Au AN ARRAINDSAD FLORIDA TOVWA ilar
“eonraBuTiOns i
oe AR INDIA .
Oi (Eu NSLA
Barney L. Lipscomb, Editor
Botanical Research Institute of Texas
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©SIDA, CONTRIBUTIONS TO BOTANY
VOLUME 20, NUMBER 1, PAGES 1-460
19 Juty 2002
COPYRIGHT 2002
BOTANICAL RESEARCH INSTITUTE OF TEXAS
PRINTED IN THE UNITED STATES OF AMERICA
ISSN 0036-148
TABLE OF CONTENTS
KEY TO THE SPECIES OF ERICACEAE OF BOLIVIA, INCLUDING TWO NEW SPECIES
JAMES L. LUTEYN—
POACEAE: LOLIINAE: SECT. SUBULATAE)
—
TWO NEW SPECIES OF FESTUCA FROM SOUTH AMERICA
DANIEL STANCIK AND Pau M. PeTeRSON—21
A NEW SPECIES OF ERIOGONUM (POLYGONACEAE) FROM SOUTH TEXAS
Guy L. Nesom—31
A NEW VARIETY OF IPOMOEA COSTELLATA (CONVOLVULACEAE) FROM THE EDWARDS PLATEAU REGION OF TEXAS
Roser J. O'KENNON AND Guy L. Nesom—39
ane soe CAESPITOSA VAR, WILLIAMSIAE, VAR, NOV. (MALVACEAE)
OEL H. HOLMGREN—47
THE PRESENT STATUS OF LEDERMANN’S ApriL RivER LOCALITIES IN PAPUA New GUINEA
W. TAKEUCHI AND M. GOLMAN—55
BIOSYSTEMATIC ANALYSIS OF THE THELESPERMA SUBNUDUM COMPLEX (ASTERACEAE)
Curtis J. HANSEN, LOREEN ALLPHIN, AND MicHact D. WINDHAM—71
LIATRIS GHOLSONII (ASTERACEAE: EUPATORIEAE), A NEW BLAZING STAR FROM THE APALACHICOLA RIVER BLUEES
AND RAVINES IN FLORIDA
LORAN C. ANDERSON—97
A NEW SPECIES OF GONOLOBUS (APOCYNACEAE: ASCLEPIADEAE, GONOLOBINAE) FROM SOUTHERN Costa RICA
ALEXANDER KRINGS—105
A NEW SPECIES OF MEDINILLA (MELASTOMATACEAE) FROM ANAMALAI HI LS, SOUTH INDIA
N. SASIDHARAN AND P. SWANAPAL—109
NEW TAXA OF CRATAEGUS (ROSACEAE) FROM THE NORTHERN OKANAGAN—SOUTHWESTERN SHUSWAP DIVERSITY CENTER
J.B. PHiPPS AND RJ. O'KENNON—
nee
NEW COMBINATION IN XYLORHIZA (ASTERACEAE: ASTEREAE
Guy L. Nesom—145
LECTOTYPIFICATION OF PSEUDOGNAPHALIUM BIOLETTH (GNAPHALIEAE: ASTERACEAE)
Guy L.N
BOERHAVIA COULTERI VAR. PALMERI, A NEW VARIETAL COMBINATION FOR BOERHAVIA (NYC TAGINACEAE)
OF SOUTHWESTERN NORTH AMERICA
RICHARD SPELLENBERG—151
LECTOTYPIFICATION OF MENTZELIA TEXANA AND M. LINDHEIMERI (LOASACEAE) WITH AN ASSESSMENT
OF THEIR BIOLOGICAL STATUS
BILLIE L. TURNER—157
a
NOMENCLATURAL CHANGES AND INNOVATIONS IN PANICUM AND DICHANTHELIUM (POACEAE: PANICEAE
OBERT W. FRECKMANN AND MICHEL G. LELONG—161
A NEW SPECIES OF SENECIO (ASTERACEAE) FROM DURANGO, MEXICO
B.L. TURNER—175
Two NEW SPECIES OF WEINMANNIA (CUNONIACEAE: CUNONIEAE) FROM SOUTHERN ECUADOR
ZACHARY S. ROGERS—179
THE TAXONOMY AND MORPHOLOGY OF MACRANTHERA FLAMMEA (OROBANCHACEAE)
Jit D. ALFORD AND LoRAN C. ANDERSON—189
THE GENUS CAMPOMANESIA (MYRTACEAE) IN ATLANTIC RAINFOREST FRAGMENTS IN SERGIPE, NORTHEAST REGION OF BRAZIL
Myrna F. LANDIM AND Lestle R. LANDRUM—205
BIOLOGICAL STATUS OF HEDYOTIS NIGRICANS VAR. GYPSOPHILA (RUBIACEAE) IN TEXAS
B.L. TURNER—215
DOCUMENTED CHROMOSOME NUMBERS 2002: 1. CHROMOSOME NUMBER OF STENANTHIUM (LILIALES: MELANTHIACEAE)
AND ITS SIGNIFICANCE IN THE TAXONOMY OF TRIBE MELANTHIEAE
Wenpoy B. ZOMLEFER AND GERALD L. SMITH—
DOCUMENTED CHROMOSOME NUMBERS 2002: 2. CHROMOSOME NUMBERS IN NORTH AMERICAN SPECIES OF CERASTIUM
(CARYOPHYLLACEAE
JK. MoRTON—-227
ra
ETHNOBOTANY/ SYSTEMATICS
Usos DE MELASTOMATACEAE EN EL ECUADOR
ALINA FREIRE FIERRO, DIANA FERNANDEZ, AND CATALINA QUINTANA—233
THE IDENTITY OF EAGLEWOOD (GYRINOPS, THYMELAEACEAE), A NEW ECONOMIC RESOURCE FOR PAPUA NEw GUINEA
W. TAKEUCHI AND M. GOLMAN—261
FLoristics/SYSTEMATICS
VASCULAR PLANTS OF KONZA PRAIRIE BIOLOGICAL STATION: AN ANNOTATED CHECKLIST OF SPECIES
IN A KANSAS TALLGRASS PRAIRIE
E. GENE TOWNE—269
THE VASCULAR FLORA OF THE CERRADO IN EMAS NATIONAL Park (GOIAS, CENTRAL BRAZIL)
Marco ANTONIO BATALHA AND FERNANDO ROBERTO MARTINS —295
RUST FUNGI (UREDINALES) OF NORTHWEST oe
José R. HERNANDEZ AND JOE F. HENNEN 13
ADDITIONS TO THE CYPERACEAE AND JUNCACEAE FLORA OF THE RIO MAYO REGION, SONORA, MexICO
Eric H. ROALSON, THOMAS R. VAN DEVENDER, AND ANA Lilla REINA G.—339
fe -ELIMINAR DE LAS aa Y AGAVACEAS DE LA SIERRA “EL VIEJO,” SONORA, REGION PRIORITARIA
PARA LA CONSERVACION EN MEX
CARLOS G. VELAZCO MACIAS AND vee J. ALANIS FLORES —349
VASCULAR FLORA OF BIG LAKE Bortom WILDLIFE MANAGEMENT T AREA, ANDERSON COUNTY, TEXAS
Kay M. FLEMING, JASON R. SINGHURST, AND Watter C. HoLmes—355
deci OF XERIC SANDYLANDS IN THE POST OAK SAVANNA REGION OF EasT TEXAS
AR. MAcRoserts, MICHAEL H. MACRoBERTS, AND JAMES C. CATHEY-—373
EFFECTS OF FIRE ON TWO PITCHER PLANT BOGS WITH IMENTS ON SEVERAL RARE AND INTERESTING PLANTS
Eric L. KEITH AND N. Ross Carric-—387
NATIVE FLOWERING PLANT SPECIES NEW OR OTHERWISE SIGNIFICANT IN KENTUCKY
Rosert F.C. NACZI, RONALD L. Jones, F. JOSEPH Metzeler, MARK A. GORTON, AND TIMOTHY J. WECKMAN—397
NEW RECORDS IN ASTERACEAE FOR ALABAMA AND ARKANSAS
Guy L. Nesom—403
SIX NON-NATIVE SPECIES NEWLY DISCOVERED IN THE IOWA VASCULAR FLORA
ALLISON W, Cusick —405
NOTEWORTHY VASCULAR PI ANTS FROM ARKANSAS. l|
ERIC SUNDELL, R. DALE THOMAS, CARL AMASON, AND Curls Dorritt—409
COMMELINA BENGHALENSIS (COMMELINACEAE) NEW TO NORTH CAROLINA AND AN UPDATED KEY TO CAROLINA CONGENERS
ALEXANDER KRINGS, MICHAEL G. BURTON, AND ALAN C. YorkK——419
AGROSTIS AVENACEA (POACEAE: POOIDEAE): FIRST RECORD FOR THE MEXICAN FLORA
Arey NAVA-Roso, MaricELA GOMEZ-SANCHEZ, AND Manuel GONZALEZ-LEDESMA—423
AN UPDATE TO THE VASCULAR FLORA OF CALCASIEU PARISH, LOUISIANA
Ray NEYLAND—431
NEW VASCULAR PLANT RECORDS FOR TEXAS
Guy L. NESoM AND RoserT J. O’KENNON—435
NOTES ON THE FLORA OF TEXAS WITH ADDITIONS AND OTHER SIGNIFICANT RECORDS. II
LARRY E. BROWN AND |. SANDRA ELSIK—437
ERRATUM—4.45
RUPERT BARNEBY AWARD—446
BOOK REVIEWS AND NOTICES 30, 46, 104, 114, 148, 156, 160, 178, 188, 232, 268, 312,348, 372, 396, 408,
430, 434, 447
INDEX TO NEW NAMES AND NEW COMBINATIONS IN SIDA 20(1), 2002.
BOERHAVIA COULTER! (HOOK. F.) S. WATS. VAR. PALMERI (S. WATS.) SPELLENB., COMB. NOV.—153
CRATAEGUS ATROVIRENS J.B.PHipeS & O'KENNON, SP. NOV.—141
CRATAEGUS CASTLEGARENSIS J.B PHippS & O'KENNON, SP.NOV.— 121
CRATAEGUS ENDERBYENSIS J.B. PHipPS & O'KENNON, SP. NOV.— 136
CRATAEGUS OKANAGANENSIS VAR. WELLSII J.B. PHIPPS & O'KENNON, VAR, NOV.— 132
CRATAEGUS ORBICULARIS J.B. PHIPPS & O’KENNON, SP. NOV.— 138
DiMuaee SHUSWAPENSIS J.B. PHipPS & O'KENNON, SP. NOV,— 128
CHANTHELIUM SE USTIFOLIA (HITCHC. IN B.L. Ros. & FERNALD) FRECKMANN & LELONG, STAT. NOV.— 166
DICHANTHELIUM SECT. See FRECKMANN & LELONG, SECT. NOv.— 164
DICHANTHELIUM SECT. ENSIFOLIA (HITCHC.IN B.L. Ros. & FERNALD) FRECKMANN & LELONG, STAT. NOV.— 166
DICHANTHELIUM SECT. LANCEARIA (HITCHC. IN B.L. Ros. & FERNALD) FRECKMANN & LELONG, STAT. NOV.— 166
DICHANTHELIUM SECT. LANUGINOSA (HITCHC. IN B.L.Rob. & FERNALD) FRECKMANN & LELONG, STAT. NOV.— 166
DICHANTHELIUM SECT. LINEARIFOLIA FRECKMANN & LELONG, SECT. NOV.— 164
DICHANTHELIUM SECT. MACROCARPA FRECKMANN & LELONG, SECT. NOV.— 165
DICHANTHELIUM SECT. NUDICAULIA HITCHCOCK & CHASE EX FRECKMANN & LELONG, SECT. NOV.—165
DICHANTHELIUM SECT. OLIGOSANTHA (HIiTCHC. IN B.L.Ros. & FERNALD) FRECKMANN & LELONG, STAT. NOV.— 166
DICHANTHELIUM SECT. PEDICELLATA HITCH. & CHASE EX FRECKMANN & LELONG, SECT. NOV.—165
DICHANTHELIUM SECT. SPHAEROCARPA (HITCHC. IN B.L.Rop. & FERNALD) FRECKMANN & LELONG, STAT. NOV.— 166
DICHANTHELIUM SECT. STRIGOSA FRECKMANN & LELONG, SECT. NOV,—
DICHANTHELIUM ACICULARE SUBSP, ANGUSTIFOLIUM (ELLIOTT) FRECKMANN & LELONG, COMB. & STAT. NOV.— 167
DICHANTHELIUM ACICULARE SUBSP. FUSIFORME (HITCHC.) FRECKMANN & LELONG, COMB. & STAT. NOV.—167
DICHANTHELIUM ACICULARE SUBSP. NEURANTHUM (GRISEB.) FRECKMANN & LELONG, COMB. & STAT. NOV.—167
DICHANTHEI 1UM ACUMINATUM
NUM (SCRIBN.) FRECKMANN & LELONG, COMB. & STAT. NOV.— 167
DICHANTHELIUM ACUMINATUM SUBSP. FASCICULATUM oe FRECKMANN & LELONG, COMB & STAT.. NOV.— 167
ICHANTHELIUM ACUMINATUM N.EX NASH) FRECKMANN & LELONG, COMB. & STAT. NOV.—167
DICHANTHELIUM ACUMINATUM SUBSP. LEUCOTHRIX ees FRECKMANN & LELONG, COMB. & STAT. NOV.—167
DICHANTHELIUM ACUMINATUM SUBSP. LINDHEIMERI ee FRECKMANN & LELONG, COMB. & STAT. NOV.—168
DICHANTHELIUM
SH) FRECKMANN & LELONG, COMB. & STAT. NOV.— 168
DICHANTHELIUM ACUMINATUM SUBSP. SERICEUM ene FRECKMANN & LELONG, COMB. & STAT. NOV.—168
DICHANTHELIUM ACUMINATUM SUBSP, SPRETUM (SCHULT.) FRECKMANN & LELONG, COMB. & STAT. NOV.— 168
DICHANTHELIUM ACUMINATUM SUBSP. THERMALE (BOL.) FRECKMANN & LELONG, COMB. & STAT. NOV.—168
DICHANTHELIUM CHAMAELONCHE ( TRIN.) FRECKMANN & LELONG, COMB. NOV.— 168
DICHANTHELIUM CHAMAELONCHE SUBSP. BREVE (HITCHC. & CHASE) FRECKMANN & LELONG, COMB. & STAT. NOV.— 168
DICHANTH SUBSP. ASHEI (G, PEARSON EX ASHE) FRECKMANN & LELONG, COMB. & STAT. NOV.— 169
DiaNTHELON COMMUTATUM SUBSP. EQUILATERALE (SCRIBN.) FRECKMANN & LELONG, COMB. & STAT. NOV.—169
DICHANTHELIUM COMMUTATUM SUBSP. JOORI (VASEY) FRECKIMANN & LELONG, COMB. & STAT. NOV.— 169
DIC HANTHEHIN DICHOTOMUM SUBSP. LUCIDUM (ASHE) FRECKMANN & LELONG, COMB. & $1 69
(ASHE) FRECKMANN & LELONG, COMB. & STAT. NOV.— 169
EX ELLIOTT) FRECKMANN & LELONG, COMB & STAT. NOV.—169
DICHANTHELIUM DICHOTOMUM SUBSP. NITIDUM Sit ae KMANN & LELONG, COMB. & STAT. Nov.—169
DICHANTHELIUM DICHOTOMUM 5 ASHE) FRECKMANN & LELONG, COMB. & STAT. NOV.—170
DICHANTHELIUM DICHOTOMUM SUBSP. YADKINENSE (ASHE) FRECKMANN & LELONG, COMB. & STAT. NOV.—170
ey ENSIFOLIUM SUBSP. CURTIFOLIUM eek FRECKMANN & LELONG, COMB. & STAT. NOV.—170
SH) FRECKMANN & LELONG, COMB. & STAT. NOV.— 170
DicaAicTHELION OVALE SUBSP. PRAECOCIUS (HITCHC. & Ci ae FRECKMANN & LELONG, COMB. & STAT. NOV.—170
ee,
nr
NA
DICHANTHELIUM OVALE SUBSP. PSEUDOPUBESCENS (NASH) FRECKIVANN & LELONG, COMB. & STAT. NOV—170
DICHANTHELIt UBSP. VILLOSISSIMUM (NASH) FRECKMANN & LELONG, COMB, & STAT. NOV.—17
DICHANTHELIUM PORTORICENSE SUBSP. PATULUM (ScRIBN. & err.) FRECKMANN & LEL LONG, COMB. NOv.—170
DICHANTHENIUM STRIGOSUM SUBSP. GLABRESCENS (GRISEB.) FRECKMANN & LELONG, COMB. & STAT. NoOV.—171
(TRIN.) FRECKMANN & LELONG, COMB. & STAT. NOv.—171
DICHANTHELIUM TENUE (MUHL.) FRECKIMANN & LELONG, COMB, NoV,—171
ERIOGONUM RIOGRANDIS NESOM, SP. NOV.—
FESTUCA CUZCOENSIS STANCIK & PM. PETERSON, SP. NOV.—22
FESTUCA TOVARENSIS STANCIK & P.M. PETERSON, SP. NOV. 24
GONOLOBUS TENUISEPALUS KRINGS, SP. NOV.—
IPOMOEA COSTELLATA TORREY VAR. EDWARDSENSIS O'KENNON & NESOM, VAR. NOV.—39
LIATRIS GHOLSONII L.C. ANDERSON, SP. N 8
MEDINILLA ANAMALAIANA SASIDHARAN : SUJANAPAL, SP. NOV—110
PANICUM SECT. ANTIDOTALIA FRECKMANN & LELONG, SECT. NOV.—163
PANICUM SECT. HEMITOMA HitcHc. & CHASE EX FRECKMANN & LELONG, SECT. NOV—163
PANICUM AMARUM SUBSP. AMARULUM (HITCHC. & CHASE) FRECKMANN & LELONG, COMB. & STAT. NOV.—171
PANICUM ANCEPS SUBSP. RHIZOMATUM (HITCHC. & CHASE) FRECKMANN & LELONG, COMB. & STAT. NOv.—-171
PANICUM CAPILLARE SUBSP. HILLMANII (CHASE) FRECKMANN & LELONG, COMB, & STAT. NOV.—17
PANICUM DICHOTOMIFLORUM SUBSP. BARTOWENSE (SCRIBN. & MERR.) FRECKMANN & LELONG, COMB. & STAT. NOV.—171
RITANORUM (SVENSON) FRECKMANN & LELONG, COMB. & STAT. NOV.—171
Snide HALLII SUBSP. FILIPES anc FRECKMANN & LELONG, COMB, & STAT. NOV.—172
PANICUM HIRTICAULE SUBSP. SONORUM (VASEY) FRECKMANN & LELONG, COMB. & STAT. NOV.—
PANICUM HIRTICAULE SUBSP. STRAMINEUM (HITCHC. & CHASE) FRECKMANN & LELONG, COMB. & STAT NOV.—172
FRECKMANN & LELONG, COMB. & STAT. NOV.—172
=
PANICUM PHILADELPHICUM SUBSP. GATTINGERI (NASH
PANICUM PHILADELPHICUM SUBSP. LITHOPHILUM (SWALLEN) FRECKMANN & LELONG, COMB, & STAT. Nov.—172
PANICUM RIGIDULUM SUBSP. ABSCISSUM (SWALLEN) FRECKMANN & LELONG, COMB, & STAT. NOV.—172
PANICUM RIGIDULUM SUBSP. COMBSII (SCRIBN. & C.R. BALL) FRECKMANN & LELONG, COMB, & STAT. NOV.—172
PANICUM RIGIDULUM SUBSP. ELONGATUM (PURSH) FRECKMANN & LELONG, COMB, & STAT. NOV—172
PANICUM RIGIDULUM SUBSP. PUBESCENS (VASEY) FReCKMANN & LELONG, COMB. & STAT. NOV.—173
PSEUDOGNAPHALIUM BIOLETTH A. ANDERS. [NOM. NovJ—149
SENECIO SANDERSIANA B.L. TURNER, SP NOV.—175
SIPHONANDRA BOLIVIANA LUTEYN, SP. NOV.—
SPHAERALCEA CAESPITOSA MLE. JONES VAR. WILLIAMSIAE N.H. HOLMGREN, VAR. NOV.—49
Aiea
THELESPERMA PUBESCENS DORN VAR. CAESPITOSUM (DORN) C.J. HANSEN, STAT. NOV.—91
THELESPERMA WINDHAMII C.J. HANSEN, NOM. ET STAT, NOV.—
THEMISTOCLESIA UNDUAVENSIS LUTEYN, SP. NOv.—15
WEINMANNIA BRADFORDIANA Z. oni) v.—180
WEINMANNIA CONDORENSIS Z. ROGERS, SP. NOvV.—183
XYLORHIZA LINEARIFOLIA (T.J. Wars.) NESomM, COMB. NOV.—145
¥
ae
KEY TO THE SPECIES OF ERICACEAE OF BOLIVIA,
INCLUDING TWO NEW SPECIES
James L.Luteyn
Institute OF VS EIGN evan)
The New IR DOLUTHCGT OGIGET)?
Bronx, New York 10458. 5126, U.S.A.
jluteyn@nybg.org
ABSTRACT
A review of the Ericaceae of Bolivia is herein presented. Nineteen genera and 53 species are native;
two genera and 19 species are endemic; 18 additional species are shared by Bolivia and adjacent Peru:
one species is occasionally cultivated. Most taxa are found in the wet, cool habitat, montane ‘ ‘Yungas”
of the eastern slopes of the Cordil me nee paneling pebween 2000 m and aoe m elevation.
land illustrated.
Siphonandra boliviana Luteyn and T
A key in English and Spanish to the 19 genera and 53 native species is provided. The names of all
taxa of Ericaceae attributed to Bolivia are accounted for.
RESUMEN
Se presenta una resena de las especies de Ericaceae de Bolivia. Diez y nueve géneros y 53 especies son
nativas, dos Baie y 19 es son endémicas, os cae adicionales son cormparudas por Bo-
]
La
livia y lay
en habitats montanosos, hitmedes y fies de las “Yungas” de la vertiente oriental de la Cordillera
Real, entre 2000 y 4000 m de elevacion. También se describen y se ilustran Siphonandra boliviana
Luteyn y Themistoclesia unduavensis Luteyn. Se presenta una clave en inglés y en espanol para los 19
géneros y las 53 especies nativas. Los nombres de todos los taxones de Ericaceae atribuidos a Bolivia
estan justificados.
INTRODUCTION
The Ericaceae are a large, cosmopolitan family of over 125 genera and 4500 Spee
cies, which inhabit the temperate regions of the world and montane areas in
tropical latitudes (Luteyn 1998, in press). In the Neotropics, the Ericaceae are
composed of 46 genera (70% endemic) and about 800 species (ca. 94% endemic),
although generic limits in the Andean Vaccinieae are still poorly understood
(Kron et al. 2002; Luteyn 1997, 2001). Neotropical Ericaceae arean ee -cen-
tered family, adapted to moist, open, cool montane environments. Overall spe-
cies richness in the Neotropics increases nearer the Equator, with the highest
species numbers concentrated in Colombia and Ecuador between 1000 m and
3000 m. Nineteen genera and 53 species are native to Bolivia. Two of those gen-
era and 18 of the species are endemic, while 18 additional species have ranges
that overlap between Bolivia and adjacent Peru and one species range overlaps
between Bolivia and extreme northwestern Argentina; one species is occasion-
ally cultivated. In Bolivia, the Ericaceae occur almost exclusively in the wet,
SIDA 20(1):1 - 20. 2002
2 BRIT.ORG/SIDA 20(1)
cool, montane cloud forests, mostly in the “Yungas” regions of the eastern slopes
of the Cordillera Real, ranging between 2000 m and 4000 m elevation. One of
these species (Satyria neglecta) ranges primarily below 1000 m elevation, 12
species range primarily between 1000 m and 2000 m, 18 species range prima-
rily between 2000 mand 3000 m, and 20 species range primarily between 3000
mand 4000 m. Agarista boliviensis and Gaylussacia cardenasii, in contrast,
occur in relatively dry montane regions in south-central Bolivia at about 1200-
2500 m elevation. Although Bolivia isa very large country within the Neotropics,
it lies at the southern end of the distributional range of the family and many of
its mountainous regions are dry. Therefore, relatively few Ericaceae occur in
Bolivia compared to countries in the Northern Andes and adjacent southern
Mesoamerica. There are no economic uses known for the Ericaceae of Bolivia.
There are very few general publications about the flora of Bolivia and only
the out-dated checklist of Foster (1958) gives an idea of the numbers of species
of Ericaceae that occur there. Killeen et al. (1993) is misleading for Ericaceae,
since virtually all the taxa in Bolivia are shrubs not trees. There is, however, a
project currently being organized by the Missouri Botanical Garden in collabo-
ration with all the major herbaria in Bolivia and the New York Botanical Gar-
den that has begun a modern checklist of the plants of Bolivia. With regards to
the Ericaceae of Bolivia, only the larger overall treatment of the tribe
Thibaudieae (=Vaccinieae) by A.C. Smith (1932) gives details of some of the
Bolivian blueberries. The most up-to-date list of Ericaceae of Bolivia, along with
some descriptions and photos, may be found on the website “Neotropical Blue-
berries” (Luteyn 1998). A list of all species of Ericaceae attributed to Bolivia,
including the current status of names given by Foster (1958), is herein presented
inan Appendix. The largest herbarium collection of Ericaceae in Bolivia is found
in La Paz, at the National Herbarium (LPB); there is also a moderate sampling
in Cochabamba (BOLV). The geographical regions of Bolivia in greatest need of
more collecting and with greatest possibilities for more Ericaceae records would
be the areas northeast of La Paz towards the frontier with Peru.
Neotropical Ericaceae, in general, have undergone dynamic speciation and
extensive adaptive radiation due to their ecological and life-form plasticity, colo-
nization abilities, adaptation to epiphytic habits, and co-evolution with hum-
mingbirds (Luteyn 2002). Given the high diversity and endemicity within
neotropical Ericaceae, along with high levels of habitat alteration, protection of
Andean montane ecosystems should become a priority for their conservation.
FAMILY DESCRIPTION AND KEYS TO THE BOLIVIAN SPECIES OF ERICACEAE
ERICACEAE
Terrestrial or epiphytic shrubs, subshrubs, sometimes lianoid, rarely trees, often
rhizomatous; indumentum when present of simple unicellular hairs, that are
usually translucent or whitish, these sometimes glandular tipped, or often (es-
LUTEYN, ERICACEAE OF BOLIVIA 3
pecially on leaves beneath) of multicellular, multiseriate, swollen-headed, usu-
ally glandular, hair-like structures called fimbriae which are brownish to red-
dish-brown. Leaves alternate, simple, usually petiolate, exstipulate but rarely
bud scales appearing pseudostipular; lamina coriaceous to membranous, ever-
green, the margin usually entire but sometimes serrulate-crenate, the venation
pinnate or plinerved; leaf scars usually witha single vascular bundle scar, nodes
usually with one trace and one gap. Inflorescences axillary, racemose, panicu-
late, fasciculate, or flowers solitary; individual flowers pedicellate or rarely
sessile in axils of small or large, deciduous or persistent floral bracts; pedicel
bibracteolate; bracteoles persistent, small or large. Flowers bisexual, actinomor-
phic or slightly zygomorphic, 5(-7)-merous, obdiplostemonous, hypogynous
or epigynous and with a biseriate perianth, typically without floral odors, rarely
with extrafloral nectaries, the superior-ovaried genera pollinated by bees and
the inferior-ovaried genera by hummingbirds; aestivation valvate, imbricate,
or reduplicate; calyx continuous or articulate with the pedicel, synsepalous, the
sepals rarely fleshy and accrescent to the fruit, the hypanthium when present
terete, angled, or winged; corolla membranous to thick-carnose, polypetalous
or more commonly sympetalous, cylindric, campanulate or urceolate, terete or
angled to winged opposite the lobes; stamens 8-10(-14), in 2 whorls, usually
twice as many as the petals or rarely just as many, equaling the corolla in over-
all length or 1/2-1/3 the corolla length, equal with each other or alternately
unequal, borne on the edge of an obscure to prominent nectariferous disc; fila-
ments equal or unequal, usually straight or rarely S-shaped (geniculate), ligu-
late but sometimes basally dilated, sometimes also basally papillose, distinct
or connate, with or without spurs, shorter or longer than the anther; anthers
inverting during development, 2-celled, equal or unequal, often distally with 2
distinct or connate tubules or terminal awns, sometimes provided with abaxial
spurs; disintegration tissue present or lacking; thecae smooth to coarsely granu-
lar, the base rounded to apendiculate; tubules when present conical and rigid
or cylindric and flexible, of equal or ca. 1/2 the diameter of the thecae, longer to
shorter than the thecae; dehiscence introrse by longitudinal or more typically
by apical to subapical clefts or pores, rarely perfectly terminal; pollen grains in
tetrahedral tetrads; carpels +-5(-10), fused; ovary superior or inferior, usually
with as many locules as carpels or with twice as many locules as carpels or
rarely loculate in lower portion and 1|-locular above; placentation axile, rarely
intruded parietal; ovules numerous per locule or rarely solitary, anatropous to
campylotropous with a single integumentary layer; style single, fluted, hollow;
stigma simple but occasionally weakly lobed. Fruit a loculicidal or septicidal
capsule, berry, or drupe, with a usually persistent, rarely accrescent and fleshy
calyx; seeds small, ca. 1-1.5 mm long, usually numerous (1 per locule in
Gaylussacia), winged or tailed only in Bejaria, sometimes enclosed in a muci-
laginous sheath, the testa thin with elongated or isodiametric cells, the en-
BRIT.ORG/SIDA 20(1)
dosperm fleshy, the embryo straight, usually white or sometimes green. Chro-
mosome numbers: x=6, 8, 11, 12(2), 13, 19, 23.
Ericaceae comprise ca. 125 genera and 4500 species worldwide, and are
cosmopolitan with the exception of Antarctica. Nineteen genera and 53 spe-
cies are native to Bolivia. Rhododendron simsii, native to China, is sometimes
cultivated in the montane areas throughout the Neotropics (including Bolivia),
but is not treated here.
ENGLISH KEYS
Notes on using the keys
The keys below are based primarily on herbarium specimens, although living
(in situ and greenhouse-grown plants), alcohol-preserved material, and photo-
graphs taken from field or greenhouse-grown material have been used. Floral
measurements are taken from herbarium material at anthesis unless otherwise
stated: colors are from fresh material observed by the author unless otherwise
stated; if a range of measurements is not available, the known measurement is
preceded by the abbreviation “ca.” (about); calyx limb length includes measure-
ment of the lobes, and anther length includes thecae and tubules.
KEY TO THE SPECIES OF BOLIVIAN ERICACEAE WITH SUPERIOR OVARIES
. Corolla with petals separate; fruit a septicidal capsule
Bejaria aestuans
. Corolla with petals fused; fruit a loculicidal capsule or berry.
2. Sta ens with fila ent niculate anther \/ ithout te rminal awns, and witho
one disintegration i tissue on ileal side Agarista sien
2. Stamens with fi i with terminal awns, and with white dis-
integration tissue on abaxial side.
3. Fruit a berry, rarely calyx becoming fleshy at base but never surrounding the
berr Pernettya prostrata
3. Fruit a capsule, surrounded by the fleshy, accrescent calyx (calyx rarely not
fleshy).
4. lee solitary in axils of normal (or only slightly reduced) ieee
_ Thick-stemmed shrub to 5 m tall; leas broade
yellowish-gre
ultheria ania var. secunda
5. Thin- sernmed aie to 0.4 (rarely 1-2) m area urceolate to
near| It glob se, inflated at the base and ery constricted at the throat,
ae ce rose-red
4. Flowers in axillary racemes.
aultheria vacinioides
6. Abaxial surface of lamina and entire inflorescence tomentose-lanate
Gaultheria eriophylla var. mucronata
6. Abaxial surface of lamina and inflorescences glabrous or sparsely to
densely pubescent but never tomentose-lanate.
7. Young twigs and inflorescences picuously strigose with straight,
rigid, appressed hairs, usually so dense as to obscure me lamina
usually densely and persistently reddish-strigose bene
8. Ca lyx geRIONs corolla glabrous to yey hairy, never densely
Io}
g ; Qlabrous OF ery weakly short-pi
LUTEYN, ERICACEAE OF BOLIVIA 5
inflorescen ngested (glomerate) at anthesis with the flowers
broadly overlapping Gaultheria glomerata
8. Calyx and corolla densely strigose-hirsute with ferruginous
subsetose, rarely glandular hairs; ovary densely short-white pilose
or cinerous; inflorescence not congested at anthesis, the flowers
idely spaced Gaultheria bracteata
7. Young twigs and inflorescences glabrous or variously spreading pu-
bescent, but not appressed-strigose with straight, rigid hairs; lamina
glabrous or variously pubescent, but 7 strigose beneath.
9. Repent, rhizomatous subshrub, 0.1—-0.2 m tall; leaf pe usuall
bovate with the apex rounded aultheria hapalotricha
9. Erect, subshrubs to shrubs, 0.1-8 m tall; leaf ari cee but
only rarely obovate, the apex usually acute to acuminate
10. Leaf lamina usually distinctly and prominently reticulate-
veined on both surfaces, the base usually acute to rounded;
inflorescences clustered at eas ups and seen ex-
SS
ceeding the leaves in length; cal andular
rolla white, pilose te Gaultheria reticulata
oO
. Leaf lamina not conspicuously reticulate-veined on both sur-
faces, the base usually rounded and deeply cordate; inflores-
>
5
corolla red, glabrous within
Gaultheria erecta
KEY TO THE SPECIES OF BOLIVIAN ERICACEAE WITH INFERIOR OVARIES
. Stamens strongly unequal with filaments or anthers, or filaments and anthers al-
alla Aa COr clea rea unequal.
2. Fil nnate over entire length; anthers with tubules widening
dist ie
3. Leaves elliptic-lanceolate, 6-10 cm long, 1.5-3 cm broad, basally obtuse to
unded, 3-plinerved; inflorescence densely pilose (except corolla); aoe
ie 21mm long; corolla ca.11 mm long Satyria boliviana
3. Leaves valine oblong, 3-6 cm long, 1.2-1.8 cm broad, basally cuneate to
subattenuate, pinnately veined; inflorescence glabrous; pedicels 6-12 mm
long; corolla ca.6 mm lon Satyria neglecta
2. Filaments unequal, distinct or partially connate; anthers with sides parallel, not
widening distally.
4, Anthers equal; stamens 1/2-1/3 as long as corolla; floral bracts rarely large
and showy but if so then early deciduous.
5. Leaves ae veined Orthaea pinnatinervia
5. Leaves plin
6. Stems, ine leaves, rachis, pedicels, and corolla pilose Orthaea rusbyi
6. Plants essentially glabrous.
7. Corolla 11-12 mm long Orthaea constans
7. Corolla 15-33 mm lon
8. Staminal filaments distinct.
9. Rachis 3-5 cm long; pedicels 8-13 mm long___ Orthaea weberbaueri
9. Rachis ca. 1 cm long; pedicels ca. 7 mm long Orthaea ignea
8. Staminal filaments connate
BRIT.ORG/SIDA 20(1)
10. Staminal dehiscence by terminal or subterminal pores Orthaea
boliviensis
10. ee a ee ae lateral clefts Orthaea ferreyrae
Anti as corolla or rarely 1/2-2/3 the corolla oe
fora bracts usually aces and rae usually persistent through anthes
. Inflorescence a pan
_ Inflorescence a race
12. Calyx densely coll the matted hairs persistent; leaves persistently
h endishia pubescens
| ane martii
soft-pilose beneat Cav
12. Calyx glabrous to pilose, but never woolly and the hairs never mat-
ted; leaves glabrous to glabrate Cavendishia bracteata
1. Stamens equal with filaments and anthers of equal lengths (rarely anthers incon
spicuously alternately unequal)
eoles located at apex of pedicel and surrounding calyx (and sometimes
lower corolla).
14. Leaves less than 5 mm broad.
15. Corolla campanulate-cylindric, 10-14 mm long, white sometimes tinted
Disterigma pernettyoides
___ Disterig
ma
with pin
15. ene subcylindric to somewhat urceolate, 6-9 mm long, re
empetrifolium
14. Leaves more than 5 mm broa
16. Corolla 10-12 mm long, naouly cylindric, red
Disterigma pallidum
16. Corolla 4.5-7.5 mm long, subcylindric to ovate-urceolate, white to red.
17. Flowers 4 lla subcylindric, rarely puberulous within;
staminal aament distinct Distelgme alaternoides
tT) A te-ljr
AI
17. Flowers usually 5
ceolate, conspicuously pilose within; staminal filaments panes
terigma ovatum
13. Bracteoles located well below apex of pedicel, or if apical then not ee ng
ie Tubules elongate, thin, very graceful, about half as wide (or less) as thecae;
dehiscence pores perfectly terminal.
19. Filaments distinct; rachis to 0.5 cm long; calyx ca. 9 mm long; corolla
35-37 mm long Siphonandra magnifica
19. Filaments connate;rachis 2-5 cm long;calyx 7-8 mm long; corolla 25—
48 mm long.
20. Corolla ca.25 mm long
20. Corolla 43-48 mm long
18. Tubules about as wide as thecae, or if narrower then proportionally muc
se than thecae; ne by lateral slits or elongate cle
. Thecae conspicuously | rigid, elongate- to short-coni-
- stamens often 1/3-1/2 as long as corolla.
es Stiff coriaceous; pedicels 20-40(—55) mm long; calyx lobes
aa ae in number, often in a state of fusion, 1— A mm long;
corolla 18-35 mm long;stamens 9-15 mm long _____
Leaves chartaceous to soft coriaeous; deel 10- 14 mm ine
calyx lobes lacking or 5, apiculate and less than 0.4 mm bas
rolla 25-40 mm long; stamens 8-10 mm lon ammisia pauciflora
21. Thecae smooth to minutely papillate; tubules foabe ae elon-
gate to short; stamens usually as long as corolla.
iphonandra elliptica
Siphonandra boliviana
N
MN
LUTEYN, ERICACEAE OF BOLIVIA 7
23. a articulate with pedicel.
24. Corolla elongate- niDulen -)11-27 mm long lly carnose
eee a lobes te-fil and inconspicu-
ous with regard to anthers
25. Corolla 20-27 mm long; staminal filaments distinct.
26. Leaves truncate or subcuneate at base; calyx smooth,
not papilose at base Thibaudia macrocalyx
26. Leaves attenuate at base; calyx strongly papillose at
base Thibaudia axillaris
25. Corolla 7-13 mm long; staminal filaments connate.
27. Leaves obtuse, rounded to subcordate at base; twigs,
leaves, and inflorescences (including rachis, pedicels,
calyx and sometimes corolla) densely white pilose
Thibaudia densiflora
27. Leaves cuneate, attenuate or subattenuate at base;
plant surfaces glabrous or ete
28. Leaf sai acute; calyx 5.5-6 mm oa the lobes
deltate, ca. 1 mm long Thibaudia regularis
28. es pen obtuse; calyx 2.5-3.5 mm long, the lo
piculate, less than 1 mm long Thibaudia crenulata
24. Corolla Pees to campanulate, 7-12 mm long, normally of
thin texture and membranaceous, the lobes imbricate:; fila-
ments proportionally long with regards to anther length.
29. Ovary with a ee ovule in each of the 10 locules; fruit a
drupe with 10 pyren Gaylussacia cardenasii
29. Ovary with few to numerous ovules in each of the 5(-
locules; fruit a many-seeded berry.
30. Leaves glabrous or lacking subfasciculate hairs be-
neath; corolla cylindric-urceolate, white to pinkish
Vaccinium amie
30. Leaves with subfasciculate hairs beneath; corolla
te-campanulate, green aaiue cdependene
23. Calyx continuous with pedicel, the pedicel not jointed at apex
31. Calyx conspicuously angled to 5-winged; corolla terete or
angled
32. Calyx angled opposite the lobes Polyclita turbinata
32. Calyx angled to winged alternate with the lobes.
33. Leaves linear, ca. 2 mm broad, 1-nerved Rusbya taxifolia
33. Leaves mostly ovate,greater than 1 cm broad, multi-nerved.
34. Plants with pilose-hispid habit (including twigs,
petioles, leaves beneath, racemes, pedicels, and
calyx); corolla cylindric, terete throughout, 9-10
mm long, sparsely pilose distally Themistoclesia
peruviana
7
Ww
d
. Plants with glabrous habit (although leaves be-
neath with brownish fimbriae); corolla urceolate-
turbinate, conspicuously swollen at base and nar-
rowed at throat, bluntly 5-angled, 8-11 mm long,
Themistoclesia unduavensis
\
»
glabrous
31. Calyx terete; corolla terete.
8 BRIT.ORG/SIDA 20(1)
35. Corollas usually large, carnose to coriaceous, (0.6-)1.5—5
e long, if less than 1 cm then staminal tubules
twice as long as thecae;staminal tubules 2-5 times longer
than thecae; seeds with white embryos
36. Corolla 0.6-0.8 cm long, white to pinkish _____-s Demosthenesia
cei
36. Corolla 1.7-5 cm long, red.
Corolla 2-3 cm long and 3-8 mm diam., slightly
zygomorphic Demosthenesia mandonii
37. Corolla 3.5-5 cm long and 7-10 mm diam., acti-
nomorphic mosthenesia spectabilis
Ww
al
Corollas small, thin-membranaceous, up to ae mm long,
but if longer then filaments proportionally much longer
than anthers; staminal tubules about equalling anthers;
seeds with green embryos.
38. Flowers usually in few- to many-flowered fascicles or
racemes, rarely solitary; pedicels slender but not prop-
a cernuous; filaments usually shorter than anthers.
_ Inflorescence a fascicle of 3-6 flowers, the rachis
one Diogenesia boliviana
. Inflorescence a raceme of up to 18 flowers, the
rachis 2.5-6 cm long Diogenesia racemosa
38. Flowers 1-2 per axil; ian usually relatively long and
thin, filiform, cernuou arely absent flowers
sessile); filaments pe nae than anther
40. Flowers sessile Sphyrospermum sessiliflorum
40. Flowers conspicuously pedicellate.
4]. Leaves suborbicular to oblong-ovate, (0.7-)
0.9-1.5(-1.8) cm long, the apex rounded or ob-
tuse; flowers usually extending well beyond
the leaves; corolla 4-6 mm long; stamens 4
Sphyrospermum buxifolium
. Leaves ovate to ovate-lanceolate, usually con-
spicuously longer than broad, (1-)2-3.5
5) cm long, the apex obtuse to acute, some-
times shortly and bluntly acuminate; flowers
rarely extending beyond the leaves; corolla
(4-)5-7(-9) mm long; stamens 4-5 or 8-10
Sphyrospermum cordifolium
By
—
SPANISH KEYS
Notas sobre la utilizacion de las claves
Las siguientes claves se basan principalmente en ejemplares de herbario, aunque
también en material vivo (in situ y de plantas de invernadero), material
preservado en alcohol y fotografias tomadas en el campo y en invernaderos.
Las medidas florales se tomaron del material de herbario en la antesis a menos
que se diga lo contrario; los colores se tomaron del material vivo observados
por el autor a menos que se diga lo contrario; si no se tiene un intervalo de
LUTEYN, ERICACEAE OF BOLIVIA
medidas, entonces las medidas conocidas se presentan precedidas por la
abreviacion “ca.” (cerca); la longitud del limbo del caliz incluye la medida de los
lébulos y la longitud de la antera incluye las tecas y los tubulos.
CLAVE PARA LAS ESPECIES BOLIVIANAS DE ERICACEAE
CON OVARIO SUPERO
. Corola con pétalos separados; fruto una capsula septicida
ee con pétalos unidos; fruto una capsules loculicida ouna baya.
. Estambres con filamento les y sin tejido
blanco desintegrado en el lado abaxial Agarista es
2. Estomac con lime Hes rectos; anteras con aristas terminales y con teji
Bejaria aestuans
blar el lado abaxial.
3, ruts una oo raras veces el caliz se vuelve carnoso en la base pero nunca
rodea lab Pernettya prostrata
Fruto un ul
1a Cd ,rodeada por el caliz carnoso y acrescente (caliz raras veces
no eer
| ri | | H | 1 +
idas)
5 Arbusto de tallo grueso hasta Sr m de altura; corola campanula
ncha en el apice, verde-amarillenta eria ae var.secunda
5: ae de tallo delgado hasta 0,4 (raras veces 1-2) m de altura;
nte subglobosa, inflada en la base y muy
contraida en la cre rosada a rosado-roja
4, bas en racimos axilar
corola urceolada a estrecham
Gaultheria vaccinioides
pe alias abaxial de la lamina y toda la inflorescencia lana
me Gaultheria siaphvlla var.mucronata
6. ea en de la lamina y la inflorescencia glabras 0 esparcida a
densamente sane pero nunca lanado-tomentosas.
mas jovenes e inflorescencias Boh spice tamicnte estrigosas con
eee rigidos, adpresos tan densos que ocultan
las superficies; lamina en general densa y persistentemente rojizo-
eae en el envées.
ri glabro; corola glabra a beanies ee nunca
nsamente estrigosa en to rt glabr il
'
( |
og
yc Uae plies en vel spice, fafoescencia congestionada
i* |
Gaultheria glomerata
hirer n |
i Jo
4 |
. Calizy
subsetosos landul
|
tricomas cortos, blancos. ocinéreos;inflorescencia no congestionads
en la antesis, las flo
espaciadas ome bracteata
7, Ramas jovenes e i noe fo) ce nce
peuroads espaciad
Pall
I estrigosa con pelos rectos,
g ariadamente pubescente pero no estrigosa
en fale
9. piesa: rizomatosos rastreros, 0,1—-0,2 m de altura; lamina de
la hoja generalmente obovada con el apice redondeado |
eria
hapalotricha
9. Subarbustos a arbustos erectos, 0,1-8 m de altura; lamina de la
hoja de forma variada pero raras veces obovada, el Apice
generalmente agudo a acuminado.
BRIT.ORG/SIDA 20(1)
10. Lamina de la hoja en general marcada y prominentemente
reticulado-nervada en ambas caras, la base generalmente
aguda a redondeada; inflorescencias ieee en la punta
de las ramas y CONS spicuam nt cediendo la ongitu tud de las
hojas; caliz y corola ae corola plat nca, pie por
ntro aultheria reticulata
. Lamina de la hoja no conspicuamente Pnrene en
ambas superficies, la base generalmente redondeada y
profundamente cordada;inflorescencias eriaues alo largo
de las ramas, sin exceder la longitud de las hojas;caliz y corola
a menudo pubescente-glandulosos; corola roja, glabra por
ntro
oO
Gaultheria erecta
CLAVE PARA LAS ESPECIES BOLIVIANAS DE ERICACEAE CON OVARIO INFERO
1.
Ectambres desi | fil tosoant fi| alternamente
hs = t v4
desiguales en forma conspicua.
2. Filamentos igquales y connados en toda su longitud; anteras con tubulos
ensanchandose distalmente
3. Hojas eliptico-lanceoladas, 6-10 cm de largo, 1,5—3 cm de ancho, obtusas a
redondeadas basalmente, 3-plinervias; inflorescencia densamente pilosa
oe en la corola); pedicelos 16-21 mm de largo; corola ca. 11 mm de
larg Satyria boliviana
3: ee eliptico-oblongas, 3-6 cm de largo, 1,2-1,8 cm de ancho, cuneadas a
subatenuadas basalmente, pinnatinervias; inflorescencia iuaemaee 6-
12 mm de largo; corola ca.6 mm de lar Satyria neglecta
2. Filamentos desiquales, libres o ine connados; anteras con los lados
paralelos, sin ensancharse distalmente.
4. Anteras iguales; estambres 1/2-1/3 tan largos como la corola; bracteas florales
pero si asi,entonces deciduas tempranamente.
Orthaea pinnatinervia
raras veces grandes y
5. Hojas pinnatinervias
5. Hojas plinervadas.
6. Tallos, peciolos, hojas, raquis, pedicelos y corola pilosos Orthaea rusbyi
6. Plantas esencialmente glabras.
7. Corola 11-12 mm de largo Orthaea constans
7, Corola 15-33 mm de largo
8. Filamentos estaminales libres.
9. Raquis 3-5 cm de largo; pedicelos 8-13 mm de largo Orthaea
weberbaueri
9. Raquis ca. 1 cm de largo; pedicelos ca.7 mm de largo ___ Orthaeaignea
8. Filamentos estaminales conna
10. Dehiscencia estaminal por poros terminales o subterminales __ Orthaea
boliviensis
. Dehiscencia estaminal au suturas laterales Orthaea ferreyrae
4. Anteras deAceles estambres tan largos como la corola o raras veces 1/2—2
3 de la longitud de ésta; bracteas ae generalmente grandes y vistosas,
Sau eeed persistentes durante la antesis.
. Inflorescencia en panicula
ep eraiias en racimo.
aliz densamente lanoso, los |
oo. suaves persistentes por el envés Cavendishia pubescens
Cavendishia martii
| dad | ho} con
LUTEYN, ERICACEAE OF BOLIVIA
12. Caliz glabro a piloso p nca lanoso y los pelos nunca enredados
noes gables a cldalescenies. nis endishia bracteata
1. Estaml de ig ngitud (raras an
alternamente desiguales en ee inconspicua).
13. ia localizadas en el apice del pedicelo y rodeando el caliz (y algunas
eces la parte baja de la corola).
i Hojas menos de 5 mm de ancho.
15. Corola Hise 10-14 mm de largo, ple algunas
veces tenida de ro erigma aes anes
15. Corola ee a algo urceolada, 6-9 mm de =e roja Disterigma
See hiaian
14. Hojas mas de 5 mm de ancho.
16. Corola 10-12 mm de largo, estrechamente cilindrica, roja Disterigma
pallidum
16. Corola 4,5-7,5 mm de largo, subcilindrica a ovado-urceolada, blanca a
fe Flores meras;corola be ease raras veces pubérula por dentro;
filamentos estaminales libre Disterigma alaternoides
17. Flores generalmente 5-meras (algunas veces 4-meras): corola
ovado-urceolada, conspicuamente pilosa por dentro; filamentos
estaminales cohesionados en la base Disterigma ovatum
13. Bractéolas localizadas muy por debajo del apice del pedicelo o si apica
5 sin abrazar el céliz.
18. Tubulos alargados, delgados, casi tan ancho como la mitad (o menos) de
la teca; dehiscencia por poros perfectamente terminales.
19. Filamentos libres; raquis os 0,5 cm de largo; caliz ca.9 mm de largo;
corola 35-37 mm de lar Siphonandra magnifica
19. Filamentos connados; oe 2-5 cm de largo; caliz 7-8 mm de largo;
corola 25-48 mm de largo
20. Corola ca.25 mm 4e largo
Siphonandra elliptica
20. Corola 43-48 mm de largo Siphonandra boliviana
18. Tubulos casi tan anchos como la teca o si mas rene entonces
roporcional mucho mas cortos que la teca; dehiscencia por suturas
laterales o hendiduras elsigaeas.
21. Teca conspic
conicos; estambres a menudo 1/3-1/2 tan largos como la corola,
22. Hojas rigidamente coridceas; pedicelos 20-40(-55) mm de largo;
lobulos del caliz prominentes, 3-5,a menudo algo fusionados, 1-3
e largo; corola 18-35 mm de largo; estambres 9-15 mm de
sammisia guianensis
m de
1 | id alaragados a cortament
largo Ps
. Hojas cartaceas a suavamente coridceas; pedicelo 10-14 mm
largo; |6bulos del caliz ausentes 6 5, apiculados y menos de 0,4
e largo; corola 25-40 mm de largo; estambres 8-10 mm de
lar Psammisia pauciflora
21. Teca is a SIMI ane te Rapllosar tubulos flexibles, cilindricos
alargados nte tan largos como la corel
23. Cal articulado con el pedicis
rola alargada, tubular, (7-)11-27 mm de largo,normalmente
carnosa O coriacea, los [6bulos valvados; filamentos cortos e
inconspicuos con respecto a las anteras
N
i)
12 BRIT.ORG/SIDA 20(1)
25. Corola 20-27 mm de largo; filamentos estaminales libres.
26. Hojas truncadas o subcuneadas en la base; caliz liso
no papiloso en la base __ Thiba udia macrocalyx
26. Hojas atenuadas en la base; caliz fuertemente papilos¢
en la base Tnibavels axillaris
ol Fa | aye : |
Corola 7-13
27. Hojas obtusas sae eeaes a subcordadas en la base;
NO
cal
ramas, aces e ee epocliatae ae hae bau:
pedicelos, cali algu 1aS 3
blanco-pilosas Thibauidia densiflora
27. Hojas cuneadas, atenuadas o subatenuadas en la base;
superficies de la planta glabras 0 esencialmente glabras.
28. Apice de la hoja agudo; caliz 5,5-6 mm de largo,
los l6bulos deltados, ca. 1 mm de largo Thibaudia
regularis
28. Apice de la hoja obtuso;caliz 2,5-3,5 mm de largo,
los I6bulos apiculados, menos de 1 mm de largo
hibaudia crenulata
24, aol ole dices a campanulada, 7-12 mm de larg
normalm t leigada'/ meimiorenacee ie
b 105; fil ionalmente larg specto
ala longitud de las oe
29, Ovario con 10 ldculos, un solo évulo por léculo; fruto una
drupa con 10 pirenos Gaylussacia cardenasii
29, Ovario con 5(-10) ldculos, pocos a numerosos Ovulos en
cada ldéculo;fruto una baya con muchas semillas.
30. Hojas glabras 0 sin pelos subfasciculados en el enves;
corola cilindrico-urceolada, blanca a rosada Vaccinium
floribundum
30. Hojas con pelos subfasciculados en el envés; corola
rotado-campanulada, verde Vaccinium dependens
23. Caliz continuo con el pedicelo, no articulado.
31. Caliz conspicuamente angulado hasta 5-alado;corola terete o
angulado.
32. Cadliz anqulado opuesto a los lobulos Polyclita turbinata
32. Caliz angulado a alado alterno con los lobulos.
33. Hojas lineares, ca. 2 mm de ancho, 1-nervias Rusbya taxifolia
33. Hojas principalmente ovadas, mas de 1 cm de ancho,
multinervias.
34, Plantas con habito hispido-piloso (incluyendo
ramas, pec envés de las hojas, racimos,
pedicelos y caliz);cor Sater riers eet
terete, 9-10n mm de largo, espaciadamente pilosa
en el parte distal Se peruviana
34. Plantas con habito glabr elen e las
hojas con fimbrias de color eee sous
urceolado- De a coh wasceunel hinchada
la base \ nla garganta,ampliamente
5- angulada, 8- i mm ae largo, glabra Themistoclesia
unduavensis
5 ee
LUTEYN, ERICACEAE OF BOLIVIA
13
31. Caliz terete; corola terete.
5. Corola agen jalnicnte belie carnosa a coriacea, (0,6-)1,5
. cmomasd simenos de 1 cm entonces los tubulos
estaminales dos veces tan tae como las tecas; tubulos
estaminales 2-5 veces mas grandes que las tecas; semillas
con embriones blancos.
36. Corola 0,6-0,8 cm de largo, blanca arosada ___ Demosthenesia
pearcei
1s)
5.
36. Corola 1,7—5 cm de largo, roja.
37. Corola 2-3 cm de largo y 3-8 mm en diam.,
ligeramente zigomorfa Demosthenesia mandonii
37; sees 3,5-5 cm de largo y 7-10 mm en didm.,
orfa Demosthenesia spectabilis
Corola pequena, ee membranacea, hasta 10 m
oO si es mas larga entonces los ee
proporcionalmente mucho mas largos que las anteras;
tbs + 7 In i | { | + ill
embriones verdes.
38. Flores generalmente en fasciculos 0 racimos de pocas
a muchas flores, raras veces solitarias; pedicelos
delgados pero no propiamente péndulos; filamentos
generalmente més cortos que las anteras
39. Inflorescencia en fasciculos de 3-6 flores, sin raquis
Diogenesia boliviana
39. Inflorescencia en racimos de hasta 18 flores, el
Aad 2,5- ot cm ae argo eedenesta racemosa
. Flores 1 2 por |
largos y | delgados, filiformes, péndulos o raras veces
Ww
ioe)
largos que las anteras.
40. Flores sésiles oe sessiliflorum
40. ores conspicuamente pediceladas.
1. Hojas suborbiculares a ae ovadas, (0,7-)
0,9-1,5(-1,8) cm de largo, el apice redondeado
Ar fl | + + 1ek J
mucho més alla de las hojas; corola 4-6 mm
delargo; estambres 4 —_ opoyrespernum buxifolium
Hojas ovadas a ovad
a
o
=.
\ceoladas, en general
conspicuamente mas ae que anchas, (1- )
2-3,5(-5) cm de largo, el dpice obtuso a agudo,
algunas veces corta y llanamente acuminado;
flores raras veces extendiéndose mas alla de
las hojas; corola (4-)5-7(-9) mm de lar
estambres 4-5 u 8-10 phy ieo ania cordifolium
NEW SPECIES
Siphonandra boliviana |uteyn, sp. nov. (Fig. 1). Type: BOLIVIA. DEPTO. LA PAz. Prov.
Bautista Saavedra: Charazani, W of Chullina, 3400 m, 1 Aug 1994 (£D), B. Herzog
H200 (HOLOTYPE: NY; ISOTYPES: LZ, s.n.).
14 BRIT.ORG/SIDA 20(1)
ica staminum filamentis connatis nec distinctis, rhachidibus 2-5 nec 0.5 cm longis, et ab
—
AS. magni
S.elliptica corollis 45-47 nec ca. 25 mm longis differt.
Shrub (size unknown); mature branches terete, glabrous, the bark exfoliating
in thin strips; twigs subterete, sometimes shallowly angled or ribbed, short-
pilose with white hairs; buds axillary, the scales 2, valvate, ca. 2mm long, short-
pilose. Leaves alternate, coriaceous, elliptic to oblanceolate, 2.5-5.5 cm long, 1-18
cm broad, basally cuneate, apically broadly acute to nearly obtuse, marginally
entire, essentially glabrous above or sparsely short-pilose proximally along
midrib, sparsely pilose beneath especially along midrib, also provided with
reddish-brown, basally swollen, glandular fimbriae beneath; pinnately nerved
—
with 4-6 secondary veins anastomosing near margin, the midrib and second-
ary veins impressed above and raised beneath, the reticulate veins plane to
slightly impressed above but inconspicuously raised beneath; petiole rugose,
subterete, broadly flattened above, 4-5 mm long, Inflorescence axillary, race-
mose, ca. 20-flowered, apparently nodding; rachis subterete, striate to angled,
densely short-pilose with white hairs, at least 5cm long (still in bud, apparantly
still elongating); floral bracts ovate, acuminate, 3-4 mm long, densely short-
pilose; pedicels subterete, striate to angled, densely short-pilose as rachis, l1-13
mim long; bracteoles located in proximal 1/3 pedicel, similar to floral bracts, 2-
3 mm long, Flowers: calyx articulate with pedicel, 7-8 mm long, densely short-
pilose as rachis; hypanthium cylindric, ribbed, 4-5 mm long, rounded at base;
limb spreading-campanulate, 3.8-4.5 mm long; lobes 5, deltate, acute, 15-2 mm
ong; sinuses obtuse; corolla of 5 fused petals, long-cylindric, 43-48 mm long,
6-7 mm diam., short-pilose throughout, with white hairs, the lobes 5, deltate,
acute, ca. 2mm long: stamens 10, equal, ca. 32 mm long; filaments connate, gla-
brous, ca. 7 mm long; anthers ca. 28 mm long; thecae granular, ca. 6 mm long,
incurved at base: tubules thin, delicate, about half as wide as thecae, ca. 22 mm
long, dehiscing by perfectly terminal, flaring pores; ovary S-loc ular; style about
equaling corolla. Berry not seen.
Distribution —Endemic to Bolivia and known only from the type collection,
which was made ina Weinmannia forest.
Etymology—The species is named for the country Bolivia to which itisendemic.
Siphonandra is a small, distinctive, high-elevation genus of three species:
S. elliptica which is common and ranges from south-central Peru to northern
Bolivia, S. magnifica which is endemic to Bolivia and is herewith maintained
despite the fact that the type and only specimen was destroyed during World
War II, and the new species herein described 5. boliviana, for which only the
type collection is known. With only one extant collection of the latter two species,
it is difficult to assess the relationships between any of the three species in this
genus. Nevertheless, S. boliviana is easily distinguished morphologically from
the other two species by the characters mentioned in the key and diagnosis.
—
LUTEYN, ERICACEAE OF BOLIVIA
sett gate
EAN. een eae
Fic. 1. Sip dra boliviana | A. Habit. B. Portion of stem showing put ¢ sealed floral bract,
pedicel acauiee calyx and daenile D.L
show-
HT £+ : ldah: fan + llart?
ing side, fro
meaner unduavensis | uteyn, sp. nov. (Fig. 2). aie BOLIVIA. DeEpto. LA PAZ.
v. Nor Yungas: trail to Rio Coscapa, ca. 2.5 km E of Unduavi, along new rd to
ae and ca. 6 km W of Cotapata, ca. 16° 17'S, a 53'W, 3200-3350 m, 19 Mar
2000 (ED, J.-L. Luteyn, E. Ann Powell & S. Beck 15471 (HOLOTYPE: NY: ISOTYPES: AAU,
CAS, hK, UPB, MO, TEX, — 6 others to be distributed by LPB).
] ];
A a : 1 1 Tal | ]
hispido, COFOl
et glace nec pilosa Acts aliter ater
Rhizomatous, terrestrial or epiphytic subshrubs, to 30 cm tall: mature stems
terete, striate, glabrous, the bark cracking longitudinally but not exfoliating;
16 BRIT.ORG/SIDA 20(1)
fic. 2. Th . ee ) spy A Habj ‘san 1 £] £ 4 £ d il hid scales. B. Portion
a eee C Infl howing twig, floral bracts pedicel, bracteoles, calyx, and corolla. D
; Py ra " L . vos
J
1 | +1 f |
bo |
c L
of stamens. G
g back fit and side views. (A fan Solomon 18195; B-G from type collection).
twigs subterete, flattened to broadly and obtusely ribbed, weakly striate, gla-
brous; buds axillary, the scales 2, valvate, ovate, acuminate to acicular, 4-5 mm
long. Leaves alternate, coriaceous, slightly bullate, clustered near apex, ovate to
elliptic-ovate, 4.4-9 cm long, 1.4-4.5 cm broad, basally rounded to obtuse,
apically long-acuminate, marginally entire, glabrous on both surfaces but pro-
vided with brownish, glandular fimbriae beneath; 3-5-plinerved with inner
nerves arising 7-11 mm above base, midrib, lateral nerves, and reticulate vein-
letsimpressed above and raised beneath; petioles rugose, subterete, broadly flat-
tened above, 3.5-5 mm long, glabrate. Inflorescences axillary but usually from
older. leafless nodes, flowers 1-2(-3) per node, when 2-3 then with very short
LUTEYN, ERICACEAE OF BOLIVIA 7
rachis; rachis (when present) subterete, 1-4 mm long, glabrous, obscure: floral
bract ovate, acute, somewhat cucullate, ca. 2mm long, marginally glandular-
limbriate, glabrous; pedicels slender, subterete, striate, sharply angled distally,
14-18 mm long, glabrous; bracteoles nearly basal, ovate, acuminate ca. 3 mm
long, glabrous but marginally glandular-fimbriate. Flowers: calyx continuous
with pedicel, 6-8.5 mm long, essentially glabrous; hypanthium obpyramidal,
strongly 5-winged, 3.5-5 mm long, sparsely short-pilose along proximal por-
tions of wings; limb spreading, ca. 3-3.5mm long: lobes 5, broadly ovate, sharply
and shortly acuminate, ca. 2-2.8 mm long, sparsely short-pilose along margins;
sinuses acute; corolla of 5 fused petals, carnose, urceolate-turbinate, broadly
swollen basally, broadly and bluntly 5-angled, 8-11 mm long, 7-12 mm diam.
at base, glabrous, reddish-orange, the lobes 5, reflexed, deltate, acute, ca. 1.5mm
long, green in bud but becoming light green to white at anthesis; stamens 10,
equal, nearly as long as corolla, ca.8 mm long; filaments distinct, sparsely short-
pilose, ca. 2.5-3 mm long: anthers ca. 7-7.5 mm long; thecae smooth, incurved
at base, ca. 3.5 mm long; tubules distinct to base, ca. 3.5-4 mm long; ovary 5-
locular; style about equaling corolla. Berry not seen.
Distribution.—Endemic to Bolivia and known only from the province of
Nor Yungas, between Unduavi and Cotapata, at ca. 3000-3500 m. The plants
are found in the wet, moss-covered, cloud forest associated with Clusia,
Weinmannia, Hedyosmum, Brunellia, Miconia, and Desfontainea.
Etymology.—The species is named for the small village of Unduavi, close to
the locality from which all collections have been made.
Themistoclesia is a neotropical genus of ca. 25 species, ranging from Costa
Rica and Panama through the Andes of South America from Venezuela into
northern Bolivia. Themistoclesia unduavensis may be distinguished by its rhi-
zomatous, subshrub habit, somewhat bullate leaves, 1-3-flowered, axillary and
often ramiflorous inflorescences, carnose corollas that are bluntly 5-angled and
basally swollen, and reddish-orange corollas with green to white lobes. Indi-
vidually each of these characters is either absent or uncommon amongst the
other species in the genus, and the combination of characters effectively isolate
this species from all others.
—
—
Additional collections examined: BOLIVIA. La Paz. Nor Yungas: between Unduavi and Cotapata, as-
cending Paramo Yunqueno, 3200 m, 8 Apr 1996 (1), Beck 22680 (NY, LBP): Unduavi, 3000 m, Jul 1965
(1), Braun 37 (US); 4 km E of Unduavi Nuevo, mule trail of Rusby, 3300 m, 20 Mar 1988 (f1), Grifo &
Solomon 1005 (BH, NY); trail to Rio Coscapa, ca. 3 km E of Unduavialong new rd, 5.6 km W of Cotapata,
300-3500 m, 5 May 1990 (f1), Luteyn & Dorr 13476 (NY, LPB, plus 5 to be distributed): 1.4 km E of
Cotapata, 3200 m, 20 Mar 1988 (f1), Solomon 18195 (MO, NY).
APPENDIX
List of all species of Ericaceae attributed to Bolivia, including current status of
names given by Foster (1958). [Key: names appearing in italics are currently
18
BRIT.ORG/SIDA 20(1)
synonyms; names appearing in Roman are currently accepted; ! = new species,
BO = species endemic to Bolivia (or nearly so); * = species endemic to northern
Bolivia and adjacent central to southern Peru; ** = species not found in Bolivia;
CULT = introduced ornamental species|]
Agaricta holi
viensis (Sleumen Judd BO (but see
1990)
Judd and Hermann
Befar
eee
aestuans
Befaria denticulata Remy = Bejaria aestuans
Befaria glauca Bonpl. = Bejaria aestuans
Befaria glauca var. coarctata (Bonpl.) Mansf. &
Sleumer = Bejaria aestuans
Befaria ee var. glandulosa Mansf. & Sleumer
ejaria aestuans
Behan glauca var. setosa Mansf. & Sleumer =
Bejaria aestuan
Befaria glauca vat. anes Mansf. & Slemuer
Bejaria aestu
Befaria ee ee & Endl. = Bejaria
viensis - Fedtsch.& Basilevsk. = Bejaria
aestuans
Befaria parvifolia Rusby = Bejaria aestuans
Bejaria aestuans L
Cavendishia acuminata (Hook.) Hemsl. =
ckmanniana Hoerold =
Cavendishia bracteata (Ruiz & Pav. ex J.St.-Hil.)
Hoerold
Cavendishia martii (Meissn.) A.C.Sm. *
Cavendishia paniculata Rusby = Cavendishia
martil
Cavendishia pubescens (Kunth) Hemsl.
Cavendishia chan ae var. boliviensis Hoerold =
Cavendishia pubescens
Cavendishia sillarensis eres = Cavendishia
bracteata
Cavendishia_ strobilifera
Cavendishia bracteata
(Kunth) Hoer. =
n L Pritt ea | ees
Ceratostema serratum Britton = Thibaudia
macrocalyx
Chupalon viridiflorum Kuntze = Cavendishia
martil
Clethra spp. = Clethraceae
Demosthenesia fabulosa (Slemuer) A.C.Sm.= ?
(type and only specimen destroyed during
orld War Il; protologue insufficient for ge-
@
= ‘ie
mination)
Se nes graebneriana (Hoerold) A.C.Sm.
= D.mandonii
ii
mosthenesia mandonii (Britton) A.C.Sm. *
SR pearcei (Britton) A.C.Sm. BO
Demosthenesia spectabilis (Rusby) A.C.Sm. *
a
Diogenesia boliviana (Britton) Sleumer BO
Diogenesia racemosa (Herzog) Sleumer BO
eee canes haun Nied.
aaa ium (Benth)
—
fe sm. = eee alaternoide
Disterigma empetrifolium pe Drude
Disterigma ovatum (Rusby) S.F.Blake *
Disterigma pallidum A.C.Ssm.B
Disterigma pernettyoides (Griseb. ex Wedd.)
Nied. *
saaercbect ek bolivianum (Britton) Herzog =
en oliviana
ne ean sa racemosum Herzog =
Diogenesia racemosa
Gaultheria anastomosans (L.f.) Kunth =not
found in Bolivia, only in Colombia and Ven-
ezuela
Gaultheria barosmoides Rusby = Gaultheria
vaccinioides
ultheria eee DC. = Gaultheria
glo mera
Gaultheria ae (Cav.) G.Don *
Gaultheria pele Willd. var. secunda (Remy)
Luteyn
theria conferta Benth. = Gaultheria
astomosans
] difolia Kunth = Gaultheria erecta
Gaultheria erecta Ven
Gaultheria eriophylla (bars ) Sleumer ex Burtt var.
mucronata (Remy) Luteyn *
Gaultheria formosa Remy = Gaultheria erecta
Gaultheria glabra DC. = Gaultheria reticulata
Gau
+ o>
Kunth
Gaultheria glomerata (Cav.) Sleumer
Gaultheria hapalotricha A.C.Sm.
LUTEYN, ERICACEAE OF BOLIVIA
Gaultheria mucronata as = Gaultheria
ophylla var. mucrona
Gaul nena x WH Gault-
a pi gee Benth. = Gaultheria
ceed remyana A.C.Sm. = Gaultheria
hylla var. mucronata
eae reticulata Kunth
Gaultheria rufescens DC. = Gaultheria bracteata
Gaultheria saxicola Wedd. = Gaultheria
ccinioides
eine secunda Remy = Gaultheria buxifolia
r.secunda
ene serrulata Herzog = ? Gaultheria
vaccinioides x G. erecta (see Luteyn 1995)
Gaultheria tetriches Rusby = Gaultheria erecta
Gaultheria tomentosa Kunth *
Gaultheria vaccinioides Wedd. *
Gaylussacia cardenasii A.C.Sm. BO
Gaylussacia pseudogaultheria Cham.& Schlechtd
Rhododendron simsii Planchon CULT
Rusbya boliviana Britton = Diogenesia boliviana
Rusbya pearcei Britton = Demosthenesia pearcei
Rusbya taxifolia Britton BO
Satyria boliviana Luteyn BO
Satyria neglecta A.C.Sm. BO
Siphonandra boliviana Luteyn ! BO
Siphonandra elliptica (Ruiz & Pav. ex G.Don)
Klotzsch *
slacti magnifica Sleumer = ? (type and only
a n destroyed during World War II, but
“ ogue sufficient for determination
ae A.C.Sm.= Siphonandra elliptica
wi
ee robusta Rusby = Sphyrospermum
cordifo
sat Ale buxifolium feo & Endl.
Sphyrospermum cordifolium h.
ve in sessiliflorum ees BO
Themistoclesia peruviana A.C.Sm. *
ound in Bolivia, only in Brazil
Hornemannia boliviensis Kuntze = Thibaudia
boliviensis
1 L ee ee ee . 1 A ‘ | eae
Orthaea boliviensis B. ae & Basilevsk. BO
Orthaea constans A.C.Sm
Orthaea pinnatinervia Mansf. *
Orthaea rusbyi Luteyn BO
Orthaea weberbaueri Hoerold *
Pernettya densa Rusby = Pernettya prostrata
sas phyllyraefolia (Pers.) DC. =not found
n Bolivia, only Argentina and aie
Pe aes piesa (Cav.) D
Pernettya prostrata var. ie aad (D
= Pernettya prostra
— prostrata var. purpurea (D.Don ex
G.Don) Sleumer = Pernettya prostrata
ae schizostigma Rusby = Pernettya
rostrata
C.) Sleumer
Polyclita turbinata (Kuntze) A.C.Sm. BO
Psammisia elliptica (Rusby) A.C.Sm.= Psammisia
pauciflora
Psammisia guianensis Klotzsch
Psammisia pauciflora Griseb. ex A.C.Sm.
Themistoclesia unduavensis Luteyn ! BO
Thibaudia axillaris Rusby BO
Thibaudia boliviensis (Kuntze) Hoerold =
ibaudia crenulata
Thibaudia crenulata Remy *
Thibaudia densiflora (Herzog) A.C.Sm. BO
Thibaudia macrocalyx Remy BO
Thibaudia oblongifolia Remy = Cavendishia
pubescens
Thibaudia regularis A.C.Sm.*
Vacciniopsis ovata Rusby = Disterigma ovatum
Vacciniopsis tetramera Rusby = Disterigma
alaternoides
Vaccinium cece (G.Don) Sleumer *
Vac
dy Dunal =not in Bolivia,
on a Per
Vaccinium ee ifolium Kunth = Disterigma
empetrifolium
Vaccinium epacridifolium Benth. = Disterigma
empetrifo
Vaccinium floribundum Kunth
Vaccinium floribundum var. ramosissimum
(Du wo SHEE ae ou earel tle
dai.
