh 46108-2228, U.S.A. University of Wyoming
rabelergumich.edu Laramie, Wyoming 82071-3165, U.S.A.
rhartman@uwyo.edu
ABSTRACT
We compare the treatment ui = le in ls EA North a (b & Hartman 2005), with that in the Synoptical
Flora of North A (R g troduced species by subfamily, genus, and location of
npe or where first noted.
RESUMEN
Comp 1 : Fs ne ee yo] 1 yll la Flora of North A ica (Rabeler & H 2005). con el de la Synoptical Flora
of North A ira (Robi 1897), iend dici ] p i 3 i lucid I ili género, y localización
del d brimient donde f la por primera vez.
Publication of the Caryophyllaceae in Flora of North America [FNA] (Rabeler & Hartman 2005) presented
the opportunity to compare those results with previous efforts. Our treatment is the first to include all taxa
accepted for North America north of Mexico since that by B.L. Robinson (1897) in Synoptical Flora of North
America.
The family treatment for FNA was the combined effort of ten authors, two of whom also served as
editors. Four subfamilies were recognized in which 286 species and 54 infraspecific taxa belonging to 37
genera were treated in 212 pages of illustrated text.
By contrast, the Caryophyllaceae in Synoptical Flora of North America was published solely by B.L.
Robinson. lt first appeared as two papers (Robinson 1893, 1894) “preliminary to [the] treatment” in the
Flora and then in the second fascicle of the Synoptical Flora (Robinson 1897). Three subfamilies containing
167 species and 53 infraspecific taxa belonging to 19 genera were treated in 48 pages sans illustrations.
The Illecebraceae (Paronychioideae) were placed far from the Caryophyllaceae, following G. Bentham and
J.D. Hooker's (1862-1883) classification, and apparently were never published. Based on members of this
subfamily that were known by 1890, we estimate that 7 genera, 24 species, and 1 infraspecific taxon would
have been included in the Paronychioideae (and the tallies for the Caryophyllaceae), expanding the treat-
ment by several pages (Table 1).
The Caryophyllaceae worldwide contain about 3000 species in 83 (Silene s.l.) or 89 (Silene s.s.) genera
(Rabeler & Hartman 2005). We followed a four-subfamily classification which emphasizes morphological
features but is yet to be adequately tested by molecular analyses. The four subfamilies are: Alsinoideae (30
genera, 1040 species), Caryophylloideae (26 [20] genera, 1500 species), Paronychioideae (17 genera, 200
species), and Polycarpoideae (16 genera, 210 species). The most commonly cited classification (Bittrich 1993)
includes three subfamilies, treating the Polycarpoideae as a tribe within the Paronychioideae. Preliminary
evidence indicates that the Alsinoideae and the Caryophylloideae form a monophyletic group within the
family separate from members of the other two subfamilies (Nepokroeff et al. 2002; Fior et al. 2006).
The balance of this paper will compare the two treatments of Caryophyllaceae of North America (Rabeler
& Hartman 2005 and Robinson 1897) by subfamily and summarize the increases in both native and alien
members. Our analyses take into account differences in taxonomic rank, although subspecies and varieties
are combined under the collective term, infraspecific taxa.
J. Bot. Res. Inst. Texas 2(2): 1285 — 1289. 2008
1286 Journal of the Botanical R h Institute of Texas 2(2)
rH ET Kien £, Lin aL A fihaf
Tase 1. A compari fil ber of g (9), species (sp), and infraspecific ) by y yophy
Subfamily Robinson 1894, 1897 Rabeler & Hartman 2005
Alsinoidea 5 g/ 77 sp + 28 infra 16 g/ 137 sp + 32 infra
bu" 8 9/ 68 sp + 19 infra 8 9/89 sp + 14 infra
Paronychioideae 7 g/ 24 sp + 1 infra* 6 g/ 32 sp + 4 infra
Polycarpoideae 6 g/ 22 sp 4- 5 infra 7 g/ 28 sp + 4 infra
*projected tally — see text
At the subfamily level, most of the differences involve the Alsinoideae (Table 1). Changes within this
subfamily (stipules absent; sepals not awned, free or only basally connate; petals usually present; fruits
capsules) encompass most of the new generic alignments and new species, both native and introduced.
Robinson followed a broad concept for Arenaria and Stellaria. The narrower concepts we employ recognized
six genera (Arenaria, Eremogone, Honckenya, Lepyrodictlis, Minuartia, and Wilhelmsia) in the former genus and
three (Myosoton, Pseudostellaria, and Stellaria) in the latter.
By contrast, the Caryophylloideae (stipules absent; sepals not awned, fused basally into a cup or tube;
petals present, often conspicuous; fruits capsules) treatment deviated the least. The major differences are
Robinson's segregation of Lychnis from Silene (Morton 2005b preferred a more inclusive circumscription)
and the addition of 10 new native species of Silene.
The differences in the Paronychioideae (stipules present; sepals often awned, distinct or scarsely fused;
petals absent; fruits utricles) treatments involve the addition of ten new taxa of Paronychia and the transfer
of Scleranthus to the Alsinoideae.
Within the Polycarpoideae (stipules present; sepals rarely awned, distinct or scarsely fused; petals
absent or often present; fruits capsules), major changes involve the addition of the genus Polycarpaea, two
new alien species of Spergula, and five additional species of Drymaria.
The net change in the number of taxa between the two treatments is shown in Table 2. As noted above,
the increase in the number of genera (26 to 37) mostly reflects differences in generic concepts. Change in
the number of species (185 to 286) suggests that the number of species has increased by 50 percent; this is
somewhat inflated since some of Robinson's infraspecific taxa are now treated as species (e.g., in Cerastium).
The very similar numbers of infraspecific taxa (53 vs. 54) is somewhat deceiving. Besides elevation of some
taxa to the species level, others are no longer recognized (e.g., the five varieties of Silene douglasii that Robin-
son recognized). On the other hand, 22 varieties and subspecies in the FNA oe 6 Den eto
since 1894. Given these caveats, the total taxa comparison (species + inf t
although increasing by about 40 percent since 1894, probably es the actual dae
CHANGES INVOLVING NATIVE TAXA
Since the beginning of North American botany, most floristic attention has been on collecting and identify-
ing our native flora. The distribution of new native species by genus is shown in Table 3. One native genus
(Geocarpon in 1914) and 49 native species have been described since 1894. Not only are 63 percent of these
species in the Alsinoideae, but 45 percent are within Arenaria s.l.
The majority of the new native species were first discovered in the western United States; not entirely
unexpected given that, prior to 1890, much of the West was poorly known botanically. In descending order,
California (12), the southwest United States (10), Texas (8), Alaska (4), and the Pacific Northwest (3) account
for 37 of the 49 species. The remaining 12 initially were discovered in northeastern North America (5),
southeastern United States (3), Russia (2) and the central United States (2).
CHANGES INVOLVING INTRODUCED TAXA
Efforts to document our alien (introduced or exotic) flora more precisely than *occurring as a weed here and
there" (Gypsophila muralis L., Gleason and Cronquist 1991) have increased in recent decades. Rabeler has
TABLE 2. TI ber of g species, and infraspecifi
percent increase, respectively.
Robinson Rabeler & Hartman 96 increase
Genera 26 37 42
Species 191 286 50
Infraspecific 53 54 1
Total taxa 244 340 39
Tage 3 Th L £ . . f... E B by DE ily ig
Alsinoideae (31): Arenaria (4), Cerastium (5), Eremogone (5), Geocarpon (1), Minuartia (11), Pseudostellaria (2), Stellaria (3)
Caryophylloideae (10): Silene (10)
Paronychioideae (5): Paronychia (5)
Polycarpoideae (3): Drymaria (2), Spergularia (1)
been involved in reporting new introductions of species and distribution records for Caryophyllaceae over
the past 29 years (Rabeler 1980). Likewise, botanists at the Rocky Mountain Herbarium (see Hartman and
Nelson 2008) have made a concerted effort to document invasives during more than 54 major inventories
throughout the Rocky Mountains (over 840,000 new numbered collections since 1978, both native and in-
troduced) These initiatives long preceded the recent piqued interest in exotics. Alien taxa are very important
in the Caryophyllaceae. Of the 37 genera in the 2005 treatment, all members of 15 of those genera (41%) are
introduced to North America; this includes six of the eight genera in Caryophylloideae.
The distribution of new introduced species by genus is shown in Table 4. Three genera (Polycarpaea,
Lepyrodiclis, Moenchia) did not appear in Robinson (1897) despite Moenchia erecta (L.) G. Gaertn., B. Mey. &
Scherb.having been collected near Baltimore and Philadelphia in the 1830s (Rabeler 1991). Thirty species,
one-half of them members of the Alsinoideae, were first reported since 1897; seven of these were documented
in North America subsequent to 1960.
Knowing when and where an introduction first arrived in North America is nearly impossible unless
it was introduced intentionally. For example, Petrorhagia prolifera (L.) PW. Ball and Heywood was brought
to North America around 1800 and appeared in John Bartram's garden in Philadelphia (Rabeler 1985). Of
course, the actual documentation of the plant may be long after its introduction.
Where documented, the first collections of *new" introductions were chiefly from coastal states, especially
port areas. They include the southeastern United States and Texas (8), California (6), the Pacific Northwest
(6), and northeastern North America (4). Only three species appear to have been gathered first in an inland
state or province (Moehringia triphylla (L.) Clairv. in Ohio, Cerastium diffusum Pers. in Missouri, and Silene
sibirica (L.) Pers. in Saskatchewan).
Some recently discovered aliens, especially those first collected in th theastern United States, have
become widespread in eastern North America (e.g., Stellaria pallida (Dumort.) Crépin, Cerastium pumilum
Curtis, and C. brachypetalum Pers. Of the aliens included in Robinson (1897), some have become widespread.
Gypsophila paniculata L. noted as *doubtful in Manitoba" now is a noxious weed there as well as in California
and Washington (Rice 1997 onwards). Holosteum umbellatum L., in Robinson's time known only around
Philadelphia, Pennsylvania, is now documented from 41 states or provinces (see esp. Shinners 1965).
The occurrence of two alien species has diminished sharply. Agrostemma githago L. and Vaccaria his-
panica (Mill.) Rauschert were formerly very common in wheat fields. As agricultural practices advanced,
seed purity increased and transport of propagules of these species declined. In both cases, the range maps
in the FNA treatments (Thieret 2005; Thieret and Rabeler 2005) are probably best described as historical
1288 J lof the Botanical R h Institute of Texas 2(2)
TABLE 4 i peci y ily and g
Alsinoideae (15): Cerastium (6), Lepyrodiclis (1), Moehringia (1), Moenchia (1), Sagina (2), Stellaria (4)
Caryophylloideae (14): Dianthus (2), Gypsophila (2), Petrorhagia (2), Saponaria (1), Silene (7)
Paronychioideae (4?): Paronychia (2), Herniaria (1), Scleranthus (1?
Polycarpoideae (6): Drymaria (1), Polycarpaea (1), Spergula (2), Spergularia(2)
1 E 1 : £4] t
maximal distributions. While very few recent collections
Agrostemma githago remains on the noxious weeds lists of 13 states or Doris (Rice 1997 onwards).
WHAT MIGHT THE FUTURE HOLD?
We believe that there are additional native Caryophyllaceae yet to be described in North America. Nineteen
of the 286 species in the FNA treatment were described since 1960. Two additional species have been de-
scribed, both from California, since the 2005 treatment; Silene salmonacea TW. Nelson, J.P. Nelson and S.A.
Erwin (Nelson et al. 2006) and Eremogone cliftonii Rabeler and R.L. Hartman (Rabeler & Hartman 2008).
Likewise, the number of exotics are bound to increase. With seven species in the 2005 treatment first
reported since 1960, it seems likely that as botanists take our dynamic alien flora more seriously, the tally
will rise.
In addition to new discoveries and distributional records, additional study of the relationships of the
taxa likely will change the treatment of the Caryophyllaceae. Many such questions are discussed in the
FNA treatment (Rabeler & Hartman 2005) and could form the basis of interesting studies. As examples, do
Stellaria dicranoides and Stellaria fontinalis really belong in other genera (Morton 2005a) and should Silene
and/or Minuartia be treated as multiple genera as discussed by Morton (2005b) and Rabeler et al. (2005)
respectively.
ACKNOWLEDGMENTS
We thank the authors of the FNA treatments who helped make this comparison possible and Arnold Tiehm
and John K. Morton for their constructive comments on an earlier draft of this paper.
REFERENCES
BENTHAM, G. AND J.D. Hooker 1862-1883. G plant d | vata.
difinita 3 vols., London, A. Black et al.
BrrricH, V. 1993. Caryophyllaceae. In: K. Kubitzki J. G. Rohwer, and V. Bittrich., eds. The Families and Genera of Vas-
cular Plants. Flowering Plants, Dicotyledons, Magnoliid, Hamameliid and Caryophyllid Families. 2:206-236.
Fior, S., PO. Karis, G. CASAZZA, L. MINUTO, AND F. SALA. 2006. Molecular phylogeny of the Caryophyllaceae (Caryophyl-
lales) inferred from chloroplast matK and nuclear rDNA ITS sequences. Amer. J. Bot. 93.399-41 1.
Gueason, H.A. AND A. CRONQUIST. 1991. Manual of Vascular Plants of Northeastern United States and adjacent Canada,
2™ ed. New York Botanical Garden, New York.
HARTMAN, R.L. AND B.E. EN di General information for floristic proposals. [The Boiler Plate]. http:/Awww.rmh.
Uw lu/ I f forFloristicsP roposals. .pdf (22 March 2008 )
Morton, J.K. 2005a. 22 Stellaria In: Flora of North America Editorial Committee, eds. Flora of North America North
of Mexico. Oxford University Press, New York and Oxford. 5:96-114.
Morton, J.K. 2005b. 36. Silene. In: Flora of North America Editorial Committee, eds. Flora of North America North
of Mexico. Oxford University Press, New York and Oxford. 5:166-214.
NELSON, T.W, J.P. NELSON, AND S.A. Erwin. 2006. A new species of Silene in the Sil
from the Klamath Mountains of Shasta-Trinity National Forest, Trinity County, California. Madroño 53:72-76.
NePoknoerr, M., W.L. Wacner, R.K. RABELER, E.A. Zimmer, S.G. WELLER, AND A.K. Sakai. 2002. Relationships within Caryo-
L Í
omplex (Caryophyllaceae)
Dahal 3 Ll pt , Caryopl I in Narth A 1] 1289
phyllaceae inferred from moiecular sequence data. Botany 2002 Program Abstracts: 105. Available at: http://
bsa2002.scientific-conference.net/sympos12/abstracts/6.shtml (25 March 2008).
RABELER, R.K. 1980. Petrorhagia prolifera, a naturalized species in Michigan. Michigan Bot. 19:83-88.
RABELER, R.K, 1985. Petrorhagia (Caryophyllaceae) in North America. Sida 11:6-44,
RaseLER, R.K. 1991. Moenchia erecta (Caryophyllaceae) in eastern North America. Castanea 56:150-151.
RABELER, R.K. AND R.L. HARTMAN (eds.). 2005. 43. Caryophyllaceae. In: Flora of North America Editorial Committee, eds
Flora of North America North of Mexico. Oxford University Press, New York and Oxford. 5:3-215.
RABELER, R.K, R.L.HARTMAN, AND F.H. UTECH. 2005. 24. Minuartia. In: Flora of North America Editorial Committee, eds.
Flora of North America North of Mexico. Oxford University Press, New York and Oxford. 5:116-136.
RABELER, R.K, AND R.L. HARTMAN. 2008 ["2007"]. Eremogone cliftonii (Caryophyllaceae), a new species from California.
Madrofio 54:329-333.
Rice, PM. 1997 onwards. INVADERS Database System. University of Montana, Missoula, MT. http://invader.dbs.
umt.ecu (22 March 2008).
Rosinson, B.L. 1893. The North American Sileneae and Polycarpeae. Proc. Amer. Acad. Arts 28:124-155. [also as
Contr. Gray Herbarium 5].
Robinson, 8.L. 1894. The North American Alsineae. Proc. Amer. Acad. Arts 29:273-313. [also as Contr. Gray Her-
barium €].
Rosson, B.L. 1897. Order XVIII. Caryophyllaceae. In: B.L. Robinson, ed. Synoptical Flora of North America. Vol. |
(1, fasc. 11):208-255.
SHINNERS, L.H. 1965. Holosteum umbellatum (Caryophyllaceae) in the United States: Population explosion and
fractionated suicide. Sida 2:119-128.
Tuer, JW, 2005. 37. Agrostemma. In: Flora of North America Editorial Committee, eds. Flora of North America
North of Mexico. Oxford University Press, New York and Oxford. 5:214-215.
THIERET, J.W. AND R.K. RAseLER, 2005. 31. Vaccaria. In: Flora of North America Editorial Committee, eds. Flora of North
America North of Mexico. Oxford University Press, New York and Oxford. 5:156
1290 J | of the Botanical R h Institute of Texas 2(2)
BOOK REVIEW
Jos C. Truett. 1996. Circling Back: Chronicle of a Texas River Valley. (ISBN 0877455317, pbk.). University
of Iowa Press, 119 West Park Road, 100 Kuhl House, Iowa City, Iowa 52242-1000, U.S.A. (Orders:
www.uiowapress.org, 800-621-2736, 800-621-8476 fax). $19.95, 230 pp., photographs, 6" x 9".
Back home for the funeral of his grandfather in 1967, Joe C. Truett thought long and hard about the world of the Angelina River valley
in East Texas that Corbett Graham bad known and he himself had glimpsed. His vision resuited in this portrait of a place over time,
its layers revealed by his love and care and curiosity. He celebrates his family's heritage and the unspoiled natural world of the Piney
Woods without nostalgia.
O 4 ja] of I Lol
1 Laa A Y PEA nth 1 f 1 Ea: A Je ee J aes poet law A ff, E
tnat ne had E g ping
activities on fish, wildlife, and Baba Disillusionment built over ti
preventing environmental degradation. He began to look for other channels that would be more effective. Land of Bears and Honey was
one result of that search and Circling Back is a continuation of that search.
Part = ii n nu anc RUE Series, a nn is Ms HIDE out, lucid, and should be included in any public or college
library. Iti y Gary L. Jennings, Botanical Research Institute of Texas, 500 East
4th Street, Fort Worth, ma 76102-4025, USA.
J. Bot. Res. Inst. Texas 2(2): 1290. 2008
THE BIG THICKET: PRISTINE WILDERNESS
OR ARCHEOLOGICALLY UNDERSTUDIED?
Michael! H. MacRoberts and Barbara R. MacRoberts
Bog Research, 740 Columbia, Shreveport, Louisiana 71104, U.S.A. and
Herbarium, Museum of Life Sciences
Louisiana State University in Shreveport
Shreveport, Louisiana 71115, U.S.A.
ABSTRACT
Received opinion is that the Big Thicket of east Texas inhabited until tl ly ni ] tury. It was a pristine wilderness.
Thi : ; El 41 T 1 1 ¿ol A ns an] 1
A11153 VIC YW PE Vy
RESUMEN
La opinión aceptada es que el Big Thicket del este de Texas estuvo deshabitado hasta principios del siglo diez y nueve. Era una tierra
virgen pristina. Este punto de vist tá apoyado ni en d jueolégi i históricos
Cozine (2004) and Callicott et al. (2006) maintain that the Big Thicket of east Texas was a wilderness prior
to 1800. No one ever lived there. However, they do not cite any archeological sources in support of this
position. We therefore undertook a study to determine the validity of their position because we previously
had discovered that many pronouncements about the Big Thicket are incorrect (MacRoberts & MacRoberts
2004, 2007; MacRoberts et al. 2007) and because in the “great wild debate" many authorities maintain
that the North American continent was not a pristine wilderness in 1492 but instead was a landscape that
had been highly modified by Native Americans (Callicott & Nelson 1998; Denevan 1998; McCann 19994,
1999b; Bonnickson 2000; Vale 2000; Williams 2000; Pyne 2001; Kay 2002; Mann 2005). However, some
want it both ways. While Callicott et al. (2006:28) agree that colonists and explorers did not find a *howl-
ing wilderness" but a landscape “as much a cultural artifact as the one they left behind,” they also state that
"of course, throughout North America tracts of wilderness were ... interspersed with more or less intensely
peopled places." According to them, one such wilderness was the Big Thicket (Callicott et al. 2006: 29). Na-
tive Americans made only occasional hunting forays into it and the Spanish found it inhospitable.
A survey of the archeological literature on Native American presence in the Big Thicket, including the
Big Thicket National Preserve, shows that archeological work is virtually non-existent (Shafer 1968; Shafer
et al. 1975; McClurkan 1968; Story 1990b; Bousmann et al. 2004; Ricklis 2004). Compared with other
regions of Texas, the archeological coverage of the Big Thicket is minuscule (Perttula 2004). Ricklis (2004)
makes this clear in his recent review of southeastern Texas archeology in which he barely mentions any-
thing except coastal areas. We, therefore, contacted Texas archeologists to confirm our findings and obtain
current information on archeology in the Big Thicket. Victor Galan (pers. comm.) wrote that of the surveys
that have been conducted in the Big Thicket region, few cover enough area with adequate testing methods,
still fewer have good stratigraphic information, and even fewer yet have been thoroughly excavated. One of
the findings of early archeological surveys in the Big Thicket region was that there appeared to be relatively
few sites (Shafer et al. 1975). Whether this paucity of sites was due to masking by dense ground cover, or
was an accurate reflection of limited human use was not clear (Shafer et al. 1975; Story 19902). Years later,
Shafer (pers. comm. March 13, 2008) states:
T 1»?
“Given what I know now ...I am convinced that issed a lot of sites in the Big Thicket. A bi Ed visibility, and the
[2] otf y
th did tü } 1 i 1 Also T a A fih : well
Shafer (pers. comm., March 17, 2008) is currently of the opinion that the Big Thicket is archeologically
understudied. There is, however, enough archeological evidence to indicate that man has been in the Big
J. Bot. Res. Inst. Texas 2(2): 1291 — 1293. 2008
1292 ical R h Institute of Texas 2(2)
Thicket since Paleoindian times (at least since 8,000 years ago) (Bousmann et al. 2004; Victor Galan pers.
comm.).
The Spanish left a voluminous record of their presence in eastern Texas (Bolton 1914, 1915; Swanton
1942, 1946; Freeman 1990). The Spanish encountered Hasinai and Caddo tribes in the region of Nacogdo-
ches and Natchitoches, and Attacapan tribes between Nacogdoches and the coast, mainly in the San Jacinto
and Trinity river drainages (Bolton 1914, 1915; Swanton 1942, 1946; Perttula 1992). However, the Spanish
had no interest in the eastern part of the Big Thicket (Newton, Jasper, Tyler and Hardin counties) because,
rather than colonization, their aim was to prevent French encroachment by establishing forts and missions
in areas where the French threatened. The French did not threaten the Big Thicket region, but threats existed
north of the Big Thicket at Natchitoches and on the coast on the lower Trinity River. The Spanish repeatedly
traveled across the western part of the Big Thicket (Polk, Liberty, San Jacinto counties) on their overland
trips from Nacogdoc! their presidio and mission on the lower Trinity River (the Spanish appear to have
never used waterways for travel). Consequently, the Spanish did not avoid the Big Thicket any more than
they avoided other regions of Texas; they simply had no interest in the area.
In summary, the paucity of archeological research prevents any detailed understanding of Native
American occupation de anión impact on the Big Thicket. Until the Big Thicket is seriously explored archeo-
logically, our und g of Native American impact there will remain minimal. This, in turn, means that
an understanding of Big Thicket “biocomplexity” (interrelation of “coupled human and natural systems”)
(Callicott et al. 2006; Liu et al. 2007) will ultimately have to include the archeologist.
ACKNOWLEDGMENTS
Victor Galan, Deep East Texas Archeological Consultants, and Harry Shafer provided information on
Big Thicket archeology, directed us to some of the Big Thicket archeological literature, and reviewed the
paper.
REFERENCES
Bouton, H.E. 1914. Athanase de Mezieres and the Louisiana-Texas frontier 1768-1780. Arthur H. Clark Co.,
Cleveland.
Botton, H.E. 1915. Texas in the Middle Eighteenth Century. University of Texas Press, Austin.
BoNNICKSEN, T.M. 2000. America's ancient forests: from ice age to the age of discovery. J. Wiley, New York.
BOUSMANN, C.B., B.W. Baxer, AND A.C. Kerr. 2004. Paleoindian archeology in Texas. In: T.K. Perttula, ed. The prehistory
of Texas. Texas A & M Press, College Station. Pp. 15-97
CaLucorr, J.B. AnD M.P. NeLson (eds.). 1998. The great new wilderness debate. Univ. Georgia Press, Athens.
CaLuicoTT, J.B., M. Aceveno, P. Gunter, P. HARCOMBE, C. LINDQUIST, AND M. Monticino. 2006. Bi plexity in the Big Thicket.
Ethics, Place €: Environm. 9:21-45,
Cozine, J.J. 2004. Saving the Big Thicket: from exploration to preservation, 1685 — 2003. Univ. North Texas Press,
Denton.
Denevan, W.M. 1998. The pristine myth: the landscape of the Americas in 1492. In: Callicott, J.B. and M.P. Nelson,
eds. The great new wilderness debate. Univ. Georgia Press, Athens. Pp. 414—442.
FREEMAN, M.D. 1990. Cultural history of Europeans, Africans, and their descendents. In: D.A. Story, J.A. Guy, B.A.
Burnett, M.D. Freeman, J.C. Rose, D.G. Steele, BW. Olive, and KJ. Rinehard, eds. The archeology and bioarche-
ology of the Gulf Coastal Plain. Vol.2. Research Series, No. 38. Fayetteville, Arkansas Archeological Survey. Pp.
367-384
Kay, C.E. 2002. Are ecosystems structured from the top-down or bottom-up? A new look at the old debate. In:
C.E. Kay and R.T. Simmons, eds. Wilderness and political ecology: aboriginal influences and the original state
of nature. Univ. Utah Press, Salt Lake City. Pp. 215-237.
Liu, J. (AND 15 others). 2007. Complexity of coupled human and natural systems. Science 317:1513-1516.
MacRoserTs, M.H. AND B.R. MacRoserts. 2004. The Big Thicket: typical or atypical? East Texas Hist. J. 42:42-51.
f the Big Thicket 1293
a Li
MacRoserTs, M.H. AND B.R. MacRoserTs. 2007. Phytogeography of the Big Thicket, east Texas. J. Bot. Res. Inst. Texas
1:1149-1155.
MacRoserts, M.H., B.R. MAcRoBERTS, AND R.G. Kauwskv. 2007. Vascular plant species/area relationships (species rich-
ness) in the West Gulf Coastal Plain: a first approximation. J. Bot. Res. Inst. Texas 1:57/-583.
MANN, C.C. 2005. 1491: new revelations of the Americas before Columbus. Knopf, New York.
McCann, J.M. 1999a.. The making of the pre-Columbian landscape. Part |. The environment. Ecol. Restorat. N.
Amer. 17:15-30.
McCann, J.M. 1999b. The making of the pre-Columbian landscape. Part Il. The vegetation, and implications for
restoration for 2000 and beyond. Ecol. Restorat. N. Amer. 17:107-119.
McCLurkan, B.B. 1968. Livingston Reservoir, 1965-1966: Late archaic and neo-American occupation. Pap. Texas
Archeol. Salvage Project 12:1-115.
PeRTTULA, T.K. 1992. The Caddo Nation. Univ. Texas Press, Austin.
PerTTULA, T.K. 2004. (ed.). The prehistory of Texas. Texas A & M Press, College Station.
Pyne, S.J. 2001. Fire, a brief history. Univ. Washington Press, Seattle.
Rickus, RA. 2004. The archeology of the Native American occupation of southeast Texas. In: T.K, Perttula, ed. The
prehistory of Texas. Texas A 8: M Press, College Station. Pp. 181-202.
SHarer, H.J. 1968. Archeological investigations in the San Jacinto River basin, Montgomery County, Texas. Pap.
Texas Archeol. Salvage Project 13:1-82.
SHAFER, H.J., E.P. BAXTER, T.B. STEARNS, AND J.P. Derinc. 1975. An archeological assessment of the Big Thicket National
Preserve. Report 19: Anthropology Laboratory, Texas A € M University.
Story, D.A. 1990a. Environmental setting. In: D.A. Story, J.A. Guy, B.A. Burnett, M.D. Freeman, J.C. Rose, D.G. Steele,
B.W. Olive, and KJ. Rinehard, eds. The archeology and bioarcheology of the Gulf Coastal Plain. Vol.1, Research
Series, No. 38. Fayetteville, Arkansas Archeological Survey. Pp. 5-26.
Story, D.A. 1990b. Cultural history of the Native Americans. In: D.A. Story, J.A. Guy, B.A. Burnett, M.D. Freeman,
J.C. Rose, D.G. Steele, B.W. Olive, and KJ. Rinehard, eds. The archeology and bioarcheology of the Gulf Coastal
Plain. Vol.1, Research Series, No. 38. Fayetteville, Arkansas Archeological Survey. Pp. 163-366.
SWANTON, J.R. 1942. Source material on the history and ethnology of the Caddo Indians. Smithsonian Institution,
Bureau Amer. Ethnology Bull. 132. Washington, D.C.
SWANTON, J.R. 1946. The Indians of the southeast United States. Smithsonian Institution, Bureau Amer. Ethnology
Bull. 137. Washington, D.C.
Vale, T.R. 2000. The pre-Columbian landscape of the United States: pristine or humanized? In: TR. Vale, ed. Fire,
native peoples, and the natural landscape. Island Press, Washington. Pp. 1-39.
WiLLiams, J.M. 2000. Book review: Bonnicksen, T.M. 2000. America's ancient forests: from ice age to the age of
discovery. Conservation Ecol. 4:2.
1294 J l of the Botanical R h Institute of Texas 2(2)
BOOK REVIEW
Jim WARK AND JOSEPH A. MUSSULMAN. 2004. Discovering Lewis and Clark from the Air. (ISBN: 978-0878424894,
pbk.). Mountain Press Publishing Company, PO. Box 2399, Missoula, Montana 59806, U.S.A. (Orders:
www.mountain-press.com, 800-234-5308). $40.00, 272 pp., color illustrations, 9 1/2" x 9 1/2".
A LEE | 1 1 T; XX 1 11 L [LT LAA. A | 3 ff. Lal É d Clark Expedition
F CE dorus P »
TI is foll 1 from the air, uo rend by step-by-step di i f thei te choi d explorati
The idea of the book was simple enough. The authors fea a small plane over th te of the C f Di and took aerial
Y é
hotos of places along the trail. The principal impression one gets from these photos is how completel land f 200 years ago
Pp 8 P 8 P pletely P
has vanished—dammed up, paved over and plowed under. The rivers of Lewis and Clark's time (and it was mostly a river oe are
drowned under the slack water of d of dams, 2M Vae they are not MA they are often channelized, strai: s and oth-
ya ae
erwise "improved." S prise tl I g: Beaverhead R Tobacco Root
Mountains, and Bi M ins. Each photo is illumi 1 by lysis by tl hors, accompanied B A quotations from
the Lewis and Clark journals.
py g, as in asa n id and index. It would be a good addition for public
| 11 1;1 3 1 1 A 1 1 r
Street, Fort Worth, Texas 76102-4025, U.S.A.
L. Jennings, Botanical Research Institute of Texas, 500 East 4th
is
J. Bot. Res. Inst. Texas 2(2): 1294. 2008
DIFERENCIAS EN EL USO DE PLANTAS ENTRE HOMBRES Y MUJERES
EN UNA COMUNIDAD DE PIE DE MONTE DEL NORTE DEL ECUADOR
David Suárez Duque
Corporación Grupo Randi Randi
uito, ECU,
davsua2@yahoo.com
RESUMEN
En est idad col gi 298 especies de plantas, de estas el 31% f ñaladas por hombres, el 40% por mujeres y
1 2 oa Be J y T = T LE A 1 1 1 1 E E £1 D ]
su trabajo y "M Alincluir una variab! ial ] género, en i igaci de etnobotáni puede mej l entendimient
de los ecosistemas, su biodiversidad y la gente que los usa.
ABSTRACT
y 298 species of pl identified by local inf Of these, 31% are used by men, 40% by women and the rest
by both. Men and 's] ledge and f pl infl 1 by work and gender roles. I believe that the inclusion of social
variables h a gender i hno! ical h, imp 1 1 ling al Ah find; ity.
INTRODUCIÓN
La Etnobotánica es un campo interdisciplinario, considerado como una herramienta indispensable para
establecer estrategias de conservación y manejo (Cunningham 2001). Permite de manera rápida, conocer
las especies más utilizadas, formas de usos y otros datos relacionados con la interacción de las personas
con las plantas (Plotkin 1995). En la mayoría de trabajos etnobotánicos, los botánicos y botánicas han
dado gran importancia a la flora y han dejado de lado las características socioeconómicas y culturales de
sus informantes (Arango 2004). Entre los aspectos sociales que se ha dejado de analizar se encuentran las
diferencias de género en el conocimiento, uso y control de las plantas.
El género según Rodríguez et. al. (2004) se refiere a las relaciones construidas socialmente entre mu-
jeres y hombres, culturalmente e institucionalmente incorporadas, que varían por sitio, situación contexto
y tiempo. E] incluir esta variable social en investigaciones sobre etnobotánica, puede permitir visualizar la
diferencia que existe en el conocimiento y usos que hombres y mujeres dan a las plantas.
El conocimiento diferente que hombres y mujeres pueden tener sobre el uso de las plantas, no implica,
segün Joekes (2004), una línea de género un se, si no más bien conocimiento e por las oe de
uso, roles y labores que hombres y mujeres desempefian en su comuni
basadas en la division social del trabajo, deis que hombres y mujeres podrían tener conocimientos
especiales de la flora de entorno (Agarwal 1994).
Alincorporar la variable de género, en la presente i
de pie de monte en el norte del Ecuador, se buscaba mostrar re diferencias que hay en el uso y conocimiento
de plantas, entre hombres y mujeres.
t } tani n uma camiunidad calona
METODOLOGIA
La comunidad de San Jacinto de Chinambi, donde se realizó la investigación, está ubicada en el norte
del Ecuador, en la provincia del Carchi, entre el territorio del pueblo indígena Awá y el bosque protector
Golondrinas, en el pie de monte de la cordillera occidental de los Andes. En este lugar viven 30 familias de
descendencia: Awá, afroecuatoriana proveniente del Valle del Chota, y mestiza procedente de varios lugares
J. Bot. Res. Inst. Texas 2(2): 1295 — 1308. 2008
1296 Journal of the Botanical R h Institute of Texas 2(2)
de la Sierra del Ecuador y del sur occidente de Colombia. Los primeros asentamientos colonos se dieron
hace casi cincuenta años (Comunidad de San Jacinto 2005).
El territorio de la comunidad de San Jacinto está atravesado por el río Chinambí, formado por el aporte
de varias quebradas, que nacen en la zona alta. A su alrededor se encuentran zonas planas que a medida
que se alejan del río, forman pendientes más pronunciadas. La parte sur del territorio ha sido utilizada para
la producción agropecuaria, mientras que la norte, donde se encuentran las pendientes más acentuadas,
esta representada por remanentes de bosques. La cabecera norte del territorio tiene una cuchilla desde
donde se extiende el territorio comunal (Comunidad de San Jacinto 2005). La altitud de los territorios en
la comunidad de San Jancito de Chinambí va desde los 960 msnm hasta los 2400 msnm, y pertenece a las
formaciones vegetales de bosque siempre verde pie montano (Cerón et. al. 1999), bosque siempre verde montano
bajo y de neblina montano (Valencia et. al. 1999).
Basado en el hecho de que la comunidad se originó por asentamientos de personas colonas, se decidió
tomar como unidad de análisis e interacción la finca!, que es un espacio social y económicamente construido,
donde se puede encontrar un mosaico de ecosistemas. La investigación fue realizada en nueve fincas de
diferentes familias, ubicadas al azar dentro del territorio de la comunidad. Con cada familia se trabajó dos
días completos, un día con el hombre y al otro con la mujer. Para lograr el apoyo durante todo el día y no
interferir con sus tareas diarias, a cada participante se le pagó el diario o jornal (salario por un día de tra-
bajo en la comunidad). En el caso de las mujeres, para que ellas pudieran participar y no se preocupen de
incumplir con sus labores diarias en sus hogares, se pagó a otra persona, para que ese día cuide a las hijas
e hijos pequeños y prepare la comida para toda la familia. Este pequeño detalle ratificó que las mujeres
en la comunidad de San Jacinto, como las de otras partes del Ecuador, en un mismo día cumplen doble
trabajo (reproductivo y productivo); algunas también realizan actividades comunitarias y religiosas como:
organización de festejos comunales, reuniones escolares y del cabildo entre otras. Todas estas actividades
hicieron que el tiempo que las mujeres pudieran aportar a la presente investigación fuese menor.
Los recorridos fueron planificados por las y los encuestados en la noche del día anterior y siempre
se trató de no influenciar en la decisión de las personas sobre el sector a visitar al día siguiente. Con los
recorridos se recolectó plantas en los huertos, remanentes de bosque, jardines e incluso en las macetas que
cuidan en sus casas. Con las personas encuestadas se anduvo por los lugares donde recolectan o cultivan
sus plantas.
Para registrar los usos de las plantas se utilizó una encuesta en la que se anotó algunas características
como: lugar donde se encontraba, parte de la planta que se usa, si la especie era cultivada o silvestre. Además
de estos datos, se apuntó el hábito, la presencia de látex, resina, flores o frutos. A cada planta mencionada
por las personas de la comunidad se le asignaba un número de encuesta a partir del 001. De cada especie se
colectó entre dos y cuatro especímenes que fueron prensados en pro MNA y preservados en cone 2
7096 (Cerón 2003). Cada muestra colectada se registró en el catá ampo desde el dec
DS 1593 al 1925, estas fueron identificadas por comparación con b a que descansa en el Herbario
Nacional del Ecuador (QCNE). Los especímenes de esta investigación reposarán en el QCNE y CHEP.
—
RESULTADOS Y DISCUSIÓN
En la comunidad de San Jacinto se registraron 298 especies de plantas usadas, de éstas el 3196 fueron
señaladas por hombres y el 4096 por mujeres, las restantes especies utilizaban ambos. Las familias más
importantes por número de especies usadas por las mujeres son: Solanaceae, Araceae, Lamiaceae, Poaceae,
Asteraceae, Fabaceae y Urticaceae, mientras que los hombres utilizan las familias Moraceae, Myrtaceae,
Lauraceae, Meliaceae, Rubiaceae, Solanaceae y Actinidaceae. Hombres y mujeres coincidieron en el uso de
las siguientes familias en orden de importancia: Poaceae, Asteracea, Rutaceae, Meliaceae, Euphorbiacae,
Clusiaceae y Lamiacae. El listado de las especies esta en el apéndice 1.
£&
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Suárez, Dif i H de plant tre hom i 1297
En las 18 personas de la comunidad de San Jacinto de Chinambi encuestadas se registró un promedio
de 46,6 plantas usadas por persona (Fig. 1). A pesar que las mujeres de la comunidad conocen en promedio
más usos de las plantas (54,7 plantas), que las que conocen los hombres (38,5 plantas), la diferencia no
es estadísticamente significativa (Anova F- 2,58; p-0,127). Una cosa interesante a observar en la figura 1,
es que a pesar de que los hombres conocen menos plantas que las mujeres, el conocimiento entre ellos es
más homogéneo. Entre las mujeres encuestadas hay algunas que en su casa solo hacen trabajos domésticos,
mientras que otras por ser madres solteras tiene que administrar la finca, y cuidar a sus hijos-as. Esta difer-
encia de trabajo hace que en ellas el conocimiento también sea variable (Fig.1). Mientras que la mayoría de
hombres se dedican sólo a trabajos agropecuarios y de extracción de madera por eso su conocimiento puede
ser más homogéneo.
La mayoría de plantas que utilizan los hombres son silvestres, mientras que las mujeres utilizan más
plantas cultivadas (Fig. 2). El hombre utiliza más plantas que esta extrayendo de ecosistemas naturales,
mientras que las mujeres están utilizando más especies de ecosistemas antropogénicos (agroecosistemas).
Por ende la mayor parte de las plantas que las mujeres utilizan están en el huerto (zona de cultivos con
parches de bosque) dentro de la finca. Mientras que los hombres utilizan plantas del bosque (Fig. 3). Esta
diferencia estadísticamente significativa del lugar en donde extraen las plantas (Anova F-10,172; p«0,05),
está ligada también al trabajo que cada persona realiza. Incluso hay mujeres que nunca visitan la zona de
bosque comunitario, por varias razones, entre las que ellas mencionaron estaban: la lejanía, falta de interés o
el excesivo trabajo en su casa. Mientras que los hombres tienen mucho interés en la zona del bosque, porque
aún es posible encontrar allí, especies de madera fina o suelo apto para la agricultura.
El huerto es la parte de la finca donde se mezclan fragmentos de bosques en regeneración, zonas de
pasto, cultivos y la casa de la familia, es un mosaico de hábitats donde las personas colonas está manejando
sus recursos. Este lugar según Álvarez (2005) y Rocheleau et. al. (2004), es un ámbito fundamentalmente
femenino y un espacio de replicación de la cultura, pero no siempre las mujeres pueden tomar decisiones
sobre él. Además las mujeres mantienen en la casa, ya sea en maceteros o en el jardín (a un lado de la casa)
varias especies de plantas, traídas de otros sectores fuera de San Jacinto o del bosque local, esto lo hacen
para: alegrar la casa o tenerlas más cerca. Mientras que los hombres no acostumbran a mantener plantas en
este espacio (Fig. 2). Esta necesidad de las mujeres de mejorar el entorno de la casa, también fue mencionada
por mujeres colonas brasileñas en la investigación Kainer & Duryea (1992) y por mujeres indígenas Zenúes
en Colombia (Álvarez 2005).
Las mujeres usan más hierbas (Fig. 4) que los hombres, mientras que ellos usan más árboles que ellas.
Esta diferencia significativa de los hábitos de las plantas usadas (Anova F-9,088; p«0,05), muestra que
hombres y mujeres ven a los ecosistemas de manera diferente. Por ejemplo mientras las mujeres están
usando más el sotobosque (arbustos, hierbas), los hombres conocen y usan más especies del dosel (árboles).
Aunque esta apreciación no es absoluta, concuerda con la encontrada por Phillips & Gentry (1993) en Perú,
que los hombres conocen más especies de bosque (árboles), mientras que las mujeres conocen más hierbas
(plantas no boscosas).
A] preguntarles a los hombres y las mujeres, que parte de la planta utilizan, respondieron en la mayoría
de casos que todo el individuo. En el caso de los hombres al utilizar la madera del árbol, tienen que extraer
a todo el individuo, a pesar de que sólo utilizan la parte comercial del tronco y el resto la dejan para que
se descomponga en el bosque, o dependiendo de la madera la utilizan para leña. Mientras que las mujeres,
como la mayor parte de plantas que utilizan son hierbas, ellas también extraen todo el individuo, es obvio
que arrancar una hierba, es menos destructivo que talar un árbol. Dependiendo de la intensidad de uso, el
utilizar toda la planta podría ser considerado como un uso destructivo, y extraer solo una parte como uso
no destructivo, o sostenido (Caniago & Siebert 1998; Correte et. al. 2005). A pesar de que en la presente
investigación no se consideró esta idea al momento de diseñarla, es conveniente mencionar que Caniango
& Siebert (1998), manifiesta que si el uso de toda la planta de una especie es intensivo, puede ocasionar la pérdida
de la especie.
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En relación a los usos de las plantas, bci existe una diferencia por género (Anova F=6,433; p«0,005),
los hombres j un mayor nú t uso maderable, mientras que las mujeres mencio-
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naron más especies con uso alimenticio (Fig. 5). Estos concuerdan con los encontrados por Arango
(2004) en Colombia y Figueiredo et. al. (1993) en Brasil. Estos autores-as sugieren que esta diferencia en el
conocimiento de plantas está ligada a las actividades que hombres y mujeres tienen en su comunidad.
Es notable la diferencia que existe entre hombres y mujeres en el conocimiento y uso de plantas or-
namentales, para ellas el tener plantas “bonitas” cerca de la casa es importante, porque hace a la casa más
acogedora. Mientras que para los varones no son importantes las especies ornamentales, porque no tiene
valor comercial aum y ellos no están peus en el arreglo del hogar. Esta respuesta de las mujeres de San
Jacinto sobre las plant uerda con lo manifestado por mujeres de Cachoeira en Brasil, en
la investigación d: Kainer & Duryea (1992). Algunos varones que mencionaron especies silvestres con uso
ornamental, estaban pensando en manejar estas plantas, pero hasta el momento no habían hecho intentos
de plantarlas fuera del bosque.
En cuanto a las plantas con uso medicinal las mujeres conocen más especies que los hombres, esta
diferencia también la mencionan Caniago & Siebert (1998) en un trabajo realizado en Indonesia. Este con-
traste según Rocheleu et. al. (2004) se debe a que la responsabilidad del cuidado de la salud de las familias
rurales, está siempre a cargo de la mujer. Algunas de las plantas medicinales que ellas mencionaron dentro
de la encuesta, no son nativas de San Jacinto, más bien fueron traídas por las personas cuando colonizaron
la zona (Ocimum campechianum Mill., Salvia alvajaca Oerst., Ruta graveolens L., Verbena litoralis Kunth, entre
otras). Esto concuerda con lo realizado por los habitantes de una comunidad rural de Conceicáo-Acu que al
colonizar llevaron consigo las especies medicinales HR ues más utilizadas (Correte et. al. 2005).
Cuando se preguntó sobre la dosis o posología de alg medicinales, algunas de las entrevis-
tadas, no sabían, porque no había lado la especie, pero conocían por tradición su uso. Al preguntarles,
porque no las usaban algunas de ellas decían: que era más fácil y seguro salir al centro de salud, cuando alguien
estaba enfermo-a, mientras que otras decían que usaban las penas hasta e salir a ver al médico, o cuando las
medicinas no funcionaban bien. Casi todas las plantas medi las por los hombres se encuentran
en el bosque, incluso muchas están ligadas a cicatrización o el alivio de las quemaduras producidas por la
motosierra. Un ejemplo es la resina de Cyathea pallescens (Sodiro) Domin, que aparece cuando se corta a esta
planta para facilitar las labores de extracción de madera, como esta planta produce una resina gelatinosa,
en el caso de sufrir una quemadura con la motosierra, ellos se colocan sobre la parte quemada esta resina,
para aliviar el ardor y seguir trabajando.
Al categorizar las especies encontradas en San Jacinto, según los criterios de origen: endémica, nativa,
cultivado o introducida al país, propuestos por Jørgensen & León-Yánez (1999) y Valencia et. al. (2000)
se encontró que la mayoría de plantas que utilizan las personas son especies nativas del Ecuador (Fig. 6),
a pesar que también usan especies introducidas. Las mujeres son las que más especies introducidas usan,
muchas de las cuales son ornamentales. Un problema con estas plantas ornamentales e introducidas, es que
con el pasar de los años, algunas de estas especies se pueden convertir en plagas dificiles de erradicar. Hasta
el momento este dw no es preocupante en la zona de estudio, pero existen muchos casos en Ecuador de
especies han rtido en plagas (como en las Islas Galápagos y otras áreas naturales),
pero no se logró ae en la literatura si estas plantas fueron introducidas por hombres o mujeres. Lan-
tana camara L. es un ejemplo interesante de diseminación de especies ligadas al género, ya que esta especie
fue llevada por la esposa del gobernador de Ceylon, por los colores de las flores, desde Brasil hasta su jardin
en Indo-China. De su jardin esta planta se diseminó por toda la India e Indo-China ocasionado algunos
problemas a la flora nativa (Morton 1994). En San Jacinto las mujeres no usan muchas plantas nativas del
bosque, y no se registró ni una sola especie endémica para el Ecuador, que es usada exclusivamente por
estas mujeres. En cambio los hombres de esta comunidad usan más plantas nativas y endémicas, porque
aün siguen extrayendo recursos del bosque nativo. Las especies endémicas para el Ecuador que utilizan las
personas de San Jacinto están mencionadas en el cuadro 1.
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Las personas colonas a pesar de que han sido tachados siempre de destructoras de los ecosistemas
naturales, o como personas con pocas tradiciones, tienen un importante conocimiento sobre las plantas que
es necesario rescatar. Además este saber se va acrecentando, según van pasando los años de asentamiento
de la comunidad, ya que la gente cada nuevo dia trata de entender mejor el nuevo ecosistema donde vive,
para sacarle más provecho. Segün Joekes (2004) las mujeres y hombres aprenden lo que necesitan saber, lo cual
depende de su relación práctica con el recurso mismo en términos de la división genérica del trabajo que se da en la
localidad. El entender como usan estas personas las plantas, puede dar pautas para proponer estrategias más
prácticas para la conservación de la biodiversidad, especialmente en lo referente a domesticación y manejo
de algunas especies.
Las personas de la comunidad de San Jacinto conocen diversas especies de plantas que las encuentran
en diferentes lugares en su comunidad. El conocer cada uno de estos ecosistemas y su biodiversidad depende
de los roles y el trabajo que desarrolla cada una de las personas en su familia y comunidad. Esto ratifica
lo propuesto en otras investigaciones sobre el tema (Arango 2004; Figueiredo et. al. 1993; Joekes 2004;
Rocheleau 1998) El incluir una variable social como el género, en investigaciones de botánica o de ecología,
pueden llevar a tener un mejor entendimiento de los ecosistemas, su biodiversidad y la gente que los usa.
Aportando de esta forma importante información para su conservación y manejo.
Cuaoro 1. Es din: IO J J I bdo C buste:
Familia Especie Categoría UICN
Actinidiaceae Saurauia herthae Sleumer LC
Bromeliaceae Pitcairnia ferre -ingramiae M. Lutter & Dalistione VU
Melastomataceae Conostegia centronioides Markgr LC
Melastomataceae Meriania drakei (Cogn.) Wurdack LC
Moraceae Ficus lacunata Kvitvik VU
Passifloraceae Passiflora montana Holm-Niels & Lawesson EN
LC: Preocupación menor VU: Vulnerable EN: En Peligro
APÉNDICE 1
LISTA DE ESPECIES USADAS POR HOMBRES Y MUJERES DE SAN JACINTO DE CHINAMBI
G=Informante (H-Hombre, M=Mujer), HA=Hábito (A=Árbol, Ab=Arbusto, H=Hierba, He=Hemiepífita, L=Liana, E=Epifita; tomados
de: pa P. y S. León-Yánez 1999 y Valencia et. al. 2000), Uso PS Mec men Aa=Alimento
de animales, MeA=Medicina de animales, 0r=Ornamental, Ot=Usos variados), N ncuentran
las plantas que no tenían nombre común pero si uso y las plantas que tenían un nombre pers era particular x la persona
encuestada). En esta lista no se incluyen las morfoespecies sin nombre científico o género.
Nombre científico G HA Nombre común Uso
Abutilon dianthum C. Presl H Ab Jarabe M
Acalypha diversifolia Jacq. HyM Ab Pigua Me
Acmella brachyglossa Cass. HyM H Botoncillo Me, A
Acmella dla (Walter) Rich HyM H Botoncillos Me
Acrostichum H H elecho Á
i m) pizophylum Cass HyM H Mama Juana MeA, Me
Aechmea strobilacea L.B. Sm M H Pita A
Agave americana L HyM H Cabuya Ot
Ager Ab "Sin nombre" Me
Allamanda cathartica L. M Ab "Flor amarilla" Or
Aloe vera (L.) Burm. f. HyM H Sábila Or
Alternanthera mexicana (Schltdl.) Hieron. M H scancel Me
Anacardium occidentale L. H A “Sin nombre” M, Aa
Ananas comosus (L.) Merr. HyM H Piña A
Suárez, Dif. i | d pl t tre homt y j
Nombre científico G
Annona montana Macfad. HyM
Annona muricata L. M
Anomospermum chloranthum Diles M
Anthurium pier Linden HyM
Arachis hypogaea L. M
Axonopus micay eod HyM
ipd Sp M
accharis trinervis Pers. HyM
Pod corosilla H. Karst H
Bactris gasip M
Begonia fischeri Schrank M
Begonia parviflora Poepp. & Endl. H
Beilschmiedia costaricensis (Mez & Pittier) C.K. Allen M
Bidens cynapiifolia HyM
Billia columbiana dan & Linden HyM
Bixa orellana HyM
Borojoa sp. M
Piu Sis L. M
arborea (L) Lagerh. M
eo pui D. Don M
Brunfelsia chiricaspi Plowman H
Bryophyllum pinnatum (Lam.) Oken M
Bunchosia argentea Jacq.) DC. HyM
Calathea metallica Planch. & Linden M
Calathea crotalifera S Watson M
Calathea marantifolia Standl HyM
Calyptranthes cf. plicata McVaugh H
mpomanesia lineatifolia Ruiz & Pav. HyM
Canna indica L. M
Capsicum annuum L. M
Cardamine bonariensis Pers, HyM
Carica microcarpa Jacq HyM
Carica papaya L H
Casearia sp H
Castilla sp M
Cecropia sp. H
Cedrela montana Moritz ex Turcz. HyM
Centropogon cornutus (L.) Druce HyM
Pei dium ambrosioides L. M
Chusquea sp. HyM
Chon pubescens Vahl H
Citrus sp. M
inus dictos (Christm.) Swingle HyM
Citrus maxima (B x Rumph.) Merr. HyM
Citrus medica L. HyM
Citrus reticulata Blanco HyM
Cladocolea sp. M
Cleome hassleriana Chodat M
Clusia magnifolia Cuatrec. HyM
Codiaeum sp M
Coffea arabica L HyM
Coix lacryma-jobi M
Colocasia esculenta(L.) Schott HyM
Columnea medicinalis (Wiehler) L.E. Skog & LP. Kvist H
Conostegia centronioides Markar. HyM
Cordia sp. H
Nombre común
Guanábana
Corazón de la selva
aní
Pasto Micay
Chilca morada,
Chiica, Chilca blanca
"Sin nombre"
Chonta Duro
Chulco
Peluca, Jabón
María
Hoja blanca
Hoja de Bijao
Pisulde
Guayabilla, Guabilla
Alcira
Ají
Berro
Col de monte
Papaya
Guión
Caucho
Yarumbo
HIS Cedro
Gallito
Limón Meyer
Limón rea
Naranja, Naranja agria
imón
Mandarina
Suelda
Pestaña de Ángel
Madroño de monte
“Hojas con puntos amarillos”
Café
San Pedro
Papa balsa o de monte
Ataja sangre
Palo amarillo
Huevo de pava
1303
- A l af tha Ratanical D h Institut f Texas 2(2)
Nombre científico G HA Nombre comün Uso
Cordia hebeclada I.M. Johnst. M A Hojarasca M
ordia mexiana I.M. Johnst. H A Huevo de pava Aa
Cortaderia d oo Stapf HyM H Saboya, Sobretana A, Aa
ostus did usby H H Caña agria Ot
Costus lim un HyM H Caña agria Me
Coussapoa eds Mildbr. H He Matapalo M
Coussarea latifolia Stanal. HyM Ab Murciélago M
Cucurbita moschata — V allo A
Cupania cinerea Poe HyM A Guabo de mona Sabalito M, A
Cyathea caracasana loth D Domin M A “Sin no Or, Ot
Cyathea pallescens (Sodiro) D H A Helecho Me
Cyclanthera pedata (L) uM HyM V Achogcha A
Cymbopogon citratus (DC.) Staff M H Hierba Luisa Ot, Me
Dacryodes cupularis Cuatrec. H A Copal M
Daucus carota L. HyM H Zanahoria A
Dendropanax macrocarpus Cuatrec. H A M
de cf seguine Jacq.) Schott M H Hoja blanca Or
Diplazium trianae (Mett.) C. Chr. M H Hielo del monte A
Drymonia turrialvae Hansa. H H “Sin nombre” Or
Elateriopsis spectabilis Mart. Crov. M V ena Ot
Eleocharis elegans (Kunth) Roem. & Schult. HyM H Totora, Totorilla Me,Ot
Endlicheria cf browniana Mez H A Yalte M
Eschweilera aff integrifolia (Ruiz €: Pav. Ex Miers) H A Tete M, Aa
ut
Eschweilera cf pittieri R. Kunth HyM A Pilche M
Eschweilera cf. rimbachii Standl. HyM A anelo M
nu in rimbachii Standl. H A Tete, Canelo M
Eugen H A Arrayan M
es mili Des Moul. M Ab “Flores rojas con espinas" Or
. HyM A Chaparral A
Euterpe precatoria Mart. HyM A almito A
Faramea fragrans Standl. M Ab Café silvestre A
Ficus sp. HyM He atapalo M
Ficus andicola Standl. H He Matapalo M
Ficus cf carica L. H A Higüerillo Me
Ficus wales Kvitvik H A Higuerón M,Aa
Ficus pertusa Lf. M He Matapalo M
rcinia E (Kunth) Hammel HyM A Madroño de monte A, Ot
Guadua agustifolia Kunt HyM A Guadúa Ot
Guarea cartaguenya Cuatrec HyM A Chalde M
Guarea cf. polymera Little HyM A Sabalito Ot
Guarea kunthiana A. Juss. A aie Eig eee
Guarea macrophylla Vahl M A "Sin nombre M
Guarea polymera Little HyM A Copal M
Guatteria megalophylla Diles H A Tete
Guzmania lingulata (L.) Mez H He Vicundo pequeño Or
Gynerium sagittatum (Aubl) P. Beauv. M H aña Blanca A
dyosmum cuatrecazanum Occhioni H A Guayusa A, Me
Hedyosmum cosilla Solms M Ab Guayusa Me
bread an M H Girasol Or
Heliconia s HyM H Platanillo Or
Heliconia a Dodson & A.H.Gentry M H Platanillo Ot
Heliconia id nano & G.L. Mcrales H H Platanillo Or
Hemarthria ir) Stapf & CE. Hubb. HyM H Pasto tropical Aa
Altos pe Ruiz & Pav. M H Gallo de monte A
Heteropsis H He Vena de mimbre Ot
Suárez, Dif i | le plant tre | j
Nombre científico G
Hibiscus rosa-sinensis L. M
AO jid (Thunb) Ser. M
Hyeron H
Hyeronima macrocarpa Müll. Arg. H
Hyptis verticillata Jacq. HyM
onn balsamina L. M
Inga HyM
Inga p lora Benth. HyM
Inga edulis Mart HyM
Inga marginata Willd. H
Inga thibaudiana DC. M
Ipomoea batatas (L) Lam. M
Iresine herbstii Hook. M
Iriartea deitoidea Ruiz & Pav. H
Justicia pectoralis Jacq. H
Lantana camara L. HyM
Lantana s M
Lycopodielia Mis (L) Pic. Serm. M
Mangifera indica HyM
Manihot sp. M
Manihot esculenta Crantz HyM
Marrubium vulgare L. H
Matisia soegengii Cuatrec. HyM
Medicago sp. M
Melia azedarach L. M
Melissa officinalis L. M
Mentha pulegium L. M
Mentha spicata L M
Meriania aff drakei (Cong.) Wurdack H
ut 5 SD. HyM
a sp. HyM
Mas sp. H
Naucleopsis sp. H
Nectandra acutifolia jn & ~ Mez HyM
Nectandra guadaripo H
Nectandra purpurea us & pir Mez H
Nectandra reticulata (Ruiz & Pav.) Mez HyM
Nicotiana tabacum M
Ochroma Leite cer ex Lam.) Urb. M
Ocimum campechianum Mill M
Ocotea insularis M Mez H
untia s M
Opuntia ficus-indica (L.) Mill. M
Otoba gordoniifolia (&.DC.) A.H. Gentry H
Otoba gracilipes (A.C. 5m.) A.H. Gentry H
Pachystachys lutea Nees M
aspalum dilatatum Poir. HyM
Paspalum pi ded ia ex Flüggé HyM
Siem x iculat HyM
Passiflor M
rius ecol Sims M
Passiflora maliformis L. M
Passiflora montana Holm-Niels & Lawesson HyM
in | palenquensis Holm-Niels. & Lawesson HyM
Pelargonium H
NUR D GIU (L) L'Hér. M
o
o
EA A Gedeon SD Ge ow GegBOege BO To E
o [on
LO IDA» ILAILI-
>>>
oo
[n
LOILDILIIIDx»P»
Nombre comün
Flor rosada, F. blanca
Mo
Verbena, Chivo
Amerindia, Miramelindo
u
Guaba, Guaba de monte
Guabo silvestre
Guabillo
Guaba silvestre
Camote
"Hoja verde-blanca"
Pambil
Contra del día
Yuca Amarilla
Yuca
Amor seco
Zapote de monte
Hierba buena
“Sin nombre"
Tipo
Banano, Plátano, orito
Guayabillo
Guid
Aguacatillo
Aguacate de monte, Jigua,Yalde
abaco
Palo Balso
Albaca
Mal de aire
Cactus
Penca
Sangre de gallina
Sangre de gallina
Flor amarilla
Pasto miel
pedes
cillo
Ad silvestre
uyá
Granadilla de hueso
Granadilla de hueso
Granadilla
Chupana
Geránio
1305
1306
Nombre científico
Pennisetum purpureum Schumach.
Pentagonia cf. grandiflora Standl.
Persea americana Mill.
Pfaffia aurata (Mart) Borsch
Phaseolus sp.
Pope us verrucosum L. Mathieu ex Schott
Phyllanthus anisolobus Müll. Arg
Pilea fasciata Weld.
iper Sp.
Piper aduncum L.
ici ottoniifolium C. DC.
poen ae ~ H. Luther & Dalström
Plantago m
Pourouma Me Mart.
Pouteria sp.
Psammisia ulbrichiana Hoerold
Psidium acutangulum DC.
Psidium guajava
Psychotria alleni Standl.
Psychotria pilosa Ruiz & Pav.
a podo,
i nchospora watsonii (Britton) Davidse
Rollinia andicola Maas & Westra
Rosa hibrida L.
Rubus E Schltdl.
Rubus niv b
Rubus piene Poir.
Ruta graveolens
Saccharum officinarum L.
Salvia alvajaca Oerst.
Sansevieria cf. e Prain
Saurauia bullosa W
Saurauia cf. diae uh Spreng.
Saurauia herthae Sleu
Schizolobium a a ) S.F. Blake
Senna reticulata (Willd.) H.S. Irwin & Barneby
Sida poeppigiana (K. Schum.) Fryxell
Sloanea 2e Rugby
mila
me ai febrifuga Kunth
Solanum ind Dunal
Solanum be Cav.
Solanum iesus rsicum L.
Solanum nigrescens M. Martesns & Galeotti
Solanum quitoense Lam.
Solenostemon sp.
Sorocea steinbachii C.C. Berg
Stachytarpheta Ee (Rich.) Vahl
wietenia macrophylla
uud pace eo
Tagetes
Tapira g ao Aubl.
Terminalia amazonia (J.F. Gmel.) Exell
a o (Decne.) C.F. Reed
phylla S
nealmia o (Ruiz & Pav.) Poepp. & Endl.
Nombre común
Darliz, Puntero
Aguacate
Pilpe
Fréjol tumbe
Corazón morada
Cordoncillo, Cueche,
Matico de monte
Guabiluca
Santa María
Flor roja
Llantén
va
Caimito
Picaflor
Guabillo de hoja larga
Nacedero
Helecho
San Juanito
Anona
Rosa
Mora de monte
Mora silvestre
uda
Caña de azúcar
Albaca, Linaza
Hoja larga
Dalia, Moquillo
M O
Grillo Blanco
Hierba Mora
Tomate de árbol
Tomate de riñón
Hierba mora
Rabo de ratón
Caoba
Camote Morado
Hierba de gallinazo
Cedrillo
oble
R
Culantrillo, Culantro del monte
itute of Texas 2(2)
M
M, Ot
Or
M
A
Ot
Me,MeA,Ot
Ot, Me
Ot
Suárez, Dif. | uso de plant tre hombres y mujeres 1307
bg científico G HA Nombre común Uso
eobroma cacao L. M A Cacao A
v ndsia sp. HyM E Vicundo Or
Tithonia diversifolia (Hemsl.) A. Gray HyM H Tres dedos Me
radescantia zanonia (L.) Sw. M H igua Me
Trichilia cf laxipaniculata Cuatrec. H Á Chalde M
Trichilia surinamensis (Mic) C. DC. H A Guabo M
Triplaris americana L. HyM A Cucharrillo M
Turpinia occidentalis (Sw.) G. Don H A Sabaleta silvestre Ot
Urena lobata L. HyM Ab Abrojo Aa, Ot
mas caracasana (Jacq) Gaudich. ex Griseb. M Ab Ortiguilla, Ortiga Ot
M H Ortiga de monte, O. negra Me
te litoralis Kunth HyM H Verbena Me
Vernonia sp. HyM Ab Diazmaco Ot
Vetiveria zizanioides (L.) Nash M H Pasto Vetiver Aa
Virola elongata (Benth.) Warb. HyM A Brazilargo M, Aa
Xanthosoma sp. HyM H amacho
Yucca cf aloifolia L. M A "Sin nombre" Or
Zanthoxylum riedelianum Engl. H A Naranjuelo M
M H Maíz, Morochillo A
AGRADECIMIENTOS
A las familias de San Jacinto de Chinambi por compartir su conocimiento, en especial a Silvio Lara por su
apoyo en la coordinación de la logística para el trabajo de campo. Al proyecto Tejiendo Redes financiado por
el International Development Research Center de Canadá que ejecuta la Corporación Grupo Randi Randi
por el financiamiento para la ejecución de la investigación. Al QCNE y CHEP por abrir sus puertas a esta
investigación. A Danilo Gordillo y Nadia Ruiz, que participaron en la investigación desde sus disciplinas.
A Tatiana Castillo, Verónica Cortez, Gabriela Cruz, Segundo Chimbolema y Telma Paredes que apoyaron
en diferentes fases del proyecto. A Susan Poats, María Cuvi, Diana Fernández, Marcelo Silva, Georgiana
Bráulete, Jorge Caranqui, Maria Isabel Ríos y Carla Gavilanes por su apoyo, cometarios y sugerencias a la
investigación y al artículo. Además agradezco por sus sugerencias y comentarios Alina Freire-Fierro y al
otro revisor (anónimo).
REFERENCIAS
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Joekes, S. 2004. Género y Subsistencia en el norte de Pakistán. En: Vázquez, V. €: M. Velásquez, comp. Miradas al
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de México, México.
JORGENSEN, P. v S. LEÓN-YANEz (eds). 1999. Catalogue of the vascular plants of Ecuador. Monogr. Syst. Bot. Missouri
Bot. Gard. 75:1-1182
Kaner, K. v M. Duryea. 1992. Tapping women's knowledge: Plant resource use in extractive reserves, Acre, Brazil.
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eds. Ethnobotany: evolution of a discipline. Dioscorides Press, New York.
PHILLIPS, O. AND A.H. Gentry. 1993. The useful plants of Tambopata, Perú: |. Statistical hypotheses tests with a new
quantitative technique. Econ. Bot. 46:15-32
Roprlcuez, G., M. Blanco y F. AzorerrA. 2004. La diversidad hace la diferencia. Acciones para asegurar la equidad de
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Sustentables con Equidad de Género. Universidad Autónoma de México, México.
VALENCIA, R, C. Cerón, W. PALACIOS Y R. Sierra. 1999. Formaciones vegetales de la sierra del Ecuador. En: Sierra, R, ed.
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TREE CANOPY RESEARCH AND STUDENT EXPERIENCES
USING THE DOUBLED ROPE CLIMBING METHOD
Courtney M. Kilgore Harold W. Keller!
Department of Biology and "ad la
University of Central Misso
Warrensburg, Missouri 64093, A
Sydney E. Everhart
Department of Plant Pathology
University of Georgia
Athens, Georgia 30602, U.S.A.
Melissa S. Skrabal
Maple Woods Community College St. M vu pil cab Center
Mathematics/Science Division uth Frem
Eu yd Ms U.S.A.
Eds " Biology and na —
University of Central Miss
E Missouri Ho "m
Angela R. Scarborough Kenneth L. Snell
Department of Biology and Earth Science
Uni
iversity of Central Missouri
Warrensburg, Missouri 64093, U.S.A.
Kansas City, Missouri 64156, U.S.A.
Charly Pottorff
Charly5 International
ost Office Box 1282
Manhattan, Kansas 66505, U.S.A.
Joseph S. Ely
Department of Biology and Earth Science
University of Central Missouri
Warrensburg, Missouri 64093, U.S.A.
ABSTRACT
Students from the spin of Central Mi i explored tl ftu S A ins N
d
North Carolina, Daniel B 1F K
] Park in Tennessee
1
y, and Big Oak Tree State Park a Ha Tonka S Parl
and Pertle Springs
in Missouri from the summer of 2000 to 2007. The Doubled Rope Nisi cs Method (DRCM) was n to ne more an e Ara
trees without injury. This included fes vine Te and also 49 tree speci 11
, usually g
more than 30 m in total height. 5 id mandatory bing l held at Pertle Springs. The Wes of this
1 1 1 z ir J A eL : : f+} TARD(CM pa] A 1
4 Es i e rr r
11 1 A A Dl 1 T al Dl I +} Dijk] Phase AA vg E A F +]
r r 4 4, O D
DRCM and h d iated with climbing gear, Di sites, and climbing di d in detail P pui gear and knots
used in the DRCM 1 ibed lill lwi 1 ith the cli i 4 aerial
reproductive structures, and SOS in vertical transects from ne near eae level to 6 the treetop. The sampling lorca student
field pin iences, y of wW my y I ), tree canopy Bn soin coe and a
led +h de Wf, o A + PRU 1 MIiAAIL e
and education g
tili 1 nr j t amined th di rs n H4 :
Multi Dies importance yptog y y coa
mosses, wens. lichens, and fers, O j fi | including mollusks, nematodes, and tardigrades were also
noted T 1; 1 TWD HA Ez J 1 1 s g AE d T 1
O Da T e i4 [24 r
ask questions d develop! 1 1 ill i 1 led Mike hidenh
LI d dE fr o r
RESUMEN
E pe E E pea es Pe pe | ^ T de TAa Y E A 1 A And 1 1D. MI 1 “Great t Smoky Mountains" en Ten-
nessee y Carolina del Norte, Bosque Nacional * Daniel Boone” en Kentucky y ae Estatal “Big Oak Tree," Parque Estatal "Ha Ha
onka"
y Pertle Springs en Missouri, desde el verano del 2000 al verano del 2007. E
qur Ac do ENN árhal E E PERE fein E einn a
do de cuerda doble para escalar (MECD) se usó para
E 341 A S PPM jT. AQ i de ekal
7 : r
a du D exceden los 60 cm de diámetro a la altura del pecho y más de 30 m de la altura total y tres especies de enredaderas.
1 Ri; 1 Axl 1 iil Acita d tn tra hai TA inf, 165 A
? 3 Ey J r
obtener ayuda financiera y una esas della del MECD y 1 iencias de i igaci de 1 li I ]
por una Fase de Aventura, una Fase de Laboratorio, y una rer de Publicación. Las ventajas y d ¡jas del MECD y riesgos asociados
con el alae de e sitios e ue y bol El equipo de escalar y los nudos usados
: " 1 qe: 4 55,3 3 1 1 A +
en el MECD j I p ,las
estructuras reproductoras aéreas, y criptóg; icales desde el nivel del suelo 1 la copa de l boles. El protocolo
del muestro 1 H 7 d ld he td J (TY 1 j Tw T UNE t
r AA r d 7
£T.
3
, 500 E 4" Street, Fort Worth, Texas 76102-4025, U.S.A.
J. Bot. Res. Inst. Texas 2(2): 1309 — 1336. 2008
1310
AA L E : 1
Colegio de £ jemp p I
en la ocurrencia y la imy ia de criptóg t macromicetos, musgos, hepáticas, líquenes, y
4
los incluidos i cos Wendtsdos y tardigrados. pias oe pu E investi-
i la
1 : 15: 1
helechos y
A 23
o
1 d i ld 1 de arb ly para earen haser
X e o
preguntas, y d llar las hipótesis q A imi de la biosfera.
doubled limi hod (DRCM), Great Smoky Mountains National Park, Tennessee and North Carolina,
I REY
Daniel Boone National Forest, Rentados Pertle Springs, Missouri, myxomycetes, tree canopy, tree climbing
INTRODUCTION
This paper describes the climbing methodology, sampling protocol, and student research experiences used
to survey and inventory tree canopy biodiversity in the Great Smoky Mountains National Park (GSMNP) of
Tennessee and North Carolina, Daniel Boone National Forest (DBNF) in Kentucky, and Big Oak Tree State
Park, Ha Ha Tonka State Park, and Pertle Springs in Missouri. This research project began the summer of
2000 and has included more than 20 undergraduate and master's degree level students at the University of
Central Missouri (UCM).
Landscape vistas in GSMNP make this the most popular national park with over 11 million visitors
annually. Scenic mountains, waterfalls, valleys, and the mixed diversity of deciduous and evergreen trees
highlight brilliant fall leaf colors, old growth forests host champion-size trees, and summer haze and storms
create beautiful cloud fields in high elevation areas of the Park (Fig. 1A—H). Located just a short distance to
the north of GSMNP, the DBNF in Kentucky has a rugged topography that features scenic land formations
and waterfalls, especially at Cumberland Falls State Park (Fig. 11-K
The purpose of this paper is to provide information on how to obtain financial grant support and the
additional narrative details and color images that document the terminology of climbing gear used to gain
tree canopy access. Also provided are detailed descriptions of climbing trees using the Doubled Rope Climb-
ing Method (DRCM), and a body of literature that describes tree canopy research in temperate forests. The
field experiences designated here as the Adventure Phase used the DRCM to gain access to the tree canopy.
The advantages of DRCM serve as an example of tree canopy access that will help others with planning and
executing tree canopy exploration. Results are summarized and in some cases reported for the first time.
National Science Foundation Small Grant for Exploratory Research
This research project would not have been possible without the financial support of the National Science
Foundation (NSF) Small Grant for Exploratory Research (SGER) through the Biodiversity Surveys and Inven-
tories Program, Division of Environmental Biology. When applying for such a grant the appropriate Program
Officer should be contacted to determine if the project proposal idea meets the specific SGER criteria, and
in our case, the application of new expertise or new approaches to established research topics. Our research
idea of accessing the tree py using the DRCM to explore and collect , macrofungi, lichens,
liverworts, mosses, and ferns in the Great Smoky Mountains National Park (GSMNP) appeared to match
Fic. 1. AH n e I y AA H Ai : 1D. LL A A rt fC atli nl g T. r +. +h | I R f J f. Yall J I
£ H a IM | TL f. J J [| J A MA + J 1 A th
area and Tou in me discovery of the new W'spedes Marheg arboricola. = Treetop iew along Ridge Mountain Road overl g Cades Cove, D
fter af in sl <E Clingman's Dome, the | ig! int in T t 2,025 m, showi g y lead IF fir trees caused
L y +L Dal y Jal 1? i an | H : E Fr Palla i I { g L n : g r LAS - HI r rl g. G.
Call I | g 0 I g I tA fal + | I L I Le J £E aL r J r L| n J H I ive, old growth, tulip poplar
p +. J Alatan E fal EL I L £^ kel I L g al + I-K Kantucky sitas J n ID FEL | Da-1JD I
Kilgore et al. T —áÓ 1311
-
—
MOUNTAINS =
^& NATIONAL PARK “
ZE
GREAT SMOKY
n
AN INTERNATIONAL BIOSPHERE DESE AVE
Rock
R
1312 J l of the Botanical R h Institute of Texas 2(2)
this criterion. We proposed to go where no one had gone before in the tree canopy at GSMNP and to make
observations and collections of targeted groups of cryptogams (Keller 2005a, 2005b).
The program officer requested that HWK prepare a proposal, limited to five pages, entitled "SGER-RUI:
Tree Canopy Biodiversity (myxomycetes, macrofungi, mosses, liverworts and lichens) in the Great Smoky
Mountains National Park." Also included, was a clear statement of why this proposed research should be
considered exploratory and high risk, and the significance of its potential impact on the discipline or field.
It was important that the applicant work closely with the program officer since the SGER ls undergo
an internal merit review by NSF staff. In our case the draft narrative, budget details, and biographical sketch
were previewed before submission by the NSF program officer. The maximum budget was $100,000, but
the usual average amount was less for a project period of one to two years. Certain promising SGER projects
approved by the program officer and division director may be extended for up to six additional months
and supplemented with up to $50,000 in additional funding. Most of the budget supported student wages,
climbing equipment and supplies, travel, and meals. In most cases lodging was provided by Discover Life
in America in the GSMNP and the United States Forest Service in the DBNF
METHODOLOGY
Methods to Access the Tree Canopy
Tree canopy exploration reached new heightsi in the 1970s with a variety of canopy access techniques modi-
fied from mountain climbing methods including the use of polyester ropes with a high tensile strength along
with various ascenders (Nadkarni 1988; NERIS 1994; Nadkarni 1995; Lowman & Wittman 1996). Early
canopy access usually resulted in a solo researcher climbing a tree using a method such as the single rope
technique (SRT), cherry pickers, or ladders (Lowman 2004). The single rope technique was the first rope
technique used to access the canopy in the 1970s (Lowman 1994; Lowman & Bouricius 1995; Lowman 2004).
This method allows a researcher to sample a vertical transect into the canopy that can be replicated at other
sites (Lowman & Wittman 1996; Lowman 1999). Rope studies with single climbers g 1 small
data sets (Nakarni 1995). oa dic, ropes can be used to attach structures in the canopy suci as collapsible
Tiat
platforms used to observe arboreal | over a period of time. This allows ple scientists to perch
in the canopy and observe animal activity with minimal disturbance. The SRT cannot be used to access the
upper branches in the canopy as noted by early arborists (Nadkarni 1988; Moffet & Lowman 1995).
Information about different methods of accessing the tree canopy are described in various books and
journal articles as well as part of a lecture given by H. Bruce Rinker (2004) at a symposium entitled “Tree
Canopy Biodiversity in the Great Smoky Mountains National Park” presented at the Mycological Society of
America 2004 annual meeting held jointly with the North American Mycological Association. Chapter 1 by
Moffett and Lowman (1995) includes tree canopy access techniques used for scientific studies, and Table
I evaluates the different access methods assigning an assessment value from 1 (least desirable) to 10 (most
desirable). The information contained in this table will help tree canopy biologists select the best access
methods for their own special research projects. Updated progress in canopy walkway construction and
canopy cranes is included in Chapters 23 and 25 (Lowman & Rinker 2004) and emphasizes how canopy
access techniques will impact future sampling, hypotheses testing, and facilitate ecotourism in the tree
canopy.
Ground based methods oí canopy research are useful in collecting data for mobile or sensitive species
and demanding projects such as insect surveys. In addition, radio telemetry can be used to study arboreal
animals, colorful paint can be applied to arboreal reptiles, and arthropods can be collected en masse using
insecticidal fogging (Moffet & Lowman 1995). Traps can be hoisted in the canopy using rope and pulley
systems; for example, a team of researchers from UCM used “composite flight-intercept traps” (Fig. 2A) at
Big Oak Tree State Park, Missouri to survey insect biodiversity (Wilson et al. 2003)
Recent canopy research favors a more collaborative approach and focuses on methods such as towers,
canopy walkways, cranes, and the canopy raft and sled, all of which can accommodate several scientists at
h and student i 1313
Kilaore et al.. Ti
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banned 1966, Atlanta, Georgia.
a time (Lowman 2004). These methods usually sample from the outer limbs or periphery of the tree, not the
trunks, whereas rope climbing allows the researcher to be in direct contact with the trunk of the tree. Tow-
ers represent a permanent structure for canopy research; however, few trees are within reach from a single
point. Canopy walkways and cranes are attached to towers and provide a larger sample area (Lowman 1994).
Canopy walkways are often more affordable than cranes, which cost between one and five million dollars
(Lowman 2004, and are often used for ecotourism and conservation (Lowman & Bouricius 1995; Lowman
et al. 2002). The canopy raft is of French design and effectively creates a platform lifted by a dirigible to the
treetop canopy from which scientists can study the canopy/atmosphere interface (Lowman 1994; Nadkarni
1995; Lowman & Wittman 1996; Lowman et al. 2002; Lowman 2004). A smaller raft that is trailed behind
the dirigible, called a sled, is used to skim the canopy allowing scientists to collect leaves, epiphytes, vari-
ous flowers and their pollinators (Lowman 1994; Nadkarni 1995; Lowman & Wittman 1996; Lowman et
al. 2002; Lowman 2004). Other more daring methods of cooperative canopy exploration include ultralight
planes (Lowman & Wittman 1996).
Methods to access the tree canopy have not generally complied with Occupational Safety and Health
Administration (OSHA) regulations (Grushka et al. 1999). A team of researchers developed the first OSHA
1314 J l of the Botanical R h Institute of Texas 2(2)
regulated fall arrestance canopy access system to be used by scientists at the University of Columbia Bio-
sphere 2 Center for Research and Education in Oracle, Arizona (Grushka et al. 1999). Although risky, canopy
research is essential as biologists race to survey species in dwindling forests worldwide. It is estimated that
over half of the world’s species of plants and animals occur in the tree canopies (Nadkarni 1995; Lowman
2004). Canopies also play an integral role in nutrient cycling, are a major source for photosynthesis, and
serve as a carbon sink for atmospheric CO, (Lowman 1994; Lowman & Wittman 1996; Lowman 2004).
Canopy access methods should be considered when creating a research project. Studies can involve
sessile organisms, mobile organisms, or interactions of sessile and mobile species within the tree (Lowman
1999, Lowman 2004). Sessile organisms such as epiphytes that grow along the trunk and inner branches of
the tree are the easiest to study with rope methods in the canopy and represent fewer logistical constraints.
However, certain plant organs such as flowers and buds produced on the tips of branches and uppermost
branches of the canopy are more difficult to reach with rope climbing methods and would therefore be better
sampled using a crane, walkway, or raft (Lowman 2004). Another way to access mature, aerial reproductive
structures (cones and dried fruits) on branches in the outer tree canopy used limb walking with the DRCM
(Kilgore 2008).
ible using dirigibles, cranes with gondolas, cherry pickers, elevated platforms
or walkways, inflatable a pecoris tree houses or bridges, ladders, towers, boats, sleds, pole climb-
ing spurs and belt (Fig. 2B,C), free hand or rope climbing methods. Even so, the current literature does
not include a detailed discussion of the advantages and disadvantages of the DRCM, as a useful method for
conducting canopy research, the description and use of the climbing equipment (ropes, climbing saddles,
knots), safety precautions as these apply to tree canopy survey research, and a stepwise *how to" approach
of the climbing methodology. Previous publications include general information about the DRCM (Snell &
Keller 2003; Keller et al 2003; Keller 2004a, 2004b, 2005a; Everhart & Keller 2008; Everhart et al. 2008;
Kilgore 2008) used mostly in the GSMNP.
Requirements for nme and the Tree Climbing School
Selection of student li first involved national recruitment efforts including [ g
in national newsletters, on our web site: http://faculty.ucmo.edu/myxo/, on electronic bulletin boards, and
making announcements at regional and national professional scientific meetings such as the Association
of Southeastern Biologists and Mycological since! of America apu meditiga: Local recruitment efforts
involved displaying flyers and posters in university hall I bing images and scenic color
images from past field experiences, presenting pace seminars, mund delivering announcements in
biology classes. Three phases of this research proe were b eU emphasized; the Adventure Phase (tree
climbing and sampling); the Laboratory Phase ( d moist chamber cultures); and Publication
Phase (poster and oral platform presentations alone with space team written articles for newsletters, or
refereed journal papers based on individual student research projects).
Prospective students were interviewed to determine their interest and qualifications to complete all
three phases of the tree canopy biodiversity research project. A student profile included the following in-
formation: field experiences such as hiking, backpacking, and camping in remote areas; athletic activities
that involved team sports or activities, strength building exercises, repelling, rock or wall climbing, and
tree climbing using rope systems; ane ude! (biology preference), undergraduate classes, grade point
average, and biology j including use of computer software, database spreadsheets,
microscopes, digital cameras, use col rosaria maps, and global positioning systems; future career in-
terests, and especially interest in graduate school. Strength building exercises to increase upper body arm
strength and body form were important in successfully reaching upper parts of the tree canopy. Students
with research projects already planned in advance that included tree and vine species targeted for sampling
were more successful in the Publication Phase.
Each qualified student was required to sign a Release and Acknowledgment of Risks Agreement prior
to attending the tree climbing school. This document described the risks involved in travel to, from, and
Kilgore et al., Ti Í ] student i 1315
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1316 Journal of the Botanical R h Institute of Texas 2(2)
within forested areas, hiking in remote areas, dangers such as stinging insects, wild animals, poison ivy, and
using ropes to climb trees. In addition, each student was required to be more than 18 years of age and have
medical insurance. Contact information was provided in case of medical emergency. Additional optional
information included any physical condition that might increase the risk of being in the field for prolonged
periods of time, and the endurance required to hike long distances and climb trees. Emphasis was given to
team sports and cooperative activities since the Adventure Phase was a team effort that required living and
working in close quarters. We had a minimum of 4 to 8 participants in the Adventure Phase for the GSMNP
and DBNF field trips.
A two-day tree climbing school was held in late April or early May at Pertle Springs, Warrensburg,
Missouri (Fig. 3A-H). Charly Pottorff, a professional arborist from Manhattan, Kansas, was the first tree
climbing instructor (Fig. 3A,D,E,F,H). He has been a professional arborist since 1968 and also a member of
the Kansas Arborist Association and I ional Society of Arboriculture. His involvement for many years
in popular regional Tree Climbing Jamboree competitions as an organizer, convener, and practitioner of
rope climbing techniques has resulted in being on the cutting edge of climbing techniques and equipment
design and safety. The combination of his physical exercise routines and tree climbing expertise allowed
him to serve as a role model for oe next generation of tree canopy climbers.
Instructors led 1 physically demanding to ensure that each student
climber had developed he: physical strength and conditioning to safely climb 30 m or more into the tree
canopy. This included a series of exercises that helped build upper body strength. Indoor exercises included:
pull ups with palms up or down on the bar, cliff hangers with hands facing one another with palms on the
bar (Fig. 3A), finger tip push ups, leg lifts, and the ee ao — an axe ee finger walk (Fig. 3C) that
increases finger strength. Outd ran stadium dleg strength. Rope
climbing practice at Pertle Springs included using the cba equipment and gear (knot tying Fig. 3B,)
and demonstrations by the instructors (Figs. 3D,E; 7A,E). Climbers were supervised by the instructor on
their first climbing attempt (Fig. 3F). Once in the tree canopy, climbers practiced using lanyards, knots used
in advancing, relaxing in the saddle, and standing on a branch to rest and adapt to the canopy height (Fig.
3G). A group picture was taken at the completion of the tree climbing school (Fig. 3H). Only the students
who successfully completed the training course on the basic DRCM were considered for the project.
The Adventure Phase, Laboratory Phase, and Publication Phase had to be satisfactorily completed to
meet the objectives of this project (Counts et al. 2001; Keller & Skrabal 2002; Keller et al. 2002; Snell 2002;
Keller & Snell 20022; Keller & Snell 2002b; Snell & Keller 2001; Snell & Keller 2003). Each student kept a
daily journal of observations that in some cases were used in research papers. Students were given writing
assignments as part of team-writing projects. These narratives were collated into popular articles published
in different newsletters (Counts et al. 2000; Henley et al. 2000; Keller & Skrabal 2002; Keller et al. 2002;
Skrabal et al. 2001; Keller & Everhart 2007; Kilgore 2007; Kilgore & Keller 2007; Keller et al. 2008; Kilgore
& Keller 2008).
Advantages and Disadvantages of the Doubled Rope Climbing Method (DRCM)
The two basic rope climbing systems most often used are the single and doubled rope methods (Jepson
2000). Advantages and disadvantages are associated with both techniques. Advantages of the single rope
technique (SRT) include quick canopy access because the rope can pass over several tree crotches without
impeding climbing, and it is safe and efficient when practiced correctly using back-up ascenders and de-
scending devices (Dunlap 2002). A disadvantage of SRT is that the climber cannot advance higher in the
canopy by advancing the rope from the original installation point. The climber can only advance if a separate
climbing line is used (Jepson 2000). Another disadvantage of SRT is that several pieces of equipment are
needed, including ascenders, pulleys, and stir-ups, which are a more mechanized mode of ascending into
the tree canopy. If any of these mechanical devices are lost the climber cannot complete the climb (Charly
Potorff, pers. comm.).
An advantage of using the DRCM for canopy research in the backcountry is that the necessary equip-
Kilgore et al., T | | student i 1317
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1318 Journal of the Botanical R hinstitute of Texas 2(2)
ment is minimal so that each climber can pack the entire set of climbing gear into a standard backpack.
The use of additional equipment such as ascending and descending devices is optional (Adams 20072;
Adams 2007b). To minimize the necessary climbing equipment, it is recommended that the climber rely
on the climbing rope, split tail, safety rope, and his or her knowledge of the proper climbing knots used to
access the tree canopy. This provides more mobility when hiking over long distances and allows quick use
at specific sites. In addition, the DRCM is a non-invasive method that allows the climber to advance the
rope to higher branches, often reaching near the top of the tree (Keller 20042). The DRCM requires more
strength than the SRT and climbers must d in top pay condition to eo master this technique.
Some researchers may see this as a di d a hindrance to completing tree canopy research in the
backcountry. However, it is never a disadvantage to be in good physical condition when conducting tree
canopy research in the backcountry and using DRCM was an incentive for the students to be in excellent
health and develop athletic skills.
An advantage for both rope methods is that they ffordable than other methods such as canopy
walkways, towers, or cranes (Moffett & Lowman 1995). Ropes are also easily transported to different study
sites, allowing researchers to sample from trees that are geographically separated rather than grouped to-
gether as in walkways, towers, or cranes; however, several research climbers are needed to sample a large
number of trees.
Tree Climbing Hazards
Hazards to climbers can be avoided with pre-climbing safety inspections (Fig. 4A—E). Jepson (2000) includes
an entire chapter on pre-climbing safety inspections. Safety inspections should be done on equipment, the
climbing site, and the tree. Possible hazards in the equi] include frayed ropes or lanyards, inconsistent
diameter of ropes, heat damage on ropes or lanyards, unsealed rope ends, cracks in safety helmets or eye
protection gear, holes in gloves and clothing, tears or broken stitching in the climbing saddle, elongated
holes in the climbing saddle waist buckle, and cracks or distortion in D-rings or carabiners that do not
lock properly. Climbing gear should be free of dirt and debris to avoid jamming carabiners and possible
loosening of knots.
Each study site must be inspected for animal activity, such as bees in the trees (Fig. 4A) hornet nests in
the ground, venomous snakes (Fig. 4B), and also bear activity, wild pigs, raccoons, and possibly mountain
lions. When climbing in the mountains, it was important to be aware of terrain by looking for steep slopes,
boulders, loose rocks, landslide areas, and mountain creeks. Poisonous plants are often common on dis-
turbed sites, so the climber must know how to identify poison ivy, poison oak, poison sumac, and stinging
nettle.
Stormy weather and lightning pose serious hazards. Lightning is a deadly threat when working in the
tree canopy (Fig. 4C). Weather forecasts in the GSMNP were unreliable since they were not specific to any
particular region of the park. It frequently rains heavily in one part of the park with little or no rain in other
parts. At lower elevations, movement of storms can be observed at fairly long distances. This allowed the
climber sufficient time to complete work and exit the tree safely. Higher altitudes made it difficult to tell
exactly how far away a potential storm was located. Furthermore, rain is also a hazard because wet ropes
are more likely to slip, tightening knots and making ascension more difficult. Wet branches reduce traction,
making it more difficult to climb into and advance within the tree. Ground sites become wet and slippery,
increasing the chance that a climber would fall. Climbing before, during, and after thunderstorms and rain
should be avoided.
Climbers should be aware of any electrical hazards such as power lines and grounded electrical lines
(Fig. 4D). Septi t d drain fields should also be avoided. The climber should be careful not to launch
the towbap over a limb in the direction of any vehicles, buildings, or people. Hikers and bystanders should
be kept away from the climbing area, illa public curiosity and education were encouraged, so if a hiker
d work for the moment to explain our activities and recommend
came into the area, all team
rli
a safe distance for viewing.
Kilgore et al., Ti py | | student experi 1319
The climbing tree also poses potential hazards to the climber. These include signs of damage such as
lightning damage on the trunk (Fig. 4C), broken limbs, dead trees and branches (Fig. 4E,) cavities at the
base of the tree, poison ivy, thorns, honey bees in cavities, presence of wood rotting fungi, and loose bark
(Fig. 1H). When foliage is dense in the canopy, broken limbs may not be visible. Binoculars help locate hard
to see hazards, especially dead branches or damaged sections of the tree. Prior to climbing, the climber tests
the sturdiness and integrity of the limb. If the limb bends or makes a noise indicating it is damaged, the
limb should be avoided.
Tree Climbing Equipment and Gear
The Big Shot.—The Big Shot is an oversized slingshot that has a 2.4 m metal pole with a rubber gripper
attached to the bottom end and a detachable head consisting of a forked metal frame on the top end. At-
tached to each end of the forked head are two strands of durable elastic rubber tubing that are joined in the
middle with a sling that cradles the throw bag, similar to a slingshot. This metal pole can be separated into
two 1.2 m sections that can be tied and carried on top of the backpack and assembled on site when ready to
use (Fig. 5A,B). The Big Shot is used to launch the throwbag with the throwline over the desired climbing
limb, thereby allowing climbing ropes to be installed in the upper canopy.
Throwbags.—Throwbags are made of thick woven material or leather, double stitched at the seams
and filled with non-lead-based pellets. Throwbags usually come in two shapes: a tightly filled, aerodynamic
torpedo-shape, and a partially filled tear-drop-shape for maneuvering through narrow crotches or inclined
branches. Throwbags should be inspected before use to make sure the stitched seams have not broken. A
metal O-ring is located at one end for throwline attachment. Throwbags weigh 227 g, 340 g, 397 g, or 454
g and each weight was able to reach different heights in the tree (Fig. 5D). The lightweight throwbags (227
g) were used more frequently by female tree climbers, reaching heights up to 25 m.
Throwline.—The throwline (also called slickline) is a small-diameter (approximately 1.75 mm), light-
weight line that is yellow or multicolored (Fig. 5B; 6]; 7A—D) and is designed to be the first line installed
in the tree. The throwline is tied to the throwbag O-ring with an Anchor Hitch knot that is dressed so no
protruding ends wedge in crotches or hook on branches. A cloth storage bag approximately 14 cm deep and
10 cm wide is used to store the throwline to keep it from tangling. One end of the throwline is attached to
the bottom of the storage bag and the other end is attached to the throwbag. Usually a throwline of 60 m
is used, which means the shooter can reach a tree crotch or limb about 30 m high. Continual usage of the
throwline often results in kinks that must be removed by tying and stretching the line between two trees.
The ends of the throwline often frayed and this was fixed by melting the ends using a match or butane
lighter.
Climbing ropes.—Ropes are available in different colors and lengths (Fig. 5A,B). Climbing ropes of 36
m in length are coiled in about 1 m lengths and a gasket hitch knot used to keep the rope from uncoiling
(Fig. 6L). Once installed, the rope is limited to 18 m in height since it is doubled over a branch or crotch.
Climbing ropes are a bright orange or a white color, 16 stranded, 1.3 cm diameter, with a tensile strength of
3,175 kg. One end has a spliced “eye” so a carabiner could be used to “tie in" (Fig. 6J) instead of tying a knot
to the climbing saddle O-ring. The other end of the climbing rope has spliced ends with whipped twine to
prevent the ends from fraying (Fig. 6A). Another white rope 60 m in length is used to reach a total height
of 30 m after installation over a branch. The objective is to get the climbing rope installed at the highest
possible point so advancing higher in the tree canopy takes less time. Split tails were approximately 1.5 m
to 2 m long and were made the same as the climbing ropes (Fig. 5A).
Ropes are the climbers' lifeline and must meet minimum Occupational Safety and Health Administra-
tion standards that include abrasion resistance and a minimum breaking strength (dry test) of 2,449 kg.
Ropes should be inspected before use to make sure that possible weakness due to abrasion, cuts from knife
use or sharp bark edges, and accumulation of dirt have not weakened the rope and increased the risk of
breakage.
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1320 Journal of t te of Texas 2(2)
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saddles have an extra wide 15.2 cm back support that is foam filled and incased in soft brown leather (Fig.
3E,F). The saddle can be adjusted to the waist and padded leg straps adjusted to the thighs with buckles.
The leg straps of this saddle are sometimes uncomfortable in the groin region after prolonged periods in the
tree canopy. Each saddle has two D-rings for carabiners or to tie the climbing rope and split tail, two O-rings
for lanyard attachment, and also two snaps for equipment such as a knife for removing bark samples, a reel
bound tape for height measurements, storage bag for throwbags and throwlines, and water bottles. Leather
saddles weigh approximately 3.2 kg and increase the weight carried in the backpack.
The butterfly saddle is lighter weight at approximately 1.8 kg and is easier to carry (Fig. 3B,D,G; 5B;
7F-1,K,L). The padded backrest is 18 cm in height giving back support to the climber when in the saddle
for longer periods of time. The belt and leg loops buckle with a smooth-action quick release. This saddle has
two D-rings, two small O-rings and three cloth attachment loops on the back rest. Female student climbers
prefer the butterfly saddle because it is more comfortable.
Each saddle provides freedom of movement in a horizontal or upside down position that relaxed tired
legs. Both hands were free to handle a heavy-bladed knife to collect bark samples, or to use a 20X hand lens
to scan the surface of the bark, or use a whistle to communicate with other climbers and the ground crew
(Fig. 5C).
h and student experi 1321
Kilgore et al., T py
Lanyards (safety ropes).—This personal safety rope is used as an additional “tie in" while gathering
bark samples or as a single tie in when advancing the climbing rope higher in the tree (Fig. 5A; 7F-L, K,D.
Lanyards consist of a thick rope, connecting devices on each end with carabiners or a rope snap, and a
lanyard adjuster to tighten the rope around the tree trunk. Sometimes it is difficult to secure the lanyard
because of the large diameter of the tree trunk. Climbers usually use their legs to pull themselves as close to
the tree trunk as possible and swing the rope around the trunk, catching the opposite snap end with their
feet. The lanyard snaps are attached to the two D-rings on the climbing saddle or to carabiners.
Head gear.—Several types of head gear are worn, but clearly the best is a form fitting helmet without
a brim or edge. Brightly colored red, blue, orange, or white helmets make it easy to track the climber high
in the tree canopy, especially when thick foliage obscures the view from ground level. A helmet is worn by
both the climber and ground crew members. Helmets protect the climber from falling objects such as debris
from limbs, falling tree bark, and branches. Chunks of bark sometimes fall from above the climber, espe-
cially in old growth champion-sized trees, and the combination of gravity and height can result in serious
injury to anyone underneath. Special tree climbing helmets made of durable hard plastic that is lightweight
(approximately 0.5 kg) with an inner padded lining feature an external chin sd to prevent the helmet
from becoming detached. Ventilation holes along the side prevent i 1 the head band
Sometimes a bandana is worn to prevent slippage of the helmet (Fig. 5A,B; 7F-I K).
Gloves.—The climbers must wear gloves to protect their hands from rope burn, to assure a firm grip
on the climbing rope, to prevent scratches from sharp edges of bark, and to protect fingers when using a
sharp-bladed knife when collecting bark samples. The palm is made of a latex coated, rubberized, and
roughened surface and the backside from breathable cotton and polyester fibers (Fig. 6G-I; 7C,D,F,H,1,L).
Knots used in the Doubled Rope Climbing Method
Six basic knots are used: the Anchor Hitch, Figure Eight stopper knot, Blake’s Hitch, Half Hitch, Monkey Fist,
and Gasket Hitch (Fig. 6A—L). These knots, except the Half Hitch, are described in detail with illustrations
in Jepson (2000). Toss (1990) presents an excellent, illustrated guide to tying knots. The website, http://
www.iwillknot.com, gives animated examples of how to tie various knots and serves as a valuable resource
for novice knot tyers (Hudson 2008).
Anchor Hitch.—This attachment knot is used to secure the throwline to the throwbag (Fig. 6A—C). It
can also be used to tie one end of the rope to the D-ring or a carabiner on the climbing saddle if the climb-
ing rope lacks an eye splice. This knot must be finished with a Figure Eight stopper knot to ensure that
no creeping occurs. Creeping is the term used when the rope slowly moves through the knot, eventually
untying the knot.
Figure Eight.—This attachment knot is a type of stopper knot (Fig. 6D—F) used as an added safety
measure for the climbers to prevent the climbing rope from creeping through the attachment hitches, such
as the Anchor Hitch and Blake's Hitch (Fig. 6C,F). This knot is easy to tie and resembles a figure eight when
tied correctly.
Blake's Hitch.—This knot is a type of friction hitch, also known as a climbing hitch (Fig. 6G-I) and
serves to grab the rope when it is under tension. When tension is lessened, the hitch can be moved up or
down the rope. The Blake's Hitch knot is a variation of the Tautline Hitch, but is considered to move more
smoothly on the rope with less friction to the climbing rope, making it more desirable as a climbing knot.
The Tautline Hitch is also notorious for creeping and also binds and tightens to the rope, requiring frequent
adjustments during climbing. Although the Blake's Hitch is more tedious to tie than the Tautline Hitch, it is
a more functional climbing knot. A Figure Eight stopper knot is always tied to the tail of the Blake's Hitch
to prevent any creeping (Fig.6F).
Half Hitch.—This knot is used to attach the throwline to the climbing rope before hauling the climbing
line over the limb. Four or five Half Hitches are tied to one end of the climbing rope and will tighten when
the throwline is pulled (Fig. 6J). Most importantly, they do not interfere with the climbing rope sliding over
the crotch of a limb.
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1322
Monkey Fist.—This knot is considered a type of throw knot (Fig. 6K). It is used for throwing one end
of the climbing rope over a limb when advancing in the canopy. Throwing this knot works best when the
climber is advancing short distances in the canopy. It can be tedious and awkward to throw, as the climber
is usually forced to throw the knot straight overhead. Once the knot has successfully passed over the limb
it unravels sending the rope back to the climber. If the Monkey Fist knot is unsuccessful, the climber may
opt to use a throwbag attached to the throwline to gain access to the upper limbs in the canopy.
Gasket Hitch.—This knot is used when coiling the climbing rope for storage after use (Fig. 6L) and
allows the climber to hang the rope up to dry or for storage. If tied successfully, the climber should be able
to toss the coiled rope without it uncoiling.
Climbing Procedure
The tree climbing procedure (Fig. 7A-L) began with a pre-climbing inspection to search for any dangers
that might impact climbers. The buddy system was used, where each climber had a ground crew member,
consisting of one or more individuals instructed in climbing techniques, safety procedures, and collecting
protocols. Every member of the climbing team (climb o prouade crew members) wore a hardhat and fol-
]
lowed proper safety procedures. Ground crew members for double-checking the pre-climbing
inspection, maintaining the throwline and the climbing ipe. at the base of the tree, communicating with
the climber, and supplying the climber with equipment such as hammers and collecting bags. Eye contact
was maintained with the climber to ensure safe conditions, to recover bags of bark collected and dropped
to the ground, to collect bark samples at 2 meters in height, to record data on a data sheet, and to manage
the perimeter of the tree, thus keeping the area free of debris and hikers away from the hazardous zone.
Each climber wore gloves, safety glasses, and a protective helmet when climbing the tree and taking
samples. Safety glasses kept debris out of the climbers' eyes and the helmet protected the climber from
falling branches and limbs. The climber wore a long-sleeved shirt, long pants, sturdy hiking shoes, helmet,
and a climbing saddle designed for arborists. The long-sleeved shirt and pants prevented abrasion to the
skin while maneuvering in the tree, through branches, and also helped prevent insect bites. Ground crew
members also wore helmets to protect them from falling branches, limbs, throwbags, collecting bags, and
any equipment that needed to be tossed from the canopy by the climber.
When the tree was declared safe for climbing, a 227 to 454 g pellet-weighted throwbag was attached
to a lightweight throwline and either thrown (Fig. 7A) or shot over the crotch of the desired tree limb using
the "Big Shot" (Fig. 7B). The throwline must be free of knots and neatly coiled into a bag placed approxi-
mately one meter in front of the Big Shot. This area must be free of leaves, twigs, and branches to minimize
drag when shooting. Preparation to shoot required the throwbag be positioned in the sling and the gripper
attached to the sling pulled taut (Fig. 7B). To prevent misfire, the throwbag, throwline and O-ring must be
carefully positioned in-line. There was a potential risk of backfiring and hitting the shooter in the face if
the Big Shot, throw bag, and throwline were not properly aligned.
One way to ensure success when shooting the throwline and throwbag over the limb was to focus on
the target and visualize the throwbag moving over the crotch. Shooting the Big Shot can be done two ways:
single-hand method or double-hand method. The single-hand method is best for individuals with strong
upper-body strength and allows maximum accuracy by stabilizing the pole with the free hand. The double-
hand method can achieve higher shots than the single-hand method but compromises accuracy with both
hands being used to pull the sling (Fig. 7B).
Even when the throwbag was successfully shot over the desired crotch of the limb, it does not always
reach the ground due to friction caused by rubbing against bark, limbs, vines, and leaves. To correct for this,
a variety of techniques were used to lower the throwbag to the ground. The first technique was to whip the
throwline up and down in a po Su b d This was usually effective when the throwline was
stuck in a tree *v" crotch as it lifts the tl th tch, allowing the throwbag to descend. Another
method was called the "strumming" technique (Fig. 7C,D). This method was akin to drawing a bow and
consisted of pulling the throwline towards the torso with two fingers while keeping the line outstretched
Kilgore et al. Ti
Pd y
1323
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1324 Journal of the Botanical R h Institute of Texas 2(2)
with the other hand. By quickly releasing the throwline with the two fingers, a wave of vibrations was sent
along the throwline causing it to move toward the ground. When the throwbag reached the ground, the
throwline was moved as close to the trunk as possible by using a circular, whip-like arm movement.
Once the throwline was set, it was tied onto the larger climbing rope using a series of half hitches (Fig.
6J) and used to pull the climbing rope over the desired limb. A limb was strongest where it was attached
to the tree trunk and therefore it was essential that the climbing rope be positioned as close to the trunk
as possible. A whip-like motion was used to move the climbing rope over the limb and next to the trunk.
Once the rope was installed, the climber prepared to ascend. Critical supervision by the ground crew was
essential as the climber tested the safety of the limb by hanging and bouncing using full body weight with
the climbing rope.
Climbing ropes were tied to a climbing saddle using an Anchor Hitch knot or attached using the eye
splice to a special D-ring in a process called “tying in" (Jepson 2000) (Fig. 6A—C,G-I; 7E). The working end
of the climbing rope was the end attached to the climbing saddle. For right-handed climbers, the working
end of the rope was attached to the D-ring on the left side of the harness and became shorter as the climber
advanced higher. The running end of the climbing rope was the end that the climber pulled down in order
to advance into the tree canopy and became longer as the climber advanced higher. It ran along the right
side of the climber's body and was attached to the saddle via a smaller rope, usually around 1.5 m in length,
called a "split tail” (Fig. 5A; 6A-C; 7E). The split tail was attached to another D-ring using the eye splice or
tied using an Anchor Hitch (on the right side of the saddle for right-handed climbers) and tied to the run-
ning end of the rope with a Blake's Hitch knot (Fig. 6G-D.
The Blake's Hitch knot was moved when the climber pulled down on the running end of the rope with
the right hand and used the left hand underneath the knot and pushed up, sliding the slack split-tail upward.
The climber then used his/her body to facilitate pushing up on the Blake's Hitch knot through a rhythmic
thrusting motion of the hips or by bouncing his/her feet off of the tree trunk (Fig.7 G,H). Thrusting and
bouncing motions lowered the tension on the climbing rope so that the Blake's Hitch was moved up the
slack rope. When the climber was not advancing, the weight of his/her body caused the Blake's Hitch knot
to tighten so that it did not slide.
Females on the tree canopy climbing team used a special foot loop technique that allowed them to
use their lower body strength to help pull down on the running end of the climbing rope while pushing
up on the Blake's Hitch knot (7F,H). The running end of the rope was curved into a loop and held in the
climber's right hand. After the climber pushed the foot loop down with either foot and pulled the running
end of the rope down with her right hand, she pushed up on the Blake's Hitch with her left hand. This foot
loop procedure is described in greater detail in Kilgore (2008); other foot loop techniques are described in
Jepson (2000).
A safety rope, also called a lanyard, was used to secure the climber to the trunk or around a branch
while he/she was sampling bark or when the working end of the climbing rope is detached from the climbing
saddle while the climber was advancing higher in the canopy. To “safety in” at the point of rope installa-
tion, the climber installed a short lanyard (more than two meters long), over a branch or around the trunk
for an additional safety measure or while maneuvering (Fig. 7K). The lanyard was fastened onto the front
attachment point or onto D-rings located on the side of the saddle. The lanyard length was easily adjusted
using a mechanical locking system. Lanyards were frequently used in conjunction with the climbing rope to
displace pressure points of the climbing saddle, to reduce swinging during ascension, or to pull the climber
into reach of grapevines.
Advancement from lower limbs to upper limbs in the canopy required the climber to unhook the
working end of the rope from the D-ring, lengthen the working end, and tie a special Monkey Fist knot in
the working end (Fig. 6K). Another way was to throw the throwline and throwbag over a limb higher in the
canopy much like you would from ground level and then attach the throwline to the working end of the
climbing rope using a series of half hitches. Once the climber had thrown the Monkey Fist knot over the
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1326 J lof the Botanical R h Institute of Texas 2(2)
desired limb higher up in the canopy, or attached the climbing rope to the throwline and hauled it over the
limb, the working end of climbing rope was reattached to the saddle.
The DRCM allowed climbers the freedom to use both hands to reach and sample tree bark in a vertical
transect from near ground level to near the apex of a tree (Fig. 7J,K) It was important to remember how
high the rope was secured in the tree, especially after advancing into the upper canopy. If the climber was
higher than half the length of the rope, the climber ran out of rope during descent. When this happened,
the climber stopped at one of the lower limbs, re-attached the rope to a lower limb and descended again.
Descending from the canopy required the climber to slowly pull down on the Blake's hitch knot while hold-
ing onto the running end of the climbing rope to control speed (Fig. 71). It was important to control speed
while descending so that the rope and gloves were not burned due to friction in too rapid a descent.
Bark collection at 3 m increments up to 35 m took three hours or longer, depending on the difficulty of
removing bark, maneuvering around limbs, difficulty reaching grapevines, and advancing higher to reach
leaf voucher specimens from the outer canopy. Although the saddle was padded, extended periods of time
hanging in the saddle caused pressure points to develop and legs to go numb. Numb legs were revitalized by
inverting the body or walking up the trunk and leaning back so the body was positioned with the legs above
the head. The climber could also stand on a limb or lie back on a limb to reduce pressure on the legs.
The DRCM was most efficient for sampling organisms from the trunk of the tree or from the parts of
the limbs closest to the trunk. However, samples may need to be collected from the outer limb and requires
the climbers to use a method called *limb walking" (Fig. 7L). This required the climber to advance to the
apex of the canopy and secure the rope around the trunk of the tree and at least one limb for stability. The
climber then determined the straigl path through the canopy to the target limb below. Once the climber
descended to the target limb, the rope was tightened so the climber could stand on the limb. Facing the
trunk of the tree, the climber proceeded to walk backwards while holding onto the rope and gradually slid
the Blake's Hitch down the rope to keep tension on the rope. When the climber reached the desired distance
from the trunk the targeted aerial structures near the outer canopy were sampled. To return to the trunk,
the climber stood on the limb and slid the Blake's Hitch up the rope maintaining rope tension.
Student field research experiences and activities
This project required team effort (Fig. 8A—J). The “ground crew" included multidisciplinary experts who
served as mentors for the students. These experts gave special lectures, slide shows, and field demonstra-
tions during evenings or on rainy days on the targeted groups of organisms to aid the student climbers in
the recognition and collection of specimens and suitable bark samples. Each evening, students sorted bark
samples, separating mosses, liverworts, and lichens and prepared voucher herbarium specimens for the
tree species (Fig. 8A). Other evening group activities that helped break the work routine and keep morale
high were playing cards, baseball, Frisbee, yard darts, and inviting park personnel and friends for a group
supper cooked by the students.
Field research was en ided into 2 three-week sessions nim. the pu from June to a Taai
climbed for six consecutive days with one day of free time.
at night with a flashlight with a focused spotlight that enabled students to collect 2 DE: on niche un-
derside of decaying logs. Tiny myxomycete sporangia in various stages of development glisten and become
more conspicuous at night when a directed light beam highlights areas of the log difficult to observe in
daylight hours. This was especially true for extensive fruitings of different species of Cribraria and Echinos-
telium minutum de Bary on decaying conifer logs (Keller 2004a). These night-time forays with flashlights
led to the first observations of slugs feeding on the immature fruiting body stages of myxomycetes (Keller
& Snell 2002b). Trail excursions sometimes led to unusual observations such as a millipede feeding on an
immature stage of a myxomycete fruiting body (Fig. 8B).
One of the most remarkable discoveries was a new species of myxomycete, Diachea arboricola H.W.