= Vaccinium floribundum
Vaccinium marginatum Dunal = Vaccinium
floribundum
20 BRIT.ORG/SIDA 20(1)
Vaccinium penaeoides Kunth = Disterigma — Vaccinium polystachyum Benth. = Vaccinium
empetrifolium floribundum
ACKNOWLEDGMENTS
Stephan Beck encouraged me to write this paper. [thank the Herbario LPB (La
Paz, Bolivia), Jim Solomon, the Missouri Botanical Garden, and the late David N.
Smith for facilities and logistical support. Thanks and acknowledgments are
also given to the Jessie Smith Noyes Foundation and the National Science Foun-
dation under whose grants field work was undertaken and this paper was writ-
ten. [thank Stella Sylva for translating the key into Spanish and computer help,
Bobbi Angell for the beautiful illustrations, and Carmen Ulloa U. and Walter
Judd for critically reviewing the manuscript.
REFERENCES
Foster, R.C. 1958. A catalogue of the ferns and flowering plants of Bolivia.Contr.Gray Herb.
184:1-223. [Ericaceae, pp. 152-154]
Jubb, W.S. and PM. Hermann. 1990. Circumscription of Agarista boliviensis (Ericaceae). Sida
14:263-266.
Kiteen, TJ.,E.Garcia E.,and S.G. Beck (eds.). 1993. Guia de arboles de Bolivia. Herbario Nacional
de Bolivia and Missouri Botanical Garden, La Paz and St. Louis. [Ericaceae, pp. 279-285]
Kron, K.A., E.A. Powett, and J.L. Lutevn. 2002. Phylogenetic relationships within the blueberry
tribe (Vaccinieae, Ericaceae) based on sequence data from matK and nuclear riboso-
mal ITS regions, with comments on the placement of Satyria. Amer. J. Bot.89:327-336.
Luteyn, J.L. 1995. Gaultheria. Pp. 384-488. In: J.L. Luteyn, ed. Ericaceae—Part Il. The
superior-ovaried genera (Monotropoideae, Pyroloideae, Rhododendroideae,
Vaccinioideae p.p.). Fl. Neotrop. Monogr. 67:1—560.
Luteyn, J.L. 1997. A review of and taxonomic realignments within the neotropical genus
Macleania (Ericaceae: Vaccinieae). BioLlania, Edicion Esp. No. 6:455-465.
Luteyn, J.L.1998. Neotropical blueberries:The plant family Ericaceae. www.nybg.org/bsci/
res/lut2.
Luteyn, J.L.2001.Two new species and two new combinations in Mesoamerican Ericaceae.
Brittonia 53:437-446,
Luteyn, J.L. 2002. Diversity, adaptation, and endemism in neotropical Ericaceae: Biogeo-
graphical patterns in the Vaccinieae. In: K. Young, C. Ulloa Ulloa, J.L.Luteyn and S.Knapp,
eds. Plant evolution and endemism in Andean South America. Bot. Rev. 68:55-87.
Lureyn, J.L. (In press). Ericaceae (Heath Family). In: N.P. Smith, S.V. Heald, A. Henderson, S.A.
Mori and D.W. Stevenson, eds. Flowering plant families of the American Tropics. Princ-
eton University Press/New York Botanical Garden Press, Princeton, NJ and Bronx, NY.
Smity, A.C. 1932. The American species of Thibaudieae. Contr. U.S. Natl. Herb. 28:311-547.
TWO NEW SPECIES OF FESTUCA FROM SOUTH AMERICA
(POACEAE: LOLIINAE: SECT. SUBULATAE)
Daniel Stancik Paul M. Peterson
Department of Botany Department of Systematic Biology - Botany
Charles University of Prague National Museum of Natural History
Benatskd 2, 128 01 Prague Smithsonian Institution
CZECH REPUBLIC Washington, DC 20560-0166, U.S.A.
dan_stancik@yahoo.com peterson.paul@nmnh.si.edu
ABSTRACT
Two new species, Festuca cuzcoensis Stancik & PM. Peterson and F t is Stancik & PM. Peterson,
from the Andes of Bolivia and Peru are described and illustrated. The two new species appear to be
closely related to Festuca flacca Hack. ex E.B. Alexeev from Ecuador. A key to the South American
species of Festuca sect. Subulatae is given.
RESUMEN
Se describen e ilustran dos especias nuevas de Festuca de los Andes de Bolivia y Pert: F cuzcoensis
Stancik & PM. Peterson y EF tovarensis Stancik @ P.M. Peterson. Las nuevas especies se consideran
estrechamente relacionadas con la especies F flacca Hack. ex F.B. Alexeev de Ecuador. Ademas se
presenta una clave para la determinacion de las especies de Festuca sect. Subulatae en Suramérica
While making determinations of South American material in the United States
National Herbarium the senior author recognized several specimens with
unique features. The two new species discussed here appear to be members of
Festuca subg. Subulatae (Tzvelev) E.B. Alexeev sect. Subulatae. The position of
sect. Subulatae in subg. Subulatae is clear, however, the relationships with other
sections in this subgenus are unresolved. Alexeev (1980, 1982, 1986) recognized
three sections in subg. Subulatae: sect. Subulatae, sect. Elmera EB. Alexeev, and
sect. Glabricarpae E.B. Alexeev. Alexeev (1986) and Clayton and Renvoize (1986)
also recognized two other subgenera in these flat-bladed South American spe-
cies of Festuca: subg. Subuliflorae E.B. Alexeev and subg. Obtusae E.B. Alexeev.
Aiken’s et al. (1997) treatment included species of subg. Subuliflorae and subg.
Obtusae, sensu Alexeey, in subg. Subulatae sect. Obtusae, and moved a single
species from sect. Elmera,sensu Alexeev, to sect. Subulatae. In addition, Lu (1992)
described sect. LongiglumesS.L. Lu to include some Chinese species within subg,
Subulatae. Clearly, there are considerable differences of opinion among promi-
nent agrostologists as to possible taxonomic relationships among these Festuca
species.
Section Subulatae consists of about 25 species from North and South
America, Asia, and Africa (Aiken et al. 1997; Alexeev 1977, 1980, 1988; Lu 1992:
Stancik ined.). Individuals of Festuca sect. Subulate are typically loosely tufted
SIDA 20(1): 21 - 29. 2002
22 BRIT.ORG/SIDA 20(1)
with extravaginal innovations lacking cataphylls, with flat blades without well-
developed ribs, and have relatively large, open panicles with numerous branches.
In this section the spikelets are typically lanceolate with two short, entire, and
narrow glumes that are membranous to coriaceous/membranous. The coria-
ceous/membranous lemmas are lanceolate, entire, and usually long-awned or
rarely awnless. Anthers are short (1-2 mm long) and the ovary hairy, rarely gla-
brous. The two new species discussed here clearly are members of subfamily
Pooideae, tribe Poeae, and subtribe Loliinae (Soreng et al. 2001). With the addi-
tion of these two new species in South America, Festuca sect. Subulatae con-
sists of the following eight species: F cochabambana E.B. Alexeev, FE coromotensis
Briceno, F cuzcoensis, F flacca Hack. ex E.B. Alexeev, F. parodiana (St.-Yves)
Nicora, F sodiroana Hack. ex E.B. Alexeev, F tovarensis,and F. ulochaeta Nees ex
Steud.
Festuca cuzcoensis Stancik & P.M. Peterson, sp. nov. (Fig. 1). Type: PERU.
DEPARTAMENTO CUZCO. Provincia Urubamba: above Ollantaytam| shade, high
up ravine, 3600 m, 5 Dec 1923, A.S. Hitchcock 22532 (HOLOTYPE: US- er
Festuca ulochaeta auct. non Nees ex Steud: Hitchcock 1927, 321 p.p. Macbride 1936, 123 p.p. Tovar
1S
90 p.p.
Haec species a Festuca ulochaeta Nees ex Steud. et F tovarensis Stancik & P.M. Peterson culmis
ee, 12 non 2-4) et aristae brevi [3-4 (-5) non 5-12 mm] differt. A Festuca ulochaeta Nees
cud. et F coc cae bana E.B. Alexeev ligula folii 2-3 (non 0.2-2) mm longa distinguitur
Perennial, loosely tufted. Culms 80-110 cm tall, erect, glabrous, with 6-12 nodes
in basal half, the internodes short. Sheaths membranous, brown, more or less
pubescent, margins free; innovations extravaginal; auricles absent. Ligules 2-3
I acuminate, dentate. Blades 12-15 cm long, 3-9 mm wide,
flat, green, scabrous on ribs of abaxial surface. Panicles 20-25 cm long, 5-10 cm
wide, open; branches pendant, scabrous. Spikelets 8-10 mm long; florets 3-4
(perfect); rachilla 12-14 mm long, puberulent. Glumes narrowly lanceolate,
coriaceous, purplish, scabrous, apex acute; lower glume (2-)2.5-4.5 mm long, l-
veined; upper glume 4-6 mm long, inconspicuously 3-veined. Lemma 7-8.5mm
long, lanceolate, chartaceous to membranous, 3- or 5-veined, purplish-green;
apex scabrous, entire; awn 3-4(-5) mm long, scabrous, straight. Palea almost as
long as lemma, 2-keeled, the keels finely scabrous, deeply 2-dentate. Lodicules
oblong, 2-dentate. Stamens 3; anthers (1.2-)1.5-2 mm long. Ovary apex with a
few hairs. Caryopses lanceolate.
Distribution and habitat.—This species was collected in Andean forest zone
of Peru and northeastern Bolivia between 3200-3850 m.
Etymology.—The specific epithet refers to the type locality region, ie, De-
partment Cuzco in southern Peru.
mm long,
——
Additional s ined: BOLIVIA. Departamento La Paz: Unduavi, in Walden, 3300 m, 12
‘eb 1907, fue htien 6415 (US). PERU. Departamento Ancash. Prov. Yungay: Huascaran National Park,
\
“4 ‘ Re
PARTANCERIN! 2002
t
f a blad D. Spikelet
A +e
|. Lodicules. J. Pistil. K
|
k 2253). A. Habit. B. Inflorescence. C. Sheath, ligule
Fic. 1. Fest
daitd
E. Floret. F. Lemma, ventral view. G. P.
Stamen. L. Caryopsis, dorsal view. M. Caryopsis, ventral view.
BRIT.ORG/SIDA 20(1)
Llanganuco sector, between Chinancocha and Pucayacu (77° 39’ W-09' 05'S), 3700-3850 m, 7 May
1985, Smith 10517 (USM); Smith 10551 (USM), Smith 10561 (USM). Proy. Huari: Huascaran National
Park, Quebrada Pachachaca, a lateral valley : Quebrada Rurichinchay (77° 16’ W-9" 23'S), 3840-
3870 m, 13 Jun 1986, Smith etal. 12632 (USM). 1 t to Cuzco. Prov. Calca: 7 km SE of Lares on
road towards Pampa Corral and Calca (13" 08' 49.9" S-72' 00' 34.2" W), 3620 m, 17 Mar 2002, Peterson
ulio-Rodriguez 16582 (PRC, US, USM). Departamento Huancavelica. Proy. Tayacaja: Near
a
& R ef
Salcabamba, 3200 m, 13 Apr 1962, Tovar 3612 (USM).
Festuca tovarensis Stancik @ P.M. Peterson, sp. nov. (Fig. 2). Type: PERU.
Departamento Huancavelica, Provincia Tayacaja: Chuspi-Hda. Tocas, entre
Colchabamba y Paucarbamba, monte bajo, 2800 m, 22 Apr 1954, O. Tovar 2057
(HOLOTYPE: US-2181286!; soTYPE: USM),
Festuca ulochaeta auct. non Nees ex Steud: Tovar 1993, 90 p.p. Brako L. et al. 1993, 950.
Haec species a Festuca ulochaeta Nees ex Steud. lemmate breviore (5.5-6.5 non 6-9 mm), arista 5-7
5) mm longa et recta (non flexuosa) differt. A Festuca flacca Hack. ex E.B. Alexeev ligula folii
(non 7-1
perioribus 3.5-5 (non 2.3-3.5)
~2 (non 2-4.5) mm longa, truncata et ciliata (non lacerata), glumis su
mm et antheris 1.5-1.6 (non 0.9-1.2) mm distinguitur.
Loosely tufted perennials. Culms 70-90 cm tall, erect, glabrous, with 2-4 nodes
in basal half. Sheaths membranous, brown, margins free; innovations extrav-
aginal; auricles absent. Ligules 1-2 mm long, membranous, truncate, margins
ciliate. Blades 8-15 cm long, 1.5-4.5 mm wide, linear, flat, green, scabrous with
ribs on abaxial surface. Panicles 15-20 cm long, 7-10 cm wide, open; branches
pendant, scabrous. Spikelets 7.5-9.5 mm long; florets 3 (perfect); rachilla 1.1-1.4
mm long, puberulent. Glumes narrowly lanceolate, coriaceous, purplish, gla-
brous, apex acute (sometimes scabrous): lower glume 1.5-1.8 mm long, l-veined;
upper glume 3.5-4.5(-5) mm long, 1-3 veined. Lemma 5.5-6.5 mm long, lan-
ceolate, chartaceous to membranous, 3-veined, purplish-green, scabrous; apex
entire; awn 5-7 mm long, terminal, scabrous, straight. Palea as long as lemma,
2-keeled, the keels scabrous; apex hairy, deeply 2-dentate. Lodicules ca. 0.8 mm
long, lanceolate, acuminate. Stamens 3; anthers 1.5-1.6 mm long. Ovary sparsely
hairy near apex. Caryopses lanceolate; hilum linear, 2/5-1/2 of total length.
Distribution and habitat—This species is known only from the Andean
orest zone of Peru, at 2500-3250 m.
Etymology.—The specific epithet honors the eminent Peruvian botanist and
—
agrostologist, Oscar Tovar Serpa.
Additional specimens examined: PERU. Departamento Ayacucho. Proy. Huanta/La Mar
Volken-Nebel-Buschwald, 37 km to Ayna, 3250 m, 23 Mar 1977, Ellenberg 7024 (USM). annueai
Cajamarca. Prov. Cajamarca: Road to Yumagual, 2500 m, 26 Jun 1966, Vega 249 (USM).
DISCUSSION
In South America sect. Subulatae is represented by eight native species (Table
|). In this table we compare the salient morphological features that distinguish
these taxa, including the North American species, FE subulata Trin. All the spe-
=
SS
SSS
7 J
2.002,
\ AR face RINT
pega
IF
f a blade. C Spikelet D. Floret. E Lemma,
with lodicules, pistil, and stamens. H. Lodicules. |. Pistil. J. Stamen. K. Cary-
.
nA
t
2057). A. Habit. B. Sheath, ligule
ventral view. F. Palea with stamen.G. Palea
dorsal view. L. Caryopsis, ventral view.
opsis,
Taste 1. Morphological comparison of the South American species of Festuca sect. Subulatae and the North American Festuca subulata.
Taxon F.coromotensis F.cochabambana F.ulochaeta F.cuzcoensis __F. tovarensis F. flacca F.sodiroana F.parodiana F.subulata
geographical
distribution Venezuela Bolivia Argentina, Peru, Peru Ecuador Ecuador, Argentina = North
Brazil, Bolivia Colombia America
Colombia,
Venezuela
culm nodes 3-5 3-4 2- 6-12 2-4 3-7 2-4 5-7 3-4
ligule size (mm) 0.5-1 0.2-0.7 0.3-1(-2) 2-3 1-2 3-35 0.5-1.1 1-2 0.5-1
form of ligule truncate truncate truncate acuminate truncate acuminate truncate truncate truncate
spikelet size (mm) 11-15 9-12 8-10 7.5-9.5 7-8 75-9 10-13 8-10
number of florets 3-4 4-5 3-5 3-4 3 2-4 4-5(-6) 4-6 3-4
lower glume size (mm) 2.5-3 1.8-2.2 2.5-3.5(-4 (2-)2.5-4.5 1.5-1.8 1.2-2.4 1.2-1.8 2-2.5 2.5-4
upper glume size (mm) 4.8-5.5 2.7-3.2 3.5-4.5 4-6 3.5-4.5(-5) 9 2.5-3.5 2.0-2.9 3-4 3.5-5
lemma size (mm) 8-8.5 5.5-6.2 6-8(-9) 7-85 5.5-6.5 6.5-7 5-65 7-8 6-8
awn size (mm) 0-0.5 6-12 7-15 3-A(-5) 5-7 9-12 awnless awnless 5-10
form of awn Straight straight flexuous straight straight straight awnless awnless Straight
anthers size (mm) 1.2-2 1-1.2 1.1-1.5 (1.2-)1.5-2 1.5-1.46 1.1-1.4 0.8-1.2 1.3 1.5-2
tip of ovary glabrous hairy sparsely sparsely sparsely sparsely glabrous hairy hairy
hairy hairy hairy hairy
(L)0Z vaIs/DuO'LINa
AMAEDICA 7
cies are morphologically very similar to one another. However, F ulochaeta can
be easily separated from the remaining seven species by having long-awned
lemmas with the awn flexuous whereas the other taxa have straight awns or
awnless lemmas. Festuca cuzcoensis and F tovarensis are possibly most closely
related to F flacca since all three share lanceolate spikelets of the same size, 2-
4 florets per spikelet, straight awns 3-12 mm long, and small anthers 1.1-2 mm
long. Festuca cuzcoensis differs from F tovarensis by having 6-12 nodes per culm
(2-4 nodes in F tovarensis), acuminate ligules (verses truncate), lower glumes
(2-)2.5-4.5 mm long (verses 1.5-1.8 mm), lemmas 7-8.5 mm long (verses 5.5-6.5
mm), anda pierous ovary (verses sparsely hairy). Festuca parodiana shares
the following characteristics with FE sodiroana: truncate ligules, 4-6 florets per
spikelets, and awnless lemmas. However, F parodiana can be separated from F
sodiroana by having a hairy (glabrous in E sodiroana) ovary, lower glumes 2-
2.5 mm long (verses 1.3-1.8 mm), upper glumes 3-4 mm long (verses 2.2-2.9
mm), lemmas 7-8 mm long (verses 5-6.5 mm), and 5-7 nodes per culm (verses
2-4). Festuca coromotensis is almost awnless (mucronate) but differs from F
sodiroana and F parodiana by having large, scabrous lemmas 8-9 mm long.
There appears to be a geographic component as well since the principal area of
distribution for F ulochaeta is southeastern Brazil and northeastern Argentina
with new range extensions reported from the Cordillera Oriental of Colombia
and Cordillera de Mérida in Venezuela (Stancik 2001). The remaining seven
species occur exclusively in the montane forest zone of the Andes in small en-
demic populations ranging from Argentina in the south to Colombia and Ven-
ezuela in the north. Festuca elviae Briceho, a Venezuelan species, is also a pos-
sible member of Festuca sect. Subulatae. However, further study is needed to
confirm its inclusion in this section.
KEY TO THE SPECIES OF FESTUCA SECT. SUBULATAE IN SOUTH AMERICA
. Lemmas awnless or with a mucro to a: mm long.
2. Lemmas 8-9 mm long, densely scabrous F. coromotensis
2. Lemmas 6.5-8 mm long, aes or papillate.
3. Culms with 2-4 nodes; lower glumes 1.2-1.8 mm long; upper glumes 2.0-2.9
mm long; lemmas 5—6.5 mm long; ovary glabrous F. sodiroana
3. Culms with 5-7 nodes; lower glumes 2—2.5 mm long; upper glumes 3-4 mm
long; lemmas 7-8 mm long; ovary hairy F. parodiana
1. Lemmas with awn 3-15 mm long.
4. Lemma awns markedly flexuous, 7-15 mm long F. ulochaeta
4. Lemma awns — 3-12 mm lonc
5. Sheaths pubescent F. cochabambana
5. Sheaths seats rarely scabrous.
6. Lower glumes (2-)2.5-4.5 mm long; lemma awns 3-4 mm long
F. cuzcoensis
6. Lower glumes 1.2-2.4 mm long; lemma awns 5-12 mm long.
28 BRIT.ORG/SIDA 20(1)
7. Ligules 2-3.5 mm long; upper glumes 2.5—3.5 mm long; lemma
awns 9-12 mm long F. flacca
7 Ligules 1-2 mm long; ur upper glumes 3.5—4.5(—5) mm long; lemma
awns 5—7 mm long F, tovarensis
ACKNOWLEDGMENTS
We wish to thank the Smithsonian Institutions, Fellowships and Grants for
supporting a research visit by the senior author to the United States National
Herbarium. The senior author would like to thank ICETEX for financial sup-
port in Colombia, the Instituto de Ciencias Naturales de la Universidad Nacio-
nal de Colombia for use of their study facilities, and the Grant Agency of the
Czech Republic (Grant GACR No. 42-201174) for financing fieldwork in Ecua-
dor. We thank curators from the following herbaria: AAU, B, COL, LPB, MA, PR,
PRC, QCA, QCNA, VEN, and W. Appreciation is extended to Alice R. Tangerini
for providing the illustrations, Dan Nicolson for reviewing the Latin diagnoses,
and Susan Pennington for technical support. Gerrit Davidse, Stephan L. Hatch,
Robert D. Webster, and Barney Lipscomb are thanked for reviewing the manu-
script on short notice.
REFERENCES
Aiken, S.G., MJ. Datiwitz, C.L. MCJannet, and L.L. Consaut. 1997. Biodiversity among Festuca
(Poaceae) in North Raita anode evidence from DELTA and clustering programs,
and an INTKEY package for interactive, illustrated identification and information re-
trieval. Canad. J. Bot. 75:1527-1555.
Atexeev,E.B.1977.On the systematics of Asian fescues (Festuca). 1.subgenera Drymanthe
Subulatae, Schedonorus, Leucopoa. Bjull. Moskovsk. Obsc. Isp. Prir. Otd. Biol. 82:95-102.
Acexeev, E.B. 1980. Festuca L. subgenera et sectiones novae ex America et Mexica. Novosti
Sist. Vyss. Rast. 17:42-53.
Acexcev, E.B. 1982.A new section and three new species of the genus Festuca from Mexico
and Central America. Bot. Zhurn. (Moscow & Leningrad) 67:1 289-1292,
Atexeey, E.B. 1986. Festuca L. (Poaceae) in Venezuela, Colombia et Ecuador. Novosti Sist.
Vyss. Rast. 23:5-23.
Acexeev, E.B. 1988.Genus Festuca (Poaceae) in Japonia, Peninsula Coreana et Insula Taiwan.
Novosti Sist.Vyss. Rast. 25:5-27.
Brako, L. and J.L. Zarucci. 1993. Catalogue of the flowering plants and gymnosperms of
Peru. Monogr. Syst. Bot. Missouri Bot. Gard. 45:1— 1286.
Crayton, W.D. and S.A. Renvoize. 1986.Genera graminum. Grasses of the world. Her Majesty's
Stationery Office, London. Pp. 93-94.
HitcHcock, A.S. 1927. The grasses of Ecuador, Peru, and Bolivia.Contr.U.S. Natl. Herb. 24:291—-
DOO:
Lu, L.S. 1992. New taxa of Festuca L. from China. Acta Phytotax. Sinica 30:529-540.
Maceripe, J.F. 1936. Gramineae, Flora of Peru. Field Mus. Nat. Hist., Bot. Ser. 13:96-261,
>
Sorena, R.J.,G. Daviose, PM. Peterson, F.O. Zutoaca, EJ. Juoziewicz, and T.S. Fitcueiras. 2001. Cata-
logue of New World grasses (Poaceae), suprageneric classification. http://
mobot.mobot.org/W3T/Search/nwgc.html
Stancik, D.2001.New records of the genus Festuca L. (Poaceae) for Colombia and Venezu-
ela. Caldasia 23:337-339.
Tovar, O.1993.Las gramineas (Poaceae) del Peru. Ruizia 13:1—480.
TUrPE, A.M.1969.Las especies argentinas de Festuca (excluidas las patagonicas). Darwiniana
15:189-283.
BRIT.ORG/SIDA 20(1)
BOOK REVIEW
James D. Mausetu, Roerto Kirs_inc, and Cartos Osto.aza. 2002. A Cactus Odyssey.
(ISBN 0-88192-526-8, hbk.). Timber Press, Inc., 133S.W. Second Avenue, Suite
450, Portland, OR 97204-3527, US.A. (Orders: wwwtimberpress.com, 800-
327-5680, 503-227-2878, 503-227-3070 fax). $39.95, 306 pp, 191 color pho-
tos, +4 maps, 6" x 9"
—
This book describes the history /evolution of cacti in South America showing the dynamic variety ol
sitats from high desert to low rainforest. A Cactus Odyssey is a fine addition to botanical literature
neriences taken from the extensive careers of the authors in the region, along with nu-
ha
based on ex]
merous field trips in Bolivia, Argentina and Peru between the years 1995 and 2000. The hard work
and combined expertise of three botanists, James Mauseth from Texas, Roberto Kiesling from Ar-
gentina, and Carlos Ostolaza from Peru led to the publication of this tour through the parts of South
America that supports diverse cactus populations. Along with the unusual plants that are studied,
the authors’ interactions with the local peoples that are encountered, provides interesting perspec-
tives on the areas where they traveled. A point of particular interest is the wide range of morphologi-
cal differences among these cacti found in very different places. Dr. Mauseth has done extensive work
“the Cactaceae. In the book, many points are made illustrating
on the anatomy and physiology o
why certain cacti grow as they do, and why they are so well adapted to their sometimes harsh sur-
roundings. The geological history of the Americas is also shown as a key factor in the evolution ol
Cactaceae.
Written not only for botanists, the terminology and amount of technical detail are well pre-
sented, giving the reader good definitions and descriptions. Included in the explanation of why the
Cactaceae are only native to the Americas, the authors tell the reader what the common traits are
among all cacti; those being the presence of areoles, clusters of spines, betalain pigments, and the
haracteristic structures of flower, fruit and seed. Those traits combine to make the cacti very difler-
rom other continents. A discussion of the water storage system in
)
ent from similar looking plants
ifferent tissues related to the environment, give the reader insight
and the special modifications of ¢
to the incredible process of survival that the cacti have developed living in such extreme environ-
ments. The herbarium specimen, seed and other collections taken for lab examination have given the
rors thousands of hours of future lab work, as well as potential reams publishable data
The book includes excellent photography; some very impressive photographs taken at high
aul
elevation show hillsides otherwise barren except for compact groups of densely spined cacti. The
differences in growth habits from the high elevation to middle and low elevation populations are
-} > ith +} -
Ips wn othet
significant. Major c cen in soil type and weather, as well as the benelicial relationsh
plants, increase the ability for cacti to have more vigorous growth. “Nurse” plants range from trees
ss
and shrubs in woodlond areas, to tall grasses in prairies, to rain forests trees where the cacti are ¢
phytes on branches along with bromeliads. An interesting note taken of the cacti that are shown is
growing saguaro cacti in Arizona. Close to 18 differ-
how many of them look very similar to the tal
ent genera are shown as tall, branching plants, on Ww ue eenes ies aes are very di fferent
‘oe
from their North American cousins. The efforts of
through parts of the world seldom seen by travelers. thie is enlishienine work eee habitat
and aspects of a spectacular plant family and its evolution never before considered by most people
—Justin Allison, Botanical Research Institute of Texas, Fort Worth, TX 76102-4060, U.S.A.
SIDA 20(1): 30. 2002
A NEW SPECIES OF ERIOGONUM (POLYGONACEAE)
FROM SOUTH TEXAS
Guy L.Nesom
Botanical Research Institute of Texas
509 Pecan Stre
Fort Worth, TX 7610 ae USA,
ABSTRACT
{
lants previously identified as E. multiflorum Benth.
ple
Ther new species is known from 17 countiese! the Rio Grande plains in south-central Texas. Eriogonum
fers from E. multiflorum in its broader leaves with auriculate-subclasping bases (vs.
attenuate, non-clasping leaf bases) and its involucres with more numerous flowers (22-28(-36) vs.
(-15) flowers). The two taxa are primarily allopatric and sharply Scout OHOn>) in morphology.
Putative intermediates have been observed in three counties. Both inct from the ap-
I
parently closely related E.annuum Nutt. in their ee peduncles, sceay tomentose and glabres-
cent involucres, and cordate perianth lobes.
RESUMEN
a] ] fe ]
multiflorum
se ies eeceus sp. Nov. se segrega cle
Benth. L a especie se conoce de 17 condad os de las Ilanuras de Rio Grande en el sur-centro de
Texas. Eo riograndis difiere de E. multiflorum en sus hojas mas anchas con bases auriculado-
semiabrazadoras (en vez bases de las hojas atenuadas, no ane antes) y sus EEvoleros con flores mas
numerosas (22-28(-36) vs. 6-12(-15) flores). Los d fuertemente
oe en su Aner pleeta: Se han observado iaiermienios putativos en tres condados. Ambas
muy relacionado E.annuum Nutt. Por sus aa a mas
as involucros esparsamente tomentosos y lobulos del perianto glabrescentes, y corda
The Texas species of Eriogonum Michx. were treated in detail by Reveal (1968,
1970), who observed (1968, p. 203) that E. multiflorum Benth. and E.annuum
Nutt. “are the most frequently collected species in the state and collectively the
most widely distributed species.” Together, they constitute subg. Micrantha
(Benth.) Reveal (Reveal 1969). Reveal provided a set of detailed morphological
contrasts to distinguish these two taxa, noting that their difference in perianth
lobe shape allows easy identification. He did not call attention to significant
infra-specific variation in either species.
The present study finds that plants pony identified as Eriogonum
multiflorum can be separated into two, morphologically distinct sets of popu-
lations. Those occupying the largest part of the range in Texas, along with those
from peripheral localities in northwestern Lousiana, southwestern Arkansas,
and south-central Oklahoma (Fig. 2), are identified here as typical E.
multiflorum. The type collection of E. multiflorum (Bentham, Trans. Linn. Soc.
London 17:413. 1837) was made in 1833 by Thomas Drummond (Drummond
SIDA 20(1): 31 - 38. 2002
32 BRIT.ORG/SIDA 20(1)
“between Brazoria and San Felipe de Austin,” perhaps in what is now Austin
County (from where a modern collection of E. multiflorumalso has been made).
Anillustration based on the Drummond type material, presumably at K (Hook.
Icon. Pl. 3, t. 250. 1840) shows leaves with non-clasping bases and the descrip-
tion notes “Involucres with about 6-9 flowers”—these features (see below) along
with the general habit unambiguously establish the identity of the species.
Observation of additional type material of E. multiflorum (GH photocopies.
isotype and syntypes fide annotations by J. Reveal in 1967) cor roborate this.
Plants identified as FE. multiflorum in the Rio Grande plains of southern Texas
represent a previously undescribed spec
ala a riograndis Nesom, sp. nov. (Figs. 1, 2). Typr: U.S.A. TEXAS. SAN PATRICIO
J mi E of Sinton, Welder Foundation peas Area, near headquarters build-
a grassy field along swale a in ie 28 Oct 1958, FW. ¢ Guild 8510 (HOLO-
BRIT-SMUI ISOTYPES: TAES!,
Eriogono multifloro Benth. similis duratione annua, caulibus foliosis tomentosis, periantho stipitato,
blongi-cordatis; differt foliis majoribus ad basim auricu
et lobis externis perianthii o ati-
subamplectentibus et involucris floribus plus numerosis.
Annual or biennial herbs from a taproot, eglandular. Stems erect, 2.5-6(-H, ~20
fide Reveal 1970, Richardson 1995) dm tall, usually unbranched until the inflo-
rescence, closely but loosely gray-white and persistently woolly-tomentose.
Leaves mostly on the lower 3/4-4/5 ol the stem, basal not persistent, alternate
oblong-obovate to oblong-elliptic or oblong, epetiolate and shallowly auricu-
late-subclasping, apex rounded to obtuse, 15-40(-45) mm long, 10-22(-25) mm
wide, margins flat or narrowly revolute, entire but often closely undulate, up-
per surface lightly but persistently woolly-tomentose, lower surface densely
and persistently white- to tawny-tomentose. Inflorescences cymoid, usually
relatively compact. Involucres on peduncles 2-5 mm long, turbinate to turbi-
nate-campanulate, 2-2.8 mm high, 2-3 mm wide, 5(-6)-lobed, externally
sparsely but persistently tomentose to glabrate, internally densely tomentose.
Flowers 22-28(-36) per involucre, exserted from the involucre on filiform
pedicels. Perianth bright white to pinkish, maturing or drying orangish, exter-
nally glabrous, internally sparsely tomentose; perianth lobes dissimilar, outer
lobes basally cordate and broadly oblong to oblong-elliptic, 2-2.7 mm long, L.5-
22 mm wide, midrib thick from base to apex, inner lobes linear, 0.1-0.2 mm
wide. Fruits dark brown to red-brown, |.9-2.2 mm long, not winged, glabrous.
Habitat and phenology.—Fields, pastures, roadsides, mesquite prairies, oak
woodlands, and other open habitats, sandy soil; (Aug-)Sep-Nov(-Dec).
Distribution.—The distribution of Eriogonum riograndis (Fig. 2) corre-
sponds to phytogeographic pattern #17 of Sorrie and Weakley (2001): South
Texas-Northeast Mexico Mesquital. They note that “The northern portion is
commonly denoted as the South Texas Plains, but ecologically the whole area
a
—
NESOM, A NEW SPECIES OF ERIOGONUM FROM TEXAS
Fic. 1 U-ahi¢e Pe pe eee | Pay aig - ee es L ! , BRIT)
34 BRIT.ORG/SIDA 20(1)
Io |. intermediate A
° ;
Ajalg ) 100 200
x 4 | E. riograndis 4
(am miles
105° 100° . 95° 30°
Zee | £ |
listributi f Eriog jograndi d E. multiflorum. Records are from BRIT-SMU, CAMU, OKLA,
ms, Tee: UL and VDB.0 y bol p 1c f, II n ° hj ty | lind f Conn tah
is more properly termed the Tamaulipan Scrub region, or Tamaulipan Mesquital,
due to the dominant shrubby vegetation, especially Prosopis glandulosa Torr..
(Sorrie & mee ley 2001, p. 61). The distribution of the recently described
Pseud austrotexanum Nesom (Nesom 2001) also fits the same phy-
togeographic pattern. Reveal (1970) noted that E. multiflorum occurs in Mexico;
presumably this would be E. riograndis in the interpretation here, but a speci-
men documenting the Mexican distribution has not been seen in the present
study
Etymology.—The appositive epithet refers to the central geographic feature
in the range of the new species, the Rio Grande (river).
Additional S.A. Texas. Aransas Co.: +.7 mi E of Rockport, 19 Sep 1936, Parks
and Cory 20336 (TAE S); Aransas Refuge, 30 Sep 1944, Cory 45936 (TAES): St. Joseph Island, irregular
dunes beyond back beach, 7 Nov 1964, Andrews 72 (1 EX), Aransas Wildlife Refuge, sandy soil at
Equipment Shed, 18 Sep 1968, Fleetwood 9317 (LL). sandy soilalong Hwy 35ca 5 mi Nol Rockport, 26
Sep 1981, Brown 5421 (TAES), 1/2 miN of Goose Island State Park on ne Road 13, common in clear
ing in oak woodland, 29 Dec 1982, Fryxell 3836 (BRIT, TEX). Brooks Co. 3 mi Eof Encino, 6 Jul 1935
[sterile], Parks and Cory 141421 (TAES); 14 mi S of Falfurrias, sandy oak reeion 15 Sep 1942, Lundell
inston 541455
LU
11941 (LL); a few mi F of Jim Hogg Co. line on Rd 755, loose sand prairie, 13 Sep 1954, Jo
NESOM, A NEW SPECIES OF ERIOGONUM FROM TEXAS 35
(TEX); 4 mi SE of King Ranch, Encino Division Headquarters, in deep, loose sand blow-out area, 18
Nov 1954, Gould and Morrow 6705 (SMU, TAES, TEX); 15 mi E of Hebbronville on State Hwy 285, in
Pleistocene gravel, 25 Nov 1962, Dohnke 6 (SMU, TEX); 12 mi E of Hebbronville, Hwy 285, 10 Nov
1962, Solis 48 (TAES, TEX), 4.6 mi S of FR 2191 (extension) from its ject with TX 285, Mariposa ranch
gate, SE of Falfurias, open hillside to edge of oak mott i: paeED sendy saul : Oct 1990, S.& G. Jones
6081 (V DB). Calhoun Co.: Matagorda Island, 1955, Balls.n nd, access road from
air base HQ to beach at end of island, ca 2 mi E of base, al 1973 [not f Tver Hartman 3722
(TEX). Duval Co.: Benavitas, 340 ft, 12 Aug 1941, Fisher 41101 (TEX). Hidalgo Co.: of Edinburg, 11
Nov 1942, Walker 126 (LL); ca. 30 mi N of Edinburg, in sand, 23 Aug 1944, a 44,276 (SMU);
par Ranch, 10 mi N of Edinburg, in sand, 2 Nov 1973, Everitt s.n. (SMU); La Reforma Training
ea (Texas Nati Guard), N end of Sector C, openings in mixed mesquite-black Be ush Sha ane
on ae level cee 6 Oct 1993, Carr 13177 (TEX); La Reforma Training Area (Ti
along main N-S road, 1.4 road mi N of gate 24, grazed grassland on gently rolling upl ae with scat-
tered mesquite, 255 ft, 250ct 1994, Carr 14302 (TEX). Jim Hogg Co.: State Hwy 285 E of Thompsonville,
in light ane loose sand, 11 Nov 1962, Ramirez, Alva, and McCart 8708 (SMU, TAES, TEX): 10 mi N
of La Gloria on Texas Farm Rd 1017, 15 Oct 1990, Miller, Brant, & Noyes 5811 (VDB): E side of FM 1017,
3.0 road mi S of jet with smaller road at Agua Nueva, ca. 450 ft, occasionally mown grassland in deep
loose sandy soil, roadside, 7 Oct 1993, Carr 13208 (TEX). Jim Wells Co.: sandy loam in dry lake,
Romarsid Ranch, | Nov 1943, Freeborn 142 (TEX). Karnes Co.: 2 miS of Karnes City, frequent in dry,
deep sandy soil, fallow field, 27 Oct 1952, Johnson 1010 (LL, SMU, TAES); 3 mi NE of Kenedy, dry
Escondido Creek, infrequent in dry sandy loam soil, 25 Jul 1954 [very early bud], Johnson 1618 (SMU).
Kenedy Co.: 18.7 mi N of Raymondville, frequent along road in sandy soil, 2 Dec 1945, Cory 51495
(SMU); Norias Division of King Ranch, Saltillo Pasture, 14 Sep 1953, Johnston s.n.(TAES, TEX), 17 Sep
1953, Johnston s.n. (TAES, TEX), 24 Nov 1953, Johnston s.n. (SMU, TAES, TEX); 18 mi S of Riviera, dune
area, 12 Oct 1952, Morrow and Nord 19 (TAES): Norias Division of King Ranch, =F corner of ranch,
open sandy coastal acre 23 Sep 1958, Lundell and cla 5155 (BRIT, LL); 3.4 mi N of Armstrong,
fine gray sand, 6 Nov 1953, Shinners 17082 (SMU). Kleberg Co.: Padre Island, 17 o 1940, Cory 36795
(SMU), Cor ene Cory 36797 (TAES), Cory ee Cory 36799 (TEX); Kingsville, sum-
mer 1940, Sinclair s.n. (TEX); Laureles Division of King Ranch, loose sand, 15 Sep 1953, Ta nston s.n.
(TEX), 29 Nov 1953, Johnston s.n. (TEX); Brooks County line on Road No. 285, loose sand salar
prairie, 14 Sep 1954 sie lata 541513 ie TEX); Padre Island National Seashore, end of service road
todump area, small st ne zed dunes near margin on brackish back island pond, 14 Oct ee emke
3009 (TEX). N o Bay, 5 Oct 1922, Tharp 1552 (TEX); Corpus Christi, 18 Dec 1935 [past
fruit], ie Tessar Mustang Island, loose sand, 26 Oct 1954, Hildebrand 82 (TEX),
Mustang Island, sand dunes 300 yds from Gulf, 11 Nov 1965, Crutchfield 1027 (LL); Mustang Island
cemetery, Port Aransas, sandy soil near fence, 14 Oct 1967, Gillespie 110 (TEX); eure au
rock B ee area, west side of island, 29 Jul 1967, Gillespie 200 (TEX); Mustang Island State Par ’ of
~ Rd 5 O ft SW of Corpus Christi Pass, dry sand along elevated et: in tidal flat, oo {t, 20
Oct 1989, oa 10158 (TEX). Refugio Co.: Black Jacks below Austell, 4 Dec 28, Phipps s.n. (TEX). San
Patricio Co.: 1.8 miSW of Aransas Pass, 19 Sep 1936, Parksand Cory 20338 a ES): 4 mi SW of Ingleside,
19 Sep 1936, Parks and Cory 20339 (TAES); 2 mi SE of Ingleside in deep fine sand, 2 Oct 1950, Jones 383
(SMU); 1 miS of Ingleside in deep fine sand, 8 Jun 1951 [pre-flowering], Jones 565 (SMU); Welder Wild-
life Refuge ca. 13 mi E of Sinton, high banks above Aransas River in Mare Trap Pasture, occasional in
tight sandy loam, 29 Sep 1956, Rowell 5212 (SMU). Webb Co.: 3 mi S of Mirando City at Los Ojuelos, in
grayish black loose sand, LO Oct 1961, Magnon and Rodriguez 14 (SMU). Willacy Co.: Yturria, 50 ft, 6
Aug 1924 [not in nee Runyon 665 (TEX); along hwy from Yturria Station north, 24 Sep 1937,
Runyon 1808 (TEX); Yturria, open dry sandy ground, 10 m, 22 Sep 1939, Runyon 4237 (TEX);
haa le, a 1941, Shiller 759 (TAES, TEX): Sauz Ranch, sand, 23 Nov 1953, Johnston and Davis
sn. (SMU, TAES, TEX). Zapata Co.: 7 mi N of Zapata, US Hwy 83, 15 Nov 1961, Munoz, McCart, and
Cabrera 39 aa TEX).
36 BRIT.ORG/SIDA 20(1)
Eriogonum multiflorum and E. riograndis can be distinguished by a simple vi-
sual inspection relying primarily on leaf morphology. A count of flowers per
involucre, usually requiring low magnification, confirms their distinction.
Morphological differences between the two are summarized in the following
contrast.
. Leaves oblong-obovate to oblong-elliptic or oblong,
5-4(-4.5) cm long, 10-22(
25) mm wide, auriculate-subclasping at the base, upper surfaces persistently
mentose to weakly glabrescent, usually grayish; flowers 22—28(— 36) pet nvohit
ee onum riograndis
. Leaves mo tly narrow! ellipti ~--lanceolate to narrowly lanc eolte 2s 5( oe cm lonc Ng,
3—9(-15) mm wide
e, attenuate at the base, rarely rounded, but not auriculate-
subclasping, upper surfaces glabrescent to glabrate, commonly reddish; flowers
(6-)9-12(-16) per involucre _ riogonum multiflorum
These two taxa are similar in habit and overall morphology, and they appar-
ently
>
nave a sister relationship. Eriogonum riograndis is similar to E.
multiflorum in its annual duration, lealy and tomentose stems, cymose inflo-
rescence with pedunculate involucres, stipitate perianth internally pubescent
but externally glabrous, and dissimilar perianth lobes (outer oblong-cordate,
inner linear). They have similar phenologies and both occur in sandy habitats.
Asa pair, these taxa differ from E. annuum in their oblong-cordate outer peri-
anth lobes (vs. obovate and basally attenuate) and sparsely tomentose, com-
monly glabrescent involucres (vs. densely and a rsistently tomentose). The in-
volucres of E.annuumalsoare on shorter peduncles (subsessile to 1-3 mm long),
O
{0
=
ten giving the inflorescence branches a somewhat secund appearance, and
each involucre produces 25 or more flowers (similar to those of E. riograndis).
Eriogonum riograndis and E. multiflorum are primarily allopatric in geo-
graphic range (Fig, 2), but E. multiflorum has been documented within the range
of E. riograndis.
Zapata Co. E. riograndisand E. multiflorum both have been collected here.
Plants of the latter (Hamby 1711, TEX)are typical in leaf morphology and [lower
number, as are plants of E. riograndis (citation above)
A few plants with apparently intermediate features have been collected
(Fig. 2).
Dimmit Co. (18 mi EF of El Indio, Miller et al. 5775, TAES): leaves narrow,
sessile but f lowers 36 per involucre. This plant has leaves of E. multiflorum but
a high number of flowers like E. riograndis. Hoglund s.n. (Carrizo Springs, TEX)
has narrow, sessile leaves but 20-21 flowers per involucre
Karnes Co. (2 mi Sof Karnes City, Johnson 1010, LL, SMU, TAES): leaves rela-
tively narrow and strongly glabrescent but those on the upper 2/3 of the stem
basally truncate to subauriculate and subclasping; involucres 18-27-flowered
(tending toward intermediate in flower number). These plants are identified
here as E. riograndis. Another from Karnes Co. Johnson 1618, SMU) is [orally
=
—
NESOM, A NEW SPECIES OF ERIOGONUM FROM TEXAS 37
immature but similar in leaf morphology to Johnson 1010, but the tendency for
smaller number of flowers and the sessile lower leaves indicate that genes from
E. multiflorum may be present.
Wilson Co. (Kicaster, Parks Rx 3016, TEX): upper leaves clasping to
subclasping, flowers ca. 18-22. This plant is similar to those of Johnson 1010 in
Karnes Co., apparently intermediate. Parks and Cory 11800 (E of Floresville,
TAES) has narrow sessile leaves but 24 flowers. Parks 5355 (Kicaster school,
TAES) is typical E. multiflorum.
Choice of rank.—Conspicuous morphological and geographic discontinuity
between Eriogonum multiflorum and E. riograndis is the predominant feature
of their contrast. Putative morphological intermediates suggest that hybrid-
ization may occur where they come into geographically close contact, but no
zone of intergradation exists to suggest that significant gene flow occurs be-
tween the two taxa. It might be argued that their apparent sister relationship
should be recognized by treating them as varieties within a single species, but
the degree of morphological differentiation between species in any given ge-
nus (including Friogonum) is variable, and the discontinuity documented here
(with inference of genetic isolation) provides justification for treating E.
multiflorum and E. riograndisat specific rank, consistent with species concepts
in many other genera.
ACKNOWLEDGMENTS
Iam grateful to the staff of TEX-LL for their help during a recent visit there,
TAES for a loan of specimens, Emily Wood (GH) for sending photocopies of
type material of E. multiflorum, Amy Buthod at OKL for helpful information
on Oklahoma distribution, staffs of CAMU and OKLA for sending photocopies
of Oklahoma collections, and to Jim Reveal for comments on the manuscript.
Linny Heagy provided the fine illustration; a discussion with Ted Barkley re-
garding the choice of epithet was helpful.
REFERENCES
MacRoserts, D.T. 1989.A documented checklist and atlas of the vascular flora of Louisiana.
Louisiana State Univ. Shreveport, Bull. Mus. Life Sci. No. 9.
Nesom, G.L. 2001.Pseudognaphalium austrotexanum (Asteraceae: Gnaphalieae), a new spe-
cies from southeastern Texas and adjacent Mexico. Sida 19:507-511.
ReveAL, J.L. 1968. Notes on the Texas Eriogonums. Sida 3:195-205.
Reveal, J.L. 1969. The subgeneric concept in Eriogonum (Polygonaceae). In: J. Gunckel, ed.
Current topics in plant science. Academic Press, New York. Pp. 229-249.
Reveal, J.L. 1970. Eriogonum. In: D.S. Correll and M.C. Johnston. Manual of the vascular plants
of Texas. Texas Research Foundation, Renner. Pp.510-516.
RICHARDSON, A. 1995. Plants of the Rio Grande Delta. Univ. of Texas Press, Austin.
38 BRIT.ORG/SIDA 20(1)
Situ, E.B. 1988. An atlas and annotated list of the vascular plants of Arkansas. Dept. of
Botany and Bacteriology, Univ. of Arkansas, Fayetteville.
Sorrit, B.A. and A.S.Weaktey.2001. Coastal plain vascular plant endemics: Phytogeographic
patterns. Castanea 66:50-82.
A NEW VARIETY OF IPOMOEA COSTELLATA
(CONVOLVULACEAE) FROM THE
EDWARDS PLATEAU REGION OF TEXAS
Robert J.O’Kennon and Guy L.Nesom
Botanical Research Institute of Texas
509 Pecan Street
Fort Worth, 1X 76102-4060, U.S.A.
ABSTRACT
Ipomoea costellata var. edwardsensis, var. nov., is described, illustrated, and mapped. It apparently is
endemic to the Edwards Plateau region of Texas, where it is known from six counties. It is set apart
from I. costellata elsewhere in its range by its combination of shorter peduncles and bright white,
nearly rotate flowers with fewer and deeper lobes.
RESUMEN
Se describe, se ilustt Ly fi tellata vat. upalastiale var. nov. Es be gra
Ca |
ndémica de | ion de | t paaids de Texas, d e separa
de I. costellata Sor su combinacion de pedtinculos mas cortos y flores blanco brillante, casi rodadas
con menos ld6bulos y mas profundos
Plants of a small-flowered, pedatisect-leaved morning glory from the Edwards
Plateau region of Texas are identified as Ipomoea costellata Torrey but are dis-
tinct from other plants of the species in morphology, phenology, and geogra-
phy. The Edwards Plateau plants are here recognized as a distinct variety.
Ipomoea costellata Torrey var. edwardsensis O’Kennon & Nesom, var. nov. (Figs.
. TyPe: U.S.A. TEXAS. TRAVIS CO: Colorado River, Montopolis Bridge, 8 Nov
1934, B.C. Tharp s.n. (HOLOTYPE: TEX).
lis brevioribues |]
Differt a I. costellatae Torrey sensu la
corollis albis rotatis lobis paucioribus profundioribus. . .
Plants annual herbs from a filiform taproot, at first erect, becoming prostrate or
clambering on low vegetation, slightly twining at the very tips. Stems simple or
with 1-5 (or more) branches originating ca. 2-4 cm above the base, terete, 3-60
cm long, 0.5-1 mm in diameter, branching, green, glabrous. Leaves pedatisect,
2-3.5 cm wide, petioles 7-25 mm long, nearly even in length or decreasing
slightly from base to apex of stem, glabrous, ultimate leaf segments 7-9(-1),
linear to linear-lanceolate, unequal, the longest 15-24 cm long, 1.5-3 mm wide,
outer shorter than the inner and usually lobed from near the base, green above
and beneath, glabrous, margins entire, sparsely hispid-ciliate, apices acute, api-
culate. Flowers solitary or less commonly paired; peduncle spore to sHeay
decurved or ascending, terete, filiform, 3-22 mm long, green, g parsely
SIDA 20(1): 39 - 45. 2002
40 BRIT.ORG/SIDA 20(1)
a
\ GF”
mA
YAY
\i
i
aoe
es if] | ie Hi fyi ef pars J 4 ef 4 NL O1TIC\
Fic. 1 ( )
€] Hat
edwardsensis
=
Fig. 2. Photographs of Ip
flowers (from Nesom FW99 and O’Kennon
O’KENNON AND NESOM, A NEW VARIETY OF IPOMOEA COSTELLATA FROM TEXAS 41
hispid-pilose; pedicel 6-8 mm long, slightly thicker than the peduncle, sepa-
rated from the peduncle by a pair of awl-shaped bracts, glabrous to sparsely
hirsute-pilose; sepals equal, subimbricate, elliptic to elliptic-lanceolate, 4-5.2
mm long, 1.5-2 mm wide, midrib green, low-carinate, smooth to muricate, mar-
gins entire, hyaline, the apex rounded to retuse, apiculate; corolla tubular to
long-campanulate, all parts bright white, glabrous, 8-11 mm long, tube 4-5 mm
long, 2mm in diameter, limb with 5 oblong-ovate, apiculate, spreading lobes
2-2.5mm long, flowers opening with lobes nearly erect, spreading at right angles
at full anthesis, closing erect past maturity; stamens subequal, white, filaments
pubescent along whole length with long, viscid trichomes; style white, 4.5 mm
long (equaling the filament length), glabrous, stigma white, slightly bilobate,
apparently in contact with the anthers. Fruits bilocular dehiscent capsules 5-6
mm in diameter, tan upon drying, chartaceous, glabrous; seeds 4, dark brown
to blackish, obovoid, 3-angled,ca. 3mm long, 2 mm wide, densely and minutely
strigose.
Additional collections examined. United States. Texas. Bexar Co.: Government Canyon State Natu-
ral Area, Eside of Wildcat mien E edge of San Geronimo on ad or W edge of Helotes Quad, ‘guess-
timated’ lat/long—29° 33'5 98° 45'00" W, elev. 1200-1240 ft, rare, two area seen among sparse
herbaceous vegetation in very mote dark brown clay in natural openin n Cretaceous lime tone
bedrock exposed on flattest part of ridgetop, associates include ee vaginatus, Bothriochloa
ischaemum var. songarica, Croton monanthogynus, Senna lindheimeriana, 24 Oct 1995, Carr 15050
(TEX). Burnet Co.: N of RM Road 1431, 1.8 road mi WNW of US Rte 281, Marble Falls Quad, local in
thin, fairly dry, organic-sandy soil in unshaded weather pit on low granite outcrop, ca. 1500 ft elev.
annual, stems twining only at tips of taller plants, 18 Aug 1988, Carr 9136 with Kutac, Lynch, and
Brown (TEX); RM Road 1431, L7 mi W of jct with US Rte 281 in Marble Falls, area of exposed granite,
N side of road across from large quarry, 23 Oct 2001, Nesom FW99 and O’Kennon (BRIT, TEX). Gillespie
Co.: Onion Creek Bluff, 400 m E of FM 783 in NW Gillespie Co., scarce in limestone crevices 10 m
above Onion Creek, 19 Oct 1990, O’K n 8125 (BRIT). Llano Co.: near the summit of Dutch Moun-
EX
tain, ca. 1.5 mi N of Enchanted Rock, ae vine, 29 Sep 1976, Butterwick and Lamb 3303 (TEN).
Travis Co.: Austin, Colorado River below Dam, 29 Sep 1929, Ecology Class s.n. (TEX); Austin, Onion
Creek, 15 Oct 1929, Whitehouse W-29-3 (TEX); McKinney Falls State Park, S of Onion Creek at mouth
of Williamson Creek, frequent, locally abundant in thin soil in weather pits on exposed calcareous
bedrock, ca. 100-300 ft, 29 Oct 1985, Carr 7050 (BRIT, TEX). Uvalde Co.: 2.1 mi W of jct of FM 127 and
FM 1049 on FM 127, dry limestone ledge, 2 Nov 1985, Keeney 5371 (BRIT).
Etymology.—The epithet alludes to the location of the plants on the Edwards
Plateau.
Distribution, habitat,and phenology.—Collections at hand indicate that Ipo-
moea costellata var. edwardsensis is distributed over the Edwards Plateau, al-
though the plants are apparently uncommon and inconspicuous. Shinners’
treatment of Texas Ipomoea (1970) noted only that I. costellata occurs in the
trans-Pecos region, although in 1960 he annotated the three early collections
from Travis County (1929 and 1934, cited above) as I. costellata; McDonald (1995)
did not map or cite any collections of I. costellata from the Edwards Plateau. We
suspect that searches for var. edwardsensis will broaden its known distribution
42 BRIT.ORG/SIDA 20(1)
—
but confirm that it is endemic to the Edwards Plateau. Limestone ledges and
crevices, and thin soil of weather pits in bedrock of limestone and granite, lOO-
1500 ft elev. Flowering (August-)September-November. We observe that var.
edwardsensis apparently is particularly sensitive to grazing.
Ipomoea costellata (excluding var. edwardsensis) occurs in Texas in the trans-
Pecos region and in Webb Co. (Fig. 3), continuous with its distribution in the
Mexican states of Chihuahua and Coahuila. It is recorded from all of the north-
ernmost states of Mexico (Sonora, Chihuahua, Coahuila, Nuevo Leon, and
Tamaulipas) except Baja California and continues southward to Guatemala (see
McDonald 1995, Fig. 2). Austin and Huaman (1996) reported its occurrence also
in Venezuela but not Guatemala.
A recent taxonomic revision (McDonald 1995) treats Ipomoea costellata and
seven closely related species within Ipomoea sect. Leptocallis (G. Don) J.A. Mc-
Donald, and another species has been recently added (Austin & Tapia Munoz
2001). Ipomoea costellata is distinct within sect. Leptocallis in its annual,
taprooted habit and small flowers (ca. | cm long). Ipomoea costellata alone in
the section is autogamous—self-pollination is facilitated by the stigma in con-
—
tact with stamens. Variety edwardsensis also has this reproductive arrangement
and is obviously closely related to the rest of the species, but its geographic sepa-
ration and set of differences (couplet below) provide a reasonable basis lor for-
mal taxonomic recognition.
Peduncles 3-22(-30) mm long, peduncle plus pedicel mostly shorter than or equal to
the subtending leaf, ca. equal the petiole length; corollas bright white, with five,
blong-ovate lobes; flowering (Aug-)Sep-Nov Ipomoea costellata var. edwardsensis
Peduncles (15-)27-70 mm long, peduncle plus pedicel mostly 1.5—3 times longer than
the subtending leaf; corollas pink, purple, bluish, white, or yellow, with ten, shal-
lowly rounded lobes; flowering un—)Jul-Oct __ Ipomoea costellata var. costellata
oO
CO >
Ipomoea flowers are fragile and commonly are not useful for taxonomic dis-
tinctions in pressed specimens. We emphasize the short peduncle length, espe-
cially of mature fruits, as the most obvious and easily observed feature to dis-
tinguish var. edwardsensis. Scattered plants of L costellata sensu lato (var.
costellata) may have peduncles approaching the short length of var.
edwardsensis, but the latter (as a population system) is set apart from rest of
the species by this feature.
The flower color of Ipomoea costellata was described by McDonald (1995)
as “blue, or rarely yellow throughout.” Other sources note the following: “pale
lavender” (Texas— Whitehouse 17154, SMU), “pale pink” (Texas—Turner et al.
53447, SMU), “reddish” (Texas—Keough 227, TEX), “purple” (Texas—Butterwick
and Lamb 1759, TEX), “rose-purple” (New Mexico—Martin & Hutchins 1981),
lavender (photo of Texas plant—Rickett 1969), “limb pale pink, throat and tube
near white” (New Mexico—Spellenberg 3852, LL), “lavender or purplish, tube
paler” (Sonora—Wiggins 1964), “tube lavender, purple near tips, yellow-white
O’KENNON AND NESOM, A NEW VARIETY OF IPOMOEA COSTELLATA FROM TEXAS 43
@ Ipomoea edwardsensis
© Ipomoea costellata
Fic. 3. Distribution of Ipomoea costellata var. edwardsensis and |. costellata var. costellata in Texas. The range of var.
costellata continues westward to Arizona and southward to Guatemala (see text). Records are from BRIT-SMU and
TEX-LL.
at base” (Chihuahua—Henrickson 7557, TEX), “yellow tube, purple-red limb”
(Chihuahua—Henrickson 7692, TEX), “cream with green-yellow throat”
(Coahuila— Wendt 1783, TEX), and “pale blue” (Chiapas—Breedlove 52353, TEX).
Plants in the vicinity of Edo. Mexico produce yellow flowers (according to Mc-
Donald) and have been segregated as I. painteri House—this taxon was included
in the synonymy of I. costellata by McDonald but treated as distinct by Rico R.
(1985), who described the flower color of I. painteri as white or creamy. Plants
in Nuevo Leon and Tamaulipas also apparently produce white flowers and are
further distinguished within the species by thick taproots and apparently pe-
rennial duration; peduncles of these plants are long.
The corolla morphology of var. edwardsensis also is distinctive from any
we are aware of within Ipomoea costellata and certainly from populations of
the species on the periphery of var. edwardsensis in Texas and northern Mexico.
44 BRIT.ORG/SIDA 20(1)
In var. edwardsensis, the limb is deeply and relatively narrowly lobed, and at
full anthesis, the lobes spread at nearly right angles to the tube (Fig. 2). The line
drawing (Fig. 1) shows a flower as it appears in early anthesis, just opening. Flow-
ers of L.costellata elsewhere in its range apparently have ten, shallow lobes and
the limb is funnelform, widening gradually toward the apex (Rickett 1969, pl.
99; Warnock 1977, Fig. 6, p. 178)—“corolla campanulate ... | subentire, scant-
ily 10-lobate,” as ee by McDonald (1995, p. 106).
Variety edwardsensis is clearly a ‘satellite’ of the larger and variable I.
costellata sensu lato. Recognition of var. edwardsensis emphasizes its relative
internal consistency and its morphological and geographical distinction from
the rest of the species. Other recognizable geographic variants, particularly as
distinguished by flower color, have been formed within I. costellata, this pro-
cess perhaps quickened by the apparent tendency for autogamy. Formal recog-
nition of the Edwards Plateau populations implies that other geographic seg-
ments of I. costellata might also be justifiably recognized (McDonald pers.
comm.), and we agree with this.
McDonald (1995) did not provide a formal statement of his concept of va-
rietal versus specific rank within sect. Leptocallis, but several species in his treat-
ment include geographically distinct varietal taxa separated by one to several
apparently non-intergrading morphological characters. In contrast, Yatskievych
and Mason (1984) recognized two varieties within I. tenuiloba Torrey (of sect.
Leptocallis) that are morphologically intergrading, this taxonomy accepted by
McDonald. Yatskievych and Mason also observed that I. tenuiloba, I capillacea
(Kunth) G. Don, and L. plummerae A. Gray (including L patens (A. Gray) House)
constitute “a very close-knit species complex.” With further consideration of
the rationale for applying ranks within sect. Leptocallis, we believe that the
Edwards Plateau variety of L costellata eventually may be treated at higher rank.
ACKNOWLEDGMENTS
We are grateful to the staff of TEX-LL for their help during recent visits. An-
drew McDonald and Dan Austin reviewed the manuscript and we are grateful
to Andrew McDonald for a discussion of variation in I. costellata. Linny Heagy
provided the fine illustration.
REFERENCES
Austin, DF. and Z. Huaman. 1996.A synopsis of Ipomoea (Convolvulaceae) in the Americas.
Taxon 45:3-38.
Austin, D.F. and J.L. Tapia Munoz. 2001. Ipomoea sororia (Convolvulaceae), a new species
from Yucatan, Mexico. Sida 19:807-810.
McDonato, AJ. 1995. Revision of Ipomoea section Leptocallis (Convolvulaceae). Harvard
Pap. Bot. 6:97-122.
Maatin, W.C. and C.R. Hurcuins. 1981. A flora of New Mexico. Vol. 2. J.Cramer, Vaduz
O’KENNON AND NESOM, A NEW VARIETY OF IPOMOEA COSTELLATA FROM TEXAS 45
RickeTT, H.W. (ed.).1969.Wild flowers of the United States:Volume 3: Texas (part 2). McGraw-
Hill Book Company, New York.
Rico Ropricuez, L. 1985./pomoea.|n: Rzedowski, J.y G.C.de Rzedowski, eds. Flora fanerogamica
del Valle de México. Diseno Editorial: Myriam Cerda. Pp. 250-256.
SHINNERS, L.H. 1970. Convolvulaceae (excluding Cuscuta).|In:D.S. Correll and M.C. Johnston.
Manual of the vascular plants of Texas. Texas Research Foundation, Renner. Pp. 1241-
1254.
Warnock, B.H. 1977. Wildflowers of the Davis Mountains and Marathon Basin, Texas. Sul
Ross State University, Alpine, Texas.
Wicains, |.L. 1964. Flora of the Sonoran Desert. In: F. Shreve and I.L.Wiggins.Vegetation and
flora of the Sonoran Desert. Stanford Univ. Press, Stanford, CA.
YATSKIEVITCH, G. and C.T. Mason. 1984. A taxonomic study of Ipomoea tenuiloba Torrey
(Convolvulaceae), with notes on related species. Madrono 31:102-108.
46 BRIT.ORG/SIDA 20(1)
BOOK REVIEW
STEPHEN ANDERTON. 2001. Urban Sanctuaries: Peaceful Havens for the City Gardener.
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SIDA 20(1): 46. 6. 2002
SPHAERALCEA CAESPITOSA VAR. WILLIAMSIAE,
VAR. NOV. (MALVACEAE)
Noel H. Holmgren
New York Botanical Garden
Bronx, NY 10458-5126, U.S.A.
ABSTRACT
In Railroad Valley of east-central Nevada isa less densely pubescent, thinner-leaved, and somewhat
smaller-flowered version of Sphaeralcea caespitosa of west-central Utah. This variety is described as
new, S. caespitosa var. williamsiae N.H. Holmgren.
—
Key worps: Malvaceae, Great Basin flora, Sphaeralcea caespitosa var. williamsiae
RESUMEN
En el valle Railroad del centro-este de Nevada esta una version as eis ed ee con
pubescencia menos densa hojas finas, y fl tro-oeste de Utah. Se
una nueva variedad, S. cde spitosd var. wi ‘amisine ste Holmgren.
The genus Sphaeralcea, consisting of about 40 species, is found in temperate
and warm temperate parts of North and South America. It is a difficult genus
taxonomically with poorly delimited species. Frequent hybridization and back-
crossing may be the culprits. In North America there are about 26 species with
the greatest concentration in southwestern United States and northern Mexico.
Arizona should be considered as the center of diversity for the North American
members with about 16 species, and the concentration decreases nearly expo-
nentially with distance in all directions. The Intermountain Region, an area
more than two-and-a-half times larger than Arizona, is home to 12 species with
only three extending north of the region: S. grossulariifolia (Hook. & Arn.) Rydb,,
S.munroana (Douglas ex Lindl.) Spach ex A. Gray, and S. coccinea (Nutt.) Rydb.
The latter two species extend to Canada and S. coccinea reaches the furthest
east for the genus, extending into southern Manitoba, western Minnesota, west-
ern lowa, Kansas, Oklahoma, and western Texas.
Although Kearney’s (1935) revision of the North American species is now
67 years old, it has held up relatively well, which is remarkable for such a com-
plex genus. His concepts of the species are still accepted, but perhaps reluc-
tantly by many flora writers, in the absence of a modern assessment of the all
the North American species. The only modifications come from the seven new
taxa that have since been added, five of which are in the Intermountain Region.
Welsh, in his study of the genus for the Utah Flora (Welsh et al. 1993), added
four of them, S. psoraloides S.L. Welsh (1980), S. janeae (S.L. Welsh) S.L. Welsh
(1980, 1998), S. moorei (S.L. Welsh) S.L. Welsh (Welsh 1980; Atwood & Welsh, in
SIDA 20(1): 47 - 54. 2002
48
BRIT.ORG/SIDA 20(1)
press), and S.grossulat iifolia var. fumariensis S.L. Welsh (Welsh & Atwood 2001);
and a fifth from just south of the Utah border in northwestern Arizona, within
the Intermountain Region, S. gierischii N.D. Atwood & S.L. Welsh (In press).
Two other new additions are species from south of the region, S. polychroma La
Duke (1985) and S. reflexa Fryxell, Valdés-Reyna & Villarreal (991).
In my study of the genus for Intermountain Flora | have found the need to
expand S. caespitosa M. E. Jones to accommodate a disjunct population from
Railroad Valley in Nye County, Nevada, which can be distinguished asa separate
variety. The species is closely related to S.ambigua A. Gray, differing from it most
significantly in its shorter stature. The geographical ranges of the two varieties
of S.caespitosa lie to the north of the northern limits of S.ambigua in Utah and
eastern Nevada and to the east of its most northern limits in western Nevada
and eastern California. A historical scenario for how the two varieties evolved
is not implicit from their geographical setting. The two exist in similar desert-
valley habitats separated by about 160 km of four major north-south oriented
mountain ranges alternating with three relatively broad valleys. Their close
similarity may be indicative of recent evolution from an element that migrated
northward during a warmer Holocene episode, and their subsequent isolation
has been long enough (in this interpretation) for some genetic drift to occur.
The two varieties of S. caespitosa can be distinguished from each other and
from S.ambigua by the characters in the following key.
1. Plants 2.5-8 dm tall; calyx 8-16 mm Ss ie column (3.5-)4—5 mm long;
schizocarp of 8-12(-14) mericarps; seeds m long; southern half of Nevada
and southwestern and northwestern U ss a eae from the distributions of
the 5. caespitosa varieties), southward through southeastern California and
ern Arizona to Baja California
1. Plants small,0.3-2.5 dm tall;calyx 10-18 mm long; staminal columr 1n 3.5-9 mm long;
schizocarp of 11-14 oe seeds 1.9-2.3 1
and central-western Uta
S.ambigua
ym long; east-central Nevada
S. caespitosa
2. Calyx 13-18 mm long; a 16-23 mm ong staminal column 3.5-6 mm long;
leaves thickish, grayish-green, densely pubescen tally with stellate hairs over
lapping and concealing or nearly edie ‘i aren plants 0.3-1.5(-1.7)
dm tall; endemic to the western Utah desert in southwestern Millard County
and adjacent northwestern Beaver County, at 1600-2000 m elevation var. caespitosa
. Calyx 10-14 mm long; petals 15-20 mm long; staminal column 6-9 mm long,
mature leaves relatively thin greenish moderately pubescent vi ith a considerable
amount of surface exposed; plants 0.7-2.5 dm tall; endemic to Railrc
ad
Valley in northeastern Nye County, hoe 1400-1600 m elevation var. williamsiae
Sphaeralcea caespitosa ME. Jones
Perennial herb, 0.3-1.6(-2.5) dim tall, arising from a woody, branched caudex
surmounting a taproot: herbage grayish-green with a dense pubescence or pale
greenish with a moderately dense pubescence, the hairs stellate, the rays of the
hairs spreading in several different directions; stems few to several, ascending
HOLMGREN, SPHAERALCEA CAESPITOSA VAR. WILLIAMSIAE 49
or erect, unbranched; leaves all cauline, petiolate, the petioles 1-3.5(-7) cm long,
the blades of the leaves at midstem larger than the lower, 1.5-4.5(-5) cm long
and about as wide, deltate, ovate, or suborbicular, the base cordate, truncate, or
broadly cuneate, the margin coarsely crenate or dentate-crenate, and sometimes
obscurely 3-lobed, palmately veined, the veins prominent beneath and slightly
channeled above; stipules filiform or linear, inflorescence few-flowered, the lower
{lowers racemose or solitary in leaf axils, the upper ones often in thyrsoid clus-
ters, the rachis, pedicels, and calyces pubescent as below; pedicels 3.5-13 mm
long, divaricately ascending, with stipular bracts at the base; involucral bracts
filiform, pale brownish; calyx 10-18 mm long, the lobes lanceolate to ovate, acute,
2/3-3/4 of the calyx length; petals 15-23 mm long, the short claw ciliate, the
blade obovate, reddish-orange (grenadine); staminal column 3.5-9 mm long,
stellate pubescent, bearing anthers at the apex; styles with capitate stigmas;
schizocarp of 11-14 mericarps forming a ring 7.5-8.5 mm in diam., each mericarp
2-seeded, 3-5.5 mm high, 2.2-4.6 mm wide, rounded dorsally and rounded to
obtuse apically, coarsely reticulate on the sides of the indehiscent lower 1/3
and smooth-sided on the upper 2/3, pubescent on back; seeds 1.9-2.3 mm long,
reniform, dark brown to black, minutely puberulent in patches.
Sphaeralcea caespitosa M.E. Jones var. caespitosa (Fig. 1A-D). Sphaeralcea
caespitosa M.E. Jones, Contr. W. Bot. 12:4. 1908. Type: UTAH. BEAVER Co.: “Wa Wa.
west of Frisco, at Dry Station, ... on very poor volcanic soil covering lava with a
shallow coat, and in crevices of ce rocks where there is a little soil,” 6000 ft, Bea-
ver Co., Utah, 25 Jun 1906, M.E. Jones s.n. (HOLOTYPE: POM: isotypes: NY!, US)
Perennial herb 0.3-L7 dm tall; herbage grayish-green, densely pubescent, the
stellate hairs on mature leaves usually overlapping and concealing the leaf sur-
face; leaves with petioles 1-3.3 (-5) cm long, the blades thick, 1.5-4.5 cm long
and about as wide, deltate, ovate, or suborbicular, the base truncate to broadly
cuneate, the margin coarsely crenate and sometimes obscurely 3-lobed; calyx
13-18 mm long; petals 16-23 mm long; staminal column 3.5-6 mm long:
mericarps each 4.5-5.5 mm high and 3.2-4.6 mm wide, rounded apically.
Habitat.—Shallow, gravelly alluvial soils, mainly from the Sevy Dolomite
Formation (Welsh 1993), and also from calcareous formations, often growing
in shadscale, rabbitbrush, or matchweed associations, ranging from (1400) 1600
to 2000 m elevation.
Distribution.—West-central Utah in the valleys and foothills of southwest-
ern Millard County and adjacent Beaver County, in and around the Desert Range
Experimental Station (Fig. 2).
Phenology.—Flowering from late May to June, fruiting from mid June to
August.
Sphaeralcea caespitosa M_E. Jones var. williamsiae N.H. Holmgren, var. nov. (Fig.
1E-H). Type: U.S.A. NEVADA. NYE Co: Railroad Valley, just W of Lockes (Black
50 BRIT.ORG/SIDA 20(1)
3mm
A-D. Spt J * ME] aespitosa.A.Habit, B. | {ck . gt ),C.Caly 4} +lot
Y Mericarp. a H. caespitosa var. wiamside N.H. Holmgren. E. Habit, F. Leaves G.c d bractlets, H. Mericarp
Bobbi A sed on:A,C. B. Maguire 20861, B. (left to right) RC Holmgren 265,B Maguire
20876, 5.L. Welsh & M. Chatterley 19530, D.R.C. Holmgren 265, E, G.5.L. Welsh 20579, F, H.K.H. Thorne & B.T. Welsh 957.