Keller & M. Skrabal. Melissa Skrabal observed the myxomycete plasmodial tracks and beautiful iridescent
sporangia from 9-24 meters on the bark surface and fissures of a white oak tree (Keller & Skrabal 2002;
Kilaore et al. T
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1328 Journal of f Texas 2(2)
Keller et al. 2002; Keller et al. 2004) (Fig. 8 C,D). This was an example of a woman in science doing field-
work that required strength, agility, and athleticism. She represents a role model for other young women to
follow in her footsteps. She also designed and sketched our tree canopy biodiversity logo which was made
into a cloth patch to provide research team members, volunteers, park personnel and interns, reporters, and
friends with a memento of our tree canopy biodiversity research project. The logo recognizes the support
of NSF and BS&I and our home institution Central Missouri State University (now UCM). More than 100
of these patches were distributed and are posted on bulletin boards, worn on blazers or jackets, or to help
identify gear bags or backpacks (Fig. 8D. A book entitled Fungi introduces a new series of Ranger Rick Books
for children in grades 3—5. Including a section entitled Exploring for Fungi that features Melissa Skrabal col-
lecting lichens and Diachea arboricola high in the tree tops in CMT (Carson 2003).
The NSF-Research Experience for Teachers Program facili ional devel pmer ent of ae E
on the cutting edge of science through partnerships ca local KE districts and universities (I
al. 2005). Trish Smith, a Warrensburg Middle School seventh grade life science teacher, along with students
and faculty from UCM, participated in a summer tree canopy Mo m in the GSMNP (Fig. 8F).
The project created a website at http:// burg.k12.mo Piadventure/ where the
field Adventure Phase "Exploring Life in the Forest Canopy,” represented the first tier of the ¡Adventure
website. This innovative wel bled students and teachers to experience tree py research and learn
about the All Taxa Biodiversity Inventory as part of the Adventure Phase (Fig. 8E). This was followed by the
Laboratory Phase where students observed moist chamber cultures with wet bark that enabled students to
observe a living miniature ecosystem composed of myxomycetes, fungi, lichens, mosses, liverworts, green
algae, cyanobacterial algae, myxobacteria, tardigrades, insects, nematodes, and possibly other invertebrates
(Fig. 8G). The students found several rare myxomycete species such as Echinostelium arboreum H.W. Keller
& T.E. Brooks, known only from a few locations in the world.
The second tier of the website at http://warrensburg.k12.mo.us/iad ^whatis.html was aniAdven-
" +] lxi is Quid te d i 1 +] 1 1 and mmtenm fa mntentorich ectarvline 1
I g y the direction y , using
ilable on the Internet, including that provided real-world data and primary literature
sources. This activity was designed to help students discover how to use and access data and information
on the Internet, and to solve problems and make choices. Students were expected to develop their own
research questions, design their own experiment or investigation using the specimens and collected data.
This subsequently led them to the Publication Phase, where they were expected to create poster presenta-
tions shared with parents and the school community. These classroom activities and website experiences
encouraged secondary students to choose field biology as their future career (Smith & Keller 2004).
Student Research Special Recognition
Kenneth (Kenny) L. Snell was a graduate of the first tree climbing school and served as an instructor in later
years (Fig. 6K, 7E). His leadership and mentorship in the field was a valuable resource for future student
climbers. He was the project leader beginning the summer of 2000 and created the tree and myxomycete
database for future research projects. His master’s thesis (Snell 2002) resulted in the first tree canopy paper
published on Myxomycetes (Snell & Keller 2003). Discover Life in America highlighted his accomplish-
ments at UCM in the lead article published in the All Taxa Biodiveristy Inventory Quarterly (Keller 2002b).
Kenny received two UCM awards, the Reid Hemphill O ling Scholar for his scholarship, research, and
citizenship; and the university-wide Graduate Thesis Award for the best graduate thesis Te the year 2002.
Erica E. Parker was a McNair Scholar Award Recipient at UCM. This program prepares first-generation,
low-income, undergraduate college students for doctoral study. Approximately 24 students were selected
as eligible McCAP participants. She was awarded first place for the best research paper (Parker & Keller
2003)
Angela R. Scarborough was selected as a McNair Scholar that resulted in her being featured on the
front cover of the McNair Journal (Scarborough 2005) (Fig. 8H); and her research paper presented at the
h and student i 1329
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Kilgore et al.. TI
3 7
Association of Southeastern Biologists (ASB) 2006 Annual Meeting won the Outstanding Microbiology
Award (Scarborough 2006a) and her poster won the Elsie Quarterman-Catherine Keever Award for the best
ecological poster (Scarborough 2006b).
Sydney E. Everhart, a UCM graduate student, won the Outstanding Microbiology Award for the best
paper presentation at the ASB 2007 Annual Meeting (Everhart et al. 20072), and also the Elsie Quarterman-
Catherine Keever Award for the best ecological poster (Everhart et al. 2007b). Her master's thesis (Everhart
2007) submitted to UCM was recognized by the University Research Council as the best graduate student
thesis in 2008.
Courtney M. Kilgore received the Nahm Award for the Outstanding Graduate Student from the UCM
College of Science and be nd The selection criteria included a major in a graduate program within the
College and d in leadership, scholarship, and citizenship. She also was the winner
of the Mycological Society of America official conference t-shirt design for the annual 2008 meeting held
at Pennsylvania State University. Four edible, mushroom cultivars were included in the winning design,
Shiitake (Lentinula edodes), Hen of the Woods (Grifola frondosa), button mushroom and Portabella white
and brown variety (Agaricus bisporus), and the velvet foot mushroom (Flammulina velutipes), and five molds
around the circular border: Aiternaria, Aspergillus, Fusarium, Penicillium, and Trichoderma. The design style
was reminiscent of early taxonomical drawings and scientific illustration with a classic art nouveau border
to frame the design.
Special Media Highlights
One of the objectives of this tree canopy project was to involve volunteers, park interns, undergraduate
and graduate students, in interpretive exhibits, news media coverage (print and television), publication of
technical articles in peer reviewed journals and also in popular magazines that sent a powerful conserva-
tion message for biodiversity. The grants acknowledged here for the period 2000 to 2008 resulted in 20
refereed papers in scientific journals, 15 articles published in newsletters, mostly in the Mycological Society
of America newsletter “Inoculum,” the “ATBI Quarterly" from Discover Life in America, and "What's Up” from
the International Canopy Network. The majority of these articles were on the front cover as the lead article.
Abstracts (74) were represented by power point and poster given at many different professional
meetings. Four master’s degree theses were completed during this period.
The media coverage gave the public a better understanding of the occurrence and importance of mostly
cryptogams such as myxomycetes, macrofungi, mosses, liverworts, lichens, and ferns and observations on
invertebrates, including insects, mollusks, and tardigrades. Media attention highlighted the DRCM and re-
porters interviewed students and project team members. Interpretive exhibits, newspaper articles, popular
books for children, and television feature stories based on our tree canopy exploration and discoveries in
the GSMNP were described by Keller (2004a).
Rock Creek National Park, located in Washington D.C., hosted the first 24-hour BioBlitz held May 18-19,
2007, jointly sponsored by the National Geographic Society (NGS) and the National Park Service (NPS);
Fig. 8J). Student climbers demonstrated the use of the DRCM when sampling bark for myxomycetes. The
BioBlitz which began in 2007, will be held annually in urban areas of NPS units for the next 10 years with
the goal of increasing public awareness through the documentation of species inventory, public outreach
activities, and science education for all age groups (Kilgore 2007; Kilgore & Keller 2007). This first BioBlitz
drew media coverage that resulted in a photo of Sydney E. Everhart on the front page of the Metro section
of the May 19", 2007 issue of the Washington Post. Perched in her climbing saddle in the top of a white oak
tree canopy, she collected bark samples that were later transported to the laboratory and cultured in moist
chambers for myxomycetes and other organisms. Video of this event is available for viewing at this URL
address: http://video.nationalgeographic.com/video/player/specials/films-specials/, The NGS video page will
load and the featured video will begin to play. In the dialogue box that says “search all videos” enter “Rock
Creek Bioblitz” and press “GO.”
1330 J | of the Botanical R h Institute of Texas 2(2)
More recently, National Geographic Television produced two films “BioBlitz Rock Creek Park 2007"
and “Smoky Mountains Treetop Exploration," that appeared as part of the Wild Chronicles series on Public
Broadcasting Stations (PBS) nationwide. The former as Episode #236, August, 2007 and the latter as Episode
#318, February, 2008, and had a running time of seven minutes for each episode. Boyd Matson was the
host and narrator. National Geographic Society Mission Programs supports pioneering research and field
expeditions through programs such as the Committee for Research and Exploration.
During July of 2007, National Geographic Television Producer Jason Orfanon shot 10 hours of film
footage over a five-day period in the GSMNP. The storyline documented the exploration of the tree canopy
using the DRCM by a research team of two University of Central Missouri graduate student climbers, SEE
and CMK, who demonstrated how to access, climb, and gather bark samples with myxomycetes, macrofungi,
lichens, mosses, and liverworts from the tree canopy. Harold W. Keller coordinated the ground crew and
served as the principal investigator for the research project entitled “RUI: Biodiversity and Ecology of Tree
Canopy Biota in the Great a e opos ud d is available for viewing at this URL ad-
dress: http://video.nati specials/. The NGS video page will
load and the featured video will MM to play. In A daek box that says “search all videos” enter “Smoky
Mountain Tree Canopy” and press “GO.” Films will appear as blocks. Click the second block that is called
“In Search of Slime” which has a beautiful color image of a sessile, densely netted, plasmodiocarpous fruit-
ing body with a conspicuous bluish iridescence of the rare myxomycete Didymium perforatum Yamash.
In addition to the video, Matson al nducted a 20-minute interview with CMK and HWK about their
tree canopy adventures, which aired on National Geographic Weekend. This is a new radio program hosted by
Boyd Matson that highlights stories of “exploration to the far corners of the planet and the hidden corners
of your own backyard.” It airs on radio stations on Saturdays and Sundays.
RESULTS
Tree and Vine Species Climbed and Sampled
Tree and vine species represented 52 taxa and more than 500 individual trees were climbed over the eight
year period of this project. Some species of trees are represented by many individual trees, for example,
Acer rubrum, A. saccharum, Fraxinus americana, Juniperus virginiana, Liquidambar styraciflua, Liriodendron
tulipifera, Picea rubens, Pinus echinata, P. strobus, Platanus occidentalis, Tsuga canadensis, and grapevines, Vitis
aestivalis, and V. vulpina. Some of the tree species were scattered, fewer in number, and difficult to find,
but the species given here occurred in greater numbers, were closer together, easier to locate and climb,
and larger. These trees met our climbing criteria for bark sampling. Trees were located using trail guides,
vegetation maps, consultation with local residents, park and state officials, and questioning hikers on the
trail (Keller 2006).
Tree and vine species listed alphabetically: Abies fraseri (Pursh) Poir., Acer negundo L., A. rubrum L.,
A. saccharinum L., A. saccharum Marsh., Aesculus flava Aiton, A. octandra Marsh, Ampelopsis cordata Michx.,
Betula alleghaniensis Britton. B. lenta L., Carya alba (L.) Nutt., C. cordiformis (Wangenh.) K. Koch., C. glabra
(Mill) Sweet, C. illinoinensis (Wangenh.) K. Koch., Cercis canadensis L., Cornus florida L., Diospyros virgi-
niana L., Fagus grandifolia Ehrh., Fraxinus americana L., F. pennsylvanica Marsh., F. profunda (Bush) Bush,
Halesia carolina L., Juglans nigra L., Juniperus virginiana L., Liquidambar styraciflua L., Liriodendron tulipifera
L., Magnolia acuminata (L.) L., Nyssa sylvatica Marsh., Picea rubens Sarg., Pinus echinata Mill., P. strobus L.,
Platanus occidentalis L., Populus deltoides Bartram ex Marsh., Prunus serotina Ehrh., Quercus alba L., Q. falcata
Michx., Q. michauxii Nutt., Q. montana Wild., Q. muehlenbergii Engelm., Q. prinus L., Q. rubra L., Q. velutina
Lam., Robinia pseudoacacia L., Sorbus americana Marsh., Taxodium distichum (L.) Rich, Tilia americana L., T.
heterophylla Vent., Tsuga canadensis (L.) Carr., Ulmus americana L., U. rubra Muhl., Vitis aestivalis Michx., and
V. vulpina L
Myxomycete Species on Trees and Vines
Corticolous myxomycetes complete their entire life cycle only on the bark of living trees and vines (Keller
Kilgore et al., Ti h and student i 1331
re LI
& Braun 1999; Everhart & Keller 2008). Some corticolous myxomycete species were collected in the field
directly on the bark p ne trees and vines and some were harvested from laboratory moist chamber cul-
tures. Bark cl d pH were important factors in th d distribution of
on living, healthy trees and vines (Snell & Keller 2003; Keller 2004a; dd & Keller d Scarborough
2006a; Keller & Everhart 2008; Everhart et al. 2008). The more prod t for i
species are paced Nere with the total number of species find to date. These numbers were pia
from previ (Martin & Alexopoulos 1969; Keller & Braun 1999; Parker & Keller 2003; Parker
& Keller 2005; Scarborough 2005; Bcaiboroue t 2006b; Snell & Keller 2003; Snell et al. 2003; Everhart &
Keller 2008; Everhart et al. 2008; Kilgore 2008). The number of different myxomycete species on trees and
vines are listed from highest to lowest, with number of species in parentheses: Juniperus virginiana (54), Acer
rubrum (49), Quercus alba (43), Liriodendron tulipifera (41), Vitis aestivalis (39), Fraxinus americana (31), Vitis
vulpina (25), Pinus strobus (24), Ulmus americana (20), Acer saccharum (17), Tsuga canadensis (17), and Pinus
echinata (14). All other trees had fewer species than listed here and Abies fraseri had no myxomycete species
above diameter at breast height.
The mean pH is given for the above listed tree species from the highest pH to the lowest. Juniperus vir-
giniana had the highest pH (7.3) bark values, highest water absorption capacity, and the highest corticolous
myxomycete species diversity based on our field and laboratory tree canopy studies to date (Keller & Braun
1999; Keller 2004a). Ulmus americana (7.0) and Fraxinus americana (6.7) also had thick bark with high wa-
ter absorption capacity (Parker & Keller 2003). Quercus alba (5.7), Vitis vulpina (5.5), Acer saccharum (5.5),
Liriodendron tulipifera (4.9), Vitis aestivalis (4.8), Acer rubrum (4.7), Tsuga canadensis (4.1), Pinus strobus (3.8),
and Pinus echinata (3.8) represent the rest of the tree and vine species (Snell & Keller 2003; Everhart et al.
2008; Kilgore 2008). The last three tree species had the lowest pH and lower number of my ycete species
in part due to the lower pH and lower water absorbing capacity. This trend of a lower pH range associated
with especially resiniferous gymnosperm tree bark, resulted in the lowest number of myxomycete species.
FUTURE DIRECTIONS
The results and scope of this project demonstrate that the DRCM is an alternative way to study tree canopy
biota. Nevertheless, additional research is needed to answer questi bout organisms in hard to reach places.
For example, limb walking using the DRCM facilitates sampling from the trunk axis to the outer periphery
of the tree. Studies using DRCM and limb walking are capable of sampling aerial reproductive structures,
such as gymnosperm female cones (Pinus spp.), pods from Cercis canadensis, ball-like fruiting structures of
Liquidambar styraciflua and the long pods of Catalpa speciosa (Warder) Warder ex Engelm. (Kilgore 2008).
Further research in the tree canopy is needed to document the vertical distribution of lichen growth
forms (crustose, foliose, and fructicose) and lichen species among these three growth habits from the tree
base at ground level to the treetop (Ciegler et al. 2003; Lumbsch et al. 2005; Fanning et al. 2007; Keller et
al. 2007).
Little i g
tree trunks. Most of the species found to date in the tree canopy also occur on ground sites. This is another
group that should be targeted for more tree canopy survey and inventory (Davison & Keller 2004).
The presence of basidiomycetes and ascomycetes on the bark surface and in tree crotches should be
targeted for collection and identification. No ascomycetes were collected and identified in our bark cultures,
but they were present nevertheless. Only five basidiomycetes were collected on healthy living trees-certainly
a scanty harvest.
The total height and diameter of the tree should be supplemented with core samples to determine the
actual age by counting tree rings. The actual age of the tree is lacking in most studies and this piece of data
would match size and age of trees and the time needed for organisms to colonize the bark surface. This
invasive technique would require special permission from the National Park Service or the United States
Forest Service, or be conducted on private property.
| bout the distributi f mosses and li (I l JÉ vertical l
1332 | of the Botanical R h Institute of Texas 2(2)
Bark samples were prepared in moist chamber cul i liately or within several months after field
collection. Laboratory p ls should include exactly how old the bark samples were when first wetted in
moist chamber cultures. Storage at room temperature in sample bags enclosed inside of a large paper bag
and tightly sealed for 12 to 24 months or longer increases the chances of filamentous mold contaminants,
especially species of Trichoderma, mucoraceous species, Aspergillus species, and unidentified white molds.
Myxobacteria appear quite frequently in moist chamber cultures of bark from living trees and vines
and represent a group of understudied organisms with potentially unidentified species which are strictly
arboreal. Bark and grapevine samples from the tree canopy also have nematodes and tardigrades, and these
organisms mostly are known from ground level sites. The role of nematode interactions with myxomycete
plasmodia needs further study based on recent results (Kilgore & Keller 2008).
Observation and collection of snails and slugs along the tree k was possible using the DRCM. Slugs,
Philomycus carolinianus (Bosc) and P. flexuolaris Rafinesque, were documented at heights up to 14 m on the
trunk and around a treehole filled with water (Keller & Snell, 2002b). Snails with shells, for example, Me-
sodon normalis (Pilsbry) about the size of a quarter, and Anguispira jessica Kutchka about the size of a dime,
appeared to move from ground level up to 15 m in the former and 24 m in the latter. The DRCM is the best
way to record slug and snail movements in the tree canopy and determine if any species are arboreal. Little
is known about tree canopy slugs and snails (Thomas Watters and Dan Dourson, pers. comm.)
Epiphytic plants in the tree canopy should also be studied. Pleopeltis polypodioides (L.) Andrews & Wind-
ham (the resurrection fern), was found in large conspicuous clumps along the trunk and branches at lower
heights on a national champion tree, Fraxinus profunda (Bush) Bush in Big Oak Tree State Park, Missouri
(Keller pers. obs.). This fern was observed i with the me dd and Mn Pre may be seen with
ground-based binoculars higher in the canopy. 1 tes el n the sharp-
est eyes and binoculars, and only the DRCM made it dures to see these ms higher in the tree canopy.
The typically lithophilic Polypodium appalachianum Haufler & Windham (Rock Cap Fern) was discovered
as a tree canopy epiphyte 35-40 m above ground on a horizontal branch in a champion-sized Liriodendron
tulipfera in the GSMNP. Even though this fern had leaves up to 16 cm long and 5 cm wide, it could not be
seen with ground-based binoculars because of the height and location on the upper side of a horizontal
branch (Keller et al. 2003). Tiny epiphytic plants that include orchids and other vascular plants may represent
arboreal species that can be detected only by using the DRCM. Vertical transects of the bark surface can be
scanned from the base to the top of the tree canopy visually or using a 20x hand lens for tiny organisms.
Certain corticolous myxomycetes only are known from the tree canopy such as Trabroohsia applanata
H.W. Keller. This myxomycete species and many others should be cultured from spore to spore and DNA
analysis determined to correctly classify and develop phylogenetic relationships instead of morphospecies
concepts (Keller 1996; Keller 2005c; Keller & Everhart 2008).
Many more studies using rope climbing methods are needed to determine if arboreal biota exist in the
tree canopy of temperate forests. Research projects using the DRCM will help the next generation of tree
canopy biologists to explore, ask questions, and develop hypotheses that will increase our knowledge of the
biosphere.
ACKNOWLEDGMENTS
Our sincerest gratitud to all ofthe student climbers who participated in this project. Faculty and student
colleagues in the Deraan of Biology and Earth Science at UCM provided encouragement and logistical
support that made this project possible. The success of this tree canopy project depended on numerous in-
dividuals, including local residents who knew where to find a champion tree or grapevine and professionals
who offered their enia in taxonomic identification and reviewing papers for publication.
Ricardo P. 1 the English abstract into Spanish. Keith Langdon and Paul Super from GSMNP
gave assistance in obtaining collecting permits, maps and directions where to find trees, and facilitated
permission for National Geographic to produce a television film about our tree canopy biodiversity research
Kilgore et al., T h and student i 1333
project. Jeanie Hilten assisted with equipment needs, logistics, park directions, and lodging at GSMNP as
part of Discover Life in America. Carolyn and Reid Franks generously offered their home and friendship to
us on numerous occasions while we conducted research in GSMNP. David Taylor, botanist for the USFS at
DBNF, assisted in obtaining collecting penu lodging facilities, and finding suitable tree climbing sites.
John Perry, Glen Dandeneau, and the at the Berea College of Forestry helped us locate tree
sites in the Berea College Forest. Tim Smith at the Missouri Department of Conservation and Mike Currier
at the Missouri Department of Natural Resources assisted with obtaining collecting permits for collecting
sites in Missouri.
Many individuals accompanied the tree canopy team on expeditions to GSMNP or DBNF, offered their
expertise in a particular field, and participated as ground crew members. These individuals included: Alex
Ciegler, Paul Davison, Uno Eliasson, Thomas Gaither, Diane Nelson, Kenneth Nelson, Jay Raveill, Trish and
Stan Smith, and Ted Stampfer. Stephen Wilson assisted with canopy flight-intercept insect traps in GSMNP
and Big Oak Tree State Park, served on graduate student thesis committees, edited theses, and reviewed
papers for publication.
Trish Smith involved seventh grade life science students at Warrensburg Middle School with the Re-
search Experience for Teachers project by culturing myxomycetes in the classroom, developed an interactive
website for students to participate in their own tree canopy adventure, and invited members of the UCM
tree canopy team to visit the classroom to help identify organisms in moist chamber cultures. Jean Gerrath
and Jay Raveill assisted students in identifying grapevine and trees vouchers respectively. Gabriel Moreno
produced valuable scanning electron micrographs of myxomycete species that had unusual characteristics.
Uno Eliasson aided in collecting and identifying myxomycete specimens that were new records or had
unusual characteristics, and he reviewed papers for publication. Karl and Jeanne Braun and Ted Stampfer
reviewed numerous papers for publication.
Robert Breshears, Mike Ferro, Jim Murray, and Glenda Carmack volunteered their photographic skills
and accompanied team members into the field on numerous occasions to visually record the DRCM. Many
of their photographs appear in publications produced by the tree canopy team. Photo credits: James Murray
= Figs. 1A,B,F,G, 3H, 5A, 7E,J, 8A,C; Courtney M. Kilgore = Figs. 1C,K, 4C; Harold W. Keller = Figs. 1D—
E,H-7, 3G, 4D-E, 5B,D, 6J,L, 7F-LL, 8F—G,I; Glenda Carmack = Figs. 2A, 3B,D,F; 4A, 5C, 6A-F,K, 7A, 8E;
Jerry Coule = Fig. 2B-C; Mike Ferro = Fig. 3A,C,E; Kenny L. Snell = Fig. 4B; Robert Breshears = Figs. 6G—I,
7B—D,K, 8J; Stan Smith = Fig. 8B; Sydney E. Everhart = Fig. 8D; Bryan Tebbenkamp = Fig. 8H.
Lisa Schmidt and Terry McNeeley of Instructional Design at UCM contributed many hours of their
technical ti d support assisting with posters, power point presentations, web site construction, and
other media a profesional conferences. Mike Griefe highlighted our projects through University Relations
at UCM.
Nalini Nadkarni and Scott Hollis of the International Canopy Network regularly published articles in
the ICAN newsletter "What's UP”? Jinx Campbell, Current Editor of the Mycological Society of America's
newsletter “Inoculum” along with past editors Richard Baird, Donald Ruch, and Lorelei Norvell all featured
our tree canopy articles. Jason Orfanon, NGS television producer, spent time in the field with us in GSNNP
to film our research for the television series “Wild Chronicles.” Boyd Matson interviewed two of the authors
on the radio show “National Geographic Weekend.”
This project was financially supported in part by the National Science Foundation Division of Envi-
ronmental Biology Awards #0079058 and #0343447, Discover Life in America Awards #2001-26, #2002-17,
and #2004-6, National Geographic Society Committee for Research and Exploration Award #7272-02, the
Missouri Department of Natural Resources for the tree canopy research in Big Oak Tree State Park, Missouri,
the United States Department of Education McNair Scholars Program, and the UCM Summer Undergraduate
Research and Creative Projects Program.
The authors wish to thank H. Bruce Rinker, Lafayette Frederick, and Dennis L. Miller for their careful
and meticulous reviews of an earlier version of the manuscript.
1334 J lof the Botanical R h Institute of Texas 2(2)
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ARISTOLOCHIA SCHULTZEANA (ARISTOLOCHIACEAE) NEW TO ECUADOR
Douglas H. Goldman
Harvard iugis ue
22 Divinity Av
Cambridge, NOM is Dm USA
dgoldmangfas.harvard.edu
ABSTRACT
Aristolochia schultzeana is a rare, low-elevation, humid, pluvial forest species. It was previously known only from Colombia but is now
reported for Ecuador
RESUMEN
Aristolochia schult i le los bosques húmedos y pluviales de baja elevación. Previamente, esta esp
solamente de Colombia, pero ahora se reporta para Ecuador.
In July 2007, while I was teaching a neotropical plant diversity class for Arizona State University's Andes
and Amazon Field School, in the western part of the Amazonian region of Ecuador, a student produced a
strange, spectacular and relatively large Aristolochia flower from a forested area at the edge of a nearby vil-
lage. The flower was reminiscent of a Nepenthes or Sarracenia leaf, with an elongated floral tube terminated
with two limb lobes, a small lower (anterior) lobe and a large, erect and expanded upper (posterior) lobe
that was yellow with red-brown veins to the inside and with a margin of fleshy, forked, black fimbriae (Fig.
1). Unpleasantly scented, reminiscent of dead fish, I received the flower slightly dehydrated with the upper
limb lobe hanging over the mouth of the floral tube. After taking several high-resolution digital photographs
(300 dpi), the flower was placed in a plastic bag with some water, successfully rehydrating it and returning
the posterior limb lobe to a vertically-erect position. Thereafter, several more photos were taken, including
dissections and close-ups of the flower. Several days later I was taken to the actual plant in the field, which
was a large vine with cordate leaves, growing just inside the edge of a young forest. It was not flowering
or fruiting at the time. I was unable to obtain necessary collection and plant-export permits prior to my
visit to Ecuador, therefore no herbarium specimen was made of this plant. It was not until several months
later, using the photographs, that I identified this species as Aristolochia schultzeana, later confirmed by
Favio González at the Herbario Nacional Colombiano. Furthermore, while researching this identification
I learned this was apparently the first report of this species for Ecuador and only the third report for this
species overall, suggesting that this is a species of striking global rarity. While photographing this plant I
was unaware that I was documenting something of any significance, as my photos at the time were for the
benefit of my curiosity alone.
A
Voucher: ECUADOR. Provincia del N p d t . north facing slop
h rainf i l ] to put cut wood; ca. icd mS of thé well- traveled, paved iat near the S side of the Rio
e 1 r Fr 4
NI 1 SL Win) :]1 IT with in 1na TCI f Te ena; elev ca 440m;
ca. 0103: 01.9"S, 77°41'06.5"W; UTM 18M 201152, 9883757N (WGS84), 18 Jul 2007, D. Goldman 3990 (BH, COL, GH, QCA).
The voucher consists of a printed set of 28 digital color photographs of the one flower found of this species,
all but one photograph taken by myself, each printed on paper 25 cm x 18 cm, with scale indicated, all
bound as a unit and mounted or contained on a single herbarium sheet. In addition, this photo-set has been
deposited in the botany libraries at Harvard University (both printed and on a compact disk), and is also
available for viewing in Morphbank, an archival, web based, public-access database of images of biological
specimens for scientific study (www.morphbank.net).
Aristolochia schultzeana O.C. Schmidt is a humid-tropical rainforest species that previously had been
considered endemic to southern Colombia, where apparently it has been collected only twice (González
J, Bot. Res. Inst. Texas 2(2): 1337 — 1339. 2008
1 Laféha Dotaniz4l D h Institut. f Texas 2(2)
1338
Fic. 1. Fresh flower, July 18, 2007. Photo credit: Kenny Richards.
1990). Hohene (1942) mentioned this species was present in Bolivia, but he did not cite any Bolivian speci-
mens. The type locality is to the east side of the Andes in Colombia, near Umbría in the Departamento del
Putumayo, 0°54'N, 76°10'W, 325m altitude, Oct-Nov 1930, G. Klug 1786 (B [holotype, destroyed], BM, F, GH,
K, MICH, NY [lectotype, designated by González 1990: 125], S, US). The second collection from Colombia
Goldman, Aristolochia schult to Ecuador 1339
is relatively close to the Pacific coast, near Llorente, Tumaco, Departamento del Nariño, 8 Sep 1976, O. de
Benavides 628 (COL). See González (1990) for a summary of this species in Colombia. Because this species
is known, although sparingly, from within about 50 km of the Ecuadorean border, its discovery in Ecuador
has been anticipated for some time (González 1994, 1999). Although it could have been expected that this
species would first be found in Ecuador very close to the Colombian border, this newest record represents a
range extension or perhaps a disjunction, found a minimum of about 160 km from of the Colombian border
and approximately 270 km from nearest Colombian site for this species (in Putumayo). This suggests that
the species could be more widespread than expected in the region, and its occurrence adjacent to a well-
traveled, paved road suggests that it is more common than previously considered but simply overlooked, or
that it flowers infrequently.
Aristolochia schultzeana is similar to another, but more widespread, tropical rainforest species, A. didyma
S. Moore, found in scattered localities throughout northern South America (Foster 1958; González 1994;
Hoehne 1942; Macbride 1937). Both species have large, cordate leaves, lack pseudostipules, have flowers
of similar overall shape, fimbriate upper limb-lobe ornamentation, and floral coloration and patterning.
However, A. didyma has a syrinx (an internal, downward projection of the floral tube into the utricle) which
A. schultzeana lacks. Furthermore, the upper (posterior) limb lobe of A. didyma is deeply bifid apically with
a tiny mucro in the sinus, whereas A. schultzeana has an undivided upper limb lobe with an elongate (7-15
mm) apical mucro. See Hoehne (1942) and González (1994) for a full comparison and illustrations of these
two species.
Some local residents near the field site in Ecuador commented to me that this plant can be propagated
from cuttings. This suggests that people in Santa Urcu and other nearby Quichua communities may have
specific uses for this plant, although an exact use was not mentioned. Aristolochia spp. are commonly used
in Ecuador for digestive-related ailments or occasionally malaria (Ríos et al. 2007), particularly in the Ama-
zonian region.
ACKNOWLEDGMENTS
I am gratefui to Ricardo Licui, Kenny Richards and Tod Swanson for their assistance in locating and intro-
ducing me to this plant, Kenny Richards for his field photograph of the flower, Tod Swanson and The Andes
& Amazon Field School of Arizona State University for providing me the opportunity to visit this region,
Carla Gavilanes, Susan Poats, Rodrigo Sierra, Scott Walker & Elizabeth Wentz for valuable geographic
information, Favio González (COL) for confirming the identification of this species, Dan Fulop for help
with the Spanish resumen, Luis Mendonca de Carvalho for translation of portions of Hoehne (1942), Mario
Blanco and an anonymous reviewer for valuable comments on this manuscript, and Hobbes Goldman for
assistance throughout.
REFERENCES
FosTER, R.C. 1958. A catalogue of the ferns and flowering plants of Bolivia. Contr. Gray Herb. 184:1—223.
GoNzÁLEZ, F.A. 1990. Aristolochiaceae. In: J.O. Rangel, A. Cadena, G. Correal and R. Bernal, eds. Fl. Colombia
12:1-184.
GONZALEZ, F.A. 1994. Aristolochiaceae. In: G. Harling and L. Andersson, eds. Fl. Ecuador 51:1-42.
GONZALEZ, RA, 1999. Aristolochiaceae. In: PM. Jorgensen and S. León-Yánez, eds. Catalogue of the vascular plants
of Ecuador. Monogr. Syst. Bot. Missouri Bot. Gard. 75:255-256.
HotHNE, F.C. 1942, Aristolochiaceas. In: F.C. Hoehne, ed. Flora Brasilica 15(2). Instituto de Botánica, São Paulo, Brasil.
MACBRDDE, J.F. 1937. Aristolochiaceae, Flora of Peru. Field Mus. Nat. Hist., Bot. Ser. 13:431-443,
Ríos, M., H.B. PEpERsEN, MJ. Kozio, AND G. GRANDA. 2007. La colección etnobotánica del Herbario QCA. In: M. Ríos,
M.J. Koziol, H.B. Pedersen and G. Granda, eds. Plantas útiles del Ecuador. Aplicaciones, retos y perspectivas.
Ediciones Abya-Yala, Quito, Ecuador. Pp. 111—640.
1340 J | of the Botanical R h Institute of Texas 2(2)
BOOK REVIEW
STEPHEN S. WHITE AND MARSHA V. GALLAGHER (Eps.), with translation by WILLIAM J. Orr, PAUL SCHACH, AND DIETER
KancH. 2008. The North American Journals of Prince Maximilian of Wied: May 1832-April
1833. (ISBN-10: 0806138882; ISBN-13: 978-0806138886, hbk.). University of Oklahoma Press, 2800
Venture Drive, Norman, Oklahoma 73069-8218, U.S.A. (Orders: www.oupress.com, 800-627-7377).
$85.00, 544 pp., color and black and white illustrations, 12" x 8".
Made f. | Bod , the North A i 1 f list Pri Maximili
e] r o
of Wied in 1832-34 he £i ientific exploration of the Missouri River's vee? reaches since the epic journey of Lewis and Clark
almost ao years earlier. Maximilian’s j It until now. This up UU quality, oversized
volume, the first of a dn: set, nee on the Maximilian Peder C lesion at Joslyn Art M
ethnographer lex] I he A i Westinthe 1830s provide
]uable infi i bout the Plains Indi ime, Maximilian i fth ll state of Neuwied and served in the
Prussian army. H d k ex] in Brazil in 1815— Rd id in ads wn in 1832-34. On the latter journey, he took with
him the S Karl Bod it] I peor encountered. NA trav ni Min Boston SISSE along the
Ohio River to St. Louis, Missouri, from where they 1l ] he Mi i, Nebraska
South Dakota, North pd and Montana. The westernmost point they aus on their n was Fort McKenzie, a trading post
Maxi lan, Hidatsa, and other Indian tribes of the Missouri
E
in central Montana
River area, while Bod le! Ioda f 1 E Ad ings of
] g B hald: Cl : £411 1 21,1]
Many exhibitions and books have WE Josly d Karl
Bodmer. Less attention has been paid to Prince Qi ane manuscript journals, which de di collectively called Bs quinis
H1
Indians i ds encountere
Y J
These dich each containing about 300 pages filled , places, flora, fauna, and events,
ritten bv him in a now obsolete German script and ülisqated with his ink and watercolor anos are the subject of a complex,
ira ear trees project being carried out by Joslyn's Durham Center for Western Studies in partnership with the University of
Oklahoma Press
lume 1 covers the bns of the expedition, commencing with a stormy Maia across the e on eee with the
tier t f St. Louis. Maxi primary goal had always been the Western wild is
journey there: the belated delivery of essential supplies, wariness of a cholera epidemic affecting chosen routes of travel, and iis own
illness that required a En dalam in New oe n In the mun id in the east, Bodmer oie m in
A
his in as ud. £ g Į purty
and p king the frontier. His exquisit positi ffer a fascinating window on a brash new nation, from its
] y M +1 " 1 ] ; ; f. £Tad: 3 : :
g 8 p Illinois
Maximilian’ 1 Bod "s pi f invaluabl dress, culture, and f l Indian
tribes that subsequently almost demei owing to disease, war, and white akan t on their lands. Maximilian wrote up his
observations in Reise in das innere Nord-America in den Jahren 1832 bis 1834, 2 vol. (1839—41) (Travels in the Interior of North America). An
Tz Tiat 1 E £ + Fata MES field j 1 I 2131: à D, pl of th pen MA Lift A 1 Di; T. Their Final Days of
8
Glory (1976).
The North Ámeri anj qe 1 ff, 1 bl Y iew ftt Mi 1 li ] 1 ] 1 1
it i 1 tati faithfully reproduces
rr
history of the oe West. n meet account, newly translated
s 11 asides, and appendices. This first volume, which covers May
Maximilian's 110 vatercolors as
n to April Bis documents Maximilian’ voyage to North America and his first encounters with Indians upon reaching the West.
y western American dd m a a of mns value. I am looking forward with
It i I d fauna and a general index
great anticipation to E oe volumes g
Recommended for all | ] of the ENVÉS or Western art.—Gary L. Jennings, Botanical
Research Institute of Texas, 500 East 4th Street, Fort Worth, Texas 76102-4025, US. A.
J. Bot. Res. Inst. Texas 2(2): 1340. 2008
ASCLEPIADOIDEAE (APOCYNACEAE) COMO MALEZA EN EL CULTIVO DEL
AGAVE TEQUILERO (AGAVE TEQUILANA VAR. “AZUL”)
Irma López-Muraira Rubén Iruegas
Instituto Tecnológico de Tlajomulco, Jalisco, Km. 10 Dupont, S.A. DE CV.
Carr. Tlajomulco San Miguel Cuyutlán, Tlajomulco de Zúñiga Homero No. 206 Col. Chapultepec-Morales
Jalisco, 0) MÉXICO, DF. CP. 11570
lopezmurairaghotmail.com ruben.iruegas-1Emex.dupont.com
Juan Florencio Gómez Leyva
Instituto Tecnológico de Tlajomulco Instituto Tecnológico de Tlajomulco
Jalisco, Km. 10 Carr. Tlajomulco San osi Cuyutlán Jalisco, Km. 10 Carr. Tlajomulco San Miguel Cuyutlán
Tlajomulco de Zúñiga, Jalisco, MEXICO Tlajomulco de Zúñiga, Jalisco, MÉXICO
ee o jfgleyva@gmail.com
RESUMEN
Identificamos 15 pecies de herba del bfamilia Asclepiadoid 51 3 ho : Me z
azul (Agave tequilana F.A.C. Weber var. “azul”) en el Estado de Jalisco, México. D le | peci hábi pad pi
el despliegue natural de las Deas del due lo que retrasa su crecimiento. Así mi t lto endemi 11 especies
exclusivas a México familia Asclepiadoid Itivo de agave tequil
F
ABSTRACT
We identify 15 species of herbs in the subfamily Bi onion jie in 65 SE points D fields cultivated with pau agave oe
ee i onde F. E C. nin ar. "azul" Us the state oe Mé I bing habi
g Likewise, we find a high endemi th 11 of the wee e restricted to
Mexico. This is the first study abo weedy | f the subfamily Asclepiad found g g I g p
INTRODUCCION
El Tequila es originario de Jalisco y su producción esta basada en el cultivo del agave azul (Agave tequilana
F.A.C. Weber var. “azul”), de donde se obtiene la materia prima para la elaboración de esta bebida. Como otros
cultivos, el agave puede ver mermado su rendimiento por la acción de varios factores bióticos como plagas,
enfermedades causadas por microorganismos y en especial por la interferencia ocasionada por maleza.
El agave por ser un cultivo semiperenne y de crecimiento lento, cuyo ciclo de producción va de seis a
nueve años, es más propenso a la invasión por la maleza que otros cultivos; especialmente en los primeros
cuatro años de crecimiento. Además, al no estar bajo un sistema de irrigación sufre “stress” hídrico durante
los meses de estiaje (Salamanca 2006).
Las plantas invasivas que afectan la producción del agave pueden ser de 3 tipos: a) aquellas que surgen
durante el temporal de lluvias y que están caracterizadas por hierbas anuales de diversas familias; b) las que
germinan durante la época de otoño-invierno y, c) la maleza perenne, que es la que compite durante más
tiempo con el cultivo de agave. La maleza perenne esta compuesta por plantas de especies leñosas, algunas
de las cuales son de hábito trepador.
La subfamilia Asclepiadoideae son un buen ejemplo de la exitosa adaptación de vegetales trepadores,
ya que en la búsqueda de los requerimientos de agua y nutrientes no solo interfieren en el cultivo compit-
iendo por estos requerimientos; sino también por luz y espacio, ya que al desarrollar un rápido crecimiento,
acompañado del engrosamiento de sus tallos impiden el natural despliegue de las hojas del agave, logrando
así retrasar su crecimiento y normal desarrollo (López-Muraira et al. 2007).
Debido al desconocimiento de las plantas invasivas en el cultivo del agave, es importante conocer la
diversidad de estas comunidades con la finalidad de entender su estado actual. Este trabajo reporta la pres-
J. Bot. Res. Inst, Texas 2(2): 1341 — 1345. 2008
1342 J | of the Botanical R h Institute of Texas 2(2)
encia de las especies de la subfamilia Asclepiadoideae que actúan como malezas en el cultivo del agave en
el estado de Jalisco, México.
REVISION DE LITERATURA
La definición más aceptada de maleza o mala hierba es “planta que no se desea tener en un lugar y tiempo
determinado,” se considera a la maleza como plantas silvestres, no cultivadas, que invaden campos de cul-
tivo o que viven a la orilla de los caminos (Rojas 1984) y en el agave puede interferir en los rendimientos,
y ya seca, sigue constituyendo un riesgo al acentuar las heladas y como perdida total del agave en caso de
incendios (Luna 2003).
Varias especies de Asclepiadoideae conocidas como lechosas (milkweed) han sido registradas como
malezas en diversos cultivos. Así, Morrenia odorata Lindl. ha sido reportada por Futch & hall (2003) para
cítricos en Florida y Ampelamus albidus (Nutt.) Britton para los cultivos de maíz y soya en Virgina, USA por
Bradley & Hagood (2001). En Australia, Grice & Martin (2005) reportan a Calotropis procera (Aiton) WT.
Aiton y C. gigantea (L.) W.T. Aiton como maleza en pastizales y Weston et al. (2005), mencionan a Vincetoxi-
cum rossicum (Kleopow) Barbar. (= Cynanchum rossicum (Kleopow) Borhidi)) como una maleza introducida
de Europa hacia Norteamérica, que se está expandiendo rápidamente. Pero sin duda las especies del género
Asclepias representan a las malezas más conocidas del grupo de las Asclepiadoideae, ya que de acuerdo con
la NAPPO (North American Plant Protection Organization, 2003) hay cerca de 100 especies pm con
áreas cultivadas, especialmente cereales y cultivos forrajeros, así como en áreas no cultivadas desde Canadá
hasta México.
Aunque autores como Hernández et al. (2003), Monroy et al. (2005) y López-Muraira e Iruegas (2006),
registran en los cultivos de agave la presencia de diversas especies de malezas trepadoras, entre ellas Ipomoea
purpurea (L.) Roth, I. pedatisecta M. Martens & Galeotti, Cucumis anguria L., C. dipsaceus C.G. Ehrenb., Cy-
clanthera dissecta (Torr. & A. Gray), Echinopepon floribundus (Cogn.) Rose, E. milliflorus Naudin, E. racemosus
(Steud.) C. Jeffrey, Sicyos deppei G. Don y iie Miu Nana, sin embargo, solo López-Muraira e Iruegas
en ea 5 especies de la familia A ii Decne., Gonolobus sororius A. Gray,
ep ata H.B.K., Matel icol W. D. Stevens y Mellichampia rubescens A. Gray, esto se debe a que
muchas veces las malezas tespadoras: se confunden entre sí ya que no existe información suficiente para la
identificación de éste grupo de plantas.
MATERIALES Y METODO
El trabajo se desarrolló del 2005 al 2006 en muestreos realizados de manera aleatoria a través de campos
agaveros de uno a seis años de edad ubicados en 65 localidades del estado de Jalisco. Los muestreos se re-
alizaron en diferentes municipios, comprendidos en las 5 regiones principales productoras de agave: Región
Altos Sur, Región Ciénega, Región Valles, Región Centro y Región Sierra de Amula, ubicadas entre los 20?
10. m N y 104°14.040 W, hasta los 21?05.294 N y 102?48.802 W.
Los fueron identificados usando las claves taxonómicas de Rzedowski y Rzedowski 2001; Juárez-
ne y Leda 2003; Rapini 2000; Standley y Williams 1969; Standley 1924 y por comparación con ejem-
plares depositados en el herbario IBUG, siguiendo el arreglo filogenético propuesto por Liede-Schumann et
al. 2005 y Rapini et al. 2006.
RESULTADOS
En los 65 sitios de muestreo realizados durante 2005 y 2006, se colectaron e identificaron 15 especies de
plantas de la subfamilia Asclepiadoideae, de las cuales 12 son de hábito trepador, 2 de crecimiento erecto y
solo una de ellas presenta hábito rastrero. De acuerdo a Rapini et al. (2006) y Liede-Schumann et al. 2005, se
registran 4 subtribus: Asclepiadinae, Cynanchinae, Oxypetalinae y Gonolobinae. Según Rapini et al. (2007)
el grupo MOG compuesto por las subtribus Metastelmatinae, Oxypetalinae y Gonolobinae son de origen
netamente neotropical y la ausencia de especies de la subtribu Metastelmatinae en el presente trabajo se
I 6áne7z-Muraira atal Acrlaniadaid [| l li al A + A]
I j | g ] 1343
Tasta 1. Especies de Ascelepiadoideae reportad | | cultivo del ag
Especie Subtribu Origen Habito
Asclepias curassavica L. Asclepiadinae América tropical Erecto
Asclepias linaria Cav. Asclepiadinae Norteamérica Erecto
ynanchum ligulatum (Benth.) Woodson Cynanchinae Endémica de Méxi Trepador
= Mellichampia rubescens A. Gray
Cynanchum foetidum (Cav) Kunth Cynanchinae Endémica de Méxi Trepador
Dictyanthus pavonii Decne. Gonolobinae Endémica de Méxi Trepador
Funastrum elegans Schitr. Oxypetalinae Endémica de Méxi Trepador
Funastrum pannosum ar Schltr. Oxypetalinae Endémica de Méxi Trepador
Gonolobus barbatus (H.B.K.) Gonolobinae Mesoamérica Trepador
Gonolobus lali Co R. Br. ex Schult. Gonolobinae Endémica de México Trepador
— Gonolobus erianthus Dec
E ui BL. EE & Greenm. Gonolobinae Endémica de Méxi Trepador
Gonolobus uniflorus H.B.K. Gonolobinae Endémica de Méxi Trepador
Labidostelma Emus Schltr. Gonolobinae Mesoamérica Trepador
— Matelea quirosii
Macroscepis obovata H.B.K. Gonolobinae Endémica de Méxi Trepador
atelea nummularia Decne. Gonolobinae Endémica de Méxi Rastrero
Matelea sepicola W.D. Stevens Gonolobinae Endémica de México Trepador
debe a que este grupo esta circunscrito a Sudamérica, principalmente Brasil y las islas del Caribe, mientras
Oxypetalinae, aunque se distribuye también para Sudamérica, su género Funastrum se dispersa mas hacia
Centro y Norteamérica, por su parte Gonolobinae, aunque de amplia distribución neotropical se dispersa
del noreste de Sudamérica a México hasta Norteamérica, pero no en las islas del Caribe. Las especies de
Asclepiadoideae se enlistan el la Tabla 1 y sus fotografías estan en la Figura 1.
DISCUSION
Es notorio encontrar la presencia de 15 especies de Asclepiadoideae como malezas trepadoras en el cultivo
del agave, ya que por ser un cultivo perenne, con poca área foliar y un manejo inadecuado de maleza,
no solo estas especies sino otras malezas de hábito trepador han sabido adaptarse y transformarse en un
problema para los agricultores agaveros. Por otra parte, es imp falar todas las especies reportadas
en el presente estudio son de origen netamente americano y por eso el gran endemismo registrado en el
presente estudio con 11 de las 15 especies reportadas como endémicas de México, algunas observaciones
hechas durante el transcurso de la presente investigación nos indican que de manera din este grupo de
plantas soporta el manejo de control por parte del agricultor agavero a b
control manual y mecánico.
r O L
CONCLUSIONES
Se reporta la presencia de 15 especies de Asclepiadoideae de las cuales 11 son endémicas a México. De
acuerdo a la literatura, por su hábito de crecimiento y las observaciones de campo, a estas especies se les
considera como malezas en el cultivo de agave.
AGRADECIMIENTOS
Agradecemos el apoyo recibido de los curadores del herbario IBUG por permitirnos consultar su acervo
botánico.
1344 i Hof tha Batanical D h Institute of Texas 2(2)
E
Asclepias linari
J
4
Gonolobus uniflorus
Matelea sepicola
López Muraira et al : Rerl I ladaid l r| Ine dal Ag * A] 1345
REFERENCIAS
AAG HA J IHiH
BRADLEY, K.W, P. Davis, AND E.S. HAGoob 2001. Identification and control of honeyvi
(Nutt) Britt. in Virginia. Publication 450-139W. Plant pathology, cor and weed science.
FurcH, STEPHEN H. AND DaviD W. Hatt. 2003. Identification of vine weeds in Florida Citrus. University of Florida,
Extension IFAS. HS926.
Grice, A.C. AND T.G. Martın. 2005. The management of weeds and their impact on biodiversity in the rangelands.
The CRC for Australian Weed Management, Townsville.
HÉRNÁNDEZ, M.L., I.G. LÓPEZ-MURAIRA v A.E. FLoREs-SuAREz. 2003. Agroecología de maleza en el cultivo de Agave tequilana
en las regiones de Amatitán-Tequila y Tlajomulco de Zúñiga, Jalisco, México. XXIV Congreso Nacional de la
Ciencia de la Maleza, Manzanillo, Col., México.
JUAREZ-JAIMES, V. Y L. Lozapa. 2003. Asclepiadaceae. In Flora del Valle de Tehuacán-Cuicatlán. Fascículo 37. Instituto
de Biología, UNAM, México.
LiEDE-SCHUMANN, S., A. Rapini, D.J. GOYDER, AND M.W. Chase. 2005. Phylogenetics of the New World Subtribes of
Asclepiadeae (Apocynaceae-Asclepiadoideae): Metastelmatinae, Oxypetalinae, and Gonolobinae Syst. Bot.
30:184-195.
LÓPEZ-MURARA, l.G. v R. IRuEGAs. 2005. Plantas trepadoras en el cultivo del (A ilana Weber) en Amatitán
y Tequila, Jalisco. XXVI Congreso Nacional de la Ciencia de la Me Chapala: Jo México.
LÓPEZ-MURAIRA, [.G. v R. Iguecas. 2006. El Complejo de maleza conocida como Chayotillo en el cultivo del agave
en Jalisco, México. XXVII Congreso Nacional de la Ciencia de la Maleza, Ensenada, Baja California Norte,
México.
LÓPEZ-MURAIRA, I.G., H.R. IRUEGAS Y J.F. Gómez Levva. 2007. Maleza de la familia Convolvulaceae en el cultivo del agave
(Agave tequilana). XXVIII Congreso Nacional de la Ciencia de la Maleza, Mazatlán, Sinaloa, México.
Luna, H.G. 2003. Guía fitosanitaria del cultivo de agave. Agroentorno. 1* Edición, Guadalajara, Jalisco, México.
Monroy, B., E. Pimienta, S.A. HURTADO, Y J.G. GALINDO. 2005. Efecto competitivo de la maleza sobre el crecimiento y
la respuesta fisiológica de agave (Agave tequilana Weber var. azul) en la región de Tequila, Jalisco, México.
Scientia-CUCBA 7(2):113-130.
NAPPO, 2003. Pest Fact Sheet. Asclepias syriaca L. NAPPO - PRA / Grains Panel Pest Fact Sheet - Asclepias syriaca
L. North American Plant Protection Orgaization.
RAPINi, A. 2000. Sistemática: Estudos em Asclepiadoideae (Apocynacea) da Cadeia do Espinhaço de Minas Gerais.
Tese (Doutorado) Instituto de Biociencias de Universidade de Sao Paulo. Departamento de Botanica.
RAPINI, A, M.W. Chase, AND T.U.P. Konno. 2006. Phylogenetics of South American Asclepiadoideae (Apocynaceae)
Taxon 55:119-124.
RAPINI, A., C. VAN DER BERG, AND S. LIEDE-SCHUMANN. 2007. Diversification of Ascl
World. Ann. Missouri Bot. Gard. 94:407—422.
Rozas, G.M. 1984. Manual teórico práctico de herbicidas y fitorreguladores. 2a Edición. Ed. LIMUSA Mex.
RzeDowski, G.C. v J. Rzepowski. 2001. Flora Fanerogámica del Valle de México. Instituto de Ecología A.C.
SALAMANCA, M. 2006. Cambios en la estrategia de manejo de maleza en el agave tequilero. Simposio Prioridades
Fitosanitarias Actuales en el cultivo del Agave tequilana Weber var azul. XXIX Congreso Nacional de Control
Biológico. Manzanillo, Col., México. 107-109
STANDLEY, PC.1923. Trees and shrubs of Mexico. (Fagaceae to Fabaceae) Smithsonian Institution. USNM. Contr.
US. Natl. Herb. 23(2):171-515,
STANDLEY, P. AND L.O. WiLiams. 1969. Flora de Guatemala. Asclepiadaceae. Field Mus. Nat. Hist, Bot. Ser. 24(3):
407-472.
Weston, L.A. J.N. Barney, AND A. DiTommaso. 2005. A review of the biology and ecology of three invasive perenni-
als in New York State: Japanese knotweed (Polygonum cuspidatum), mugwort (Artemisia vulgaris) and pale
swallow-wort (Vincetoxicum rossicum) Pl. Soil 277:53—69.
(Apocynaceae) in the New
Journal of t! IR h Institute of Texas 2(2)
1346
BOOK REVIEW
PETER BERNHARDT. 2008. Gods and Goddesses in the Garden: Greco/Roman Mythology and the Scien-
tific Names of Plants. (ISBN 0-8135-4266-9, hbk.). Rutgers University Press, 100 Joyce Kilmer Way,
Piscataway, New Jersey 08854-8099, U.S.A. (Orders: www.rutgerspress.rutgers.edu, 800-848-6224).
id 239 pp., MO 9 1/2" x 6 1/2".
1 1 ] 1:
Th:
and those who may fave just casually wondered how plants got their names. Peter Bernhardt of St.Louis Tay and the Missus
tis th
nins So has NOM us qua books that piq Epogen In vi one he Eu the
f1 of nymphs, satyrs, and centaurs, and th 1
Lo] L3 4 i 4 4
€. 1 ¿3 A+? 1 1 s T n 1 1 1 ] +1 J
r r e e e O
Le | a e | 1 f s 1 For example, z T 341 1 3 fi 1
trees or woody vines (Daphne, Cayo) iet demigods and iue men oai into herbs pn svildilowere" (Crocus, RO
In his Mix: ee p gy will not unpece th
] ] ach ac Native A +1 Tul mens : 41 1 dus 5 1:1
> ? 4 o
figures in plant taxonomy.
T } m RI: lthat includes pl taxonomists, and f h, and another
eo r r e 4
f sci ifi f pl } his i i g ] labl ] f y personal or public library. -Joann
Karges, Librarian (retired), Texas Christian Ui Lil Box 298400, Fort Worth, Texas 76129, U.S.A.
J. Bot. Res. Inst. Texas 2(2): 1346. 2008
THE SPECIES OF FUMARIA (PAPAVERACEAE) IN TEXAS, INCLUDING
F. DENSIFLORA, FIRST RECORDS FROM NORTH AMERICA
Monique Reed Richard V. Lansdown
Herbarium, Biology Department 45 The Bridle
U 3258 Stroud
Texas A&M University Gloucestershire GL5 4SQ
College Station, Texas 77843-3258, U. S. A. England, U.K.
Tim C.G. Rich
Department of Biodiversi 1S ic Biology
rsity
National Museum of Wales
Cardiff CF10 3NP Wales, U.K.
ABSTRACT
Examination of specimens of Fumaria from Texas reveals that Fumaria officinalis, F. parviflora, and F. densiflora have been collected in
the state; the latter has not previously been recorded from North dem Notes on bi EE the species known from Texas are
neni aon with a list of representative specimens. Herl th America should be re-examined for
1
li sted for the region.
RESUMEN
E examen ES e de Fumaria de T l ł gid l do F ia officinalis, F. adii y F. densiflora; esta
Norte Ámérica. Se aport i I 5 listinguir | i Texas, junto con
esas
1
Tid pr ncia
la región
The genus Fumaria (formerly placed in the Papaveraceae) comprises some fifty species (Lidén 1986) of an-
nual herbs native to Europe, Africa, and Asia, though many species are now naturalized elsewhere in the
temperate zone. The plants are similar in aspect to the more familiar Corydalis, having spreading to ascend-
ing stems, finely divided leaves, and racemes of strongly zygomorphic, spurred flowers. Fumaria differs in
having a caducous style (Lidén 1986); white to pink or purplish flowers, often with darker petal tips; and
single-seeded fruits.
Fumaria has been known in Texas since at least 1927 [F. officinalis L. subsp officinalis. Texas. Goliad
Co: Roadside, Mar 1927, Williams 73 (TEX).] Correll and Johnston (1970) listed both F. officinalis and F.
parviflora Lam for the state, as did Jones, et al. (1997). Boufford (1997) listed F. officinalis, F. capreolata L.,
and F. vaillantii Loisel. for North America and only F. officinalis for Texas. He did not include F. parviflora for
North America, stating that it occurs in North America only as a waif. Since then, most Texas collections
have been “pigeonholed” into F. officinalis. Billie L. Turner annotated many sheets as F. officinalis in 2000,
and Tuner et al. (2003) mapped only F. officinalis for the state.
In 2007, Richard Lansdown examined TAMU holdings of Fumaria and detected F. densiflora among them.
[Texas. Medina Co: Hondo; Weedy vacant strip at jct. Ave H and 18" Street, 16 Mar 2004, Reed, Walters, &
Corbett 2746 (TAMU)].
We subsequently borrowed specimens of Fumaria from Texas from BRIT/SMU, TAES, TEX/LL, and
SBSC. Careful examination, with reference to the works of Lidén (1986) and Sell (1993) and to European
herbarium specimens, confirmed the presence of multiple collections of F. officinalis, F. parviflora, and F.
densiflora from Texas. In one case, two species were collected from the same location within two weeks of
one another.
] Bot "M Tayac 2(2): 1347 — 1351. 2008
1348
Additionally, all three species are represented by material collected since 1990. Some populations may
be persisting, but it is probable that additional introductions continue to occur.
Identifying specimens of Fumaria can be problematic, especially with dry material. Characters tradi-
tionally used to separate species are not always reliable (Lidén 1986). Flower color is not definitive, since
white flowers may turn pink after pollination; the flowers increase in size during development; and bract
vs. pedicel length is variable. The relative sepal length and width, length of fully mature flowers, and fruit
characteristics are the most reliable when separating the species known from Texas.
The following key will separate the species known from Texas.
1. Sepals broader than the corolla F. densiflora
1. Sepals usually narrower than the corolla (occasionally nearly as broad).
2.. Fruit broader than long, often truncate or even depressed apically (may be apiculate in subsp. wirtgenii,
but not keeled or ridged); flowers pink, 7-9 mm long F. officinalis
2. Fruit narrower than long, semewhat keeled apically, often appearing apiculate if viewed from the side;
flowers white at anthesis, (4.5-)5-6 mm long F. parviflora
A more confident identification of the three species known from Texas can be made by comparing suites
of characters (largely from Lidén 1986) as given below. Note that in all species the corollas may be darker
(pink to purple or black) at the tips.
Fumaria densiflora DC. Bracts usually longer than the fruiting pedicels; fruiting pedicels strongly thick-
ened; sepals 2-3(3.5) mm long, 1.5-3 mm broad, usually orbicular to ovate, wider than the corolla, giving
unexpanded racemes a papery aspect; corolla 6—7 mm long, pink; fruit apically rounded to obtuse, slightly
keeled, rugulose, 2-2.3 mm long and broad (Fig. 1). Images of the Medina Co. plants may be viewed at
http://www.csdl.tamu. A EPUM A NDS
1 11
narrower than the corolla;
Fumaria officinalis L. Bracts Y? as longas to as]
corolla (6.6—)7-9 mm long, pink when well -developed; fruit da T long, rugulose, the apical pits shal-
low and rounded: subsp. officinalis. Sepals (1.6—)2.5—3.5 mm long, 1.5(-1.7) mm broad; fruit 2(-2.4) mm
long, (2.25-)2.5 mm wide, apically truncate and appearing depressed or emarginate when viewed from the
side: subsp. wirtgenii (Koch) Sell. Sepals smaller, 1.5-2 mm long, 0.75-1 mm long; corolla 7-8 mm long,
paler than in subsp. officinalis; fruit rounded, gly rugose, apically essentially subtruncate, rounded
and minutely apiculate when viewed from the side, but not keeled, ridged, or emarginate (Fig. 2).
Fumaria parviflora Lam. Bracts usually relatively broad and longer than the pedicels; fruiting pedicels
often strongly thickened; sepals 0.5—0.75(-1.8) mm long, 0.4—0.75(-1.6) mm wide, not broader than the
corolla; corolla (3.9-)5-6 mm long, white before pollination and often fading pink afterwards; fruit 1.6-2.3
mm long, 1.6-2.5 mm wide, usually longer than wide, apically subglobose to nearly acute, keeled, pointed
as seen from the side, the apical pits broad and squarish (Fig. 3).
A FINAL NOTE
It must be emphasized that all Texas—and indeed all North American collections!—of Fumaria represent
naturalized, recently introduced, single-event/impermanent, waif, or otherwise non-native populations.
Thus, any new collection of Fumaria ought to be identified using keys which include all Fumaria species.
That is to say, a key to European, Asian, and African species ought to be used in addition to a key designed
for a particular region. For example, in addition to the species listed here or present in the key in Boufford
(1997), F. bastardii Bor. and F. reuteri Boiss. (2 F. martinii Clay) have also been recorded in North America
(Boufford 1997.) It is entirely possible that additional species will be und in Texas in the future. It is also
i would reveal the presence
likely that a thorough re-examination of all North American herb
of more taxa than the three listed by Boufford (1997).
r
Reed et al., Fumaria species in Texas 1349
J 1 £r. LEES | “e D r] iv MAZA 1002) cartel
Fic. 1. Flower, fruit
ADDITIONAL REPRESENTATIVE SPECIMENS FROM TEXAS
Fumaria densiflora DC.
Bexar Co.: San Antonio, N side Loop 410 between Espada Rd and Roosevelt Rd (U.S. 281), 1 Apr 1969, MacEwan s.n. (SMU). Dimmitt
Co.: Crystal City, Mar 1931, Parks s.n. (TAES); 2% mi S of Crystal, 1 Apr 1973, Sturm s.n. (SMU). Montgomery Co.: Kingwood, King-
wood Drive, 15 bin 1987, Ora s.N., S Travis Co.: Austin, near ein Toi Museum, 26 Mar 1956, McCart 5526 (SMU);
Austin, Col Bridge, 19 Apr 1957, M 555 f Texas, Austin, 8 Mar
1955, Anderson s.n. (TEX). Wharton. Co.: 0-0.5 mi ni E of Kriegel Rd, N of Caney Cree S of Hwy 1301, 15 Feb 1990, Hanka 2 (TAMU),
Boling, Luper Ranch, 6 Feb 1971, Fleetwood s.n. (TEX).
Fumaria officinalis L.
Note. —We attempted to name the better material to subspecies following Lidén (1986). Lidén (1986) and
Sell (1993) define the subspecies in different ways, and the cl they use do not always correlate well,
so the determinations are provisional. Boufford (1997) noted the occurrence in North America of plants
conforming to subsp. wirtgenii but did not recognize infraspecific taxa in his account. The specimen cited
1350 J lof the Botanical R h Institute of Texas 2(2)
D. J I£ Edn Y cartel
Fic. 2. FI 1 fruit of Fi ia officinalis. A. subsp. officinalis. B. subsp. wirtaenii
F H Y
ri A £, L- £r. : c" n J J £ 1:42 fanor\
Fic. 3
i )
near the beginning of this paper belongs to subsp. officinalis. The two below were undeterminable as to
subspecies.
Gillespie Co.: Fredericksburg, 18 May 2003, Lindeman s.n. (BRIT). Smith Co.: Old Experimental Station 1 mile N by E of Troup (aban-
doned 1931), 9 Apr 1949, Cory 55568 (SMU)
Fumaria parviflora Lam.
Bexar Co.: San Antonio, N side Loop 410 between Espada Rd and Roosevelt Rd (U.S. 281), 13 Apr 1969, MacEwan s.n. (SMU). Dimmitt
Co.: Carrizo Springs, 29 May 1948, Hudspeth 216 (BRIT). Hidalgo Co.: McAllen, railroad tracks between Weyheuser and Texas Motor
Trailer Court, 3 Apr 1965, Fleetwood 8067 (TEX); McAllen; N McColl Rd between Violet and Nolana Streets; 26 Feb 1992, Halse 4440
(BRIT, TAES); Weslaco, 18 Feb 1939, Friend 27851 (TAES); Weslaco, 18 Feb 1942, Friend 1949 (SMU); Weslaco, 18 Feb 1942, Friend
1950 (TEX). Travis Co.: NW corner of SE b pa at oe e edid Treatment en ca. 1.5 air mi N 32? W of jct. of FM. 973
and TX. Rt. 71, 19 Apr 1983, Carr 7283 (BRIT). Wi Mill, f SW corner of Lake Creek Parkway
and RM 620, 18 May 1982, Carr 3969 (BRIT).
ACKNOWLEDGMENTS
The authors would like to thank the curators and staff of BRIT/SMU, TAES, TEX/LL, and SBSC for the loan
of material; Mike Powell for searching the SRSC collection for Fumaria; Magnus Lidén, David Boufford, and
Reed et al., Fumaria species in Texas 1351
Jacqueline van de Veire for helpful reviews and comments; Magnus Lidén for permission to reprint his excel-
lent drawings; and Robert Corbett for the images which appear in the referenced online image gallery.
REFERENCES
Bourronp, D.E. 1997. Fumaria. In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America
North of Mexico. 12+ vols. New York and Oxford. 3:356-357.
Corrett, D.S. AND M.C. JoHNston. 1970. Manual of the vascular plants of Texas. Texas Research Foundation,
Renner.
Jones, S.B., J.K. Wierz, ano PM. Montcomery. 1997. Vascular plants of Texas: a comprehensive checklist including
synonymy, bibliography, and index. University of Texas Press, Austin.
Liven, M. 1986. Synopsis of Fumarioideae (Papaveraceae) with a monograph of the tribe Fumarieae. Opera Bot.
88:1-133.
Se, PD. 1993. Fumaria L. In: Tutin, T.G., Burges, N.A., Charter, A.O., Edmondson, J.R., Heywood, V.H., Moore, D.M,
Valentine, D.H., Walters, S.M. ano Webb, D.A., eds. Flora Europaea, 2nd edition. Cambridge University Press,
Cambridge. 1:306-311.
TURNER, B.L., H. NichoLs, G. Denny, AND O. Doron. 2003. Atlas of the vascular plants of Texas, vol 1. Sida, Bot. Misc. No.
24. Botanical Research Institute of Texas, Fort Worth.
1352 J | of the Botanical R h Institute of Texas 2(2)
BOOK REVIEW
Tony RoDD AND JENNIFER STACKHOUSE. 2008. Trees: a Visual Guide. (ISBN 978-0-520-25650-7, hbk.). Uni-
versity of California Press, 2120 Berkeley Way, Berkeley, California 94704-1012, U.S.A. (Orders: www.
ucpress.edu, orders@cpfsinc.com, 800-777-4726). $29.95, 304 pp., color photographs, tables, glossary
and index, 11" x 9".
a ans dt Vos ing! 1 sel y 1 td ad lt - 1 ail h EN 1 1 ff List 1 Authors Rodd
have gi I ler af: ic guid ll thing ix di section "A avers ee trees,’ Led readers to the
defi iti f hati 1tl ranoing
g I ranging
from how basic parts function t to Pad E occurs, 11 ival in ext i tal conditi i tedi t
two. Closely related diversity i n ene are d in section three. The tree diversity and
lso includ i 1 pi lad orld, including information on such
trees as sir cedar of poi n. Sequoia, E many more.
insection four. J i ] hiel P h hah;
The ecology of trees and speci hip th i t. as ie as some al animal species f. li h e
are presented. Ten community ty} included in the book. An f e discussed is the Savannas, gaa
in Africa, which is inhabited by trees such as Umbrella Acacias and aude such as Patas Kn and giraffes, which have unique
relationships with plant species in the region.
The
maining sections are "Trees and the human world" and "An indispensable resource" which explore the many uses humans
have for trees and the i ft The uses of tree and tree products by man are astounding: food, building supplies,
medicine, music, art, a oria. tools, fibers and more. The authors nis ud a pues such as biodiversity,
flood protection, and soil stabilization, which many readers may not h
F
Trees: a Visual Guide serves as a beautiful [ I ft t kind and the ap Planet. ne Rodd and
Stackhouse h written an easy to UM text w ith color photogr aphs which tr uly he i I : a Visual
23 3 f, Ss TR ff, f, f. 1 1 A J: 14h; t O A O pR
E Lee LucReyaoo,
Botanical Research Institute dí Texas, 500 East 4th Street, Fort Worth, Texas 76102-4025, U.S.A.
J. Bot. Res. Inst. Texas 2(2): 1352. 2008
NOTEWORTHY COLLECTIONS FROM THE SOUTHEASTERN UNITED STATES
Bruce A. Sorrie Richard J. LeBlond
s of North bine Herbarium University - North Carolina Herbarium
orth Carolina Botanical Garden North Carolina Botanical Garden
a p North dis 27599-3280, U.S.A. Chapel Hill, mue Carolina 27599-3280, U.S.A.
bruce.sorrie@ncmail net RichardLeBlondacharter.net
ABSTRACT
Taeld 1 E p A A Jioc 1 1 TENUES RUE Je faroa : Aonlinje di yperus grayi Dichan-
thelium annulum, Echinod llus, Eleocl ii, Fimbristyli aan nana, Hypericum lloydii, Juncus dudleyi,
Juncus interior, Juncus wd n pun Juncus al iade m mie o drummondii, Panicum dichotomiflorum
Var. puri tanorum
Polyg 1 j r. attenuata, pice human. Sclerolepis
Q 1 e 1 1
S
2
d
TTey
uniflora i
wo » ? od 2. r r
ced Jnr Ac F Agalinis filifolia Aletris lutea, Boltonia diffusa, Paena pumila. J ticulat DE
4
multifida, Polygonum glaucum, Yucca gloriosa.
RESUMEN
¡AS A | a pps A A | 1-3 PE = 1 1
A 7 <A nic Ar H +
23 especies: Ag Cy ype-
Lu
rus grayi, Dichantheli lum, Echinod llus, Eleochari l ii, Fimbristylis sch aylussacia nana, Hypericum aa
Juncus dudleyi, Juncus interior, ies diis Tus Lin in Juncus E oa oi moneda drummondii, Panic
ttenuata, Sagittaria chap-
ci
dichotomiflorum puritanoru
E
1 las
F
T, pe Jat
manii, n aa Diriailuria ijuva, ia viridiflora. De ot
ado
: Agalinis filifolia Aletris lutea Boltonia diffusa va pumila, ] Leucospora
multifida, Polygonum glaucum, Yucca ao
INTRODUCTION
We have conducted botanical fieldwork and herbari tudies in tl theastern United States coastal plain
since 1990. In part, this work was directly related to our occupation as biologists with the North Carolina
Natural Heritage Program, in part as a result of contract work to produces status pu of rare plant taxa
for the U.S. Fish & Wildlife Service, and in part related to ongoing Research Associates
at the University of North Carolina herbarium (NCU). In addition, the senior author has searched many
national and local herbaria while preparing county-level range maps for over 1000 endemic coastal plain
taxa. These studies have yielded a large number of noteworthy records, some of which have been published
(e.g., Sorrie & Leonard 1999; LeBlond & Sorrie 2001).
In the course of this work we visited over two dozen major herbaria with significant holdings of coastal
ts for those which hold vouchers cited herein). These efforts have resulted
Anl
plain plants (see
in the discovery of new state recoja for twenty three species of vascular plants in the southeastern United
States. For eight other species, the vouchers cited below represent significant range extensions or repre-
sent the only known current records in specific states. Scientific names follow published volumes of Flora
North America; otherwise they follow Weakley (2007). Herbarium acronyms are those of Holmgren et al.
(1990)
ACANTHACEAE
Yeatesia viridiflora (Nees) Small € Dicliptera halei Rm A widespread but sparsely distributed species
of the Gulf Coastal Plain from Florida and Georgia to eastern Texas. The map in
Hilsenbeck (1989) depicts a hich al lud tof Alabama, Mississippi, and Louisiana,
plus a tiny portion of meca The latter state was listed by Small (1896), but we have seen no specimen
J. Bot. Res. Inst. Texas 2(2): 1353 — 1361. 2008
1354 J | of the Botanical R h Institute of Texas 2(2)
and Y. viridiflora does not appear in the Tennessee Atlas (Chester et al. 1997). In Alabama (Alabama Flora
Committee 2004) and Louisiana (Thomas & Allen 1996) this species is decidedly local, being absent from
large portions of each. We can find no records for Arkansas (Smith 1988) and Mississippi (Kartesz 1999);
the following are the first records for these states.
ucher specimens: ARKANSAS: Dallas Co.: fertil ded hillsides, Thatcher's Bluff, P.O. Sardis, elev. 250, 27 Jun 1964, D. Demaree
50592 (BRIT). MISSISSIPPI: Wayne Co.: Bucatunna Creek ca. 7 mi E of Waynesboro, T8N R5W, 11 Oct 1970, K.E. Rogers 4777 with
R. Watson (MISS)
AGAVACEAE
Yucca gloriosa L. This species of maritime habitats is uncommon from North Carolina to Georgia, and
rare from Florida to Louisiana. This is apparently the third county of occurrence in Florida (Atlas of Florida
Vascular Plants 2007) and the only one from the west coast of the peninsula.
Voucher specimen: FLORIDA: Pinellas Co.: occasional in foredunes, west coast of Caladesi Island, 2 Jun 1963, R.H. Wagner 876
(DUKE)
ALISMATACEAE
Echinodorus tenellus (Mart.) Buch. (= E. parvulus Engelm.). Pymes and n ed ies map this diminu
tive annual in ten states (but read Va. for Vt.). However, it ted by Tatnall
(1946) for Delaware, Jones (2005) for Kentucky, Thomas and Allen (1993) for Louisiana, Steyermark (1963)
for Missouri, Stone (1911) for New Jersey, Taylor and Taylor (1994) for Oklahoma, and LeBlond and Sorrie
(2001) for North Carolina. The following are the first published records for Arkansas and Mississippi.
Voucher specimens: ARKANSAS: Baxter Co.: WSW of Mt. Home, small stock pond along roadside, 26 Oct 1974, B.L. Lipscomb 992
(BRIT). MISSISSIPPI: Lamar Co.: 3 mi W of Purvis, abundant on edge of lake, 29 Aug 1973, K.E. Rogers 9223 (GA, NCU, VDB); same
location (Mossy Pond), 1 Sep 1998, B.A. Sorrie 9924 with S.W. Leonard (GH).
Sagittaria chapmanii (J.G. Smith) C. Mohr. Haynes and Hellquist (2000) treat this entity as S. graminea
Michx. ssp. chapmanii (J.G. Smith) R.R. Haynes and C.B. Hellquist, but analyses of allozyme variation by
Hauber and Lege (1999) argue convincingly that S. graminea and S. platyphylla (Engelm.) J.G. Smith are more
closely related to each other than either is to S. chapmanii. We therefore follow Weakley (2007) in treating S.
chapmanii as a full species. This species is notable among our Sagittaria in having branched inflorescences.
Haynes and Hellquist (2000) map it in South Carolina, Georgia, Florida, and Alabama; LeBlond and Sor-
rie (2001) document it from North Carolina; Majure (2007) reports it from Mississippi. These are the first
records for Louisiana, Texas, and Arkansas
Voucl i ARKANSAS: Bradley Co.: Johnsonville Prairie, 5.9 mi SW of Jol ill ] 1, Sec. 23, T16S, ROW,
flat hen prairie and woods, 5 me 1984, R.D. Thomas 90284 (NLU); Calhoun Co.: along dirt road in flat pine woods 1 mi S of
Arkansas 4, ca. 6 mi W of Hampton and US 167, 30 May 1990, R.D. Thomas 118250 (NLU). LOUISIANA: Allen Parish: wood road 1 mi
W of highway 165, 1 mi N of Oakdale, 17 Aug 1979, S. Darwin 1973 with E. Sundell (NLU); Beauregard Parish: sandy roadside 3 mi S of
De Ridder, 11 Jul 1970, Northwest State College Biology Class s.n. (NLU); Bossier p ca. 2 mi N a pda water of ditch along road,
7 Aug 1938, D.S. a 10131 ee H.B. Correll (DUKE); Livingston P ji , SE of Holden, 19 Mar 1975,
M. Allen 642 I ] ypress Branch, 4 mi W of Mora, 30 Ap 1972, R.D. Thomas 28600 (NLU);
Rapides Parish: 5 mi S of Pineville, shallow water of roadside pond in pine woods, 21 Oct 1966, J.W. Thieret 25012 (DUKE); St. Tam-
many Parish: pine hin » d iiam 41, E n HE Bayou, N of Hickory Fire Tower, 19 Jun 1976, R.D. Thomas 49523 (NLU); ca.
5 mi NW of Slidell, di 26, 12 Jun 1968 JW. Thieret 29206 (DUKE). TEXAS: Grimes Co.: near highway
1774, 7 mi SE of Plantersville, Rb in ditcl ding water , 1 Jul 1969, J.J. Sperry s.n. (BRIT).
ASTERACEAE
Boltonia diffusa Elliott. Radford et al. (1968) include this species within their concept of B. caroliniana
(Walter) Fernald, but Karaman-Castro and Urbatsch (2006) recognize them as distinct species and map
B. diffusa in North Carolina and southward. We are aware of only a single historical specimen in North
Carolina: H.E. Ahles 19986 (NCU) from Stanly County in 1956. The following vouchers provide current
documentation. Since all records are from highly disturbed habitats, the native/alien status of this species
in the state is uncertain.
c H I1 ADI 1, Al I. £: +l +l + Ilantad Chat 1355
Vi TNA- A € L 3 1 1231
, and loblolly pine plantation,
SR 1121 N of Leggett Branch, 11 Oct 2007, B.A. Sorrie 12047 (NCU); Moore Co.: abundant in dry clearing in floodplai
just downstream from confluence with Nick's Creek, 21 Aug 2007, B.A. Sorrie 12003 (NCU, US.
Sclerolepis uniflora (Walter) B.S.P. A coastal plain endemic of periodically inundated wetlands, this species
ranges from New Hampshire to Florida and Alabama (Lamont 2006). Sclerolepis uniflora is not included for
Louisiana by Thomas and Allen (1993) or for Mississippi by Kartesz (1999). The following ds establish
its presence in Louisiana and ——
Voucher specimens: LOUISIANA: St. Tamman: P hall ater, woods road through wet slash pine-
pond cypress forest, is Wetlands ccn 2 Jun 1999, L. Smith s.n. n. (LSU, NLU). MISSISSIPPI: George Co.: 9.1 mi E of route
63 on US 98, savanna-like area, 26 May 1973, D. Boufford 9441 with H.E. Ahles (GH).
CYPERACEAE
Cyperus grayi Torr. Gray's umbrella-sedge occurs in dry sand d coastal d ly along the
Atlantic and Gulf Coastal Plain from New H hire to Alabama (Kartesz 1999) and Misas: (Radford et
al. 1968). Itis indicated as absent from Florida by Kartesz (1999), Tucker, Marcks, and Carter (2002), USDA
NRCS (2007), and the Atlas of Florida Vascular Plants (2007). This is the first record for Florida.
Voucher specimen: FLORIDA: Liberty Co.: Apalachicola Bluffs and Ravines Preserve near Bristol, 4 Jun 1991, RJ. LeBlond 2171
(FSU)
Eleocharis engelmannii Steud. A widespread species in the United States and southern Canada, but rarely
collected in the southern Atlantic and Gulf Coastal Plains. Thomas and Allen (1993) do not include it for
Louisiana; this is the first record for that state.
Voucher specimen: LOUISIANA: Jeff Davis Parish: along West Bayou Lacassine, 23 Sep 1967, J.W. Thieret 27938 (DUKE).
1
Fimbristylis schoenoides (Retz.) Vahl. This fimbry sedge, introduced from Asia, has been 1 from
Georgia and Florida west to Louisiana (Kral 20022). This is the first known North Carolina collection.
Voucher specimen: NORTH CAROLINA: Hyde Co.: e Island, in maritime wet grassland along N side of NC 12, 0.1 mi E of
peH National Seas! I lary sign just SE ke village; 17 Oct 2005, RJ. LeBlond 6154 (NCU).
eJ
Fuirena pumila (Torr.) Spreng. This specialist of ponded depressions, sinkhole ponds, and glacial kettle-
hole ponds is scattered along the Atlantic and Gulf states from Massachusetts to Florida and Texas, with
disjunctions to the Great Lakes region. Majure (2007) reports it from two Mississippi counties. This is the
third record for Mississippi.
Voucher specimens: MISSISSIPPI: Lamar Co.: Mossy Pond, 3 mi W of Purvis, 1 Sep 1998, B.A. Sorrie 9928 with S.W. Leonard (GH,
MISS, NCU)
>
Lipocarpha drummondii (Nees) G.C. Tucker. Tucker (2002) does not include Louisiana in the range of
this diminutive annual, and Thomas and Allen (1993) do not include it. The specimen cited below differs
from L. micrantha (Vahl) G.C. Tucker in possessing second scales 0.7 mm long. This is the first record for
Louisiana.
Voucher specimen: LOUISIANA: Caddo Parish: ] ] edge of d along railroad, Section 6, ca. % mi SE of Reiser, 7 Jul 1970,
J.W. Thieret 32137 (NCU).
Rhynchospora cephalantha A. Gray var. attenuata Gale. Kral (2002b) synonymizes this variety under
nominate R. cephalantha, which ranges from New York to Florida and Texas. However, LeBlond in Weak-
ley (2007) recognizes it as distinct, due to its narrower and shorter achenes and its tendency to produce
hemispherical heads (vs. subglobose to globose). In addition, var. attenuata inhabits wooded streamheads,
baygalls, and semi-open seepage slopes, rather than savannas, marshes, and other open habitats occupied
by var. cephalantha. In our experience var. attenuata is not a rare plant, but merely overlooked. Gale (1944)
1356 J lof the Botanical R h Institute of Texas 2(2)
cites specimens of var. attenuata from North Carolina, South Carolina, Alabama, and Mississippi. Thomas
and Allen (1993) do not mention this variety, even in synonymy, for Louisiana. This is the first record for
that state.
1 dul lor Teal fu PAIT DA NI
Voucher specimens: LOUISIAN
Forest, ca. 1.5 mi E 22s e Church and Louisiana route e 10, 7 Sep 1987, R.D. Thomas 101449 with C.M. Allen (NLU).
ERICACEAE
Gaylussacia nana (A. Gray) Small. Luteyn et al. (1996) list this shrub for Alabama, Florida, Georgia, and
Tennessee. This species is not listed for North Carolina (Radford et al. 1968), Mississippi (Kartesz 1999),
or Louisiana (Thomas and Allen 1993); these are the first records. The Louisiana specimen does not give
parish name and it is unlikely to have been collected in Orleans Parish, but rather in adjacent St. Tammany,
LM or basa Parishes where suitable pinelands occurred.
LOUISIANA: “New Orleans” 1832, T. Drummond s.n. (GH). MISSISSIPPI: Harri Co.: Handsboro, low wet pine
woods, 29 May 1951, D. Demaree 30621 (GH); Mississippi City, low wet woods, 1 Jun 1951, D. Demaree 30677 (DUKE); sandy open pine
ridges, Mississippi City Post Office, 9 Jun 1954, D. Demaree 35539 (GH); DeSoto National Forest, N of Krohn cemetery, W of Old Biloxi
Road, rolling dry longleaf pine upland, 18 Apr 1997, B.A. Sorrie 9158 (GH, MISS, NCU); DeSoto National Forest, N of White Star Road,
hilltop under longleaf pine, oak, and dogwood, 22 May 1997, B.A. Sorrie 9248 (MISS, NCU); Jackson Co.: Ocean Springs, 4 Apr 1889,
E.S. Earle s.n. (DUKE); O Springs, 18 Jun 1895, J. Skehan s.n. (GH); Fontainebleau Point, 11 Jun 1954, D. Demaree 35561 (GH); e
Creek NE of Mor An local in ps od woods near low bog, 15 Apr 1964, S. McDaniel 4185 (FSU, IBE); NORTH CAROLIN.
New Hanover Co.: pine/ ] Il habitat W of Wilmington Beach, 22 Jun 1997, RJ. LeBlond 4761 with A.S. Weakley CE
same location, 23 Oct 2002, RJ. LeBlond 5748 (NCU). The 1997 record was usly reported as G. tomentosa (A. Gray) Pursh ex
Small by LeBlond and Sorrie (2001).
Kalmia angustifolia L. The two eastern North American "sheep laurels," Kalmia angustifolia and K. caro-
lina Small, were treated as full species by Southall and Hardin (1974) but as varieties of K. angustifolia by
Ebinger (1974). The ranges of these taxa overlap in a small area of southeastern Virginia. Recent field and
herbarium studies suggest that hybridization is rare, which supports species status (Sorrie and Weakley
ms.). Heretofore, K. angustifolia had not been reported from North Carolina (Radford et al. 1968); the fol-
dd ee documentation.
INA: Gates Co.: Wyanoke area, 3.5 km NE of Riddicksvill fd dd its, abundant
"e species, 9 Aug 2005, J. Townsend 3486 (NCU, VPD; same data but 3.9 km NE of Riddicksville, J lowhsend 3487 (NCU,
VPD
HYPERICACEAE
Hypericum lloydii (Svenson) W.P. Adams. This dwarf shrub occurs in the lower piedmont and upper
coastal plain from North Carolina to Alabama. The Atlas of the Virginia Flora (2007) does not include H.
lloydii; this is the first record for that state.
Voucher specimen: VIRGINIA: Mecklenl C dside of Virgini 58, 7 Jul 1967, W.D. Seaman 7460 (NCU).
e [s]
JUNCACEAE
Juncus articulatus L. A northern species that in eastern United States ranges south to Pennsylvania and
New Jersey, with several records in the mountains of Virginia, West Virginia, and Kentucky (Brooks and
Clemants 2000). The following provides label data for a curiously disjunct record from the seacoast of North
Carolina. Although the specimen was annotated correctly in 1954 by W.T. Batson and included in Radford,
Ahles, and Bell (1968), details of this record have not been published previously.
Voucher specimens: NORTH CAROLINA: Dare Co.: marsh at Cape Hatteras, 23 May 1954, A.E. Radford 8188 (NCU).
Juncus dudleyi Wiegand. This species of calcareous soils is not mapped in the deep southern United States
by Brooks and Clemants (2000). However, the Alabama Flora Committee (2004) maps it in Bullock and
Montgomery Counties. Thomas and Allen (1993) omit it from Louisiana; the following is the first docu-
mentation for Louisiana.
[4 H JA 14^DI l M 1. f 41 "I 4 TT d Chag 1357
Voucher specimens: LOUISIANA: Ouachita Parish: cleared lots and ds off Lakeside St. in new Waterside Park Subdivision, E of
Monroe, 7 May 1967, R.D. Thomas 2614 (NCU); Winn Parish: Cedar Creek, 3 mi E of Winnfield, open Gleditsia wood, 24 Apr 1955,
G.R. Cooley 3686 wiih L.J. Brass (NCU).
Juncus interior Wiegand. This is a widespread midwestern species of prairies and prairie-like habitats.
Brooks and Clemants (2000) map it in adjacent Louisiana, Arkansas, and Tennessee, but not Alabama and
Mississippi. The Alabama Flora Committee (2004) maps it in Barbour County; Kartesz (1999) does not map
it in Mississippi. This is the first record for Mississippi and second for Alabama.
Voucher specimens: ALABAMA: Perry Co.: four mi S of Marion on highway 5, open prairie, 15 May 1969, J.L. Thomas 2066 (NCU).
MISSISSIPPI: Bolivar Co.: 5 mi W on Old Pace Road, P.O. Cleveland, 26 May 1960, H.L. Jacob 1351 (NCU).
Juncus militaris Bigelow. Thi ] species of lakes and pond g th to Delaware, Maryland,
and northern Indiana (Brooks and Clemants 2000). This is the first record for North Carolina, a disjunc-
tion of some 380 km. Plants are abundant in a dense but narrow band in 0.3—0.7 m of water in a very large
Carolina a lake.
Vi W. C Park, SE side of Lake Phelp hallow water
with caló aquaticum and Xyris smalliana, 4 Sep: 2005, B.A. Sorrie 11715 with L. Mize, J. Mize (GH, NCSC, NCU, US).
Juncus nodatus Coville. This species of the southern midwest United States is notable for the ring-like
bands on leaves at each cross-partition. These are the first records for Mississippi.
| MISSISSIPPI: Jacl C ditch, Ocean Springs, 9 May 1970, K.E. Rogers 3246-B (NCU); Rankin Co.: E shor
of Paket Reservoir between Safe Harbor Mari d Pelahatchie Bay area, locally abundant, 7 Oct 1972, K.E. Rogers 8812-A nd
Juncus subcaudatus (Engelmann) Coville & S.F. Blake. This species is a relative of J. canadensis J. Gay,
but nearly confined to the Appalachian Mountain system. Radford et al. (1968) do not include it for South
Carolina; this is the first record.
Voucher specimen: SOUTH CAROLINA: Oconee Co.: Whitewater River NW of Jocassee, 2 Sep1956, A.E. Radford 17875 (NCU).
LAMIACEAE
Pycnanthemum flexuosum (Walter) B.S.P. This coastal plain endemic ranges from Virginia to Alabama
(Kartesz 1999). This is the first record for Mississippi. The specimen was determined by H.A. Gleason in
1960 as P. setosum, a closely related congener, but the specimen possesses narrow leaves characteristic of P.
flexuosum (note: the drawings of these two species are reversed in Gleason 1952). It was on this collection
that Kartesz (1999) based his Mississippi occurrence of P. setosum; therefore the latter record should be
deleted, leaving P. setosum with a range of New Jersey to Florida.
Voucher specimen: MISSISSIPPI: Pearl River C ly grassy flat along railroad in cleared longleaf pineland, 2 mi S of Pica-
yune, 26 May 1956, J.D. Ray, Jr. 6488 (NCU).
LENTIBULARIACEAE
Utricularia juncea Vahl. A New World species of wet acidic seepages, pine savannas, pond margins, and
boggy wetlands, U. juncea in North America is restricted to the coastal plain from Long Island, New York
to southern Florida and eastern Texas. This is the first record for Maryland, the species being omitted by
Brown and Brown (1984).
Voucher specimen: MARYLAND: Prince George Co.: Suitland Bog, 13 Aug 1921, P.T. Newbold 708 (MO).
LILIACEAE
Aletris lutea Small. A widespread species of pine savannas and flatwoods, ranging from southern Georgia
to south Florida to southeastern Louisiana. According to Thomas and Allen (1993), this is the first record
from west of the Mississippi River.
Voucher specimen: LOUISIANA: Vernon Parish: Kisatchie National Forest, silt loam, level, burned over, T1S R8W Sec.14, 2 May
1940, E.L. Stone 21 (FSU).
1358 J | of the Botanical R h Institute of Texas 2(2)
OROBANCHACEAE
Agalinis divaricata (Chapman) Pennell. Outside of Florida, this species was heretofore known only from
three southern Alabama counties (specimens at FSU, GH, NCU). Jones and Coile (1988) do not include it
for Georgia; this is the first published record for that state.
Voucher specimen: GEORGIA: Decatur Co.: in dry sandy soil, E side of airstrip road at L/S [Lake Seminole], 15 Sep 1973, A. Gholson
s.n. (FSU)
Agalinis filifolia (Nuttall) Raf. This species ranges throughout Florida and southeastern Georgia. Pennell
(1929) cited an old Gates specimen from Alabama that lacked location data; the Alabama Flora Committee
(2004) mapped it in Baldwin Md These are the first modern records for Alabama.
i ALABAMA: Baldwin C ls N of sand d If of Mexico, highway 180 between Gulf
Shores Bud Fort Morgan State Park, 7 Oct 1967, M. H. ire 3850 (NCU); same place and m B.E. Wofford 10192 (NCU); Bon Secour
National Wildlife Refuge, W of Little Lagoon, 29 Sep 1997, B.A. Sorrie 9573 with RJ. LeBlond (AUA, GH, NCU).
PLANTAGINACEAE
Leucospora multifida (Michx.) Nuttall. Leucospora prefers moist to wet sandy or muddy shores and depres-
sions, including disturbed habitats. It is widespread in eastern North America, but rare along the Atlantic
Coastal Plain, with the North Carolina population disjunct from the panhandle of Florida. This is the only
known specimen for North Carolina. Kartesz (1999) maps it in North Carolina and cites Weakley (1997)
as the source; we here give full details of this record.
Voucher specimen: NORTH CAROLINA: Carteret Co.: Cherry Point Marine Corps Outlying Field Atlantic, NW of the community of
Atlantic, in borrow pond within NW corner of runway complex, 2 Sep 1993, RJ. LeBlond 3544 (NCU)
POACEAE
Dichanthelium annulum (Ashe) LeBlond. Within the Dichotoma group, D. annulum is one of the most
distinct taxa (LeBlond 2001), with its relatively tall stature, soft villous foliage, short hairy upper internodes,
minute ligules, and bearded nodes. Freckmann and Lelong (20032) synonymize it under D. dichotomum (L.)
Gould ssp. mattamuskeetense (Ashe) Freckmann & Lelong, but that taxon differs in its Pare leaves, larger
spikelets, coastal plain distribution, and affinity for wet acidic soils. Dichanthelium ges from New
Jersey to Missouri, south to Georgia and Mississippi; it inhabits open woodlands, glades, and barrens, often
in high pH substrates. This represents the first collection in North Carolina since 1968.
Voucher specimen: NORTH CAROLINA: Stanly Co.: Morrow Mountain State Park, north slope of Biles Mountain, common in
graminoid dominated glade-like woodland under Quercus alba, Q. stellata, Carya glabra, Cercis canadensis, Pinus echinata, 1 Jun 2007,
B.A. Sorrie 11963 (NCU)
Panicum dichotomiflorum Michx. var. puritanorum Svenson. This represents the first nd report
of this rare grass from Delaware and possibly the entire Delmarva Peninsula. Panicum dich iflorum var.
puritanorum ranges along the coastal plain from Nova Scotia and New Hampshire south to Virginia, with
disjunct occurrences around Lake Michigan in Indiana and Illinois. It is found primarily in wet sands and
organic soils of seasonally exposed pond and lake shores. It was previously reported from Delaware by R.A.
Ralph in an undated manuscript at the University of Delaware (Kartesz 1999). This manuscript was prepared
sometime prior to 1992 while Ralph was a student at the College of Marine Studies in Lewes (McAvoy pers.
comm.). No voucher specimen for this report has been located. Plants from Delaware northward typically
have slender culms 0.3—6 dm long and leaves 1-8 mm wide. Plants with floral characters of var. puritanorum
but with stout culms 0.8—2 m long and leaves 7-25 mm wide (similar to var. dichotomiflorum) occur in bot-
tomlands in southeastern Virginia, and have been recognized as P. dichotomiflorum var. imperiorum Fernald
(Fernald 1942). These plants are currently regarded as belonging to var. puritanorum by Weakley (2007) and
presumably are the source of the inclusion of Virginia within the range by Freckmann and Lelong (2003b).
The Virginia plants need further study.
Cc t I1 ADI d, al le £ el el $ llnttad Chat 1359
Voucl peci DELA RE: S Co.: Huckleberry Pond, SE of Milford and NW of Broadkill Beach, 17 Sep 1991, R.J. LeBlond
2554 with B.A. Sorrie and W.A. McAvoy (DOV).
POLYGALACEAE
Polygala leptostachys Shuttleworth. This essentially Floridian species is rare outside of that state, known
from only a few specimens in Georgia (Jones and Coile 1988) and Mississippi (Kartesz 1999). This is the
first published record for Alabama. Al Schotz (pers. comm.) informed us that he has found it in Baldwin
and Geneva Counties.
Voucher specimen: ALABAMA: Escambia Co.: Conecuh National Forest, N of road to Hines Lake, dry to xeric knoll in longleaf pine-
wiregrass-shrub flatwoods, 6 Aug 1996, B.A. Sorrie 8974 (AUA, GH).
POLYGONACEAE
Polygonum glaucum Nutt. This plant of maritime fored d accreting beaches ranges from Massachu-
setts to north Florida, almost exclusively on the Atlantic coast. The Atlas of Florida Vascular Plants (2007)
maps it only in Duval and Gulf Counties. This is the second record for the Gulf coast of Florida.
Voucher specimen: FLORIDA: Franklin Co.: Saint Vincent Island, open dry sand dunes, 25 Jun 1936, D.S. Correll 5583 (DUKE).
ACKNOWLEDGMENTS
We wish to thank curators and staff of the following herbaria for access to specimens in their care: BRIT,
DOV, DUKE, FLAS, FSU, GH, IBE, LSU, MISS, NCSC, NCU, NLU. We also thank Alan Weakley and the
Flora of Virginia Project for continued funding of herbarium research. David Rosen and Steven Hill signifi-
cantly improved the manuscript.
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Alabama.
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ATLAS OF THE VIRGINIA FLORA, 2007. www.biol.vt.edu/digital atlas.
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Alismatidae, Arecidae, Commelinidae (in part), and Zingiberidae. Oxford University Press, New York.
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Chester, E.W., B.E. WoFFORD, AND R. KRAL. 1997. Atlas of Tennessee Vascular Plants. Vol. 2, Dicotyledons. Misc. Pub. No.
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Esincer, J.E. 1974. A systematic study of the genus Kalmia (Ericaceae). Rhodora 76:315-398.
FERNALD, M.L. 1942. Additions to the flora of Virginia. Rhodora 44:380—381.
FRECKMANN, R.W. AND M.G. LeLoNG. 2003a. Dichanthelium. Flora of North America north of Mexico. Vol. 25, Magno-
liophyta: Commelinidae (in part): Poaceae, part 2. Oxford University Press, New York.
FRECKMANN, R.W. AND M.G. LELoNa. 2003b. Panicum. Flora of North America north of Mexico. Vol. 25, Magnoliophyta:
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Gate, S. 1944. Rhy pora, section Eurhynchospora, in Canada, the United States, and the West Indies. Rhodora
46:89-278, interrupted pagination.
GLEASON, H.A. 1952. The new Britton & Brown illustrated flora of the northeastern United States and adjacent
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Hauser, D.P. AND L. Lect. 1999. A survey of allozyme variation among three members of the Sagittaria graminea
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Haynes, RR. AND C.B. HeLLauist. 2000. Alismataceae. Flora of North America north of Mexico. Vol. 22, Magnoliophyta:
Alismatidae, Arecidae, Commelinidae (in part), and Zingiberidae. Oxford University Press, New York.
HiLsENBECK, R.A. 1989. Taxonomy of Yeatesia (Acanthaceae). Syst. Bot. 14:427-438.
1360 J lof the Botanical R h Institute of Texas 2(2)
HoLMGREN, P.K., N.H. HOLMGREN, AND L.C. Barnett. 1990. Index herbariorum, 8^ ed. New York Botanical Garden,
Bronx.
Jones, R.L. 2005. The plant life of Kentucky. University Presses of Kentucky.
Jones, S.B., JR. AND N.C. Cole. 1988. The distribution of the vascular flora of Georgia. Dept. of Botany, University of
Georgia, Athens.
KARAMAN-CASTRO, V. AND L.E. UngArscH. 2006. Boltonia. Flora of North America north of Mexico. Vol. 20, Magnoliophyta:
Asteridae (in part): Asteraceae, part 2. Oxford University Press, New York.
KARTESZ, J.T. 1999. A synonymized checklist and atlas with biological attributes for the vascular flora of the Unites
States, Canada, and Greenland. First edition. In: Kartesz, J.T. and C.A. Meacham. Synthesis of the North American
Flora, Version 1.0. North Carolina Botanical Garden, Chapel Hill, North Carolina.
KraL, R. 2002a. Fimbristylis. Flora of North America north of Mexico. Vol. 23, Magnoliophyta: Commelinidae (in
part): Cyperaceae. Oxford University Press, New York.
KraL, R. 2002b. Rhynchospora. Flora of North America north of Mexico. Vol. 23, Magnoliophyta: Commelinidae
(in part): Cyperaceae. Oxford University Press, New York.
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Asteraceae, part 3. Oxford University Press, New York.
LeBLono, RJ. 2001. Taxonomy of the Dichotoma group of Dichanthelium (Poaceae). Sida 19:821-837.
LEBLOND, R.J. AND B.A. Sorrie. 2001. Additions to and noteworthy records for the flora of the coastal plain of North
Carolina. Castanea 66:288-302.
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of the southeastern United States. Castanea 61:101-144.
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Carolina Press, Chapel Hill.
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manuscript, Department of Botany, University of Delaware, Dover, Delaware.
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405-410.
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the author
SORRIE, B.A. AND S.W. LEONARD. 1999. Noteworthy records of Mississippi vascular plants. Sida 18:889-908.
SORRIE, B.A. AND A.S. WeAKLEY. Notes on Kalmia angustifolia and K. carolina (Ericaceae) in Virginia and the Carolinas.
Unpubl. ms.
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90:1-23.
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TATNALL, R.R. 1946. Fiora of Delaware and the Eastern Shore. Soc. Nat. Hist. Delaware.
TAYLOR, RJ. AND C.E.S. TAYLOR, 1994. An annotated list of the ferns, fern aliies, gymnosperms, and flowering plants
of Oklahoma. 3rd edition. Biology Dept. Herbarium, Southeastern Oklahoma State Univ., Durant.
Thomas, R.D. ano C.M. ALLEN. 1993. Atlas of the vascular flora of Louisiana. Volume 1: ferns & fern allies, conifers,
& Monocotyledons. Louisiana Dept. Wildlife & Fisheries and The Nature Conservancy, Louisiana Field Office,
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Thomas, R.D. AND C.M. ALLEN. 1996. Atlas of the vascular flora of Louisiana. Volume 2: Dicotyledons. Acanthaceae
to Euphorbiaceae. Louisiana Dept. Wildlife & Fisheries and The Nature Conservancy, Louisiana Field Office,
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c e 1 $8 ADI d, N le £, "I al. " Iletens] Cent 1361
Tucker, G.C. 2002. Lipocarpha. Flora of North America north of Mexico. Vol. 23, Magnoliophyta: Commelinidae
(in part): Cyperaceae. Oxford University Press, New York.
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versity of North Carolina, Chapel Hill.
J lof the B ical R h institute of Texas 2(2)
1362
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J. Bot. Res. Inst. Texas 2(2): 1362. 2008
ANALYSIS OF THE COMPLETENESS OF VASCULAR PLANT RECORDS
IN FLORIDA
Justin K. Williams Anica Debelica
poa of Biological Sciences Department of Biology
Sam Houston State University Baylor University
Huntsville, = 77341-2116, USA. One Bear Place # 97388
Waco, Texas 76798-7388, U.S.A.
ABSTRACT
Using the speci lationship f la S=CA?, we predicted the species ricl bvaseula pi f I Nn A M M The
vl eee i n f de 1 1 ial’ 4 1 1 f de y gathered
hom licer records. Results indicate that 6796 of the Flanda counties have documented P Baa a that d or
exceeded preattiov Y ues The ud. counties TUR of the Florida) 1
onstitute data
constitute data
O E
Key Wonps: biodiversity, Florida, herbarium, inventory, mapping, modeling, species area relationship, species richness, vascular
RESUMEN
área S=CA?, se predijo 1
1 J 4 A
1
id A 1 1 1 1 T t t dad (339. de Florida) car n delad tari a dad
id £
d ies y por ello constituyen lagunas de datos en los inventarios florísticos del estado.
t r
Williams and Lutterschmidt (2006) presented a herbarium assessment model that provides a mathematical
and objective approach to identifying data gaps in documented species richness determined from a state’s
herbarium collections. The model (here coined Collections Records Assessment Model or CRAM) compares the
documented species richness value for counties within a state (determined from a database of herbarium
records) to the predicted species richness value (determined by the species area relationship formula).
Results generated isolate a county by determining if its documented species richness value falls above or
below predicted species richness. A county with documented species richness below the predicted value
is considered under collected and constitutes a data gap in the state’s records. Using CRAM, Williams and
Lutterschmidt (2006) analyzed Texas and demonstrated that for much of the state’s counties (88%) docu-
mented species richness values fell below predicted values. Given that documented species richness is a
direct function of collections (Preston 1948; Williams & Lutterschmidt 2006), counties with species rich-
ness values below predicted richness are counties with little or no specimen collecting thus representing
data-gaps in the state’s herbaria.
A state requires two data sources to perform CRAM. The first data source is a database or Atlas of a
substantial Gf not entire) portion of a state’s herbarium specimens. The database should be constructed in
such a way that the number of species documented per county can be reported. Data source two requires
that a state have a substantial number (15+) of published floristic inventories that represent a varying array
of area size (from State level down to a few hectares). Information needed from each published inventory
is the area of the study site and the number of species reported from that area. These values are necessary
for ad the constants C and z used in the Do. area o formula S=CA*. Species-area
relationship i ] las "one of community ecology "s few] (S I 76). ” The species- area relation-
ship simply states uñas as area increases species richness increases B 4 Lomolino 1998). The species
area relationship can therefore be used to estimate or predict the number of species within a given area. In
J. Bot. Res. Inst. Texas 2(2): 1363 — 1371. 2008
1364 Journal of the Botanical R h Institute of Texas 2(2)
the formuia S = species richness, A = Area, C = the number of species when area equals 1, this is defined by
the intercept of the slope for known values (those gathered from the floristic inventories) and z - the rate of
increase in species as area increases, this is defined by the slope of the known values (those gathered from
the floristic inventories; MacArthur & Wilson 1967). After surveying the literature and state herbaria it was
determined that Florida possessed substantial data in both sources. Consequently, we present an analysis
of the documented floristic completeness for the state of Florida.
METHODS AND MATERIALS
Published checklists and floras for regions with defined boundaries within the state of Florida were identi-
fied (Table 1) using the FloraS of North America project (Qian et al. 2007). From each checklist the number
of species was recorded and the geographic flora-coverage area (kilometers?) was calculated. Both metrics
were log transformed and entered into a database. The database was imported into SPSS 10.1 and linear
regression was used to determine the relationship between species richness (dependent variable) and area
(independent variable). From this analysis both the slope (z value) and the intercept (C value) for vascular
plants in Florida were determined (Fig. 1).
To determine predicted species richness for each individual county in Florida the C and z constants
were then applied to the Arrhenius log-log (log S = log C + log A*) model with A representing the area in square
kilometers for each of the 67 counties (Table 2) in Florida.
We then accessed the Atlas of Florida Vascular Plants (Wunderlin & Hansen 2008) datab d recorded
the taxa richness reported from each county in Florida. Note that we reported here taxa richness rather than
species richness. This is because: 1) Wunderlin and Hansen (2008) were thorough in their documentation
of both species and their intra-specific taxa (both sub-species and varieties) and 2) it is the actual number
of biological entities that we are concerned with rather than their systematic position. Whether a taxon is
recognized as a species or a variety does not diminish the fact that it ecologically unique from other taxa
and occupies space within the designated area under study.
RESULTS AND DISCUSSION
The constants z (0.17) and C (245) for vascular plants in Florida were determined using linear regression
(Fig. 1) of geographical area and documented species richness values gathered from the 27 representative
floristic inventories in Florida (Table 1). The determined value of z is within the accepted range of z values
(0.12—0.17) for vascular plants within continents (MacArthur & Wilson 1967). The rather high z for Florida,
in comparison to the entire range for terrestrial plants, is readily explained by the tropical physiognomy of
the state. C indicates a species richness of 245 species for any given square-kilometer area in Florida. Us-
ing the obtained C and z values, we then predicted species richness for each of the 67 counties in Florida
(Table 2).
A cubic lysis was used to compare documented (r° = 0.165) and predicted species richness
(re = 0.999) io. each of the 67 counties in Florida (Fig. 2). Counties with documented species richness that
approximate (here determined as within 9596 of predicted richness) or exceed predicted species richness
fall near, on or above the predicted regression line; counties that have an under representation of species
richness fall well below the predicted regression line (Fig. 2). This cubic regression model allows curators
and researchers to identify counties that are under collected/documented. Our results indicate that 22 (or
32.896) of the 67 counties in Florida fall well below the predicted line and are, therefore, considered under
collected counties (Table 2 and Fig. 3). The total area (38240 km?) occupied by these under collected coun-
ties covers 26.5 96 of Florida. Compared to the state of Texas, with only 29 out of the 254 counties well
collected, the state of Florida is very strong in the documentation of vascular plant richness. It is suggested
however, that effort be made to improve the collection and documentation of species richness for the under
collected counties in Florida, specifically Baker, Glades, Hendry, Lafayette, Okeechobee, and Union Cos.
(Table 2) which have documented less than 7096 of the predicted species for their county.
rin In.k.al: ,S H >L fo PI J 1365
4.00-
3.507
)
3.0071
Log | (Species Richness
3
f
2.00-
1 y 17 y T E 5 1 1
-4 00 -2.00 0.00 200 4.00 6.00
Log (sq km)
Fic. 1. Logarithmic relationship I pecies rict ] geographi Dot: each of the 27 ch klist and floras list Ai iiia
ificant relationship pecies rict { phical area (F = 39.737, df= 25, — 0.614). ricas
It is recognized that additional checklist for Florida probably exist, most likely in the form of unpub-
lished government reports, and that those used (Table 1) for this study are only published representatives
of the potential literature for Florida. Regardless, we feel that those used in this study represent well the
degree of variation needed to determine credible C and z va In particular the floras used represent an
astonishing array of area size, ranging from th tate with 144557 km? to a mere 0.01 km". In addition,
when C is calculated using only 14 of the floras (selecting every other e ese arranged from largest to
smallest starting with the entire state) the C value is 253, only an eight specie ifference from the 245 used
for this analysis. Indeed, eight species can make a substantial ene: when factoring in an increase in
area, however when using predicted species richness val d from a C value of 253 to determine the
completeness of the Florida plant record, there is no substantial difference in the counties isolated as when
using a C value of 245. We do accept that over time, and with it the consequent accumulation of additional
data, the values presented in this study (Table 2) may and probably will change. We stress, however, that it
is not our aim to present fixed species richness values per county, but rather to provide benchmark values
that can be used in focusing collection/documentation effort.
Williams and Luttershmidt (2006) demonstrated that in T
in general greater in counties that had herbaria and lower in counties without ba The implication is
A ter i in)
values WIC
J lof the B ical R h Institut f Texas 2(2)
1366
Taste 1. Published values of species richness fi lar pl | associated geograpt
Numberof Area (km2) Generalized Location Citation
Species
4144 144557.60 Florida State Wunderlin & Hansen 2008
943 582.75 Merrit Island, Brevard Co. Poppleton et al. 1977
899 46.54 Jonathan Dickinson State Park, Martin Co. Roberts et al. 2006
726 116.81 Myakka River State Park, Sarasota and Manatee Cos. Huffman & Judd 1998
604 9.06 ichetucknee Springs, Suwannee and Colombia Cos. Herring & Judd 1996
576 124.84 Waccasassa Bay, Levy Co. Abbott & Judd 2000
540 9.25 O'Leno State Park and northeast River Rise State Preserve Tan & Judd 1995
Alachua and Columbia Cos.
523 9.79 Little Manatee, Hillsborough Co Myers & Wunderlin 2003
480 3724 Timucuan ecological and historic preserve, Duval Co. Zomlefer et al. 2007
477 326.34 Fakahatchee Strand State Preserve, Collier Co. Austin et al. 1990
466 105.15 Bull Creek, Osceola Co. Huck 1979
449 20.20 Cedar Key, Levy Co. Amoroso & Judd 1995
422 56.20 Paynes Prairie, Alachua Co. Fasley & Judd 1990
393 743 Dog Island, Franklin Co. Anderson & Alexander 1985
372 730.38 Biscayne National Park, Miami-Dade Co. Stalter et al. 1999
360 9.32 Manatee Springs, Levy Co. Gulledge & Judd 2002
356 0.88 Gold Head Branch Ravine and del Uplands, Clay Co. White € Judd 1985
336 0.34 The Hammock, Dunedin, Pinellas Genelle & Fleming 1978
323 4.22 Gulf ae National Seashore, deu Key, Escambia Co. Looney et al. 1993
290 3.65 Fort DeSoto Park, Pinellas Co. Thorne 1995
289 9.51 Little Talbot Island, Duval Co. Easley € Judd 1993
237 1.27 Fort Matanzas National Monument, St. Johns Co. Zomlefer et al. 2004
233 0.46 Blowing Rocks Preserve, Martin Co. Richardson et al. 1992
180 0.75 Atlantic University Ecological Site, Broward Co. Austin 1990
168 810.67 Ocala National Forest, Marion Co. Mohlenbrock 1976
108 0.01 Turtle Mound, Volusia Co. Norman 1976
63 0.03 Chicken Key, Miami-Dade Co. Guala 1993
that botanists tend to concentrate their specimen collections in localities near their work or home base. The
phenomena of collecting near biological stations was termed the "Collector's Syndrome” by Soberon et al.