Rock Station), 34.5 air km SW of Currant, 38° 33'13'N, 115° 46'34"W, T8N R55E S15,
4850 ft, 28 May 2001, N.H. Holmgren & PK. Holmgren 14322 (HOLOTYPE: NY};
ISOTYPES: BRY| RENO! UTC).
AS} Ic <a var. caespitosa foliis tenuioribus, modice (non dense) pubescentibus atque
oH breviore, 1O- ve mm (non 13-18 mm) longa differt.
Perennial herb 0.7-2.5 dm tall; herbage greenish, moderately pubescent; leaves
with petioles 1.5-3.7(-7) cm long, the blades relatively thin, 1.5-3.5(-5) cm long,
1.5-3(-4) cm wide, deltate or ovate, the base cordate to truncate, the margin
coarsely toothed or dentate-crenate and sometimes obscurely 3-lobed; calyx 10-
14 mm long; petals 15-20 mm long; staminal column 6-9 mm long; mericarps
3-5.5 mm high, 2.2-3.5 mm wide, obtuse apically.
Habitat Shallow gravelly soils on alluvium and valley fill, in greasewood
and shadscale communities, growing with Artemisia spinescens, Atriplex
HOLMGREN, SPHAERALCEA CAESPITOSA VAR. WILLIAMSIAE 51
|
NEVADA
Millard Co.
Nye Co
Lf
| wy | Beaver Co.
119°W Vv
37°N ——
Sphaeralcea caespitosa
@ var. caespitosa
¥ var. williamsiae
— a
Fic.2.M f N d d Utah, showing the distributi f Spt I it it dS. caespitosa var
} t l
p
williamsiae.
confertifolia, Ephedra viridis, Eriogonum shockleyi, Hilaria jamesii,and Kochia
americana, ranging in elevation from 1400 to 1600 m elevation.
Distribution.—Railroad Valley, northeastern Nye County, Nevada.
Phenology.—Flowering from May to June, fruiting from mid May to July.
Etymology.—It gives me special pleasure to name this lovely desert mallow
in honor of Margaret Jensen Williams (1917-2000). Sphaeralcea ranked among
the top of the many native Nevada genera for which Margaret had great fond-
ness, and it is apparent from her many collections of this taxon that she re-
garded it as being particularly special. As seen by the tributes to her (Tiehm
2000), Margaret touched many lives.
Her sense of adventure lured her to many parts of the world, but she hada
special bond with the Great Basin, especially Nevada. She always lived in Ne-
vada, born in Gardnerville to ranchers, John and Wilhelmina Jensen. She earned
her bachelor’s degree at the University of Nevada, Reno, in chemistry and a
master’s degree in mathematics. She married her Chemistry Professor, Loring
Williams, and together raised to two children. She was a math instructor at the
52 BRIT.ORG/SIDA 20(1)
University until her appointment was terminated by bureaucratic nepotism
rules. From 1959 until retirement in 1981, she taught 4th grade children in a
Reno elementary school.
At home, in the early 1950s, she began to concentrate her enormous energy
on gardening, at first growing the standard nursery-trade plants. As time passed,
she became more and more sophisticated in her choice of garden subjects, es-
pecially alpine rock garden plants. It was not long before she was an active
member of American, English, and Scottish rock garden societies. Her interests
in the Nevada flora were spurred on by the writings in the society journals of
Dwight Ripley and Rupert Barneby on their botanical excursions throughout
the Great Basin. This deepening interest led her to take classes in plant tax-
onomy and field botany. At first she was just collecting seeds for exchange with
rock garden friends in the U.S. and abroad. Then in the early 1970s, John Tho-
mas Howell of the California Academy of Sciences inspired her to collect plant
specimens.
In 1975 she founded the Northern Nevada Native Plant Society, which she
presided over for several years, first as President, then as Executive Director. Her
charisma, strong will, passion for plants, and contagious enthusiasm drew ina
large and active membership, and the Society continues to thrive.
Her legacy lives on through the many research collections filed through-
out many herbaria. Her collections filed at the New York Botanical Garden are
especially valuable in our research on the Intermountain Flora (Cronquist et al.
1972-1997). She was involved in one way or another in collecting 12 specimens
that are now types for species or varieties. Where she was not listed as the prin-
cipal collector, it can be assumed that Margaret was the one who enabled the
collection to be made. This attractive new variety joins Polyctenium williamsiae
Rollins, Eriogonum ovalifolium var. williamsiae Reveal, and Astragalus
convallarius var. margaretae Barneby as honorific eponyms.
Ona personal note, my wile Pat and | willalways treasure our many memo-
ries of Margaret. She supported our Intermountain Flora research in so many
ways, from providing food and lodging in her home, transporting us to and from
the airport and the RENO herbarium, to being a prompt and delightful corre-
spondent.
Paratypes: Nevada. Nye Co.: all in Railroad Valley: low volcanic hills | mi NE of Currant, 23 May 1982,
RC. Barneby 17821 (NY); 20 mi SW of Currant, 14 May 1941, A. Eastwood & J.T. Howell 9426 (US),
UTC); T8N R55E S21 (NW1/4), +900 ft, + Jun 1991, L. Grover s.n. (RENOV;: 50.2 km road distance 5S of
Currant on the valley road, TSN R56EF $3, 38° 19'18"N, 115° 39'38"W, 1495 m, 7 Jun 2000, N.H. Holmgren
& PK. Holmgren 13840 (NY); along U.S. Hwy 6, S side of the highway and at the SW edge of Black
Rock Station (Lockes), T8N R55E S15, 38 3314"N, 115 46°34" W, 1480 m, 7 Jun 2000, N.H. Holmgren &
PK. Holmgren 13845 (BRY!, NY!, RENO! UTC); W side of Grant Range, ca. | road mi NE of Blue Eagle
Spring along road to Currant, T8N R57E SI, 4800 ft, 12 May 1987, T. Knight 1557 (NY); 0.75 mi N of
kes Ranch, 21 May 1945, B. Maguire & A.H. Holmgren 25128 (NY! UTC): U.S. Hwy 6, ca. 20 mi S of
Currant, 30 May 1986, H.C. Stutz 94320 (BRY!); Cherry Creek Road, 1.6 mi to a small knoll N of the
Loc
HOLMGREN, SPHAERALCEA CAESPITOSA VAR. WILLIAMSIAE 53
road, T5N R56E $22, 5200 ft, 2 Jun 1980, K.H. Thorne & B.T. Welsh 942 (BRY!); | mi NE of Currant,
white knoll on N side of road, TION R58E $9, 5250 ft, 3 Jun 1980, K.H. Thorne & B.T. Welsh 957 (BRY!,
NY); ca. 22 mi SW of Currant, T8N R56E S1, 4800 ft, 1 Jul 1980, B.T. Welsh, K.H. Thorne & S.L. Welsh
413 (BRY!, RENO); 1 mi NE of Currant, TION R58E S9 (NW/NW), 5300 ft, Sarcobatus-Hilaria-
Chrysothamnus community, gravelly substrate, 23 May 1981, S.L. Welsh 20579 (BRY!, NY!, RENO);
near Hwy 6, SW of Lockes, T8N R55E 821, 5300 ft, 20 Jun 1980, M. J. Williams 80-164-1(NY!, RENO):
ca. mi NE of Currant, near Hwy 6, TION R58E S4, 5300 ft, 29 Jun 1980, M.J. Williams 80-166-1 (RENOD):
3.4 mi SW of Lockes, NW of Hwy 6, T8N R55E S19, 5300 [t, 9 May 1980, MJ. Williams, S. Cochrane &
A. Tiehm 80-8-3 (RENO}); 3.5 mi SW of Lockes on U.S. Hwy 6, then NW for 1.4 mi, T8N R54E $25, 5350
ft, 9 May 1980, MJ. Williams, S. Cochrane & A. Tiehm 80-9-3 (RENOD); near U.S. Hwy 6, 0.4 mi NE of
Currant, TLON R58E S4, 5150 ft, 26 May 1979, MJ. Williams & J. S. Holland 79-54-7 (BRY!, RENO): US.
Hwy 6, near Mile 119, N of Currant, TION R58E $4, 5300 ft, 23 May 1981, MJ. Williams & A. Tiehm 81-
18-1(RENO!): S of Lockes, 1.6 miS of U.S. Hwy 6, T8N R55E $27, 4770 ft, 23 May 1981, Mj. Williams &
A. Tiehm 81-19-7 (RENOD); ca. 3 mi SW of Lockes, T8N R55E $30, 5300 ft, 28 May 1978, MJ. Williams
& M. Williams 78-75-1 (RENO! UTC).
—
ACKNOWLEDGMENTS
My deepest gratitude goes to Patricia K. Holmgren for her careful reading of the
manuscript, assistance in field work, dealing with collecting permits, and in
procuring loans. lam grateful to Arnold Tiehm for his prompt and informative
responses to my queries, to Bobbi Angell for her expert illustrations in Fig. 1,
and to Patricia Eckel for her rendering of the Latin diagnosis, and to Paul Fryxell
and Steven Hill for their thoughtful and constructive reviews. I thank the cura-
tors of the herbaria at BRY, RENO, US, UT, and UTC for being gracious hosts in
their herbaria and, in some instances, for lending me their specimens. The re-
search for this paper was in conjunction with my ongoing research on the In-
termountain Flora, which has, during my research on the Malvaceae, been sup-
ported by The New York Botanical Garden with additional funding from the
Mary Flagler Cary Charitable Trust, the George S.and Dolores Doré Eccles Foun-
dation, the Harriet Ford Dickenson Foundation, The Andrew W. Mellon Foun-
dation, and Mrs. Nicholas J. Sakellariadis.
REFERENCES
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West, U.S.A. vols. 1, 3a, 3b, 4, 5,6. New York Bot. Garden Press.
Fryxet, PR. A., J. Vatoés Reyna, and J.A. Vittarreal QO. 1991. A new species of Sphaeralcea
(Malvaceae) from Coahuila, Mexico. Southw. Naturalist 36:358-360.
Kearney, |. H. 1935. The North American species of Sphaeralcea subgenus Eusphaeralcea.
Univ. Calif. Publ. Bot. 19(1):1-127.
La Duke, J.C. 1985. A new species of Sphaeralcea (Malvaceae). Southw. Naturalist 30:433-
TicHm, A. 2000. Tribute to Margaret: Margaret Jensen Williams 1917-2000 [followed by
other tributes]. Northern Nevada Native Plant Soc. Newsl. 26(7):1-19.
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Wetsh, S.L. 1980. Utah flora: Malvaceae. Great Basin Naturalist 40:2 7-37.
Weis, S.L. 1993. Malvaceae. In: Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. Higgins. A
Utah flora. Ed. 2. Print Services, Brigham Young University, Provo, Utah. Pp. 473-479.
Weish, S.L. 1998. An undescribed Astragalus (Leguminosae) from southern Utah, a new
subsection of the genus, and validation of the combination Spt cea j (Welsh)
Welsh. Great Basin Naturalist 58:386-389.
Wetsh, S.L. and N.D. Atwoop. 2001. New taxa and nomenclatural proposals in miscella-
neous families-Utah and Arizona. Rhodora 103:71-95.
THE PRESENT STATUS OF LEDERMANN’S APRIL RIVER
LOCALITIES IN PAPUA NEW GUINEA
W. Takeuchi M.Golman
Botanical Research Institute of Texas Divisional Manager, PNG National Forest Authorit ity
HES Ie SOCGG NO MAUR nGHRGTe Service PNG National Forest Ser
La riul 1 PNG Forest Research Institute PO. Box 5055, Boroko —
PAPUA NEW GUINEA
Lae, Morobe Province 411, PAPUA NEW GUINEA
ABSTRACT
The April River of East Sepik Province has received lit ttenti i | l ul f
the early 1900s. However the social, logistical, and vegetation status of April environments present
considerable opportunities for floristic discovery and documentation.
Key Worps: botanical survey, Kaiserin-Augusta Expedition, type localities, Papua New Guinea.
ABSTRACT (JAPANESE)
EDEMP LDA=—FAMBCSPODDSHK, Rey PMOL SY ILUN— thE, KA
CKSRIORRURDT OIC LMEAENTOAMOK. Cy TITERO, BANC MDS
SMIREBET SCH CS. KTRKAMSCLOMECHS, ELC. BHOMECLOTS
bh, CEHKR., FRRMR*ADILOOR* OMMPME CMI SMEBCRERITS, ARERR
pS Gist tHE | OR ARE AEA DNS REESE ICRU CLUS TEMS DMRS,
TOK IGO PAS (MELANESIAN TOK PISIN)
April wara long Is Sepik Provins i bin kisim liklik luksave tasol long taim bilong Geman long 1900
ikam inap long nau. Tasol, bikos long dispela hap, igat bikpela na gutpela bus, diwai, graun na tu laif
bilong ol man na meri ino gat bikpela senis, wei dispela i bringim ol saveman na meri long lukluk
moa long wok painim out igo insait long ol bus na diwai long hap bilong Apri
—
wara,
INTRODUCTION
Of all the exploratory surveys conducted in New Guinea during the last 100
years, the German Kaiserin-Augusta Expedition of 1912-13 stands out as being
particularly significant. During that expedition, botanist Carl Ledermann trav-
eled extensively through the south Sepik basin with a multidisciplinary con-
tingent, eventually collecting a total of 6,639 angiosperm numbers over a pe-
riod of 18-19 months. Originating at an early point in the exploration of the
Papuasian flora, a disproportionate number of Ledermann’s collections were
designated as type specimens. yea) of his localities have never been revisited.
The unfortunate circumstances of Ledermann’s labors are an enormous obstacle
to modern scientific inquiry, for the main German sets were destroyed in the
SIDA 20(1): 55 — 70. 2002
56 BRIT.ORG/SIDA 20(1)
1943 fire at Berlin Herbarium, effectively erasing the primary basis for the iden-
tification of numerous plant species. Veldkamp et al. (1988) have provided an
informative discussion of the relevant issues surrounding the loss of such a criti-
cal botanical corpus.
One of the most productive parts of the Kaiserin-Augusta itinerary was
the series of collecting trips made by Ledermann to the April River (Aprilfluss).
The Aprilfluss yielded a bountiful harvest of new species, many of which have
unfortunately proven impossible to evaluate without the original collections.
With modern assessment indicating the apparent existence of a zone of floris-
tic endemism centered on the Hunstein-April region, it has become particu-
larly desirable to revisit the classical localities and secure new material. Such
efforts would enable neotypification of many species whose types were lost at
Berlin (e.g., Tapeinochilos hollrungii K. Schum. cf. Gideon 1998). Although a
list of plants requiring new collections is reportedly being compiled (Veldkamp
et al. 1988), no summaries are yet available.
The first serious reexamination of Ledermann’s sites occurred in 1966, with
the CSIRO-sponsored expedition to Mt Hunstein by R. Hoogland and L. Cra-
ven. In 1989a multi-institutional survey returned to the Hunstein Range under
the auspices of the National Geographic Society, obtaining 1,200 more botani-
cal numbers (cf. Bakker 1994 fora general account). The first author, one of five
botanists on that expedition, independently returned to the Hunstein summit
in July-August 1990 and to the nearby Waskuk Hills in August 1994 and June
1995. Additional collections from the April-Sitipa drainages were made in Sep-
tember 1990, and from the April River serpentine zone on the south side of Mt
Hunstein, in July 1995 (Fig. 1). The most recent trip to the subdistrict occurred
in August 2001 (cf. Takeuchi &@ Golman 2002; Takeuchi & Renner 2002), from
which much of the information in the following narrative has been taken.
—
THE SOCIAL ENVIRONMENT
The indigenous population of the Hunstein subdistrict is spread diffusely
through the region in small clan-based villages. Ambunti is the largest settle-
ment: with a district headquarters, tradestores, an aid ee two schools, several
churches, and a grass airstrip. Further upriver and within the 24 ae Suna
the average settlements are much smaller, often consisting of as
family. At Garuka (Waskuk Hills) for example, the total adult sence was
exactly ten during the time of the 1995 survey. The larger villages like Baglam
and Melawei have about a hundred inhabitants.
—
In the Sepik region, river-borne traffic is the major means of communica-
tion, trade, and transport. Most of the population are thus concentrated near
the water. This also affords the people convenient access to fish and saksak (made
from the starchy pith of the Metroxylon palm) which together comprise the
bulk of the average villager’s diet. A limited amount of subsistence slash and
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58 BRIT.ORG/SIDA 20
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burn gardening provides dietary variety. Sweet potatoes, taro, bananas, and
coconuts constitute the main crops, but plantings are generally not a principal
source of village sustenance. For the most part, the April River populace are
semi-nomadic hunter-gatherers.
Like the Sepik River, the April is a meandering silt-laden waterway which
often changes course after severe floods. A recently introduced fish from South
America (the pacu, erroneously identified as a piranha in the local media) has
now entered the April drainage and is rapidly becoming the most commonly
caught fish for consumption. Villagers report that the appearance and prolifera-
tion of the pacu have been accompanied by disappearance of many native fish
species, particularly the endemic species of Sepik catfish (Arius spp, cf. Allen
1991). The pacu has reportedly been involved in several fatal attacks on bathers,
but such anecdotal reports are often exaggerated and their veracity is uncertain.
Socioeconomic development in the April region is virtually nonexistent.
The local economy is primarily subsistence in nature, and cash incomes are
opportunistic. Money is occasionally earned from the sale of wood carvings,
crocodile skins, birdwing butterflies, cacao, and from seasonal sources in the
ecotourism niche market. Very recently, a lucrative trade has also emerged in-
volving Gyrinops ledermannii Domke (Thymelaeaceae), with Indonesian buy-
ers paying up to PNGK 800-1,000 (USD 265-335) for each kg of black wood.
The arborescent species is endemic to the upper Sepik (Ding Hou 1960), and
has become a significant income source for landowners due to consumer de-
mand for the wood’s pleasing fragrance, a development reminiscent of the san-
dalwood trade. Known as ‘may-hasei’ in the Parenemo language, G. ledermannii
is common on the southwest side of Kamelsrticken and is also distributed on
slopes facing the April River, particularly near Etappenberg (Takeuchi & Gol-
man 2002). For many decades the species was known only by Ledermann’s type
gathering from the May River, but current indications suggest the tree prob-
ably ranges even into West Papua (ibid.).
In spite of income from such incidental sources, by Western standards the
overall status of the people would be regarded as impoverished. Due to the in-
accessibility of health services, mortality rates are high, especially from ma-
laria and infections, and the local inhabitants frequently seek palliatives and
remedies from whatever natural resources are at hand. Skin diseases like ring-
worm and scabies often affect entire villages.
April River culture is an improbable assortment of incongruities and ap-
parent contradictions. Villagers have adopted western standards of dress, yet
still retain their traditional lifestyle and customs. Many of the residents speak
standard English, often freely intermixed with their tribal tokples (Baihenemo,
Parenemo, Waskukil) in ad hoc combination. Although Christianity in its many
variants is dominant throughout the region, belief in sorcery is closely inter-
woven through the Christian ethic. Sudden deaths are nearly always attributed
59
to black magic or to secret murders (sanguma) facilitated by plant poisons.
‘Ukhop pok’ (Anodendron oblongifolium Hemsl; Apocynaceae) is the most com-
monly mentioned agent in sanguma killings. There is also a widespread con-
viction in the existence of spirits (masalai), believed to pervade all aspects of
the natural world and which form an inseparable part of the villagers’ meta-
physical world-view.
The population growth rate for East Sepik Province is currently 2.9% per
annum, a relatively high figure, but below the Mamose! regional average of 3.3%
(National Statistical Office 2001). Most of the provincial population is concen-
trated in the built-up areas near the coast, and the interior sections such as April
River consist primarily of wilderness habitat. Due to the low population densi-
ties in this part of PNG, there are no subsistence-based threats to the environ-
ment. However, because the entire region along the April-Salumei is one of PNG’s
most economically depressed areas, landowners are understandably attracted by
the inducements offered by logging and mining interests. A major portion of the
district has been proposed as a timber concessional area under the Forest Man-
agement Agreement protocols (cf. Papua New Guinea Forest Authority 1998).
Reconciling the conflicting desire for socioeconomic development on one hand,
with conservation concerns on the other, will be a major challenge for commu-
nity planners. Many of the relevant issues are presently being addressed by the
World Wildlife Fund (WWF), through the establishment of an ICAD (Integrated
Conservation and Development) program in the Hunstein area and the related
demarcation of a Wildlife Management Area (WMA; Fig. 1). The Village Devel-
opment Trust is also contemplating start-up of a comparable program in the
Bugabugi territory and the Waskuk Hills (Aung & Kisokau pers. comm.). In the
past, investigators moved freely throughout the area, subject only to approval
from the customary landowners. With the recent and impending developments,
this is likely to change. Intervention by nongovernment organizations (NGOs)
invariably results in new requirements which can complicate scientific access
and research. The complications are offset however, by the real and perceived
benefits which NGO association can deliver to affected communities.
Land ownership and the subsequent control of access to forest resources
are traditional and patrilineal. Nearly all territories fall under clan jurisdiction,
so discretion must be exercised during plant collecting in order not to trespass
on land unapproved for gathering. A gratuity paid to the landowner(s) is often
necessary to ensure approval for plant collecting.
THE LOGISTICAL ENVIRONMENT
In contrast to the many obstacles which confronted the first explorers, access to
the upper Sepik has improved to such an extent that the region is now a major
'Mamose is the administrative unit composed of West Sepik, East Sepik, Madang, and Morobe Provinces.
60 BRIT.ORG/SIDA 20(1)
ecotourist destination. Until recently, ocean-going tourist vessels made regu-
larly-scheduled luxury cruises as far upstream as Ambunti. Although travelers
can still expect numerous delays arising from conditions endemic to develop-
ing countries, entry to the April basin has been considerably eased by the new
airfields at Bugabugi and Niksak.* Both airfields are small grass strips main-
tained by New Tribes Mission, and can accept only the single-engine Cessna
206, wit
can also be made via the airfield at Ambunti, which can accommodate larger
—
h maximum lift capacity of 5 passengers or 400 kg. Otherwise access
twin-engined Islanders capable of carrying 19 passengers or 1,600 kgs of freight
(Missionary Air Fellowship). From Ambuntia visitor would ordinarily proceed
to the April River with dugout canoes powered by Yamaha outboard motors
(15, 30, or 40 hp motors are the usual ratings). The fuel consumption of a 30 hp
motor ona roundtrip between Ambunti and Bugabugi will be at least 140 li-
ters. In the upper Sepik, fuel is problematic, being either unavailable or prohibi-
tively expensive.
The buttress ridges in the vicinity of Bugabugi (Pinape, Natawe, and
Okahsa), lead directly to Mt Hunstein and represent the easiest path to the top.
In 1989, the National Geographic expedition approached the summit from the
north, which is actually the longest and most logistically difficult land route.
Ledermann’s two trips to Hunsteinspitze had come the same way, from Ambunti
and through Wasui-Wagu Lagoon (cf. Steenis-Kruseman 1950: 317-18). The
entry via Wagu was similarly taken by Hoogland and Craven in 1966 (Craven
pers. comm.). The quickest access however, is to follow the south-descending
ridges from the April River, preferably starting at Natawe (Fig. 2). With the newly
operational airfield at Bugabugi, the latter route can now be used to reach the
Hunstein crestline within two days of departure from the provincial seat at
Wewak. The south-side approach provides the added advantage of passage
through unexplored ultrabasic habitats.
Field conditions in the April drainage are onerous. The heat and humidity
are stultifying during dry periods and the wet season is accompanied by tor-
rential downpours after which the river is hazardous at many places along its
course. Prolonged visits are a health risk. The physical difficulties are aggra-
vated by the absence of any service infrastructure throughout the region, ex-
cept for basic facilities at missionary stations. Over much of the year the mos-
quito and leech populations are exceptionally troublesome even by the
standards of an alluvial tropical environment. When it is not raining, horse-
flies are annoyingly plentiful, especially in areas where pigs are present in the
forest. Poisonous snakes have been responsible for a substantial number of
deaths in the village communities
the Ra
the enigmatic localities from Ledermann’s Aprilfluss itine
Niksak is 18 | th of Lordberg
ry, and ld be the only cost-effective point of access in any future attempt at revisiting the latter site.
Fic. 2 Phot hi £ . N
~ bined) J J*t
A: Okahsa, tthao |t £ Ea nthill fehLnu “Do Fore-
J
4 £thoe Anvil Bi R- Ct
(on slopes behind the ridgeline). C: the distinctive double
peaks of Kamelsriicken (the high peak is the Mt Samsai of
modern maps). Ledermann’s Bambusberg and Lordherg are
hohind th
ina direction ca halfway
between Okahsa and Etappenberg.
62 BRIT.ORG/SIDA 20(1)
THE FLORISTIC ENVIRONMENT
The vegetation along the April River consists of three principal formations. The
swamp forest (with several facies including levee forest, meander scrolls, swales,
etc.) is the dominant community on the flood plain. Typically waterlogged or
inundated during rainy periods, the swampy habitat is dominated by Metroxylon
sagu Rottb., Campnosperma brevipetiolata Volk., or herbaceous vegetation (espe-
cially Saccharum robustum Brandes & Jeswiet ex Grassl: cf. Warner & Grassl 1958).
On better drained alluvium the forest is more structurally developed, giving rise
toa second association with many macrophyllous taxa (e.g, Artocarpus commu-
nis J.R. & G. Forst., Caryota rumphiana Mart, Pangium edule Reinw., Sterculia
macrophylla Vent., and Nauclea orientalis L.). These communities have charac-
teristically high proportions of robust climbers along the edges, particularly Ca-
lamus, Korthalsia, and to a lesser extent Freycinetia, and the interior sections
often support a luxuriant shrub layer. On the Hammermaster and Saunders
(1995a, b) forest classification system, the riverine vegetation is described as an
open forest on low elevation plains and fans, or as swamp woodland (forest code
‘Po’ and ‘Ws’ on overlays SB 54-4 and SB 54-3 to the Ambunti and Mianmin
1:250,000 Australian Survey Corps topographic sheets). Paijmans (1976) provides
a concise overview of the vegetation in such environments.
At several places along its course, the April River skirts the base of serpen-
tine/ultramatic foothills and there is a sharp floristic break as the swamp and
alluvial associations are replaced by foothill forest. The contrasting features of
the forest facies on ultramafics are: 1) a pronounced reduction in climbers (e.g,,
Calamus is virtually absent), 2) a more varied canopy structure, including a
noticeable reduction in large-leaved taxa, 3) increased floristic richness, most
apparent inan cae w moe is not subject to the periodic flooding charac-
teristic of the alluvial flats, 4) a ‘clean’ visual aspect to the community, with
good visibility through the forest because of the absence of climbers and com-
parative lack of epiphytic growth on tree boles, 5) a very obvious change in
taxonomic composition, many taxa being common on the ultramafic slopes
but absent from immediately adjacent alluvial forest. When observed from the
outside, the ultramafic community is similar in physiognomy and structure to
a montane forest, except for the paucity of epiphytes. Based on air photo inter-
pretation, the April foothill zone is a medium-crowned low altitude upland for-
est (code ‘Hm’ in Hammermaster & Saunders 1995a), a vegetation type which
is probably Papuasia’s richest forest structural category (Louman & Nicholls
1995). Although an unpublished report (Sohmer et al. 1991) provides a brief
botanical description of the Hunstein Range, there is still nothing in the way of
a comprehensive account for the April basin.
The forest on ultramafic foothills is easily accessed near Bugabugi? (Pinape,
Natawe and Okahsa: in sequential order upriver), though the best section is situ-
=,
—
63
ated at the base of the Hunstein Range between Gahom and Okahsa, close to
the junction of the April and Sitipa rivers (Figs. 3, 4). A conspicuous feature of
the hill community is the presence of Agathis labillardieri Warb.emergents on
the ridge buttresses and crestlines. Agathis attains heights exceeding 50 m, and
the April provenances can be seen from the river towering over the surround-
ing canopy. The populations are of commercial significance, so not unexpect-
edly, has attracted the attention of logging and ecoforestry operators. Due to
the steep and rocky slopes, Bugabugi villagers do not establish subsistence gar-
dens in the foothill habitat, and the slopes are thus remarkably free of human
disturbance. However, many understories near the river are being disrupted by
domestic and wild pigs. Large seral gaps are also scattered through the forest as
a result of natural disturbances from landslides, lightning strikes, and
windthrows. For a lowland environment, the frequency of lightning-induced
gaps is surprisingly high, an indication of the intense electrical and thunder-
storm activity in the Hunstein Range.
During the 1998 el Nino disturbance, the Hunstein subdistrict and the en-
tire basin to the south was severely affected by drought. The Sepik became dry
in many sections even though it represents the largest catchment system among
PNG’s rivers. At Yambon gate, the Sepik current normally accelerates through
a narrow channel renowned for treacherous whirlpools (cf. Townsend 1968),
yet villagers were able to walk across to the opposite bank during the 1998
drought. In spite of such conditions, and unlike many other forested sites in
PNG, April River environments did not experience any fires during the last dis-
turbance. However a large tract downstream of Bitara has been recently flat-
tened by cyclonic-force winds, and the forest canopy in a swath ca. | km wide
completely removed. From current events, it is apparent that vegetational his-
tories in this basin are at least partly determined by catastrophic events (cf.
Johns 1986).
Since the time of its initial exploration, the Hunstein subdistrict has been
judged a particularly significant hotspot for diversification and endemism. The
biodiversity status of the area has assumed legendary proportions, even though
the perception of unusual richness can be regarded as a natural outcome of
physical and historical considerations.
The geological complexity of the April-Hunstein territory is a major con-
tributing factor in the diversification of its flora. Alluvium, colluvium, sand-
stone, shale, mafic/ultramafic metamorphics, igneous, clay, and coralline lime
*Bugabugi is 1 2 km NE of the classical locality of Etappenberg, the latter representing one of Ledermann’s
cae sites on Kamelsrucken : -amel Back; on some maps now as Samsai Mt) of the Aprilfluss inal
| pay. There is some inconsistency in the use of the loc
name ‘Samsai’ or Saisaie because the name is Ree by Wagu and some Bugabugi villagers to Mt. eae
(Hunsteinspitze), and is used with that reference, for example, in the Hoogland and Craven collections from 1966,
64 BRIT.ORG/SIDA 20(1)
stone substrates occur throughout the area in patches of varying size (Davies
& Hutchison 1980a, b). There is thus a highly fragmented pattern of contrast-
ing substrates over the terrain, reflecting the mani
old geological origins of the
present territory and capable of supporting specialist taxa in a complex mo-
saic. Superimposed on the foundation of edaphic environments is an excep
tional annual rainfall estimated at 7,000-9,000 mm (ibid.). High rainfalls are
directly related to floristic diversification (Gentry 1988) so perhumid condi-
tions in the April basin have probably played a major role in the development of
a speciose flora. When the dynamic fragmentation of the ecosystem is then
considered, over presently observable scales ranging from lightning strikes to
catastrophic storms, it is not difficult to imagine how a great number of plant
species can be accommodated in side by side fashion over relatively small dis-
tances. The heterogeneity of April habitats also needs to be viewed in terms of
the evidence regarding correlations between plant distributions and phases of
tectonic accretion (cf. Welzen 1997). Part of the district’s reputation as a zone of
floristic peculiarity (Veldkamp et al. 1988) is due to its easternmost position in
the terrane complex which constructed most of northern West Papua (cf,
Pigram & Davies 1987). The Hunstein flora is probably best understood as an
extension of the West Papuan flora. During the time of Davies and Hutchison
(1980b) the Hunstein-April-Salumei petrology was already known to extend
TAVCIICUI AR
£41
Interior perspective OT
substrates. Stem densities are high.
Fic. 4.
66 BRIT.ORG/SIDA 20(1)
westwards rather than eastwards. If the division between PNG and West Papua
were to be based on geophysical rather than political criteria, the border would
be at April-Salumei rather than Sandaun.
Based on the relationship to tectonics, future distributional records of
Hunstein/April endemics will occur to the west, as the Papuan affinity clari-
fies through further exploration of West New Guinea (Indonesian West Papua).
At present, the Indonesian side is even more inadequately surveyed than PNG,
and the historical disparity in documentation between the two halves is partly
responsible for the perception of unusually high endemism in the Hunstein/
April region. The newly discerned disjunction of Myrmephytum in the Vogelkop
and Hunstein areas ebb pers. comm.), of Faika in West Papua and the April
River (Takeuchi & Renner 2002), and the probable occurrence of Gyrinops
ledermannii in Jayapura (Takeuchi & Golman 2002) are suggestive of the phy-
togeographic connections, and of the effect which historical patterns in floris-
tic documentation have on assumed distributions (Figs. 5, 6).
5
DISCUSSION
Nearly 90 years after the Kaiserin Augusta Expedition, many of the environ-
ments along the April River are still unexplored. The localities at the
Expedition’s farthest penetration into the south basin, near the headwaters of
the Bamali tributary, have never been revisited. Of these, Lordberg is probably
the most intriguing, but any effort focused on the Divide south of Niksak will
undoubtedly reap huge rewards. The recent discoveries from Bugabugi exem-
plify the possibilities. Although the 6,639 collections taken by Ledermann rep-
resent an unprecedented achievement for his time, even a personal effort of that
intensity cannot provide the desired level of sampling saturation for sucha flo-
ristically rich region. A collections program delivering at least 10,000 new num-
bers is much needed.
The factors militating in favor of future survey success in the April drain-
age can be recanted: 1) forest communities in the region are in an exceptional
state of preservation, 2) traditional lifestyles and ethnobotanical competences
are still retained by local villagers, who are thus a valuable source of the folk-
lore knowledge being rapidly lost in other districts, 3) improved site access to
the interior basin will enable adoption of cost-effective logistical schedules, 4)
research conducted by various investigators since 1989, has created local aware-
ness of the work done by scientists, which will facilitate future inventories.
No collections effort conducted under normal constraints, can achieve 100%
coverage of a targeted flora. There is always something which will be overlooked
on account of rarity, phenology, localized distribution, logistical limitations,
etc. Even the collections obtained by Ledermann during his Sepik travels are
not such a superlative effort as may appear at first glance. The 6,600+ numbers
from the Kaiserin Augusta survey amount to approximately 400 numbers per
(det. by $.S. Renner)
& Hatus.) Philipson. in tk
ava L
Fic
68 BRIT.ORG/SIDA 20(1)
Fic. 6.As for figure 5, showing leaves, branchlet 1 fruits (leaf blad 10 cm wide). The April River Faik
J+ AA aes (r€ Tal 1
re a PNG aeneric
~s
month ona time-averaged basis (excluding Ledermann’s 2 months of inactiv-
ity due to sickness). By the productivity standards of contemporary collectors
working under rapid assessment protocols, this is a below-average result. The
significance of the Ledermann collections are another matter altogether how-
ever. And in fairness, the logistical environment of his time was undoubtedly
much more limiting than now. The difficulties of prolonged expeditionary
schedules also should not be underestimated. Anyone with knowledge of
Papuasian forest environments can appreciate the hardships in being exposed
to tropical conditions for the duration which Ledermann was subjected to.
Some indication of the spottiness of Ledermann’s sampling coverage can
be gauged by results from the 1966 and 1989 expeditions to Mt. Hunstein. Sev-
eral scores of new taxa have been described from the later visits, so clearly the
original effort was not very comprehensive. To what extent does that situation
apply to even more remote sites, such as Niksak-Lordberg? On the basis of ac-
cumulating evidence showing that the richest floristic interval in Papuasia Is
the premontane zone (Takeuchi & Golman 2001), the best environments within
the April region are probably the ones at the Central Divide, and these have
never been seen since Ledermann’s time. The diversity potential of this flora
has yet to be plumbed.
ACKNOWLEDGMENTS
We thank parataxonomist A. Towati of Ambunti, the Glei villagers of Waskuk,
and the Galiase and Kokomo clans, for their assistance during our visits to the
upper Sepik. Hitofumi Abe (Ecosystem Research Group, University of Western
Australia) wrote the Japanese translation. Figure | was adapted from Geographic
Information System downloads from the PNG Forest Authority. Reviewers
David Middleton (Harvard University Herbaria) and an anonymous referee,
made helpful comments on the draft.
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BIOSYSTEMATIC ANALYSIS OF THE THELESPERMA
SUBNUDUM COMPLEX (ASTERACEAE)
Curtis J. Hansen! and Loreen Allphin2 Michael D.Windham
“ore, of Botany and Range Science Garrett needa
Brigham Young University Utah Museum of Natural History
Provo, UT 84602, U.S.A, University of Utah,
Salt Lake City, UT 84112, U.S.A.
ABSTRACT
Ch li 1 phyl i | { morphology, allozymes, and nrDNA-ITS sequence
data were used to ee eee relationships in the Thelesperma subnudum complex. Of spe-
cial interest is the placement of three globally rare members of the complex: und les eas rma cdespitosum,
1]
I. pubescens, and T. subnudum var.alpinum. The first two rare taxa yield nly diploid (n=12) chromo-
some counts; T. subnudum var. alpinum is consistently ee (n=24). Thelesperma subnudum
th diploid (n=12) and tetraploid (n=24) populations. Pee ieee
ses of individual data sets ia support (bootstrap 88%-99%) T. subnudum var. alpinum as be
more Cosa related to T. pubescens and z cae es than to T. subnudum var. Sabai. ome
otstrap 100%) for the clade of T. pubescens, T. caespitosum,
and T. stubpudum var. niet um. nm Beliguany ev cer suggests that T. subnudum var. alpinum may be
an allotetraploid resulting from hybridization between T. pubescens and T. subnudum. Based on the
available evidence, we propose the following nomenclatural changes: Thelesperma pubescens Dorn
Var. caespitosum (Dorn) C.J. Hansen, stat. nov, and Thelesperma windhamii C.J. Hansen, nom. et stat. nov.
Key Worps: Thelesperma, Asteraceae, nrDNA-ITS sequences, phylogenetics, and systematics.
RESUMEN
tiidi
Se han usa cromosomicos y analisis filogenéticos de datos morfolégicos, alozimas y de
secuencias de nrDNA-ITS para tratar de resolver las relaciones taxonomicas dentro del complejo
Thelesperma subnudum. De especial interés es la posicion taxonémica de los tres miembros del
ee Thelesperma caespitosum, T. pubescens y T. subn guia var. Bey considerados
Losd 12), mientras
ai le T. cecibieies var. alpinum s se mostré cous antement: como tetrap lide (n = 24). pees
ides ( ae Pele
is fuertemente coe
subnudum \ i
cl Ges Tee | are ] qf. : J
resultados ne g
88%-99%) el agrupamiento de T. subnudum vat. alpinum con af reaps y T. pues que su
Pes aie con pauer pelt var. subnudum
I re Re var. alpinum, T.caespitosum y T. artist
Beeeneie preliminares aren que T. subnudum var. alpinum podria ser un alotetraploide
resultante de la hibridacion entre T. subnudum y T. pubescens. De acuerdo con la evidencia disponible
proponemos los siguientes cambios nomenclaturales: Thelesperma pubescens Dorn var. caespitosum
(Dorn) CJ. Hansen, stat. nov. y Thelesperma windhamii CJ. Hansen, nom. et stat. nov.
PALAsRAs CLave: Thelesperma, Asteraceae, nrDNA-ITS secuenciacion, filogenética y sistematica.
‘Current Address: Biological Sciences Department, Auburn University, Auburn, AL 36849, U.S.A.
Author for correspondence: (email; loreen_woolstenhulme@byu.edu)
SIDA 20(1): 71 - 96. 2002
72 BRIT.ORG/SIDA 20(1)
—
Thelesperma Lessing (Asteraceae: Heliantheae, Coreopsidinae) comprises about
15 species, mostly herbaceous perennials (annuals and subshrubs rare), native
to south-central and western North America and warm-temperate South
America (Melchert 1963; Bremer 1994a). The genus is thought to be monophyl-
etic based on shared characters such as strongly dimorphic involucral bracts,
scarious-margined inner involucral bracts that are connate from 1/5to 1/2 their
lengths, opposite leaves, and pappi that are absent or composed of hispid or
serrulate awns (Bremer 1994a). Previous taxonomic studies of Thelesperma have
focused on morphology (Shinners 1950a, b; Alexander 1955) and cytology
(Melchert 1963; Greer 1997; Greer & Powell 1999) and were not explicitly phylo-
genetic. Nevertheless, past work on the genus has identified some species groups
that may be monophyletic. One of these is the Thelesperma subnudum com-
plex, which is restricted to the Colorado Plateau and areas adjacent to the Rocky
Mountains in western North America (Fig. l). This group includes live com-
monly accepted taxa: T. subnudum A. Gray var. subnudum, T. subnudum A. Gray
var. alpinum S.L. Welsh, T. pubescens Dorn, T. cdespitosum Dorn and T.
marginatum Rydb. Features used to distinguish members of the T. subnudum
complex are given in Table I.
Delimitation of taxa within the Thelesperma subnudum complex has var-
ied, and at least three classifications have appeared (Table 2). Dorn (1990), who
discovered and named T. pubescens and T. caespitosum, recognized each as dis-
—
tinct species. He treated T. marginatum as a separate species and accepted
alpinumasa variety of T. subnudum. In the 2nd edition of A Utah Flora, Welsh
et al. (1993) classified both T. caespitosum and T. subnudum var. alpinum as va-
rieties of T. subnudum: T. marginatum and T. pubescens were not treated because
the former is not known from Utah and discovery of the latter in Utah post-
dates publication of the flora. Cronquist et al. (1994) recognized only T.
subnudumand T. pubescens at the species level; T. caespitosumand T. subnudum
var. alpinum were included within T. pubescens, and T. marginatum was treated
asa variety of T. subnudum.
These divergent classifications result from differing interpretations of the
available morphological and cytological information. Additional genetic data
(e.g.,allozyme and DNA) are needed to allow an informed choice between com-
peting taxonomies. The need to pursue these studies has been heightened re-
cently by conservation concerns. Thelesperma pubescens, T.caespitosum, and T.
subnudum var. alpinum are rare taxa restricted to a few localized populations
(Fig. 1). All three have been listed as potentially endangered (category 2) by the
US. Fish and Wildlife Service (U.S. Dept. of Interior 1985, 1993). The accumula-
tion of genetic data is an important step in determining their eligibility for fed-
eral protection under the Endangered Species Act.
Our objectives in this study are threefold: 1) to develop baseline genetic data
for members of the Thelesperma subnudum complex, 2) to analyze the data in
HANSEN ET AL
50° nw +
aereneoete eres, (@)
ot 0 = 105. W
SS +4]
re aN Ss Ma
FoR a 8
C U
A
: a
aie a no a
Lf. marginatun
vie nudum
var. subnudun [J
var. alpinun
T. ecaespttosum A&A
T. pubescens ©
Fic. 1. Distribution of taxa in the Theless difi Dorn 1990).
74 BRIT.ORG/SIDA 20(1)
he
=
}
Thelesperma subnudum complex. Modified from Dorn 1
Taste 1. Selected piece ecological, and distributional features distinguishing taxa of
990.
Taxa Leafsegments Pubescence Heads Rootstock Habitat Elevation (m)
Tsubnudum — long,broad ~~ none |-several, somewhat common, 1050-2310
radiate creeping — sandy soils
(rarely)
discoid
T.subnudum — short,narrow lower 1(2), discoid somewhat — specialized, 1830-2680
var. alpinum stems creeping, Carmel
& leaves branched — Form. Navajo
caudex Sandstone
I.marginatum long,broad = none |-several, somewhat common, 2130-2835
discoid creeping sandy out
wash
I. pubescens — short,narrow_ leaves 1(2),discoid branched — specialized, 2430-2715
caudex weathered
Bisho
p
conglomerate
specialized, 1220-2680
ower caudex whitish shale
blade)
-caespitosum short, narrow petioles 1(2), discoid branchec
(
=
concert with morphology to produce an explicit phylogenetic hypothesis for
these taxa, and 3) to compare that hypothesis to existing classifications and
propose any desirable taxonomic changes. Through this process, we hope to
shed light on the relationships and possible origin of T.subnudum var. alpinum,
the rarest and most enigmatic member of the group.
MATERIAL AND METHODS
Determination of Chromosome Numbers/Ploidy Level
Ploidy levels were determined for all taxa in the T. subnudum complex using a
combination of chromosome counts and analyses of allozyme banding pat-
terns. For chromosomal observations, capitula (in bud) were obtained from field
populations and fixed in Farmer's solution G parts ethanol:l part glacial acetic
acid), Chromosome counts were made from meiotic figures obtained using stan-
dard squash techniques and acetocarmine staining (Turner & Johnston 1961;
Strother 1972). Preparations were examined under phase contrast on a Zeiss
Axioplan2 Microscope. Images of chromosome squashes were saved electroni-
cally using Zeiss Image® software from Carl Zeiss, Inc.
In populations for which flower buds were unavailable, ploidy level was
determined by careful examination of allozyme banding intensities across a
variety of enzymes (Danzmann & Bogart 1982a; Dessauer & Cole 1984; Pryer
& Haufler 1993). Because allozyme markers are additive, codominant, and
HANSEN ET AL., DIVOTOICMAILS UF 75
Taste 2. Previous taxonomic treatments of Thelesperma subnudum and allied taxa.
Dorn (1990) Welsh et al. (1993) Cronquist et al. (1994)
T. subnudum var. subnudum T. subnudum var. subnudum T. subnudum var.subnudum
T. subnudum var. alpinum T. subnudum var. alpinum T. pubescens
T. pubescens Not treatec T. pubescens
l. caespitosum T. subnudum var. caespitosum T. pubescens
T.marginatum Not treated T.subnudum var. marginatum
inherited in a Mendelian fashion, the expressions of alleles at various ploidy
levels are expected to be proportional to their gene dosages. In our study, chro-
mosomally documented diploids in Thelesperma always showed balanced band
patterns at heterozygous loci. If a heterozygote showed unequal band intensi-
ties of 3:1 (ina monomeric enzyme) or 9:4:] (in a dimeric enzyme), the particu-
lar individual always proved to be tetraploid (the only type of polyploid en-
countered during our study). Thus, chromosomally unknown populations
could be assigned toa specific ploidy level based on the presence or absence of
unbalanced heterozygous allozyme patterns.
Morphological Data
Our coding of morphological character states is based on observation of ap-
proximately 300 herbarium sheets from ASC, BRY,GH, MONTU, NMU, NY, RM,
UT, and UTC (herbarium designators follow Holmgren et al. 1990). These data
were supplemented by information obtained from Melchert (1963), Welsh (1983),
Dorn (1983, 1990), Jansen et al. 1991), Ryding & Bremer (1992), Welsh et al. (1993),
Cronquist et al. (1994), and Karis & Ryding (1994). Ingroup taxa included T.
subnudum var. subnudum, I. subnudum var. alpinum, T. pubescens, T. caespitosum,
and T. marginatum, plus the related species T.filifolium (Hook.) A. Gray, T. longipes
A. Gray, and T. megapotamicum (Spreng.) Kuntze. Bidens has been identified as
a possible sister genus to Thelesperma (Ryding & Bremer 1992) and two species
of that diverse genus, B. cernua L. and B. frondosa L., were chosen as outgroup
taxa. A total of 16 characters (14 binary and 2 multi-state) was included in the
morphological analysis (Tables 3, 4).
—
~—
nr DNA-ITS Sequence Data
Samples used in the DNA study are identified by their GenBank accession num-
bers in Table 5. ITS sequences for the outgroups Bidens cernua and B. frondosa
were obtained from GenBank; voucher data for these collections can be found
in Ganders et al. (2000). The ITS sequences for all Thelesperma taxa were ob-
tained directly by extracting total DNA from the leaf tissue of dried specimens
using a basic CTAB extraction protocol (Hillis et al. 1996). Two different indi-
viduals from each taxon were sampled in order to check for intraspecific differ-
ences. The ITS-1 @ 2and 5.85 regions were amplified using the polymerase chain
—
76 BRIT.ORG/SIDA 20(1)
Paste 3. Morphological characters and character states used in phylogenetic analysis. All characters
treated as unordered.
1. Herbs (0
2. Perennial (0); annual (1).
3. Plants with taproots, sometimes creeping (0); fibrous roots (1); rhizomes (2).
4. Plants without a branching caudex (0); plants with a branching caudex (1).
Ww
subshrubs (1
a=
5. Plants not marcescent basally (0); plants marcescent basally (1).
6. Leaves cauline - leaves basal, much reduced distally (1).
7. Leaves simple, or dentate only (0); leaves pinnately or ternately lobed (1).
8. nee re ae leaf petioles or leaf margins pubescent (1); leaves pubescent
throughout (2).
9. Inner involucral bracts essentially free (QO); inner involucral bracts connate 1/5 or more (1).
10. Inner involucral bracts without scarious margins (0); inner involucral bracts with scarious
margins (1)
1. Capitula with ray flowers (0); capitula without ray flowers (1).
12. Disc florets equally lobed (0); disc florets unequally lobed (1)
13. Cypselae straight (0); cypselae incurved (1).
14. Cypselae pubescent throughout (0); cypselae glabrous or apically pubescent only (1).
1
]
Ss
5. Pappus present (0); pappus absent (1).
6. Stems pubescent (Q); ctems glabrous (1).
Taste 4. Morphological data matrix (Oh taxa in Bidens and Thelesperma. Characters as listed in Table 3.
Variable characters are coded as‘v
Character Number
12345 67891 111111
Taxa 0 123456
Bidens cernua 01100 00000 vOO00 0
ele frondosa a 00 01000 10000 0
i t 0001111111 101111
Th a erma filif ues Ol oe
Thelesperma longip 10000 11011 10110 1
Thelesperma marginatum 0000 11011 101101
Thelesperma megapotamicum )01011 111101
Eneiepena panera re 11211 101111
ma suon Q0000 11011 vO110 1
ae ienieie var.alpinum 00011 11011 101101
reaction (PCR) and two primers ina 1 ratio. Primers used in PCR amplification
and DNA sequencing were ITS-4 (White et al. 1990) and a modified sequence of
ITS-5 (White et al. 1990). The latter, designated ITS-I, has the sequence: GTC CAC
TGA ACC TTA TCA TTT AG (L. Urbatsch, pers. comm.). The following PCR cy-
cling protocol was used in amplification: a 95°C hot start for 12 min. 40 cycles of:
95°C for 20 sec., 54°C for 30 sec., and 73°C for 1 min: a 4 min. final extension at
HANSEN ET AL., DIVOTOTCNIALINOG UF 77
73°C; and ending with a hold at 4°C. To confirm base positions, the ITS-] & 2
and intervening 5.8S regions of Thelesperma were sequenced in both directions.
The contigs, or unidirectional sequences, were assembled using Sequencher
(Gene-Codes, Ann Arbor, MI) and visually checked. Base positions that gave
equally strong, contrasting signals on both strands were coded as ambiguous
according to the IUPAC-IUB ambiguity code set included in the program
Sequencher. All characters were coded as unordered, and gap characters (“-”)
were treated as missing data rather than a fifth character state (Baldwin 1993).
Allozyme Data
An electrophoretic survey was performed to analyze allozyme variation at a
variety of enzyme loci. A total of 765 plants were sampled from 25 natural popu-
lations (about 30 plants/population) representing all taxa in the Thelesperma
subnudum complex, T. longipes A. Gray, and one outgroup population of Bidens
cernud L. (Table 5). Fresh leaf material was collected in the field and kept on ice
until returning to the laboratory (2-3 days maximum). Tissue was then ground
using a mortar and pestle in the phosphate-PVP extraction buffer of Soltis et al.
(1983). Extracts were absorbed onto wicks cut from Whatman 3MM filter pa-
per and stored at -80°C until electrophoresis.
A total of 19 enzymes was surveyed electrophoretically for variability. Six
enzymes, representing 11 putative loci, provided consistent, interpretable re-
sults. Two buffer systems were used to resolve these enzymes in 11.5% (w/v)
starch gels. A tris-citrate/borate buffer system (System 6 of Soltis et al. 1983)
provided good resolution for leucine aminopeptidase (LAP), phospho-
glucoisomerase (PGI), phosphoglucomutase (PGM), and triosephosphate
isomerase (TPI). A pH 7.5 modification of the morpholine-citrate buffer system
of Odrzykoski and Gottlieb (1984) was used to resolve malate dehydrogenase
(MDH) and shikimate dehydrogenase (SkDH). Staining schedules and proto-
cols followed Soltis et al. (1983) and Murphy et al. (1996).
Genotypes were inferred directly from electromorphs observed on the
stained gels, based on the assumption that enzyme substructure and compart-
mentalization parallel those observed in other flowering plants (Gottlieb 1981).
When enzymes showed more than one allozyme locus, the most anodal (fast-
est migrating) locus was designated number I, the next fastest number 2, etc. If
more than one allele was present at a locus, the most anodal allele was desig-
nated a, the next fastest b, etc.
In a departure from common practice, the allozyme alleles detected at each
locus were coded as present or absent for each taxon and included ina phyloge-
netic analysis. Although advocated by some (Mickevich & Johnson 1976;
Mickevich & Mitter 1983; Buth 1984), this approach has been criticized by
Swofford et al. (1996) because it violates the assumption of character indepen-
dence. Similar treatment of allozyme data from Mimulus sect. Erythranthe
78 BRIT.ORG/SIDA 20(1)
Taste 5, Populations used in morphology, allozyme, and nrDNA-/TS studies of Thelesperma. Letters
before collection numbers identify the following collectors: H = CJ. Hansen; S = CJ. Stubben; W =
M. D.Windham; Wo = L. Allphin Woolstenhulme. All vouchers are deposited at the University of
Utah herbarium (UT) unless otherwise indicated by upper case herbarium designators (based on
Holmgren et al. 1990) following the collection numbers. x = no allozyme samples.
Taxa (Code) Geographic origin & voucher Allozyme Sources for DNA
sample (GenBank #)
size
T. caespitosum (Thcal) Utah: Duchesne Co.: 30 #AYO17361
H and S 97-80
(Thca2) Wyo: Sweetwater Co.: 30 #AYO1 7360
Hand § 97-9]
T. filifolium
var. filifolium (Thfi2) Tex: Potter Co.: x #AYO17365
Higgins 16391 (BRY)
var. intermedium (Thfil) Wyo:Goshen Co x #AYO1 7364
Nelson 25622 (RM)
T longipes (Thlo2) NewM: Dona Ana Co. $ 385 29 #AYO17355
(Thlo1) Tex: Culberson Co.: Higgins x #AYO17354
17688 (BRY)
Te marginatum (Thma2) Wyo: Fremont Co.:H 97-134 30 #AYO1/357
(Thmal) Wyo: Natrona Co. Hartman x #AYO1/356
38509 (RM)
T.megapotamicum — (Thmel) = NewM: Sandoval Co.: x #AYO17366
Atwood et al. 24016 (BRY)
(Thme2) = Utah: San Juan Co.: Atwood x #AYO1/36/7
22534 (BRY)
T. pubescens Wyo: Sweetwater Co.: 29
Hand § 97-101
(Thpul) Wyo: Uinta Co..H and $ 97-171 30 #AYO17358
(Thpu2) = Wyo: Uinta Co..H and § 97-117 30 HAYO17359
T.subnudum Ariz: Coconino Co.: Wo 31
and H 97-141
Ariz: Coconino Co.: Wo 33
and H 97-142
Ariz:Coconino Co.: Wo 35
and H 97-143
Ariz: Mohave Co.: W 97-330 30
(Thsu2) Ariz: Mohave Co.:H et al. 97-43 39 #AYO17351
Nev: Clark Co.: W 98-247 2)
Utah: Carbon Co.: Wand 30
Heckel 2417
Utah: Duchesne Co.: H and 30
Nielsen 97-71
Utah: Emery Co.:H and Wi 97-66 30
Utah: Garfield Co.: H 97-79 30
continued
HANSEN ET AL., DIVOTOTCMATING UF 79
Taxa (Code) Geographic origin & voucher Allozyme Sources for DNA
ample (GenBank #)
size
Utah: Grand Co.: H and W 97-69 30
Utah: Uintah Co.: H and 30
Nielsen 97-72
(Thsu4) Utah:Washington Co.: 36 #AYO17352
Hetal. 97-45
Utah: Wayne Co.:H and W 97-57 30
Utah: Wayne Co.:H and W 97-68 23
var. alpinum (Thwil) Utah: Wayne Co.:H 97-73 31 #AYO17362
Utah: Wayne Co.:H 97-74 30
(Thwi2) Utah: Wayne Co.: Anderson x #AYO17363
92? (BRY)
Bidens cernua (Bice) Utah: Utah Co.:H 97-144 30
Ganders et al. 2000 x #U67098
Bidens frondosa (Bifr) Utah: Utah Co.: Welsh 608 (BRY) x
Ganders et al. 2000 x #U67094
(Windham unpubl.) produced a phylogenetic tree highly concordant with in-
formation from other sources, suggesting that these data may contain a strong
phylogenetic signal in spite of their perceived limitations. The allozyme analy-
sis undertaken here is presented as an experiment to further assess the value of
enzyme data in phylogenetic reconstruction.
Phylogenetic Analyses
All phylogenetic analyses were performed using the computer program PAUP
(Phylogenetic Analysis Using Parsimony, version 3.1; Swofford 1991) utilizing ran-
om stepwise addition. Only the shortest trees were retained in each search.
Allcharacters were considered unordered and given equal weights, with multi-
state characters treated as ‘uncertain. Characters and character states were
tracked, organized, and manipulated using the computer program MacClade
3.0 (Maddison & Maddison 1992). Nodal support in each topology was deter-
mined by100 bootstrap replicates (BS; Felsenstein 1985), as well as by calculat-
ing Bremer support values (BV; Bremer 1988, 1994b).
Separate and combined phylogenetic analyses were conducted on three
data sets: morphology, nrDNA-ITS sequences, and allozymes. Examining all
relevant data in a combined fashion can produce a more robust estimate of
phylogeny than separate analyses by maximizing congruence among different
sources of data in phylogenetic inference (Hillis 1987; Kluge 1989; de Queiroz et
al. 1995; Nixon & Carpenter 1996). To estimate levels of congruence among data
sets, incongruence length differences (ILDs) were calculated (Mason-Gamer &
—
80 BRIT.ORG/SIDA 20
=
1)
Kellogg 1996; Johnson & Soltis 1998). This index measures the amount of extra
homoplasy that results from the combination of two or more data sets, as de-
scribed by Mickevich & Farris (1981) and Farris et al. (1994, 1995). The ARNIE
program in the Random Cladistics software package (Siddall 1995) was used to
determine the significance of ILDs. a-values less than 0.05 were considered sul -
ficient evidence to reject the null hypothesis of data set homogeneity. Com-
bined analyses included only those taxa common to all phylogenetic data sets.
RESULTS
Chromosome Data
All individuals sampled from populations of T. pubescens and T. caespitosum
proved to be diploid with chromosome counts of n = 12 (Table 6; Figs. 2A-B).
Individuals sampled from populations of T. subnudum var. alpinum consistently
were tetraploid with n = 24 (Fig. 2C). Our analyses revealed that some Colorado
Plateau populations of T. subnudum var. subnudum are diploid (n=12; Fig. 2D)
whereas others are exclusively tetraploid (n=24; Fig. 2E). Although the geo-
graphic ranges of these cytotypes overlap and both appear to be common
(Windham, Hansen, ee seus unpubl. data), we have yet to identify a locality
where they occur together. Both ploidy levels of T. subnudum var. subnudum are
represented in the morphological, DNA, and allozyme analyses that follow.
A chromosome number was not obtained directly for [. marginatum, and
no previously published counts were found in the Index to Plant Chromosome
Numbers. However, an analysis of allozyme banding intensities in our collec-
tion from Fremont Co., Wyoming, provided strong evidence that the plants at
this locality are diploid. This sample of T. marginatum showed only balanced
heterozygote ae patterns, as is expected in diploid organisms (Danzmann
& Bogart 1982b; Wendel & Weeden 1989) and observed in all chromosomally
verified diploid populations of Thelesperma.
—
Morphological Data
Parsimony analysis of lo morphological characters yielded two most-parsimo-
nious trees, the strict consensus of which is shown in Figure 3. Relative to the
outgroup species chosen, Thelesperma was strongly supported as a monophyl-
etic group (BV=6, BS=100%). The first branch within the Thelesperma clade pro-
ducesa trichotomy that separates I. megapotamicumand T.filifolium (both with
unequally lobed disc florets and a well developed pappus) from the other spe-
cies. The clade encompassing the remaining taxa is weakly supported (BV=l,
BS=70%), branching to form a polytomy in which each taxon (with the excep-
tion noted below) occupies its own unresolved branch. The only deviation from
this pattern is the grouping of IT caespitosum, T. pubescens, and T. subnudum var.
alpinumina single clade with strong support (BV=2, BS=88%). Within this clade,
the last taxon is moderately supported as sister to [. caespitosumand T. pubescens.
HANSEN ET AL
81
Tase 6. Chromosome counts on taxa belonging to the Thelesperma subnudum complex. Apparent
first counts for a taxon are marked by a double asterisk following the relevant name. Letters before
collection numbers identify the following collectors: H = CJ. Hansen; S = C.J. Stubben;W = M.D
Windham; Wo = L. Allphin Woolstenhulme.
Thelesperma caespit Dorn (= T. pubescens Dorn var. caespitosum (Dorn) C.J. Hansen)**
ae Utah Duchesne Co. 1.2 mi up jeep road to Anthro Mtn. from Chokecherry
ane (17S, R4W, S18): H and § 97-80 (UT)
n=12 Wyo — Sweetwater Co. ca. 3.3 mi past bridge on dirt road to Scott’s Bottom
SE of Green River (T18N, R106W, S31); H and § 97-9]
(UT)
Thelesperma pubescens Dorn (= T. pubescens a var. pubescens)**
=12 Wyo — Sweetwater Co. ca.0.3 mi S of switchback on dirt road leading up to
ane Mtn. Summit (T13N, R112W, S10) H and § 97-
101 (UT)
n=12 Wyo — UintaCo. ca. . 6 mi past left fork is Bc Creek Mtn. (T13N,
13W,S3);H and $ 97-111 (UT)
n=12 Wyo Uinta Co. ie summit of Hickey a radio tower (T13N,
114W, $13); H and S 97-117 (UT)
Thelesperma subnudum Gray
=12 Ariz. Mohave Co. W of Wolf Hole Valley ca. 3.85 km ENE of Mustang
Knoll (T39N, R12W, S31); Ha, 8, W and Wo 97-43 (UT)
n=24 Utah Washington Co. W base of Smithsonian Butte (1425, R11W, $21);H, S,
Wand Wo 97-45 (UT)
Thelesperma subnudum Gray var. alpinum Welsh (= T. windhamii CJ. Hansen)**
Utah Wayne Co. SW of Teasdale near the base of Boulder Mtn.
(1295, R4E, S20); W 93-144 (UT)
n=24 Utah Wayne Co. ca.0.6 mi W of State St.in Teasdale on road to
ee Creek (T29S, R4E, $21); H 97-73 (UT)
n=24 Utah Wayne Co. ca.0.8 mi S of SR 24 on dirt road to Government
Creek ae R4E, S18): 97-74 (UT)
ITS Sequence Data
Of 663 characters (aligned length), 61 were variable, and 6 were phylogeneti-
cally informative. The sequence of T. caespitosum was incomplete with approxi-
mately 87 bp missing compared to the other sequences. Possible non-specific
amplification of the ITS region in T. caespitosum resulted in a double signal on
the chromatogram. Multiple attempts to re-extract DNA and obtain a clearer
signal failed. To determine if there was a loss of phylogenetically informative
characters in the missing region of T. caespitosum, two different sequence length
scenarios were analyzed. The first scenario involved aligning all sequence
lengths equal to that of T. caespitosum (.e., no gaps in T. caespitosum but with
an 87 bp truncation in all other taxa). The second scenario was to align full
sequence lengths in all taxa except I’ caespitosum (ie.,87 bp gapin TL. caespitosum
only). In both analyses, the same single most-parsimonious tree was obtained
82 BRIT.ORG/SIDA 20(1)
na cart
Bae
so
Pnae 2
> Dh . L £ Lb £, £ * ohana Tha] n J I
Fic
>
—
gral | p ) T. pubescens, n=12
(= var. pubescens, B) T. caespitosum, n=12 (= T. pubescens var. caespitosum), C) 7. subnudum var. alpinum, n=24 (= T.
windhamii), D) T. subnudum, n=12, E) T. subnudum, n=24.
HANSEN ET Al 33
— F pribescens
88 | _—____— T. caespitosum
——— T. subnudum
var. alpinum
T. marginatum
T. sttbnudum
|
|
100
peeeemencaie | bee T. longipes
T. megapotamicum
1. filifolium
ee arene ieee eee — B. frondosa
B. cernua
Fic.3. Strict £7 . . | rn hal oe op e4
Bremer support values below. Length= 19, Cl= 0.947, RI= 0.941.
(Fig. 4). Furthermore, bootstrap support only differed by 1-2 percent, being
slightly higher in the second scenario. These results suggest that very few phy-
logenetically important characters are located in the 87 bp gapof T.caespitosum
and subsequent combined analyses utilized the second scenario alignment.
The topology of the single most-parsimonious tree from the ITS sequence
analysis was similar to that derived from morphology ( Figs. 3, 4). The mono-
phyly of Thelesperma is again supported by a 100% bootstrap value; Bremer
support in the ITS data is significantly higher (BV=16). The only topological
difference between the morphological and ITS trees involves the placement of
[ filifolium. In the ITS analysis, this species forms a clade with all Thelesperma
other than T. megapotamicum on a relatively well supported branch (BV=2;
BS=86%). As in the morphological tree, all taxa on this branch form an unre-
solved polytomy with the exception of T. pubescens, T. caespitosum, and T.
subnudum var. alpinum. Support for this clade is significantly higher in the ITS
tree (BV=5; BS=99) and, once again, T. subnudum var. alpinum receives moder-
ate support as the sister taxon of T. caespitosum and T. pubescens. A comparison
of pair-wise distances based on the aligned ITS sequences shows very little di-
vergence among these rare taxa (Table 7).
Allozyme Data
Missing allozyme data made up 5.3% of the total data matrix due to poor stain-
ing resolution of SKDH and PGlin populations of T. marginatum and T. longipes.
The results of a phylogenetic analysis based on presence/absence data yielded
84 BRIT.ORG/SIDA 20(1)
7. pubescens
85
2
95 eS mT CC SPILOSUM
5)
a TP subndum
var. alpinum
== Fos deters : annem 7" marginal
79
| > | ee = T. subnudum
79
3 -
——- ~~ T. longipes
= eee B. cernua
Fic 4 Single most. ; ‘ous tree f hausti ITS { ly. Bootst |
above the line, Bremer support values are below. Length= 70, Cl= 0.986, RI= 0.968.
a single most-parsimonious tree (Fig. 5). Relative to the outgroup species Bidens
cernua, the subset of Thelesperma taxa included in the allozyme analysis form
two distinct clades. One consists of the three rare endemics, which show a strong
association (BV= 5, BS=95%) comparable to that observed in the morphological
and ITS trees (Figs. 3, 4). Once again, T. subnudum var. alpinum is sister to T.
caespitosum and T. pubescens, and there is increased support for this topology
(BV=2: BS=85%). The other Thelesperma clade recovered in the allozyme analy-
sis (Fig. 5) consists of T. marginatum, T. subnudum, and T. longipes, which group
together with moderate support (BV=2; BS= 79%). Nearly identical support
(BV=3; BS=79%) exists for the placement of T. marginatum as the sister taxon to
T.subnudum and T. longipes.
_
Combined Data Set Analysis
Statistical comparisons of trees resulting from the individual data sets revealed
that they were highly congruent and amenable to being combined in a single
analysis. This combined analysis of morphology, allozymes, and nrDNA-ITS
sequences resulted in a single most-parsimonious tree, the topology of which
was identical to that of the analysis based on allozymes alone (Fig. 6). Boot-
strap and Bremer support values for the (marginatum (subnudum/longipes))
clade are not significantly different from those derived from the allozyme analy
sis. Support for the (alpinum (pubescens/caespitosum)) clade is increased by
combining data sets. The association of these three taxa is supported by a 100%
ootstrap estimate, and the Bremer support value increases to 12. Statistical
Taste 7. Pair-wise divergence in nrDNA-ITS sequences for Thelesperma and Bidens. Percent divergence ab diagonal; total number of nucleotide differences
below diagonal (generated in PAUP 3.1). Codes for taxa provided in Tasie 6
Thsu2 Thsu4 Thiol Thlo2 Thmal Thma2 Thpul Thpu2 Thcal Thca2 Thwil Thwi2 Thfil Thfi2 Thmel Thme2 Bice Bifr
Thsu2 0 0.40 0.60 0.80 0.80 1.4 1.0 1.2 1.0 0.80 0.80 0.40 0.40 1.0 1.0 6.3 a7
Thsu4 =O 0.40 0.60 0.80 0.80 14 1.0 1.2 1.0 0.80 0.80 0.40 0.40 1.0 1.0 6.3 5.f
Thiol 2 2 0.20 0.80 0.80 1.4 1.0 1.2 1.0 0.80 0.80 0.40 0.40 1.0 1.0 6.3 5.9
Thlo2 3 S) 1 1.0 1.0 1.6 1.2 1.5 1.2 1.0 1.0 0.60 0.60 1.2 1.2 6.5 6.1
Thmal 4 4 4 5 0 1.8 14 15 14 1.2 1.2 0.80 0.80 14 14 6.7 6.1
Thma2 4 4 4 5 0) 1.8 14 1.5 14 1.2 12 0.80 0.80 14 14 6.7 6.1
Thpul 7 8 9 9 0.40 0.50 0.40 0.60 0.60 1.6 1.4 2.0 2.0 71 6.7
Thpu2 5 5 5 6 7 7 2 0 0 0.20 0.20 1.2 1.0 1.6 1.6 6.9 6.5
Theal 5 5 5 6 6 6 2 0 0 0.20 0.20 k2 1.0 2.0 2.0 6.9 6.4
Thea2 5 5 5 6 7 7 2 0 0 0.20 0.20 1.2 1.0 1.6 1.6 6.9 6.5
Thwil 4 4 4 5 6 6 3 1 1 1 0 1.0 0.80 14 14 6.7 6.3
Thwi2 4 4 4 5 6 6 3 ] ] 1 0 1.0 0.80 1.4 14 6.7 6.3
Thfil 2 2 2 3 4 4 8 6 5 6 2 ) 0.20 1.0 1.0 6.3 5.9
Thfi2 2 2 2 3 4 4 7 5 4 5 4 4 ] 1.0 1.0 6.3 Ke)
Thmel 5 5 5 6 7 7 10 8 8 8 7 i 5 5 0 6.1 af
Thme2 5 5 5 6 7 7 10 8 8 8 7 i 5 5 0 6.1 Daf
31 31 32 33 33 a5 34 28 34 33 33 31 31 30 30 Do
D
a
=
i)
(oe)
NO
co
No
\O
WW
Oo
Ww
cS
WW
ce
(oS)
Ww
Ww
NO
N
OV
(ee)
i)
oS)
ne
ww
No
Ne)
NR
Ke)
NR
co
No
[ee]
27
W 1d NISNVH
86 BRIT.ORG/SIDA 20
=
—
)
pte —— _T. pubescens
94
4
100 | born FE CAC SPIOSUM
12
— se ; a i — T. subnadium
var. alpinum
= os tee era —— 7. marginatum
75 |
2 | ——— anne T. subnudum
78
3
ara ————— T. longipes
eee eae ane ——_B. cernua
Fic. 5. Single Rose parsimonious t tree from an Le search based on allozyme data only. Bootstrap values are
above the line gth= 75, C= 0.827, RI= 0.690.
support for the placement of T. subnudum var. alpinum as sister to the other
two taxa increases as well (BV=4; BS=94%).
DISCUSSION
We gathered molecular and morphological data, which were analyzed sepa-
rately and in combination to obtain the best estimate of the phylogeny of the T.
subnudum complex. Tree topologies were highly concordant, and the single
most-parsimonious tree from the combined analysis was more resolved and
exhibited improved nodal support over any of the individual analyses.
The level of congruence among data sets in this study indicates t
allozymes contain valuable phylogenetic information that can be recovered
through parsimony analysis. This suggests that concerns regarding the inde-
pendence of characters (Swofford et al. 1996) should not disqualify allozymes
from playing at least a limited role in phylogenetic studies. Because this ap-
proach is relatively untested, however, we will refrain from placing undue em-
phasis on the allozyme tree in the following taxonomic discussion. This applies
to the combined analysis as well, because the topology of that tree may be un-
duly influenced by the relative abundance of informative allozyme characters.
below addresses only those patterns independently
—
at
iu
The taxonomy outlinec
observed in all data sets.
The relationships portrayed in our phylogenetic trees can be summarized
as follows. Thelesperma pubescensand If. caespitosum are closest relatives, form-
ing a clade in all analyses and showing very little divergence from one another
—_
HANSEN ET Al 87
———— T. pubescens
pa
Pes | ——————— TT. caespitosum
i 7. subd
ar. alpinum
————— TT. marginatum
2 — - T. subniudium
100, +— ; ———— T. longipes
T. filifolium
ee ee aera ene? T. megapotamicum
= aan eeeee rere B. frondosa
eee gee Se : —_—— B. cermua
A ITCNNA Ast +
VINA Udld otto.
p val bove the line, B pport val below. Length= 137, Cl= 0.905, RI= 0.772.
Fic. 6. Single most i i derived fi bined lysis of all morpholoay.
J r f Aa
Baie
(Tables 4 & 7). Another relationship consistent across all analyses is the place-
ment of I. subnudum var. alpinum as the sister taxon to the T. pubescens/
caespitosum clade. Bootstrap support for this topology ranges from 88% in the
morphological analysis to 100% in the combined analysis. Relationships among
other taxa of the Thelesperma subnudum complex are poorly resolved, with the
aforementioned clade forming a polytomy with T. subnudum, T. marginatum,
and T. longipes in the morphology and ITS analyses (Fig, 3). In the allozyme and
combined analyses (Fig. 4), those three taxa form a separate, moderately sup-
ported clade with T. marginatum basal to T. subnudum and T. longipes. These
results are not wholly concordant with any of the proposed classifications of
the T. subnudum complex summarized in Table 2, suggesting that nomencla-
tural changes are warranted.