(1996) and later "The Botanist Effect" by Moerman and Estabrook (2006). Indeed, within Florida the coun-
ties with lower than expected species richness val ones without herbaria (Fig. 3). It is acknowledged,
however, that there are plenty of counties without herbaria that have documented species richness greater
than predicted values (Fig. 3). Pautasso et al. (2007) suggest that species richness values are higher in some
counties not because of the efforts of botanical collecting, but rather because humans tend to concentrate
in areas with higher bio-diversity (more plant species - more humans = more botanist = herbaria). They
demonstrated a positive correlation for all the counties in the Continental United States, showing that
Human population increased with an increase in plant species richness. The question of whether botanist
drive documented species richness or species richness drives human settlement is a “Chicken and Egg
argument that is better argued elsewhere. However based on our research we do believe that documented
species richness is a product driven by the efforts of botanist and not vice versa.
This paper is one of many in the recent effort to develop models that predict species richness values
for the vascular flora of the United States. The methods use a variety of techniques including GIS modeling
(Iverson & Prasad 1998; Jarnevich et al. 2006) and species area relationship (SAR) (McNeill & Cody 1978;
Buys et al. 1994; Williams & Lutterschmidt 2006; Qian et al. 2007). The aim in developing such models is
to provide benchmark SR values that can be used for a variety of conservation efforts including assessing
the completeness of museum collections (Williams & Lutterschmidt 2006; demonstrated here), determining
Tage 2. List of the 67 H | g | 2008) hness (g | from the formula S=CA3),
95% error.
County Documented Predicted Area Documented taxa
tana richness species (km. mi.) greater than
richness predicted richness?
Alachua 1547 943 2264 TRUE
Baker 583 880 1516 FALSE
998 921 1978 TRUE
Bradford 528 780 759 FALSE
Brevard 1174 968 2637 TRUE
Broward 1039 997 3122 TRUE
Calhoun 1070 875 1469 TRUE
Charlotte 772 906 1796 FALSE
Citrus 1137 879 1512 TRUE
Clay 985 884 1557 TRUE
Collier 1247 1090 5246 TRUE
Columbia 947 928 2064 TRUE
eSoto 670 893 1651 FALSE
Dixie 735 908 1823 FALSE
Duval 1231 923 2004 TRUE
Escambia 1586 899 1715 TRUE
Flagler 777 851 1256 FALSE
Franklin 1531 869 1410 TRUE
Gadsden 1238 861 1337 TRUE
Gilchrist 609 804 904 FALSE
Glades 490 923 2004 FALSE
ulf 761 871 1436 FALSE
Hamilton 579 860 1333 FALSE
arde 664 893 1651 FALSE
Hendry 527 989 2985 FALSE
Hernando 1218 849 1239 TRUE
Highlands 1095 970 2663 TRUE
Hillsborough 1608 973 2722 TRUE
Holmes 602 851 1250 FALSE
Indian River 742 857 1303 FALSE
Jackson 1460 950 2372 TRUE
Jefferson 935 883 1548 TRUE
Lafaye 444 868 1406 FALSE
Lake 1232 957 2469 TRUE
Lee 1275 929 2081 TRUE
Leon 1686 900 1727 TRUE
Levy 1236 984 2897 TRUE
Liberty 1345 935 2165 TRUE
Madison 642 905 1792 FALSE
Manatee 1120 916 1919 TRUE
Marion 1248 1044 4089 TRUE
Martin 995 872 1439 TRUE
Miami-Dade 1676 1083 5040 TRUE
Monroe 1135 964 2582 TRUE
Nassau 929 896 1688 TRUE
Okaloosa 1201 954 2423 TRUE
Okeechobee 563 923 2005 FALSE
Orange 1100 949 2350 RUE
Osceola 837 1013 3424 FALSE
Palm Beach 1042 1085 5113 TRUE*
1368 J lof the Botanical R h Institute of Texas 2(2)
Taste 2. continued
County ocumented Predicted Area Documented taxa
taxa richness species (km. mi.) greater than
richness predicted richness?
Pasco 1117 917 1929 TRUE
Pinellas 1173 774 725 TRUE
Polk 1294 1076 4855 TRUE
Putnam 1029 912 1870 TRUE
Santa Rosa 1270 968 2634 TRUE
Sarasota 995 876 1480 TRUE
Seminole 887 787 798 TRUE
St. Johns 857 886 1577 TRUE*
St. Lucie 686 876 1483 FALSE
Sumter 925 869 1413 TRU
Suwannee 606 904 1781 FALSE
| 805 972 2699 FALSE
Union 458 754 622 FALSE
Volusia 1311 981 2857 TRUE
Wakulla 1262 885 1571 TRUE
Walton 1315 974 2739 TRUE
Washington 837 878 1502 TRUE*
the degree of invasiveness (Jarnevich et al. 2006), and determining the degree of sampling effort needed to
complete a survey (Palmer et al. 2002). We believe that such models are important in directing future col-
lecting, research and granting efforts, as well as laying the foundation for testing theoretical models.
ACKNOWLEDGMENTS
We thank William I. Lutterschmidt for helpful comments and the initial review of the manuscript and
Richard P. Wunderlin for providing critical comments during the review process.
REFERENCES
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—
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Wurre, D.L. ano W.S. Jupo. 1985. A flora of Gold Head Branch ravine and adjacent uplands, Clay County, Florida.
Castanea. 50:250-261.
WiLLIAMS, J.K. AND W.I. LurreRsCHMIDT. 2006. Species-area relationships indicate large-scale data gaps in herbarium
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and K. N. Campbell (application development), Florida Center for Community Design and Research] Institute
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ZOMLEFER W.B., D.E. GIANNASI, AND W.S. Jupo. 2007. A floristic survey of Timucuan ecological and historic preserve,
Duval Co. J. Bot. Res. Inst. Texas 1:1157-1178.
1372 I nf 4h D HN E. PES
Journal of i f Texas 2(2)
b
BOOK REVIEW
DAVID CUTLER, Teo BOTHA, AND DENNIS STEVENSON. 2008. The Virtual Plant (Network CD-ROM). (ISBN 978-
1405169707, hbk.). Wiley-Blackwell Publishing, 2121 id ie na lowa 50014-8300, U.S.A.
(Orders: www.blackwellpublishing.com, orders lac] hi
p g.com, 800-862-6657).
$300.00.
that improves flotation in aquatic plants The ae Plant ee isa god source e of isual supr ] kes this approach
bd. en buo An Applied VANS When used; ina ird d one Mp gir the entire lab experience to be mo
enriching for both teacher and student, i 1, real-world microsco
E £ git Py:
Also included on the CD are a number of useful p j ] be ed zd adapted for ais purposes and more.— Natalia
Quinteros, Graduate Student, Texas Christian Universit 1B ical R h Institute of Texas, Fort Worth, Texas, U.S.A
From the Hae ue "The Virtual pei ro ROM is a oe to ps book idi nam an a ES ior Mure
This Manon version at he cD- ROM contains s high-quality Pa Ware ing electron on images. This key
S LS 1 J k f {+l 4 es t
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4 DEN Mox! Tem3Senls0az
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nth he CD-ROM, king thi 1 indisy bl for | d laboratory
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J. Bot. Res. Inst. Texas 2(2): 1372. 2008
SPECIES RICHNESS OF VEGETATIONAL AREAS OF TEXAS:
A FIRST APPROXIMATION
Michae! H. MacRoberts and Barbara R. MacRoberts
Bog Research, 740 Columbia, Shreveport, Louisiana 71104, U.S.A. and
erbarium, Museum of Life Sciences
Louisiana State University in t
Shreveport, Louisiana 71115, U.
ABSTRACT
Using the Atlas of the Vascular Plants ici Texas is ie peru n M assi area map of Texas, we analyzed the species rich-
lly rich and that richness does not correlate with
1
ness of the different areas of Texas.
primary productivity or size of area. oe iur the ratio of ts to di is not the same in the eastern and western parts of
1 - 1
the state. Other Ein also west Hut not in the east; E are nd more
1 i } f Asteraceae and F
common in the east than in the west;
equal in bth i aa W ] litional ional f Texas probably should be reassessed and perhaps a simpler
ME :
oo eo
vegetation p hat i t
RESUMEN
Usando el Atlas of the Vascular Plants of Texas y el mapa De s de áreas d VEgel ion me Texas, se ha on la ri-
A en iaa de Hs Entes áreas iu Texas. E
Al 4 "rece t 1 1 do J Taa x1 J; 14 dx
F E
ps a del estado. O bié tes: Las P dominan l iledó ]
(C 1 ^ 1 f. 1
T 1
oeste pero no en el este; 1 te; las Cactaceae son mucho más comunes en el
1 4 ] ca din 1 3
oeste que en el aste; Las A
is e 1 ET pa p m
y quizás elaborar un mapa de veg ple q g la actual
E
información sobre la distribución.
INTRODUCTION
It is well known that the different vegetational areas of Texas (sometimes called vegetational regions) are
not equally rich in species. For example, Diggs et al. (2006) have indicated that, while east Texas has 3402
taxa, north central Texas has only 2376 taxa and that the Great Plains, although one-fifth the land mass
of the United States, has only 3067 taxa. The latitudinal gradient in species richness is well known and is
as true of plants as it is of animals, but little is understood about the factors determining species richness
(Owen 1990; Thorne 1993; Ziv & Tsairi 2004; MacRoberts et al. 2007; Qian et al. 2007).
The Turner et al. (2003) Atlas of the Vascular Plants of Texas is primarily useful as a source of distribu-
tional information for individual species. However, it (and to a certain extent the Hatch et al. 1990 Checklist
of the Vascular Plants of Texas) can be used, with caution, as data for other types of studies. In this paper, we
use Turner et al. (2003) to determine the species richness of vegetational areas across Texas.
METHODS AND LIMITATIONS
The most often used vegetational areas map of Texas is shown in Figure 1 (Gould 1962, 1975; Correll &
Johnston 1970; Jordan et al. 1984; Hatch et al. 1990; Telfair 1999; Turner et al. 2003; Diggs et al. 2006).
Using the Turner et al. (2003) Atlas, we counted all species that occurred in each vegetational area. This
was accomplished by drawing each area on a clear plastic sheet of transparency film the same size as the
maps in the Atlas and overlaying the transparency on each of the approximately 5030 maps. If the species
(dot) occurred within the vegetational area, it was counted as occurring in that area whether or not it was
predominately found in another area. We did not question the validity of the traditional vegetational areas
nor did we question the dots on the distribution maps in the Atlas but accepted them, recognizing that
J. Bot. Res. Inst Texas 2(2): 1373 — 1379. 2008
1374 Journal of the Botanical R h Institut
f Texas 2(2)
DALLAM ; MANS-
vous
| lennan PORO
D en
pe q unl d
LC E gt FLA
| | EN 1
PARNER i meal | WALL
PALO-E PARKER |
: i
ISTEPHENS eie | P
e
far aniano) ERATH o ] Eus y >
, id
>
[emm msl T
Sa
EN | eir Taal
NG bu ^ w T
Y. b DAVIS ; A.
N “I
\ = | BREWSTER
nul
- s
Nu a E E ye. call T
X uxo ee
1. Pineywoods em] PN ae ha
2. Gulf Prairies and Marshes X 6 TUN mre
3. P k Savanna ne owa | ERR
4. Blackland Prairies T
5. Cross Timbers and Prairies esa; Hae en
6. South Te ins po aedi
7. Edwards Pla A,
8. Rolling Plains
9. High Plains
10. Trans-Pecos
Fic. 1. Vegetational areas of Texas (from Hatch et al. 1990).
the distribution maps are incomplete and there are undoubtedly mistakes in them, and that the traditional
vegetational areas are not universally accepted (MacRoberts & MacRoberts 2003a). We did not exclude
non-native species but assumed, probably incorrectly, that they would be about equally frequent in each
area; also recognizing that non-natives are part of the flora and are here to stay. We included data only at
the species level. There are many sources of error in the data from uneven collecting (areas most heavily
collected are near universities with herbaria) to the fact that Turner et al. (2003) *in positioning of dots
within counties, if only a single collection was noted ... usually placed the dot in the center of the county
concerned (except in the Trans Pecos region....).” We, therefore, assigned species to areas conservatively:
if a dot occurred on the vegetational area boundary we either checked with other sources (e.g., Hatch et al.
1990) to see if the problem could be resolved or counted it as occurring only in the areas already represented
by the species. That is, we assumed that it was on the side of the county that was in the vegetational area
already occupied. Thus, if an east Texas species occurred as far west as Travis County, and there was a single
dot in the center of Travis County where the Edwards Plateau and the Blackland Prairies meet, then the
Mar-Dal 4 IAA. -Dal- ts, $ > teh in T, 1375
species was not counted as occurring in the Edwards Plateau vegetational area but only in the area or areas
to the east. We did not encounter many of these border problems, but to have an independent assessment of
the accuracy of the Turner et al. (2003) maps, we compared them with the reported distribution of species
in Hatch et al. (1990) where there is no ambiguity in vegetational area reported. For this comparison, we
used the Pteridophytes and Gymnosperms in one sample and the Cyperaceae in another. We found a very
strong positive correlation between the Turner et al. (2003) and Hatch et al. (1990) samples (R? - 0.8648
and R? = 0.8023, respectively), supporting our use of the Turner et al. (2003) data as satisfactory for the
task at hand. In addition, the sample taken from the Turner Atlas is over 5000 species and ten vegetational
areas, which is a substantial amount of data. We, therefore, do not believe that the problem of “dot place-
ment" introduces a great deal of error as compared to other problems, such as uneven collecting among the
different vegetational areas.
We looked at the distribution of several plant families (Poaceae, Cyperaceae, Asteraceae, Cactaceae,
and Fabaceae) by vegetational area to see if any patterns of plant distribution were evident.
We used Owen and Schmidly's (1986) above-ground primary productivity (primary productivity index)
data for Texas to see if this correlated with species richness. We averaged their data for each vegetational
area to obtain a single figure for comparison with our richness numbers.
RESULTS AND DISCUSSION
The species area/vegetational area lt ized in Table 1. Texas vegetational areas are not equally
species rich. The Gulf Prairies and Marshes are the richest, the Trans-Pecos is second, and the Pineywoods
is third. The High Plains are the least rich.
Species ricl lat ly very weakly and negatively (r, = -0.3697) with size of vegetational area.
The Edwards Plateau ranked first in size but fourth in richness; the Rolling Plains ranked second in size but
ninth in richness, and so on.
Species richness correlates only very weakly but positively (r, = 0.3455) with primary productivity.
Gulf Prairies and Marshes rank first in species richness but third in productivity. Trans-Pecos ranks second
in species richness but tenth in productivity, and so on. These findings agree in general with those of Owen
(1990), who found for mammals evidence contradictory to the hypothesis that greater productivity is as-
sociated with greater species richness. Samples at other scales than those used by Owen or ourselves might
show different results, but this was not tested.
Breaking down the major figures somewhat, the monocot/dicot/Gymnosperm-Pteridophyte ratios
fell into two groups across Texas. For the eastern parts of Texas, monocots varied between 2996 and 3096,
dicots between 6796 and 6996, and Gymnosperms and Pteridophytes between 296 and 396. For the western
part of Texas, monocots varied between 19% and 24%, dicots between 74% and 77%, and Gymnosperms
and Pteridophytes between 296 and 4%. There is a significant difference (chi square = 116, 3df, p = .00001)
between these groups. Interesting here is that the number of grass species by vegetational area appears to
be practically the same ranging from 11.296 for the Pineywoods to 16.696 for High Plains, with the others
grouped closely between these (Table 2). However, when monocots alone are considered, grasses dominate
the west Texas vegetational areas but do not dominate the east Texas vegetational areas. In the four east-
ernmost areas of Texas, grasses constitute less than 5096 (range 36.996 to 47.896) of the monocot flora, but
in the six westernmost areas, they constitute more than 5096 (range 52.896 to 68.596). Cyperaceae show the
opposite trend (Table 3) and are much more common in the east than in the west. Asteraceae show only a
mild trend toward being more common in west Texas than in east Texas (Table 4). Fabaceae show virtually
no east-west trend (Table 5), but Cactaceae show a clear east-west trend (Table 6).
The reason for the differences in species richness among areas is not easily understood (Owen 1990;
Withers et al. 1998; Qian et al. 2007, see also discussion in Diggs et al. 2006; MacRoberts & MacRoberts
2008 in press), but presumably a combination of complicated interrelated factors (elevation variation,
precipitation and its seasonality, soil diversity, temperature extremes and averages, sunshine, geological
1376 J | of the Botanical R h Institute of Texas 2(2)
l If ge of pteridophytes, gymnosperms, monocot dicots, total species, area in millions ofha., and productivity ff getati Area data from
Hatch et al. (1990). Primary productivity index from 0 | Schmidly (1986).
Vegetation region Ptero. & Gymno. Monocots Dicots Total Area Productivity
Gulf Prairies & Marshes 43 (2%) 667 (29%) 1607 (69%) 2317 4.1 1905
Trans-Pecos 95 (4%) 411 (1996) 1688 (77%) 2194 73 295
Pineywo 58 (3%) 655 (30%) 1440 (6790) 2153 64 2161
Edwards Plateau 70 (3%) 482 (23%) 1535 (74%) 2087 10.3 906
Blackland Prairie Al (2%) 582 (29%) 1391 (69%) 2014 5.1 1848
Post Oak Savanna 38 (2%) 67 (30%) 1258 (68%) 1863 28 1981
Cross Timbers 38 (2%) 420 (24%) 1271 (74%) 1729 6.2 1397
South Texas Plains 28 (2%) 353 (23%) 1169 (75%) 1550 8.5 948
Rolling Plains 19 (2%) 284 (24%) (75%) 1205 9.7 794
High Plains 19 (2%) 216 (24%) 658 (74%) 893 79 529
Taste 2. Num! f ge of P by vegetational
Vegetational Area Number of species Percentage of total Percentage of
species in area monocots only
Pineywood 242 112 36.9
Gulf Prairies & Marshes 319 138 47.8
Blackland Prairies 277 13.8 476
Post Oak Savannah 246 13.2 43.3
Edwards Piateau 263 12.6 54.6
Cross Timbers 222 128 52.8
South Texas Plains 224 14.5 634
Trans-Pecos 254 11.6 61.8
Rolling Plains 168 13.9 59.2
High Plains 148 16.6 68.5
Tag 3. N ge of Gr y vegetational
Vegetationa! Area Number of species Percentage of total Percentage of
species in area monocots only
Pineywoods 205 9.5 313
Gulf Prairies & Marshes 168 73 25.2
Blackland Prairies 164 8.1 282
Post Oak Savannah 163 8.7 28.7
Edwards Plateau 93 44 193
Cross Timbers 90 5.2 214
South Texas Plains 58 37 16.4
Trans-Pecos 57 26 13.9
Rolling Plains 52 43 18.3
High Plains 32 36 148
complexity, etc.) is responsible, none of which is easy to measure. Intuitively, it would seem that areas with
relatively high seasonal temperatures and rainfall, e.g., the Pineywoods, would have high species richness.
At the same time, the environmentally diverse Trans-Pecos, with its great topographical relief and diverse
habitats that range from deserts to wooded mountain slopes, also would support a high number of species.
Areas with moderate to low habitat diversity, moderate to low rainfall, and i liate temperatures might
be expected to have fewer species.
Mar-Dal 4 Ja4.-Dal ts, 5 : Ha! in T, 1377
Taste 4. Number and percentage of Asteraceae by vegetational area.
Vegetational Area Number of species Percentage of total Percentage of
species in area dicots only
Pineywoods 248 115 17.2
Gulf Prairies & Marshes 265 114 16.5
Blackland Prairies 255 12.7 183
Post Oak Savannah 226 12.1 18.0
Edwards Plateau 178 133 18.1
Cross Timbers 235 13.6 184
South Texas Plains 210 13.5 18.0
Trans-Pecos 327 149 1933
Rolling Plains 179 148 19.8
High Plains 146 16.3 222
Taste 5. Number and y ge of Fabaceae by vegetational area.
Vegetational Area Number of species Percentage of total Percentage of
species in area dicots only
Pineywoods 162 7.5 113
Gulf Prairies & Marshes 167 72 10.4
Blackland Prairies 154 7.6 11.1
Post Oak Savannah 149 7.9 11.8
Edwards Plateau 164 79 107
ross Timbers 147 8.5 11.5
South Texas Plains 137 88 11.7
Trans-Pecos 159 7.2 94
Rolling Plains 106 8.7 11.6
High Plains 74 82 112
Taste 6. Number and percentage of Cactaceae by vegetational area,
Vegetational Area Number of species Percentage of total Percentage of dicots
species in area only
Pineywoocs 5 0.2 0.3
Gulf Prairies € Marshes 17 0.7 1.1
Blackland Prairies 8 04 0.6
Post Oak Savannah 7 04 0.6
Edwards Plateau 45 2.2 29
Cross Timbers 18 1.0 14
South Texas Plains 30 25 3.3
Trans-Pecos 75 34 44
Rolling Plains 18 1.5 2.0
High Plains 14 1.6 2.1
In the course of this work we recognized that plant species do not appear to pay much attention to
vegetational area boundaries. The vast majority of species are not confined to any one area but spill out into
adjacent areas. Thus, the most often used Texas vegetational area map (Figure 1), and its many derivatives
and modifications, does not appear to be very accurate; it is probably no more accurate than any of the ecore-
gional maps so far produced (see MacRoberts & MacRoberts 2003a for a discussion of various vegetational
schemes for the West Gulf Coastal Plain). For example, east Texas, although it is almost always mapped
1378 Journal of tanical R h Institute of Texas 2(2)
as several vegetational areas, has no floristic breaks (MacRoberts & MacRoberts 2003b). The Pineywoods
grades into Post Oak Savannah, which in turn grades into Blackland Prairie, and southward into the Coastal
Prairies and Marshes. These regions share 9896 of their flora (MacRoberts & MacRoberts 2004). Only a few
areas in Texas might be true breaks, one such being the southeastern part of the Edwards Plateau where the
Balcones Escarpment seems to be a floristic barrier. However, the Edwards Plateau grades northward into
the Cross Timbers, Rolling Plains, and High Plains, and westward into the Trans-Pecos. The main floristic
break in Texas is right down the middle of the state in a 300 km wide ecotone between about 96? and 99* W
longitude (MacRoberts & MacRoberts 2003b, see also McLaughlin 2007). Consequently, we believe that it
is time to re-think vegetational area mapping for Texas in light of the extensive collecting that has occurred
over the past half-century and to use the total flora as a basis for establishing vegetational areas.
ACKNOWLEDGMENTS
Amanda C. Lewis, Red River Watershed Management Institute, Louisiana State University in Shreveport,
aided with the figure and table. Robert Kalinsky aided with the statistics. Billie Turner and an anonymous
reviewer provided many helpful comments on the paper.
REFERENCES
Correll, D.S. AND M.C. JOHNSTON 1970. Manual of the vascular plants of Texas. Texas Research Foundation, Renner,
Texas.
Dicas, G.M, B.L. Lirscoms, M.D. REED, AND R.J. O’KENNON. 2006. Illustrated flora of east Texas. Sida, Bot. Misc. 26:1—
1594,
GouLp, EW. 1962. Texas plants—a checklist and ecological summary. Texas Agric. Exp. Sta. Misc. Publ. 585:1-
112.
Goutp, FW. 1975. The grasses of Texas. Texas A&M University Press. College Station.
Hatch, S.L., KIN. GANDHI, AND L.E. Brown 1990. Checklist of the vascular plants of Texas. Texas Agric. Exp. Sta. Misc.
Publ. 1655:1-158.
JORDAN, T.C., J.L. BEAN, AND W.M. HoLmes. 1984. Texas: A geography. Westview Press, Boulder, Colorado.
McLAuGHLIN, S.P. 2007. Tundra to tropics: the floristic plant geography of North America. Sida, Bot. Misc. 30:1-
58.
MacRostnrs, M.H. AND B.R. MacRoserts. 2003a. West Gulf Coastal Plain ecoregions. Sida 20:1247-1276.
MacRoserts, M.H. AND B.R. MacRoserts. 2003b. The east-west transition of flora in Texas: a biogeographical analysis.
Sida 20:1693-1700.
MacRoserts M.H. AND B.R. MacRoserts. 2004. The post oak savanna ecoregion: a floristic assessment of its unique-
ness. Sida 21:399-407.
MacRoserts, M. H., B.R. MacRosERTS, AND R. G. KauiNskv. 2007. Vascular plant species/area relationships (species rich-
ness) in the West Gulf Coastal Plain: a first approximation. J. Bot. Res. Inst. Texas 1:577—583.
MacRoserts, M.H. AND B.R. MacRoserts. 2008. The Big Thicket as floristically unique habitat. J. Bot. Res. Inst. Texas.
In press.
Owen, J.G. 1990. Patterns of mammalian species richness in relation to temperature, productivity, and variance
in elevation. J. Mammology 71:1-13.
Owen, J.G. AnD DJ. ScHMIDLY. 1986. Environmental variables of biological importance in Texas. Texas J. Sci.
38:99-117.
Qian, H., J.T. FRIDLEY, AND M.W. PaLmer 2007. The latitudinal gradient of species-area relationships for vascular plants
of North America. Amer. Naturalist 170:690-701.
TELFAIR, R.C. Il. (ed.) 1999. Texas wildlife resources and land uses. Univ. of Texas Press, Austin.
Thorne, R.F. 1993. Phytogeography. In: Flora of North America Editorial Committee, eds. Fl. North America
1:132-153. Oxford Univ. Press, New York and London.
RA Dal 4 IRA DAL ts, Sr H He! in T; 1379
Turner, B.L., H. NichoLs, G. Denny, AND O. Doron. 2003. Atlas of the vascular plants of Texas. Sida, Bot. Misc. 24:
1-888.
Wiruens, M.A., M.W. Palmer, G.L. Wane, PS. Wurre, AND PR. Neal. 1998. Changing patterns in the number of species in
North American floras. In: T.D. Sisk, ed. Perspectives on land-use history of North America: a context for un-
derstanding our changing environment. U.S.G.S., Ole ua Division. Reston, Virginia. Pp. 23-32.
ZN, Y. AND A. Tsaini. 2004. The species richness-productivity rel p: time to stop searching for a"true" pattern?
Community Ecology 5:177-180.
1380 Journal of the Botanical R h Institute of Texas 2(2)
BOOK REVIEW
RONALDO BARTHEM AND MICHAEL GOULDING. 2007. An Unexpected Ecosystem: the Amazon as Revealed by
Fisheries. (ISBN 978-9972-2912-4-1, pbk.). Missouri Botanical Garden Press, PO. Box 299, St. Louis,
Missouri 63166-0299, U.S.A. (Orders: www.mbgr info, 314-577-9594 fax). $35.00, 241 pp., color
photographs and illustrations, 9" x 10 1/2"
L 1 a 1 CAATTE Jr “47 ; 1 Lal ld E B voa: 1 fl H
r r e i
inf d to the rest of the world. Although I feel this book is f l 1 tł ientific mind, it is filled much
infi ion that will al in tł ler iust looking fi ] inf i l book about fish of the Amazon River
J O o
Á TT ld $ 1 ana 1 1 1 1 11 T 1 11 +1 1 LE H 1 Mis wr ey a Suy
L r O i r e
River. The book also d great job at describing both the good fishing methods and, unfortunately for the ecosystem, the bad fishing
n 1 id f. E J fol o4 s £, * 1 HE J «1 E: EE PERSE h A 1 D 1
methods. gar g particularly y gray I
book. They too help ext te tł ge being told in the text. I do feel the text is hard to follow in a sense of understanding all of
the information, but it does a good jo! 1 ibing h inf ion f 1 A O
my favorite features of the book is the descripti f the 29 un food fishes. Each fish has its own detailed de-
M md its PHA with detailed, color coded maps, as well g y patterns, sp g periods, and much more. Each
i quc 1 hal ela thi 1
r O 4 o
Not only will An nera E tem ed ] he fish in the Amazon River, but it will also "s inne much about bas
conservation of the river itself and much of the forest se it. I anes highly recommend th
the Amazon Rainforest. It is will enjoy sharing at your di table.— Kyle Rasmussen,
Intern, Botanical Research Institute of Texas, 500 East 4th Street, Fort Worth, Texas 76102-4025, U.S.A.
J. Bot. Res. Inst, Texas 2(2): 1380. 2008
CAREX BREVICULMIS (CYPERACEAB),
NEW TO THE FLORA OF NORTH AMERICA
Lucas C. Majure Charles T. Bryson
Florida Museum of Natural History and USDA-ARS
Department of Botany Southern Weed Science Laboratory
ju a ibd PO. Box 350
PO. Box Stoneville, Mississippi 38776, U.S.A.
Gainesville, pun > 1, U.S.A. charles.bryson@ars.usda.gov
Imajure@ufl.edu
ABSTRACT
brevicul j Asia, Australia, and the Indian Subcontinent, was pees E in uA in ee dud
idR U.S.A., at two areas in the oF of Meridian. Further
this s specie North i R i }
to a — A vane NEA Bons oni to year, and could oe peek We pae a detailed descriotiom and
Tux
E
RESUMEN
Carex breviculmis, especie nativa de Asia, Australia, y el Nuus d due adds en E ape Pop en N apo este
Trabajo
paa d ions de dan le EE. UU. a principios de 200
Aol»nna 1
Js: 1
Nb ME no era conocida de la
flora de Norte Améri tes de este d brimiento. Varios viaj iti 1 u tar bi pi
* J hl s ae A qu T
INTRODUCTION
Carex breviculmis R. Br., Ao-suge or blue sedge, was described from Australia in 1810 (Moore & Edgar 1976;
Walker 1976). It is a widespread species native to the far-east and is known from east Afganistan, Australia,
China, India, Japan, Korea, Malaysia, Nepal, New Guinea, New Zealand, Pakistan, Russia, and Taiwan
(Hooker 1894; Goncharov et al. 1964; Ohwi 1965; Moore & Edgar 1976; Walker 1976; Koyama 1978;
Egorova 1999; Kukkonen 2001). This species is a perennial that inhabits *open fields" and "sandy coasts"
(Ohwi 1965), "exposed grassy slopes and water courses" (Goncharov et al. 1964), *open grasslands, forest
margins" and other disturbed areas (Walker 1976; Koyama 1978), and mountainous areas with altitudes
ranging from 2000—3900 m (Hooker 1894; Kern 1976; Kukkonen 2001). Carex breviculmis forms a complex,
which can be found from "NW Himalaya to Japan and through Malaysia to Tasmania and New Zealand"
(Kukkonen 2001).
Carex breviculmis is highly variable (Walker 1976; Koyama 1978). A number of species have been
described within this complex that are now considered conspecific with C. breviculmis, e.g., C. leucochlora
Bunge, C. royleana Nees (Ohwi 1965; Koyama 1978) or as distinct species (Egorova 1999; Kukkonen 2001).
Additionally, numerous subspecies and varieties have been recognized in order to describe this variation
(Ohwi 1965; Merrill 1968; Walker 1976; Koyama 1978). Egorova (1999) treats C. breviculmis and C. leucochlora
as distinct species and separates them based on culm height, with C. breviculmis having culms from 1-2 cm
in height and many, distinct veins on the perigynia. Egorova (1999) also states that the type specimens of
C. royleana and C. hypochlora Freyn, are "identical" to C. leucochlora. Kukkonen (2001) places C. brevilculmis
ssp. royleana in synonymy under C. royleana, but mentions that the C. breviculmis comple, wich meee
C. leucochlora, needs revision. Obviously, there is a great deal of confusion about
within this group. Based on inspection of specimens of C. breviculmis and C. leucochlora at MO and based
J Bot Rec. inst Taxas 2(2): 1321 —1387. 2008
1382 Journal of the Botanical R h Institute of Texas 2(2)
on the morphology of our specimens and current nomenclature in Tropicos (2008), we determined that our
collections most closely correspond with C. breviculmis.
SPECIES DESCRIPTION
Carex breviculmis belongs to the species rich Old World section Mitratae and can immediately be distinguished
from all other species in North America except Carex caryophyllea Latourrette by the tubercle-like structure
at the achene apex. Prior to our discovery, the only species of Carex within this section known to U.S.A.
was C. caryophyllea. Carex caryophyllea, an introduction from Europe, was discovered in Massachusetts in
the northeastern U.S.A. in 1826; it is also found in D.C. and Maine (Standley 2002). Carex breviculmis is
easily distinguished from C. caryophyllea by its caespitose habit and lowest inflorescence bract longer than
the inflorescence. Carex caryophyllea is colonial from creeping rhizomes and has an inconspicuous lower
inflorescence bract shorter than the inflorescence. In Mississippi, the habit of C. breviculmis is similar to
species in section Acrosystis, but can be distinguished by its brown plant bases and long awned, green and
hyaline pistillate scales when compared to the purple plant bases and acute, purple, green, and hyaline
pistillate scales of C. albicans (willd.) Spengel var. australis (L.H. Bailey) Rettig and C. nigromarginata Schwe-
initz. In vegetative plants, C. breviculmis might be confused with C. blanda Dewey or C. leavenworthii Dewey,
both weeds of turf, lawns, flower beds, and disturbed sites (Bryson 1985; Bryson & Carter 2008). The leaf
color in vegetative C. breviculmis plants is a darker green than in C. leavenworthii and the widest leaf is much
narrower than the widest leaf in C. blanda. The following description is based on specimens collected in
Meridian, Mississippi during 2007 and 2008.
Carex breviculmis R. Br. Prodr. 242. 1810. (Figs. 1-2).
Plants densely caespitose from short rhizomes and fibrous roots; culms trigonous, 15-42 cm long, 0.7-0.9
mm wide, antrorsely scabrous; leaves dark to lime green, shorter than the culms, blades 3-20 cm long,
1.0-2.5 mm wide, leaf bases chestnut to light brown; inflorescence 3-4 spikes, pistillate spikes 23, 5.1-12
mm long, 3-4 mm wide, perigynia elliptic, 2.53.0 mm long, 0.8-0.9 mm wide, pubescent, 3-5 nerves
per face, beak 0.8 mm long, bulging midway, pistillate scales 4.4—4.8 mm long, 0.7-1.2 mm wide, ovate to
obcordate, acute to awned, awns 0.5-2.8 mm long, antrorsely scabrous, longest proximal; staminate spike
2.3-10 mm long, 1.2-1.8 mm wide, staminate scales 3.0-3.4 mm long, 0.8-1.1 mm wide, ar te orawn
to 0.5 mm long, antrorsely scabrous; bracts subtending lowest two pistillate spikes, sheaths 3-20 mm long,
front hyaline, bract blades 1.2-10 cm long, 0.8-4.0 mm wide, proximal bract longer than inflorescence;
achenes 1.4-1.6 mm long, 0.5-0.6 mm wide, trigonous, brown; stigmas 3, 0.5-0.7 mm long; style 0.5—0.7
mm long, swollen at base, base persistent forming swollen tubercle-like structure at achene apex; anthers
3, 2-2.2 mm long, 0.05 mm wide, filaments 2.8-3.2 mm long.
COLLECTIONS AND HABITAT DESCRIPTION
Carex breviculmis was first collected on March 23, 2007 from the Gypsy Cemetery in Meridian, Lauderdale
County, Mississippi (Fig. 3). In 2007, the Gypsy Cemetery was heavily mowed during the summer and most
of the C. breviculmis was observed growing under moderately shaded conditions beneath a large Quercus
nigra L. near a fenceline with much less maintenance than the interior of the cemetery. During a visit to the
Gypsy Cemetery in May 2008, C. breviculmis was observed in shaded and open areas growing in Cynodon
dactylon (L.) Pers. and Stenotaphrum secundatum (Walter) Kuntze turf and around grave markers. Additional
specimens were collected on May 5, 2007 from a site adjacent to Interstate 20 and a railroad that runs
perpendicular to Interstate 20 through Meridian. At this site, pua were growing in the middle of an old,
rarely used trail that parallels the railroad under shaded condi in sandy, mesic soils, where the canopy is
dominated by trees and shrubs approximately 15 years old. The vegetation is dominated by Albizia julibrissin
Durazz., Carex basiantha Steudel, C. digitalis Willd. var. floridana (L.H. Bailey) Naczi & Bryson, C. kraliana
Naczi & Bryson, Carpinus caroliniana Walter, Cercis canadensis L., Dichanthelium commutatum (Schult.) Gould
ssp. joorii (Vasey) Fernald, Fraxinus pennsylvanica Marsh., Hypericum hypericoides (L.) Crantz, Liriodendron
1383
£^» Mast. A e
TOT NUI LH AMEKA
(ctb) and A and C-F drawn from Majure 2334 (ctb).
Fic. 1. Hl
| of the Botanical R h Institute of Texas 2(2)
1384
r >——
d
dl
Ld
nos
banc
d
eher
buses d
Aube
ad
d
Mete,
m$
Lans d
d
—
—
3
IPP
Imis R. Br. coll ] in Lauderdale C ty, Mississippi, Majure 2334: A pl
Fic. (
C. base of plant.
tulipifera L., Lonicera japonica Thunb., Luzula echinata (Small) FJ. Herm., Morella cerifera (L.) Small, Rubus
trivialis Michx. and Symphyotrichum lateriflorum (L.) A Lóve & D. Lóve. On December 23, 2007, when the
first two sites were revisited, the densely caespitose clumps of this Carex species were still dark green and
had not been killed by winter frosts.
Early in March 2008 and again in May 2008, C. breviculmis was collected at yet another cemetery in
Meridian and from a highly disturbed vacant lot behind a fast-food restaurant. At the second and larger
cemetery, Magnolia Cemetery Perpetual Care, thousands of C. breviculmis plants were observed in open and
RA j Ip ; ( } iculmi for Nor th America ids
1
4,200 Kilometers
Fic. 3. The locati f C breviculmis R. Br. in Lauderdale C y, Mississippi, U.S.A
shaded areas, even growing out of cracks in cement, around large headstones and around poorly mowed
areas under large Magnolia grandiflora L. trees. The soils at both cemeteries ranged from sand to sandy loams,
while those from along the trai! and behind the fast-food restaurant were loam to silty loam soils.
We have observed a high degree of morphological variation in this species, even within the same
population. Plants growing out of cement, headstones or in compacted, drier soils tended to be smaller than
plants which were found near tree bases or in slight depressions. Also, the “lazy” culms of plants growing in
open, short lawn areas tended to fall over and lay flat on the ground, whereas those growing next to some
structure (cement, base of tree, etc.) tended to remain nearly erect or merely droop but not fall to ground
level. It appeared that this could be advantageous to the species when growing in heavily mowed areas, in
that sexual reproduction still could take place under such conditions.
Voucher specimens. U.S.A. MISSISSIPPI. Lauderdale Co.: Meridian at Gypsy Cemetery off of 8th St. at the junction with 40" Ave.
32.36384°N, 88.71913°W, 23 Mar 2007, L.C. Majure 2119 with C. Doffitt & J. Hill (MISSA, USMS, herb. Bryson); 10 May 2008, L.C.
Majure 3214 with C.T. Bryson (FLAS, TENN, US); C.T. Bryson 22690 with L.C. Majure (BRIT, CHSC, DAV, DOV, DSC, FSU, GH, IBE, JSU,
MICH, MISS, MISSA, MMNS, MO, NY, SWSL, TENN, ae USCH, USMS, VDB, VPI, VSC, WIN, herb. Bryson); S of 1-20 E and Merid-
ian adjacent to RR track just W of Mt. Bart of frontage road. from 31* Ave. S 32.34856°N, 88.71904°W, 5 May 2007, L.C.
Majure 2334 (DOV, FLAS, MISS, MMNS, MO, SWSL, VSC, herb. Bryson); 10 May 2008, L.C. Majure 3215 with C.T. Bryson (FSU, NCU,
US, USMS, WIS); C.T. Bryson 22697 with L.C. Majure (BRIT, CHSC, DAV, DOV, DSC, FSU, GH, IBE, JSU, MICH, MISS, MISSA, MMNS
MO, NY, SWSL, TENN, UARK, USCH, USMS, VDB, VPI, VSC, WIN, herb. Bryson); Meridian, Magnolia Cemetery Perpetual Care off
of 23", Ave. ca. 0.5 km N of 20" St. and ca. 200 m E of northernmost entrance 32.38801°N, 88.69965°W, 15 March 2008, L.C. ee
2877 (FLAS, MISSA); 10 May 2008, L.C. Majure 3213 with C.T. Bryson (BM, FSU, GA, LSU, TEX, US, USMS, WIS); C.T. B 22
L.C. Majure (BRIT, CHSC, DAV, DOV, DSC, FSU, GH, IBE, JSU, MICH, MISS, MISSA, MMNS, MO, NY, SWSL, TENN, UARK, USCH,
USMS, VDB, VIP, VSC, WIN, herb. Bryson); Meridian, Magnolia Cemetery Perpetual Care off of 23 Ave. ca. 0.5 km N of 20* St. and
ca. 250 m E from southernmost entrance = 38571N, 88. oo 10 May 2008, C.T. Bes a uds L.C. tdi dd ide Mur
ff4AfE R North adj t hind Arby
herb. Bryson); Meridian,
E of 18" Ave. S; 32.35774°N, 88. 68607°W, 10 May 2008, L.C. Majure 3198 with C.T. Bryson (FLAS, MISSA); C.T. Baise 22705 with L.C.
Majure (SWSL, herb. Bryson).
I UnfehkhrD tsi D h A
1386 Journal oft t f Texas 2(2)
DISCUSSION
The source of introduction of C. breviculmis into the southeastern U.S.A. can only be speculated at this point.
The city of Meridian was established in 1860 and has been an important railway center, where current in-
tersecting railroads joined the city to the south as far as Mobile, AL and to the west as far as Vicksburg, MS
(Allgood 1983). The C. breviculmis discovered in Meridian could have come into the country via goods being
shipped from coastal ports into the interior U.S.A. by railway. One population of C. breviculmis is directly
adjacent to the main railroad running through the city, and the other populations are located close to the
downtown area, where the railway is not very far away. Also, the Meridian Air Force Base (Key Field) is
within 1 km of one population of C. breviculmis. From the base, flights come and go from all over the world.
Key Field is currently the home of the Air National Guard 186% Air Refueling Wing and the 185" Army
Support Facility. It is possible that this species could have been introduced inadvertently as a *hitchhiker"
on air-crew, troops, airplane machinery or cargo from Asia.
Another potential source of entry is the Rose Hill Cemetery, more commonly known as the Gypsy
Cemetery, in Meridian. This cemetery has been the burial site for the Gypsy royal family since 1915 and
has received many international visitors (Turnage 1999). The once Queen and King of the Gypsies in the
U.S.A. and other Gypsy royalty are buried in the cemetery. Clothing and shoes of visitors or items brought
to the cemetery for placement on gravesites could have been contaminated with seeds of C. breviculmis.
Carex breviculmis is cited as a weed (Moore & Edgar 1976; Moody 1989; Simpson & Inglis 2001; Global
Compendium of Weeds list 2007; Bryson & Carter 2008) and has not been recorded rnamen
& Carter 2008) and does not seem to be in the nursery trade in the U.S.A. (A. A. Reznicek personal comm.).
In the four localities where C. breviculmis has been found in Meridian, three of the sites contain relatively
large populations ranging from thousands of plants at one cemetery to hundreds of plants at the other two,
smaller sites. This species appears to be weedy in two of the largest cemeteries in Meridian, where it was
common in C. dactylon and S. secundatum turf. Based on our observations, the size of populations, and the
distance between sites, we suspect that C. breviculmis has been in the Meridian area for several decades. Ad-
ditional surveys are needed to determine the distribution of C. breviculmis in Meridian and adjacent areas.
Further dispersal of achenes and live plants is likely on contaminated mowing, construction, and grave
digging equipment.
ACKNOWLEDGMENTS
We thank the University of Florida Herbarium (FLAS) for invaluable library access and MO for specimen
loans of Carex breviculmis (including C. leucochlora) and other closely related species in sect. Mitratae. We also
thank Monica Arakaki for reviewing our Spanish abstract. We also thank Richard Carter, Gary N. Ervin,
Robert F.C. Naczi, A.A. Reznicek, and David J. Rosen for helpful comments and reviews of the manuscript.
This work was supported in part by funding from the USGS Biological Resources Discipline @04HQAG0135)
to Gary N. Ervin, Mississippi State University.
REFERENCES
A GLoBAL Compendium Of Weeps. 2007. (http://www.hear.org/gcw/). Accessed 1 Oct 2007.
Au coob, V.C. 1983. Soil survey of Lauderdale County, Mississippi. United States Department of Agriculture, Soil
Conservation Service, Washington, D.C.
Bryson, C. T. 1985. Carex weeds of Mississippi. J. Miss. Acad. Sci. 30:10.
Bryson, C. T. AND R. CARTER. 2008. The significance of Cy Weeds. In: R.F.C. Naczi and B.A. Ford, eds. Sedges:
uses, diversity, and systematics of the Cyperaceae. Monogr. Syst. Bot. Missouri Bot. Gard. 108. St. Louis, MO.
Pp. 15-101.
Ecorova, T.V. 1999. The sedges (Carex L.) of Russia and Adjacent States (within the limits of the former USSR).
Missouri Botanical Garden Press. St. Louis.
GONCHAROY, N.F., KRECHETOVICH, V.l., KRISHTOFOVICH, A.N., KUZENEVA, O.l., Larin, I.V., PALIBIN, L.V., RosHevITz, PY., SERGIEVSKAYA, L.P,
1 Bryson, Carex breviculmi for North America 1387
SHISHKIN, B.K., SHTEINBERG, E.l., TSINZERLING, Y.D., AND YUzEPCHUK, S.V. 1964. In B.K. SHISHKIN, ed. Flora of the U.S.S.R. Vol. ill.
Israel Program for Scientific Translation Ltd., Jerusalem.
Hooker, J.D. 1894. The flora of British India. Vol. VI, Orchidaceae to Cyperaceae. L. Reeve and Co, London.
Kern, J.H. 1976. Cyperaceae. In: C.G.GJ. Van Steenis ed. Flora Malesiana, Noordhoff International Publishing,
Leyden, The Netherlands. Pp. 435-753.
Koyama, T. 1978. Cyperaceae. In H.I. Li, T. Lin, T. Huang, T. Koyama and C.T. DeVol, eds. Flora of Taiwan Vol. V,
Angiospermae. Pp. 191-372.
KUKKONEN, |. 2001. Cyperaceae. In S.I. Au and M. Qaiser, eds. Flora of Pakistan. No. 206. University of Karachi, Karachi
and Missouri Botanical Garden Press, St. Louis.
Merrit, E.D. 1968. An enumeration of Philippine flowering plants. Vol. | A. Asher & Co., Amsterdam.
Moony, K. 1989. Weeds reported in rice in south and southeast Asia. International Rice Research Institute,
Manila.
Moore, L. B. AND EDGAR, E. 1976. Flora of New Zealand. Vol. H. A.R. Shearer, Government Printer, Wellington, New
Zealand.
Onwi, J. 1965. In: FG. Meyer and E.H. Walker, eds. Flora of Japan. Smithsonian Institution, Washington, D.C.
Simpson, D.A. AND C.A. INaus. 2001. Cyperaceae of economic, ethnobotanical, and horticultural importance: a
checklist. Kew Buil. 56:257—360.
STANDLEY, L.A. 2002. Carex (sect. Mitratae). In: PW. Ball, K. Gandhi, RW. Kiger, D. Murray, J.L. Zarucchi, A.A. Reznicek,
and J.L. Strother, Flora of North America, vol. 23. Oxford Univ. Press. Newtons Pp. 553-554,
Tropicos. 2008. Missouri Botanical Gardens web site, http://www.tropicos.org/Name/9908579. Accessed May
31, 2008.
Turnace, L. 1999, Meridian, Mississippi — Graves: King and Queen of the Gypsies. http:/www.roadsideamerica.
com/tip/785. Accessed May 31, 2008.
WaLker, E.H. 1976. Flora of Okinawa and the Southern Ryuku Islands. Smithsonian Inst. Press, Washington, D.C.
1388 J lof the Botanical R h Institute of Texas 2(2)
BOOK REVIEW
KELLY ALLRED. 2008. Flora Neomexicana, I: the Vascular Plants of New Mexico. An Annotated Checklist
to the Names of Vascular Plants, with Synonymy and Bibliography. (no ISBN, hbk.). Published
on demand m author. (Orders: www.lulu.com). $60.00, 617 pp. 6" x 9".
Having} : 1 hahh iated catalog of Col lo FI Weber & ma re LE cen
L2 1 Es 1 1
M numeral "T" following the title.
ul pans, bn: at least lichens ind Spe Let us hope that the reject prospers. Most popular esa books create a
I vascular pl Il of botanical biot
Tis 1 $ i T MA 1 1 h d P 1 e d
fth ft j its geolcgical history, soils, ecoregions i andal f lyi 1 f
[24 O A u O * e T EG O
A1 A 1 312 1 za4 bh f el e E, 1. MA ba m A a ] +1 D LT AL prts J 1 1
y y Survey gh 1915 MN
the lives of the prominent field pa who scoured the terrain. A special section lists species of special concern, dde aliens,
tati J 1 * 1 :
ds, Į topics.
The checklist treats e major groups of plants: Ferns and Fern ned Gy mnosperms, Dicots, and Monocots, each with their
families in alphabetical order. There is a page-index to the f. genera. The ca Ge while necessarily D: the scientific
hat the author prefers, does not decree its proper usage, for th New Mexico,
citing the Hu in which they have been used. Tus one is is to select th iti pihen user's taxonomic philosophy. For
example, in the Ásteraceae, under the ni l li Under each genus, one or more citations
lead one to pertinent papers. R ly abbreviated) justify inclusi fth the list
Unfortunately, in these times, such log bsol it is published. The Vus PNG America, now in pub-
lication is. ill y for f 1 j d professional—to decide ta
where the n discipli f cladisti d molecular biolog boldly decree the validity of ging inf i i
with phylogeny I hold oa floras by to bon as i ny can with traditional phological fi implyi lationships. The Pa
g gl ] jority of practical botanists. We find it interesting to seu
what the new school ying al luti y descent, b hard for practiti heir findi
method.
In ine I feel ed ick: dica onene in such a state of flux, I can Hence) E the diis in s field to select
I d by their After all
of a point of view. They are not etched in stone. And the Catalog of the New Mexico Flora is precisely what the practitioner RR to do
satisfactory wor
J cy a oe E 1 1
a es oe its bn) may be NA isan bots o one. How many
opy literature. All of the names listed here
L
hi At
copy
are backed up with hers i hert
My critici i hey simply speak to things that I wish were anaes or omitted. Most of tl Id have doubled the
size of the book. I mi ing ing | I deplore t! habit of “common name” for all species. Who will ever
use them? Lloyd Shinners Id in his grave. Instead of all tl ious “common” names, I would have been more interested
in seeing the Spanish names oe common xD i is uses medicinal or ini S Iam d to see iu the phy P ae. tax-
onomies created by cladi g y ji were the last I miss
literature citations of the families, genera, and species. I miss speci itations for species | f ly one or a few collections,
and the herbaria housing the vouchers.
The interior western states are a remnants e a brun iesu We a the geal herbaria, a comrades-in-arms,
amateurs i E on the earlier
settled east and west coasts. The area is nene, th f activity small, ou the iuis ru yielding m new a and ca Here
f 1 d 1 1 [y : 341 F
many of us have : pp g g g
M 1 d kicking Jct : 1 141 1 J 1 fend 1 L d William
A. Weber, Prof. Emeritus, Curator, University of Colorado tama. Boulder, Colorado 80309, U.S.A., bill weber @colorado. S
J. Bot. Res. Inst. Texas 2(2): 1388. 2008
VASCULAR FLORA OF C. BICKHAM-DICKSON/RED RIVER EDUCATION AND
RESEARCH PARK, CADDO PARISH, LOUISIANA: AN OXBOW LAKE COMMUNITY,
WITH COMMENTS ON EXOTIC/NATIVE SPECIES RATIOS
Michael H. MacRoberts and Barbara R. MacRoberts Gary M. Hanson
Bog Research, 740 Columbia, Shreveport, Louisiana 71104 U.S.A. Red River Watershed Management Institute
Herbarium, Mu f Life Sciences Louisiana State University in Shreveport
Louisiana State University in Shreveport One University Place
University Place Shreveport, Louisiana 71115-2399 U.S.A.
Shreveport, Louisiana 71115-2399 U.S.A.
Red River Watershed Management Institute
Louisiana State University in Shreveport
One University Place, Shreveport, Louisiana 71115-2399 U.S.A.
ABSTRACT
A floristic inventory was conducted as part of a brad assessment o the a eri kson/Red River Education and Research
Park, Caddo Parish, Louisiana. The goal was to bow lake area. An additional goal was
to provide an overview of disturbance history and the exotic species of the flora. The flora consists of 318 species of which 68 (21%)
are not natives.
RESUMEN
lizó un i io florísti parte d evaluación biológica del C. Bickham-Dickson/Red River Education and Research
f
Park, "Cada Parish, Louisiana. El objeti un listado base de la flora de esta área lacustre. Un objetivo adicional fue aportar
una visión de conjunto de la historia de las perturbaciones de la flora y especies exóticas. La flora consiste en 318 especies de las que
68 (2196) no son nativas.
INTRODUCTION
River systems have und lrasti lification since the beginning of agriculture. For example, the Nile
and its flood plain have been under intense human pressure and, consequently, modification for thousands
of years (De Villiers 2000). In the New World, both Native Americans and Europeans often selected rivers
and their floodplains for farms, plantations, and cities. Whether we speak of oe or St. ani we can
lification early y on. Today,
see that these sites were desirable and came under human occupation and thus
there are few major river systems or even parts of major river systems that approach Sue conditions
(Noss et al. 1995; Outwater 1996; DeVilliers 2000).
The Red River is no exception. Caddo Indians inhabited its floodplain and followed later by European
settlers and their African slaves (Perttula et al. 2008). The first ecologist/botanist to study the Red River
was Peter Custis, naturalist on the ill-fated 1806 Red River Expedition (Flores 1984; MacRoberts et al.
1997; MacRoberts & MacRoberts 2004; Hardy 2008). Only two of Custis plant specimens survive, but the
descriptions provided by both Custis and Thomas Freeman (leader of the expedition) are tantalizing: prai-
ries, swamps, cedar forests, and canebreaks lined the Red River. The next important ecological information
about the Red River's ecology was the land survey of the 1830s (MacRoberts & MacRoberts 2005). A recent
conference, The Freeman and Custis Red River Expedition of 1806: Two Hundred Years Later, focused attention
on the Red River and Upper West Gulf Coastal Plain and concluded that very little is known biologically
about it and that no inclusive study has been undertaken to elucidate the ecology of this system since the
early nineteenth century (Bragg 2008; MacRoberts & MacRoberts 2008).
In this paper, we undertake to describe the floristics of the C. Bickham-Dickson/Red River Education
and Research Park (Park hereafter), a highly modified oxbow system on the Red River floodplain. We also
J. Bot. Res. Inst. Texas 2(2): 1389 — 1406. 2008
1390 Journal of the Botanical R h Instit f Texas 2(2)
briefly consider the relationship between site disturbance and native/non-native (exotic) species ratios:
"non-native" or "exotic" referring to introduced species from outside the United States.
There were very few exotic species in North America in 1491. Only cultivars brought to North America
from Central America and Mexico were imports. Now, 20 to 35 percent (depending on author) of the 20,000
odd species found in North America are exotics from all parts of the world (Stuckey & Barkley 1991).
While there is an extensive literature on exotic species (Elton 1958; Simberloff 2004; Sax et al. 2005)
and much data on the relationship between percentage of exotic species in a flora and the overall size of the
flora, there is little quantitative data and even less experimental data on the relationship between disturbance
history and the exotic/native flora. Nonetheless, it is widely recognized that highly disturbed sites such as
cities and urban parks have a high percentage of exotics (ca. 20-60%); wl listurbed sites have much
lower percentages (ca. 0-10%) (Fox & Fox 1986; McIntyre & Lavorel 1994; Basinger et al. 1997; Stalter et
al. 1997; Stapanian et al. 1998; Lonsdale 1999; McKinney 2002, 2006; Harcombe et al. 2007). Percentage
of exotics, therefore, can be used as an indicator of degree of site disturbance (Gilliam 2007).
STUDY SITE
The Red River originates in the Great Plains and flows 2,200 km east and southeast until it reaches the
Mississippi River in central Louisiana. Red and brownish red silt and clay particles, eroded from mainly
Triassic and Permian rock, give the river its distinctive color. From north central Texas to the Mississippi
River, the river is sluggish with a low-lying, flat, extensive, Pleistocene/Holocene floodplain valley five to
thirty km wide with oxbows, sloughs, and backwater swamps. Flanking the floodplain at its lower reaches
are woodlands dominated by oak, pine, and hickory. As the glaciers receded and sea levels rose, the gradient
of the river lessened, causing it to braid and meander widely, resulting in deposition of alluvium that we see
today. The floodplain slope is gradual (Newkirk & Mueller 1980; Triska 2008).
Hunters entered the Red River area at least 12,000 years ago near the end of the last glaciation, long
before the present climate and biota were established. Horticulture came much later, probably no more
than 3000 years ago. The effect of Native Americans on the ecology is not known, but there is considerable
debate on this matter (e.g., Mann 2005). The Red River was first seen by European explorers in the sixteenth
century. Its lower reaches were colonized by the French in the eighteenth century, but because the “Great
Raft”—an immense logjam—blocked the river, its upper part was not fully explored until the nineteenth
century (Flores 1984). In 1806, the Freeman and Custis expedition went through and around the raft to near
the present day border of Oklahoma and Arkansas (Flores 1984; MacRoberts & MacRoberts 2004; Hardy
2008). Marcy and McClellan (1854) completed the exploration of the river in the mid-nineteenth century.
The raft, which extended in the mid-eighteenth century from Natchitoches to Shreveport was cleared by the
1850's but immediately reformed above Shreveport and grew by accumulation of driftwood to almost the
Arkansas border by 1873. The raft dammed the river, causing extensive flooding of low-lying areas and the
creation of raft-lakes. Cottonwood, cypress, and red cedar apparently made up the bulk of the raft, and older
sections were covered with herbaceous plants and shrubs. The raft was finally completely cleared by 1873,
with the result that raft-lakes drained and the River was navigable. Rapid exploitation of the floodplain and
the establishment of farms, plantations, towns, and cities followed (Humphreys 1971; Flores 1984; Triska
1984, 2008; Bagur 2001).
Over the last two centuries, the river and its floodplain ecosystems have undergone drastic modifica-
tion. Much of the landscape was converted to farms and plantations and eventually the river was modified
by leveeing, straightening, and damming. The floodplain was severely modified by agriculture and urban
sprawl. The once continuous forest of varied communities that covered hundreds of thousands of hectares
is now virtually gone. Cypress swamps, willow riverbank shrublands, and cottonwood forests hang on
while some floodplain communities such as cedar forests, cane brakes, and prairies have vanished entirely
(MacRoberts et al. 1997; MacRoberts & MacRoberts 20082).
Although floodplains in general have received substantial phytogeographical and ecological attention
MacRoberts et al., Vascular flora of C. Bickham-Dickson/Red River Park 1391
and there are numerous generalized descriptions and classifications according to hydrologic conditions,
soils, and vegetation (e.g., Sharitz & Mitsch 1993; Messina & Conner 1998; Mitsch & Gosselink 2000), the
Red River floodplain is poorly known, especially floristically (see Newkirk & Mueller 1980). Aside from
checklists of parishes and communities along the Red River, its last floristic survey was in 1806. It has
never been the subject of an ecological assessment (MacRoberts & MacRoberts 2008b). The most important
literature dealing with local ecology is small. MacRoberts (1979) and MacRoberts and MacRoberts (2006)
produced a flora of Caddo Parish. Van Kley and Hine (1998) described the wetland vegetation of Caddo
Lake—a remnant raft-lake. There have been several studies of floodplain forests in eastern Texas (e.g., see
literature in Mundorff 1998). MacRoberts and MacRoberts (2005), using original land plats, described the
1830s woody vegetation of Caddo Parish. Diggs et al. (2006) and Van Kley (2006) summarized the com-
munity types of eastern Texas and western Louisiana. Ware (1956) briefly described the vegetation on a
sand bar near Natchitoches, and Teague and Wendt (1994), concentrating on Bossier and Caddo parishes,
conducted the first parish-wide comprehensive survey of high quality natural communities in Louisiana,
only two of which were floodplain communities. Two non-quantified reports round out the list: Palmer
(1923) provided an anecdotal report of the Red River forest at Fulton, Arkansas, and the Henderson State
University Biology Department (1979) prepared a modest “biological inventory of the Red River waterway”
for the Corps of Engineers.
The Park is located between the campus of Louisiana State University in Shreveport and the Red River
(Sexton-Gordon 2004) (Fig. 1). It is 236 ha and includes an 81 ha oxbow lake created in the spring of 1945
by the change in course of the river during a flood. The Park ranges in elevation from about 44 to 49 m above
sea level. It periodically floods and its oxbow lake is only a few meters deep. To the west is a man-made
levee with adjacent barrow-pit and to the east is the Red River, which is now impounded and is in reality a
linear lake. Natural levees occur adjacent to the river and the oxbow. The floodplain is about 15 km wide
at this point. The Park was once within the area of the Great Raft. The part of the raft adjacent to the Park
was cleared between 1833 and 1837.
The Park shows the classic geologic structure of river meander systems and oxbow lakes (Sharitz &
Mitsch 1995). The river channel meanders through the floodplain, transporting, eroding, and depositing
alluvial sediments. Natural levees adjacent to the channel result from deposition of coarser sands when
floods overtop the banks. Point bars form from sedimentation on the convex side of river curves and soon
support vegetation, which stabilizes the soils. At the same time, on the concave (cut bank) side of the river
the relatively fast water moving cuts away the bank. s the river eae across the floodplain, meander
scrolls (ridges and swales) form, giving th j y. Oxbow lakes form when the meander
cuts off at its neck, and a new channel forms. The deposed ei are generally described as “riverwash:”
“recently deposited, slightly acid to moderately alkaline sand, silt, and clay.” “Most of the material is sandy,
but there are many thin layers of clay and silt. The profiles are not uniform from place to place. At frequent
intervals during floods, fresh material is deposited or several feet of material is washed away. In this man-
ner the upper layers change from one year to another. The slopes are uneven. The slope range is generally
from less than 1 percent to 3 percent, but there are some short slopes of as much as 5 percent” (U.S.D.A.
Soil Conservation Service 1959:41—42).
The land that encompasses the Park first came under individual ownership with the 1801 Spanish
Land Grant to Francois Grappe, son of a French father and Caddoan mother, born at La Harpe’s Post north
of the Great Bend probably in what today is Bowie County, Texas. Grappe was a trader, interpreter, guide,
and cattle runner. He was centrally involved in the Freeman and Custis Red River expedition as guide and
interpreter or supplier of interpreters and guides to the ill-fated expedition. The grant given to Grappe con-
sisted of four leagues along the Red River in what is today south Caddo Parish. The grant was recognized in
1835 in the Caddo Cession and passed to Grappe’s sons, who in turn sold it to Jahiel Brooks, Caddo Agent,
who in turn managed to lose it to creditors by the 1850’s. Neither Grappe nor his sons ever lived on this
land nor is there any evidence that they exploited it in any way. The first white settler of what is today the
1392 the Botanical R h Institute of Texas 2(2)
Fic. 1 Anal Diei} mal ¡Dar D: EA Pr 1D h Dl 4 CI A I ppr c IT CA CL I Photo
DL L nes £1 or aD: Minos I Dahi- Dales? ISIS
Park appears to have been Samuel Norris. Norris, since the 1820's, had lived on "Rush Island”—a part of
the Grappe claim, either in the Park or just south of it. What Norris used the land for is not entirely clear
but apparently hogs, cattle, and farming were involved. His claim was eventually disputed and it too went
to creditors at the same time as did Brooks’. By the 1890s, the land had become part of the Fullilove Planta-
tion (O'Pry 1928; Flores 1984; Henrici 1985). The site has subsequently been cleared, grazed, plowed and
planted with forage crops, and used as a horse stables and riding area.
In 1839 at the time of the General Land Office land survey, the Red River was a little to the east of its
present position at the Park. By 1890, it had meandered halfway across the Park and was moving steadily
west. In 1945, its channel ran along what is now the oxbow lakes in the Park. But during the spring of that
year, the river overflowed and cut a new channel through the narrow neck of land to the east of the park and
established its present channel (Advisory Committee Report 2002). Thus, geologically the area of the Park
is not old. With the exception of those soils on its extreme western edge, all the soils have been reordered
over the last two centuries.
Construction of the Red River Navigation Project, which was authorized by Congress through the Rivers
and Harbors Act of 1968, led to the development of five locks and dams downstream. When the Corps of
Engineers completed the last structure, Lock and Dam 5 about 30 km downstream in late 1994, the average
river level at ilie Park was increased to a post-construction pool stage of 147 feet above mean sea level. The
lake level is currently erratic and depends on river flood stage and other internal factors.
Teague and Wendt (1994) give additional background information on climate, geology, and history of
the area. Prior to the construction of locks on the river in the 1990s, July to December was the period of
lowest water, and January to June the period of highest water. Rainfall is approximately 115 cm per annum.
The City of Shreveport and the State of Louisiana jointly purchased the site in 1980 and named it C.
MacRoberts et al., Vascular flora of C. Bickham-Dickson/Red River Park 1393
Bickham Dickson Park. The Park was used solely for recreational activities such as sport fishing and boating
until the mid-1980s when Louisiana State University in Shreveport began biological and geological research
at the site. In 2000, the City entered into an agreement with Louisiana State University in Shreveport un-
der which the University would be allowed to utilize the park as a "living laboratory" for natural resource
education and research. Under the agreement, Louisiana State University in Shreveport would manage the
Park as Red River Education and Research Park. Research at the Park, which also includes the development
of innovative environmental monitoring technologies, has been conducted as part of the ongoing environ-
mental studies at Louisiana State University in Shreveport, and other academic and research endeavors of
the university.
The Red River floodplain vegetation in and surrounding the Park consists primarily of what has been
called *batture" (Lester et al. 2005), "Populus deltoides temporary flooded forest alliance" (Weakley 1998), or
seasonally flooded river floodplains, and semi-permanently flooded swamps (Van Kley 2006). From one side
of the Red River floodplain to the other, the following woody species occur: cottonwood (Populus deltoides),
green ash (Fraxinus pennsylvanica), sycamore (Platanus occidentalis), black willow (Salix nigra), hackberry
(Celtis laevigata), boxelder (Acer negundo), pecan (Carya illinoinensis), rough-leaf dogwood (Cornus drummon-
dii), persimmon (Diospyros virginiana), mulberry (Morus rubra), bois d'arc (Maclura pomifera), swamp privet
(Forestiera acuminata), cypress (Taxodium distichum), button bush (Cephalanthus occidentalis), honey locust
(Gleditsia triacanthos), and elm (Ulmus spp.). Historically, there were virtually no oaks (Quercus spp.), pines
(Pinus spp), sweetgum (Liquidambar styraciflua), magnolia (Magnolia grandiflora), or hickories (Carya spp.)
other than pecan on the floodplain. The species directly adjacent to the river itself were typically also at
the farthest edge of the floodplain just before the uplands are reached. At the floodplain edge, there is an
abrupt elevation change (10 to 30 m) and a vegetational transition in which pines (Pinus), oaks (Quercus),
hickories (Carya), magnolia (Magnolia), and sweetgum (Liquidambar) dominate and floodplain species drop
out. These species were present in 1830, and they are present today (MacRoberts & MacRoberts 2005). The
edge of the floodplain is often marked by a bayou (e.g., Bayou Pierre) just before the uplands and the floristic
transition.
METHODS
Beginning in February 2003 and continuing through November 2007, we regularly surveyed the Park for
flora by walking through all areas. Specimens collected, identified using local Floras (e.g., Smith 1994;
Diggs et al. 2006), and deposited at Louisiana State University in Shreveport herbarium. Nomenclature
generally follows Kartesz and Meacham (2005). We surveyed the Park for community types.
We reviewed the botanical literature for studies of the flora of mm land use areas SR as parks,
$
ic ratios
research stations, natural areas, counties, states, etc., to obtain data on d
The review was not exhaustive but only meant to sample such areas. We confine our list to studies conducted
during the last decade because, as Small and McCarthy (2001) point out, there are an ever increasing number
of exotics in the flora.
RESULTS AND DISCUSSION
We found 318 species at the Park, 68 (2196) of which were exotics. These are listed in the Appendix.
The main plant associations that we have separated in the Park are:
Willow forest.—Willow (Salix) forest, which sometimes includes sycamore (Platanus occidentalis) and
cottonwood (Populus deltoides), occupies moist to wet soils in periodically flooded river and lake edges and
other shallow wet areas. It is this community that colonizes sandbars and other recently scoured areas.
Lake side/inundated non-scoured areas.—This is a continuation of the previous community but is
associated with periodically flooded/wet areas such as e edges and swales. Many species occur in this
area including willow (Salix), button bush (Cephalantl talis), boxelder (Acer negundo), cy press (Taxo-
dium distichum), and sycamore (Platanus ee in small numbers. Grasses and sedges and many other
1394 Journal of the Botanical R h Institute of Texas 2(2)
herbaceous plants also occur. Here also is vegetation growing on floating logs, tree stumps, and the base of
those woody species that inhabit lake margins, such as button bush.
Open water with floating vegetation.—Duckweed (Lemna, Spi
green matt on the lake surface. Mosquito fern (Azolla caroliniana) can be common. Water-lotus (Nelumbo lutea)
is dense in one area of the oxbow lake. Coontail (Ceratophyllum demersum) and hydrilla (Hydrilla verticillata)
occur in many areas, usually in shallow water where they can become very dense. Algae sometimes form
mats on top of shallow-growing Ceratophyllum.
Sandbar.— Because the river is now controlled by dams and locks, natural sandbars do not form. Old
aerial photographs show sandbars adjacent to the Park. Willow (Salix nigra and S. exigua) forest is the com-
munity that first colonizes sandbars.
Cottonwood-sycamore-hackberry-ash-pecan forest.—This forest type domi he middle to higher
elevations of the Park, notably the ridges and levees. Any combination of these species, with lesser amounts
of honey-locust (Gleditsia triacanthos), bois d'arc (Maclura pomifera), and boxelder (Acer negundo), can occur.
Willow is also found here in swales. The understory is predominately rough-leaf dogwood (Cornus drum-
mondii). The herbaceous groundcover is sparse, but poison ivy (Toxicodendron radicans) and Juncus and sedges
can be common.
Open (Old) fields.—Open fields make up a large p ge ofthe Park. These were previously pasture
and were planted with legumes and grasses for hay and pasturage. This community contains the highest
number of exotic species. The fields typically have scattered cedar (Juniperus virginiana), bois d'arc (Maclura
pomifera), and honey-locust (Gleditsia triacanthos).
Ruderal.—The Park contains other areas, such ds/road sides, mowed areas, ragweed (Ambrosia tri-
fida) “forests,” and in some places, dodder (Cuscuta) covers large areas. These areas are highly disturbed.
We found many studies of undisturbed through highly disturbed sites in which the total flora and
number of exotics were given (Table 1). We graph these in Figure 2 by land use type and size of area simply
to make the data in Table 1 more accessible
As we have pointed out, received wisdom is that there is a correlation between percentage of exotics in
a flora and site disturbance (McIntyre & Lavorel 1994; Stapanian et al. 1998; Withers et al. 1998; Lonsdale
1999; McKinney 2006; Gilliam 2007 and references therein). Predictors of exotic richness include years of
European settlement, disturbance, and surrounding population density (McKinney 2002). Cities present
dramatic examples of this with their precipituous loss of native species and gain of exotic species (McKin-
ney 2006). Large areas such as states are highly disturbed and have high percentages of exotics (generally
between 15 and 35 percent)(Rejmanek & Randall 1994): eastern Texas, for example, adjacent to Caddo Parish,
has 18 percent exotics and Arkansas and Louisiana have 24 and 25 percent, respectively. States along the
eastern and western coasts of North America are the most disturbed (longest occupied) and have the most
exotics (Withers et al. 1998; Palmer 2008). Pennsylvania, for example, an eastern state much longer occu-
pied by European settlers than eastern Texas, Louisiana, or Arkansas, has 37 percent exotics. County-sized
areas tend to have slightly fewer exotics than states but basically rank with them. Madison County, Texas,
has 14 percent exotics; Caddo Parish, Louisiana, has 17 percent; Giles County, Tennessee, has 21 percent.
Natural areas and nature preserves tend to have fewer exotic species than do state and national parks (4 to
13 percent versus 12 to 17 percent, respectively). Heavily used urban parks have the most exotics (19 to 60
percent). Only very small areas remain without major disturbance and exotics; these are the sites frequently
chosen by ecologists to study "natural" communities. Such small areas, often less than a hectare, may show
no disturbance and may be entirely free of exotics ( 0 to 2 percent) even when surrounded by disturbed
areas (Gilliam 2007 and references therein).
On the basis of the data in Figure 2, it would appear that only small landscapes remain in natural
condition (i.e. without disturbance or exotics). All large landscapes ultimately include disturbed areas and
presumably, therefore, exotics. But, size matters little as data on parks suggest: Central Park in New York City
has 6096 exotics and the tiny Castillo San Marcos Monument has 4296. While it would be ideal to develop
ola ti 11r Antinio
MacRoberts et al., Vascular flora of C. Bickham-Dickson/Red River Park
1395
TABLE 1. Sy | ]
PARKS Species Area (sq km) exotics % Source
Central Park, NY 583 341 60 DeCandido et al. 2007
Castillo San Marcos Mt, FL 137 0.09 42 Zomlefer & Giannasi 2005
B-D Park, Caddo Parish, LA 318 2.36 21 This study
Middlesex Fells Park, Boston, MA 400 0.69 26 Harcombe et al. 2007
Pelham Bay Park, New York, NY 956 34 Harcombe et al. 2007
Highland Heights Comm.Park, OH 403 0.42 34 Jog et al. 2005
Shaker Median Park, OH 298 14 47 Delong et al. 2005
Botany Garden, Houston, TX 446 0.5 19 Harcombe et al. 2007
Mount Vernon 270 0.8 26 Wells & Brown 2000
Fort Washington Nat. Park, MD 973 19.56 25 Steury & Davis 2003
STATES
Pennsylvania 3318 119236 Coxe et al. 2005
3187 153270 24 Diggs et al 2006
Louisiana 3269 134226 25 Thomas & Allen 1993-1998
Fast Texas 3402 162134 18 Diggs et al. 2006
COUNTIES
Greene, PN 890 1500 20 Coxe et al. um
Oktibbeha, MS 1148 1185 15 Leidolf et al. 200
Caddo, LA 1405 2283 17 DU E nee 2006
dui AR 1110 2023 12 Marsico
Giles 1208 1582 21 Est M
UN Ms 797 1732 16 Martin et al. 2002
Pike, AL 1190 1759 20 Diamond 2
Yalobusha, MS 805 305 15 Denley et al. 2002
Madison, TX 1071 1217 14 Neill & Wilson 2001
STATE PARKS
Myakka River State Park, FL 731 16.86 12 Huffman & Judd 1998
Little Manatee State Park, FL 523 98 17 Myers & Wunderlin 2003
Fall Creek Falls, TN 879 89 13 Fleming & Wooford 2004
NATURAL AREAS
Turkey Creek, Big Thicket, TX 691 31.5 Brown et al. 2
Hickory Creek, Big Thicket, TX 401 28 4 MacRoberts et e 2002b
Big Sandy Creek, Big Thicket, TX 693 58 7 Brown et al. 2006a
Lance Rosier, Big Thicket, TX 694 101 10 Brown et al. 2006b
Rattlesnake Falls, TN 627 0.63 13 Estes & epis d
Big Branch, 563 58.7 13 Rosen et al. 2
Starkey Wilderness FL 483 449 9 Furguson & mom 2006
Big Lake Bottom WMA, TX 459 16.85 5 Fleming et al. 2002
Beadle Barrens Preserve, IL 306 0.04 7 Fdgin et al. 2005
Dean Hills Nature Preserve, IL 313 0.3 6 Feist et al. 2004
Stephenson Memorial Forest, KY 538 0.51 10 Thompson & Fleming 2004
Ouachita Mountains Biol. Sta., AR 337 1.54 7 MacRoberts et al. 2005
Pushmataha WMA, 447 76.9 7 al LA 2006
Cooters Bog, Vernon Parish, LA 135 0.03 1 rts € MacRoberts 1993
Big Thicket, Wet Savanna, TX 135 0.06 0 MacRoberts € MacRoberts 1998a
Muck Bog, Anderson Co., TX 136 0.05 2 MacRoberts & MacRoberts 1998b
Kisatchie Prairie, Winn Parish, LA 137 0.03 1 MacRoberts & MacRoberts 1996
Beech Slope, Sabine Co., TX 120 0.03 1 MacRoberts & MacRoberts 1997
Longleaf pine uplands, LA 158 0.004 0 MacRoberts et al. 2002a
Engeling WMA, Anderson Co., TX 930 44.65 6 Singhurst et al. 2003
1396 J | of the Botanical R h Institute of Texas 2(2)
Relationship of Exotic Species to Area
at Selected Sites
70 4 > - i
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R? = 0.0614
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@ Parks — AStates & Counties iB State Parks © Natural Areas — 9 C. Bickham-Dickson Park
Fic. 2. Relationship of exoti i t selective sites, Data from Table 1.
MacRoberts et al., Vascular flora of C. Bickham-Dickson/Red River Park 1397
a large data set comparing natural verses disturbed areas, size of area, history and time since disturbance
began, the scope of this paper excludes that possibility. Nonetheless, there are such data sets being devel-
oped, for example, The Floras of North America project by Michael Palmer at Oklahoma State University
(Palmer 2008)(not to be confused with the Flora of North America project) (see also McKinney 2002, 2006
for literature).
In the case of C. Bickham-Dickson/Red River Education and Research Park, the anthropogenic distur-
bance is great: it was a plantation and horse stables and is now a city park and a research area for a university.
It has been cleared of canopy vegetation, and some sections have regenerated canopy. It has been grazed; it
has been hayed and plowed. Part of it is mowed. There are old buildings. There are paved and dirt roads.
There is a pier, and fishermen use both the bank and boats. Vehicles leave the roads, and off-road vehicles
strip the ground of vegetation. Picnickers use the area, as do bird watchers. And the Park is surrounded by
an ever increasing urban landscape: suburbs, a university, and lake-front housing. Today, the level and flow
of the Red River is artificially controlled by the Corps of Engineers’ lock and dam system, and in 2007 and
2008, most of the Park was flooded for several months causing die-off of many plant species not adapted to
prolonged submersion. Twenty-one percent of the flora is exotic, a figure at the lower end of parks in our sample
(Table 1), but many of these species are aggressive invaders: hydrilla MO Ej verticil lata), Chinese tallow
tree (Sapium secs E privet (Ligustrum sinense), Japanese h ), alligator
, Burmuda grass (Cynodon dactylon), and Johnson grass (crum Walepense)
But the worst is oe to come. In neighboring lakes, ponds, and sloughs other aggressive invaders are
poised: giant salvinia (Salvinia molesta D.S. Mitch.) has just reached Caddo Parish and is choking nearby
lakes; water-lettuce (Pistia stratiotes L.), a native of the Gulf coast, has recently arrived in Caddo Parish and
undoubtedly will find its way into the Park; water hyacinth (Eichornia crassipes (Mart.) Solms) is in the Red
River adjacent to the Park and it is only a matter of time before it invades; and brittle water nymph (Naja
minor All.) has nd been one in Caddo Parish.
reconstruct what has happened to the Park, we do not know its “pristine”
ecology and, A what percent of native species has been lost. All we can reconstruct is the native tree
flora from the1830s land plat data (MacRoberts & MacRoberts 2005). These data indicate that, while we
still have the same native tree species we did in 1830, we also have some new ones, notably exotics such as
Chinese tallow tree (Sapium sebiferum), silktree (Albizia julibrissin), and chinaberry (Melia azedarach).
As long as the Park is in a continual state of disturbance largely because of conflicting social interests
and thus uses (Corps of Engineers, recreation, homeowners, business, state, city, and university), invasive
weeds, native and exotic, will find the area ideal and continue to accumulate to the detriment of native
species. Therefore, we can expect the Park to shift farther from a native state to a weedy and exotic state.
Whether the current pressure on the Park will result in a catastrophic change in the ecosystems—some
would say that it already has—or gradual change, remains unknown (Scheffer et al. 2001; Liu et al. 2007).
But that the Park will change is certain, notably if managers attempt drastic intervention; for example, to
control aggressive weeds such as hydrilla, when no one knows how much resilience the system has at pres-
ent or, for that matter, if the system or parts of it has already shifted to a new equilibrium. In that case it
will likely be impossible to restore to a previous state (Scheffer et al. 2001).
APPENDIX 1
Vascular plants found at Bickham-Dickson/Red River Education and Research Park. An asterisk * = exotic.
Numbers without collector are collection numbers of B.R. and M.H. MacRoberts. Other collectors are
named. Specimens are deposited at Louisiana State University in Shreveport herbarium (LSUS). Common
names are in parentheses after collection numbers.
Acanthaceae Ruellia caroliniensis (J.F. Gmel.) Steud., 6802 (Carolina wild
Justicia ovata (Walt.) Lindau, 6486 (loose-flower water- petunia)
willow)
1398
Aceraceae
Acer a L., 6014, 6161, 6462, 6831 (boxelder)
Alismataceae
RD cordifolius (L.) Griseb., 6643, 6796, 6984 (creep-
in rhead)
Sagittaria graminea Michx., 6233, 6998 (grass-leaf arrow-
head)
Amaranthaceae
*Alternanthera philoxeroides (Mart.) Griseb., 6443 (alligator
weed)
Amaranthus hybridus L., 6978 (amaranth)
nacardiaceae
Toxicodendron radicans (L.) O.Ktze., (not collected) (poision
om
in tainturieri Hook., 6019 (hairy-fruit chervil)
Cynosciadium digitatum DC., 6447 (finger dogshade)
Hydrocotyle umbellata L., 6229 (many-flower marsh-pen-
nywort)
iei verticillata Thunb., 6645 (whorled marsh-
ennywork)
E um L., 6661 (Canadian black-snakeroot)
e arvensis (Huds.) Link, 6426 (spreading hedge-parsley)
Trepocarpus aethusae Nutt. ex DC., 6442, 6449 (whit-
enymph)
Apocynaceae
Apocynum cannabinum L., 6430, 6969 (Indian hemp)
Aquifoliac
E d Walt 6212, 6669 (deciduous holly)
llex opaca Ait., 5980 (American holly)
Asteraceae
Acmell sitifolia (Lam.) R.K. Jansen, 6837 (opposite-leaf
UMP od
spotflower)
Ambrosia artemisiifolia L, 7001 (annual ragweed)
Ambrosia psilostachya DC., 6847, 7000 (perennial ragweed)
Ambrosia trifida L., 6762 (great ragweed)
Baccharis halimifolia L., 6759 (groundseltree)
Bidens laevis (L.) B.S.P, 6987, 6988 (smooth beggarticks)
*Carduus nutans L., 6489, 6659 (nodding ai iL
P Eun pilosa Nutt, 6424 (soft golden-ast
oclinium coelestinum (L.) DC., 6818 (blue Me n
a canadensis (L) Cronq., 6678, 6679, 6779 (Canadian
horseweed)
Coreopsis tinctoria Nutt., 6508 (golden tickweed)
Dra da Aaa (Vahl) Cass., 6433 (clasping cone-
flow
Eclipta cM (L) L, 6835 (false daisy)
Elephantopus carolinianus Raeusch., 6791 (Carolina elephant-
oot)
Erigeron philadelphicus L., 6170 (Philadelphia fleabane)
Erigeron strigosus Muhl. ex Willd., 6431, 6675, 7002, 7003
(prairie fleaban
Eupatorium capillifolium (Lam.) Small, 6778 (dog-fennel)
Eupatorium serotinum Michx., 6795, 6819 (late-flowering
thoughtwort)
—
f Texas 2(2)
SUMIGI vi
Gaillardia aestivalis (Walt) Rock, Burden s.n. (fireweed)
Gaillarida pulchella Foug., 6027 (firewheel)
Gamochaeta pensylvanica (Willd.) Cabrera, 6018 (Pennsylvania
everlasting)
RECON amará Aia jue 6483 (yellowdicks)
m.) Britt. & Rusby, 6677, 6685, 6803,
6825 LSE es
iva annua L., 6985 (annual marsh-elder)
Krigia caespitosa (Raf) Chambers, 5972, 7369 (weedy dwarf-
delion)
Lactuca canadensis L., 6649, 6758 (florida blue-lettuce)
Mikania scandens (L.) Willd., 6634 (climbing hempvine)
Packera obovata (Muhl. ex Willd.) W.A. Weber & A. Love, 5944
(round-leaf groundsel)
Pluchea odorata (L.) Vass., 6787, 6777 (sweetscent)
Pseudognaphalium obtusifolium (L.) Hillard & Burtt., 6849
(blunt-leaf rabbit-tobacco)
Pyrrhopappas pauciflorus (D.Don) DC., 7641, 7360, 6198
delion
Rudbeckia hirta L, 6481 (black-eyed-susan)
*Senecio vulgaris L., 5945 (old-man-in-the-spring)
Smallanthus uvedalia (L) Mackenzie ex Small, 7180 (bear's-
foot
Solidago lol L. pu scabra Torr. & Gray, 6832, 6951
(Can
*Sonchus asper (L ) Hill 5936, 5957 (spiny-leaf sow-thistle)
Symphyotrichum lateriflorum (L.) A. & D. Love, 6957, 6972
(American aster
hyotrict (Ell) Nesom, 6999 (fragile-stem
American aster)
Symphyotrichum subulatum (Michx.) Nesom, 6821, 6965
(seaside American aster
*Taraxacum officinale G.H. Weber ex Wiggers, 5960 (common
delion)
Vernonia baldwinii Torr, 6798 (western ironweed)
Xanthium strumarium L., 6226, 6661, 6663 (cockleburr)
Azollaceae
Azolla caroliniana Willd., 5935 (Carolina mosquito fern)
Berberidaceae
*Nandina domestica Thunb., 6465 (sacred-bamboo)
Bignon
Camps pra (L) Seem. ex Bureau, 6179, 6652 (trumpet
reeper
ragin
"eoropum UTER L, 6995, jen (Indian heliotrope)
5,6007, 7367 (large-seed
forget-me- not)
Brassicaceae
*Brassica rapa L., 5981 (rape)
*Capsella bursa-pastoris if Medik., 5939, 5958 (shepherd's
purse)
Lepidium virginicum L., 6421 (poorman's-pepperwort)
Sibara virginica (L.) Rollins, 5938 (Virginia winged- LN
Campanulaceae
Lobelia cardinalis L., 6989 (cardinal-flower)
MacRoberts et al., Vascular flora of C. Bickham-Dickson/Red River Park
ere perfoliata (L.) Nieuwl., 6025, 6422 (clasping-leaf
s-looking-glass)
Caprifoliaceae
*Lonicera japonica Thunb., 6192 (Japanese honeysuckle)
Sambucus nigra L., 6188, 6660 (black elder)
aryophyllace
“ena serlo L., 6034 (thyme-leaf sandwort)
ium glomeratum Thuill., 5940, 5964 (sticky mouse-ear
RM
Sagina decumbens (Ell.) T. & G., 5968, 6032 (trailing pearl-
wort)
*Stellaria media (L.) Vill, 5965 (common chickweed)
Ceratophyllaceae
Ceratophyllum demersum L., 6956 (coon's-tail)
Commelinace
*Commelina lid L, 6828 (Asiatic dayflower)
lina virginica L., 6824 (Virginia dayflower)
Convolvulaceae
*Convolvulus arvensis L, 6651, 6765 (field bindweed}
Dichondra carolinensis Michx., 6005 (Carolina d 's-foot)
Ipomoea cordatotriloba Dennst., 6511, 6676 (t
Ipomoea lacunosa L., 6816, 8062 (whitestar)
Cornaceae
Cornus drummondii C.A. Mey., 5998, 6512, 6760 (rough-leaf
dogwood)
Cucurbitaceae
*Cucumis melo L., 6808 (cantalope)
Cupressaceae
podes virginiana L., 5979, 5989 (eastern red-cedar)
Taxodium distichum (L.) L.C. Rich., 6514 (cypress)
Cuscutaceae
Cuscuta gronovii Willd., 6815, 6961, 8042 (dodder)
Cuscuta indecora Choisy, 6807, 6841 (large-seed dodder)
Cyperaceae
Carex cherokeensis Schw., 6009, 6165 (Cherokee sedge)
Carex crus-corvi Schuttlew. ex Kunze, 6438 (raven-foot
sedge)
Carex decomposita Muhl., 7429 (cypress-knee sedge)
oe flaccosperma Dewey, 6184, 6217, 6008, 6162 (thin-fruit
sedge)
Carex frankii Kunth, 6833 (Frank's sedge) [C aureolensis Steud.
NA]
Carex leavenworthii Dewey, 6021, 6154 (Leavenworth's
sedge
Carex retroflexa Muehl. ex Willd., 6174 (reflexed sedge)
Cyperus acuminatus Torr. & Hook. ex Torr, 6450, 6492, 6636
taper-tip flat sedge)
pode croceus Vahl., 6492, 6630, 6671,6677, 6834 (Baldwin's
sedge)
*Cyperus difformis L., 6993 (variable flat sedge)
Cyperus erythrorhizos Muhl., 6644 (red-foot flat sedge)
Cyperus esculentus L., 6857 (chufa)
Cyperus odoratus L., 6839, 6782, 6983 (rusty flat sedge)
*Cyperus rotundus L., 6425 (purple flat sedge)
1399
Eleocharis obtusa (Willd.) J.A. Schultes, 6448, 6515 (blunt
ike-r
Eleocharis palustris (L.) Roemer & J.A. Schultes, 6231, 6429
mon spike-rus
Fimbristylis autumnalis (L) Roemer & J.A. Schultes, 6773
(slender fimbry)
Fimbristylis vahlii (Lam.) Link, 8040, 8066 (fimbry)
Fuirena simplex Vahl., 6673 (western umbella sedge)
Rhynchospora corniculata (Lam.) Gray, 6982 (short-bristle
horned beak sedge)
Ebenaceae
Diospyros virginiana L., 6468, 6670 (common persimmon)
Equise
Dis i L., 6003, 6487 (tall scouring-rush)
Euphorbiaceae
Acalypha rhomboidea Raf., 6800 (common three-seed-
mercury.
Chamaesyce nutans (Laq.) Small, 6648, 6801, 6823, 6845,
6955 (eyebane
Chamaesyce prostrata (Ait.) Small, 6958 (prostrate sandmat)
Croton capitatus Michx., 6687, 6817 (hogwort)
Croton Min L., 6688 (vente-conmigo)
Croton monanthogynus Michx., 6763, 6846 (prairie-tea)
EE dentata Michx., 6844 (toothed spu d e)
*Euphorbia helioscopia L. 5961 (mad-woman's-milk)
Phyllanthus caroliniensis Walt., 6981 (Carolina rm
*Sapium sebiferum (L.) Roxb., 6196 (Chinese tallow)
Fabac
ius ori Durazz., 6463 (silktree)
A osa L., 6793, 7357 (false indigo-bush)
Cercis ub L., 6232 (redbud)
D / illi, is (Michx) M M. ex Bi Rob.& Fern. 6478
(prairie bundle-flower)
Desmodium canescens (L.) DC., 6827, 6766 (hoary tick-
trefoil)
Gleditsia triacanthos L., 6186 (honey-locust)
Glottidium vescarium (Jacq.) Harper, 6775, 6822 (bagpod)
*Medicago arabica (L.) Huds., 6176, 7359 (spotted medick)
*Medicago minima L., 5978, 7640 (medick
*Medicago lupulina L., 5977 (medick)
*Medicago polymorpha L., 6031 (tooth medick)
Mimosa strigillosa T. & G., 6420, 6657 (powderpuff)
Rhynchosia minima (L) DC., 6809, 6859, 6991 (least snout-
mm
bean)
Sesbania herbacea (P. Mill) McVaugh, 6855 (peatree)
Strophostyles helvula (L.) Ell, 6683, 6772 (trailing fuzzy-bean)
*Trifolium campestre Schreb., 5988, 6030 (lesser hop clover)
*Trifolium pratense L., 6187, 6466 (red clover)
*Trifolium repens L., 6153, 7361 als Con
*Trifolium resupinatum L., 6022 (re r
Vicia ludoviciana Nutt., 6177, 7364 (Louisiana Vete]
*Vicia sativa L., 5962 (garden vetch
*Vicia villosa Roth, 6185 (winter vetch)
Fagaceae
Quercus nigra L., 5975 (water oak)
Quercus virginiana Mill., 5953 (live oak)
1400
Gentianaceae
Centaurium muehlenbergii (Griseb.) W. Wright ex Piper, 6435
onterey centuary)
Sabatia campestris Nutt, 6495 (Texas-star)
Geraniaceae
Geranium carolinianum L., 5985, 7363 (Carolina crane’s-bill)
*Geranium dissectum L., 6159, 7362 (cut-leaf crane's-bill)
Hydrocharitace
*Hydrilla ptas (Lf) Royle, 6513, 7464b (water-thyme)
Limnobium spongia (Bosc.) Steud., 7936, 7937 (frog's bit)
Iridaceae
Sisyrinchium rostulatum Bickn., 6227, 7370 (annual blue-
eyed-grass)
Juglandaceae
Carya illinoinensis (Wangenh.) Koch, 6470 (pecan)
Juncaceae
Juncus bufonius L., 6445 (toad rush)
Juncus dichotomous Ell., 6182 (forked rush)
Juncus marginatus Rostk., 6432, 6493 (grass-leaf rush)
Lamiaceae
Hedeoma hispida Pursh, Burden s.n. (rough hedeoma)
*Lamium ampexicaule L., 5937 (giraffehead)
*Lamium purpureum L., 5967 (red henbit)
Salvia lyrata L., 6180 (lyer-leaf sage)
Stachys crenata Raf., 7647 (shade betony)
Teucrium canadense L., 6640 (American germander)
Lemnaceae
Lemna spp. 6234, 6861 (duckweed)
Spirodela sp., 6234, 6860 (duckmeat)
Wolffia sp., 6862 (watermeal)
Lentibulariaceae
Utriculara gibba L., 7181 (humped bladderwort)
Liliaceae
*Allium vineale L., 6157 (crow garlic
*Liriope muscari (Dcne) Bailey, 6829 (big blue lilyturf)
*Narcissus sp., 5955 (daffodil)
Nothoscordum bivalve (L.) Britt., 5943 (crowpoision)
Loganiaceae
Mitreola petiolata (Gmel. Torr. & Gray, 6638, 6672, 6781, 6814
(lax hornpod)
Lythraceae
Ammania coccinea Rottb., 6788, 6789, 6997 (valley redstem)
Lythrum alatum Pursh, 6783 (wing-angle loosestrife)
—
Malvace
Hibiscus pe All, 6641 (halberd-leaf rose-mallow)
ibiscus moscheutos L., 6639 rd -eyed rose-mallow
*Sida rhombifolia L., 6647 (tea
Sida spinosa L., 6641, 6785 (prickly Cuni
Meliaceae
*Melia azedarach L., 6183 (china-berry)
=
Menispermaceae
Cocculus carolinus (L.) DC.,,6469, 6658, 6971 (Carolina coralbean)
f Texas 2(2)
Molluginaceae
*Mollugo verticillata L., 6810, 8063 (green carpetweed)
Moraceae
Maclura pomifera (Raf) Schneid., 6191 (bois d'arc)
*Morus alba L., 5996, 6194, 7358 (white mulberry)
Morus rubra L., 6194 (red mulberry)
Myricaceae
Morella cerifera (L.) Small, 7041 (southern bayberry)
Nelmbonaceae
Nelumbo lutea (Willd.) Pers., 7428 (yellow-lotus)
Oleaceae
Forestiera acuminata (Michx) Poir., 6214 oo
Fraxinus pennsylvanica Marsh., 6682 (gree
*Ligustrum sinense Lour., 6166, 6650 (Chinese privit)
Onagraceae
Gaura mollis James, 6223 (velvetweed
Ludwigia decurens Walt., 6790 (wing-leaf primose-willow
Ludwigia leptocarpa (Nutt.) Hara., 6786, 6836, 8058 (angle-
stem primose-willow
Ludwigia peploides (Kunth) Raven, 6451 (floating primose-
willow)
Ludwigia repens J.R. Forest., 6633 (creeping primose-willow)
Oenothera biennis L., 6794 (King's cureall)
Oenothera laciniata Hill, 5995, 6171, 7366 (cut-leaf evening-
=
=
primrose)
Oenothera speciosa Nutt., 6434, 7365 (pinkladies)
Ophioglossaceae
Ophioglossum crotalophoroides Walt., 6026 (bulbous adder's-
tongue)
rchidaceae
Spiranthes laciniata (Small) Ames, 6500 (lace ladies-tresses)
Oxalidaceae
Oxalis corniculata L., 5947, 6656 (creeping yellow wood-
sorrel)
Passifloraceae
Passiflora incarnata L., 6970 (purple passion-flower)
Phytolaccaceae
Phytolacca americana L., 6761 (pokeweed)
Pinaceae
Pinus taeda L., 5982 (loblolly pine)
Plantaginaceae
Plantago aristata Michx., 6494, 6509 (Mexican plantain
Plantago heterophylla Nutt., 6023 (slender plantain)
Plantago virginica L., 5994 (pale-seed plantain)
Platanaceae
Platanus occidentalis L., 6002 (sycamore)
=
Poaceae
ee glomeratus (Walt.) B.S.P., 6963 (bushy
bluestem)
Andro, o virginicus L., 6996 (broom-sedge)
Aristida oligantha Michx., 6843, 7004 (prairie three-awn)
*Briza minor L., 6172 (lesser quaking grass
MacRoberts et al., Vascular flora of C. Bickham-Dickson/Red River Park
*Bromus catharticus Vahl., 6190, 6028 (rescue grass)
*Bromus japonicus Thunb. ex Murr., 6457 Eee brome)
Cenchrus spinifex Cav., 6504 (coastal sandbur
*Cynodon dactylon Pers., 6458, 6689 Mid ass)
Dichanthelium aciculare (Desv. ex Poir) Gould & Cork 7652
(narrow-leaf panic grass)
E shot n l /C-l IE
g id, 6491 (manyflow-
ered rosettegrass)
Digitaria ciliaris (Retz) Koel., 6768, 6811 (southern crabgrass)
*Digitaria ischaemum (Schreb.) Schreb. ex Muhl., 6962
smooth crabgrass)
*Echinochloa colona (..) Link, 6646 (jungle-rice)
Mou muricata (Beauv.) Fern., 6854 (rough barnyard
m" walteri (Pursh) Heller, 6840
*Fleusine indica (L) Gaertn., 6770 (Indian g grass)
Elymus virginicus L., 6156, 6655 eae wild De
Eragrostis secundiflora J. Presl., 6631, 7005 (red love grass)
Festuca paradoxa Desv., 6193 (clustered fescue
Leersia lenticularis Michx., 6959, 6976 (catchfly grass)
Leersia oryzoides (L) Sw., 6967 (rice cutgrass
Leersia virginica Willd., 6964 (white grass)
Leptochloa panicea (Retz) Ohwi, 6767 (needle spangletop)
Leptochloa panicoides (Presl.) Hitch., 6858 (Amazon spangle-
top)
*Lolium perenne L., 6173 (perennial rye grass)
Panicum capillare L., 6820, 6840, 6994, 8060 (common panic
grass)
Fal Aiek
6680, 6966, 6842 (fall panic
grass)
Panicum rigidulum Bosc. ex Nees, 6780, 6838 (red-top panic
grass)
*Paspalum dilatatum Poir., 6456 (golden crown grass)
Paspalum distichum L., 6635, 6856 (joint crown grass)
*P, ipee notatum Fluegge, 6452, 6503, 6662, 8044 (Bahia
gra
Paspalum setaceum Michx., 6501, 6505 (slender crown
gra
*Paspa e urvillei Steud., 6974 (Vasey's grass)
Phalaris re Walt., 6216, 6476 (May grass)
*Poa annua L., 5984 (annual blue grass)
Setaria ipid (Poir.) PEN 6769 (marsh bristle
grass)
*Setaria pumila (Poir.) Roemer & Schutes, 6418, 6973 (yellow
bristle grass
*Sorghum halepense (L.) Pers., 6467 (Johnson grass)
Sphenophoiis obtusata (Michx.) Scribn., 6497, 6498 (prairie
wedgescale)
Triplasis purpurea (Walt.) Chapm., 6812, 6851 (purple
andgrass)
Tripsicum dactyloides (L.) L, 6979 (eastern mock grama)
*Tritic estivum L., 6485 (bread wheat)
Zizaniopsis miliacea (Michx.) Doell. & Asch., 6439, 6992
(marsh-millet)
Polygona
Sumi ovata (Walt) Shinners, 6681 (American buck-
wheatvine)
1401
Polygonum hydropiperoides Michx., 6852 (swamp smart-
Polygonum scandens L., 6975 (climbing false buckwheat)
Polygonum setaceum Baldw., 6642 (bog smartweed)
Polygonum virginianum L., 6830 (jumpseed)
*Rumex crispus L., 5959 (curly dock)
*Rumex hastatulus Baldwin, 6017 (heart-wing sorrel)
Portulaceaceae
Portulaca oleracea L., 6784 (little-hogweed)
Potamogetonaceae
Potamogeton nodosus Poir, 6632, 6666 (long-leaf pond-
weed)
Primulaceae
Anagallis minima (L.) Krause, 6033 (chaffweed)
Samolus valerandi L., 6444 (seaside brookweed)
Ranunculaceae
Anemone berlandieri Pritz., 5954 (ten-petal thimbleweed)
Myosurus minimus L., 5974 (tiny mousetai
*Ranunculus muricatus L., 5948 (spring-fruit buttercup)
Ranunculus pusillus Poir., 6220 (low spearwort)
*Ranunculus sardous Crantz, 5986, 6440, 7368 (hairy but-
tercup)
Rhamnaceae
Berchemia scandens (Hill) Koch, 5993, 6215 (Alabama
supplejack)
Rosaceae
Crataegus spathulata Michx., 6181 (little-hip hawthorn)
*Duchesnea indica (Andr) Focke, 5952 (Indian strawberry)
Geum canadense Jacq., 6482 (white avens
Prunus angustifolia Marsh., 5987 (Chickasaw plum)
*Pyrus calleryana Dcne., 5976 (Bradford pear)
Rubus trivialis Michx., 5990 (southern dewberry)
Rubiaceae
Cephalanthus occidentalis L 6637 (button bush)
Diodia teres Walt., 6686, 6813 (poorjoe)
Diodia virginiana L., 6446 (Virginia buttonwood)
Galium aparine L., 6016 (sticky-willy)
Galium tinctorium (L) Scop., 6428 (stiff marsh bedstraw)
Houstonia micrantha (Shinners) Terrell, 5942, 5969 (southern
bluet)
uet
Houstonia pusilla Schoepf, 5941, 5971 (tiny bluet)
*Sherardia arvensis L., 5949, 5963 (blue field-madder)
Spermacoce glabra Michx., 6668 (smooth false buttonweed)
Rutaceae
Zanthoxylum clava-herculis L., 6475 (Hercules’-club)
Salicaceae
Populus deltoides Bartr. ex Marsh., 6013, 6164 (cottonwood)
Salix exigua Nutt., 6665, 8043a, 8043b (narrow-leaf willow)
Salix nigra Marsh., 6000, 6020, 6168, 6474, 6479, 7959, 7960
(black willow)
—
apindaceae
Cardiospermum halicacabum L., 6776, 8065 (love-in-a-puff
1402
Saururaceae
Saururus cernuus L., 6490 (lizard's tail)
Scrophulariace
Agalinis ll (Nutt.) Small
false foxglove)
Leucospora multifida (Michx.) Nutt., 8041 (narrow-leaf cono-
bea)
Britt., 6805, 8059 (prairie
*Mazus pumilus (Burm.f) Steenis, 5983 (Japanese mazus)
Mecardonia acumnata (Walt) Small, 6853 (axil-flower)
Mimulus alatus Ait., 6799 (sharp-wing monkey-flower)
Nuttallanthus texanus (Scheele) D.A. Sutton, 5992 (Texas
toadflax)
"Veronica arvensis L, 5970 (corn speedwell)
Veronica peregrina L., 5946, 5973, 7371 (neckweed)
"Veronica persica Poir., 5966 (bird-eye speedwell)
Smilacaceae
Smilax bona-nox L., 6220, 6488 (Fringed Greenbrier)
Solan
esca pe Nutt., 6653, 6667 (long-leaf ground-
cher
rry
Solanum carolinense L., 6419 (Carolina horse-nettle)
Solanum elaeagnifolium Cav., 6496 (silver-leaf nightshade)
Typhaceae
Typha latifolia L, 7451 (cat-tail)
I Inftha Dat taal D hi
f Texas 2(2)
Ulmus alata Michx, 6218 (winged elm)
Ulmus americana L., 6001 (American elm)
Urticaceae
Boehmeria cylindrica (L) Sw., 6797, 6968 (small-spike false
ettle)
Valerianaceae
Valerianella radiata (L.) Dufr., 5991 (beaked cornsalad)
Verbenaceae
Phyla lanceolata (Michx.) Greene, 6473, 6960 (northern
frogfruit)
ogiru
*Verbena MER ie 6471 (Brazilian vervain)
Verbena mall, 6 6804 (Texas vervain)
Verbena jd L., (white vervain)
Violaceae
Viola bicolor Pursh, 5951 (field pansy)
Viola sororia Willd., 5950 (blue violet)
Viola sagittata Ait., 6006 (arrow-leaf violet)
Vitaceae
Ampelopsis arborea (L.) Koehne, 6477 (peppervine)
Ampelopsis cordata Michx., 6480, 6654 (heart-leaf pepper-
vine)
Parthenocissus quinquefolia (L.) Planch., 6460 (Virginia-
creeper)
Vitis riparia Michx., 6189, 6197, 6986, 5997 (river-bank
)
Ulmaceae grape
Celtis laevigata Willd., 6163 (hackberry)
ACKNOWLEDGMENTS
This project was supported by grants from Louisiana Board of R Support Fund, the J. Bennett Johnston
Foundation, and the Environmental Protection Agency, Begin. 6, Dallas Office, Louisiana Department of
Environmental Quality. Jennifer La Pierre and Jennifer Cook, Media and Public Relations, LSUS, and Cran
Lucas, Biology depu supplied and developed the figure. Amanda C. Lewis aided with the electronic
preparation of ipt and Robert Kalinksy aided with Figure 2. Two anonymous reviewers provided
Eu
many useful comments.
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A FLORISTIC SURVEY OF VASCULAR PLANTS OVER PARTS
OF NORTHEASTERN NEW MEXICO
Michael H. Schiebout and Donald L. Hazlett Neil Snow!
School of Biological Sciences School of Biological Sciences
University of Northern Colorado University of Northern Colorado
Greeley, Colorado 80639, U.S.A. Greeley, Colorado 80639, U.S.A.
ABSTRACT
A floristi leted i ] w Mexico, including all of Harding, Mora, and San Miguel counties, most of Colfax
County, and M SINE north of Interstate 40. Prior to this study the approximately 41,000 km? (ca. 15,800 mi?) study area, much
of which is private land, had not been systemically surveyed for its plant diversity. A total of 9,619 specimens were collected during
2004 and 2005, which included 93 families, 437 genera, 911 species, and 956 taxa. Rorippa teres, Chenopodium berlandieri var. berland-
ieri, Artemisia ludoviciana var. incompta, Tetraneuris torreyana and Rosa x harisonii (escaped from cultilvation) were confirmed as state
y
records. Approximately 9.3% of the flora (89 taxa) was non-native, including 11 species listed as “noxious” by the state of New Mexico.
o e uin an overview of ne and Beology, a discussion of vegetation types are included. The annotated checklist
tions, counties MCA Mus one voucher per county cited, habitat
types, and elevational fi I Speci lati y collected habitats suggest strongly
that additional taxa will i found in each TM A Sorenson index was used to calculate ilorisue similarity of the Kiowa National
Grasslands in New Mexico (within the study area) with that of the Pawnee and Comanche National Grasslands in Colorado. The first
analysis, which mendes " p taxa, indicated that the a Separated ds ix greatest gsm E the least samen hs
second analysis, whicl abc y values,
eo
between the e Grasslands.
RESUMEN
11 Aina fl : 13] T T ] te Ha NT Ms f. 1] A 1, 1 dad de Colfax
F L ?
Harding, Mora, Quay (al norte), y San Miguel d las ér 1 imi 2004 y 2005. Antes de este estudio esta área de 41,000
1 2 eg AnA :23 q 7 1 A hal Ans 7 pa] detall H la di idad icf. ted
TE i fr El oO Ly
pl 1 Tin 1] a 10 1 A i 1 z J J 1 E A oct tuidi 254 ZI. an E. e Ma , 437
epu 911 a y E taxa. También el total 89 pl óticas ( ivas), lo a es aproximadamente 9.3% de
la flora total. D Rorippa teres, TO berlandieri var. berlandieri, Artemisia ludoviciana var. incompta,
Tetraneuris torreyana, y Rosa x harina TAE inci vez pus el estado de Nuevo México. a pe exóticas había 11 especies
designadas para el estado de Nuevo Méxi Están incluidos | y datos del Eus geología,
ups » i au Las cifras más ae = estas aun con los números n ida nombres iS autorías ud BN
Enl i
e r X
1 1 1 4 + J EC Af: > A : > A 1 1 eae | 4 4 Se hicieron
o a L
4 slicice del indice de & (m PR Epes : Santee Jada idad de plantas en la Pradera Nacional del Kiowa en Nuevo
México con las diversidades de plantas que ocurren en las DIR b dM del Pawnee y del Comanche en Colorado. El primer
C
análisis dd tanto las E nativas como las p ión. Como iacu que cuanto más n un área n
u d tr j El E d álisis i 1 y 6 l planas nativas y
1 "E 1 B r 411 22d A : ia 1 1 1
4 e e L x- L3 e
pero no en todas de las praderas estudiadas.
INTRODUCTION
The Rocky Mountains in the United States have been the site of over forty intensive floristic survey dating
back to the early 1980s (e.g., Nelson 1984; Marriot 1985; Kastning 1990; Hartman 1992; Snow 1994—95;
Hartman & Nelson 1998; Elliot 2000; Holt 2002; Reif 2006). Data from floristics projects are critical for
determining emergent properties of taxa, such as assessing which are endangered or threatened, and habitat
“Present address: Herbarium Pacificum, Bishop Museum, 1525 Bernice St., Honolulu, HI 96817, U.S.A.
2Author for correspondence: neil.snow@bishopmuseum.org
J. Bot. Res. Inst. Texas 2(2): 1407 — 1447. 2008
1408 J lof the Botanical R h Institute of Texas 2(2)
data provide important insights about how best to protect such species (Ertter 2000). Floristic studies also
faciliate the tracking of vegetative change through time (Stein & Davis 2000), which is increasingly valuable
given the extensive urbanization, overgrazing, and natural disasters (fire, flooding, diseases of forests) that
have significantly changed much of the American landscape. Only floristics data can track the historical
spread and relative abundance of non-native taxa, many of which are decreasing the commercial and private
value of many lands (D'Antonio & Meyerson 2002). In addition, distributional data from floristic surveys
can guide the restoration of degraded landscapes using native taxa (Stein & Davis 2000). Plant surveys may
even be used in regional land planning (Neely et al. 2006) even though many counties in the United States
have species lists that greatly underestimate the true number of plant taxa from those areas (Moerman &
Estabrook 2006; Norris et al. 2001)
Comparing present and past plant distributions also allows biologists to evaluate the effects of long-
term ecological changes. For example, the Floristic Quality Index (Northern Great Plains Floristic Quality
Assessment Panel 2001) gives each plant species from a given area a value from 0 to 10 based on its ecological
importance (Andreas et al. 2004). When plants from a specified area are viewed cumulatively, a value can be
calculated that helps determine how the area should be ranked in terms of regional conservation priorities.
As average global temperatures continue to increase and geographical distributions of species shift, floristic
studies can be used to make informed decisions about the conservation of particular species.
Floristic surveys are particularly valuable because they summarize distributional data at relatively
localized scales. An incomplete (and thus erroneous) knowledge of the distribution of a species can cause
problems in the listing of plants as threatened or endangered in two ways. First, a species may be considered
endangered erroneously due to limited knowledge of its true distribution and abundance. Expensive pro-
tective policies could be enacted unnecessarily due to formal listing on Endangered or Threatened lists, as
well as causing unnecessary litigation (Ertter 2000). Second, a lack of (or limited) knowledge of taxonomic
diversity, as for example how morphologically similar species are distinguished and how their distributions
differ, may exclude taxa that need genuinely need protection (Ertter 2000). Given the large number of newly
described plant taxa from North America over the past three decades, including many from New Mexico
(Hartman & Nelson 1998), additional survey work is imperative if we expect to provide maximal protection
for genuinely rare and threatened taxa.
The most important results of floristic studies, however, are the discovery of plant taxa new to science
during intesive (Hartman & Kirpatrick 1986; Wilken & Hartman 1991; O'Kane & Reveal 2006) or routine
collecting (Reveal 2002; Holmgren & Holmgren 2002). In light of such discoveries and the fact that large
parts of North America (especially Canada and Alaska) have low collecting densities, the need for additional
floristics work is clearly warranted in many areas.
New Mexico is one of the least explored states botanically in the United States, with perhaps only Ne-
vada being less intensively surveyed overall among the mainland western states. Prior to this study, which
was carried out primarily by the first author (Schiebout 2006), the High Plains region of northeastern New
Mexico had been inadqueately surveyed. Evidence in support of this observation came from records in the
database at the University of New Mexico, Albuquerque (UNM), which in late 2003 had accessioned and
databased fewer than 900 total specimens from Union, Colfax, Harding and Quay counties (T. Lowrey,
UNM herbarium curator, pers. comm. 2003). Although that figure now exceeds 3250 specimens (exclusive
of specimens from this project) it still reflects a relatively low collection density for this expansive area.
The goal of this study was to document the floristic diversity of vascular plants from a large part of the
northeastern counties of New Mexico. Two Du nee were among the objectives. The first was to collect ex-
tensively on private lands, given that pri ( lly cattle ) are among the least-collected
portions of the western United States. This oo was is deemed important becase most of the study area is
privately held ranchland and because most ranches have one or more wetland areas that harbor plant taxa
unique to those habitats. The second objective was to focus on aquatic and riparian habitats, which typically
include taxa with narrow ecological distributions, and because aquatic and riparian habitats represent a
small portion of the study area.
Schiebout et al., V. lar plants of northeastern New Mexi 1409
r
Boundaries of Study Area.—The study area is spread over a six-county area in northeastern New
Mexico (Fig. 1) that we estimate encompasses approximately 41,000 km? (ca. 15,800 mi’). It includes most
of Colfax, Quay north of Interstate 40, and virtually all of Union, Harding, Mora, and San Miguel counties
(Fig. 1). The survey area included elevations from 1100 to 2650 meters and focused mostly on the plains and
foothills. The western boundary generally was the lower foothills of the Southern Rocky Mountains, although
a few collections extended up to 2650 meters in Colfax County (Schiebout 2006; Table 2). A floristic study
in progress that encompasses the headwaters and drainage of the Vermejo River and higher elevations of
Colfax County and adjacent areas is in progress (B. Legler, pers. comm. 2008). The eastern boundary of the
area was the Texas/Oklahoma border. The northern edge was the state of Colorado, whereas the southern
boundary extended to approximately 35°15'N latitude (Fig. 1).
Geography and Geology.—The study area can be divided into three biogeographical regions: the high
plains of New Mexico, encompassing the eastern half of the study area, the central plains of New Mexico,
comprising the middle region of the study area, and the mountainous region occupying the western part of
the study area (Dick-Peddie 1993). The high plains and the central plains together are part of the Southern
Great Plains, whereas the mountainous region is on the east slope of the Sangre de Cristo Range of the
southern Rocky Mountains (Chronic 1987).
Geologically, the study area is located east of the Rio Grande Rift. Moving east from the Rift towards
Texas and Oklahoma, the landscape includes expansive plains, rolling hills, low bluffs and river valleys
(Chronic 1987). Sedimentary rocks from the Permian and Miocene epochs are the most common in this
area. Much of the surface region is covered with Permian limestone, part of the erosion-resistant San Andres
limestone. Other common rocks in northeastern New Mexico include red Triassic sandstones and siltstones
of the Chinle formation, and Dakota sandstone and Mancos shale formations from the Cretaceous period.
Miocene-Pliocene gravel, part of the Ogallala formation, can also be found at the surface in many parts of
northeastern New Mexico (Chronic 1987).
The study area contains relatively few prominent geological structures. However, faulting, folding, and
some warping occur in the sedimentary layers. Much of the relief in the area was formed from the downcut-
ting of the Canadian River and its tributaries. The Canadian River, which has its headwaters near Raton Pass
in northern Colfax County, flows south through east-central New Mexico to the Conchas Reservoir in San
Miguel County. Thereafter it flows east across the Texas Panhandle into Oklahoma. The river cuts a 300—400
meter deep gorge in Mills Canyon through the Tertiary and Mesozoic strata of the Las Vegas Plateau section
of the Great Plains and the Canadian Escarpment (Wisniewski & Pazzaglia 2002), where the river divides
Mora and Harding counties.
In the northern part of the study area, remnants of hundreds of small basaltic to nephelinitic volcanoes
are ies (U.S. Geologic Service 2006). pone of the more eroded volcanoes date to the Tertiary period,
tones are from the Pleist to Recent epochs (Chronic 1987). The newer volcanoes
have experienced relatively little erosion. Volcanic activity of the more recent epochs left broad lava flows
over large areas of northern Union and Colfax counties, some of which extend into southern Colorado.
Climate.— Climate varies relatively little throughout the study area. Annual precipitation ranges from
39.3—45.9 cm (Table 1). Precipitation increases slightly from southeast to northwest. This change is seen by
comparing data from Cimmaron, Las Vegas, and Raton in the north to stations located in the eastern region,
such as Conchas Dam and Clayton (Table 1).
The wettest months are July and August, when moisture from the Gulf of Mexico moves in a southeast-
erly circulation pattern (Rogash 2003). As the moisture-laden air ascends to higher elevations it condenses
and falls as precipitation.
Temperatures likewise vary along a northwestern to southeastern gradient. The average annual maxi-
mum temperatures in the southeast average above 22.3°C, whereas annual maxima of 17.5-18.3°C occur
in the northwest (Table 1), Elevation changes account for most of this variation, given the relatively small
differences in latitude. In general, altitudes are highest in the northwest and lowest in the southeast. The
1410 J | of tl f Texas 2(2)
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Taste 1. Climatic data from 10 weath ions in nortl New Mexico (Wi Regional Cl Center 200
Location Ave. Annual Max. Ave. Annual Min. Ave. Annual Vn Precipitation
(Altitude in meters) n p 1971-2000 Temp. (C) 1971—2000 (cm) 1971—200
AMISTAD (1376) 214 42 399
BELL RANCH (1372) 22.7 43 393
CIMARRON (1891) 183 0.6 45.9
CLAYTON (1512) 19.4 4.1 40.1
CONCHAS DAM (1294) 22.9 6.8 394
DES MOINES (2018) 17.5 2.1 458
LAS VEGAS (2013) 182 20 42.9
MOSQUERO (1665) 19.8 3.3 41.9
SPRINGER (1979) 20.2 0.6 43.1
44.3
RATON (1806) 18.1 -0.2
Schiebout et al., Vascular plants of northeastern New Mexi 1411
warmest months are July and August, whereas the coldest temperatures occur from December through
March.
Vegetation Classification.—The study area is located predominantly within the North American Prairie
Province, with portions of the western region within the Rocky Mountain Province (e.g., Takhtajan 1986).
Ecologists have classified the area in several ways. The Rocky Mountain Province is sub-classified into the
Intermountain Valleys and Lower Mountain Slopes, which include the Pinyon-Juniper Woodlands (West
& Young 2000) and the Southern Rocky Mountains (Peet 2000). The North American Prairie Province is
sub-classified into a number of major grassland types. Shortgrass prairie dominates the study area (Sims and
Risser 2000). Dick-Peddie (1993) described seven land cover classes located in northeastern New Mexico,
including coniferous and mixed woodland, plains mesa grassland, montane grassland, montane scrub, ur-
ban or farmland, subalpine coniferous forest, and juniper savanna. The classification of Dick-Peddie (1993)
was followed broadly, but some additional categories were added for this project. The broad categories used
to summarize distributions herein are grasslands, coniferous woodlands, wetlands, disturbed areas, and
miscellaneous. Broad categories are subdivided as evident in Table 2.
METHODS
Plant collections were made at 249 sites across the study area during the late spring and summer during the
relatively wet years of 2004 and 2005. In light of the large amount of privately held land in the study area,
mostly cattle ranches, a concerted effort was made to collect on private lands. Efforts were made to focus on
wetland areas (broadly construed) on the private properties. Eighteen landowners granted permission to
collect from their properties, which resulted in 3012 specimens, or nearly one third of the total collections.
Microsoft Access was used to generate lists of taxa collected from each private property, which have been
sent to each landowner in gratitude for their cooperation and assistance.
Colleting permits were secured for the Kiowa National Grassland (FS-2700-5), nine New Mexico state
parks (SU-05-06-ALL-02), and the Maxwell and the Las Vegas National Wildlife Refuges (22580-05-LV03
and 2005—01, respectively).
The general collecting p ] ially followed Hartman (1992). In brief, entire days were spent
collecting plants in flower or fruit from selected sites. Since most of the study area was located on the high
plains, where the number of roads is generally adequate and remote mountainous areas absent, collections
could be made more rapidly than for similar projects with greater amounts of mountainous terrain and fewer
roads. Specimens sometimes were pressed the day of collection, but usually were placed on ice overnight and
pressed the next day. Pressed specimens were dried immediately on portable wooden driers using 100-watt
bulbs for two to four days. Over 150 days were devoted to collecting and pressing over two seasons. The
249 total collection sites were chosen to reflect the diversity of topography, geology, habitat types, and other
factors, as well as locations on public or private lands. Unlike the procedure discussed by Hartman (1992),
no attempt was made to collect based on predetermined criteria regarding the ideal number of collection
points per linear mile. The first set of specimens is deposited at the University of Northern Colorado (GREE),
along with photocopies of field notebooks. The largest number of duplicates resides at the University of New
Mexico (UNM), and another large set was deposited at the Rocky Mountain Herbaruim at the University of
Wyoming (RM).
References used most frequently for making identifications included: Flora North America Editorial
Committee (1993+), Great Plains Flora Association (1986), Martin & Hutchins (1980, 1981), Ivey (2003), Diggs
et al. (1999), Harrington (1964), Weber & Wittman (2001), Isely (1998) and Cronquist et al.(1972-1997).
Cox (2001) was used to assist with identifications of non-native taxa in New Mexico. The non-native nox-
ious plants of New Mexico were identified using USDA PLANTS (United States Department of Agriculture,
Natural Resources Conservation Service 2006).
The initial nomenclatural authority for this project was Snow & Brasher (2004), which has been replaced
by Snow (2007) each of which was based largely on the APGII classification (APG 2003; see also Stevens
1412
TABLE 2. |
J
oftwo types (e.g. Lacustrine- Plains mesa grassland).
Vegetation Type Sites Taxa Elevation (M)
Total Taxa
Grasslands
Plains-Mesa grassland 82 434 44 1150-2500
Plains Mesa grassland-Pinyon-juniper woodland 3 125 13 1350-1700
Plains-Mesa grassiand-Riparian 20 353 36 1350-1600
Plains-Mesa grassland-Rocky slopes 3 76 8 1150-1400
Montane grassland 1 4 04 2600-2650
Montane grassland-Lacustrine ] 123 12 2250-2300
Montane grassland-Riparian 2 136 14 2350-2600
Foothills grassland 2 78 8 1850-2000
Foothills grassland-Artificial impoundments 1 61 6 2000-2050
Coniferous woodlands
Pinyon-juniper woodland 8 181 18 1450-1950
Pinyon-juniper woocland-Oak woodland 2 13 1 1950-2050
Pinyon-juniper woodland-Mixed shrubland 1 8 1 1600-1700
Pinyon-juniper woodland-Ponderosa pine woodland 2 47 5 1750-1950
Pinyon-juniper woocland-Rocky Wash 1 25 3 1750-1800
Ponderosa pine woodland 6 177 18 1200-2350
Ponderosa pine woodland-Riparian 3 122 12 1800-2350
Mixed coniferous woodland 1 26 3 2400-2450
Mixed coniferous woodland-Riparian 2 104 11 2250-2300
Deciduous woodlands
Riparian woodland 3 116 12 1350-1450
Riparian woodland-Rocky Wash 1 93 9 1550-1600
Oak wood land 3 168 17 1550-2100
Oak wood land-Riparian 2 90 9 2350-2400
Shrublands
Mixed shrubland 8 222 22 1250-2000
Mixed shrubland-Lacustrine 1 35 4 1750-1800
Wetlands
Riparian 19 471 48 1100-2500
bou Rocky Wash 2 93 9 1350-1400
Lacustrine 5 140 14 1800-2450
o ae grassland 2 56 6 1800-1850
Palustrine 2 29 3 1300-1900
Disturbed
Artificial impoundments 9 240 24 1150-2050
Roadside 48 392 40 1150-2500
Miscellaneous
Badlands 1 9 1 1150-1200
Rocky Wash 1 42 4 1500-1550
Rocky Slope 1 45 5 1350-1400
2001). Specimens that did not key out easily or match herbarium specimens particularly well were marked
with “aff.” (affinities to). Species that may represent state records are prefixed with a number sign (#) in
front e oe oe name.
1 for the five most frequently collected habitat types (plains
mesa ned roadside, riparian, pinyon-juniper associates, plains mesa grassland-riparian) to assess the
Schiebout et al., V. lar nlant« of north Mau Mavi ‘aig
F
thoroughness of the sampling regime. Sequential visits to a habitat type are recorded on the x-axis. The
cumulative number of species collected in each habitat type is recorded on the y-axis.
RESULTS
Over 9,600 specimens were collected, which include the following personal collection numbers: M. Schiebout
(117-2278, 2295-3113 and 3135-9151); N. Snow (9279-9526, 9564-9936), and approximately 55 miscellaneous
collections of D. Hazlett. Relatively small outliers of the Carson National Forest and Santa Fe National Forest
occurring in the study area were bypassed, given recent floristic work on the Carson National Forest by Jill
Larson (unpublished) and the Santa Fe National Forest by Reif (2006).
Vegetation types
To better understand the distribution of individual taxa, the study area was partitioned into a number of
vegetation types (Table 2) and a record was made for each taxon of the vegetation types in which it was col-
lected. The vegetation types, although necessarily somewhat arbitrary and broadly defined, none the less are
useful to land use managers, ecologists, and other workers. They are described in the following paragraphs,
including some of the more commonly encountered taxa in many.
Grasslands.—Plains Mesa Grassland is the most abundant type of grassland in New Mexico (Dick-
Peddie 1993). It encompasses about two thirds of the study area, extending from the eastern border of New
Mexico west to the foothills of the southern Rocky Mountains. As with other grasslands of central North
America, it has a large representation of taxa from Poaceae and Asteraceae. A total of 82 collecting sites
were made from Plains Mesa grassland, which yielded 434 taxa (Table 2). Some of the taxa collected most
commonly included: Aristida purperea var. longiseta, Bouteloua curtipendula, Bouteloua dactyloides, Bouteloua
gracilis, Cheatopappa ericoides, Chenopodium incanum, Chenopodium pratericola, Cirsium ochrocentrum, Elymus
elymoides var. brevifolius, Engelmannia pinnatifida, Gl nin ede Var. Epinal ad. Guara coccinea,
NA 1
Hymenopappus filifolius var. polycephalus, ij , Ratibida colum-
nifera, Solanum elaeagnifolium, Solanum rostratum, Sphaeralcea co coccinea var. coccinea, Tetraneuris scaposa var.
scaposa, Thelesperma meg , Xanthisma spinulosum and Zinnia grandiflora
Montane praselanded were dla to include areas above 2,000 m elevation. This type of grassland
was rare, restricted to the western part of the study area, and generally was associated with wetland habitats.
Montane grasslands yielded 4 taxa (Table 2).
Foothills grasslands are located up to 2,000 m; samples were collected from these sites only twice for
this study, yielding 78 taxa (Table 2).
Coniferous Woodlands.—Pinyon-juniper woodlands are dominated by Pinus edulis and either (or
both) Juniperus scopulorum and J. monosperma. The most abundant herbaceous species were Berlandiera
lyrata, Bouteloua dactyloides, Chaetopappa ericoides, Elymus elymoides var. brevifolius, Glandularia bipinnatifida
var. bipinnatifida, Melampodium leucanthum, Oenothera albicaulis, Tetraneuris scaposa var. scaposa and Vulpia
octoflora. The 8 collection sites from this habitat yielded 181 taxa (Table 2).
Ponderosa pine woodland is dominated by Pinus ponderosa. Other types of vegetation often grew with
the pine, frequently including species of oak (Dick-Peddie 1993). The most abundant native Species were
Erigeron flagellaris, ee pobycephatus, Juniperus scopulorum, d Eaton Pinus edulis, Poa
pratensis, Q var. o TI ly encountered
were Cynoglossum offi inale, Dactylis gl ta and Melilotus officinalis. The 6 collection sites from this habitat
were represented by 177 taxa (Table 2).
Mixed coniferous woodland is a mixture of coniferous species including Abies, Picea, Pinus and Pseu-
dosuga. This vegetation type occurs in the western and northern parts of the study area at elevations above
approximately 2400 m. The single collection visit yielded 26 taxa (Table 2).
Deciduous Woodland.—Riparian woodland occurs on floodplains adjacent to watercourses. Species
of Salix and Populus species dominated this vegetation type. The 3 collection sites from riparian woodlands
yielded 116 taxa (Table 2).
1414 J lof the Botanical R h Institute of Texas 2(2)
Oak woodland forest or shrubland is dominated by Quercus gambelii, Q. grisea and/or Q. x undulata. The
3 collection sites from this vegetation type included 168 taxa (Table 2).
Shrubland.—Mixed shrubland consists of shrubs (and sometimes low-growing trees) and included
species of Quercus, Cercocarpus, Rhus and Artemisia. The most abundant species were Artemisia ludoviciana,
Berlandiera lyrata, Juniperus monosperma, Quercus grisea, Quercus x undulata, Rhus aromatica var. trilobata,
Senecio flaccidus var. douglassii, Solanum elaeagnifolium, Thelesperma megapotamicum and Zinnia grandiflora.
The 8 collection sites in this habitat yielded 222 taxa (Table 2).
Wetlands.—Riparian vegetation occurs in areas with seasonally saturated soils, or in sand or rocks ad-
jacent to normal watercourses. The most abundant species were Bromus japonicus, Eleocharis palustris, ellen
RI
smithii, Equisetum laevigatum, Erigeron divergens var. divergens, Melilotus albus, Poa pratensis,
trilobata, Shoenoplectus pungens and Tragopogon dubius. The 19 collections nude in riparian habitats ae
471 taxa (Table 2).
Lacustrine areas occur along the margins of natural ponds or lakes. The most abundant species col-
lected from this vegetation type were Astragalus bisulcatus var. bisulcatus, Convolvulus arvensis, Chaetopappa
ericoides, Eleocharis palustris, Erigeron canus, Erigeron colomexicanus, Glandularia bipinnatifida var. bipinnatifida,
Linum lewisii var. lewisii, Melilotus officinalis, Poa pratensis, Taraxacum officinale and Tragopogon dubius. The
five collection sites from lacustrine areas included 140 taxa (Table 2).
Palustrine areas included wet meadows, marshes, groundwater springs or other natural areas of stand-
ing water. Two collection sites in paulustrine areas resulted in 29 taxa (Table 2).
Disturbed Areas.—Artificial impoundments included the margins of artificial ponds and reservoirs
such as Clayton Lake, Ute Lake and Conchas Lake. The most abundant species collected from these sites
were Aegilops cylindrica var. cylindrica, Berlandiera lyrata, Calylophus hartweggii subsp. pubescens, Cirsium
undulatum, Elymus elymoides var. brevifolius, Gaura parviflora, Glandularia bipinnatifida var. bipinnatifida,
Melampodium leucanthum, Melilotus officinalis, Tamarix ramosissima, Tragopogon dubius, Xanthisma spinulosum
and Zinnia grandiflora. The nine collections sites at artificial impoundments yielded 240 taxa (Table 2).
Roadside habitat was designated mainly for areas between paved roads and their disturbed shoulders
and adjacent fencelines. The most abundant species were Chaetopappa ericoides, Convolvulus arvensis, Elymus
elymoides var. brevifolius, Elymus smithii, Gaura coccinea, Glandularia bipinnatifida var. bipinnatifida, Lappula
occidentalis var. cupulata, Linum lewisii var. lewisii, Machaeranthera tanacetifolia, Melilotus officinalis, Ratibida
columnifera, Sphaeralcea coccinea var. coccinea, Tetraneuris scaposa vax. scaposa, Tragopogon dubius and Xanthisma
spinulosum. A total of 48 collection sites from roadside habitats yielded 392 taxa (Table 2)
Miscellaneous Areas.—Badlands are highly eroded areas that have nutrient-poor soils and support
relatively little plant life. Only 1 site, surrounded by Plains-Mesa grassland, was clearly of this type; it yielded
9 taxa (Table 2).
Rocky washes consist of rocky drainages where water flows only briefly in plains or foothills. The sole
rocky wash habitat site yielded 42 taxa (Table 2).
Rocky slopes, as defined here, are rocky areas where no single vegetation type dominates. The 1 col-
lection site from this habitat type which occurred in plains-mesa grassland, yielded 45 taxa (Table 2).
Numerical Summary of Vascular Plant Taxa Collected.—Figures reported here are tentative, and
readers will note instances where we indicate that the initial determinations may be incorrect and need
further study. The time and resources were not available to update tentative, initial identifications.
This study documented 93 families, 437 genera, 911 species (including those of hybrid origin), and 45
infraspecific taxa, for a total of 956 taxa of vascular plants (Table 3). The most species-rich families were
Asteraceae, Poaceae, and Fabaceae (Table 4), which together accounted for ca. 4096 of the taxa collected. The
diversity for Amaranthaceae is higher than typically reported regionally (Taylor 2000; Arnett 2002; Reif 2006)
because we followed APGII by including a in Amaranthaceae. Likewise, Plantaginaceae now
includes many genera formerly included in S , most notably the species-rich genus Penstemon.
Finally, Boraginaceae now includes species foamed included in Hydrophyllaceae, although the number of
borage taxa overall are only minimally enlarged by taxa formerly in Hydrophyllaceae.
Schiebout et al., Vascular plants of northeastern New Mexico 1415
Taste 3. $ y oft llected by hig
Families Genera Species and Infraspecific Total Taxa
nothospecies
Fern Allies 2 2 > 0 5
Ferns 3 5 9 1 10
Gymnosperms 3 6 10 0 10
Angiosperms 85 424 887 44 931
Total 93 437 911 45 956
Taste 4. Plant families i | New Mexico with the g ber of Table 5. Plant g ith the g | | M
Family Number of Percentage of Genus Number of Number
Taxa Total Taxa Species of Taxa
Asteraceae 184 19.2 Astragalus 17 17
oaceae 128 13.4 Carex 16 16
Fabaceae 71 74 Penstemon 16 16
Brassicaceae 37 3.9 Asclepias 13 13
Rosaceae 31 32 Potentilla 11 11
Plantaginaceae 28 29 uncus 11 11
Cyperaceae 27 28 Erigeron 11 12
maranthaceae 27 28 Chenopodium 10 11
Boranigaceae 25 2.6 alea 10 12
Ranunculaceae 25 2.6 10 10
Artimisia 9 12
Oenothera 9 11
Twelve genera were represented by 10 or more taxa, including (in decreasing order of abundance) As-
tragalus, Carex, Penstemon, Asciepias, Potentilla, Juncus, Erigeron, Chenopodium, Dalea, Solidago, Artemisia, and
Oenothera (Table 5). These genera compare similarly to similar studies from nearby areas to the north (Taylor
2000; Arnett 2002), although Dalea and Oenothera are somewhat less common in other floristic studies from
the Southern Rocky Mountians. Dalea becomes a more common floristic element in the Chihuahuan Desert
to the south, and some taxa in the genus appear to be near their northernmost range at higher elevations in
the study area.
Some taxa common to abundant relatively nearby to the north in Colorado appeared to be much less
common in the study area. For example, Elaeagnus angustifolia, which is a noxious weed in Colorado, was
not commonly encountered in the study area. Cirsium arvense, another noxious weed common in Colorado
and elsewhere, was found only in isolated patches of disturbed areas.
Five taxa are confirmed as state records for New Mexico: Rorippa teres, Chenopodium berlandieri var.
berlandieri, Artemisia ludoviciana var. incompta, Hue torreyana and Rosa x harisonii. The lattermost is a
tate records for New Mexico are reported but these
horticultural taxon escaped from cultivation. Six p
require additional confirmation. Although a umber of significant range extensions were noted for some
taxa considered to be noxious weeds (Schiebout 2006), this paper does not address range extensions.
DISCUSSION
This project survey represents the first extensive floristic summary of the plant diversity occurring in this
part of northeastern New Mexico based on focused field work and documented vouchers. Some specimens
were difficult to identify with a high level of confidence irrespective of which references were used. This was
partially due to the project having been carried out in a smaller herbarium and frequently having limited
1416 J | of the Botanical R h Institute of Texas 2(2)
(or no) comparative material. Mentzelia and Hymenopappus include species that are not easily identified in
this region given existing resources, nor did infraspecific key easily for Heterotheca villosa, Aristida purpurea,
Tetraneuris acaulis and Artemisia ludoviciana.
No species listed as threatened or endangered were collected or observed during this study, which
corroborates the perceived rarity of such taxa in the study region.
Approximately 275 taxa were collected only once, whereas approximately 115 were collected only twice.
(Tentative figures are given because a few specimens remain indetermined). These numbers, along with the
species accumulation curves (below), suggest strongly that further surveying is needed to fully document
the plant taxa occurring in the area and in the various habitats. Additional collections from earlier in the
flowering season (March and April) will likely yield additional records of native (e.g., Leucocrinum montanum,
Townsendia grandiflora) and non-native (e.g., Chorisora tenella, Ranunculus testiculatus) specimens. Likewise,
additional fieldwork in September and October likely would increase the number collections and diversity
of taxa with late-summer flowering and fruiting schedules.
Species Accumulation Curves.—Species accumulation curves were analyzed for five habitat types to
assess the thoroughness of the sampling regime (Fig. 2). The curves reflect the total number of taxa known
from a habitat type as each habitat type site is visited sequentially. Species accumulation curves were calcu-
lated for plains mesa grassland, plains mesa grassland-riparian, riparian, pinyon juniper associates (including
pinyon juniper woodland, pinyon juniper woodland oak woodland ecotone, pinyon juniper woodland mixed
woodland ecotone, pinyon juniper woodland ponderosa pine woodland ecotone, pinyon juniper woodland
rocky wash ecotone, plains mesa grassland and pinyon juniper woodland) and roadside collections.
A curve that leveled off towards the right as an increasing number of sites were collected within a
habitat type would suggest that relatively few additional taxa would likely be recorded from a particular
vegetation or habitat type with further collecting, and that the collecting regime was beginning to paint
a fairly complete picture of diversity for a given habitat type. However, all of the curves reflected a steady
upward trend when the study ended (Fig. 2). The rates of increase for plains mesa grassland and roadside
areas were decidedly less than for the other three, which remained steep, and which suggest many more
collections will be needed needed from these habitat types before the floristic diversity associated with each
is well characterized. Although the curve associated with pinyon juniper associates flattened somewhat in
its middle, the collection rate accelerated again in the latter collecting sites.
Table 6 reflects the average number of new taxa collected per site for five habitat types overall, during
the first and second havles (respectively) of the total visitations to each habitat type. The two habitats with
the greatest number of additional species per new site were the plains mesa grassland-riparian and riparian.
In addition, the riparian habitat showed a greater increase in number of additional species per site visited for
the second half of the collection episodes compared to the first half (Table 6). One probable partial explana-
tion for this observation involves the late flowering periods of many Cyperaceae and Juncaceae, which are
common in riparian areas. Another reason may be that relatively few habitats that generally support these
families were visited during the first collection season. In general, all vegetation types indicated incomplete
collecting (Fig. 2, Table 6). In particular, riparian vegetation appears to need the most additional attention
for future collecting (Table 6).
Non-native and invasive taxa.—Non-native or invasive plant taxa were represented by 89 species,
or approximately 9.3% of the taxa. The percentage of non-native taxa is presented for five of the most com-
monly encountered habitats (Table 7) shows that roadsides had the greatest percentage of non-natives at
11.7 percent. This result was expected given that roadsides are corridors for the spread of invasive plants.
The figure of approximately 9.3% non-native taxa is slightly less than the figure of approximately 11% pre-
viously reported for New Mexico (Cox 2001), and compares similarly to other recent floristic studies from
the southern Rocky Mountain area (Hazlett 2004; Arnett 2002). However, small towns and the disturbed
habitats typically associated with them, such as vacant lots, were not targeted in this survey. Additional
collecting in incorporated areas likely will yield additional records of non-native taxa.
Schiebout et al., V. lar plants of northeastern New Mexi 1417
Species Accumulation Curves |
Riparian
1
1
us D SEED
; GERE D
[^7] Be AA E |
2 o seo [| ll. Roadside |
E LT a” a
53 zo
- 8 di ph mon.” Pinon Juniper Associates
> & a ls
| $^ |
| E E T | =e Plains Mesa Grassland - |
E — se M ——— POETE Riparian |
=
o om Plains Mesa Grassland |
|
ES c
1 6 11 16 21 26 31 36 41 46 51 56 61 66 71 76 81 |
Number of Collection Sites
Fic 2 Sj i lati for fi ftl fi ] tly | i hahi yp
TABLE 6. Th g ber of llected for the first time during ea ive visit for five Tas 7. The percentage of non-native species collected across five
habitat types. Total re | ftaxa collected geforthefirstti visits habitat types.
over two years. First Half re, | | llected for the first ti gefi
half of the total number of visits; Second Half t! I | half of all visi Vegetation type Collected
Non-Native (%)
Habitat Type Total First Half Second Half
Plains Mesa Grassland 9.7
Plains mesa grassland 5.3 6.3 43 Pinyon Juniper Associates 9.9
Plains messa Grassland- Riparian 17.6 22.9 124 Riparian 10.0
Pinyon juniper woodland- associates 15.5 20.2 112 Roadside 11,7
Roadside 82 12.3 4.0 Plains Mesa Grassland-Riparian 11.0
Riparian 248 19.1 29.9
At the time of this study the State of New Mexico officially listed 35 taxa as invasive and noxious weeds
(USDA 2006). Of these, 11 were collected in at least one county in the study area. Five were classified as
Class C noxious plants: Aegilops cylindrica, Convolvulus arvensis, Elaeagnus angustifolia, Tamarix ramosissima,
and Ulmus pumila. Class C species are widespread in New Mexico (DuBois 1999). Three were classified as
Class B noxious plants: Carduus nutans, Cirsium vulgare, and Conium maculatum. Class B species are only
located in certain portions of the state and should be treated as Class A plants in regions from which they
are absent (DuBois 1999). Three were classified as Class A noxious plants: Linaria dalmatica subsp. dalmatica,
Cardaria draba, and Cirsium arvense. Class A species are either absent from New Mexico or have a very limited
distribution in the state (DuBois 1999),
Comparing the distribution of plants collected to the distribution maps located at the New Mexico State
Weed Information Web Site (NMSU 2005), five species included new county records. The range of Aegilops
cylindrica now includes Colfax, Mora and Union Counties; that of Lepidium draba and Cirsium vulgare now
include Mora County; that of Linaria dalmatica subsp. dalmatica now includes Colfax County; and the range
of Ulmus pumila now includes Mora County (Table 8).
Species of Special Concern.—Few plant taxa in northeastern New Mexico are federally listed as
threatened or endangered. From the six county study area only 10 species are designated as rare by the New
] Inf tha Bataniral D h Institute of Texas 2(2)
1418
Tase 8, Declared noxious weeds collected from the study area, country distribution in study area, and New Mexico state weed status (New Mexico State University Weed
Information 2005).
Scientific Name County Distribution in Study Area State Weed Status
Aegilops cylindrica Colfax, Mora, San Miguel, Union C
arduus nutans Colfax, Mora, , San Miguel B
Cirsium arvense Colfax A
Cirsium vulgare Colfax, Mora B
onium maculatum olfax B
Convolvulus arvensis Colfax, Harding, Mora, San Miguel, Union C
Elaeagnus angustifolia San Miguel, Qua C
Lepidium draba Mora, San Miguel, Union B
Linaria dalmatica subsp. dalmatica Colfax A
Tamarix ramosissima Colfax, Harding, Mora, San Miguel, Quay, Union C
Ulmus pumila Mora, San Migel, Union C
Mexico Rare Plant Technical Council (1999; Tables 9 and 10). Of these, one collection was made of Astragalus
wittmannii. Three other species were collected that are being monitored by the New Mexico Natural Heritage
program include Euphorbia strictior, Astragalus shortianus, and Penstemon auriberbis.
Floristic Comparisons.—Taxa from the Kiowa National Grassland portion of this survey were compared
to the published results of the Comanche (Hazlett 2004) and Pawnee National Grasslands (Hazlett 1998). The
Pawnee is located in northeastern Colorado, whereas the Comanche Grassland is in southeastern Colorado.
These locations were EOS for comparison because they are similar ecosystem types (shortgrass prairie)
with similar elevati g a latitudinal gradient with nearly the same longitude. The Pawnee Grassland is
located between 40?36' and 41?00' latitude. The Comanc he Grassland is located between 37?00' and 38?00'
latitude. The Kiowa Grassland is located between 35°45’ and 36?45' latitude. A Sgrensen’s Similarity Index
was used to determine floristic similarities between the three grasslands (Serensen 1948).
Similarity between two areas is expressed by the quotient of similarity (qs), wherein (qs) = 2 (C)/A + B,
where A = the number of species of sample A, B = the number of species of sample B, and C = the number of
species common to both A and B (Barbour et al. 1987). This formula provides a measure of taxa in common
between two areas, but does not account for relative abundance of species. The higher the value of qs, the
greater the number of Sere taxa between the two areas.
hat included all taxa, a second analysis compared areas after the non-native
In addition t
taxa had been removed hom the data. Including the second dd c is of considerable interest in light of
current needs, particularly at the State level, to monitor and control the spread of noxious plant species.
Evaluating the relative abundance of non-native plants in this manner may provide insights into the control
of invasive species (Pysek et al. 2005).
The results indicate a clear latitudinal variation in species distributions (Table 11). The closer the loca-
tions are to each other the more taxa they share. For example, the relatively adjacent Pawnee and Kiowa have
an Index of Similarity of 0.46, whereas that for the relatively distant Kiowa and Comanche is 0.62 (Table 11).
The similarity of the Pawnee and Comanche is 0.51, a value intermediate between the Grasslands separated
by the greatest distance (Table 11).
Given that many i ive species hav
DAC exist to pa A on the a ey a that exclude non-native taxa may portray the
ts. For example, the inclusion of exotic spe-
cies may artificially inflate the sim) between the Kew a and Pawnee, since their exclusion leads to a lower
similarity value (Table 12). d die pos of whether exoti included (Table 11) or excluded
(Table 12), grasslands ne another ihan those further pu (Kiowa with Pawnee).
An unexpected result was obtained for qe Kiowa and Comanche Grasslands when exotic species were
tensive distributi in North America and that relatively few
Schiebout et al., Vascular plants of northeastern New Mexico 1419
T |
Rare Pant Council 1999),
Taxa County Distribution
Astragalus wittmannii Colfax, Harding, Mora
Calochortus gunnisonii var. perpulcher Colfax, Mora, San Miguel
inium robustum Colfax, Rio Arriba, Sandoval, Taos
Delphinium sapellonis Bernalillo, Los Alamos, Mora, San Miguel, Sandoval, Santa Fe
Erigeron subglab San Miguel, Taos
Eriogonum aliquantum Colfax
Grindelia acutifolia Colfax
Hackelia hir Colfax, Mora, Rio Arriba, San Miguel, Santa Fe, Taos
Ipomopsis sancti-spiritus San Miguel
Packera spellenbergii Harding, Union
Salix arizonica Mora, Rio Arriba, Taos
Tale 10 D ES g fi | | > ta J £ | ly by Ha día [ g H D n. f. il 1999).
Taxon USFWS State USFS Natural Heritage Global
of NM NM Rank
Astragalus wittmannii SoC SoC Sen 53 3
Calochortus gunnisonii SoC SoC Sen 54? G5T4?
var. perpulcher
Delphinium robustum SoC SoC Sen 54? G5T4?
Delphinium sapellonis SoC SoC Sen S? G2?
Erigeron subglaber Soc Soc Sen $3 G3
riogonum aliquantum SoC SoC 53 G3
Grindelia acutifolia SoC SoC S2 G2
Hackelia hi SoC SoC S? G4?
Ipomopsis sancti-spiritus E E E S1 G1
Packera spellenbergii SoC SoC Sen S2 G2?
Salix arizonica SoC SoC Sen $1 G2G3
TABLE 11. Sorensen’s MEN AT Index for Í federal g lands includi g I Taste 12. Serensen's Floristic Simil Index for tt federal g lands incl g ly
+ il dd me Rold sk: g I | L | £ +h FI + 4. n x Bold "| | dA g } £
diagonal are Sgrensen's Floristic Index of Similarity. the diagonal are Sorensen’s Floristic Similarity Index.
oman awnee Kiowa Comanche Pawnee
Grassland Grassland Grassland Grassland Grassland Grassland
Kiowa Grassland 448 300 223 Kiowa Grassland 397 256 166
Comanche Grassland 0.62 513 263 Comanche Grassland 0.65 387 207
Pawnee Grassland 0.46 0.51 521 Pawnee Grassland 4] 0.52 406
removed (Table 12). In thi the similarity index increased with the removal of exoti pecies. A potential
explanation for this result could be that t i y be restricted to specific habitat types associ-
ated with just one of the Grasslands. For example, ihe. Canadian River basin may support unique non-native
taxa neca EN its riparian habitat. Unknown edaphic, climatic, or biogeographic barriers may also prevent
moving farther north into the Comanche Grassland, or likewise from moving farther south
r
1420 J | of the Botanical R h Institute of Texas 2(2)
from the north into the Kiowa Grassland. The same result was obtained between the Pawnee and Comanche
Grasslands when non-native taxa were excluded, although the increase in similarity index was only 0.01.
CONCLUSIONS
This floristic study has been the first to sy ically inventory a large part of northeastern New Mexico that
focused particularly on the plains and mesas. Despite securing over 9,600 specimens, the average collection
density for the area is still not particularly high. We have not attempted to accurately estimate the collection
density given the many recent collections from northern New Mexico by colleagues (mostly) at the University
of Wyoming, some of which overlap (especially in Colfax County) this study area. Despite the limited cover-
age in many areas, we estimate this proejct has documented in excess of 9096 id the taxa present in the study
area. A priority to focus much ofthe collecting on privately held lands ived, although each property
typically was visited only once or a few times. We believe that much more field Work will be needed to fully
characterize the plant diversity of northeastern New Mexico and that future collecting should continue to
focus on prviately held lands and riparian areas, both of which will reveal many new records.
ANNOTATED CHECKLIST
The annotated checklist that follows summarizes information about each plant taxon collected from the
study region. It also includes a few taxa known from the area but not collected in this study (e.g., Packera
spellenbergii). Abbreviations used in the annotated list follow immediately below. Nomenclatural authorities
generally follow Snow (2007). In light M dius add of alternative names for some taxa, the
most commonly used synonym i n brackets following details of the species. However,
some generic changes are so commonly accepted "m Physaria instead of Lesquerella) that synonomys for
these taxa have not been included.
dedo of information in the Annotated east for each taxon.—Latin binomial (with in-
, if appropriate) Authority: total number of coll y (voucher number with collector
[no NR prior to number - Michael Schiebott NS = Neil Snow, DH- Donald Hazlett, BR = Brian Reif]);
vegetation types; elevation range in meters.
f
Key to symbols prefixing some taxa Abbreviations for Vegetation Types
SR = state record mog = montane grassland
it = potential state record fog = foothills grassland
l! = non-native (Cox 2001) pjw = pinyon-juniper woodland
I! = non-native noxious (USDA, NRCS 2006) ppw = ponderosa pine woodland
* = affinity towards a particular species mcw = mixed coniferous woodland
= affinity at the infraspecific level riw = riparian woodland
~ = species tracked by New Mexico Natural Heritage Pro- © oaw = oak wood land
gram mxs = mixed shrubland
& = known from study area but not collected rip = riparian
County abbreviations lac = lacustrine
CO = Colfax pal = palustrine
HA = Harding ari = artificial impoundments
MO = Mora rds = roadside
QU = Quay bdl = badlands
SM = San S rsl = rocky slope
UN = Uni row = rocky wash
FERN ALLIES Selaginellaceae
Equisetaceae Selaginella densa Rydb.: 7; HA (3057), SM o UN 2 66 B);
a arvense L.: 6; CO Lr MO (4729); rip-ppw, rip, Fs, pjWerow, MEWTIP, TIPApmo;
mo , rip-ppw; 1800-2600 Selaginella mutica D. C. Eaton ex Underw. var. B oid 3; HÀ
T pan Braun: 34; pa (6724), HA (6579), MO (DH- T 18h SM (8904); mxs, rip; 1,600-19
(4723), SM (6169), UN (6807); rip-ppw, oaw-rip, a rip, Selaginella una dij Hieron: 4; HA (5449) UN 8694) rip,
mog-rip, rip-pmg, pmg, rds, oaw, ari; 1400-2400 mew-tip, pmg-rip, row-rip: 1350-1750m.
Schiebout et al., V. | pl Ic nf north Mau; M
FERNS
pean
E eptennriondke (L) Hoffm.: 1; HA (DH-11716);
1750-1
Dryopteridaceae
A fragilis (L.) Bernh.: 2; CO (6655); rip, mew; 2400-2450
T neomexicana Windham: 1; HA (1333-A); pjw-row;
1750-1800 m.
Woodsia oregana D. C. Eaton subsp. cathcartiana (B. L. Rob.)
indham: 3; HA (458), UN (3235); ari, ppw-pjw, pmg-rip;
*Woodsia aff. oregana D. C. Eaton subsp. cathcartiana (B. L.
Rob.) Windham: 1; UN (5165); mcw-rip (on basalt).
Fieriüdcese.
i R DIAS J} |
coc chiensis: 1; SM (4788); oaw; 1950-2000 m
Cheilanthes eatonii Baker: 6; HA (3053), MO (DH- 12183), SM
(8868), UN (3233); pjw-row, mxs, ari, rip, rsl; 1500-1850 m.
Cheilanthes on Moore: 4; HA (457), MO (DH-12181), UN
(5164); ppw-pjw, mcw-rip, rip-row, rsl; 1250-1750 m.
Cheilanthes n o 3; HA (5448), SM (3713) ; rip, pjw-
row, pjw;
"o o Tenn 3 SM (4838), UN (2483); mxs,
w, rip; 1
m uA duco 1, MO (DH- 12180); row, 1500-1600 m.
GYMNOSPERMS
Cupressaceae
Juniperus communis L. var. A Pursh: 2; CO (4697); rip-
ppw, rip; 1800-245
Juniperus monosperma cdi Sarg.: 38; HA (4409), MO
(487), QU (3681), SM, (6078) UN (5804); pmg-pjw, rip, mxs,
pmg, as pjw-row, mxs-lac, ppw, ari, oaw, rds, pjw,
iw, rsl, r du rip-row; 1100-2200 m
Juniperus scopulorum Sarg.: 39; CO (6721) HA (8435) MO
(6244) SM (8953) UN (5179); rip-ppw, ppw, rip, rds, mew,
ppw-pjw, fog-ari, pjw, rip-oaw, caw, rip-pmg, fog, pma,
rip-row, rds-riw, ari, pjw-pmg; 1550-2450 m.
Ephedra
Ephedra mini. On 4; ee (4493), SM (6083); ari, mxs,
pmg, ppw; 1150-1
Pinaceae
Abies concolor (Gord. & e Hildebr: 2; CO (4715); rip-ppw,
mog-rip;
Picea engelmannii ie ex Engem var. engelmannii: 1; CO
(9126); mog-rip; 2550-2600
"E pedis. p edis 2; oo. (6576), MO (3361); rip;
Ti Bs mE 28; CO (6743), HA (468), MO (3158), SM
(4820), UN (5754); rip-ppw, rip, pow, pow-pjw, pjw-row,
mxs, mxs-lac, e pjw, riw, mcw-rip, rsl, pmg,-pjw,
rsl-riv; 1450-2350
Pinus ponderosa Drs ex P. € C. Lawson var. scopuloru
Engelm.: 24; CO (5320), HA (5458), MO (4752), SM (741),
UN (5174); oaw-rip, mcw-rip, rip-ppw, rip, ppw, mew, rip,
1421
bjw, pjw-row, ppw-pjw, mxs-lac, rip-pmg, mog-rip, lac;
1550-2450 m.
Pseudotsuga menziesii (Mirb.) Franco var. glauca (Beissn.)
Franco: 7; CO (5319), MO (4767), UN (5175); oaw-rip,
mog-lac, opw, mew, lac, rip-ppw, mcw-rip; 2250-2450 m.
ANGIOSPERMS
Adoxaceae
Sambucus racemosa L. var. microbotrys (Rydb.) Kearn. & Peeb.
:2; CO (6671); rip, mcw-rip; 2250-2450 m.
Agavaceae
Leucocrinum montanum Nutt. ex A. Gray: 1; UN (4218); rip-
oaw; 1900-1950 m.
Yucca baccata Torr. var. baccata: 5; CO (7932), HA (DH-12146),
MO (6376), SM (8906); rip, ppw, rip-ppw, mxs; 1550-2350
* "e aff. elata Engelm.: 4; HA (4975), SM (4849); rip, pmg,
rds; 1300-1400 m. According to one of the reviewers, "Y
elata does not occur E of Hun Co. in Ea iud
TED paniculat
INA | | i | J
vai usty ECO LE do
of Y. E Y intermedia var. ramosa, and roces
hybrids involving various taxa including Y elata. None o
these are concepts have strong support. This section of
the genus has poorly resolved species boundaries, but
yuccas in this study area are not known to fall in the ac-
cepted circumscription of Y elata”.
Yucca glauca Nutt. var. glauca: 23; CO (6513), HA (5005),
MO (2259), QU (4460), SM (6035), UN (5608); ppw, mxs,
pmg, rip, pmg-pjw, pjw, rds, ari, prng-rip, riw, row, rsl-riv;
1150-2350m
Yucca neomexicana Wooton & Standl.:
2000-2050 m
Alismataceae
ane gramineum Lej: 1; CO (9025); lac; 1800-1850 m.
held. 2; UN (8038); oaw, ari; 1550-1600
1; SM (7093); ari;
a
m.
Sagittaria latifolia Willd.: 1; MO (DH-12107); 1500-1600 m.
Alliaceae
Allium cernuum Roth: 9; CO (8766), MO (8812), SM (8929),
UN e an rip, mog-lac, mog-rip, fog-ari, oaw, fog;
1550-260
Allium esas Regel: 4; MO (6394), QU d 9804), UN
3581); pmg, rip-ppw, rip-pmg; 1400-2350
Allium geyeri S. Watson var. geyeri: 4; CO o UN (5147);
mcw-rip, ppw; 2250-2350 m
Allium Ecl S. o var. tenerum M. E. Jones: 1; CO (5305);
w-rip; 2350-240
Am perdulce S. V. sal var. perdulce: 14; HA (5840), MO
(4557), QU (3664), SM (3921), UN (3172); pmg, rip-pmg,
pmg-pjw, mxs-lac, rds, ari, pjw; 1250-1750 m
Amaranthaceae
! Amaranthus albus L.: 1; UN (1531), pmg, 1500-1550 m.
Amaranthus arenicola |. M. Johnst.: 7; HA (8258), UN (7625),
pmg, pmg-rip; 1300—1550 m. Most collections were from
sandy soils
1422
A Watson: 1; HA (7762); pjw; 1750-1800
| Pa hybridus L.: 6; CO (8999), MO (8585), SM
(8940), UN (7998); lac, rip, fog-ari, pmg-rip, oaw; pmg;
1400-20
Amaranthus palmeri S. Watson: 6; HÀ usd UN (7720); rip-
pmg, pmg, rip-row; 1350-1500
! Amaranthus retroflexus L.: 2; UN roe see 1400-1550 m.
One collection was made on lava substra
Atriplex argentea Nutt. var. argentea: 1; HA a rip-pmg;
di canescens (Pursh) Nutt. var. canescens: 18; CO (6790),
A (6592), MO (3166), SM (3928), UN (1160); rds, lac, pmg,
ari, ppw, pmg-lac, rip, rip-row, piw, fog, rda-rip, rsl, rsl-riv;
1350-1650 m
! eed gibum L.: 28; CO (8991), HA (8306), MO
(8494), SM (7044), UN (6863); lac, ppw, mog-rip, rip, rip-
omg, rip- rds, ari, fog-ari, riw, pmg, oaw, fog, pmg;
1350-2
ae a afi edi L:3; UN (1650) CO (5511), HA (717);
mis png: oe bs
ar. berlandieri: 1; UN (2605
5 pg; ES 1550 m. lected o on B substrate. All
Allred 2005, 2006
FNA A ea) confirm tns asa uis record
C
Val ex
Asch: 8; CO (8987), HA (7195), UN (2649); lac, rds, pmg, ari,
rds-rip; 1400-2500 m
I* Chenopodium aff. ci (L) Asch.: 1; SM (7034); ari;
2000-2050 m.
Chenopodium desiccatum A. Nelson: 17; HA (714), SM (8926 A),
N (241); pmg, fog-ari, ari, rip-orng, rip; 1350-2050 m.
* Chenopodium aff. desiccatum A. Nelson: 1; CO (8718); rds;
2450-2500 m.
Chenopodium fremontii S. Watson: 3; HA (1335), bis I A),
UN (1170); fog, rip-ppw, pjw-row; 1750-23
* Chenopodium aff. fremontii S. Watson: 3;CO ed HA (514),
UN (5176); rip, pjw, mcw-rip; 1550-2450
! beoe Dm L. var. glaucum: 1; SM (7035 A); ari;
le crop s um L. var. glaucum: 1; CO (8986);
lac;
Crrpodun incanum (S. Watson) A. Heller var. incanum: 45;
H 58), SM (5935), UN (952); rip-pmg, rip, pmg, pjw,
mxs, row, fog, rds, pmg-pjw, pmg-pjw; 1150-1850 m.
Chenopodium leptophyllum (Moq.) Nutt. ex S. Watson: 10; HA
(6593), MO (7816), QU (2698), UN (7702); rip, pjw, rip-pmg,
pma, rds; 1300-1800 m
Chenopodium neomexicanum Standley: 1; MO (8829); rip;
2450-2500 m.
Chenopodium pratericola Rydb.: 51; CO (7157), HA (4933), MO
(507) SM 2302), UN (7336); O ari, mog-rip, rds, pmg
rip, mxs, riw, row; 1300-2600 m. [= C. pratericola Rybd.]
* Chenopodium aff. pratericola B: 1; HA (5043); pmg;
1250-1300 m. [=C.
Chenopodium simplex (Torr) Raf.: 4; CO (7508), MO (8493), UN
(2072); mog-lac, rip, rip-pmg; 1390-2300 m.
>
T
[| laf el Dat * Al D hi PA
f Texas 2(2)
* Chenopodium aff. simplex (Torr.) Raf.: 2; CO (5338), MO (2264);
lac, rds; 1800-1950 m
Cycloloma atriplicifolium (Spreng.) J. M. Coult.: 2; HA (8311),
UN (2969); pmg; 1300-1500 m. Specimen collected from
sandy soil.
Froelichia gracilis (Hook.) Moq.: 13; HA (3041), MO (8513), UN
60); mxs, rip, pmg, pmg-pjw, rip-row; 1400-1600 m.
| Kochia scoparia (L) Schrad.: 16; CO (7165), HA (7295), MO
(2256), SM EA UN (7967); ari, lac, rip-pmg, pmg, rds,
So a
—
e]
(P M & Smit: 10; CO (6498),
HA (7225), SM (8125), UN (1651) , pmg, rip-pmg, rip,
mg-pjw, rsl, rds; 1350- 20 50m
votar (Schult) Greene: 1; HA (4625); pmg-lac;
- m. Specimen was collected from sandy soil.
! stl australis R. Br. 45; HA (7223), MO (8583), QU (2702),
M (8189), UN (7385); pmg, rip-pmg, pjw, rip, mxs, rds,
m ari, ari, row, pmg-rsl, rip-row; 1150-2050 m. [= Salsoa
tragus L]
! ies collina Pall: 2; HA (522), MO (DH-12187); pjw, rip;
Bu oo (Torr) Rydb.: 1; MO (8501); rip; 1550-1600
m.
Anacardiaceae
Rhus aromatica Aiton var. trilobata (Nutt.) A. Gray ex S. Wat-
son: 65; CO (7923), HA (5085), MO (7843), QU (3673), SM
(6022), UN (5681); rip, ppw, pmg, pmg-pjw, mxs, ppw-pjw,
rip-row, ari, rds, b riw, pmg-rip, caw, rsl, row, rip-oaw,
rsl-riv; 1100-23
Tu microphylla SAU ex A. Gray: 4; SM a QU (9685),
2605); mxs, pmg, p ppw, rds; 1200-13
Ib) O (6735),
HA (5440), MO (4728), SM (61 14), UN dei rip-ppw, oaw-
rip, rip-ppw, mcw-rip, i2 lac, rip, ppw-pjw, mog-rip, pjw;
rip-pmg; 1400-2400
~
Apiaceae
Ammoselinum popei Torr. & A. Gray: 2; SM (NS-9630); pmg,
pw; 1200-1300 m.
Angelica grayi (J. M. Coult. & Rose) J. M. Coult. & Rose: 1, UN
(7983); oaw; 1550-1600 m.
Berula erecta (Huds.) Coville var. incisa (Torr) Cronquist: 2; HA
(8293 A), UN (8008); pmg, oaw; 1300-1600 m
Cicuta maculata L. var. angustifolia i 3; CO (7918), UN
(7981); rip, mog-rip, oaw; 1550-2
I! Conium maculatum L.: 2; CO s p (6139); rds, rip;
350m
E s (Pursh) Raf: 3; UN (3595); rip-pmg, rds;
-1600
Dum Em (J. M. Coult. 8: Rose) Dorn: 4; CO (6704);
rip-ppw, rip, mew- pa mog-lac; 2250-24
Cymopterus montanus lorr. & A. Gray: 10; HA "" 9), QU (NS-
9805) SM pae P (4229); pmg-pjw, rip-pmg, pmg-lac,
jw, ri w, rds, pmg; 1350-1950 m
Harbouria trachypleura (A. Gray) J. M. Coult. & Rose: 11; CO
(4687), HA (8640), MO (6403), SM (6152); rip-ppw, pmg,
mog-rip, ppw, lac, rip, ari, pjw, pjw-ppw; 1800-2400 m.
Schiebout et al., V. lar plants of northeastern N
Heracleum sphondylium L. var. lanatum Mss d Dorn: 3; CO
6673), MO (8852); rip, rip-ppw; 2300-2
Ligusticum porteri J. M. Coult. & Rose: 3; e E oaw-rip,
-rip, mog-lac; 2250-2600
i eed (Torr) DC: 1; UMS mcw-rip; 2150-
men collected in basalt substrate.
| mos sativa iis 1; CO (8762); rds; 2450-2500 m.
—
Apocynaceae
Apocynum androsaemifolium L.: 2; HA (DH-
rip; 1550-2350 m.
— sable 6; CO (6725), HA oe UN (5658);
p-ppw, pmg, mxs, pmg-rip, riw; 1300-
ud X forbundun Greene: 1; jt a oaw;
1550-16
aceso 2; HA (8246); UN (2034); omg, pmg-rsl;
1300-1400
11729), MO (6411);
Asclepias open (Decne) Woodson var. asperula: 7; HA
(4443), MO (4545), QU (NS-9692), SM (4835), UN (6971);
pm S, mxs, pjw, ari, pmg; 1350 ;
Asclepias aaa na Woodson: 17; HA (5370), MO (7830),
QU ae SM (8901), UN (5621); mxs, rip, pmg, ari, rds,
rip; 1350-1900 m
Asis n Engelm. ex Torr: 4; CO dii HA (NS-
N (5213); rds, omg, rsl-riv; 1400-2000 m
Ni latifolia (Torr) Raf: 28; HA (7773), QU (2710), SM
5960), UN (5820); P id mxs, rds, pmg-rip, riw, rip,
row, rip-row; 1300-18
Asclepias macrosperma en 6; HA (6597), MO (8598), SM
8899); rip, mxs; 1450-1900
Asclepias oenotheroides Cham. E Schltdl: 1; UN (2152); rsl;
~
T
0-1 ;
dpi ii (A. Gray) Vail: 10; HA a mi ~ pjw,
w, pmg-rip, pmg, rds, rip-row; 1350
du E 4 CO (7162), HA ns th qo ari,
lac, pal, oaw; 1400-2000 m
da ccs dd (A. M Vail: 17; CO (7158), HA (6601),
O (8 M (5973), UN (1721); ari, lac, rip, pjw, pmg,
rds, fog-ari; 1300-2000
Asclepias tuberosa L. var. n (Woodson) Shinners: 1; SM
(6299); rip-pmg; 2200-2250 m.
Asclepias viridiflora Raf: a HA (5861), UN (5565); omg, rip,
-rip, ari; 1400-1
UE crispum ele Schltdl: 3; HA (3038), SM (6031),
N (5675); mxs, riw; 1250-1550
Araceae
Lemna minor L.: 3; SM (6127), UN (6799); rip, ari, riw; 1400-2000
m.
*Lemna aff. minor L.: 1; UN (5189); mcw-rip; 2150-2200 m.
Asparagac
paraguas. 3; CO (4281); lac, pmg, ari; 1750-2000
Aces mieu L. var. lanulosa (Nutt.) Piper: 19; CO (8736),
A (5101), MO (8799), SM (7100), UN (7986); rds, rip,
rip- see m mog-lac, rip, ari, rip-pmg, mcw-rip, oaw;
1550-2
1423
ipeo ius (Pursh) Raf. var. aan * ES - d UN (4217);
og-lac, mog-rip, rip-oak; 1
ino artemisiifolia L:1;SM cei = (9090008 m.
Ambrosia confertiflora DC.: 29; HA (7296), MO (2236), SM
(8182), UN (8012); rip- 2s rip, pmg, pjw, mxs, rds, oaw,
fog, pmg-rsl; 1350-2000
i Que DC.: i HA (7215), MO (8531), QU
(2 M (8108), UN (7706); pmg, rip, rds, pmg-rip, ari,
s rip-row; 1300-2500 m
Ambrosia tomentosa Nutt.: 8; CO (8743), E (8808), SM (7101);
rds, ari, lac, rip, fog-ari; 1800-25
Ambrosia trifida var. texana Scheele: 4; m (1851), UN (7969);
rds, oaw, rip; 1350-23
d dire e dracunculoides (DC) Nutt.: 1; UN (2391),
Antennaria a cm 3; CO (6720), HA (5456); rip,
Antennaria e s 17; CO (7898), HA (NS-9495), MO
( M (3707), UN (4235); pmg, rip, oaw-rip, ppw,
mog-lac, mog, mew, lac, mog-rip, rip-ppw, pjw, mcw-rip,
rip-oaw, eiw; 1550-2650 m
ii rosea Greene: 3; CO (4685), MO (3355); rip-ppw,
-rip, rip; 1800-2500 m
T na rosulata Rydb: 1; UN (1014); pmg; 1500-1550 m.
E la ~ 1; HA; pmg; 1300-1350
ollected in sand
pem minus Bernh. 3; um (7497), MO (8514), UN (7974);
mog-lac, rip, oaw; 1550-2300 m.
Artemisia bigelovii A. Gray: 1; MO (DH-12079); 1700-1800 m.
eo campestris L. var. o (Nutt.) M. Peck : 3; CO
; rds, mog-rip;
a aff. campestris L.: 1; UN o. pmg; 1400-1450 m.
acen conu A. W. Wood ex Carruth: 6; HA (DH-1 1708),
2250), UN (1440); rds, rip-pmg, pmg; 1300-1950 m.
"ms dracunculus L.: 6; CO (7926), MO (8809), UN (7977);
rip, oaw, mog-rip, rds; 1550-2600 m
* Artemisia aff dracunculus L.: 1; QU (2692): rds; 1300-1350 m.
Artemisia filifolia Torr.: 30; CO (6631 B), HA (8398), MO (7839),
SM (8122), UN (1993); rip, omg, pmg-rip, rds, rip-row;
1300-2450 m.
Artemisia franserioides Greene: 1; SM (BR-8872); 2800-2900 m.
Artemisia frigida Willd.: 17; CO (9028), HA (7749), MO (8819),
SM ea UN (8682); lac, rip, rds, us rip, pjw, pmg, rip,
ri, mxs, pmg-rip, oaw, fog; 1500-
Artemisia z frigida er 1; HA (757); pmg; pan 1850 m.
onqui
9; CO (7905), HA (3045), "o a Qu (2695), a 600,
UN (2143); rip, ppw, mxs, rds, rsl, rip-row; 1250-250
This report is a dips pol A co ones ee
529
pers. comm., 2008; Flora North America /ol )come
from Baca and pi s counties in ic 2 a
Artemsisa ludoviciana Nutt. var. latiloba Nutt.: 1; CO (9121);
mog-rip; 2550-2600 m. FNA lists this name as a synonym
of var. candicans (Rydb.) H. St. John. According to FNA
the nearest known occurrences are Utah and Wyoming,
although the forthcoming Floristic Synthesis of Kartesz
(pers, comm, 2008) has records for this taxon under the
subspecific epithet candicans (Rydb.) Keck as occurring
1424
SOS ane west — the o area in Guadalupe and
espec
e ludoviciana tt. er 24; CO (8787), HA
5 MO (8580), SM p UN (8028); rds, rip, ppw,
mog- da pjw-row, omg, ari, oaw, pmg-pjw; 1400-2350
m.
* Artemisia aff. tridentata Nutt.: 3; HA (725); pmg; 1750-1850
m.
Artemisia tridentata Nutt. var. tridentata: 3; HA (8627), UN
(1439) pig, rip-pmg; 1700-1850 m.
| Q V, AC
L. Welsh: 2; MO (8579), UN (1994); rip, rds; 1550-1700
a ln DC: 3; HA (8297), QU (6985), SM d
, pmg; 1150-1800 m
mud IE 2 Gray: 3; HA (8401), UN (8712); rip,
-rip; 1400-
up ih Wash A. a E HA (4418), SM (6046 A), UN
739); pmg-pjw, pmg, rip-pmg, mxs, riw; 1250-1450 m.
ms dissecta (A. Gray) Britton: 1; HA (DH-12195); pmg; 1700
mauriopsis dissecta (A. Gray) Rydb.]
Bahia a A. Gray: 3; QU (6983), SM (6058); ari-rds, mxs;
-15 :
Berlandiera iyrata Benth.: 68; HA (4407), MO (4567), QU (4477),
M (6091), UN (5545); pmg-pjw, pmg, rip-pmg, pal,
rip, mxs, rds,ari, lac, oaw, pjw, riw, pmg-rsl, rds, rip-row;
^
Bidens pU 1; EN (2536), 2. row; 1350-1400 m.
MO (8529); rip; 1550-1600
Atte
moa brachyphylla (A. Gray) A. Gray: 5; HA (8390), MO
-12188), SM (8875), UN (1611); rip, mxs, pmg-pjw, rsl;
1450-1900 m.
Brickellia is (Torr. & A. Gray) A. Ed var. californica: 1;
mg-fip; 1500-1550
Brickellia paro (L) w var. id (Wooton
Stand.) B. L. Turner: 2; CO (9027), UN (2854); lac, pmg;
28
Hm i
o
Brickellia grandiflora (Hook) Nutt. var. grandiflora: 2; CO (7921);
rip, mog-rip; 1550-2600 m.
I Carduus nutans L.: 11; CO (7473), MO t SM (6158); ppw,
lac, rip, ari, pmg, fog-ari; 1550-2300
Centaurea americana Nutt.: 1; SM (7023); iis 1200-1250 m.
[-Plectocephalus americana (Nutt.) D. Don in R. Sweet]
| Centaurea biebersteinii DC: 1; CO (7901); rip; 1550-1600 m.
Chaetopappa ericoides (Torr.) G. L. Nesom: 92; CO (4672), HA
(4308), MO (2239), QU (4465), SM (4821) UN (5229); rip-
ppw, is rds, pmg-lac, fog, pmg, pal, rip-pmg, rip, pjw, mxs,
ari, oaw, pmg-pjw, rai-pmg ; 1150-2050 m
Cnnsanterum leucanthemum L.: 1; CO (5317); oaw-rip;
2350-2400 m. [=Leucanthemum vulgare Lamark]
chsh rns (A. Gray) Greene var. wee (Wooton
wa S. i PAKE L d (2221); pings
adii Jrbatsch,
R. P. Roberts & Neubig]
Chrysothamnus vaseyi (A. Gray) Greene: 1; HA
—
8371); rip;
-1
0m.
! Cichorium intybus L: 1; SM (7046); ari; 2000-2050 m.
f Texas 2(2)
l! Cirsium arvense (L.) Scop.: 4; CO (7155); ari, lac, mog-lac,
-rip; 1800-2600 m.
Cirsium s rum A. Gray: 49; HA (7274), MO (2260), QU
M (2338), UN (7638); rip-pmg, mxs, rds, ari, rsl,
p w, rip, fog, row, rip-row; 1150-2000 m
Cirsium bs (A. Gray) Petrak subsp. parryi: 3; CO (6688); rip,
mog-lac; 1600-2450 m
Cirsium scariosum Nutt. var. coloradense D. J. Keil: 1; CO (9125);
mo 2550-2600 m
Cirsium undulatum (Nutt) Spreng. var. undulatum: 61; CO
36), HA (5465), MO (4578), QU (4484), SM (6145), UN
(5629); rip-ppw, lac, rds, ppw, rip, mog-lac, ari, mog-rip,
mxs, pmg, pjw, pmg-rip, fog-ari, riw, pmg, oaw, pmg-rsl,
rip-row; 1350-2600 m
I rir vulgare (Savi) Ten.: " CO (7941), MO (478); rip;
Conyza canadensis (L) Cronquist var. canadensis: 20; CO (9037),
HA F MO (8833), SM (8210), UN (8708); lac, ari, rip-
w, rip, omg, mxs, fog-ari, rds; 1150-2500
e pa L: 1; UN (4033); rds; 1350- ve m.
m tinctoria Nutt. var. tinctoria: 2; MO (DH-11733);
650m
Deis runcinata (E. James) Torr. & A. nds var. oo (Nutt.) B.
Boivin: 2; HA (4901), pal, rip; 1300-13
Crepis runcinata (E. James) Torr. & A. s var. runcinata: 1; CO
9062); mog-rip; 2550-2600 m.
Cyclachaena xanthifolia (Nutt.) Fresen.: 4; HA (8321), MO
(8845), UN a pmg, rip, oaw; 1300-2500 m. [=Iva
xanthifolia Nutt.]
Dietaria ae an Gray) D. R. Morgan & R L. Hartman var.
bigelovii: 1; CO (9050); mog-rip; 2550-2600 m. [FMachaer-
ntheram n bigelovii (A. Gray) Greene)
Dieteria canescens (Pursh) Nutt. var. canescens: 1; CO (8784);
rds; 2450-2500 m. [=Machaeranthera canescens (Pursh)
A. Gray var. canescens]
eod canescens (Pursh) Nutt. var. glabra (A. Gray) D. R.
n &R. L. Hartman: 1; SM (2331); rds; 1950-2000 m.
[= aa canescens (Pursh) A. Gray var. glabra)
L papposa (Vent.) Hitchc.: 9; HA (3051), MO (DH-
91), UN (1515); mxs, pmg, pmg-pjw, pmg-rsl, rds;
n 700 m.
Engelmannia pinnatifida A. Gray ex Nutt.: 87; CO (5504), HA
(5916), MO (8509), QU (6989), SM (6138), UN (5707); rds,
pmg, pmg-lac, rip, mxs, pjw, ppw, ari, pmg-rip, riw, row,
pjw-pmg, fog, pmg-rsl; 1150- 2200 m.
Ericameria nauseosa (Pall. ex Pursh) G. L. Nesom & G. I. Baird
abrata (A. Gray) G. L. Nesom & G. |. Baird: 4; CO
pees UN (8006); rip, lac, caw, pmg-pjw; 1550-1700 m.
Frigeron bellidiastrum Nutt. var. bellidiastrum: 15; HA (4946),
QU (3656), SM (3809), UN (5616); pmg, rip, rip-pmg, rds;
1250-1550 m. Many collected from sandy soils.
Erigeron canta Gray: 10; CO (4636), HA (5873 A), 5M (4520),
mg, lac, rds, fog, pmg- lac; 1 050.
g icanus A. Nelson: 57; CO (5127), HA (5899),
MO (6232), SM (6153 B), UN (5737); mxs, lac, rds, pmg-lac,
pmg, fog, mxs-lac, rip, ppw, oaw, pjw, rip-pmg, riw, ari,
pjw-ppw, pjw-oaw; 1400-2200 m.
F4
lampy
Schiebout et al., V. lar plants of northeastern N
Erigeron coulteri Porter: e (6679), rip; 2400-2450 m.
Erigeron eae . Gray var. divergens: 28; CO (4289),
HA (5429), MO mon QU (3687), SM (6165), UN (3217);
= rip-ppw, rds, opw, mog-lac, rip, mxs, pmg-rip, pjw,
mg, ari, fog-ari; 1300-23 Om
Es flagellaris A. Gray: 29; CO (4637), HA (5441), MO
4761), QU (2689), UN (7979); pmg, mcw-rip, rip, oaw-rip,
rip-ppw, mog-lac,ppw, rip-pmg, lac, mog-rip, rds, oaw;
1300-2400 m.
Erigeron leiomerus A. Gray: 1; CO (3431); mcw; 2400-2450 m
Erigeron dl w Var. oe 2; CO (9307), HA (4439);
pmg-pjw, pmg; 14
Men pectus T DC: 2; CO (7576); mog-lac, mog-rip;
0-2600
& Ta a Cronquist: Not collected but known from
tudy area in San Miguel county (Tables 10, 11).
* Erigeron aff. subtrinervis Rydb.: 4; CO (7885), MO (8797); rip,
ac, mog-rip; 1550-2600 m
— subtrinervis Rydb. ex usd & Britton var. subtrinervis:
O (6663); rip; 2400—245
BOE. Nutt. ex DC.: 5; HA 230 SM ali B), UN (5805);
g, rds, pmg, rip-pmg; 1300-1500
Evax verna Raf: 6; HA (4367), En ree i: pma, pmg-pjw,
, mxs; 1300-1450 m.
Gaillardia pinnatifida Torr: 28; HA (4601), SM (8422), UN (1359);
mg-lac, rip, mxs, rds, pjw, rip-pmg, riw, rsl, pmg, pmg-rsl,
rip-row; 1350-1850 m.
Gaillardia pulchella Foug. var. pulchella: 25; HA (4969), QU
(4488), SM (6012); rip, SACR rip-omg, ari, ari-rds, rds, mxs,
rsl-pmg, ppw; 1150-17
~ ee d xm e CO (8730), UN (2734); rds;
-2 . [See note in next entry.]
# Bon inornata Greene: 6; MO (DH-12220), UN (2048);
pmg-rsl, rds, rip-pmg, pmg; 1300-1650 m. Some authors
consider G. acutifolia and G. inornata synonyms of G.
hirsutula Hook. & Arn. e.g. :
Grindelia nuda A.W.Wood var. aphanactis (Rydb.) G. L. Nesom:
34; CO (5518), HA (396), MO (7798), QU (2707), SM (7060),
UN (7365); rds, ari, ppw, rip, pjw, rip-pmg, pmg, fog-ari,
rsl, pmg-pjw, rip-row; 1350-2500 m. [See note under G.
squarrosa.
Grindelia nuda A.W. Wood var. nuda: 12; CO (9009); HA (8372);
MO (2255); SM (8130), UN (6932); lac rip, pmg, rds pmg-
rip; 1350-2000 m. [See note under G. squarrosa.]
E tie (Pursh) Dunal var. squarrosa: 6; CO (7596),
mog-lac, ppw, rds, mog-rip, oaw; 1550-2600 m.
a me ors synonymize G. nuda under G. squarrosa.
Gutierrezia sarothrae (Pursh) Britton & Rusby: 41; CO (8985),
HA ( , MO (7815), QU (2682), SM B UN (1538);
lac, rds, mog-rip, pmg, rip, pjw, mxs, for-ari, fog, pmg-rsl,
ee pmg, i TOW; Ton 2500 m
AM y 5: HA (7276) , QU (6990), SM
(8209); rip- pmg, pmg, pmg, rds; 1150-1 400 m
Helenium autumnale L. var. montanum (Nutt) Fernald: 1; MO
(8851); rip; 2450-2500 m
Helianthella parryi A. ae 1; co (7877); rip; 1550-1600 m.
Helianthus annuus L.: 38; CO (6763), HA (8635), MO (8839),
QU (2699), SM m UN (7394); rds, lac, ari, pow, pmg,
3
fe]
a
Re
=
1425
rip, mxs, o ari, pmg-rip, oaw, pmg-pjw, fog, row;
1300-2500
Helianthus ann ps 3; SM (7039), UN (5220); ari, fog-ari,
veamos Nutt. var. fallax: 17; CO (9095), HA (8327),
, QU o SM (8105), UN (8663); mog-rip,
pe pmg, rds, pmg-pjw; 1350-2600 m
TE pea ed var. pendlar (Heiser) B. L Turner:
27; - ae SM (4889), UN (5626); pmg, mxs, pal, rds,
mxs pmgrrip, rip; 1150-2000 m
ie AME. Nutt. subsp. subrHtomboideus (Rydb.)
O. Spring € E. E. Schilling: 1; CO (8717), rds, 2450-2500
m. [-Helianthus rigidus (Cass.) Desf. var. subrhomboideus
(Rydb.) Cronquist]
Heliomeris multiflora Nutt. var. multiflora: 4; CO (8727), MO
(8798); rds, rip; 2300-2500 m. [=Viguiera multiflora (Nutt.)
S. F. Blake]
Heliomeris multiflora Nutt. var. nevadensis (A. Nelson) W.F. Yates:
3; CO (9096), UN (7992); mog-rip, rds, oaw; 1550-2600
m. [-Viguiera multiflora (Nutt) S. F. Blake var. nevadensis
A. Nelson]
Heliopsis helianthoides (L.) Sweet var. scabra ae jg
3; CO (7512), UN (8036); mog-lac, rds, oaw;
m. [=Heliopsis helianthoides (L.) Sweet var. d =
R. Fisher) Steyerm.]
iios pie oe and Standl.: 1; MO (DH-12205); rsl;
re sy pinnatum Cav.:
1500-1600
Eo canescens (DC) Shinners: 5; HA (444), MO (2248),
(8024); Bw. PU A oaw, ape oe 1950 m.
^: 10; HA (8363),
UN (7680); rip, rip- oma, a row, pmg; 1300 -1500m
* Heterotheca aff. villosa (Pursh) Shinners: 1; UN (1882); rds-rip;
; HA (DH-12210); pjw;
1 —-1600
Heterotheca villosa (Pursh) Shinners var. foliosa (Nutt.) Harms:
12; CO (7910), HA (7756), MO s E en UN (568);
ip, pjw, pmg, rip-pmg, rds; 11
? E rd villosa (Pursh) See Var. a foliosa (Nutt.)
N (2789); pmg; 1400-1450 m
TE villosa (Pursh) Shinners var. minor (Hook.)
Semple: 18; CO (9088), HA (8391), SM (6161), UN (1056);
mog-rip, rip, pjw-row, fog-ari, rip-pmg, ari, row, pmg, rds;
1300-2250 m.
? Heterotheca villosa (Pursh) Shinners var. aff. minor (Hook)
semple: 2 HA oe UN (2847); mxs, , pmg; 1400-1550 m.
(A. Gray) Semple:
18; CO (9007), SM (8889), UN (1 zu e lac, mxs, riw,
pmg, fog, rip-pmg, rip-row; 1350-2500
Hymenopappus filifolius Hook. var. cinereus (e Ib) I. M. Johnst::
6; HA (4600), MO (6185), e (4511), UN (3956); pmg-lac,
w, lac; 1550-220
"merca flavescens A. es var.flavescens: 17; HA (4328),
QU (6 Ee SM (4876), UN (5759); pmg, ari, rds, riw, rip;
11 Ds
TEC. newberryi (A. Me oe 2; CO (6684),
MO (6404); rip, iu PW: v s
Lh !
F
"m ) B.L. Turner:
1426
74; CO (5513), HA (4442), MO (4561) QU (4487), UN (5235);
rds, ppw, pmg-pjw, dp mxs, if pma, rip, pjw, ari, riw,
oaw, rsl, rip-row; 1150-
Hymenopappus ein E À co (6455), UN (2028); fog,
omg-rsl; 1350-2000
Hymenoxys odorata DC.: e HA Sen SM (3858), UN (2573);
pmg, rip-pmg, rds; 1350-1500
TE cce richardsonii (Hook) E var. floribunda (A.
Gray) Parker: 1; SM (2322), rds; 1900-1950 m
wo A (Hook.) a var. oe 4;
; pp, rip, mog-lac;
mes du 9; n (5341), HA uo P mo. lac, pmg,
ari, rip, mxs-lac; 1
Lactuca canadensis E var. longifolia piss Farw.:2; CO (7518
63); mog-lac, rds; 1950-
! Lactuca serriola L.: 22; CO (8994), e (8296), MO (8841), SM
(8962), UN (8050); lac, rds, omg, rip, pjw, rip-pmg, fog-ari,
aw; 1300-2500 m
8 Liatrus punctata Hook. var. mucronata (DC) B. L. Turner in B.
L. Turner et al.: 7; CO (8770), UN (1773); rds, pmg-rip, pmg;
1400- 2500 m. This specimen needs more attention. Not
previously known in NM [=L. mucronata DC]
Liatris Rer Hook. var. punctata: 10; CO (9029), HA (8271),
QU (2676), UN (2581); lac, pmg, rds, rip-pmg; 1300-1850
Š
zd
ead
m.
Lygodesmia juncea (Pursh) D. Don ex Hook.: 14; CO (9010),
A (6587), MO (8571), UN (2394); lac, rip, mxs, pmg, rds,
pmg-rip; 1400-1950 m
j (Pursh)D.D H
pmg, pal, rip; 1300- poH
k.:4; HA (5872);
* Ivaodesmia aff. j
83; CO (6759), HA
(5403), MO (4560), QU (3682), SM (4879), UN (5553); rds,
lac, mxs, rip-pmg, pmg-pjw, rip, pmg-lac, pmg, ari, pjw,
riw, rsl-pmg; 1250-2000 m
Melampodium decanta tom 8, A. Gray: 90; HA (4438), MO
(7809), QU (4485), SM (6081), UN (5715); pmg-pjw, pmg,
pmg-lac, rip, mxs, pjw, ppw-pjw, pjw-row, mes-lac, ari,
rds, oaw, rip- a riw, ame pjw, pmg-rsl, fog, rip-row,
rsl-riv, ppw; 1
Go a e eae Don in R. Sweet: 5; CO (9044),
O (8537), UN (1764); mog-rip, rip, rds; 1550-2600 m.
actuca oblongifolia Nutt]
Backera ae (A. Gray) W. A. Weber & A. Lave: 1; CO (7868);
rip; 1550-1600 m
Packera neomexicana | (A. Gray) W. A. Weber & A. Lóve var.
mutabilis (Greene) W. A. Weber & A. Lóve: 26; CO (5490),
HA (3307), MO (6184), SM (6151), UN (3951); rds, rip-ppw,
oaw-rip, fog, pmg, mxs-lac, ppw, rip, lac, rip-pmg, pjw,
—
S; -
Packera plattensis (Nutt) W. A. Weber & A. Love: 9; HA (5394),
SM (6 6), UN (5197 A); mxs, pmg-lac, mcw-lac, rip, pjw,
pw, rds; 1550-2000 m.
& Toa HH TUE (T. M. Barkley) C. Jeffrey: Not collected
but known from study area in Union and Harding coun-
ties (Tables 10, 11).
Packera tridenticulata (Rydb.) W. A. Weber & A. Lóve: 24; CO
(5337), HA (5890), MO (4569), SM (3781), UN (5196); lac,
I l af alk D sonal D h Partit t
f Texas 2(2)
rip-ppw, rip, lac, pmg, mxs-lac, rds, rip-pmg, mcw-rip,
jw-pmg, pjw-oaw; 1350-2450 m.
i sphacelata (Nutt. ex Torr.) Cory: » HA (5041), UN
00); pmg, rip-pmg, rip, rds; 1250-15
E confertum A. Gray: 2; HA ed vs (8601); rip;
1550-1600 m
Parthenium incanum Kunth: 1; HA (NS-9464); rsl-riv; 1550-
1600 m.
Pectis angustifolia Torr. var. angustifolia: 4; UN (2187); pmg, rds,
rip-row; 1300-1550 m.
Pericome caudata A. Gray: 3; HA sid 12121), CO (7872), MO
DH-12186); rip, rsl; 1550-17
Tan aff. oppositifolia ee Ib. ex Britton: 3; HA
(4305), MO (8516), UN (204); omg, rip; 1350-1600 m.
Picradeniopsis oppositifolia (Nutt.) Rydb. ex Britton: 27; CO
(6446), HA (8624), SM (8951), UN (1511); fog, mxs, pmg,
fog-ari, riw, rds, rip-pmg, pmg-pjw, pjw-mxs; 1300-2050
m
Picradeniopsis woodhousei (A. Gray) Rydb.: 17; HA (5870), MO
(8505), UN (5541); pmg, rip, pal, pmg-rip, rds, rip-row;
50m
Pseudognaphalium canescens (DC.) W. A. Weber: 1; HA (DH
peudognaphalur stramineum (Kunth) W. A. Weber: 6; HA
(7221), SM (8095), UN (7692); omg, rip, pmg-rip, ari;
1350-1600 m.
ee tagetina (Nutt.) Greene: 20; HA (9434), MO (4558),
pia (4457), a RE UN (2740); rip, pmg, mxs, rds, ari,
p-pmg; 1
hs EORUM (D. Don) DC.: 1; MO (6351), rds,
0-23
Pyrrocoma crocea (A. Gray) Greene var. crocea: 1; MO (BR-8424);
ff (Nutt.) W 2, Granrl-O1- :CO (67 86),
HA (6607), MO (8805), QU (4461), SM (5953), UN (5217);
rds, mog-rip, ari, ppw, rip, pmg, rip-pmg, pjw, ppw-
pjw, mxs, fog-ari, riw, oaw, fog, row, pmg-rsl, pmg-pjw;
Ratibida tagetes (E. James) Barnhart: 36; CO (9004), HA (8295),
O (7822), QU (2690 B), SM (5992), UN (7363); lac, pmg,
rip-pmg, pjw, rip, ppw-pjw, rds, ari, fog-ari, oaw, pmg-pjw,
fog; 1300-
Rudbeckia hirta L. var. pucherima Farw.: 4; CO (7449); ppw,
rip, mog-lac, mog-rip; 1 0
Rudbeckia laciniata L. var. Res (A. Nelson) Cronquist: 9; CO
d i MO (8830), UN (7997); m ppw, rip, mog-lac, mog-
p, ri w, oaw; 1550-2500
! o laciniato L.: 24; CO EU HA (4612), MO
(4554), SM (6135), UN (5522); pmg, lac, pmg-lac, rds,
mxs-lac, ppw, oaw, rip, ari, fog-ari, rip-pmg, pjw, rip-oaw;
1350-2050 m.
Senecio bigelovii A. npe var. ie la ~ 2; CO (7578), SM
); mog-lac;
ios crassulus A. i " En n oaw-rip, mcw;
2350-2450 m
Se cotilla Roue var. a (Rydb.) Greenm.: 1;
CO (9053); mog-rip; 2550-2
Schiebout et al., Vascular plants of northeastern New Mexico
Senecio flaccidus Less. var. douglasii (DC.) B. L. Turner & T. M.
Barkley: 27; HA (5397); MO (8582), QU (4478), SM (2310),
UN (5745); mxs, pmg, rip, ari, id riw, rip-pmg, pmg-pjw,
row, fog, rsl, rip-row; 1150-20
Senecio pudicus Greene: 2; CO a rip, ppw; paa Aster
ascendens Lindl: 1; UN (8035); oaw; 1550-1 m. [
Symphyotrichum ascendens (Lindl.) Nesom]
Senecio riddellii Torr. € A. Gray: 3; HA B0. MO (7823), UN
(5746); mxs, pmg, riw; 1400-1600 m. [= S. spartoides Torr.
& A. Gray var. fremontii (Torr. & A. Gray) Greeenman]
d bos aff. oo Torr. & A. Gray: 2; UN (3024), omg,
1400-145
Senecio orga dual 1; CO (6657); rip; 2400-2450 m.
So e c anadensis: 3; CO (7914), MO (8843);
OQ-rip; ne ~ m.
sooo altissima L. var. gilvocanescens (Rydb) Semple: 11;
O (9002), UN (6940); lac, rds, pmg, oaw, rip-pmg, fog;
moe 2500 m. [= Solidago canadensis var. gilvocanescens
Rydb.
* Solidago aff. canadensis L.: 1; UN (2521); rip-row; 1350-1400
m
Solidago gigantea Aiton: 3; CO (7445), SM (8914), UN (7685);
xs, rip-pmg; 1450-2300 m
Solidago missouriensis Nutt: 1; CO (7895), rip; 1550-1600 m.
E a a 2; UN (8053), HA (8621); oaw, pmg;
U
Lo]
50-185
ii sit aff. mo Bartl.: 4; HA (1343), UN (2594); pjw-row,
un 1 - 800. Union County plants collected on lava
is
Solidago p du (A. Gray) Greene: 1; CO (9082), mog-rip;
# sold igi; L var. rigida: 2; CO (8786 A), MO (8842);
m. [= Oligoneuron ridigum (L.) Small subsp.
P Es
T pei Kunth var. simplex: 1; CO (7571), mog-lac;
Solidago dee Nutt. var. pallida Porter: 1; CO (9116); mog-
rip; 2550-2600 m
Solidago velutina DC. subsp oe (A, Gray) Semple: 1;
UN (2450); rip; 1350-1
! Sonchus asper (L.) Hill: 21; io p HA (4347), MO (8527),
E 4486), SM (3784), UN (5683); lac, ppw, rip-pmg, mxs,
mg, ari, riw; 1150-2300 m.
E mt pauciflora (Torr.) A. Nelson: 9; HA (8267),
SM (5997), UN (5721); pmg, ari, mxs, riw, pmg-pjw;
1250-1850 m
* Stephanomeria aff. pauciflora (Torr.) A. Nelson : 3; HA (5892),
UN (1537); pmg-pjw, pmg; 1500-1750 m
Symphyotrichum ericoides (Lindl.) Newsom: 1; UN (2622); pmg;
1500-1550 m. Collected from a lava substrate. [=Aster
ericoides L. var. ericoides]
ido aff. ericoides (Lindl.) Newsom: 2; CO (8990),
37); lac, pjw; 1750-1800 m
o falcatum (Lindl.) Nesom: 1; CO (1839), rds;
—2350 m. [=Aster falcatus Lindl. var. commutatus (Torr.
A. G. Jones]
Symphyorichum fendleri (A. Gray) Nesom: 1; UN (1539); pmg;
0-1700 m. [=Aster fendleri A. Gray]
1427
E Hd aff. frondosum: 1; MO (8517); rip; 1550-1600
Smphyonctun laeve (L) Lóve & Lóve var. geyeri (A. Gray)
Nesom: 2; CO (1822), MO MM rds, rip; 2300-2500 m.
ee dae vis L. var. geyeri
A porteri (A. E Nesom: 1; HA (DH-12074);
1700-1800 m. [=Aster porteri A. Gray]
i Ls pod aff. porteri A. Gray: 1; HA (466); pow-pjw;
opt Pd Cavanilles: 1; HA (DH-12211); pjw;
1500-1600 m
! Taraxacum officin ake Weber ex F. H. Wigg.: 40; CO (4270), HA
3), MO (4749), SM (4508), UN (608); lac, rds, oaw-rip,
rip, mcw-rip, pmg, rip-ppw, ari, pmg-lac, ppw, mcw, mog,
mxs-lac, mog-rip, pjw, rip-pmg, oaw; 1300-2650 m.
Tetradymia canescens DC.: 1; CO (7866); rip; 1550-1600 m.
e acaulis (Pursh) Greene var. acaulis: 39; CO (5493),
HA (4606), MO (6186), SM (4530), UN (4249); rds, lac, rip-
ppw, id -lac, M mxs, ppw, lac, ari, rip-oaw, pmg-pjw,
E rip; 1350- ae
FEO aespiti
a A. Nelson: 1
CO (3413), HA (4331), UN (3565) fog, s mxs, rip, Es
rip-pmg, pmg-lac, pjw, pmg; 1350-2500
Tetraneuris d (A. Gray) Greene: 3; SM TUM mxs, rds;
letraneuris scaposa (DC) Greene var. scaposa: 73; HA (5021),
MO (8512), QU (4476), SM (4823), UN (5228); pmg, pmg-
pjw, pal, rip, pjw, mxs-lac, rds, dos TOW, ari, mxs, riw, pmg-
rip, row, rsl, rsl-riv, rsl-pmg, ppw; 1150-1800 m.
SR Tetraneuris torreyana (Nutt. o 2; HA (NS-9523),
SM (3709); pjw-pmg; 1700-1800 m. To the best of our
| ledge this is the fi I | l, asit was
not reported by Flora of North America or Kartesz (pers.
comm., 2008). Heil and O'Kane also report it for Apache
O, AZ A (21: 451) reports it from CO
Thelesperma filifolium (Hook) A. Gray var. intermedium (Rydb,)
Shinners: p a eke QU (9785); ari, mxs, pmg, rip-pmg,
bal, ppw;
mE. megapotamicum (Spreng.) Kuntze: 103; CO
(7911), HA (6569), MO (4577), QU (4495), SM (4798),
UN (5640); rip, fog, pmg-pjw, pmg, mxs, pjw, rds, ppw,
ari, oaw, fog-ari, rip-pmg, riw, rsl, pmg-rsl, row, rip-row;
50m
25
Tekspemosurudum Gray: 1; HA (DH-11644); 1800-1850
Poop acerosa (DC) Strother: 1; SM (NS-9665); ppw;
Om.
ecd ja Nutt.: 2; HA (8636); pmg; 1800-1850
m.
| Tragopogon dubius Scop.: 90; CO (5502), HA (6606), MO (496),
QU (4481), SM (4874), UN (5223); rds, lac, pmg, rip-ppw,
mcw-rip, rip, mog-lac, ari, ppw, mog-rip, fog, pmg-lac, mxs,
pal, pjw, fog-ari, pmg-rip, riw, oaw, row; 1150-2500 m.
! Tragopogon porrifolius L: 4; MO (4556), UN (3544 A); rds,
g; 1350-1700 m.
Verbesina encelioides (Cav.) Benth. € Hook. f. ex A. Gray var.
exauriculata B. L. Rob. & Greenm.: 6; MO (8528), SM 30
UN (2105); rip, rds, omg, rip-pmg, rip-row; 1300-1950 m.
1428
Vernonia marginata (Torr.) Raf: 3; UN (7679); rip-pmg, rip,
rip-row; 1350-1500 m
Xanthisma spinulosum (Pursh) D. L. Morgan & R. L. Hartman:
10; CO (5348), HA (4932), MO (7820), UN (5205); omg, lac,
rip-pmg, riw; 1300-1850 m. [=Machaeranthera pinnatifida
(Hook) Shinners var. glaberrima (Rydb)) B. L. Turner & R.
L. Hartman]
Xanthisma orc (Pursh) D. L. Morgan &R. L. Hartman
pe D. L. Morgan & R. L Harman n MO (DH-
para lada xa B. L. Turner & R. L Hartman]
Xanthisma spinulosum (Pursh) D. L. Morgan &R. L. Hartman
var. spinulosum: 105; CO (6447), HA (5844), MO (4553),
QU (4491), SM (5926), UN (8023); pmg, rds, ari, lac, fog,
pal, rip-pmg, rip, pmg-pjw, mxs, oaw, riw, row, pmg-rsl,
rip-row, ppw, rsi-pmg; 1150- 2300 m. [=Machaeranthera
pinnatifida (Hook) Shinners vat. pinnatifida]
ids d L.: 10; CO (8995), HA (8264), MO (8559),
09); lac, pmg, rip-row, rip, rip-pmg, pmg-pjw, pmg-
B E 850 m.
Zinnia grandiflora Nutt: 76; CO (1811), HA (5017), MO (2246),
QU (4475), SM (4861), UN (5568); rds, pmg, mxs, pal, pmg-
pjw, rip, pjw-row, ari, fog-ari, riw, pmg-rip, rsl, row, fog, rip,
rip-row; 1150-1950 m
Berberidac
Berberis Kup Gray: 3; CO (6716); rip-ppw, rip; 1600-2350
m.
Mahonia fremontii (Torr) Fedde: 1; SM (NS-9922); pjw;
1900-1950 m
E (Lindl) G. Don: 2; CO (7889); rip, mcw;
550-
Betulaceae
Alnus incana (L.) Moench var. occidentalis (Dippel) C. L.
Hitchc.: 3; CO (7902), MO (6390); rip, mog-rip, rip-ppw;
1550-2700 m.
Betula occidentalis Hook.: 2; CO (6703), rip-ppw, rip; 2300-2450
m
Boraginaceae
Cryptantha cinerea (Greene) Cronquist var. jamesii Cronquist:
15; i d QU (NS-9820), UN (5612); pmg, pmg-rip,
rds; 1
Copan eels (Torr. & A. Gray) Greene var. elachan-
tha |. M. Johnst.: 32; HA (4433), QU (3670), SM (6067), UN
dM pmg-pjw, pmg, E aa lac, rip, rip-pmg, mxs-lac,
ari, rds, mxs, riw; 1150-18
Cryptantha fendleri (A. s Gee 1; SM (8895); mxs;
-1 .
Mri minima Rydb.: 10; HA (5378), QU us SM (3884),
UN (3 mxs, pmg, rds, rip-pmg; 1200-1850
meu rds (Greene) Payson: 12; bs (6449), HA
(8619), UN (5751); fog, pmg, pjw, riw, ari, oaw, rds;
1400-1850 m.
*Cryptantha aff. thyrsiflora (Greene) Payson: 1; UN (1136); fog;
1850-1 ]
! Cynoglossum officinale L.: 14; CO (5314), MO (6405), UN (8030);
oaw-rip, rip-ppw, mcw- dd mog-lac, rip, pow, mog-rip,
rip-oaw, Caw; 1550-240
I Ingel Dat ai D h a t,
f Texas 2(2)
Ellisia nyctelea (L.) L.: 2; UN (4206); rip-oaw, mcw-rip;
1 950 m.
Hackelia floribunda (Lehm) |. M. Johnst.: 3; CO (7542); MO
(8853); mog-lac, rip; 2250-2500 m
& Hackelia hirsuta (Wooton & Standley) I.M. Johnston: Not
collected but known from study area in Colfax, Mora, and
: Miguel county (Tables 10, 11).
Heliotropium convolvulaceum (Nutt) A. Gray var. convolvulace-
A (8350), UN (8657); rip, pmg-rip; 1300-1600 m.
ipi ind curassavicum L. var. obovatum DC.: 1; CO (9031
A); lac ;
cocoa fendleri (A. Gray) A. Heller var. fendleri: 2; CO
(5 w-rip, mcw-rip; 2250-
o in (S. Watson) Greene var. cupulata (A.
. C. Higgins: 47; CO (5122), HA (4431), MO (4571),
al (3671), SM (4824), UN (5814); rds, lac, fog, prng-lac,
pmg-pjw, rip-pmg, mxs-lac, ari, rip, pjw, pmg, rip-oaw,
rsi-pmg; 1100-2200 m
? Lappula occidentalis (S. Watson) Greene aff. var. cupulata (A.
Gray) L. C. Higgins: 1; UN (3185); rds; 1550-1600 m.
f XE T X b. fc X. A +p a
HA (NS-9510), SM (NS-9862); pjw-pmg, pjw; 1350-1950.
Lappula occidentalis (S. Watson) Greene var. occidentalis; 24;
CO (6783), HA (4319), MO (6389), QU (4473), SM (4791), UN
(5750); rds, mog-rip, pmg, rip, mxs-lac, ari, oaw, rip-pmg,
W, riw, pjw-oaw; 1400-2300 m
eae squarrosa (Retz) Quin 3 CO (9054), MO (3168);
-rip, rip-row; 2550-260
Lithospermum incisum Lehm.: A ta (3414); HA (6578); QU
(NS-9693), SM (3345); UN (5580); pmg-lac, rip, rds, mxs-
lac, pmg, pjw, pmg-rip, rip-oaw, ari, pjw-ppw, pjw-oaw;
1150-2050 m
Lithospermum multiflorum Torr. ex A. Gray: 6; CO (6702), MO
el rip-ppw, ppw, mog-lac, rip, mog-rip; 1550-2350
BEL aff. multiflorum Torr. ex A. Gray: 1; HA (5025);
mg, 1250-1300 m
Mertensia franciscana A. Heller: 6; CO eds oaw-rip, mcw-rip,
rip-ppw, rip, mog-lac; 1800-
Mertensia lanceolata (Pursh) A. DC.: io (5239), MO (4720),
UN (4193); mcw-rip, rip-ppw, mcw, o mog-rip, mcw-
rip, rip-oaw, rds, pmg, oaw; 1800-
Nama hispidum A. Gray: 3: CO A ET pum SM (1291);
mogrrip, rds, ari; 1150-260
Onosmodium molle Michx. var. e OEA (Mack.) I.
Johnst.: 5; E (7567), UN (6958); mog-lac, riw, pmg, ari;
1400-230
Phacelia alba PE 1; MO (6408); rip-ppw; 2300-2350 m.
Phacelia integrifolia Torr. var. integrifolia: 12; HA (4341), SM
a UN (3522); rip-pmg, pmg, rip, pal, mxs; 1300-1450
Pia ON & A. Gray: 12; QU (4469), SM (3765); ari, rds,
rip-pmg, pjw, bdl, rsl-pmg; 1150-1500 m.
Brassicaceae
| Alyssum desertorum Stapf: 1; CO (4666); rip-ppw; 1800-1850
m
Schiebout et al ,V | lant:
—
| ice parviflorum M. Bieb. var. micranthum (C. A. Mey.
orn: 5; CO (3446), SM p -9864), UN (3462); rip, mew,
xa pjw-oaw, pjw; 1900-2450 m.
Aes hirsuta (L) Scop. var. pnacapa (M. Hopkins) Rollins:
50); rip, ppw; 20 0m
! Barbarea vulgaris R. Br: 5; CO (5263), MO e mcw-rip,
rip-ppw, rip, pmg-lac, mog-rip; 1550-2400
d ien (S. Watson) W. A. Weber: 1; M (NS-9915);
w; 2000-2050 m
Boe holboeli (Hornem.) var. pinetorum (Tidestr) Dorn: 1;
SM (NS-9847); rip; 1800-1850 m.
! Camelina microcarpa Andrz. ex DC.: 15; CO (5495), MO (6206),
SM (4532), UN (1778); rds, e pmg, mog-lac, ppw, ari, pjw,
oaw, rip-oaw; 1550-2300
! Capsella bursa-pastoris (L.) vem 10; CO (4706), MO (4724),
ipa 7 o UN (5187); rip-ppw, oaw-rip, mcw, mog-rip,
-rip, rds; 1700-2500 m.
| Crespo e (Pall.) DC: 2; UN (4241); rip-oaw, rds;
This may be more common than sug-
ested here En to its early flow
vá (L) Dumort.: 1; UN 2 rds; 2000-2050
—
m.
Descurainia incana (Bernh. ex Fisch. & C. A. Mey.) Dorn var.
i O (7523); mog-lac; 2250-2300 m
* Descurainia aff. incana (Bernh. ex Fisch. & C. A Mey.) Dorn
var. in O (7587); a lac; 2250-2300 m
c incana (Bernh. ex Fisch. & C. A. Mey.) Doth var.
incis CO (7884), SM e 9604), UN (1772); rip, rds;
Descurainia pinnata (Walter) Britton subsp. halictorum (Co-
ckerell) Detling: 46; CO (4294), HA (4619), MO (3155),
QU (3655), SM (4793), UN (4244); lac, pmg, s
mxs, rip-pmg, pmg-pjw, rds, rip, ari, oaw, pjw, ri
bdl,ppw; pjw-oaw; ón 2050 m. = D
var. osmiarum (Cockerell) Shini
| Descurainia sophia (L Webb ex Piatti 31; CO (5359), HA
(3285), MO (4562), SM (4840), UN (5531); lac, pmg, rip-
ppw, as pmg-lac, mxs, rip, ari, piw, pmg-rip, riw, rip-oaw;
1150-2050 m.
Dimorphocarpa wislizeni (Engelm.) Rollins: 10; HA SAM
(4492), SM (8203); prng, rip, ari, rds; 1150-1350
Draba reptans (Lam) Fernald: 1; UN (3213); pmg; d 1600
m.
Erysimum asperum (Nutt) DC.: 51; CO (6634), HA (4599), MO
(2230), SM (3931), UN (142); pmg, rip, oaw-rip, mog-
lac, rds, is ues ppw, pjw, pmg-rip, rsl, rip-oaw;
1400-24
m d E reum (Nutt) DC.: 1; CO (3398); pmg;
800-1850
Ro aff. m um (Douglas ex ee Greene: 2; SM
(8869), UN do ROS rds; onn 2000
LA
. argill
(Greene) RJ Davis 5; CO (6739), MO (6368); rip- DOW. ppw,
g-rip; 2250-2600 m.
a capitatum (Douglas ex Hook.) Greene var. capi-
u ¡CO (5494), o MO (6335), SM (2300), UN
a rds, pmg, pmg-rip, oaw, fog; 1350-2300 m.
1429
! Erysimum repandum L: 2; MO (3149), UN (4221); rip, rip-oaw;
Hesperidanthus linearifolius (A. Gray) Rydb.: 6; CO (9078), HA
(1303), MO (6210), mog-rip, fog, pmg, pjw-row, mxs,
Ha 1500-2600 m. [= Thelypodiopsis linearifolius (A :
ay Al-Shehbaz]
^ Pa aff. linearifolius (A. Gray) Rydb.: 1; SM (4830);
mxs; 1750-1800 m
! Hesperis matronalis L.: 1; CO (5318); oaw-rip; 3350-2400 m.
Lepidium densiflorum Schrad. var. densiflorum: 28; HA (5024),
MO (4563), SM (3696), UN (5642); pmg, mxs, pmg-pjw, pal,
rip-pmg, mxs-lac, rds, rip, oaw, pjw, riw; 1250-2250 m.
tl Lepidium draba L: 5; MO (4727), UN (4240), SM (9935); rds,
mog-rip, 3 rip-oaw, rip; 1350-2400 m. [= Cardaria
draba (L) D jo
Lepidium ramo m A. Nelson var. bourgeauanum
(Thell.) EIE e eN (6586), QU (3651), SM (3910), UN
(3506); rip, pal, pmg-pjw, pmg, rip-pmg, bdl, ppw, rds;
1150-1750 m
? Lepidium ramosissimum A. Nelson aff. var. bourgeauanum
(Theil) Rollins: 1; SM (NS-9678); ppw; 1200-1250 m.
Physaria fendleri (A. Gray) O'Kane & Al-Shehbaz: 10; QU (3667
B), SM (6069); ari, mxs, rds, rip-pmg, pjw, pmg, ppw;
1100-1500 m
d intermedia (S. Watson) O’Kane & Al-Shehbaz: 2; SM
rip; po 2000 m.
ied Manque A. Gray) O'Kane & Al-Shehbaz: 9; CO
4688), HA (4579), MO (3152), UN (4242); rip-ppw, oaw-
rip; pmg-lac; mxs-lac; rip, rip-oaw, mcw-rip, rds, omg;
Physaria ovalifolia (Rydb.) O'Kane & Al-Shehbaz: 15; HA (4444),
O (3159), QU (4459), SM (NS-9664), UN (6826); prng-pjw,
pjw, pmg, rip-row, ari, pmg-rip, rds, rip-pmg, bdl, ppw;
-1
! Nasturtium officinale R. Br: 4; SM (6269), UN (4234); rip-pmg,
mcw-rip, rip-oaw, oaw; 1550-2250 1 m.
Noccaea ind (A. Gray) Holub: 8; CO (5260), SM (BR-7259),
(NS-9304); rip, mcw-rip, mog-lac, ppw, mcw, pmg;
m
450 m.
m sinuata (Nutt) Hitchc: 7; CO (4274), E. n UN
(8566); lac, pmg-lac, prng, rip, pmg-rip; 1550-1850
m teres MA Stuckey: 1; MO eroan l 2 2500
m. The m.
2008) are o Co., AZ and BR d cd
counties,
L
Ires ] 5.1 INI (20705
Hi
g, rds; 1550-2000
m.
Stanleya E ia. Britton var. pinnata: 1; CO (6789);
ds; 1800-18
Thelypodium mearum D Endl. ex Walp. var. integri-
hal : 3; HA (DH 72), MO (DH-12106), UN (1595);
Tepe wrightii A. "id subsp. weg 1; HA pie rip;
ecim from s soil.
! resp arvense L.: 6; ae (6228). HA mA a (3154),
N (4237; rip-ppw, mog-lac, mxs, rip, rip-oaw, oaw;
oo m.
1430
Cactac
Conphanto Moi (Nutt.) Britton & Rose var. arizonica
(Eng T. Marshall: 2; HA (422), UN (5582); pmg,
1400- 1800m
T Aa coccineus Engelm. var. coccineus: 1; SM (3702),
HA NS 28), pw, BIN prg; 1709 om
] 4f
jd SM (3900),
pjw; 1 450-15001 m.
cn d bachii (Terscheck ex Walp.) F. Haage var.
us (Britton & Rose) L D. Benson: 1; QU (NS-9681);
pmg; mob
w viridiflorus Engelm. var. o 3; CO (4271
58); lac, rip-pmg; 1400-1
Opuntia eeu Im-Dyck var. exeun 3; CO (6490),
SM ); ppw, mxs, ari; 1250-205
Orpuntia je (Haw) DC. var. m. 7; HA (6588), SM
(8151), UN (7335); rip, pjw, mxs, rip-pmg, pmg; 1400-1800
m. [Cylindropuntia imbricata (Haw.) Kunth var. imbricata]
T phaeacantha Engelm.: 15; CO (7144), HA (4934),
(6205), SM (5928), UN (5583); ari, omg, rip-pmg, pmg-pjw,
pal, mxs, ppw, rds, oaw; 1300-2250 m. Material rede
for varietal determination; see next two entrie
Opuntia phaeacantha Engelm. var. dela x co (6767),
A (5004), UN (5650); rds, omg, riw; 1350-
Opuntia polyacantha Haw. var. polyacantha: 4; dal a 6), HA
(5062) SM (6129); ppw, rip, rip-pmg, pmg; 1300-2350
m
mo
Campanulaceae
Campanula parryi A. Gray var. parryi: 1; CO (9046); mog-rip;
—2600 m
Campanula rotundifolia L: 7; CO (6626); rip, mog-rip, rds, ppw,
mog-lac; 1550-2600 m
Triodanis perfoliata (L.) neni 2; SM (6121); rip, rip-pmg;
nnabac
cats ERU L; Š a (8323), UN (1207); pmg, rip, rds,
Celtis reticulata Torr.: : E ne -9459), UN (3574); rip-pmg, rds,
rsi-riv; 1400-1600 m
Humulus lupulus L. var. neomexicanus A. Nelson & Cockerell:
2; CO (7897), MO (8791); rip; 1550-2500 m.
Caprifoliac
! Lonicera ae 1; SM (3368); ale 2000-2050 m. Speci-
en was collected from cla
Symphoricarpos occidentalis E ie CO (9092), MO (8820),
N (7984); mog-rip, rds, rip, oaw; 1550-2600
* a. i occidentalis Hook: 1; CO (25) mog-
lac; 2250-23
Valeriana E E Gay 1; MO (3357); rip; 2450-2500 m
Valeriana edulis Nutt. ex Torr. & A. Gray var. edulis: 4; CO (6731),
MO (4772); rip-ppw, mog-rip, mog-lac, lac; 2250-2450
m
Caryophyllaceae
Cerastium brachypodum ala ex A. Gray) B. L. Rob.: 1; MO
(4719); mog-rip; 2350-240
f Texas 2(2)
JUuiTdi Yi
fe B g. subsp. vulgare (Hartm.) Greuter
& Burdet: 1; CO (6697); rip-ppw; 2300-2350 m
Cerastium nutans Raf. var. nutans: 2; MO (6333), SM (3695); rds,
pjw; 1900-2300
Eremogone fendleri d Gray) Ikonnikov: 8; CO (7414), MO
(61 bn SM (6111); ppw, rds, mog-lac, rip, mog-rip; 1550-
m. [=Arenaria fendleri A. Gray var. fendleri]
Tuum pide (Fenzl) Farw. var. texana (B.L. Robins.)
A (723); pmg; 1750-1800 m
E Vids (Torr. & A. Gray) Nutt. ex A. Nelson: 1; UN
(1956); rds; 1700-1750 m
Paronychia jamesii Torr. & A. Gay 7; HA (8618), QU oes
8192), UN (2904); pmg, mxs, rds; 1150-1850
T sessiliflora te 6; HA (8617), UN m pmg,
, Isl, fog; 13
messer ide 2 W. A. Weber & R. L. Hartm.: 1;
; mcw-rip; 2250-2300 m.
Pe anti E p (6162); rip; 1950-2000 m
d^ d Poir: 2; CO (7555); mog-lac, oaw-rip; 2250- 2400
Gime SUBE Hook. subsp. hallii (S. dE C. L. Hitchc: 1;
O (9064); mog-rip; 2550-2600
Ceratophyllaceae
Ceratophyllum demersum L.: 1; UN (7378); ari; 1550-1600 m.
Cleomaceae
Cleome serrulata Pursh: 4; UN (8681); pmg-rip, rip, rip-row;
1350-1550m
Polanisia dodecandra (L.) DC. var. trachysperma (Torr. & A. Gray)
H. Iltis: 15; HA (6576), MO (7701), UN oe rip, pmg,
pmg-rip, fog, rds, rip-row; 1350-1750
Commelinac
ea b 18; HA (3086), MO (DH-12215), 5M (8132),
UN (2982); mxs, rip, pmg-rip, rds, pmg, pmg-pjw, rip-row;
Tradescantia occidentalis (Britton) Smyth: 25; HA (4944), QU
(3654), SM (8873), UN (5734); omg, rip, rds, mxs, rip-pmg,
riw, pmg-rip, riw, pmg-pjw; 1150-1900 m.
Convolvulaceae
s sepium (L.) R. Br. var. qs oe N. H.
Holmgren: 1; SM (3903); pjw; 1450-1
T. 58; CO (5342), SES MO (8560),
M (4518), UN (5735); lac, oaw-rip, rds, lac, pmg, mog-lac,
ari, fog, ppw, pmg-lac, rip, ppw, fog-ari, rip-pmg, pjw, riw,
oaw; 1300-2500 m.
Convolvulus equitans Benth.: 21; HA (4306), SM (5933), UN
(7633); ea pjw, rip, rds, rip-pmg, pmg-pjw, row, rip-row;
1350-1750
* Cuscuta aff. ae Engelm.: 1; UN (2948); pmg; 1450-1500
E cosas. "i bea Choisy: 2; HA (8322), UN (1957); pmg,
D
Cusc a "Mes 1; UN (DH-1181 1); pus el
Em alsinoides (L.) L.: 2; UN (998); pmg, 1
Evolvulus nuttallianus Schult.: 16; HA (5003), Mo Pues SM
(3896), UN (5549); pg, rip, pjw, rip-pmg, riw, pmg-rip;
1250-1750 m
Schiebout et al., V. pu cpu A d AT i RA
n uc vu A. Gray: 1; MO (DH-12189); rip;
1500—160
ar us Egle Hallier f.: 1; HA (DH-1218); rip; 1500-1600
oon leptophylla Torr: 34; HA (6611), MO (8575), QU
91), SM (8200), UN (7711); rip, mxs, pjw, pmg, rip-pmg;
1150—1800 m.
Cornaceae
Cornus sericea L. var. sericea: 1; CO (7899); rip; 1550-1600 m.
Crassulaceae
Sedum integrifolium (Raf.) A. Nelson: 1; CO (6669); rip;
2400—2450
Cucurbitacea
E E Kunth: 28; HA (6573), MO (7842), QU
(2711), SM (6015), UN A rip, pmg, rds, mxs, rip-pmg,
riw, ari, fog; 1300-
Cyclanthera us (Torr. EE Gray) Arn.: 2; SM (8129); pmg-
rip; 1400-1450 m
peraceae
Bolboschoenus maritimus (L.) Palla subsp. dera (A. Nelson)
öve & D. Love: 6; CO (9024), HA (DH-12062), MO (7848),
SM (8982), UN (7395); lac, pmg, fog-ari, ps 2050m
Carex aquatilis Wahlenb. var. aquatilis: 1; CO (5276 A); mcw-
rip; 2250-2300 m.
Carex brevior (Dewey) Mack. ex Lunell: 6; CO (7608), HA (5473),
SM (6174), UN (5808); mog-lac, rip, rip-pmg, mcw-rip;
1300-2300 m.
Carex emoryi Dewey: 3; CO (5276 B), SM (9852), UN (4264);
mcw-rip, rip-oaw, rip; 1800-2300 m
Carex gravida L. H. Bailey var. lunelliana (Mack) F. J. Herm.: 4;
MO (4758), UN (6811); mog-rip, ari; 1550-2400 m.
Carex hystricina Muhl. ex Willd.: 1; UN (8066 A); oaw;
550-1600 m.
Carex pellita Muhl. ex Willd.: 8; CO (6750), HA (5476), MO
(4760), UN en rip-ppw, mog-lac, ppw, mog-rip, rip,
ari; 1550-2500 m. [Carex lanuginosa Michx]
uL. Dewey: 1; MO (6388), rip-ppw; 2300-2350
m.