The Thelesperma pubescens/T. caespitosum clade
Dorn (1990) recognized both taxa in this clade at the species level; Cronquist et
al. (1994) combined them (and T. subnudum var. alpinum) under the name T.
pubescens. Welsh et al. (1993) were mute on the matter of T. pubescens (which
was not known to occur in Utah at the time the flora was published), but prob-
ably would consider it a variety of I. subnudum, the treatment accorded to T.
caespitosum. The last classification is the least practicable in light of the data
presented in this study. To treat T. caespitosum as a variety of T. subnudum and
maintain monophyly, our data (Figs. 3, 4) indicate that other species would have
to be subsumed within T. subnudum as well. These include T. marginatum, T.
88 BRIT.ORG/SIDA 20(1)
longipes, and possibly even T. filifolium. We consider such a species concept,
encompassing taxa not previously included in the T. subnudum complex, to be
unacceptably broad.
Our data indicate that Thelesperma pubescens and T.caespitosum are sister
taxa showing minimal genetic divergence. They differ by a single morphologi-
cal character involving the distribution of pubescence on the leaves (Tables 3
& 4; Dorn 1990). Recent collections from near Anthro Mountain on the West
Tavaputs Plateau in Duchesne Co., Utah (BRY- Goodrich et al. 25159) provide
additional insight on their relationship. In that population, individual plants
range from having strictly petiolar pubescence (T. caespitosum type) to com-
plete leaf blade pubescence (TL. pubescens type). Whether a result of incomplete
primary divergence or secondary convergence resulting from hybridization or
selection, there is complete intergradation. It appears that distribution of pu-
bescence, in the absence of correlated characteristics, is probably a tenuous foun-
dation for recognizing species in the T. subnudum complex.
Under the Biological Species Concept (Mayr 1942), T. caespitosum and T.
pubescensare “groups of actually or potentially interbreeding populations” that
probably should be recognized as conspecific. Invoking the Cohesion Species
Concept (Templeton 1989), T. pubescens and T. caespitosum also qualify as con-
specific based on phenotypic similarities (cohesion) due to gene flow by means
of es iaperpEee tine Because these two taxa lack unique, diagnosable sets
of morphological and molecular characteristics and don’t appear to have sepa-
rate evolutionary trajectories, they also would be considered a single species
under the Phylogenetic Species Concept (Cracraft 1983; Davis & Nixon 1992).
With regard to the caespitosum/pubescens clade, our data are most con-
gruent with the classification presented by Cronquist et al. (1994). In that treat-
ment, T. pubescens is considered specifically distinct from T. subnudum, with T
caespitosum included within T. pubescens. We propose two modifications to this
classification. First, we exclude T. subnudum var.alpinum from synonymy with
T. pubescens for reasons that will be discussed below. Second, we propose to rec-
ognize T. caespitosum asa variety of T. pubescens. Excluding the intergradient
population on Anthro Mountain, Utah, there is a definite correlation among
pubescence, substrate, and geography (Table 1; Fig. 1). We feel that the incipient
—
divergence in this clade is best recognized at the varietal level.
Origin and relationships of Thelesperma subnudum var. alpinum
This taxon appears to be exclusively tetraploid (Table 6), a point that is central to
hypotheses regarding its evolutionary origin and to its classification. Tetraploids
typically are derived from diploids through the incorporation of additional sets
of chromosomes (see Harlan & De Wet 1975), and they are informally grouped
according to the similarity of their constituent genomes (Crawford 1989). Polyp-
loids containing genomes that are very similar chromosomally and genetically
HANSEN ET AL., BIOSYSTEMATICS Ur 89
(usually derived from within a single species) are considered autopolyploids.
Because of genetic similarity to their progenitor diploids, autopolyploids are
rarely given species status (e.g., Mosquin 1966). Polyploids that contain well-
differentiated genomes SORTED: obtained through hybridization between differ-
ent species) are considered allopolyploids and usually treated as distinct spe-
cies. The proper classification of T. subnudum var. alpinum thus hinges on
whether it proves to be an autopolyploid or an allopolyploid.
Classifications of this taxon by Dorn (1990) and Welsh et al. (1993) imply
that its closest relative is Thelesperma subnudum, and suggest that it may be an
autopolyploid derived from within that species. Two lines of evidence refute
that putative relationship. Although the taxon shows some morphological simi-
larities to T. subnudum (Dorn 1990), the ITS sequence data clearly indicate a
closer relationship to T. pubescens and T. caespitosum. In fact, all data sets devel-
oped during this study support the placement of T. subnudum var. alpinum as
sister to these taxa, not I. subnudum var. subnudum (Figs. 3 & 4). Further evi-
dence that var. alpinum is not an autopolyploid derivative of T. subnudum comes
from the fact that known autopolyploids apoun in this species and do not re-
semble var. alpinum. These undisputed lyploidsin T.subnudum
identical to diploid T. subnudum in morphology and ITS sequences (Table 7). In
our analyses, they were recognizable only through chromosome studies or the
detection of unbalanced heterozygosity in allozyme markers shared exclusively
with diploid T. subnudum.
Despite significant genetic similarity, italso seems unlikely that Thelesperma
subnudum var. alpinum is an autopolyploid derivative of either T. pubescens or T.
caespitosum. It varies toward T.subnudumin some morphological characters, most
notably the somewhat creeping rootstock. The two taxa are similar enough to
convince Dorn (1990) to maintain them as varieties of a single species. The pres-
ence, in most individuals, of a PGI-1 allele otherwise found only in T. subnudum
and I. marginatum provides further evidence of genetic links to taxa outside the
I. pubescens/caespitosum clade. Variety alpinum also shows several
autapomorphic traits that distinguish it from T. pubescens and T. cdaespitosum,
including pubescent flowering stems and unique allozyme variants.
We suspect that T. subnudum var. alpinum may be an allotetraploid result-
ing from hybridization between diploid T. subnudum and either T. caespitosum
or T. pubescens. This would explain the pattern of shared characters and appar-
ent morphological intermediacy that has led to such divergent classifications
(Dorn 1990 vs. Cronquist et al. 1994). It also would explain allozyme banding
patterns at the PGI-1 locus, where most individuals of var.alpinumare heterozy-
gous for alleles derived from the pubescens/caespitosum and subnudum/
marginatum clades respectively. At this one locus, var. alpinum approaches fixed
heterozygosity, one of the genetic hallmarks of allopolyploidy. The absence of
fixed heterozygosity at other allozyme loci may be due to homoeologous chro-
90 BRIT.ORG/SIDA 20(1)
mosome pairing or extensive gene silencing (Windham 1988). Over time, al-
lopolyploids lose expression of duplicated parental genes through various ge-
netic processes, especially null mutations (Roose & Gottlieb 1976; Werth &
Windham 1991). Given enough time, the polyploid taxon becomes genetically
“diploidized” (Grant 1981).
A parallel process may explain the lack of diagnostic T. subnudum ITS se-
quences in var. alpinum. Recently formed allopolyploids would be expected to
show the ITS sequences of both diploid parents (Soltis et al. 1995; Cook & Soltis
1999: 2000; Gernandt & Liston 1999). With time, however, ITS loci often experi-
ence concerted evolution, which randomly eliminates one of the parental se-
quences (Sang et al. 1995; Wendel et al. 1995; Polanco et al. 1999). Under this sce-
nario, var. alpinum still could be an allopolyploid hybrid in which the original
T. subnudum sequence has been lost to concerted evolution.
Each of the possible evolutionary origins (two autopolyploid and one al-
lopolyploid) of Thelesperma subnudum var. alpinum discussed above would
supporta different classification for this taxon. The hypothesis that var. alpinum
isan autopolyploid derived from T. subnudum, the only scenario congruent with
the classifications of Dorn (1990) and Welsh et al. (1993), can be rejected. To
uphold alpinumasa variety under T. subnudum and still maintain monophyly,
we would have to expand the species definition of T. subnudum to include the
entire complex plus T. longipes and, possibly, T. filifolium (Figs. 3 & 4). In our
opinion, lumping half of the species in the genus Thelesperma into one species
is not a desirable solution.
Our data are not sufficiently robust to distinguish between an autopolyp-
loid origin of var. alpinum from within the T. pubescens/caespitosum clade or
an allopolyploid origin through hybridization between members of that clade
and I. subnudum. Nevertheless, we can propose a classification that would be
|] a
—
—
phylogenetically congruent of which scenario proves more plausible.
To include var. alpinum within T. pubescens (as done by Cronquist et al. 1994)
would be cladistically indefensible if alpinum subsequently is shown to be an
allopolyploid. However, if we anticipate that the latter hypothesis is more plau-
sible and recognize var. alpinumasa distinct species, such a treatment remains
valid in the event that alpinum maintains its current position as the basal
branch of the T. pubescens/caespitosum clade. Variety alpinum is morphologi-
cally distinct from the other members of this clade and, because of its polyp-
loid chromosome number, is probably genetically isolated from the strictly dip-
loid taxa. Thus, it can be recognized as a species under the Biological Species
Concept (Mayr 1942) as well as the Phylogenetic Species Concept (Cracraft 1983;
Davis & Nixon 1992: Davis 1996).
_
Other taxa in the Thelesperma subnudum complex
The two remaining taxa typically assigned to this complex are T. subnudum
HANSEN ET AL., DIVOTITEMAILS UF 91
and T. marginatum. Dorn (1990) treated them as distinct species; Cronquist et
al. 1994) considered the latter a variety of T. subnudum. These taxa are part of
an unresolved polytomy in the morphological and ITS data sets (Fig. 3), but
form a moderately supported clade with T. longipes in the allozyme and com-
bined trees (Fig. 4). The placement of T. subnudum as sister to T. longipes in-
stead of I’ marginatum in the latter trees suggests a relationship at odds with
previous classifications, which exclude T. longipes from the T. subnhudum com-
plex. This result should be confirmed by additional studies before taxonomic
revisions are proposed. Even if the allozyme data are discounted, there still is
no support for expanding T. subnudum to include T. marginatum as proposed
by Cronquist et al. 1994). Recognition of T. marginatum as a variety of T.
subnudum would require its placement as sister to T. subnudum in a phyloge-
netic analysis. That these two taxa do not form a clade in any of our analyses
suggests that they should continue to be treated as separate species. Each has a
unique, diagnosable set of morphological and molecular characteristics indica-
tive of a distinct evolutionary trajectory, thus satisfying the definition of a phy-
logenetic species (Davis & Nixon 1992).
In order toimplement the classification outlined above, two nomenclatural
innovations are necessary:
Thelesperma pubescens Dorn var. caespitosum (Dorn) CJ. Hansen, stat. nov.
BASIONYM: Thelesperma caespitosum Dorn, Madrono 37: 293. 1990. TyPE: U.S.A.
WYOMING. Sweetwater Co. TI8SN, RLO6W, SEL/4 of SE1/4 of Sect. 3l and SW1/4 of
SW1/4 of Sect. 32,5 km SE of Green River, barren white shale ridge, 1890 m, 22
Jun 1988, Dorn 4948 (HOLOTYPE: RM)).
Thelesperma windhamii C.J]. Hansen, nom. et stat. nov. BASIONYM: Thelesperma
subnudum A. Gray var. alpinum Welsh, Great Basin Naturalist 43: 369. 1983. TYPE:
US.A. UTAH. Wayne Co.: T285, R4E, 513 (NEI1/4), 3 mi due N of Bicknell, bristle-
e forest on multicolored clay hills, 2745 m, 20 Jul 1980, Atwood and Th-
ompson 7646a (HOLOTYPE: BRY!).
This new name for I. subnudum var. alpinum honors the junior author, M.D.
Windham, who proposed and co-directed the study. We chose not to raise the
epithet alpinum to species rank because the taxon never occupies truly alpine
habitats and most populations occur at moderate elevations (ca. 2200 m) in
semi-desert regions.
KEY TO THE TAXA IN THE THELESPERMA SUBNUDUM COMPLEX (ASTERACEAE)
; cm long;stems mainly
9-35(-50) cm tall, scattered Foner: a Re retuhat creeping ae rootstock;
rays present or absent; plants widely distribut
2. Ray florets se (rarely absent), scarious margins of inner involucral bracts
mostly 1-1.5 mm or more wide; Nevada, Utah, Arizona, New Mexic
Colorado T. subnudum
2. Ray florets absent, scarious margins of inner involucral bracts mostly 0.5-1 mm
wide; Wyoming, Montana, Alberta, Saskatchew T.marginatum
92 BRIT.ORG/SIDA 20(1)
. Leaves or lower stems pubescent;leaves 1.5-4 cm pues mainly 3-19 cm tall,
clustered on a thick, branching caudex with old, persistent leaf bases; ray florets
absent; plant distribution restricted,
Pappus a toothed crown; lower portion of flowering stems pubescent; plants
: ) T. windhamii
—
QU
from Wayne Co.,, Uta
3. aes absent; lower portion of flowering stems glabrous; plants from NE Utah
SW Wyomin
5 Leaves ees scent throughout; plants appearing gray-green __ T. pubescens
var. pubescens
4, Leaves pubescent only on petioles or (rarely) on the proximal portion of the
blade; plants appearing green T. pubescens var. caespitosum
It is hoped that this revision of the Thelesperma subnudum complex will re-
solve some of the taxonomic confusion in the group. Although the treatment of
T. pubescens and T. caespitosum as conspecific makes the aggregate taxon less
rare, long-term monitoring and land management still will be necessary. All
three rare taxa are restricted to specific substrates, and populations of T.
pubescens near Green River and Hickey Mountain in Wyoming are endangered
due to off-road vehicle use and oil and gas development. Populations of T.
windhamii located west of Teasdale, Utah, are similarly threatened by off-road
vehicle use.
To improve phylogenetic resolution, future studies should sequence more
rapidly evolving regions of the Thelesperma genome, such as the external tran-
scribed spacer regions of ribosomal DNA (Baldwin & Markos 1998). Artificial
hybridization studies also might prove useful for revealing reproductive barri-
ers and species boundaries. Ultimately, research should be expanded to include
all taxa of Thelesperma. Deciphering relationships within Thelesperma will aid
in identifying possible sister genera and thereby contribute to our knowledge
of higher level relationships in the Coreopsidinae and Heliantheae.
ACKNOWLEDGMENTS
We thank Chris J. Stubben for help in locating and collecting populations of
rare Thelesperma and for collecting samples of T. longipes from New Mexico.
We also thank Lowell Urbatsch for providing ITS primer sequences. We are
grateful to John Strother and Leigh Johnson for their reviews and helpful com-
ments on earlier versions of the manuscript. We also give special thanks to
Marcos Losada for his Spanish translation of the abstract. This study is based
ona M.S. thesis by the senior author done at Brigham Young University.
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=
LIATRIS GHOLSONI (ASTERACEAE: EUPATORIEAE), A
NEW BLAZING STAR FROM THE APALACHICOLA RIVER
BLUFFS AND RAVINES IN FLORIDA
Loran C. Anderson
Department of Biological Science
Florida State University
Tallahassee, FL 32306-4370, U.S.A.
ABSTRACT
jan
A new species of Liatris (Asteraceae: Eupatorieae,) is described from Liberty County, Florida. Pre-
sumably its closest relative is L. gracilis, from which it differs in morphology, chemistry, and ecology.
The species is diploid (n = 10).
Key Worps: Asteraceae. Liatris. blazing star, endemics
RESUMEN
Se describe una nueva especie de Liatris (A ae: Eupatorieae,) del condado de Liberty, Florida.
Probablemente su faerie mas cercano es L Pali de quien difiere en morfologia, quimica, y
ecologia. La es} 5 (n = 10)
The upper Apalachicola River region in the Florida panhandle has long been
known for its diverse and unique flora. Asa Gray (1875) visited the area to ob-
serve Iorreya taxifolia Arn. E.E. Callaway (1966), theologian, amateur arche-
ologist, and one-time gubenatorial candidate in Florida, cited the unusual flora
in support of his claim that the original Garden of Eden was located there. Fur-
ther, he suggested the Torreya tree provided the Biblical “gopherwood” used in
constructing Noah’s ark. Although his ideas did not gain wide acceptance, many
endemic taxa are nonetheless known from the area (Clewell 1977; Myers & Ewel
1990). It is the center of one of five “hot spots” for biotic rarity and richness in
the continental United States (Stein et al. 2000). Now, a new species of Liatris
can be added to the list of rare endemics found there.
For many years, local botanists, such as R.K. Godfrey and A.K. Gholson (pers.
com.), have been aware of some unusual populations of blazing star in ravines
and along the bluffs of the Apalachicola River in the Florida panhandle. The
plants would most probably “key out” to Liatris gracilis Pursh, but they do not
fit well.
Herzet al. 1984a) studied the chemistry of Liatris gracilis. Later, his team
(Herz et al. 1984b) studied the “bluffs blazing star” and found that, in contrast
to authentic L. gracilis, which contained an unusual group of germacrenolides,
it possessed a distinctive group of heliangolides.
The Apalachicola bluffs plants are distinctive morphologically and chemi-
SIDA 20(1): 97 — 103. 2002
98 BRIT.ORG/SIDA 20(1)
cally but have not been given nomenclatural recognition. The most recent sum-
mary of the genus (King & Robinson 1987) makes no mention of this taxon;
neither does Wunderlin (1998) in his guide to the flora of Florida. It is here named
Liatris gholsonii to honor Angus K. Gholson of Chattahoochee, Florida, an in-
defatigable field botanist and astute student of the local flora. Together, he and
Godfrey collected an extensive population series of this new species.
METHODS AND MATERIALS
Vegetative and floral features of the new species and of Liatris gracilis were
measured from the extensive population samples vouchered at FSU. Flowering
heads were softened in a weak solution of NaOH in 50% ethanol to return tis-
sues to their hydrated size and to facilitate dissection. Meiotic chromosome
counts were determined from fresh flower buds preserved in Carnoy’s solution
(6:3:1 parts alcohol: chloroform: acetic acid). Gholson’s personal herbarium is
here termed “AKG’; it will eventually become part of FLAS.
=4
SPECIES DESCRIPTION
Liatris gholsonii L.C. Anderson, sp. nov. (Fig. 1). Typr: UNITED STATES. FLORIDA.
Liberty Co.: — upper slopes of No Name Ravine on the Nature Conservancy's
Apalachicola Bluffs and Ravines Preserve, TIN, R7W, SW1/4 Sec 7, 13 Sep 2001,
L.C. Anderson 19932 (HOLOTYPE: BRIT; IsOTYPES: FSU, MO, NY)
Herbae perennes plus minusve pubescentes et glanduloso-punctatae, caulibus +.5-18.5 dm longis,
eramosis infra ucla aes iam. Folia cane a ce petiolata 8-25 cm longa, up oblongo-
=)
nce aus 27mm orescentiae
if
J: 112 }
vel reflexis 2-26 mm longis. Bracteae florales
nee sessiles, anguste diigteae 2 5.9 mm alone: ve, L8-9 mm latae. Capitula turbinata 6-10 mm
longa, phyllariis 8-15 plerumque acutis ad acuminatis. Flosculi disci 3-6, corollis infundibuliformibus
7-10 mm longis roseis ad purpureis. Achaenia +.5-4.8 mm longa |0-costata
barbellato 5-6 mm longo.
pubescentia, pappo
aa
Perennial herbs, with spherical corm-like rootstocks 2-2.5 cm wide; radical
eaves (usually absent at anthesis) long-petiolate, 22-25 cm long, blades entire,
narrowly elliptic-oblong, 20-26 mm wide, glabrate, finely glandular-punctate
on both surfaces. Stems (4.5-)5.5-12(-18.5) dm long, terete, striate, unbranched
below the inflorescence, pubescent with hairs appressed or spreading. Cauline
—
leaves alternate, entire, numerous, much longer than internodes, punctate; lower
leaves (8-)13-22(-25) cm long, the petioles (2.5-)4-7(-10) mm long, ciliate, blades
elliptic to oblong-lanceolate, tapering at base and tip, (8-)11-22(-27) mm wide,
midveins raised and sparsely villosulous; leaves gradually reduced in size and
becoming sessile toward the inflorescence, grading to the leafy floral bracts
with little change in size or shape. Inflorescence elongate, determinate, race-
mose (rarely paniculate in damaged plants or frequently so in garden plants),
pedicels spreading or reflexed at anthesis, pubescent, (2-)5-12(-26) mm long
on heads toward base of inflorescence. Lowest floral bracts sessile, narrowly
ANDERSON, A NEW SPECIES OF LIATRIS FROM FLORIDA 99
Fic. 1. Plant of Liatris gholsonii (Godfrey 76707.1, a relatively small plant with few leaves to facilitat ing) and
enlargement of one head from Anderson 19932.
elliptic to lanceolate, (2-)2.5-4.5(-5.9) cm long, (1.8-)2-6(-10) mm wide. Heads
turbinate-cylindric, (6-)6.5-8.5(-10) mm long, phyllaries (8-)9-12(-15), green
to dark purple, ovate to oblong, acute to acuminate (rarely obtuse), glandular-
punctate, + ciliate, margins hyaline, white or roseate. Disk flowers 3-5(-6); co-
rollas (6.7-)7-8.5(-10) mm long, narrowly funnelform, sparsely glandular punc-
tate, throat pubescent internally, the tube ochroleucous or pinkish, the lobes
pink to purple, slightly spreading to reflexed, (2.1-)2.5-3.6(-4.6) mm long: bifid
styles pink, (11-)12-13(-14) mm long, anthers (2.5-)3.5-4(-4.4) mm long, with
retuse appendages 0.2 mm long; mature achenes 3.5-5 mm long, 10-ribbed, gray
except for raised, dull green ribs, narrowly turbinate, densely hispidulous, the
100 BRIT.ORG/SIDA 20(1)
hairs 0.4-0.5 mm long, pappus strongly barbellate, 5-6 mm long, white, tinged
with purple to give a dull rosy appearance. n = 10
Phenology.—Primarily flowering from September to mid-October, a very few
precocious plants blooming in early July and a few blooming in early November.
Habitat.—The species occurs in loamy sand or gravelly sand mostly in deep
shade of deciduous tar ands on upper slopes of bayhead ravines or along the
less shady ecotone between the woodland at tops of ravines adjoining more open
scrub oak woodland. It also occurs in xeric to sub-mesic woodland and more or
less open sites on bluffs facing the Apalachicola River or on sandy ridge tops
om
—_
between ravines.
Associated woody species include: Asimina parviflora (Michx.) Dunal,
Callicarpa americana L., Carya pallida (Ashe) Engelm. & Graebn., C. tomentosa
(Poir. in Lam.) Nutt., Cornus florida L. , Diospyros virginiana L., Forestiera
godfreyi L.C. Anderson, Fraxinus americana L., Ilex opaca Ait., Juniperus
virginiana L. , Magnolia grandiflora L., Osmanthus americanus (L.) Benth. &
Hook.ex A. Gray, Ostrya virginiana (Mill) K. Koch, Persea bor bonia (L.) Spreng.,
Pinus glabra Walt., Prunus alabamensis Mohr, Quercus alba L., Q. geminata
Small, Q. hemisphaerica Bartr, Q. nigra L., Rhus copallina L., Sebastiania
fruticosa (Bartr.) Fern., Sideroxylon lanuginosum Michx,, Vaccinium arboreum
Marsh., and Viburnum dentatum L. A few herbaceous species are associates in
the shaded sites, such as Chasmanthium sessiliflorum (Poir.) Yates, Michella
repens L., Smilax pumila Walt, Solidago arguta Aiton, and Spigelia marilandica L.
Additional specimens examined (collections listed generally from north to south in occurrences):
FLORIDA. Liberty Co.: Torreya State Park: 1 Oct 1972, R.K. Godfrey 72216 (FSU), pine-hard wood for-
est, 8 Oct 1972, R.K. Godfrey 72317 (FLAS, FSU); grassy trail in half shade, 9 Oct 1978, R.K. Godfrey
76707 (FSU-population sample of 16 sheets); Weeping Ridge. 21 Sep 1983, R.K. Godfrey 80984 (FSU).
gravelly sandy soil, 12 Oct 1983, R.K. Godfrey 81020 (FSU); just S of park entrance, 17 Oct 1988, R.K.
Godfrey 8293] (FSU); between picnic area and stone bridge, 24 Sep 1983, A.K. Gholson 9348 (AKG).
Rock Hill, 27 Sep 1969, R.K. Godfrey 68853 (FSU); near Rock Bluff, 13 Oct 1956, Redfearn & Godfrey
1026 (FSU, NCU). tributary off Spotts Branch, Upper Sweetwater Creek, 6 Jul 1989 (few in bloom this
date), R.K. Godfrey 83342 (FSU); N of Sweetwater at “old camp torreya,” 21 Sep 1983, R.K. Godfrey
80982 (FSU): ridge NW of Sweetwater, 5 Oct 2001, L.C. Anderson 20018 (FSU, FTG, Us, VDB), 16 Nov
2001, L.C — 20075 (FSU), L.C. Anderson 20077 (FSU); Apalachicola River bluff W of
Sweetwater,l6 Nov 2001, LC. Anderson 20081 (FSU): near rte 270 between rte 12 and Sweetwater, 26
Sep 1980. R.K. Gow y 78179 (FSU); No Name Ravine, 5 Oct 2001, L.C. Anderson 20009 (FSU), culti-
vated in nature garden at Apalachicola Bluffs and Ravines Preserve headquarters building, 5 Oct
2001. L.C. Anderson 20002 (BRIT, FSU, MO, NY, NCU, US), 20003 (BRIT, FSU, MO, NY); Beaver Dam
Creek, 1.6 mi from rte 6 on rte 271, 15 Sep 1988, R.K. Godfrey 82866 (FLAS, FSU); 1.65 mi from rte 6 on
rte 270, 27 Mar 199] (vegetative), R.K. Godfrey 84073 (FSU), | mi from rte 12, 30 Sep 1983, R.K. pai
80995 (AKG, FSU); arm of Little Sweetwater Creek, in shade, 30 Sep 1983, R.K. Godfrey 80992 (FSU, 4
sheets), in sun, 80993 (FSU-4 sheets, UWFP); head of Little Sweetwater Creek near junction rte a
and rte 12, 21 Sep 1983, R.K. Godfrey 80975 (FSU); Alum Bluff, 10 Nov 1963, D.B. Ward 3727 (FLAS), in
shade, 3 Oct ee . iS Ge dfrey 8IOOI(FSU, 2 sheets), in sun, 81002 (FSU-3 sheets, UWFP): + Apr 1990
(vegetative), R.K. Godfrey 83776 (FSU): in open, 2 Oct 1983, A.K. Gholson 10643 (AKG, FSU), ins
2 Oct 1983, A.K.G Shaler 10644 | AKG, FSU): ridge by Kelley Branch, in shade, 2 Oct 1983, R.K. Godfrey
a
ade,
ANDERSON, A NEW SPECIES OF LIATRIS FROM FLORIDA 101
> insun. RK. sles 80998 (FSU, 4 sheets), 2 Oct 1983, AK. Gholson 10642 (AKG), 24 Sep
L.C. Anderson 11834 (FSL
DISCUSSION
Gaiser (1946) recognized 10 series of species in her monograph of Liatris. Series
Graminifoliae (with obtuse, ciliolate phyllaries as in L. gracilis) and series
Pauciflorae (which has generally larger heads with acuminate phyllaries that
lack cilia) are relevant to this discussion. The phyllaries of L. gholsonii are cili-
ate (as in Graminifoliae) but usually acuminate (as in Pauciflorae). The dis-
tinctiveness of the two series was already diminished when Godfrey (1961) de-
scribed L. provincialis; that Florida panhandle endemic has strongly acuminate
but ciliolate phyllaries.
Gaiser (1950) studied the cytology in these series and found the species were
all diploids (n = 10). The chromosomes were small with uniform karyotypes.
For Liatris gholsonii, meiotic chromosome counts from buds of Anderson 19932
were n = 10; the chromosomes were also very small, so chromosome numbers
and karyotypes are not distinctive features taxonomically.
The new species appears most closely related to Liatris gracilis; both have
small heads on relatively long, usually spreading pedicels. The two differ in
their ecology, chemistry (Herz et al. 1984a, b), and morphology. Typical habitat
for L. gholsonii is on more or less mesic, shaded ravine slopes and bluffs; only a
few plants occur in full sun (in openings in the canopy from tree-fall or along
the ecotone between the ravine and the adjacent sandhills), whereas L. gracilis
typically occurs in sunnier, more xeric, open flatwoods and savannas. The two
species are not sympatric, but L. gracilis does occur nearby in the same county.
Foliage is perhaps the most distinguishing feature separating the two spe-
cies. Basal leaves on young plants are broadly elliptic in Liatris gholsonii (as in
Godfrey 84073), whereas they are narrowly lanceolate in L. gracilis (see photo-
graph in Gaiser 1950). Flowering plants of L. gholsonii tend to retain more of
the basal and lower cauline leaves, which are also wider (mostly 11-22 mm wide)
than those of L. gracilis (4-13 mm wide). The transition between cauline leaves
and floral bracts is gradual in the new species, and the floral bracts are mostly
2-6 mm wide, whereas they are narrower (1.0-1.8 mm wide) in L. gracilis.
One might think the differ in foliage could be products of the species’
differing habitats. In general, plants of the same or related species grown in full
sun have smaller, narrower leaves than those found on plants grown in the shade.
However, the larger leaves of L. gholsonii are not due to habitat. The few collec-
tions of L. gholsonii taken in full sun were more robust plants that had shorter
internodes, average-sized cauline leaves for the species, but wider floral bracts
than plants from shadier sites. Plants grown in full sun tended to have more
phyllaries in their involucres (and thus be more like those of L. gracilis) than
those grown in shade.
—
BRIT.ORG/SIDA 20(1)
Personnel of the Apalachicola Bluffs and Ravines Preserve established a
wildflower garden in full sun near their headquarters building. The garden soil
was enriched with mushroom compost and was given supplemental water
Seeds of Liatris gholsonii from No Name Ravine were sown in the garden in
1997, and many plants became established. These plants are somewhat atypi-
cal in
—_—
raving mostly two or three stems per rootstock and large, paniculate in-
florescences. These “full sun” plants still had the characteristic leaves and flo-
ral bracts of L. gholsonii.
Several features of floral morphology tend to separate to two species, but
variation is sufficient within each species that no single feature provides abso-
lute identification. For example, phyllaries in the new species tend to be acumi-
nate, but some plants have phyllaries that are only acute (or rarely obtuse), and
phyllaries in Liatris gracilis are usually very obtuse (rounded apically), but a
few specimens have phyllaries that are acute or rounded but mucronate. Sum-
mation of some floral measurements from over 30 samples per species is in Table
|. Plants of L. gholsonii tend to have shorter involucres (shortest in the genus)
with fewer phyllaries, but their corollas and achenes tend to be larger than those
of L. gracilis.
Differences between the two species in floral morphology (Table 1) ar
present (though minimal is some cases). The salient distinguishing features
between the two taxa are mostly vegetative; the following key couplet provides
a useful summary:
Lower floral bracts over 2 mm wide; lower cauline leaves elliptic, mostly over 12 mm
wide; phyllaries usually acur lis ere) / obtuse
Lower floral bracts under 2 mm v
Liatris gholsonii
anceolate, mostly ly under
> lower cauline leaves
12 mm wide; phyllaries es ane rarely acute
_ Liatris gracilis
Now the Florida panhandle has two endemic species of Liatris; L. provincialis
is found in Franklin and Wakulla counties and is considered endangered in
Florida (Coile 2000). The range of L. gholsonii is even more restricted; it ranges
about 14 km from Torreya State Park to just south of Alum Bluffs and occurs on
bluffs or in ravines only a short distance eastward from the Apalachicola River.
The most distant population from the river is found in the headwaters of
Sweetwater Creek about 9 km from the river. This species is clearly a candidate
for listing as endangered.
-_—
ACKNOWLEDGMENTS
Staff of The Nature Conservancy’s Apalachicola Bluffs and Ravines Preserve
provided some logistical support. Mark Garland supplied the Latin diagnosis.
Ken Womble scanned the specimens into Adobe Photoshop for the illustration.
Guy Nesom, Bob Kral, and A.B. Thistle and anonymous reviewers provided help-
ful suggestions on the manuscript.
ANDERSON, A NEW SPECIES OF LIATRIS FROM FLORIDA
Taste 1. Averages (and ranges) of selected floral features in Liatris gholsonii and L. gracilis.
Liatris gholsonii
Liatris gracilis
Involucral length, mm 6.96(6.0-10.0) 7.62(5.5-10.0)
Phyllary number 10.69(8-15) 12.33(9-16)
Flower number 4.06(3-5) 4.30(3-7)
Corolla length, mm 7 84(6.8-9.8) 7.69(6.0-9.7)
Corolla lobe length, mm 3.09(2.1-4.6) 2.82(2.0-3.9)
Anther length, mm 3.80(2.6-4.5) 2.98(2.0-3.9)
Achene length,mm 4.07(3.5-5.0) 3.40(2.8-4.4)
REFERENCES
CaLtaway, E.E. 1966.In the beginning: creation, evolution, Garden of Eden, and Noah’s ark.
Carlton Press, New York.
Ciewett,A.F.1977.Geobotany of the Apalachicola region. Pp.6-15.In:R.J.Livingston and E.
A. Joyce, Jr, eds. Proceedings of the Conference on the Apalachicola Drainage System.
Fla. Dept. Natural Resources, Marine Research Laboratory. Fla. Marine Res. Publ. 26.
Coie, N.C. 2000. Notes on Florida's endangered and threatened plants. Florida Dept. Agric.
Consumer Serv., Div. Plant Industry-Bot. Sect. Contr. 38, Gainesville.
Gaiser, L.O. 1946. The genus Liatris. Rhodora 48:165-183, 216-263, 273-326, 331-382,
393-412.
Gaiser, L.O. 1950. Chromosome studies in Liatris. |. Graminifoliae and Pauciflorae. Amer. J.
Bot. 37:414-423.
Goorrey, R.K. 1961. Liatris provincialis, sp. nov. (Compositae), endemic in western Florida.
Amer. Midl. Naturalist 66:466-470.
Gray, A. 1875.A pilgrimage to Torreya. Amer. Agric. 34:266-267.
Herz, W., K. Watanase, and J.F. Blount. 1984a. Stereochemistry of chapliatrin and its conge-
ners from Liatris gracilis. Phytochemistry 23:373-382.
Herz, W., K. Waranase, R.K. Gorey, and J.F. Blount. 1984b. 3-germacren-6,12-olides from an
undescribed Liatris taxon. Phytochemistry 23:599-606.
KiNG, R.M. and H. Rosinson. 1987. The genera of the Eupatorieae (Asteraceae). Monogr. in
Syst. Bot., Missouri Bot. Gard, 22.
Myers, R.L.and J.J. Ewet. 1990. Ecosystems of Florida. Univ. Central Florida Press, Orlando.
STEIN, B.A., LS. Kutner, and J.S. Abas. 2000. Precious heritage, the status of biodiversity in
the United States. Oxford Univ. Press, New York.
WUNDERLIN, R.P. 1998. Guide to the vascular plants of Florida. Univ. Press of Florida, Gainesville.
BRIT.ORG/SIDA 20(1)
BOOK REVIEW
Grorce E. Burrows and Ronacp J. Tyrt. 2001. Toxic Plants of North America. (ISBN
0-8138-2266-1, hbk.). lowa State Press, 2121 State Avenue, Ames, IA 50014,
)S.A. (Orders: 1-800-862-6657, 1-515-292-3348 fax: wwwiowastatepress.com ).
$174.95, 1350 pp, 8 color plates, numerous line drawings, 8 1/2" x II”,
this is! Itis a a (nearly = ny aa 8 Ibs) treatment of the plants in
ombined insights
ate a breadth
ks of L.H. Pammell
What a magnificent book
North America that are known t
of the two authors, a veterinary fonicclowtce and plant systematist ee
and depth of coverage that make this work a worthy successor to the earlier boo
(1911) and J.M. Kings bury (1962).
The work organiz ed for easy use. There is a brief introductory chapter, which includes a de-
lightful dedication to the late Dr. J.W. Dollahite of Texas A&M University, who made important con-
tributions to veterinary toxicology in through the middle 20" century. Then comes the alphabeti-
cally arranged treatments for 75 families of plants that are known to be poisonous. The treatments
or each genus includes a consideration of the species.
it
are divided into genera, and the treatment f
Nomenclatur forms to current floristic usage in North America. There is a general description
of the plant and a statement of range and ea that are designed to permit ready identification.
Each toxic plant or group of closely aes: ts is gi discussion of the disease problems, dis-
hol ts trol. The page margins are wide, and printec
ease genesis, clinical signs, |
there are line drawings and gener alized distribution maps for each plant. Short phrases in boldface
noting the topics in the text are also given in the hoe Structural formulas are provided for the
various toxins that are discussed in the text. A nice touch is the colored photographs of 64 toxic
plates in the center . the enue
jen
plants, Ses on 8 colorec
er the families of known toxic concern, there is a |
ted toxicity. Then comes a short discussion on plant identifica:
tailed chapter on 36 other families
that are | doubtful or merely suspec
tion, a list of plants by their adverse effects, a list of plants of concern for dogs, cats, and other pets, an
extensive glossary, and an index.
Each family treatment has its own list of references, and the lists are extensive and current. For
example, the family Euphorbiaceae, which includes numerous toxic plants, has 194 entries in the list
of references. These come from the oo ee toxicological, and botanical literature, and
gthy reference lists are given for other families
most of the references are from the ep ast 25 5 yea
as well, which make the book a rich ae tool for researchers. A cursory pass-through showed
all references to be from the English language literature.
A particularly attractive feature is the simple accessibility of the contents. A pee needing
information about a toxic plant (such information is often needed in a hurry) can locate the informa
tion ag wn rapid) and oe nat is remarkable suc ; a ee pacomipaseine book.
Il know of things that the
f
g
yA
re
ane a have included! but had ~ authors opted for een a book would never have
ompleted. Instead of a piddly quibble here and there, we can all be grateful that the work was
com aoe and made available to the great community of consumers of botanical and toxicological
ey, Botanical Research Institute of lexas, barkley@britorg
information.— [heodore M. Bark
SIDA YA 20(1): ): 104, 2002
A NEW SPECIES OF GONOLOBUS
(APOCYNACEAE: ASCLEPIADEAE, GONOLOBINAE)
FROM SOUTHERN COSTA RICA
Alexander Krings
Herbarium, Department of Botany
North Carolina State ple
Raleigh, NC 27695-7612, U.S.A.
Alexander_Krings@ncsu.edu
ABSTRACT
A new species of G bed, resulting from collections from southern Costa Rica.
RESUMEN
Se describe una especie nueva deG lobus co ltadod 1 l i del sur de Costa
Rica.
Recent study of lianas and vines in southern Costa Rica resulted in a prelimi-
nary checklist of seventy species in thirty-two families from the Las Cruces
Biological Station (Krings 1999). Ongoing work with collections resulting from
this study revealed a new species of Gonolobus Michx. (Apocynaceae:
Asclepiadeae, Gonolobinae).
seas Secor ted eae sp. nov. (Fig. 1). Type: COSTA RICA. PUNTARENAS:
Brus, San Vito de Java, Estacion Biologica Las Cruces, 30 m downhill from
= Marker Ft 59, 1200 m elev. 11 Mar 1997, Krings 274 (HOLOTYPE: F; ISOTYPES: CR,
NCSC, US
Species nova Gonolobus distinguibili a Gonolobus jaliscensis B.L. Rob. & Greenm. petioli c. 4-4.3 cm
longi; sepala lineare vel anguste lanceolata, supra oo subtus dense pilosa, 5-7.2 mm; corollae
purpureis, lobi deltatis, supra puberulis, subtus pilos
Twining vine; indument of short and long _— the short hairs straight, pointed
or sometimes capitate, to 0.4 mm long, borne primarily along the stem, midvein,
and secondary veins, long hairs straight to flexuously curved, (2-)2.8-3.5(-4.3)
mm long, borne along stem, veins, and over leaf surfaces; stems densely yellow-
ish-clear pilose or hirsute; latex white, abundant; petioles 4-4.3 cm long, densely
yellowish pilose to hirsute; leaves opposite, 12-14 cm long, 6.8-8 cm wide, ovate,
densely yellowish pilose to hirsute on both surfaces, the bases cordate to some-
what auriculate, frequently oblique, the apices acuminate; inflorescences axil-
lary, umbelliform, densely 9-15-flowered, the peduncles 0.5-1 cm long; pedicels
7-9 mm long, densely yellowish pilose to hirsute, the hairs to 3 mm long; flow-
ers mildly fetid; sepals 5, linear to narrowly lanceolate, 5-7.2 mm long, pilose or
hirsute on the outer surface, glabrate on the inner surface; corolla campanulate
SIDA 20(1): 105 - 108. 2002
106 BRIT.ORG/SIDA 20(1)
Lah i bien et Al er | DI 1 aL 6
C. Inflorescence: D
fever’ in ee E. ied, flattened flower F. ewig ee in a ora and anes appendages, G. Dexall of
a rr
Pollinarium. All based on Krings 274 (HT: F; IT CR, NCSC, US).
to suburceolate, 5-lobed, the tube 1-2.5 mm long, the lobes deltate, uniformly
flat, dark purple to dark brownish-red, 3-4 mm long, conspicuously shorter
than the sepals, long yellowish pilose to hirsute below, the hairs to 2.8mm long,
coarsely villosulous-strigillose above; faucal annulus 5-lobed, ca. 0.05-0.1 mm
thick, glabrous; corona 1, 5-lobed, fused at the base, adnate to and surrounding
KRINGS, A NEW SPECIES OF GONOLOBUS FROM COSTA RICA 107
the gynostegium, the lobes deltoid to rhomboid, essentially as high as the style-
head, 0.4-0.5 mm high; anthers with fleshy, obdeltate, dorsal appendages;
pollinaria horizontal, ca. 0.5 mm long; follicles unknown.
Gonolobus comprises about 150 species limited to the New World tropics
and subtropics (Stevens 2001). Gonolobus tenuise palus appears to be closely re-
lated to Gonolobus jaliscensis B.L. Rob. @ Greenm., sharing indument of short
and long hairs, densely multi-flowered, umbelliform inflorescences, small co-
rollas (ca. 8 mm in diam.), and similar coronas and anther appendages.
Gonolobus jaliscensis, however, exhibits petioles shorter (to 2.2 cm long), sepals
shorter (only to 2 mm long), corollas greenish to yellowish (purple in G.
tenuisepalus), and anther appendages mushroom-shaped. With the addition
of G. tenuisepalus, nine species of Gonolobus are known from Costa Rica (two
yet undescribed, see Stevens 2001). An updated key follows:
1. Corollas oe within or with barbed lobes.
2. A and adaxial leaf surfaces glabrous, exce a Daluae ane iene
ae principal veins and in the axils g
ages = 14mm long Cee ban: Kunth
2. poate and arena leaf surfaces puberulent; faucal annulus ciliate ae
by the barbellate hairs of fthe corolla li | ‘ I
ages ca.0.9 mm long erik aul Hemsl.
1. nae ee glabrous or sees but not barbellate or barbed.
rolla purple, the lobes 4 mm long or less; faucal annulus 0.05-0.1 mm vale
penal a 5-lobed ridge; nee appendages obdeltate, ca. 0.5 mm long
Gonolobus enisepals Krings
3. Corolla greenish or yellowish (sometimes purple in G. sp. and G )
lobes = 6 mm long; faucal annulus various; anther appendages ei G sp. B)
OR = 0.3 mm long and variously shaped, the apices entire, bifid, or with a central
ee
. Outer ene Surface with trichomes — 1.5 mm long; anther appendages
bifid olobus spss ooa ae ) W.D. Stevens.
4. Outer petal surface glabrous or the oe = 0.4 mm in leng nther
appendages entire, mamillate tipped, or lacking, but not i d.
Corolla lobes = 15 mm long; anther appendages subquadrate, 1-2 mm
long Gonolobus albo ee Woodson
5. Corolla lobes S 12 mm long; anther appendages various or lacking.
6. Corolla purple, the lobes 7.2-9.5 mm long, the apices concave; faucal
annulus an erect, 5-lobed tube, 0.7-1.2 mm long; anther appendages
pee Gonolobus sp.B
G 2enist >|lowish (sometimes purple in G. sp.D), the lobes of
various ie ngths, a nar; ie (anna various, sometimes an indistinct
ridge (e.g, G. chiriquiensis); anther appendages p
7. Leaf bases truncate to rounded; anther ene rounded, with a
short, central, mammillate tip Gonolobus chiriquensis
Woodson) Woodson
7. Leaf bases cordate; anther appendages various.
8. Corolla purple, brownish, or green, the tube 1.7—2.7 mm long, the
108 BRIT.ORG/SIDA 20(1)
lobes 6.6-11.7 mm long, 3.2-4.7 mm wide; gynostegium oe
the stipe 0.3- - 6 mm long; corona cup-shaped onolobus sp. D
8. Corolla green, the tube 3.7-5.7 mm cone, the lobes 12.2-18. a mm
long, 7.8-12.5 mm wide; g essentially sessile; corona
not cup-shaped sass denticulatus (Vahl) W.D. Stevens
ACKNOWLEDGMENTS
Collections at Las Cruces Biological Station were made possible thanks toa travel
grant and assistantship provided by the College of Forest Resources at North
Carolina State University and fellowships in Tropical Biology from the Organi-
zation for Tropical Studies (OTS). | thank: OTS for permission to conduct re-
search at Las Cruces and for help with collecting permits, Luis Diego Gomez for
support and insight during my stay, Paul Fantz for help with the Latin diagno-
sis, Mary E. Endressand J. Francisco Morales for thoughtful reviews of the manu-
script, and the following herbaria and their curators (particularly Robert Wilbur,
William Burger, and Pedro Acevedo) for access to their collections: DUKE, F US.
REFERENCES
Krinas, A. 1999.An annotated preliminary checklist of the dicotyledonous lianas and vines
from the Las Cruces Biological Station, Costa Rica. Sida 18:1247-1258.
Stevens, W.D. 2001. Asclepiadaceae. Mongr. Syst. Bot. Missouri Bot. Gard. 85:234—270.
Woopson, R.E., Jr, R.W. ScHery, and D.L. Speman. 1975. Flora of Panama. Part VIII. Family 163.
Asclepiadaceae. Ann. Missouri Bot. Gard. 62:103-156
A NEW SPECIES OF MEDINILLA (MELASTOMATACEAE)
FROM ANAMALAIT HILLS, SOUTH INDIA
N.Sasidharan and P. Sujanapal
Kerala Forest Research Institute
Peechi 680 653
Kerala, INDIA
ABSTRACT
An undescribed species of the genus Medinilla Gaud., from the Anamalai Hills, South India, is de-
scribed and illustrated.
RESUMEN
Una especie no descrita del género Medinilla Gaud., de Anamalai Hills, sur de India, se describe y se
ilustra.
INTRODUCTION
Medinilla Gaud., an Afro-asiatic genus is estimated to have 430 species
(Veldkamp pers. comm.). The genus has a bicentric distribution pattern due to
the movement of landmasses during the late Cretaceous period from a
Gondwanian origin (Nayar 1972). In Asia, the distribution ranges from Indian-
Himalaya southwards to Sri Lanka, Myanmar, Thailand, Cambodia, Vietnam,
Malay Peninsula eastward through the islands of the Malay Archipelago, New
Guinea down to Northern Australia, and is adapted to warm humid
paleotropical climate. More than 300 species are reported from the Indo-Ma-
laysian region (Nayar 1966). Malayan regions and Madagascar are the principal
centers of diversity and endemism- Philippines: 80 species, Madagascar: 70 spe-
cies, Borneo: 48 species (Perrier 1951; Regalado 1990, 1995). Nayar (1972) has
analyzed the distribution pattern of Asiatic Melastomataceae and recognized
four centers. The Western Ghats is included in the Deccan-Ceylon center (Pen-
insular India and Sri Lanka) of species formation. Among the 9 Indo-Sri Lankan
species, 5 are reported from Peninsular India and Sri Lanka and others are con-
fined to the subtropical Himalayas and Assam hills. Medinilla beddomei Clarke
and M. malabarica Bedd. are exclusive to Southern Western Ghats and the third
species, M. fuchsioides Gard., formerly considered endemic to Sri Lanka, was
recently reported from Southern Western Ghats (Shetty & Karthikeyan 1976).
During explorations in the windward region of Southern Western Ghats,
interesting specimens of Medinilla were collected from three localities of Anamalai
Hills. The specimens did not agree with the species known from India and Sri Lanka.
Further study with literature on Medinilla (Bremer 1987; Perrier 1951; Regalado
1990, 1995) shows that the specimens have more affinity to Indo-Sri Lankan
—
—
SIDA 20(1): 109 - 113. 2002
BRIT.ORG/SIDA 20(1)
—
species and is allied to Medinilla malabarica Bedd. The specimens were further
compared with the collections from Indo-Malaysia in the Kew Herbarium and
no matches were found. It is described and illustrated here as a new species.
Medinilla anamalaiana Sasidharan \ Sujanapal, sp. nov. (Fig. 1). Typr: INDIA.
KERALA: Palakkad District, Parambikulam rs Sanctuary, Shek 2 mudi 1200
m, | May 1999, P Sujanapal 19431 (HOLOTYPE: MH; ISOTYPES: C “ALLA RI, Ld.
Medinilla qnamalai
7
ina M. ma laban icae os in pape omnium oe innovatiorumque rulo-
brunneorum, sed | gulari oliis 3-costatis, apice
a1] =}. | eh i eyes bee | : - } | Bi 14.
VEL
obtuse acuminato, inf
floribus
breve pedicellatis gerentibus distinguendum. In M. malabarica rami annotini teretes cnet folia
crasse coriacea 5-costata, apice acuto vel obtuso, flores in cymis pedunculatis gerentes.
Epiphytic subshrubs; branches fleshy, acutely quadrangular or
subquadrangular, sparingly branched, sometimes rooting from lower nodes,
smooth, on drying forms a winged appearance; stem 20-35 cm long, young
shoots reddish-brown. Leaves membranous, opposite pairs unequal, sometimes
equal, larger one 7-11.5 x 3-5cm, smaller 4-6 x 2.5-3.5cm, elliptic-oblong, base
subcordate or obtuse, apex obtusely acuminate, acumen sometimes twisted,
margin entire, 3-nerved, lateral ribs near the margins, faint towards apex; peti-
ole 2-4 mm, flattened, glabrous. Flowers 4-merous, 2-5 in axils or from leafless
nodes in horizontal row, sometimes 2 or 3 on a short stalk (1-2 mm); pedicel
with 2 minute bracteoles at base, jointed, 2 mm at anthesis, elongating to 4mm
in fruit; hypanthium obovate, subquadrangular, 3 mm across, thinly
puberulous, shortly narrowed at mouth with 8 minute teeth; petals 5-7 x 4mm,
rose-pink, obliquely obovate, minutely apiculate; stamens 8, filaments 3 mm
long; anthers 2 mm long; ovary 1.5 mm across, obovoid, 4-celled, connected to
hypanthium with 8 membranous appendages; style to 7 mm long; stigma mi-
nutely capitate, papillate. Berries 4mm across, globose, glabrous; crowned by a
small hypanthial lim b; seeds many, yellowish-brown, |.5 x 1mm, smooth, ovoid,
concave on the side of raphe.
Medinilla anamalaiana is allied to M. malabarica Bedd. in the general ap-
pearance of the leaves and reddish-brown tender shoots. However, it can be dis-
tinguished from the latter by the succulent quadrangular or subquadrangular
branches, 3-nerved membranous leaves with obtusely acuminate apex and
short pedicelled flowers in axils or from leafless nodes in horizontal rows.
Medinilla malabarica Bedd. has terete woody branches, 5-nerved thickly co-
riaceous leaves with acute or obtuse apex and flowers in pedunculate cymes.
Note.—The floral structure of Medinilla appears to be uniformly similar
in most of the species and is not much diagnostic value. Because of the high
rate of endemism in Medinilla most of the workers
—
ao
followed geographic distri-
bution in addition to morphological or vegetative characters in their regional
revisions or floristic works. Our taxonomic concept for describing M.anamalaiana
also follows the same approach in addition to morphological characters.
SASIDHARAN AND SUJANAPAL, A NEW SPECIES OF MEDINILLA FROM INDIA
Fic. 1. Medinill laiana Sasidt 2 Cyianana lA. fl
ig; B. flower; C hypanthium; D hypanthium cut
open; E. flower L.S; F. anther; G. pistil; H. fruit; |. Seed.
112 BRIT.ORG/SIDA 20(1)
Flowering and Fruiting.—January-June.
Etymology.—The specific epithet of the new species is derived after its type
locality, the Anamalai hills.
Distribution and Status.—The species is so far restricted to the evergreen
forests, with an altitude ranges from 400-1300 m in the windward side of
Anamalai hills.
Habitat and Ecology.—An epiphyte in the crevices of trees in the middle
stratum of evergreen forests. This species is often found associated with epi-
phytes like Aeschynanthus perrottetii A. DC. Common tree associates are
Gymnanthemum arborea (Buch.-Ham.) H. Rob, Elaeocarpus glandulosus Wall.
ex Merr, Iurpinia malabarica Gamble, Drypetes wightii (Hook. f.) Pax & Hoffm.,
Palaquium ellipticum (Dalz.) Baill, Calophyllum polyanthum Wall. ex Choisy,
Mesua thwaitesii Planch. & Triana, etc.
—
PARATYPES: INDIA. Kerala. Palakkad District: Parambikulam Wildlife Sanctuary, Puliyala 1000 m, 3
Apr 2000, P Sujanapal 30043 (KFRD. Pathanamthitta District: Goodrical Range, Moozhiyar 500 m,
23 May 1984, N Sasidharan 3142 (KFRI). Idukki District: Periyar Tiger Reserve, Vallakkadavu 900 m,
3 Jun 1995, Jomy Augustine & KP Rajesh 15429 (KFRD
—
KEY TO SOUTH INDIAN SPECIES OF MEDINILLA
1. Flowers in horizontal rows or on 1-2 mm long peduncle; petals 5-7 x 4 mm
M. anamalaiana
Soe in pedunculate cymes; peduncle 5-12 mm long; petals 10-14 x 8-10 mm.
. Leaves orbicular, fleshy; branchlets 2-3 mm thick, climbing M. beddomei
oe elliptic or elliptic-oblong, subcoriaceous; branchlets more than 5mm thick,
not climbing, clump forming.
3. Petiole 4—7 mm long; lateral nerves arises from the base of the lamina
M. malabarica
3. Leaves sessile or subsessile; lateral nerves arises ca. 2 mm above the base of
the lamina M. fuchsioides
ACKNOWLEDGMENTS
We are grateful to Wildlife Wing of Kerala Forest Department for the fi-
nancial assistance; Director, Kerala Forest Research Institute, for providing fa-
cilities and Wildlife Wardens and Staff of the Parambikulam Wildlife Sanc-
tuary and Periyar Tiger Reserve. Sincere thanks are also due to J.F Veldkamp,
Rijksherbarium, Leiden for critical comments and Latin diagnosis, V.B. Sajeev
for illustrations. The first author is thankful to E. Nic Laughadha, Kew Her-
barium for the help rendered in referring the relevant Medinilla specimens.
REFERENCES
Bremer, K. 1987. Melastomataceae In: M.D. Dassanayake and FR. Fosberg, eds. A revised
handbook for the flora of Ceylon. Amerind Publishing Co. New Delhi.Vol. 6, pp.
157-240.
SASIDHARAN AND SUJANAPAL, A NEW SPECIES OF MEDINILLA FROM INDIA 113
Nayar, M.P. 1972. Centres of development and pattern of distribution of the family
Melastomataceae in Indo-Malesia. Bull. Bot. Surv. India 14:1-12.
Nayar, M.P. 1966. Contributions to the knowledge of Indo-Malaysian and other Asiatic
Melastomataceae. Kew Bull, 20:235-244.
PerrieR DE LA BatHie, H. 1951. Melastomatacees In: H. Humbert, ed. Flore de Madagascar et
Comores Familie 153:1-326.
REGALADO, J.C. Jr. 1990. Revision of Medinilla (Melastomataceae) Borneo. Blumea 35:5-70.
REGALADO, J.C. Jr. 1995. Revision of Philippine Medinilla (Melastomataceae). Blumea 40:
113-193.
SHeTTy, BV. and S. KarTHIKeYAN. 1976. Medinilla fuchsioides Gaertn. (Melastomataceae)—A
new record for India. Bull. Bot. Surv. India 18:215.
——
BRIT.ORG/SIDA 20(1)
BOOK REVIEWS
Dennis W. WoobLanp. 2000. Contemporary Plant Systematics. Third Edition. (ISBN
1-883925-25-8, hbk.). Andrews University Press, 213 Information Services
Building, Berrien Springs, MI 49104-1700, 616-471-6134. (Orders:
aupress@andrews,edu. http://www.andrews.edu/press). Price not given,
569 pp., numerous line drawings, 7" x 10 1/2'
Contemporary Plant Systematics is written with the undergraduate or serious amateur botanist in
and are more rigorous and con-
an
mind. Often texts on this subject are directed at a graduate audience
iin less “practical” information. Too often this subject is misunderstood and just a black box of “use-
ormation to the layperson. Dennis opens this box to view by including such practical topics
less” in
as: basic rules for constructing and using dichotomous keys, a how-to on collecting, handling, and
pre-
preserving specimens, and the role of botanical gardens in society. | really liked the chapter on
serving plant diversity that included such topics as: value of plant species to humans, how species
become pit sees enetic engineering and PION and how you can help endangered wild
gered, g g
A onthe relevance of systematics to society ending with the most prac-
stan
tical a all: job gppouiinities and qualifications. T he book stays true to its title and includes the im-
portant contemporary topics of phylogenetics and molecular systematics. Of course, the bulk of the
« deals with the classification of vascular plants beginning with the Pteridophytes. Dennis has
; from around the world to give a less North American-centric view of
boo
been careful to include group
the vascular plant kingdom with each group illustrated with botanical line drawings. My complaint
white. This gives
with the text is this: though it contains few photographs they are all in black anc
the text a rather outdated appearance. Tasik fhe text comes with a CD photo atlas of 4,700 color
images | feel a more contemporary text layout would benelit this text.—Robert J. George, Botanical
Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A., reeorge@ britorg
KATHRYN BERNICK. 1998. Basketry & Cordage from Hesquiat Harbour. (ISBN O-
7718-0525-9, pbk.). Royal British Columbia Museum, PO. Box 9815 Stn Prov,
Govt, 675 Belleville Street, Victoria, British Columbia, VSW 9W2, Canada.
$14.95, 152 pp., 114 figures, 6" x 9"
The Hesquait are a subdivision of the people of the Northwest a isl Suite Area and are plated to
the aboriginal inhabitants. The study of the plant-fiber
of Hersquait Harbor was undertaken as part of a larger project to aeonsUnee burial practices and
site use over time.
| icul ly d ibed {ill | by drawings all the techniques of manu-
The author has
facture of the recovered artifacts, and the material composition and plant names are given. In addi-
ied by use. Finally, there is a com-
tion aid are ae aoe of the actual objects, which are classif
of the findings and ae for further study
i
Jenth - | elarit 7
Ol
Taine a book for the general reader or primarily for botanists, but th
addition to the library of archaeologists, sce ae and basketry
the wor : make ita aul able
G Fort Worth, lexus.
t} “te :
enthusiasts. INGUICF Lit burg,
SIDA 20(1): 114. 2002
NEW TAXA OF CRATAEGUS (ROSACEAE)
FROM THE NORTHERN OKANAGAN-SOUTHWESTERN
SHUSWAP DIVERSITY CENTER
J.B.Phipps' R.J.O’Kennon
The University of Western Ontario ‘Botanical ens Institute of Texas
— of Plant Sciences n Street
1151 Richmond St. ae ae 102-4060, U.S.A.
London, Ontario, N6A 587, CANADA okennon@brit.org
jphipps@uwo.ca
ABSTRACT
Five new species and one new variety of Crataegus (Rosaceae) are described for the northern
Okanagan area of British Columbia. They are C. castlegarensis (ser. Douglasianae), C. shuswapensis @
ser. ian and C. piles la nensis var. wellsii, C. enderbyensis, C.orbicularis and C. atrovirens
(all ser. Purpureofructi). This generates a rich western North American center of diversity for
ee ik 13 species so far known from this newly recognized center. Four of the species de-
scribed here are endemic to the northern Okanagan but C. castlegarensis ranges widely to northern
Oregon, northeastern Utah and western Montana while C. okanaganensis var. wellsii extends into
the Okanogan of Washington. All the new taxa are illustrated and mapped. Revised keys for the
relevant series are provided.
Keyworps: Crataegus, Rosaceae, new taxa, systematics, Okanagan, diversity center.
RESUMEN
Se describen cinco especies nuevas y una variedad de Crataegus (Rosaceae) en el norte de la region
e Okanagan en la Columbia Britanica. Son C. castlegarensis (ser. Douglasianae), C. shuswapensis (
ser. Douglasianae), y C. es a var. wellsii, C. enderbyensis, C. Sadie ioe igre a
ser. Purpureofructi). Ee — un centro de diversidad rico en el oeste de Norte América para
Gralaegus con 13 esp ‘idas hasta el momento en este nuevo centro. a AHO de las especies
lémi del le Okanaga pero tl g Oregon
noreste de Utah y oeste de Montana, mientras que C. okanaga nensis var. wellsii se extiende por el
Okanoga de Washington. Todos los nuevos taxa se ilustran y cartografian. Se aportan claves revisadas
para las series relevantes.
The northern Okanagan of British Columbia and the southwestern part of the
adjacent Shuswap drainage is the richest area for Crataegus in North America
west of the Rocky Mountains. In addition to the five new species described here,
there are another three (Phipps & O’Kennon 1998) recently described species
(C.okennoniiJ.B.Phipps, C.okanaganensisJ.B.Phipps & O’Kennon and C. phippsii
O’Kennon), as well as four long-ago described species, C. chrysocarpa Ashe (incl.
C. piperi Britton = C. columbiana Howell, sens. auctt.), C. macracantha Lodd. ex
Loud., C. douglasii Lindl, C. suksdorfii (Sarg.) Kruschke, plus the introduced C.
monogyna Jacq., fora total of 13 species. This is enough to recognize a local diversity
SIDA 20(1): 115 — 144. 2002
116 BRIT.ORG/SIDA 20(1)
center for Crataegus (Fig. 1), by far the richest in western North America for an
area of this size. This diversity center is mapped in more detail in Fig. 2 where
many of the collection localities cited in this paper can be located. The contrast
with earlier understanding of Crataegus in British Columbia is striking. T.M.C.
Taylor (1973) listed only three species for the province, the native C. douglasii
and C. columbiana and the introduced C. monogyna, although the illustration
labelled C. columbiana is clearly C. macracantha as we have noted previously
(Phipps 1998).
The native hawthorns of the Okanagan-Shuswap belong toa diverse range
of series, not necessarily particularly closely related to one another, viz.:
Douglasianae, Purpureofructi, all the species of which occur in the region, and
Macracanthae and Rotundifoliae (sens. auctt.). The reasons for this rich variety
of hawthorn in the northern Okanagan and southwestern Shuswap drainage
cannot be spelled out precisely but presumably relate to the location of former
glacial refugia, as well as the benign climate. It is, of course, clear that wherever
the relevant refugia were at a given time they must have been constantly relo-
—
cated by changes of Pleistocene climate. In this mountainous region, hawthorns
are virtually restricted to the valley bottoms, become rarer southwards (the
southern Okanagan is much drier) and are nearly absent from the forest envi-
ronment of the middle slopes. In the somewhat mesic area of the northern
Okanagan, which is the area of greatest natural diversity, natural hedgerows
dominated by hawthorns are a feature of the agricultural landscape and haw-
thorns also occur along smaller watercourses and ditches as well as forming
downslope hillside thickets in some moister sites. The annual rainfall here is
around 500-600 mm but falls off to about 200 mm in the southern Okanogan.
In this paper we describe the new taxa C. castlegarensis (ser. Douglasianae),
C. shuswapensis (@ ser. Douglasianae), C. okanaganensis var. wellsii, C. orbicu-
laris, C.enderbyensis and C. atrovirens (all ser. Purpureofructi). Updated keys
are produced for the series in question. All species are illustrated with line il-
lustrations. Distribution maps of the new taxa were created by Range-Mapper
(Tundra Vole Software, Fairbanks, AL) from files of latitude-longitude coordi-
nates and then artistically improved by Susan Laurie-Bourque using Corel Draw.
The discoveries are the result of several years of fieldwork by the authors in the
mid-1990s and 2001-2 plus further discoveries in 2000 by the first author with
his wife Sheila and son William. The collections of May, 2002 substantially in-
crease the number of flowering specimens previously available for study and
from which the cited material derives.
—
TAXONOMY
In this part we set out the new taxa in the context of others in the same series by
providing a key to series and references to descriptions of the already known
PHIPPS AND O’KENNON, NEW TAXA OF CRATAEGUS FROM CANADA
| ies
DENSITY OF CRATAEGUS TAXA IN
N, AMERICA |
GRID
NES = LIMITS OF CRATAEGUS DISTRIBUTION
Ss
|
|
NO. OF SPP, OR SPP. GROUPS/540 KM*
HEAVY LI
x wo 200 Males
ener ne eanenene te
200 Kilemeter |
> ° 190 E
Lambert Reimythat fqva-Arta Proecton _ ae
- ; 0 20
Fic. 1. Locator map for the Okanagan diversity center. Diversit
western Shuswap; B = northwestern Montana; C = intermou
y cent gan—south
ntain Colorado.
118
BRIT.ORG/SIDA 20(1)
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oe
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ye
7
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eae
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Fic. 2. Detail map o
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with its ep
is, C. orbicularis, C.
icenter at Enderby. Unlabelled lines are 500m
h j 1 C. atrovirens
con
us
P
PHIPPS AND O’KENNON, NEW TAXA OF CRATAEGUS FROM CANADA 119
species in these series. In the cases of C. douglasii, which has been much con-
fused with other hawthorns in the area, and C. suksdorfii, both of which, as far
as we know, lack modern, detailed species descriptions, we also provide full
English descriptions. We also provide amended series descriptions. The depth
{ lobing of leaves is referred to by term ‘leaf incision index’ or‘LII CIFT in latin).
Series Douglasianae (Rehder ex C.K. Schneid.) Rehder.
Bushes to small trees; l-year old wood generally tan to deep purple-brown;
thorns mostly 1.5-2.5 cm long, dark and shiny when young. Leaves deciduous:
petioles fairly short; blades 3-5(-7) cm long, broadly elliptic or elliptic to some-
what obovate in general shape, shallowly (0-)2-3 lobed per side, the lobes ob-
tuse to subacute; margins toothed; venation craspedodromous to nearly fully
camptodromous in some unlobed forms; petioles and leaf teeth tips somewhat
glandular. Inflorescences 8-20 flowered; branches glabrous or pubescent,
bracteolate; anthesis early. Flowers 12-18 mm diam.; calyx-lobes subentire,
small; stamens 10 or 20, anthers pink; styles 3-5. Fruit smallish, 0.6-1.0 cm long,
ellipsoidal to subspherical, black or dark purple at full ripeness, hairy or gla-
brous; nutlets 3-5, dorsally furrowed, lateral faces pitted.
This is a small series of + widespread ‘black-fruited’ hawthorns of the
northwestern USA and adjacent Canada with four species, one of which, C.
douglasii, extends to the Great Lakes area. Series Douglasianae are notable for
their laterally pitted nutlets, short thorns and dark purple to black fruits at full
maturity in which they resemble Eurasian ser. Nigrae. We add one new species
to this group and amend the key.
KEY TO SPECIES
—
Stamens 20;leaf-blades often unlobed, if lobed, lobes generally obtuse to subacute;
when lobeless venation semi-camptodrom 1. C.suksdorfii
. Stamens 10; leaf-blades usually lobed; oe of leaves subacute to acute; venation
craspedodrom
2. Inflorescence ns anches glabrous; thorns on twigs always a fruit longer than
broad, ae chestnut, dark ea ple or black in late Au
=
3. Thorns 2-3.5 cm long; flowers 12-15 mm diam; fruit aan usually ia
oie or even black by i ee very dae aie at this time 2.C. douglasii
3. Thorns 1.5-2 cm long; flowers 15-18 mm diam; fruit ampulliform, chestnut
or burgundy in late August, eeeciin not very bloomy 3.C. okennonii
2. Pr creer branches and nypantnln m + pubescent; thorns on twigs occa-
sionally paired or in triads; fruit subglobose to depressed, passing through crim-
son-lake (purplish-crimson) to burgundy in late August 4.C. castlegarensis, sp. nov.
1. Crataegus suksdorfii (Sarg.) Kruschke, Milwaukee Public Mus. Public. Bot.
3:163. 1965.
Crataegus douglasti Lindl. var. suksdorfii Sarg., Bot. Gaz. 44:65. 1907.
Bushes, often 3-5 m tall, perhaps larger; one-year old twigs tan, older grayish;
thorns 1.5-2.5 cm long, recurved or straight, glossy dark brown when young.
120 BRIT.ORG/SIDA 20(1
Leaves deciduous: petioles 0.75-L5 cm long, glabrous, eglandular westwards, in
Montana glandular, blades + rhomb-elliptic to ellipt-obovate in general shape,
3-6 cm long, tapered to the base, unlobed to irregularly sharply to very bluntly
2-3 lobed per side; margins toothed, the teeth in Montana glandular; venation
craspedodromous to semi-campdodromous in blunt-lobed western specimens;
appressed hairy above, + glabrate below except pubescent near the veins in
Oregon. Inflorescences 7-15 flowered, branches glabrous, punctate, bearing
small, early caducous, linear, membranous, gland-margined bracteoles. Flow-
ers ca. 15-17 mm diam. hypanthium glabrous externally; calyx-lobes ca. 4 mm
long, triangular, abaxially glabrous, margins minutely and sparingly glandu-
lar-serrate: petals + circular, white; stamens 20, anthers pink; styles 4-5. Fruit |
cm long, subglobose to broad-ovoid in the few specimens seen, black at matu-
rity, dull vinous purple a month before; calyx-lobes reflexed; nutlets 4-5, dor-
sally grooved, sides erose.
Crataegus suksdorfii occurs mainly along the coast and ranges from California
northwards to Vancouver Island, the Queen Charlotte Islands and coastal Alaska
to around Anchorage. It is also scattered sporadically inland east to Montana.
—
Crataegus suksdorfii is by far the most northerly occurring Crataegus in North
America at 61’ N lat. It is diploid according to Brunsfeld and Johnson (1990).
In the relatively few specimens I have studied, the fruit is smaller and more
globose than in C. douglasii. It is most convincingly differentiated from C.
douglasii by stamen number and perhaps by fruit shape and early color, but
like C. douglasii, it also has a variety of leaf shapes and a venation type which
lie generally outside the range of that species. The glandular leaf-teeth and
petiole make the Montana specimens distinct. Some of the deeply dissected,
pointed lobed, + delt to rhomb-ovate leaved forms from the western US appear
very distinctive.
2. Crataegus douglasii Lindl. Bot. Reg. 21: pl. 1810. 1835.
Crataegus columbiana Howell, FL Northw. Amer. 1:163. 1898.
Bushes 4-6 m tall, thorny; thorns + stout, 1.5-3.5 cm long, straight or slightly
recurved, deep brown and shiny when young; one-year old twigs deep tan to
dark mahogany, somewhat shiny when young, older branches gray to dark gray,
often showing brown or alternatively tan-brown eastwards. Leaves deciduous;
petioles 0.75-1.5 cm long, slightly glandular and pubescent in youth; blades 4-
7 cm long x 2-3cm wide, somewhat variable in shape but generally elliptic to
broad-elliptic or sub-rhombic, generally with 2-4 coarse lobes per side, alter-
natively elliptic-obvate with much smaller lobes; margins coarsely and sharply
toothed, the teeth tipped with small glands when young; venation
craspedodromous with 4-5 lateral veins per side; quite densely appressed short-
pubescent above, below usually glabrous except on the veins. Inflorescences
10-25 flowered; branches usually glabrous, bearing caducous, membranous,
PHIPPS AND O’KENNON, NEW TAXA OF CRATAEGUS FROM CANADA 121
linear, gland-margined bracteoles. Flowers 12-15 mm diam.; hypanthium ex-
ternally glabrous; the calyx-lobes very short, 3-4 mm long, fairly broad trian-
gular, adaxially glabrous, the margins with a few tiny glands distally; petals +
circular, white; stamens 10, anthers pink: styles 3-4. Fruit 6-8 mm diam., ellip-
soid, dull black when fully ripe, though vinous when younger, generally very
bloomy until maturity; calyx-lobe remnants short, blunt, reflexed; nutlets 3-4,
dorsally grooved, laterally excavated.
Douglas hawthorn is a common, somewhat variable bush of southern and
western British Columbia up to the panhandle of Alaska which ranges south
to northern California (Shasta Co.) and inland to the Rocky Mountains of Idaho,
Montana and sw Alberta. Further east, disjunct populations occur in the Cy-
press Hills (AB & SK), Black Hills WY & SD), and around the northern Great
Lakes. In western North America Crataegus douglasii is found mainly near
water in drier areas but in more mesic localities ina good variety of open wood-
land. In the Great Lakes area it mainly occurs in old pastures and on fence lines.
Crataegus douglasii is distinguished from C. suksdorfii which has 20 sta-
mens, semi-camptodromous venation and little lobed, + obtuse leaves, and from
the taller C. okennonii which has generally shorter thorns, usually straight
single trunk, ampulliform-orbicular fruit and crimson-lake autumnal foliage
color. Crataegus douglasii is mainly tetraploid (Brunsfeld et al. 1990).
3. Crataegus okennonii J.B. Phipps, Sida 18:170. 1998.
This species commenced flowering 10-14 days later than C. douglasii in the
northern Okanagan in 2002.
4. SS castlegarensis J.B Phipps & O’Kennon, sp. nov. (Fig. 3). Type: UNITED
TES. IDAHO. Lemhi Co: US 93, E side, 14.5 rd. mi N of Salmon, roadside ditch next to
ene field, alt. ca. 3950 ft, bush, 5 m tall, foliage dull, dark, coriaceous, fruit strongly
clustered, red-burgundy, with short pedicels, 20 Aug 1996, J.B. Phipps & O’Kennon 7396
(HOLOTYPE: UWO; IsoTyYPES: BRIT, CAN, MO, MONTU, TRT, UBC, V, WS).
lel : 1
: rutices, 2.5-5.0 m alti, unc
2-3cm longae, mediana nade rectae vel leviter recurvatae, nitenter atrobrunneae, apice atro in
ee Folia decidua; petioli 0.75-1.5 cm longi, varie pilosi, eglandulosi; laminae 3.5-6.0 cm longae,
oblanceolatae vel ovato-rhombeae in forma generali; apex late triangularis, basis late cuneata vel
cuneata; vix lobata 3-4 lobatae per aoe eliduantum coriaceae; venatio . craspedodroma, 4-5
nervatae pe if aliquantum
glabrescens in vena atc: abaxialis rare puboreens a glabra venis eee. Inf fein ae 8-20
florae; pedicelli leviter vel dense pubescente linearibus, membranaceis glandulo-
marginatis. Flores ca. 12-15 mm vel iam. SE Berumaue glabrur um extrinsecus; lobi See 3
1¢
mm longi tri angulares | serrati; petala + I
4. Fructus 10-12 mm diam., orbiculares ee oblati, recedentes i in bjunctans cum pedicellis,
sparsim pilosi, carmesini pur] p reflexi,
+ erosi; pyrenae 3-4, dorsaliter sulcatae BR racist lil | f |
Shrubs, 2.5-5.0 m tall with one or more erect stems; — spreading; l-year
old twigs brown; thorns 2-3 cm long, of medium stoutness, straight to slightly
122 BRIT.ORG/SIDA 20(1)
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PHIPPS AND O’KENNON, NEW TAXA OF CRATAEGUS FROM CANADA 123
recurved, shiny dark or reddish-brown with blackish tips when young. Leaves
deciduous; petioles 0.75-1.5 cm long, eglandular, variably hairy; blades 3.5-6.0
cm long, oblanceolate to ovate-rhombic in general shape; the tip broad-trian-
gular, base broad-cuneate to cuneate; scarcely lobed to 3-4 lobed; margins
toothed, teeth with fine gland-tips when young; venation craspedodromous,
each side 4-5 nerved; conspicuously appressed hairy above when young, some-
what glabrescent except on the mid-nerve above, thinly pubescent to glabrous
below except on the veins, somewhat coriaceous. Inflorescences 8-20 flowered:
branches thinly to densely pubescent, bearing caducous, linear, membranous,
gland-margined bracteoles. Flowers ca. 12-15 mm diam, hypanthium gener-
ally glabrous externally; calyx lobes 3 mm long, triangular, remotely glandu-
lar-serrate; petals + circular; stamens 10, anthers pink; styles 3-4. Fruit 10-12
mm diam., + orbicular, often somewhat oblate, recessed at junction with pedicel,
thinly pilose, crimson turning to reddish-plum or ultimately blackish purple;
calyx-lobes patento-reflexed, + erose; nutlets 3-4, dorsally grooved, lateral faces
usually pitted.
Crataegus castlegarensishasa wide range (Fig. 4) from the John Day (South
Fork) R. basin of Oregon and the Okanagan of British Columbia eastwards to
the Flathead drainage of northwestern Montana, thence south down the moun-
tains to northeastern Utah and west-central Wyoming. We have also recently
recorded it from the Cypress Hills of Alberta and Saskatchewan but those speci-
mens are not cited. It is most abundant north of Kelowna, British Columbia
and in the Flathead drainage in Montana and is found in brushy places in the
valleys, particularly near ditches and natural waterways. Fine specimens of this
species occur on the grounds of Selkirk College, Castlegar, British Columbia,
where several were collected for us by Peter Wood, then a professor at that insti-
tution. It was here that the authors first became aware of the distinctness of
this taxon and we are pleased to name it after the town of Castlegar.
Like a good number of Crataegus species from the Pacific Northwest C.
castlegarensis has been collected but overlooked and the few specimens that we
have encountered other than our own collections have been variously assigned
to C. douglasii and C. columbiana. Crataegus castlegarensis is apparently the
most closely related of ser. Douglasianae to C. douglasii. It differs in the fruit
shape (¢ orbicular/turbinate in C. castlegarensis; ellipsoid in C. douglasii), fruit
color at comparable period of ripening (often dull plum blotched green early,
then burgundy to crimson, finally deep purple and not very bloomy in C.
castlegarensis; starting about the third week of August as for the former species,
usually already full black and remaining very bloomy unless abraded in C.
douglasii); inflorescence branches and hypanthium, also, often fruit, proximally
(pubescent to dense pubescent in C. castlegarensis, glabrous in C. douglasii).
Pubescence of parts may abrade through the season and one may have to look
124 BRIT.ORG/SIDA 20(1)
110°. -105°
a i
ad
Fic. 4. Distrit +, £ ‘i } ic ¢ Ilatad A
very carefully to discern it on fruiting pedicels. The junction of the fruit to the
pedicel, often recessed like an apple, is also a characteristic of the new species.
It came into flower about a week later than C. douglasii in the northern
Okanagan in 2002.
fey
A very interesting feature that we record is branched thorns (double to
quadruple) on the young twigs. This feature is not known to us in other haw-
thorns except for the mutation of C. monogyna,cv. Horrida and occasional speci-
mens of C. enderbyensis. On some of the bushes this multiple thorniness was
abundant, in others more sporadic. We did not deliberately collect for this fea-
ture, indeed were not conscious of its relationship with C. castlegarensis early
on, so it is possible that a more thorough check in the field of many of the col-
lected bushes may have yielded more material like this. Branched thorn speci-
mens are asterisked in the list of exsiccatae.
Additional specimens examined: CANADA. BRITISH COLUMBIA: Castlegar, N side of Frank Beinc
rive, ca. LO ft down the embankment below the roadway, alt. 1550 [t, slender hawthorn, thorns 2.5-
2.7 cm long, anthers creamy pink, 15 May 1995, Peter Wood 1(UWO). Castlegar, top of Frank Beinc
er
er
Rd. to Selkirk College, opposite airport, scrubby small ravine, alt. 487 m, bush, 5 m tall, narrow-lan-
ceolate leaves % obtuse lobes, purple fruit in dense clusters, 26 Sep 1993, J.B. Phipps & RJ. O’Kennon
PHIPPS AND O’KENNON, NEW TAXA OF CRATAEGUS FROM CANADA 125
6823* (BRIT, CAN, ID, MONTU, RM, UBC, UT, UWO, V). Pleasant Valley Rd. NNE of Vernon, E side of
road, ca. 0.5 mi N of Vowle’s property, in bottom of ravine in apple orchard, 5 m bush, black fruit, 3-
forked thorns, 25 Sep 1993, J.B. Phipps & RJ. O’Kennon 6818’ (BRIT, UWO). Castlegar, Selkirk College,
top of wooded steep slope near Peter Wood's olfice, alt. LOO0 ft, collection of different leaves from 4m
bush, 27 Sep 1993, ].B. Phipps & RJ.O’Kennon 6825 (UWO). Okanagan Valley, ca.8 km NNE of center
of Vernon, Pleasant Va me Rd., poune igen s residence, hedge at S boundary of Burke’s PYO, alt. 380
, dense, broad, mature hedg y hawthorns, bush 2.5 m tall, anthers 10, pink, 02 May
1994, J.B. Phipps 6881* (BRIT, ‘UBC. UWO). on amalka L. Provincial Park, along road ca. 25 m SW of
red gate, brushy roadside, alt. ca. 1450 ft, bush, 5 m tall, fruit burgundy turning black, foliage large
(equals JBP 6888), 18 Aug 1994, J.B. Phipps & RJ. O’Kennon 6980* (UWO). Northern Okanagan,
Kalamalka Provincial Park, r eae near red gate, bush, 6 m tall, fruit ie + spherical to short
turbinate, locally abundant, 26 Aug 2000, J.B. & S.R. Phipps 8187* (UWO). Castlegar, top of Frank
om
ie
Oo.
Beinder Rd. to Selkirk College, opposite ean in bottom of are roadside ravine, alt. 487 m
bush, 5 m, stamens 5, anthers small, pink, (-JBP & RO’K 6823), 06 May 1994, J.B. Phipps 6925 (UBC.
UW0O). Castlegar, top of Frank Beinder Rd. to Selkirk College, opposite airport, bush 2 m tall, half-
way aa ace before ‘curve road’ sign, stamens 5, anthers pink, 06 May 1994, mee J.B.6926 (UWO).
7 SW of Princeton on BC3, edge of Typha marsh on SE side of hwy, alt. 2800 ft, bush, 3.5 m
tall, a orbicular, burgundy, beginning to be ee by slnpuiese lvs thin, coriaceous, shiny (=/BP
& RO’K 6774), 17 Aug 1994, J.B. Phipps & RJ.O 6972 (BRIT, CAN, MONTU, TRT, UBC, UWO,
V). Castlegar, Frank Bender me sais slopes “ith aspens, ca. 250 m E of campus gate, alt. 1550
ft, tree,6m tall, fruit li 23 Aug 1994, J.B. Phipps & RJ.O’ Kennon
7053 (BRIT, UBC, UWO). Castlegar, Fri ane Beinder Drive. sc ae slopes with aspens, ca. 75m
campus gate & behind pine, alt. 1550 ft, bush 2.5 m tall with purple-black globose fruit, 23 Aug cot
J.B. Phipps & RJ. O’Kennon 7055 (BRIT, UBC, UWO). Castlegar, Frank Beinder Drive, opposite airport
entrance, at top of little ravine virtually within reach of road, alt. 1600 ft, bush, 5 m tall, fruit orbicu-
lar, eke eek a reddish plum, 23 Aug 1994. J.B. Phipps & RJ.O’Kennon 7054 (BRIT, UWO,
V). road mi SW of Princeton on BC3. edge of large dried up marsh on E side of hwy, bush 10 ft
tall eee reddish, fruit black-burgundy, 23 Sep 1993, J.B. Phipps & R,J.O’Kennon 6774 (UWO). Sa
Arm, large hawthorn pasture just N of railroad and west of central business section, alt. ca. 1900 ft,
bush, 4 m tall, fruit deep burgundy, + spherical, 20 Aug 1994, [.B. Phipps & RJ.O’Kennon 7030 (UWO).
Vernon, Y Lake, open aspen grove near pond, 10 May 1970, T.M. Brayshaw & D. Laidman s.n.* (UWO
V)
mon
numerous triple and aoe thorns—J.B.P.
93, 4 mi S of 45°N lat sign and 18 road mi S of Salmon, 4600 ft,
gravelly banks of Salmon R, ae 5m tall, 17 May 1997, J.B. Phipps & RJ. O’Kennon 7532 (BRIT, UT,
UWO). US 93,17.5 mi Sof Salmon, brushy banks of Salmon R.. bush, 4 m tall, flowers 16 mm diam., 17
May 1997, J.B. Phipps & RJ. O’Kennon 7535 (BRIT, ID, UWO); US 93, 14.5 mi N of Salmon, E of side
road, fenceline against pasture, alt. 4300 ft, bush, 5m tall, inflorescence very pubescent, tight cluster
Q
aoa"
a)
of flowers, flowers diameter greater than or equal to 15 mm, anthers 10, pink, some paired thorns,
(equals JBP 7396), 17 May 1997, J.B. Phipps & RJ. O’Kennon 7537* (BRIT, ID, MONTU, UWO); US 93, 17
rd. mi N of Salmon, brushy banks of South Fork, Salmon R., alt. 3900 ft, bush 4 m tall, 17 May 1997,
J.B. Phipps & RJ. O° Senmon Le reaee ID, MONTU, UT, UWO). US 93, at pee of gravel oa near
Salmon River, ca. 18 rd. mi S of Salmon and ca. 4 mi Sof 45° N lat. sign, alt. 4600 ft, several-stemmed
bush 5 m tall, _ cna ripe: green-light red-dull red-burgundy, + oblate- ae ey with ie
depression at pedicel, lvs coriaceous, blue-green, broad, lobed, frt. in dense clusters, short icin
small ee bracteoles, 20 Augl996, J.B. Phipps & RJ. O’Kennon 7394 (BRIT, ID, MONTU, UT, UWO);
US 93,8 mi N of Salmon, 2.5 mi S of Tower Creek, in field, alt. ca. 4300 ft, dense bush, 4m ae leaves
ee blue-green, somewhat cupped, fruit orbicular i ee pedicel, green to dull
red, 20 Aug 1996, J.B. Phipps & RJ.O’Kennon 7395 (ID, UT, UWO); U 4rd.mi N of Salmon, west
side, roadside ditch, alt. 3900 ft, bush, 2 m tall, fruit dull purple- a oe ne 1996, J.B. Phipps & RJ.
O’Kennon 7397 (UWO). Payette Co. Hwy. 53 ere in floodplain of Payette River, ca. 9 rd. mi SE of
126 BRIT.ORG/SIDA 20(1)
Payette, alt. 2300 ft, slightly swampy fields, 27 Aug 1994, J.B. Phipps & RJ. O’Kennon ne (UWO).
Washington Co.: US 95 at Monroe C es ca. a mi ° of aie (MP 88.4), alt. ca. 2700 [t, brushy area,
bush, 5m tall, fruit large, spherical, red y,27 1994, J.B. Phipps & | O’Kennon
7095 (BRIT, ID, MONTU, RM, UT, UWO). MONTANA: Flathe ad Co.: Hwy 35, ca. 0.5 mi SE of Creston,
alt. 3050 ft, dense bush 3 m tall, fruit orbicular, dull red, 25 Aug ood |B. Phipps & RJ.O’Kennon 7071
(MONTU, RM, UWO). Junction of state road 35 and unpaved road to Fish Hatchery, small ae
thicket in old pasture, alt. ca. 2800 ft, bush, 3 m tall, flowers at popcorn, anthers pink, 20 May 196
J.B. Phipps & RJ. O’Kennon 7589 (MONTU, UWO). Flathead River Valley, 1 mi E of 206, ca. 3 mi SE
Columbia Falls center on dirt extension of S end Berne Jensen) Rd., 3100 [t, large 6 m bush near
pond, (=/BP & RO’K #6838), stamens 10, anthers pink, coll. 8 May, buds forced, 08 May 1994, ].B. Phipps
6938 (BRIT, ID, MONTU, RM, UBC, UWO),. Flathead River Valley, | mi E of 206, 3 m SE Columbia
Falls center at extreme S end of Jensen Rd., by pond, alt. 3100 ft, 5 m bush, fruit orbicular, purple-
black, 27 Sep 1993, ].B. Phipps & RJ. O’Kennon 6838 (BRIT, ID, MONTU, RM, UBC, UwO). wens Co.:
State Road 35 E of Polson, between Turtle Lake Road and Ducharme, Indian Agency lands with stream,
mature ponderosa pine and mown grass, along water edge, alt. 2950 ft, bush, 6 m tall, twigs with
some bi-thorns, in bud, anthers pale pink, 19 May 1997, ].B. Phipps & RJ.O’Kennon 7581* (UWO); Old
Freight Rd., off US 93 W of St. Ignatius, hawthorn thickets in creek beds between dry grassland, at
roadside, alt. 2900 ft, bush 4 m tall, leaves rhomb-ovate, little lobed, badly rusted, fruit broad-
ampulliform to suborbiculate, red to purple-black, common, Aug 21 1996, J.B. Phipps & RJ.O’Kennon
7409 (BRIT, MONTU, RM, UWO); Old Freight Rd., off US 93 W of St. Ignatius, hawthorn thickets in
creek beds between dry grassland, at roadside, alt. 2900 ft, bush, 4 m tall, leaves rhomb-ovate, little
lobed, badly rusted, fruit broad-ampulliform to suborbiculate, red to purple-black, common, 21 Aug
1996, J.B. Phipps & RJ. O’Kennon 7410 (UWO); ca. 0.5 km up Old Freight Rd. from US 93, hawthorn
thickets along creek, alt. 3000 ft, 17 May 1997, J.B. Phipps & RJ. O’Kennon 7552 (MONTU, UWO); Old
Freight Road off Hwy 93, within sight of Hwy 93, milepost 30.4, 3 mi N of Ravalli, 10 pink anthers,
15-22 flowers per inflorescence, pedicels pubescent, calyx glabrous, medium thorns, 18 May 1995,
RJ. O’Kennon 13162 (CUWO); Old ee Road oe iy 93, acne oo of | oy 93, milepost 30.4, 3
mi N of Ravalli, 17-22 flowers per inflores ) pin] calyx glabrous
looks like a 18 May 1995, RJ.O’Kennon ea WO): Old Breioht Ros ad on Hwy 93, within sight
of Hwy 93, milepost 30.4, 3m N of Ravalli, 17-22 flo vers per inflorescence 10y kantl s, full flower.
pedicels and calyx pubescent, 18 May 1995, RJ. O’Kennon 13165 (UWO). Lineal Co.: Hwy 93, be-
tween Roosville &@ Eureka, W side of rd., ca. 1.0 mi N of junction with 37, seepage slopes below irriga-
tion channel, on hillside, alt. 2850 ft, bush 4 m, crus-galli like foliage, frt. orbicular, crimson chang-
ing to nie 24 Aug 1994, J.B. Phipps & RJ. O’Kennon 7062 (BRIT, MONTU, UWO). Sanders Co.:
Little Bitterroot R. near confluence with Flathead R., alt. 2700 {[t, bush 4 m tall, fruit + or-
bicular, a red, 21 Aug 1996, J.B. Phipps & RJ. O’Kennon 7418 UD, MONTU, UWO); 7 km direct NE
of center of Camas Prairie Basin on road from Little Bitterroot Valley, close to Schmilz Lo ac
cess road, seepage zone in prairie, alt. 3000 ft, bush, + m tall, anthers 10, pink, 18 May oe Phipps
& RJ. O’Kennon 7576 (BRIT, MONTU, UWO), at Little Bitterroot R., at eastbound road crossing ex-
actly 20 km SE of Hot Springs, Evan Melton ranch, thickets at edge of water, alt. ca. 2800 ft, bush, 4m
tall, anthers 10, pink, 18 May 1997, J.B. Phipps & RJ. O’Kennon 7571(BRIT, MONTU, RM, UWO). Ca. 24
km (direct) SE of Hot Springs, hawthorn thickets along banks of Little Bitterroot R., alt. 2800 ft,
bush, 3 m tall, anthers 10, pink, 18 May 1997, J.B. Phipps & RJ. O’Kennon 7570 (BRIT, MONTU, RM,
UWO); ca. 24 kim (direct) SE of Hot Springs, hawthorn thickets along banks of Little Bitterroot R.,
alt. 2800 ft, bush, 3 m tall, anthers 10, pink, 18 May 1997, J.B. Phipps & RJ. O’Kennon 7569 (BRIT, ID,
U, UT, UWO); hawthorn thickets with pasture along Little Bitterroot R., ca. 24 km SE of Hot
BD
»
Springs, Melton Sr. (grandfather) ranch, alt 2750 ft, bush 3 m tall, leaves broad, fruit burgundy, or-
bicular, Aug 211996, J.B. Phipps & RJ.O’Kennon 7423* (UWO), hawthorn thickets with pasture along
Little Bitterroot R.,ca.24 km SE of Hot Springs, Melton Sr. (grandfather) ranch, alt 2750 ft, bush 3.5m
tall, foliage intermediate, fruit burgundy, orbicular, 21 Aug 1996, J.B. Phipps & RJ. O’Kennon 7425
PHIPPS AND O’KENNON, NEW TAXA OF CRATAEGUS FROM CANADA 127
(BRIT, UWO); banks of Little Bitterroot R.ca. half mi S of confluence with Flathead R., bush 3 m tall,
18 May 1997, J.B. Phipps 7561 & RJ. O’Kennon* (MONTU, UWO); hawthorn thickets along bank of
Little Bitterroot R., abt 24 km direct SE of Hot Springs, 2800 ft, bush, 3 m tall, 18 May 1997, J.B. Phipps
& RJ. O’Kennon 7566 (BRIT, MONTU, UWO); hawthorn thickets along bank of Little Bitterroot R.,
abt 24 km direct SE of Hot Springs, 2800 ft, bush, 3 m tall, 18 May 1997, J.B. Phipps & RJ. O’Kennon
7568 (BRIT, MONTU, UWO),; crossing of Little Bitterroot R. exactly 20 km SE of Hot Springs, thickets
at edge of water, Evan Melton ranch, 2800 ft, bush 4 m tall, 18 May 1997, J.B. Phipps & RJ. O” Kennon
7573 (BRIT, MONTU, RM, UWO); main road SE from Hot Springs along Little Bitterroot River at 5m
due N of major NS/EW bend (near location of 7420) at Malinak mailbox, alt 2800 ft, two haw ee
pees ditch on west side of road, healthy bush, 4 m tall, 21 Aug 1996, J.B. Phipps & RJ. O’Kennon 7427
O), Hot Springs-Perma Rd., Camas Prairie Basin, ca. 4 mi S of Camas Prairie, alt. 2850 ft, large
a 6m tall, fruit burgundy-red, 21 Aug 1996, J.B. Phipps & RJ. O’Kennon 7429 (UWO); Little Bitter-
root River at eastbound road crossing exactly 20 km SE of Hot Springs, road from Hot Springs to
Evan Melton ranch, alt. 2790 ft, open thicket of hawthorns in pasture near creek, bush 3 m tall, |
blue-green, fruit deep burgundy, orbicular, with recessed pedicel, locally frequent, 21 Aug 1996, J.B.
Phipps & RJ. O’Kennon 1420 (MONTU, RM, UWO). OREGON: Grant Co.: John Day Valley, S fork,
Humphreys, small tree, 30 Apr 1925, L.F Henderson 5146* (CAS, DS, photo UWO). UTAH: Cache Co
Sof Hyrum Reservoir at 7800, off 4008S, tenceline, S side of dirt road, alt. 4800 ft, bush, 5 m tall. eal
flowers, LO pink anthers, 15 May 1997, |.B. Phipps & RJ. O’Kennon 7528 (BRITID,UT,UWO): E side of
south end of Hyrum Reservoir across field from intersection of 7000W and 4008, hedgeline, alt. 4800
ft, bush, 4 m tall, flowers small, anthers 10, deep pink, 15 May 1997, J.B. Phipps & RJ. sa 1527
(BRIT, ID, MONTU, NY, RM, UBC, US, UT, UWO). Weber Co.: Huntsville, on N side of U
after it turns E of Pineview Reservoir, roadside with ditch backing mee fields, a : m tall, but
early, white, forced, 10 pink anthers, 4 green styles, inflorescences more floriferou-s and dense than
C. rivularis, 15 May 1997, ].B. Phipps & R,J. O’Kennon 7526 (UWO); UT 39 ca. 1/2 mi after it turns E, E
of Pineview Reservoir, roadside with ditch (water flowing) backing on to irrigated field, 2 adjacent
bushes, ca. 6 m tall, eas foliaged, Ivs. blue-green, young twigs & thorns say olive-brown, frt.
orbicular, recessed at pedicel contact, green tur a light to dull red to vinous to black, on same tree
18 Aug 1996, J.B. Phipps & RJ.O’Kennon 7389 (BRIT, ID, MONTU, RM, UT, UWO); ravine to E of SR.
167 (Trappers’ Loop Rd.), along creek, alt. 5200 : bush 4 m tall, abundant, 22 May 1997, J.B. Phipps &
RJ. O’Kennon 7608 (BRIT, MONTU, UT). WASHINGTON: Okanogan Co.: Chopaka Rd., 2.75 mi from
Loomis-Nighthawk Rd. junction, 28 Aug 1994, J.B. Phipps 7106 & RJ. O’Kennon (UWO). Spokane Co.:
S of Deer Park, 0.5 mi Sof milepost 179 on US 395, 30 Sep 1993, .B. Phipps 6861 & RJ.O’Kennon(UWO).
Whitman Co.: Staley Rd., 2 km NE of Chambers, floodplain of small creek, dense small tree 4 m tall,
fruit orbicular, red changing to burgundy, 27 Aug 1994, J.B. Phipps 7097 & RJ. O’Kennon* (BRIT, UBC,
S, WTU). WYOMING: Lincoln Co.: US 89, S of Thayne and 0.4 mi N of Salt River, floodplain
srasties with hawthorn thickets, alt. 5950 ft, bush, 4 m tall, inflorescence pubescent, oven 10,
pink, in green bud, forced, M ee 1997, ].B. Phipps & RJ. O’Kennon 7597* (BRIT, MONTU, RM, UWO);
US 89S of Thayne at 0.4m Salt River crossing, fenceline in flood plain, alt. 5950 ft, = 3.5m
tall, (= JBP 7462), in tight a Brake 16 May 1997, |.B. Phipps & RJ. O’Kennon 7531* (RM, UWO); US
89, S of Thayne and 0.4 mi N of Salt R. crossing, 2.5 mi N of junction with 238, fencelines in flood-
plain, alt. 5950 ft, bush, 3.5 m tall, leaves deep green, somewhat lobed fruit red to plum-red, orbicular,
held erect, 25 Aug 1996, J.B. Phipps & RJ. O’Kennon 7462 (BRIT, UWO). Teton Co.: Targhee National
Forest, US 89, 5.4 mi S of Hoback Jct., flats of Snake River, alt. 6200 ft, dense brushy area with tall
trees of cottonwood and pine, bush, 6 m tall, foliage narrow, fruit spherical, colors greenish to red to
plum to burgundy, 25 Aug 1996, |.B. Phipps & R.J. O’Kennon 7460 (BRIT, MONTU, RM, UT, UWO).
Targhee National Forest, flats of Snake R., 5.4 mi S of Hoback Jct., alt. 6200 ft, 21 May 1997, J.B. Phipps
7593a* & RJ.O’Kennon (UWO). Washakie Co.: US 16, 1 mi E of Tensleep, river flats of Tensleep R., in
sagebrush country, alt. 4550 [t, bush, 4 m tall, badly rusted, leaves thin, (one bush was non-rusted),
24 Aug 1996, j.B. Phipps & RJ. O’Kennon 7456 (UWO)
fruit suborbicular, plum-colorec
128 BRIT.ORG/SIDA 20(1)
Incertae Sedis
Crataegus shuswapensis J.B. Phipps & O’Kennon, sp. nov. (Fig. 5). Type: CANADA;
BRITISH COLUMBIA: Northern Okanagan, Enderby, ca. 200 m E of bridge over
Shuswap River on Mabel Lake Rd., bush, 3.5 m tall, fruit deep purple (=JBP ae
20 Aug 1994, J.B. Phipps & RJ. O’Kennon 7009 (HOLOTYPE: UWO; ISOTYPES: BR
CAN, DAO, TRT, UBC).
Frutices ad 5 m alti; ramuli unius anni glandulacei vel atrobrunnei; spinae 2-3 cm longae, + validae,
rectae vel leviter recurvatae, in juventute atrobrunneae. Folia decidua; petioli 1.5-2.25 cm longi,
sparsim glandulo-maculati, leviter pubescentes: laminae 5-7 cm longae, rhombeae in forme eee
lobis acutis, ca. ai lobatae aa ae cum a | — ee venatio craspedodroma, venis secondariis +5
per | presso-pubescens, abaxialis glabra.
ini lereseentise 12-18 florae; ramuli glabri, cum ale bracteolis caducis linearibus, glandulo-
marginatis. Flores ca. 15 mm diam., hypanthium extrinsecus glabrum; lobi caleycis 4-5 mm longi,
p24 i E a J F g
triangulares, minute glandulo-marginati; petala circularia; stamina 1015-18), antheris pallide vel
clare roseis vel rare tyriis; carpelli et styli 2-3. Fructus 12 mm longi, + late-ellipsoidei, in juventate
roseo-vinosi, atropurpurei in maturitate; reliquiae calycis adsunt, patento-recurvatae; pyrenae 2-3,
dorsaliter sulcatae, in lateribus planae.
Bushes to 5.5 m tall: one year old twigs tan to deep brown; thorns 2-3 cm long,
stoutish, straight or slightly recurved, very dark brown when young. Leaves
deciduous; petioles 1.5-2.25 cm long, sparsely gland-dotted, slightly pubescent;
laminas 5-7 cm long, rhombic in general shape; ca. 4-lobed per side, LII to 25%,
venation craspedodromous, with 4-5 pairs of secondary veins; lobes sharp;
margins remotely serrate; somewhat sparsely appressed-pubescent above, gla-
brous below. Inflorescences 12-18 flowered, the branches glabrous, some linear
caducous gland-margined bracteoles present. Flowers ca. 15 mm diam.; hy-
panthium glabrous externally; calyx-lobes 4-5 mm long, triangular, minutely
panne ee petals circular; stamens 1015-18), anthers pale to bright pink
or rarely puce; carpels and styles 2-3. Fruit + broadly ellipsoidal, 12 mm long,
claret color at first, deep purple at maturity; calyx remnants present, patento-
recurved; nutlets 2-3, grooved dorsally, smooth on sides.
Crataegus shuswapensis is locally common near Enderby in the northern
Okanagan and southwestern Shuswap drainage, British Columbia. Collection
localities may be identified, at least approximately, on figure 2.
This is a very distinctive hawthorn on account of its rhombic, wide and
quite deeply and sharply lobed leaf-blades, stamens with anthers pink or some-
times puce and fairly large suborbicular to broad-ellipsoid purple fruit. The
flowers, and to some extent the fruit, are much like C. douglasii but other char-
acteristics, the rather long stout thorns, leaf shape, fruit color about the third
week in August, 2-3 styles and nutlets point in a different direction. Flowering
material with 10 pink stamens from near Polson, Montana, may be the same
—_—
and fruiting specimensare required to verify this. This species flowers synchro-
nously with C. castlegarensis in the northern Okanagan.
129
PHIPPS AND O’KENNON, NEW TAXA OF CRATAEGUS FROM CANADA
1cm.
15 stamen form. Scale bars
Fic. 5. Li
130 BRIT.ORG/SIDA 20(1)
Additional specimens examined: CANADA. BRITISH COLUMBIA: Valley of the Shuswap, 8.0 km EF
OE es little to E of Brash Creek, road-edge, woodland § side of road, shade, alt. 1270 ft, 4 m tall
large bush in shade of balsam poplar, black fruit, 20 Aug 1995, J.B. Phipps 7172 (UWO), Northern
Okanagan, Spallumcheen Municipality, Marshall Road (dead end gravel drive off Mcleery Rd.), alt
1300 ft, bush, 2.5 m tall, fruit — slum-purple, ellipsoid, 20 Aug 1995, J.B. Phipps 7169 (BRIT, UBC,
UWO, V); Mabel L. Rd., N side, ca. 150 m F from Shuswap River bridge, 3-3.5 m tall, anthers exhibit
slight pink tinge, 15 May 1995, Ls 7 D. Donovan 2755 (UWO); Near Enderby, S end of 974A, E side of
Hwy 97A, ca. 150 m from Mabel L. Rd., forest margin, alt. ca. 1250 ft asl, ca. 2.5 m tall, 15- : pink
anthers, spindly, leaves larger, lighter green (shady location?), 15 May 1995, L.S.& D. Donovan 2754
(UWO),; Near Enderby, 5 end of 97A, E side of Hwy 97A, ca. 150 m from Mabel L. Rd., forest margin,
alt. ca. 1250 ft asl, ca. 3m tall, anthers pink to red, 15-18 stamens, 15 May 1995, L.S. & D. Donovan
2753 (UWO); S end of 97A, near Enderby, E side of Hwy 97A, ca. 150 m from Mabel L. Rd, forest
margin, ae ca. 1250 ft as... ca. 1.75 m tall, 15-18 stamens, deeply lobed leaves, anthers pink, 15 May
1995, L.S. & D. Donovan 2752 (UWO), Rd. to Mabel L. ca. 1.2 mi E of Enderby and between 25 m and
200 m W of “Autobody collision and repairs” sign, hedgerow, alt. 1275 ft, bush 3 m tall, fruit bur-
gundy-red, 20 Aug 1994, ].B. Phipps & RJ. Okennon 7015 (BRIT, UWO); 1.25 mi E of Enderby on Mabel
L. Rd., ca. 200 m E of “Autobody Collision, Repairs & Painting” sign, 1275 {t, hedgerow, bush, 5.5 m
tall, 20 Aug 1994, J.B. Phipps & RJ. O’Kennon 7018a (BRIT, UWO); Shuswap Valley, ca. 8 km E of cen-
ter of Enderby on road to Kingfisher, S side of road, 1270 ft, hedge backing small woodlot, bush, 4m
tall, stamens 10, anthers pink, (-JBP & RO’K 6815), 05 May 1994, J.B. Phipps 6922 (BRIT, CAN, UBC,
UWO, V); Shuswap Valley, road to Kingfisher, ca. 2 km E of Enderby, opposite sign on N, “Autobody
—_
Collision, Repairs & ee _alt. 1275 ft, hedgerow S side of road, bush 5 m, ca. 18 puce anthers,
a JBP & RO’K 6810), 05 May 1994, Phi PPs, J.B.6917 (BRIT, UBC, UWO); Shuswap valley, just E
of Enderby, across bridge, N side of road at edge of swamp forest, alt. ca. 1250 ft, bush, 3 m tall, thorns
medium, dark green foliage, stamens 15, anthers pale pink, 05 May 1994, |.B. Phipps 6910 (BRIT, UWO),
Northern Okanagan, Spalumcheen Municipality, Stepney Cross Rd., lowest point, large bush, 4 m
1, 26 Aug 2000, J.B. Phipps 8182 & S.R. Phipps (UWO).
co
et)
—
ser. Purpureofructi J.B. Phipps G O’Kennon.
Bushes or small trees, generally 3-6 m tall; thorns 2-5 cm long, slender to quite
stout, shiny dark brown to purple or black at 2 yrs, usually quite numerous;
extending twigs thinly pubescent or glabrous; at 1 year deep reddish-brown or
mahogany, older dark gray. Leaves deciduous; petioles 1/4-1/2 length of blade,
1-3 cm long, eglandular or with some small black glands, glabrous to thinly
pubescent while young; blades 3-8 cm long, + rhombic to elliptic-oblong, ovate-
rhombic or occasionally suborbiculate in general shape; acute at the apex, cu-
neate to somewhat rounded at the base; 1-3(-4)-lobed per side, the lobes ob-
scure and shallow to well-defined and acute and with max. L Il ca. 20% mate
]
serrate except at the base, the teeth with small gland
4-6(-7) per side, + impressed in the more coriaceous-leaved species; usually
scabrate-pubescent on the upper surfaces young, this variably glabrescent
(abrading with age), below glabrous on the surface, pubescent or not on the
veins; markedly coriaceous to thin; autumnal color generally brilliant. Inflo-
rescences 8-25 flowered; branches glabrous or thinly pubescent, bearing cadu-
cous, membranous, linear, gland-margined bracteoles. Flowers 12-18 mm diam.;
—
PHIPPS AND O’KENNON, NEW TAXA OF CRATAEGUS FROM CANADA 131
hypanthium externally glabrous to pubescent; calyx-lobes 2-4 mm long, nar-
row, abaxially glabrous, margins finely glandular-serrate; petals + circular,
white; stamens (5)10-12, anthers white or pink; styles 2-4(-5). Fruit 8-10 mm
diam., + orbicular to ellipsoidal or ampulliform, mostly red in late August, deep
purple or deep reddish-purple (often nearly black) in late September; calyx-
lobes narrow or broad, reflexed to erecto-patent, thinly hairy to glabrous; nut-
lets 2-4(-5), dorsally furrowed, sides plane to quite deeply pitted.
Six species are now recognized, found from the Okanagan Valley of British
Columbia and Washington east to northwest Montana. They are all locally com-
mon in suitable habitats. We add three new species and one variety to this se-
ries and emend the key.
These include some very handsome hawthorns, particularly in late sum-
mer and autumn when their usually abundant fruit turns from bright red to
deep purple and the leaves manifest brilliant coloring in bronze, burgundy, yel-
low etc. These hawthorns have very high ornamental potential but are not yet
cultivated outside botanic gardens and arboreta.
KEY TO SPECIES OF SERIES PURPUREOFRUCTI
. Leaf-blades rhomb-elliptic to elliptic in general shape, lobes + sharp.
2. Calyx-lobes in fruit us. reflexed, if erecto-patent, very narrow;anthers pink or white;
leaf- blades various but never suborbiculate, only rarely broad-elliptic.
3. Anthers pink; anthesis later than C. macracantha; leaves thin; thorns some-
what slender 1. C. williamsii
3. Anthers generally cream or white; anthesis with or somewhat before o
macracantha; leaves coriaceous; thorns usually + stout
4. Leaf-blades 3.5-6 cm long, coriaceous, without strongly impressed vena-
tion above, flattish to strongly concave, generally mid-green until fall; hy-
panthium glabrous or hairy 2.C. okanaganensis [incl. var. wellsii]
4. Leaf-blades 4-8 cm long, softly coriaceous, with distinctly impressed ve-
nation above, flattish to convex, generally deep green until ee
glabrous 3.C,. enderbyensis, sp. nov.
. Calyx-lobes in fruit sub-erect, broad, herbaceous; anthers pink; leaf blades ellip-
tic to broad-elliptic to suborbiculate, sometimes ovate
. Leaf-blades elliptic to broad-elliptic or reece aie green; fruit ca.8 mm
eee glabrous, oblong-suborbicular reddish in August, plum in September
inflorescences glabrous 4.C C arbiculatis sp.nov.
5. Leaf-blades elliptic to ovate in general shape, deep green; fruit 13-15 m
long, hairy,ampulliform to salsa deep iid in late August, nearly aN
in September; infloresences pubes .C. atrovirens, sp.nov.
1. Leaf-blades broad-ovate to ovate- ae ic in genta shape, lobes subacute to
obtuse 6. C. phippsii
NO
1. Crataegus williamsii F-ggl., Bull. Torrey Bot. Club 36:641. 1909. See also J.B.
Phipps, Canad. J. Bot. 76:1889. 1998
2. Crataegus okanaganensis J.B. Phipps @ O’Kennon, Sida 18:178. 1998.
132 BRIT.ORG/SIDA 20(1)
KEY TO VARIETIES
Leaf-blades 1.2-1.6 x as long as wide, sharply lobed; fruit flask-shaped 2a. var.okanaganensis
Leaf-blades1.6-2 x as long as wide, shallowly lobed in distal half only or occasionally
essentially unlobed; fruit subspherical 2b. var. wellsii, var. nov.
2a. Crataegus okanaganensis var. okanaganensis.
2b. serie ha okanaganensis var. wellsii ].B. Phipps @ O’Kennon, var. nov. (Fig.
6). TypE: UNITED STATES: WASHINGTON: Okanogan Co.: Palmer Lake, N side; large
bu . ; ; m tall, stamens 5, anthers ivory, (equals J.B.P. & RJ.O’K. 6875), + May 1994,
J.B. Phipps 6905 (HOLOTYPE: UWO; |sOTYPES: BRIT, CAN, DAO, UBC, US, WS).
Frutices grandes vel arbores parvae ad 6 m alti, saepe cum truncis
atrospadicei; spinae 2- “5 cm ie in juventute nitenter atro-brunneae, ovals recurvatae. Folia
decidua; petioli 1-15 cm longi, in juventute supra glandulosi saltem et pope laminae 3.5- i
cm longae, ellipticae vel ellipt oblongae vel ellipti ITICO rhombe ae vel elliptico
saepe leviter contractae in petiolum vel cum basi obtusa; apices acutae; saepe + cum bien as
>ramuli unius anni
parallelis in parte media; latera nonlobati vel lobis distinctis 1-3, latere distal angen) loborum
plerumque + erpendiealea ad venam li venis
secondariis principalibus +-5 per latus; facies aasials prev iter eas serrata in ei:
abaxialis glabra venis exceptis. Inflorescentiae 8-15 f1 pilosi, |
Aneatibus, membranaceis, glandulo-marginatis. Flores ca. 15mm diam, se aun sup glabrum
et subter vill . lobi calycis anguste-triangulares, tenuiter g
(5-)10-12, antheris plerumque eburneis; car ai et styli 2-3(-4). pacing ca. 10 mm diam., + orbicu-
lares, subter pilosi, sanguinei in Augusto exeunte, in maturitate atropurpurei; lobi calycis triangu-
]
lares, plerumque reflexi; pyrenae 2-3, dorsaliter sulcatae planis vel vadositer foveolatis.
Large shrubs to 6 m tall, often with multiple trunks; | year old twigs mahogany;
thorns 2-5 cm long, shiny dark brown when young, stout, recurved. Leaves de-
ciduous; petioles 1-1.5 cm long, glandular above at least when young, pubes-
cent when young; blades 3.5-5cm long, elliptic or ellipt-oblong to elliptic-rhombic
or elliptic-ovate in general shape; apex acute, base often slightly tapered into
petiole or obtuse; often + parallel-sided at broadest part, unlobed or with 1-3
distinct lobes, distal side of angle of lobes usually + perpendicular to mid vein;
margins serrate; venation craspedodromous with 4-5 main secondary veins per
side; short-appressed serrate when young above, glabrous below, except along
veins. Inflorescences 8-15 flowered; branches thinly hairy, with early caducous,
glandular bracteoles. Flowers 15 mm diam.; hypanthium externally glabrous
above and villous below; calyx-lobes narrow-triangular, finely glandular, pet-
als + circular; stamens (5-)10-12, anthers usually ivory; carpels and styles 2-
3(-4). Fruit ca. 10 mm diam., + orbicular, hairy below, dull deep red at late Au-
gust, deep purple at full ripeness; calyx-lobes triangular, usually reflexed; nutlets
2-3, dorsally sulcate, sides plane or shallowly pitted.
Crataegus okanaganensis var. wellsii is found in the Okanagan valley of
British Columbia and Washington and is common around Vernon (Fig. 7). Ab-
original people in Washington recognized this taxon as “stemoq’w” according
to R. May’s collection notes (cited specimen May 258RM from WA).
This variety was first collected by T.C. Wells at Kalamalka Provincial Park,
PHIPPS AND O'KENNON, NEW TAXA OF CRATAEGUS FROM CANADA
Ss
BON Q SP
zh : ce" 3)
% @ A
sal © a \W /gymag ’
cape a A
yA me AAR Ng i
\ Of
% yee
a eal .
“Sy Patent ff
NIG:
yb rs
Fic. 6. Line drawi fC f j IIsii. Scale bar = 1 cm.
134 BRIT.ORG/SIDA 20(1)
123° ™7120° Mtr ™114°
‘N
1
517
Fic. 7. Distributi f h is var. wellsii from collated records.
near Vernon. In its modal form it is very distinct with its unusual leaf-shape
(quite differently lobed from var. okanaganensis). It also differs from var.
okanaganensis in its orbicular fruit, fewer nutlets, fewer stamens (in the type),
and crimson (or pink-burgundy) fall color versus copper-bronze in var.
okanaganensis. Fruit may be a somewhat different color than nearby var.
okanaganensis at the same time in late August, although in the third week of
August they are brilliant scarlet. The most extreme form, virtually thornless, was
seen at Palmer Lake, northern Washington. In spite of the several differences
from var. okanaganensis the presence of intermediates prevents a higher rank.
Additional speci camined: CANADA. BRITISH COLUMBIA: Ca. 10 km N of Vernon center on
dirt road, N paand into O'Keele rr nn off 97A at LL A. Cross Rd., boundary hedge along ditch, com-
mon component, 3.5m bush, fruit burgundy, 25 Sep 1993, J.B. Phipps & RJ. O’Kennon 6804 (UWO).
Kalamalka Lake Prov. Park, 2 km SE of Vernon, at southern termi a Kidston Road, alt. 440 m, in
damp low-lying area, in open ae 5 m shrub, few thorns, 5 stamens, cream eee anthers), 16
May 1993, 1.C. Wells & MLE. Wells 1799 (CUWO). Kalamalka La
southern terminus of Kidston Road, alt. 440 m in damp low-lying area, in open ane 4m shrub, 5
stamens, cream colored anthers, common, 16 May 1993, T.-C. Wells & M.E. Wells i (UWO). Kalamalka
_ake Prov. Park, 2 km SE of Vernon, at southern terminus of Kidston Road, alt. 440 m in damp low-
me
ke Prov. Park, 2 km SE of Vernon, at
ying area, in open thicket, 4 m shrub, 5 stamens, essentially thornless, 16 May 1993, T.-C. Wells & MLE.
PHIPPS AND O’KENNON, NEW TAXA OF CRATAEGUS FROM CANADA 135
Wells 1796 (UWO). as Calder property, E of 97, ca.9 km NNE of Vernon, scrubby partly cleared
hillside, alt. ca. 474 m, young 3 m bush, purple fruit, 26 Sep 1993, J.B. Phipps & RJ. O° ae 68194
(UWO). Okz fo ae ca. 10 km N of Vernon off 97A NW bound along dirt road into O'Keefe
Ranch, at L. and A. Cross Road, alt. 1270 ft, boundary hedge along ditch, on SW side, bush, : m tall,
stamens 10, pre pale pink (=].B. Phipps & R.J.O° a 6804), 03 May 1994, J.B. Phipps 6893 (UWO).
Kalamalka Lake Provincial Park, ca. 10 km SSE of center of Vernon, near parking lot after yellow
gate, alt. ca. 1375 ft, large bush 6 m tall, 5 ivory stamens, (= J.B. Phipps & RJ. O’Kennon 6798), 03 May
1994, J.B. Phipps 6885 (TRT, UBC, UWO, V). Ca.10 km N of Vernon center on dirt road N into O'Keefe
Ranch off 97A at L. & A. Cross Road, alt. 1270 ft. boundary hedge along ditch, common component,
3.5m bush, fruit burgundy, 25 Sep 1993, ].B. PI hee RJ.O’Kennon 2050 WO). Shuswap Valley, just
E of Enderby, across bridge, S side of road, top of river bank, alt. ca. 1250 ft, bush 4 m tall, in bud,
stamens 5 or 10, anthers ivory, 05 May 1994, . picid UW0O). Kalamalka L. Prov. Park, 2 km SE
of Vernon, Kidston Rd. entrance at red gate, with Amelanchier alnifolia, Symphoricarpus albus, Pinus
ponderosa, open thickets, alt. 550 m, 4 m tree, shiny scarlet fruit, common, 18 Aug 1987, T.-C. Wells 689
(UBC # 19422, UWO). Kalamalka L. Prov. Park, 2 km SE of Vernon, Kidston Rd. entrance at red gate,
with Amelanchier alnifolia, Symp! albus, Pinus ponderosa, open iene alt. 550 m, 4m
tree, shiny scarlet fruit, common, 118 Mo 1987, IC wells 90(U BC # 194423, UWO). Shuswap Valle
just E of ween across bridge, S side of road, top of river bank, alt. ca. 1250 . bush 4 m tall in late
bud, stamens | anthers ior ie May 1994, ].B. Phippr6915(0W WO). Kalamalka L. Prov. Park, 2 km SE
of Vernon, at S terminus of Kidston Rd. red gate, with Amelanchier alnifolia us GLDUS
Pinus ponderosa, open thickets, 550 m, 3 m shrub, heavily fruited, shiny, Qaeos rt, common, TC
Wells 691 (UUWO, UBC 194424): Aug 18. 1087. Shuswap Valley, just E of Enderby, across bridge, S side
of road, top of river bank, alt. ca. 1250 ft, bush 6 m tall, in late bud, stamens 10, anthers ivory, 05 May
re PaaEes 6916 (UWO). Spallamcheen Municipality, E side 97A 1/2 mi N of intersection with
97, scrubby slopes rich in a bush 6m sey red fruit, most abundant Crataegus here, 19 Aug
1994, J.B. Phipps & RJ. O’Kenn 99 (BRIT, UWO, V). Ca. 1 mi S of Enderby, N-trending, downslope
gravel driveway bordered with er en i: 1300ft, bush, 4 m tall, fruit burgundy, 20 Aug 1994,
J.B. Phipps & RJ. O’Kennon 7006 (BRIT, UBC, UWO). Ca. 10 km N of Armstrong, W of Indian Reserva-
tion, alt. ca. 1550 ft, long farm drive with Crataegus hedgerow, bush 2 m tall, fruit in various stages of
red, 21 Aug 1994, |.B. Phipps & RJ. O’Kennon 7035 (UWO). Okanagan Valley, Spallumcheen Munici-
pality, L.& A. Cross Road, extreme E end, just W of Hwy 97A, alt. 1270 ft, along driveway N into farm,
this species abundant here, shrub 4 m, rather dense fruit, blood-red, + spherical, J.B. Phipps 7161(CAN,
ID, TRT, UBC, UWO, V); Aug 17, 1995. Northern Okanagan, Spallumcheen Municipality, Otter Lake
Cross Rd., just N of Hales Rd., 19 Aug 1995, J.B. Phipps 7162 (TRT, UBC, UWO, V). Northern Okanagan,
Spallumcheen Municipality, Marshall Rd., alt. 1300 ft, natural hedgerow, 20 Aug 1995, J.B. Phipps &
RJ.O’Kennon 7170(CAN, TRT, UWO). Between Vernon & Armstrong, intersection of Otter Lake Cross
Road & Hales Rd., alt. ca. 1300 [t, in partial shade of Ponderosa Pine and Douglas fir forest, 2 m tall,
fresh anthers pink, May 15, 1995, L.S. & D. Donovan 2749 (UWO). Shuswap Valley, just B of Enderby,
across bridge, W side of road, 250 m[ alt. ca. 1250 ft, level open fielc g roadside
across from cattle feeder, bush 3.5 m tall, ee ie cream, 15 May 1995, LS meg D. ‘Gone 2741
(UWO). Kalamalka L. Provincial Park, bushy slopes above red gate, in seepage zone, 1550 ft, same sp.
as Wells 689-691, bush 4 m tall, | shining, mid-green, ie bright red with long ca-
lyx-lobes, 18 Aug 1994 a B. Phipps & RJ.O’Kennon 6982 (CAN, UBC, UWO, V). Just E of Enderby a ter
bridge over Shuswap R., rough grass near water, S side of road, alt. 1250 : eice of water near oxbow
of Shuswap R., bush 5 m stall burgundy fruit, 25 Sep 1993, J.B. Phipps & RJ. O’Kennon 6809 (UBC,
UWO).E side of 97A, 1/2 mi N of intersection with 97, scrubby slopes with long cleared area, bush 4
m tall, fruit + spherical, plum-colored, common here, 19 Aug 1994, J.B. Phipps & RJ. O’Kennon 6997
(BRIT, TRT, UBC, UWO, V). Ca. 1 mi S of Enderby, gravel driveway to NE starting at Indian Cemetery,
alt. 1300 ft, bush, 3.5 m tall, fruit bright red, ae scarce here 20 Aug 1994, J.B. Phipps & RJ.
O’Kennon 7007 (UWO). Ca. | miS of Enderby, gat Indian Cemetery, alt.
136 BRIT.ORG/SIDA 20(1)
1300 ft, bush, 3.5 m tall, fruit + orbicular, dull red, a lew here, 20 Aug 1994, [.B. Phipps & RJ.O’Kennon
7005 (CAN, UBC, UWO, V). Mabel Lake Rd., 1.3 mi E of Enderby center, opposite house with solid
wooden fence, alt. 1275 ft, bush, 5 m tall, fruit e ca red, - Aug les Phipps & RJ. ene
7020(UWO). Kalamalka L. Provincial Park, bru | alt. 1600 [t,
bush, 4 m tall, leaves coriaceous, shining, ae green, ane bright a with lens elie lobes, : Aug
1994, J.B. Phipps & RJ. O’Kennon 6983 (BRIT, UBC, UWO). Northern Okanagan, ae Mu-
Marshall Rd. fruit scarlet, spherical, 20 Aug 1995, /.B. Phipps 7171 (UW
WASHINGTON: Okanogan Co.: US 97, 1.1 km S of Ellisforde,
ak in gr - field (mostly C. macracantha), alt. 275 m, bush, 4 m tall, dense, fruit deep red, +
round, 23 Aug 1995, J.B. Phipps 7176 ee WS, WTU): Westside Rd. ca. 4 ae N of Tonasket, huge
hawthorn stand along riverside, 975 t., bush, 7 m tall, fruit burgundy, spherical, 30 Sep 1993, |B.
Phipps & RJ. O’Kennon 6868 (BRIT, U fs WS, WTU): Palmer Lake, N side, alt. 1150 ft: thornless 7 m
bush, fruit burgundy, 1 Oct 1993, |.B. Phipps & RJ. O’Kennon 6873 (UWO, WS, WTU); Palmer Lake, N
end, open hawthorn thicket along gravel beach, just E of Chopata Lodge, alt. 1150 ft, bush 7 m tall,
thornless, burgundy fruit, | Oct 1993, J.B. Phipps & RJ. O’Kennon 6875 (BRIT, UWO, WS, WTU); Palmer
Lake, N side, anthers ivory, 4 May 1994, J.B. Phipps 6095 (UWO, WS, WTU). Spokane Co.: Spokane, 03
Sep 1896, C.V. Piper 2387 (WS). Stevens Co.: West End, Wellpinit, variety with maroon, barely hairy,
softer berry, 12 Sep 1976, Kk. May 258RM (V).
aes n hedge over RR
3. Crataegus enderbyensis J.B. Phipps & O’Kennon, sp. nov. (Fig. 8). Type: CANADA:
BRITISH COLUMBIA: just E of eer after bridge over Shuswap R., S side of road,
rough grass near water, alt. ca. 1250 ft, large bush 5m tall, burgundy-red fruit, 25
Sep sie Phipps & RJ. O” eee 6808 (HOL OTYPE: UWO,; ISOTYPES: BRIT, CAN,
DAO, L
br unnel, veteriores atro-
Frutices grandes vel arbores parvae, 5-6 m alti li unius anni purpureo
cinerei; spinae 2.5-3.5 cm longae, crassae, recurvatae, nitenter spadiceae in juventute. Folia decidua;
petioli 1-2 cm longi, + shi laminae 4-8 cm mee ovato-rhombeae vel late ellipt-rhombeae in
orma Salsity forma pl - basis late cuneata vel + rotudata; latera ca. 3-lobatae per
latus, [FI max. ca. 20%; venatio craspedodroma nec venis per latus distincte impressis; margines
ae ee textura coriacea, et color atrovirens in maturitate. facies adaxialis appresso-
eae aie abaxialis + gl: abra, coloribus in praeclaris. Inflorescentiae 15-25 florae; ramuli
villosi, teolis caducis, linearibus, membranaceis, glandulo-marginatis. Flores 15 mm diam.; hy-
panthium extrinsecum nee apr, lobi ca alyels anguste-triangulares, tenuiter glandulo-marginati,
abaxialiter glabri 10, antheris cremeis; carpelli et styli 3. Fructus 8-10 mm
diam., ellipsoideo- ere saepe in corymbo magno { .cremei in juventute, tum armeniaci
tum coccinei, in maturitate purpuret; ae calycis angustae, es pubescentes:; pyrenae 3,
dorsaliter sulcatae, lateribus planis vel vadositer foveatis.
Large bushes or small trees to 5 or 6 m tall; l-year old twigs purple-brown, older
dark gray; thorns 2.5-3.5cm long, stout, recurved, shiny mahogany-brown when
young. Leaves deciduous; petioles 1-2 cm long, + glabrous; blades 4-8 cm long,
ovate-rhombic to broad elliptic-rhombic in general shape; base broad-cuneate
to + rounded; with ca. 3 lobes per side, largest LII ca. 20%; margins finely toothed;
venation craspedodromous, distinctly impressed, with (3-)4-5 lateral veins per
side; leathery and dark green at maturity, also rather convex; appressed-pubes-
cent above, + glabrous below except on the + pubescent veins, brilliantly
colored in fall. Infloresences 12-25 flowered; branches villous, bearing cadu-
cous, linear, membranous, gland-margined, bracteoles. Flowers ca. 15 mm diam.;
137
PHIPPS AND O’KENNON, NEW TAXA OF CRATAEGUS FROM CANADA
is, Scale bars = 1 cm.
BRIT.ORG/SIDA 20(1)
hypanthium glabrous externally; calyx-lobes narrow triangular, finely gland-
margined, glabrous abaxially; petals + circular; stamens 10, anthers cream; car-
pels and styles 3. Fruit 8-10 mm diam,, ellipsoid-urceolate, often in a large, +
pendant corymb, cream when young, turning to apricot-colored then red, at
full maturity purple; calyx-lobe remnants narrow, reflexed, pubescent; nutlets
3, dorsally furrowed, their sides plane or shallowly pitted.
Crataegus enderbyensis is presently only recorded from the Armstrong-
Enderby region of the upper Okanagan of British Columbia where it is known
from a good number of specimens. Collection localities may be identified, at
least approximately, on figure 2.
This species is easily recognised in the field by its handsome, softly leath-
ery, somewhat drooping, convex, dark green foliage, brilliantly colouring in fall,
with clearly impressed venation, as well as by its usually large fruiting corymbs
with their at first very pale, pendant fruit which become Bae nt red by late
ae and ay turn purple-black. It is closely related to C. okanaganensis.
—
Add | ined: CANADA. BRITISH COLUMBIA: sale Okanagan, Spallumcheen
t, hedges, bush, 3.5 m tall, fruit
Muntetpalaiy. Powerhouse Rd., E side, near Stardel Drive, alt. ca. 135(
ellipsoid, orange-red, 19 Aug 1995, J.B. Phipps 7103 (OWO). Between ae & Armstrong, intersec-
tion of Otter Lake Cross Road and Hales Road, alt. ca. 1300 ft, in partial shade of Ponderosa Pine and
Douglas Fir forest, bush 2 m tall, anthers cream, {ls. not quite out, 15 May 1995, L.S.& D. Donovan 2750
(UWO). Between Vernon & Armstrong, intersection of Otter Lake Cross Road & Hales Road, alt. ca.
1300 ft, in so : ide of Ponderosa Pine ae cha ‘ir forest, 5m tall, fresh anthers cream col-
ored, 15 May 1 _S. & D. Donovan 2747 (UWO). Between Vernon & Armstrong, intersection of
Otter Lake Cross on & Hales Rd., ditch & road ne backing mesic forest, NW side of road near little
valley, alt. 1300 ft, forest margins, mesic, bush 3.5 m tall, fruit cream turning salmon, 21 Aug 1994, J.B.
Phipps, & RJ. O’Kennon 7032 (UWO). Ca. 200 m E of bridge over Shuswap Rd. on road Enderby to
Mabel Lake, S side of road, alt. 1250 ft, (equals J.B. Phipps & RJ. O’Kennon 6808), bush 5 m tall, nearest
road, {rt. elliptical, pendant, shades of red (pale to deep), 20 Aug 1994, J.B. Phipps & RJ. O’Kennon
7010 (BRIT, UBC, UWO). Back Enderby Rd., ca. 4 mi SE of Enderby, ca. 200 m N of creek crossing, alt.
ca. 1450 ft, dense hedges E side of road, bush, 5 m tall, fruit pale, 21 Aug 1994, J.B. Phipps& RJ.O’Kennon
7040 (UWO). Ca. 5 km ENE of Armstrong on E-W aligned rd. just ENE of Armstrong airport,
hedgerows, S side of road, alt. 1450 {t, bush 5 m tall, 21 Aug 1994, ].B. Phipps & RJ. O’Kennon 7042
(UWO). Mabel L. Rd, 1.25 mi FE of Enderby ca. 200 m E of “Autobody collision, repairs & painting
leaves with deeply impressed veins, very coriaceous,
sign,” alt. 1275 ft, hedgerow, bush, 5.5 m tall,
fruit unripe, cream-salmon, 20 Aug 1994, J.B. Phipps & RJ. O’Kennon 7018 (BRIT, UWO). Shuswap
Valley, just E of Enderby, across bridge, S side of rd. on mound by roadside depression, alt. ca. 1250 ft,
bush 5 m tall, 10 stamens, ivory anthers, in bud, 05 May 1994, J.B. Phipps 6911 (UBC.UWO)lalso J
6914,6915,6916 (UWO)|. Ca. L7 mi E of Enderby on Kingfisher Rd, on S side at beginning of S-bend,
of prominent hill, fence line of road backed by woodland, alt. 1280 ft, bush 3 m tall, foliage colors
autumnal, fruit purple, 25 Sep 1993, |.B. Phipps & RJ.O’Kennon 6813 (UWO). Ca. 17 mi E of Enderby
inglfisher Rd on S side at beginning of S-bend, $ of prominent hill, fence line of road backed by
woodland, alt. 1280 ft, bush 4 m tall, fruit purple, foliage deep green, 25 Sep 1993, J.B. Phipps & RJ.
VKennon 6812 (UBC, UWO).
—_
~
4. Crataegus orbicularis J.B. Phipps & O’Kennon, sp. nov. (Fig. 9). Type: CANADA:
BRITISH COLUMBIA, Northern Okanagan, Spallumcheen Municipality, Back Enderby
Rd., ca. 4 mi SE of Enderby, ca. 200 m N of creek crossing, alt. ca. 1450 ft, dense
PHIPPS AND O’KENNON, NEW TAXA OF CRATAEGUS FROM CANADA 139
hedges E side of road, bush 4 m tall, large, orbiculate leaves, fruit pale deep red, 21
Aug 1994, J.B. Phipps & RJ. O’Kennon 7039 (HOLOTYPE: UWO; ISOTYPES: BRIT, CAN,
MO, TRT, UBC)
er l; : es ] cat |
Frutices, late-patentes, 2-4
3-5 cm ones aes rectae vel puma kak recurvatae. ae decidua; panic ‘25- 20 cm
1
longi, adaxialiter breviter pilo if | g | mivoneee elliptic ae
vel fere circulares, | r basem i
venis secondatils 5- ae 7) per latus tobi 3- a lacus breves, triangulares IFI = 10%; ee cum
dentibus | pilosa, abaxialis sparse pilosa,
miagh pubescentibus in venis. Inflorescentia 10-18 florae, rammalis subglabris distaliter, sparsim
linearibus | i gl andulo marginatis. Floresca. 18mm
diam.; fgee extrinsecum oe glabrum; lobi calycis 2-3 mm longi, anguste-triangulares,
tenuiter glandulo-marginati; stamina 10, antheris roseis, car pen = ee 4 te Fructus 8 mm diam.,
oblongi, glabri, coccinei vel purpurei, in inf i, @ Erector
patentes; pyrenae 3-4, dorsaliter sulcatae, lateribus + pas vel vadositer ers
Wide-spreading bushes, 2-4 m tall; | year old twigs strong mid-brown, branches
dull gray; thorns 3-5 cm long, slimmish, straight or more usually slightly re-
curved; Leaves deciduous, petiolate; petioles 1.25-2.0 cm long, adaxially shortly
hairy, usually eglandular; blades mainly 5-8 cm long, broadly elliptic to nearly
orbicular, the broader ones curved across the base, the narrower somewhat at-
tenuated; venation craspedodromous, with 5-6(-7) secondary veins per side;
with 3-4 short triangular lobes, LI] < 10 %; margins with very small teeth ex-
cept near the base; shortly appressed hairy above, thin-hairy below, more pu-
bescent on veins. Inflorescences 10-18 flowered; branches sub-glabrous distally,
sparsely villous proximally, bearing caducous, linear, membranous, gland-mar-
gined bracteoles. Flowers ca. 18 mm diam.; hypanthium sub-glabrous externally;
calyx-lobes 2-3 mm long, narrow-triangular, finely gland-margined; stamens
10, anthers pink; carpels and styles 3-4. Fruit 8 mm diam., oblong, glabrous,
red, later changing to plum when fully ripe, in showy infructescences; calyx-
lobes very conspicuous, erecto-patent; nutlets 3-4, dorsally sulcate, their sides
+ plane or shallowly and smoothly pitted.
This is a very attractive species so far only known within a few miles of
Enderby, B.C. and easily recognized by the foliage shape, thorn-type and fruit
(color, conspicuous calyx-lobes). Crataegus or bicularis stands out in early spring
with its light green, tinted eau-de-nil, foliage and handsome clusters of large
flowers. Collection localities may be identified, at least approximately, on fig-
ure 2
Idi |: CANADA. BRITISH COLUMBIA: Nort! Okanagan, Spallumcheen
Municipality, Back Enos) Road, hedges, E side, alt. ca. 1450 ft, same as J.B. Phinpseonel O’Kennon
7039 : vear, bush 3.5 m tall, fruit lake, 19 Aug 1995, /.B. Phipps 7166 (BRIT, MONTU,
9 but
UBC, UWO). Northern Okanagan, Spallumcheen Municipality, Mountain View Road, N end, E side,
alt. ca. 1500 ft, 19 Aug 1995, J.B. Phipps 7165 (BRIT, UBC, UWO, V). Ca. 10 km N of Armstrong, W of
Indian Reservation, alt. ca. 1550 ft, long farm drive with Crataegus hedgerow, bush, 2 m tall, fruit in
various shades of red, 21 Aug 1994, J.B. Phipps & RJ. O’Kennon 7036 (BRIT, UBC, UWO, V). Mabel
Lake Rd, 1.3 mi E of Enderby center, opposite house with solid wooden fence, alt. 1275 ft, bush, 3 m
140 BRIT.ORG/SIDA 20(1)
Fic. 9. Line drawi f Crat bicularis. Scale bars = 1 cm.
PHIPPS AND O’KENNON, NEW TAXA OF CRATAEGUS FROM CANADA 141
tall, foliage large, fruit pale-medium red, 20 Aug 1994, J.B. Phipps & RJ.O’Kennon 7019(BRIT, MONTU,
UBC, UWO, V). Shuswap Valley, ca. 2.8 km E of Enderby on Kingfisher Rd., in hedgerow on S side
road opp. house with wooden fence on N side, alt. ca. 1280 ft, bush 3 m tall, large flowers, 10 stamens,
pink anthers, (possibly =].B. Phipps & R.J.O’Kennon 6813), 05 May 1994, J.B. Phipps 6920(UWO). North-
ern Okanagan, Spallumcheen Municipality, Back Enderby Rd., 21 Sep 1998, WJ.T. Phipps a & c(UWO).
5. at atrovirens J.B. Phipps & O’Kennon, sp. nov. (Fig. 10). Type: CANADA:
TISH COLUMBIA: Northern Okanagan, Spallumcheen Municipality, Otter Lake
ae Road, 22 Aug 2000, J.B. Phipps 8171& S.R. Phipps (HOLOTYPE: UWO; ISOTYPES:
RIT, CAN, MO, TRT, UBC, US).
| | is velerectis. R liel i pul illi unius
Frutices 2-5 malti
annus brunnei nitentes, veteriores atrocinerei, spinae 3-5 cm longae, numerosi, + rectae vel leviter
curvatae, nitentes atrobrunneae, ad apices nigrae, veteriores atrocinereae. Folia decidua; petioli
sores 15-20% illius laminarum, dense spUnesesutes in sulco adaxdal, in maturitate sparse
1_A(
pubescentes, eg glandulosi vel cum paucis, , par +-6(-7) cm
nee anguste-ovatae ad late elliptico- ionibess vel late e Tliptienss in forma generali; apex acutus
vel subacutus, basis cuneata; latera cum 3-4 lobis ea ube acutis vel eBELSIS: IFI sinus maximi ca.
10%; margines tenuiter serrati, dentibus acutis vel subacu g apices;
] J ~ 1s ] salir } ] = 7
venatio i ane oper
maturitate, abaxialiter turitat lor atrovirens et + coriacea, colore in autumno
Bieneulaces: Inflorescentiae 8- DoH reas pam E Geno Sam pubescentes. Flores 10-13 mm diam;
riangulares, marginibus glandulo-serratis
ad a laciniatis; petala alba; stamina 10, eae roseis; styli - 3 eae 13-15 mm alti, 10
mm lati, + ellipsoidei bro, + villosi;
caro dulcis, aurantiaca; lobi calycis va dep prominentes eho rectit pyrenae 2-3, dorsal iter sulcata, in
lateribus foveolatis.
Shrubs, 2-5 m tall, with multiple, mainly suberect to erect stems. Extending
branches of the season pubescent; at 1 yr glossy brown; older dark gray; thorns
3-5 cm long, numerous + straight to very slightly curved, at 2 yr. glossy dark
brown, black at the tip, older deep gray. Leaves deciduous; petioles 15-20% length
of blades, densely pubescent at maturity in the adaxial groove, sparsely pubes-
cent elsewhere, eglandular to with a few small glands mainly distally; blades
4-0(-7) cm long, narrow-ovate to broadly ellipt-rhombic or broad-elliptic in
general shape; tip acute to subacute, base cuneate; sides with 3-4 small sub-
acute to obtuse lobes, LIl of largest sinus ca. 10%; margins finely toothed, teeth
acute to subacute, bearing minute glands at their tips; venation
craspedodromous, 3-5 veins per side; adaxially with scabrous appressed pu-
bescence, abrading with age, abaxially nearly glabrous; dark green and some-
what coriaceous at maturity, in fall turning bronze-brown. Inflorescences 8-15
flowered; branches witha fairly dense spreading pubescence. Flowers 10-13 mm
diam.; hypanthium externally pubescent; calyx lobes narrow-triangular, mar-
gins glandular-serrate to glandular-laciniate; petals white; stamens 10, anthers
pink; styles 2-3. Fruit 13-15 mm high, 10 mm wide, + ellipsoid, burgundy in
late August, blackish-purple a month later; variably hairy; flesh soft orange
color; calyx-lobes very prominent, suberect; nutlets 2-3, dorsally grooved, pit-
ted on the sides.
BRIT.ORG/SIDA 20(1)
Fl ilable wt he drawi 1. Scale bars =
I
Crataegus atrovirens is now known from about 20 collections and many
more specimens on the ground from an area of about 10 miles north and south
of Enderby, British Columbia, where it is locally common to abundant in natu-
ral hedgerows and other brushy sites in the valley. Collection localities may be
identified, at least approximately, on figure 2. This taxon appears quite distinct
from other sympatric kinds and is often recognizable in habit alone.
Of species known to date from the northern Okanagan, C. shuswapensis,
also newly described in this paper, is the one most likely to be confused with C.
PHIPPS AND O’KENNON, NEW TAXA OF CRATAEGUS FROM CANADA 143
atrovirens. The coloring of C. shuswapensis is similar, but it has a finer, more
wide-branching growth habit, and a glabrous inflorescence, redder fruit in the
fourth week of August, much smaller and more recurved calyx-lobes in fruit,
whose margins rather than being serrato-laciniate, are merely minutely glan-
dular. In addition, the leaves of C. shuswapensis are more sharply lobed.
Crataegus atrovirens is probably more closely related to C. okanaganensis, but
is easily differentiated from that species in the field by the much darker color-
ing of foliage, fruit and bark, as well as by its pink anthers. Compared to C. or-
bicularis the fruit is conspicuously larger and more hairy in C. atrovirensand is
soft and ripe in the fourth week of August when it is hard and unripe in C.
orbicularis. It isa dull hawthorn, sombre in appearance, contrasting markedly
with more flashy congeners such as C. okanaganensis. It appears to be more
xeromorphic than other fairly large Okanagan hawthorns.
dditional specimens examined: CANADA. BRITISH COLUMBIA: Okanagan Valley, NNE of Vernon,
Pleasant Valley Rd., hedge behind Vowle’s residence, alt. 380 m, natural hedge, bush 3 m tall, 19 Aug
1994, J.B. Phipps 6996 & RJ. O’Kennon (UWO). Northern Okanagan, Spallumcheen Municipality,
Stepney Cross-Road, bush 5 m tall, 26 Aug 2000, J.B. Phipps 8184 & S.R. Phipps (UWO). Northern
Okanagan, Spallumcheen Municipality, Back Enderby Rd., ca. 1 mi N of Send, bush 5 m tall, 26 Aug
2000, J.B. Phipps 8181& S.R. Phipps (UWO, V). Northern Okanagan, Spallumcheen Municipality, Back
Enderby Rd., 1/4 mi from S end, bush, 2.5 m tall, 26 Aug 2000, J.B. Phipps 8179 & S.R. Phipps (UBC,
UWO). Northern Okanagan, Spallumcheen Municipality, Back Enderby Rd., near S end, bush, 3 m
tall, J.B. Phipps 8180 & S.R. Phipps (CAN, UWO). Northern Okanagan, Spallumcheen Municipality,
Marshall Rd., bush, 4 m tall, J.B. Phipps 8186 & S.R. Phipps (UWO). Northern Okanagan, Hwy.
Vernon, ca. 2 mi W of Lumby, bush, 12 [t tall, 26 Aug 2000, J.B. Phipps 8188 & S.R. Phipps (UWO).
Shuswap Valley, ca. 3miNNE of Enderby, Anderson Rd., bush, 3 m tall, 26 Aug 2000, J.B. Phipps 8185
& S.R. Phipps (BRIT, UWO). Enderby, 6 Aug 1898, J.R. Anderson s.n.(UWO, V). Hwy 97, ca. 2.5 km S of
Enderby (S of Indian Cemetery), roadside, E side, backing alfalfa field, alt. 1320 ft, dense shrub, 3 m
tall, coriaceous leaves, deep green, fruit purple, abundant here, 20 Aug 1994, J.B. Phipps & RJ.O’Kennon
7008 (UWOQ).