Carex occidentalis L. H. Bailey: 12; CO (5321), SM (3755), UN
ds 65); oaw- is pmg, He ~ rip, ppw, pjw, rip-pmg,
cw-rip, oaw
*G SN occidentalis | H. e 1; CO (5360); lac; 1800-1850
m inops dd subsp. heliophila (Mackenzie) Crins 4; CO
(65 O (4765); mcw, ppw, lac, rip-ppw; 2000-2450 m.
Eo pensyhanica Lam. var. digyna Boeck]
* Carex aff. i Subsp. come (Mackenzie) Crins.: 1;
MO (4764); lac; 2400-245
^. L INI 7E 7
, "
Carex petasata Dewey: rip, ari; 1550-1600
a a Boott: 1; HA (DH-11664); 1750-1800 m.
Carex siccata Dewey: 1; CO (6749); rip-ppw; 2300-2350 m.
Carex sprengelii Dewey ex Spreng.: 1; CO (3443); mcw;
Carex stenophylla Wahlenb.: 6; CO (4710), MO (3359), UN
(5148); rip-ppw, rip, mew, mog-rip, mcw-rip; 1550-2500 m.
1431
Cai lata Boott: 1; MO (8857 Aj; rip; 2450-2500 m
Corer piace Michx.: 7; HA (5472), MO (8503), SM (8983),
UN (8071); rip, ppw-pjw, fog-ari, ari, oaw; 1550-2050 m.
de eth Muhl.: 3; HA (4926); UN (7387); pmg,
Om.
Then edens L. var. leptostachys Boeck.: 1; MO (8569);
rip; 1
Cyperus 2 1; MO (DH-12214); rip; 1550-1650 m.
a cd 10; HA (8357), E i 12197), UN
p, mxs, rip-pmg, pmg; 1300-1
opes ad L: 1; HA (DH-12213); rip, e 1500- 1700
(Willd) Schult.: 1; UN (1629); pmg-pjw;
TR aff. ot
1650-1700 m.
Eleocharis palustris (L) Roem. & Schult.: 52; CO (5332), HA
(4616), MO (4759), SM (6171), UN (4263); lac, mcw-rip, rip,
mog-lac, ppw, mog-rip, pmg-lac, pmg-pjw, pmg, ppw-
pjw, mxs- lac, rip- uo ari, fog-ari, oaw; 1300-2600 m.
Schult) Link ex Bluff & Fingerh.: 1;
CO (9038); lac; 1800- n m. [=Eleocharis coloradoensis
(Britt.) Gilly]
Schoenoplectus acutus (Muhl. ex Bigelow) A.Lóve & D. Lóve
var. acutus: 3; CO (9006), HA (DH 12125), SM (8964 B);
lac, fog-ari; 1600-2050 m. Reported but not evidently
confirmed I NM, if correct these specimens will confirm
its presen
? Schoenoplectus pungens (Vahl) Palla var. aff. pungens: 29; HA
(5469), MO (8498) QU (NS-9798), SM (3874), UN (5744);
rip, pal, pmg-rip, pmg, ari, riw; rds; 1300-2050 m. More
recent information suggests these are probably variety
longispicatus (Britt) S. G. Sm., since var. pungens is not
nown from NM or adjacent states.
Schoenoplectus tabernaemontani (K. C. Gmel) Palla: 17; CO
(5364), HA (5464), MO (8539), QU (4506), SM (6167), UN
(4266); lac, rip, ppw-pjw, rip-ppw, ari, fog-ari, pmg-rip,
mcw-rip, oaw-rip, oaw; 1150-2 i
Scirpus pallidus (Britton) Fernald: 6; CO (8782), HA (5474), MO
(8847), UN (8068); rds, mog-lac, rip, ppw-pjw, rip, oaw;
1550-2500 m
Elaeagnaceae
I! Elaeagnus angustifolia L.: 3; A (NS-9706), SM (6018); mxs,
fog-ari, pmg; 1150-
Euphorbiaceae
Argythamnia mercurialina (Nutt.) Müll. Arg.: 1; UN (7693); rip-
pmg; 1450-1500
Chamaesyce fendleri (Torr. & A. Gray) Small: 23; HA (5073), MO
(780 2 3 (2673), SM (8909), UN (5540); da ll mxs, rip,
pmg-rip, pmg-pjw, rsl-riw, ppw; 1
cromaence ind (Engelm) Small: 1; UN D rip-row;
Came obscene (Engelm.) Small: 1; HA (8385); rip;
m. Specimen collected from sandy soil.
Bs (Engelm. Small: 6; HA P n (NS-9734),
SM (6065), UN (6945); omg, mxs, rds; 1150-1650 m
Chamaesyce missurica (Raf) Shinners: 12; HA me UN (7624)
pmg, rip-pmg; 1300-1700 m.
1432
Chamaesyce serpyllifolia (Pers.) Small: 3; HA (7726), MO (8564);
pjw, rip; 1550-1791 m
Midi stictospora (Engelm.) Small: 2; HA (DH-11656),
N (2746); rds; 1300-1850 m
Croton texensis (Klotzsch) Müll. Arg: 37; HA (8320); QU (2669);
SM (5994); UN (2101); pmg, rip-pmg, mxs, rds, ari, prg-rsl,
rip, rip-row; 1150-1600 m.
Euphorbia brachycera Engelm.: 2; HA (DH-11784), UN (9334);
mxs; 1600-2000 m.
SR £uphorbia dentata Michx. var. dentanta: 20; HA (6602), MO
(7814), QU (2667), SM (8948), UN (1637); rip, rip-pmg, rds,
fog-ari, rsl, pmg-rsl, rip-row; 1300-2050 m. As best we can
determine our report is a state record and represents a
southwestward expansion of the nearest portion of the
range from southeastern CO (Las Animas and Baca cos.)
and Cimarron Co., OK.
Euphorbia marginata Pursh: 24; HA (8634), MO (8577), UN
(7626); pmg, pjw, rip, rip- Ma ari, rds, fog, row, pmg-rsl,
pmg-pjw, rip-row; 1300-1
Euphorbia spathulata Lam.: 9; $^ dm QU (NS-9793), SM
: 39); pmg-pjw, rds, pjw, pmg, rsl-pmg, ppw; 1150-1450
~ aptat strictior Holz: 4; HA (5064), SM (8202); pmg;
50-1350. Most specimens collected from sandy soil.
siae sylvatica Garden ex L. subsp. sylvatica: 6; HA (5038),
N (982); pmg; 1250-1600 m. Specimens collected from
andy soils.
Tragia ramosa Torr.: 6; HA (6551), MO (8506), QU (NS-9725), UN
(5591); rip, mxs, pmg-rip, riw, pmg; 1150-1600 m.
Fabaceae
Amorpha canescens Pursh: 5; HA (4401), MO (6381), SM
(6300), UN (8014); pmg-pjw, rip-ppw, rip-pmg, ari, oaw;
] 350m
Amorpha fruticosa L.: 3; SM (8127); pmg-rip; 1400-1450 m.
Astragalus adsurgens Pall. var. robustior Hook: 3; MO (6198),
U ; ppw, rds, rip-oaw; 1900-2200 m. [=A. laxmannii
o var. ii qu Ish]
is Douglas ex G. Don: 3; CO (5246), UN (5199);
p, pmg; 1800- 2300 m
Pa o (Hook.) A. Gay var. bisulcatus: 10; CO
(5330), HA (4614), SM (4523), UN (5200); pmg, lac, pmg-lac,
mcw-rip; 1500-1850 m
Astragalus crassicarpus Nutt. var. crassicarpus: 5; HA (5055), SM
(NS-9936), UN (3511), pmg, rds; 1300-1850 m.
m drummondii Douglas ex iu 2; CO (7504), UN
mog-lac, rip-oaw; 1900-230
Aerial E Nutt.: 2; CO (5128), j ieee rds, pmg;
E
E tum ^ Gray var. hallii. 2; UN (4251); rip-oaw, oaw;
1550-1950 m.
Astragalus MB Douglas ex AK var. m M. E.
ones: 2; SM (3904); ppw, pjw; 1200-1
irum jouflokds Hook.: 3; HA (DH- ae B (NS-9927),
N (3205 Bj; pmg, pjw; 1800-1950 m
Asi missouriensis Nutt. var. missouriensis: 30; CO (5487),
HA (4611), MO (3161), QU (NS-981 7), SM (4515), UN (3252);
f Texas 2(2)
rds, pmg-lac, pmg, pmg-pjw, rip, mxs-lac, rip-row, lac, pjw,
pm , mxs, ari; 1400-2050 m
Astragalus mollissimus Torr. var. mollissimus: 33; HA (5842), QU
(3675), SM (3836), UN (6820); pmg, pmg- E rip, ari, oaw,
png: rip, nu rip- row, rs rsi-riw, v bdl, BI 150-2000 m
DC B y:3 QU
(NS-9700), SM (NS-9926); pmg, rds, } pjw; 1150-1950 m.
Astragalus puniceus Osterh. var. puniceus: 1; CO (9060); mog-
rip; 2550-2600 m
* Ac tragalus iS
à
EB
D
Pursh: 1; HA (DH-11750); 1850-1900
Astragalus racemosus Pursh var. racemosus: 6; CO (5267), SM
d UN (3545); mcw-rip, rds, pmg, oaw; 1350-2300
- Angas shortianus Nutt.: 1; UN (3478); rds; 1500-1550
ill tenellus Pursh: 1; HA (4615); pmg-lac; 1800-1850
m. Specimen collected from sandy soil.
~ Astragalus wittmannii Barneby: 1; HA (DH-11709), pmg;
1800-1850 m. Collected from limestone substrate.
Dalea aurea Nutt. ex Pursh: 15; HA (8426), UN (6967), rip, omg,
rip-pmg, ari, pmg-rsl, rsl, i 1350-1700 m.
* Dalea aff. candida Michx.: 1; UN (2358); rds; 1400-1450 m.
Dalea candida Michx. var. oligophylla (Torr) Shinners: 31; CO
(7929), HA (3042), MO (7837), QU (2684), SM (8181), UN
(7334); rip, pjw, pmq, rip-pmg, mxs, ari, rds, oaw, pmg-rsl,
fog, rip-row; 1350-2050 m.
Dalea enneandra Nutt.: 9; HA (8358), MO (7819), QU (7006),
SM (8194), UN (2505); rip, pmg, rds, rip-pmg, rsl, rip-row;
1150-1600 m
Dalea formosa Torre 14; HA (NS-9470), QU (4482), SM (9649);
ari, mxs, Caw, pjw, rsl-riw, pjw-pmg, rsl-pmg, pmg, ppw,
rds; 1100-1800 m.
i jamesii Mt & A. Gray: 13; HA (4395), SM (4851),
pmg-pjw, pmg, rip, mxs, rds, oaw, pjw, pmg-rip,
a 1250- 2000 m
Dalea lanata Spreng. var. lanata: 1; HA (8369), rip; 1300-1350
m.
Dalea nana Torr. var. carnescens (Rydb.) Kearney & Peebles:
1; HA (3063 B); mxs; 1500-1550. Collected from a rocky
substrate.
Dalea nana Torr. & A, Gray var. nana: 4; HA (8256); UN (2845);
pmg, rip-pmg; 1300-1550 m
Dalea purpurea Vent. var. arenicola (Wemple) Barneby: 10;
CO (6766), HA (7219), UN (1883); rds, pmg, mxs, rip-pmg;
1350-1850 m.
* Dalea aff. purpurea Vent.: 3; HA (451), UN (1095); ppw-pjw,
pmg, rds; 1350-1850 m.
Dalea purpurea Vent. var. purpurea: 19; CO (5479), HA (5090),
QU (4496), SM (3922), UN (1597); rds, m pmg-pjw, ari,
mxs, pjw, pmg-rip, oaw; 1300-2500
? E ok Vent. aff. var. a 1; UN (3994); rds;
m. de d Gray) Shinners: 10; HA (8455), MO (8515),
QU i 97 ps SM (5996), UN (6849); rip, pmg, ppw-pjw, rds,
150-1850m
i so (Nutt.) Spent var. villosa: 1; UN (1017); pmg;
550m
Schiebout et al., V. lar plant
Desmanthus cooleyi (Eat.) Trel.: 20; HA (8386), MO (7835), QU
is - SM (6002), UN (7715); rip, pmg, mxs, ari, rds, rip-
250-2 00 m
Il. ex B. LL Rob. & Fernald:
; UN (2441); rip; 1350-1400 m.
! Ec triacanthos L.: 2; HA (529), UN (2401); pjw, rip;
600m
Chom BAM Nutt. ex Pursh: 17; CO (6756), HA (4904),
O (8 se SM (7118), UN (5673); rip, a pmg, pal,
rip-ppw, p-pmg, riw, oaw; 1300-
Hedysarum ils Nutt. var. boreale: 4; HA (ou rip, pmg,
jw-mxs; 1600-1850
Hoffmanseggia glauca Pies Eifert: 10; HA (4398); SM
(5924); UN (2069); pmg-pjw, rip-pmg, rds, pmg-rsl, pmg;
Lathyrus eucosmus Butters & H. St. John: 9; CO (7545), MO
ind SM (4810), UN e A rip, caw, rip-pmg,
jw, rds, rds-riw; 1600-225
Lathyrus vont (S. wth te: 2; CO (4679), MO (4738);
rip-ppw, mog-rip; 1800-
Lathyrus oe (ine var. leucanthus (Rydb.) Dorn.: 2;
CO (5279), UN (4239); mcw-rip, rip-oaw; 1900-2300 m.
ls taxon boundaries of this s others were discussed
M by Johnson & Allred 200
mi polymorphus Nutt. var. incanus (J. G. c e ~ ex
Rydb.) Dorn: 1; UN (3205 A), pmg; 1550 -16
Lupinus argenteus Pursh var. argenteus: 2; CO e mog-lac,
r Om.
ipis em Pursh var. argophyllus (A. D S. M 3;
CO (8772), UN (NS-9310); rds, ppw; 2200-
xU plattensis S. Watson: 7; CO (5133), n m UN
(5575); rds, pmg-lac, fog, pmg, pmg-rip; 1350-2000 m.
Lupinus pusillus d var. S iid 3; HA (5387), MO (3145);
mxs, rip; 1550-16
! Medicago lupulina E n CO (4705), HA (5451), MO (4730),
SM (8937), UN (4232); rip, rip-ppw, mcw-rip, mog-lac,
ppw, mog-rip, ppw, ppw-pjw, mxs-lac, mog-rip, lac,
rds, fog-ari, pmg-rip, pjw, pmg, mcw-rip, rip-oaw, oaw;
1400-2500 m
| Medicago polymorpha L.: 1; CO (5488), rds; 1900-1950 m.
| Medicago sativa L: 13; CO (6784), HA (5460), MO (8834), SM
(7070), UN (705); rds, ari, rip, pjw, oaw; 1350-2500 m
! Melilotus albus Medik.: 35; CO (8998), HA (6604), MO (8565),
SM (8918), UN (7323); lac, rip, rip-pmg, ppw-pjw, pjw, pmg,
mxs, ari, rds, pmg, pmg-pjw, fog, rip-row; 1300-2500 m
| mie officinalis (L) Pall: 79; CO (5516), HA (4323), MO
(4546), QU (4501), SM (6049), UN (5237); rds, lac, rio-ppw,
pmg, ari, Mes -lac, ppw, mog-rip, fog, pmg-lac, mxs, rip,
pjw, ppw-pjw, fog-ari, rip-pmg, oaw; 1150-2500 m.
Mimosa in Ortega var. biuncifera (Bentham)
Barneby: 3; HA (4954 B), UN (861); omg, rds; 1300-1400 m.
Mimosa borealis A. Gray: 6; HA (5032), QU (4483), SM (NS-
9648), UN (2135); pmg, ari, riw, rsl, ppw; 1150-1400 m
os d Pursh var. bigelovii A. Gray: 11; CO (5124)
A (5057), UN (3208); rds, rip-ppw, ppw, rip, pmg;
re 900 m.
Oxytropis lambertii x Oxytropis sericea: 1; CO (9114); mog-rip;
2550-2600 m
1433
cen Nutt. var. sericea: 9; CO (5358), HA (3291), MO
(62 ee e s lac, rip-ppw, mog-rip, rds, ari, mcw-rip,
pmg; 1
pedamelan aro (Pursh) J. ita 2; CO (7550),
mog-lac, oaw; 1550-
Ta a (To orr. & A. Gray) an le HA (4312), QU
(NS-9716), SM (8195), UN (6870); omg, rip, pmg-rip, rds;
1100-1550 m
Prosopis glandulosa Torr. var. glandulosa: 13; HA (6565),
QU (4489), SM (6001); rip, pmg, rip-pmg, mxs; ari, rds;
1150-1600 m.
Psoralidium lanceolatum (Pursh) Rydb.: 6; CO (6474), HA (4985),
37); ppw, rip, mxs, pmg-rip, pmg; 1350-2050 m.
Psoralidium tenuiflorum (Pursh) Rydb.: 76; CO (7472), HA
(5915), MO (6241), QU (2701), SM (8917), UN (7307); ppw,
lac, fog, pmg, pjw, rip, mxs, rds, ari, rip-pmg, riw, oaw,
-rsl, rsl, row, M row; 1250-2000 m.
Robinia neomexicana A. Gray var. neomexicana: 11; CO (5327),
HA (5399), MO eet UN (8060); oaw-rip, mog-lac, rds,
mxs, rip, oaw; 1450-2350 m
| Robinia pseudoacacia L.: 2; HA (510), UN (2862); pjw, pmg;
-1
Senna roemeriana (Scheele) Irwin: 6; SM (5930); rds, mxs, ari;
0-1500 m.
Sophora nuttalliana B. L. Turner: 40; CO (4640), HA (4396), MO
(4551), QU (4467), SM (6155) UN (5755); pmg, lac, pmg-
pjw, rip-pmg, mxs, pmg-lac, rip, pjw, mxs-lac, rds, rip-row,
ari, fog-ari, riw; 1150-2050 m.
Thermopsis montana Nutt. var. abel (A. Nelson) Dorn:
1;UN ); rip-oaw; 1 om
Thermopsis montana Nutt. var. montana : 7; CO (4703), MO
(4774); rip-ppw, mcw-rip, ppw, mog-lac, mog-rip, lac,
~
aw; 1800-260
! Trifolium dinde L.: 6; CO (6648), MO (8813), rip, mog-lac,
, rds, rip-ppw; 2250-2600 m
| um ae L:7; CO (7556), MO (814) E. oe rip,
mog-lac, mog-rip, rip-ppw, mcw-rip; 1
TN americana Muhl. ex Willd. var. americana: Hs E (ie
MO (8793), UN (5201); rip-ppw, rip, mcw-rip, pow, mog-
lac; 1800-2500 m
Vicia americana Muhl. ex Willd. var. minor Hook.: 21; CO (4642),
MO (4773), SM (6150), UN (1720); pmg, lac, ppw, mog-rip,
rds, rip-ppw, ppw, rip, rip-pmg, pmg-pjw, rip-oaw, pjw-
; -2450 m.
Vicia ludoviciana Nutt. var. ludoviciana: 12; CO (4301), HA
(4584), SM (3793), UN (4130); lac, omg, pmg-lac, rip-pmg,
pmg-pjw, rds, mxs-lac, pjw; 1350-1950 m
Fagaceae
Quercus gambelii Nutt: 21; CO (6442), HA (9491), MO (4725),
SM (9839), UN (8059); fog, rip-ppw, oaw-rip, rip-mcw, ue
mog-lac, rip, pow, mog-rip, rip-ppw, oaw; 1550-250
m.
Liebm.: 6; HA (4411), MO (4550), SM (8892), UN
(572 6); pmg-pjw, mxs, rds, riw, rip; 1350-1900 m
Quercus X undulata Torr.: 23; HA (5459), MO (6207), SM
(4792), UN (1753); rip, pjw-row, ppw-pjw, lac, ppw, rip-
1434
row, oaw, mxs, rip-pmg, pjw, riw, rds, pmg-pjw, rsl-riv;
m
135 :
* Quercus aff. X undulata Torr.: 2; HA (3312), SM (3708); mxs-lac,
pjw; 1750-1
Gentianace
Centaurium e (Buckley) Fernald; 2; HA (7241), MO
(DH-11731); rip-pmg; 1350-1400 m. [=Zeltnera calyca
(Buckley) Mansion]
Frasera speciosa Douglas ex Griseb.: 3; CO (7581); mog-lac,
ppw, oaw; 1550-2350 m
Gertionaparyi Engelm: 1; HA (8326); omg; 1300-1350. Speci-
collected from sandy soil.
en detonsa (Rottb.) Ma var. elegans (A. hu N.H.
Holmgren: 1; CO (9056); mog-rip; 2550-2600
Geraniaceae
! Erodium cicutarium (L) UHér. ex Aiton: 23; CO (4681), HA
(4605), MO (6211), SM (3721), UN (1438); rip-ppw, mcw;
pmg-lac, mxs, ppw, mxs-lac, rds, Me = ari, fog-ari, pjw,
rip-pmg, pmg, iu pur 1550-24
Geranium atropurpureum A. Heller var. p— 5;
CO (6734), MO in UN (9324); rip-ppw, rip, rds, pow;
2150-2500 m.
Geranium caespitosum E. James var. caespitosum: 9; CO
6658), UN (7993); rip, ppw, mog-lac, rds, mog-rip, oaw;
50-2600m
a
Un
Geranium richardsonii Fisch. & Trautv.: 2; CO (6659); rip, mog-
rip; 2400-2600 m
Grossulariaceae
ji aureum ai var. aureum: 3; UN (6846); ari, rip-oaw;
50-1950
Ne cereum ee 7; CO (4674), UN (3255); i ppw, mog-
ip, mew, mcw-rip, rds, ari, ppw; 1600-2600
Ribes leptanthum A. Gray: 4; CO (3427), SM e UN (4213);
mcw, rip, rip-oaw, oaw; 1900-2450 m.
Hydrangeaceae
Jamesia americana Torr. & A. Gray var. americana: 1; CO (6630);
rip; —2450 m.
Philadelphus microphyllus A. Gray: 1; CO (7912); rip; 1550-1600
m.
Hyperic
dise Ho. L.: 1;CO (7583); mog-lac; 2250-2300 m.
Iridaceae
Iris missouriensis Nutt.: 11; CO (5298), HA (4624), MO (4735), SM
-9897); oaw-rip, mcw-rip, pur He P ppw, mcw;
pmg-lac, mog-rip, oaw, rds; 170
Sisyrinchium demissum Greene: 4; CO ean, a (5142); oaw-
rip, rip, mcw-rip, pmg-rip; 1500-2450
* Sisyrinchium aff. montanum Greene: » MO (4768), lac;
2
Juncaceae
Juncus arctici r. balticus (Willd. Trautv.: 24; CO (4282),
HA pue MO (4757, SM (7078), UN (4262); lac, mcw-rip,
pmg, rip-ppw, rip, ari, mog-lac, ppw, ud rip, pal, pjw,
oaw, rds, pmg-rip, rip-oaw; 1300-2600
Juncus bufonius L.: 1; MO (8596); rip; 1550- m
f Texas 2(2)
Juncus dudleyi Wiegand: 5; CO (7483), HA (476), MO (8859), UN
; ppw, ppw-pjw, rip, pmg-rip; 1400-2500 m.
"UHR a d SM did ^r pw; 1900F du
ar. .L. Hitchc.: 6;
CO s MO aes), SM (6320); ppw, mog- -lac, mog-rip,
rip, rip-pmg; 220
Juncus filiformis L.
1550-2400 m.
Juncus M a 7; HA (5454), MO (8504), SM ~
U , bjw, pjw-row, rip-pmg; 1400-2200
Juncus D 2; CO (9146); ppw, mog-rip; 2250- UT
600 m
x do (5329), UN (3262); oaw-rip, ari;
m.
Juncus nodosus L.: 1; CO m HA (DH-11718), UN (8067);
pw, oaw; 1550-2
Juncus tenuis Willd.: 1; itt (7384) ari; 1550-1600
Juncus torreyi Coville: 8; HA n MO (8502), k (7689);
p-pmg, rip, ari; 1300-1600
n comosa E. Mey.: 1; CO (6639) rip; 2400-2450 m.
Lamiaceae
ifl N O (7564), MO (8724), SM
(6113); mog- -lac, rds, rip, rip- DD 1950-2500 m.
Hedeoma drummondii Benth.: 2; HA (8610), SM mm pmg,
mxs; 1 900m
| Marrubium vulgare L.: 16; CO (7876), HA (4977), MO (6195),
SM n UN (1201); rip, -lac, pmg, mxs, rds, ppw,
mg, ari, oaw; 1300-220
TREE arvensis L.: 8; CO is P (8309), MO (8794), UN
il mog-lac, mog-rip, ppw, pmg, rip, oaw; 1300-2600
T fistulosa L. var. menthifolia (Graham) Fernald: 4; CO
66), mog-lac, ppw, rip, rds; 1550-2350 m
pisi ose Nutt.: E HA (4359), MO (6338), QU (4497),
SM (5995), UN (5699); rip-pmg, rds, ari, rip-pmg, riw, pmg;
11 is 2300 m
Monarda ei L. var. occidentalis (Epling) Palmer &
Steyerm.: 2; HA (5067); pmg, rip; 1300-1350 m
a me 3; CO (6709); rip-ppw, rip, ppw; 2250- 2450
T reflexa Hornem.: 17; HA (3077), MO (8530), SM (8955),
UN (1203); mxs, rip, fog-ari, pmg, rds, rip-pmg, row, pmg-
pjw; 1300-2050 m
Stachys pilosa Nutt. var. pilosa; 2; CO (7875); rip, ppw;1550-
2300 m. These specimens were misidentified originally
as S. palustris L, which occurs in the northcentral and
northeastern US.
Teucrium laciniatum Torr.: 36; HA (5904), MO (2231), QU (4462),
SM (6094), UN (5555); pmg, pmg-pjw, mxs, rip, rds, ari, pjw,
pmg-rip, riw, oaw; 1150-2200 m
Liliaceae
Calochortus gunnisonii S. Watson var. gunnisonii: 1; CO (7529);
om
& ibus oiii S. Watson v Var. ape Puen? Cockerell:
collected but olfax, Mora
a Can Min 27l Inti
(Tables 10, 11).
Linacea
Linum aristatum BE 2; SM (3808), UN (5731); rip-prng,
riw; 1400-14
Schiebout et al., Vascular plants of northeastern New Mexico
Linum berlandieri Hook. var. berlandieri: 1; UN (5206); pmg;
1450-1500 m.
Linum lewisii Pursh var. lewisii: 38; CO (4298), HA (4592), MO
(8789), SM (4509), UN (4245); lac, rip-ppw, rds, ppw, mog-
rip, n lac, omg, mxs- as rip-pmg, rip-oaw, row,
pjw-pmg, pjw; 1350-26
M pratense (Norton) T 1; QU (NS-9808); pmg;
—1450 m.
em (Engelm,) A. Heller: 12; HA (4594), MO (2240),
QU (NS-9780), SM pum UN (3560); pmg-lac, rip, pmg,
rds, mxs, pjw; 1150-19
Linum ican Pursh var. compactum (A. Nelson) C. M. Ro-
gers: 7; CO (6433), HA (5837), QU (NS-9790), UN (354);
1300-2000 m.
Linum rigidum Pursh var. rigidum: 4; MO (6199), SM (7094), UN
(6956); ppw, ari, rip-pmg, pmg; 1450-2200 m.
Loasaceae
Cevallia sinuata Lag.: 1; QU (7005), ari-rds; 1150-1200 m.
Mentzelia decapetala (Pursh ex Sims) Urb. & Gilg ex Gilg: 2; MO
8604), UN (2161); rip, rsl; 1350-1600 m
* Hence dad (Nutt.) A. Gray: 1; QU (4474 A); ari;
1100-
Mentzelia Ped (Nutt. A. Gray var. multiflora: 9; CO (6477),
QU (6980), SM (7119), UN (7978); ppw, ari-rds, ari, rds, oaw,
pmg, row; 1150-2050
? Mentzelia multiflora (Nutt.) A. var. aff. multiflora: 1; QU
(NS-9680); omg; 1150-1200
Mentzelia nuda (Pursh) Torr. & A. = var. nuda: 4; QU (4474 B),
SM (5979), UN (1776); ari, pmg, rds; 1150-2000 m.
* Mentzelia aff. nuda ind Torr. & A, Gray var. nuda: 1; CO
(5349), lac; 1800-1850
di nuda (Pursh) Torr. z A Gray var. stricta (Osterh.) Har-
n: 34; CO (7890), HA (7184), QU (6988), SM (5956),
p rip, pmg, rip-pmg, rds, rsl, fog, pmg-rip, rip-row;
Mentzelia nuda (Pursh) Torr. & A. Gray aff. var. stricta (Osterh.)
LE 3; QU (2693), UN (2785); rds, pmg; 1300-1450
Mene ced Nutt. ex Sims: 1; HA (6567); rip;
Nun e DETENER Nutt. ex Sims: 1; MO (8597); rip;
1550-1600 m.
Mentzelia reverchonii (Urb. & Gilg) H. J. Thomps. & Zavort: 2;
HA (7222), UN (1198); pmg, rds; 1350-1650 m.
Malvaceae
aod parvulum A. Gray: 3; SM
-1450 m.
cali involucrata (Torr. & A. Gray) A. Gray var. involucrata:
N (6843); ari, oaw, rds; 1550-2000 m
Ne 11; CO (4704); HA (5867), MO (8511),SM
(8933), UN i) ii ~ pmg, lac, rip, fog-ari, mcw-rip,
rip-pmg; 14
Sidalcea bos n i vat. ec 3; CO (6722); rip-ppw,
rip, mog-rip; 1550-2600 m.
Sphaeralcea angustifolia (Cav) G. Don: 19; MO (DH-12060),
Al SM (4858), UN (2620); pmg-pjw, pmg, rds,
mxs, pjw, rip-pmg, riw, rip, pmg-rsl, ppw; 1200-1500 m.
=
6066), H IN [57720 , riw;
1435
Sphaeralcea coccinea (Nutt.) Rydb. var. coccinea: 75; CO (5361),
HA (4404), MO (4544), QU (2679), SM (4831), UN (6896); lac,
pmg, rds, fog, pmg-lac, pmg-pjw, pmg- ds mxs, rip, pjw,
riw, mcw-rip, oaw, rsl-pmg, ppw; 1150-2200 m
iic iiia A. Gray var. iit A Ai (5398) MO
(8581), S 332); mxs, rip, rds;
Mart
Proboscidea m (Mill) Thell subsp. /ouisianica: 2; HA
UN (7647); pmg; 1300-1500 m. Collected from
2 do E
Melanthia
Veratrum calor Durand: 2; CO (5272); mcw-rip, mog-
lac; 2250-2300 m
Zigadenus elegans Pursh: 2; CO (7536); mog-lac; 2250-2300
m.
Moraceae
! Maclura pomifera (Raf) C. K. Schneid.: 1; HA (DH-12056);
1500-1550 m.
! Morus alba L.: 1; HA (NS-9458); rsl-riw; 1550-1600 m.
Myrsinaceae
Lysimachia ciliata L: 2; CO (7402); ppw, mog-lac; ppw, mog-
lac; 2250-2300 m
Nyctagin
Abronia fora Nutt. ex Hook. var. fragrans: 23; HA (5027),
QU (4500), SM “i UN (625); rds; pmg, rip, rip-pmg,
ari; e 1600
Mirabilis albida We Heimerl: 2; CO (1830 A), HA (8328);
, pmg, 1300-2500.
* Mirabilis aff. hirsuta (Pursh) MacMill.: 2; UN (7332); rip-pmg,
mg; 1400-1550 m.
Mirabilis linearis (Pursh) Heimerl: 38; CO (6788), HA (5053),
MO (7825), QU (2666), SM (8944), UN (5678); rds, pmg, rip,
pmg, ari, fog-ari, rip-pmg, riw, oaw, fog, rsl, row, rip-row;
1300-2050 m
icis multiflora (Torr.) A. Gray var. multiflora: 6; CO (6469),
e uU SM (4877), UN (2162); ppw, rip, rds, pmg-pjw,
(zs
2 13
Mirabilis d (Michx) MacMill.: 2; UN (5688); riv, oaw;
Oleaceae
ics aaa Nutt. var. mls 2; HÀ (8472), SM
(3 rip, rip-pmg; 1400-1
Tus vus dus L: 1; HA (NS- D ok -riw; 1550-1600 m.
Onagrace
Coo hare (Benth.) P. H. Raven subsp. fendleri (A.
Gray) To id H. Raven: 2; SM (8920), UN (3952); fog-
ri, pmg; 1 :
? o Ec (Benth.) P. H. Raven subsp. aff. fendleri
(A. Gray) Towner & P. H. Raven: 2; UN (3212), QU (NS-9727);
pmg; 1150-1600 m. Collected from rocky substrate.
Calylophus hartwegii (Benth.) P. H. Raven subsp. pubescens (A.
Gray) Towner & P. H. Raven: 18; HA (4415), QU (4458), SM
(5982) UN (4151); 1150-1750 m.
alylophus lavandulifolius (Torr. & A. Gray) P. H. Raven: 23; CO
(5482), HA (733), QU (NS-9813), SM (8205), UN (1521);
I
1436
rds, pmg-lac, a do -rsl, rip-pmg, pmg-pjw, rip-oaw,
rip-row; 1150-1
Eo o 2 P. H. Raven: 38; HA (4898), QU
(2686), UN (6848); pal, omg, rds, riw, pmg-rip, ari, fog;
uen atu Raf. var. glandulosum (Lehm.) Dorn: 4; CO
800); mog-lac, rip; 2250-
a coccinea mites Pursh: 105; CO (5123), HA (5087), MO
(4549), QU (4499), SM (5931), UN (5749); pmg, rds, fog,
ppw, pal, mxs, rip-pmg, rip, pmg-pjw, pjw, rds, ari, oaw, lac,
pjw, riw, mcw-rip, row, rip-row, rsl-pmg; 1100-2250 m
Gaura uia Douglas ex Lehm: 33; CO (6762), HA (7253),
MO (2261), QU (7008), SM (5985), UN (1668); rds, lac, ari,
ip, pmg, ppw, mxs, fog-ari, oaw; 1150-2200 m
[=Gaura mollis Jame s]
Gaura villosa Torr. var. villosa: 13; HA (4940), QU (7007), UN
(317); pmg, rds, pmg-rip; 1150-1550 m
Ludwigia peploides (Kunth) P. H. Raven: 1; UN (7370); ari;
SI
Oenothera albicaulis Pursh: 46; CO (5496), HA (5843), MO
(3147), QU (4468), SM (6064), UN (5771); rds, rip-ppw,
pmg, rip-pmg, rip, pmg-pjw, mxs, pmg-lac, mxs-lac, ari,
w, riw; 1100-2250 m.
d Too z oo Pursh: 2; MO (6342), SM (2299); rds;
mE canescens Torr. a dl 3; CO (DH-11766), UN
(6813); ari, pmg; 1400-
Oenothera cespitosa Nutt. var. pM 1; SM (NS-9642); ppw;
1200-1250 m
Oenothera cespitosa Nutt. var. o (Rydb.) Cronquist:
1; CO (5289); oaw-rip; 2350-24
Oenothera coronopifolia Torr. & A. pun 17; CO (7459), HA
(6582), MO (8815), SM (8870), UN dd Hi mog-rip,
fog, pmg, rip, rds, mxs, rip-pmg; 1250
Oenothera elata Kunth var. hirsutissima (A. E ay ex TR )
Cronquist: 7; CO (7543), HA (8423), MO (8491), UN (2438);
mog-lac, rip, ppw, ari; 1450-2300 m
a ms Kunth var. aff. hirsutissima (A. Gray eu Wat-
on) Cronquist: 1; MO (6382); rip-ppw; 2300-2350
T engelmannii (Small) Munz: 2; HA (4938), UN "m
pmg, pmg-rip; 1300-1400 m.
uve flava (A. Nelson) Garrett: 5; HA (5431), MO (3146),
M (3348), UN (5184); mxs, mxs-lac, rip, omg, mcw-rip;
;
Oenothera pallida Lindl. var. runcinata (Engelm.) Cronquist: 1;
(DH-11735); 1600-1650 m
no villosa Thunb. var. strigosá (Rydb.) Dorn: 3; CO
N (8034); rip, mog-rip, oaw; 1550-2600
m. rugis var. ee (Rydb,) Dom: 1 MO
7845); pmg; 1550-1600
# Oenothera villosa (Kunth) P. " Raven var. villosa: 1; MO (8825);
rip; 2450-2500
Orchidaceae
Corallorhiza sp.: 1; MO (4766); lac; 2400-2450 m
Corallorhiza striata Lind!.: 2; CO (2661); rip-ppw, mog-lac;
1800-2300 m
f Texas 2(2)
Platanthera purpurascens (Rydb.) Sheviak & W. F. Jenn.: 1; CO
9042); mog-rip; 2550-2600 m.
Orobanchaceae
Castilleja o Gray var. integra: 32; CO (4689), HA (4432),
MO (4566), SM (7107), UN (3946); rip-ppw, mog-rip, fog,
E ae mxs, rip, pjw-row, mxs-lac, rds, x row, ppw, lac,
w, rip-pmg, rip-oaw, pmg; 1350-2600 m.
Castileja an icm ex Hook. var. miniata: 1; CO (6690);
; 2400-2
Castilleja sessiliflora ee 5; HA (4590), QU oo SM (4525);
-lac, pjw, lac, pmg, rip; 1100-1
Orobanche ludoviciana Nutt. var. pero (Nutt) Beck: 7;
HA (8240), SM (6059), UN (993); pmg, mxs, rip-pmg;
-1650 m.
Orthocarpus luteus Nutt.: 2; CO (9073), UN (8002); mog-rip,
oaw; = :
Pedicularis procera A. Gray: 5; CO (5273), MO (8803); mcw-rip,
mog-lac, mog-rip, rip; 2250-2600 m.
Oxalidaceae
Oxalis corniculata L.: 4; SM (6143), UN (3243); rip, rip-pmg, ari;
1400-2000 m.
Oxalis violacea L.: 1; CO (6629); rip; 2400-2450 m.
Papaveracea
a Gray: 11; CO (5114), MO id UN (692);
rds, fog, ppw, pmg, oaw, row; 1450-2
Argemone polyanthemos c G.B. SS. 7; UN (5569);
mg-rip, pmg; 1350-1600 m
Argemone squarrosa cin var. squarrosa: 6; UN (6953); pmg,
m. A number of collections were
e from lava e
E aurea Willd. var. aurea: 6; CO (5294), MO (3362);
ae 5-9410); caw-rip, mcw-rip, mew, rip, oaw, pmg;
0-2500 m.
? e aurea Willd. var. aff. aurea: 1; MO (8510); rip;
0-1
Corydalis aurea Willd. var. occidentalis Engelm. ex A, Gray:
10; CO (4675), MO (3141), QU (NS-9740), SM (3832), UN
156); 1150-1850 m
! Eschscholzia californica Cham. subsp. californica: 1; SM (NS-
9626); pmg; 1250-1300 m.
Phrymaceae
Mimulus glabratus Kunth var. jamesii (Torr. & A. Gray ex Benth.)
A. Gray: 1; UN (5193); mcw-rip; 2100-2200 m.
Plantaginaceae
* decias aff. parviflora Lindl.: 1; CO (9031 B); lac; 1800-1850
Md plantaginea (E. James) Rydb.: 2; MO (4736); mog-rip,
rip-ppw; 2300-2400 m. [=Synthris plantagenia (E. James)
th
Benth.]
ll Linaria dalmatica (L.) subsp. dalmatica: 1; CO (9057);
mog-rip;
Nuttallanthus texanus s D.A. Sutton: 4; SM (6286), UN
(3228); rip-pmg, pjw, ari; 1550-2250 m. [=Linaria canaden-
) Dum. Cours. var. texana (Scheele) Pennell]
E albidus Nutt.: 1; UN (3493); pmg; 1500-1550 m.
Schiebout et al., Vascular plants of northeastern New Mexico
Penstemon e o HA d SM (4842), UN (5554);
pmg, rds, pmg-rip; 1300-1
Penstemon ongustiolus ud ex a hy var. Dun (A. Heller)
Rydb: 1; CO (5115); rds; 1850-1
~ Pen emon auriberbis Pennell: Me n (4427), UN (4164);
pmg-pjw, pmg, pmg-rip, riw, rds; 1300-1650
ind barbatus (Cav.) Roth var. torreyi (Benth.) h Gray: 13;
O (6647), HA (6561), MO (7803), SM (8898), UN (7964);
ii rip- iiA mog-lac, rds, pjw-row, ppw-pjw, pmg, mxs,
w; 1550-2450 m
Person oe Pennell: 3; UN (8193) pmg, rds; 1500-2000
ha os fendleri A. Gray: 2; HA (4598), SM (4527); pmg-lac,
lac; 1800-1850 m
Penstemon glaber Pursh var. brandegeei (Porter) Freeman: 3;
CO (7501), UN (8022); mog-lac, rds, oaw; 1550-2500 m
Penstemon gracilis Nutt.: 1; CO (7531); mog-lac; 2250-2300 m.
Penstemon inflatus Crosswhite: 1; CO (6633); rip; 2400-2450 m.
Penstemon jamesii Benth: 28; CO (5484), HA (5900), MO (4564),
a ), SM (6159), UN (6954); rds, ea ppw, pmqg, ari,
, rip-pmg, pjw-pmg; 1150-2
pres rydbergii A. Nelson var. dl 2; 16 (6511), UN
4210); ppw, rip-oaw; 1900-2400 m.
Penstemon secundiflorus Benth: 19; HÀ (765), QU (NS-9760),
S HE N (9424); prng, rip, rds, mxs, pjw, rip-pmg,
pj w-pmg, rsl-pmg; 1150-1850 m
CAM panic Rydb.: 2; HA (5104), UN (5529); rds;
Penstemon strictus Benth.: 2; CO (7444); ppw, mog-rip;
2300-2600 m.
Penstemon e Gray var. virgatus: 1; CO (9094); mog-rip;
2550-2600
Pntagoerepo or 2; SM (6110), UN (5619); rip, prng-rip;
Wenig ca L.: 9; CO (4682), HA an MO (8576),
17); rip-ppw, rip, rds, ari; 155
! POEM major L. var. major: 2; MO HE rip, pmg;
1550-1600 m
=
Plantago patagonica Jacq.: 47; CO (5121), HA (5845), MO
(4541), QU (6992) SM (6063), UN (5556); fog, pmg-lac, omg,
rds, s rip-pmg, pjw, ari, mxs, ari, pjw, riw, oaw, rip-oaw,
rsl-omg, ppw; 1100-2300 m
Veronica americana Schwein. ex Benth.: 1; co (7558); mog-lac;
0-2300 m.
Veronica catenata Pennell: 12; HA (4974), MO (8796), SM
(3846), UN (5665); rip, rip-pmg, riw, mcw-rip, oaw, ari;
—2500 m
Veronica eerie L. var. xalapensis (Kunth) H. St. John & F. W.
Warren: 6; HA (3325), UN (5815); 1300-1800 m
! Veronica pne L. var. humifusa (Dickson) Vahl: 1; CO
(3408); pmg-lac; 1800-1850 m.
Poaceae
Achnatherum hymenoides (Roem. & Schult.) Barkworth: 40;
CO (6779), HA (5389), MO (7856), SM (4534), UN (2882);
rds, fog, ppw, mxs, rip, pmg, mxs, lac, ari, pmg-rip, oaw,
pjw-mxs; 1250-2050 m. [=Oryzopsis hymenoides (Roem.
& Schult] Ricker ex Piper]
1437
Achnatherum robustum (Vasey) Barkworth: 10; CO (8744),
ae 30), UN (1804); rds, rnog-rip, ari, fog-ari, oaw, pmg;
vae cylindrica Host: 6; CO (5500), MO (6225), SM (4539)
N (5534); rds, ppw, lac, pmg-rip, oaw; 1400
Agropyron triticeum Gaertn.: 1; SM (NS-9846); rip; T ie
m. [=Eremopyron triteceum (Gaertn.) Nevski]
Agrostis exarata Trin.: 5; CO (7954), HA (8484), SM (8883), UN
(7314); rip, mog-rip, rip, mxs, rip-pmg; 1 0m
ind md Willd.: 2; CO (7946); rip, mog-lac; 1250- 1600
TEL stolonifera L: 11; CO (7950), HA (7255), MO (8865),
S ); rip, mog-lac, rds, ppw, mog-rip, rip-pmg, pmg,
mxs; 1350-2500 m
k: aequalis var. aequalis: 2; CO (7603), MO
8861); mog-lac, rip; 2250-
Andropogon gerardii o 7; HA (7789), MO (8551), QU
(2719), UN (7320); pjw, rip, rds, oaw, rip-pmg, pmg, rip-
row; 1300-1800 m.
un hallii Hack.: 12; HA (465), MO (8867), UN (7669);
mg, ppw-pjw, rip, rip-pmg, row, rds; 1300-2500 m.
mes ascent L:8 E - 036); pmg-tsl, rip-pmg, rds,
, rip-row; 1300-17
m His Hee oen ex Willd.: 1; HA (8374); rip;
300-1350m
No
Un
o
o
us a Vasey 11; UN (2043); pmg-rsl, pmg, rds, rip;
mos pee Nutt. var. longiseta (Steud.) Vasey: 86; CO
(5510), HA (5853), MO (7855), QU (7013), SM (8216), UN
(6919); rds, pmg, rip-pmg, rip, pal, pjw, pjw-row, mxs, -
ari, riw, pmg- row, a pmg-tsl, ppw; 1150-2200
Aristida purpurea r. purpurea: 29; CO (6777), P
5380), QU Dn EM on UN (6839); rds, mxs, a
pmg-pjw, ari, pjw, oaw, pmg-rip, rip-row; 1300-1
Aristida purpurea Nutt. var. wrightii D iE 6; HA M
SM (6109); rip, pmg, mxs; 1300-1
! Avena fatua L.: 1; UN (1672), rds; mu Dm
¡elias om (Torr) Nash: 2; CO (9132); mog-rip,
soos bain (Lag.) Herter: 2; SM (7028), UN (2555);
| m M (L) Keng var. songarica (Rubr. ex
Fisch. & C. A. Mey) Celarier & Harlan: 16; HA (8652), QU
ons, "i 2039) pmg, rds, pmg-rip, pmg-rsl, rsl, rip-row,
rip; 1
oe nos (DC) Herter subsp. torreyana (Steudel)
Allred & Gould: 49; HA (6617), MO (8540), QU (7011), SM
(8144), UN (7667); rip, pmg, pjw, rip-pmg; rip, rds, mxs, ari,
mg-rsl, pmg-pjw, row; 1150-2000 m. Specimens were
identified initally as B. saccharoides.
cim springfieldii (Gould) Parodi: 6; HA (3109),
SM (8159), UN (1933); mxs, rip-pmg, rds, rsl, rip-row;
~
Bouteloua O (Michx) Torr. var. caespitosa Gould
&K a: 5; QU Q718), SM (8885), UN (2037); rds, mxs,
p pmg; 1300-2000 m
S iun E (Michx.) Torr. var. curtipendula: 62;
1438
HA (7781), MO (8544), SM (8967), UN (7318); pjw, pmg,
ne P. row, mxs, rip, rds, fog-ari, ari, les rip, oaw,
w, fog, rsl, rip-row; 1150-2050
sora ip (Nutt) J.T. Columbus: ay CO (4268),
5833), MO (2266), SM (8134), UN (6891); lac, pmg,
a rip, mxs, rip- Pa pjw, riw, pmg-pjw, rsl, fog, omg-rsl,
rip-row; 1150-2250
Bouteloua eriopoda e 4; HA (3102), MO (DH-12185),
SM ); mxs, pmg-rip; 1400-1650
Bouteloua gracilis (Kunth) Lag.: 78; CO (9018), HA (7782), MO
(8863), QU (2715), SM (8970), UN (7310); lac, mog-rip, rds,
pjw, pmg, rip, rip-pmg, pjw-row, mxs, fog-ari, oaw, row,
fog, pmg-pjw, pmg-rs!; 1300-2
Bouteloua hirsuta Lag. var. hirsuta: Ps P (8478), MO (DH-
12057), SM (8160), UN (2123); rip, s = pmg, pjw, mxs,
rds, pmg-pjw, rsl, rip-row; 1150-19
Bromus anomalus Rupr. ex E. iub E p ene HA (562);
rds, mog-rip, ppw-pjw; 1700-
! Bromus catharticus Vahl: 22; CO n ps (5887), MO (8856),
o (NS-9801), SM (2341); rds, ia mog-rip, mxs, pjw,
xs-lac, rip, oaw, pjw; 1100-2600
eos ciliatus L: 4; CO (7610), HA [E 9492); mog-lac, rip,
r 350
=
! Bromus inermis m var. inermis; 33; CO (5131), HA (4620),
MO id - SM (4538), UN (1742); rds, rip, rip-ppw, ppw, ari,
fog, p ac, pjw-row, lac, fog-ari, pmg; 1550-
e yis ca ex Murray: 41; CO (5362), HA (4993),
O (490), QU (NS-9830) SM (2342), UN (5224); lac, rds, ari,
ppw, a rip, rip-pmg, riw, oaw, rsl-pmg, rip-row;
1350-2
ion antes (Shear) Rydb.: 9; CO (6689), HA (4388), MO
74), SM (8888), UN (6929); rip, rds, rip-pmg, mxs, riw,
pmg; A
| Bromus tectorum L.: 42; CO (5316), HA (5415), MO (6422), QU
(NS-9709), SM (5961), UN (263); rip-ppw, oaw-rip, pmg,
ee mxs, rds, mog-rip, oaw, rip-pmg, pjw, mcw-rip, riw,
w-ppw; 1150-2400 m
T eni stricta (Timm) Koeler subsp. inexpansa e
Gray) C.W. Greene: 1; CO (9134); mog-rip; 2550-2600
Cenchrus longispinus (Hack.) Fernald: 14; HA (8285), MO o
SM (5975), UN (2018); pmg, mxs, rip, ari, pmg-rip, pmg-rsl,
rds, rip-row; 1250-1600 m.
Cenchrus spinifex Cav.: 1; QU (2714); rds; 1300-13
Chloris verticillata Nutt.: 22; HA (5044), QU (2712), a m
N (1200); pmg, rds, mxs, fog-ari, um rip, rsl, prng-rsl,
rds, rip, rip-row, rsl-riw; 1250-2
! ond d (L) Pers, var. a 2; SM (6097); mxs;
~
Docs Eee L.: 20; CO (5322), MO (4755), SM (7129),
e 5); oaw-rip, rip, rip-ppw, rds, ppw, vu rip, lac, ari,
mg, pmg, pjw, pjw-oaw; 1200-2450
Danthonia dnd Scribn.: 2; CO (7606); la ppw;
0-2350
ry D P (1 P Roa iv ev Roam & Schult: CO
ppw; 2250- 2300 m.
~
7461);
Cl |
scribnerianum
(6154); rip, rip-pmg;
(Nash) pu 3; HA (5446), SM
1700-2
f Texas 2(2)
Digitaria californica (Benth.) Henrard: 1; HA (9475); rsl-riw;
-1
Distichlis spicata (L) Greene var. stricta (Torr) ete 4: HA
(4391), MO (7854); rip-pmg, pal, pmg; 1350-
# | Echinochloa crus-pavonis (Kunth) J.A. eim var. Crus-
pavonis: 1; UN (257); pmg; 1450-1500 m. The specimen
should be re-examined given the infrequency of the
variety in the North America
! Echinochloa crus-pavonis (Ku th) s chult. var. macra (Wie-
gand) Gould: 8; QU (4503), hy 6965) UN a ari, rds,
rip-pmg, pmg, pmg-rsl, rip-row; 1150-15
Echinochloa BM (P. Beauv) Fernald var. qup
Wiegand: 10; HA (8379), SM (8167), UN (7665); rip, rip-pmg,
jw, pmg, fog-ari, mxs, oaw; 1300-2050 m
ee muricata (Beauv.) Fern. var. muricata: ii: UN (7405);
i; 1550-1600 m
T canadensis L.: 49; CO (7948), HA (6623), MO (7851),
SM (7131), UN (327); rip, mog-lac, ari, rds, ppw, mxs, rip,
pmg, ppw-pjw, pmg, pmg-rip, riw, oaw, rds, oaw, pmg-rsl,
rip-row; 1300-2500 m.
! Elymus d (Host) Runem. var. elongatus: 1; SM (8974);
000-2050 m. [=Leymus racemosus (Lam.) Tzev.]
ds oe (Raf.) Swezey var. brevifolius (J. G. Sm.)
n: 127; CO (5132); HA (4622), MO s QU (4502),
e ps UN (5219); rds, lac, ari, mog-lac, mog-rip, rip,
fog, pmg-lac, ppw, pmg-pjw, pmg, pmg-rip, pjw, mxs,
mxs-lac, oaw, fog-ari, riw, mcw-rip, row, rip-row; rsl-pmg;
pjw-ppw; 1150-2600 m.
| Elymus giganteus Vahl.: 1; CO (6758); rip; 2150-2200 m.
[=Leymus racemosus (Lam.) Tzvelev]
Elymus glaucus Buckley var. glaucus: 1; UN (8082); oaw;
Elymus lanceolatus (Scribn. & J. G. Sm.) Gould var. lanceolatus:
; UN (8086); oaw; 1550-1600 m.
Elymus smithii (Rydb.) Gould: 74; CO (6774), HA (6618), MO
(8858), SM (8981), UN (5526); rds, ari, mog-lac, mog-rip,
ppw, fog, pmg-lac, mxs, rip, pjw, pmg, rip-pmg, pal, fog-
ari, riw; 1300-2600 m.
Elymus ae (Link) Gould ex Shinners var. trachycaulus:
18; CO (8740), SM (7027), UN (389); rds, mog-lac, lac, rip,
ari, omg-rip, oaw, rip-pmg, pmg; 1400-2500 m.
! Eragrostis cilianensis (All.) Vignolo ex Janch.: 6; SM (8133), UN
(2067); -rip, pmg-rsl, rds, rip-row; 1300-1500 m.
Ae curtipedicellata m " » (5506), SM (6057), UN
7662); rds, mxs, rip-pmg; 1
| ud curvula (Schrad.) es var. confer Stapf: 1; UN
(5595); pmg-rip; 1400-14
eis pilosa (L.) P. ie var. ee 2; MO (8554), SM
75); rip, fog-ari; 1550-2050 m. o reported in
pecimens otherwise know
Eragrostis secundiflora J. Presl subsp. d Coros S. D. Koch:
9; HA (8336), UN (872); pmg, mxs; 1300-1600
Eragrostis sessilispica Buckley: 4; HA (5098), UN oe pmg;
0-1600 m. Specimens collected from sandy soil.
"e trichodes (Nutt.) A. W. Wood: 5; HA (1352); pjw-row,
mxs, rip; 1500-1 :
Erioneuron losa (Buckley) Nash: 39; CO (4630), HA (4337),
QU (4504), SM (6105), UN (5706); prng, fog, mxs, pmg-pjw,
r
Schiebout et al., Vascular plants of northeastern New Mexico
i E i eo riw, row, pmg-rsl, rsi-riw, pjw, ppw;
tes en arizonica Vasey: 1; CO (6547); ppw; 2300-2350
E idahoensis Elmer var. idahoensis: 1; CO (9133); mog-
rip; 2550-2600 m.
Festuca thurberi Vasey: 1; MO (6365); rds; 2250-2300 m.
Glyceria did (Nash ex Rydb.) M. E. Jones: 2; CO (9143), UN
mog-rip, caw; 1550-2600 m
Glyc sid udis. Watson: 1; MO (2862); rip; 2450-2500 m
Glyceria striata (Lam.) Hitchcock var. stricta (Scribn.) Fernald:
60); mcw-rip, oaw; 1550-1600 m.
Hesperostipa comata (Trin. & Rupr.) Barkworth var. comata: 22;
CO (7953), HA (4907), QU (3677), UN (1983); rip, pmg, pal,
mxs, ari, rds, pmg-rip, riw; 1250-1750 m. [SStipa comata
Trin. & Rupr.]
Hesperostipa m (Trin. & Rupr) Barkworth var. interme-
dia (Sc
rion. & Tweedy) Barkworth: 1; UN (4188), pmg,
1450- 1500 m.
(Thurl 7; CO (5508),
HA (4621), MO (6252), QU (NS- 9731), SM Rue UN (4015);
rds, omg, pmg-lac, pmg-pjw, opw, mxs, pjw, rip-pmg,
pjw-mxs, rsl-pmg; 1150-2200 m. [=Stipa Hu Euh.
(Thurb. ex Coult) Scribn.]
Hesperostipa spartea (Trin.) Barkworth: 1; CO (7589); mog-lac;
250-2300 m. [=Stipa spartea Trin.]
Hilaria jamesii (Torr) Benth.: 50; CO (5344), HA (5048), MO
(4575), QU (NS-9713), SM (3938), UN (1229); lac, rds, fog,
pmg, rip-pmg, pmg- d rip, mxs, pjw, riw, row, pmg-rsl,
rslpjw-mxs; 1150-18
Hilaria mutica uc) eth 6 a
pmg; 1250
Hordeum D e L:22; o mw HA (5442), MO (7857), SM
(707 gd UN (5694); pal, pmg- b lac, pg, rip, mog-lac,
, Tip, ari, rip-pmg, riw
TR Mus Nutt.: 53; CO n HA (6608), MO (4576),
QU (3652), SM (5966), UN (5527); rds, rip, mxs, pmg,
pmg-pjw, mxs-lac, oaw, rip-pmg, pjw, riw, rsl-pmg, ppw;
1150-2250m
! Hordeum UTIR L.: 6; CO (5501), SM o UN (4204); rds,
rip-ppw, lac, rip-pmg, pjw; 1450-
Koeleria macrantha (Ledeb.) Schult.: 12; CO (7958) SM (8879),
; rip, ppw, mog-lac, rds, mog-rip, mxs, rip-pmg,
oaw; 1550-2600 m
J hloa fi (L) Kunth subsp. fascicularis (Lam.) N. Snow: 3;
SM (8166), UN (7401); mxs, ari, rip-pmg; 1450-1600 m.
Lycurus phleoides Kunth: 8; HA (7790), SM ird UN (1036);
bjw, ppw-pjw, mxs, pmg, rip-row; 1350-1
A setosus (Nutt) C. Reeder: 10; HA (8650), p (8552),
); pmg, ppw-pjw, rip, rip-pmg, rsl, pmg-pjw,
e 1350-1850 m.
Melica porteri Scribn.: 1; CO (7600); rue 2250-2300 m.
Munroa squarrosa (Nutt) Torr: 1; CO (DH-1 1765); 1850-1900 m.
E. arenacea (Buckley) Hitchc.: 1; QU (2717 B); rds;
6054), UN (348); mxs, rds,
<=
Mosa DM Buckley: 12; HA (7206), QU (2717 A),
(2881); omg, rip, rds, rsl, row, pmg-rsl; 1300-1750 m.
1439
Muhlent x Trin.) Parodi: 3; CO
(9020), UN (2125); La rip-pmg, B 1300-1850 m
* Muhlenbergia aff. asperifolia (Nees & Meyen ex Trin.) Parodi:
1; CO (9021); lac; 1800-1850 m.
Ee (Nutt) Hitchc.: 3; CO (9135), HA (1351);
p, rds, pjw-row; 1750-2600
Mutientergiapungens Tut 1; ic rds; 1700-1750
a
n repens (J. Presl) Hitchc.: 1; SM (NS-9896); pjw;
icis ds torreyi (Kunth) Hitchc. ex Bush: 11; HA (750), UN
); pmg, pmg-pjw, rip, omg, rds, fog; 1300-1900 m.
ey wrightii Vasey ex J. M. Coult.: 2; CO (1850), UN
(1502); rds, rip-pmg; 1700-2350 m.
Nassella viridula (Trin) Barkworth: 3; MO (2278), UN (1739);
; ; m. [=Stipa viridula Trin.]
Panicum bulbosum Kunth: 1; SM (8142); pmg-rip; 1400-1450
m
Panicum capillare L.: 7; CO (9017), HA (DH-12134), MO
(2271), ui | UN (2765); lac, rip, rds, fog-ari, pmg;
Panicum n a var. hallii: 10; HA (6619), UN rip,
rip-pmg, pmg-rsl, rds, row, pmg, rip-row; 1300- :
Panicum obtusum Kunth: 36; HA (7279), MO E QU
(2716), SM (6108), UN (8704); rip-pmg, pjw, rip, pmg,
mxs, p rds, ari, fog-ari, riw, rsl, row, pmg-rsl, pmg-pjw;
1250-2050 m.
Panicum ene L.: 12; CO (7478), HA (8481), MO (7852), UN
(6921); ppw, rip, rip-pmg, pmg, rip; 1350-2300 m
did seen Michx.: 5; HA (8378), UN (1031); rip, pmg,
rip-row
! Phleum pee 9; CO (7588), MO (8855), UN (709); mog-
lac, ri w, rds, mog-rip, rds; 1550-2600 m
PE pru. (Trin. & Rupr.) Barkworth: 7; CO
(6747), UN (5153); rip-ppw, rip, mog-lac, ppw, mcw-rip,
ari; 1550-2350 m. [=Oryzopsis micrantha (Trin. & Rupr)
=
Poaarida Vasey: 2; CO (8722), SM (6166); rds, rip; 1950-2500 m.
Poa bigelovii Vasey & Scribn.: 5; HA (5395), MO (3157 B), SM
44); mxs, mxs-lac, rip, pjw, rip-pmg; 1550-1950 m
Poa fendleriana (Steud.) Vasey: 14; CO (5323), MO (4754),
SM (3748), UN (3620); oaw-rip, ari, mcw, mog-rip, lac,
rip, pjw, mcw-rip, rds, pmg-rip, pjw-oaw, pjw-ppw;
1500-2500 m
* Poa aff. fendleiana (Steud.) Vasey: 1; CO (6692); rip;
240
Poa interior s 2; CO (7417), UN (4257); ppw, rip-oaw;
1900-
Popa. CO (5325), HA (5443), MO (4780), SM (3880),
N (8 oaw-rip, mcw-rip, mog-lac, rip, rip-pmg, lac,
oaw; 1400-2450 m.
! Poa ie L: 58; CO (6752), HA (4992), MO (4753), SM
(4536), UN (4258); rip-ppw, rip, omg, rds, oaw-rip, lac, ari,
ppw, fog, pmg-lac, pal, mxs, rip-pmg, ppw-pjw, mxs-lac,
Ds mcw-rip, oaw; 1300-2450 m
a aff. pratensis L.: 1; UN (5161); mew- id 2100-2200 m.
fois trivialis L.: 1; SM (6182); rip; 1950-200
1440
Poa wheeleri Vasey: 2; CO (9034), UN (5156); lac, mcw-rip;
1800-1850 m. [=Poa nervosa (Hook.) var. wheeleri (Vasey)
C. L. Hitchc]
| Polypogon monspeliensis (L.) Desf: 16; CO (9032), HA (5385),
O (7858), QU (7009), UN (6802); lac, mxs, rip, pmg-rip,
g, rds, riw, ari; 1150-1850 m
Schedonnardus paniculatus (Nutt) Trel: 19; CO (7415), HA
d 787 : SM (6101), UN (1992); ppw, rip-pmg, pjw, pmg,
ari, oaw, pmg-rsl, rds; 1300-2300 m
E. arundinacea (Schreb) Dumort.: 10; CO (4631),
MO (6224), SM (4806), UN (3985); oaw, pmg, rds, rip-ppw,
ppw, pjw, rip-pmg, rip-oaw, pjw-oaw; 1650-2350 m.
[=Festuca arundinacea Schreb]
Cir Die oparium (Michx.) Nash var. scoparium: 18; CO
(7479), HA (6620), MO (8558), SM (8880), UN (1997); ppw,
mog-rip, rip, pmg, mxs, rip-pmg, pmg-pjw, rds, rip-row;
1300-2300 m
pcd (Scribn. & Merr.) K. Schum: 13; HA (6599), QU
M (8138), UN (2558); ME mxs, rds, mxs, pmg-rip,
mg, e rsi-riw; 1150-1
! mi: viridis (L) P. Beauv.: 12; o (9023), QU (2723), SM
(8971), UN (2878); lac, ari, ppw, rds, fog-ari, oaw, rip-pmg,
pmg; 1300-2050 m.
Sorghastrum nutans (L) Nash: 7; MO (DH-12083), a (8882),
4); mxs, rip-pmg, pmg, rip; 1300-1900
! Sorghum halepense (L.) Pers.: 7; HA (8475), QU 004 SM
(7030); UN (3031); rip, pmg, rds, pmg-rip; 1150-1500 m.
m obtusata (Michx.) Scribn. var. major (Torr.)
K : 3; HA (4987), UN (5676); rip, riw, pmg-rip;
Du 450 m. Further study may reveal these specimens
as S. intermedia (Rydb.) Rybd
v obtusata (Michx) Scribn. var. obtusata: 5; CO
n 63), HA (6622), MO (8557), UN (5767); ari, rip, riw;
—2000 m.
us airoides (Torr) Torr: 19; CO (6773), HA (4994), MO
(2274), SM (8145); UN (294); rds, ppw, ari, rip, pmg, pal,
rip-pmg, pjw; 1300-2300 m.
Sporobolus cryptandrus (Torr) A. Gray: 40; CO (1849), HA (8377),
O (2273), QU (2729), UN (1080); rds, rip, pmg, mxs, m
row, pmg-rsl, pmg-rip, pmg-pjw, rip-row; 1300-2350
* Sporobolus aff. cryptandrus ~ A. Gray: 2; HA (8480), p
(2465 B); rip; 1350-1
Sporobolus giganteus Nash: 3; on (DH-12066), UN (918); prng-
rip, ping, een
Trid & Standl.: 4; HA (7200 B), SM
(3829); pmg, bno: rip; 1350-1450m
Trisetum montanum Vasey var. montanum: 1; SM (BR-4124);
2600-2700 m.
| Triticum aestivum L.: 3; CO (4632), SM (7029), UN (4187); pmg,
e — À
Vulpia octoflora (Walter) Rydb.: 45; HA (4989), MO (4542 A), QU
(3649), SM (3743), UN (5539); rip, rip-pmg, pmg, pmg-lac,
pmg-pjw, pal, mxs, ari, rds, pjw, bdl; 1100-1950 m.
Polemoniaceae
Collomia linearis Nutt.: 1; CO (7565); mog-lac; 2250-2300 m.
Ipomopsis aggregata (Pursh) V. E. Grant subsp. candida (Rydb,)
f Texas 2(2)
V. E. E D. Grant: 4; CO (8767), UN (7963); rds, oaw;
1550-2500
Ipomopsis edm (Pursh) V. E. Grant subsp. formosissima
(Greene) a 6; CO (7458), MO (8802), UN (7976); ppw,
rip, mog-rip, mog-lac, oaw; 1550-2500 m
E laxiflora (J. M. Coult) V. E. Grant: 17; HA (4956), QU
(NS s SM (6089), UN (5783); omg, rip, rip-pmg, mxs,
d mg-pjw, bdl; 1150-1750 m
TRAE longis qa : A all subsp. longiflora: 2; SM
(8874); mxs, rds; 1850
& Ipomopsis cl Wilken E iss: Not isis
ut! Tables
10,11)
Ipomopsis spicata (Nutt.) V. E. Grant subsp. spicata: 1; HA (DH-
11749); 1850-1900 m
Phlox nana Nutt: 12; SM (6125); lac, rip, ari, rds, rip-pmg, pjw,
pjw-oaw, pjw-ppw; 1800-2250 m.
Polemonium foliosissimum A. Gray: 1; CO (7502); mog-lac;
2250-2300 m
Polygalaceae
Polygala alba Nutt. var. alba: 27; HA (4307), QU (4471), SM
(6004), UN (6844); ji d pmg-pjw, ari, rds, pjw, omg-
rip, pmg-rsl; 1150-1750
Polygonaceae
Ed alatum Torr. var. alatum: 14; CO (6705), HA (7743),
O (6208), UN (8048); rip-ppw, ppw, rds, rip, fog, pjw, ari,
rip-pmg, caw, Png: pas 2500 m
Eri 3:HA (8265), SM (8199), UN (7627);
rip-pmg; rds; rip-row; 1300-1850 m
& o aliquantum Reveal: Not collected but known
study area in Rid county (Tables 10
Eriogonum jamesii Ben ar. jamesii: 26; CO (7 (7 455), HA
7239); MO [A m (8912), UN (1565); ppw, rip, rds,
mog-rip, a mxs, ari, pomg-rip, fog, pmg-pjw, rsl, rip-
row; 1350-2600 m.
? Eriogonum jamesii Benth. var. aff. jamesii 1; UN (7673); rip-
pmg; 1450-
Eriogonum lachnogynum Torr. ex Benth.: 9; HA (8616), QU
(2691), UN (1561); pmg, rds, pmg-pjw, rsl; 1350-1850 m.
Eriogonum lonchophyllum Torr. & A. Gray var. fendlerianum
Benth.) Reveal: 1; UN (7966); oaw; 1550-1600 m.
Eriogonum tenellum Torr. var. tenellum: 11; HA (7741), QU
(4480), SM (8916), UN (5722); pjw, ari, pmg, mxs, riw, pmg-
ip, pmg-pjw, rip-row, pjw-pmg; 1150-1900 m
? Eriogonum tenellum Torr. var. aff. tenellum: 1; HA (441); ppw-
pjw; 1700-1750 m.
lopia o ) Á. Löve: 3; MO (8801), UN (1736); rip,
rds; 19 0 m.
Persicaria jud L. var. emersua (Michx.) Hickman: 1; HA
(786); pal; 1850-1900 m
Persicaria amphibia L. var. stipiilacea (Coleman) Hara: 1; UN
—
I
| Fa
(7382); rip-pmg.
Persicaria e (Raf) Nieuwland: 1; HA (DH-12208); ari;
1700
| E patho (L) A. Gray: 7; CO (8996), HA (7728),
MO (8844), SM (8100), UN (7396); lac, pjw, rip, pma-rip,
ari, oaw; 1400- 2500m
Schiebout et al ,V | lant
! Persicaria maculosa A. Gray: 1; MO (8600 BJ; rip; 1550-1600
ia arenastrum Jord. ex Boreau: 4; HA (4976), UN
(7301); rip, pmg, rip-pmg; ioe 1550 m
! Polygonum aviculare L.: 1; SM (8117); pmg- rip; 1400- 1450 m.
Polygonum erectum L.: 3; HA (7247), UN (2813); rip-pmg, omg;
1350-1450 m.
Rumex altissimus A. W. Wood: 13; CO (7878), HA (5434),
M (3786), UN (5671); rip, mxs, rip-pmg, riw, pmg, ari;
1300-1600 m.
! Rumex crispus L.:12; CO (6651), HA (4986), MO (8602), SM
MM UN (8700); rip, rip-ppw, ppw-pjw, pmg, rds, ari,
w, oaw; 1300-2450 m
Rumex hymenosepalus Torr.: 5; HÀ (4965), QU (9781), SM
(3849), UN (5769); rip, rip-pmg, pmg, bdl; 1150-1350 m.
Rumex maritimus L. var. fueginus (Phil.) Dusén: 1; UN (7377);
ari; 1550-1600 m
Rumex salicifolius Weinm. var. triangulivalvis (Danser) J. C.
Hickman: 5; CO (7154), MO (8603), SM (7105), UN (8046);
ari, rip; caw; 1550-2 :
| Rumex stenophyllus Ledeb.: 6; CO (9036), HA a SM (8924),
UN (6805); lac, rip, fog-ari, ari; 1300-205
* Rumex aff. verticillatus L.: 1; UN (7374); ari; ea 1600 m.
Pontederidaceae
Heteranthera limosa (Sw.) Willd.:
1500-1600 m
=
1; HA (DH-12205); lac;
Portulacaceae
! Portulaca oleracea L.: 1; UN (2189), pmg; 1400-1500 m.
Portulaca pilosa L.: 1; MO (DH 12181), rip; 1500-1600 m
Phemeranthus parviflorus (Nutt.) Kiger: 7; HA (448), MO (DH-
12209), boi (2070); ppw-pjw, mxs, rip, rip-pmg, pmg;
13 750 m. Flora North America (4:493) adopts a
broader oct Sabe. which we d here, but
o now are often treated as P
confertiflorus (Greene HRA
Potamogetonace
Eoo loss Raf. var. foliosus: 1; HA (DH-12054);
1500-16
EU. m Poir: 1; UN (7389), ari; 1550-1600 m.
Primulaceae
Androsace occidentalis jud 3; CO (5244), MO (3358); mcw-
rip, mew, rip; 2250-250
Androsace septentrionalis L.: P 1 (5299), SM (6130); rip, oaw-
rip, rip-ppw; 1800-2450 m.
Ranunculaceae
Aconitum columbianum Nutt. var. columbianum: 1; CO (9052);
Anemone canadensis L: 1; MO (6396); rip-ppw; 2300-2350 m.
Anemone o A. id 3; CO (6737); rip-ppw, mog-lac,
pw;
Anemone patens i var. SUM da Pritz.: 4; CO d MO (4742);
mcw-rip, mog-rip, lac, omg;
Aquilegia coerulea E. James var. coerulea: " CO (7487); mog-
lac; 2250-2300 m.
Clematis columbiana (Nutt) Torr. & A. Gray var. columbiana:
1441
4: MO a UN (5137); mog-rip, lac, rip, mcw-rip;
235 =
Clematis ssa i var. hirsutissima: 2; CO (9300); rds,
Clematis d Na 3; CO (7580), SM (7108); mog-lac,
rip, ari;
Clematis scottii de à CO (5306); oaw-rip, mcw-rip, mog-
lac; 2250-2400 m.
! Consolida ajacis (L) Schur: 1; CO (7141); ari; 1950-2000 m.
Rip nuttallianum Pritz.: 1; CO (9076); mog-rip;
-2300 m
cena ramosum e 3; CO (7539), MO (8846); mog-lac,
rip; 2250-2500 m
Q. Molnhini } Nec I i}
known from
study area in Colfax county (Tables 10, 11).
& Delphinium sapellonis Tidestrom: Not collected but known
from study area in Mora county (Tables 10, 11).
Delphinium virescens Nutt.: 18; HA (5412), QU (NS-9774),
SM (4881), UN (5656); pmg, mxs, rip, rds, rip-pmg, riw;
150-2250
Mo 7 virescens Nutt.: 1; UN (3576); rip- E 1400-
m imen collected on rocky subst
me minimus L.: 1; CO (4269); lac; 1800- pe m. mes
men collected from alkaline soil.
Ranunculus cymbalaria Pursh: 4; HA Ss di (4741), UN
(5192); rip, mog-rip, mcw-rip; 1300-2
Ranunculus gmelinii DC.: 1; UN (5190); mcw-rip; ; 2100- 2200
m. Collected from a basalt substrate.
Ranunculus inamoenus Greene var. inamoenus: 6; CO (5313);
UN ecc oaw-rip, mcw, mcw-rip, rip-pmg, Caw;
1400-245
Ranunculus d METER 4; CO (7438 A); MO (6418); ppw,
mog-rip, rip-ppw; 2250-2400 m
Ranunculus ranunculinus (Nutt.) Rydb: 1; MO (4732); mog-rip;
2350-2400 m.
Ranunculus sceleratus L. var. multifidus Nutt.: 9; SM (6117), UN
(5718); ari, rip, rip-pmg, riw; 1300-2000 m
| priis ME Crantz: 1; CO (4665); rip-ppw;
00- :
DR ae Engelm. ex A. Gray: 10; CO (5308), MO
(4748); oaw-rip, rip-ppw, rip, mcw-rip, mog-lac, ppw,
mog-rip; 22 .
* Thalictrum aff. fendleri Engelm. ex A. Gray: 1; CO (4693); rip-
ppw; 1800-1850 m
Rhamnace
Liber ue Gray: 1; MO (6380); rip-ppw, 2300-2350 m.
Rosacea
Agonist Michx.: 5; CO (6744), MO c UN (8013);
rip-ppw, mog- lac, p.e oaw; 1550-2500
Amelanchier an olia xM de var. alnifolia: 1;
O (BR-8275); 2800- 2900 m
Cocca montanus Raf. var. montanus: 21; CO (6640), HA
(6556), MO (i05. SM (NS-9591), UN (3996); rip, rip-ppw,
rds, ppw, pjw-row, mog-rip, rip-row, oaw, rsl, mxs, pjw-
mxs; 1350-2450 m.
n Meque 1; CO (NS-9283); caw; 1950-2150
1442
Fallugia paradoxa (D. Don) Endl. ex Torr: 11; CO (6701), HA
(5393), MO (7806), SM e x ~ rip-ppw, rip, mxs,
in-row, rds, fog; 1
Fragaria Vs 1; CO Du x p 245 0m
Fragaria virginiana Mill: 3; CO (5259), UN (4219); mcw-rip,
rip- caw oaw; 1900-2300 m
Geum Jacq.: 2; CO (7886
Geum macrophyllum Willd. var. perincisum (Rydb.
(6745) rip-ppw; 2300-2350 m
XB. A. Heller var. dumosus: 2;
CO (6642); rip, ius 2250-2450 m
Physocarpus monogynus (Torr) J. M. Coult.: 3; CO (6680), MO
(6379), UN (5139); 2100-2450 m.
Potentilla anserina L.: 1; MO (4740); mog-rip; 2350-2400 m.
Potentilla arguta Pursh var. convallaria (Rydb.) T. F. Wolf: 1; CO
61); mog-lac; 2250-2300 m
iis concinna Richardson
54); mog, rip; 2450-2650
PEIUS o L.: 1; CO oe rib; 2400-2450 m.
Potentilla gracilis Douglas ex Hook. var. pulcherrima (Lehm.)
Fernald: 4; CO (7485); rip, mog-lac, pmg, oaw; 1950-2450
); pow, rip; 1550-2300 m.
Raup: 1; CO
io
: 2; CO (3350), MO
m.
Potentilla hippiana Lehm. var. hippiana: 6; CO (7448), MO
(8828); ppw, rip, mog-lac, mog-rip, rds; 1550-2600 m.
subsp. monspeliensis (L.) Asch. & Graebn.:
5; CO (7150), MO (8562), SM (7049), UN (7373 B); ari, rip,
rip-ppw; 1550-2350 m.
Potentilla paradoxa Nutt.: 1; SM (7048); ari; 2000-2050 m.
Potentilla pensylvanica L. var. ee 3; CO (7883), MO
(6353), UN (5181); 1550-2
Potentilla Bp Nutt. var. eae (Engelm. ex Lehm.) S.
Wat 9; HA (5413), MO (6379) SM (3816), UN (5778);
mxs, rip, rip-pmg, ari, pjw; 1300-1950 m. Specimens col-
lected from sandy soils.
Potentilla subviscosa Greene: 1; SM (3739); pjw; 1900-1950 m.
Prunus americana Marshall: 5; CO (1833), HA (8312), UN
(4238); rds, pmg, rip-oaw, n mew-rip; 1300-2350 m.
ien id a L. var. melanocarpa (A. Nelson) Sarg.: 14; CO
(529 A (443), MO os UN (4224); oaw, ppw, rip,
eu oaw-rip, mog-lac, rds, mcw, ppw-pjw, mog-rip,
mcw-rip; 1700-2450 m
Rosa arkansana Porter var. arkansana: 1; UN (4189); rds-riw;
1879 m.
Rosa woodsii Lindl. var. ultramontana (S. Watson) Jeps.: 16;
CO (6715), HA (5455), MO (8827), SM (3711), UN (1729);
rip-ppw, oaw-rip, rip, rds, mog-lac, ppw, mog-rip, pjw,
ri , 0aw; 1550-2500 m.
SR ! Rosa x harisonii Rivers: 1; CO (4651); pmg; 1750-1800 m.
According to Kartesz (pers. comm., 2008), the nearest
ud of this escaped ornamental are Uintah Co,
and Mahaska Co, lA,
ius a 2; SM (3733); pjw, rip-pmg; 1900-2250
+ Rubus aff. idaeus L.: 1; SM (BR-3227); 2500-2700 m.
Rubus idaeus L. var. strigosus (Michx) Maximowicz: 1;
(6683); nip; 2400-2450 m. [=A. i. var. aculeatissimus Ee
&Tiling
f Texas 2(2)
Rubus neomexicanus A. Gray: : os om UN (4208); pjw,
mcw-rip, rip-oaw; 1900-19
* Rubus aff. E eden A. pu 1; SM (3705 BJ; pjw;
1900-1950 m.
Rubus peru Nutt. var. parviflorus: 1; HA (DH-11715);
1750-1800 m
=
Rubiacea
compat. 1; CO (7880), SM (3810); rip, pmg; 1350-1600
T boredle L.: 10; CO (7881), SM (BR-6980), MO (6400), UN
(5171); rip-ppw, rip, mog-lac, pow, mcw-rip; 1550-2450
m.
li asperrimu Gray) Higgins &
S. L. Welsh: 6; CO Bp HA (5430); SM oe UN (4212);
aw-rip; rip-ppw, mog-lac, mxs, rip-pmg; 1450-2400 m.
ems triflorum Michx.: 1; SM (BR-4212); 2500-2600 m
Hedyotis nigricans (Lam.) Fosberg var. nigricans: 9; HA (8260),
QU (2677), UN (1952); pmg, mxs, rds, rip-pmg, ari;
Houstonia rubra Cav.: 1; SM (NS-9875); pjw; 1600-1650 m.
Ruscaceae
Maianthemum racemosum (L) Link var. amplexicaule (Nutt.)
Dorn: 1; CO (7575); mog-lac; 2250-2300 m
Maianthemum stellatum (L) Link: 5; CO (4665) MO (4734
rip-ppw, oaw-rip, mcw-rip, mog-rip; 1800-2600
Nolina texana S. Watson: 8; HA (6585), MO on 12148), SM
iden rip, mxs, pjw, D m rds; 1450-1650 m. Disagree-
rthis is distinct from N. greenei
aT
me
S. m ex Tet
utaceae
Ptelea trifoliata L.: 5; HA (8393), MO (509), QU (NS-9735); pmg,
rip, pjw-row, pjw-mxs; 1150-1800 m
Salicaceae
* Populus aff. x acuminata Rydb; 1; HA (DH-H2006-01); rip;
1675 m.
Populus angustifolia E. James: 1; SM (BR-8670); 2650-2700 m.
Populus deltoides W. Bartram ex Marshall var. occidentalis
Rydb.: 14; CO (5331), HA (NS-9440), QU (NS-9694), UN
(2442); rip, pmg, lac, oaw, ari, rip-pmg, rds; 1300-2000 m.
Collections of Hazlett s 2006-01 through 04) in Mills
Canyon, Harding Cou may represent hybridization
between P deltoides EE P angustifolia
Populus tremuloides Michx.: 3; CO (6645), UN (4256); rip, mcw-
rip, rip-oaw; 1900-2450 m.
Salix amygdaloides Andersson: 10; CO (5266), HA (DH-2006-
07), MO (4744), QU (NS-9689), SM (3822), UN (5614); mcw-
rip, mog-rip, rip-pmg, riw, rip-oaw, pmg; 1150-2300 m.
* Salix aff. amygdaloides Andersson: 2; UN (7399); ari;
1550-1600 m
& Salix arizonica Dorn: Not collected but known from study
area in Mora county (Tables 10, 11).
Salix bebbiana Sarg.: 1; SM (4156); 2600-2700 m.
Salix exigua Nutt. var. exigua: 31; CO (5265), HA (5369), MO
(8816), SM (4814), UN (5169); mcw-rip, rip-ppw, mog-lac,
mog-rip, rip, pmg-lac, ppw, mxs, ppw-pjw, pmg, ozw, ari,
rip-oaw, riw, rds, pmg-rip; 1350-2500 m.
Schiebout et al., V. J Inte nf anc FERME TENET
Salix exigua Nutt. var. pedicellata (Andersson) Cronquist: 3; HA
(6596), UN (6823); rip, riw, ari; 1350-1600 m
Salix lasiandra Benth. var. caudata (Nutt) Sudw.: 1; HA (NS-
9488); rip; 1600-1700 m. [SS. lucida Muhl. subsp. caudata
(Nutt.) E. Murray]
Santalaceae
Arceuthobium vaginatum (Willd.) J. Presl var. m
(Engelm.) Cronquist: 1; MO (4778); lac; 2400-2450
Comandra umbellata (L.) Nutt. var. pallida (A. DC.) M. E. pm
4; iu HA ta 9517), SM (3885), UN (3561); rip-ppw,
pjw 1400-1850 m.
Porn juniperinum da 3; SM (4818); mxs, rip;
Sapindaceae
Acer glabrum Torr. var. glabrum: 1; CO (6643); rip; 2400—2450
m.
? Acer aff. negundo L.: 1; MO (DH-12115); 1500-1600 m.
Acer negundo L. var. interius (Britton) Sarg.: 1; SM (NS-9848);
rip; 1800-1850 m
Acer negundo L. var. violaceum (Kirchn.) Jaeger: 1; HA (8392);
rip; 1450-1500 m
Sapindus saponaria L. var. drummondii (Hook. & Arn.) L. D.
Benson: 1; HA (DH-11725); 1500-1600 m
Saururaceae
Anemopsis californica Hook. € Arn.: 1; SM (6294); rip-pmg;
2200-2250 m
Saxifragaceae
Heuchera parvifolia Nutt. ex Torr. & A. Gray: 4; CO (4686), MO
(4763); UN (5150); 1800-2450 m
Saxifraga bronchialis L. var. austromontana (Wiegand) Piper
ex G. N. Jones: 1; CO (6637); rip; 2400-2450 m.
Scrophulariacea
ure pm Pursh: 1; CO (7491); mog-lac;
WE thapsus L.: 27; CO (6674), HA (7750), MO (8561),
SM (7110), UN (7406); rip, rds, ppw, mog-lac, pjw, pmg,
fog-ari, ari, pmg-rip, oaw; 1350-2500 m
Simaroubaceae
! Ailanthus altissima (Mill) Swingle: 1; HA (NS-9457); rsl-riw;
155 600m
Solana
catracho oe (Juss.)
eae HPLC convida: (Moric.) Britton: 22; HA (4309), QU
(2674) SM (3775); UN (5792); pmg, pmg-pjw, rds, mxs,
rip-pmg, pjw, pmg-rsl, pow; 1150-1550 m
Chamaesaracha coronopus (Dunal) A. Gray: 11; HA (5376),
UN (2057); mxs, pmg-pjw, rip-prng, pmg, rds, pmg-pjw,
rip; 1350-1850 m.
! Datura quercifolia Kunth: 2; HA (8329), UN (2658); pmg;
1300-1500 m
| Datura stramonium L: 1; UN (590); pmg; 1400-1450 m.
dr Aena Miers: 2; HA (NS-9526), MO (7833); pmg;
D'Arcy: 1; CO (9033); lac;
me
"EU OR Gray: 1; SM (4826); mxs; 1750-1800 m.
1443
e hederifolia A. Gray var. comata s n 4; HA
07), (7972); pmg, oaw, rip-pmg; 1
iv hederifolia A. Gray var. fendleri (A. e conus 3;
A (7734), UN (2429); rip, pjw, mxs; 1350-
id Pas a De Nees: 1; UN E. rip-pmg;
Es
ENS. is fee rs 7; HA (5076), UN (5572);
; ip;
Physalis virginiana Mill.: 7; HA qm MO (8519), UN (1732);
prar rds, a row; 2350-2000 m
r) Raf: 24; CO (4267, HA (4366), QU (9791),
UN (2211); i; n pi rip-pmg, pmg-lac, mxs, rds, fog,
pmg, ppw;
Solanum americanum m a UN (8051) oaw; 1550-1600 m.
A revie is suggests the identify of this o is S.
igru which if correct means it is introduc
Solanum ERE Cav.: 88; HA (4365), MO 25 QU
(2680), SM (2295), UN (5231); rip-pmg, pmg, mxs, pmg-
bjw, rip, id pjw, rds, ari, fog-ari, riw, pmg-rsl, rsl-riw;
1150-1900
uu rostratum Duna: 48; HA (8620), QU (2694), SM (8923),
N (7634); pmg, pjw, mxs, rds, fog-ari, riw, pmg-rip, pmg-
i pmg- pjw, fog, —— eee 2050 m.
Tamaricaceae
!! Tamarix ramosissima Ledeb.: 32; CO (5357), HA (5423), MO
(7794), QU (4505), SM (6028), UN (5775); lac, ari, ppw, rip,
mxs, rip-pmg, pmg, ari, rds, riw, rip-row; 1150-2050 m
Theophrasta
de E Kunth: 1; HA (8293 B); pmg; 1300-1350
Typhaceae
Typha angustifolia L.: 10; CO (5363), HA (8383), MO (8587), UN
(1912); lac, rip-pmg, rip, ppw-pjw, oaw, rds; 1300-1850
m.
x os aff. UE L: 2; HA (481), SM (8110); ppw-pjw,
mg-rip; 1 750m
d latifolia Hi M MO (8848), SM (7037); rip, fog-ari, ari;
2000-2500
Ulmaceae
!! Ulmus pumila L.: 5; MO (7834), SM (7103), UN (3949); omg,
ari, rip; 1250-2050 m.
Urticaceae
Urtica dioica L. var. procera (Muhl. ex Willd.) Wedd.: 7; CO (5255),
MO (8821), UN ~ mcw-rip, mog-lac, rip, rip-ppw, oaw,
ari; 1550-2500
Verben
Hp ‘pinnate (Nutt.) Nutt. var. bipinnatifida: 91;
(4273), HA (5910), MO (2232), QU (2668), SM (4516),
UN (5218); lac, fog, pmg, pmg-lac, mxs, rds, pmg-pjw,
rip-pmg, pjw, mxs-lac, rip, ppw, ari, fog-ari, oaw, rsl, row,
-oaw; 1150-2200 m
Glandularia bipinnatifida (Nutt.) Nutt. var. ciliata (Benth.) B. L.
Turner: 3; SM (7053), UN (4089); ari, rds, pjw; 1600-2050
—
m.
Phyla cuneifolia (Torr) Greene: 18; CO (4653), HA (4362), SM
1444
(7111), UN (5660); gs ari, rip-pmg, mxs, rip, pjw, riw,
pmg-pjw; 1350-
! Phyla nodiflora (L.) Greene: aN (5974); ari, mxs; 1250-1300
m.
Verbena bracteata Lag. & Rodr.:19; CO (9016), HA (4581), MO
Dat *54l D h lec
f Texas 2(2)
Journal of tl
Viola nuttallii Pursh: 5; CO (5297), UN (4216); oaw-rip, mcw-rip,
A 0m
Viola pedatifida G. Don: 3; CO (5288); oaw-rip, mcw-rip, mcw;
2250-2450m
Vitaceae
(8520), UN (7379); lac, rds, pmg-lac, pjw, rip, omg, rip, ari,
pmg-rip, rip-row; 1300-2500 m
Parthenocissus vitacea (Knerr) Hitchc.: 13; CO (4691), HA (7242),
j MO
Verbena macdougalii A. Heller: 7; CO (7915), MO (8837), SM
(8533), SM (6144), UN (5167); rip-ppw, ppw, rip-pmg,
pw-pjw, rip, pmg, oaw, pjw, mcw-rip; 1350-2250 m.
Vitis s Raf: 5; HA (528), SM (3705 A), UN (5551); pjw,
(8958), UN (1781); rip, mog-lac, mog-rip, fog-ari, ari, rds;
a MENOR E - s; 1400-2250 m
Verbena plicata Greene: 2; HA (7724), SM (5918); pjw, rds; « rm aff. bs Raf: 1; HA (8473); rip; 1450-1500 m.
1400-1800 m
Viol Zygophyllaceae
es l . Kallstroemia parviflora Norton: 1; HA (DH-12149); 1500-1600
Hybanthus verticillatus (Ortega) Baill: 1; HA (6563); rip; m
30-1600 m. Krameria lanceolata Torr.: 13; HA (5034), QU (4470), SM (6016),
Viola canadensis L.: 4; CO (5245); mcw-rip, rip-ppw, oaw-rip,
oaw; 1800-2400 m.
; pmg, mxs, ari, pjw, pmg-rip; 1150-1550 m.
! Tribulus terrestris L.: 6; HA (8330), SM (8114), UN (1027); pmg,
rip; 1300-1550 m.
ACKNOWLEDGMENTS
This paper represents part of a M.S. thesis submitted by the first author to the University of Northern Colo-
rado. The first author would like to thank the numerous private landowners, Kiowa National Grasslands,
the U.S. Fish and Wildlife Service, and the NM state parks, for permission to collect from their properties.
Our sincere appreciation is extended to those who made a special effort to assist in this project, including:
Blair Clavel, Dan Garcia, Patty Hoban, Charles Jordan, Keith Long, Roy Mitchell, Jim O'Bryant, Bruce Rob-
inson, Joe Rodriguez, and Nancy Wells. Pam Smith (GREE) and Anton Reznicek (MICH) helped identify
some specimens of Cyperaceae; Ron Hartman (RM) reviewed many specimens of Caryophyllaceae and
Apiaceae. John C. Moore served as a thesis committee member for the first author. Ben Legler (RM) kindly
transported duplicate specimens from GREE to Albuquerque and Laramie and helped import the data into
a Specify database. Most of the original specimens at GREE were mounted by B. Williams, M. J. Niemann,
and M. Orris. Appreciation is extended to curators at the universities of New Mexico, Colorado-Boulder,
and Wyoming, as well as the Denver Botanical Gardens, whose collections were critical for confirming many
collections. The third author thanks John Kartesz for sharing several draft versions of the Floristic Synthesis
of North America to help ascertain state and county records. We apologize for those whose names that may
have been unintentionally omitted. The first author would also like to express gratitude for funding of this
project from The New Mexico Native Plant Society, The Garden Club of America, University of Northern
Colorado through the Winchester Fellowship, Flood and Peterson Insurance Grant, and the Graduate
Student Association. Support for publishing costs is gratefully acknowledged from the U.S. Department of
Education GAANN fellowship (grant + P200A060315-08) program to the School of Biological Sciences at
UNC and from the Rocky Mountain Herbarium, University of Wyoming. The manuscript benefitted from
the suggestions of S.L. O'Kane, Jr., K. Heil, B. Lipscomb, and the extensive and extraordinarily useful com-
ments of an anonymous reviewer.
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IR h Institute of Texas 2(2)
BOOK NOTICE
Davin S. INGRAM, DAPHNE VINCE-PRUE, AND PETER J. GREGORY (eds). 2008. Science and the Garden, the Scien-
tific Basis of Horticultural Practice, 2^4 Edition. (ISBN 978-1405160636, pbk.). Wiley-Blackwell
Publishing, 10475 Crosspoint Boulevard, Indianapolis, Indiana 46256, U.S.A. (Orders: www.wiley.com,
877-762-2974). $49.99, 368 pp., 135 illustrations, 7 1/2" x 9 3/4".
Contents:
Diversity in the Plant World
Know Your Plant: Structure and Function
Reproduction: Securing the Future
Naming Plants
Selecting and Breeding Plants
Soils and Roots
Soil Cultivation and Fertility
The Plant's Environment: Light and Water
Raising Plants from See
Propagating Plants Vegetatively
hape and Size
Colour, Scent and Sound in the Garden
Climate, Weather and Seasonal Effects
Gardening in the Greenhouse
The Diversity of Undesirables
Controlling the Undesirables
Maturation, Ripening and Storage
Conservation and Sustainable Gardening
Gardens and the Natural World
Gardens for Science
J 4 sel 2 11 1 1 f. J
WILIL Jal y plant science
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or r horticultural science course. Ones reason is that ae authors do a very Lue Job. of OVE in aio terms the ar or
Journal of Environmental Quality
] conservation has greatly enhanced this new edition: it
A + : J D f 1 hind:
“A 1 1 A =
is a ‘must ad for all interested in horticulture did gardening.”—John Macleod: RHS Professor of Horticulture
h 1 1 1
From the publisher: “M ional gardening |
seed Sowing and ae cuttings. This ‘Boole is DE in explaining in o terms some of the science that anderes these
are plants green? ae shoud one cut pu a d iei when dame cuttings? Why
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practices I pally a book of "Why' - y
d h water? But it i a on to n with Hs "p g l I
TI id he basi 1f ioni f l , nomenclature
genetics and plane e sel — d oes factors ee Eu ae a propagation e production, pest
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and disease control, Į g
Now with full PE aud th d editi ides tl with completely revised and updated cl r 1
first edition, with new io ~ clearer focus on the topics, and four new d mne. ipo matters an is BE p
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, namely: Diversity in the Plant World; C
the xou World; and Gardens for Science."
J. Bot. Res. inst. Texas 2(2): 1448. 2008
CHECKLIST OF THE VASCULAR PLANTS OF
FAYETTE COUNTY, PENNSYLVANIA
Cynthia M. Morton Loree Speedy
Section of Botany Section of Botany
Carnegie Museum of Natural History Carnegie Museum of Natural History
4400 Forbes Avenue 4400 Forbes Avenue
Pittsburgh, Pennsylvania 15213, U.S.A. Pittsburgh, Pennsylvania 15213, U.S.A.
ABSTRACT
Fayette County has been botanized since the early 1800s; however, a checklist of the county's vascular flora has not previously been
compiled. A search of herbaria augmented by targeted fieldwork has resulted in a list of 1,249 taxa of native and naturalized plants
comprising 525 genera and 134 families. The six families with the largest ipia js i wn are Asteraceae, Poaceae, Cyperaceae,
Rosaceae, Pede and Lamiaceae. Carex, Viola, Rubus, Polygonum and Solid There are approximately 201
non hat have been introduced mainly from Europe and UM This dedii ee 10 species of Lycophytes, 2
species of Equisetophs tes, 42 species of Polypodiophytes, 10 species of Gy I , and 1,149 species of Angiosperms. Eighty-five
species have global or state ranking
RESUMEN
El condado de Fayette County ha sido herborizado desde el comienzo de E 1800s; sin m no n sido paco a
un listado de la flora vascular del MU Una búsqueda en | di
resultado una lista de 1,249 y ] den 525 géneros y y 134 familias, iss seis familias con
el mayor nú de especi nas steraceae, Poaceae, Cyperaceae, Beine Fabaceae y Lamiaceae. hes ess pin NS dai y
Solidago son los cinco EA géneros. Hay aproximadamente 201 vas que han
Europa y Eurasia. Este listado reconoce 10 especies de Lycophytos, 2 especies de Equisetophytos, 42 o de Polypodiophytos, 1 10
especies de Gimnospermas, y 1,149 especies de Angiospermas. Ochenta y p gog
Site Description
Fayette County, Pennsylvania, is located in the southwestern corner of the state (Fig. 1). It is bordered by
Westmoreland County to the north, Somerset County to the east, Washington and Greene Counties to the
west, and the states of West Virginia and Maryland to the south. The entire western boundary of the county
is the Monongahela River. The Youghiogheny River enters on the southwestern side of Pennsylvania on the
border between Fayette and Somerset counties and flows northwest through a gap in the Chestnut Ridge to
Connellsville and then to McKeesport.
Fayette County receives total annual precipitation of approximately 39 inches along the Monongahela
River to 42 inches at the higher elevations of Chestnut Ridge and Laurel Hill. The precipitation is usually
well distributed throughout the year with February usually the driest month and June the wettest. Average
seasonal snowfall is 31 inches annually with the ground being snow-covered about 30 days per year.
The average date of the last 32 degree temperature in the spring is usually April 28", and the first frost
of the season is October 14". The length of the growing season, from the last killing frost in spring to the
first killing frost in autumn, is about 169 days (Uniontown, Fayette County Climate, www.ems.psu.edu/
PA Climatologist/uniontown).
Sophie's Woods within Friendship Hill National Historic Site has been reported by Jeffrey D. Wagner
and Robert Coxe of the Western Pennsylvania Conservancy as an old growth red oak-mixed hardwood
forest. In their own words, "Forests of such maturity are unusual for the county and the region in general.
Dominants include red oak (Quercus rubra), white oak (Q. alba), scarlet oak (Q. coccinea), tulip tree (Lirioden-
dron tulipifera), and shagbark hickory (Carya ovata). Major shrubs include mountain laurel (Kalmia laifolia),
spicebush (Lindera benzion), and American witchazel (Hamamelis virginiana). Herbs in this forest are sparse
J. Bot. Res. Inst Texas 2(0: 1449 — 1474. 2008
1450 J l of the Botanical R h Institute of Texas 2(2)
Fayette County A,
Pennsylvania ri
dy
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| RN e i 10
Miles Kilometers
bal
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Fayette City} ye Twp
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mainly composed of Virginia creeper (Parthenocissus quinquefolia), round-leaved violet (Viola rotundifolia),
false solomon's seal (Smilacina racemosa), and sessile bellflower (Uvularia sessilifolia). Save for the conspicu-
ous absence of American chestnut (Castanea dentata), this forest is probably similar to those forests seen by
Albert Gallatin when he lived here and George Washington when he surveyed the area" (Wagner & Coxe
Ferncliff Peninsula is a unique habitat encompassing a 100-acre peninsula with many rare and un-
usual plants for Pennsylvania. The Youghiogheny River brings seeds from West Virginia and Maryland into
the gorge that is warmer than the surrounding area, allowing plants to grow in areas where they normally
would not occur. The area was designated as a National Natural Landmark by the U.S. Congress in 1973
and was named a State Park Natural Area in 1992. The peninsula is shaded by oaks, hickories, tulip trees,
and beech. Rare plants include: Barbara's buttons (Marshallia grandiflora), Carolina tassel-rue (Trautvetteria
caroliniensis), Slender blue iris (Iris prismatica), Autumn willow (Salix serissima), and buffalo-nut (Pyrularia
pubera). In addition, plant fossils such as Lepidodendron from 365-million-year-old club mosses are plentiful.
Morton and Speedy, Vascular plants of Fayette County, Pennsylvania 1451
Boulders from the size of basketballs to houses occur on the waterfront while white waters make it one of
the most scenic of all Eastern riverfronts (Palmer 1984).
The soils of the county consist of five main series: Gilpin-Wharton-Ernest association, Guernsey-
Westmoreland-Clarksburg association, Dekalb-Hazleton-Cookpart association, Upshur-Albrights association,
and Monongahela-Philo-Atkins association (Kopas 1991).
The Gilpin-Wharton-Ernest association is widely distributed. In the east it occupies the areas between
Chestnut Ridge and Laurel Hill, while in the west it extends from Point Marion to Perryopolis and from
Point Marion to Laurelville. Th il d 1 moderately to well-drained. They consist of level to very
steep soils underlain by acid shale and some sons bedrock on uplands. This soil association makes up
about 37 percent of the county. Portions of this association are strip-mined producing highly acidic soils,
however, this association also has some of the better farming soils of the county.
The Guernsey-Westmoreland-Clarksburg association consists of rounded hilltops and a series of benches
located along the slopes. These soils are deep and poorly to well-drained. They consist of moderately steep
soils influenced by limestone strata in the bedrock on uplands with medium to moderately fine texture.
This soil association makes up about 27 percent of the county. Areas of strip mine spoil, mine dumps, and
urban land occur. Dairy and general farming are dominant.
The Dekalb-Hazleton-Cookpart association is located on and along Chestnut Ridge and Laurel Hill
and along the Youghiogheny River. Most of the association is very stony. These soils are moderately deep
to deep and moderately to well-drained. They are coarse to medium in texture and level to very steep soils
underlain by bedrock that is dominantly acid sandstone on uplands. This soil association makes up about
25 percent of the county. Most of this association contains forestland.
The Upshur-Albrights association consists of soils formed in materials weathered from red shale, lime-
stone, and some sandstone. It occurs along the upper slopes of Chestnut Ridge and Laurel Hill. These soils
are deep and poorly to well-drained and occur on gently sloping to very steep uplands. This soil association
makes up about 7 percent of the county. Much of this association is forested with a few large stone quarries
present.
The Monongahela-Philo-Atkins association consists of soils formed in deep alluvial deposits that are
common along the Monongahela and Youghiogheny Rivers. These soils are deep and moderately to poorly
drained and are nearly level to sloping soils on stream terraces and flood plains. This soil association makes
up about 4 percent of the county. Parts of this association contain the county's community and industrial
development (Kopas 1991).
Fayette County is situated in the Laurel Highlands of southwestern Pennsylvania; the terrain consists
of rolling hills and mountains. The eastern part of Fayette County is mountainous with Chestnut Ridge
rising to heights of more than 2,900 feet. The west of Chestnut Ridge is a rolling and hilly plateau. Much of
Fayette County was once underlaid by coal but most of it has been mined. Elevations range from a high of
2,900 feet to a low of 290 feet above sea level.
All of Fayette County is in the Monongahela River watershed, which is part of the Ohio and Mississippi
River watersheds. Two of the principal tributaries of the Monongahela River, the Youghiogheny and Cheat
Rivers, flow through Fayette County.
The Monongahela subbasin has a total drainage area of 2737 square miles. It includes the Pennsylvania
portion of the Monongahela River, from West Virginia and Maryland to Pittsburgh. The Pennsylvania por-
tion of the subbasin encompasses all of Fayette County, and parts of Greene, Washington, Westmoreland,
and Somerset Counties.
The Monongahela subbasin in Fayette County is divided into the following major watersheds: Lower
Youghiogheny River, Middle Monongahela River, Upper Monongahela River and the Upper Youghiogheny
River watersheds.
The Lower Youghiogheny River watershed has a total drainage area of 478 square miles and its major
streams include Sewickley Creek, Jacobs Creek and the lower portion of the Youghiogheny River. The Middle
1452 J | of the Botanical R h Institute of Texas 2(2)
Monongahela River watershed has a total drainage area of 509 square miles and its major streams include
the middle portion of the Monongahela River, Pigeon Creek and Redstone Creek.
The Middle Monongahela Watershed shows many extremes in water quality due to both man-made
and natural causes. The greatest water problem is the pollution of the water resources by the drainage from
coal mining operations, and the hundreds of abandoned oil and gas wells that are not properly sealed.
The Upper Monongahela River has a total drainage area of 377 square miles and its major water ways
include Whitely Creek, Dunkard Creek, and the uppermost Pennsylvania portion of the Monongahela River.
The Upper Youghiogheny River has a total drainage area of 384 square miles and its major streams include
the Indian Creek, Laurel Hill Creek and the upper portion of the Youghiogheny River.
Watershed conditions need to be evaluated to detect if biodiversity is increasing or decreasing in these
areas. Such comparisons E be ien important in mining sites because they will reflect the interac-
tions of many ofthe other i llution and soil changes, which can be utilized to make better
management decisions (Pennsylvania DEP vaad Notebook 2006).
History
Early Indian traders, land agents, and explorers were the first Europeans to blaze the way into Fayette
county. In the mid-18" century, rivers were the only easily accessible means of transportation upon which
the French took routes down from Canada. In 1754, George Washington led an expedition into the region
which ended in defeat by the French and Indians at Fort Necessity. In 1758 after the British won the French
and Indian War, settlers came back into the region to initiate subsistence farming. The Braddock Road from
Cumberland northward from Fort Necessity down to Gist Plantation and on through Connellsville were the
main roads in Fayette. A connector was needed to the west and Col. Burd supplied that in 1759, with the
help of the Indian chief Nemacolin. The connector was from Gist's Plantation to Brownsville. These routes
provided the basic network that the National Road would later follow.
In the 1770s, the territory that would become Fayette County was claimed by both Pennsylvania and
Virginia. The claim was finally settled in 1780 when the Mason-and-Dixon Line was extended to the original
limit of Pennsylvania's land grant, giving Virginia frontage on the Ohio River.
Fayette County was created in September 1783 from the southern portion of Westmoreland County.
The county was named for Marquis de la Fayette, a young Frenchman who assisted Gen. George Washington
during several major battles of the American Revolution.
Fayette County's geographic location and the wealth of its natural llectively responsible
for producing two major historical periods. The first is the transportation via “road and river era" and the
second is the “coal and coke era.”
Industrialization came at the turn of the 19^ century with iron furnaces being built in the mountains
to harvest deposits of iron ore and to utilize the ling forest as a wood source for charcoal. The glass
industry was first established when Albert Gallatin opened his New Geneva glassworks in 1794. Within
the next 20 years, numerous glass factories were erected at various points along the river, which brought
national recognition to the upper Monongahela River valley. The National Road, built in 1815 opened up
the Northwest Territory, allowing travel into areas that were previously inaccessible by river.
Vast deposits of bituminous coal underlying much of Fayette County resulted in the development of
an internationally renowned coal and coke industry around the turn of the 20% century. Thousands of Eu-
ropean immigrants came to Fayette County to work in the mines. However, by the 1950s most of the large
Fayette County mines were finished and the county's economy took a major downward turn. Tourism and
industrialization are the current growth prospects.
The county is divided into 42 municipalities with the majority of the population residing in the urban
and suburban areas. The first census taken in 1790 documented that Fayette County had a population of
nearly 13,325. As of 2000, the population was 148,644. According to the U.S. Census Bureau, the county
has a total area of 2,067 km? (798 mi?), 2,046 km? (790 mi?) of it is land and 20 km? (8 mi?) is water (U.S.
Census Bureau 2007).
RA nyt, Ic J , WV. I I $ er tta f. ty, Pennsylvania 1453
Major Collectors
Collecting in Fayette County for the Carnegie Museum Herbarium began in the early 1800s and has continued
to the present. Most of the collectors have been members of the Botanical Society of Western Pennsylvania
and the staff of the Section of Botany at Carnegie Museum. Botanists who have made significant contribu-
tions to the knowledge of the Fayette County are J. Bright, W.E. Buker, R. Coxe, L.K. Henry, O.E. Jennings,
J.A. Shafer, and L. Speedy.
METHODS
This checklist was compiled by searching the herbarium of the Carnegie Museum of Natural History (CM),
Academy of Natural Sciences of Philadelphia (PH) and John Franklin Lewis Herbarium at the California
University of Pennsylvania (CUP). Herbarium collection information was obtained from the Morris Ar-
boretum (MOAR). Six specimens were examined and included from the Academy of Natural Sciences of
Philadelphia (PH) and six specimens from the John Franklin Lewis Herbarium at the California University
of Pennsylvania (CUP). Fieldwork was conducted targeting underrepresented areas of the county. Most of
the collections were made during the last two years, but some date from the late 1940s and early 1950s,
and a few are from 1873 and 1883. Rhoads and Block (2007) was the primary source for plant identifica-
tion. For generic and species names, we have generally followed Kartesz (1999). Authorities are abbreviated
according to Brummitt and Powell (1992).
RESULTS AND DISCUSSION
This list includes the names of all native and naturalized species known to occur in Fayette County. It
includes a total of 134 families, 525 genera, and 1213 species. The number of families, genera, species and
non-native are similar to the recently reported taxa for Crawford, Greene and Washington Counties (Table
I). The six families with the largest number of species are Asteraceae, Poaceae, Cyperaceae, Rosaceae, Fa-
baceae and Lamiaceae. Carex, Viola, Rubus, Polygonum, and Solidago are the largest genera. This checklist
recognizes 10 species of Lycophytes, 2 species of Equisetophytes, 42 species of Polypodiophytes, 10 species
of Gymnosperms, and 1,149 species of Angiosperms. There are approximately 201 non-native species that
have been introduced mainly from Europe and Eurasia. Eighty-five species have global or state ranking. Of
the 85 plants that have global or state ranking, four, including, Aconitum reclinatum, Delphinium exaltatum,
Scutellaria saxatilis, and Vitis rupestris, have a global ranking of G3 or vulnerable status. Three plants, Mar-
shallia grandiflora, Potamogeton tennesseensis, and Spiraea virginiana, have a global ranking of G2, and one,
Malaxis bayardii, is globally ranked as G1G2. The remaining 85 plants are either a G4 or G5 status, indicating
an apparently secure or secure condition globally.
Aconitum reclinatum belongs to the buttercup family. It is a perennial herb from fascicled roots, with
cauline leaves that are deeply divided and coarsely toothed. The flowers are white, grouped in an inflorescence
at the end of the stem and lack spurs. White monkshood flowers from June to September. There are only
two species of Aconitum in Pennsylvania. The second species, A. uncinatum has blue flowers, therefore white
monkhood is easy to identify when in flower. It grows in moist areas along streams, in coves, or in seeps
within mixed forests. It is threatened by logging activity and may be disturbed by drainage or disturbance
of wetland habitats.
Delphinium exaltatum has alternate leaves that are palmately divided into 3—5 segments, 15 cm long,
silvery-green abaxially and pubescent above and below. The basal lobes of some of the larger leaves are
divided again and make the blade appear 5-lobed. The flowers are in terminal racemes. The 4 purplish-blue
petals are dimorphic with the upper two expanded at the base and forming a spur. The lateral petals are
reflexed and have long white hairs. The stamens are approximately 30 in number with yellow pollen. The
5 sepals of this plant are irregular. This species is readily confused with another similar taxon, Delphinium
tricorne, however D. tricorne is a smaller plant, which flowers much earlier than D. exaltatum). By July, when
D. exaltatum is beginning to flower, D. tricorne has already set fruit. Most of the collections are from rich
1454 J | of the Botanical R h Institute of Texas 2(2)
Tagle 1. R hecklists produced since 2005, including total number of speices, families, genera; non-native and global rare species found.
Total non-native Families Genera Rare G1-3
Crawford' 1168 208 134 485 3
Fayette 1214 201 134 522 8
Greene ? 890 179 118 438 1
Washington? 982 203 125 472 1
'Morton, Speedy and Bissell (2007), *Coxe, et al. (2005), and ¿Morton and Speedy (2007).
shaded woods and rocky limestone bluffs. This species is quite conspicuous and may be subject to casual
picking or dug for gardens.