6. Crataegus phippsii O'Kennon, Sida 18:185, 1998.
ACKNOWLEDGMENTS
The line drawings were prepared by Susan Laurie-Bourque of Hull, Quebec who
is thanked by the authors for the beauty of her realisations and her meticulous
attention to detail. Susan Laurie-Bourque also artistically improved the rather
crude output of Range-Mapper by using Corel Draw. The work was largely
funded by grants to the first author by the National Sciences and Engineering
Research Council of Canada. We also wish to thank Laurie Donovan of
Okanagan University College, Vernon, British Columbia, for collection of criti-
cal flowering material at times when we were unable to be present and Antony
Littlewood, Dept. of Classical Studies, The University of Western Ontario, for
critically checking the Latin diagnoses.
144 BRIT.ORG/SIDA 20(1)
REFERENCES
Brunsretp, $.J.and FD. Jonson. 1990. Cytological, morphological, ecological and pheno-
logical support for specifc status of Crataegus suksdorfii (Rosaceae). Madronho 37:
274-282.
Puipes, J.B. 1998. Introduction to the red-fruited hawthorns (Crataegus, Rosaceae) of west-
ern North America. Canad. J. Bot. 76:1863-1899.
Pripes, J.B.and R.J. O’KENNoN. 1998. Three new species of Crataegus (Rosaceae) from west-
ern North America: C. okennonii, C. okanaganensis and C. phippsii. Sida 18:169-191.
Taytor, T.M.C. 1973. Crataegus. In:The rose family of British Columbia. British Columbia Pro-
vincial Museum, Dept. of Recreation and Conservation. Handbook no, 30:26-32.
NEW COMBINATION IN XYLORHIZA
(ASTERACEAE: ASTEREAE)
Guy L.Nesom
Botanical Research Institute of Texas
09 Pecan Stree
Fort Worth, TX 76102-4060, U.S.A.
ABSTRACT
Xylorhiza glabriuscula var. linearifolia is treated within Xylorhiza at specific rank as Xylorhiza
linearifolia (TJ. Wats.) Nesom, comb. nov, a ROT macaneal id is sBeOnTP a isolated from X.
Lal ; ] =
1
buti | PI hy bridize with lia var. imberbis.
ABSTRACT
Xylorhiza glabriuscula var. linearifolia se trata dentro Xylorhiza a nivel especifico como Xylorhiza
linearifolia (T.]. Wats.) Nesom, comb. nov. Xylorhiza linearifolia esta aislada geograficamente de X.
glabriuscula sensu stricto, pero se solapa y aparentemente se hibrida con X. tortifolia var. imberbis.
Watson (1977) described Xylorhiza glabriuscula Nutt. var. linearifolia TJ. Wats.
from the region of Moab in east-central Utah. Cronquist (1994) observed that it
“appears to form a distinctive local taxon with a range more than 60 km long” and
treated it at specific rank, but he placed it within a broadly defined Machaeranthera.
Welsh et al. 1993) followed Watson’s treatment. For the treatment of Asteraceae in
the developing Flora of North America, this taxon is recognized at specific rank
and the following combination is required to place it within Xylorhiza.
Xylorhiza linearifolia (TJ. Wats.) Nesom, comb. nov. BASIONYM: Xylorhiza
glabriuscula Nutt. var. linearifolia TJ. Wats., Brittonia 29:215. 1977. Machaeranthera
linearifolia (TJ. Wats.) Cronq., Intermountain Fl. 5:274. 1994. Type: UNITED
STATES. UTAH. Grand Co: 6 mi NW of Moab on US Hwy 160, 21 May 1971, TJ.
Watson 679 (HOLOTYPE: TEX!: ISOTYPES: mene GH, MO! MONTU, NY, UC).
Xylorhiza glabriuscula, X. linearifolia, and X. venusta (M_E. Jones) Heller are
closely morphologically similar among themselves and presumably are closely
interrelated. All three have diploid populations; X. venusta and X. linearifolia
also have tetraploid populations (Watson 1978). Within X. linearifolia, the two
cytotypes “grow intermixed over the small range of the taxon” (Watson 1978):
four populations of diploids and four of tetraploids were found, all within Grand
County, Utah. According to Stockton (1983), the diploids and tetraploids of X.
linearifolia differ in minor ways, but the present study (following Watson) finds
no variation that would require further nomenclatural differentiation
Watson (1977, p. 215) observed that Xylorhiza linearifolia (as var. linearifolia)
“overlaps morphologically” with X. glabriuscula (var. glabriuscula) but the
SIDA 20(1): 145 - 147. 2002
BRIT.ORG/SIDA 20(1)
146
tifolia var. imberbis. Outlines
TEX-LL, MO, and published sources, prima-
if,
lia, X. glabriuscula, X. venusta, and X. t
Fic. 1.G
ALL
JMU,
PDIT_CA
1a77\.6
\ }
+4] (1988) Th A
7
5 Ferry, 6 Jun 1927, Cottam (2671 (LL!).
1
Co., Lee
NESOM, NEW COMBINATION IN XYLORHIZA 147
present study notes that X. linearifolia similarly overlaps with X. venusta.
Xylorhiza linearifolia differs from both taxa in its leaf morphology, as outlined
by the following key.
1. Leaves linear-oblong, truncate or rounded-auriculate at the base Xylorhiza linearifolia
1. Leaves oblanceolate to oblong-spatulate, tapering at the bas
2. Stems a) for more than 2/3 of their length, peduncles oe 6 cm long; leaves
wide; involucres 7-14 mm long, 12-25 mm wide le ioe
2. Stems lea in the lower 1/2-2/3 of their length, peduncles 6-20 cm; leaves
15 mm wide; involucres 12-20 mm long, 20-35 mm wide chi venusta
Xylorhiza glabriuscula and X. venusta have slightly overlapping ranges in
northeastern Utah and northwestern Colorado (Fig. 1) and intermediates, pre-
sumably of hybrid origin, occur in that region (Watson 1977). The range of X.
linearifolia is essentially imbedded within that of X. venusta, but no evidence
of hy bridization between these two taxa has been reported. Xylorhiza linearifolia
apparently is reproductively isolated from both of its close relatives.
In contrast to the isolation of Xylorhiza linearifolia from X. glabriuscula
and X. venusta, the range of X. tortifolia var. imberbis (Crong.) TJ. Wats. con-
tacts that of X. linearifolia from the south and these two taxa grow intermixed
within a few localities. “In these zones of contact, intermediates (suggesting
hybridization) are found” (Watson 1977, p. 215)
Stockton (1983) noted that Xylorhiza linearifolia is intermediate in mor-
phology and habitat between X. glabriuscula sensu stricto and X. tortifolia var.
imberbis and suggested that X. linearifolia may be “a stabilized hybrid deriva-
tive” between them, presumably from an earlier time when the putatively pa-
rental species were sympatric. Further evidence that would test this hypoth-
esis is lacking.
ACKNOWLEDGMENTS
lam grateful to the staffs at MO and TEX-LL for help during recent visits there
and to John Semple and Eric Lamont for review comments.
REFERENCES
Avesee, B.J.,L.M. SHuttz, and S. GoopricH. 1988. Atlas of the vascular plants of Utah. Utah Mu-
seum of Natural History, Univ. of Utah, Salt Lake City.
Cronauist, A. 1994. Intermountain flora. Vol.5, Asterales. New York Botanical Garden, Bronx.
Stockton, T. 1983. The origin of Xylorhiza glabriuscula var. linearifolia (Asteraceae). Amer. J.
Bot. 70(5), pt. 2[abstracts]:103.
WATSON, TJ. Jk. 1977. The taxonomy of Xylorhiza (Asteraceae—Astereae). Brittonia 29:199-2 16.
Watson, T.J., Jr. 1978. Chromosome numbers in Xylorhiza Nuttall (Asteraceae — Astereae).
Madrono 25:205-210.
Wetsh, S.L.,N.D. ATwoon, S. Gooprich, and L.C. Hicains (eds.). 1993. A Utah flora (ed. 2, revised).
Print Services, Brigham Young University, Provo, Utah.
148 BRIT.ORG/SIDA 20(1)
BOOK REVIEW
B.G. Bowes. 2000. A Color Atlas of Plant Structure. (ISBN 0-8138-2093-4, pbk.).
lowa State Press, 2121 State Avenue, Ames, IA 50014, U.S.A. (Orders: 1-800-
862-6057, 1-515-292-3348 fax; ea A eR AEE $49.95, 192 pp,
numerous color plates, 7 3/4" x 101/4".
The amount of information generated in the many fields of plant science is growing at an exponen-
tial rate. The detailed study of plant anatomy should be a vital component in the training for any
student of plant biology. However, with the advent of new analytical techniques in molecular biol-
ogy, biochemistry, and physiology, many students cannot find the time in their program of study to
include a course in basic plant anatomy. In fact, in some pperany preer ams it is even difficult to finda
students with a solid background.
faculty member who has sufficient tra
This book provides a long- ee solution to both of idles: problems. One of the most difficult
tasks for instructors teaching beginning ae students is to teach them how to understand plant
anatomy using microscopic cross-sections of the various organs. Part of that difficulty lies either
with the student's inexperience in sng microscopes, or in the poor quality of microscopes used in
teaching many undergraduate laborato
van Bowes’ Color Atlas of Plant Structure is a eemean illustrated and concise introduc-
ive)
A
ree
he fascinating world of plant anatomy. Each chapter consists of several pages of text, with
eav-
tion tot
abundant references to illustrations on the following pages. The text is concise and to the point,
ing much of the detailed description to be presented with the illustrations Chemises: Those illus-
—
QP
with a wealth of information. $ ctures referred toi I num-
bered, not ea or labeled, which makes each photograph easier to pert
he organization of the book follows that ae in many basic botany courses. The first
chapter provides an overview of the major Divisions of vascular and non-vascular plants, and con-
cludes with a brief introduction to the gross morphology of flowering plants. Chapters 2-4 provide
a discussion of plant cells, tissues and embryology. Chapters 5-8 deal with leaf, stem, root, and floral
anatomy.
The illustrations themselves are beautifully done, and each is labeled to indicate the type of
microscopy used to create the photograph. There are two or three photographs on each page, and the
captionsa are at ranged insucha way that itise easy to determine maiclicap ue n refers to which photo.
1]
The only difficulty this reviewer had with the illustrations was that occasionally the numbers refer-
ring to ty points, were difficult to see on the photographs.
is book is a wonderful reference for anyone iyeies in a plant sciences. It should be a
ae text or at least an available reference in all basic botany a nomy classes, and deserves
a place in the personal DPEADES of EDOtaN StS horticulturaiss ni alt and natural history artists
pn Trock, Collecti ‘al Research Institute of Texas, Fort Worth, TX 76102-
4060, U.S
SIDA 20(1): 148. 2002
LECTOTYPIFICATION OF PSEUDOGNAPHALIUM BIOLETTII
(GNAPHALIEAE: ASTERACEAE)
Guy L.Nesom
Botanical Research Institute of Texas
9 Pecan Street
Fort Worth, TX 76102-4060, U.S.A.
Pseudognaphalium biolettii A. Anderb. [nom. nov], Opera Bot. 104:147. 1991.
naphalium bicolor Bioletti, Erythea 1:16. 1893 [non Gnaphalium (Helichrysum)
bicolor (Lindl.) Schultz-Bip., Bot. Zeit. 3:171. 1845; nec Gnaphalium bicolor Franch.,
J. Bot. (Morot) 10:411. 1896]. Type: U.S.A. California. Without designating a type,
Bioletti 1893, p. 17) noted that “The description is drawn up from dried specimens
from San Diego and from a plant growing in the Garden of Native Plants at the
University of California. .. Common in low thickets among the coast hills about
San Diego, according to Prof. Greene. ...” At UC, there are at least two specimens of
Gnaphalium bicolor that might represent the material referred to by Bioletti: San
Diego, 25 Mar 1891, G.W. Dunn s.n. (UC 31964); San Diego Co, Witch Creek, 1893,
R.D. Alderson s.n. (UC 31965). Two ND-G specimens, however, apparently more
clearly represent authentic type material—they are labeled “Gnaphalium bicolor,
Bioletti, Erythea 1.16. Berkeley, Bot. Garden, 1892,” without other information (lec-
totype, designated here: ND-G 59203, photocopy’; isolectotype: ND-G 59205, pho-
tocopy!). These two large branches were broken above the base, without roots, and
their similarity suggests they may have been collected from a single plant. The
morphology of the ND-G types indicates that this species consistently has been
identified correctly.
Perennial herbs, taprooted, fragrant, the stems 2-7(-12) dm tall, much-branched
at the base and commonly lignescent, eglandular, persistently above,
glabrescent or not below. Leaves oblanceolate to oblong-oblanceolate, mostly
1.5-5(-8) cm long, 4-10(-15) mm wide, the upper becoming lanceolate, widely
to narrowly auriculate-clasping, not decurrent, strongly bicolored, bright green
and usually densely glandular above, persistently white-tomentose beneath,
sometimes the upper leaves glandular but those toward the base eglandular.
Capitula 5-5.5(-6) mm high; phyl laries sl ny opaque-white sometimes slightly
pinkish, strongly graduated, apex rounded to nearly truncate, apiculate. Pistil-
late flowers 41-73. Bisexual flowers 5-13. Cypselae shallowly ridged, not papillate.
Pseudognaphalium biolettii occurs in the southern/southwestern half of
California and in adjacent Mexico (Baja California and Baja California Sur), at
elevations of 5-1200 meters (under 600 meters in California) on rocky slopes,
sandy plains with Larrea, coastal strand, matorral, and chaparral.
ACKNOWLEDGMENTS
Barbara Hellenthal provided information and photocopies of the relevant ND-G
SIDA 20(1): 149 — 150. 2002
150 BRIT.ORG/SIDA 20(1)
specimens of Gnaphalium bicolor and John Strother provided information re-
garding material of G. bicolor at UC and JEPS. Review comments of K. Gandhi
and J. McNeill also are much appreciated.
REFERENCE
Biotetti, F.T. 1893. Two new Californian plants. Erythea 1:16-17.
BOERHAVIA COULTERI VAR. PALMERI, A NEW VARIETAL
COMBINATION FOR BOERHAVIA (NYCTAGINACEAE) OF
SOUTHWESTERN NORTH AMERICA
Richard Spellenberg
Bic log yy L CPU tment
w Mexico State University
i Higa ne 88003-8001, U.S.A.
ABSTRACT
Boerhavia coulteri (Hook. L)S. Wats. var. palmer (S. Wats.) Spellenb., comb. nov, based on B. spicata
meri S. Wats., f from the v icinity of Guaymas, Sonora, Mexico, is propos ed fora Boerhavia
Choisy var. pa
that differs from the var. coulteri in its small fruits and flowers, and its with sparsely-fruited spicate
orescences. The variety mh from the Channel Islands of California to the Big Bend kaegon of
inf
Texas near the Rio Grande, and is very common on the Sonoran Desert. The two varieties are often
sympatric, and they eee to a slight extent. A key is presented to distinguish the species of
Boerhavia with spicate inflorescences in the United States.
RESUMEN
Boerhavia coulteri (Hook. f.) S. Wats. var. le ri(S. te aint comb. nov., basada en B. spicata
Choisy var. palmeri S. Wats., de la vec G nora, México, se propuso para una forma
de Boerhavia que difiere de la var oe cou ite ri por: su foe y ries pequenos, y su frutos esparcidos
en inflorescencias espiciformes. La anales de California hasta la
region del “Big Bend” del Rio Bravo del Norte de Texas, y es muy comun en el Desierto Sonorense. Las
variedades a menudo son simpatricas, y se intergradan un poco. $ | dara distinguit
£
t
las especies con inflorescencias espigadas de Boerhavia de los Estados Unidos de América
The species of Boerhavia with spicate inflorescences have proven taxonomi-
cally difficult for the past century, the variation presented by the complex re-
sulting in very different taxonomic treatments. At one extreme is that of
Standley (1911) where nine species were recognized. In 1918 Standley submerged
B. watsoni as a synonym of B. coulteri, but maintained the other taxa he had
recognized. At the other extreme is Reed’ s (1969) treatment, in which five of the
species Standley recognized [B. coulteri (Hook. f.) S. Wats., B. rosei StandL, B.
spicata Choisy in D.C, B. torreyana (S. Wats.) Standl., B. watsoni Standl.] were
placed as synonyms in one variable species, B. spicata. Most authors of floras
that include all or part of the Sonoran Desert region have followed Standley
(1918) in recognizing B. coulteri, and also B. spicata, and B. torreyana if they were
present in the area of concern (Spellenberg 1993; Martin @ Hutchins 1980;
Kearney & Peebles 1964; Munz & Keck 1968; Wiggins 1964). In contrast, Felger
(2000) recognized some plants included in B. coulteri, as broadly constructed
by those authors, as B. spicata var. palmeri S. Wats. It is this taxon that is recog-
nized as a variety of B. coulteri in this paper.
SIDA 20(1): 151 -— 155. 2002
152 BRIT.ORG/SIDA 20(1)
While studying the complex for an upcoming treatment for the Flora of
North America North of Mexico, I noted that B. torreyana was conspicuously
papillate in the sulci between the ribs of the fruit, a character previously un-
published. None of the other Boerhavia taxa in the complex are at all papillate.
Watson (1889) cited no specimens when he coined the name Boerhavia spicata
var. torreyana S. Wats., but specimens from western Texas collected during the
mid-19th century Boundary Survey, very likely seen by him, have papillate fruits
(specimens from Torrey Herbarium, NY, have been seen; relevant specimens
from GH on loan to ENCB have not been seen). The character of papillae on the
fruit is consistent in populations from western Texas, New Mexico, southeastern
Arizonaat higher elevations, throughout much of Arizona north of the Mogollon
Rim, and extreme south-central Utah (= B. spicata by Welsh et al. 1987). On very
similar appearing plants from the Sonoran Desert region, some of which are
identified as B. torreyana in collections, but which actually represent B. coulteri
var. palmeri (as proposed below), papillae are absent and sulci are much narrower
and less rugose. That B. torreyana is clearly distinguishable from other spicate
Boerhavia has helped to clarify the complex. This is particularly so in Arizona
and Sonora where three or four very similar taxa occur within the region.
In 1889 Watson transferred Senkenbergia coulteri Hook. f.to Boerhavia, and
he stated that the collection upon which the name was based was Coulter 1425.
He noted that the perianth was one line in length (= 1/12", ca. 2 mm) and that
fruits were clavate, 1-15 lines long, truncate, and acutely angled. Only one such
specimen exists in the Hooker Herbarium at Kew. It is labeled as having come
from “Alta Sonora, Mexico,” and is here considered the holotype (photo and
photocopies at NMC). Its fruits are clearly in irregularly fasciculate clusters in
the spicate inflorescence, and they are truncate at the apex. On the same page
of his 1889 publication, Watson stated that his new variety, Boerhavia spicata
var. palmeri, based on the collection Palmer [41 from Guaymas, Sonora, in 1887,
had perianths 0.5 line long, and fruits clavate-oblong, a line long, obtuse, ob-
tusely angled. He also noted the inflorescence to be “open and very slender.”
Palmer I4lis represented by two specimens at GH! (on loan to ENCB, lectotype
not selected; fragment at DS!). In raising this taxon to specific status, Standley
(1909) had to adopt a new name (B. watsoni) because the binomial B. palmeri S.
Wats. was occupied. He noted that Arizona plants were placed there somewhat
doubtfully, suggesting that geography would place them with B. coulteri. Rela-
tive to other plants in the complex, the plants he identified as B. watsoni have
smaller, more widely spaced, less clustered fruits (if at all clustered), and seem
to have had stems that were decumbent-spreading.
It is proposed here that the populations Standley recognized as B. watsoni,
and that Watson described as B. spicata var. palmeri, are part of the B. coulteri
complex and can be recognized at the varietal level. The following combina-
tion is proposed:
SPELLENBERG, A NEW VARIETAL COMBINATION FOR BOERHAVIA 153
Boerhavia coulteri (Hook. f.) S. Wats. var. palmeri (S. Wats.) Spellenb., comb. nov.
Boerhavia spicata Choisy var. palmeri S. Wats., Proc. Amer. Acad. 24:70. 1889.
The differences the two varieties of B. coulteri that Watson, and later Standley,
noted by their classifications are very apparent in the field in their extreme
states, but intergradient plants occur. Occasionally B. coulteri var. coulteri and
var. palmeri are intermixed in populations (e.g., Spellenberg & Zucker 12954 =
var. palmeri, 12955 = var. coulteri, AZ, Pima Co., ca. 10 air km SW of Marana,
Anway Rd. 1 km S of Avra Valley Rd., 16 Aug. 1999; NMC). In the two mixed
populations | have seen, the taxa differed in habit, the stems of var. coulteri de-
cumbent at the immediate base, then strongly ascending or erect, contrasting
to the more decumbent-spreading habit of var. palmeri, but habit is very likely
to be influenced by ecological conditions and density of populations. Some
mono-varietal populations of the var. coulteri have decumbent-spreading stems.
The var. coulteri also has larger flowers, a denser inflorescence, and fruits in
irregular fasciculate clusters, whereas the var. palmeri usually has minute flow-
ers, and well-spaced, smaller fruits, the clusters much less pronounced. Plants
are most often present in monomorphic populations representing only one
variety. The flower of B. coulteri var. palmeri is illustrated in Spellenberg (2000,
fig. 1), but is labelled as B. torreyana; the complex was not understood at the
writing of that article.
Infl of all the spicate species (B. coulteri included) I have observed
are visited by various insects in the morning when flowers are open and pro-
ducing nectar, the insects moving from inflorescence to inflorescence, and from
plant to plant. Larger-flowered species are visited more frequently (Spellenberg
2000). In mixed populations, particularly, inter-taxon pollination is very likely,
and hybridization is presumed possible. Both varieties of B. coulteri are weedy,
responding positively to disturbance, and with the advent of increased human
impact in arid areas of the Southwest, may have come into increased contact.
Boerhavia coulteri var. coulteri is geographically the more restricted of the
two varieties in the southwestern United States (ranges in Mexico have not been
determined), occurring from the eastern to the western borders of Arizona,
south of the Mogollon Rim. A population discovered in south-central New
Mexico in a well-collected area and may be the result of a recent introduction
(N.M., Dona Ana Co.,e Las Cruces along University Ave.e of Telshor Drive, 1 km
e of IH-25, in gravel planting strip between sidewalk and road, 24 Aug 2001,
Spellenberg and Brouillet 13267, DUKE, MT, NMC). The var. palmeri occurs
throughout the Sonoran region in the United States, from southeastern Califor-
nia to southeastern Arizona. It occurs on Santa Catalina Island in the Channel
Islands of California (Standley, 1909, cited as B. watsoni, Orcutt s.n., 12 Sep. 1889,
US)., where it may have been introduced. In Texas the var. palmeri is known
from a few collections on or near the Rio Grande, the eastern most being from
the eastern side of present day Big Bend National Park (M.S. Young, s.n., muddy
BRIT.ORG/SIDA 20(1)
bank of Rio Grande, near San Vicente [5 mi SW of Boquillas], 20 Aug 1915, TEX
[2], LL [1]. On one of these Texas collections, all identified originally as B.
torreyana, is pencilled, “but looks just like B. spicata Choisy Arizona,” the writer
probably alluding to its similarity to B. coulteri var. palmeri of Arizona, as here
recognized. Because this is the first report of this taxon for Texas, the two other
collections seen are cited here: L.C. Hinckley 1505, Presidio Co., Capote Canyon
1 mi. below falls, 28 May 1941, US; C-H. Muller 8438, Presidio Co., along washes
of rolling Rio Grande Plain, 4 mi SE of Ruidosa, 28 Jul 1945, LL. In southwest-
ern New Mexico the var. palmeri is occasional and weedy; in the vicinity of Las
Cruces it is always in disturbed areas. It may be introduced there, but if so, it is
long-established in the region (e.g., Wooton & Standley 3905, NM, Dona Ana Co
mesa W of Organ Mts., 4000 ft, 5 Oct 1907, LL, TEX, US).
The spicate taxa of Boerhavia in the United States may be distinguished by
the following key. In this key the range of fruit length-to-width ratios is given
in the format Cength/diam. = 1.3-2.D.
1. Fruits usually 4-angled
—
rarely 5-angled), broadly obovoid (length/diam. = 1.3-2.1
inflorescence usually densely glandular villous, without sticky intern
—_
odal bands
B. wrightii A. Gray
Fruits 5-angled, broadly to narrowly obovoid (length/diam. = 1.7—3.1); branches of
inflorescence glabrate to puberulent but not glandular, usually with sticky intern-
odal bands.
2. Surface of sulci of mature fruit papillose, otherwise smooth or somewhat rug-
ose, the edges of the ridges along sulci strongly ru
B. torreyana
2. Surface of sulci not papillose, instead
ooth or sometimes rugose, the edges
of the ridges along sulci not rugose to aout rugo
3. Fruits broadly obovoid (length/diam. = 1.7-2.1), sually overlapping in inflo-
rescence; sulci and ribs slightly rugose; sulci usually about 0.5 times as wide
f ribs B. spicata
. Fruits narrowly obovoid or obpyramidal (length/diam.= 2.13.1), eveiiapping
in inflorescence or remote; sulci and ribs slightly rugose t oth; sulci 0.1-
mes as wide as base of ribs.
ioe)
4 Paneer with fruits often a 50-100% of their length, com-
only + asciculate in group
s of 2-4, each fruit 2
oat trunc ate, many conic, or ne oe B. coulteri var. coulteri
4. Inflore tl ome lapping 1-50% of their an
aiaale 2-3 ina cluster,each nuit 2-2. 4 mm long, occasionally longer
usually rounded apically
2.5-3,.6 mm long, truncate,
B. coulteri var. palmeri
ACKNOWLEDGMENTS
| acknowledge with gratitude the help of Brian Stannard (kK), who sent photo-
copies anda photographic print of the holotype of Senkenbergia coulteri Hool
f. lalso thank curators of herbaria who loaned specimens for this study, and to
Richard Felger for various e-mail “chats” regarding this taxon
SPELLENBERG, A NEW VARIETAL COMBINATION FOR BOERHAVIA 155
REFERENCES
Feicer, R.S. 2000. Flora of the Gran Desierto and Rio Colorado of northwestern Mexico.
Univ. Arizona Press, Tucson.
KEARNEY, T.H.and R.H. Peestes. 1964. Arizona flora, 2nd.ed., with supplement. Univ. Calif. Press,
Berkeley.
Martin, WLC. and C.R. Hutcuins. 1980. A flora of New Mexico, vol. 1.J. Cramer, Vaduz.
Munz, PA. and D.D. Keck. 1968. A California flora, with supplement. Univ. Calif. Press, Berke-
le
Reep, C.F. 1969. Nyctaginaceae. In: C.L. Lundell, ed. Flora of Texas, vol. 2(1). Texas Research
Foundation, Renner. Vol. 2(1), pp. 151-220.
SPELLENBERG, R. 1993. Nyctaginaceae.In:J. Hickman, ed. The Jepson manual: higher plants of
California. Univ. Calif. Press, Berkeley. Pp. 768-774.
SPELLENBERG, R. 2000. Blooming “behavior"in five species of Boerhavia (Nyctaginaceae). Sida
19:311-323.
StanbLey, PC. 1909. The Allioniaceae of the United States with notes on Mexican species.
Contr. U.S. Natl. Herb. 12:303-389.
STANDLEY, PC. 1911. The Allioniaceae of Mexico and Central America. Contr. U.S. Natl. Herb.
13:377-430.
STANDLEY, PC. 1918. Allioniaceae. North America flora 21:171-254. New York Botanical Gar-
den, Bronx.
Watson, 5. 1889.VI.Contributions to American botany. 1.Upon a collection of plants made
by Dr. E. Palmer, in 1887, about Guaymas, Mexico, at Muleje and Los Angeles Bay in
Lower California, and on the island of San Petro Martin in the Gulf of California. Proc.
Amer. Acad. Arts 24:36-87.
Wetsh, S.L., N.D. ATwooo, S. Gooprich, and L.C. Hicains (eds.). 1987. A Utah flora. Great Basin
Naturalist Mem. 9. Brigham Young University, Provo.
Wicains, |.L. 1964. Flora of the Sonoran Desert. In: F. Shreve and I.L.Wiggins, Vegetation and
flora of the Sonoran Desert. Stanford Univ. Press, Stanford. Pt. 2, pp. 180-1669.
156 BRIT.ORG/SIDA 20(1)
BOOK REVIEW
RICHARD L. Duse. 2001. Turfgrasses: Their Management and Use in the Southern
Zone, Second Edition. (ISBN 1-58544-161-9, pbk.) Texas AGM University
Press. John H. Lindsey Building, Lewis Street, 4354 TAMU, College Station,
TX 77843-4354. (Orders: www.tamuedu/upress, 979-458-3982, 800-826-
8911, 979-847-8752 fax). $27.95, 336 pp, 51 color photos, 15b/w photos, 68
line drawings, 7" x 9",
Although monotypic plantings of an exotic grass pce might ek fear or loathing in those con-
=
cerned with biodiversity, few w« uld di pute the impact nomy orout quality
of life. Gulf courses, ae a ee landsca ipesand interstate roadways are all depen-
dent i g furferasses capitalizes on their importance
and seeks vid hensive inf tion on how to grow and maintain lawn
This eoaee ete ises a revision ol i edition in 1989. Chapters include the contributions of
turfgrasses, the turfgrass plant, tl tof turfgrasses, southern turfgrasses, speci-
fication for sir aiaes es een cultural nies weed control in turf, turfgrass diseases, man-
aging turfgrass insects and turfgrass maintenance programs. The second chapter gives a brief, yet
clear introduction to grass morphology. In following chapter, Duble elaborates on the growth and
development of turfgrasses, covering such topics as seed germination, See, ~~ Vv igor, leaf
t slays ively
growth and thatch, and cultural San affecting root oan th. Chapter four i
to southern turfgrasses, dividing them into warm-s season. For each group of grasses
(example: Bermudagrass), he adlndes information on the origin and distribution of species, a spe-
cies esoayuon, ee and uses, horticultural varieties, prepagalion. management and pest
problems. W ron cultural practices, the author incl like mowing,
water management and feviilization, A handy worksheet for water budwering paides the reader in
estimating total cost for irrigation. Special cultural practices address vertical mowing, aerification
spiking and topdressing, to name a ii The section on weec e control nenuns undesirable plants in
turf with both descriptions and li gg control. After reading this
book, one should be able to identify and eat turf diseases and insects since the chapters pertaining
o these topics seem comprehensive. The final chapter has the commercial turf manager in mind
a topics like putting greens and o ing fields.
Overall, this is a well-written and thorough tome. | have but a few quibbles. First, many of the
ds a bit dated. [ noticed only a handful of the citations were published after 1990
with the pour published before 1980. Thus, it seems that some of the material presented herein is
outdated. Turfgrass is big business and big money and the demand for current information should
be reflected with research. While the both the black and white and color photos are clear and repre-
sentative, the line drawings of turfgrass species and turl weeds are of poor quality. The drawing of a
staminate Buffalograss (p. 72), for instance, could easily be mistaken for Bouteloud and Kentucky
bluegrass (p.84) looks more like Johnsongrass. It is likely that the target audience would not be fa-
ee with all the species discussed in this book. A clear illustr. could be a strong confirmation
for identification.
This book is appropriately written with the commercial turfgrass manager in mind. Those
working in this field should not be without a copy of this book.—Amy Trauth Nare, Botanical Re-
search Institute of Texas, Fort Worth, TX, 76102-4060, U.S.A, amy_trauth@yahoo.com
alll
SIDA 20(1): 156. 2002
LECTOTYPIFICATION OF MENTZELIA TEXANA AND
M. LINDHEIMERI(LOASACEAE) WITH AN ASSESSMENT
OF THEIR BIOLOGICAL STATUS
Billie L. Turner
Plant Resources Center
Biolabs 311, University of Texas
Austin, TX 78712, U.S.A.
and
Box 1110, Alpine, TX 79831, U.S.A.
billie@uts.cc.utexas.edu
In their monograph of the family Loasaceae, Urban and Gilg (1900) treated the
genus Mentzelia as having 36 taxa, two of these, M. lindheimeri and M. texana,
proposed as new. The purpose of the present paper is to typify each and provide
an evaluation of their biologic status.
Mentzelia lindheimeri Urban & Gilg, Monogr. Loas. 54. 1900. Type: US.A. TEXAS:
UVALDE Co.: Jun 1885, Reverchon 1650 (LECTOTYPE, here designated: MOI:
ISOLECTOTYPE: US).
Urban and Gilg in their protologue of this species cited five specimens gath-
ered by five different collectors, three of these from Texas, two from northern
Mexico. All of the specimens cited by these two authors were said to be from
European herbaria, namely “Berol., Boisser-Barbey, Canby, Lips., Paris.,J.D. Smith,
Vindobon.” | have examined duplicates of all of the sheets concerned and select
the following from among these as adequate lectotypes: U.S.A. Texas: Uvalde
Co.: Jun 1885, Reverchon 1650 (lectotype: MO!; isolectotype: US!). Darlington
(1934) also listed this collection as belonging to his concept of M. lindheimeri.
Urban and Gilg listed the locality of Reverchon 1650 as “in planitiebus arenosus
montis Upper Guadelupe” but the locality on the lectotype is that given here.
Ideally, a lectotype should be selected from among the specimens exam-
ined by its original author or authors, but the European collections cited by
Urban and Gilg, who worked out of the herbarium at Berlin (B), were unavail-
able to me. Because of this, I find it appropriate (as permissible under Art 9.10 of
the current Code), to designate lectotypes from American syntypes. | have also
applied this same reasoning in the typification of the following taxon.
Mentzelia texana Urban & Gilg, Monogr. Loas. 52.1900. Without specific locality,
May-Oct, 1849, Wright 210 (LECTOTYPE, here designated: GH; ISOLECTOTYPES: GH,
Us),
In the protologue of this taxon, which I consider to be a synonym of M.
lindheimeri, Urban and Gilg noted that, so far as known, it was confined to
SIDA 20(1): 157 — 159. 2002
158 BRIT.ORG/SIDA 20(1)
Texas, citing three specimens: Drummond 91 (from sandy soils between Victo-
ria and Gonzales, Texas; Trecul 1220 (dry sites between Turkey Creek and the
San Pedro [Devils River| and Wright 210 (in part, mixed with M. oligosperma
Nutt.). They also noted that material of their new species, presumably those
cited, were based upon specimens “ in herb. Berol., Boisser-Barbey, Candolle,
Delessert, Kew.,, Paris., Vindobon.”
Darlington (1934) retained the species, citing two of the above mentioned
type elements (Drummond 202 and Wright 202). She also cited three additional
specimens from the state of Pueblo, Mexico. | take the latter to be
misidentifications of the more southern M. hispida Willd., or some close rela-
tive having relatively short petals. Correll and Johnston (1970), following
Darlington, retained M. texana in their treatment of Mentzelia for the flora of
Texas.
Inarecent study of Mentzelia for Texas | have examined duplicates of two
of the three collections cited by Urban and Gilg in their protologue ( Drummond
9land Wright 210) and have selected Wright 210 as an appropriate element for
lectotypification, largely because duplicate specimens of this collection are
currently housed at GH! and US!
The type locality of M. texana is moot. According to label data on Wright
210 (US!) the specimen concerned was collected “May-Oct, 1849.” But other than
this time span, no other information relating to its collection is given. Accord-
ing to Johnston (1940), on June 28, 1849, Charles Wright collected an unnamed
species of Mentzelia (Wright's field number 54D on the hills of Turkey Creek
in western Uvalde County. Since Gray often renumbered and/or combined
Wright's collections (cf. Johnston 1940), it is likely that Wright 210 is the num-
ber assigned by Gray to Wright's field collection 541, at least |encountered no
Wright collection so numbered in the present study.
The positioning of M. texana in synonymy with the simultaneously pub-
lished M. lindheimeri is based upon the study of numerous specimens from
ten or more herbaria, and field work over a number of years in the areas con-
cerned. Urban and Gilg, in their key to species, distinguished M. texana from
M. lindheimeri by its shorter petals (7-9 mm vs. 12-14 mm)and sessile or shortly
pedicellate fruits (vs always clearly pedicellate). | find these two quantitative
characters to be quite variable within the M. lindheimeri complex and have
little hesitancy in reducing M. texana to synonymy. Finally, it should be noted
that Darlington (1934) listed the lectotype of M. texana (Wright 210) among his
“Specimens examined:” as a sheet of M. lindheimeri, which agrees with the
present authors disposition.
—
REFERENCES
Corrett, D.S. and M.C. JoHNsToN. 1970. Manual of the vascular plants of Texas. Texas Res.
Foundation, Renner
TURNER AND M. LINDHEIMERI 159
JOHNSTON, ILM. 1940. Field notes of Charles Wright for 1849 and 1851-52. Typed copy with
commentary by |.M. Johnston, Gray Herbarium (duplicate copy at TEX).
Dartincton, J. 1934. A monograph of the genus Mentzelia. Ann. Missouri Bot. Gard. 21:
103-226.
Urean, |.and E. Gita. 1900. Mentzelia, in Monographia Loasacearum. Nova Acta Acad. Caes.
Leop.-Carol. Germanica Nat. Cur. 76:22-356.
160 BRIT.ORG/SIDA 20(1)
Book RECEIVED/ NOTICE
Annotations and comments by the Editor
DaPHNE GAIL FAutin, DouGLas J. FutuyMA, and H. BRADLEY SHAFFER (eds). 2001. Annual
Review of Ecology and Systematics. Volume 32, 2000. (ISBN 0-8243-1432-
8, hbk; ISSN 0066-4162). Annual Reviews Inc., 4139 El Camino Way, P.O.
Box 10139, Palo Alto, CA 94303-0139, US.A. (Orders: www.AnnualReviews.org,
800-523-8635, 650-493-4400, 650-424-0910 fax). $140.00 hbk., 614 pp.
oxo.
Readers of Sida will find a few articles of interest in this Annual Review of Ecology and Systematics.
Vo
uting Authors, and Cumulative Index of Chapter Titles (Vols. 28-32).
ume 32 for the year 2001 has 19 articles followed by a Subject Index, Cumulative Index of Contrib-
Contents.—
. Chemical detection of natural enemies by arthropods: An ecological perspective
Cc }
d
: hromos meiotic drive
Environmental influences on regional deep-sea species diversity
The physiology of life history trade-offs in animals
Saas
Urban ecological systems: Linking terrestrial ecological, physical, and socioeconomic compo-
nen of metropolitan areas
6. Same? in freshwater invertebrates
Annlied evolution
be
8. Mistletoe—a Key stone resource in forests and woodlands worldwide
9. The role of disturbance in the ecology and conservation of birds
10 proaches to the study of territory siz pend shape
Ll. The population biology of invasive species
12. Streams in the urban landscape
13. Integrating function and ecology in studies of adaptation: Investigations of locomotor capacity
as a model iow
l4. TI th conifer family . History, status, and value for palaeoenvironmental
A
15. The units of selection on mitochondrial
lo. Evolutionary patterns sia aes THasi Therapsids
17. Ecology, conservation, and public po
18. Male-killing, nematode infections, bacter nee infection, and virulence of cytoplasmic bac-
teria in the genus Wolbachia
19. Biospheric trace gas fluxes and their control over tropospheric chemistry
SIDA 20(1): 160. 2002
NOMENCLATURAL CHANGES AND INNOVATIONS IN
PANICUM AND DICHANTHELIUM (POACEAE: PANICEAE)
Robert W.Freckmann Michel G. Lelong
Department of Biolo Department of Biological Sciences
University of Wisconsin-Stevens Point University of South Alabama
Stevens Point, WI 54481 U.S.A. Mobile, AL 36688 U.S.A
ABSTRACT
oy
Infrageneric taxa in Panicum and Dichanthelium have a long history of usage, but at least eight of
these taxa have not been validly published previously. The new taxa published here are Panicum
sect. Antidotalia, P. sect. Hemitoma, Dichanthelium sect. Clandestina, D. sect. Linearifolia, D. sect.
Macrocarpa, D. sect. Nudicaulia, D. sect. Pedicellata, and D. sect. Strigosa. es two new combin
tions, made for a forthcoming treatment of these two genera, are as follows: D. sect. Angustifolia; D.
sect. Ensifolia; D. sect. Lancearia; D. sect. Lanuginosa; D. sect. Oligosantha; D. sect. Sphaerocarpa; D.
aciculare subsp. angustifolium, subsp. fusiforme, subsp. neuranthum, D. acuminatum subsp.
columbianum, subsp. fasciculatum, subsp. implicatum, subsp buco: subsp MRGREUB EN: subsp.
tongiigulatum, SUPSP: sericeum subsp spretum subsp thermale. D breve;
ashei, subsp equilaterale subsp. joori; D.dichotomum subsp. haetaui. subsp.
C
ee ae aa ees SUED: nitidu, suDSp Seales subsp. yadkinense; D.
ubsp. praecocius, subsp
if
pieudbpubecccne subsp villosissimu: me portoricense subsp. ne D. strigosum subsp.
elabtescens subsp ! p.amarulum; Hareapssun=p) a comiabatt
I . il ii: P dich ifl | PET sUDSE 3
Pate bsp. sonorum Cres stramineum, P. p} bsp. gattingeri, eubsp
lithophilum; and P. as ae subsp. abscissum, subsp. com bsii, subsp. elongatum, subsp. pubescens
RESUMEN
Los taxones ieee en Panicum y Dichanthelium han sido usados durante mucho tiempo,
pero por ee menos ocho de estos taxones no han sido aeistes validamcas: Los nuevos taxones
aqui publicados son Panicum sect. Antidotalia, P sect. Hemitoma, Dic] t. Clandestina,
sect. ee ee . sect. Macrocarpa, D. sect. Nae Dz sect. Pedicellata, y D. sect. Strigosa.
Cacia y dos nuevas combinaciones, designadas en un futuro tratamiento d géneros,
son: D. sect. Angustifolia; D. sect. Ensifolia; D. sect. Lancearia; D. sect. Lanuginosa; D. sect. ee
D. sect. Sphaerocarpa; D.sect. Angustifolia; D. sect. Ensifolia; D. sect. Lancearia, D. sect. Lanuginosa; D
sect. Oligosantha, D. sect. Sphaerocarpa;, D. aciculare subsp. ae ium, oe fusiforme, subsp.
neuranthum, D. acuminatum subsp. columbianum, subsp. fasciculat ubsp. implicatum, subsp
leucothrix, subsp. li aac es longiligulatum, subsp. sericeum, ae sp. spretum, saben:
thermale; D.chamaelonche y subsp. breve; D. Conia tam subsp. ashei, subsp. equilaterale, subsp
i: D. dich I je subsp tt t subsp. microcarpon, subsp. nitidum
joort
subsp. rodnokense, scbep: yadkinense; D. ensifolium subsp. curtifolium,; D. oligosanthes subsp.
scribnerianum; D. ovale subsp. praecocius, subsp. pseudopubescens, subsp. villosissimum, D.
portoricense subsp. patulum; D strigosum subsp. ee subsp. leucoblepharis; D. tenue; P.
amarum subsp. amarulum; P. anceps subsp. rhizomatum, P. capillare SUpeP: hilmanit P.
lichotomifl sp. bartowense owe ‘puritanoram, P. hallii subsp. filipes; P. hirt
sonorum, subsp. stramineum; P bsp. gattingeri, subsp. lithophilum; y Prieihilien
subsp abscissum subsp oe ee subsp aoaeaann cabo pubescen
SIDA; 20 1): 161 — 174. 2002
162 BRIT.ORG/SIDA 20(1)
In preparing the treatment of Panicum and Dichanthelium for Volume 25 of
The Flora of North America (Barkworth et al. leds, in prep.), we find it neces-
sary to describe eight new infrageneric taxa and make fifty-two new combina-
tions. Editorial policy for this volume requires that infraspecific taxa be treated
as subspecies rather than as varieties.
INFRAGENERIC NAMES
The genus Panicum, as recognized by Linnaeus (1753), included 20 species in
two groups: Spicata and Paniculata. Most of these species have been tranferred
subsequently to other genera, including Digitaria, Echinochloa, Oplismenus,
Paspalum, Pennisetum, and Setaria. The segregation of Dichanthelium from
Panicum by Gould (1974) on characters of anatomy and photosynthetic path-
ways is supported by a recent molecular phylogeny of the Panicoideae (Giussani
et al. 2001).
Bentham and Hooker (1883) treated Panicum as a relatively broad genus
and published several infrageneric taxa. Nash (1903) treated Panicum ina nar-
rower sense. He named and described nine unranked groups of Panicum in
Small, Flora of the Southeastern United States. Even though no infrageneric rank
was indicated, these names were validly and effectively published, and eight of
these names can serve as basionyms in Panicum or Dichanthelium. Hitchcock
revised the Gramineae for Robinson and Fernald (1908) in Gray’s Manual of
Botany, ed. 7. He divided Panicum in this treatment into two sections, the rank
indicated by the double-s sign, which Robinson and Fernald define on p. 32 as
indicating a section. He divided section Eupanicum into six unranked names,
presumably subsections, and added 14 names, presumably series, within the
“subsection” Dichotoma. Although these 20 infrasectional names are unranked,
these names were also validly published and can also serve as basionyms.
Hitchcock and Chase (1910) added 13 additional unranked groups in Pani-
cumand effectively published descriptions of these groups. Several subsequent
taxonomists (e.g. Fernald 1936, 1938, 1947; Hsu 1905) have treated their group
names as sectional or subsectional names. Hitchcock and Chase, however, ex-
plicitly stated (page 17) that they did not consider these groups to be formal
taxonomic entities. They used the same group names in their later publications
(Hitchcock 1913, 1915, 1935, 1951: Hitchcock and Chase 1915) but never suggested
that they had changed their minds with respect to the taxonomic status of the
groups. Article 34.1 of the International Code of Botanical Nomenclature
(Greuter et al. 2000) states that a name is not validly published when it is not
accepted by the author in the original publication. It seems likely that, had
Hitchcock changed his mind regarding these group names, he or Chase would
have included that information in one of their later publications for Hitchcock
was a very active participant in the development of the nomenclatural code
(Hitchcock 1919; Lawrence 1951) and hence must have been aware of the need
~s
NOMENCLATURAL 163
tomakeaclear statement of sucha change of mind. Therefore, if the Hitchcock
and Chase groups added in the 1910 paper are to be recognized as formal taxa,
they must subsequently be validly published. We have indicated that we are
deriving our section names from these group names in the 1910 paper by using
“Hitchcock & Chase ex Freckmann & Lelong” for those groups which retain
the Hitchcock and Chase group name and circumscription.
Pilger 1931) named new sections in Panicum, indicating these as “nov. sect.,”
meeting all requirements that apply to a 1931 publication. He also apparently
published validly, but inadvertently, as sections two of the Hitchcock and Chase
groups by reference to the Hitchcock and Chase names in the 1910 and 1915
papers, even though he did not indicate these as “nov. sect.”
Hsu (1965) is often cited as validating Hitchcock and Chase names or as
publishing new combinations based on these names, but no types were cited,
as required after 1 January 1958 by Article 37.1 of the Code. Gould (1974) raised
subgenus Dichanthelium to a genus, making new combinations for 17 species
and three varieties, but he did not treat subgenera or sections. Brown in Brown
and Smith (1975) elevated two sections of Panicum to subgenus status in
Dichanthelium. Zuloaga (1987) returned to a broader concept of Panicum and
published four new sections, each with a Latin description and citation of a
type. He also changed the status of three of Hitchcock’s group names to sec-
tions, citing types and validly publishing these names.
NEW TAXA
Panicum sect. Antidotalia Freckmann & Lelong, sect. nov. Typus: Panicum
antidotale Retz
Plantae perennes rhizomatibus crassis. Culmi erecti duri robusti, e nodis medius ramosi. Ligulae
mebranaceae ciliatae. Paniculae pyramidalis, plus minusve laxae, multiflorae, spiculis distincte
pedicellatis. Spiculae ovoideae glabrae; gluma inferior circa dimidiam longitudinem spiculae
attingens, 3-5 nervia; gluma superior et lemma inferior 7-nervia; anthoecium inlerius staminiferis;
anthoecium superius glabrum, nitidum
Plants perennial, with thick rhizomes. Culms erect, robust, hard, branching at
the middle nodes. Ligule a ciliate membrane. Panicle pyramidal, more or less
lax, many-flowered, with spikelets on long pedicels. Spikelets ovoid, glabrous;
first glume about 1/2 as long as the spikelet, 3- or 5-nerved; second glume and
lower lemma 7-nerved; lower floret staminate; upper floret smooth, shiny. x = 9.
Panicum sect. Hemitoma Hitchc. & Chase ex Freckmann & Lelong, sect. nov.
Typus: Panicum hemitomon Schult
D] ee 1 ]
| Culmierecti vel reclinati, saepe sterilis. Ligu-
L
lae menbrapecene ciliatae vel beens Paniculae angustae; rami pauci erecti spiciformes spiculis
secundis subsesillibus. Spiculae anguste ovoideae vel lanceolatae lateraliter compressae, glabrae.
Anthoecium inferius staminiferis; lemma superior tenuis, flexilibus, albida, involuta non nisi prope
basin.
164 BRIT.ORG/SIDA 20(1)
Plants perennial, semi-aquatic, with extensive rhizomes or stolons. Culms erect
or sprawling, often sterile. Ligule a ciliate or lacerate membrane. Panicle nar-
row; branches few, spiciform, erect, with subsessile to short-pedicelled, secund
spikelets. Spikelets narrowly ovoid to lanceolate, laterally compressed, glabrous.
Lower floret staminate; upper lemma thin, flexible, whitish, rolled over the palea
—_
only near the base.
Dichanthelium sect. Clandestina Freckmann & Lelong, sect. nov. Typus: Panicum
clandestinum L.
Plantae | it is vel caudicibus. Culmi 50-150 mm alti; culmiautumnalie nodis
I
13 Ree | a a | { t im . Ae ] ] 7 =e
il | ramulosi Sae Pe if
Es qT I
intra foliorum vaginas. Folia basilaria rosulam hiemalem formantes. Vagina tumida, maculata,
papillosa-hispida vel pubescentia et viscida, ore constricta. Lamina basilaria grandis, ovata-lanceolata;
laminae caulinae 6-14. Spiculae ellipsoideae vel ovoideae, acutae, 2.2-3.6 mm longa, nervis
prominibus.
Plants perennial, froma thick rhizome or caudex. Culms 50-150 cm; autumnal
culms with branches arising from the middle and upper nodes, rebranching
and often forming dense axillary fascicles, the panicles enclosed in the sheaths.
Basal leaves forming a winter rosette. Sheath usually swollen, mottled, papil-
lose-hispid or pubescent and viscid, constricted at summit; basal blades large,
ovate-lanceolate; cauline blades 6-14. Spikelets ellipsoid to ovoid, pointed, 2.2-
3.6 mm long, with prominent veins.
Dichanthelium sect. Linearifolia Freckmann & Lelong, sect. nov. Typus: Panicum
linearifolium Scribn. ex Nash.
Plantae perennes caespitosae. Culmi erecti vel effusi vel cernui; internodia inferna brevissima;
internodia supera saepe manopere elongata. Culmi autumnali e nodis prope basin ramosi, paniculas
condensatas minus quam 5 em super terram vel culmos steriles formantes. Laminae basilares
angustae, erectae vel ascendes, laminas caulinas infernas simulans; laminae superae plerumque
erectae, rigidae, 1-5 mm latae, 15-60-plo longiora quam latiora. Spiculae ovoideae, ellipsoideae, vel
obovoideae, 2.0-4.3 mm longa, glabrae vel pubescentae.
Plants perennial, tufted. Culms erect to spreading or drooping, the lower inter-
nodes very short, the upper 2-4 internodes often much elongated. Autumnal
culms branching from the nodes near the base, producing condensed panicles
within about 5cm of the ground or sterile shoots. Basal blades narrow, erect or
ascending, resembling the lower cauline ones; cauline blades usually erect, stiff,
the upper ones 1-5 mm wide and 15-60 timesas long. Spikelets ovoid, ellipsoid,
or obovoid, 2.0-4.3 mm long, glabrous or pubescent.
Hitchcock and Chase (1910) used the group name “Depauperata” for most
of the species in this section. However, because Pilger (1931) published section
Depauperata for certain Old World species of Panicum and because
Dichanthelium is often included in Panicum, we preter to avoid creating a hom-
onym if these genera are combined.
iu
NOMENCLATURAL 165
Dichanthelium sect. eis Freckmann & Lelong, sect. nov. Typus: Panicum
macrocarpon LeConte ex
Plantae perennes rhizomatibus brev ibus uel caudicibus, raro ehizomia es Elongavibis: ‘Colm
| sparse ramosi, {
principales 20-110 cm
culmi secundarii a autumnali, parviores. Folia basilaria rosulam hiemalem formantes.
Foliorum vaginae glabrae vel pe eae non eee laminae caulinae 3-7. Paniculae saltem
partim exsertae Anthoecium l
dD)
I ial, froma knotted ae a caudex, or rarely an elongated rhizome.
Main culms 20-110 cm, ascending to erect, sparsely rebranching, not producing
dense axillary fascicles; additional culms (sometimes arising in autumn) smaller.
Basal leaves forming a winter rosette. Sheaths glabrous or pubescent, not viscid;
cauline blades 3-7. Panicle at least partly exserted. Lower floret often staminate.
Nash (1903) published the infrageneric name Latifolia within Panicum,
indicating that this name was based on Panicum latifolium L. However, it is
evident from his description “much branched shrubs...outer [spikelet] scales
woolly-tipped”) that this name was based on a misapplication of Panicum
latifolium, referring instead to Panicum (=Lasiacis) divaricatum L. According
to Art. 7.4 of the Code a name is typified by the type of the basionym, even
though it may have been applied to a taxon now considered not to include that
type. However, in the case of an unranked infrageneric name, a modern author
may publish a new name at a definite rank. We prefer to avoid confusion over
the application of Latifolia by basing this section name on a different type.
_—
ie sect. Nudicaulia Hitchcock & Chase ex Freckmann & Lelong,
ct. nov. TyPus: Panicum nudicaule Vasey.
diy | Lali ] t ] Als
Plantae perennes caesptosae. Culmi 20-60 cm alti, i
] ]
]
TO e basin ramosi. Laminae lanae el rigide involutae,ascen andes
redactae. Folia basilaria rosulam hiemalem formantes. Panicula ious Ses pauc cil lora. Spiculae
anguste ellipsoideae, 2.4-2.9 mm longa, glabrae.
Plants perennial, tufted. Culms 20-60 cm, weakly ascending, glabrous; autum-
nal culms branching from the base. Blades flat to stiffly involute, ascending,
crowded at the base, the uppermost ones much reduced. Basal leaves not form-
ing a distinct winter rosette. Panicle long-exserted, few-flowered. Spikelets nar-
rowly ellipsoid, 2.4-2.9 mm long, glabrous.
Dichanthelium sect. Pedicellata Hitchc. @ Chase ex Freckmann & Lelong, sect.
nov. Typus: Panicum pedicellatum Vasey.
Plantae perennes basibus duris cormiformibus et rhizomatibus gangliiformibus. Culmi erecti vel
effusi; culmi autumnali divaricate ramosi ante maturitem paniculae ad finem decumbes. Folia
] ee leah; ] ; licul } + i : -7] +1;
Oo I c
Spiculae obovoideae vel obpyriformes, 3.2-4.4 mm longa, papillatae pubescentae, basi attenuatae.
Anthoecium superius ad apicem puberulum, acutum vel attenuatum.
Plants perennial, from a hard, corm-like base and a knotted rhizome. Culms
166 BRIT.ORG/SIDA 20(1)
erect or spreading; autumnal culms freely divaricate-branching before the pri-
mary panicle matures, ultimately decumbent. Basal leaves not forming a win-
ter rosette. Ligule a ciliate membrane. Blades papillose-ciliate. Spikelets obo-
void to obpyriform, 3.2-4.4 mm long, papillose-pubescent, attenuate at the base.
Upper floret pointed to attenuate, puberulent at the apex.
Dichanthelium sect. Strigosa Freckmann & Lelong, sect. nov. Typus: Panicum
strigosum Muhl. in Elliott.
Plantae perennes dense caespitosae. Culmi principales graciles; internodia inferna breves; 3-5
les
internodia supera elongata. Culmi autumnali pulvinum densum formantes. Laminae mo
pallidovirens vel flavovirens, plerumque marginibus ciliatis. Laminae basilares grandes laminas
caulinas infernas simulans. Spiculae late ellipsoideae vel obovoideae, L.1-2.3 mm longa, glabrae vel
pubescentae.
Plants perennial, densely tufted. Main culms slender, the lower internodes short,
the upper 3-5 internodes elongated. Autumnal culms forming a dense cushion.
Blades soft, pale green or yellowish, usually with ciliate margins. Basal blades
large, scarcely separable from the lower cauline blades. Spikelets broadly ellip-
soid to obovoid, 1.1-2.3 mm long, glabrous to pubescent.
Bentham and Hooker (1883) created the section Laxiflora to include cer-
tain Old World species of their subgenus Eupanicum, but not including P.
laxiflorum Lam. To avoid creating a later homonym if Dichanthelium is treated
as part of Panicum, we are basing the section name on Panicum strigosum Muhl.
in Elliott, rather than P laxiflorum.
NEW COMBINATIONS
Dichanthelium sect. Angustifolia (Hitche. in B.L. Rob. @ Fernald) Freckmann
& Lelong, stat. nov. BASIONYM: Panicum (unranked) Angustifolia Hitche. in BLL.
Rob. & Fernald, Gray’s Man. Bot. Ed. 7:106. 1908. Type: Panicum angustifolium Elliott.
Dichanthelium sect. Ensifolia (Hitchc.in B.L. Rob. @ Fernald) Freckmann &
Lelong, stat. nov. BASIONYM: Panicum (unranked) Ensifolia Hitche. in B.L. Rob.
& Fernald, Gray’s Man. Bot, ed. 7:113. 1908. Tyre: Panicum ensifolium Baldwin ex
Elliott.
Dichanthelium sect. Lancearia (Hitche. in B.L. Rob. & Fernald) Freckmann &
Lelong, stat. nov. BASIONYM: Panicum (unranked) Lancearia Hitche. in B.L. Rob.
& Fernald, Gray’s Man. Bot, ed. 7:114. 1908. Type: Panicum lancearium Trin.
Dichanthelium sect. Lanuginosa (Hitchc. in B.L.Rob. & Fernald) Freckmann &
Lelong, stat. nov. BASIONYM: Panicum (unranked) Lanuginosa Hitche. in B.L. Rob.
& Fernald, Gray’s Man. Bot, ed. 7:109. 1908. Type: Panicum lanuginosum Elliott.
Dichanthelium sect. Oligosantha (Hitchc. in B.L.Rob. & Fernald) Freckmann &
Lelong, stat. nov. BASIONYM: Panicum (unranked) Oligosanthia Hitche. in B.L. Rob,
& Fernald, Gray’s Man. Bot, ed. 7:114. 1908. Type: Panicum oligosanthes Schult.
Dichanthelium sect. Sphaerocarpa (Hitchc. in B.L.Rob. & Fernald) Freckmann
Lelong, stat. nov. BASIONYM: Panicum (unranked) Sphaerocarpa Hitchce. in BL.
Rob. & Fernald, Gray’s Man. Bot, ed. 7:113. 1908. Type: Panicum sphaerocarpon Elliott.
FRECKMANN AND LELONG, NOMENCLATURAL 167
Dichanthelium aciculare subsp. angustifolium (Elliott) Freckmann & Lelong,
comb. & stat. nov. BASIONYM: Panicum angustifolium Elliott, Sketch Bot. S. Caro-
lina & Ga. 1:129.1816. Panicum nitidum var. angustifolium (Elliott) A.Gray, N. Amer,
Gram. 2:112.1835. Chasea angustifolia (Elliott) Nieuwl, Amer. Midl. Naturalist 2:64.
1911. Dichanthelium angustifolium (Elliott) Gould, Brittonia 26:59. 1974. Panicum
aciculare var. angustifolium (Elliott) Wipff & S.D. Jones, Phytologia 77:457. 1994.
TYPE: U.S.A. SOUTH CAROLINA: May, S. Elliott s.n. HOLOTYPE: CHARL, fragment USI).
mh
Dichantheli ] bsp. fusiforme (Hitche.) Freckmann & Lelong, comb.
& stat. nov. BASIONYM: Panicum fusiforme Hitchc., Contr. U.S. Natl. Herb. 12:222.
1909. Dichanthelium fusiforme (Hitchce.) Harvill, Castanea 42:177. 1977. TYPE:
CUBA: western Cuba, 1863, C. Wright 3454 (HOLOTYPE: GOET, fragment USI).
Panicum neuranthum var. ramosum Griseb., Cat. Pl. Cub. 232. 1866. Dichanthelium aciculare var.
ramosum (Griseb.) Davidse, Novon 2:104. 1992.
Dichanthelium aciculare subsp. neuranthum (Griseb.) Freckmann & Lelong,
nov. BASIONYM: Panicum neuranthum Griseb., Cat. Pl. Cub. 232.
1866. TYPE: CUBA: eastern Cuba, 1860, C. Wright 3453 (HOLOTYPE: US 2209346)).
Panicum pinetorum Swallen, Proc. Biol. Soc. Wash. 55:93. 1942.
theli i (Scribn.) Freckmann & Lelong,
comb. & stat. nov. BASIONYM: Panicum columbianum Scribn., US.D.A. Div. Agrost
Bull. 7:78, f.6.1897. D ) Freckmann Phytologia
39:270. 1978. Panicum acuminatum var. columbianum (Scribn.) Lelong, Brittonia
36:270. 1984. TyPE: U.S.A. DISTRICT OF COLUMBIA: 14 Jul 1894 EL. Scribner s.n. (HO-
LOTYPE: US 743463).
Dichanthelium acuminatum subsp. fasciculatum (Torr.) Freckmann & Lelong,
Stat.. nov. BASIONYM: Panicum dichotomum var. fasciculatum Torr., Fl.
North. & Mid. U. S. 1:145. 1824. Panicum huachucae var. fasciculatum CTorr.) FT.
Hubb., Rhodora 14:171.1912. Panicum lindheimeri var. fasciculatum (Torr) Fernald,
Rhodora 23:228. 1921. Panicum lanuginosum var. fasciculatum (Torr.) Fernald,
Rhodora 36:77. 1934. Dichanthelium lanuginosum var. fasciculatum (Torr)
Spellenb. Madrono 23:145. 1975. Dichanthelium acuminatum var. fasciculatum
(Tor r.) Fr eckmann, Phytologia 48:108. 1981. Panicum acuminatum var. fasc iculatum
(Torr.) Lelong, Brittonia 36:269. 1984. TYPE: U.S.A. NEW JERSEY: sandy fields, J. Torrey
s.n. (HOLOTYPE: NY1).
Dichanthelium acuminatum subsp. implicatum (Scribn. ex Nash) Freckmann
Lelong, comb. & stat. nov. BASIONYM: Panicum implicatum Scribn. ex Nash
in Britton & A. Br, Ilust. Fl. 3:498, f. 267a. 1898. Panicum unciphyllum var.
implicatum (Scribn. ex Nash) Scribn. @ Merr. Rhodora 3:123. 1901. Panicum
lindheimeri var. implicatum (Scribn. ex Nash) Fernald, Rhodora 23:228. 1921. Pani-
cum lanuginosum var. implicatum (Scribn. ex Nash) Fernald, Rhodora 36:77. 1934.
Dichanthelium acuminatum var. implicatum (Scribn. ex Nash) Gould & C.A. Clark,
Ann. Missouri Bot. Gard. 65:1126. 1979 [imprint date 1978]. Panicum acuminatum
var. implicatum (Scribn.ex Nash) Beetle, Phytologia 48:192. 1981. TYPE: U.S.A. MAINE:
low marshy ground, Cape Elizabeth, 26 Jul 1895, FL. Scribner s.n. (HOLOTYPE: US
7428961).
Dichanthelium acuminatum subsp. leucothrix (Nash) Freckmann & Lelong,
.& stat. nov. BASIONYM: Panicum leucothrix Nash, Bull. Torrey Bot. Club
24:41. 1897. Dichanthelium leucothrix (Nash) Freckmann, Phytologia 48:101. 1981.
168 BRIT.ORG/SIDA 20(1)
Panicum dcuminatum var. leucothrix (Nash) Lelong, Brittonia 36:271. 1984. TYPE:
U.S.A. FLORIDA: Lake Co. low pineland at Eustis, 16-31 Jul 1894, G.V. Nash s.n. (HO-
LOTYPE: NYY)
Dichanthelium acuminatum subsp. lindheimeri (Nash) Freckmann & Lelong,
.@ stat. nov. BASIONYM: Panicum lindheimeri Nash, Bull. Torrey Bot. Club
24:196. 1897. Panicum lanuginosum var. lindheimeri (Nash) Fernald, Rhodora
36:77. 1934. Dichanthelium lindheimeri (Nash) Gould, Brittonia 26:60. 1974.
Dichanthelium lanuginosum var. lindheimeri (Nash) Freckmann, Phytologia
39:270. 1978. Dichanthelium acu Hes m var. lindheimeri (Nash) Gould & C.A.
Clark, Ann. Missouri Bot. Gard. 65:1127. 1979 [imprint date 1978]. Panicum
acuminatum var. fees (Nash) | aes Phytologia 48:193. 1981. Panicum
lindheimeri var. typicum Fernald, Rhodora 23:227. 1921. Type: U.S.A. TEXAS: springs,
banks of the Guadeloupe, near New Brauntels, 1846, FJ. Lindheimer 565 (HOLO-
; 1
Dichantheli i bsp.1 latum (Nash) Freckmann & Lelong,
comb. & stat. nov. BASIONYM: t Panic m ae rte Nash, Bull. Torrey Bot. Club
26:574.1899. Dichantheli cuminatum var. longiligulatum (Nasl hn) Goul d&CaA.
Clark, Ann. Missouri Bot. “Gard. 65:1127-1128. 1979 limprint date 1978].
Dichanthelium longiligulatum (Nash) Freckmann, Phytologia 48:102. 1981. Pani-
cum acuminatum var. longiligulatum (Nash) Lelong, we 30:270. 1984. TYPE
U.S.A. FLORIDA: Apalachicola, 1892, G.R. Vasey (HOLOTYPE: NY
Dichanthelium acuminatum subsp. sericeum (Schmoll) ecteaas & Lelong,
c . & stat. nov. BASIONYM: Panicum ferventicola var. sericeum Schmoll,
Madrono 5:92. 1939. Dichanthelium lanuginosum var. sericeum (Schmoll) Spellenb.,
Madrono 23:150. 1975. Dichanthelium acuminatum var. sericeu m scones
Freckmann, Phytologia 48:107. 1981. Type: U.S.A. WYOMING: Mi h Hot Springs,
Yellowstone National Park, in wet ground about hot springs, A. Nelson 6037 (HO-
LOTYPE: RM 20108).
bani aes acuminatum subsp. spretum (Schult.) Freckmann & Lelong,
. & stat. nov. BASIONYM: Panicum spretum Schult., Mant. 2:248.1824.
i hanthelium spre (Schult.) Freckmann, Phytologia 48:102. 1981. Tyre: U.S.A.
New England, moist ground, Muhlenberg’s Panicum No. 37 (HOLOTYPE: PH).
Dichanthelium acuminatum subsp. thermale (Bol.) Freckmann & Lelong, comb.
Stat. nov. BASIONYM: Panicum thermale Bol. Proc. Calif. Acad. Sci. 2:181. 1862.
Dichanthelium lanuginosum var. thermale (Bol.) Spellenb., Madrono 23:151. 1975.
Dichanthelium acuminatum var. thermale (Bol.) Freckmann, Phytologia 48:107.
1981. TYPE: U.S.A. CALIFORNIA. Sonoma Co. northern part of Sonoma County, on
hot rocks and in hot water flowing from Geyser Springs and Geyser Mountains,
H.N. Bolander 3941 (HOLOTYPE: GH)
Dichanthelium chamaelonche ( Trin.) Freckmann G Lelong, comb. nov. BASIONYM:
Panicum chamaelonche Trin., Gram. Panic. 242. 1826. Type: U.S.A. “AMER. BOR.”
Fnslins.n. (HOLOTYPE: LE; fragment and photograph US!).
—
——
Dichanthelium chamaelonche subsp. breve (Hitchc. @ Chase) Freckmann &
_elong, comb. & stat. nov. BASIONYM: Panicum breve Hitchc. & Chase, Contr.
U.S. Natl. Herb. 15:271. 1910. Dichanthelium dichotomum var. breve (Hitchc. &
Chase) Gould & CA. Clark, Ann. Missouri Bot. Gard. 65:1120. 1979 [imprint date
1978]. Panicum chamaelonche var. breve (Hitche. & Chase) Lelong, Brittonia 36:267.
NOMENCLATURAL 169
1984. Dichanthelium ensifolium var. breve (Hitche. &@ Chase) B.F. Hansen &
Wunderlin, Ann. Missouri Bot. Gard. 75:1646. 1989 [imprint date 1988]. Panicum
ensifolium var. breve (Hitchc. & Chase) Wipff & S.D. Jones, Phytologia 77:460. 1994
: ORIDA: Jensen, low pine woods between scrub hills, among pal-
metto, 5 Apr 1906, A.S. Hitchcock 734 (HOLOTYPE: US 558435).
Dichanthelium commutatum subsp. ashei (G. Pearson ex Ashe) Freckmann &
Lelong, comb. & stat. nov. BASIONYM: Panicum ashei G. Pearson ex Ashe, J. Elisha
Mitchell Sci. Soc. 15:35 1898). Panicum con fe var. ashei(G. Pearson ex Ashe)
Fernald, Rhodora 36:83 (1934). TYPE: U.S.A. NEW YorRK: Ithaca, Jul 1898, W.W. Ashe
s.n. (LECTOTYPE [ISOTYPE] US!, [designated by Hitchc. & Chase, Contr. US. Natl. Herb.
15:301. 1910.)
Dichanthelium tat b ilaterale (Scribn.) Freckmann & Lelong,
comb. & stat. nov. BASIONYM: Panicum equilaterale Scribn., U.S.D.A. Div. Agrost.
Bull 11:42. 1898. Type: U.S.A. FLORIDA: Lake Co.: scrubby hammock lands, Eustis,
16-25 Aug 1894, G.V. Nash 1674 (LECTOTYPE: US 7439291 [designated by Hitchcock
& Chase, Contr. U.S. Natl. Herb. 15:310. 1910.)
Dichanthelium commutatum subsp. joori (Vasey) Freckmann & Lelong, comb.
stat. nov. BASIONYM: Panicum joori Vasey, U.S.D.A. Div. Agrost. Bull. 8:31.1889.
Type: U.S.A. LOUISIANA: East Baton Rouge Par, near Baton Rouge, creek bank, in
dense tufts, 1 Oct. 1885, J.F Joor 39 (HOLOTYPE: US!). Panicum divergens Kunth in
H.B.K. has been used by Zuloaga et al. (1993) to replace P. commutatum. However,
Davidse & Zuloaga (2000) in proposing rejection of P. divergens note that the prov-
enance of the type collection is uncertain and that the name had never been ap-
plied ade) to P commutatum.
Dichanthelium dichot subsp. lucidum (Ashe) Freckmann & Lelong, comb.
stat. NOV. BASIONYM: Panicum lucidum Ashe, J. Elisha Mitchell Sci. Soc. 15:47.
1898. Panicum dichotomum var. lucidum (Ashe) Lelong, Brittonia 36:265. 1984.
Dichanthelium lucidum (Ashe) LeBlond, Sida 19(4):831. 2001. TyPE: U.S.A. NORTH
my INA: deep shady swamps bordering Lake Mattamuskeet, 10 Jun-6 Jul 1898,
Ashe s.n. ‘i ECTOTYP oe US! [designated by Hitchcock & Chase, Contr.
US. tr Herb. 15:198. 1910.)
Dichanthelium dichotomum subsp. mattamuskeetense (Ashe) Freckmann &
elong, comb. & stat. nov. BASIONYM: Panicum mattamuskeetense Ashe, J. Elisha
aoe Sci. Soc. 15:45. 1898. Panicum dichotomum var. mattamuskeetense (Ashe)
Lelong, Brittonia 36:265. 1984. Dichant pal mattamuskeetense (Ashe) LeBlond,
Sida 19:832. 2001. Type: U.S.A. NORTH CAROLINA: around Lake Mattamuskeet, 10
Jun-6 Jul 1898, W.W. Ashe and G. Pearson s.n. vA ECTOTYPE: US 2808955! [designated
by Hitchcock & Chase, Contr. U.S. Natl. Herb. 15:186. 1910.)).
Dichanthelium dichotomum subsp. microcarpon (Muhl. ex Elliott) Freckmann
& Lelong, comb. & stat. nov. BASIONYM: ee m mic alas Muhl. ex Elliott,
Sketch Bot. S. Carolina 1:127.1816. Type: U.S.A. ¢ RGIA, A.A. Baldwin s.n. (LECTO-
TYPE: CHARL [designated by Hitchc. & Chase, a U.S. Natl. Herb. 15:181.1910.)).
Dichanthelium dict subsp. nitidum (Lam.) Freckmann & Lelong, comb.
& stat. nov. BASIONYM: Panicum nitidum Lam., Tabl. Encycl. 1:172.1791. Panicum
dichotomum var. nitidum (Lam.) Alph. Wood, Class-book bot., ed. s.n. [b], 786. 1861.
Dichanthelium dichotomum var. nitidum (Lam.) LeBlond, Sida 19(4):829. 2001.
Type: U.S.A. “CAROLINA”: eastern Carolina, J. Fraser s.n. (HOLOTYPE: P fragment US!).
170 BRIT.ORG/SIDA 20(1)
Dichanthelium dichotomum subsp. roanokense (Ashe) Freckmann & Lelong,
comb. & stat. nov. BASIONYM: Panicum roanokense Ashe, J. Elisha Mitchell Sci.