Scutellaria saxatilis is a member of the mint family. The genus Scutellara is very distinctive and easily
identified by the protuberance on the upper lip of the corolla. This species is a stoloniferous plant with up-
per cauline leaves that are broadly lanceolate to triangular, and cordate at the base. The leaves are coarsely
crenate, 4—5 cm long, with a long petiole. The flowers of S. saxatilis are in panicles. Most of the collections
are from semi-shaded areas in dry woods or, occasionally, in moist areas along streams. This species is not
common, and due to its inconspicuous nature it may be overlooked. Its recovery potential is unknown.
Vitis rupestris is a member of the grape family. It is a bushy shrub-like grape that is seldom climbing.
Tendrils are lacking or if present, reddish and only found on the uppermost nodes. The leaves are reniform,
coarsely toothed and pubescent below, usually along the veins. The pith is interrupted by a diaphragm at
each node. The V. rupestris flowers from early April to mid-June. Fruits are black berries. It grows in sandy
deposits on rocky river banks.
The Large-flowered Marshallia, Marshalli diflora, is in the family Asteraceae. The leaves are narrow
and tapering with three parallel veins. Near the base the leaves are petiolate, but near the apex the leaves
are sessile. The flowers are small tubular, white or light blue, and clustered in disc-shaped heads at the apex
of each main stem. Marshallia grows in wet forests or meadows and on stream banks throughout central
Appalachia and into the southern Appalachians from southern Pennsylvania to Eastern Tennessee. It is a
perennia! herb flowering from early June to late July.
Potamogeton tennesseensis is a submersed and floating-leaved aquatic with very slender, terete stems.
The floating leaves are about 2-4 cm long and 5-13 mm wide while the submersed leaves are 0.2-2 mm
wide with 1-3 major veins. The overall small size of P. tennesseensis, in combination with the acute floating
leaves, render the species reasonably distinctive from other Potamogeton species. The species flowers from
May through September and fruits from July to October. The species occurs in ponds and streams.
The Appalachian spiraea, Spiraea virginiana, is in the Rosaceae. It is a clonal shrub reaching a height
of 3 meters. The leaves are long, narrow, and taper to the base with nearly smooth edges. They are whitish
underneath. The flowers are small with 5 white petals. The flowers are grouped into large showy clusters,
up to 5 inches wide at the end of the stems. Spiraea virginiana flowers from late June to early July and fruits
from July into early September. It typically grows along gravel bars and banks of swift flowing streams and
scoured river banks that experience periodic flooding. The species is currently known from seven states
from Pennsylvania to Alabama. It has become so rare and individual populations are so isolated that it is
not known to set fertile seed due to lack of cross-pollination.
Malaxis bayardii, Bayard's adder's mouth, is one of the smallest of our native orchids. There are only
about a half dozen known populations of this plant worldwide. The orchid has been reported to occur in
dry, open woods, shale barrens and sandy pine barrens. The plant flowers from mid-July to August with
fruits persisting into September. It is about 4 to 8 inches tall with one oval green clasping leaf. At the top of
the stem is a spike of about 150 tiny yellow-green flowers that mature from the top down. The lip petal is
divided into two lobes. Malaxis unifolia is similar in appearance but is typically found in swamps, bogs and
AA T IC J Vi | | + £r. n fF. " A
| y, y Pennsylvania 1455
wet habitats. In addition, M. unifolia has a densely flowered inflorescence with flowers that mature from the
bottom up.
There are 5 taxa in the Fayette County flora that are listed by the Pennsylvania Department of Agri-
culture (2007) as noxious weeds: Cirsium arvense (Canadian Thistle), Cirsium vulgare (Bull or Spear Thistle),
Datura stramonium (Jimsonweed), Lythrum salicaria (Purple Loosestrife), and Rosa multiflora (Multiflora
Rose). Other species considered serious invasives in Pennsylvania's native ecosystems are: Acer platanoides
(Norway Maple), Acer pseudoplatanus (Sycamore Maple), Ailanthus altissima (Tree-of-Heaven), Alliaria peti-
olata (Garlic Mustard), Berberis thunbergii (Japanese Barberry), Celastrus orbiculatus (Oriental Bittersweet),
Chelidonium majus (Greater Celandine), Conium maculatum (Poison Hemlock), Coronilla varia (Crown Vetch),
Elaeagnus umbellata (Autumn Olive), Euonymus alatus (Winged Euonymus), Festuca elatior (Tall Fescue), Hes-
peris matronalis (Dame's Rocket), M QE E (Border Privet), Pension vu “gare (Common Privet),
QW ii (Morrow’s
Lonicera japonica Japanese Honey I kii (Amur H
Honeysuckle), Microstegium vimineum Upas Stilt Grass), Oniithozalum ünbellatum (Star-of-Bethlehem),
Pastinaca sativa (Wild Parsnip), Phalaris arundinacea (Reed Canary-grass), Polygonum cuspidatum (Japanese
Knotweed), Polygonum sachalinense (Japanese Knotweed), Potamogeton crispus (Curly Pondweed), and Spiraea
japonica (Japanese Spiraea) (DCNR 2004).
While this checklist probably does not include of every species in Fayette County, it is the most com-
prehensive list presently available.
ANNOTATED CHECKLIST OF THE SPECIES OF
FAYETTE COUNTY, PENNSYLVANIA
Taxa are listed according to the following format: taxon name, author(s), ((year) collector and number)
global: state ranking [Synonyms], non-native source and herbarium location if not CM. We follow the state
(S) and global (G) ranking systems developed by The Nature Conservancy (1996). The global numbers are
designated from 1 (critically imperiled) to 5 (secure). Other notations include SH, which denotes historical
occurrence, and SR, which indicates reported without persuasive documentation. Synonyms are included
for names not in common usage in the state or regional manuals. In cases where there was more than one
specimen present in the collection, recent collections of current collectors for the western Pennsylvania
region were cited.
Families, genera, and specific and infraspecific taxa are arranged alphabetically within vascular plant
groups Angiosperms, Gymnosperms, Lycophytes, Equisetophytes, and Polypodiophytes.
ANGIOSPERMS Alismataceae
Alisma subcordatum Raf., 2006 L. Speedy LS-06-846
Sagittaria australis (J.G.Sm.) Small, 2006 L. Speedy and M.
Bowers LS-06-805
Acanthaceae
Justicia americana (L.) Vahl, 2006 L. Soeedy LS-06-842
Aceraceae Sagittaria g Michx. var. graminea, 2006 L. Speedy and
Acer negundo L. var. negundo, 2006 L. Speedy LS-06-343 D. Sofran LS-06-933
Acer nigrum Michx. f, 2007 L. Speedy and J. Nusser 07-605 Sagittaria latifolia Willd., 1977 M. Polechko s.n.
Acer pensylvanicum L, 2006 L. Speedy LS-06-143 Sagittaria rigida Pursh, 1977 M. Polechko s.n.
Acer platanoides L., 2007 L. Speedy 07-1631 a
Acer pseudoplatanus L., 1935 R. Chinn s.n. Euro
Amaranthaceae
Acer saccharum Marsh. var. saccharum, 2006 ' “speedy LS- Amaranthus hypochondriacus L., 1886 J.A. Shafer s.n. Africa
06-144 Anacardiaceae
Acer saccharinum L., 2006 L. Speedy LS-06-106 Rhus copallinum L. var. latifolia Engl. 2007 L. Speedy 07-
Acer spicatum Lam., 1981 FH. Utech 81-038 1029
rubrum L. var. rubrum, 2007 L. Speedy LS-07-20-1 Rhus glabra L., 1953 LK. Henry and WE. Buker s.n.
Acer rubrum L. var. trilobum Torr. & Gray ex K. Koch, 1902 JA. — Rhus typhina L., 2006 L. Speedy LS-06-340
Toxicodendron radicans (L.) Kuntze ssp. radicans, 2007 L.
Åcoracède Speedy 07-1853
Toxicodendron vernix (L) Kuntze, 1922 E.H. McClelland s.n.
a
Acorus calamus L., 1977 M. Polechko s.n. India
1456
Annonaceae
Asimina triloba (L.) Dunal, 2006 L. Speedy LS-06-107
Apiaceae
Angelica atropurpurea L., 1948 W.E. Buker s.
Angelica a Michx., 2006 L. P and J. Nusser
LS06-661
Angelica venenosa (Greenway) Fern., 2007 L. Speedy 07-
1532
Cicuta maculata L., 2007 L. Speedy 07-1023
Chaerophylium procumbens (L.) Crantz var. procumbens, 2006
L. Speedy LS-06-104
Conium maculatum L., 1901 J.A. Shafer s.n. Europe
Cryptotaenia canadensis (L.) DC., 1999 R.B. mS s.n.
Daucus carota L., 1981 EH. Utech 81-214 Eur.
Erigenia bulbosa (Michx.) Nutt., 2006 L. oy LS-06-43
G5:54
Heracleum maximum Bartr,, 2006 L. Speedy L506-342-1
Hydrocotyle americana L., 2007 L. Speedy and M. Bowers
8
7-
Osmorhiza EE (Michx.) C.B. Clarke, 2006 L. Speedy
-06-17
Osmorhiza AE (Torr) DC., 2006 L. Speedy LS-06-172
E a ) Raf., 2006 L. Speedy and M. Bowers LS-
Psa sativa a 1981 EH. Utech 81-212 Eurasia
cula canadensis L. var. canadensis, 2007 L. Speedy and
: 8
Sanicula marilandica L., 2007 L. Speedy LS-07-257
Sanicula odorata (Raf) K.M. Pryer & L.R. Phillippe, 2006 L.
Speedy and D. Sofran LS-06-212
Sanicula trifoliata Bickn., 2007 L. Speedy 07-613
Sium suave Walt., 2007 L. Speedy 07-1031
Taenidia integerrima (L) Drude, 2007 L. Speedy and M. Bow-
ers 07-685
Thaspium barbinode (Michx.) Nutt., 2006 L. Speedy and D.
Sofran LS-06-211
Thaspium trifoliatum (L) Gray var. aureum Britt, 1959 W.E.
Zizia aptera (Gray) Fern., 1959 W.E. Buker s.n.
Zizia aurea (L) W.D.J. Koch, 2007 L. Speedy and M. Bowers
LS-07-239
Apocynaceae
Amsonia tabernaemontana Walt. var. tabernaemontana, 1941
J. Lewis s.n.
Apocynum androsaemifolium L., 1941 O.E. Jennings s.n.
Apocynum cannabinum L., 2006 L. Speedy LS-06-363
Vinca minor L., 2007 L. Speedy 07-1723 Europe
Aquifoliaceae
llex laevigata (Pursh) Gray, 1933 J. Bright 8642
montana Torr. & Gray ex Gray, 2006 L. Speedy LS-06-405
llex verticillata (L.) Gray, 2007 L. Speedy 07-1615
Nemopanthus mucronatus (L.) Loes., 1946 OF. Jennings s.n.
Araceae
Arisaema dracontium (L.) Schott, 1923 EH. McClelland s.n.
es triphyllum (L) Schott ssp. pusilum (Peck) Huttleston,
905 OE. Jennings s.n.
f Texas 2(2)
Arisaema Mein (L) Schott E A (Britt) Hut-
tleston, 2006 L. Speedy LS
Arisaema Nw o ssp. ME 1995 B.L. Isaac
and J.A. Isaac 7074
Orontium aquaticum L., 2006 L. Speedy LS-06-164 G5:S4
Peltandra virginica (L) Schott, 1991 S.A. Thompson 8912
Symplocarpus foetidus (L.) Salisb. ex Nutt., 2007 L. Speedy
LS-07-18
ralia
sali nudicaulis L, 2006 L. Speedy LS-06-200
ssp. racemosa, 2006 L. Speedy and J. Nusser
and M. Bowers LS- 06-59
T spinosa L., 2006 L. a LS-06-832
Panax quinquefolium L., 2006 L. Speedy and J. Nusser and M.
Bowers L 95
Panax trifolius L., 2006 L. Speedy LS-06-141
Aristolochia
Aristolochia E Lam., 1992 J.A. Isaac 3920
ae serpentaria L, 1905 J.A. Shaferand Kinzer, G.E.
mus canadense L., 2006 Speedy, L. LS-06-52
Asclepiadaceae
Asclepias exaltata L., 2007 L. Speedy and M. Bowers 07-692
Asclepias incarnataL. ssp. incarnata, 1981 F.H. Utech 81-126
Asclepias purpurascens L., 1999
Asclepias quadrifolia Jacq., 1942 O. E. Jennings s
Asclepias syriaca L., 2006 L. Speedy and Nusser, i LS- 06-581
Asclepias tuberosa L. ssp. tuberosa, 1981 EH. Utech 81-047
Asteraceae
Achillea millefolium L. var. millefolium, 2007 L. Speedy LS-07-
401 Europe
Ageratina altissima (L) King & H.E. pu var. altissima, 2006
. Speedy and D. Sofran LS-06-93
Ambrosia o L. var. elatior T Descourtils, 2007 L.
Speedy 07-
Ambrosia trifida i var. trifida, 2006 L. Speedy LS-06-849
Anaphalis margaritacea (L.) Benth., 2007 L. Speedy 07-1829
A ial Mii lioica ( )B 2006
IL
L. Speedy LS-06-110
Antennaria neglecta Greene, 2007 L. Speedy LS-07-364
Antennaria parlinii Fern. ssp. fallax (Greene) Bayer & Stebbins,
95 . Beer s.n.
Antennaria pa Fern. sp parlinii, 2007 L. Speedy LS-07-
187
Antennaria plantaginifolia (L) Richards., 1934 E.H. Graham
ae solitaria Rydb., 1999 S.P. Grund 2141 G5:S1
Anthemis arvensis L., 1924 J. Bright s.n. Europe
Anthemis cotula L., 1901 J.A. Shafer s.n. Europe
Arctium minus Bernh., 1991 F.H. Utech 91-1341 Eurasia
Arctium tomentosum P. Mill, 1984 S.A. Thompson and J.
Loughlin 1881 Eurasia
Arnoglossum atriplicifolium (L.) H.E. Robins., 2006 L. Speedy
LS-06-1011
Arnoglossum muehlenoergii ER H.E. Robins., 1996
S. Grund and J.L. Smith 1
Pennsvlvania
oF Ld
Artemisia annua L., 1916 J. Bright s.n. Eurasia
Artemisia vulgaris L. var. vulgaris, 2004 E. Zimmerman 292
urasia
Bellis perenis L., 1985 D. Krueger #25 pu CUP
Bidens aristosa (Michx.) Britt., 2000 R. Coxe
Bidens bipinnata L., 1970 WE. Buker s.n.
Bidens cernua L., 2006 L. Speedy LS-06-965
Bidens frondosa L., 2007 L. Speedy and et al. 07-1606
Bidens tripartita L., 2006 L. Speedy LS-06-851
Bidens vulgata Greene, 1916 O.E. Jennings and G.K. Jennings
s.n.
Brickellia eupatorioides (L) Shinners var. eupatorioides, 1928
Centaurea biebersteinii DC., 1981 EH. Utech 81-428 Europe
Centaurea jacea L., 1999 S.P. Grund 2387 Europe
Centaurea nigra L., 1920 J. Bright s.n. Europe
Cichorium intybus L., 2006 L. Speedy LS-06-573 Europe
Cirsium altissimum (L.) Spreng., 1994 J.A. Isaac 6072
Cirsium arvense (L.) Scop., 1987 R.S. Mulvihill and VJ. Mosca
48 Eurasia
Cirsium muticum Michx. 2007 L. Speedy 07-1545
Cirsium pumilum (Nutt) Spreng., 1947 OE. Jennings s.n.
Cirsium vulgare (Savi) Ten, 1931 OE. Jennings s.n. Eurasia
io y coelestinum {L.) DC., 2006 L. Speedy LS-06-978
ae a (L) Cronq. var. canadensis, 2007 L. Speedy
07-1812
Coreopsis lanceolata L., 1999 R.B. Coxe s.n.
Coreopsis tinctoria Nutt. var. tinctoria, 2 E E. Jennings s.n.
Coreopsis tripteris L., 2007 L. Speedy 0
Crepis capillaris (L.) Wallr., 1987 A.W. ers m 20 Furope
Doellingeria infirma (Michx) Greene, 1942 J. Bright 19041
[Aster infirmus Michx.]
Doellingeria ee (P. Mill) Nees var. umbellata, 2006 L.
Erechtites beca (L) Raf. ex DC. var. hieraciifolia, 1970
Erigeron annuus (L.) Pers., 1999 R.B. Cox
ood philadelphicus L. var. id 1999 R.B. Coxe
B pulchellus Michx. var. pulchellus, 1939 O.E. Jennings
s.n.
Erigeron strigosus Muhi. ex Willd. var. strigosus, 1999 R.B.
oxe s.n.
Eupatorium altissimum L., 2006 L. Speedy LS-06-833
Eupatorium perfoliatum L. var. perfoliatum, 2007 L. Speedy
07-1055
Eupatorium rotundifolium L. var. ovatum (Bigelow) Torr., 2006
L. Speedy LS-06-839 G5:S3
Eupatorium serotinum Michx., 2006 L. Speedy LS-06-850
Eupatorium sessilifolium L. var. sessilifolium, 1940 O.E. Jen-
nings s.n.
Eutrochium fistulosum (Barratt) E. Lamont, 1981 FH. Utech
81-433
Eutrochium maculatum (L) E. Lamont var. maculatum, 2001
J. Polonoli FG89
Eutrochium purpureum (L.) E. Lamont var. purpureum, 2006 L.
Speedy LS-06-1006
1457
Eurybia ird. (L) Nesom, 2006 L. Speedy LS-06-945 [Aster
diva L]
Eurybia e (L.) Cass., 2006 L. Speedy LS-06-838 [Aster
macrophylla L.
Eurybia radula (Ait. Nesom, 1961 D.H. Ross 34 G5:S2 [Aster
radula Ait.)
a schreve (Nees) Nees, 2006 L. Speedy ana J. Nusser
63 [Aster schreberi Nees]
En a pd (L) Greene var. graminifolia, 2007 L.
Speedy 07-1622
Galinsoga quadriradiata Cav., 2004 E. Zimmerman 291 Central
and Sout ic
go purpurea (L) Cabrera, 1939 O.E. Jennings and
KJ N.
ah uliginosum L., 1987 A.W. Cusick 27122 Europe
Hasteola suaveolens (L.) Pojark., 1903 O.P. Medsger s.n.
Helenium autumnale L. var. autumnale, 2007 L. Speedy 07-
173
Helianthus decapetalus L., 2006 L. Speedy and D. Sofran
$-06-726
Helianthus divaricatus L., 1951 L.K. Henry and W.E. Buker s.n.
Helianthus giganteus L., 1950 L.K. Henry and W.E. Buker s.n.
Helianthus hirsutus Raf, 2007 L. Speedy LS 07-1537 G5:S2
Helianthus microcephalus Torr. & Gray, 2006 L. Speedy LS-
6-95
Helianthus strumosus L, 1951 L.K. Henry and W.E. Buker s.n.
Helianthus tuberosus L., 1957 L.K. Henry s.n.
Heliopsis helianthoides (L) Sweet var. helianthoides, 2006 L.
d D. Sofran LS-06-725
Hieracium caespitosum Dumort., 2007 L. Speedy LS-07-404
Furope
Hieracium gronovii L., 1961 W.E. Buker
Hieracium Mic L. var. fasciculatum [use Lepage, 1942
nnings s.n. G5:S3
NH. paniculatum L., 2006 L. Speedy and J. Nusser
S-06-673
Hieracium scabrum Michx. var. scabrum, 1966 W.E. Buker s.n.
Hieracium venosum L., 2006 L. Speedy LS-06-553
ochaeris radicata L., 2004 E.A. Zimmerman 222 Eurasia
lonactis linariifolius (L.) Greene, 2001 J. Polonoli FG91
A biflora (Walt.) Blake var. biflora, 1961 D.H. Ross 9
actuca biennis (Moench) Fern., 2007 L. Speedy 07-1702
T tuca canadensis L., 1953 L.K. Henry and F.H. Beer s.n.
Lactuca oe (L) Gaertn. var. floridana, 2007 L. Speedy
7-1
Lactuca hs Muhl. ex Nutt. var. sanguinea (Bigelow) Fern.,
1900 J.A. Shafer s.n. G5?:S3
e. communis L., 1999 S.P. Grund 2353 Europe
d vulgare Lam., 2007 L. Speedy LS-07-409
e [Chrysanthemum leucanthemum L.]
laris scariosa (L.} Willd. var. scariosa,1917 B.H. Patterson s.n.
hasta pad Beadle. 8: FE. Boynt., 2001 J. Polonoli
a acuminata (Michx) Greene, 1987 A.W. Cusick 26997
Aster acuminata Michx.
Packera qurea E A. & D. Love, 2006 L. Speedy LS-06-155
[Senecio aureus L]
1458
Packera obovata (Muhl. ex Willd.) W.A. Weber & A. Love, 2006 L.
LS-06-184 [Senecio obovatus Muhl.ex Willd]
Packera paupercula (Michx.) A. & D. Love, 1915 B.H. Patterson
s.n. [Senecio paupercula Michx.]
Prenanthes alba L., 1916 J. Bright s.n
Prenanthes altissima L., 2006 L. Speedy LS-06-981
Prenanthes crepidinea Michx., 2007 L. Speedy LS-07-121
Prenanthes trifoliolata (Cass.) Fern., 1947 W.E. Buker s.n.
Pseud hali ii( , 1933 L. Lincoln
s.n. [Gnaphalium macounii Greene]
Pseudognaphalium obtusifolium (L.) Hilliard & Burtt ssp. obtusi-
folium, 195 . Buker s.n. [Gnaphalium obtusifolium L]
oe hirta L. var. hirta, 1965 H.H. Fong and FH. Pettler
M. Fylypiw WP-1143
Mod fitta L. var. pulcherrima Farw., 1999 R.B. Coxe s.n.
Rudbeckia laciniata L. var. laciniata, 1999 R.B. Cox
Rudbeckia triloba L. var. triloba, 1950 L.K. Henry and W.E.
er sn.
Sericocarpus asteroides (L.) B.S.P, 1923 J. Bright s.n.
Silphium perfoliatum L. var. perfoliatum, 1941 OE. Jennings
s.n.
APAM cid E val rada a F Lewis 518
|, 2007 L. Speedy
07-1446-1 G4G5: SRIPolymnia uvedalius Ù LJ
Solidago aitissima L., 2006 L. Speedy LS-06
Solidago arguta Ait. var. arguta, 1987 AW. aa 26995
Solidago bicolor L., 2006 L. Speedy LS-06-830
Solidago caesia L., 2007 L. Speedy 07-1653
Solidago canadensis L. var. canadensis, 1981 EH. Utech 81-
496
Solidago canadensis L. var. hargeri Fern.,1910 OE. Jennings
nd G.K. Jennings s.n.
Solidago erecta Pursh, 1901 J.A. Shafer s.n.
Solidago ee È; A L. Speedy 07-1577
Solidago gigantea Ait., 1942 O.F. ae s.n.
sie. a ju 1965 LK. H
Solidago nemoralis Ait. var. a 2006 L. Speedy LS-
06-9
Solidago patula Muhl. ex Willd. var. patula, 2007 L. Speedy
07-1657
Solidago rugosa P. Mill. ssp. aspera, (Ait.) Cronq, 1950 L.K. Henry
and W.E. Buker s.n.
Solidago rugosa P. Mill. ssp. rugosa var. rugosa, 1987 A.W.
Cusick 26919
Solidago rugosa P. Mill. ssp. rugosa var. sphagnophila Graves,
945 OE. Jennings s.n
Solidago speciosa Nutt. var. speciosa, 1915 J. Bright s.n.
G5T5?:SNR
Solidago squarrosa Nutt., 1929 O.E. Jennings s.n.
Solidago uli E Nutt. var. linoides e & ne 1984 S.A.
and J.M. Loughlin 1889
Solidago ces Nutt. var. uliginosa, P 3 A and
Botanical Society of Western PA LS-06-1041 G4G5:$3
um rd Muhl. ex Willd. var. ulmifolia, 1945 O.E.
Jenn N.
Sonchus pin L.) Hill, 1924 J. Bright s.n. Europe
m cordifolium (L.) Nesom, 2006 L. Speedy LS-
6-946 [Aster cordifolius L]
f Texas 2(2)
S / / | (Willd.) N lanceolatum
var. lanceolatum, 2007 L. Speedy and et al. 07- 1603 [Aster
lanceolatus Willd. ]
EE lateriflorum (L.) A. & D. Love var. lateriflorum,
ud L. zc 07- 1713 Dome ) Britton]
I ter) Neson l, 1956 W.E. Buker
s.n. [Aster lowrieanum Porter]
Symphyotrichum novae-angliae (L) Nesom, 2006 L. Speedy
LS-06-1015 [Aster novae-angliae L.
Symphyotrichum patens (Ait) Nesom var. patens, 1970 L.K.
enry s.n. [Aster patens Ait.]
xi dd pregona ANIC ex Za) Bou 1938
| Willd]
Spiga psu (Willd) Nesom var. pee 2006 L.
Speedy LS- 6 [Aster pilosus Willd.
Smphyochom pium (Willd.) Nesom var. pringlei (Gray)
E. Buker s.n. [Aster pilosus Willd. var. pringlei
^ Sra) SF. pos
Nesom, 1950 L.K. Henry arid WE. Bükers s.n. 1 G5: 53 use
praealtum Poir. var. angustior Wie
Symphyotrichum prenanthoides (Muhl. ex Willd.) Nesom,
2006 L. Speedy LS-06-955 [Aster prenanthoides Muhl.
ex Willd.]
Symphyotrichum puniceum (L) A. € D. Love var. puniceum,
2006 L. Speedy LS-06-960 [Aster puniceus L.]
Symphyotrichum shortii (Lindl.) Nesom, 2007 L. Speedy 07-
1629 [Aster shortii Lindl.]
Symphyotrichum undulatum (L.) Nesom, 1981 FH. Utech 81-
473 [Aster undulatum L]
Taraxacum laevigatum (Willd.) DC., 1905 O.E. Jennings s.n.
Eurasia
uocum officinale G.H. Weber ex Wiggers ssp. officinale,
K. Henry and WE. Buker s.n. pues
iuc farfara L., 1959 W.E. Buker s.n
Verbesina alternifolia (L.) Britt. ex o 207 L. Speedy
-1643
Vernonia gigantea (Walt) Trel. ssp. gigantea, 1987 A.W. Cusick
26912
Vernonia noveboracensis (L.) Michx., 2000 R.B. Coxe s.n.
a strumarium L. var. canadense (P. Mill.) Torr. & Gray,
n W. S
"i nac
ied Aum Meerb., 2007 L. Speedy and J. Nusser
7-603
Impatiens pallida Nutt., 2007 L. Speedy 07-1065
Berberidaceae
Berberis thunbergii DC., 2006 L. Speedy LS-06-344 Japan
Po thalictroides (L.) Michx., 2006 L. Speedy LS-
sesona diphylla (L) Pers. 1950 L.K. Henry and FH. Beer
Poy peltatum L., 2006 L. Speedy and D. Sofran LS-
Betulaceae
Alnus incana (L) Moench ssp. rugosa (Du Roi) Clausen, 2007
L. Speedy 07-1038
Morton and Speedy, Vascular plants of Fayette County, Pennsylvania
Alnus ae (Ait) Willd., 2006 L. Speedy LS-06-162
Betula alleg Britt. var. alleghaniensis, 2006 L. Speedy
LS-06-1013
Betula alleghaniensis Britt. var. macrolepis (Fern.) Brayshaw,
934 OL. Jennings s.n.
Betula lenta L., 2007 L. Speedy 07-1731
Carpinus caroliniana Walt. ssp. virginiana (Marsh.) Furlow, 2006
S 69
Corylus americana Walt., 2006 L. Speedy LS-06-440
Ostrya virginiana (P. Mill) K. Koch var. virginiana, 2006 L.
Speedy LS-06-32
Bignoniaceae
Campsis ok (L) Seem. ex Bureau, 1901 J.A. Shafer s.n.
Catalpa bignonioides Walt., 1940 J.F. Lewis
e od (Warder) Warder ex ue 2007 L. Speedy
Boraginaceae
Cynoglossum officinale L., 2004 E.A. Zimmerman 212 Eurasia
re virginianum L. var. virginianum, 1961 L.K. Henry
Ec aum vulgare L. var. vulgare, 1999 R.B. Coxe s.n. Europe
Hackelia virginiana (L) .M. Johnston, 2007 L. Speedy 07-
1454
Heliotropium europaeum L., 1972 A.W. Cusick 12647 Europe
Lappula squarrosa (Retz.) Dumort., 1924 J. Bright s.n. Eurasia
dirae latifolium Michx., 2006 L. Speedy LS-06-624
«S4
"dcs virginica (L.) Pers. ex Link, 1950 L.K. Henry and FH.
Myosotis laxa Lehm., 1999 R.B. Coxe s.n.
2d E L., 2007 L. Speedy 07-1019 Europe
Ono Michx. ssp. hispidissimum (Mackenzie
"ini ien im Shafer 172 G4G5T4:S1
—
Brassicaceae
Alliaria petiolata (Bieb.) Cavara & Grande, 2006 L. Speedy
LS-06-133 Europe
Arabis canadensis L., 2004 R. Coxe s.n.
Arabis glabra (L.) Bernh., 1923 OE. Jennings and E.H. Graham
Arabis laevigata (Muhl. ex Willd.) Poir. var. laevigata, 2006 L.
-06-103
arbarea vulgaris Ait. f, 1987 FH. Utech 87-128 Eurasia
Brassica juncea (L.) Czern., 1924 J, Bright s.n. Eurasia
Brassica nigra (L.) W.DJ. Koch, 1901 J.A. Shafer s.n. Eurasia
Brassica rapa L. var. rapa, 1929 J. Bright 2561 Europe
Capsella bursa-pastoris (L.) Medik, 1950 L.K. Henry and FH.
. Eurasia
Cardamine angustata O.E. Shulz, 2007 L. Speedy LS-07-103
Cardamine bulbosa (Schreb. ex Muhl.) B.S.P, 2007 L. Speedy
L5-07-182
Cardamine concatenata (Michx.) Sw., 2006 L. Speedy LS-
06-12
Cardamine diphylla (Michx.) Wood, 2006 L. Speedy LS-06-
7
13
Cardamine flexuosa With., 1975 W.E. Buker s.n. Europe
Cardamine parviflora L. var. arenicola (Britt) O.E. Schulz, 1924
1459
Cardamine pensylvanica Muhl. ex Willd., 2006 L. Speedy
LS-06-152
E rotundifolia Michx., 2007 L. Speedy and M. Bow-
Drava ve vena L, al, WE. Buker s.n. Europe
1904 OE. Jennings s.n. Eurasia
Heer matranall L., 2006 ft Speedy and D. Sofran LS-06-
226 Europe
Lepidium campestre (L) Ait. f, 1999 R.B. Coxe s.n. Eurasia
Lepidium virginicum L. var. virginicum, 1939 W.G. Burig s.n.
Raphanus raphanistrum L. var. raphanistrum, 1942 OE. Jen-
nings s.n. Mediterranean
Raphanus sativus L., 1948 W.E. Buker s.n. Mediterranean
Rorippa nasturtium-aquaticum (L.) Hayek, 2006 L. Speedy
LS-06-134
Rorippa palustris (L) Bess. ssp. fernaldiana (Butters & Abbe)
Jonsell, 1905 J.A. Shafer and G.E. Kinzer s.n
Rorippa sylvestris (L.) Bess., 1987 A.W. Cusick 27121 Europe
Sinapis alba L., 1948 W.E. Buker s.n. Mediterranean
Sinapis arvensis L., 1902 J.A. Shafer s.n. Mediterranean
Sisymbrium altissimum L, 1923 OE. Jennings and EH. Graham
s.n. Eurasia
Thlaspi arvense L., 1975 W.F. Buker s.n. Europe
Callitrichaceae
Callitriche heterophylla Pursh ssp. heterophylla, 1950 W.E.
Buker s.n.
Callitriche palustris L., 1989 C. Bier and T. Bier s.n.
Callitriche terrestris Raf., 1939 O.E. Jennings s.n.
Calycanthac
Calycanthus us L. var. floridus, 1939 O.F. Jennings s.n.
Campanulaceae
Campanula aparinoides Pursh, 1948 D.H. Krouse s.n.
a ~ americanum (L) Small, 2007 L. Speedy
m RM L., 2006 L. Speedy Fs ^: Sofran LS-06-728
Lobelia inflata L., 2007 L. Speedy 07-17
Lobelia ipi L. var. siphilitica, s L. Speedy and D.
Sofran LS-06-929
Lobelia spicata Lam. var. spicata, 1965 L.K. Henry s.
A perfoliata (L.) Nieuwl. var. perfoliata, Ht C.M.
oardman s.n.
Cannabac
a um Sieb. € Zucc., 1991 EH. Utech 91-2024
Bonus lupulus L. var. lupuloides E. Small, 2006 L. Speedy
Caprifoliaceae
Diervilla lonicera P. Mill, 1902 J.A. Shafer and O.P. Medsger
s.n.
Lonicera canadensis Bartr. ex Marsh., 1932 W.R. VanDersal s.n.
Lonicera dioica L, 1932 W.R. VanDersal s.n.
Lonicera japonica Thunb, 2007 L. Speedy 07-1805 Asia
Lonicera maackii (Rupr) Herder, 1999 RB. Coxe s.n. Asia
Lonicera morrowii Gray, 2007 L. Speedy 07-1650 Japa
Sambucus $n L. ssp. canadensis (L) R. Bolli, 2006 Speedy,
L. LS-06-34
1460
Sambucus racemosa L. var. racemosa, 2006 Speedy, L. LS06-
146-2
dissi albus (L) Blake var. laevigatus (Fern.) Blake,
6W
"n aurantiacum Bickn. var. aurantiacum, 1921 E.H.
McCleiia
Triosteum a Bickn. var. glaucescens Wieg., 1929
OE. Jennings s.n.
Viburnum acerifolium L., 2007 L. Speedy 07-1061
Viburnum dentatum L. var. dentatum, 1999 R.B. Coxe s.n.
Viburnum lentago L., 1946 O.E. Jennings s.n
Viburnum nudum L. var. cassinoides (L.) Torr. & Gray, 2006
L. Speedy and T. Pearson and D. Bryant LS-06-761 G5:S1
Viburnum prunifolium L., 2006 L. Speedy LS-06-197
Viburnum recognitum Fern., 2006 L. Speedy LS-06-434
Caryophyllaceae
Agrostemma githago L., 1984 EH. Utech 84-377 Europe
Arenaria serpyllifolia L, 1999 R.B. Coxe s.n. Europe
Cerastium eet m Baumg. ssp. eel m Greuter
& t, 1923 E.H. McClelland s.n
e gode Thuill, 1939 O.E. e s.n. Eurasia
Cerastium nutans Raf. var. nutans, 1957 L.K, Henry s.n.
Dianthus armeria L., 2007 L. Speedy LS-07-408 Europe
Dianthus barbatus L., 1999 R.B. Coxe s.n. Eurasia
Moehringia lateriflora (L.) Fenzl, 1946 H.N. Mozingo s.n.
Myosoton aquaticum (L.) Moench, 1997 S.P. Grund 1787
Europe
Paronychia canadensis (L.) Wood, 2006 L. Speedy LS-06-565
Paronychia fastigiata (Raf.) Fern. var. fastigiata, 1939 W.G.
naria firi
is L, 2007 L. Speedy VE d Europe
Silene antirrhina L, 1930 WR. VanDersal 1
s latifolia Poir., 2006 L. Speedy and K. jm LS-06-468
rope
ee nivea (Nutt.) Muhl. ex Otth, 1905 B.C. Van Pitt No. 145,
PH
Silene stellata (L.) Ait. f., 2006 L. Speedy LS-06-564
ilene vulgaris (Moench) Garcke, 1952 C.M. Boardman s.n.
Europe
Spergula arvensis L., 1905 J.A. Shafer s.n. Eurasia
Stellaria graminea L., 1946 O.E. Jennings s.n. Europe
S n Nod ein: ex ea var.longifolia, 2007 L. Speedy
sei a a ón da 2007 L. Speedy LS-07-93
rope
mos pubera Michx., 2006 L. Speedy LS-06-111
Celast
pea or Thunb., 2006 L. Speedy LS-06-950 Japan
nd Chin
Celastrus Mcr L., 1999 R.B. Coxe s.
uonymus alatus (Thunb) Sieb. 3007 L. Speedy 07-1652
cuina and Jap an
i us L, 1964 W.E. Buker s.n.
Euonymus an P Jacq. var. atropurpurea, 1935 J.F.
ssn
E TT Nutt., 1946 H.N. Mozingo s.n.
[] [pe ak Dat -ali D h een
f Texas 2(2)
Chenopodiaceae
Chenopodium album L. var. missouriense (Aellen) |.J. Bassett &
C.W. Crompton, 1950 L.K. Hen
Chenopodium botrys L., 1902 J.A. EE s.n. Euro
Chenopodium simplex (Torr.) Raf, 1940 O.E. pecus s.n.
Cistaceae
Lechea racemulosa Michx., 2006 L. Speedy LS-06-560
Clethraceae
Clethra acuminata Michx., 2006 L. Speedy LS-06-633 G4:S1
Clusiaceae
Hypericum ascyron L, 1919 O.E. Jennings and G.K. Jennings
s.n.
Hypericum canadense L., 2006 L. Speedy and K. Jurkovic
-06-680
Hypericum densiflorum Pursh, 2007 L. Speedy 07-1541
G5:53
iru ellipticum Hook., 2007 L. Speedy and M. Bowers
pecan gentianoides (L.) B.S.P, 2007 L. Speedy and Botanical
Socie estern PA 07-1553
Then hypericoides (L.) Crantz ssp. multicaule (Michx. ex
Willd.) Robson, 1915 J. Bright s.n.
Hypericum mutilum L., 2006 L. Speedy and K. Jurkovic LS-
Hypericum perforatum L., 1999 R.B. Coxe s.n. Europe
DE prolificum L., 2007 L. Speedy and M. Bowers
Te icum m Lam., 1965 L.K. Henr
Triadenum fraseri p Gleason, 2006 L. pum and K.
Jurkovic LS-06-68
Triadenum m (L) Raf, 1984 S.A. Thompson and J.
Loughlin 1888
Commelinaceae
Commelina communis L. var. communis, 2000 R. Coxe s.n.
Asia
Tradescantia virginiana L., 1985 D. Krueger 166-B, CUP
Convolvulaceae
n sepium (L.) R. Br. ssp. appalachiana Brummitt, 1981
-222
ao o (Kit.) Griseb. ssp. fraterniflora (Mackenzie &
Bush mmitt, 1950 L.K. Henry and W.E. Buker s.n.
wo arvensis L., 1948 L.K. Henry s.n. Europe
Ipomoea pandurata (L.) G.FW. Mey., 1950 W.E. Buker s.n.
Cornaceae
Cornus alternifolia L. £, 1946 H.N. Mozingo s.n.
Cornus amomum P. Mill, 2006 L. o: LS-06-1018
Cornus florida L., 2006 L. Speedy LS-0
Cornus obliqua Raf, 2007 L. Speedy Ue in
Cornus racemosa Lam., 1999 R.B. Coxe s.n
Nyssa sylvatica Marsh., 2006 L. Speedy and M. Bowers LS-
06-277
Crassulaceae
Hylotelephium telephium (L.) H. Ohba. ssp. telephium, 1999 R.B.
Coxe s.n. Eurasia [Sedum telephium L.
f.
tv. Pennsvivania
ef ES
Penthorum sedoides L., 2005 B.C. Van Pitt No. 99, PH
Sedum ternatum Michx., 2007 L. Speedy LS-07-416
ucurbita
Emo eta (Michx.) Torr. & Gray, 2007 L. Speedy
07-1
Sicyos m L., 2006 L. Speedy LS-06-947
Cuscutaceae
Cuscuta gronovii Willd. ex J.A. Shultes var. gronovii, 1999 S.P.
Cuscuta pentagona Engeim. var. pentagona, 1982 F.H. Utech
82-389
Cyperaceae
Bulbostylis capillaris (L) Kunth ex C.B. Clarke ssp. capillaries,
197 i
Carex albicans Willd ex. Spreng var. albicans, 1960 W.E. Buker
s.n.
Carex albolutescens Schwein., 1924 J. Bright s.n.
Carex albursina Sheldon, 2006 L. Speedy LS-06-563
Carex amphibola Steud., 1918 J. Bright s.n
Carex annectens (Bickn.) Bickn., 1946 O.E. Jennings s.n.
Carex argyrantha Tuckerman, 1954 W.F. Buker
Carex atlantica Bailey ssp. atlantica, e : Speedy 07-538
Carex baileyi Britt, 1930 W.R. VanDersal 1
Carex bromoides Schkuhr ex Willd. ssp. ae 1999 R.B.
Coxe s.n.
Carex careyana Torr. ex Dewey, 1997 S.P. Grund 1971
G4G5:S1
Carex cephaioidea (Dewey) Dewey, 1924 J. Bright s.n.
Carex cephalophora Muhl. ex Willd., 1908 O.E. Jennings s.n.
rex communis Ep var. communis, 2006 L. Speedy and K.
e ic LS-06
Carex crinita Lam. var. crinita, 1908 B.H. Patterson s.
Carex debiiis Michx. var. rudgei Bailey, 2007 L. Hug 07-545
Carex D Willd. var. digitalis, 2006 L. Speedy and J. Nusser
LS-0
Cal p. echinata, 1951 W.E. Buker
oe o Es 2 L. Speedy and K. ne Ls- 06-
695
Carex frankii Kunth, 1999 R.B. Coxe s.n.
Carex glaucodea Tuckerman ex Olney, 1889 J.A. Shafer s.n.
Carex gracilescens Steud., 2006 L. Speedy LS-06-620
Carex gracillima Schwein., 1905 O.E. Jennings s.n.
Carex granularis Muhl. ex Willd., 1918 J. Bright s.n.
Carex grayi Carey, 1932 J. Bright 5453
Carex gynandra Schwein., 2007 L. Speedy and M. Bowers
7-94
Carex hirsutella Mackenzie, 2003 L. Speedy and C.M. Bow-
ers s.n.
Carex hirtifolia Mackenzie, 1921 J. Bright s.n.
Carex hitchcockiana Dewey, 2006 L. Speedy LS-06-627
Carex interior Bailey, 2007 L. Speedy 07-531
Carex intumescens Rudge, 2007 L. Speedy 07- Nie
Carex jamesii Schwein., 2007 L. Speedy i S-07-
Carex laevivaginata (Kukenth.) Mackenzie, Bos L. Speedy
-06-410
Carex laxiculmis Schwein. var. copulata (Bailey) Fern., 1929
E.M. Gress s.n.
1461
Carex laxiculmis Schwein. var. laxiculmis, 1920 O.E. Jennings
s.n.
Carex laxiflora Lam. var. laxiflora, 1992 J.A. Isaac 3907
Carex leavenworthii Dewey, 1940 O.E. Jennings and J.F. Lewis
s.n.
ua E Wahlenb. ssp. leptalea, 2006 L. Speedy LS-
on "eptonervi a Fern., 2006 L. Speedy LS-06-406
in Willd., 2007 L. Speedy 07-1538
Corel lurida Vishlenib. ve L. Speedy LS-06-416
Carex pensylvanica Lam., 2006 L. Speedy LS-06-14
Carex planispicata Naczi, 1929 J. Bright 1892
Carex plantaginea Lam., 2007 L. Speedy 07-609
Carex platyphylla Carey, 2006 L. Speedy LS-06-59
Carex prasina Wahlenb., 2006 L. Speedy LS-06-359
Carex projecta Mackenzie, 1999 Grund, S.P. 2262
Carex radiata (Wahlenb.) Small, 2006 L. d LS-06-561
Carex roanensis FJ. Herm., 1929 J. Bright 1
Carex rosea Schkuhr ex Willd., 1992 J.A. Es 3902
Carex scabrata Schwein., 2006 L. Speedy LS-06-408
Carex scoparia Schkuhr ex Willd. var. scoparia, 2007 L. Speedy
and
Carex sparganioides Muhl. ex Willd. var. sparganioides, 1992
J.A. lsaac 3892
Carex stipata Muhl. ex Willd. var. stipata, 2006 L. Speedy
5-06-433
Carex striatula Michx., 2007 L. Speedy 07-616
Carex stricta Lam., 2007 L. Speedy and M. Bowers 07-832
Carex swanii (Fern) Mackenzie, 2006 L. Speedy and M. Bow-
ers LS-06-793
Carex torta Boott ex Tuckerman, 2006 L. Speedy LS-06-156
Carex tribuloides Wahlenb. var. tribuloides, 2007 L. Speedy and
Carex umbellata Schkuhr ex Willd., 1935 L.K. Henry s.n.
— Mea ll ii Willd., 2007 7 ee 07-1074
3 L. Speedy and
CM. Bow: s.n.
Carex yd Schkuhr ex Willd., 1923 J. Bright s.n.
Carex woodii Dewey, 1999 S.P. Grund wit
Cyperus bipartitus Torr., 2006 L. Speedy LS
os esculentus L. var. leptostachyus TN 7 L. Speedy
- 6
E flavescens L., 1997 S.P. Grund pus
Cyperus odoratus L., 1997 Grund, S.P.
Cyperus squarrosus L., 2006 L. bs P e 848
Cyperus strigosus L. 1999 R.B. Coxe s.n
us ium arundinaceum E Britt., 2006 L. Speedy and T.
on and D. Bryant L 6
o acicularis (L.) Re & J.A. Schultes, 1934 E.H.
Eleocharis elliptica Kunth, 1999 S.P. Grund and et al. 2275
Eleocharis Ea anri Steud., 1999 S.P. Grund and R. Coxe
eland 2359
Eleocharis aia Steud., 1949 OE. Jennings s.n.
Eleocharis obtusa (Willd.) J.A. Schultes, 2006 L. Speedy LS-
06-446
Eleocharis palustris (L) Roemer & J.A. Schultes, 1986 C. Bier
and B. Woodworth and M. Bowers s.n.
1462
Eleocharis tenuis (Willd.) J.A. Schultes var. pseudoptera (Weath-
vens.) Svens.,1940 OE. Jennings s.n.
d tenuis (Willd) J.A. Schultes var. tenuis, 1999 R.B.
Cox
Recta ms D J.A. Schultes var. verrucosa (Svens.)
Sve peed 07-540 G5T3T5:S1
E. 2006 L. Speedy
of Western PA LS-06-1040
Fimbristylis autumnalis (L) Roemer & J.A. Schultes, 1997 S.P.
Grund 2008
Ic
X
Rhynchospora alba (L.) Vahl, 2006 L. Speedy and K. Jurkovic
LS-06-6
Rhynchospora capitellata (Michx.) Vahl, 2006 L. Speedy and
K. Jurkovic LS-06-685
Rhynchospora recognita (Gale) Kral, comb. nov. ined., 1974 R.
Fulton s.n. G5?:S1
Schoenoplectus acutus (Muhl. Ex Bigelow) A. & D. Love var.
acutus, 1949 W.E. Buker s.n. G5:S2
Schoenoplectus pungens (Vahl) Palla var. pungens, 1999 S.P.
ax
N
[er
M pa punhianus (Fern.) M.T. Strong, 2006 L. Speedy
and K. Jurko -06-688
mice eh s m (K.C. Gmel.) Palla, 1977 M.
Polechko s.n.
Scirpus atrovirens Willd., 1999 R.B. Coxe
Scirpus cyperinus (L) Kunth, 2007 L. an and M. Bowers
Scirpus expansus Fern., 2007 L. Speedy and M. Bowers 07-
940
Scirpus hattorianus Makino, 1999 RB. Coxe s.n.
pode microcarpus J. & à is 1929 OL. BL s.n.
Scirpus pendulus Muhl., 1 OE. Jennin
Scirpus UR Vahl, n L. Sas n Jurkovic LS-
Dioscoreaceae
Dioscorea oppositifolia L., 2007 L. Speedy and et al. 07-1601
Chin
na
Dioscorea quaternata J.F. Gmel., 2006 L. Speedy LS-06-400
Dioscorea villosa L., 1999 R.B. Coxe s.n.
Dipsacaceae
Dipsacus fullonum L., 1991 S.A. Thompson 8905 Europe
Droseraceae
Drosera ien iy 2006 L. Speedy and T. Pearson and D.
S-06-
Drosera ee ea e var. rotundifolia, 2006 L. Speedy and K.
Jurkovic LS-06-705
Elaeagna
Elaeagnus dmm Thunb, 2006 L. Speedy LS-06-117 Asia
Ericaceae
Epigaea repens L., 2006 L. Speedy and M. Bowers m
Gaultheria procumbens L., 2006 L. Speedy LS-06
sous baccata toe K. Koch, 2006 L. m
Richards LS-06-2
"mis es L., 2006 i Speedy LS-06-415
Lyonia iiie (L) DC. var. ligustrina, 2007 L. Speedy and
M. Bowers 07-833
£al Dat * 4l D |" T
Journal Ul
f Texas 2(2)
Oxydendrum arboreum (L.) DC., 2000 R. Coxe s.n. G5:S354
Rhododendron arborescens (Pursh) Torr., 2007 L. Speedy
7-539
Rhododendron maximum L., 2006 L. Speedy and M. Bowers
-06-517
Rhododendron periclymenoides (Michx.) Shinners, 2006 L.
Speedy and M. Bowers LS-06-280
Rhododendron viscosum (L.) Torr., 2006 L. Speedy and E.
Richards LS-06-300
Vaccinium angustifolium Ait, 1995 B.L. Isaac and J.A. Isaac
Vaccinium corym L, 2006 L. Speedy LS-06-205
Vaccinium macrocarpon Ait., 2006 L. Speedy and K. Jurkovic
-06-
6
Vaccinium oxycoccos L., 2006 L. Speedy and F. Richards LS-
06-298
Vaccinium pallidum Ait., 2006 L. Speedy LS-06-206
Vaccinium stamineum L., 2006 L. Speedy and M. Bowers
6-272
Euphorbiaceae
Acalypha rhomboidea Raf., 2007 L. Speedy 07-1575
Acalypha virginica L., 1902 J.A. Shafer and O.P. Medsger s.n.
Chamaesyce maculata (L.) Small, 1999 S.P. Grund and et al.
2349
Chamaesyce nutans (Lag.) Small, 1970 LK. Henry s.n.
Euphorbia corollata L., 2007 L. Speedy and M. Bowers 07-
683
pun cyparissias L, 1915 J. Bright s.n. Eurasia
a dentata Michx. var. dentata, 2006 L. Speedy LS-
"mun ive L., 1934 EB. Burgess s.n. Europe
Euphorbia marginata Pursch, 1941 O.E. Jennings s.n.
Fabaceae
Amorpha fruticosa L., 2000 R. Coxe s.n.
Amphicarpaea bracteata (L.) Fern. var. bracteata, 2007 L.
Speedy
Amphicarpaea bracteata (L) Fern. var. comosa (L.) Fern., 1950
L.K. Henry and W.E. Buker s.n.
Apios americana Medik., 2006 L. Speedy and BSWP LS-06-
755
Astragalus canadensis L. var. canadensis, 1942 O.E. Jennings
G5:S2
Baptisia australis (L.) R. Br. ex Ait. f. var. australis, 2007 L. Speedy
07-1610 G5:S3
Baptisia tinctoria (L) R. Br. ex Ait. f, 2007 L. Speedy and M.
Bowers 07-
dd canadensis L. es uda. 2006 L. nd m mi pa 97
ni titans var.
006 L. Speedy LS- ee
coil varia L, ae L. eed Ma 627 Europe
8 LK Henry s.n.
Desmodium nena ) DC, a L. Speedy 07-1026
Desmodium glutinosum (Muhl. ex Willd.) Wood, 1950 W.E.
Desmodium laevigatum (Nutt.) D.C., 1964 J. Lewis s.n. G5:SU
Desmodium nudiflorum (L.) DC., 1957 LK. Henry s.n.
Pennsvlvania
ar ,
Desmodium UR (L) DC. var. paniculatum, 1957 L.K.
nry s.
Desmodium US Schub., 2006 L. Speedy LS-06-831
Ges triacanthos L., 2007 L. Speedy 07-1642
Glycine max (L.) Merr, 1933 J. Bright 8614 East Asia
feud nd L., 2007 L. Speedy and M. Bowers 07-677
Juhl. ex Willd., 2001 S.P. Grund and L. Smith
Europe
Lathyrus
Lespedeza capitata Michx., 1950 L.K. Henry and W.E. Buker
(L) Hornem., 2006 L. Speedy and M.
s.n.
Lespedeza frutescens
wers LS-06-800
Lespedeza hirta (L) Hornem. ssp. hirta, 1950 L.K. Henry and
W. n
spedeza repens (L) W. Bart, 1915 J. Bright s.n.
= corniculatus L., 2004 E.A. Zimmerman 217
a a L. = eiii var. occidentalis S. o 1948
. Buker
Me ps capo mur n ae S.P. Grund and et al. ~ Eurasia
Melilotus officinalis (L) Lam., 1999 R.B. Coxe s
Phaseolus polystachyios (L.) B.S.P. var. m m J.
Bright s.n. G5:S1S2
Robinia pseudoacacia L., 2007 L. Speedy 07-1694
Senna A (Fern) Irwin & Barneby, 1922 O.E. Jennings
Suc aie biflora (L.) B.S.P, 1995 J.A. Isaac 8062 65:52
m aureum Pollich, 2004 E.A. Zimmerman 218 Eurasia
Trifolium campestre Schreb., 1995 B.L. Isaac and J.A. Isaac
66 Europe
Trifolium hybridum L., 1904 O.E. Jennings s.n. Eurasia
Trifolium pratense L., 2004 E. Zimmerman 289 Europe
Trifolium repens L., 2004 E. Zimmerman 284 Europe
Vicia caroliniana Walt., 1946 H.N. Mozingo
Wisteria sinensis (Sims) DC., 2007 L. peace and M. Bowers
LS-07-249 China
Fagaceae
Castanea dentata (Marsh.) Borkh., 2006 L. Speedy LS-06-403
Fagus grandifolia Ehrh., 2007 L. Speedy 07-1580
Quercus alba L., 2006 L. Speedy LS06-210-1
Quercus bicolor Willd., 1999 R.B. Coxe s.n.
Quercus xbrittonii W.T. Davis (pro spJ, 1940 Jennings, O.E.
s.n.
Quercus coccinea Muenchh. var. coccinea, 1945 O.E. Jen-
nings s.n.
Quercus ilicifolia Wangenh., 1969 S. Chamberlain s.n.
us ee Michx., 2007 L. Speedy and M. Bowers
aas ion Engelm., 1910 OE. Jennings and G.K.
nnings s.n
Pal palustris Muenchh., 2000 R. Coxe s.n.
Quercus prinus L., 2007 L. Speedy 07-1811
Quercus rubra L. var. ambigua (Gray) Fern., 2007 L. Speedy
07-1076-1
Quercus rubra L. var. rubra, 2007 L. Speedy 07-1648-1
1463
Quercus velutina Lam., m L. Speedy and Botanical Society
of Western PA 07-15
Fumariaceae
dlumia fungosa (Ait.) Greene ex B.S.P, 2004 R. Coxe s.n.
Corydalis flavula (Raf) DC., 2006 L. Speedy LS-06- s:
Corydalis sempervirens (L.) Pers., 1992 J.A. Isaac 390
Dicentra canadensis (Goldie) Walp., 2006 L. cM LS-06-35
Dicentra cucullaria (L.) Bernh., 2007 L. Speedy LS-07-13
Gentianaceae
Bartonia virginica (L.) B.S.P, 2006 L. Speedy and T. Pearson and
D. Bryant LS-06-779
Gentiana andrewsii Griseb. var. andrewsii, 1981 F.H. Utech
81-427
Gentiana clausa Raf., 2007 L. Speedy 07-1662
Gentiana linearis Froel., 1952 L.K. Henry and W.E. Buker s.n.
Gentiana saponaria L. var. saponaria, 2006 L. Speedy and
Botanical Society of Western PA LS-06-1035 G5:S1S2
SEIS OEERE
ornnii Imhir
mu 1964 J.F. Lewis s.n. Europe
Geranium alla L 2006 L. Speedy LS-06-190
Grossulariace
Ribes pa E 2007 L. Speedy 07-624
Ribes rotundifolium Michx., 1999 R.B. Coxe s.n.
Ribes rubrum L., 1995 J.A. Isaac 7047 Eurasia
Haloragaceae
Myriophyllum heterophyllum Michx., 1997 S.P. Grund 2001
G5: S1
Hamamelidaceae
Hamamelis virginiana L., 2007 L. Speedy 07-1849
Hippocasta
ime pa ne p L. Speedy and Botanical Society of
7-1613-1
mes glabra Willd., 1957 L.K. Henry s.n.
ydrangeaceae
Deutzia scabra Thunb., 2007 L. Speedy 07-1649 Asia
Hydrangea arborescens L., 2006 L. Speedy LS-06-944
Hydrocharita
Elodea eos EIER ) St. John, 1977 M. Polechko s.n.
Vallisneria americana Michx., 2006 L. Speedy and D.Sofran
Hydrophyllaceae
io eai appendiculatum Michx., 2006 L. Speedy LS-
06-
"muni canadense L., 1997 S.P. Grund 2019
Hydrophyllum virginianum L., 2006 L. Speedy LS-06-183
Phacelia purshii Buckl., 2006 L. Speedy LS-06-180
Iridaceae
Iris germanica L, 1951 W.E. Buker s.n. Europe
Iris prismatica Pursh ex Ker-Gawl., 1942 O.E. Jennings s.n.
G4G5:S1
Iris versicolor L, 1942 O.E. Jennings s.n.
Sisyrinchium angustifolium P. Mill, 1999 R.B. Coxe s.n.
1464.
Juglandaceae
Carya alba (L.) Nutt. ex Ell., 2006 L. Speedy LS06-977-2
Carya cordiformis (Wangenh.) K. Koch, 2006 L. Speedy and
. Bowers LS-06-528
Carya glabra (P. Mill) Sweet, 1950 OE. Jennings s
Carya ovalis (Wangenh.) Sarg., 1949 OE. poss and EH.
Graham s.n.
Carya ovata (P. Mill) K. Koch, 2006 L. Speedy L506-976-1
Juglans cinerea L., 2006 L. Speedy and J. Nusser and M. Bow-
ers LS06-599-1
Juglans nigra L., 1920 O.E. Jennings s.n.
Juncaceae
Juncus acuminatus Michx., 2006 L. Speedy and K. Jurkovic
-06-684
Juncus articulatus L., 1999 S.P. Grund and et al. 2346
Juncus brachycephalus (Engelm.) Buch., 1947 OE. Jennings
Juncus brevicaudatus (Engelm.) Fern, 2007 L. Speedy 07-
Juncus bufonius L. var. bufonius, 1929 O.E. Jennings s.n.
Juncus ed J. Gay ex Laharpe, 2006 L. Speedy and T.
Pearson and D. Bryant LS-06-784
Juncus debil Gray, 2006 L. Speedy and K. Jurkovic LS-06-707
G5:S3
5:
Juncus dudleyi Wieg., 1924 J. Bright s.n
Juncus E. L. var. pylaei pia Fern. & Wieg., 2006 L.
edy and K. Jurkovic LS-06-7
Juncus efus " var. Pus Fern. m 2006 L. Speedy and
ncus ibi E 2000 R.B. Coxe
mcis subcaudatus (Engelm.) Coville & ek var. a
999 S.P. Grund and R. Coxe and E. Cope
ms: tenuis Willd., 2006 L. Speedy and M. pen LS-06-
795
Luzula acuminata Raf. var. acuminata, 2006 L. Speedy LS-
06-140
Luzula acuminata Raf. var. carolinae (S. Wats.) Fern., 1915 J.
Luzula echinata (Small) F.J. Herm., 1938 LK. Henry s.n.
Luzula multiflora (Ehrh.) Lej. ssp. multiflora var. multiflora, 2007
Speedy
LS-07-130
Lamiaceae
i nepetoides (L.) Kuntze, 1939 W.G. Burig s.n.
Ajuga ns L., 1920 J. Bright s.n. Eurasia
Sei ps (L) Benth., 1950 L.K. Henry and W.E. Buker
T hirsuta (Pursh) Benth. var. hirsuta, 2000 R. Coxe s.n.
Clinopodium vulgare L, 2007 L. Speedy and M. Bowers 07-
694
Collinsonia canadensis L., 2007 L. Speedy 07-1048
Cunila origanoides (L.) Britt., 1917 B.H. Patterson s.n.
Glechoma hederacea L., 1999 R.B. Coxe s.n. Eurasia
Hedeoma pulegioides (L) Pers., 2007 L. Speedy 07-1630
Isanthus brachiatus (L) B.S.P, 1953 L.K. Henry and FH. Beer
Lamium maculatum L., 1995 A. Genovese #12 Eurasia, CUP
Journal of tl f Texas 2(2)
Leonurus cardiaca L. ssp. cardiaca, 1981 FH. Utech 81-219
sia
Lycopus americanus Muhl. ex W. Bart., 1999 R.B. Coxe s.n.
ao pi Michx. var. uniflorus, 1999 S.P. Grund and
al. 235
bap UM L, 2006 L. Speedy and M. Bowers LS-06-
Herne cordata (Nutt.) Britt, 2007 L. Speedy LS-07-414
:51
m officinalis L., 2007 L. Speedy 07-1697 Asia
entha aquatica L, 1924 J. Bright s.n. Eurasia
Med arvensis L., 1999 S.P. Grund and et al. 2334
Mentha spicata L, 1950 L.K. Henry and W.E. Buker s.n. Eu-
rope
Mentha xvillosa Huds. (pro sp.), 1942 O.E. Jennings s.n.
Monarda clinopodia L., 2006 L. Speedy and M. Bowers LS-
06-513
Monarda didyma L., 2006 L. Speedy and M. Bowers LS-06-
512
Monarda fistulosa L. var. fistulosa, 1941 O.E. Jennings s.n.
Monarda media Willd., 2006 L. Speedy and J. Nusser LS-06-
671
Nepeta cataria L., 2006 L. Speedy L5-06-574 Eurasia
Origanum vulgare L., 1934 J.F. Lewis s.n. Mediterranean
Prunella vulgaris L. ssp. lanceolata (W. Bart.) Hulten, 2004 E.
Zimmerma 5
Pycnanthemum incanum (L.) Michx. var. incanum, 1932 J.
Bright 7521
Pycnanthemum tenuifolium Schrad., 2006 L. Speedy LS-06-
629
ET c om (Michx) Pers. var. verticillatum,
Buker
Salvia an L 1995 i L Isaac and J.A. Isaac 7552
Scutellaria elliptica Muhl. ex Spreng. var. wm 2007 L.
Speedy 07-1024
Mi A ] KASSE
|. ex Spreng. var. hirsuta (Sh
1953 W.E. Buker s
E incana Biehler, d L. Speedy LS-06-640
a o L. var. lateriflora, 2007 L. Speedy and M.
07-918
a a Riddell, 2006 L. Speedy and J. Nusser LS-
06-590 G3:51
Stachys palustris L, 1950 D.H. Krouse s.n.
Stachys tenuifolia Willd., 1976 P. Jeffrey #25, CUP
Teucriu L. var. canadense, 1999 R.B. Coxe s.n.
Trichostema dichotomum L., 1953 LK. Henry and FH. Beer
Lauraceae
Lindera benzoin (L.) Blume var. benzoin, 2007 L. Speedy 07-
1667
Sassafras albidum (Nutt) Nees., 2006 L. Speedy LS-06-149
Lemnaceae
Spirodela polyrhiza (L) Schleid., 1977 M. Polechko s.n.
Lentibulariacea
Uti ios macori Le Conte, 2006 L. Speedy and K. Jurk-
cL
Ve [| I + £r ta f
tv. Pennsylvania
ar ,
Liliaceae
Allium canadense L. var. canadense, 2006 L. Speedy LS-06-
357
Allium cernuum Roth var. cernuum, 1941 O.E. Jennings s.n.
Allium tricoccum Ait., 2007 L. Speedy LS-07-124
Allium vineale L. ssp. vineale, 1995 B.L. Isaac and J.A. Isaac
7550 Europe
Chamaelirium luteum (L.) Gray, 1955 W.E. eo s.n.
Clintonia borealis (Ait.) Raf, 1957 W.E. Buker s
Clintonia umbellulata oe Morong, 2006 hi Speedy and
ofran
Ein americanum Ker-Gawl. ssp. americanum, 2006
L. Speedy 3
Hypoxis hirsuta (L.) Coville, 1950 W.E. Buker
ue canadense L. ssp. editorum (Fern.) f 1950 WE.
Buker s.n.
A dd L. var. philadelphicum, 1947 W.E. Buker
Lum. da dl L; a L. edd and D. Sofran LS-06-731
f, 2006 L. Speedy and D. Sofran
LS-06-222
Maianthemum racemosum (L.) Link ssp. racemosum, 2006
L. Speedy and D. Sofran LS-06-219 [Smilacina racemosa
(L) Desf.
Maianthemum stellatum (L.) Link, 1999 S.P. Grund 2398 [Smi-
lacina stellatum (L.) Desf]
Medeola virginiana L., 2006 L. Speedy and K. Jurkovic LS-
06-464
Muscari botryoides (L.) P. Mill., 1960 W.E. Buker s.n. Europe
Narcissus pseudonarcissus L, 1947 W.E. Buker s.n. Europe
e umbellatum L., 1950 L.K. Henry and F.H. Beer
n. Eur
a oum (Walt) Ell. var. ieee, (J.A. & J.H.
Schultes) Morong, 2006 L. Speedy LS-06-177
Polygonatum pubescens (Willd.) Pursh, n L. Speedy LS-
06-112
pon os (Michx.) D. Don, 2006 L. Speedy LS-
06-1
thi Gawl.) Morong var. gramineum,
J. Bright s.n. G4G5: S152
Sens lanceolatus Bi ue var. lanceolatus, 1940 O.E.
Jennings and J.F. Lew
Ps erectum L., 2006 : 3 LS-06-39
Trillium grandiflorum (Michx.) Salisb., 2006 L. Speedy LS-
06-50
Trillium undulatum Willd., 2007 L. Speedy LS-07-384
Uvularia grandiflora Sm., 2006 L. Speedy LS-06-622
Uvularia perfoliata L., 2007 L. Speedy LS-07-411
Uvularia puberula Michx., 2006 L. Speedy and M. Bowers
LS-06-276
Uvularia sessilifolia L., 2006 L. Speedy LS-06-166
Veratrum viride Ait., 1999 R.B. Coxe s.n
Limnanthaceae
Floerkea proserpinacoides Willd., 2006 L. Speedy LS-06-55
Linaceae
Linum striatum Walt., 2006 L. Speedy and K. Jurkovic LS-
1465
Linum virginianum L. var. virginianum, 1941 O.E. Jennings
s.n.
Lythraceae
Cuphea viscosissima Jacq., 1919 O.E. Jennings s.n.
Lythrum salicaria L., 2006 L. Speedy LS-06-841 Europe
Magnoliaceae
Liriodendron tulipifera L., 2006 L. Speedy LS-06-1000
Magnolia acuminata (L) L., 2006 L. Speedy LS-06-566
Magnolia macrophylla Michx. 1932 O.E. Jennings s.n.
Magnolia tripetala (L.) L., 1999 RB. Coxe s.n. 65:52
Malvaceae
Abutilon theophrasti Medik, 1873 B.H. Patterson s.n. Asia
Hibiscus s L, 1883 J.A. Koch s.n. Asia
Malva moschata L., 1948 L.K. Henry s.n. Europe
Sida eee L., 1950 LK. Henry and WEE. Buker s.n.
Melastomatace
Rhexia virginica E p L. Speedy and D. Sofran LS-06-734
Menispermacea
Menispermum uds L, 2007 L. Speedy 07-1464
Mollunginace
Mollugo opel L, 1939 WG. Burig s.n.
=
Monotropaceae
Monotropa hypopithys L., 1999 S.P. Grund 2354
Monotropa uniflora L., 1951 LK. Henry and W.E. Buker s.n.
Moraceae
Maclura pomifera (Raf) Schneid., 1952 A.W. Buvinger s.n., PH
Morus alba L., 2006 L. Speedy LS-06-370 Asia
Morus rubra L. var. rubra, 1931 O.E. Jennings s.n.
Najadaceae
Najas gracillima (A. Braun ex Engelm.) Magnus, 1977 M.
Polechko s.n.
Najas minor All., 1977 M. Polechko s.n. Europe
Nyctaginaceae
Mirabilis nyctaginea (Michx) MacM. 1951 L.K. Henry and
W.E. Buker s.n
Oleaceae
Fraxinus americana L., 2007 L. Speedy and M. Bowers 07-
934
Fraxinus nigra Marsh., 1999 RB. Coxe s.n.
Fraxinus pennsylvanica Marsh., 1934 J.F. L
Ligustrum obtusifolium Sieb. & Zucc., pu uL m LS-06-
366 Japan
Ligustrum vulgare L., 1999 R.B. Coxe s.n. Europe
Onagraceae
Chamerion angustifolium (L.) Holub ssp. ci g
Ka rtesz, 1947 WE. Buker s.n.
a L. ssp. alpina, 1950 W.E. Buker s.n.
pens pond Ehrh. (pro sp), 1941 O.E. Jennings s.n.
Circaea lutetiana L. ssp. canadensis (L) Aschers. & Magnus,
1995 B.L. Isaac and J.A. Isaac 7558
dress ~ Biehler, 2006 L. Speedy and BSWP
P oS Raf, 1965 L.K. Henry s.n.
1466
Gaura biennis L., 1982 FH. Utech and M. Ohara 82-398
us ee L., 2006 L. Speedy and K. Jurkovic LS-
mens ae (L) M. Polechko s.n.
Oenothera biennis L., 1950 LK. Henry and W.E. Buker s.n.
Oenothera fruticosa E. a fruticosa, 1924 R.R. Dreisbach
3131
Oenothera fruticosa L. ssp. glauca (Michx.) Straley, 1981 F.H.
8
Oenothera perennis L., 1954 W.E. Buker s.n.
Orchidaceae
Arethusa bulbosa L., 1950 W.E. Bukerand L.K. Henry and FH.
d M. Henrici s.n. G4:S 1
Calopogon tuberosus (L.) B.S.P var. tuberosus, 1948 W.E. Buker
s.n.
Corallorhiza maculata (Raf) Raf. var. maculata, 1950 W.E.
Corallorhiza odontorhiza (Willd.) Poir. var. odontorhiza, 1968
kii cs ~ Ait, 1997 J.S. Shriver and A. Shriver and
mith 8
Option ian Salisb. var. pubescens dins : Knight,
. Shriver and A. Shriver and C. Smith 8
E spectabilis ( ) Raf, 1962 W.E. Buker s.n.
Goodyera pubescens (Willd.) R. Br. ex Ait. f, 2006 L. Speedy
LS-06-56
Isotria verticillata Raf., 2007 L. Speedy and M. Bowers 07-681
Liparis liliifolia (L) L.C. Rich. ex Ker-Gawl, 2004 E.A. Zimmer-
man 211
Liparis loeselii (L) L.C. Rich., 1965 W.E. Buker s.n.
Malaxis bayardii Fern., 1973 P. Monk s.n. G1G2:51
Malaxis unifolia Michx., 1973 P. Monk s.n
Platanthera ciliaris (L) Lindl., 1947 O.E. Jennings s.n. G5:S2
latanthera clavellata (Michx.) Luer, 1947 O.E. Jennings s.n.
Platanthera flava (L.) Lindl. var. herbiola (R. Br. ex Ait. E) Luer,
1 n.
Platanthera grandiflora (Bigelow) Lindl., 1968 A. Molans s.n.
Platanthera lacera (Michx.) G. Don, 1948 W.E. Buker s.n
Platanthera a (Pursh) Lindi, 2006 L. Speedy and J.
md peramoena e a 1996 JS. Shriver and A.
and C.Smith 6
eds opie ie | 2006 L. Speedy and K.
Jurkovic LS-06-700
¡ciel cernua S L.C. Rich., 2006 L. Speedy and Botanical
of Western PA LS-06-1038
Sponte is ~ Raf. var. gracilis (Bigelow) Luer, 1995
States mes (H.H.Eat) Ames, 1940 OE. Jennings s.n.
G5:
ee ochroleuca (Rybd.) Rybd., 1956 W.E. Buker s.n.
Orobanchaceae
Conopholis americana (L.) Wallr. f, 2006 L. Speedy and M.
Bowers LS-06-270
Epifagus virginiana (L.) W. Bart., 2006 L. Speedy LS-06-971
Orobanche uniflora L., 2006 L. Speedy LS-06-109
f Texas 2(2)
Oxalidaceae
Oxalis dillenii Jacq., 1970 L.K. Henry s.n.
Oxalis grandis Small, 1951 L.K. Henry and W.E. Buker s.n.
Oxalis montana Raf., 1886 J.A. Shafer s.n
Oxalis stricta L., 2007 L. Speedy 07-1698
Oxalis violacea L., 2007 L. Speedy and M. Bowers LS-07-245
Papaveraceae
Chelidonium majus L. var. majus, 1981 FH. Utech 81-142
Eurasia
Sanguinaria canadensis L., 2006 L. Speedy LS-06-114
Passiflor
Passiflora e L., 1969 W.E. Buker s.n. G5:S1
Phytolaccaceae
Phytolacca americana L. var. americana, 2007 L. Speedy
699
Plantagin
Plantago piam L., 2004 E. Zimmerman 285 Europe
Plantago major L, 1942 O.E. Jennings s.n.
Plantago rugelii Dcne. var. rugelii, 2006 L. cl LS-06-980
Plantago virginica L., 1938 WR. Witz
Platanaceae
Platanus occidentalis L., 2007 L. Speedy 07-1050-1
Poaceae
Agrostis capillaris L., 1946 RW. Pohl 5954 Europe, PH
Agrostis gigantea Roth, 2006 L. Speedy and K. Jurkovic LS-
06-683 Europe
Agrostis perennans (Walt.) Tuckerman, 2006 L. Speedy LS-
06-1010
Agrostis scabra Willd. 2006 L. Speedy LS-06-985
Andropogon ~ Vitman, 2006 L. Speedy and D. Sofran
-06-
Andropogon Tan (Walt.) B.S.P. var. glomeratus, 2006
L. Speedy and T. Pearson and D. Bryant LS-06-762 G5:S3
Andropogon virginicus L. var. virginicus, 2006 L. Speedy LS-
6
6-9
Anthoxanthum odoratum L. ssp. odoratum, 2007 L. Speedy
LS-07-406 Eurasia
Aristida dichotoma Michx. var. dichotoma, 1906 O.E. Jennings
and GK. J
armenia elatius (L) Beauv. ex J. & K. Presl var. elatius,
8 tech 80-153a Furope
mn xo (Thunb.) Makino, 2006 L. Speedy and D.
Sofran LS-06-931 Southeast Asia
g anadensis, 2006
L. Speedy LS-06-452
Mercede coarctata ee Eat, 2006 L. Speedy and T.
and D. Bryant L
Pea
Chasmanbhium latifolium Td Yates, 2006 L. Speedy LS-
06-631
Cinna daa L., 2006 L. Speedy LS-06-975
Brachyelytrum erectum (Schreb. ex Spreng.) Beauv., 2007 L.
-615
Brachyelytrum septentrionale (Babel) G. Tucker, 2006 L. Speedy
Bowers LS-06-861
Bromus commutatus Schrad., 1951 W.E. Buker s.n. Europe
Morton and Speedy, Vascular plants of Fayette County, Pennsylvania
Bromus hordeaceus L. ssp. hordeaceus, 1995 B.L. Isaac and J.A.
Isaac 7568 Europe
Bromus inermis Leyss. ssp. inermis var. inermis, 2004 E. Zim-
merman Europe
Bromus pubescens Muhl. ex Willd., 2006 L. Speedy and M.
L 2
Dactylis glomerata L. ssp. glomerata, 1987 FH. Utech 87-130
Europe
Danthonia compressa Austin ex Peck, 2006 L. Speedy LS-
06-558
Danthonia spicata (L) Beauv. ex Roemer & J.A. Schultes, 2006
LS-06-554
Deschampsia flexuosa (L.) Trin. var. flexuosa, 1938 OE. Jen-
nings s.n
Diarrhena americana Beauv., 2007 L. Speedy 07-1666
4751
G
eds acuminatum (Sw.) Gould & C.A. Clark var. fas-
culatum (Torr) Freckmann, 1934 OE. Jennings s.n.
LO. boreale (Nash) Freckmann, 1965 W.E. Buker
Dichanthelium boscii (Poir. Gould & C.A. Clark, 1995 B.L. Isaac
J.A. Isaac 7547
s clandestinum (L.) Gould, 2006 L. Speedy LS-
Bc m commutatum (J.A. Schultes) Gould, 1931 J.
Bright 5459
doin depauperatum (Muhl.) Gould, 1942 O.E. Jen-
ings s.n.
PI a .) Gould var. dichotomum, 2006
ae ition (L) Gould €: CA. Clark, 1947 O.E.
ings s
Diana per (Scribn. ex Nash) Gould, 1934
S Sh.
DN e ek meridionale (Ashe) Freckmann, 1955 W.E.
uker s.n.
m sabulorum (Lam) Gould & C.A Clark var. thinium
( c. & Chase) Gould & C.A. Clark, 2006 L. Speedy
E
Dichanthelium sphaerocarpon (Ell) Gould var. ee
(Scribn.) Gould & CA. Clark, 1953 W.E.
Dichanthelium sphaerocarpon (Ell.) Gould var. CODE
1931 J. Bright 5421
Digitaria ceived (Schreb.) Schreb. ex Muhl., 1995 Isaac,
J.A. 8396 Eurasia
eee Kr (L) Scop., 1997 Grund, S.P. 2010
— crus-galli (L.) Beauv., 2004 E. Zimmerman 287
Ts muricata (Beauv.) Fern. var. muricata, 1950 L.K.
Henry and W.E. Buker s.n.
Eleusine indica (L.) Gaertn.,
Tropics
Elymus hystrix L. var. hystrix, 2006 L. Speedy LS- 06- 618
Elymus repens (L.) Gould, 1928 J. Bright s.n. Eur.
Elymus riparius Wieg., 2006 L. Speedy and D. pon LS-06-
732
1915 J. Bright s.n. Old World
Elymus virginicus L. var. virginicus, 2007 L. Speedy and J.
Nusser 07-607
1467
Radios capillaris (L) Nees, 1924 J. Bright s.n
ragrostis hypnoides (Lam.) B.S.P, 2006 L. e LS-06-967
EU pectinacea (Michx.) Nees ex Steud. var. pectinacea,
915 J. Bright s.n.
Eragrostis spectabilis (Pursch.) Steud, 2006 L. Speedy LS-06-
7
Festuca subverticillata (Pers) Alexeev, 2007 L. Speedy and J.
7-597
Glyceria canadensis (Michx.) Trin., 2006 L. Speedy and K.
Jurkovic LS-06-712
Glyceria grandis S. Wats. var. grandis, 2006 L. Speedy LS-06-
436
Glyceria melicaria (Michx.) F.T. Hubbard, 2006 Speedy, L
LS-06-1004
Glyceria septentrionalis A.S. Hitchc., 1905 J.A. Shafer and G.E.
inzer s.n.
pese striata (Lam.) A.S. Hitchc., 2007 L. Vend 07-549
us lanatus L., 2006 L. Speedy LS-06-426 Eur
pes oryzoides (L) Sw., 1982 EH. Utech and M Ohara 82-
393
d ed Willd., ud " d 07-1636
S.J. Darbyshire, 1995 B.L. Isaac
and J.A. Isaac 7582 enda
Lolium perenne L. ssp. perenne, 1907 J.A. Shafer s.n. Europe
Lolium pratense (Huds.) SJ. Darbyshire, 2007 L. Speedy 07-
1715 Europe
Microstegium vimineum (Trin.) A. Camus, 2007 L. Speedy 07-
1807 Iropical Asia
Muhlenbergia frondosa (Poir) Fern., 1951 L.K. Henry and W.E.
Muhlenbergia schreberi J.F. Gmel., 1901 J.A. Shafer s.n.
Muhlenbergia sylvatica Torr. ex Gray, 2007 L. Speedy 07-1717
"oo pud (Willd.) B.S.P, 2006 L. Speedy and J.
E a id dum H.N. Mozingo s.n
Panicum dichotomiflorum Michx. var. E 2007
eedy and et al. 07-1605
Pünicum E hit 1916 OE. Jennings and G.K. Jennings
and B
T a Bernh. ex Trin,, 1919 O.E. Jennings
a fone Bosc ex Nees var. elongatum (Pursh) Le-
lo 1 OE. Jennings s.n.
Panicum ruda de ex 2 var. pubescens (Vasey) Lelong,
198 Cusick 26
Panicum ae don ex a var. rigidulum, 1929 J. Bright
2352
Panicum verrucosum Muhl., 2006 L. Speedy and D. Sofran
-06-
Panicum virgatum L. var. virgatum, 2006 L. Speedy LS-06-
988
Pennisetum glaucum (L.) R. Br, 1970 L.K. Henry s.n. Africa
Phalaris arundinacea L., 2006 L. Speedy s BSWP LS-06-749
Phleum pratense L., 1930 J. Bright 5042 Eur
Poa alsodes Gray, 2006 L. Speedy and K. mods LS-06-472
Poa annua L., 1942 OE. Jennings s.n. Eurasia
Poa compressa L., 2007 L. Speedy and M. Bowers 07-686
Europe
1468
Poa cuspidata Nutt., 2006 L. Speedy LS-06-11
Poa palustris L., 2006 L. Speedy and BSWP LS-06-751
Poa pratensis L. ssp. pratensis, 2007 L. Speedy LS-07-399
Poa m Fern. & Wieg., 2007 L. Speedy and J. Nusser
-602
Poa P Gray, 1995 B.L. Isaac and J.A. c 7089
Poa trivialis L., 2007 L. Speedy 07-546 Euro
Schizachyrium scoparium (Michx.) Nash var. DE 1982
FH. Utech and M. Ohara 82-394
Setaria faberi Herrm., 1970 W.E. Buker s.n. Eastern Asia
Setaria pumila (Poir.) Roemer & J.A. Schultes, 2006 L. Speedy
-06-
m Bn ) Beauv. var. viridis, 2006 L. Speedy LS-06-958
uid nutans (L.) Nash, 2006 L. Speedy LS-06-959
Spartina pectinata Bosc ex Link, 2006 L. Speedy and BSWP
LS-06-754
Sphenopholis nitida (Biehler) Scribn., 1920 J. Bright s.n.
Tridens flavus (L) A.S. Hitchc. var. flavus, 2007 L. Speedy 07-
1066
Polemoniaceae
Phlox divaricata L. ssp. divaricata, 1987 F.H. Utech 87-144
Phlox divaricata L. ssp. laphamii (Wood) Wherry, 2006 L.
Speedy LS-06-179
Phlox maculata L. ssp. maculata, 1981 F.H. Utech 81-119
Phlox maculata L. ssp. pyramidalis (Sm.) Wherry, 1951 L.K.
n
Phlox paniculata L., 2006 L. Speedy LS-06-630
Phlox stolonifera Sims, 2006 L. Speedy LS-06-14
MEN reptans L. var. reptans, 2007 L. m LS-07-
19
Polygalaceae
Polygala ambigua Nutt., 2000 R.B. Coxe
Polygala cruciata L. var. aquilonia E p Schub., 2006 L.
dy and T. Pearson and D. Bryant LS-06-778 G5:S1
Polygala Bus Willd., 1905 O.E. Jennings s
Polygala sanguinea L., 2006 L. Speedy and 5 Jute LS-
06-689
Polygala verticillata L. var. verticillata, 1954 W.E. Buker s.n.
Polygona
Fagopyrum Nc EE Moench, 1942 O.F. Jennings s.n.
Asia
idee sl (L) A. Love, 1982 FH. Utech and M. Ohara
397 Europe
EL aoe (Houtt.) R. Decr., 2007 L. Speedy 07-1693
Japan
Fallopia sachalinensis (F. Schmidt ex Maxim.) R. Decr., 2006 L.
Speedy LS-06-989 Japa
Fallopia scandens (L.) Holub, 2007 L. Speedy 07-1701
desata amphibia var. emersa (Michx.) Hickman, 2006 L.
and BSWP LS-06-750 i hydropiper (L.)
O ie O.E. Jennings s
Persicaria hydropiperoides Mido, pam 2007 L. Speedy
07-1635
Persicaria lapathifolia (L.) S.F. Gray, 1999 S.P. Grund and et
al. 2341
f Texas 2(2)
Persicaria longiseta (de Bruijn) Moldenke, 2006 L. Speedy
LS-06-353
Persicaria maculosa S.F. Gray, 1999 R.B. Coxe s.n. Europe
Persicaria nepalensis (Meisn.) H. Gross, 1997 A.L. Woolston
3062 Asia
Persicaria orientalis (L.) Assenov, 1944 OE. Jennings s.n. India
Persicaria pensylvanica (L.) Small, 2006 L. Speedy LS-06-844
Persicaria punctatum (Ell) Small, 2007 L. pan 07-1734
Persicaria sagittatum (L.) Gross, 2000 R.B. Cox
Persicaria M iuis E! anii 2007 L. a 07-1075
Poly 1 S.A. d 8891 Furope
Prid. cilinode mus 1999 R.B. Coxe
Rumex acetosella L., 2006 L. Speedy LS-06- D Eurasia
Rumex crispus L. ssp. crispus, 1991 S.A. Thompson 8906
urope
Rumex obtusifolius L., 2004 E.A. Zimmerman 214 Europe
Portulac
debo caroliniana Michx. var. caroliniana, 2006 L. Speedy
LS-06-1
Claytonia a L. var. virginica, 2006 L. Speedy LS-06-48
Portulaca grandiflora Hook., 1900 J.A. Shafer s.n. Argentina
Potamogetonac
sd REDE Tuckerman, 1999 S.P. Grund and
et al.2
ronca crispus L., 2006 L. Speedy and BSWP LS-06-759
Eo x diversifolius Raf. var. diversifolius, 1997 S.P.
Grund 2016
Potamogeton 1977 M. Polechko s.n.
Potamogeton foliosus Pot A foliosus, 1977 M. Polechko s.n.
Potamogeton nodosus Poir., 1977 M. Polechko s.n
Potamogeton tennesseensis Fern., 2006 L. Speedy and BSWP
LS-06-758 G2:51
Primulaceae
Anagallis arvensis L. ssp. arvensis, 1901 J.A. Shafer s.n. Eurasia
Lysimachia ciliata L., 2006 L. Speedy and M. Bowers LS-06-
523
Lysimachia lanceolata Walt., 1959 OE. Jennings s.n.
Lysimachia nummularia L., 2004 E. Zimmerman 286 Europe
Lysimachia quadrifolia L., 2007 L. Speedy and M. Bowers
07-664
7-6
Lysimachia xproducta (Gray) Fern. (pro sp), 1916 B.H. Pat-
terson s.n.
Lysimachia terrestris (L.) B.S.P, 2006 L. Speedy and M. Bowers
S-06-869
Pyrolaceae
Pyrola americana Sweet, 1976 W.E. Buker s.n
Pyrola elliptica Nutt., 1905 J.A. Shafer and G.E. Kinzer s.n.
Ranunculace
Aconitum on Gray, 1999 S.P. Grund and R. Coxe and
E. Copeland 2368 G3:S1
Aconitum uncinatum L. ssp. muticum e Hardin, 2006 L.
Speedy and BSWP LS-06-752 G4:S
dla pachypoda Ell., 1932 WR. Van m s.n.
Actaea podocarpa DC., 2006 L. Speedy LS06-996-1
f
Pennsylvania
y,
L. var. racemosa, 2006 L. Speedy and M. Bow-
ers LS-05-511 [Cimicifuga raceraosa (L.) Nutt]
dies ee L. var. quinquefolia, 2006 L. Speedy
LS-
mos virginiana L. var. virginiana, 2006 L. Speedy LS-
06-570
Aquilegia canadensis L., 1987 FH. Utech 87-173
Aquilegia vulgaris L., 1962 W.E. Buker s.n. E
Caltha palustris L. var. palustris, 2007 L. Speedy B 07-115
Clematis virginiana L., 2007 L. Speedy 07-10
Delphinium exaltatum Ait, 1994 J.D. TAA e CJ. Boget
Delphinium tricorne Michx., 2006 L. Speedy LS-06-100
Hepatica nobilis Schreber var. acuta (Pursh) Steyermark, 2006
L. Speedy
Hepatica nobilis Schreber var. obtusa (Pursh) Steyermark,
1 .K Henry s.n
Hydrastis canadensis L., 2007 L. Speedy LS-07- Yi
Ranunculus abortivus L., 2006 L. Speedy LS-06-1
jste acris L. var. acris, 2006 L. sid in 06-350
Eur
w allegheniensis Britt. 1999 R.B. Coxe s.n.
Ranunculus hispidus Michx. var. caricetorum (Greene) T. Dun-
n, 1920 OE. Jennings s.n.
Ranunculus hispidus Michx. var. hispidus, 2006 L. Speedy
-06-4C
Ranunculus hispidus Michx. var. nitidus (Chapman) T. Duncan,
1957 L.K. Henry s.n., PH
Ranunculus micranthus Nutt, 1997 S.P. Grund 1789
Ranunculus recurvatus Poir. var. recurvatus, 2006 L. Speedy
LS-06-174
Ranunculus repens L., 1999 R.B. Coxe s.n. Europe
Thalictrum coriaceum (Britt.) Smal:, 2005 C.M. Bowers s.n.
G4:52
Thalictrum dioicum L., 2007 L. Speedy LS-07-270
Thalictrum pubescens Pursh, 2006 L. Speedy and M. Bowers
LS-06-522
Thalictrum thalictroides (L) Eames & Boivin, 2006 L. Speedy
LS-06-10
Trautvetteria caroliniensis (Walt. Vail var. caroliniensis, 2007 L.
peedy 6 G5:
Rhamnaceae
Ceanothus americanus L., 1985 C.W. Bier s.n.
Frangula alnus P. Mill., 2007 L. Speedy 07-1705 Europe
Rosaceae
Agrimonia gryposepala Wallr., 2006 L. Speedy and J. Nusser
-06-582
Agrimonia parviflora Ait., 2007 L. Speedy 07-1025
Agrimonia pubescens Wallr, 1957 L.K. Henry s.n.
Agrimonia rostellata Wallr., 1953 W.E. Buker s.n.
Agrimonia striata Michx., 1999 RB. Coxe s.n.
Amelanchier arborea (Michx. f.) Fern. var. arborea, 2006 L.
Speed 9
Amelanchier canadensis (L) Medik., 1908 OE. Jennings s.n.
:SNR
Amelanchier xintermedia Spach (pro sp), 1963 OE. Jennings
1469
Amelanchier laevis Wieg., 2006 L. Speedy LS-06-130
Amelanchier stolonifera Wieg., 2007 L. Speedy 07-136
Aruncus dioicus (Walt) Fern. var. dioicus, 2007 L. Speedy
07-1660
Crataegus dodgei Ashe, 1920 OE. Jennings s.n.
Cra ud macrosperma Ashe, 2006 L. Speedy LS-06-1017
Crataegus phaenopyrum (L. f) Medik., 1946 J. Lewis s.n
Cotes puni (Wendle.f) K. Koch, 1902 W.M. ey and
C. ent and B.F. Bush 38
DUO repens L., 1954 W.E. Buker s.n.
Duchesnea indica (Andr) Focke, 2006 L. Speedy LS-06-953
Asia
Fragaria vesca L. ssp. vesca, 1955 L.K. Henry s.
ko a Duchesne ssp. virginiana, i L. Speedy
We ANM Jacq. var. canadense, 2007 L. Speedy and
J. Nusser 07-599
Geum vernum (Raf) Torr. & Gray, 1920 O.E. Jennings s.n.
Geum virginianum L., 1934 J. Bright 9651
Malus coronaria (L.) P. Mill., 2006 L. Speedy LS-06-165
Malus floribunda Sieb. ex Van Houtte, 2007 L. Speedy 07-
1463 Asia
Malus pumila P. Mill, 1934 E.H. Graham s.n. [Pyrus malus L]
rasia
Photinia iie (Lindl) Robertson & Phipps, 1946 OLE.
ennings s.n. [Pyrus floribunda Lindl.]
Photinia tance (Michx.) Robertson & Phipps, 2006 L.
Spe LS-06-163 [Pyrus melanocarpa (Michx.) Willd]
tono E s (Lam.) Robertson & Phipps, 1922 O.E. Jen-
rus floribunda Lindl.]
Ned ires (L) Maxim. var. opulifolius, 2007 L.
-1
Porteranthus trifoliatus (L.) Britt, 2006 L. Speedy LS-06-453
Potentilla canadensis L. var. canadensis, 2006 L. Speedy LS-
06-159
Potentilla norvegica L. ssp. monspeliensis (L.) Aschers & Graebn,
B. N.
Potentilla recta L., 2007 L. Speedy LS-07-402 Europe
Potentilla simplex Michx., 2007 L. Speedy and M. Bowers
LS-07-246
Prunus americana Marsh. var. americana, 1954 L.K. Henry and
Prunus avium (L.) L., 2007 L. Speedy LS-07-17 Eurasia
Prunus pensylvanica L. f. var. pensylvanica, 2006 L. Speedy
$-06-145
Prunus persica (L.) Batsch, 2006 L. Speedy LS-06-99-1 Asia
Prunus serotina Ehrh. var. serotina, 2006 L. Speedy LS-06-202
Prunus virginiana L. var. virginiana, 2006 L. Speedy LS-06-
639
Rosa carolina L. var. carolina, 2007 L. Speedy and M. Bowers
7-690
Rosa eglanteria L., 1918 J. Bright s.n. Europe
Rosa multiflora Thunb. Ex Murr, 2007 L. Speedy 07-1646
Asia
Rosa palustris Marsh., 1999 R.B. Coxe s.n.
Rosa wichuraiana Crepin, 2006 L. Speedy and M. Bowers
$-06-530 Asia
Rubus adjacens Fern., 1904 OE. Jennings s.n.
1470
Rubus allegheniensis Porter var. allegheniensis, 2006 L. Speedy
S-06-401
Rubus canadensis L., 1929 J. Bright 1866
Rubus fecundus Bailey, 1951 H.A. Davis and T. Davis 9466
Rubus flagellaris Willd., 2007 L. Speedy and M. Bowers LS-
07-248
Rubus illecebrosus Focke, 1999 R.B. Coxe s.n. Japan
Rubus invisus (Bailey) Britt, 1950 H.A. Davis and T. Davis
9166
Rubus hispidus L., 2007 L. Speedy and Botanical Soc. of W
Pa 07-642
Rubus laudatus Berger, 1955 H.A. Davis and T. Davis 10905
Rubus occidentalis L., 2006 L. Speedy and K. Jurkovic LS-06-
476
Rubus odoratus L. var. odoratus, 2006 L. Speedy LS-06-368
Rubus paganus Bailey, 1950 H.A. Davis and T. Davis 9079
Rubus pensilvanicus Poir., 2006 L. Speedy LS-06-413
Rubus permixtus Blanch., 1950 H.A. Davis and T. ~ s.n.
Rubus plicatifolius Blanch., 1905 OE. Jennings s
Rubus provincialis Bailey, 1950 H.A. Davis and T. Dy vis
Rubus pubescens Raf. var. pubescens, 2006 L. Speedy cud E:
Richards LS-06-310
Rubus pugnax Bailey, 1950 H.A. Davis and T. Davis 9174
Rubus racemiger Bailey, 1951 H.A. Davis and T. Davis ~
Rubus recurvicaulis Blanch., 2006 L. Speedy LS-06-44
Rubus roribaccus (Bailey) Rydb., 1955 H.A. Davis and Davis
10903
Rubus rosa Bailey, 1924 J. Bright s.n.
Rubus setosus Bigelow, 1951 H.A. Davis and T. Davis 9473
G5:SH
Rubus suus Bailey, 1906 O | G.K. Jennings s.n
Sanguisorba canadensis L., E 940 O. E Jennings s.n.
Sorbus americana Marsh., 1946 O.E. Jennings s.n.
Spiraea in Du Roi var. alba, 2007 L. Speedy and M. Bowers
07-
Spiraea p Du Roi var. latifolia a x 1986 C. Bier and
B. Woodworth and M. Bowers
Spiraea japonica L. f. var. o — Rehd., 2006 L.
Speedy LS-06-998 Japan
Spiraea virginiana Britt., 1901 J.A. Shafer 343 G2:SX
Waldsteinia fragarioides (Michx.) Tratt. ssp. fragarioides, 1938
L.K. Henr
Rubiaceae
Cephalanthus occidentalis L, 1999 R.B. Coxe s.n.
Galium aparine L., 2006 L. Speedy LS-06-115
Galium asprellum Michx., 1991 S.A. Thompson 8901
Galium boreale L., 1904 O.E. Jennings s.n
Galium circaezans Michx. var. circaezans, 2006 L. Speedy
LS-06-617
Galium circaezans Michx. var. hypomalacum Fern., 1935 O.E.
nnings s.n
Galium lanceolatum Torr., 1950 W.E. Buker s.n.
Galium latifolium Michx., 2004 R. Coxe s.n. G5:53
Galium mollugo L., 2007 L. Speedy 07-1696 Eurasia
Galium pilosum Ait. var. pilosum, 1948 W.E. Buker s.n.
Galium tinctorium (L.) Scop., 2007 L. Speedy and M. Bowers
07-835
f Texas 2(2)
Galium trifidum L. ssp. trifidum, 1954 W.E. Buker s.n. G5:S2
Galium triflorum Michx., 1999 S.P. Grund and R. Coxe and E.
ae mud
L., 2006 L. Speedy LS-06-113
Houstonia purpurea HE var. purpurea, 2007 L. Speedy 07-625
G5T5:S1
Houstonia serpyllifolia Michx., 2004 R. Coxe s.n. G4?:S1
Mitchella repens L., 2006 L. Speedy LS-06-148
Salicaceae
Populus alba L., 1950 W.E. Buker s.n. Eurasia
m Xcanescens (Ait.) Sm. (pro sp), 1901 J.A. Shafer s.n.
E UE Bartr. Ex Marsh. ssp. deltoides, 1940 J.F.
ewis s.n
Populus grandidentata Michx. 1999 R.B. Coxe
Populus xjackii ( cv. Balm of Gilead) Sarg., 1953 à da Henry and
Populus tremuloides Michx., 1941 O.E. Jennings s.n.
Salix bebbiana Sarg., 1947 W.E. Buker s.n
Salix eriocephala Michx., 1999 R.B. Coxe s.n.
Salix fragilis L., 2007 L. Speedy 07-1067 Europe
Salix humilis Marsh. var. humilis, 1953 L.K. Henry and WE.
Salix interior Rowlee, 2007 L. Speedy and M. Bowers 07-
1608
Salix nigra Marsh., 2006 L. Speedy LS-06-361
Salix sericea Marsh., 2007 L. Speedy and M. Bowers 07-937
Salix serissima (Bailey) Fern., 1932 W.R. Van Dersal s.n. G4:S2
Santalaceae
Pyrularia pubera Michx., 2006 L. Speedy and M. Bowers LS-
25 G5:S3
Sarraceniacea
Sarracenia Hs L. ssp. gibbosa (Raf) Wherry, 2006 L.
Speedy and E. Richards LS-06-306
Saxifragaceae
Chrysosplenium americanum Schwein. ex Hook., 2006 L.
Speedy LS-06-154
Heuchera americana L. var. americana, 1987 F.H. Utech 87-
165
Heuchera pubescens Pursh, 1968 H. iu. ns et al. WP-1350
Mitella diphylla L., 2007 L. Speedy LS-07-
d o ed (Haw.) Steud, pue L. Speedy LS-
9 G5:S
"bd iis RM L, 2007 L. Speedy LS-07-379
Saxifraga virginiensis Michx. var. virginiensis, 2007 L. Speedy
LS-07-11
Tiarella cordifolia L. var. cordifolia, 2006 L. Speedy LS-06-138
Scrophulariaceae
Agalinis purpurea (L.) Pennell, 1929 J. Bright 2376
Aureolaria flava (L.) Farw. var. flava, 1947 L.K. Henry s.n.
Aureolaria laevigata (Raf.) Raf., 2006 L. md oo
Aureolaria virginica (L.) Pennell, 2004 R. Cox
Castilleja coccinea (L.) Spreng., 2007 L. ed LS-07-362
S2
G5:
Chaenorhinum minus (L.) Lange, 2006 L. Speedy and E. Rich-
ards LS-06-312 Europe
Morton and Speedy, Vascular plants of Fayette County, Pennsylvania
Chelone glabra L., 2006 L. Speedy LS-06-1012.
Collinsia verna Nutt., 2006 L. Speedy LS-06-170 G5:S4
Digitalis purpurea L. var. purpurea, 1905 J.A. Shafer and G.E.
Kinzer s.n. Europe
Gratiola neglecta Torr., 2006 L. Speedy LS-06-437
Linaria vulgaris P. Mill, 1981 EH. Utech 81-221 Eurasia
Lindernia dubia (L) Pennell var. dubia, 2007 L. Speedy 07-
1028
Melampyrum lineare Desr. var. lineare, 1947 W.E. Buker s.n.
Mimulus ringens L. var. ringens, 2007 L. Speedy and M. Bow-
-920
Pedicularis canadensis L. ssp. canadensis, 2006 L. Speedy
9
Pedicularis lanceolata Michx., 1903 O.P. Medsger s.n.
G5:S1S2
Penstemon digitalis Nutt. ex Sims, 1952 C.M. Boardman s.n
Scrophularia marilandica L., 2007 L. Speedy 07-1455
Verbascum blattaria L., 1999 R.B. Coxe s.n. Eurasia
Verbascum thapsus L., 1953 L.K. Henry and FH. Beer s.n.
Europe
Veronica americana Schwein. ex Benth., 2006 L. Speedy
6-351
Veronica chamaedrys L., 1920 J. Bright s.n. Eurasia
Veronica officinalis L. var. officinalis, 2006 L. Speedy LS-06-
Veronica peregrina L. ssp. peregrina, 1939 OE. Jennings s.n.
Veronica serpyllifolia L. ssp. serpyllifolia, 1932 W.R. VanDersal
s.n. Europe
Veronicastrum virginicum (L.) Farw., 2006 L. Speedy LS-06-
638
Simaroubaceae
Ailanthus altissima (P.Mill) Swingle, 2006 L. Speedy LS-06-
626 Asia
Smilacaceae
Smilax glauca Walt., 2006 L. Speedy LS-06-555
Smilax herbacea L., 2007 L. Speedy 07-1030
Smilax rotundifolia L., 2006 L. Speedy LS-06-203
Smilax tamnoides L., 1934 EH. Graham s.n.
Solanaceae
Datura stramonium L., 1939 W.G. Burig s.n. exotic Mexico
oe ~ L., 1950 L.K. Henry and W.E. Buker s.n.
a i Nees var. heterophylla, 1953 L.K. Henry
Physalis longifolia Nutt. var. subglabrata (Mackenzie & Bush)
Cronq., L dy 0
Solanum carolinense L. var. carolinense, 1955 C.M. Boardman
Solanum dulcamara L. var. dulcamara, 2007 L. Speedy 07-
1708
Solanum ptychanthum Dunal, 1900 J.A. Shafer s.n.
Sparganiaceae
CHharnnnirim
, 2006 L. Speedy and T. Pearson
and D. Bryant LS-06-788
Sparganium angustifolium Michx., 1908 O.E. Jennings s.n.
G5:S2
1471
Sparganium erectum L. ssp. stoloniferum (Graebn.) Hara, 1999
R.B. Coxe s.n.
Staphyleaceae
Staphylea trifolia L., 2006 L. Speedy LS-06-105
Thymelaeaceae
Dirca palustris L., uud L. Speedy and Botanical Society of
Western PA 07-1
Tiliaceae
Tilia americana L. var. americana, 2006 L. Speedy LS-06-951
Tilia americana L. var. heterophylla (Vent) Loud., 2006
e L$-06-371
Tilia cordata Tilia cordata P. Mill, 1934 A. Conn s.n. exotic
Europe
Typhaceae
Typha latifolia L., 1991 S.A. Thompson 8888
Ulmaceae
Celtis occidentalis L., 2007 L. Speedy 07-1822
Celtis tenuifolia Nutt., 1950 W.E. Buker s.n
Ulmus americana L., 2007 L. Speedy 07-1638
Ulmus rubra Muhl., 2006 L. Speedy LS-06-63
Urticaceae
Boehmeria cylindrica (L.) Sw., 1951 L.K. Henry and W.E. Buker
s.n.
a o (L) Weddell, 2006 L. Speedy and J.
Nusser LS-0
E. venant Muhl. ex Willd., 1970 L.K. Henry s.n.
Pilea pumila (L) Gray var. pumila, 2007 L. Speedy 07-1821
Urtica ris L. ssp. gracilis (Ait.) Seland., 1946 H.N. Mozingo s.n.
Valerian
Valeriana man Michx,, 2006 L. Speedy LS-06-181
Valerianella chenopodiifolia (Pursh) DC., 1950 LK. Henry and
Valerianella umbilicata (Sullivant) Wood, 1987 FH. Utech
7-146
Verbenaceae
Phryma leptostachya L., 2006 L. Speedy LS-06-616
bbs hastata L. var. hastata, 2007 L. Speedy and M. Bow-
-862
e simplex Lehm., 1948 W.F. Buker
Verbena urticifolia L. var. urticifolia, ia S.A. Thompson
1875
Violaceae
Hybanthus concolor (T.F. Forst.) Spreng., 2006 L. Speedy
LS-06-619
Viola affinis Le Conte, 1954 L.K. Henry and W.E. Buker s.n.
Viola bicolor Pursh, 1950 L.K. Henry and F.H. Beer s.n.
Viola xbissellii House, 1955 L.K. Henry s.n.
Viola blanda Willd. var. blanda, 2006 L. Speedy LS-06-132
Viola xbrauniae Grover ex Cooperrider, 1920 O.E. Jennings s.n
Viola canadensis L. var. canadensis, 2006 L. Speedy LS-06-193
Viola cucullata Ait., 2006 L. Speedy LS-06-135
Viola xeclipes H.E. Ballard, 1942 O.E. Jennings s.n.
Viola hastata Michx., 2006 L. Speedy LS-06-131
Viola hirsutula Brainerd, 1947 W.E. Buker s.n.
1472
Viola labradorica Schrank, 2006 L. Speedy LS-06-157
Viola lanceolata L. ssp. lanceolata, 1988 R.S. Mulvihill and R.C.
| loskeyi Lloyd ss llens (B
2006 L. Speedy LS-06-167
Viola xmalteana House, 1961 L.K. Henry s.n.
Viola xpalmata L. (pro sp.), 1992 J.A. Isaac 3908
Viola xporteriana Pollard (pro sp), 1950 L.K. Henry and WE.
) M.S. Baker,
2 +
Buker s.n.
Viola xprimulifolia L. (pro sp.), 2007 L. Speedy and Botanical
Soc. of W Pa 07-648
Viola dei i var. pubescens, 2006 L. Speedy and D.
Sofran
"m os AN var. scabriuscula Schwein. ex Torr. & Gray,
2006 L S-06-23
Viola xredacta House, 1953 L.K. Henry and W.E. Buker s.n.
Viola rostrata Pursh, 2006 L. Speedy LS-06-70
Viola rotundifolia Michx., 2006 L. Speedy LS-06-94
Viola sagittata Ait. var. ovata (Nutt) Torr. & Gray, 2006 L.
peedy LS-06-208
Viola sagittata Ait. var. sagittata, 1956 W.E. Buker s.n.
Viola sororia Willd., 2006 L. Speedy LS-06-42
Viola striata Ait., 2006 L. Speedy LS-06-178
Viola tripartita Ell, 1920 O.E. Jennings and G.K. Jennings and
et. al. s.n. G5:SH
Vitaceae
Parthenocissus quinquefolia (L.) Planch., 2007 L. Speedy
07-1063
Vitis Ae vu var. bicolor Deam, 2007 L. Speedy and
M. Bowers 07-670
Vitis cinerea nee Millard var. baileyana (Munson)
Comeaux, 1934 EH. Graham s.n. G4G5TNR:SH
Vitis labrusca L., 2006 L. Speedy and K. Jurkovic LS-06-466
Vitis xnovae-angliae Fern. (pro sp), 1942 OE. Jennings s.n.
1
iparia Michx, 1942 O.E. Jennings s.n.
Vitis — anon 2001 S.P. Grund and L. Smith 2562
Vitis oe L. 2007 L. Speedy and M. Bowers 07-672
Xyridac
Xyris torta uo 2006 L. Speedy and T. Pearson and D. Bryant
LS-06-780 G5:S1
GYMNOSPERMS
Cupressa
Thuja E di L, 1985 S.A. Thompson and M. Stieber
2224
Pinaceae
Abies balsamea (L.) P. Mill, 1998 P. Ludrosky 20-11-98, CUP
Larix decidua P. Mill, 1999 S.P. Grund 2140 Europe
Pinus resinosa Ait., 1966 A. Conn s.n.
Pinus rigida P. Mill, 2006 L. Speedy LS06-449-1
Pinus strobus L., 2006 L. Speedy and J. Nusser LS06-578-1
Pinus sylvestris L., 1985 K. Peters 22-7E-86 Eurasia, CUP
Pinus virginiana P. Mill, 1941 O.E. Jennings s.n.
Tsuga canadensis (L.) Carr., 2006 L. Speedy LS-06-402
f Texas 2(2)
Taxaceae
Taxus canadensis Marsh., 1936 W. McLean s.n. G5:S3S4
LYCOPHYTES
Isoétaceae
Isoétes engelmannii A. Braun, 1933 W.R/ Van Dersal s.n.
Lycopodiac
Huperzia ane (Michx) Trevisan, 2006 L. Speedy LS-06-
993
Lycopodiella SA (L.) Holub, 2006 L. Speedy and E.
peu rds L
im clavat 6 L. Speedy LS-06-414
pane Parodi ets 1963 OE. Jennings s.n.
M ric digitatum Dill. ex A. Braun, 2006 L. Speedy and
K. Jurkovic 60
ai nes yi W.H. Wagner, Beitel & Moran, 2006 L.
Speedy and K. Jurkovic LS-06-692
Lycopodium obscurum L., 2006 L. Speedy and K. Jurkovic.
-06-462
Lycopodium tristachyum Pursh, 1946 H.N. Mozingo s.n.
Selaginellaceae
Selaginella apoda (L.) Spring, 1937 OE. Jennings s.n.
EQUISETOPHYTES
Equisetaceae
Equisetum arvense L., 2006 1. Speedy LS-06-64
Equisetum hyemale L. var. affine (Engelm.) A.A. Eat., 2007 L.
Speedy 07-1724
POLYPODIOPHYTES
Aspleniace
Asp pu montanum Willd., 1995 S.P. Grund 1024
Asplenium pinnatifidum Nutt., 1995 S.P Grund 1025 G4:S3
Asplenium d (L.) B.S.P. var. platyneuron, 2006 L.
Spee -06-
Pub istnd L., 2006 L. Speedy LS-06-991
Asplenium ruta-muraria L. var. cryptolepis (Fern.) Wherry, 1934
Asplenium trichomanes L. ssp. trichomanes, 1948 D.H. Krouse
s.n.
Asplenium xtrudellii Wherry (pro sp), 1999 S.P. Grund and
et al. 2272
Dennstaedtia
ET Mp NE (Michx.) T. Moore, 2007 L. Speedy
m P" (L) Kuhn. var. latiusculum (Desv) Under-
ood ex Heller, 2006 L. Speedy LS-06-556
Dryopteridaceae
Athyrium filix- E (L) Roth ssp. ang
2006 L. Speedy LS-06-1005
a filix-femina (L.) Roth ssp
K. Henry s.n.
owner bulbifera (0) Bein ie JA. Isaac 3918
lell, 2007 L. Speedy and
(Willd.) Clausen,
f DD {RAL Yul
ave
J. EN 07- 604
Mart J SI ly, Wy. I r| 4 £F. ben f 4 , Pennsylvania 1473
Cystopteris tenuis (Michx) Desv., 2006 L. Speedy LS-06-992 Botrychium virginianum (L.) Sw., 2007 L. Speedy and J. Nusser
Deparia acrostichoides (Sw.) M. Kato, 2006 L. Speedy LS-06- 07-598
999 Ophioglossum vulgatum L. 2004 J. Love s.n. G5:5354
Diplazium pycnocarpon (Spreng.) Broun, 2006 L. Speedy and Osmundacaae
A OS Osmunda cinnamomea L. var. cinnamomea, 2006 L. Speedy
and M. Bowers LS-06-274
Dryopteris carthusiana (Vill) H.P. Fuchs, 2006 L. Speedy LS-
d . Osmunda claytoniana L., 2006 L. Speedy and M. Bowers
Dryopteris cristata (L.) Gray, 2007 L. Speedy 07-1658 L 7
Dryopteris goldiana (Hook. ex Goldie) Gray, 2006 L. Speedy =, P ne cH RACES 2007 L. Speedy
and J. Nuss 9 2 :
Var.
; LS-06-66 e ji : i
and Botanical Soc. of W Pa 07-636
ue intermedia (Muhl. ex Willd.) Gray, 2007 L. Speedy
Polypodiaceae
Dr xu. aac (L) Gray, 1999 R.B. Coxe s.n Polypodium appalachianum Haufler € Windham, 2006 L.
Dryopteris xtriploidea Wherry, 2006 L. Speedy LS06-9401 Speedy and J. Nusser L5-06-660
Gymnocarpium dryopteris (L) Newman, 1951 LK. Henry and Polypodium virginianum L, 2006 L. Speedy LS-06-365
.E. Buker s.n. Pteridac
Matteuccia struthiopteris (L.) Todaro, 2006 L. Speedy LS-06- Adiantum CEA L, 2007 L. Speedy LS-07-410
Pellaea atropurpurea (L) Link, 1994 J.A. Isaac 6070
Thelypteridac
Phegopteris MM (Michx.) Watt, 1965 L. Louden s.n.
Phegopteris hexagonoptera (Michx.) Fee, 2006 L. Speedy
362
Onoclea sensibilis L, 1999 R.B. Coxe s.n.
Dri Th cl Dd RAIL y
var. acrostichoides,
1995 B.L. Isaac and J.A. Isaac 7084
Woodsia obtusa (Spreng. Torr. ssp. obtusa, 1950 L.K. Henry
and W
-06-1
2
Thelypteris noveboracensis (L.) Nieuwl.,, 2007 L. Speedy 07-
Ophioglossaceae 6
Botrychium dissectum Spreng., 2006 L. Speedy LS-06-972 Thelypteris palustris Scl pubescens, 2007 L. Speedy and
Botrychium lanceolatum (Gmel.) ela var. eMe M. Bowers 07-846
tum Pease & Moore, 2003 E.A. Zimm
Botrychiurn matricarifolium (A.Braun ex "d A. Braun ex
Koch, 1949 W.E. Buker s.n
ACKNOWLEDGMENTS
Our thanks go to the Wild Resource Conservation Fund of the Pennsylvania Department of Conservation
and Natural Resources for partial funding of the field work. We would also like to thank Ann F. Rhoads and
Derick B. Poindexter for reviewing the manuscript. We are also thankful to Amanda Juretic for data entry.
For field work assistance, we thank Mark Bowers, Deb Bryant and Tom Pearson, Karen Jurkovic, Darlene
Madarish, James Nusser, Dennis Sofran, Pete Woods and Ephraim Zimmerman. Landowners who provided
permission to explore their property include the Young Family, Richard Dennis, Lillian Gwosden and the
Municipal Authority of Westmoreland County.
REFERENCES
BRUMMITT, R.K. AND C.E. PoweLL, 1992. Authors of plant names. Royal Botanic Gardens, Kew.
DCNR. 2004. Invasive plants of Pennsylvania. http//www.dcnr.state.pa.us/forestry/ wildplant/invasive.aspx
Coxe, R., C.M. Morton, M.J. HAvwooD, B.L. Isaac AND J.A. Isaac, 2007. Checklist of the vascular plants of Greene County,
Pennsylvania. Sida 21:1829-1859,
FLORA OF NORTH ÁMERICA EDITORIAL COMMITTEE, EDS. 1993+. Flora of North America north of Mexico. 12+ vols. New
York and Oxford.
KARTESZ, J.T. 1999, A synonomized checklist and atlas with biological attril for the vascular flora of the United
States, Canada and Greenland. First ed. In: Kartesz, J.T. and C A. Meacham. Synthesis of the North American
Flora, Version 1.0. North Carolina Botanical Garden, Chapel Hill.
Koras, RA. 1991. Soil survey of Fayette County Pennsylvania. US Department of Agriculture. Soil Conservation
Service. Washington, D.C.
1474 J lof the Botanical R h Institute of Texas 2(2)
MoRTON, C.M. AND L. Speepy. 2007. Checklist of the vascular plants of Washington County, Pennsylvania. J. Bot. Res
Inst. Texas 1:1229-1249.
Morton, C.M., L. SPEEDY, AND J.K. BisseLL. 2007. Checklist of the vascular plants of Crawford County, Pennsylvania. J.
Bot. Res. inst. Texas 1:631—653.
PALMER, T. 1984. Youghiogheny-Appalachian River. University of Pittsburgh Press, Pittsburgh, PA pp.337.
PENNSYLVANIA DEPARTMENT OF AGRICULTURE. 2007. Pennsylvania noxious weed control list. http://www.agriculture state.
pa.us/agriculture/Cwp/view.asp?q-1 27683
PENNSYLVANIA DEPARTMENT OF ENVIRONMENTAL PROTECTION WATERSHED NoteBook. 2006 http://www.dep.state.pa.us/dep/
deputate/ watermgt/WC
RHOADS, A.F. AND T.A. BLock. 2007. The plants of Pennsylvania, an illustrated manual, second edition. University of
Pennsylvania Press, Philadelphia, Pennsylvania.
THe Nature CONSERVANCY (1996). Element ranking list—Pennsylvania Natural Heritage Program. http://www.natu-
ralheritage.state.pa.us/PlantsPage.aspx
U.S. Census Bureau. 2007. State and county quickfacts. http//quickfacts.census.gov/qfd/states/ 42/42125.html
RARE AND ENDEMIC PLANTS OF THE BIG THICKET:
A PHYTOGEOGRAPHICAL ANALYSIS
Michael H. MacRoberts and Barbara R. MacRoberts
Bog Research, 740 Columbia
Shreveport, Louisiana 71104, U.S.A. and
Herbarium, Museum of Life Sciences
Louisiana State University in Shreveport
Shreveport, Louisiana 71115, U.S.A.
ABSTRACT
f + T 1 Tie seh et iAL f T, 3 1 1 f,
he i i
[al ES O Ls r
Gulf Coastal Plain endemics. X 1s, bogs, wetland pine savannas, prairies, and 1 pt p
RESUMEN
El Big Thicket del sureste de Texas se sitúa relativamente alto con respecto a la riqueza de especies de plantas raras de Texas, y cerca
del máximo para los endemismos de la West Gulf Coastal Plain. Los arenales xéricos, pantanos, sabanas húmedas de pino, praderas, y
lugares vacios es donde están la mayoría de estas especies.
INTRODUCTION
As part of a continuing study of the biogeography of the Big Thicket region of southeastern Texas (Brown
et al. 2002, 2005, 2006a, 2006b, 2008; MacRoberts & MacRoberts 2004, 2007, 2008; MacRoberts et al.
2002c, 2007, in prep) in this paper we investigate Big Thicket rare and endemic plants.
METHODS
Using the Carr (2005) and Poole et al. (2007) annotated lists of rare Texas species and several sources for
state distribution (e.g., Turner et al. 2003; Carr 2005; Diggs et al. 2006; Poole et al. 2007), we mapped
rare species richness (defined as being the number of rare taxa per county) across Texas. The Carr (2005)
sample consists of 450 species and includes all G3/T3 and rarer taxa. We modified the Poole et al. (2007)
list to include only G2G3/T2T3 and rarer species, making that sample consist of 201 taxa. G and T ranks
are discussed in Poole et al. (2007). Using Kartesz and Meacham (2005) and the Flora of North America
(1993-2006), we mapped Big Thicket rare species across North America.
Using the MacRoberts et al. (2002b) list of West Gulf Coastal Plain endemics and a grid of 17 quadrats
of equal size measuring about 130 km on a side and data on plant distribution from Thomas and Allen
(1993-1998), Turner et al. (2003), and Diggs et al. (2006), we plotted endemic plant richness in the West
Gulf Coastal Plain.
We define the Big Thicket region as Newton, Jasper, Tyler, Hardin, Polk, San Jacinto, and northern
Liberty counties. A map showing the area constituting the West Gulf Coastal Plain is in MacRoberts et al.
(2002b, 2007) and Diggs et al. (2006).
RESULTS
Figure 1 shows the distribution of rare Texas species by county for the Carr (2005) list. Figure 2 shows the
distribution of rare Texas species by county for the modified Poole et al. (2007) list. In the Carr (2005) list,
the Big Thicket region has 31 rare species. The modified Poole et al. (2007) list has 10 rare species. Figure 3
shows the distribution of West Gulf Coastal Plain endemic species by region. The Big Thicket has 69 West
Gulf Coastal Plain plant endemics.
J. Bot. Res. Inst. Texas 2(2): 1475 — 1479. 2008
1476 J lof the Botanical R h Institute of Texas 2(2)
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DISCUSSION
In the Carr (2005) list, the Trans-Pecos region has by far the greatest number of rare species in the state,
followed by the eastern part of the Edwards Plateau, the Coastal-Bend, and southern Texas (see also Dia-
mond et al. 1997; Dobson et al. 1997). The Pineywoods region and specifically the eastern edge of the Big
Thicket (Newton and Jasper counties) follows these areas. None of the Big Thicket region rare species is
western. In the Carr oe list, all Big ood rare plants are either eastern, notably southeastern, (4596)
(e.g., Platanthera cha g , Xyris scabrifolia), or West Gulf Coastal Plain
endemies (55%) es g., Bartonia t , Rudbeckia sable Trillium pusillum var. texanum, Yucca cernua). The
communities in which Big Thicket region rare species occur are xeric sandylands (26%), bogs/wetland pine
savannas (2696), upland pineland (13%), barrens (13%), and baygalls (10%). The remaining 12% are divided
among mixed hardwood forest, swamp, and bottomland hardwood forest.
In the modified Poole et al. (2007) list, the same pattern occurs as in the Carr (2005) list except it is
less extreme. Nine of the Big Thicket rare species in this list are endemic to the West Gulf Coastal Plain
RA Dal + IRA Dak D Al J
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Fic. 2. Distrib uti f T ! by ty i g I lified Poole et al. (2007) list. The lack of f
and one is southeastern. In this list, rare Big Thicket species occur in baygalls, bogs, prairies, barrens, dry
uplands, and xeric sandylands.
The Big Thicket region is rich in West Gulf Coastal Plain endemics. It is second only to the region
just to its west centered on Montgomery, Walker, Houston, Leon, Madison, Grimes, Brazos, and Robertson
counties. The reason for this high incidence of West Gulf Coastal Plain endemics in the Big Thicket region
is the presence of xeric sandylands (MacRoberts et al. 2002a; MacRoberts & MacRoberts 2006; Diggs et al.
2006), notably at the Roy Larsen Sandylands Sanctuary (Matos & Rudolph 1985). Xeric sandylands account
for about 50% of West Gulf Coastal Plain endemics (MacRoberts et al. 2002b; MacRoberts & MacRoberts
2006) and about 59% of those in the Big Thicket region. Most of the remainder of the Big Thicket region
endemics (29%) are in bogs, wetland pine savannas, barrens, baygalls, and prairies. The higher frequency
of West Gulf Coastal Plain endemics just west of the Big Thicket is due to its slightly higher incidence of
xeric sandyland endemics combined with prairie, bog, and barren endemics.
The Big Thicket, therefore, fares well in Texas and the West Gulf Coastal Plain with respect to rare and
endemic species. It ranks in the top half for richness of rare Texas species and near the top for West Gulf
1478 J lof the Botanical R h Institute of Texas 2(2)
55
25 | 46 32
23 | 60 | 59 p
43 | 75 | 69 Y 44
Ed ES 50 Sa
34 nn
Fic. 3. West Gulf Coastal Plain endemics by quadrat.
Coastal Plain endemics, pinpointing important habitat for conservation, PRO xeric sandylands, bogs,
wetland pine savannas, prairies, and barrens. Since rare and i bject to change because
of taxonomic research, distributional discoveries, and other DOC these results are, of course, tentative
and future lists will undoubtedly provide changes in the results presented here.
ACKNOWLEDGMENTS
Amanda C. Lewis, Museum of Life Sciences, Louisiana State University in Shreveport, aided with the figures.
Jackie Poole provided many useful comments on the paper.
REFERENCES
BROWN, L.E., K. HiLL HOUSE, B.R. MacRoserts, AND M.H. MacRoserts. 2002. The floristics of Windham Prairie, Polk County,
Texas. Texas J. Sci. 54:227-240
Brown, L.E., B.R. MacRoserts, M.H. MacRoserts, PA. HARCOMBE, W.W. Pruess, I.S. ELsik, AND D. JoHNsoN. 2005. Annotated
checklist of the vascular flora of the Turkey Creek Unit of the Big Thicket National Preserve, Tyler and Hardin
counties, Texas. Sida 21:1807-1827.
Brown, L.E., B.R. MacRoserts, M.H. MacRoserts, PA. Harcomse, W.W. Pruess, l.S. ELsik, AND S.D. Jones. 2006a. Annotated
checklist of the vascular flora of the Big Sandy Creek Unit of the Big Thicket National Preserve, Polk County,
Texas. Sida 22:705-723.
Brown, L.E., B.R. MacRoserts, M.H. MacRoserts, PA. Harcomee, W.W. Pruess, 1.5. Esk, AND S.D. Jones. 2006b. Annotated
checklist of the vascular flora of the Lance Rosier Unit of the Big Thicket National Preserve, Hardin County,
Texas. Sida 22:1175-1189.
Brown, L.E., B.R. MacRoserts, M.H. MacRoserts, PA. HARCOMBE, W.W. Pruess, 1.5. ELsik, AND S.B. Waker. 2008 (in press).
Annotated checklist of the vascular flora of the Beech Creek Unit of the Big Thicket National Preserve, Tyler
County, Texas. J. Bot. Res. Inst. Texas -
Carr, W.R. 2005. An annotated list of G3/T3 and rarer plant taxa of Texas. Unpubl. report. The Nature Conservancy
of Texas, Austin, Texas.
DiamoND, D.D., C.D. True, AND K. He. 1997. Regional priorities for conservation of rare species in Texas. SouthW.
Naturalist 42:400—408.
MacRobert 1 MacRoberts, R 1 endemic plants of the Big Thicket 1479
Dices, G.M., B.L. Lirscoms, M.D. REED, AND RJ. O'KeNNoN. 2006. Illustrated flora of east Texas. Sida, Bot. Misc. 26:1-
Dosson, A.P, J.P. RobniuEz, W.M. Roserts, AND D.S. Witcove. 1997. Geographic distribution of endangered species in
the United States, Science 275: 550-553.
FLORA OF NORTH AMERICA EDITORIAL COMMITTEE (eds.). 1993-2006. Flora of North America North of Mexico. Vols. 1, 2, 3,
4, 5, 19, 20, 21, 22, 23, 24, 25, 26. Oxford Univ. Press, New York and Oxford.
KARTESZ, J.A. AND C.A. MEACHAM. 2005. Synthesis of North American flora. Version 2.0. North Carolina Botanical
Garden. Chapel Hill.
MacRoserts, B.R, M.H. MacRostnrs, AND J.C. CatHey. 2002a. Floristics of xeric sandylands in the Post Oak Savanna
region of east Texas. Sida 20:373-387.
MacRoserts, M.H., B.R. MacRogerTs, B.A. SORRE, AND R.E. Evans. 2002b. Endemism in the West Gulf Coastal Plain:
importance of xeric habitat. Sida 20:767-780.
MacRoserTs, B.R., M.H. MacRosERrs, AND L.E. Brown. 2002c. Annotated checklist of the vascular flora of the Hickory
Creek Unit of the Big Thicket National Preserve, Tyler County, Texas. Sida 20:781
MacRoserts, M.H. AND B.R. MacRoserts. 2004. The Big Td ple or ae eos Hist. Assoc. 42:42-51.
MacRosenrs, M.H. AND B.R. MacRoaenrs. 2006. Xeric t eric sandhill woodlands) of the West Gulf
Coastal Plain: a review. Unpublished report. Bon ans Natural Moda Program. Department of Wildlife and
Fisheries. Baton Rouge, Louisiana. 52 pp.
MachRosenrs, M.H., B.R. MACROBERTS, AND R.G. KaLinsky. 2007. Vascular plant species/area relationships (species rich-
ness) in the West Gulf Coastal Plain: A first approximation. J. Bot. Res. Inst. Texas 1:577-583.
MacRoserts, M.H. AND B.R. MacRoserts. 2007. Phytogeography of the Big Thicket, east Texas. J. Bot. Res. Inst. Texas
1:1149-1155.
MacRoserTs, M.H. AND B.R. MacRoserts. 2008. The Big Thicket of Texas as floristically unique habitat. J. Bot. Res. Inst
Texas 2:663-669.
MacRoserTs, M.H., B.R. MacRoserts, AND D.C. RupoLrH. In prep. Vertebrate richness and biogeography in the Big
Thicket of Texas. Texas J. Sci.
Matos, J.A. AND D.C. RuboteH. 1985. The vegetation of the Roy E. Larsen Sandylands Sanctuary in the Big Thicket
of Texas. Castanea 50:228-249
Poote, J.M., W.R. Carr, D.M. PRICE, AND J.R. SINGHURST. 2007. Rare plants of Texas. Texas A. & M. Univ. Press, College
Station.
Thomas, R.D. AND C.M. ALLEN. 1993-1998. Atlas of the vascular flora of Louisiana. 3 Vols. Louisiana Department of
Wildlife and Fisheries. Baton Rouge, Louisiana.
Turner, B.L, H. Nichols, G. Denny, AND O, Doron. 2003. Atlas of the vascular plants of Texas. Sida Bot. Misc. 24:
1-888.
1480 Journal of the Botanical R h Institute of Texas 2(2)
BOOK NOTICE
ZHENBIAO YANG (zp.). 2008. Annual Plant Reviews, Volume 33: Intracellular Signaling in Plants. (ISBN-13:
978-1-4051-6002-5, 10: 1-4051-6005-0, iva Mia Reo Publishing, 2121 State Avenue, Ames,
Iowa 50014-8300, U.S.A. (Orders: www. hing.com, orders@ames.blackwellpublishing.
com, 800-862-6657). $224.99, 430 p pp., 48 arena OM x 6".
Contents of Volume 33:
1. Transmembrane Rui in a po Kinases and their Ligands—Keiku U. Torii
2. Heteronomic G-protein er Plants—Lei Ding, Jin-Gui Chen, Alan M. Jones, and Saraah M Assmann
3. ROP/RACGTPases—Ying Fu, Duam B Ko Sane ies M
4. Mitogen-activated M Kinase Cascades in Plant g g PT Zhang
7Then.M; p, A Ci ni
5. Calcium Signals and T.
6. Paradigms and Networks for Siam in Plant Cells —Sheng Luan
7. Reactive Pun n o in Plants—Gad Miller, n. Coutu, Vladimir Shulaev, and Ron Mittler
. Lipid-Mediated Signaling — Wendy F. Boss, Daniel V. Lynch, and Xuem
9. The ms and Signal Transaction: Role and "m of e Actin- and Microtubule-Binding Proteins—Patrick J. Hussey
and Takashaki Hashimoto
10. PCI ] 1 the Ubiquitin Prot S (UBS) in Plant Development—Yair Halimi and Daniel A. Chamovitz
11. Si B ll ] Aurora P. Fernandez and Asa Strand
12. Signaling by Protein Phosshotylaion in Cell Senge aie Sasabe and Yasunori Machida
13. Guard Signaling—Yan Wu
14. The Molecular N ] Signaling—Zhizhong Gong, Viswanathan Chinnusamy, and Jian-Kang Zhu
Index
J. Bot. Res. Inst. Texas 2(2): 1480. 2008
ANNOTATED CHECKLIST OF THE VASCULAR FLORA OF THE LOBLOLLY UNIT
OF THE BIG THICKET NATIONAL PRESERVE, LIBERTY COUNTY, TEXAS
Barbara R. MacRoberts
Larry E. Brown and Michael H. MacRoberts
Herbarium, Spring Branch Science Center Bog Research, 740 Columbia
8. estview Drive Shreveport, Louisiana 71104, U.S.A.
Houston, lexas 77055, U.S.A. and Herbarium, Museum of Life Sciences
larry-theplantman@att.net Louisiana State University in Shreveport
Shreveport, Louisiana, 71115, U.S.A.
Warren W. Pruess and |. Sandra Elsik Suzanne Birmingham Walker
Department of Ecology and Azimuth Forestry Services, Inc
Evolutionary Biology 14671 State Highway 87 South
Rice University Shelbyville, Texas 75973 U.S.A.
Houston, Texas 77005, U.S.A.
ABSTRACT
An annotated, vouchered checklist is provided of the vascular plant taxa of the 222 hectare Loblolly Unit of the Big Thicket National
Preserve ir. Liberty County, Texas. Documented for the unit are 305 taxa including 34 (11%) exotics
RESUMEN
re] 1 A 1 1 14] 33>31 141 sedes EEdggTqa]
Reser Big Thicket National en el condado de liber Texas. Se P para la unidad 305 taxa incluyendo 34 (11%) exóticas.
INTRODUCTION
This paper is the 6th part of an inventory, based on available voucher specimens, of the vascular flora of the
Big Thicket National Preserve in southeastern Texas. In 5 previous papers, we provided annotated checklists
of the Hickory Creek, Turkey Creek, Big Sandy Creek, Lance Rosier, and Beech Creek units (MacRoberts et al.
2002; Brown et al. 2005, 2006a, 2006b, 2008). At this time, the Menard Creek Unit has been inventoried, and
a paper is anticipated in the near future. Here, we provide an annotated checklist of the Loblolly Unit.
SITE AND METHODS
The Big Thicket is located in the West Gulf Coastal Plain in southeastern Texas (see Diggs et al. 2006 for
literature and description). Peacock (1994) reported 12 units for the Big Thicket National Preserve. Since
then the Big Sandy Creek Corridor (1,798 hectares), the Canyonlands (350 hectares), and Village Creek
Corridor (2,059 hectares) have been added. The addition of these 3 units has added 4,207 hectares to the
34,000 reported by Peacock in 1994. However, Deanna Boensch, Big Thicket fire ecologist (pers. comm.),
indicated the total hectares are now 40,088. The units range from 222 to 10,000 hectares and are scattered
over 7 Texas counties. The Loblolly Unit consists of 222 hectares in Liberty County (Fig. 1). It is located on
the Beaumont formation on Aldine-Aris/Gyron-Aldine, Vamont, and Waller soils, and is topographically
flat, and poorly drained with very slow permeability (Deshotels 1978; Aronow 1981; Shelby et al. 1992;
Griffith 1996). Elevation is about 30 meters. After heavy rain falls, standing water is present in the flatwoods
for extended periods of time. The Unit consists of basically 1 plant community complex: Floodplain and/
or Flatland Hardwood Forest, a common association in the area (Ajilvsgi 1979; Marks & Harcombe 1981;
Bezanson 2000).
It has been suggested that a century ago the Loblolly Unit was prairie; but, because of fire suppression
and other human activities, it has become Flatland Hardwood Forest (Watson 1979; Peacock 1994; Diggs et
J Bot. Res. Inst. Texas 2/7): 1481 — 1489. 2008
: Laféha Ratanical D h Institute of Texas 2(2)
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al. 2006). However, others have questioned this interpretation maintaining that it has always been Flatland
Hardwood Forest although it may have been cleared in the past (Harcombe & Marks 1979). Our study of
the Loblolly Unit found no prairie species or any remnant prairie habitat. Therefore, the flora is clearly a
Flatland Hardwood Forest (Harcombe & Marks 1979; Marks & Harcombe 1981).
Like all units of the Big Thicket National Preserve, the Loblolly Unit has not previously been the subject
of a detailed floristic inventory.
Because our purpose is to produe a list of taxa known to occur on the Loblolly Unit, a vouchered
specimen was considered to be the only evid table for inclusion in the list. A total of 471 herbarium
specimens form the data for this report. AII Eden have been annotated by Larry Brown.
Larry Brown collected 277 specimens during 11 field days from 5 August 1978 to 14 June 2008. The
Brown et al., Vascular flora of the Loblolly Unit, Big Thicket 1483
MacRoberts' collected 127 specimens during 5 field days from 30 July 2007 to 18 May 2008. Sandra Elsik
and Warren Pruess collected 35 specimens during 2007. Susan Walker collected 23 specimens on 22 October
2007. Geraldine Watson collected 9 specimens over several years in the late 1970s and early 1980s.
In most cases, nomenclature follows Jones et al. (1997); Kartesz (1999); and Diggs et al. (2006). In cases
of multiple collections, no more than 4 are listed for each taxon.
RESULTS
The following is an annotated list of the Loblolly Unit taxa. Abbreviations used are given below.
W - Geraldine Watson. Her specimens are at TAES.
LB - Larry Brown. His specimens with collection numbers above 29,000 are at TAES and those few
with numbers below 29,000 are at SBSC.
MM - Barbara and Michael MacRoberts. Their specimens are at TAES.
SEWP - Sandra Elsik and Warren Pruess. Their specimens are at TAES.
SH - Shawn Harper, grandson of LB.
SW = Susan Walker. Her specimens are at TAES.
* = exotic
+ = see note at end of list.
Acanthaceae Araceae
Hygrophila lacustris (Schlecht. & Cham.) Nees, LB 32266 Arisaema triphyllum (L.) Schott., MM 8004
Justicia ovata (Walter) Lindau var. lanceolata (Chapm.) R.W. Ardliaceãe
Long, LB 32200, 29281, 29298; MM 7930 n oat LB 32306; MM 7895
Ruellia r (J.F. Gmel.) Steud., LB 29318; MM 8231;
LBSH 33675
Arecaceae
Sabal minor Jacq.) Pers., LB 32415
Aceraceae . .
Acer rubrum L., SEWP 3044; MM 7906, 8094 Aristolochiaceae
Aristolochia serpentaria L., MM 7990
Amaryllidac
Hymenocallis pom (Raf) Shinners, LBSH 33311 Asclepiadaceae
Asclepias perennis Walter, LB 32211, 29285; MM 7923; SW
Anacardiac
Rhus SINUS L,LB3
Toxicodendron radicans "s A ntze, LB 32254 o racea
mella DE (Lam. Jansen, LB 32206; MM 7935
Pru p | | a L, LB 32431; MM 7975
entella erecta (L.f) Fernald, MM 8012; LB 32315 Baccharis halimifolia L, MM 7998
Chaerophyllum tainturieri Hook. var. dasycarpum Hook. ex S. Conoclinium coelestinum (L) DC, LB 32269
i i mE Coreopsis tinctoria Nutt., LB 32261; MM 7926
capi tainturieri Hook. var. tainturieri, LBSH 33302 Eclipta prostrata (L) L, MM 7934, 8010; LB 5849
alae sa ne vá hyllum (Pers) Sprague ex Britton & P. Elephan no carolinianus Raeusch., LB 32425; s 7968
Wilson, Elephantopus tomentosus L, LB 32414; MM
pha
Secada dai atum Debe ee pete rl (L.) Raf, LB 32274, 3241 T 8070
Eryngium prostratum Nutt. ex DC, LB 3 Erigeron philadelphicus L, SEWP 3034
Hydrocotyle verticillata a Thunb, LB 2 m 9323 MM 791 E Eupatorium lao Grell MIG
= +£up m semiserratum DC., GW 2159
fenes LBSH id Eupatorium serotinum Michx. 2
Ptilimnium capillaceum Heels Raf, LB 32205 e E (Torr. & A. Gray) Greene ex Porter &
Sanicula canadensis L., LB 292 Britt
Apocynaceae ES Duc A (Willd.) Cabrera, SEWP 3030
Trachelospermum difforme (Walter) A. Gray, LB 32412 Gamochaeta purpurea (L.) Cabrera, LBSH 33305, 33488
Aquifoliaceae Helenium amarum (Raf) Rock var. amarum, LB 32265; MM
902
llex decidua Walter, LBSH 33298; SW 139
llex opaca Aiton, LB 32278
Krigia cespitosa (Raf) K.L. Chambers var. cespitosa, LBSH
llex vomitoria Aiton, LB 32251 dde
Mikania scandens (L.) Willd., LBSH 33666
1484
Pluchea camphorata (L.) a nu a MM 8002
Pluchea foetida (L) DC., M
Pyrrhopappus carolinianus qe DC, LB 32436
Pyrrhopappus pauciflorus (D. Don) DC., LB 29309a
Solidago altissima L, MM 7976
ps ii Mill. var. asper (Aiton) Fernald, LB 32293;
E m ) Hill, Tem ~
*Sonchus oleraceus L., M
Symphyotrichum e " Love & Love, MM 8099
iott) Nesom, LB 32270
Ve ernonia missurica Raf, MM 790 8
*Youngia japonica (L.) DC., LBSH 33308
Berberidaceae
Podophyllum peltatum L., LBSH 33507
Betulaceae
Carpinus caroliniana Walter, SW 140
Ostrya M (Mill) K. Koch, LB 32424
Bignon
Mas uo m L, LB 32252, 29317
Campsis radicans (L.) Seem. ex Bureau, LB 32443
Catalpa bignonioides Walter, LB 32442
Blechna
da areolata (L.) T. Moore, MM 7983; LBSH 33484
Boraginaceae
B o dd poc E MM 8100
li Mill, LB 29312, 5852
Myosotis macrosperma Engelm,, LBSH 33297
Brassicaceae
Rorippa sessiliflora (Nutt.) Hitchc., LBSH 33284
Bromeliaceae
Tillandsia usneoides (L.
Buddlejaceae
Polypremum procumbens L., LB 32437
Campanulaceae
Lobelia cardinalis L., MM 8009; LB 14856
L
L, SW 137
Caprifoliaceae
*Lonicera japonica Thunb., SEWP 3028
Sambucus Hu ra L. subsp. canadensis (L.) Bollii, LB 32255;
M7
Viburnum po L., LB 29305
PRA
*Cera glomeratum Thrill, SEWP 3042
LE id (L) Vill, SEWP 3041
Commelinaceae
Commelina diffusa Burm f, MM 8077
Commelina virginica L., LB 32272; MM 7925
Convolvulaceae
Dichondra carolinensis Michx., SEWP 3027; MM 7896
Ipomoea cordatotriloba vobi 904
Ipomoea lacunosa L., MM 800
f Texas 2(2)
Cor
E pus L., SEWP 3005; SW 145; LBSH 33672
Cornus foemina Mill, LB 29320, 29304; SW 138
Nyssa biflora Walter, LB 32304; LBSH 33301
Nyssa sylvatica Marsh., LB 29306; MM 8085; LBSH 33506
Crassulac
Fico le L, LB 32433, 14866; MM 7916
Cucurbitaceae
Cayaponia quinqueloba (Raf) Shinners, SW 149
Cyperaceae
Carex abscondita Mack., med ~
Carex blanda Dewey, M
Carex complanata Torr. & e LB 32248
Carex caroliniana Schwein., LBSH 33293
arex cherokeensis Schwein., LBSH 33289
Carex crus-corvi Shutlew. ex Kunze, LB 32189; SEWP 3022
Carex debilis Michx., LB 3
Carex flaccosperma n a 33487
Carex frankii Kunth, LB 32191, ae 9; MM 7927, 8230
Carex intumescens Rudge, LBSH 3348
Carex joori L H. Bailey, LB 32277, 3724; MM 7996
Carex louisianica Bailey, LB 29296, 29300, 32419; MM 7997
Carex reniformis (L.H. Bailey) Small, LBSH 33287
Carex triangularis Boeck., LBSH 33285, 33481; LB 29324
Carex tribuloides Wahlenb. var. sangamonensis Clokey, LB
32186, 32195; LBSH 33502, 33680
Cyperus acuminatus Torr. & Hook. ex Torr, LBSH 33492
*Cyperus entrerianus Bockeler, LB 32242; LBSH 33479; SW
159
=
Cyperus erythrorhizos Muhl., MM 8005 b
P ai ochraceus Vahl, Ri 7977
|, LB 32283, 32310
Can virens Michx, LB
Eleocharis engelmannii pn m 33668
Eleocharis microcarpa Torr, MM 8022
pisa montana (Kunth) Roem. & Schult, LB 32285
Eleocharis montevidensis Kunth, LB 32267; SEWP 3024
a miliacea (L.) Vahl, MM 8089
Isolepis carinata Hook. & Arn. ex Torr, LBSH 33299
+Kyllinga pumila Michx., LB 32258, 17501a; MM 7987, 8080
nchospora caduca Elliott, LB 32286, 32307, 32196, 32192
Rhynchospora corniculata (Lam.) A. Gray; LB 32245; MM
7922
Rhynchospora glomerata (L.) Vahl, LB 32301, 32317; LBSH
33678
Rhynchospora inexpansa (Michx.) Vahl, LB 32201; LBSH
3
Rhynchospora mixta Britton, LB 29295; LBSH 33480
Scirpus cyperinus L., LB 32212; SW 132
Scleria oligantha Michx., LB 32423
Scleria triglomerata Michx., LB 29286
nstaedtiaceae
Pteridium aquilinum (L) Kuhn var. pseudocaudatum (Clute)
A. Heller, LB 32416
Brown et al., Vascular flora of the Loblolly Unit, Big Thicket
Dryopteridaceae
Onoclea sensibilis L., LB 32289, 29288; MM 7984
Polystichum acrostichoides (Michx) Schott, MM 8095; LBSH
3
Ebenaceae
Diospyros virginiana L., LB 29284
Ericaceae
Vaccinium arboreum Marshall, LB 29294
Euphorbiaceae
Acalypha gracilens A. Gray, LB 32308
Phyllanthus caroliniensis en E 32411,33132
*Phyllanthus urinaria L., MI
*Triadica sebiferum (L) Small, ie € MM 7905
Fabaceae
“Albizia julibrissin Durazz., MM 8109
Centrosema virginianum (L.) Benth., GW 2156
Desmodium glabellum (Michx.) DC., GW 2157; MM 7992;
H 33494
BS
*Medicago lupulina L, MM 8
*Trifolium campestre Schreb., sn 33291
ROGO repens L, LBSH 3328
subsp T LBSH 33290
Fagaceae
Quercus alba L., MM 8101
Quercus falcata Michx., MM 8103, 8093
Quercus laurifolia Michx., LB 32275a
Quercus michauxii Nutt., LB 29283; MM 7911
Quercus nigra L., MM 8084, 8092; LB 32275
Quercus phelios L., LB 32250, 29290
Quercus shumardii Buckley, SW 135
Quercus velutina Lam., MM 8083, 8090; SW 144, 141; LBSH
677
Gentianaceae
*Centaurium pulchellum (Sw.) Druce, MM 8229
Geraniaceae
Geranium carolinianum L., SEWP 3045
Haloragaceae
Proserpinaca palustris L. var. crebra Fernald & Griscom, LB
32276; LBSH 33671
Hammamelidaceae
Liquidambar styraciflua L., LB 29291
Hydrophyllac
Hydrolea a A SW 153
Hypericaceae
Hypericum ue .) Crantz, LB el 5; MM 7915
Hypericum mutilum L., LB 32299; MM 8
Triadenum dee (J. G. Gmel.) Gleason, Hi 29293; MM 7978
ypoxidaceae
Hypoxis curtissii Rose, MM 8237
Hypoxis wrightii (Baker) Brackett, LBSH 33287
Iridac
ae
annm rosulatum E.P. Bicknell, LB 29311; SEWP 3026;
MM 8236
1485
Juglandaceae
Carya aquatica (Michx. f.) Nutt., SW 136
Carya glabra (Mill) Sweet, LB 32292; MM 8087; LBSH 33496;
SW 134
Juncaceae
Juncus acuminatus Michx., LB 32313
Juncus coriaceus Mack., LB 32243; SEWP 3043
Juncus dichotomus Elliott, LB 32267a
Juncus diffusissimus Buckley, LBSH 33490
Juncus effusus L., LB 31 a 3025
Juncus tenuis Willd., LBSH
Juncus validus Coville, LB
PE
oma hispida Pursh, LBSH 33495
as alata (Raf) Shinners, MM in
Lycopus rubellus Moenc
Lycopus virginicus L., GW 2086; vom
Micromeria brownei (Sw.) Benth. var. SUE A. Gray,
LB32207; SEWP 3037
Prunella vulgaris L., LB 29308
Salvia lyrata L., SEWP 3040
Scutellaria integrifolia L., LB 32287
cte tenuifolia Willd., LB 32190, 32260; MM 7921; LBSH
3669
Laurac
Sassafras Um (Nutt.) Nees, SEWP 3014; MM 7901, 7993
Liliaceae
Nothoscordum bivalve (L.) Britton, SEWP 3010
Linaceae
Linum striatum Walter, LB 32439
Banda
Gelsemium sempervirens (L.) Aiton f, SEWP 3015
Miro ut (J. F. Gmel) Torr. & A. Gray, LB 32188; MM
Lygodia
“ygodum je japonicum (Thunb. ex Murray) Sw. LB 29316; SEWP
3016 7909
ie
*Cuphea carthagenensis (Jacq.) J. F. Macbr., LB 32305, 5593;
MM 7928
Magnoliac
Magnolia ie L, SW 146
Malvaceae
Modiola caroliniana (L.) G. Don, SEWP 3038
Sida rhombifolia L., LB 32294; MM 7970
Melastomataceae
Rhexia mariana L. var. mariana, LB 32295
Moraceae
Morus rubra L., MM 7903
Myricaceae
Morella cerifera (L) Small, SEWP 3011
1486
Oleaceae
Chionanthus , MM 8086
Fraxinus americana L., LB 2
Fraxinus pennsylvanica te LB 32309, 32432; MM 8107;
Onagraceae
de decurrens Walter, MM 8001
Ludwigia glandulosa Hae Le 32210
us hirtella Raf., LB 3
Ludwigia octovalvis (Jac ud : A Raven; MM 7933, 7979
Ludwigia palustris (L.) Elliott, LB 3219
Ludwigia sphaerocarpon Elliott, GW 2399
Qenothera speciosa Nutt., SEWP 3035
Ophioglossaceae
Botrychium biternatum (Savogny) Underwood, MM 7995,
8104
Oxalidaceae
Oxalis corniculata L. var. wrightii (A. Gray
303 H 33304
M
B.L. Turner, SEWP
Passifloraceae
Passiflora incarnata L., E ~ MM 8098
Passiflora lutea L., LB 322
Pinaceae
Pinus taeda L., LB 32291, 32319
Plantaginaceae
*Plantago major L., LB 32184; MM 7973, 8234
Plantago rhodosperma vm LBSH 33310
Plantago virginica L., MM 8
Plata
dosi pa Pra L., LB 32420; SW 133
Poaceae
Agrostis hyemalis (Walter) Britton, Sterns € Poggenb., LBSH
33499
Axonopus fissifolius (Raddi) Kuhlmann, LB 32314, 32303a;
GW 2177
Axonopus furcatus (Flugge) A. S. Hitche., LB 32427
*Briza minor L., LBSH 33288
*Bromus japonicus Thunb. ex Murray, LB 32181
Chasmanthium laxum (L.) H.O. Yates, GW 2314
Dichanthelium acuminatum (Sw.) Gould & C.A. Clark var.
acuminatum, LB 29301a, 32320
Dichanthelium acuminatum (Sw.) Gould & C.A. ned var. lin-
Nou (Nash) ene & CA. Clark, LBSH 3348
(Schult) Gould s commu-
tatum, LB 29287, 32422; LBSH 33679
ecco oe (L) Gould subsp. dichotomum,
7988
m X pM (L) Gould subsp. microcarpon
(Muhl. ex Elliott) Freckmann & Lelong, LB 32197, 32257,
3; LB
Dichanthelium laxiflorum (Lam.) Gould, LB 32435
Dichanthelium ovale ( Elliott) Gould & C.A. ub ld villosis-
simum (Nash) Freckmann & Lelong, L
Ee E (Elliott) "ed ae 3020;
LBSH 3
f Texas 2(2)
vui ciliaris (Retz) Koeler, LB 32297
+E chloa colonum (L.) Link, MM 7980
rts indica (L.) Gaertn., MM 8067
Elymus virginicus L., LB 32203; MM 8227
e. hypnoides (Lam.) Britton, Sterns & Poggenb., GW
n ESI ME oH 2162; LB 32204; SW 158
ee Ki VE 2208
J (l " An subsp. setarius (Lam.) Mez ex
Ekman, GW 2313; MM 8067; LBSH 33670
Panicum anceps Michx. LB 32300, 32318
Panicum dichotomiflorum Michx., MM 8076
Panicum gymnocarpon Elliott, x pia 32428; MM 7925
cla rigidulum Nees, LBSH 3
Paspalum dilatatum Poir., LB 25 m 7974
+Paspalum conjugatum P.J. Bergius, LB 32303
aspalum laeve Michx. var. laeve, LB 32440
2
Paspalum nn PJ. Bergius, LB 17502
Paspalu aceum Michx., LB 32312
‘Paspalum urile Steud., LBSH 33497
Phalaris a a Nees ex Trin., LBSH 33498; MM 8235
Phalaris cna ies LBSH 33482
*Poa annua L., LBS 294
acchan pcs vie dels "iid
*Sacciolepis indica (L.) Chase,
Setaria parviflora (Poir.) ceden ie
Sphenopholis obtusata (Michx.) a ee 33500
Polygonac
Polygonum pam Meisn., MM
Polygonum hydropiperoides Michx., s T 29302; SEWP
3032; MM 8008
Polypodiac
"o a (L.) E.B. Andrews € Windham subsp.
ichauxiana (Weatherby) E.B. Andrews & Windham,
8091
Primulac
“Anagalisa aus SL, LB 29322; Pad —
subsp. | | LB 29313,
32182; SEWP 3018; MM 7920
Ranunculaceae
*Ranunculus muricatus L., SEWP 3047
Rhamnaceae
Berchemia scandens (Hill) K. Koch, LB 29321, 32322; SEWP
Rosaceae
Crataegus marshallii Eggl., MM 8097
Crataegus opaca Hook. & Arn. ex Hook, LB 32282, 29284a,
29307
*Duchesnea indica (Andr) Focke, SEWP 3033; SW 154
Prunus serotina Ehrh. var serotina, SEWP 3012; LBSH 33483
Rubus argutus Link, LB 32280, SEWP 3023, 3007; MM 7986
Rubiaceae
Cephalanthus occidentalis L., LB 32429; MM 7914
Brown et al., Vascular flora of the Lobloily Unit, Big Thicket 1487
Symplocaceae
poe virginiana L., LB 32259; ~ 7912
SEWP 304 Symplocos tinctoria (L.) L. Her, LB 29282; LBSH 33300; SW
148
Galium aparin
Galium tinctorium L., LBSH 33 om 33306
Houstonia pusilla Schoepf,, LBSH 33283a
Mitchella repens L., LB 29310
Saururaceae
Saururus cernuus L., LB 29283a; MM 7913
Thelypteridaceae
Thelypteris kunthii (Desv) Morton, LB 29281a, 32185; MM
7991
Ulmaceae
Scrophulariaceae Ulmus americana L., LB 32249; SEWP 3001; SW 150; LBSH
Bacopa monnieri (L.) Penn., LB 32262; MM 8088 6
Gratiola virginiana L, LB 31 938; SEWP 3002 Ürtitaceae
o NO Boehmeria cylindrica (L) Sw, LB 5595; MM 7894
Lindernia crustacea (L.) F. von Muell. LB 32410; MM 7969,
Valerianaceae
8081; SW 157
Lindernia dubia (L) Penn., LB 32288 Valerianella radiata (L.) Dufr., LBSH 33286
Mecardonia acuminata (Walter) Small, MM 7972 Ver
benaceae
lia americana L., LB 32281; MM 7898
i dita diflora (L.) Greene, MM 7929
Verbena brasiliensis x L, — 33667
iro procumbens (Mill) Small, SEWP 3029; MM 7919,
"E umbrosum (J.F. Api S.F. Blake, MM 8106
Mimulus alatus Aiton, MM 793
Scoparia dulcis L., LB 32296; WT a
Veronica peregrina L.var. peregrina, LBSH 33305a
Smilacaceae
Smilax bona-nox L, MM 7989; LB 32316
Verbena scabra Vahl,
Verbena urticifolia L., P is B
Violaceae
Viola lanceolata L., LBSH 33296
Viola sororia Willd. var. missouriensis (Green) McKinney, SEWP
Smilax glauca Walter, MM 8096 3009
Smilax rotundifolia L, SEWP 3008; LB 29297; MM 7985
il
Smilax smallii Morong, SEWP 3006 Vitaceae l
Ampelopsis arborea (L) Koehne, LB 32271
Solanaceae Parthenocissus quinquefolia (L.) Planch, LB 32311; MM 8079
Physalis cinerascens (Dunal) A.S. Hitchcock var. cinerascens, ^ vitis aestivalis Michx., SW 142; MM 8232
LB 29310a Vitis cinerea (Engelm.) Engelm. ex Millardet var. cinerea, LB
Solanum carolinense L., LB 32256; MM 7910 29299, 32418
Stvracaceae Vitis rotundifolium Michx, LB 32263; MM 7893
Styrax americana Lam., LB 32279 Xvri
yridaceae
Xyris ambigua Bey. ex Kunth, GW 2224; LB 32436
NOTES
Eupatorium semiserratum. This species is usually found in low moist areas which are frequent in this
Unit. See Brown et al.(2007) for a discussion of this species in Texas.
ton area (Harris and surrounding
Xall r3 |
is the most common Kyllinga i in the Hou
Kyllinon nd "n
cuales) but not present in this Unit. K. pumila is rare in the Houston area but common along C.R. 2071
that extends through the center of the Unit.
Quercus velutina. This woody plant, as of now, has not been observed as a tree in the Houston area but
grows as a small understory shade species. It is separated from the similar Q. falcata by the longer erect
branched hairs on the abaxial surface of the shade leaves. On Q. falcata these trichomes are smaller and
appressed [see the key in Smith (1994).
Echinochloa colonum. Most books use colona as the specific epithet. See Ward (2005) for this change in
orthography.
Paspalum conjugatum. Turner at al. (2003) mapped this species in Cameron County in extreme south Texas
with 1 record from Hardin County in the Pineywoods region of east Texas. Diggs et al. (2006) consider the
Hardin County specimen to be from a plant cultivated as a lawn grass. For now, the only east Texas collec-
tions of native plants are from Liberty County in the Trinity River National Refuge and the Loblolly Unit. The
1488 J | of the Botanical R h Institute of Texas 2(2)
plants were frequent along the southern edge west from the junction with C.R. 2071. Additional specimens
at SBSC are from the various units of the National Wildlife Refuge complex in Brazoria County.
DISCUSSION
There are 88 families and 201 genera for the 305 taxa Q71 native) on the Loblolly Unit list. Eight families,
Poaceae (42 taxa), Cyperaceae (36 taxa), Asteraceae (31 taxa), Scrophulariaceae (11 taxa), Apiaceae (10 taxa),
Lamiaceae (9 taxa), Fagaceae (8 taxa), and Fabaceae (7 taxa) account for 50 96 of all taxa collected.
We compared the present list of species for Loblolly Unit with the species reported by Turner et al.
(2003) in their Atlas of the Vascular Plants of Texas and found that 115 taxa, 37.7 96 of EE eed on x
Loblolly Unit list, are not reported in Liberty County in that source. This result is not surf
that, until recently, little focused collecting has been undertaken in the Big Thicket region.
While it is the case that our list is incomplete, as are all floras, we believe that we have collected prob-
ably 90+ 96 of the taxa. We estimate that the Loblolly Unit has about 281 to 290 native species/taxa. Of the
7 Units inventoried so far, the following taxa are reported only for the Loblolly Unit: Hymenocallis liriosme,
Cynosciadium digitatum, Rorippa sessiliflora, Commelina diffusa, Ipomoea lacunosa, Eleocharis engelmannii, Hy-
drolea uniflora, Hypoxis wrightii, Micromeria brownei, Ludwigia sphaerocarpa, Paspalum conjugatum, Ranunculus
muricatus, Physalis cinerascens var. cinerascens, and Verbena urticifolia. Only 14 taxa are new to this Unit. Thus
9596 of the taxa in the Loblolly Unit had already been collected in other units of the Big Thicket National
orbs ] /.
Preserve, which is to be expected since most of the other units p p Flatland Hardwood
forest communities.
ACKNOWLEDGMENTS
This study was supported in part by an All Taxa Biological Inventory (ATBD grant from the Big Thicket Asso-
ciation of Saratoga, Texas. We thank Paul Roling, liverwort and moss student of the Big Thicket; and Deanna
Boensch, Big Thicket fire ecologist, for providing information regarding the new Big Thicket Units. Also
thanks to Ray C. Telfair II and Eric Keith for their careful reading of the paper and for the improvements.
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BOOK NOTICE
SABEEHA MERCHANT, WINSLOW R. BRiGGs, AND Vicki L. CHANDLER (eds). 2008. Annual Review of Plant Biol-
ogy: Volume 59. (ISBN 978-0-8243-1346-3, hbk; ISSN 1543-5008). Annual Reviews Inc., 4139 El
Camino Way, PO. Box 10139, Palo Alto, CA 94303-0139, U.S.A. (Orders: www.AnnualReviews.org,
science@annualreviews.org, 800-523-8635, 650-493-4400, 650-424-0910 fax). $89.00 indiv., 831
pp., 7 5/8" x 9 3/8".
Contents of Volume 59:
1. Our bios with DE aca ox E. Conn
New I in Plants—AngZlique Besson-Bard, Alain Pugin, and David Wendehenne
Jander
3. Plant anio to Insect Hertivores Gra A. Howe and Georg
4. Patterning and Polarity in Seed is eae L. Bowman and Sandra K. Floyd
Chlorophyll Fluorescenc n Vivo—Neil R. Baker
/
6. Seed Storage Oil mais A. Graham
7. The Role of Pd ud in Photosynthetic Organisms: Emerging Functions for Glutaredoxins and Glutathionylation— Nicolas
Rouhier, StZphane D. Lemaire, and Jean-Pierre Jacquot
8. Algal Sensory A Hegemann
9. Plant Proteases: From BOT to TOS Mechanisms— Renier A. L. van der Hoorn
10. Gibberellin Met Shinjiro Yamaguchi
11. Molecular Basis of Plant Ar biie Yonghong Wang ijr Li
12. Decoding of Light ee 2 Plant Phytochromes and Thei Gat B d Giltsu Choi
13. Flooding Stress: Acclimations and Genetic Diversity—J. Een -Serres and LA.CJ. Vacs
14. Roots, Nitrogen ane ae and Ecosystem Services—Louise E. Jackson, Martin Burger, and Timothy R. Cavagnaro
15. A Genetic Regulatory Network in the Development of Trichomes and Root Hairs—Tetsuya Ishida, Tetsuya Kurata, Kiyotaka Okada,
wii Takuji Wada
16. ular Asp f Seed Dormancy—Ruth Finkelstein, Wendy Reeves, Tohru Ariizumi, and Camille Steber
l7. Trehalose Metabolism and Signals Matin J. Paul, Lucia F. Primavesi, Deveraj Jhurreea, and Yuhua Zhang
18. Auxin: The Looping Star in Plant Development—RenZ ee and Ben Scheres
19. Regulation of Cullin RING Ligases—Sara K. Hott dy Callis
20. Plastid Evolution— Sven B. Gould, Ross F. Waller, and Genel. McFadden
21. Coordinating Nodule Morphogenesis with Rhizobial Infection in Legumes— Giles E.D. Oldroyd and J. Allan Downie
22. pn. o ded a for the Polar Cell Growth Machinery in Pollen Tubes—Alice Y. Cheung and in -ming Wu
23.
US T Moves Center Stage—Fra anziska Turch, Fabio Fornara
24. Plant Aquaporins: Membrane al: with Multiple Integrated Functions—Christophe Maurel, Lionel Verdoucq, Doan- pM Luu,
and VZronique Santoni
25. Metabolic Flux Analysis in Plants: From Intelligent Design to Rational Engineering— Igor G.L. Libourel and Yair Shachar-Hill
26. Mechanisms of Salinity Tolerance—Rana Munns and Mark Tester
27. Sealing Plant Surfaces: Cuticular Wax Formation by Epidermal Cells—Lacey Samuels, Ljerka Kunst, and Reinhard Jetter
28. Ionomics and the Study of the Plant Ionome— David E. Salt, Ivan Baxter, and Brett Lahner
29. Alkaloid Biosynthesis: Metabolism and Trafficking—J3rg Ziegler and Peter J. Facchini
30. Genetically Engineered Plants and Foods: A Scientist's Analysis of the Issues (Part I)— Peggy G. Lemaux
Cumulative Index of Contributing Authors, Volumes 49-59
Cumulative Index of Chapter Titles, Volumes 49-59
J. Bot. Res. Inst. Texas 2(2): 1490. 2008
NEPTUNIA PLENA (FABACEAE: MIMOSOIDEAE) REDISCOVERED IN TEXAS
Alfred Richardson Ken King
Department of Biological Sciences 1008 W. 5% Street
The University of Texas at Brownsville Weslaco, Texas 78596, U.S.A.
80 Fort Brown
Brownsville, Texas 78520, U.S.A.
ABSTRACT
On October 17, 1938, Robert R llected fN 1 f hallow pond hofA gin Kenedy County,
Texas. No subsequent collections m Texas have id reported. 1 that i ber of our flora. On
October 27, 2007, a populati lity
RESUMEN
bu tA die a d Runj gió péci de Ni lena d h de Armstrong en el condado d
Kenedy, T: N d lección p ior de Tejas F ido q iembro p d
flora. i SP 2007, una pobiació icida del N. pl f del mi lug
Neptunia plena is commonly found in the West Indies, Mexico, and South America (Correll & Johnston
1970). It was introduced into India. It is a sprawling perennial which grows in ponds or damp soils. The
stems growing in water are horizontal and become covered with a brown, spongy fungal growth (when the
fungus is present); in soil, the stems are more or less erect. The younger stems produce adventitious roots
when submerged in water. The leaves are sensitive, twice compound, about 6.5 cm long, with 2 to 5 pairs
of pinnae. The petiole bears a gland positioned immediately below the lowest pair of pinnae (Fig. 2). The
flowers are in globose or ovoid heads bearing two types of flowers. The lower portion of the head bears large,
shiny, brilliant yellow staminodes; the upper portion of the head bears small whitish, normal flowers (Fig.
1). The legumes are flattened, about 2 cm long and 6 mm broad.
Neptunia plena was collected in Kenedy County by Robert Runyon on October 17, 1938. According
to label information on his specimen at TEX (R. Runyon 1959), it was growing in a shallow pond south of
Armstrong. Since that time, there have been no reports of the species from Texas. The natural conclusion
was that it was probably not a persistent part of the Texas flora. It was so noted in the Manual of the Vascular
Plants of Texas (Correll & Johnston 1970).
On October 27, 2007, we found a population growing in the median of Highway 77 in an ephemeral
pond in Kenedy county, 4.8 miles north of the Willacy County border. This is the same general area as
the Runyon collection. Growing nearby were Nymphaea elegans, Ludwigia decurrens, Eleocharis sp., Prosopis
glandulosa, and Physalis pubescens. The median landscape is mowed along the edge of the highway, but the
middle sections are left in the "wild" state to help support the development of native species. The soil type
is sandy loam with underlying clay which helps to hold water in the ponds after heavy rains. The rainfall
average is twenty six inches per year, but there are cycles of drought years followed by one or a few wetter
years. It is during these wetter years that the ponds form.
After examining other similar habitats, we found N. plena growing in several ponds near the one in
which it was first seen. Another population was found in Cameron County on Highway 1847, four miles
north of Highway 510. A specimen was not collected. In most localities, N. pubescens was found growing
nearby.
Our conclusion is that the plants emerge during the periodic wet seasons and produce flowers and
seeds. When the ponds disappear during the dry years, the plants become dormant and persist as seeds or
perhaps underground propagules.
J. Bot. Res. Inst. Texas 2(2): 1491 — 1493. 2008
1492 J | of the Botanical R h Institute of Texas 2(2)
P ~
Qe P PA l
3 >. a = - »
Fic. 2. Leaf with petiolar gland.
Di-havd AW Nantimi pl li I in Texas 1493
There are several reasons why N. plena could have been flourishing in this area without being noticed.
There has not been much field activity in the area, as evidenced by the relatively few specimens available.
Secondly, during the periodic drought years, the plants would be dormant. Also, the blooming season may
be short. We returned to the site one week later, and there were few blooms. A further difficulty is the pres-
ence of two similar Neptunias in the area. N. pubescens has an inflorescence similar to that of N. plena, but
it lacks the petiolar gland of that species. N. lutea also has a yellow globose or ovoid inflorescence, but the
flowers are all the same type. At a distance all three species would look alike.
Voucher specimen: TEXAS. Kenedy Co.: Hwy. 77, 4.8 mi N of Willacy County border, ephemeral pond in median, 27 Oct 2007, Rich-
ardson and King 3365 (BRIT, TEX).
ACKNOWLEDGMENTS
We thank Amanda K. Neill, director of the herbarium at the Botanical Research Institute of Texas (BRIT-
SMU-VDB) for information from that herbarium. Monique D. Reed provided helpful review comments.
REFERENCES
CorreLt, D.S. AND M.C. JOHNSTON. 1970. Manual of the vascular plants of Texas. Texas Research Foundation, Renner,
Texas. 1881 pp.
Diccs, Jr. G.M., B.L. Lirscom8, AND R.J. O’KENNON. 1999. Shinners & Mahler's illustrated flora of north central Texas.
Sida Bot. Misc. 16.
Turner, B.L. 1951. Revision of the United States species of Neptunia (Leguminosae). Amer, Midl. Naturalist
46:82-92
1494 lof the Botanical R h Institute of Texas 2(2)
ANNOUNCEMENTS
2008 DELZIE DEMAREE TRAVEL AWARD RECIPIENTS
The 20th Annual Delzie Demaree Travel Award was presented at the 55th Annual Systematics Symposium
(17-18 Oct 2008) at the Missouri Botanical Garden, St. Louis. Two students were presented the Travel Award:
Patricia Lu-Irving, University of Washington, Seattle and Piero Protti, University of Kansas, Lawrence.
The 2008 Travel Awards were underwritten by 1) Delzie Demaree Travel Award Endowment, 2) Mem-
bers of the Delzie Demaree Travel Award Committee, and 3) John Clayton Chapter of the Virginia Native
Plant Society.
Anyone interested in making a contribution to Delzie Demaree Endowment Fund, which supports the
travel award, may make contributions by VISA or MasterCard or by a check, payable to Botanical Research
Institute of Texas, to Barney Lipscomb, 500 E 4th Street, Fort Worth, TX 76102-4025, U.S.A. 1-817-332-
7432; Email: barneyObrit.org. Thank you.
THE 2009 APPLICATIONS FOR THE DELZIE DEMAREE TRAVEL AWARD
Applications for the 2009 Delzie Demaree Travel Award should include a letter from the applicant telling
how symposium attendance will benefit his/her graduate work and a letter of recommendation sent by the
major professor. Please send letters of application to: Dr. Donna M.E. Ware, P.O. Box 8795, Herbarium, Biol-
ogy Department, The College of William and Mary, Williamsburg, VA 23185-8795, U.S.A. 1-757-221-2799;
Email: ddmware@wm.edu. The period for receiving applications will end three weeks prior to the date of
the symposium if a sufficient number of A o are in hand at that time. Anyone wishing to apply
after that date should inquire whether ions are still being accepted before applying. The Systematics
Symposium dates for 2009 are 9-10 ache: 2009.
The Delzie Demaree Travel Award was established in 1988 honoring Delzie Demaree who attended
35 out of a possible 36 symposia before he died in 1987. Delzie Demaree was a frontier botanist, explorer,
discoverer, and teacher. His teaching career as a botanist began in Arkansas at Hendrix College in 1922. He
also taught botany at the University of Arkansas, Navajo Indian School, Yale School of Forestry, Arkansas
A&M, and Arkansas State University at Jonesboro where he retired as professor emeritus in 1953. One of
the things he enjoyed most as a botanist was assisting students with their field botany research.
J. Bot. Res. Inst. Texas 2(2): 1494. 2008
ASCLEPIAS HIRTELLA (APOCYNACEAE)
NEWLY DOCUMENTED FOR THE FLORA OF TEXAS
Matt White
882 Hwy 50
Campbell, Texas 75422 U.S.A.
mwhitegparisjc.edu
ABSTRACT
Asclepias hirtella (Pennell) Woodson is reported as new for Texas in Lamar, Fannin and Henderson counties.
ABSTRACT
Se cita Asclepias hirtella (Pennell) Woodson como nueva para Texas en los condados de Lamar, Fannin y Henderson.
Asclepias hirtella (Pennell) Woodson was not reported for Texas by Correll and Johnson (1970), Jones et al.
(1997), Diggs et al. (1999), or Turner et al. (2003). Recent field work at Camp Maxey, a Texas Army National
Guard facility in Lamar County, has revealed the presence of the species in open Schizachyrium scoparium
B where it was rare (Fig. 1). The plants were growing in soils mapped as Bernaldo fine sandy loam,
"a brown, slightly acid fine sandy loam" found on the "tops and sides of neg” Dec [oi ed aU
an underlying geology of Blossom Sand (Shelby et al. 1966). Common f
Vernonia texana, Liatris elegans var. elegans, Polygala incarnata, ix mueren hirta.
testa aad hirtella and the closely related A. longifoli Acerates (ElL) Woodson. Asclepias
hirtella is “a species of Midwestern prairies and barrens” ME 2007) and grows in bottomland as well
as upland prairies, glades, pastures, roadsides and railroads (Yatskievych 2006). It occurs from Georgia to
Louisiana, Oklahoma, Kansas, Minnesota, southern Ontario, West Virginia and Tennessee. Asclepias longi-
folia, by contrast, occurs along the Gulf and Atlantic coastal plain (Woodson 1954) from southeast Texas
through southern Delaware. Turner et al. (2003) mapped collections of A. longifolia from Newton, Jasper,
Tyler, Hardin, Liberty and Galveston counties as well as what he considered a very disjunct collection from
Fannin County. An examination of the Fannin County specimen indicates that it is actually A. hirtella, as
is an additional Henderson County collection cited below. These specimens suggest the species is rare in
northern and eastern Texas.
This species appears to occur with more regularity in nearby Oklahoma and has been collected several
times in nearby McCurtain, Pushmataha and Atoka counties (Pennell 1919; Hoagland et al. 2004). In Mc-
Curtain County the species is found at Red and Grassy Slough in the Red River Valley (Hoagland & Johnson
2004), both of which are very close to Texas.
The iie: key, taken from Woodson Sal: should allow these similar peces to be identified:
Inf | AA +1 £i IP RI .
1. numerous, y spherical, very siignty tinted | with purple
plants relatively stout Asclepias hirtella
1. Inflorescences both terminal and lateral, the lateral rather few, rather lax and hemispherical, the flower
rather liberally tinted with purple; plants rather slender ae longifolia
Voucher specimens: TEXAS. Fannin Co.: 1 mi N of Monkstown on FM 79, in open valley and grassy areas, 19 May 1963, D.S. Correll
$ H.B. Correll 27491 (LL). Henderson Co.: 9.5 mi S of Athens, low ground, sandy clay, 22 Aug 1951, L.H. Shinners 19106 (BRIT). Lamar
Co.: Camp Maxey TXARNG Training Facility in open grassland, 33.41675 N & 95.32904 W, 17 Aug 2007, M. White s.n. (BRIT).
ACKNOWLEDGMENTS
I am grateful to Natural Resource Specialist Katherine Crosthwaite of the Texas Army National Guard and
the staff at Camp Maxey for access to the site. I would especially like to thank both Walter Holmes and
J. Bot. Res. Inst. Texas 2(2): 1495 — 1496. 2008
1496 J | of the Botanical R h Institute of Texas 2(2)
Fis. 1. S f Asclepias hirtella collected from | County, Texas, 17 Aug 2007 (Matt White
s.n., BRIT).
Plain, McCurtain County, Oklahoma. Castanea 69:284-296,
Barney Lipscomb, whose careful
editing greatly improved the manu-
script. Amanda Neill found the
collection from Henderson County.
Special thanks to Bill Carr for track-
ing down the Correll and Correll
specimen from Fannin County and
for numerous suggestions on the
manuscript. Joe Jackson, a librarian
at Paris Junior College, for helping
with interlibrary loans.
REFERENCES
CorreLL, D.S. AND M.C. JOHNSTON. 1970.
Manual of the vascular plants of
Texas. University of Texas at Dallas,
Richardson.
Dicas, G.M. Jr, B.L. LIPSCOMB, AND R.J.
O'KENNON. 1999. Shinners & Mahl-
er's illustrated flora of north cen-
tral Texas, Botanical Research
Institute of Texas, Fort Worth.
HOAGLAND B.W., A.K. Burnoo, 1.H. BUTLER,
P.H.C. CRAWFORD, A.H. Upasi, W.J. Euis-
ENS, AND RJ. Tyr. 2004. Oklahoma
vascular plants database (http://
geo.ou.edu/botanical), Oklahoma
Biological Survey, University of
Oklahoma, Norman.
HOAGLAND, B.W. AND F. JOHNSON. 2004.
The vascular flora of Red and
Grassy Slough Wildlife Man-
agement Areas, Gulf Coastal
Jones, S.D., J.K. WiPrr, AND PM. Montcomery. 1997. Vascular plants of Texas: a comprehensive checklist including
synonymy, bibliography, and index. University of Texas Press, Austin.
PENNELL. F.W. 1919. Plants of southern United States. Bull. Torrey Bot. Club. 46:184—185.
ResseL, D. 1979. Soil survey of Lamar and Delta counties, Texas. Texas Agricultural Experiment Station, College
Station.
SHELBY, C.A., M.K. Pieper, D.E. Owen, TJ. FREEMAN, A.C. WRIGHT, AND V.E. Barnes. 1966. Geologic atlas of Texas, Texarkana
sheet. Bureau of Economic Geology, The University of Texas at Austin.
Turner, B.L., H. NicHoLs, G. Denny, AND O. Doren. 2003. Atlas of the vascular plants of Texas. Vol. 1. Sida, Bot. Misc.
24:1-648.
WEAKLY, A.S. 2007. Flora of the Carolina, Virginia, Georgia, and surrounding areas. working draft as of 11 January
2007. http://www.herbarium.unc.edu/flora.htm
Woopson, R.E., JR. 1954. The North American species of Asclepias. Ann. Missouri Bot. Gard. 41:1-211.
YATSKIEVYCH, G. 2006. Steyermark's flora of Missouri. Vol 2. Missouri Department of Conservation, Jefferson City.
LILAEOPSIS CAROLINENSIS (APIACEAE) A SPECIES NEW TO TEXAS
AND A KEY TO LILAEOPSIS IN TEXAS
Stephan L. Hatch A. Tucker Slack
S.M. Tracy Herbarium (TAES) LD. B Wildlife Hie e
fE Sci 1M sand Wildlife D
Texas A&M University E Texas 77640, UnA
College Station, Texas 77843-2138, U.S.A.
ABSTRACT
The occurrence of Lilaeopsis caroli is J.M. Coult. & Rose is reported new to Texas and a key to the species of Lilaeopsis in Texas is
provided.
Key Wonps: Lilaeopsis carolinensis, new to Texas
RESUMEN
Se reporta la presencia de Lilaeopsis carolinensis J.M. Coult. & Rose, en Texas y se p lave y l especies Lilaeopsis en Texas.
Lilaeopsis carolinensis was recently collected at the Texas Parks and Wildlife Department, J.D. Murphree
Wildlife Management Area, Jefferson County, Texas, in an interior ditch of compartment 2 on 15 April 2008.
The species occurred in a shallow, freshwater canal associated with Hydrocotyle sp. and Salvinia minima J.G.
Baker in a floating vegetation mat. The specimen was identified using Radford et al. (1968), Godfrey and
Wooten (1981), and Tobe et al. (1998).
This species is native to the southeastern U.S.A. (USDA-NRCS 2008) and likely a recent introduction
to Texas, not having been recorded by Correll and Johnston (1970), Correll and Correll (1975), Affolter
(1985), Hatch et al. (1990), Jones et al. (1997), or The Flora of Texas Database (2008). The following herbaria
(TEX/LL; SHSU; and ASTC) searched their collections for Texas specimens of this species without finding
specimens.
The North American distribution of L. carolinensis extends from Virginia south to Florida and west to
Louisiana [(Radford et al. (1968), Godfrey and Wooten (1981), and Tobe et al. (1998)]. State distributions
are AL, AR, FL, GA, LA, MS, NC, SC, and VA (USDA-NRCS 2008). This is the first reported and vouchered
collection of the species from Texas. This discovery in Texas is not unexpected because of its close proximity
to similar Louisiana habitats from which the species is known.
Another species of Lilaeopsis, L. chinensis (L.) Kuntze, is also known from a single Texas collection (L.
Brown & Marcus 16926) from Chambers County. This specimen was collected in 1998. The following key
will serve to identify the two Texas species. Tobe et al. (1998) and Radford et al. (1968) provide illustrations
of both species.
A KEY TO THE SPECIES OF LILAEOPSIS IN TEXAS
1. Peduncles shorter than leaves; pedicels 5-10 mm long; leaves (phyllodes) 10-30 cm long with 7-15 septa;
habitat fresh water L. carolinensis
Peduncles | | licels 3-4 long; | (phyllodes) 1-5 cm long with 4-6 septa; habitat
. chinensis
brackish water
Lilaeopsis carolinensis Coult. & Rose, (Fig. 1). CAROLINA GLasswort. Rhizomatous, perennial herbs; stems
adventitious, ca 50 cm long, horizontal, roots at nodes; leaves (phyllodes) 10-30 cm long, clavate to linear,
with 7—15 transverse septa; peduncles much shorter than leaves; pedicels 5-10 mm long; umbels 5-15
flowered, simple; sepals absent or minute; petals 5, white; fruits 272.5 mm diameter, mericarps ovate, veins
prominent; flowering period March to June.
J. Bot. Res. Inst. Texas 2(2): 1497 — 1498. 2008
1498 J | of the Botanical R h Institute of Texas 2(2)
Specimen examined. TEXAS. Jefferson Co.: Texas Parks and
Wildlife Department, J.D. Murphree Wildlife Management Area
in ditch within Compartment 2. Shallow, fresh water. 15 Apr
2008 Slack s.n. (TAES). Synonym L. attenuata auct. non (Hook.
& Arn.) Fern.
Lilaeopsis chinensis (L.) Kuntze, EASTERN GRASS-
wort. Rhizomatous, perennial herbs; stems ca. 50
cm long, horizontal, roots at nodes; leaves (phyl-
lodes) 1-5 cm long, clavate to linear, with 4—6
transverse septa; peduncles longer than leaves;
pedicels 3-4 mm long; umbels 4-10 flowered,
simple; sepals absent or minute; petals 5, white;
fruits 222.5 mm diameter, mericarps ovate, veins
prominent; flowering period March to June. Tidal
marshes, brackish. Synonym L. lineata (Michx.)
Greene. See PLANTS Database (USDA-NRCS 2008)
for the distribution.
ACKNOWLEDGMENTS
The authors thank the curators of the following
herbaria for a search of their collections for species
records in Texas; TEX/LL, ASTC, and SHSU.
Peduncle
REFERENCES
AFFOLTER, J.M. 1985. A monograph of the genus Lilae-
opsis (Umbelliferae). Syst. Bot. Mongr. 6:1-140.
CorreLL, D.S. AND H.B. CorreLL. 1975. Aquatic and wet-
land plants of the southwestern United States.
Environmental Protection Agency. U.S. Govern-
ment Printing Office, Washington D.C.
Correll, D.S. AND M.C. Johnston. 1970. Manual of the
l vascular plants of Texas. Texas Research Founda-
78 zt * tion, Renner.
: AK. 0 E WI . FLORA OF Texas DATABASE. 2008. University of Texas Her-
barium, Plant Resources Center, Austin. < http://
Ic. 1. Digital image of Lilaeopsi linensis showing im I www.biosci.ut Iu/prc/Tex.html»
and pedunde characteristics. Goprntv, R.K. AND J.W. Wooten. 1981. Aquatic and wet-
land plants of southeastern United States. Dicoty-
ledons. University of Georgia Press, Athens.
Hatch, S.L., K.N. GANDHI, AND L.E. Brown. 1990. Checklist of the vascular plants of Texas. Texas Agric. Exp. Sta. Bull
M.P. 1655. College Station.
Jones, S.D., J.K. Wiere, AND PM. Montcomery. 1997. Vascular plants of Texas. University of Texas Press, Austin.
RADFORD, A.E., H.E. Aries, AND C.R. BELL. 1968. Manual of the vascular flora of the Carolinas. The University of North
Carolina Press, Chapel Hill.
Tose, J.D., K.C. Burks, R.W. CANTRELL, M.A. GARLAND, M.E. SweeLey, DW. Hatt P. WALLACE, G. ANGLIN, G. NeLson, J.R. Cooper, D.
BickNER, K. GILBERT, N. AYMOND, K. GREENWOOD, AND N. RAYMOND. 1998. Florida wetland plants. Florida Department of
Environmental Protection, Tallahassee.
USDA-NRCS. 2008. The PLANTS Database National Plant Data Center, Baton Rouge, Louisiana. htto://plants.
usda.gov Accessed June 2008.
REVIEWERS:
VOLUME 1 (2007) AND VoLuME 2(2008)
365 reviewers: several in
dividu
als revi
iewed more than
n one manuscri
pt.
Thank you for supporting Journal of the Botanical Research Institute of Texas.
Acevedo-Rodriguez, Pedro
bib
Allorge, L.
de Assis, Marco António
Austin, Daniel F
Aymard C., Gerardo
Baeza, Carlos M.
Beard, Catherine
Blanco, Mario A.
Bogler, David J.
Booth, Helga Ochoterena
Boufford, David E.
e
Brown-Marsden, Margaret
Bryson, Charles T.
Burger, William C.
Buthod,
Butterworth, Charles A.
Amy
Campos-Ríos, María G.
Cantino, Philip D
Carlquist, Sherwin
Carr, William R
Castelo, Elen
Catharino, c; M.
Cerón, Carlos
Cerros-Tlatilpa, Rosa
Chambers, Henrietta L.
Chester, Ed
Christensen, Knud Ib
ward W.
Clancy, Keith E.
Clark, John L.
Clark, Lynn G.
Clark- ae rA A.
Clemants,
Croat, Thomas B.
Cusick, Allison W.
Daniel, P.
Darbyshire, Stephen J.
Demissew, Sebsebe
Diamond, Alvin
ipis Timothy A.
Diggs, George M
Dillon, uus Q.
Dorr, Laurence J.
Doyle, James A.
Dutton, Bryan E.
Eberwein, Roland K.
Edeoga, Hilary O.
Edwards, Christine E.
Eggli, Urs
Elisens, Wayne J.
Endress, Mary E.
Ertter, Barbara
Ervin, Gary
Espejo-Serna, Adolfo
Estrada C., A. Eduardo
Ezcurra, Cecilia
Fantz, Paul
Farruggia, Frank
Felger, Richard S.
erguson, Diane M.
Feuillet, Christian
Finot, Victor
Frederick, Lafayette
Freire-Fierro, M.Sc. Alina
Freitas, Maria de Fátima
Freudenstein, John V.
Fritsch, Peter W.
Funk, Vicki A.
Gandhi, Kanchi
García Mendoza, Abisai J.
García R., Ignacio
J. Bot. Res. Inst. Texas 2(2); 1499 — 1500. 2008
García-Armora, Pedro Ramírez
Giraldo-Cañas, Diego
Giussani, Liliana Monica
Gomesz-Sanrche
z, Maricela
González-Elizondo, M. Socorro
herel
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Guala, Gerald F
Haberle, Rosemarie (Romey) C.
Harms, Vi
Hartman, Ronald L.
Hatch, Stephan L.
Hays III, John F.
Heil, Kenneth D
Helliwell, Richard
Herendeen, Patrick
Hess, William
Heydler, Susan O.
Hind, D.J. Nicholas
Hipp, Andrew
Hodgson, Wendy C.
Holmes, Walter C.
Holst, Bruce K.
Iltis, Hugh H.
Jarvis, Charlie
Jensen, Elizabeth A.
énez, oe Rodriguez
Johnson,
Johnson, iud P
Jones, Ronald L
Jones, Thomas H.
Jorgensen, Peter M.
Judd, Walter S.
Judziewicz, Emmet J.
Kallunki, Jacquelyn A.
Kaul, Robert B
Kawasaki, Lucia
o bn Joseph H.
Kirschner, Jan
Knox, Eric
Kral, Robert
Krings, Alexander
Krosnick, Shawn
Kruse, Dale A
Kvacek, Zlatko
Lakshminarasimhan, P.
Lammers, Thomas G
Lance, Ron
cl Leslie R.
Lavin, Mat
Lawson, cen A.
E
Lidén, Magnus
Liesner, Ron
Lim, A.L.
Livshultz, Tatyana
mann, Lúcia G.
Lombarda, Julio A.
Lorence, David
Lowrey, Timothy K.
Lucas, Eve
Luteyn, James L.
MacDougal, John
MacRoberts, Michael H.
Magallanes, Arturo Solis
Magrath, Lawrence K.
Mayfield, Mark H.
Mazzola, Eugene
McCook, Lucile M.
1500
McCullough, Deanna
McDonald, Charles B
McDowell, Tim
McLaughlin, Steven P.
McNeill, John
McPherson, Gordon
Mejia Saules, Teresa
Mellicham L
Mereles (FCQ), Fátima
Miller, Dennis L.
Miller, James S.
Mora-Olivo, Arturo
Morgan, David
Mort, ME Eugene
Morton, Cynthia M.
Morton,
Mueller, Jochen
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Mygatt, Jane
Naczi, Robert F.C.
Nelson, Allan D.
Nelson, John B.
Nemoto, Tomoyuki
Nesom, Gu
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Nicolson, Dan H
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Prado, Darién E.
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Pruski, John F.
Pupulin, Franco
Quedensley, Taylor Sultan
Quintana, M.Sc. Catalina
Riser II, James P.
Roalson, Eric H.
Roberts, Roland
Rodríguez, José Delgadillo
Romero, Gustavo
Rothrock, Paul
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Ryburn, Adam K.
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Sabatier, d
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I I anf oh Dat
Sano, Paulo T
Santos-Goncalves, Ana Paula
Sasek, Thomas
Sasidharan, N.
Saunders, Richard
Shipes, Barbara Greene
Silverstone-Sopkin, Philip A.
Simoes, Andre
Sinha, Riya
Siqueiros, Maria Elena
Sivinski, Robert C.
kin Mark
Eom Laurence E.
?
Sobral, Marcos
Sosa, Victoria
Spellenberg, Richard
$ john
hn L.
Szlachetko, Dariusz L.
Talent, Nadia
Triplett, Jimmy K.
In Lhlartiénd 9)
f Texas 2(2)
Tripp, Erin
Trock, Debra K.
Turland, iq
Turner, Bi
Urbatsch, Lowell E.
Valdés-Reyna, Jesüs
Valencia A., Susana
van de Veire, Jacqueline
Vanderpool, Staria S.
VanDevender, Thomas R.
Venu, P.
Villarreal-Quintanilla, José A.
Villaseñor, Jose Luis
Voss, Edward G
Wagner, Warren L.
Walck, Jeffrey Lee
Wallace, Gary D
Weitzman, Anna L.
Wheeler, Elisabeth
White, David S
Whittemore Ala an
Whitten, William Mark
Widjaja, E
Wojciechowski, Martin F.
Woodland, Dennis W.
Worthington, Richard D.
Wunderlin, Richard P.
Zhou, Lihua
Ziljstra, Gea
zm
Zuloaga, Fernando
INDEX TO 233 TITLES WITH 324 AUTHORS:
VOLUME 1 (2007) AND VoLUME 2(2008)
We know you have a choice. Thank you for choosing and supporting
Journal of the Botanical Research Institute of Texas.
2C DNA content values in Amaranthus (Amaranthaceae)
by DonaLo B. Pratt, SHALINI N. JHANGIANI, AND ROBERT J. Wic-
GERS—2(2):1219
A floristic survey of National Park Service areas of Timucuan
Ecological and Historic Preserve (including Fort Caroline
National Memorial), Duval County, Florida by Wenoy B.
ZOMLEFER, DAVID E. GIANNASI, AND WALTER S. Jupp —1(2):1 157
A o survey of vascular plants over parts of northeastern
MicHAEL H. ScHiesout, DoNALD L. HAZLETT, AND
Ts SNOW—2(2):1407
A hil
fee tA PTUS
ae) from northeastern Mexico by Guv L. Nesom—1(2):891
A new hedge-nettle Paa o Ton "mew enor
highlands ofthe United
ern species by JoHn B. ane
A new name for the well-known i eum (Aspleniaceae
from Hale County, Alabama by Brian R. KEENER AND L.J. Dav-
ENPORT— 1 (1):103
À new species and two new combinations in Potentilla
sect. Niveae Pann by Davo F. Murray AND REIDAR EL-
ven—1(2): 811
—
A second species of Cyclopogon s.s. (Orchidaceae: Spiran-
thinae) by Eric A. CHRISTENSON—1(2):1
A synopsis of Pachyphyllum (Orchidaceae) by Eric A. Chris-
TENSON—2(1):285
A synopsis of the Mexican and Central American species
of Vaccinium S MAD by RoserT L. WILBUR AND JAMES L.
Lurern—2(1):20
Ataxonomic revision of Crataegus series 2 alas (Rosa-
ae) s B. SH AND nds do mmu 11
(Cyperaceae) from deis an first reporti from Mexico by
Davip J. ROSEN AND RICHARD CARTER— 1(1):77
sali ode (Clusiaceae), a new genus for Poecil-
uciflorum, an endemic and endangered
i Western Beds India by S. RAJKUMAR AND M.K.
pecuniary ):12
An Early C giost fossil of ibl
in rosid floral diversification by Grona O. POINAR, JR, KENTON
L. CHAMBERS, RoN B —2(2):1
Analysis of the completeness of c plant icon in
Florida by JUSTIN K WILLIAMS AND bien. E md
IH J
tor | Ya foli ai
" TAM Í "
Island, Philippines by MELANE P. MEDECILO, GEORGE E a AND
Domingo A. Mapuuip—1 (2):815
A new species of Chusquea sect. Verticillatae (Poaceae:
Bambusoideae) from Ecuador by Lynn G. CLARK, CHRISTOPHER
D. TYRRELL, Jimmy K. TRIPLETT, AND AMANDA E. FisHER—1(2):847
Anew species of Lasiolaena (Asteraceae: Eupatorieae: Gyp-
tidinae) from Bahia, Brazil = ~ Roaue, SILVANA C. FERREIRA,
AND HaRoLo RoBINSON—2(2):8
A new species of the fern g (Pteridaceae)
from southeastern Brazil by AGE SAuNo—2(2): 843
A new variety of Festuca roemeri (Poaceae) from the
California Floristic Province of North America by BARBARA
L. WitsoN—1(1):59
A new variety of Humboldtia (Fabaceae: Caesalpinioideae)
from the Western Ghats of India by PS. Upayan, KV. TUSHAR,
AND SATHEESH GEORGE— 1(1):121
A note on the type locality of um arizonica (On-
Al L AA
€) DY KATHRYN MIA
z—1(1
A on design for a light weight P ducts = press by
CHARLES T. BRYSON AND RICHARD CARTER—
A phylogeny of Bejaria (Ericaceae: a based
on molecular data by CATHERINE M. BUSH AND KATHLEEN A.
KRON—2(2):1
A phytogeographical analysis of Taxus (Taxaceae) based on
leaf anatomical Ao S RICHARD W. 2 (1):291
A ORNEJO
AND T H. Irns—2(1):75
J. Bot. Res. Inst. Texas 2(2): 1501 — 1506. 2008
ia
Eje Volcánico Transversal, México by Mana GÓMEZ- SÁNCHEZ
AND KATHIA GEORGINA TELLEZ-PIMIENTA—2(1):49
PI ) I g ap-
Rech de América del Sur by Xavier Cornejo, HuaH H. Itis,
D A. SPENCER TomB—2(1):61
icem checklist of the vascular flora of the Beech Creek
Unit of the Big Thicket National Preserve, Tyler County,
Texas by Larry E. Brown, BARBARA R. MacRosenrs, MICHAEL H.
MacRoberTs, PAUL A. HARCOMBE, WARREN VY. Pruess, |. SANDRA ELsIK,
AND SUZANNE BIRMINGHAM WALKER—2(1):65 1
Annotated checklist of the vascular flora of the Loblolly Unit
ofthe Big Thicket National Mert Liberty County, Texas
by Larry E. Brown, BARBARA R. MacRoberTs, MicHAEL H. MACR
ERTS, WARREN W. PRukss, |. SANDRA hes AND SUZANNE BIRMINGHAM
2(2):1481
WALKER—
Annotated vascuiar flora of the Dead Horse Mountains, Big
Bend iud ou Texas, NES i on mee HEU
FENSTERMACHER, A, MICHAEL Powe LL, JOE SIROTNAK, AND ren
NL LL 1):685
y
c. P
cies from Peru by ISIDORO ERE PAULM. ERES put
J. SORENG, AND SIMON L4GAARD— 1 (2):841
Araucarian source of fossiliferous Burmese amber: spectro-
GEORGE POINAR, JR, JOSEPH
B. DE AND YUYANG Wu—1(1):44
Aristidae eludendae Il: a dd of the Aristida gib-
bosa complex (Poaceae: Aristideae), including A. mar-
1502
ginalis, A orizabensis, and A
E: iai iim dra W. nM 309
by DouaLas H. GoLDMAN—2(2):1337
Asclepiadoidea (Apocynaceae) como maleza en el cultivo
del agave tequilero (Agave tequilana var. “azul”) by Irma
LOPEZ-MURAIRA, RUBÉN Da ISAAC ANDRADE GONZÁLEZ Y JUAN
FLORENCIO Gómez LEvvA—2(2):1341
Asclepias hirtella a newly documented for
the flora of Texas by MAE DIE M ):14 95
—
Vechiau Community Hip-
popotamus Sanctuary in Grane by "e ASASE AND ue A,
OTENG-YEBOAH— 1 (1):549
Boerhavia triquetra var. intermedia (Nyctaginaceae); a
new combination and varietal status for the widespread
southwestern North American B. intermedia by RicHARD
SPELLENBERG—1(2):871
Botanical composition and multivariate analysis of vegeta-
tion on the Pothowar Plateau, Pakistan by Attar A. Dasn,
SHEHZADI SAIMA, MOHAMMAD ATHAR, ATTIQ-UR-RAHMAN, AND SAEED
A. Mauk-—1(1):557
Bouteloua (Poaceae: Chloridoideae: Cynodonteae: Boute-
louinae) del noreste de México by YoLANDA HERRERA ARRIETA,
PauL M. PETERSON Y Jesus VaLpés-Reyna—2(2):917
—
Carex breviculmis (Cyperaceae), new to the flora
North America by Lucas C. MAJURE AND CHARLES T.
Bryson—2(2):1381
Caryophyllaceae in North America: change over the
last 110 years by RicwarD K. RaseLer AND RONALD L. HART-
MAN—2(2):1285
Checklist of the vascular plants of Crawford County, Penn-
sylvania by CvwruiA M. Morton, Loree SPEEDY, AND JAMES K.
BisseLL— 1(1):631
gene zh the Mov UE of lupe County, Pennsylva-
ia by Cv EE SPEEDY—2(2):1449
dei dme ped BE T Rig County, Penn-
sylvania by ou M. oe AND Loree acm (2):1229
Ch
A Isl y A ki nos 1
NANI
K. WILLIAMS AND Dawn P. DERR— 1(1):431
iUc podus names” of never ahouai poo matese
boundaries of Thevetia [e Justin K. WILLIAMS AND EM K.
STUTZMAN—2(1):4
Chromosome number for PEE Ro American
species o y JERRY
G. CHMIELEWSKI— 2(1) :487
Cinco nuevas especies de a iphone i oe Rica by
ALEXANDER RODRIGUEZ Y ALEXANDRE K. MONRO 5
Clarification of Borreria anced indi gymno-
cephala Diodi I ii B ia fl i , and
e L ** mid: 3
! ) by PIERO G. DELPRETE
AND JOSEPH H. KIRKBRIDE, JR. —2(1):305
s to the genus Festuca (Poaceae: Poeae) in
and a key to the Central American species by
Sree J on SHIRE— 1 (2):827
hi + a D A L
Correct I
? re Li
R 2(2):1233
A (D. Al |
CaCl (rOdCede@) Dy
Bua areoles in the iu subtribe Clitoriinae by
MES A, LACKEY—1(2):110
pes fungiformis gen. and sp. nov, an eupolypod
fern (Polypodiales) in Early a. Burmese amber by
GEORGE ©. Poinar, JR. AND Ron BuckLey—2(2):1175
Description of Carex klamathensis (Cyperaceae), a rare
sedge of the Klamath Region of Oregon and California,
U.S.A. by BanBARA L. WILSON, RICHARD E. BRAINERD, LAWRENCE P.
Janeway, KEL! KUYKENDALL, DANNA LYTJEN, Bruce NewHouse, Nick
Bs STEPHEN MS AND ) PETER i SUR dA
perennial in a new coastal clade in Florida by Rosin B.
oe La
z
ri +
una comunidad de pie de o del norte el eee
by Davo Suárez Duque—2(2):12
Distribution and taxonomy of idos sericeum
and S. pratense (Asteraceae: Astereae) - RONALD L. JONES,
C. THEO WirseLL, AND Guy L. Nesom—2(1):7
Distribution of Gamochaeta eoo Gnaphalieae)
in Texas, Oklahoma, Arkansas, and Louisiana by Guy L.
Nesom—1(2):1125
Distribution of hanging garden vegetation associations on
the Colorado Plateau, USA by James F. FowLer, N.L. STANTON,
AND RONALD L, HARTMAN—1(1):585
Diversidad y distribución de la flora vascular acuática de
aulipas, México by Arturo Mora-Olivo v José Luis Vil-
LASENOR—1(1):511
PEDI and floristic compe shor of e bed dale in
Jan Brazil by
the forest fragment
REGINA H.P. ANDREATA, HAROLDO C. DE Lima, ANGELA i in Vaz,
José FERNANDO A. BAUMGRATZ, AND SHEILA R. PRoriceE—2(1):575
Early historical references to the Big Thicket, Texas > MICHAEL
. MACROBERTS AND BarBARA R. MacRosERTS—2(1):66
Eoépigynia burmensis gen. and sp. nov, an E Cre-
taceous eudicot flower (Angiospermae) in Burmese
amber by GtoncE Poinar, JR, KENTON L. CHAMBERS, AND RON
BUCKLEY—1(1):91
Eragrostis (Poaceae: Chloridoideae: Eragrostideae) in Colom-
bia by Paur M. PETERSON AND Dieco GiRALDO-CAÑAS—2(2 Yu
Estudios en n Apocynaceae Neotropicales XXX: tres
vas especies andinas de Mandevilla nume
Mee by J. HANG Morus (2): 853
Ect
de Ma ndevilla hirsuta cuatro nuevas especies by J.
FRANCISCO MonALES— 1 (2):85
Exotic species of Celtis (eed in the Flora of North
merica by ALAN T. Werremore—2(1):627
Five new varieties of Agapetes (Ericaceae) from India by D.
Banik AND M. SANJAPPA—1 (2):903
pes o of Saly dry tropical forest frag-
n Central Bahia, northeastern Brazil by Domincos
E Du SILVA CARDOSO AND LUCIANO PAGANUCCI DE Qua
ROZ—
Flow diagrams for plant succession in the Middle Tennes-
see cedar glades by J.M. Baskin, E. QUARTERMAN, AND C.C.
BaskiN—1(2):1131
Index: Vols. 1 & 2
Folia taxonomica 1. Validation of two taxa from northern
South America by CHRISTIAN FeuiLeI—1(1):143
Folia taxonomica 10. New species of Nautilocalyx (Gesneri-
Episcieae) fi heV | by CHRISTIAN
/ 4
FEUILLET—2(2):825
Polls taxonomica n eo of Varronia (Cordiaceae,
1503
lilicium guajaibonense, elevated to species rank and
compared with the subspecies of Illicium cubense (ll-
liciaceae) by Watter S. Jupp, J. RICHARD ABBOTT, AND ASHLEY
Morris—2(2):799
Index of names and types in West Han Gonolobinae
1 E A e
anew name, anda
tions “by CHRISTIAN FEUILLET—2(2):837
Folia taxonomica species of Passiflora subgenus
Passiflora RUM from the Guianas Dy CHRISTIAN
FEUILLET—1 (2):8
Folia tonomia : Passiflora david mano. anew
supersection Stipulata by pue FEUILLET—1(2):895
Folia taxon 4. Conspectus of Myriopus (Heliotro-
piaceae: Poeni in n Guiana Shield by CHRISTIAN
FEVILLET—2(1):263
Folia taxonomica 5. A new name for Passiflora heterophylla
(Passifloraceae) from Cuba and Haiti by CHRISTIAN FEUILLET AND
M. MacDoucAL—2(1):267
Folia taxonomica 6. Two new species of Besleria (Gesneri-
aceae) from the Venezuelan Guayana by CHRISTIAN FEUIL-
Er—2(1):269
Folia taxonomica 7. Two new species anda new section in
E
) from tl e Guayana y
pi enezuelan
CHRISTIAN FEUILLET—2(1):2
Folia taxonomica 8. mut tecta (Passifloraceae), a new
species in subgenus o ee from the Guianas by
CHRISTIAN daro ):28
Folia taxonomica 9. New species of Passiflora subg.
Decaloba (Passifloraceae) pan northern South America
by CHRISTIAN FEUILLET AND JOHN M. MacDoucaL—2(2):817
Four new combinations in Mosiera (Myrtaceae) from the
Caribbean by Anprew SaLrwon—1(2):899
Fourteen new combinations in Sedum (Crassulaceae) by
HibEAKI OHga—1(2):889
Further transfers of glandular-pubescent species from
Chenopodium subg. Ambrosia to Dysphania (Che-
nopodiaceae) by Sercel L. MOSYAKIN AND Steven E. CLEM-
ANTS—2(1):425
Galium anglicum (Rubiaceae) new for Texas and notes on
the taxonomy of the G. parisiense/divaricatum complex
by Barney L. Lirscoms AND Guy L. Nesom—1(2):1269
Generic realignments in tribe Potentilleae and revision of
Drymocallis (Rosoideae: Rosaceae) in North America by
(1):46
cosa aE
a nov (Asteraceae: Astereae), a shale
F
kK -|
Colorado by AL SCHNEIDER
Peaay LYON, AND Guy Nesom—2(2):771
Hawthorns (Crataegus: Rosaceae) of the Cypress
Hills, Alberta and Saskatchewan by J.B. Phipps AND R.J.
O'KENNON—1 (2):1031
Heuchera woodsiaphila (Saxifragaceae), a new species
from the Capitan Mountains of New Mexico by Patrick J.
ALEXANDER-2(1):447
Hypericum adpressum (Clusiaceae; new to Arkansas and the
Ouachita Mountains, U.S.A by C. THeo Wirst.L.—1(1):713
vt
on by AL LEXANDER KRINGS—2(1):139
apela an and nomenclatural notes for
Pycnanthemum (Lamiaceae) E KENTON L. CHAMBERS AND
HENRIETTA L. CHAMBERS—2(1):193
Infraspecific taxonom nomenclature of Eleocharis
acutangula (Cyperaceae) by Davin J. ROSEN, STEPHAN L. HATCH,
AND RICHARD CARTER —1 (2):875
inventory and ce i Agave (Agavaceae) species in
Jalisco, Mexico by G O HERNÁNDEZ-VERA, MIGUEL CHAZARO
BASAÑEZ, AND ERICKA oe P: —1(1):499
Ipomoea tehuantepecensis (Convolvulaceae): a new spe-
cies from the Isthmus of Tehuantepec, Mexico by Leticia
Torres-Cotín, RAFAEL Torres-Colín, Miriam P. RAMÍREZ DE ANDA,
AND J, ANDREW McDonaLo—2(2):793
James Hinton a 2006) by George S. HINTON AND BILLIE L.
TURNER M2) Te
HL CI
County, — U.S.A. by CAROL W. WITHAM AND PETER F.
ZKA—2(2):77
Kalmiopsis fragrans (Ericaceae), a new distylous species
from the southern Cascade Mountains of Oregon by RoserT
J. MENKE AND THOMAS N. Kave—1(1):9
Kruschke names in North American Crataegus (Rosaceae)
ud J. a Phipes—1 (2): D
h American C
correction and clarifications by J.B. PHipps—2(1):473
/D nt
La familia Nymphaeaceae en el estado de Nuevo León,
México by Carlos VeLazco-Maclas, RAHIM FOROUGHBAKHCH
URNAVAB, MARCO A. ALVARADO VAZQUEZ, Y GLAFIRO J. ALANIS
HO Anos
and sp. nov., a eudicot flower
in Dominican amber showing affinities with Fabaceae
subfamily nuc EU Py GEORGE O. POINAR, JR, KENTON
L. CHAMBERS, AND ALEX E. BRowN—2(1):463
Lawrence K. Magrath (1943-2007) by Barney Lipscome
—1(1):781
x cibus and notes on Baccharis riograndensis
eae: n by Gustavo HEIDEN AND ANGELO A.
Sencar 2l 1 ):29
fal H rents ir
pyroloides
(Ericaceae) by s w Forson A AND ena E Too (1): di
Lec
Eric A. Ld 1003
in Potentilla sect. Subvisco-
sae (Rosaceae) in Aae by BARBARA ERrTER— 1(1):47
Leptochloa (Poaceae: Chloridoideae) in Colombia by Nei
Snow, Paul M. PETERSON, AND DIEGO GIRALDO-CANAS—2(2):861
Lilaeopsis carolinensis (Apiaceae) a species new to Texas
and a key to Lilaeopsis in Texas by SrePHAN L. HATCH AND A.
TUCKER Stack—2(2):1497
1504
Los encinos (Quercus: Fagaceae) de Coahuila, Mexico by
José A. VILLARREAL Q, JUAN A. ENCINA D. y MIGUEL A. CARRANZA
P—2(2):1278
Luziola subintegra (Poaceae: Oryzeae), new to Florida
e United States by Jonn M. KUNZER AND MicHAEL J.
Bop.e— 2(1):633
Lyonothamneae, a new tribe in the Rosaceae (Rosales) by
Luc BrouiLLer—2(1):385
by Xavier ConNEJo—2(1):57
Micranthes nelsoniana var. porsildiana (Saxifragaceae),
the proper name at the varietal level by Luc BrouiL-
LET—2(1):395
Minuartia drummondii (Caryophyllaceae) and Gratiola
flava (Plantaginaceae) rediscovered in Louisiana and
Gratiola bos historically in Arkansas by MicHaeL H. Mac-
Roserts, BARBARA R. MACROBERTS, CHRISTOPHER S. REID, PATRICIA L.
FAULKNER, AND uen Estes—1(1):763
Minuartia macrantha (Alsinoideae: Caryophyllaceae):
den circumscription, geographical range, and
phylogenetic i E by RoNaLb L. HARTMAN AND RICHARD K.
Rasen —2(2):12
olombia
variety in North American Crataegus (Rosaceae) by J.B.
PHipps— 1 (2):10
Molecular analysis ad Cv. Lemore, a hybrid golden-
rod (Asteraceae) by Ebwaro E. ScHILLING, James B. Beck, PATRICK
J. Cau, AND RANDALL L. SMALL——2(1):7
Molecular phylogenetic analysis of the American Sti-
peae (Poaceae) resolves Jarava sensu lato polyphyletic:
evidence for a new genus, Pappostipa by KoNsrANTYN Rom-
ASCHENKO, PAuL M. PETERSON, RoBERT J. SoRENG, NURIA GARCIA-JACAs,
OKSANA FUTORNA, AND ALFONSO SUSANNA—2(1):16
Monarda lindheimeri (Lamiaceae): new to Arkansas by
WALTER C. HOLMES AND Jason R. SiNGHURST—1 (1):717
Muhlenbergiinae (Poaceae: Chloridoideae: Cynodonteae):
from northeastern México by PAUL M. PETERSON, JESUS VALDÉS-
REYNA, AND YOLANDA HERRERA ARRIETA—1(2):933
Munrochloa, a new genus (Poaceae: Bambusoideae) with
new combination from India by MuktesH Kumar AND M.
RemesH—2(1):373
Neocalyptrocalyx morii (Capparaceae), a new species
from central ~ Guiana by Xaver CORNEJO AND HUGH H.
ILns—2(2):80
Neptunia rid (Fabaceae: Mimosoideae) rediscovered in
Texas by ALFRED RICHARDSON AND Ken Kinc—2(2):1491
New and noteworthy records of several non-native vascular
plant ips in Arkansas by Brerr E. Serviss AND James H.
Peck—2(1):637
New NU and new synonymies in the genus
Spermacoce (Rubiaceae) for the flora of Goiás and
Tocantins (Brazil) and the flora of the Guianas by Piero G.
Da RETE— 1(2):1023
New combinations in Dudleya (Crassulaceae) and Ribes
- (Grossulariaceae) by NANCY n ege 2):1015
R. Morin—1(2):1017
f Texas 2(2)
Al | "ES P e AA: " f. fec
Saxifragaceae) in North America by Luc BrouILLET AND RICHARD
CRNALL— 1(2):1019
New combinations in oo (Philadelphaceae) by
James HENRICKSON—1(2):90
New combinations in the —— vascular plant flora by
LAE eve oe Daye E Munn eH :433
bamboos of pu (P.
y C M. A. canna (1):13
New ne discoveries i Sonor Islands, Gulf of California,
cobyB R, RICHARD S. FELGER, HUMBERTO ~
a AND ADRIAN DM neue (2):12
n records of Poa (Poaceae) and Poa enu. a new
cies endemic to Chile by RoBerT J. SoRENG AND PAUL M.
o 2(2):847
New records of wetland and riparian plants in southern
California, with recommendations and additions to the
National List of Plant Species that Occur in Wetlands by
ip Steve Boro—1(1):719
RICHARD E BERNER JR
by Bruce K. HOLST AND
Marja Luca KA ): 297
New, corrected, and interesting records for the Kansas vas-
cular flora by Cates A. Morse, CRAIG C. FREEMAN, AND RONALD L.
McGREGOR—1(1):753
Nomenclatural changes in Nemacladus (Campanulaceae)
by Nancy R. Morin—2(1):397
Nomenclatural notes in North American Potentilla (Rosa-
ceae) by BARBARA EnrrER—2(1):201
Notes on Conocarpus erectus (Combretaceae) in the Baja
California Peninsula, Mexico by José Luis León-DE LA LUZ AND
RAYMUNDO DOMÍNGUEZ-CADENA—1 ( 1) ae
Notes on Lechea maritima
A. SORRIE AND ALAN S. WEAKLEY—1 (1):367
= on Phragmites australis (Poaceae: Arundinoideae)
n North America by Kristin SALTONSTALL AND DoNALD Hau-
d ):385
Notes on Texas lc (Orchidaceae) by PauL MARTIN
BRowN—1(2):12
Notes on the du mres of Xylothamia (Asteraceae:
en by inl S rd (1):145
+ Ail
) by BRucE
records. Ili by ud E Brown, Eric L. Kem}, Davio J. Rosen, AND
JOE a 12): 12
otes on the G dumosa (Eri ) by
es A Somme, AND DAN S. WO (1): 333
United States
by Bruce e SORRIE AND RICHARD J. LEBLONC—2(2):1 3
Noteworthy collections from the Yazoo- Prana Delta
egion of Mississippi by pus SkOJAG, Jr, CHARLES T. BRYSON,
AND CHARLES H. WALKER ll —1(1):769
M T En from m Florida. VIII by Loran C. ANDER-
—1(1):74
S
=
TEAM. va Sm plant collections from northwest
Louisiana by CHRISTOPHER S. REID, PATRICIA L. FAULKNER, BARBARA R.
El (1):643
Novae Gesneriaceae neotropicarum XV: Kohleria hyper-
trichosa, a new species of Gesneriaceae from north-
Index: Vols. 1 & 2
western Ecuador by JOHN LITTNER CLARK AND LAURENCE E.
Skoc— 9
Observation on vegetation changes in Cajun Prairie, a coastal
prairie flora in southwest Louisiana by CHARLES ALLEN AND
Sara THAMES—1(2):1141
Paspalum vaginatum (Poaceae), a new threat to wetland
diversity in southern Caiifornia oy RICHARD E. RIEFNER, JR. AND
J. TRAVIS CoLUMBUS— 2(1):743
Phylogenetic analyses of the genus Baptistonia (Orchi-
daceae: Oncidiinae) sensu lato vem on morphological
characters by Guy R. CHiron-—1 (2):9
iocis anda of bug America ERE ¡Prunus
FY al id
pad sequences by JoserH R. ROHRER, MEGAN A. O'BRIEN, AND
JULIE A. ANDERSON—2(1):401
Phytogeography of the Big Thicket, East Texas a 2n H.
MAcRoBERTS AND BARBARA R. MacRoserts—1(2):1 1
Praxelis clematidea (Asteraceae), a genus and species new
forthe flora of North America S RICHARD ABBOTT, C. LEÁNN
WHITE, AND S. Barry Davis—2(1):6
sut Mn cate c habitat in the
e Mountains using GIS: evidence of a new
eer population by TimotHy C. MuLLET, FRED ARMSTRONG,
BENJAMIN ZANK, AND CHRISTOPHER M. Ritzi—2(1):677
R lus ficaria (R | ), naturalized in Texas by
Guy L. Nesom—2(1):741
Rare and endemic plants of the Big Thicket: a phytogeo-
graphical analysis by MicHaeL H. MAcRoserTs AND BARBARA R.
MacRoserTs—2(2):1475
Recognition of Lechea pulchella var. ramosissima (Cista-
ceae) by Bruce A. SORRIE AND ALAN 5. WEAKLEY—1(1):369
Recognition of three taxa of eastern North American "Wald-
steinia" and their appropriate names when incorporated
into Geum (Colurieae: Rosaceae) by ALAN S. WEAKLEY AND
Kanchi N. GANDHI—2(1):414
Redescubrimiento de Axiniphyllum sagittalobum (Aster-
aceae) en la Sierra Madre del sur y notas de las especies
de este género que habitan en el estado de Guerrero,
México by ALBERTO GONZÁLEZ-ZAMORA, IsoLDA LUNA-VEGA, Y JOSÉ
Luis VILLASENOR—1(1):491
Re- AO and typification of Bana "au tolia; ma
, and M
an adventurous history by JAMES L. Lurew—2(1):243
Rediscovery of Eleocharis kleinii (Cyperaceae), an o
looked species from the highlands of FA Brazil Dn ^
Trevisan, D.J. Rosen, AND l.l. BOLDRINI—1(2):1
Rediscovery of Mirabilis hintoniorum Mycenaceae) a
striking four-o'clock endemic to the Sierra de Coalcom
Michoacán, México by Mark FISHBEIN AND VICTOR W. Med
MANN—2(1):547
Re-examination of Muhlenbergia capillaris, M. expansa,
and M. sericea (Poaceae: Muhlenbergiinae) by DaNNv J.
USTAT ON AND IE M aieo (1):85
A TE ee
, el caso de la
cuenca del río Magdalena by V. Avita- AKERBERG, B. GONZÁLEZ-
HipALGO, M. Nava-Lopez v L, ALvEIDA-LENERO—2(1):605
Reinstatement of Sagittaria macrocarpa (Alismata-
1505
ceae) by Bruce ji SORRIE, BRIAN R. KEENER, AND ADRIENNE L.
bel a Lodi
Tanta (Dial: VAFRAÁ
and Arizona and a review N of Houstonia subgenera and
sections by Enwaro E. TerreLL—1(1):109
Review of Crataegus series Apricae, ser. nov., la - flava
on by J.B. Phipps AND K.A. pd ar
Revision o | loideae)
in the Vm Indies by A ESAME Krincs—2(1): 95
Revision of Lobelia sect. Galeatella (Campanulaceae: Lobe-
lioideae) by Thomas G. Lammers—1(2):789
Revision of Siphonandra (Ericaceae: Vaccinieae), a genus
endemic i ja and Bolivia by James L. LUTEYN AND EDGARDO
M. Ortiz—2(1):249
Revision of a (Rubiaceae: Gardenieae) from the
states of Goiás and Tocantins and a new species endemic
white-sand areas in the Brazilian Cerrado by Piero G.
DELPRETE——2(2):98
c fc “e y ]
TOPHER aS. REID, eran L. tee MM R ara AND
Mae H. Mii 1251
Hedyotideae
(Rubiaceae) Thecagonum, Neanotis, Denm Kohau-
ntodon, EDWARD E. TERRELL
J
AND 6 Heo. n (1): ae
++
(Asteraceae) in Highland Balochistan, Pakistan by Sar-
FRAZ AHMAD, SHAMIM GUL, MUHAMMAD ISLAM, AND MOHAMMAD
ATHAR—1(1):569
Seed germination response of Zizania texana (Poaceae:
Oryzae) to soil inundation by M.L. AtexanbeR—2(1):673
Senecio cymbalarioides, S. subnudus, and S. subpelta-
tus, history of a muddle (Asteraceae: Senecioneae) by J.F.
VELDKAMP—2(2):1215
Silene flos-cuculi ssp. flos-cuculi (Caryophyllaceae) and Eu-
phorbia peplus (Euphorbiaceae), new to North Carolina
by Derick B. PoINDExTER—2(1):649
Species richness of vegetational areas of Texas: a first ap-
proximation by MicHAEL H. MACROBERTS AND BARBARA R. Mac-
DO rear e
I ro.
(Cyperaceae)
L Mes (UI) r
in the United States by Paur M. McKenzie, S. GALEN SMITH, AND
MARIAN A 0) id
Struc Kaieteur National
Park, Guyana by CAROL L. Keorr—2(1): 521
Talinum rugospermum (Portulacaceae) new to Oklahoma
TRICIA L. FAULKNER, BARBARA R. MacRos-
ERTS, AND MicHAEL H. MacRoserts—1(1):777
Taxonomic overview of the Heterotheca fulcrata complex
(Asteraceae: Astereae) by Guy L. Nesom—1(1):21
Taxonomy and nomenclature of Taxus (Taxaceae) by RICHARD
m. ed ub AS
by CHRISTOPHER S. REID, PA
\
ZBromopsis romu dd Genea in British Columbia,
Canada by Jerrery M. SAARELA—2(1):323
x i pi I D Ig; Marti
1506
Garcia, ec jer by WALTER S. JUDD, GRETCHEN M.
lONTA, TEODORO CLA DAN SKEAN, JR. —2(1):3
The B Thicket Born as fers wie habitat > ls
CHAEL H. MACROBERTS AND BARBAR RoBERTS—2(1):66
The Big Thicket: pristine up. or REIN
understudied? by MicHAE. H. MacRoserts AND BARBARA R.
MacRoserts—2(2):1291
The Cactaceae of the N | ipal
OR Hn l
in du y la, nie Ge
f Texas 2(2)
Three new species of Ixora (Rubiaceae) from the state of
Tocantins, Brazil by Piero G. Detprere—2(1):455
Tree canopy research and student experiences using the
doubled rope climbing method by Courtney M. KILGORE,
HaroLD W. KELLER, Sroner E. EveRHART, ANGELA R. SCARBOROUGH,
KENNETH L. SNELL, MELISSA S. SkRABAL, CHARLY POTTOREF, AND JOSEPH
S. oe. eid
and sp. nov, a fossil
3
ce in Dominican amber by Pun S Ponar JR,
RAES CALVENTE AND REGIN LENA PorscH Aeneas 529
ipii iur s Bic (Ebenaceae) in Meso-
rica by MitcHett C. Provance, IaNACIO GARCÍA RUIZ, AND
Mia SANDERS—2(2):1009
The genus Abronia (Nyctaginaceae) in Colorado, with notes
on Abronia bolackii in New Mexico by JENNIFER ACKERFIELD
AMD WILUAM i Aae 200 id 9
T1 J (iR
America by Nu Ma AND ANTHONY R. BRACH—1(1):357
The species of Fumaria (Papaveraceae) in Texas, including F.
densiflora, first records from North America by Monique
REED, RICHARD V. LANSDOWN, AND Tim C.G. RicH—2(2):1347
The taxonomy of Carex trisperma (Cyperaceae) by CHAD D.
KiRscHBAUM— 1 (1):389
The taxonomy of North American loti (Fabaceae: Loteae):
new names in Acmispon and Hosackia by Luc Broun-
Ler—2(1):387
The vascular flora of a woodland park site in east Harris
County, Texas by jn HaArcomee, l.S. Ers, W.W. PRUESS, AND
L.E. BRowN—1(1):69
The vascular flora of an Prairie: a Coastal Prairie remnant in
Brazoria County, Texas by Davip J. RostN— 1(1):679
The vascular flora of the Chunky River (Mississippi) by Lucas
C. MAJURE— 1(2):1179
The vascular flora of the Hancock Biological Station, Murray
State University, m County, Kentucky by Raten L.
THompson—1(1):60
fields, Ge orgetown,
T h 1c vascular flora of
South Carolina: a 39 year comparison by Richard STALTER,
Nu as AND MM cuia. (1):66 665
HH
p NE Olyreae), a new species from ene Gui-
ana by EMMET J. JubziEWICz AND SOL SEPsENWOL—1 (1):1
Thomas Walter Typification m Il: the known Walter types
by DanteL B. Wagp—1(1):40
Thomas Walter Typification e Ill: lectotypes and neo-
types for 20 Walter names, as recognized in the Fraser/
Walter herbarium by Danet B. Wazo—1(1):425
Thomas Walter Typification Project, IV: neotypes and epitypes
f names of genera A through C by Danie B.
Wanp—1(2):1091
Thomas Walter Typification Project, V: neotypes and epitypes
for 63 Walter names of genera D through Z by DanteL B.
WarD—2(1):47
Thomas ee Project, VI: neotypes for an ~
tional 18 Walter names by Danie. B. Waro—2(2):12
Three new names in the Caribbean flora by ALAIN n Lio-
GIER—2(2):1165
Konus L n AND ALEX E. BRowN—2(2):1
Two new Cotoneasters (Cotoneaster: MM from
Yunnan Province, China by Jeanette FRYER AND BERTIL
"MOTA Us
J 4 PA
Astereae) from southern sa i ANGUS Aero Seba
AND ILS! log BoLDRINI—2(1):45
Two new species of Cyperaceae from Peninsular India by M.A.
WADOOD KHAN AND P. LAKSHMINARASIMHAN—2(1):379
Two new species of Gratiola (Plantaginaceae) from east-
ern North America and an updated circumscription
for Gratiola neglecta by Dwayne Estes AND RANDALL L.