Soc. 15:44. 1898. Panicum dichotomum var. roanokense (Ashe) Lelong, Brittonia
36:265. 1984. Dichanthelium dichotomum var. roanokense (Ashe) LeBlond, Sida
19(4):831. 2001. Type: U.S.A. NORTH CAROLINA. Dare Co.; Manteo, Roanoke Island,
dry soil, 10 Jun 1898, W.W. Ashe s.n. (LECTOTYPE: USdesignated by Hitche. & Chase,
Contr. U.S. Natl. Herb. 15:196. 1910).
Dichanthelium dichotomum subsp. yadkinense (Ashe) Freckmann & Lelong,
comb. & stat. nov. BASIONYM: Panicum yadkinense Ashe, J. Elisha Mitchell Sci.
Soc. 16:85. 1900. Panicum dichotomum var. yadkinense (Ashe) Lelong, Brittonia
36:266. 1984. Dichanthelium yadkinense (Ashe) Mohlenbr, Erigenia 6:27. 1985.
Type: U.S.A. NORTH CAROLINA. Raleigh, May 1895, W.W. Ashe s.n. (LECTOTYPE: US
2808938! [designated by Hitche. & Chase, Contr. US. Natl. Herb. 15:195. 1910).
aout ensifolium subsp. curtifolium (Nash) Freckmann & Le ,
.& stat. nov. BASIONYM: Panicum curtifolium Nash, Bull. Torrey Bot. Club
26: 569 1899. Panicum ensifolium var. curtifolium (Nash) Lelong, Brittonia 36:266.
1984. TYPE: U.S.A. MIssISssIPPL Ocean Springs, 2 May 1898, S.M. Tracy 4598 (HOLO-
TYPE: NY).
Dichanthelium oli tl I ibneri (Nash) Freckmann & Lelong,
comb. & stat. nov, BASIONYM: Panicum scribnerianum Nash, Bull. Torrey Bot.
Club 22:421. 1895. Panicum oligosanthes var. scribnerianum (Nash) Fernald,
Rhodora 36:80. 1934. Dichanthelium oligosanthes var. scribnerianum (Nash) Gould,
Brittonia 26:60. 1974. TYPE: U.S.A. PENNSYLVANIA. Wysox, Jul 1836, J. Carey s.n. (LEC-
TOTYPE: GH [designated by Hitche. & Chase, Contr. U.S. Natl. Herb. 15:283. 1910)).
Panicum scoparium S. Watson in A. Gray, Man., ed. 6,632. 1890., nec Lam. (1798), nec Rudge (1805).
nec Elliott. 1816).
Dichanthelium ovale subsp. praecocius (Hitchc. & Chase) Freckmann & Lelong,
at. NOV. BASIONYM: Panicum praecocius Hitche. & Chase, Rhodora
8:2006. 1906. Panicum lanuginosum var. praecocius (Hitche. @ Chase) Dore in
McNeill & Dore, Naturaliste Canad. 103:562. 1977. Dichanthelium villosissimum
var. praecoc ius (Hitche. & Chase) Freckmann, Phy tologia 39:270.1978. TyPE: U.S.A
ILLINOIS. Stark Co. dry bank, near Wady Petra, 30 Jun 1900, V.H. Chase 649 (HOLO-
TYPE: US 5586861).
Dichanthelium ovale subsp. pseudopubescens (Nash) Freckmann & Lelong,
comb. & stat. nov. BASIONYM: Panicum pseudopubescens Nash, Bull. Torrey
Bot. Club 26:577. 1899. Panicum villosissimum var. pseudopubescens (Nas! n)
Fernald, Rhodora 36:79. 1934. Panicum ovale var. ed caer nace)
Lelong, Brittonia 36:271. 1984. TyPE: U.S.A. ALABAMA: Lee Co.: Auburn, 7
May 1898, FS. Earle and C.F Baker 1537 (1 een NY).
Dichanthelium ovale subsp. villosissimum (Nash) Freckmann & Lelong, comb.
Stat. nov. BASIONYM: Panicum villosissimum Nash, Bull. Torrey Bot. Club 23:149.
1896. Dichanthelium lanuginosum var. villosissimum (Nash) Gould, Brittonia 26:60
1974. Dichanthelium villosissimum (Nash) Freckmann, Phytologia 32:270. 1978.
TyPE: U.S.A. GEORGIA. Ocmulgee River swamp, below Macon, 18-24 May 1895, J.K.
Small s.n. (HOLOTYPE: NYD.
Dichanthelium portoricense subsp. patulum (Scribn. & Merr.) Freckmann &
Lelong, comb. nov. BASIONYM: Panicum nashianum var. patulum Scribn. & Merr,
NOMENCLATURAL 171
US.D.A. Dept. Agrost. Circ. 27:9.1900. Panicum patulum (Scribn. @ Merr.) Hitchc.,
Rhodora 8:209. 1906. Panicum lancearium var. patulum (Scribn. & Merr.) Fernald,
Rhodora 36:80. 1934. Dichanthelium sabulorum var. patulum (Scribn. & Merr.)
Gould & C.A. Clark, Ann Missouri Bot. Gard. 65:1113. 1979 [imprint date 1978]. TYPE:
U.S.A. FLORIDA. Manatee Co.: Braidentown [Bradenton], 3 Sep 1898, R. Combs 1296
(HOLOTYPE: US 28089821).
cameras strigosum subsp. glabrescens (Griseb.) Freckmann & Lelong,
.& stat. nov. BASIONYM: Panicum dichotomum var. lees ns Griseb,, Fl.
Beit Windies 553. 1864. Dichanthelium leucoblepharis var. gla
Gould & C.A. Clark, Ann. Missouri Bot. Gard. 65:1100 (1979) pent date 1978]
Dichanthelium strigosum var. glabrescens (Griseb.) Freckmann, Brittonia 33:457.
1981. Panicum strigosum var. ia see Srise))) Lelong, Brittonia 36:264. 1984.
TypE: JAMAICA: W. Purdie s.n. (HOLOTYPE: kK).
Dichanthelium strigosum subsp. leucoblepharis (Trin.) Freckmann & Lelong,
nov. BASIONYM: Panicum leucoblepharis Trin., Clav. Agrost. 234.
1822. Dichanthelium leucoblepharis (Trin.) Gould @ C.A. Clark, Ann. Missouri Bot.
Gard. 65:1099. 1979 [imprint date 1978]. Dichanthelium strigosum var.
leucoblepharis (Trin.) Freckmann, Brittonia 33:457. 1981. Panicum strigosum var.
leucoblepharis Trin.) Lelong, Brittonia 36:264. 1984. TYPE: U.S.A. NORTH CAROLINA,
SOUTH CAROLINA, GEORGIA, LOUSIANA: Enslin s.n. (LECTOTYPE: LE, fragment US! |des-
ignated by Hitchc. & Chase, Contr. U.S. Natl. Herb. 15:162. 1910)).
Dichanthelium tenue (Muhl.) Freckmann & Lelong, comb. nov. BAsiONyM: Pani-
cum tenue Muhl., Descr. Gram. 118 (1817). Dichanthelium dichotomum var. tenue
(Muhl.) Gould & C.A. Clark, Ann. Missouri Bot. Gard. 65:1119. 1979 [imprint date
1978]. TyPE: U.S.A: Muhlenberg Herb. 192 (LECTOTYPE: PH-M [designated by Hitche.
& Chase, Contr. US. Natl. Herb. 15:259. 1910
Panicum amarum subsp. amarulum (Hitchc. & Chase) Freckmann & Lelong,
comb. & stat. nov. BASIONYM: Panicum amarulum Hitchc. & Chase, Contr. USS.
Natl. Herb. 15:96. 1910. Panicum amarum var. amarulum (Hitchc. & Chase) PG.
Palmer, Brittonia 27:149. 1975. TYPE: U.S.A. VIRGINIA. Virginia Beach, 24 Sep 1900,
TA. Williams 3090 (HOLOTYPE: US 5927 48)).
Panicum anceps subsp. rhizomatum (Hitchc. &@ Chase) Freckmann & Lelong,
comb. & stat. nov. BASIONYM: Panicum rhizomatum Hitchc. & Chase, Contr. US.
Natl. Herb. 15:109, f. 104. 1910. Panicum anceps var. rhizomatum (Hitchc. & Chase)
Fernald, Rhodora 36:73. 1934. Type: U.S.A. SOUTH CAROLINA: Orangeburg, 18 Aug
1905, A.S. Hitchcock 450 (HOLOTYPE: US 592752!).
Panicum capillare subsp. hillmanii (Chase) Freckmann & Lelong, comb. & stat.
nov. BASIONYM: Panicum hillmaniilas “hillmani”| Chase, J. Wash. Acad. Sci. 14:345.
1934. Type: US.A. TEXAS: Amarillo, plain, 11 Aug 1918, A.S. Hitchcock 16206 (HOLO-
TYPE: US 1037542).
Panicum dichotomiflorum subsp. bartowense (Scribn. @ Merr.) Freckmann &
ng, comb. stat. nov. BASIONYM: Panicum bartowense Scribn. & Merr.,
US.D.A. Dept. Agrost. Circ. 35:3. 1901. Panicum dichotomiflorum var. pene cen
(Scribn. & Merr,) Fernald, Rhodora 38:387. 1938. TyPE: U.S.A. FLORIDA. Polk Co.: in
wet reclaimed swamps at Bartow, 29 ee 1898, R. Combs 1220 (HOLOTYPE: SUS 80538)).
Panicum di ifl subsp. p & ) Freckmann & Lelong,
ete & stat. nov, BASIONYM: Panicum dichotomiflorum var. puritanorum
172 BRIT.ORG/SIDA 20(1)
Svenson, Rhodora 22:154, f. 1-5. 1920. Type: U.S.A. MAINE: dry sandy and gravelly
beach, Half-way Pond, Barnstable, 13 Sep 1919, M.L. Fernald 306 (HOLOTYPE: GH).
Panicum hallii subsp. filipes (Scribn.) Freckmann & Lelong, comb. & stat. nov.
ASIONYM: Panicum filipes Scribn. in A. Heller, Contr. Herb. Franklin Marshall Coll.
1:13.1895. Panicum hall ii var. filipes(Scribn.) ER. Waller, South W. Naturalist 19:105.
1074. Type: U.S.A. TEXAS: growing in rich shaded ground in the upper part of the
arroyo at Corpus Christi, alt. 30 feet, 31 May 1894, A.A. Heller 1809 (HOLOTYPE: US
2463130!)
Panicum hirticaule subsp. sonorum (Vasey ) en nn & Lelong, comb. & stat.
nov. BASIONYM: Panicum sonorum Vasey in Beal, Grasses N. Amer. 2:130. 1896. TyPE:
MEXICO, Sonora: Lerdo, at the head of the Galt of c “al fornia, 1889, E. Palmer 947
(HOLOTYPE: US 2903025).
Panicum capillare var. miliaceum Vasey in Vasey & Rose, Contr. US. Natl. Herb. 1890. Pani-
cum hirticaule var. miliaceum (Vasey) Beetle, Phytologia 47:383. 1981
Panicum hirticaule su! trami (Hitche. & Chase) Freckmann S Lelong,
comb, & stat. nov. BASIONYM: Panicum stramineum Hitche. & Chase, Contr. US
Natl. Herb. 15:67, £.50.1910. Panicum capillare var. stramine um (Hitchc. & Chase)
Gould, Madrono 10:94. 1949. Panicum hirticaule var. stramineum (Hitche. &
Chase) Beetle, Phytologia 47:383. 1981 TYPE: MEXICO. SONORA: Guaymas, Jun-Nov
1887, E. Palmer 206 (HOLOTYPE: US 592753}).
Panicum | orgie ere subsp. gattingeri (Nash) Freckmann & Lelong, comb.
at. NOV. BASIONYM: Panicum capillare var. gattingeri Nash in Britt. & Brown,
st FI. 1:123. 1896. Panicum gattingeri Nash in Small, Fl. s.e. U.S. 92. 1903. TYPE:
TENNESSEE: cedar glades near Nashville, Sep, A. Gattinger s.n. (LECTOTYPE:
ea Us! |designated by Hitche. & Chase, Contr. U.S. Natl. Herb.]).
—
Panicum capillare var. campestre Gattinger, Tennessee [1]. 94. 1887,
Panicum philadelphicum — lithophilum (Swallen) Freckmann & Lelong,
comb. & stat. nov. BASIONYM: Panicum lithophilum Swallen. Proc. Biol. Soc. Wash.
54:43. 1941. Type: U.S.A. - ORGIA: on rocky slope. Stone Mountain, 23 Aug 1905,
AS. Hitchcock and A. Chase 185 (HOLOTYPE: US 952898).
a n rigidulum s subsp. abscissum (Swallen) Freckmann & Lelong, comb. &
. nov. BASIONYM: Panicum abscissum Swallen, J. Wash. Acad. Sci. 30:215, £. 4.
1940. Type: U.S.A. FLORIDA: Sebring, in dry sand at tourist camp near lake, 3 Oct
1925, P Weatherwax s.n. (HOLOTYPE: US 1259952),
Panicum rigidulum subsp. combsii (Scribn. & C.R. Ball) Freckmann & Lelong,
mb. & stat. nov. BASIONYM: Panicum combsii Scribn. & C.R. Ball, US.D.A. Div.
Agrost. Bull. 24:42, f. 16.1901 [imprint date 1900]. Panicum longifolium var. combsii
(Scribn. & CR. Ball) Fernald, Rhodora 36:69 (1934). Panicum rigidulum var. combsii
(Scribn. @ CR. Ball Lelong, Brittonia 36:263. 1984. Typr: US.A. FLORIDA. Wash-
ington Co. damp fertile flat . [low pine barrens] at Chipley, 20 Aug 1898, R.
Combs 583 (HOLOTYPE: US 805
Panicum oe subsp. Geeta (Pursh) Freckmann @ Lelong,comb. & stat.
nov. BASION oe um elongatum Pursh, Fl. Amer. Sept. 1:69. 1813 [imprint date
1814], non ee (1790). Panicum agrostoides var, Aa pe abn: Tennes-
see Agric. Exp. Sta. Bull. 7:42. 1894. Panicum rigidulum var. el Pursh) Lelong,
NOMENCLATURAL 173
Brittonia 36:263. 1984. TYPE: U.S.A. DELAWARE: ET. Pursh.n. (LECTOTYPE: K, photo US!
[designated by Hitchc. & Chase, Contr. U.S. Natl. Herb. 15:104. 1910).
Panicum stipitatum Nash in Scribn., U.S.D.A. Div. Agrost. Bull. 17 [rev. ed.}:56, f. 352. 1901.
Panicum rigidulum subsp. pubescens (Vasey) Freckmann & Lelong, comb. &
tat. nov. BASIONYM: Panicum anceps var. pubescens Vasey, U.S.D.A. Div. Bot. Bull.
8:37.1889. Panicum longifolium var. pubescens (Vasey) Fernald, Rhodora 36:69. 1934.
Panicum rigidulum var. is scens (Vasey) Lelong, Brittonia 36:263. 1984. TYPE:
S.A. ALABAMA: Mobile Co.. Mobile, C.T. Mohr s.n. (LECTOTYPE: US 80530! Idesig-
nated by Hitchc. & Chase, Cue U.S. Natl. Herb. 15:105. 1910)).
ACKNOWLEDGMENTS
We are very grateful to Mary E. Barkworth for her continuous support and en-
couragement of our efforts to produce new treatments of Panicum and
Dichanthelium and for her assistance in preparing this manuscript. We are also
very grateful to K.N. Ghandhi for his extensive and meticulous search of the
literature for infrageneric names and for his help in interpreting the applica-
tion of the Code to some difficult nomenclatural problems.
REFERENCES
BentHam, G and J.D. Hooker. 1883. Genera plantarum. Vol. 3, pt. 2, pp. 447-1258. L. Reeve &
Co., London.
Brown, WV. and B.N. Smith. 1975. The genus Dichanthelium (Gramineae). Bull. Torrey Bot.
Club 102:10-13.
Daviose, G. and F.O. Zutoaca. 2000. Proposal to reject the name Panicum divergens
(Gramineae). Taxon 49:103-104.
Feanatp, M.L. 1936. Plants from the outer coastal plain of Virginia. Rhodora 38:376-404.
Fernatp, M.L. 1938. Contributions from the Gray Herbarium of Harvard University — No.
CXXII, VII. New species, varieties and transfers. Rnodora 40:331-358.
Fernato, M.L. 1947. Additions to and subtractions from the flora of Virginia. Rhodora 49:
121-142
Gout, FW. 1974. Nomenclatural changes in Poaceae. Brittonia 26:59-60.
Greuter, W.L.,J. MCNEILL, E.R. Barrie, HM. Burvet, V. Demouuin, T.S. Fitaueiras, D.H. NicoLson, P.C. Sitva,
JE. Skos, P. TreHane, N.J. Turtano, and D.L. Hawkswortn. 2000. International code of botani-
cal nomenclature. Koeltz Scientific Books, KOnigstein, Germany.
Giussant, L.M., J.H. Cota-SANcHez, F.O. ZuLoaca, and E.A. Ketoaa. 2001.A molecular phylogeny
of the grass subfamily Panicoideae (Poaceae) shows multiple origins of C4 photosyn-
thesis. Amer. J. Bot. 88(1 1):1993-201 2.
Goutp, FW. 1974. Nomenclatural changes in the Poaceae. Brittonia 26:59-60.
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Natl. Herb. 17:181-389.
HitcHcock, A.S. 1915. Panicum L. In J.H. Barnhart, ed. N Amer. Fl. 17(3):198-288.
HitcHcock, A.S. 1919. Type-basis code of botanical nomenclature. Science 49:333-336.
BRIT.ORG/SIDA 20(1)
HitcHcock, A.S. 1935. Manual of grasses of the United States. U.S.D.A. Misc. Publ. 200.
HitcHcock, A.S. 1951. Manual of grasses of the United States. Ed. 2, Rev. A. Chase. U.S.D.A.
Misc. Publ. 200.
HitcHcock, A.S.and A.CHase. 1910.The North American species of Panicum. Contr. U.S. Natl.
Herb.15:1-396.
HircHcock, A.S.and A. Cuase. 1915. Tropical North American species of Panicum. Contr. U.S.
Natl. Herb. 17:459-539.
Hsu, C.C. 1965. The classification of Panicum (Gramineae) and its allies, with special refer-
ence to characters of lodicule, style-base and lemma. J. Fac. Sci., Univ. Tokyo, sect. 3
(Bot.), 9(3):43-150.
Lawrence, G.H.M. 1951. Taxonomy of vascular plants. Macmillan, New York.
Linnaeus, C. 1753. Species plantarum, 2 vols., Stockholm. (Facsimile edition: London, 1957-
1959).
Piicer, R. 1931. Bemerkungen zu Panicum und verwandten Gattungen. Notizblatt der
Botanischen Gartens und Museums zu Berlin-Dahlen 11 (104):237-247,
ZULOAGA, F.O. 1987. Systematics of New Word species of Panicum (Poaceae; Paniceae). In:
TR. Soderstrom, K.W. Hilu. C.S. Campbell, and M.E. Barkworth, eds. Grass systematics
and evolution. Smithsonian Institution Press, Washington, D.C. Pp. 287-306
ZULOAGA, F.O., RP. Eiuis, and O. Morrone. 1993. A revision of Panicum subg. Dichanthelium
sect. Dichanthelium (Poaceae: Panicoideae: Paniceae) in Mesoamerica, the West Indies,
and South America. Ann. Missouri Bot. Gard. 80:1 19-190.
A NEW SPECIES OF SENECIO (ASTERACEAE) FROM
DURANGO, MEXICO
B.L. Turner
Plant Resources Center
University of Texas
Austin, TX, 78713, U.S.A.
ABSTRACT
Senecio sandersiana B.L. Turner, sp. nov, is described and illustrated from near El Salto, Durango,
Mexico. It occurs in pine-oak woodlands at relatively high elevations (ca. 2720 m) and is said to be
n locally on shady slopes and along ledges. The species is believed to be closely related to S.
tolucannus DC. but differs in leaf shape and features of the capitulescence.
RESUMEN
Se describe y se ilustra Senecio sandersiana B.L. Turner, sp. nov. de cerca de El Salto, Durango, México.
Aparece en bosques de pino-encino en elevaciones relativamente altas (ca. 2720 m) y se dice que es
A ] ] ] ] a | < ] ] ] bs jon ] 1 A |
Z oO L Cc [
con S. tolucannus DC. pero difiere en la forma de la hoja y caracteristicas de la capitulescencia.
Routine identification of Mexican Asteraceae has revealed the follow novelty:
Senecio sandersiana B.L. Turner, sp. nov. (Fig. 1). TyPE: MEXICO. DURANGO: Mpio. El
Salto, “rim of plateau W of La Ciudad, 3.1 km NW of Hwy 40... off Neveros rd.
upper Arroyo Tascanto watershed,” 2720 m, 19 Apr 1999, A.C. Sanders et al. 22680
(HOLOTYPE: TEX! ISOTYPE: UCR).
Similis S. tolucanno DC. sed laminas foliorum definite cordatis, petiolis angustioribus (2-4 mm vice
3-10 mm), et pedunculis elongatis cum solum 1-2(-3) capitulis.
Perennial subscapose herbs to 60 cm high. Leaves glabrous or nearly so, 15-25cm
ong, 3.0-8.5 cm wide, forming a persistent basal rosette; petioles of well-devel-
oped leaves alate, mostly 10-15 cm long, 2-4 mm wide; blades broadly ovate to
cordate, the margins decidedly serrulate. Heads 1-3 at the apices of elongate nearly
naked scapes (bearing 2-4 much-reduced, bract-like leaves). Involucres calycu-
late, the calyculae grading into the involucral bracts; inner bracts ca. 21 ina single
series, ca.8 mm long, 1.5mm wide, sparsely glandular-pubescent along the mar-
gins, otherwise glabrous. Receptacle plane to concave, epaleate, glabrous, ca. 4
mmaacross. Ray florets 11-13, sterile; corollas yellow, glabrous; tubes ca.6 mm long;
ligules 4-6 nervate, 6-8 mm long, 2-3 mm wide. Disk florets 40-60, fertile; corol-
las yellow, glabrous, ca. 7 mm long, the tube ca. as long as the cylindrical throat,
the lobes 5, deltoid, ca. 0.5 mm long. Style branches linear, truncate, minutely
hispidulous apically. Achenes (immature) ca. 2.5 mm long, glabrous; pappus of
numerous white bristles somewhat longer than the disk corollas.
This taxon is closely related to the widespread species, S. tolucannus of the
SIDA 20(1): 175 - 177. 2002
BRIT.ORG/SIDA 20(1)
176
VACLAVIK
from holotype.
,
Fic. 1. Senecio sandersiana
TURNER, A NEW SPECIES OF SENECIO FROM MEXICO 177
Fic. 2. Distrit * Ahay} €C * 1 {rl Cees ee | \¢ Stay oe | ),and $. 4 - ( H le)
Sec. Lugentes of Senecio. It might also be compared with S. wootonii Greene of
the southwestern U.S.A. and closely adjacent northwestern Mexico. Senecio
sandersiana differs from both in having fewer heads to a capitulescence and
markedly cordate leaf blades. The distribution of these several taxa in Mexico
is shown in Fig. 2.
Etymology.—The species is named for its primary collector, A.C. Sanders,
Curator at UCR, an intrepid field worker. When informed of its possible novel
status, he remarked via email:
“I knew that 22680 was something good when I found it and so I collected
plenty. ’'d never seen anything like it and was just very interested in the plant
as an attractive object. This was the Senecio | was most interested in hearing
about....| have 5 more sheets. It was very common where | found it: | had to walk
through beds of it on the shaded slopes to find other things.”
ACKNOWLEDGMENTS
lam grateful to my wife, Gayle Turner, for the Latin diagnosis. Susan Vaclavik
provided the illustration.
BRIT.ORG/SIDA 20(1)
BOOK REVIEWS
Par HaLtipay. 2001. The Mlustrated Rhododendron. (ISBN 0-88192-510-1, hbk.).
Timber Press, Inc., 133 S.W. Second Avenue, Suite 450, Portland, OR 97204-
3527, US.A.(Orders: www.timberpress.com, 800-327-5680, 503-227-2878,
503-227-3070 fax). $69.95, 274 pp., 120 color illustrations, 4 line drawings,
81/4" x 11 3/4".
The author writes that, “this is primar uly a pees
kK of * prety Cae "It is certainly a great deal more
than that. The author, a staff member at Kew Gardens for more than 40 years, has culled from its archive
an example of almost oo subsection of every section of the genus Rhododendron. In a few cases,
ioned, or line eal were prepared from herbarium material. Hlustra-
| Li The result isa comprehensive study
new plates
specimen and a discussion of the cultivation of rhododendrons.
ages of
with anu lated text for
Of particular interest is ne act that the original illustrations were taken from the ]
Curtis's ee Magazine, which has been in existence for 213 years and has published over 11,000
e magazine, and because of the time spanned,
illustrations. There is a short essay on the history of the
lustrations themselves provide a short history of botanical illustration.
sis a beautiful book which can be enjoyed by the rhododendron enthusiast, the admirer o
—
thei
botanical illustration, and the casual reader—Ruth Ginsburg, Fort Worth, Texas.
Nancy Koper. 2000. With Paintbrush & Shovel. (ISBN 0-8139-1969-X, hbk.). The
University Press of Virginia. Charlottesville and London. (Orders:
www.upress.virginia.edu). Price not given, 280 pp., 8 photographs, | map,
222 plates. 8 1/2" x 10"
In this story of the creation, between 1935 and 1940, of the Lee Park Wildflower and Bird Sanctuary
in Petersburg, Virginia, the author has woven threads of many colors and textures into a fascinating
tapestry. Under the auspices of the Works Progress Administration, thirty-three women, black and
natural heri-
white, unemployed female heads of households, were hired to create a place where the
‘the state would be preserved. This book is both a botanical and sociological study. There are
sketches of the women who were involved, a eee of the method of work and a discussion of
the species and their habitat. There is a longer biography of the illustrator.
ped, bot: s recognized that some pl. ants and habitats were already dis
As the project devel
appearing. A herbarium was eee to preserve a record of the region, and a watercolorist was
commissioned to illustrate each species. Two hundred and twenty of these are reproduced in th
book, arranged by habitat
ane
c rn +1 ee! oe
everal levels and a tribute to the Numan spirit. Ruth
interesting book
}
Ginsburg, Fort Worth, Texas.
SIDA 20(1): 178. 2002
TWO NEW SPECIES OF WEINMANNIA (CUNONIACEAE:
CUNONIEAE) FROM SOUTHERN ECUADOR
Zachary S. Rogers
Missouri Botanical Garden
St. Louis, MO 63166-0299, U.S.A.
Zachary.rogers@mobot.org
ABSTRACT
d ic f
Two new species of Cunoniaceae, Weinmannia bradfordiana Z. Rogers and Wei
Rogers are described and illustrated. Both sclerophyllous species are known only from their type
localities and were collected from the Cordillera del Condor mountain range in the province of Morona
Santiago, Ecuador.
Key Worps: Cunoniaceae, Cunonieae, Ecuador, Neotropics, South America, Weinmannia
RESUMEN
Se describe y se ilustra Weinmannia bradfordiana Z. Rogers y Weinmannia condorensis Z. Rogers. Estas
| . ] 3] ~ J ae qi ] Se ee | 4s] . ic Ret erence Paes
C Fe y
en la region de la Cordillera del Condor en la provincia de Morona Santiago, Ecuador.
INTRODUCTION
The family Cunoniaceae is composed of 26 genera and about 300 species
(Bradford & Barnes 2001). The australly-distributed genus Weinmannia L. ac-
counts for about 150 species in five sections (Bradford 1998: Bradford 2002). All
American species are in section Weinmannia, with the greatest species rich-
ness found in northwestern South America (i.e. Venezuela, Colombia, Ecuador
and Peru) (Bernardi 1961; Harling 1999).
Weinmannia bradfordiana and Weinmannia condorensis were collected
by the late Alwyn H. Gentry (1945-1993) on his last major expedition, and were
filed in a “family indeterminate” folder for a number of years, despite the good
quality of the specimens in flower and fruit. Later, while sorting Gentry’s col-
lections, Ron Liesner (MO) came across the specimens but did not initial] y con-
sider the specimens as belonging to the family Cunoniaceae because of their
uncharacteristically short inflorescences, but he did notice a few small fruits
that looked remarkably like the septicidal capsules of Weinmannia. Ron then
showed the material to Jason Bradford, who was studying Neotropical
Weinmannia at the time, and Jason confirmed that they were, in fact,
Cunoniaceae and annotated the specimens as new species of Weinmannia (Ron
Liesner and Jason Bradford, pers. comm.).
Both species, as well as another recently described species of Weinmannia
SIDA 20(1): 179 — 187. 2002
180 BRIT.ORG/SIDA 20(1
from the Condor (Rogers 2002), have adapted extremely sclerophyllous habits
due to the very wet and windy weather conditions and the very thin, nutrient-
poor sandstone substrate. Many other woody genera on the summits of the
Cordillera del Condor have similar sclerophyllous adaptations, and many of
these species will also turn out to be new and locally endemic to the range.
Weinmannia bradfordiana and Weinmannia condorensis are known only
from their types and are probably closely related, due to their similar leaf mor-
phology and ecology and because they were collected from sites about 10 km
apart, but significant differences exist between the two including: leaf size, leaf-
let shape, number of flowers per inflorescence, and trichome density and dis-
tribution on the stems, leaves, stipules and sepals. Based upon experience with
other recognized species of Weinmannia and character variation within and
among species, the differences observed from these specimens suggest they rep-
ysis is based entirely on a subjective study of
—
resent distinct species. This ana
the type collections and available herbarium specimens, so more collections
will be needed to elucidate the patterns of variation in these species. At this
time, the type localities are unable to be revisited because the sites have been
covered by deadly land mines since the border dispute between Ecuador and
Peru in January 1995, which was about 18 months after the types were origi-
nally collected. The mine field will not be cleared for at least several more years,
but when the situation is resolved, further exploration will be needed and may
expose intermediates that support a notion expressed by one reviewer (David
Neill), who felt that both collections could represent a single species.
These new species can be easily distinguished from other Ecuadorian spe-
cies by their small trifoliolate (rarely unifoliolate) compound leaves on short
petioles that can appear to be simple and sessile to the naked eye, and by their
extremely short inflorescences (ie. pseudoracemes) due to inconspicuous pe-
duncles. The highly reduced inflorescences are very uncharacteristic of the
genus because almost all Weinmannia have elongated inflorescence axes mea-
suring several cm or more in length
Color images of the type collections can be found on the W3 TROPICOS
database at: <http://mobot.mobot.org/W3T/Search/vast.html>
=
Weinmannia bradfordiana Z. Rogers, sp. nov. (Fig. 1). Type: ECUADOR. MoRONA
SANTIAGO: Canton Gualaquiza: Campamento Achupalla, Cordillera del Condor,
15 km E of Gualaquiza, tepui-like bromeliad sward with scattered, small trees, 03°
27'S, 78° 22'W, 2090 m, 21 Jul 1993 (CL. fr), Gentry 80312 (HOLOTYPE: QCNE-092179;
ISOTYPES: AAU, GB-186976, MO-5544000, NY, US).
1ensem Hieron. et W. mariquitae Szyszyl. maxime accedit, sed a
aec species ad Weinmanniam coc
hac inflorescentia subsessili, ab illa foliis multo mUEOE PHS | ut eet sim] abet ees ouBGHS
} aT 1
propter petiolum perbrevem atque foliolis integris,
unifoliolatis sane tur, a W. fecal inflorescentia ex seauidoreente ‘ asciculis 2- ad 8-floris
versus trichomatibus plus quam 20 ciliatis
5 t
dist co
181
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ROGERS,
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inflorescences
Fic. 1. W
182 BRIT.ORG/SIDA 20(1)
Shrubs to 1m; young stems covered with dense lanate or tomentose trichomes,
the trichomes to 1.3 mm long, simple, unicellular, matted, stiff, white, nodes
densely hirsute. Leaves to 5.3 mm long, opposite, compound, trifoliolate (rarely
unifoliolate), densely congested; leaflets 2.0-4.9 x 1.4-3.3 mm, broadly obovate
to nearly orbicular, broadly boat-shaped to nearly planar, upturned at apex, all
leaflets similar in size and shape, coriaceous, thick, membranous and thin along
the margins, shiny, drying dark brown on the upper surface and light brown
on lower surface, midvein raised on upper and lower surface, thicker near the
base, secondary veins 3-5 pairs per side, slightly raised to nearly inconspicuous
on the upper surface, prominently raised on the lower surface, blade surfaces
glabrous, usually lanate or tomentose along the upper one-third of the apical
margins (usually more than 20 trichomes per apex), the trichomes are similar
to those found on the young stems, 0.4-1.0 mm long, upper surface scrobicu-
late, lower surface scrobiculate and rugose, base cuneate to slightly oblique,
margin entire, apex obtuse to slightly apiculate; petioles 0.1-0.4 mm long, re-
duced to inconspicuous or indiscernible protuberances, obscured by the dense
pubescence at the nodes, dark red or black, densely hirsute or tomentose; peti-
olules 0.1-0.4 mm long, swollen, dark red or black, glabrous, rugulose; stipules
2.2-3.6 mm x 1.8-2.7 mm, interpetiolar, paired, free, broadly ovate to suborbicu-
lar, membranous to subcoriaceous, dark red or brown, outer surface densely
lanate or tomentose, becoming more dense towards the apex and along the
margin, trichomes to 1.1 mm long, similar to those found on the young stems,
inner surface glabrous, base truncate, margin entire, apex rounded to obtuse,
caducous. Pseudoracemes paired, unbranched, densely compacted, axillary on
young growth at the terminus of the stem; peduncles to 0.9 mm long, reduced
to inconspicuous protuberances, densely lanate or hirsute; fascicles (2-)4-8-
flowered, densely compacted and congested; pedicels to 0.9 mm long in flower,
to L9 mm long in fruit, sparsely lanate or hirsute; bracteoles not seen. Flowers,
actinomorphic, bisexual, diplostemonous, recorded as white in color on the label:
sepals 4, 1.0-1.2 x 0.5-0.9 mm in flower, to 1.4 mm long in fruit, ovate, subcoria-
ceous, adaxial surface glabrous, abaxial surface densely lanate to sparsely to-
mentose, denser near the apex and along the margin; petals 4, 1.0-1.2 x 0.8-L0
mm, broadly elliptical to broadly obovate, membranous, white, midvein con-
spicuous, slightly raised and dark, glabrous, base truncate, margin entire and
ciliated along the upper portion, apex rounded; stamens 8; filaments 0.8-2.0
mm long, 0.4-0.6 mm wide at base, flattened, slender at apex, glabrous;
anthers 0.3-0.4 « 0.4-0.5 mm, orbicular, introrse, dorsifixed, longitudinally
dehiscent; nectary disk annular with 8 concrescent lobes, surrounding the ovary,
persistent in fruit; ovary 0.7-0.9 x 0.3-0.5 mm, superior, bicarpellate, syncar-
pous, red or brown, glabrous; styles 2, 0.2-0.4 mm long in flower, 0.6-0.9 mm
long in fruit, divergent near the apex of the carpels, persistent in fruit; stigma
simple, capitate. Fruits 1.9-2.4 x 14-L8 mm (length measurement not includ-
—_
ROGERS 183
ing the persistent styles), septicidal capsules, orbicular, occasionally widely
ovate, dark red or brown, costate longitudinally, glabrous; seeds not seen.
Distribution and Habitat—Weinmannia bradfordiana is known only from
the type locality, collected near one of the summits of the Cordillera del Condor
at about 2100 m elevation, and located about 15 km SE of the town of Gualaquiza.
The Condor mountain range is covered by “dwarf-forest” vegetation consisting
of many species of shrubs and small trees that have evidently adapted
sclerophyllous habits due to thin, nutrient-poor sandstone substrate, and ex-
tremely wet and windy weather on the ridge.
Etymology.—The epithet was chosen in honor of Dr. Jason C. Bradford, who
has spent a number of years in the field collecting Cunoniaceae, and who has
made valuable contributions to our knowledge of the family through his con-
tinuing research efforts.
bia cape condorensis Z. Rogers, sp. nov. (Fig. 2). TYPE: ECUADOR. MoroNna
NTIAGO: Canton Gualaquiza: Crest of Cordillera del Condor, ridge top 15 km ENE
of Gualaquiza, high montane forest and bromeliad sward. 03°22’S, 78'20’W, 2500
m, 26 Jul 1993 (£1, fr), Gentry 80465 (HOLOTYPE: QCNE-092303; ISoTYPES: AAU,
MO-5613306, NY).
Haec species ad Weinmanniam cochensem Hieron. et W. mariquitae Szyszyl. maxime accedit, sed a
hac inflorescentia paaehula ab ue Eells multo minor aus 1 ut acter SEpHeI Us. SeoeinuSave
propter petiolum
enltonolalis aisanguitur, aW. bradjordiand inflonescentia ex eseudowcems fasciculis l- ad 2-floris
esi
atque f g p versus glabris vel trichomatibus 5 vel minus ciliatis
distinguitur.
Trees to 3m; young stems covered with dense lanate or hirsute trichomes, the
trichomes to 1.1mm long, simple, unicellular, matted, stiff, white, nodes densely
hirsute. Leaves to 7.6 mm long, opposite, compound, trifoliolate (rarely unifoli-
olate), congested; leaflets 3.6-6.8 x 1.9-4.5 mm, obovate to elliptical, broadly
boat-shaped to nearly planar, upturned at apex, all leaflets similar in size and
shape, coriaceous, thick, membranous and thin along the margins, drying dark
brown, shiny, midvein prominently raised on upper and lower surface, thicker
near the base, darker than blade, secondary veins 2-5 pairs per side, raised to
nearly inconspicuous on the upper surface, more prominently raised on the
lower surface, blade surfaces glabrous, infrequently having a few sparse tri-
chomes at the apical margins (usually 5 or fewer trichomes per apex when pu-
bescent), the trichomes are similar to those found on the young stems, 0.1-0.4
mm long, upper and lower surface scrobiculate and rugose, base cuneate to
slightly oblique, margin entire, apex obtuse to slightly apiculate; petioles 0.7-
1.2 mm long, often inconspicuous, obscured by the dense pubescence at the
nodes, dark red or black, sparsely hirsute or tomentose; petiolules 0.3-0.6 mm
long, swollen, dark red or black, glabrous, rugulose; stipules 2.2-3.5 mm x 2.0-
3.0 mm, interpetiolar, paired, free, ovate to suborbicular, membranous to sub-
BRIT.ORG/SIDA 20(1)
184
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ATL ‘OL . J L «lL L t L W = | Fat A c\
the inflorescence author
J oP \ 7
coriaceous, dark red or brown, outer surface glabrous to sparsely hirsute densely
hirsute along the margin, denser near the apex, trichomes 0.6-1.0 mm long, simi-
lar to those found on young stems, inner surface glabrous, base truncate, margin
entire, apex rounded to obtuse, caducous. Pseudoracemes paired, unbranched,
densely compacted, axillary on young growth, developing at 2 or more subse-
quent nodes at the terminus of the stem; peduncles to 0.4 mm long, reduced to
inconspicuous protuberances in the leaf axils, lanate or hirsute, the trichomes
are similar to those found on the young stems; fascicles l- or 2-flowered, com-
pacted and congested; pedicels to 1.2 mm in flower, sparsely hirsute; bracteoles
not seen. Flowers actinomorphic, bisexual, diplostemonous, recorded as white
ROGERS 185
in color on the label; sepals 4, 1.6-2.4 x 0.8-2.0 mm, broadly ovate, subcoria-
ceous, adaxial surface glabrous, abaxial surface lanate, denser near the apex and
along the midvein; petals 4, 1.6-2.0 x 1.2-2.2 mm, broadly elliptical to broadly
obovate, membranous, white, midvein conspicuous, slightly raised and dark,
glabrous, base truncate, margin entire and ciliated, apex rounded; stamens 8;
filaments 0.8-2.5 mm long, 0.3-0.5 mm wide at base, flattened, slender at apex,
glabrous; anthers 0.3-0.4 x 0.2-0.4 mm, orbicular, introrse, dorsifixed, longitu-
dinally dehiscent; nectary disks annular with 8 concrescent lobes, surround-
ing the ovary, persistent in fruit; ovary to 0.9 mm long, to 0.6 mm wide, supe-
rior, bicarpellate, syncarpous, red or brown, glabrous; styles 2, to 2.1 mm long,
divergent near the apex of the carpels, persistent in fruit; stigma simple, capi-
tate. Fruits (immature) 1.8-2.5 mm long (measurement not including the per-
sistent styles), septicidal capsules, ovate, dark red or brown, glabrous; seeds not
seen.
Distribution and Habitat.—Weinmannia condorensis is known only from
the type locality, collected on a ridge-top of the Cordillera del Condor moun-
tain range at about 2500 m elevation, and located about 15 km ENE of the town
of Gualaquiza. The site, as in the case of W. bradfordiana, is covered by “dwarf-
forest” vegetation consisting of many species of shrubs and small trees with
similar sclerophyllous adaptations.
Etymology.—The specific epithet refers to locality of the type collection.
Affinities.—According to the key for the Cunoniaceae in the Flora of Ecua-
dor (Harling 1999), W. bradfordiana and W. condorensis would fit between the
couplet W. cochensis Hieron.and W. mariquitae Szyszyl. because they have small
compound leaves measuring less than 5cm in length, but W. mariquitae is very
different morphologically because it has much larger leaves with 4-8 leaflet
pairs per leaf. On the other hand, W. bradfordiana and W. condorensis can be
easily distinguished from the most morphologically similar species from Ec-
uador, W. cochensis, because they have smaller, trifoliolate (rarely unifoliolate)
leaves with smaller, entire-margined leaflets, and inconspicuous peduncles. No
other currently described species of Weinmannia in Ecuador has such highly
reduced compound leaves and peduncles.
Weinmannia bradfordiana can be vegetatively distinguished from W.
condorensis by its smaller leaves, its smaller, more broadly obovate to subor-
bicular leaflets, by the presence of 20 or more trichomes found along the apical
margins of most leaflets, and by its densely pubescent stipules. The inflores-
cences of W. bradfordiana are more congested by more numerous flowers, and
have slightly longer, more globular peduncles than W. condorensis. In contrast,
most leaflets of W. condorensis are typically larger, narrowly obovate to ellipti-
cal and glabrous. Weinmannia condorensis also has inflorescences that are re-
duced to only | or 2 flowers, and stipules that are mostly glabrous except for a
dense apical pubescence.
186 BRIT.ORG/SIDA 20(1)
Another novel and distinctive character separating the two species is that
W. condorensis has a rare inflorescence architecture, where pairs of racemes
develop at two successive nodes near the terminal end of the main stem. The
inflorescence of section Weinmannia (ic. all Neotropical species) is limited to
a pair of racemes developing from axillary buds at the most distal node, and is
a distinctive character of the section. The development of racemes at two sub-
sequent nodes has not been found in other Neotropical members of the genus,
but it has been observed in a small group of species from Madagascar and the
Comores (Bradford, pers. comm.). Patterns in inflorescence architecture are rela-
tively stable in Weinmannia, and in many cases, can be more helpful than plas-
tic vegetative characters, which often prove to be similar among many closely
related species (Bradford 1998). The interesting inflorescence architecture of
W. condorensis will definitely warrant further investigation once more collec-
tions are made available.
KEY TO MORPHOLOGICALLY SIMILAR SPECIES OF WEINMANNIA IN ECUADOR
. Leaves 10-20 mm long, yilela crenate at apex, usually 3-5 teeth, rarely 0; petioles
2-4 mm long; peduncles to 15 mm long Weinmannia cochensis
. Leaves to 7.6 mm long, margins entire; see inconspicuous to 1.2 mm long;
anon elie to 0.9 mm lone
na
rgins red by a dense apical pubescence
“(20 or more Hichomes: ae denen pubescent throughout; inflorescences
(2-)4-8-flowered, only developing at the most distal node of the terminal end
of the main stems Weinmannia ene
Leaflets narrowly obovate to elliptical, margins usually glabrous, or rarely
about 5 or fewer trichomes at the apices; stipules pubescent, nee con-
centrated along the apices and along the midvein; inflorescences 1- or 2-flow-
ered, developing at two or more successive nodes at the terminal end of the
main stems Weinmannia condorensis
N
ACKNOWLEDGMENTS
The author thanks Jason Bradford (MO-DAV), Richard Barnes (HO) and David
Neill (MO-QCNE) for their helpful review suggestions on the manuscript. The
author would also like to thank Ron Liesner (MO) and Jon Ricketson (MO) for
their comments, Roy Gereau (MO) for his preparation of the Latin diagnoses,
Barney Lipscomb (BRIT) for waving the publication costs of this manuscript,
and Uno Eliasson (GB) for expeditiously providing loan material.
REFERENCES
Bernarol, L. 1961. Revisio generis Weinmanniae. Pars |: Sectio Weinmanniae. Candollea
17:123-
Braprorb, J.C. 1998. A cladistic analysis of species groups in Weinmannia (Cunoniaceae)
based on morphology and inflorescence architecture. Ann. Missouri Bot. Gard. 85:
565-593
ROGERS 187
Braprorb, J.C. (in press, 2002). Molecular phylogenetics and morphological evolution in
Cunonieae (Cunoniaceae). Ann. Missouri Bot. Gard.
Braororo, J. C. and R.W. Barnes. 2001. Phylogenetics and classification of Cunoniaceae
(Oxalidales) using chloroplast DNA sequences and morphology. Syst. Bot. 26:354-385.
Hartine, G. 1999. Cunoniaceae. In: G. Harling and L. Andersson, eds. Fl. Ecuador 61:1-74.
Rocers, Z.S. 2002. A new species of Weinmannia (Cunoniaceae: Cunonieae) from southern
Ecuador. Novon 12:249-252.
BRIT.ORG/SIDA 20(1)
BOOK REVIEWS
HOWARD GARRETT With ODENA BRANNAM. 200L. Herbs for Texas. (ISBN 0-292-78713-
3, hbk.). University of Texas Press, PO. Box 7819, Austin, TX 78713-7819, US.A.
(Orders: 800-252-3206, 800-687-6046 fax). $60.00, 280pp. 8 photographs,
1 map, 222 plates. 81/2" x 11’.
In this fully illustrated, easy to use guide, expert advice is offered by Garrett and veteran herbalist
Odena Brannam on Bow ing nearly 150 herbs suited to Texas and Southwestern gardens. Included is
detailed infor h plant's landscape, c ana pipaiemias and ot ne: uses.
hl } { c
nla qT} ting,
Individual entries give eac
growing, harvesting, and storing it. The entries also include ideas for using each herb in gardening
and cooking, with occasional recipes, and discuss its medicinal uses. Unique is the special “insight”
entries that offers intriguing and little-known facts about the herbs. Each herb has a color photo.—
Gary Jennings, Botanical Research Institute of Texas, Fort Worth, TX 76102-4060, U.S.A.
NANcy Leys STEPAN. 200L. Picturing Tropical Nature. (ISBN 0-8014-3881-0, hbk.).
Cornell Univ. Press, 512 East State St., Ithaca, NY 14850, U.S.A. (Orders:
607-277-2211, 607-277-6292 fax). $35.00, 243 pp., numerous b&w photos,
6 x 91/4".
“Tropical nature as jungle is an artifact of history and cultur
Nancy Leys Stepan proves in a well-documentec caaea ol [how the tropics have been depicted his-
| . The focus is on Brazil (the name
This is essentially what historian
torically i in descriptive wi Itings, art literature, anc
of which comes from the name of a plant), beginning with the accounts, verbal and pictorial, of
Humboldt, Bates, Darwin, and Wallace, who described the “radical otherness” of the jungles. The
e jungle, full of lush growth, of an infinite variety of plants and animals not yet described,
of an ers for the explorer and scientist. The scenes were romantic if in reality arguments for
colonization. With the arrival of European colonists, the infiltration of these lands by armies and
their teams of medical doctors and closer association with the indigenous peoples, the perception
was soon to change, especially as tropical diseases such as malaria and yellow fever took their toll.
The jungle became “dark: disease and death lurked everywhe
kind of racial science developed along with See science. The native peoples were
es Leal al dae ae in part by such investigators as Agassiz whose intent was to refute ne
volution, and in part to show sexuality. Meo ice _ science was to prove that strange ony Sole
fo)
ries
gies were abnor ee by dise
aided in isolating the pathogens of nee and identifying their vectors and eselted a inan entirely
different form of visual iconography. The story of Carlos Chagas, his resolving the mystery of the
disease that bears his name, and his later failures is an engrossing one
The work of Roberto Burle Marx, Brazilian botanist and landscape oe is of ae inter-
est, as he embellished the cities of Sao Paulo and Rio with parks, gardet pen spaces, bringing
in native plants from the rain forest. (Heliconia burle-marxii was one a his discoveries). His con-
l lly sound, bane ser aoe and influential in concepts of landscaping.
“tn this work Stepan olfers much to the reader in social, cultural, and medical history. She also
on
peruses the s cont oncepts and ee phies of “nature” and “natural history.” Not
least are her notes nes a extensive references, which are in themselves rewarding. ‘Joann
Karges, retired librarian, Texas Christian seen Port Worth, TX.
SIDA 20(1): 188. 2002
THE TAXONOMY AND MORPHOLOGY OF
MACRANTHERA FLAMMEA (OROBANCHACEAE)
Jill D. Alford and Loran C. Anderson
Department of Biological Science
Florida State University,
Tallahassee, FL 32306-4370, U.S.A.
anderson@bio.fsu.edu.
ABSTRACT
Macranthera flammea (Bartr) Pennell is a rare, biennial speci lemi he pine f] ds of the
lower coastal pm of ne Soule astern aa nited States. Field, experimental algarden and herbarium
studies were | The taxonomic sua oe the Species is reviewed.
Recognition of swore th lan one ee in the genus is not warranted | lyx length
and lobing are due to the placement of flower units within the architect f the inflores
individual plants. A map of the species’ range is provided.
RESUMEN
A Pa fa R ) 1 : L; | Viet 5 4 ‘ jada leaniura
| I
costera inferior del sudeste de los Estados Unidos. Se hicieron estudios de campo, jardin experimen-
tal, y herbario para poner al dia la descripcion de la especie. Se revisa la historia taxonomica de la
especie. No se necesita reconocer mas de un taxon en el género ya que las diferencias en la longitud
del caliz y lobulacién se deben a la colocacién de las unidades florales en la arquitectura de la
inflorescencia en los distintos individuos. Se aporta un mapa del area de las especies.
—_—
INTRODUCTION
Macranthera is monotypic; the single species M. flammea is endemic to the
southeastern coastal plain of the United States and is commonly referred to as
“flame flower” or “hummingbird flower.” It is listed as endangered by the state
of Florida (Coile 2000). The genus was formerly placed in the Scrophulariaceae,
but recent studies have shown that family to bea heterogenous group (Olmstead
et al. 2001); the hemiparasitic taxa are better placed in the Orobanchaceae.
Here, an updated species description and range map are provided. More
detail on the life history and pollination biology is documented by Alford (2000).
NOMENCLATURAL HISTORY
The many names that have been proposed for this species indicate the diffi-
culty that early botanists had in understanding the taxon and its generic place-
ment (Table 1). William Bartram (1791) first described M. flammea as Gerardia
flammea during his travels through the southeastern states during 1773-1777.
1834, after examining an unlabeled specimen in the herbarium of the
Academy of Natural Sciences, Philadelphia (PH), Nuttall published the genus
SIDA 20(1): 189 — 204. 2002
=
190 BRIT.ORG/SIDA 20(1)
Taste 1. Synonyms for Macranthera flammea.
Synonym Year
Ge rardia flammea W. a 1791
Conradia fuchsioides 1834
Meera ene cure Benth. 1836
icranthera lecontei Torr. 1837
Russelia flammea (Bartr.) Raf. 1837
Flamaria coccinea Ra 1837
Toxopus gymnanthes Raf. 1837
Toxopus calycinus 1837
Tomilix bracteata Raf. 1837
é aSYSTOMA tubulosa Bertol. 1853
Macranthera fuchsioides var. lecontei (Torr.) Chapman 1860
Conradia lecontei (Torr.) Kuntze 1891
Macranthera flammea (Bartr.) Pennell 1913
Conradia as a dedication to the late Solomon W. Conrad, a Professor of Botany
at the University of Pennsylvania. Nuttall proposed the name Conradia
fuchsioides, because “the corolla and long exerted stamens put one in mind of
some gigantic Fuchsia” (Nuttall 1834).
Bentham (1835) applied the name Macranthera fuchsioides (Nutt.) Benth
because Conradia had been previously applied to a genus of Gesneriaceae by
Martius. The name Macranthera means “long anther.” Bentham and LeConte
were both originally cited as authors for the genus (Hooker 1835), although
Bentham (1835) felt that Torrey alone should receive credit for the generic name.
Torrey (1837) described a second species, M. lecontei (sometimes spelled
lecontii), from a specimen collected in Georgia in 1831 by LeConte. He believed
it was distinct because the segments of the calyx were entire, linear-lanceolate,
and approximately one-third the length of the corolla. In M. fuchsioides the seg-
ments of the calyx are dentately lobed and often as long as the corolla tube [on
|.eConte’s specimen (NY), Torrey applied the name Conradia lecontei Torr, along
with a note stating the segments of the calyx are entire]. The specimen itself is
damaged, but an attached illustration depicts the entire sepals. Torrey actually
published the species (1837) not as C. lecontei, but as M. lecontei, since Bentham
(1835) had determined that Conradia was already in use.
Pennell (1935) questioned the authenticity of the collection site of M.
lecontei, which was purportedly along the Altamaha River in Liberty County,
Georgia, because that site was far removed from other known occurrences of
the species. However, the species has been observed along the Altamaha River
as recently as 1997 (M. Hopkins, pers. comm.).
Rafinesque (1837) proposed many new names for Macranthera (see Table 1).
—_—
Macranthera flammea Raf. was proposed on the basis of Bartram’s original
description.
However, Rafinesque thought the genus name Macranthera was inappro-
priate because he did not believe the anthers to be of considerable length, and
he suggested the name Toxopus, meaning “incurved peduncles.” He substituted
Toxopus gymnanthes Raf. for M. lecontei Torr.and Toxopus calycinus Raf. for M.
fuchsioides (Nutt.) Benth. He further proposed the genus Iomilix meaning “cut
calyx” for M. fuchsioides, suggesting that this species be treated as either a sub-
genus or new genus because of the condition of the calyx and the presence of
what he felt to be a bifid style. He proposed the binomial Tomilix bracteata
Raf. as an additional name for M. fuchsioides (Nutt.) Benth. The new genus
Flamaria, as in Flamaria coccinea Raf., was also proposed
Bertoloni (1853) described a new genus and species, Dasystoma tubulosa,
after examination of collections in GH, NY,and PH. Chapman (1860) treated M.
leconteiasa variety of M. fuchsioides, and Kuntze (1891) resurrected the genus
Conradia on the basis of the type specimen that Torrey had originally identi-
fied as C. lecontei.
Pennell (1913) combined the two species of Macranthera into one, writing:
“this difference [in the calyx]is due wholly toage of the flower, as may be readily
seen in the field; young flowers are as in lecontei, while with age the sepals con-
tinue growing and become lobed while the corolla shrivels and shortens in situ,
so bringing about the fuchsioides state.” Pennell suggested that, because Bartram
originally described the species, the accepted name must become Macranthera
flammea (Bartr.) Pennell. He argued that Bartram named his species for its
“flame-coloured” flowers which is a fitting description. Gerardia was rejected
as a generic name because it isa synonym for Agalinis.
Later, many were concerned about the validity of Bartram’s binomials
(Fernald 1944; Rickett 1944; Merrill 1945). Rickett 1944) argued that Bartram’s
binomials are inappropriate because they appear in a publication that is not
always consistent with the Linnean system of binomial nomenclature. Merrill
(1945) defended Bartram’s binomials, writing: “the proportion of binomials to
polynomials, 358 to 2, is too great in favor of the former to justify the elimina-
tion of Bartram’s work.” Wilbur (1971) argued that, if a Bartram binomial is ac-
companied by an adequate description, the publication of the binomial should
be considered valid.
Additional controversy surrounds Bartram’s description of his collection
site. He probably confused the locality of his specimen of Macranthera with
that of the mint Calamintha coccinea (Nutt.) Benth. While traveling along the
Tensaw River near “Taensa” (near or in Baldwin Co, Alabama), Bartram (1791)
recounts the following observations: “These stony gravelly heights produce a
variety of herbaceous plants, but one in particular I shall mention on account
192 BRIT.ORG/SIDA 20(1)
of its singular beauty. I believe it isa species of Gerardia (Gerardia flammea), it
grows erect, a single stem from a root, three or four feet in height, branching
very regularly from about one half its length upwards, forming a cone or pyra-
mid, profusely garnished with large tubular labiated scarlet or flame coloured
flowers, which give the plant a very splendid appearance even at a great dis-
tance.” This description closely describes, for the most part, the form and ap-
pearance of M.flammea, but the locale and scarlet flower color are more appro-
priate for the mint Calamintha coccinea (Merrill 1945). Unlabeled specimens
of both plants were found in Bartam’s collection at the Natural History Mu-
seum in London (Pennell 1935). Bartram’s description is applicable in part to
soth plants (Table 2), but Pennell (1935) believed Bartram’s description to be
most applicable to M. flammea.
—
—
SPECIES DESCRIPTION
Macranthera flammea is generally an herbaceous biennial of the Orobanchaceae
Rosette leaves on juvenile plants are oblanceolate to spatulate. Leaves are gla-
brous with entire or pinnately lobed margins, depending on developmental
stage. Rosettes are variable in leaf number and leaf size, having 3-46 leaves per
rosette with lengths of 3-33 cm and widths of 0.8-3.6 cm (Fig. 1).
Plants may be more appropriately classified as monocarpic perennials be-
cause they may remain in the rosette phase for more than one year. Rosettes
that have attained a critical size are triggered to reproductive states by an in-
crease in day length. Before a rosette bolts into a flowering adult, it begins to
produce pinnately lobed leaves.
Reproductive plants of M. flammed are mostly 15-30 dm tall, with stems
that are obtusely four-angled and essentially nonbranching below the
candelabralike spreading inflorescence. Cauline leaves are opposite and sessile
or have short, winged petioles. Lower leaves are 8.5-16 cm long and 2-6 cm wide,
lanceolate to ovate, and pinnately lobed. Leaves are progressively reduced up-
ward along the stem; the uppermost are 4-7 cm long and |-2.5em wide, slightly
lobed, and toothed or entire. Blades are glabrous, or with minutely ciliate mar-
gins. Rosettee and lower leaves are usually shed by the time of anthesis.
The paniculate inflorescence contains long, indeterminate, racemose
branches (Fig. 2). Flowers are axillary to a reduced, usually entire, foliar bract
(see illustration in Godfrey and Wooten 1981). Pedicels are 1-2 cm long, becom-
ing reflexed in fruit. The calyx is fused with five short lobes that are equal and
linear and 7.5-15 mm long. The corolla is br a orange, Fleshy, 2-2.5 cm long,
short pubescent without, and glabrous or slightly pubescent at the base within.
A whitish nectar chamber at the corolla base is 4 mm long.
The corolla turns brown to black with age. The tube is cylindrical and much
longer (20-23 mm) than the two-lipped lobes (3-4.5 mm), with an adaxial lip
Taste 2. Similarities of Bartram’s description of Macranthera and Calamintha.
Macranthera Calamintha
Habitat wet boggy thickets stony gravelly heights
Plant branching EUnet in appa half intricately, from base
Plant shape tall pyramidal con low spreading
Plant height 3-8 feet 3-4 feet
Corolla shape tubular labiated tubular labiated
Corolla color flame colored scarlet
that is two-lobed and erect and a three-lobed, spreading abaxial lip. The four
contiguous stamens are equal in length and exerted. The filaments are orange
and become about twice as long as the corolla tube, up to 46 mm. The ovary is
ovoid and hasa slender style that is to 46 mm and a linear-clavate stigma. Ma-
ture bicarpellate capsules are 9.5-13 mm long, 7.5-10.5 mm wide, loculicidal,
and slightly pubescent. The seeds are brownish black, 2.5-3 mm long, half as
wide, irregular in shape (angular-lunate to triangular) with two or three fluted
membranous wings. Specimens of M. flammea become black upon drying (due
to presence of orobanchin or iridoid compounds as in related parasitic taxa).
The species description is verified by examination of collections (Appen-
dix 1) from major and regional herbaria: Angus K. Gholson (personal collection
designated AKG; it will eventually become part of FLAS), University of Florida
(FLAS), Florida State See (FSU), Gray Herbarium (GH), University of
Southern Mississippi (no official abbreviation, designated HATT here for con-
venience), Jacksonville State University JSU), Louisiana State University (LSU),
Missouri Botanical Garden (MO), New York Botanical Garden (NY), Old Do-
minion University (OD), Academy of Natural Sciences of Philadelphia (PH),
Smithsonian Institution (US), University of Georgia (GA), University of Missis-
sippi (MISS), University of South Alabama (USAM), Valdosta State University
(VSC), and Vanderbilt University (VDB).
The species description is also supplemented by measurements taken from
several plants at two field sites: Thomas Co., Georgia, at Greenwood or GWD
(Alford 1077 at FSU) and Liberty Co., Florida, in the Apalachicola National For-
est or ANF (Alford 1064 at FSU).
Range and Habitat—The range of M. flammea (Fig. 3) extends along the
lower coastal plain from Georgia to southeast Louisiana. This species requires
mesic habitats of bogs and thickets and is ecotonal, occurring in the sandy acid
soils of pine flatwoods along the margins of seepage slopes and shrub-tree bogs
or bays (Small 1933). It can occasionally be found in shallow water of cypress-
gum ponds or depressions (Godfrey & Wooten 1981). It does not tolerate stand-
ing water and can only be found in areas where water flow is sufficient
(Determann et al. 1997).
BRIT.ORG/SIDA 20(1)
Macranthera flammea
Fic. 1.M hera fl a | line leaf; b, branch of infl ;¢ flower with exserted stamens and style;
ie . 4 J Paes Chk - (onnn\ ‘me - =
-PUUTdI yp
Woody plants associated with M. flammea include: Acer rubrum L., Aronia
arbutifolia (L.) ELL, Clethra alnifolia var. tomentosa (Lam.) Michx., Cliftonia
monophylla (Lam.) Britt. ex Sarg., Cyrilla racemiflora L., Fraxinus caroliniana
Mill. Ilex coriacea (Pursh) Chapm., Ilex glabra (L.) Gray, Itea virginica L.,
Liriodendron tulipifera L., Lyonia lucida (Lam.) K. Koch, Magnolia virginiana
L., Myrica cerifera L., Nyssa sylvatica var. biflora (Walt.) Sarg., Persea palustris
ary
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Fic. 2
IL J 62 £ L ‘. | ° !
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raceme; C, lower axillary raceme; a, upper bud on branch; b, median bud; c, lower bud.
(Raf.) Sarg., Pinus elliottii Engelm., Pinus palustris Mill, Pinus serotina Michx.,
Rhododendron viscosum (L..) Torr, Viburnum nudum L. Herbaceous associated
taxa include: Andropogon virginicus L., Aristida stricta Michx., Arnoglossum
sulcatum (Fern.) H. Robins., Arundinaria gigantea (Walt.) Muhl., Coreopsis
gladiata Walt., Elephantopus nudatus Gray, Erianthus giganteus (Walt.) MuhL,
Eryngium integrifolium Walt., Helianthus angustifolius L., Hypericum
cistifolium Lam., Hypericum galioides Lam., Hypericum tetrapetalum Lam.,
Hyptis alata (Raf.) Shinners, Juncus validus Coville, Ludwigia hirtella Raf., Os-
munda cinnamomea L., Panicum scoparium Lam., and Pteridium aquilinum
(L.) Kuhn.
Common Name.—This species is commonly referred to as flame flower or
hummingbird flower. In at least one instance, it was referred to as Spanish prin-
cess (Pickens 1955). Flame flower is an appropriate name for M. flammea. It is
representative of the specific epithet because of the flame-like appearance of
the orange-yellow corolla (and the candelabra branching habit lends a flame
shape to the inflorescence). Hummingbird flower is less desirable as is could be
applied to any number of species that have red or orange tubular corollas.
BRIT.ORG/SIDA 20(1)
Fic. 3.0 g
EXAMINATION OF MORPHOLOGICAL VARIABILITY
Taxonomists base plant descriptions on the morphology of repeating modular
units of construction, i.e. leaves, flowers, etc. because, although the number of
units and size of the whole plant can vary greatly with age and environmental
conditions, genetic control maintains consistent size and form of modular units
(Harper 1977). Differences between species depend on the morphological form
of repeating units. Variation in characters may exist across the range of a spe-
cies, within a single population, and even within a single individual.
> as defined by Harper (1977).
Macranthera inflorescences are indetet
Meristems continue to produce new flowers and leaves, and the position of an
individual flower on the inflorescence reflects its age. By measuring changes
in floral characters through time, we determined how each individual flower
is affected by its location within the architecture of an individual plant and
her Pennell (913) was correct in stating that the differences (in calyx
—
whet
length and lobing) between the two species recognized at that time (M.
fuchsioides and M. lecontei) were in fact attributable to age.
We examined herbarium specimens (Appendix 1) to determine the extent
of variability of morphological characters across the species’ range. Characters
considered were calyx length, corolla tube length, corolla lobe length, corolla
width, pedicel length, bract length, fruit length, and fruit width. Because it was
not always possible to know which part of the plant was represented by the
herbarium specimen, we standardized measurements by choosing floral buds
at the stage of development just prior to the separation of the corolla lobes.
Lengths and width of mature fruits and, when available, the length and width
of the lower and uppermost Gust below the inflorescence) stem leaves were
measured.
To examine the extent of variability within a population and also withina
single plant, we randomly selected five plants from the established study site at
GWD in Thomas Co., Georgia, that represented the range of environmental
conditions for the site. Three flower buds (upper, median, and lower) were
marked on each of three racemes of the inflorescence (terminal, upper axillary,
and lower axillary), for a total of nine measured buds per plant (Fig. 4). Charac-
ters previously mentioned were then measured on marked buds every four days
fora period of one month. We measured fruit and foliage on 12 plants randomly
selected from the population.
Means and standard deviations were calculated for all measured charac-
ters from herbarium specimens (Table 3). Means and standard deviations of flo-
ral characters from the GWD sample population were calculated from charac-
ters measured on 28 August 1997 in order to determine the amount of variability
for characters within the plant architecture at a given time (Table 3).
The standard deviations of measured characters (Table 3) for herbarium
specimens reveal that although the floral tissue components (corolla tube) of
the flower show little variability, variability is higher in the chlorophyll con-
taining components (calyx, pedicel, and bract). The GWD population exhib-
ited more variability overall in corolla tube and calyx lengths than the her-
barium specimens because smaller and larger buds were included.
Plant 59 of the GWD sample population was selected to illustrate growth
changes over time in corolla tube length (Fig. 4) and calyx length (Fig. 5) for
each bud position of the three racemes examined. For each raceme, the corolla
tube expanded rapidly (Fig. 4). On the terminal and lower axillary racemes,
the tubes expanded to 22-24 mm before anthesis. Those of the upper axillary
raceme reached only 20-22 mm. Corolla tube width was between 4 and 5mm
for all flowers examined.
The calyces of the lower and median buds on the terminal raceme (Fig. 5,
a) had nearly reached their limits of expansion at the initial measuring, but
—
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1G.4 la . a | the lee wae I ca CAND diet A
I g August 1997.A, terminal raceme; B,
upper axillary raceme; C, lower axillary raceme.
calyx growth on the upper bud exhibited rapid elongation before reaching a
plateau. Calyces of buds on the upper and lower axillary racemes (Fig. 5, b-c)
also demonstrated rapid expansion before leveling off. The calyces of the up-
per buds were much shorter than those of the median and lower buds for all
racemes examined.
Figure 6 illustrates calyx growth for the upper buds of the three racemes
for three of the plants comprising the sample population. It shows that the up-
per axillary raceme had shorter calyces than did the lower axillary and termi-
nal racemes for plants of the sample population.
Pedicels exhibited the same growth pattern as calyces. Bracts generally did
not expand further once anthesis had occurred for an individual bud. Bracts
Table 3. Means and standard deviations for selected features of Macranthera flammea.
Herbarium specimens GWD plants
Character X S N x 5 N
calyx length,mm 10.89 3.5] 254 18.44 4.28 27
corolla tube length, mm 21.10 1.24 263 22.78 249 9
corolla lobe length, mm 3,86 0.81 240 --- —-- -
corolla width, mm 4.7) 0.78 194 4.1] 0.43 9
fruit length, mm 11.22 1.81 102 10.61 0.64 12
fruit width, mm 9.0 1.41 102 8.7 0.52 12
pedicel length,mm 15.88 3.64 220 21.88 2.27 27
bract length,mm 21.38 14.31 133 32.15 947 27
upper leaf length, cm 5.59 1.53. 93 9,59 1.47 12
upper leaf width, cm 1.8 0.9 93 3.19 0.77 12
lower leaf length,cm 12.5 3.81 28 19.72 2.03 12
lower leaf width, cm 4.21 2.26 28 4.03 0.4] 12
did exhibit very rapid growth during bud expansion, and some of the bract
lengths reached 50 mm for lower buds.
To produce more valid assessment of the amount of variation in calyx and
corolla tube lengths observed in the GWD population (Table 3), we selected buds
at various positions within the inflorescence and did not standardize to stage
of development. The mean lengths of calyx, corolla, pedicel, and bract were
greater in the GWD sample population because lower buds were included. Fruits
measured in the GWD population tended to be shorter, narrower, and less vari-
able than those in the herbarium specimens.
In general, variation was minimal in length and width of the corolla (Table
3, Fig. 4). Recent publications (Campbell et al. 1996; Smith et al. 1996; Temeles
1996) suggest that corolla width is the floral character selectively influenced
by hummingbirds, a major pollinator of M. flammea (Pickens 1927).
The position of a flower within the framework of the inflorescence reflects
its age (Harper 1977). Calyx length is determined by the position of a flower
within the framework and is therefore determined by age of the flower. Obser-
vations (Figs. 5, 6) indicate calyces reach a plateau in their growth and do not
continue to grow after flowering begins.
Pennell (1913) was correct in stating that the differences in the lengths of
calyx lobes are due to the age of flowers. However, because lobes do not grow
after anthesis, the position of the flower unit within the architecture of the in-
florescence accounts for observed differences in calyx lengths. These growth
characteristics are typical for plants with indeterminate inflorescences. Sepals
bracts, and pedicels are shorter for bud units at the tip of the raceme and longer
for those near the bottom. Internodes at the tip of the raceme are truncated,
those at the bottom elongated. Differences in the upper axillary branching po-
ey
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F J Za
upper axillary raceme; C, lower axillary raceme.
sitions can be attributed to apical dominance. The terminal racemes are always
the first to flower, followed closely by the lower axillary racemes. The racemes
flower upwards on the inflorescence so that the upper axillary racemes are the
last to reach anthesis (Fig. 2).
In general,as much variation appears to occur within a single plant or popu-
lation as throughout the entire range. Observed variation in the characters of
calyx, pedicel, and bract length can all be attributed to position effects within
the architecture of the plant. Age determines the placement of individual flower
units within an expanding inflorescence and therefore influences character
morphology.
A B
17.5-
1444
Ee E ne) oO Oo
E E
5 S
re.) Dm
& 5 12-4 Le ~-O---9
J aa
«x 12.55 Fd
> S
w
oO oO
10-
10-5 Pa upper bud —~(}— es upper bud —~(}—
we median bud - -O-— co] median bud - -O- -
$ lower bud - -O - - lower bud - -Q© - -
7.5 1 T 1 T T 8 T T T T T
8/13 8/17 8/21 8/28 9/14 8/13 8/17 8/21 8/28 9/14
1997 1997
16
Cc
_ 145 Oe Ss Oe- 2 =O
£
E
pon
>
g 125 O------6
a
Pad
>
i)
oO
10-5 F
é pper bud —-1}—
© median bud - -O-—
lower bud = -O - -
T T T T T
8/13 8/17 8/21 8/28 9/14
1997
Fic. 6. Ch i lif iti I i pl GWD during August 1997. A, termi-
nal raceme; ‘B, upper ‘axillary raceme; C, lower axillary raceme.
Observations of herbarium specimens and the GWD population do not
yield sufficient evidence of distinct variation in any character to warrant rec-
ognition of subspecific categories in M. flammea.
APPENDIX |
Specimens examined and annotated documenting known range and morpho-
logical variability of Macranthera flammea (exact localities omitted because
of rareness).
A. Baldwin Co.: 7 Sep 1912, F.W. Pennell 4553 (US, PH); 15 Oct 1940, M.G. Henry 2445 (GA),
ALAB
20 peers M.G. Henry 4935a (PH); 23 Aug 1947, M.G. Henry 4942 (PH); 16 Sep 1964, S.M. McDaniel
5314 (FSU), 11 Sep 1979, W.S. Judd 2288 (FLAS): 20 Aug L981, R. Kral 67854 (VDB). Butler Co.: 11 Oct
202 BRIT.ORG/SIDA 20(1)
1970, R. Kral 41623 (VDB). Conecuh Co.: 21 Aug 1981, R. Kral 67829 (V DB): 18 Sep 1993, R. Kral 83317
(VDB). Covington Co.: 3 Sep 1985, S.A. ae a 3011 VSU). Escambia Co.: 21 Oct 1961, R. Kral 38617
(VDB, NY): 17 Aug 1978, R. Kral 32463 (VDB). Geneva Co.: 3 Sep 1909, R. Kral 36844 (VDB). Mobile
Co.: 1872, D.C. Mohr (NY); 1878, D.C. ae (US, NY); Aug 1892, D.C. Mohr (NY): 16 Sep 1892, D.C.