SMALL—1(1):149
wo new species of Guadua (Bambusoideae: Guaduinae)
from Colombia and Bolivia by Ximena LONDONO AND ENEIDA
ZURITA—2(1):25
Two new subspecies of Sidalcea dd (Malvaceae) in
California a Steven R. HiuL.—2(2):78
A 1 oan TA
px e México by ARISA GARCÍA-MENDOZA, Gao
ANO Y ÁneL SALAZAR ÑO d € 1):79
le Costa Rica y
amá by D. ARMANDO SOTO AND A.K. Monro—2(1):41
Understand Oligactis (Asteraceae: Liabeae): the true
ntity of O. r m and O. volubilis by Dieco G.
0) 12
+ J
Var lation Hi P
. NESOM AND CALEB A. Morse—1 (1):35
Vascular flora of C. Bickham- E i River Education
and Research Park, Caddo Parish, Louisiana: an oxbow
Sim (Act. Act vb nr
y PUCICdC/ VY MUY
ratios by MicHAEL H. ipei BARBARA R. MACROBERTS, AND
ARY M. HANSON—2(2):13
Vascular flora of the p M Preserve, Ellis County, Okla-
homa by Bruce W. HOAGLAND AND Amy K. Burdop—1(1):655
Vascular plant species/area relationships (species richness)
in the West Gulf Coastal Plain: a first approximation by
MICHAEL H. e n BARBARA R. MACROBERTS, AND ROBERT G.
KaLiNSKY—1 (1):57
oni plani ype PENER, in E University of British
IBC) by Jerrery M. SAARELA
, LINDA LIPSEN,
es Y M. SAYRE, AND EE Vd MA (1):437
Vicia lathyroides (Fabaceae): new to the flora of Texas by
SONNIA Hitt, RurH Loper, JASON R. SINGHURST, AND WALTER C.
HoLmes—1(2):1253
{VW J Pr rl f,
a new 'el low
os
“the Florida panhandle by Loran C. ANDERSON A AND ROBERT
RAL—2(1):1
INDEX OF 324 AUTHORS:
VOLUME 1 (2007) AND VoLUME 2(2008)
Thank you for choosing Journal of the Botanical Research Institute of Texas.
Abbott, J. Richard —2(1): e ~ 799
Ackerfield, Jennifer —2(1):4
mad, Sarfraz—1(1):569
Alanis Flores, Glafiro J.—2(1):593
Alexander, M.L.—2(1):673
Alexander, Patrick J.—2(1):447
ees. Leñero, L.—2(1):60
, Marco A -am 593
eran Julie A—2(1):40
Anderson, Loran C.—1(1):741; 2(1):1
Andreata, Regina Helena ae 529;
2(1):575
Armstrong, Fred—2(1):677
Asase, Alex—1(1):549
Athar, Moham d 569
Ávila- eee a (1):60
Baden, iii 665
Banik, D.—1(2):90
Basañez, "iud Cr) 499
Baskin, CC—-1(2):1131
a
Baumgratz, José Famandd A—2(1):575
Beck, James B.—2(1):7
Bissell, James K.—-1(1):631
Bodle, Michael J.—2(1):633
Boldrini, Isi lob—1(2):1 119; 2(1):45
Boyd, Steve—1(1):719
Brach, Anthony R.—1(1):357
ale Richard E.—1(1):69
vo, Ángel Salazar—1 (1):79
a Luc—1(2):1019; 2(1): m 387, 395
Brown, Alex E.—2(1):463; 2(2):1167
Brown, Larry E.—1(1):693; m 1255; 2(1):
651; 2(2):1481
Brown, Paul Martin—1(2):1 2
Bryson, Charles ue 2(1):517;
2(2):1381
Buckley, Ron—1(1):91; 2(2):1175, 1183
Bush, Catherine M.—2(2):1193
Buthod, Amy K.—1(1):655
ss in d
de Moraes—1(1):529
n Domingos Benício Oliveira Silva—
2(1):
Carranza P Miguel A —2(2):12
(2
Carter, Rchare—1(0 779; 1(2): e 2(1):517
—2(1):1
Chambers, Henrietta
ee Kenton L—1(1):91; 2(1):193, 463;
2(2):1167, 1183
Chiron, Guy R.—1(2):913
Chmielewski, Jerry G.—2(1):48
Christenson, Eric A.—1(2):1001, 1003; 2(1):285
Clark, John ibm 19
Clark, Lynn G.—1(2):84
J. Bot. Res, Inst. Texas 2(2): 1507 — 1508. 2008
Clase, Teodoro—2(1):35
Clemants, Steven e
Columbus, J. Travis—2(1):74.
Cornejo, Xavier—2(1):57, 61, pn 2(2):807
Darbyshire, Stephen J.—1(2):827
—1(1):
Dasti, Altaf 557
Davenport, LJ —1(1):103
Davis, S. Barry— m
ebelica, Anica—2(2):13
Delprete, Piero Pd 1023; 2(1):305,
455; 2(2):98
Derr, Dawn P—1(1):43
Domínguez- n — ee
Duque, David Suárez—2(2):1295
Dvorsky, K.A.—1(1):171; 2(2):1 101
Edwards, Adrienne L.—1(1):345
Elsik, |. Sandra—1(1):693; 2(1):651; 2(2):1481
Elven, Reidar—1(2):81 1; 2(1):433
Ely, Joseph S.—2(2):1309
Encina D. , Juan A.—2(2):12
Ertter, Barbara—1(1):46, n am 201
Estes, Dwayne—1(1):14
Everhart, Sydney E id
Faulkner, Patricia L.—1(1):763, 777; 1(2):1251;
2(1):643
Felger, Richard 5.—
Feuillet, Chistian —1() 143; 1(2):819, 895;
- ):263, 267, 269, 275, 281; 2(2):817,
pie
Fish on Mark—2(1):547
Fisher, Amanda E.—1(2):847
Ela Berrios, Ericka—1(1):499
nseca Vaz, Angela S.—2(1):575
: EU James F—1(1):585
a Jeanette—2(1):53
utorna, Oksana—2(1):165
"ede Kanchi N.—2(1):4
Giraldo-Cañas, Diego—2(2):861, 875
Goldman, Douglas H.—2(2):1337
G yva, Juan Floren i0—2(2):1341
hez, Maricela—2(1):495
;3onzález, Isaac Andrade—2(2):1341
González-Hidalgo, B.—2(1):605
González-Zamora, Alberto—1(1):491
Gornall, Richard—1(2):1019
Gul, Shamim—1(1):569
Gustafson, Danny J.—1(1):85
Gutiérrez, Diego G.—2(2):1207
Hanson, Gary M.—2(2):1389
Harcombe, Ec. 2(1):651
Hartman, Ronald L.—1(1):585; 2(2):1225,
Hatch, Stephan L—1(2): te 2(2):1497
Hauber, Donald—1(1):38
Her 1):49
Es Arrieta, A B 933; dns 917
Hiil, Sonnia—1(2):12.
Hill, Steven mede
Hinton, George S.—1(2):1277
Hoagland, Bruce W.—1(1):655
Holmes, Mime 717; 1(2):1253
dus Bruce K.—2(1):29
Robin B.—2(2): ae
ns Bertil—2(1):53
Iltis, Hugh H.—2(1):61, 75; 2(2):807
lonta, Gretchen M.—2(1):35
Iruegas, Rubén—2(2):1341
Islam, Muhammad—1(1):569
Jacques-Hernandez, Cuauhtémoc—1(1):79
Palaos ere (1):129
Janeway, Lawren —1(1):69
Jennings, ud F—2(1):419
Jhangiani, Shalini N—2(2):1219
Jones, Ronald L.—2(1):731
Judd, Wolter S— 12AT n 2(1):35; 2(2):799
1):
Judziewicz, Emm
3
rd ala a mi. 577
Kav —2(1):297
ue zii N— sn 9
Keener, Brian R.—1(1):103, 345
255
Kilgore, Courtney M.—2(2):1309
Kincaid, erem
la. Bere (2):14
eJr p m 2(1):305
ise beu. Chad D.—1(1):389
Kral, Robert —2(1):1
Krings, Alexander—2(1):95, 139
DM Keli—1(1):69
Lackey, James A.—1(2):1101
Laegaard, Simon—1(2):841
1508
Lakshminarasimhan, a 379
León-de la Luz, José Luis—1(1):487
Liggio, Joe—1(2):125
de Lima, Haroldo Pies
Liogier, Alain H.—2(2):116
Lipscomb, Barney meer 1(2):1269
López-Muraira, Irma—2(2): 1341
Luna-Vega, Isolda—1(1):491
Luteyn, James L.—2(1):207, 243, 249
Lyon, Peggy—2(2):771
Lytjen, Danna—1(1):69
Ma, Jinshuang—1(1):357
MacDougal, John M.—2(2):817
MacRobe :577, 763, 777;
1(2):1149, 1251; 2(1):643, or 661, 665;
2(2):1291, bib dd 1475,
MacRoberts, Mi —1(1):5 e 763,777;
1(2):1149, 1251; 2(1):643, 651, ~ 665;
2(2): P a 1373, 1389, d
Madulid, Domingo A.—1(2):8
Majure, Lucas C.—1(2): 1179; aes 1381
557
rts Barhara R-—
Mckenzie, in M—1(1): 457
da Olivo, Arturo—1(1):51
orin, Na amic: 1015, M 2(1):397
ed
Morton, Pus Mcd 1(2):1229;
2(2):144
Mosyakin, ud L.—2(1):425
Mullet, Timothy C.—2(1):677
Murray, David F.—1(2):811; 2(1):433
Nava-López, M.—2(1):605
Nelson, John B.—2(2):761
Nesom, Guy L.—1(1):21, 145, 351; 1(2):891,
1125, 1269; 2(1):731, 741; 2(2):771
Newhouse, Bruce—1(1):69
O'Brien, Megan A.—2(1):401
Oteng-Yeboah, Alfred en 549
Otting, Nick—1(1):69
Peck, James H.—2(1):63
Peterson, Paul M.—1(1):35; 1(2):841, 933;
2(1):165; 2(2):847, 861, 875, 917
Phipps, J.B.—1(1):171; 1(2):1005, 1011, 1031;
2(1):473; 2(2):1101
f Texas 2(2)
Poinar Jr, George O.—1(1):91, 449; 2(1):463;
2(2):1167, 1175, 1183
Poindexter, Derick REM E
du Charly—2(2):13
Pournavab, Rahim A made
B
Powell, A. (rig :685
Provance, Mitchell C.—2(2):1009
Pruess, Warren W.—1(1):693; 2(1):651;
2(2):1481
Quarterman, E.—1(2):1
de Queiroz, Luciano Paganico amns A
Quijada-Mascareñas, Adrián—1(2):12
Rabeler, Richard K.—2(2):1225, 1285
Rahman, Attiq-ur—1(1):557
w
N
Rajkumar, S.— 9
Ramírez de Anda, Miriam ee
cr oe EOM
—1(1):763,777;1(2):1251;
p ):6 43
Remesh, M.—2 iy
Rich, Tim C.G.—2(2):13
ee m
Riefner Jr. mens 2(1):743
Ritzi, ariel M.—2(1):67
Ro Ma DAD 373; a 811
r—2(2):995
ove Joseph R.—2(1):401
Romaschenko, Konstantyn—2(1):165;
2 1233
Romero-Morales, SUL eos f 203
Roque, Nádia—2(2
Rosen, David J—10* 679, 779; 1(2):875,
1119, 1255
ríguez, Ale
Ruiz, Ignacio García—2(2):1009
Saarela, cmn 2(1):323
—1(1):55
—1 (2): 841
7
elo Alberto—2(1): :45, 291
Er Sol —1(1):1
Serviss, Brett E—2(1):637
Singhurst, Jason R.—1(1):717; 1(2):1253
):68
Skog, peice ae
:1497
Small, Randall L.—1(1):149; 2(1):7
Smith, Marian—1(1):457
Smith, S. Galen—1(1):457
Snell, Kenneth L.—2(2):1309
Snow, Neil—2(2):861,
Soreng, F "n ; 2(1):165; 2(2):
Sorrie, Bruce A.—1(1):333, 345, 367, 369;
2(2):1353
L
Soto, D. Armando—2(1
T Loree—1 (1): E T 1229; 2(2):
e Richard—1(2):871
Spjut, Richard W.—1(1):203, 291
Stalter, Richard—1(1):665
Stanton, N.L.—1(1):585
Stapleton, CM.A.—1(1):13
Steinmann, Vota 547
Susanna, Alfonso—2(1):165
Téllez-Pimienta, Kathia Georgina—2(1):495
Terrell, Edward a 109, 373
—2(1):68
A.
Torres-Colin, Leticia—2(2): 793
Torres-Colin, Rafael—2(2):793
Trevisan, R.—1(2):1119
Triplett, Jimmy K.—1(2):847
Trock, Debra K.—1(1):97
Turner, Billie Rd
Tushar, KV.—1(1):12
Tyrrell, Christopher a4 (2):847
Udayan, P5.—1(1):121
Valdés-Reyna, Jesus—1(2):933; 2(2):917
Velazco-Macías, Carlos—2(1):593
Villaseñor, José Luis—1(1):491, 511
Walker Il, Charles H.—1(1):76
Walker, Suzanne aeu MM
2):1481
Ward, Daniel B.—1(1):407, 425; 1(2):1091;
2(1):475; 2(2):127
Weakley, Alan S.—1(1):333, 367, 369; 2(1):414,
621
White, C. LeAnn—2(1):621
White, Matt—2(2):1495
hi
Wilder, Benjamin T.—1{2):1203
M Justin K.—1(1):431; 2(1):489;
(
2
Wilson, Barbars L.—1(1):49, 69
Witham, Carol W.—
Witsell, C. Theo—1(1):713; 2(1):731
Wu, Yuyang—1(1):449
Yao, George C.—1(2):815
Zank, Benjamin—2(1):67
Zika, Peter ba Ene 69; 20) 775
1(2):1157
Zurita, Eneida—2(1): 25
Index: Vols. 1 & 2 1509
BOTANICAL NAMES AND SUBJECT INDEX:
VOLUME 1 (2007) AND VoLUME 2 (2008)
New names (454) in bold face.
Abandoned rice fields—1(1):665 cct 392
Abronia—2(1):419 rigidus—
argillosa—2(1):419 Moa je
bolackii—2(1):419 utahensis—2(1):392
carletonii—2(1):419 wrightii—2(1):392
elliptica—2(1):419 a HE -veneris Herbaceous Association—
fragrans—2(1):419
glabrifolia—2(1):419 kegopoor— 1 933, 938
nealleyi—2(1):419 AFLP—1(1):389
Acaena elongata—2(1):605 coh 549; 2(2):843, 1215
capulco, Mexico—2(2):1009 Agalin
Acer p ricata—2(2):1353
negundo var. mexicanum—2(1):605 filifolia—2(2):1353
palmatum—2(1):637 Agapetes—1 (2):903
Achillea acuminata var. tipiensis—1 (2):903
alpina subsp. multiflora—2(1):442 flava var. nagensis—1(2):905
mill efolium—2(1): 605 megacarpa var. lohitensis—1(2):905
Achnatherum—2(1):165 odontocera var. edo 908
Acmispon—2(1):387 salicifolia var. glanduliflora— 1 (2):9
americanus var. o te 388 lods iss iN 130
argophyllus—2(1):38 pauciflo im 131
var. eR d ):388 Agathis—1(1):44
var. argenteus—2(1):388 iiec AU E 499; 2(2):1341
ar. fremontii—2(1):389 Agave—1(1):7
Bou 389 xsonenios- (1):79
var, A 389 subg. Littaea—1(1):79
var. notitius—2(1):3 Marginatae Au ):79
cytisoides—2(1):389 montium-sancticaroli—1(1):79
dendroideus—2(1):389 aa var. “azul’—2(2):1341
var. traskiae—2(1):389 Alabama, USA—1(1):10
var. veatchii—2(1):38¢ Alaska, USA—1(1 ne
distichum—2(1):389 Alberta, Canada—1(2):1031
flexuosus—2(1):389 Alcatraz Island, Mexico—1(2):1203
rad Aletris lutea—2(2):1353
evialatus—2(1):390 Alismataceae—1(1):345
snditorus 2 (1):39 Alocasia—1(2):815
oo ):390 nycteris—1 (2):8
e ):39 Alsinoideae anephylacse 20 1225
haydonii— 211390 Amaranthaceae—2(2):12
Diis mid Amaranthus—2(2):1219
a orbicularis 211} 390 blitum—2(1):637
tad 2(1):39 uud emarginatus—1(1):719
oe e Amazon—2(1):25
var. biolettii—2(1):391 Amazon Basin—2(1):455; 2(2):9
maritimus AS Amazonas, Venezuela—2(1): 269, 275; 2(2):825
sec ):39 Amber—1(1):91, 449; 2(1):463; 2(2):1167, 1175, 1183
ar. o 391 Amphiachyris—1(1):145
a 2(1):391 Anatherostipa—2(1):165
nevadensis—2(1):391 Andes—1(2):853; 2(1):25
var. Me 391 Andhra Pradesh, india—2(1):379
niveus—2(1):39 Angiospermae—1(1):91
ieee Anisocapparis—2(1):62
oroboides—2(1):391 speciosa—2(1):65
procumbens—2(1):39 Anotis—1(1)
var. jepsonii—2(1):392 Antennaria—2(1):487
1510
Antilles—2(1):463; 2(2):1167
Antioquia, Colombia—2(1):57
Aphanelytrum—1(2):841
ruvianum—1(2):842
pro i 841, 844
uu. ene
renenacee— 131 1(2):853, 859; 2(1):95, 139, 489;
2):1341,
Apocynoideae TE (1): c 1(2):853, 859
Aquatic vascular plant diversity—1 (1):5
Aquilegia Miis Herbaceous pe 585
ed petra
sp. septentrionalis—211) :438
ea umbrosa—2(1 ~
Arabis canadensis an ):64
Araceae—1(2):81
Araucariaceae—1 i ):449
Archeology—2(2):1291
Argentina—2(1):61, 305; 2(2):847
Aristida
gibbosa—2(1):309
gibbosa complex—2(1):309
liebmanii—2(1):309
marginalis—2(1):309
orizabensis—2(1):309
sorzogonensis—2(1):309
f. e E 322
sorzogonensis—2(1):309
Aristideae Poaceae) 201): 309
Aristolochia— 43
anukuensis—1(1):143
peltato-deltoidea—1(1):143
schultzeana—2(2):1 337
Aristolochiaceae—1(1):143; 2(2):1337
Arizona, USA—1(1):47, 109, 351, 483
ORDEN USA—1(1):713, 717, 763; 1(2):1125; 2(1):637, 761
Artemisia kruhsiana subsp. alaskana—2(1):443
Arunachal Pradesh, India—1(2): ue
Arundinoideae (Poaceae)—1(1):38
Asclepiadoideae (Apocynaceae) 201): 95, 139; 2(2):1341
Asclepias hirtella—2(2):14
Asia—1(1):91, 97, 121, iS 135, 291, 449, 557, 569, 903;
1(2):815; 2(1):53, 373, 379, 433; 2(2):1175, 1183
Aspleniaceae—1(1):103
Asplenium—1(1):103
xebencides—1(1):103
xAsplenosorus ebenoides—1(1):103
Aster—2(1):7
ic om 21,145, 351,491, 569; 1(2):891, rer 2(1):7,
5,
487,621, 731; 2(2):771, 811, 1207,
Astereae UM EE UO: 145, 351; 1(2): a —
45, 291, 731; 2(2):771
Atheropogon ed :
Atlantic Coastal Plain, USA—1(1):5
Atlantic Coastal Ridge, Florida, usas e
:529; 2(1):
marica
Atlantic Forest, South A
Atriplex polycarpa—1(1):719
Lal Datamirnl D a. een
Journal Or
f Texas 2(2)
Austrostipa—2(1):165
Axiniphyllum
corymbosun—1(1):491
pinnatisectum—1(1):491
sagittalobum—1(1):491
Baccharis
apicifoliosa—2(1):45
sect. Caulopterae—2(1):45
flexuosiramosa—2(1):48
dnd Brazil—2(1):551; 2(2):81 1, Pad
Baja California Sur, Mexico—2(2):10
Baja California, irm > p 1(2):871
Balanophoraceae—2(2):11
Balochistan, Pakistan—1 ns
Bamboo—1(1):1, 135; 2(1):25, 373
Bambusa ritcheyi—2(1):373
Bambusoideae (Poaceae)—1(1):1, 135; 1(2):847; 2(1):25,
373
Baptistonia—1(2):913
Barbieria—1 (2): Ps 01
Barrens—2(2):14
Barro Colorado and Panama—2(1):521
Bejaria—2(2
acem nae ):1
Bejarieae Ec ee 193
Besleria—2(1):269
gibbosa sect.
lares—2(1):269
neblinae—2(1):269
yatuana—2(1):271
Beta vulgaris subsp. maritima—1(1):719
Bifid petals—2(1):57
Big Bend National Park, USA—2(1):68
uos Texas, USA—1(2):1149; 205 661,665; 2(2):1291,
Neobesleria subsect. Axil-
Big Due National Preserve, USA
Beech Creek Unit—2(1):651
Loblolly Unit—2(2):1481
Bigelowia—1(1):145
Biodiversity—2(1):743
Bistorta elliptica—2(1):435
Blepharoneuron—1(2): 933,939
Blue Ridge Physiographic Province, USA—
Boehler Seeps and Sandhills Preserve, vo aa USA—
10:777
Boerhavia
Lael ch sl
triquetra—1(2):8
To var. EE (2):874
Bogs—2(2):14
Bolivar, eo POM
Bolivia—2(1):25, 61, 249
Boltonia diffusa—2(2):1353
Boraginales—2(1):263; 2(2):837
Index: Vols. 1 & 2 1511
Borinda TT 135 tetanica—1(1):69
Borreria—1(2):10 trisperma—1(1):389
ums MT var. billingsii—1(1):389
Bouteloua—2(2):917 var. trisperma—1(1):389
burkii—2(2):917 Caribbean —1 A 899; 2(1):35, 95, 139, 267, 463; 2(2):799,
tamaulipensis—2(2):917, 966
Boutelouinae (Poaceae: Chloridoideae: Cynodonte- Conophylacene—1 763; 2(2):1225, 1285
ae) —2(2):917 Cascabela—2(1
Brackish marshes—1(1):665 Cascade up USA—1(1):9; 2(2):775
Bradburia pilosa—1(1):741 Cedar glade vegetation—1(2):1131
Brasiliopuntia—1(1):529 Celtis
in 1(2):913, 1023, 1119; 2(1):45, 61, 75, 291, australis—2(1):627
5, 455, 551,575; 2(2):811, 843, 983 sinensis—2(1):627
British Columbia, Canada—2(1):323 re e 2(1):41, 207; 2(2):995, 1009,
Bromeae (Poaceae: Povideae)—2(1):323
Bromus—2(1):323 Cenosema— tao
sect. Bromeae—2(1):323 Cerbera—2(1):48
sect. Bromopsis—2(1):323 Cereus—1 o
sect. Bromus—2(1):323 Cerrado—2(2):98
sect. Genea—2(1):323 Chaetaria gibbosa—2(1):309
kalmii var. occidentalis—2(1):323 Chamaecrista adiantifolia var. o ert 521
riparius—2(1):323 Chamaesyce ophthalmica 1):6
squarrosus var. i E 323 Chaubardia hetero 10 B
Brunelliaceae—2(2):1183 Chenopodiaceae—2(1):42
Bryanthus—2(2 Sr d eus Artrosis
Bryson Field Press—2(1):5 t, Mexico—1(1):14
Burma—1(1):91, 449; 2(2): us 1183 ud ):847
China—1(1):291; 2
Chiricahua MEUS Arizona, USA—1(1):47
Chloridoideae (Poaceae)—1(2):933; 2(2):861, 875, 917
Chloris
dubia—2(2):861
truncata—1(1):719
Chondrosium
drummondii—2(2):917
foeneum—2(2):917
subscorpioides—2(2):9
Chromosome number—1 (1): y 2(1):487, 489
C. Bickham-Di R iver Ed dR h Park,
mius me 1389
29
Cactaceae—1(1):5
Caesalpinioideae fap ME 121; 2(1):463
larici ina—2 1):434
o scopulorum Herbaceous Associa-
tion—1(1):5
California, USA—1 Dos 59, 69, 719; 2(1):627, 743; 2(2):775,
783
Calyptranthes websteri—2(1):297
Camellia
japonica—2(1 ol ¡cas "
sasanqua—2(1):63 elata—1 (1):
nitidula—1
Eu m us 2(1):397
eae 2):98
Cana —1(2): 1021- US 433
Cannabaceae—2(1): 627
Capanea—2(1):19
Capitan Mountains, New Mexico, USA—2(1):447
Capparaceae—2(1):61, 75; 2(2):807
ee as 61
Capparis—2(1):61, 7
leprieurii— 2 807
Carchi Province, aoe 19
Carex—1(1):38
doc one
billingsii—1(1):401
breviculmis—2(2):1381
):22
Chunky River, 27m USA—1(2):1179
Chusquea—1(2):84
ICE a
annagardneriae—1(2):847
sect. Verticillatae—1(2):847
Cirsium rydbergii Herbaceous Association—1(1):585
Cistaceae—1(1):367, 369
Clitoria—1(2):1101
subg. Bractearia—1(2):1101
pum—1(2):1101
Clitefinae: (Fabaceae: ee (2):1101
Clusiaceae—1(1):1
Coahuila, TEE E 2(2):917, 1235
Coastal Prairie, USA—1(1):679; 1(2):1141
Coleo ee 529
Z
su g
hassei—1(1):69 | ,
klamathensis—1(1):71 Co ies ron—2(1):7
meadii—1(1):69 a uA
scabridum—2(1):75
1512
n coe
NOU cese 2
Colombia—2(1):25, 57 po :861, 875, 1207, 1337
Colorado Plateau, vr i ):58
Colorado, USA—2(1):419; ee 1225, 1407
Colurieae (Rosaceae) —2(1):4
Comanche pate oL USA—2(2):1407
Comarum—1(1):3
Combretaceae—1 a 3:48
Conocarpus erectus—1 mm
Conservation Units—1(1 ~
Convolvulaceae—2(2):79
qa “sisbergense—2 437
Cordia—2
aad 837
Cornaceae—1(1):
91
Costa Rica—2(1):41; 2(2):995
Cotoneaster—2(1):53
ser. Dielsiani—2(1):53
dielsianus—2(1):53
floridus—2(1):55
ser. Frenchetioides—2(1):53
qungbixiensis—2(1):53
ser. ped 53
N
—
cmd
=<
Un
E areoles—1 (2): a 101
Crassulaceae—1(2):889 5
Crataegus—1(2):1005, im ~ 2(1):473
aboriginum—1(2):10
Ll cine ion
a (2):1011
ser. Rode ):1
brazoria var. viburnifolia—1(2):1007
chrysocarpa
var. aboriginum—1(2):1012
var. blanchardii— 1 s 009
var. ee ):10
vigi initamina—1(2) 1008
coccinea var. pringlei—1 (2):1005
bn end she ead Pn
er. Cupressocollinae—1 (2):1056
MNA ade (2):1011
dissona—1(2):1005
divida—1(2):101 1
ferrissii—1(2):1011
flava—1(1):171
ser. Flavae—1(1):171; 2(2):1101
):1008
georgiana—1(2):1005
integriloba—1(2):1011
ser. Intricatae—1(1):171
jackii—1(2):1008
ser. Lacrimatae—1(1):171; 2(2):1101
lancei—2(2):1101, 1143
macracantha
Journal of the Botanical R h Institute of Texas 2(2)
var. divida—1(2):1012
var. A 1012
ertomentosa —1(2):1012
S db A
mollis var. incisifolia—1(2):1012; 2(1):473
er. Montaninsulae—1(2):1065
neofluvialis—1(2):1005
pertomentosa—1(2):101 1
pisifera—1(2):101 1; 2(1):473
praecox—1(2):1005
pruinosa
var. dissona—1(2):1007
var. rugosa—1(2):1012
var. virella—1(2):1013
purpurella—1(2):1066
reverchonii var. mohrii—1(2):1006
rivuloadamensis—1(2):1065
rivulopugnensis—1(2):1070
rubribracteolata—1(2):1073
R 011
schue
var. ee (2):1013
var. gigantea—1(2):1013; 2(1):473
ila-phippsi katcl i
sheridana—1(2):1031
succulenta
var. gemmosa—1(2):1013
var. neofluvialis—1(2):1007
var. pisifera—1(2):1013
ursopedensis—1(2):1031, 1081
velutina—1 (2):1005
viburnifolia—1(2):1005
viridis var. zn c
Cretacifilix—2(2):1
fungifo d 1176
Croton Toa ~ 1
Cryptogams— Pen
Cuatresia—2(1):4
a SEA
Cuba—2(1):267; 2(2):799, 1165
Cunoniaceae—2(2):118
Cyclopogon—1(2):1001
ovalifolium—1 (2):1001
ecundum—1(2):1001
Cynodonteae (Poaceae: Chloridoideae)—1(2):933; 2(2):917
Cyperaceae—1(1):69, 389, 457, 779; 1(2): 875, 1119; 2(1):379;
2(2):1381
Cyperus—2(1):379
drummondii—1(1):779
grayi—2(2):1353
karthikeyanii—2(1):379
Cypress Hills, Canada—1(2):1031
Dahlia scapigera—2(1):60
Daniel Boone National m USA—2(2):1309
Dasiphora—1(1):31
Dátil Island, "om (2):1203
Dead Horse Mountains, Texas, USA—2(1):685
Dentella—1(1):373
Index: Vols. 1 & 2
Diachea arooricola—2(2):1309
Dicerandra
eee ai): 11
subsp. Podesta A 1163
modes 2(2):1
Dichanthelium mer 353
Diodia
MN cuam. 305
schumannii—2(1):30
Diospyros
Hio deutet
sp. chiquimulensis—20) 1025
a dwyeri—2(2):1027
subsp. guanacastensis 20% 1029
031
subsp
subsp. ed rs
subsp. pedromorenoi—2(2):1036
Ms rivensis—2(2):1039
sp. veraecrucis—2(2):1044
d ):1009
subsp. balsensis—2(2):1061
e chutlensis—2(2):1065
subsp. rekoi—2(2):1071
subsp. tehuantepecensis—2(2):1074
amanap—2(2):1009
yucatanensis—
subsp. spectabilis—2(2):1086
District of Columbia, o 627
DNA content—2(2):12
DNA fingerprints—1(1):
Dominican publ 35, 463; 2(2):1 167
Doryopteris—2(2):84
ata T
sect. Doryopteris—2(2):843
jequitinhonhensis—2(2):843
trilobata—2(2):843
Doubled rope climbing method (DRCM)—2(2):1309
Dryas punctata subsp. hookeriana—2(1):441
Drying press—2(1):517
Drymocallis—1(1):31
campanulata—1(1):43
ifoli
cuneifolia
var. cuneifolia—1(1):31
var. ewanii—1(1):44
deseretica—1(1):41
glandulosa
var. reflexa—1(1):43
var. a y43
wrangelliana—1 d :43
"e var. y austinise — 1(1):3
pseudorupe
var. cru ane ie
var. saxicola—1(1):37
Dudleya abramsii—1(2):1015
subsp. costifolia—1(2):1015
subsp. setchellii—1(2):1015
Dysphania
andicola—2(1):428
eva ou
microcarpa— —2(1):42
t
truncata—2(1):427
venturii—2(1):428
Early a 91, 449; 2(2):1175, 1183
Ebenaceae—
Echinodorus Sn ee 5.5
Ecosystem transformers—
2(1):74
Ecuador—1(2):847, 853, 1001; 2 19, 75, 297; 2(2):1295,
1337
Ehrharta erecta—1(1):719
Eichhornia crassipes—2(1):637
Eleocharis
acutangula—1(2):87
subsp. acutangula MEUS 875
subsp. breviseta—1 (2):885
subsp. neotropica— 1(2):886
engelmannii—2(2): 1353
fistulosa var. E (2):875
kleinii—
liesneri—1 us ; »
Elytrigia repens—1(1):7
Endangered Species He a 1973—1(1):457
Endosperm—1(2):1101
iiir di (1):92
burmensis—1(1):92
Eperua Bun 521
Epilobium brachycarpum—1(1):719
Epiphyllum—1(1):529
Episcia—2(1):275
duidae—2(1):275
fimbriata—2(1):275
)
sphalera—2(1):27
subsect. Tr erante} 275
Episcieae ee ee
Equisetum arvense su alpestre—2(1):43
Eragrostideae (Poaceae: eben a ee 875
1513
1514
d oce 875
nsis—2(2):875
ans as ~
patula—2(2):87
ques: o
):875
Ericaceae—1(1):9, 97, 333; 1(2):903; 2(1):207, 243, 249;
Ericoideae (Fricaceae)—2(2):1193
Esmeraldas Province, ido cat 19
Estuarine wetlands—2(1):74
Ethnobotany—2(2):1295
ETS—2(1):7
Eugenia—1(2):899
A 1):299
99
1):
PS trace) 20 811
Eupatorium serotinum—1(2):1141
Euphorbia com
Eupolypod fern—2(2):1175
Euro-Mediterranean—1(1):291
Euthamia—1(1):145
graminifolia—2(1):7
Fabaceae—1(1):121; 1(2):1101, 1253; 2(1):387, 463;
2(2):1491
Fabales—2(1):385
Fagaceae—2(2):1235
Festuca—1(2):827
xgonzalez-ledesmae—1(2):832
subg. Helleria—1(2):827, 831
livida—1(2):827
longiligula—1(2):827
xmiscella—1(2):835
roemeri—1(1):59
var. klamathensis—1 (1):59
Field press—2(1):517
Fimbristylis—2(1):379
naikii—2(1):381
bein noides—2(2):1353
Sens E Florida, USA—2(1):633
Flora Caroliniana—1(1):407, 425; 1(2):1091; 2(1):475;
2(2):1279
Flora de Goids e Tocantins—2(1):455; 2(2):983
Flora of Hispaniola—2(2):1 165
Flora of North America north of Mexico—1(1):31; 1(2):811,
889, 1005, E 1, 1015, 1017, 1019; 2(1):201, 387, 395;
2(2):783, 1
Floret In 3:35
ip: pcr en 1157; 2(1):1, 633; 2(2):1163,
Flow os (2):1
Folia pl ve je 819,895; 2(1):263, 267, 269,
75, 281; 2(2):817, 8
Nd (1):431
Fort Caroline National oe ee 1157
Fossil—1(1):91; 2(1):463; 2(2):1
Fossil flower—2(2):1167
Fossilized resin—1(1):449
Four Canyon Preserve, Oklahoma, USA—1(1):655
Fragaria—1(1):34
French Guiana—1(1):1; 1(2):895; 2(2):807
):1353
parviflora—2(2):1347
Furcraea bedinghausii—2(1):605
Galactomannan—1(2):1101
Galium
anglicum—1(2):1269
oo 1269
parisien
var. ES (2):1269
var. parisiense—1(2):1269
Gamochaeta
antillana—1(2):1125
coarctata—1(2):1125
pensylvanica—1 n : 125
urp 1(2):1
Gardenieae koe E E
Gaultheria
miqueliana—1(1):97
pyrolifolia—1(1):97
LEN (1):97
Gaylussac
bigeloviana—1 H 336
dumosa 1):33
mosieri—1 Ms E
Generic realignment—1 (1):31
Gentianopsis barbata subsp. raupii—2(1):442
Geocarpon minimum—2(1):643
Geographical Information System—2(1):677
Gesneriaceae—2(1):19, 269, 275; 2(2):825
Geum—2(1):415
donianum—2(1):417
Ghana—1(1):549
Glinus ‘aclatus—10: 719
Global change—1(1):486
Gloxinieae LR LA NDA p
Gnaphalieae (Asteraceae)—2(1):48
Goias, Brazil—1(2):1023; 2(1):305; q 983
Gomesa—1(2):913
f Texas 2(2)
Gonolobinae (Apocynaceae: Asclepiadoideae)—2(1):95,
]
Gonolobus—2(1):95
bayatensis—2(1):139
broadwayae—2(1):139
ciliatus—2(1):139
Index: Vols. 1 & 2
dictyopetalus—2(1):139
variifolius—2(1):139
Gratiola—1(1):149
ebracteata—1(1):149
flava—1(1):149, 763
floridana—1(1):149
graniticola—1(1):166
heterosepala—1(1):149
neglecta—1(1):149
sect. Nibora—1(1):149
quartermaniae—1(1):163
virginiana—1(1):149
Great Smoky ibis ia Park, USA—2(2):1309
Greater Antilles—2(2):1
Grossulariaceae—1(2):1015
Guadalupe Mountains National Park, USA—2(1):677
Gua
chaparensis—2(1):31
incana—2(1):26
He eae 7
Guaduinae (P
uerrero, Mexico—1(1):491
Guiana Shield —2(1):263, 521; 2(2):837
Guianas—1(2):819, 1023; 2(1):281
Gulf Coast, North America—1(1):385
Gulf Coastal Plain (Eastern), USA—1(1):57
Gulf Coastal Plain (Western), mue 577; 1(2):1149;
2):1475
Gulf of California, Mexico—1(2):1203
Gundlachia—1(1):145
Gutierrezia—1(1):145
elegans—2(2):771
Guyana—2(1):281, 521
Gymnosperma—1(1):145
Sie db m (2):891
Habitat noceat 677
Haiti—2(1):26
Hancock a VUA Kentucky, USA—1(1):609
Hanging garden— 1(1):
Hawaiian Island, ru 789
Hawthorns—1 (2):10
Hedyotideae o F (1):373
Hedyotis—
ekmanii—2(2):1165
Hedysarum boreale subsp. dasycarpum—2(1):441
Heliotropiaceae—2(1):263
Heliotropium—2(1):264
Hesperostipa—2(1):165
Heterotheca
arizonica—1(1):27
1515
foliosa—1(1):21
fulcrata—1(1):21
var. amplifolia—1 m
var. arizonica—1(1):2
var. fulcrata—1(1): ~
viscida—1(1):21
zionensis—1(1):21
Heuchera
sect. Holochloa subsect. Cylindricae—2(1):447
woodsiaphila—2(1):448
Himalayacalamus
asper—1(1):135
planatus—1(1):137
Hinton, James (memorium)—1(2):1 277
Hippopotamus—1(1):54
Hispaniola—2(2):1165
Hosackia—2(1):387
crassifolia var. otayensis—2(1):388
oblongifolia var. cuprea—2(1):388
stipularis var. ottleyi—2(1):388
Houstonia—1(1):109
subg. Chamisme—1(1):109
subg. Ericotis—1(1):109
subg. Porotis—1(1):117
prostrata—1(1):109
purpurea—1(1):109
Huachuca Mountains, Arizona, USA—1(1):47
Hualapai Vgl Arizona, USA—1(1):47
Humboldtia—1(1):121
a var. raktapushpa—1(1):121
Hurricane Hugo—1(1):665
Hydrilla verticillata—2(1):637
Hydrocotyle dea (1):741
Hydrophytes—1(1):7
Hylocereus—1(1): A
Hypericum
adpressum—1(1):713
lloydii—2(2):1353
Ibatia ere i 139
Iceland—2(1):43
ii ^
Illiciu
ieee subsp. guajaibonense—2(2):799
guajaibonense—2(2):800
India—1(1):121, 129; 1(2):903; st 373, 379
menor Highlands a ):76
Invasive plants—2(1):743
Ipomoea
concolora—2(2): x
conzattii—2(2):79
subg. Eriospermu —2(2):793
ser. a e ):793
tehu o 793
de Mexico—1(2):12
ITS—2(1):
TEn :455
1516
araguaiensis—2(1):456
spes 456
irwinii—
Ixoroideae TOER :455
Jacaima
costata var. NA 139
s 2(1):1
Jalisco, Modo 111459 2(2):1009, 1341
Jarava 1):165
Jarava subg. Pappostipa—2(1):16
Jepson Manual, The—1(2):1017; 2(1): i 397, 783
Juncaceae—2(2):775
Juncus
articulatus—2(2):1353
sect. Sed E 775
digitatus—2(2):77
dudleyi—2(2): de
interior—2(2):1353
militaris—2(2):1353
nodatus—2(2):1353
e 1353
triformis—2(2):77
Kaieteur National Park, Guyana—2(1):521
Kalmia angustifolia—2(2):1353
Kalmiopsis
fragrans—1(1):10
leachiana—1(1):9
Kansas, USA—1(1):753
Karnataka, India—2(1):379
Kentucky Lake, USA—1(1):609
Kentucky, USA—1(1):609; 2(2):1309
Kiowa National Grasslands, USA—2(2):1407
Klamath (Siskiyou) Mountains, m ):9
Klamath Region, USA—1(1):59
Koeleria pyramidata subsp. in 434
Kohautia—1(1):373
Kohleria—2(1):19
hypertrichosa—2(1):20
Kruschke, Emil P—1(2):1011; 2(1):473
Kwakwani, Guyana—2(1):521
Lachnociona terriae—2(2):1183
poe d ms E E
erriae—2(2):1
Lake damas da USA—2(1):633
Lake Wales Ridge, Florida, USA—2(2):1163
Lamiaceae—1(1):717; 2(1):193; 2(2):761, 1163
Mp ics 2(1):464
M :464
inc 36 ):8
TONS cee NEP 2
Laubertia—1(1):431
contorta—1(1):431
Leaf blade anatomy—2(1):495
Lechea
maritima—1(1):36
var. Won 367
pulchella amosissima —1(1):370
san- ES
i. ee eee
nen ~
Lepismium—1
Leptochloa—2(2): ie
Leucospora multifida—2(2):1353
Liabeae (Asteraceae)—2(2):1207
Liabum meridense—2(2):1207
Lilaeopsis carolinensis—2(2):1497
Limnanthaceae—1(2):1017
Limnanthes—1(2):1017
alba
subsp. gracilis—1(2):1017
subsp. parishii— 1(2):1017
douglasii subsp. striata—1(2):1017
Limnorchis
aquilonis—2(1):434
huronensis—2(1):434
Limonium
indet.—1(1):719
ramosissimum—1(1):7
Lipocarpha duondi2 1353
Lobelia—1(2):78
sect. Liar m A
gaudichaudii—1(2):80
ini de ):80 B
p. gloria- cad 800, 805
Jub (2):806
wahiawa— 1(2):802
Lobelioideae (Campanulaceae)—1(2):78
Lone Mesa State Park, Colorado, vues 771
Loropetalum adea 637
Loteae (Fabaceae)—2(1):38
Louisiana, o 763; 1(2):1125, 1141,
2(1):643; 2(2):1389
mL (1):118
etrandra—1(1):109
puse i o 633
Lycurus— , 941
pes minds Rosaceae} —2(1) 385
Lyonothamnus—2(1):38
Macleania
coccinea—2(1):243
£T, >
Texas 2(2)
1251;
43
Madrean Archipelago (sky islands), North America—1(1):47
Magdalena River, Colombia—2(1):60
Magrath, Lawrence K. cU ie
Maharashtra, India—2(1): 379
Malvaceae 83
Mancos Shale—2(2):771
Mandevilla—1(2):853, 859
Pr im: (2):853
frigida—
a a: "n
horrida—1(2):863
inexperata—-1(2):865
lojana—1(2):855
Index: Vols. 1 & 2 1517
megabracteata— 1(2):866 Minuartia
similaris—1(2):867 austromontana—2(2):1225
Mangrove—1(1): 7 drummondii—1(1):763
Mapping—2(2):13 filiorum—2(2):1225
Marsilea cece 20 637 macrantha—2(2):1225
Matelea—2(1):13 muriculata—1(1):763
ostata var. goodfriendii—2(1):150 muscorum—1(1):763
dictyopetala—2(1): 130 rossii-stricta—2(2):1225
a 51 rubella—2 E A
pubescens—2(1):13 stricta—2(2):12
Morea eec tus 1003 Mirablis UNUM ME
Medranoa—1(1):14 Mississippi Natural Heritage Program, USA—1(2):1179
a Mississippi, USA—1(1):769; 1(2):1179; 2(2):1381
palmeri—1(1):148 Mississippi, Yazoo- Md Delta Region, USA—1(1):769
Au s 148 Missouri, USA—2(2):13
purpusii—1 (1):14 Mizoram, pua.
Melastomataceae—2(1 is Modeling—2(2):1363
Meliosma antioquiensis—2(1):57 Mogolon Rim, Arizona, USA—1(1):47
Mesechiteae (Apocynaceae: Apocynoideae)—1(2):853, — Monarda lindheimeri—1(1):717
859 Monilicarpa—2(1):67
Mesoamerica—2(2):1009 brasiliana—2(1):71
Mexico—1(1):47, 79, 109, 145, 431, 486, 491, 499, 511, 779; tenuisiliqua—2(1):70
1(2):827, 871, 891, 933, 1203; 2(1):207, 495, 547, 593, Monopyle—2(1):19
605; 2(2):793, 917, 1009, 1235, 1341 Moraceae—2(2):1167
Mexico City, Mexico—2(1):605 Morisita community 22 index—1(1):585
Mexico, Valley of—2(1):605 Morphometrics—
Michoacan, Mexico—2(1):547 Morro Alto Mouráo, Bum 575
Miconieae ir o M 35 Mosiera—1(2):899
Micranthes—1(2):10 androsiana—1(2):899
r Sot A cuspidata—1(2):899
calycina—1(2):102 gracilipes—1(2):899
ferruginea—1(2):1020 xerophytica—1(2):900
foliolosa—1(2):1020 Mudflats—2(1):743
gormanii—1(2):1020 Muhlenbergia—1(2):933
hieraciifolia subsp. dete D :439 capillaris—1(1):85
idahoensis—1(2):1 expansa—1(1):85
lyallii subsp a cana :440 glomerata var. ramosa—1 (2):933
merkii— 2(1):440 sericea—1(1):85
mexicana—1(2):1020 spiciformis—1(2):933
nelsoniana Muhlenbergiinae (Poaceae: Chloridoideae: Cynodonte-
subsp. aestivalis—2(1):440 ae)—1(1):85; 1(2):933
var. aestivalis—1(2):1020 Munrochloa—2(1):374
var. carlottae—1(2):1020 ritchiei—2(1):376
var. cascadensis—1(2):1020 Murray State University—1(1):609
subsp. insularis—2(1):440 Myanmar—1(1):91, 449; 2(2):1 ~ 1183
var. insularis—1(2):1020 Myrcia subcordifolia—2(1):30
var. pacifica—1(2):1020 Myriophyllum spicatum—2(1): a
var. porsildiana—2(1):395 Myriopus—2(1):263
nudicaulis—1(2):1021 candidulus—2(1):264
petiolaris—1(2):1021 uc da
porsildiana—2(1):440 paniculatus—2(1):26
razshivinii—1(2):1021 var. spigelii oe a 264
redofskyi—2(1):441 volubilis—2(1): 264
tempestiva—1(2):1021 Myrtaceae—1 (2):899; 2(1):297
tischii— 1(2):1021 Myxomycetes—2(2):1 309
tolmiei—1(2):1022
unalaschcensis—1(2):1022
Mimosoideae (Fabaceae)—2(2):1491
Minas Gereis, Brazil—2(2):843
Nagaland, India—1(2):903
Nash Prairie, Texas, USA—1(1):679
Nassella-Amelichloa clade—2(1):165
1518
National List of Plants that Occur in Wetlands, USA—
1(1):719
National Park Service, USA—1(2):1157
Natural Municipal Park of Prainha, Brazil—1(1):529
Nature Conservancy, The—1(1):655, 777
Nautilocalyx
vestitus—2(2):833
Neanotis—1(1):373
californicus—2(1):397
glanduliferus
var. australis—2(1):397
var. o ):397
orientalis—2(1):39
rubescens var. Min ):39
secundiflorus var. robbins—20) 399
S s
lla: 398
Neocaypuocaha a 807
Neonesomia—
iG. 853, 859, 1023; 2(1):285, 305; 2(2):1207
Nepal—1(1):135
Neptunia plena—2(2):1491
Nevada, USA—1(1):351
New Manual of ~ of the Central Rocky Moun-
tains—2(1):20
New Mexico, cam 351; 2(1):419, 447; 2(2):1407
Nicoraepoa—2(2):847
Norma Oficial ea (1):491; 2(1):605
North America—1(1):31, 59, 109, 149, 291, 333, 357, 385;
(2):871, 1005, 1011, 1019, 1031, 1149; 2(1):201, 323
387, 397, 401, 415, 425, 473, 487, 621, 627; 2(2):775,
hi ie 917, 1009, 1235, 1285, 1341, 1347, 1353, 1363,
407
à
North e a 345; 2(1):649; 2(2):1309
Norway—2(1):43
Nuclear magnetic
Nuevo Leon, Mo (2): 891, 933; 20): 593; 2(2): 917
Nuphar—2 `
adven po ):59
yciaginaceae 10s a ;2(1):419, 547
Nymphaea—
dap e
elegans—2(1):593
mexicana—2(1):593
Nymphaeaceae—2(1):593
Oak-hickory forest—1(1):609
Oaxaca, Mexico—2(2):793, 1009
Oenothera arizonica—1(1):483
Oklahoma Natural Heritage inventory—1(1):65
Oklahoma, USA—1(1):655, 777; 1(2):1125; 2(2): a
fTexas 2(2)
Oldenlandia—1(1):373
strigulosa—1(1):373
Oldenlandiopsis—1(1):373
Oligactis—2(2):1207
subg. Andromachiopsis—2(2):1207
al 207
boyacensis—2(2):1207
garcia-barrigae—2(2):1207
subg. Oligactis—2(2):1207
sessiliflora—2(2):1207
valeri—2(2):1207
volubilis—2(2):12
Olyreae (Poaceae: dcr
Onagraceae—1(1):
Oncidiinae (Orchidaceae)—1 (2):913
Oncosiphon piluliferum—1(1):719
Open woods—2(1):661
Opuntia—1(1):529
Orchidaceae—1(2): 913, 1001, 1003, 1265; 2(1):285
Orchidotypus—
Oregon, USA—1(1):9, 59, 69
Orthaea laurifolia—2(1):244
Oryzeae (Poaceae)—2(1):633, 673
Ottelia ipic E (1):741
Ottleya—2(1):387
Ouachita Nounans or USA—1(1):713
Oxalidales—2(2):1
Oxbow NE 2
Oxytenanthera—2(1):373
Pachyphyllum—2(1):285
Capitulum—2(1):288
rchidotypum—2(1):287
Packera hyperborealis subsp. wrangelica—2(1):443
Pakaraima cqui uuo
Pakistan—1(1):557
Pan de as ae Cuba—20 799
Pa —2(1):41, 521; 2(2):1009
Panarctic Flora Checklist —2(1): 433
Panarctic Vascular Plant Flora —2(1):433
Panay Island, Philippines—1(2):815
Paniceae (Cyperaceae)—1(1):69
Panicum
PME dL
dichotomifloru
d dehoromfom—1(9 719
sae ot
a
os (19:71 :
er labradoricum "ar :438
Medis a 13
Pappostipa—2(1): P
me
ar. digona—2(1):184
var. precordillerana—2(1):184
atacamensis—2(1):1 Em
barrancaensis—2(2):12
braun-blanquetii—2(1 x 83
chrysophylla—2(1):185
var. cordillerarum—2(1):185
Index: Vols. 1 & 2
var. crispula—2(1):185
f. ipis n
f. modica 2185
sect. Chrysovagnatae 211) 184
chubutensis—2(1):18
var. AS an 85
frigida—2(1):185
var. parvispicula—2(1):185
e ie ):183
humilis—2(1):18
Var. dececne-20) 185
n eM 3:18
o
f. NE a m 84
var. anomala—2(1):184
Pl R
nana—2(1):184
Marr. ):18
sect. Pappostpa—2 182
parodiana—2(1
n
ruiz-lealii—2(1):183
E 183
sorianoi—2(1):18
speciosa—2(1):18
f. uc nds m 82
var. atuelensis—2(1):182
var. breviglumis—2(1):182
var. ciliata —2(1):1 "
f. horrida—2(1):18
var. au cU 182
var. media—2(1):182
var. parva—2(1):182
vaginata—2(1):182
var. argyroidea—2(1):183
f. rigida—2(1):183
vatroensis—2(1):184
Paraguay—2(1):61
p tor
que d Central America—2(1):41
Paspalum—2(1)74
urvillei—1 (1): 25 id 1141
vaginatum—2(1):74
Passiflora
angusta—2(2):818
arta—1(2):819
supersect. Auriculata—2(2):817
coccinea—1(2):819; 2(1):281
supersect. Coccinea—1 (2):824; 2(1):281
compar—1 (2):821
cryptopetala—2(2):817
davidii—1(2):896
subg. Decaloba—2(2):817
foetida—1(1):143
. orinocensis—1(1):144
(1):267
subg. Passiflora—1(2): 819, 895
rufa—2(2):82
aa Stipulata—1(2):895
Calopathanthus—1(2):895
sect. o
2):895
sect. Granadillastrum—1(2):895
sect. Kermesinae—1(2):895
sect. NE (2):895
— 28
Su PRU ):27
1519
sect. Tre
dd um HG 143; 1(2):819, A 267; 2(2):817
wnee ooer ~ 2(2):14
EE Uia Bo 1449
Pentodon—1(1):37
Pereskia—1(1 E
Periandra—1(2):1101
Pertle Springs, Missouri, USA—2(2):1309
Peru—1 ad 841,853; 2(1): 75, 249
Petrador
Sa 1
var. graminea—1(1):351
a mila—1(1):351
Phaseoleae (Fabaceae)—1(2):1101
Phellodendron—1(1):357
Philadelphaceae—1 (2):901
Philadelphus—1(2):901
microphyllus
var. argyrocalyx— 1(2):901
var. madrensis—1(2):901
r. stramineus—1(2):901
Philpoines 12 815
Phragm
d: uy (1):385
Sl bsp americanus—1(1):385
subsp. berlandieri—1(1):387
iis NL :401
hylogenetic "os 1193
iege (2):913; 2
Phytogeographical cr um 1475
Phytogeography—1(1):389; 1(2):1149
Pilea—2(2):995
alfaroana—2(2):995
longbracteolata—202): 1002
moragan a
Ec 1:52
Pinar del Rio, ai sae
Piptochaetium—2(1):165
Plant biodiversity—1(1):549
Plant communities—1(1):557
1520
Plant press—2(1):517
Plant succession—1(2):1 131
Plantaginaceae—1(1):149, 763
lumerieae ( Ifoi 2(1):
Poa—2(2):847
hachadoensis var. pilosa—2(2):847
:84
Poaceae—1(1):1, 59, 85, 135, 385; 1(2):827, 841, 847, 933;
2(1):25, 165, 309, 323, 373, 495, 633, 673, 743; 2(2):847,
,917, 1233
Poeae (Poaceae —1 (2):827
Poecilon
P 129
pauciflorum—1(1):129
Poicilla
re 39
mnifolia—2(1):13
pocilopsis aspire 201): 139
idi
Pooideae Mage M 323
Portulacaceae—1(1):777
Potaro River, Guyana—2(1):521
iip 31; bd 811; 2(1):201
albiflor
anserina be yukonensis—2() 44]
arenosa subsp. chamis —1(2):813
sect. Aureae—1(1):47
concinna var. divisa—2(1):201
sect. Concinnae—2(1):201
sect. c 201
holmgrenii—1(2):8
hyparctica x sio (2):813
jepsonii—2
sect. scole: 201
litoralis—2(1):20
o ( l ):4
sect. Mufa 20 201
nivea—1(2):8
var. dis iM
sect. Niveae—1(2):811; 20 201
pensylvanica—2(1): 201
vium—2(1):201
eo —1(1):47
pseudosericea—2(1):201
ramulosa—1(1):47
rhyolitica—1(1):50
r. chiricahuensis—1(1):52
sect. Rubricaules—2(1):201
sect. Subjugae—2(1):201
RUN (1):47
bviscosae—1(1):47
sect. pim 201
sud var. ene ul
Prtranrill
Potentilleae 1(1
Pothowar Plateau, Pakistan—1(1):557
Prairies—2(2):1475
Praxelis clematidea—2(1):621
Prestonia—1(1):431
nus
m m
americana—2(1
aoa. i
gracilis—2(1):401
hortulana—2(1):401
maritima—2(1) 2401
(1):401
ect. Prunocerasus—2(1):401
On y401
subcordata—2(1):401
bellata—2(1):401
calciolum—2(2):1
TN S UEa na e 719
Pteridaceae—2(2):843
Ptilagrostis—2(1):165
Puccinellia phryganodes
subsp. neoarctica—2(1):435
SU bsp. apu
NR
ect. Ce 193
ee 193
Californicae—2(1):195
sect. ct apii at 193
flexu 2(2):13
sect. Mc ce M 193
sect. Nudae—2(1):1
y
sect. Pycnanthemum—2(1):193
virginianum—2(1):193
Quarterman, Sem 1131
Quercus—2(2):12
ie cde Es
Quillaja—2(1):385
Quillajaceae—2(1):385
Raddiella
Repas 1
minim ):1
V "um
ae
Ranunculus
ficaria—2(1):741
subsp. bulbifera—2(1):741
laxicaulis—1(1):741
412
he
f Texas 2(2)
Index: Vols. 1 & 2
sub borealis
subsp. pumilus—2(1):437
ubsp. villosus—2(1):438
Raphiolepis indica—2(1):637
Rauvolfoidae (Apocynaceae)—2(1):489
Regional is relationships—2(1):685
Rhipsalis—1(1):529
Rhododendron tomentosum subsp. decumbens—
ad cephalantha var. att pla ipid 1353
Ribes oxyacanthoides var. cognatum—
Rincon eae! ud b UE 47
ne de Jan 2(1):575
S " d 119
Riparian—1(1):719
Riparian forests—2(1):521
Rocky Mountains, USA—2(2):1225
Roraima sandstone—2(1):521
Rosaceae—1(1):31, 47, 171; 1(2):811, 1005, 1011, 1031;
2(1):53, 201, 385, 401, 415, 473; 2(2):1101
tribus Lyonothamneae—2(1):385
subseries Robustae—2(2):1124
subseries aM 108
Rosales—2(1):38
Rosid—2(2):118 ex
Rosoideae (Rosaceae)—1(1):31
Rubiaceae—1(1):109, 373; 1(2):1023, 1269; 2(1):305, 455;
Russia—2(1):433
Rutaceae—1(1):357
Sabiaceae—2(1):57
Sagittaria
chapmani—2(2):1353
Saline prairie—1(2):1251
Salt marsh—2(1):743
Salvia toaensis—2(2):1165
Salvia toaensis—2(2):1165
Salvinia molesta—1(1):719
San Esteban island, Mexico—1(2):1203
San Jacinto River, Texas, USA—2(1):661
San Marcos River, Texas, USA—2(1):673
San Pedro Nolasco Island, Mexico—1(2):1203
Santa Catalina Mountains, Mila JUSA—1(1):47
Santa Catarina, Brazil—1(2):1
Santa Rita Mountains, Pen iain
Saskatchewan, Canada—1(2):1
19
Since i acd 102): 1251
Sa ne ME pA ~ 1251; 2(1):395, 447
Saxifragales—2(2):1
Schoenoplectus E CAS
Sclerolepis uniflora—2(2):1353
S ides aut lis subsp. pratensis—2(1):443
Seashore—2(1):74
Seasonally Sy Topi Forest—2(1):551
Sedge—1(1):38
Sedge ecology—1(1):389
Sedum—1(2):889
lanceolatum var. subalpinum—1(2):890
laxum
var. eastwoodiae—1 (2):889
var. flavidum—1(2):889
var. heckneri—1(2):889
var. latifolium—1(2):890
obtusatum
var. boreale—1 (2):889
var. paradisum—1 (2):889
var. retusum—1(2):889
oreganum var. tenue—1(2):890
radiatum
var. ciliosum—1(2):889
var. depauperatum—1 (2):88
stenopetalum var. monanthum—1 A 890
wrightii
var. iris bium ie 889
ar. or ):88
Seed era ee
Seed dispersal—1 ee
Seed germination—2(1):673
Senecio
ampullaceus—2(1):643
subpeltatus—2(2):1215
Senecioneae (Asteraceae)—2(2):1215
Seriphidium quettense—1(1):569
Serpentine endemic—1(1):69
Serra da Tiririca Mountains, Brazil—2(1):575
Serra da Tiririca State Park, Brazil—2(1):575
Serra do Orobó Mountains, Brazil—2(1):551
Setaria
adharerens—1(1):719
megaphylla—1(1):719
Sibbaldia—1(1):31
n m 31
Sidalcea—2(2):78
ickmanii
subsp. fiapensis ete 783
subsp iensis—2(2):786
Sierra de Baoruco, Dominican Republic—2(1):35
Sierra de Coalcomán (1):547
Sierra Kunkaak, Mexico—1(2):12
Sierra Madre del Sur, Mexico—1 (1):491
Sierra Martin Garcia, Dominican Republic—2(1):35
Silene
Michoacán, México—
flos-cuculi subsp. flos-cuculi—2(1):649
involucrata subsp. furcata—2(1):436
soczavana var. macrosperma—2(1):436
3
Sinaloa, Mexico—1 (1):109; 2(2):1009
Sino-Himalaya—1(1):135
Siphonandra—2(1):249
boliviana—2(1):249
elliptica—2(1):249
1521
1522
magnifica—2(1):249
nervosa—2(1):256
nta- oo 256
lupa ):55
Solanaceae—2(1): 2
Solidago
altissima—1(2):1141
multiradiata a arctica—2(1):443
ptarmicoides—2(1):7
Solidaster (Solidago x rT 7
Sonora, Mexico—1(1):109
Sonoran a ae Gulf Coast subdivision—1(2):1203
—1(1):145
Sonoran D
Sonoran amen M cum. 1203
S@renson index—2( d
South Africa—2(2):12
South Am MN 143; 1(2):819, 841, 847, ed 895,
13, 1001, 1023, 1119; 2(1):19, 25, 45, 57, 61, 75, 249,
263, 269, 275, 281, 291, 297, 305, 455, e 551, 575;
2(2):807, 811, 817, 825, 837, 843, 847, 861, 875, 983,
1207, 1295, 1337
South Carolina, USA—1(1):345, 665; 1(2):1269; 2(2):1279
Southern m Mountains, USA—2(1):649
Spartina—2(1):74
Species area psa
Species ratios d Lo Ud
Species richness—1(1):577; 2(2):1363, us
Spermacoce—1(2):1023
burchellii—1(2):1025
crispata—1(2):1025
delicatula—1(2):1025
paraensis—1(2):1027
pulchristipula—1(2):1027
pi
o (2):1028
spicata—1 (2):10
Sphagnum bog—1(1):389
Spiranthes brevilabris—1(2):12
Spiranthinae ae si ana 1001
Stachys—2(2):761
iltisii—2(2):761
Stipa speciosa—2(1):165
Stipeae (Poaceae)—2(1):165
Stuckenia filiformis subsp. borealis—2(1):433
Succession—1(2):1141
Symphyotrichum
pretense—2(1):731
sericeum—2(1):731
Synoptical Flora of bui America—2(2):1285
Syrmatium—2(1):38
Talamanca, Costa Rica—2(1):41
Talinum rugospermum—1(1):777
£T, ^f^
lexas 2(2)
Tamaulipas, Mexico—1 (1): 79, 511; 1(2):933; 2(2):917
Taxaceae— 1 (1): p 291
Taxus—1(1):203,
Taree i ):266
brevifolia—1(1):291
var. polychaeta—1(1):217
var. reptaneta—1(1):219
caespitosa
var. angustifolia—1(1):268
var. latifolia—1
A
ar. adpressa—1(1):274
var. minor—1(1):274
ontorta var. mucronata—1(1):260
nac 1 xen
globosa—1(1):29
var. ica (1):224
kingstonii—1(1):240
mairei var. speciosa—1(1):246
var. NM 279
Var. digi 279
var. nana—1(1):28
Tennessee, Inner Central la eu USA—1(2):1131
TUNE Man quels du 2(2):13
r. 2(1): 637
Tertiary —2(1 ):463; 2(2):1167
ygia
Tetraz
elaeagnoides—2(1):35
eee =
ralongicollis—2(1):35
Texas, qiu ):679, e 717, 763, 779; 1(2):1125, 1149,
1253, 1255, 1265, 1269; 2(1):419, 651, 661, 665, 673,
677, 685, 741; 2(2):1291, 1347, 1373, 1475, 1481, 1491,
Thecagonum—1(1):373
strigulosum—1(1):377
Thevetia
ect. Thevetia—2(1):489
Thibaudia laurifolia—2(1):243
Thicket—2(1):661
Thomas Walter Typification Project—1(1):407, 425; 1(2):1091;
2(1):475; 2(2):1279
Threatened plant—1(2):815
Thunbergia alata—2(1):637
Thurovia—1(1):145
Timucuan Ecological and Historic Preserve, Florida, USA—
2):1157
Tocantins, Brazil
1(2):1023; 2(1):305, 455; 2(2):983
Index: Vols. 1 & 2
a nd 983
a 988
oa ):26
Tracheid pits—1(1):44
Transmexican Volcanic m
Tree canopy—2(2):1309
Tree climbing—-2(2):1309
Trochanthera—2(2):1168
lepidota—2(2):1168
Ls Specimen catalogue—1(1):437
mpqua National Forest, USA—1(1):9
e. of British eli Herbarium—1(1):437
Urban runoff—1(1):71
s id
USA—1(1):9, 29, 47, 59, 69, 103, 109, 171, 345, 351, 367, 369,
407, 425, 457,483, 577, 585, 609, 631, 655, 665, 679, 693,
713, 717, 719, 741, 753, 763, 769, 777, 779; 1(2):789,
1091, 1125, 1131, 1141, 1149, 1157, 1179, 1251, 1253,
1255, 1265, 1269; 2(1):1, 419, 433, 447, 475, 627, 633,
637,643, 649,651,661,665, 673, 677, 685, 731, 741, 743;
2(2):761, 771, 775, 1101, 1163, 1225, 1279, 1291, 1347,
1353, 1363, 1373, 1381, 1389, 1407, 1449, 1475, 1481,
1491, 1495, 1497
Utricularia juncea—2(2):1353
Vaccinieae (Ericaceae)—2(1):243, 249
Vaccinium—2(1):207
beamanianum—2(1):219
campanense—2(1):234
jason 220
uliginosum subsp. vulcanorum—2(1) :442
Vahlodea AE subsp 2(
Validation—2(1): we
Varronia—2(2):83
bullata sees — ce 837
cremersii—2(2):83
marioniae—2(2): e
Vegetation analysis—1(1):557
on communities—2(1):685
eae zones—1(2):1131
Vegetational areas—2(2):1373
Venezuela—2(1):61, 75, 269, 275; 2(2):1207
1523
Venezuelan Guayana—2(1):275; 2(2):825
Veracruz, Mexico—2(2):1009
Verbena brasiliensis—1(2):1141
Vicia
Ei E 1253
sativa
Viola CUM
Violaceae—2(1):677
Virginia, USA—1(1):367
A :415
doniana—2(1):415
REN.
subsp. doniana—2(1):415
var. E u 5
parviflora—2(1):4
Wechiau Community — Sanctuary, Ghana
—1(1):549
Wechiau, E cms
West im. 899; ES 95, 139, 267, 463; 2(2):799,
eee a nde 121, 129
Wetlands—1(1):5
Indicator s (1):719
Pine savannas—2(2):1475
Plant list—1 n 9
White sands—2(2):98
Wildland-urban e EN idus 743
Wissadula hernandioides—1(2):12
Witheringia—2(1):41
Wyoming, USA—2(2):1225
Xeric O 1475
Xeromorphy—2(2):1
ea ae e
Xylovirgata—1(1):145
Xyridaceae—2(1):1
Xyris panacea—2(1):1
Yazoo- rd rs deal Mississippi, USA—1(1):769
oun Peninsula Verum 1009
Yucca gloriosa—2(2):1353
flora—
Zizania texana—2(1):673
454 NEW NAMES AND NEW COMBINATIONS:
VOLUME 1 (2007) AND VoLUME 2 (2008)
Achillea alpina subsp. multiflora (Hook) D.F Murray & Elven,
com 2
Acmispon americanus var. helleri (Britton) Brouillet, comb.
nov.—2(1):388
Acmispon argophyllus (A. Gray) Brouillet, comb. nov.—
2(1):388
Acmispon argophyllus var. adsurgens (Dunkle) Brouillet,
comb. nov.—2(1):388
Acmispon argophyllus var. argenteus (Dunkle) Brouillet,
comb. nov.—2(1):388
Acmispon argophyllus var. fremontii (A. Gray) Brouillet,
omb. n 1):389
Acmispon argyreus (Greene) Brouillet, comb. nov.—
2(1):389
Acmispon argyraeus var. multicaulis (Ottley) Brouillet,
comb. nov.—2(1):389
Acmispon argyraeus var. notitius (Isely) Brouillet, comb.
nov.—2(1):389
Acmispon cytisoides (Benth.) Brouillet, comb. nov.—
2(1):389
1524
Acmispon dendroideus (Greene) Brouillet, comb. nov.—
2(1):
:389
Acmispon dendroideus var. traskiae (Noddin) Brouillet,
comb. nov.—2(1):389
Acmispon dendroideus var. veatchii (Greene) Brouillet,
comb. nov.—2(1):389
Acmispon distichum (Greene) Brouillet, comb. nov.—
2(1):389
Acmispon flexuosus (Greene) Brouillet, comb. nov.—
2(1):389
Acmispon glabrus (Vogel) Brouillet, comb. nov.—2(1):389
Acmispon glabrus var. brevialatus (Ottley) Brouillet, comb.
nov.—2(1):390
Acmispon grandiflorus (Benth.) Brouillet, comb. nov.—
2(1):390
Tera | 4 f, 1n All
comb, nov.—2(1):390
Acmispon greenei (Wooton & Standl.) Brouillet, comb.
nov.—2(1):390
Acmispon haydonii (Orcutt) Brouillet, comb. nov.—
2(1):390
Acmispon heermannii (Greene) Brouillet, comb. nov.—
2(1):390
Acmispon heermannii var. orbicularis (A. Gray) Brouillet,
comb. nov.—2(1
Acmispon intricatus (Eastw.) Brouillet, comb. nov.—
2(1):390
Acmispon junceus (Benth.) Brouillet, comb. nov.—
2(1):391
Acmispon junceus var. biolettii (Greene) Brouillet, comb.
nov.—2(1):391
gi td Men var. brevivexillus (Ottley) Brouillet,
comb. n 1):391
s den mearnsii (Britt) Brouillet, comb. nov.—2(1):391
Acmispon mearnsii var. equisolensis (J.L. Anderson) Brouil-
let, comb. nov.—2(1):391
Acmispon micranthus (Nutt. ex Torr. & A. Gray) Brouillet,
comb. nov.—2(1):391
Acmispon nevadensis (S. Watson) Brouillet, comb.
nov.—2(1):391
Acmispon nevadensis var. davidsonii (Greene
comb. nov.—2(1):391
Acmispon niveus (S. Watson) Brouillet, comb. nov.—
2(1):391
Acmispon nudatus (Watson) Brouillet, comb. nov.—
2(1):391
—
Brouillet,
Kunth) Brouillet, comb. nov.—
—
Acmispon oroboides
2(1):391
Acmispon procumbens (Greene) Brouillet, comb. nov.—
2(1):392
Acmispon procumbens var. jepsonii (Ottley
comb. nov.— 2
pd Lies s ex Torr. & A. Gray) Brouillet,
comb —2(1):39
Psi odes im Brouillet, comb. nov.—2(1):392
Acmispon strigosus (Nutt. exTorr. & A. Gray) Brouillet, comb.
nov.—2(1):392
Brouillet,
=
Acmispon utahensis (Ottley) Brouillet, comb. nov.—
2(1):392
Acmispon wrightii (A. Gray) Brouillet, comb. nov.—
2(1):392
Agapetes acuminata var. tipiensis Banik & Sanjappa, var.
v.—1(2):903
Agapetes flava var. nagensis Banik & Sanjappa, var.
nov.—1(2):905
Agapetes megacarpa var. lohitensis Banik & Sanjappa, var.
nov.—1(2):905
pus nov.—1(2):908
Agapetes salicifolia var. glanduliflora Banik & Sanjappa,
var. nov.—1(2):910
Agasthiyamalaia S. Rajkumar & Janarth., gen. nov.—
1(1):130
In: “e
(Bedd.) S. Rajkumar €: Janarth.,
comb. nov.—1(1):131
icr montium-sancticaroli García-Mend., sp. nov.—
1):79
"meus nycteris Medecilo, Yao & Madulid, sp. nov.—
1(2):817
Anisocapparis X. Cornejo & H.H. Iltis, gen. nov.—2(1):62
Anisocapparis speciosa (Griseb.) X. Cornejo & H.H. Iltis,
comb. nov.—2(1):65
Aphanelytrum peruvianum Sánchez Vega, P.M. Peterson,
Soreng & Laegaard, sp. nov.—1(2):842
lis (N. Busch) Elven
& D.F. Murray, comb. nov.—2(1):438
iris s petraea subsp. umbrosa (Turcz. ex Steud.)
D.F. Murray, comb. nov.—2(1):439
d dde sorzogonensis f. orizabensis (E. Fourn.) Strahan &
ini: cil nov.—2(1):322
, Sp. nov.—1(1):143
(Rydb) D.F Murray &
A vt H L L | [|
Elven, comb. et stat. nov.—2(1 ):44
Asplenium tutwilerae B.R. Keener E L.J. Davenport, sp.
nov.—1(1):104
Baccharis apicifoliosa A.A. Schneid. & Boldrini, sp. nov.—
2(1):45
Baccharis flexuosiramosa A.A. Schneid. € Boldrini, sp.
nov. —2(1):48
Besleria neblinae Feuillet, sp. nov.—2(1):269
Besleria yatuana Feuillet, sp. nov.—2(1):271
on elliptica (Willd. ex Spreng.) V.V. Petrovsky, D.F. Murray
& Elven, comb. nov.—2(1):435
Boerhavia Nap var. intermedia (M.F. Jones) Spellenb.,
comb. nov. 2):874
Bouteloua tamaulipensis c - Pierce ex D. Pacheco & Co-
lumbus, sp. nov.—2(2):9
Cal
Elven, , comb. et stat. nov.—2(1):434
priiis websteri B. Holst & M.L. Kawasaki, sp.
1):297
p. laricina (Louis-Marie)
e a (O.W. Knight) C.D. Kirschbaum, comb. et stat
nov.—1(1):401
Carex klamathensis B.L. Wilson €: L.P. Janeway, sp. nov.—
10:71
Index: Vols. 1 & 2
Chusquea annagardneriae "E Clark, C.D. Tyrrell, Triplett &
A. Fisher, sp. nov.—1(2):84
Colicodendron bahianum : Cornejo & H.H. litis, sp.
(1):76
Colicodendron valerabellum H.H. Iltis, T. Ruiz & G.S. Bunting,
Sp. nov.—2(1):82
Coptidium x spitsbergense (Hadac) Elven, comb. nov.—
437
Cotoneaster floridus J. Fryer & B. Hylmó, sp. nov.—2(1):55
Cotoneaster qungbixiensis J. Fryer & B. Hylmó, sp.
nov.—2(1):53
Cotoneaster series Sterniani J. Fryer & B. Hylmó, ser.
nov.—2(1):53
Crataegus series Apricae J.B. Phipps, ser. nov.—1(1):173
Crataegus aquacervensis J.B. Phipps & O’Kennon, sp.
nov.—1(2):1061
Crat |
ganin le
g.) J.B. Phipps, comb.
nov.—1(2):1007
Crataegus chrysocarpa var. E (Sarg.) J.B. Phipps,
comb. et stat. nov.—1(2
Crataegus chirysocarpa ve var. prascor (Sarg.) J.B. Phipps,
mo et m nov.—1(2):10
J.B. Phipps, var.
nov.—1 (2):1008
Crataegus coccinea var. pringlei (Sarg.) J.A. Macklin & JB.
Phipps, comb. et stat. nov.—1(2):1005
Crataegus cupressocollina J.B. Phipps & O'Kennon, sp.
nov.—1(2):1056
Crataegus series Cupressocollinae J.B. Phipps & O'Kennon,
ser. nov.— 5
Crataegus lancei J.B Phipps, sp. nov.—2(2):1143
Crataegus mollis var. incisifolia Kruschke, var. nov.—
1(2):1012; 2(1):473
Crataegus series Montaninsulae J.B. Phipps & O'Kennon,
ser. nov.—1(2):1065
Crataegus dede var. ar (Sarg.) J.B. Phipps, comb.
et stat. nov.—1(2
E. purpurea n Phipps & O'Kennon, sp. nov.—
1(2):10
I ut:/D
) J.B. Phipps, comb.
et stat. nov.—1(2):1006
Crataegus rivuloadamensis J.B. Phipps & O'Kennon, sp.
nov.—1(2):1065
Crataegus rivulopugnensis J.B. Phipps & O'Kennon, sp.
nov.—1(2):1070
Crataegus rubribracteolata J.B. Phipps & O'Kennon, sp.
nov.—1(2):1073
Crataegus schuettei var. gigantea Kruschke, var. nov.—
1(2):1013; 2(1):473
Mas sheila-phippsiae var. cor "emm J.B.
O'Kennon, var. nov.—1(2):10
Eu: dcr J.B.Phiops a O'Kennon, sp.
nov.—1(2):10
Crataegus Mun var. velutina (Sarg.) J.B. Phipps, comb. et
stat. nov.—1(2):1009
Cretacifilix G.O. Poinar & R. Buckley, gen. nov.—2(2):1
Misi oe G.O. Poinar & R. Buc an Sp.
nov.—2(2):1
1525
Cuatresia amistadensis D.A. Soto & A.K. Monro, sp.
nov. —2(1):41
Cyclopogon secundum Christenson, sp. nov.—1(2):1001
Cyperus aan Wad. Khan & puse, sp. nov.—
2(1):37
Dra modesta (Huck) Huck, comb. et stat. nov.—
2(2):1163
Diospyros acapulcensis subsp. chiquimulensis M D ~
vance, l. García 8: A.C. Sanders, subsp. nov.—2(2):10
Diospyros acapulcensis subsp. dwyeri M. m Froverce l.
García & A.C. Sanders, subsp. nov.—2(2):10
Diospyros acapulcensis subsp. guanacastensis M.C. Pro-
vance, |. García & A.C. Sanders, subsp 2):1029
Diospyros acapulcensis subsp. loe M.C. Pro-
vance, |. García & A.C. Sander
Diospyros acapulcensis subsp. nicaraguensis (Stand).
M.C. Provance, |. García & A.C. Sanders, comb. et stat.
nov.—2(2):1033
Diospyros acapulcensis subsp. pedromorenoi M.C. Pro-
vance, |. García & A.C. Sanders, subsp. nov.—2(2):103
Diospyros acapulcensis subsp. rivensis M.C. Provance, |.
García €: A.C. Sanders, subsp. nov.—2(2):1039
Diospyros acapulcensis subsp. veraecrucis (Standl.)
M.C. jS |. García & A.C. Sanders, comb. et stat.
nov.—2(2):10
Diospyros e Due subsp. pics A Provance, |. García
nders, subsp no 2(2):1
S subsp nov.—
Diospyros aequoris subsp. chutlensis M. eee l.
Nd & A.C ander, subsp. no —2(2):10
.ex
enworth) M. C E I. García & A.C. Sanders mis et
stat. nov.—2(2):1069
Diospyros a subsp. rekoi ae : M.C. Provance, |.
García & A.C. Sanders, comb. et stat. nov.—2(2):1071
Diospyros aequoris subsp. OS M.C. Pro-
vance, |. García & A.C. Sanders, subsp. nov.—2(2):1074
Diospyros yucatanensis subsp. spectabilis dise
M.C. pid ha García & A.C. Sanders, comb. et
nov.—2(2):10
yoy jeq is Salino, sp. nov.—2(2):
Dryas punctata subsp. hookeriana (Juz.) Jurtz, comb.
nov.—2(1):441
liz || (C. Hitrkhr Y Err+, a+ et combh
ov.—1(1):43
Drymocallis cuneifolia var. ewanii (D.D. Keck) Ertter, stat. et
como. nov.—1(1):44
Drymocallis deseretica Frtter, sp. nov.—1(1):4
Drymocallis glandulosa var. reflexa (Greene) ca comb.
V.—1(1):43
Drymocallis glandulosa var. viscida (Parish) Ertter, stat. et
comb. nov.—1(1):43
madii glandulosa var. wrangelliana (Fisch. & Avé-
Lall.) Ertter, comb. nov.—1(1):43
Drymocallis lactea var. austiniae (Jeps.) Ertter, comb.
nov. —1(1
Drymocallis pseudorupestris var. crumiana D.D. Keck ex
Ertter, var. nov.—1(1):39
1526
Drymocallis oe var. saxicola Ertter, var.
1):37
nov.—
Dudleya abramsii subsp. costifolia (Bartel & Shevock)
Moran, comb. nov.—1(2):1015
Dudleya abramsii subsp. setchellii (Jeps) Moran, comb.
nov.—1(2):1015
Dysphania andicola (Phil) Mosyakin 8: Clemants, comb.
nov.—2(1):428
iunio bonariensis (Hook. £) Mosyakin & Clemants,
comb. no 428
n puranti (Aellen) Mosyakin & Clemants, comb.
—2(1):42
Ma UE (Hauman) Mosyakin & Clemants,
comb. nov.—2(1):429
Dysphania dissecta (Moq.) Mosyakin 8: Clemants, comb.
nov.—2(1):429
l'a" L H J (| F. Si A Y L;
& Clemants, comb
nov.—2(1):428
Dysphania mandonii (S. Watson) Mosyakin & Clemants,
NUS Lanes 42
OM DI LA RA L:
& Clemants,
“Comb: nov.—2(1):42
Dysphania microcarpa NERIS Mosyakin & Clemants, comb.
nov.—2(1):428
Dysphania orna (Aellen) Mosyakin & Clemants, comb.
nov.— —2042
D Colla) Mosyakin & Clemants, comb
nov— —2(1):42
Dysphania o (Speg.) Mosyakin 8: Clemants,
comb. nov.—2(1):428
Dysphania procera (Hochst. ex Moq.) Mosyakin & Clemants,
comb. nov.—2(1):429
Dysphania pusilla (Hook. f) Mosyakin & Clemants, comb.
nov.—2(1):427
Dysphania retusa (Juss. ex Moq.) Mosyakin € Clemants,
comb. nov.—2(1):4
Rican 2 a Wilson) Mosyakin & Clemants,
comb. n
Don sooana m Mosyakin & Clemants, comb.
nov.—2(1):42
+ (Cr Al ARA Les
& Clemants, comb
nov.—2(1):429
Dysphania tomentosa (Thouars) Mosyakin & Clemants,
comb. nov.—2(1):428
gon dece truncata (PG. Wilson) Mosyakin €: Clemants,
comb 1):427
Dysphania venturii (Aellen) Mosyakin & Clemants, comb.
nov.—2(1):428
Eleocharis acutangula subsp. breviseta D.J. Rosen, subsp.
v.—1(2):885
g p. neotropica D.J. Rosen, subsp
nov.—1(2):886
Eoépigynia Poinar, Chambers & Buckley, gen. nov.—
Eoépigynia burmensis Poinar, Chambers & Buckley, sp.
1):9
Episcia duidae Feuillet, sp. nov.—2(1):275
Episcia rubra Feuillet, sp. nov.—2(1):277
[| L af ob D ID h TRES
J i i f Texas 2(2)
LOC
ter & Elven, comb. et stat. nov.—2(1):433
Erigeron heleniae Nesom, sp. nov.—1(2):891
ugenia concava B. Holst & M.L. Kawasaki, sp. nov.—
2(1):299
Festuca m -ledesmae S.J. Darbyshire, nothosp.
nov.—1(2):832
Fugenia (grossa B. A L. Kawasaki, sp. nov.—2(1):299
g S.J. Darbyshire, sp. nov.—1(2):827
F i lla SJ. Darbyshire, nothosp nov.—1(2):835
Festuca roemeri var. klamathensis B.L. Wilson, var.
nov.— 5
deis naikii Wad. Khan € Lakshmin., sp. nov.—
2(1):38
I . fc LAN C la R8, M/aalil comb
af
OV. ):336
Modus imi a subsp. raupii (A.F. Porsild) Elven,
comb.n 1):442
Geum donianum (Tratt.) Weakley €: Gandhi, comb. nov.—
2(1):4
Gratiola a D. Estes, sp. nov.—1(1):166
Gratiola quartermaniae D. Estes, sp. nov.—1(1):163
Londoño & Zurita, sp. nov.—2(1):31
Guaduai Incana Londoño, sp. nov.—2(1):26
Gutierrezia elegans Al Schneider & P. Lyon, sp. nov.—
2(2):771
pea ekmanii ann nom. nov.—2(2):1
dasycarpum (T. E D.F. Murray
& Elven, comb. et stat. ROV AUT :441
Heterotheca arizonica (Semple) Nesom, comb. et stat.
nov.—1(1):
Heterotheca nitidula (Woot. & Standl) Nesom, comb.
nov.—1(1):24
Heuchera woodsiaphila | PJ. and sp. nov.—2(1):448
sp. nov.—1(1):137
Hosackia crassifolia var. -otayensis (Moran ex Isely) Brouillet,
comb. nov.—2(1):38
g ) Brouillet, comb
nov.—2(1):388
ind coa var. ottleyi (Isely) Brouillet, comb.
Ov.—
Houstonia is Porotis Terrell, subg. nov.—1(1):117
mboldtia brunonis var. raktapushpa P5. Udayan, KV.
^ shar & Satheesh George, var. nov.—1(1):121
lllicium guajaibonense (Imkhanitskaya) Judd & Abbott,
comb. et stat. nov.—2(2):800
Ipomoea tehuantepecensis L. Torres-Colín, R. Torres-Colín,
M.P Ramírez & J.A. McDonald, sp. nov.—2(2):793
Ixora araguaiensis Delprete, sp. nov.—2(1):456
ixora congestiflora Delprete, sp. nov.—2(1):456
Ixora irwinii Delprete, sp. nov.—2(1):459
asi digitatus C. idu & Zika, sp. nov.—2(2):775
inke & Kaye, sp. nov.—1(1):10
| seminuda (Trautv) Elven, comb
2
WU. I : Cp paa |
nov.— 434
Kohleria hypertrichosa J.L. Clark & L.E. Skog, sp. nov.—
2(1):20
Index: Vols. 1 & 2
Lachnociona G. > Poinar, KL. Chambers & R. Buckley, gen.
nov.—2(2):11
Lachnociona Em O. Poinar, KL. Chambers & R. Buckley,
p. nov.—2(2):1184
Lasiambix Poinar, Chambers & Brown, gen. nov.—2(1):464
Lasiambix dominicensis Poinar, Chambers & Brown, sp.
nov.—2(1):464
Lasiolaena lychnophorioides Roque & H. Rob., sp.
nov.—2(2):812
Lechea pulchella var. ramosissima (Hodgdon) Sorrie &
Weakley, comb. nov.—1(1):370
Ds alba Pd gracilis (Howell) Morin, comb. et
stat. nov.—1(2):10
Limnanthes alba sb parishii (Jeps) Morin, comb. et
stat. nov.—1(2):101
Limnanthes hri subsp. striata (Jeps) Morin, comb.
et stat. nov.—1(2):1017
Limnorchis aquilonis (Sheviak) Rebrist. & Elven, comb.
nov.—2(1):434
Limnorchis huronensis (Nutt.) Rebrist. & Elven, comb.
nov.—2(1):434
Lobelia gloria-montis subsp. longibracteata (Rock) Lam-
mers, stat. nov,—1(2):806
Lobelia xkauaiensis (A. Gray) A. Heller, stat. nov.—1(2):801
Lobelia koolauensis (Hosaka & Fosberg) Lammers, comb.
et stat. nov.— 803
hi Lammers, sp. nov.—1 (2):802
J.F. Morales, sp. nov.—1(2):853
Mandevilla frigida J. F. Morales, sp. nov.—1(2):855
Mandela horrida J. i cloves sp. nov.—1(2):863
rales, sp. nov.—1(2):865
dep lojana JF. ME sp. nov.—1(2):855
evilla megabracteata J.F. Morales, sp. nov.—
Lobelia
HII + "EN A
Mandevilla similaris J.F. Morales, sp. nov.—1(2):867
Matelea costata var. goodfriendii (Proctor) Krings, comb.
nov.—
Matelea dictyopetala (Urb. & Ekman) Krings, comb.
nov.—2(1):130
Matelea proctori Krings, nom. nov.—2(1):151
Matelea pubescens (Griseb.) Krings, comb. nov.—2(1):131
Medranoa johnstonii (G.L. Nesom) G.L. Nesom, comb.
nov.—1(1):148
Medranoa palmeri (A. Gray) G.L. Nesom, comb. nov.—
1(1):148
Medranoa Eo lee (S.F. Blake) G.L. Nesom, comb.
nov.—1(1):1
Medranoa LR (Brandeg.) G.L. Nesom, comb. nov.—
1(1):148
Meliosma antioquiensis X. Cornejo, sp. nov.—2(1):57
Micranthes bryophora (A. Gray) Brouillet & Gornall, comb.
nov.—1(2):1020
Micranthes calycina (Sternb.) Gornall & H. Ohba, comb.
nov.—1(2):1020
g ) Brouillet & Gornall, comb.
nov.—1(2):1020
Micranthes foliolosa (R. Br.) Gornall, comb. nov.—
1(2):1020
1527
Micranthes gormanii (Suksd.) Brouiliet & Gornall, comb.
IN) 1020
| longi folia (E
gl. 8: Irmsch.)
Elven 8: D.F. Murray, comb. nov.—2(1) 439
Micranthes idahoensis (Piper) Brouillet & Gornall, comb.
nov.—1(2):1020
Micranthes lyallii ad ipis vira & Savile) Elven &
D.F. Murray, comb. nov.—
Micranthes merkii s ex inet Elven & D.F. Murray,
comb. nov.—2(1):440
Micranthes mexicana ~ & Irmsch.) Brouillet & Gornall,
com IR —1(2): 10
RA: H | "" I. /C;
\
h. &C.A. Mey)
Elven & D.F. Murray, comb. nov—2(1 ):440
Micranthes nelsoniana subsp. insularis (Hultén) Elven &
D.F. Murray, comb. nov.—2(1):440
Micranthes nelsoniana var. aestivalis (Fisch. & C.A. Mey)
Gornall & H. Ohba, comb. et stat. nov.—1(2):1020
Micranthes nelsoniana var. carlottae PE & ~ Gor-
nall & H. Ohba, comb. et stat. nov.—1(2
Micranthes nelsoniana var. cascadensis m & Savile)
Gornall & H. Ohba, comb. et stat. nov.—1(2):1020
Micranthes nelsoniana var. insularis (Hultén) Gornall & H.
Ohba, comb. et stat. nov.—1(2):1020
Micranthes nelsoniana var. pacifica (Hultén) Gornall & H.
ba, comb. et stat. nov.—1(2):1020
iiis edipi var. eae oo & Savile)
nall & H. Ohba, comb. nov.—2(1
ice nudes (D. Don) Pd A H. Ohba, comb.
nov.—1(2):1
Micranthes petiolaris (Raf.
nov.—1(2):
Micranthes duc (Calder & Savile) Elven & D.F. Murray,
comb. nov.—2(1):440
re pii x Yu. Zhmylev) Brouillet 8: Gornall,
—1(2):10
rl eee en Elven & D.F. Murray, comb.
nov.—2(1):441
Micranthes tempestiva ed & Denton) Brouillet &
Gornall, comb. nov.—1(2):10
Micranthes "D (Skelly) ale & Gornall, comb.
nov.—1 (2):10
Mcd foni dnd & A. Gray) Brouillet & Gornall,
comb 1(2):10
Brouillet & Gornall, comb.
—
Pe pee (Sternb.) Gornall & H. Ohba,
02
Menillcatpa * conejo d H. E Iltis, n nov.—2(1):67
C), X. Cornejo & H.H. Iltis,
comb. nov.—2(1):71
Monilicarpa tenuisiliqua (Jacq) X. Cornejo & HH. Iltis, comb.
nov.—2(1):70
Mosiera androsiana (Urban) Salywon, comb. nov.—
1(2):899
Mosiera cuspidata (Alain) Salywon, comb. nov.—1(2):899
Mosiera gracilipes (Alain) Salywon, comb. nov.—1(2):899
Mosiera xerophytica (Britton) Salywon, comb. nov.—
1(2):900
Munrochloa M. Kumar €: Remesh, gen. nov.—2(1):374
1528
Munrochloa ritchiei (Munro) M. Kumar & Remesh, comb.
nov.—2(1):376
Myrcia subcordifolia B. Holst & M.L. Kawasaki, sp. nov.—
2(1):303
esae candidulus (Miers) Feuillet, comb. nov.—
2(1):264
M maculatus (Jacg.) Feuillet, comb. nov.—
2(1
myropus pem (Chamisso) Feuillet, comb. nov.—
2(1):26
Feuillet,
—
Myriopus acude var. spigeliiflorus (A. DC,
omb. nov.—2(1):264
Nautilocalyx crenatus Feuillet, sp. nov.—2(2):825
Nautilocalyx pusillus Feuillet, sp. nov.—2(2):827
Nautilocalyx roseus Feuillet, sp. nov.—2(2):830
Nautilocalyx ruber Feuillet, sp. nov.—2(2):830
Nautilocalyx vestitus Feuillet, sp. nov.—2(2):833
Nemacladus australis (Munz) Morin, comb. et stat.
nov.—2(1):398
Nemacladus calcaratus Morin, so. nov.—2(1):399
Nemacladus californicus (A. Gray) Morin, comb. nov.—
2(1):397
Nemacladus orientalis (McVaugh) Morin, comb. et stat.
nov.—2(1):398
Nemacladus secundiflorus var. robbinsii Morin, var.
nov.—2(1):39
Nemacladus tenuis (McVaugh) Morin, comb. et stat.
nov.—2(1):398
Nemacladus tenuis var. aliformis Morin, var. nov.—
2(1):398
Neocalyptrocalyx morii X. Cornejo & H.H. Iltis, sp. nov.—
2(2):807
Orthaea laurifolia (M. Martens & Galeotti) Luteyn, comb.
nov.—2(1):244
eel sect. Capitulum E.A. Christ. sect. nov.—
eum sect. ee (Kraenzl) E.A. Christ.,
ae NOTA ):28
P lis sul lica (Jurtz, Korobkov
& VV. Petrovsky) Jurtz., Korobkov &VV. Petrovsky, comb.
nov.—
Papaver labradoricum (Fedde) Solstad & Elven, comb. et
stat. nov.—2(1):438
Pappostipa (Speg.) Romaschenko, PM. Peterson & Soreng,
mb. et stat. nov—2(1):181
Pappostipa ameghinoi (Speg.) Romaschenko, comb.
ov.—2(1):184
Pappostipa ameghinoi var. digona (Parodi) Romaschenko
comb. nov.—2(1):184
ee ameghinoi var. precordillerana (F.A. Roig)
ko, comb. nov.—2(1):184
haies atacamensis (Parodi) Romaschenko, comb.
nov.—2(1):184
Pappostipa barrancaensis (F.A. Roig) Romaschenko, comb.
nov.— 2(1):184; 2(2):1233
[| Il ESL Das ni D A
f Texas 2(2)
Pappostipa braun-blanquetii (F.A. Roig) Romaschenko,
comb. nov.—2(1):183
Pappostipa chrysophylla (E. Desv) Romaschenko, comb.
nov.—2(1):185
Pappostipa chrysophylla var. cordillerarum (Parodi) Ro-
maschenko, comb. nov.—
Pappostipa chrysophylla var. Eee (Kuntze) Romas-
ds ena dogm 2(1):18
f. minuta (FA.R
1D | 1
comb. nov.—2(1):1 85
Pappostipa chrysophylla f. modica (F.A. Roig) Romas-
chenko, comb. nov.—2(1):185
Pappostipa sect. Chrysovaginatae Romaschenko, sect.
nov.—2(1):184
Pappostipa chubutensis (Speg.) Romaschenko, comb.
nov.—2(1):185
Pappostipa chubutensis var. hirsutissima (FA. Roig) Ro-
maschenko, comb. nov.—2(1):185
ipsia frigida (Phil) Romaschenko, comb. nov.—
LM bs
J H H la /D iD L |
comb. nov—2(1 ):185
Pappostipa oo (Pilg.) Romaschenko, comb.
nov.—2(1):183
Pappostipa ins (Cav.) Romaschenko, comb. nov.—
2(1):185
n 1D L I
T
comb. nov.—2(1):185
Pappostipa humilis var. ruiziana (Parodi) Romaschenko,
comb. nov.—2(1):185
Pappostipa ibarii (Phil) Romaschenko, comb. nov.—
2(1):184
Pappostipa ibarii var. anomala (Parodi) Romaschenko,
comb. nov.—2(1):184
[D i^n | |
comb
nov.—2(1):184
Pappostipa Prid (FA. Roig) Romaschenko, comb.
nov.—2(1):18
pci is major (Speg.) Romaschenko, comb. et stat.
nov.—2
o has (F.A. Roig) Romaschenko, comb.
nov.—2(1):183
ee nana (Speg.) Romaschenko, comb. nov.—
2(1):18
a nicorae (F.A. Roig) Romaschenko, comb.
nov.—2(1):183
Pappostipa sect. Pappostipa, sect. nov.—2(1):18
Pappostipa parodiana (F.A. Roig) lo comb.
ov.—2(1):183
Pappostipa patagonica (Speg.) Romaschenko, comb.
nov.—2(1):183
Pappostipa ruiz-lealii (FA. Roig) Romaschenko, comb.
nov.—2(1):1
Pappostipa semperiana (F.A. Roig) Romaschenko, comb.
nov.—2(1):183
Pappostipa sorianoi (Parodi) Romaschenko, comb.
nov.—2(1):184
Index: Vols. 1 & 2
Pappostipa speciosa (Trin. & Rupr.) Romaschenko, comb.
nov.—2(1):182
Pappostipa speciosa f. abscondita (F.A. Roig) Romaschenko,
comb. nov.—2(1):182
Pappostipa speciosa var. atuelensis (F.A. Roig) Romas-
chenko, comb. nov.—2(1):182
Pappostipa speciosa var. breviglumis (Parodi) Romas-
chenko, comb. nov.—2(1):182
Pappostipa speciosa var. ciliata (FA. Roig) Romaschenko,
comb. nov.—2(1):182
Pappostipa speciosa f. horrida (F.A. Roig) Romaschenko,
comb. nov.—2(1):182
Pappostipa speciosa var. manqueclensis (F.A. Roig) Romas-
chenko, comb. nov.—2(1):182
Pappostipa speciosa var. media (Torres) Romaschenko,
comb. nov.—2(1):182
Pappostipa speciosa var. parva (F.A. Roig) Romaschenko,
comb. nov.—2(1):182
Pappostipa vaginata (Phil. Romaschenko, comb. nov.—
2(1):182
Pappostipa vaginata var. argyroidea (F.A. Roig) Romas-
chenko, comb. nov.—2(1):183
Pappostipa vaginata f. contracta (F.A. Roig) Romaschenko,
comb. nov.—2(1):183
Pappostipa vaginata var. dilatata (F.A. Roig) Romaschenko,
comb. nov.—2(1):183
Pappostipa vaginata f. inmersa (F.A. Roig) Romaschenko,
comb. nov.—2(1):183
Pappostipa vaginata f. laevis (F.A. Roig) Romaschenko,
comb. nov.—2(1):183
Pappostipa vaginata f. rigida (F.A. Roig) Romaschenko,
comb. nov.—2(1):183
Pappostipa vatroensis (F.A. Roig) Romaschenko, comb.
nov.—2(1):184
Passiflora angusta Feuillet & J.M. MacDougal, sp. nov.—
2(2):818
Passiflora arta Feuillet, sp. nov.—1(2):819
Passiflora compar Feuillet, sp. nov.—1(2):821
Passiflora davidii Feuillet, sp. nov.—1(2):896
Passiflora foetida var. orinocensis (Killip) Feuillet, stat.
nov.—1(1):144
Passiflora insueta Feuillet & MacDougal, nom. nov.—
2(1):267
Passiflora rufa Feuillet & J.M. MacDougal, sp. nov.—
2(2):822
Passiflora tecta Feuillet, sp. nov.—2(1):281
Passiflora sect. Trematanthera (Leeuwenb.) Feuillet, sect.
nov.—2(1):277
Philadelphus microphyllus var. argyrocalyx (Wooton)
Henrickson, comb. & stat. nov.—1(2):901
Philadelphus microphyllus var. madrensis (Hemsley)
Henrickson, comb. & stat. nov.—1(2):901
Philadelphus microphyllus var. stramineus (Rydb.) Hen-
rickson, comb. & stat. nov.—1(2):901
Phragmites australis subsp. berlandieri (E. Fourn.) Saltons-
tall & Hauber, comb. nov.—1(1):387
Pilea alfaroana Al. Rodr. & A.K. Monro, sp. nov.—2(2):995
Pilea gamboana Al. Rodr. & A.K. Monro, sp. nov.—2(2):997
1529
Pilea herrerae Al. Rodr. & A.K. Monro, sp. nov.—2(2):1000
Pilea longibracteolata Al. Rodr., A.K. Monro & L. Acosta, sp.
nova.—2(2):1002
Pilea moragana Al. Rodr. & A.K. Monro, sp. nov.—2(2):1004
Poa pfisteri Soreng, sp. nov.—2(2):850
Potentilla anserina subsp. yukonensis (Hultén) Soják ex
Elven & D.F. Murray, comb. nov.—2(1):441
Potentilla arenosa subsp. chamissonis (Hultén) Elven & D.F.
Murray, comb. nov.—1(2):813
Potentilla demotica Ertter, sp. nov.—1(1):53
Potentilla holmgrenii D.F. Murray & Elven, sp. nov.—
1(2):811
Potentilla hyparctica subsp. elatior (Abrom.) Elven & D.F.
Murray, comb. nov.—1(2):813
Potentilla jepsonii Ertter, nom. nov.—2(1):202
Potentilla rhyolitica Ertter, sp. nov.—1(1):50
Potentilla rhyolitica var. chiricahuensis Ertter, var.
nov.—1(1):52
Puccinellia phryganodes subsp. neoarctica (Á. Lóve & D.
Lóve) Elven, comb. et stat. nov.—2(1):435
Puccinellia phryganodes subsp. sibirica (Hadac & Á. Lóve)
Elven, comb. nov.—2(1):435
Pycnanthemum sect. Californicae K.L. Chambers € H.L.
Chambers, sect. nov.—2(1):195
Raddiella vanessiae Judz., sp. nov.—1(1):1
Ranunculus subborealis subsp. pumilus (Wahlenb.) Elven,
comb. nov.—2(1):437
Ranunculus subborealis subsp. villosus (Drabble) Elven,
comb. et stat. nov.—2(1):438
Rhododendron tomentosum subsp. decumbens Elven &
D.F. Murray, comb. nov.—2(1):441
Ribes oxyacanthoides var. cognatum (Greene) Morin,
comb. et stat. nov.—1(2):1015
Rosaceae tribus Lyonothamneae Brouillet, tribus nov.—
2(1):385
Rosaceae subseries Robustae J.B. Phipps, subser. nov.—
2(2):1124
Rosaceae subseries Tenues J.B. Phipps, subser. nov.—
2(2):1108
Salvia toaensis Alain, nom. nov.—2(2):1165
Scorzoneroides autumnalis subsp. pratensis (Hornem.)
Elven, comb. nov.—2(1):443
Sedum lanceolatum var. subalpinum (Fród.) H. Ohba, comb.
& stat. nov.—1(2):890
Sedum laxum var. eastwoodiae (Britton) H. Ohba, comb. &
stat. nov.—1(2):889
Sedum laxum var. flavidum (Denton) H. Ohba, comb. &
stat. nov.—1(2):889
Sedum laxum var. heckneri (M. Peck) H. Ohba, comb. & stat.
nov.—1(2):889
Sedum laxum var. latifolium (R.T. Clausen) H. Ohba, comb.
& stat. nov.—1(2):890
Sedum obtusatum var. boreale (R.T. Clausen) H. Ohba,
comb. & stat. nov.—1(2):889
Sedum obtusatum var. paradisum (Denton) H. Ohba, comb.
& stat. nov.—1(2):889
Sedum obtusatum var. retusum (Rose) H. Ohba, comb. &
stat. nov.—1(2):889
1530
Sedum oreganum var. tenue (R.T. Clausen) H. Ohba, comb.
& stat. nov.—1(2):890
Sedum radiatum var. ciliosum (Howell) H. Ohba, comb. &
stat. nov.—1(2):889
Sedum radiatum var. depauperatum (R.T. Clausen) H. Ohba,
comb. & stat. nov.—1(2):889
Sedum stenopetalum var. monanthum (Suksd.) H. Ohba,
comb. & stat. nov.—1(2):890
Sedum wrightii var. densiflorum (R.T. Clausen) H. Ohba,
comb. & stat. nov.—1(2):889
Sedum wrightii var. priscum (R.T. Clausen) H. Ohba, comb.
& stat. nov.—1(2):889
Sidalcea hickmanii subsp. napensis S.R. Hill, subsp.
nov.—2(2):783
Sidalcea hickmanii subsp. pillsburiensis S.R. Hill, subsp.
nov.—2(2):/86
Silene involucrata subsp. furcata (Raf) V.V. Petrovsky & Elven,
comb. et stat. nov.—2(1):436
Silene soczavana var. macrosperma (A.E. Porsild) V.V. Petrov-
sky, D.F. Murray & Elven, comb. et stat. nov.—2(1):436
Silene villosula (Trautv.) V.V. Petrovsky & Elven, comb.
nov.—2(1):437
Silene violascens (Tolm.) V.V. Petrovsky & Elven, comb.
nov.—2(1):437
Siphonandra nervosa Luteyn & E.M. Ortiz, sp. nov.—
2(1):256
Siphonandra santa-barbarense Luteyn & E.M. Ortiz, sp.
nov.—2(1):256
i Itiradiata subsp. arctica (DC) Korobkov & Elven,
comb. et stat. nov.—2(1):443
Spermacoce burchellii (E.L. Cabral & Bacigalupo) Delprete,
comb. nov.—1(2):1025
Spermacoce crispata (K. Schum.) Delprete, comb. nov.—
1(2):1025
Spermacoce delicatula (E.L. Cabral) Delprete, comb.
nov.—1(2):1025
Spermacoce dimorpha (J.H. Kirkbr.) Delprete, comb.
nov.—1(2):1025
Spermacoce incognita (F.L. Cabral) Delprete, comb.
nov.—1(2):1025
Spermacoce irwiniana (E.L. Cabral), Delprete, comb.
nov.—1(2):1025
Spermacoce multiflora (DC.) Delprete, comb. nov.—
1(2):1026
Spermacoce paraensis (E.L. Cabral & Bacigalupo) Delprete,
comb. nov.—1(2):1027
Spermacoce pulchristipula (Bremek.) Delprete, comb.
nov.—1(2):1027
Spermacoce semiamplexicale (E.L. Cabral) Delprete, comb.
nov.—1(2):1028 ;
Spermacoce spicata (Miq.) Delprete, comb. nov.—
1(2):1028
Spermacoce tocantinsiana (E.L. Cabral & Bacigalupo) Del-
prete, comb. nov.—1(2):1028
Stachys iltisii J. Nelson, sp. nov.—2(2):761
Stuckenia filiformis subsp. borealis (Raf) Tzvelev & Elven,
comb. nov.—2(1):433
Taxus biternata Spjut, sp. nov.—1(1):266
Cnlidaan
Journal of the Botanical Research Institute of Texas 2(2)
Taxus brevifolia var. polychaeta Spjut, var. nov.—1(1):217
Taxus brevifolia var. reptaneta Spjut, var. nov.—1(1):219
Taxus caespitosa var. angustifolia Spjut, var. nov.—
1(1):268
Taxus caespitosa var. latifolia (Pilger) Spjut, comb.
nov.—1(1):269
Taxus canadensis var. adpressa (Hort. ex Carriére) Spjut,
comb. nov.—1(1):274
Taxus canadensis var. minor (Michx.) Spjut, comb.
nov.—1(1):274
Taxus contorta var. mucronata Spjut, var. nov.—1(1):260
Taxus florinii Spjut, sp. nov.—1(1):222
Taxus globosa var. floridana (Nutt. ex Chapm.) Spjut, comb.
nov.—1(1):224
Taxus kingstonii Spjut, sp. nov.—1(1):240
Taxus mairei var. speciosa (Florin) Spjut, comb. et stat.
nov.—1(1):246
Taxus obscura Spjut, sp. nov.—1(1):235
Taxus phytonii Spjut, sp. nov.—1(1):237
Taxus suffnessii Spjut, sp. nov.—1(1):226
Taxus umbraculifera var. hicksii (Hort. ex Rehder) Spjut,
comb. nov.—1(1):279
Taxus umbraculifera var. microcarpa (Trautv.) Spjut, comb.
nov.—1(1):279
Taxus umbraculifera var. nana (Rehder) Spjut, comb.
nov.—1(1):281
Tetrazygia paralongicollis Judd, lonta, Clase & Skean, sp.
nov.—2(1):35
Thamnocalamus chigar (Stapleton) Stapleton, comb.
nov.—1(1):139
Thamnocalamus nepalensis (Stapleton) Stapleton, stat.
nov.—1(1):139
Thamnocalamus occidentalis (Stapleton) Stapleton, stat.
nov.—1(1):139
Thecagonum strigulosum (DC) Terrell & H. Rob., comb.
nov.—1(1):377
Tocoyena arenicola Delprete, sp. nov.—2(2):988
Trochanthera G.O. Poinar, K.L. Chambers & A.E. Brown, gen.
nov.—2(2):1168
Trochanthera lepidota G.O. Poinar, K.L. Chambers € A.E.
Brown, sp. nov.—2(2):1168
Vaccinium beamanianum Wilbur & Luteyn, nom. nov.—
2(1):219
Vaccinium campanense Wilbur € Luteyn, sp. nov.—
2(1):234
Vaccinium chihuahuense Wilbur € Luteyn, sp. nov.—
2(1):220
Vaccinium uliginosum subsp. vulcanorum (Kom.) Alsos &
Elven, comb. et stat. nov.—2(1):442
Vahlodea latifolia subsp. paramushirensis (Kudó) Elven,
comb. nov.—2(1):435
Varronia bullata subsp. humilis (Jacq.) Feuillet, comb.
nov.—2(2):837
Varronia cremersii (Feuillet) Feuillet, comb. nov.—
2(2):838
Varronia marioniae (Feuillet) Feuillet, comb. nov.—
2(2):839
Xyris panacea L.C. Anderson & Kral, sp. nov.—2(1):1
AVAILABLE TITLES FROM THE BRIT PRESS
SBM 01 | A Quantitative Analysis of the Vegetation on the Dallas County White Rock Escarpment
| SBM 02 | The Vascular Flora of St. Francis County, Arkansas
| SBM 04 | Asteraceae of Louisiana
SBM 05 | The Genus Mikania (Compositae: Eupatorieae) in México
SBM 06 | Frontier Botanist William Starling Sullivant's Flowering-Plant Botany of Ohio
SBM 07 | A Taxonomic Revision of the Acaulescent Blue Violets (Viola) of North America
~ SBM 08 | Aster & Brachyactis (Asteraceae) in Oklahoma
SBM 09 | The Genus Mikania (Compositae: Eupatorieae) in the Greater Antilles
SBM 10 | Checklist of the Vascular Plants of Tennessee
SBM 11 | Text Annotations & Identification Notes for Manual of the Vascular Flora of the Carolinas
SBM 12 | The “El Cielo" Biosphere Reserve, Tamaulipas, México
SBM 13 | Flora de Manantlan
SBM 14 | Niebla & Vermilacinia (Ramalinaceae) from California & Baja California
SBM 15 | Monograph of Northern Mexican Crataegus (Rosaceae, subfam. Maloideae)
SBM 16 | Shinners & Mahler's Illustrated Flora of North Central Texas
SBM 17 | The Grasses of Barbados (Poaceae)
SBM 18 | Floristics in the New Millenium: Proceedings of the Flora of the Southeast U.S. er
SBM 19 | Emanuel D. Rudolph's Studies in the History of North American Botany
SBM 20 | Generic Conspectus of the Tribe Asterae (Asteraceae) in North & Central America
SBM 21| A Comparative Checklist of the Plant Diversity of the Iwokrama Forest, Guyana
SBM 22 | Lloyd Herbet Shinners: By Himself
SBM 23 | Taxonomy, Distribution, & Ecology of the Genus Phaseolus (Leguminosae)
SBM 24 | Atlas of the Vascular Plants of Texas, Volume 1 & 2
SBM 25 | Los Géneros de Léguminosas del Norte de México
SBM 26 | Illustrated Flora of East Texas, Volume 1
SBM 27 | The Genus Psychotria (Rubiaceae) in the Philippine Archipelago
SBM 28 | Wild Flowers of Mombacho (Nicaragua)
SBM 29 | Muhlenbergia (Poaceae) de Chihuahua, México
SBM 30 | Tundra to Tropics: The Floristic Plant Geography of North America
OTHER PUBLICATIONS
Keys to the Vascular Plants of the Black Gap Wildlife Management Area, Brewster County
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Mosses of Texas: A Manual of the Moss Flora
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Eriocaulaceae of Continental North America, North of México
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Trees in the Life of the Maya World
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Violets (Viola) of Central & Eastern United States: An Introductory Survey
6
The Ecology of our Landscape: The Botany of Where we Live
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