Mohr (US); 30 Aug 1912, F.W. Pennell 44 oo US, PH); 31 Aug 1912, FW. Pennell 4459 (PH); 31 Aug
1912, FW. Pennell 4462 (PH); 4 Sep 1912, F.W. Pennell 4534 (MO, NY, PH): Jul 1919, E,W. Graves 1310
(MO); Aug 1919, E.W. Graves (US); 3 Sep 196, M.G. Lelong 4041 (USAM): 26 Aug 1909, H.D. Moore 621
(VDB); 3 Sep 1972, M.G. Lelong 6800 (USAM, VDB). Washington Co.: 21 Sep 1909, R. Kral 37233 (FSU):
3 Sep 1970, R. Kral 41062 JSU, VDB): 9 Sep 1970, H.D. Moore 1005 (VDB). County unknown: Gates
(NY); A. Gray (GH): J.-H. Redfield (MO).
FLORIDA. Bay Co.: 30 Oct 1957, M.G. Henry 6753 (PH); 3 Sep 1958, R.K. Godfrey 57619 (FSU).
Calhoun Co.: 6 Sep 1964, R.K. Godfrey 64605 (FSU): 18 Sep 1964, 8.M. McDaniel 5230 (FSU, VDB).
Escambia Co.: 8 Aug 1955, F.S. Ford 4314 (FLAS), 21 Jul 1950, P.L. Redfearn & R. Kral 2727 FSU), 9 Sep
1980, J.R. Burkhalter (FLAS), 1L Sep 1982, ].R. Burkhalter 8470 (MO). Franklin Co.: 20 Sep 1912, FW.
Pennell 4681 (GH, NY, PH); 22 Sep 1968, A.K. Gholson 3133 (AKG); 7 Sep 1970, A.K. Gholson (AKG); |
Sep 1976, L. Conde (FLAS); 12 Sep 1984, L. C. Anderson 7573 (FSU), 11 Sep 1985, L.C. Anderson 8798
“SU). Gadsden Co.: Sep 1835, Chapman (NY, US), 1836, Chapman (GH); 10 Jan 1909, Chapman (GH):
30 Aug 19306, H. Foster 120 (FLAS). Holmes Co.: 3 Sep 1942, LT. Nieland (FLAS); 27 Aug 1947, M.G.
Henry 4967 (PH). Jackson Co.: 21 Aug 1971, LJ. Musselmann & G.S. Waggonner 4393 (FSU, GH, NY,
ODU, GA, VDB); 10 Oct 1978, A.K. Gholson 7416 (AKG), 5 Oct 1979, A-K. Gholson 8004 (AKG): I Sep
1983, AK. Gholson 10629 (AKG). Leon Co.: 17 Sep 1960, HLL. Stripling 904 (FSU, 4 plants); 2 Jun 1970,
RJ. Nielsson (FLAS). Liberty Co.: 3 Sep 1955, R.F. Thorne & R.A. Davidson 16831 (FLAS, FSU, GA); LI
Sep 1974, L.C. Anderson 3817 (FSU); 15 Sep 1979, R.K. Godfrey 77250 (FSU); 15 Sep 1979, A.K. Gholson
7985(AKG);7 Nov 1987, P.W. Alcorn 378 (FLAS); 13 Sep 1989, S.L. Orzell & E.L. Bridges 12054 (GA); 22
Oct 1993, L.C. Anderson 14594 (FSU); 13 Sep 1997, N. Jordan 16 (FSU); 27 Sep 1995, J.D. Alford 220
FSU); 29 Sep 1997, J.D. Alford 1064 (FSU). Okaloosa Co.: 3 Oct 1949, S.C, Hood 3214 (FLAS): 19 Oct
oa R.K. Godfrey 71005 (FSU), 2 Sep 1970, A.K. Gholson 7985 (AKG), 2 Sep 1976, R.K. Godfrey
75446(FSU); 16 Sep 1993, L.C. Anderson 14558 (FSU). Santa Rosa Co.: 9 Sep 1912, F.W. Pennell 4564
(MO, PH); LO Sep 1912, F.W. Pennell 4595 (GA, PH, US); 25 Aug 1957, E.S. Ford 5403 (FLAS): | Nov 1969,
S.M. McDaniel 12519 (FSU, VDB); 2 Sep 1976, R.K. Godfrey 75445 (FSU); 29 Sep 1990, [.R. Burkhalter
23487 (FSU). Walton Co.: 2 Oct 1901, A.H. Curtiss 6922 (GA, GH, MO, NY, US); 23 Sep 1956, R.K. Godfrey
& R. Kral 55157 (FSU, GA); 25 Aug 1906, D.B. Ward 6006 (FLAS); | Oct 1970, H.A. Davis 15418 (FLAS.
GA, VDB), 6 Sep 1972, E.H. Sargent & H.A. Davis 16029 (MO). County unknown: Chapman (NY); Moht
(NY
Cs
GEORGIA. Bulloch Co.: H. Hall (GA). Crisp Co.: 4 Sep 1954, W. H. Duncan 18189 (GA, VSC).
Early Co.: 12 Aug 1947, RF. Phorne (GH, US); 25 Aug 1948, R.E. Thorne & W.C. Muenscher 8591(GA).
Emmanuel Co.: 2 Sep 1968, J.R. Bozeman 11727 (PSU, GA, GH, HATT, MISS, MO, NY, ODU, VDB, VSC);
22 Aug 1983, S.B. Jones 23813 (FSU, GA). Thomas Co.: 1897, $.M. Tracy (MO), 12 Sep 1998, J.D. Alford
1077 (FSU). Turner Co.: 21 Sep 1975, W.R. Faircloth 7893 (VSC). Ware Co.: 25 Aug L946, F.S. Sargent 122
(US). Wilcox Co.: 28 Aug 1964, R. Kral 22278 (V DB). Worth Co.: 25 Aug 1947, RF. Thorne 6362 (NY).
County unknown: fe nee {12395 (MO)
LOUI any Parish: 1846, ( Chapman (GH). 1846, Dru d (GH); 1909, A. Gray
(GH); 1] Seb mee G. “i ac, z 2911(LSU);,7 Sep 1985, A.W. Lievens 1003 (L SU): 7 Sep 1985, L.E. Urbatsch
(LSU). Washington Parish: 27 Sep 1936, C.A. Brown 6737 (LSU); 9 Sep 1962, CA. Brown 17584 (LSU); 31
Aug 1967, LJ. Mussellmann 1243 (MO), 28 Oct 1967, J.W. Thieret 28304 {FSU, GH); 16 Aug 1970, D.
Ziegler (MO); 13 Sep 1975, B.D. Thomas 46210 (VDB); 22 Oct 1975, Rich 209 (ODU); 20 Aug 1987, N.M.
Gilmore 3288 (LSU).
MISSISSIPPI. Forrest Co.: 1+ Aug 1947, M.G. Henry 4888 (PH): 7 Sep 19605, ].W. Carter 941 (HAT TD;
3 Oct 1970, K.E. Rogers 4679 (V DB); 15 Sep 1974, J. W. Wooten 2519 (USM), 24 July 1976, J.A. Barnes 648
(VDB). George Co.: 30 Sep 1995, S.W. Leonard 9173 (FSU, 26 plants). Hancock Co.: 17 Aug 1969, FH.
Sargent 9885 (MO, VDB); 16 Sep 1970, S. B. Jones 20312 (FSU, GA, MISS, VDB); 16 Sep 1970, FH. Sargent
10390 (FSU); 28 Aug 1972, H.L. Clark & K. Clark (FSU, HATT); 29 Aug 1977, M. Brooks (FSU). Harrison
16 Sep 1885, J.H. Redfield ie pa 16217 (GH, MO, NY, US); 18 July 1950, G.L. Webster & R.L.
mae 3460 (GH, NY, US); 18 Aug 1950, D. Demaree (VDB); 16 Aug 1952, D. Demaree 32853 (GH); 19
Aug 1971, C.A. Brown 22040 ca eee al Co.: 11 Sep 1889, F.S. Earle (NY); 16 Aug 1947, MG.
Henry 4921 (PH); 18 Aug 1949, D. Demaree 28202 (NY); 20 Aug 1949, D. Demaree 28300 (NY, VDB),
24 Aug 1951, D. Demaree 31281 (V DB); 17 August 1952, R.L. Diener 509 (MO); 24 Aug 1953, R.B. Chan-
nell 343 (VDB); LL Sep 1975, M. Arguelles 1455 (VSC). Lamar Co.: 14 Aug 1927, F. Cook (US). Layton
Co.: 15 Aug 1925, F. Cook (US). Marion Co.: 8 Aug 1955, J.D. Ray (GA, GH, NY); 5 Sep 1963, R. Kral 19389
(VDB). Pearl River Co.: 19 Sep 1982, S.M. McDaniel 26507 (FSU). Perry Co.: 15 Aug 1947, M.G. Henry
4904 (PH); 4 Oct 1995, S.W. Leonard 9181 (FSU, 28 plants). Stone Co.: 24 Aug 1953, D. Demaree 34081
(GH, VDB); 8 Oct 1966, T.M. Pullen 661140 (UM). Winston Co.: 6 Sep 1932, F.H. Sargent (NY). County
unknown: 30 Aug 1941, G.A. Girhart 87 (NY).
ACKNOWLEDGMENTS
We thank the Greentree Foundation for allowing access to the Big Woods at
Greenwood Plantation. Julie and Leon Neel provided valuable logistical sup-
port in the field. We thank Jean C. Putnam-Hancock and Shirlann Strickland
for use of botanical illustrations. Nancy C. Coile, Sarah Matthews, A.B. Thistle,
and an anonymous reviewer provided helpful suggestions on the manuscript.
REFERENCES
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flammea (Batr.) Pennell. M.S. Thesis, Florida State Univ., Tallahassee.
Bartram, W. 1791. Travels in North & South Carolina, Georgia, East and West Florida. James
and Johnson, Philadelphia, PA.
BentHam, G. 1835. Synopsis of the Gerardiaeae, a tribe of the Scrophulariaceae. Compan-
ion Bot. Mag. 1:198-212.
BerTo.onl, A. 1853. Memorie della Accademia della Scienze dell’ Instituto di Bologna 4:75.
Campsett, D.R., NM. Waser, and M.V. Price. 1996. Mechanisms of hummingbird-mediated
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Carin, L.G. 2000. Field guide to the rare plants of Florida. Florida Natural Areas Inventory,
Tallahassee.
CHapman, A.W. 1860. Flora of the southern United States. Ivison, Blakeman & Co., New York.
Cotte, N.C. 2000. Notes on Florida’s endangered and threatened plants. Contribution no.
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DeTERMANN, R., L.K. KigkMaN, and H. Nourse. 1997. Plant conservation by propagation. Tipularia
12:2-12
Feanao, M.L. 1944. Overlooked species, transfers and novelties in the flora of eastern North
America. Rhodora 46:1-21.
Goorrey, R.K.and J.W.Wooten. 1981. Aquatic and wetland plants of the southeastern United
States: dicotyledons. University of Georgia Press, Athens.
Harper, J.L. 1977. Population biology of plants. Academic Press, New York.
204 BRIT.ORG/SIDA 20(1)
Hooker, W.J.1835.Notice concerning the late Dr. Drummond's collections, made chiefly in
the southern and western parts of the United States. Hooker's Companion Bot. Mag.
Kuntze, O. 1891, Revisio Generum Plantarum 1:459.
Merritt, E.D. 1945. In defense of the validity of William Bartram’s binomials. Bartonia 23:
10-35.
Nurtait, T. 1834. A description of some of the rarer little known plants indigenous to the
United States, from the dried specimens in the herbarium of the Academy of Natural
Sciences in Philadelphia. J. Acad. Nat. Sci.Philadelphia 7:88.
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tegration of the Scrophulariaceae. Amer. J. Bot. 88:348-361.
Pennett, EW. 1913. Studies in the Agalinanae, a subtribe of the Rhinanthaceae. Bull. Torrey
Bot. Club 40:119-130.
Pennett, FW. 1935. The Scrophulariaceae of eastern temperate North America. The Acad-
emy of Natural Sciences of Philadelphia, Philadelphia.
Pickens, A.L. 1927. Unique method of pollination by the ruby-throat. Auk 47:346-352.
Pickens, A.L. 1955. The bird-flower as the apex of floral color display. Castanea 20:1-18.
Rarinesaue, C.S. 1837. New flora and botany of North America 2: 58-72. Philadelphia.
Rickett, H.W. 1944, Legitimacy of names in Bartram’s “Travels.” Rhodora 46:389-391.
SMALL, J.K. 1933. Manual of the southeastern flora. 1972 Reprint. Hafner, New York.
Situ, C.E., J.T. Stevens, EJ. Temeces, PW. Ewato, RJ. Hesert, and R.L. Bonkovsky. Effect of floral
orifice width and shape on hummingbird-flower interactions. Oecologia 106:482-292.
Temetes, EJ. 1996. A new dimension to hummingbird-flower relationships. Oecologia
105:517-523.
Torrey, J. 1837.An account of several new genera and species of North American plants.
Ann. Lyceum Nat. Hist. New York 4:80. pl. 4.
Wipur, R.L. 1971. A reconsideration of Bartram’s binomials. J. Elisha Mitchell Sci. Soc. 87:
56-73.
THE GENUS CAMPOMANESIA (MYRTACEAE) IN ATLANTIC
RAINFOREST FRAGMENTS IN SERGIPE, NORTHEAST
REGION OF BRAZIL
Myrna F.Landim! Leslie R.Landrum
Departamento de Biolog Department of Plant Biology
Arizona State University
Universidade Federal de Sergipe
49100-000, Sao Cristovao, SE, BRAZIL Tempe, AZ 85287-1601, U.S.A.
fs.br les. landrum@asu.edu
andim@u
ABSTRACT
The Atlantic forest is one of the ecosystems of Brazil that has suffered the greatest destruction and it
I ates wl he Atlantic forest has
is sll imperiled, especially in the Northeast. Sergipe is one of the st
been most greatly reduced, but it still has some fragments varying in size and degree of conserva-
tion. Despite their importance, botanical studies within these fragments are still greatly needed. In
this paper we report for the first time the occurrence of four species of Campomanesia in Sergipe: C.
C. viatoris. ove aa oaueny thought to be known from
| and is found in forest
dromaticad, . dichotoma, C. guaviroba, and
f Alagoas, hasa relatively
onl
and restinga areas of different districts. Campomanesia Just roba is seperted for the first time from
f these remnants may contribute
the Northeast of Brazil. We hope that studies on th
to the local preservation of this ecosystem and help Fei I
phasize the need for a greater collecting effort, lies on plant taxonomy, and the training of
new specialist
programs of forest restoration. We em-
RESUMO
ist s brasileiros mais destruidos e ainda hoje ameagado, sendo a
A mata atlantica é umd
sua devastacao mais pronunciada no Nordeste do Brasil. Um dos estados com menor area de mata
atlantica, SenEIDE apieeenta ainda hoje alguns fragmentos dont com tamanho eran as
11¢ dos |
Arlavels Apesar de sua import ancia, os es
corréncia inédita
CULISC! Vata ao
insuficientes. Nesse trabalho registi de qu
rgipe: C. aromatica, C. dichotoma, C. nance viatoris. Esta on eur te ences onhecidz
aay por oe cases ambas em Alagoas, apresenta uma distribuicdo relativamente am - no
areas de mata e restinga de diversos municipios. Registra-se, ainda, a
regiao Nordeste. Espera-se que estudos sobre a
e Campomanesia
ee
Ss
sta
ocorréncia a C. aie ba a primeira vez na
s fragmentos possam contribuir para a }
ye de recomposicao florestal. Salienta-se a necess
de mais estudos sistematicos e da formacao de novos oe
sreservacao desse ecossistema a nivel re-
ade de u
von e subsiia oe
S nso
inten
Key worps: Campomanesia, Myrtaceae, Sergipe, Atlantic rainforest fragments
INTRODUCTION
The objective of this note is to report the occurrence in Sergipe, Brazil of four
species of Campomanesia Ruiz et Pavén (Myrtaceae), usually known locally as
“suabirobas’ or a variant of this name (e.g., guavirobas, gobiraba, gabiroba). None
Universitat Bremen, UFT-AG Heyser, D-28359, Bremen, GERMANY. Email i-bremen.de
SIDA 20(1): 205 — 214. 2002
206 BRIT.ORG/SIDA 20(1)
a
of these species, nor any other species of Campomanesia, was known for Sergipe
W a Landrum (1986) monographed the genus, and one species (C. guaviroba)
has not been reported belore in Brazil’s Northeast region (Maranhao, Piaui, Ceara,
Rio Grande do Norte, Paraiba, Pernambuco, Alagoas, Sergipe, Bahia). The fact
that recent floristic exploration can reveal so much is an indication of the im-
portance of continuing efforts in this field.
Despite the great biological diversity of the Brazilian coastal forests, little
of its original cover still persists. At the time of the European discovery such
forest covered about 1,000,000 km? of Brazil, while today its area has been re-
duced to an estimated 5% (Consorcio Mata Atlantica 1992), 6% (Prance et al.
2000), or 8% CUINPE and IBAMA 1990) of its original size. In spite of being pro-
tected by law, destruction of coastal forest still continues. The remaining areas
consist of fragments varying in size and degree of conservation and are “typi-
cally small, isolated and highly disturbed” (Viana et al. 1997). Ranta et al.1998)
recently studied the degree of fragmentation in the Northeast Atlantic forest
of Brazil,and emphasize the importance of fragment size and shape for the long-
term survival of biodiversity. Smaller fragments and more irregularly shaped
fragments have a higher proportion of their areas in edges, which are more vul-
nerable to plant extinction. Because fragmentation in Sergipe has progressed so
far, extinction is more imminent.
The historical pattern of almost complete elimination of the native forest
covering in Brazil’s Northeast is explained by the relative gentle relief of the
landscape and ease of access to all areas. Farther south, the mountainous ter-
rain of the Serra do Mar makes access more difficult, and thus greater portions
of Atlantic rainforests there are relatively protected. Sergipe, the smallest of the
Brazilian states, originally had about 41% its area covered by forests (Campos
1912), but its forest covering today is estimated to be less than 1%. For this rea-
son, Sergipe has been excluded in some mappings of the remnants of Atlantic
forest carried out at the national level. However, floristic surveys in areas of
Atlantic forest in the state (Landim et al. 1998) have shown it to be a forest with
unique floristic and structural characteristics. Additional studies are urgently
needed that contribute to knowledge of the forest’s composition, structure and
dynamics, with the aim being a knowledge base for reforestation initiatives.
——
The family oe is mainly a tropical and subtropical family
(Heywood 1993), with about 3500 species and approximately 100 genera, with
two main centers of development: tropical America and Australasia (Barroso et
al. 1991). It isa family of great importance in the neotropical forests, being one
of the dominant families in Atlantic coastal forest (Barroso & Peron 1994; Leitao-
Filho 1993; Mori et al. 1983; Peixoto & Gentry 1990; Reitz et al. ses Ina study
of Atlantic forest in Sao Paulo, Mantovani (1993) found 38 tree species of
Myrtaceae (21% of the total). He considers Myrtaceae to be “the most c
—
Naracter-
istic family of arboreal species of Atlantic forest in the south coast of the State
of Sao Paulo, with floristic and structural importance.”
In general, species of Myrtaceae are quite common in the Atlantic forest
areas of the Northeast, (Siqueira 1994), and this has proved to be true the frag-
ments studied in Sergipe. In a phytosociological study in the Mata do Crasto, in
the Municipality of Santa Luzia do Itanhy (Landim et al. 1998), this family ac-
counted for 10.3% of the sampled trees, and was the sixth in basal area, and
fifth in importance value index (LVL).
Voucher specimens of collections described below are deposited in the her-
baria of the Department of Biology of the Federal University of Sergipe (ASE),
the University of Brasilia (UB), and Arizona State University (ASU).
RESULTS
Description of Campomanesia
Campomanesia can be distinguished from other genera of Myrtaceae by: 1) ovary
with (3-)4-18 locules (Fig. 1D); 2) ovules several per locule, biseriate, all or all but
one aborting in each locule; 3) locule-wall in the mature fruit chartaceous to
slightly woody, strongly glandular, serving asa false seed coat (Fig. 1C). In Sergipe,
ampomanesiacan be distinguished from most other genera of Myrtaceae when
in flower by its 5-merous flowers and inflorescences of solitary flowers or dicha-
sia. Psidium L. and Calycolpus O. Berg can be confused with Campomanesia in
flowering specimens but generally have coriaceous leaves, whereas
Campomanesia usually has submembranous to membranous leaves.
Campomanesia is quite distinctive in fruit. There seem to be several seeds
in each fruit arranged ina ring, each with a glandular covering (Fig. 1D). How-
ever, each of these “seeds” is a locule, and usually only some of them havea seed
inside. In others the ovules have all aborted. Other genera of Myrtaceae either
have few seeds (e.g., Lor 2 in Myrcia, Eugenia, or Calyptranthes) or the seeds are
numerous and hard (e.g., Psidium, Calycolpus). In no other American genus of
Myrtaceae do the seeds have a glandular covering.
The bark of the trunk of Campomanesia is usually rough with numerous
narrow, papery plates, unlike most other genera that have smooth to scaly bark.
The lateral veins of the leaves are generally prominent and broadly arch near the
ler veins often show an intricate reticulate pattern (Fig. 2).
—
—
margin and the sma
KEY TO THE SPECIES OF CAMPOMANESIA IN SERGIPE
—
Calyx nearly closed in bud; inflorescence a dichasium with peduncle 2-6 cm long
Campomanesia dichotoma
1. et open in bud; inflorescence uniflorous or a dichasium with peduncle up to
cm long.
2. Hypanthium attenuate at base, constricted near apex; anthers 1-2 mm long
Campomanesia viatoris
BRIT.ORG/SIDA 20(1)
y : Mer yuddAdia’ VON i, NI Mic
¥ ee \) FEST Z / 7 G uy hh Lath
7A : ey 2. ts Y Cay
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CH? AOL .
f x Wee
ih, . wt oh
: , ‘ RUAN Ed
u . ° oue ME as
(fe M piace e
/ 7 foc we i t a
a . ne , pure ry May + !
wae They ry, | vethye : A
es owe Lo vt ics CT . RAND
\ é ) Pad bE Depo L Viies HG Rae He
ae ‘ ; ‘44, Wier? YY Orns
a rytly a ae a” ed nee NP,
‘5 ee
( 24 BASS
YAEL ; if Wee Z i : ‘te
Ag al Yates 23 AS CEES
S ‘ Chee y eae Re
Re Rapa “SY i)
SY Na ee ak aged
4 Kt ‘ i
SAN tg es
SSN ee
= at Byes ‘ ‘ae
Raut a ‘,
a a aes Di PNR sy)
BS i sae
Sy 4 Ri
val
A * BF
sw \
4
Fic. 1. A, Flower bud of Campomanesia viatoris. B, Flower bud of C. guaviroba
1 a | 1 Th ah Sut rT + Np ft +) £ £, H
e ieanl lee ep
glandular locular ; t having 13 locules, each with a glandular
wall, [ ly rel £ | em dinky P| ip] +} | y f : a £. * = | y L ry th
one place. Upper bar = 5 mm for A & B. Lower bar = 4 mm for C and 10 mm for D.
2. Hypanthium neither attenuate at base nor constricted near apex; anthers 0.5—1
mim long.
. Calyx-lobes hemiorbicular to oblong-truncate,
leaves immature at anthesis; fruit bl
).7-2.5 times as long as wide;
ack _ Campomanesia aromatica
3. Calyx-lobes broadly triangular or broe 0.9 times as long as
idly rounded, 0.4
wide; leaves mature at anthesis; fruit yellow or orangish Campomanesia guaviroba
Campomanesia aromatica (Aubl.) Grisebach
Shrub or tree 2-20 m high; leaves i ature at anthesis, elliptic, ovate, |
anceolate,
PN ee
ee
eee
as
se Pep Stare
ists SSS aae
B
ee thiandn 0 | VR Se |
J
Fic. 2. A, ical leaf of j biroba st ingi i k
+ F loaf chawinag datail of tion (Land 2490, ASU)
or oblanceolate, 4-12(-15) cm long, 2-4(-7.5) cm wide, 1.7-4 times as long as
wide; peduncles 0.3-2.6 cm long, uniflorous; bracteoles 1-4 mm long; calyx-
lobes hemiorbicular to oblong-truncate, 2-4+.3 mm long; hypanthium obconic
to campanulate, 1.5-2.5 mm long; stamens 80-90, 4-7 mm long; anthers 0.5-0.8
mm long; ovary 4-6-locular; ovules 4-7 per locule; fruits ca. 1 cm long, black.
Specimens examined. Mun. Salgado: Capoeira de mata subperenifolia, 18 Dec 1981, G.N. Silva 39 (ASE).
Mun. Lagarto: Caatinga hipoxerofila, 7 May 1982, FE. Carneiro 374 (ASE). Mun. Itabajana: Capocira de
caatinga hipoxerofila, 20 Jul 1988, M. Fonseca 857 (ASE). Mun. Sta. Luzia do Itanhy: Mata Atlantica,
Mata do Crasto, 3 May 1995, M. Landim 354 (ASE, ASU, UB).
Although species of Campomanesia are usually known locally as “gabiroba” or
variations of this name, a specimen collected in the caatinga region was called
“cadeia brava.”
In his revision of this genus, Landrum (1986) cited only a few collections
of C.aromatica in the Northeast, namely in the states of Maranhao, Ceara, Rio
Grande do Norte, Paraiba and Bahia. The species also occurs in Bolivia (Landrum
1986), an interesting disjunction of over 2000 km. In the present study, it has
210 BRIT.ORG/SIDA 20(1
been found in areas of coastal forest (Mata Atlantica) as well as caatinga, a
shrubby vegetation of dry areas. A greater sampling effort is needed to better
understand the distribution of this species.
Campomanesia dichotoma (O. Berg) J.R. Mattos
Shrub or tree up to LO m high; leaves mature at anthesis, elliptic, elliptic-ob-
long, ovate or suborbicular, 3-9.5 cm long, 1.5-6 cm wide, l-2.3 times as long as
wide; peduncles 2-6 cm long, bearing 3-15 flowered dichasia; bracteoles ca. 2
mm long; calyx nearly closed in the bud, the calyx-lobes 1-2 mm long, the ca-
lyx tearing between the lobes at anthesis; hypanthium (including closed ca-
lyx) 5-8 mm long; stamens 200-350, 3-10 mm long; anthers 0.8-1 mm long;
ovary 6-10-locular; ovules 7-18 per locule; fruits ca. | cm long.
Specimens examined: Mun. Santa Luzia do Itanhy: Mata Atlantica, Mata do Crasto, 19 Apr 1995, M.
Landim 333 (ASE, UB), 3 May 1995, M. Landim 356 (ASE, ASU, UB), 12 Jun 1995, M. Landim 406 (ASE,
ASU, UB), 24 May 1990, M. Landim and V.M. Schettino 920 (ASE, ASU)
The common name “bacalhau’ is used for this species as well as the more com-
mon generic designation of “gobiraba.”
Campomanesia dichotoma is known from the coastal region of the North-
east of Brazil and the state of Rio de Janeiro (Landrum 1986). So far in Sergipe
examples have been found in only one fragment of Atlantic coastal
orest.
Campomanesia guaviroba (A.P. de Candolle) Kiaerskou
Tree up to ca. 12 m high; leaves mature at anthesis, mainly elliptic, less often
ovate, lanceolate, suborbicular, or ovate, 4-13 cm long, 1.7-8 cm wide, 1.4-3 times
as long as wide; peduncle 0.3-2.5 cm long, uniflorous; bracteoles ca. 4 mm long:
calyx-lobes broadly triangular or rounded, L.5-3 mm long; hypanthium obconic
to campanulate, 4-6 mm long; stamens 250-500, 3-9 mm long; anthers 0.5-1
mm long; ovary 7-14-locular; ovules 13-20 per locule; fruit 1-3.5 cm long, yel-
low or ee — 1B).
a Luzia tanhy: Mata ie Mata do Crasto, 3 May 1995, M.
im 346 (ASE, U 25 an es M. a 906 (ASE, ASL
Specim
Lane
Until recently Campomanesia guaviroba was known only from Brazil’s Atlan-
tic forest from Espirito Santo to Rio Grande do Sul and adjacent Argentina and
Paraguay (Landrum 1986). The collections cited above extend the range far to
the north (ca. 1200 km) and are the only ones known to us from the Northeast
of Brazil. Recently a collection has extended the known range to Bolivia [Santa
Cruz, Velasco Province, Parque Nacional Noel Kempff Mercado, 5 km S del
campamento Las Gamas (14° 48'14"S, 60° 23'59"W), 850 m, A Rodriguez & J.
Surubi’ 558 (ASU)|. It thus has a disjunct distribution similar to that of C.
dromatica.
In one area of Sao Paulo (Mantovani 1993) this species was represented by
only two individuals, having an LV. of 0.58. Apparently rare, it was not found
in the forest gap areas in the same study, although other Myrtaceae were, e.g.,
211
Eugenia, Marlieria, Calyptranthes, Myrciaand Gomidesia. This may mean that
it is less tolerant of disturbance (anthropogenic or not) and thus is at a higher
risk of going extinct through habitat disturbance and fragmentation.
Campomanesia viatoris Landrum
Shrub or tree to ca. 12 m high; leaves mature at anthesis, elliptic to ovate, 3-13
cm long, 2-6.5 cm wide, 1.5-2 times as long as wide; peduncle 0.5-2 cm long,
uniflorous or bearing a 3-flowered dichasium; bracteoles ca. 3 mm long: calyx-
lobes truncate-auriculate, 1-3 mm long; hypanthium 8-10 mm long, attenuate
at base, constricted near summit of ovary; stamens ca. 300, 4-8 mm long; an-
thers ca. 1-2 mm long; ovary 7-8-locular; ovules 11-16 per locule; fruit globose
except for an attenuate base, up to ca. 2.5 cm in diameter. (Fig. 1 A).
Specimens examined. Mun. Sto Amaro das Brotas: restinga, 10 Dec 1981, G. Viana 294 (ASE). Mun.
Sao Cristovao: restinga, 21 Mar 1985, G. Viana 1112 (ASE). Mun. Pirambu: Reserva Biologica Santa
Isabel, dunas no km 11 de praia na direcao de Ponta dos Mangues, 28 Jan 1992, C. Farney et al. 2944
(ASU). Mun. Santa Luzia do Itanh ade 2.5 km do Distrito de Crasto, na estrada para Sta. Luzia
do Itanhy, 27 Nov 1993, A.M.A. Amorim et al. 1500 (ASU); Mata Atlantica, 7 Apr 1995, M. L selon ane
(ASE, ASU, UB), 3 May 1995, M. Landim 355 (ASE, ASU, UB), 5 Jul 1995, M. Landim 456 (ASE,
UB); Mata Atlantica, Mata do Crasto, 7 Apr 1995, M. Landim 285 (ASE, ASU, UB), 16 Dec 1996, M.
Landim 1108 (ASE, ASU). Mun. Sao Cristovao: restinga, 9 Apr 1996, M. Landim 904 (ASE, ASU). Mun.
Areia Branca: Mata, Serra de [tabaiana, 28 Mar 1997, A. Vicente 7 (ASE).
a
me
The common names for this species are “guabiraba,” “gabiroba,” “gabirobinha,”
“gobirabinha,” and “gobiraba.”
Until recently only two collections for this species were known, the type
and one other, both from Alagoas (Landrum 1986). The type was collected by
Gardner over 150 years ago and the additional specimen has no date. Since the
species was thought to be extinct or near extinction, Landrum (1986) urged
botanists in Alagoas to search for this rare species. The collections cited above
from Sergipe indicate that Campomanesia viatoris is at least locally common
in the state and additional specimens have also been found in Bahia. [e.g., Mun.
Apora, 31 km S of Olindina along highway BR-116, 310 m, | Apr 1976, G. Davidse
et al. 11758 (ASU).
The anthers of Campomanesia viatoris anthers are unusually elongate and
similar to those of C. laurifolia. This unusual shared anther morphology may
indicate an interesting pollination mechanism deserving more careful study.
DISCUSSION
Phytogeographic analysis of species of Myrtaceae is greatly hampered by the
difficulty in identifying material, due to the fact that American species of
Myrtaceae often are very similar in the majority of their characters (Mc Vaugh
1968), and because of the cryptic nature of the characters used to identify the
genera (Landrum and Kawasaki 1997). Some floristic and/or phytosociologi-
cal studies list great numbers of species or individuals of Myrtaceae, but au-
thors often are unable to identify them to species or genus (e.g., Guedes 1992). In
212 BRIT.ORG/SIDA 20(1)
a study of the flora of Atlantic rainforest, Myrtaceae are cited as the family with
the greatest number of unknown specimens (Siqueira 1994).
Ecological studies of the Northeast of Brazil will require many more spe-
cialists to identify the great number of collections unknown to species, genus,
or even family (Siqueira, 1994). Three conditions need to be improved: 1) there
needs to be more collecting in general as this paper demonstrates, 2)
monographers must study the flora of the Northeast and include specimens
from the regional herbaria; and 3) there should be a greater investment in train-
ing more specialists.
The absence of reports of Campomanesia in Sergipe in Landrum’s 1986 re-
vision of the genus reflects the low sampling effort in the state until recently. It
is still difficult to describe the distribution of the species in the state because
they are known from so few collections. The present study indicates the exist-
ence of a flora that is still relatively unknown in the fragments of Atlantic
rainforest in Sergipe and indicates the importance of the exploration and con-
servation of these areas.
The sympatric occurrence of these four Campomanesia species in one frag-
ment, the Mata do Crasto, in Santa Luzia do Itanhy, a coastal region in the south
of the Sergipe, provides opportunities for more studies. These might involve the
identification of or eee between these four species in flowering
and fruiting phenology, pollinators and fruit predators, and reproductive iso-
lating mechanisms that may have led to speciation.
Finally, it is worth noting that even in regions with forest coverings insuf-
ficient to be registered in some national surveys, such as Sergipe, the forest rem-
nants that do exist are the last representatives of an entire ecosystem, includ-
ing plants, animals, fungi, and micro-organisms. These fragments, mostly of
medium to small size, are the best possible estimation of the original ecosys-
tem that will ever be available. Studies on the composition of these fragments
can contribute to the preservation of that ecosystem at the regional level and
will be useful for future reforestation programs. In order to avoid the drastic
loss of large portions of biodiversity, the establishment of reserves is necessary,
at least for some of these fragments, and conservation management that in-
cludes corridors between fragments as well as the involvement of neighboring
human communities in this process is desirable
ACKNOWLEDGMENTS
Reviewers R. Schmid and N. Snow offered many helpful suggestions for which
we are quite grateful. We thank Eladio dos Santos for his help with fieldwork.
REFERENCES
Barroso, G.M.,A.L. Peixoto, C.F.L. IcHaso, C.G. Costa, E.F. Guimaraes, and H.C. Lima. 1991. Sistematica
de angiospermas do Brasil. Vol. 2. Universidade Federal de Vicosa, Vicosa.
213
Barroso, G.M. and V. Peron. 1994. Myrtaceae. In: M. P.M. Lima and R.R. Guedes-Bruni, orgs.
Reserva ecoldgica de Macaé de Cima, Nova Friburgo, RJ. Aspectos floristicos das
espécies vasculares., Vol. 1.Pp. 261-302.
Campos, G. 1912. Mappa florestal. Rio de Janeiro, Servigo Geoldgico e Mineraldgico do
Brasil. Consdorcio Mata Atlantica/UNICAMP, 1992. Reserva da Biosfera da Mata Atlantica:
Plano de Acao. Referéncias basicas, 1.Campinas.
Gueoes, M.L.S. 1992. Estudo floristico e fitossocioldgico de um trecho da Reserva ecologica
da Mata de Dois Irmaos Recife - Pernambuco. Tese de Mestrado, Universidade Federal
Rural de Pernambuco, Recife.
Heywoop, V.H. (ed.) 1983. Flowering plants of the world. Oxford University Press, New York.
INPE (INSTITUTO NACIONAL DE Pesquisas Espacials) and |BAMA (INSTITUTO BRASILEIRO DO Meio AMBIENTE E
bos Recursos Naturals RenovAvels) 1990. Atlas dos remanescentes florestais do dominio
da Mata Atlantica. A web based version is available at http://
www.sosmatatlantica.org.br/atlas2001/index.asp.
LAnoim, M.F., D.M. Santos, C.S. AMarat, and M.M. Maia. 1998. Fitossociologia de um
remanescente de mata atlantica em Sergipe, Mata do Crasto. XLIX Congresso Nacio-
nal de Botanica. Sociedade Botanica do Brasil, Salvador. Resumos
LanpbruM, L.R. 1986. Campomanesia, Pimenta, Blepharocalyx, Legrandia, Acca, Myrrhinium,
und Luma (Myrtaceae). Fl. Neotrop., Monogr. 45. The New York Botanical Garden, New
York.
Lanbrum, L.R. and M.L. Kawasaki. 1997. The genera of Myrtaceae in Brazil: an illustrated syn-
optic treatment and identification keys. Brittonia 49:508-536.
LeiTAo-FitHo, H.F. (org.). 1993. Ecologia da Mata Atlantica de Cubatdo (SP). Editoras UNESP/
UNICAMP.
McVauch, R. 1968. The genera of American Myrtaceae—an interim report. Taxon 17:354-
Mantovani, W. 1993. Estrutura e dindmica da floresta atlantica na Juréia, lguape-SP. Tese de
Livre Docéncia. Universidade de Sao Paulo, Instituto de Biociéncias, Sao Paulo.
Mori, S.A.,B.M. Boom, A.M.CarvalHo, and T.S. Santos. 1983.Ecological importance of Myrtaceae
in an eastern Brazilian wet forest. Biotropica 15:68-70.
Peixoto, A.L.and A. Gentry. 1990. Diversidade e composicao floristica da mata de tabuleiro
na Reserva Florestal de Linhares (Espirito Santo, Brasil). Revista Brasil. Bot. 13:19-25.
Prance, G.T., H. Beentue, J. DRANSFIELD, and R. JOHNS. 2000. The tropical flora remains
undercollected. Ann. Missouri Bot. Gard. 87:67-71.
Ranta, P., T. Blom, J. Niemeta, E. Joensuu, and M. Sitonen. 1998. The fragmented Atlantic rain
forest of Brazil: size, shape and distribution of forest fragments. Biodiversity and Con-
servation 7:385-403.
Reitz, R., R.M. Kcein, and A. Reis. 1978. Projeto madeira de Santa Catarina. Sellowia 30(28-
30):1-320.
Siqueira, M.F. 1994. Andlise floristica e ordenacdo de espécies arbdreas da mata atlantica
através de dados binarios. Tese de mestrado, Universidade Federal de Campinas,
Campinas.
214 BRIT.ORG/SIDA 20(1)
Viana, V.M., A.AJ. Tasanez, and J.L.F. Barista. 1997. Dynamics and restoration of forest frag-
ments in the Brazilian Atlantic moist forests. In: W.F. Laurance and R.O. Bierregard, eds
Tropical Forest Remnants—Ecology, Management, and Conservation of Fragmented
Communities. University of Chicago Press, Chicago, IL. Pp. 351-365.
BIOLOGICAL STATUS OF HEDYOTIS NIGRICANS VAR.
GYPSOPHILA (RUBIACEAE) IN TEXAS
B.L. Turner
Plant Resources Center
University of Texas
Austin, TX, 78713, U.S.A.
Terrell (2001) presented a taxonomic study of the genus Stenaria (= Hedyotis
nigricans and closely related cohorts, sensu Turner 1995). In this he recognized
H. nigricans var. gypsophila as occurring in Trans-Pecos, Texas, citing among
his “selected representative specimens” four sheets (cf. Fig. 1), two from
Culberson Co., a north-south ridge of limestone outcrops extending for ca. 50
miles from near Van Horn, Texas to near the New Mexico border, and one from
the same mountain range in closely adjacent Hudspeth Co.; the fourth collec-
tion (Webster 4501, MICH) was said to be from the “Madera Canyon” in igneous
soils of Jeff Davis County.
Terrell distinguished H. nigricans var. gypsophila (Fig. 3) from the rest of
his concept of the wide-ranging, highly variable H. nigricans by the following
couplet:
1. Leaves more or less elliptical, 2.5—3.5(—4) times longer than wide; Mexico and south
west Tex var. ness
1. es eer filiform, linear, narrowly lanceolate, or oblanceolate, more than 4 time
a than wide; ate 2-8 mm long (var. nigricans and closely ae cohorts]
Careful scrutiny of the above key will show that the only meaningful charac-
ter used by Terrell to define his “var. gypsophila” is leaf shape. Indeed, Terrell,
himself (2001), states that “the most conspicuous character [distinguishing be-
tween these elements] is the elliptical leaves on rather small plants..[and] I can-
not find any other characters significantly different from var. nigricans...”
Regardless, | would like to place on record here that I take many, if not
most, of the specimens cited by Terrell (2001) as occurring outside of the re-
gional distributions of H. n. var. gypsophila (Turner 1995) to be misidentifications
of H. n. var. nigricans, these but leaf and/or habital forms occurring among oth-
erwise typical populations of var. nigricans.
This can be readily attested to by reference to the single sheet (Webster 4501)
of H. nigricans var. gypsophila cited in the above introduction. A duplicate of
this collection at SRSC (Fig. 2) is clearly linear-leaved and could not be identi-
fied as var. gypsophila by Terrell’s key to varieties. Additionally, its habit and
floral features do not conform to those of var. gypsophila as described by Turner
(1995). Indeed, I have reexamined 200 or more specimens of H. n. var. nigricans
SIDA 20(1): 215 — 220. 2002
216 BRIT.ORG/SIDA 20(1)
mens of H. nigricans var
Fic. 1.G lized distribution of Hedyotis nigricans var. nigricans in Texas (sensu Turner 1995); dots represent speci-
=. Wwsonnt\.t| I L L x | Lm ae | Pay 44 4 - £u
a A ve Ft f
J JSF
nigricans var. nigricans.
from Trans-Pecos, Texas (LL, SRSC, TEX) and find not a single plant referable to
H.n. var. gypsophila, including most of the specimens so cited by Terrell (2001).
He further ventures that “I [Terrell] consider the taxon as misnamed, as I
recorded only one collection out of 49 Mexican collections from 19 herbaria
that mentioned a gypsum substrate. It may be noted that the type specimen
came from a gypsum habitat.”
However, had Terrell examined the 49 sheets of this taxon on file at LL,TEX
he would have found 12 sheets with labels noting their occurrence on gypsum
substrates, and most of the rest from areas known to possess such soils. The
truth is that most early collectors did not know the difference between gypsum
substrates (CaSO4) and limestone substrates (CaCO4), or else they ignored such
distinctions. The highly endemic flora occurring on gypsum substrates need
TURNER, BIOLOGICAL STATUS OF HEDYOTIS NIGRICANS VAR. GYPSOPHILA
eT, H TEX)
The Univers
Hed
8 ;
var. nigricans
Det: BL, Turner, 9 Mar 1999
+2 as
PLANTS OF TEXAS }
UNIVEROITY OF MICHIGAN SUMIILER £1640 COURSE IN TAXONOMIC BOTANY
meratuuren wr tee Uuivenanr? namacooe
Houstonia nigricans (Lam,) Fern,
Teewting, Teel oe)
: ‘ Way
Shanwix Nighcans Vr - gypsoph a.
zhas 15 ene 3 : ry
Vonstuse WV
BW ao)
JEFF DAVIS CO.: Davia Mountains,
Madera Canyon, common in rocky
areas under junipers; corolla
white.
26 July 1952
Graoy L. werster, no. 4501
Fic. 2. Hedyotis nigricans var. nigricans ( 1995); duy f Webster 4501 ( g
ila Unlat
sensu Turner
£eh
rT . . bor:
HGULCT Io pictured in Fig. 3.
nigricans var. avnsonh
J F
not be reiterated, this phenomenon being well documented by Powell and
Turner (1977) and Turner and Powell (1979).
It should be noted that Terrell seemingly ignored my published comment
(Turner 1993) that “This taxon [var. gypsophila] is represented by 45 or more
BRIT.ORG/SIDA 20(1)
N
i )
y a '
aa
it
K
a
qnief
Plybelegy a 491 1f Mae
HOLOTYPE OF Hedets Vigricang vv
tala PS.L Town ev
De 4a
(LL, PrN)
The Un Texas Hert
Hedyotis nigricans (Lam.) Fosberg
var, gypsophila B.L. Turner
Determined by B.L. Turner, 1995
To
Heo ou!
Plants of Nuevo Leon
10; Le co oF 2 Mexico
Se Shout a Hig CReS
L$ G.B.Hinton 18
Cc + TEX
)
‘a Ny Sante Rita 2370 m,
be —
\ ‘H. et al 11-vi-82
coaM
“a Sparse pine woods~~
gypsum hillside
ZAG. TAMPS
ie Colonies, Fl. white,
University of Texas « Austis (LL. TEX ae
aA
iii
(TEX); according to Turner (1995) populational units of this taxon
Coahuila, Nuevo Leon, San Luis Potosi and closely adj ‘
Fic. 3. Holotype of Hedyotis nigricans var. gypsophila
pai t J IIF
southernmost Tamaulipas, Mexico.
TURNER, BIOLOGICAL STATUS OF HEDYOTIS NIGRICANS VAR. GYPSOPHILA 219
collections at LL, TEX, mostly obtained from gypseous soils in the state of Nuevo
Leon.” This is not to say that occasional plants of var. gypsophila in north cen-
tral Mexico might not approach var. nigricans in this or that character, but such
“intermediates” do not occur in Texas. Indeed, var. nigricans can be found in
relatively close proximity to var. gypsophila in Nuevo Leon without clear inter-
mediates, the former occurring on calcareous soils (e.g., Hinton et al. 24351,
24575, 25940, etc., all TEX), the latter on gypseous soils (e.g., Hinton et al. 24373,
27183, etc., TEX). In fact, my initial taxonomic sense was to recognize var. gypso-
phila as specifically distinct, but such treatment would deny the occasional
intermediates, these mostly occurring in regions of near geographical contact.
It should be emphasized that var. gypsophila is distinguished from var.
nigricans by more than habit, leaf shape and calyx vestiture. The upper leaf
surfaces of the former are more nearly papillose; those of the latter having sur-
face-sculpturing more like that of the epidermis of a peanut. In short, there are
a syndrome of characters which provide for the recognition of both taxa, al-
though the occasional plant from this or that population might lack one or an-
other of the characters concerned.
Such identification problems are touched upon by numerous workers, in-
cluding my own (Turner et al. 1988). Leaf shape and habit vary considerably
among populations of var. nigricans in Texas. Thus, heavily browsed plants will
appear much branched from the base, and newly produced basal shoots will
possess somewhat broader leaves. No doubt such plants occasioned the attri-
bution of var. gypsophila to Texas, and probably those Mexican collections cited
by Terrell as occurring outside the distribution of var. gypsophila sensu the
present author.
Finally, Lcan’t help but note that I currently live among populations of H.
nigricans var. nigricans in Trans-Pecos, Texas, and have examined numerous
populations from throughout this region and nearly all consist of plants iden-
tifiable as var. nigricans as defined by both Terrell and myself, except as how
one might let the occasional over-browsed plant cloud one’s identification.
~~
ACKNOWLEDGMENTS
It isa pleasure to acknowledge the input of two reviewers, Guy Nesom and Piero
Delprete, whose comments greatly improved the present contribution.
REFERENCES
Powett, A.M.and B.L. Turner. 1977. Aspects of the plant biology of the gypsum outcrops of
the Chihuahuan Desert . Natl. Park Service Trans. & Proc., Ser. 3:315-325.
Terre, E.E. 2001. Taxonomy of Stenaria (Rubiaceae: Hedyotideae), a new genus including
Hedyotis nigricans. Sida 19:591-614
Turner, B.L. 1995. Taxonomic overview of Hedyotis nigricans (Rubiaceae) and closely allied
taxa. Phytologia 79:12-21
220 BRIT.ORG/SIDA 20(1)
Turner, B.L. and A.M Powelt. 1979. In: Goodin and Northington, eds. Deserts, gypsum and
endemism. International Center of Arid and Semiarid Land Studies. Texas Tech Univ.,
Lubbock.
Turner, B.L., M.W. Turner, and J. S Saisie 1988. Bourne analyses and new combi-
nations in Psil halodes (Asteraceae, Heliantheae).Phytologia
65:231-239.
i Za
DOCUMENTED CHROMOSOME NUMBERS 2002: 1.
CHROMOSOME NUMBER OF STENANTHIUM (LILIALES:
MELANTHIACEAE) AND ITS SIGNIFICANCE IN THE
TAXONOMY OF TRIBE MELANTHIEAE
Wendy B. Zomlefer Gerald L.Smith
Plant Biology Department = = ——~—-s &Biolo
University of Georgia High Point ney
2502 Plant Sciences Box 3591, University Station
Athens, GA 30602-7271, U.S.A. High Point, NC 27262, U.S.A.
ABSTRACT
Mitotic ch for S hiumd dS. lei hoides (2n = 20), reported here for
Es first time, mpaten the muamber Day pushy cecumentce for S.gramineum, the only other species in
e significance of chromosome num-
Bets in the Melanthiaceae, particularly as generic sy Seeeeyee ee in tribe Melanthieae, is discussed.
RESUMEN
El d itdticos de St thiumd y S. leimanthoides(2n = 20), reportados
por anes vez ad concuerdan con el numero previamente documentado para S. gramineum, la
unica otra esp te género recircumscrito recientemente basandose en datos moleculares. Se
ae te aqui el significado del numero de cromosomas en Melanthiaceae, particularmente como
sinapomorfias genéricas en la tribu Melanthieae.
The Melanthiaceae sensu the Angiosperm Phylogeny Group (APG 1998) com-
prise five tribes (11-16 genera; ca. 154-201+ spp.) of predominately woodland
and/or alpine poe herbs occurring mainly in the temperate to Arctic zones
of the Northern I | with one species of Schoenocaulon extending into
South America; Zomlefer et al. 2001). Of concern here are two genera of the
tribe Melanthieae, Stenanthium sl. and Zigadenus s.l. As traditionally and
broadly circumscribed, both genera form poorly defined complexes i
in Zomlefer 1997). Stenanthium s.l. has sometimes been divided into two gen-
era: a monotypic Stenanthium (S. gramineum) and Stenanthella (2-4 spp.);
Zigadenus has a more complex taxonomic history involving several proposed
segregates, although contemporary botanists have typically accepted only the
monotypic segregate Amianthium with the remaining ca. 19 species maintained
in Zigadenus s.l.
The circumscription and relationships of genera within the tribe
Melanthieae were recently evaluated using parsimony analyses of ITS (nuclear
ribosomal) and trnL-F (plastid) DNA sequence data (Zomlefer et al. 2001). Based
on the cladograms generated in this study, Zigadenus sl. is polyphyletic and
SIDA 20(1): 221 - 226. 2002
=
222 BRIT.ORG/SIDA 20(1)
Stenanthium s.l. is biphyletic, with Stenanthium ss. and Stenanthella embed-
ded within two different “Zigadenus” clades. In overview, the species of
Stenanthiumand Zigadenus as traditionally circumscribed, form five strongly
supported clades that correlate with distinctive geographical distribution, chro-
mosome number, and certain morphological characters. The Stenanthium clade,
comprising Stenanthiumgramineum (Stenanthiumss.), Zigadenus densus,and
Z. leimanthoides, is defined by the morphological synapomo! phies of a slen-
der (cylindrical) bulb and one obscure (or lacking) gland per tepal. However,
the chromosome number for this group, 2n = 20, has been documented only for
Stenanthium gramineum (Miller 1930; Saté 1942: as S. robustum, see Fernald
1946). Since chromosome number is a significant feature for genera of tribe
Melanthieae, 20 was predicted as the likely mitotic oo for the cere two spe-
cies now translerred to the recircumscribed St | t lensum
and S. leimanthoides (Zomlefer et al. 2001; Zomlefer & Judd 2002).
a)
MATERIALS AND METHODS
Several plants of Stenanthium densumand S. leimanthoides were collected dur-
ing April to June 2001 (see Table 1) and transplanted to pots maintained at the
Plant Biology Dept. Greenhouse Facility at the University of Georgia and the
Biology Dept. Greenhouse at High Point University. Dividing root tip cells were
prepared for examination according to general protocols outlined by Flory and
Smith (1980), Jones and Luchsinger (1986), and Smith (1984). Once the plants
were well-established, actively growing root tips were harvested at noon and
treated with 0.2 % colchicine for + hours, rinsed in distilled water, and then
fixed in Carnoy’s solution (3 ethanol: | acetic acid) overnight or longer. Follow-
ing this fixation, the roots were rinsed in distilled water, hydrolyzed in 10N
HCl at 43° C for 25 minutes, macerated with a dissecting spatula on a glass
microscope slide, and stained with 1% aceto-orcein. After application of a cover
slip, the slide was gently heated with an alcohol lamp, placed between blotters,
and subjected to additional pressure. Slides were mounted in euparal for future
reference. Well-spread metaphase chromosomes were traced under a Leica
DMLB Research Microscope witha camera lucida attachment. Herbarium speci-
men vouchers (Table 1) are deposited at GA.
RESULTS AND DISCUSSION
The mitotic chromosome number of 2n = 20 for Stenanthium,as circumscribed
by Zomlefer and Judd (2002), is verified for all species (Table 1, Figs. 1-3). Chro-
mosome number is a useful taxonomic character for genera within tribe
Melanthieae (see Fig. 4), especially the synapomorphic 2n numbers of 20 for
Stenanthium and 22 for Toxicoscordion. A probable base chromosome number
of x = 8 is often cited for the tribe Melanthieae (Sen 1975; Tamura 1995; Lowry et al.
Taste 1.Distinguishing features of Stenanthium densum and S.
and voucher specimens for the chromosome numbers reported in this study. Bas
223
leimanthoides (from Zomlefer 1997),
ed on reports of
morphologically intergrading populations in eae aMicD edie 1984), some authors (e.g,,
Sorrie & Weakley 2001) regard these taxa as co
and phenology, Small
oa and Oceanoros).
nspe cific
some
rlap i In habitat, morphol-
1903) considered these taxa as two separate monotypic genera
Taxon Distribution, habitat, Locality & collection Mitotic
distinguishing characters, number (voucher chromosome
& phenology location) number (2n)
Ster anthiim 7,
(Desr.) Tomer & Judd se VA toeLA FLORIDA. Franklin Co. 20
Flatwoods, bogs E side of FL 65, 1.1 mi
Plants delicate, racemes N of Ft. Gadsden
simple, flowers perfect Creek, 4 Apr 2001
Apr-Ma Zomlefer 782 (GA)
FLORIDA. Liberty Co.: 20
W side of FL 379, 0.2
mi N of Forest Service
Rd. 180,4 Apr 2001,
Zomlefer 783 (GA)
Stenanthium leimanthoides Wider range: NY to ne TX GEORGIA. Taylor Co.: 20
(A.Gray) Zomlefer & Judd — Pinelands, bogs, montane 4.3 miN of Butler,
marshes, swamps sa mi E of GA137,
Plants strikingly more ae) 2001, Zomlefer
robust, racemes 92 (GA)
compound, flowers cEORG Taylor Co.:
perfect + staminate 1 miS of Butler, 20
(plants andromonoecious) 1 Whitewater
May—June Creek, W side of GA
137,18 Jun 2001,
Zomlefer 797 (GA)
1987; Zomlefer 1997), and multiples of this number (or perhaps of 4) are preva-
lent in the group (Fig. 4): Anticlea (which includes Stenanthella),
Schoenocaulon, and sister groups Amianthium and Veratrum (including
Melanthium). Unfortunately, due to the small chromosome size of members of
the tribe (ca. 2.0-4.0 um in length), the few karyotype studies (e.g., Lee 1985) are
not detailed enough to infer a possible mechanism of chromosomal evolution,
although the chromosome numbers (Fig. 4) indicate the prevalence of polyp-
loidy and/or aneuploid variation. The functional outgroup for Melanthieae (see
Zomleter et al. 2001), Z. glaberrimus (Zigadenus ss.), hasan unpublished count
of 2n = 52, tentatively reported by Preece (1956). Probable base numbers for the
other immediate outgroup taxa of the tribe Melanthieae (the remaining four
tribes of the Melanthiaceae) are five and 15, respectively, for sister taxa Parideae
BRIT.ORG/SIDA 20(1)
1 a Y Z on
~, ~ aA a ~»
ay Vu, We
‘ mf »
J 1’ t J
“Pr -— »
I f the tk ies of Stenanthium, all 2n = 20.1.5. gramineum [modified from Saté
a Fig. 58]. 2.5. densum (Zomlefer 782). 3. 5, leimanthoides (Zomlefer 792).
2n= x=
Amianthium (1 sp] 32 8
Veratrum S.I. [20-45 spp] 16, 32, 8
64, 80, 96
Stenanthium (3 spp] 20 10
.
Anticlea [ca. 11 spp.] 32 8
Toxicoscordion (ca. 8 spp] 22 11
Schoenocaulon [24 spp.] 16 8
Zigadenus S.S. [1 sp] S27 i
Fic.4. Phyl icrelationshi 1 ck bers for the g f tribe Melanthi Thecadodram
kh a ITvT< dt I F Bf } L
inoilifiad oli Zomlefer et al (2001), i
ences suminatlzeet in uzomielet (1997). A
Zigadenus
I aia tain Gonaniiiiini Anticlea, Toxicoscordion, and
Ivnhyletic Zinnd, |. (Zomlefer & Judd 2002)
r Pere?
(Trilliaceae) and Xerophylleae, and 17 and possibly six, respectively, for sister
taxa Heloniadeae and Chionographideae (Lowry et al. 1987; Zomleter 1996, 1997).
Confirmation of the chromosome number for Z. glaberrimus (a possible
polyploid) and the reassessment of the base number for the tribe merit further
investigation (Zomlefer in prep.), especially in relation to the probable base
numbers of the other tribes. Available data, however, already demonstrate that
chromosome number strengthens support for the monophyly of Toxicoscordion
and Stenanthium, as recircumscribed. Chromosome counts for the superficially
similar plants in the Melanthieae validate chromosome number as a signifi-
_
225
cant character, useful for elucidating and predicting groupings and relation-
ships. Variation in chromosome number has also proven phylogenetically sig-
nificant in other monocot clades, such as Polygonatum/Polygonateae (Tamura
1993; Tamura et al. 1997a, b).
ACKNOWLEDGMENTS
Stephanie Bergamo, Richard Carter, Angus Gholson, Jr. David E. Giannasi, Mary
Jo Godt, and Elissa Totin enthusiastically assisted WBZ in the field; S. Bergamo
also shared her time and expertise with the laboratory work. We are grateful to
Andrew W. Tull (Plant Biology Dept. Greenhouse Facility, University of Geor-
gia) for maintaining plants collected in the field; Walter S. Judd and Douglas E.
Soltis for helpful comments on the manuscript; and William Carromero for the
Spanish translation of the abstract. Funds provided by the University of Geor-
gia Department of Plant Biology financed the collecting trips associated with
this project.
REFERENCES
ANGIOSPERM PHYLOGENY GROUP [APG]. 1998. An ordinal classification for the families of flower-
ing plants. Ann. Missouri Bot. Gard. 85:531-553.
FernaAto, M.L. 1946. ldentifications and reidentifications of North American Plants. Rhodora
48:137-162.
Flory, W.S. and G.L. SmitH. 1980. The chromosomes of Habranthus martinezii, H. robustus,
and their F, hybrid. Plant Life 36:54-62.
Lee, N.S.1985.A cytotaxonomic study of the Korean Veratrum species. Korean J. Pl. Taxon.
15:155-161.
Lowry, PP., P.GotpsLaTT, and H. Tose. 1987. Notes on the floral biology, cytology and embry-
ology of Campynemanthe (Liliales: Campynemataceae). Ann. Missouri Bot. Gard. 74:573-
576.
Jones, S.B.and A.E. LUCHSINGER. 1986. Preparing squashes of plant root tips. In: Plant system-
atics, 2nd ed., McGraw Hill, New York. Pp. 181-182.
McDearman, W.W. 1984. Systematics of Zigadenus densus and Z. leimanthoides and phylo-
genetic implications of breeding systems in the Veratreae. [Abstract]. ASB Bull. 31:71.
Miter, E.W. 1930. A preliminary note on the cytology of the Melanthioideae section of
the Liliaceae. Proc. Univ. Durham Philos. Soc. 8:267-271.
Preece, S.J. 1956. A cytotaxonomic study of the genus Zigadenus. Ph.D. dissertation. State
College of Washington, Pullman.
Sen, 8. 1975. Cytotaxonomy of Liliales. Feddes Repert. 86:255-305.
Sato, D. 1942. Karyotype alteration and phylogeny in Liliaceae and allied families. Jap. J.
Bot. 12:57-161.
Smait, J.K. 1903. Flora of the southeastern United States.”Published by the author,” New
York, New York. Pp. 250, 252.
226 BRIT.ORG/SIDA 20(1)
Smit, G.L. 1984. Revision of Piptocarpha. Ph.D. dissertation. University of Georgia, Athens.
Sorrie, B.A.,and A.S.Weaktey.2001.Coastal plain vascular plant endemics: phytogeographic
patterns. Castanea 66:50-82.
Tamura, M.N. 1993. Biosystematic studies on the genus Polygonatum (Liliaceae) Ill. Mor-
phology of staminal filaments and karyology of eleven Eurasian species. Bot. Jahrb.
Syst. 1 1a] =26.
Tamura, M.N. 1995. A karyological review of the orders Asparagales and Liliales
(Monocotyledonae). Feddes Repert. 106:83-111.
Tamura, M.N., M. Ocisu, and J.-M. Xu. 1997a. Heteropolygonatum, a new genus of the tribe
Polygonateae (Convallariaceae) from west China. Kew Bull. 52:949-956.
Tamura, M.N., ALE. ScHwarzeach, S. Kruse, and R. Reski. 1997b. Biosystematic studies on the
genus Polygonatum (Convallariaceae) IV. Molecular phylogenetic analysis based on
restriction site mapping of the chloroplast gene trnk. Feddes Repert. 108:159-168.
Zomterer, W.B. 1996. The Trilliaceae in the southeastern United States. Harvard Pap. Bot.
1(9):91-120.
Zomterer, W.B. 1997. The genera of Melanthiaceae in the southeastern United States. Har-
vard Pap. Bot. 2:133-177.
Zomterer, W.B. and W.S. Juop. 2002. Resurrection of segregates of the polyphyletic genus
Zigadenus 5.|.(Liliales: Melanthiaceae) and resulting new combinations. Novon 12:299-
308
Zomterer, W.B., N.H.Wituiams, W.M. Wuirten, and W.S. Juop. 2001.Generic circumscription and
relationships in the tribe Melanthieae (Liliales, Melanthiaceae), with emphasis on
Zigadenus: evidence from ITS and trnl-F sequence data. Amer. J. Bot. 88:1657-1669.
DOCUMENTED CHROMOSOME NUMBERS 2002: 2.
CHROMOSOME NUMBERS IN NORTH AMERICAN SPECIES
OF CERASTIUM (CARYOPHYLLACEAE)
J.K. Morton
Department of Biology
University of Waterloo
00, Ontario, Canada NAL 3G1
=
Water
ABSTRACT
Chromosome numbers are reported for 85 collections of Cerastium (Caryophyllaceae) from conti-
nental North America belonging to 22 taxa. First published reports include those for C. nutans var.
objectum Kearney & Peebles (2n = 34) and C. texanum Britton (2n = 36). First reports for material of
North American origin include C. pee Tolm. (2n = 108), C. brac hypodum ( Eng. ex Gray) BLL.
Robinson (2n = 34), C. dichotomum L. (2n = 38), C. diffusum Pers. (2n = 72), C. fischerianum Ser. (2n =
66 & 72), C.glomeratum Thuill. (2n = 72) and C. semidecandrum L. (2n = 36)
RESUMEN
Se citan los numeros cromosomaticos de eciccoiones de Cerastium (Caryophyllaceae) de Norte
América continental } 22t los primeros recuentos de C.
nutans var. objectum ane & Peebles ce 6 y C. texanum poe (2n = 36). amines recuentos
n materiales de origen norteamericano de C. bialynickii Tolm. (2n
Gray) B.L. Robinson (2n = 34), C. dichotomum L. (2n = 38), C a ee Pers, (2n =72), oe aes
Ser. (Qn = 66 & 72), C glomeratum Thuill. (2n = 72) y C. semidecandrum L. (2n = 36).
INTRODUCTION
The purpose of this paper is to put on record previously unpublished data on
chromosome numbers in the species of Cerastium, both native and introduced,
occurring in continental North America.
METHODS AND MATERIALS
Mitotic chromosomes were studied in root-tip preparations. Plants were raised
in the greenhouse, either from seed or from material transplanted from the wild.
Root-tips were taken from actively growing plants, prefixed for about one hour
in a saturated aqueous solution of paradichlorobenzene and fixed in 1:3 glacial
acetic acid and ethanol (absolute). They were hydrolyzed in N hydrochloric acid
at 60°C for l1O—15 minutes and squashed in aceto-carmine. In a few cases flower
buds were used for counts of meiotic chromosomes. The buds were fixed in
Carnoy’s Fluid (acetic acid l:chloroform l:ethanol 3). Developing anthers were
teased out and squashed in aceto-carmine or aceto-orcein. The cultivated plants
were grown to maturity and voucher specimens taken for comparison with
those collected in the wild. Voucher specimens are retained in the author's re-
SIDA 20(1): 227 — 231. 2002
228 BRIT.ORG/SIDA 20(1)
Taste 1. Chromosome numbers in Cerastium. Collection numbers prefixed with NA are J.K. Morton
or J.K. Morton and Joan M.Venn collections.
Taxon Locality & collection number Chromosome no. (2n)
C.alpinum L. Nfld: Plies n a nins - 72
subsp. alpinum St. Anthony NA52, VA527/5
N.W.T: Gt. Slav ive shake at 16059 72
Ont.: Lake Superior — Leadman 72
Is, NA4532
Ont.: Hudson Bay — SE of Winisk 108
Sims 2304
Que. James Bay — Gt.Whale Rivet 72
Forest & Brisson s.n.
C. alpinum L. Ont.: Hudson Bay — Cape 104+2
subsp. lanatum (Lam.) Cesati Henrietta Maria Winterhalder 3512
C.arcticum Lange Nun.: Baffin Is.- Fox 2 Dewline 108
Site, Parmalee & Seaborn 4052a
C. beeringianum Cham. & Schitdl. Alta.: Kananaskis to Coleman ve /_NA4679 72
Alta. Ja co at. Pk. — Wilcc 72
Morton s
Alaska: er Hwy. — Chicken, NA2122 72
Alaska: Alaska Pen.— Ugashik, 72
Talbot 103-25
B.C.: Alaska Con — Summit Lake, 72
NA14139& NA14140
Calif: Alpine Co. — ound Top Mt., /2
Stebbin
Colo.: F aan Mt. me P NA/334 72
Colo.; Mt. Evan 34 72
Nfld. Northern - seas NAI2405 72
Nfld. Cape St. George, NA5409 72
N.Mex.: Sante Fe Basin, NA7144 72
ae ae —rd to Flat 72
aes Spicer 1638
N.W.T.: Mackenzie Dist. — Canada 72
Tungsten Mine, Cody & Spicer 17702
N.W.T.: oo Dist.— June Lake, 72
Cody 1/
ee! Pen. — Ste. Anne des 72
Montes, NA2977
Que. Gaspe Pen. — Perce, NA4128 2
Wyo.: Park Co. — Snow Lake, Scott 9 72
Wyo.: Beartooth ve Beartooth - 72
NA4805
Yukon: — Hwy. — North Fork 72
Pass, NA
Yukon: A 7 Hwy. — Koidern, NA2075 72
& NAA
MORTON, CERASTIUM CHROMOSOME NUMBER
TaBle 1. cont.
Taxon
Locality & collection number
Chromosome no. (2n)
C. bialynickii Tolm.
C. brachypetalum Pers.
subsp. tauricum (Sprengel) Murb.
C. brachypodum (Eng. ex Gray)
B.L. Robinson
C. dichotomum L.
C. diffusum Pers.
C. dubium (Bast.) Guépin
C. fischerianum Ser.
C. fontanum Baumg. subsp.
vulgare (t fart Lillal 1) J! eutel
& Burdet
C. fontanum Baumg. subsp.
vulgare (a glandular variety)
C.glomeratum Thuill.
Yukon: Taylor Hwy. - on Alaska border,
NA? 163
Nun.: Devon |s., Bliss s.
Nun.: Axel Heiberg Is. he Zf2
Miss.: Lee Co. — Tupelo, NA 16924
N.C.: Wilkes Co. - rte 421, 36 mi W
of Winston Salem, NA4385
N.C: Rutherford Co.— US-74 between
Rutherfordton & Lake Lure, NA4479
N.C.: Claiborne Co.— nr Tazewell,
NA14988 & NA14991
Ariz.: Apache Co. — White Mts., NA7075
Ariz. Apache Co.— S of Show Low,
A16/11
Wash.: Walla Walla Co. — Walla Walla,
Old s.n
Calif: Mendocino Co. — Fort Bragg,
Morton s.n.
Idaho:Gem Co.- Emmett, Old s.n.
B.C. Queen Charlotte Is.— Tasu Mt.
at ca. 2000 ft., NA13402
B.c.: Queen Charlotte Is.— Tasu on
mine workings near sea level, NA13440
B.c.: Vancouver, NA 1350
B.C.: Queen Charlotte Is. a, NA 13403
N.Mex.:Catron Co.— Mogollon,
a 12546
nt.: Bruce Co. - Inverhuron Provincial
oe NA1493
B.C.: Stewart Hwy. - on W side of
summit, NA13668
Man.: Cowan — on rte 10 E of Swan
River, NA4549
Ont.: Lanark Co. -— nr Almonte, NA2722
Ont.: Bruce Pen. — Tobermory, NA3702
Calif: Mendocino Co.— Westport,
Morton s.n,
Ont.: Middlesex Co.— London,
Oldham 9097
Yukon: Dawson City, NA2172
72
108
108
90
88 alson = 44
TABLE 1. cont
BRIT.ORG/SIDA 20(1)
Taxon
Locality & collection number
Chromosome no. (2n)
C. nutans Raf.
C. nutans Raf. var.
objectum Kearney & Peebles
C. pumilum Curtis
C. semidecandrum L.
C. terrae-novae Fern. & Wieg.
C. texanum Britton
C.tomentosum L.
C. velutinum Raf.
C. velutinum Raf.“v
C. viride Heller (C. arvense L
maximum (Hollick & Britt)
ho)
subsp.
ar. villosissimum"
Alta.rte 16, Vermilion, NA3796
Alta.: WNW of Edmonton -
Lac St. Anne, Morton s.n.
N.Mex.: Grant Co. — NM-90
Emory Pass
Tenn.: Gatlin burg, NA4443
_W of
N.Mex.:Catron Co.— Gila Nat. Forest,
Willow Ck. campground, Pinkava
etal. 12636
B.C. Vancouver Is.— Saanichton, NA13548
Ind.: Huntington Co. — 1-69 at exit 73,
NA1S024
N.C. aaa a ~ Maxton nr
Laurinburg, NA4408
Va.: ean NA260?
B.C.:Victoria - McNeil Bay, NA13565
Ont.: Bruce Co
Ont.: Essex Co.
NA4478
Nfld.: Bonne Bay — Table Mt., NA12183
Ariz.: Pima Co. — Kitt Peak, NA11233
Ariz. Pima Co. — Nogales, Sycamore
Canyon, Churchill 70-604
8
— |Inverhuron, Morton Sols
Point Pelee, NA3540 &
Ont.: Bruce Pen.— N of Wiarton, Morton s.n
Mo.: Jefferson Co. — De Soto, NA19069
N.Y: New York City — Staten Is., NA3580
Ohio: Erie Co.— Catawba Is., Sandusky,
NA3660
Ont.: Essex Co. — Pelee Is., Morton s.n.
Ont.: Essex Co. — Point Pelee, NA4476
Pa. Chester Co. — Nottingham, NA35/la
Pa.: Chester Co. - Lees Bridge, NA37/0
Calif: Marin Co. — Pt. Reyes, NA19200
Calif: dee el Co.— Fort Bragg,
Stebbins
Calif: Sonoma ee
Stebbins s.n
Oreg.: Clatsop Co. -
Chambers s.n.
Oreg.: Clatsop Co.
Chambers 3160
Bodega Beach,
Cannon Beach,
Onion Pt.,
MORTON, CERASTIUM CHROMOSOME NUMBER 231
search collections and partial sets of duplicates have been distributed to WAT,
MICH, DAO and VDB. Collection numbers prefixed with NA are J.K. Morton or
J.K. Morton and Joan M. Venn collections (Table 1).
DISCUSSION
Many of these chromosome counts confirm those from previous workers. How-
ever, the following appear to be the first published counts for their respective
taxa: Cerastium nutans var. objectum - 2n = 34 and C. texanum - 2n = 36. That
for C. terrae-novae - 2n = 108, is the first definitive count for that species though
Bocher (1977) reported a chromosome number of “lower than 72 (perhaps 68).”
The following chromosome counts appear to be the first published counts
for North American material of these alien species: C. dichotomum - 2n = 38, C.
diffusum = 2 72, C.glomeratum -2n=72 and C. semidecandrum - 2n = 36.
Amongst native species that also occur in other regions, that for C.
bialynichii - 2n = 108, is new. Cerastium bialynickii has not previously been
reported from North America, having been originally described from Siberia,
where it is reported to have a chromosome number of 2n = 72. It isa very com-
pact and hispid relative of C. beeringianum with obtuse sepals.
Also our count for C fischerianum is the first from this continent. The pres-
ence of two different chromosome numbers in adjacent populations of this spe-
cies is unusual. The differences appear to be genuine, for the preparations were
carefully checked. The explanation may be that the population at sea level, with
2n = 66, was growing in a weedy situation on mine tailings. As C. fischerianum
frequently intergrades with C. beeringianum it is possible that these plants were
of hybrid origin.
The count for C. brachypodum (2n = 34) is the first for North America, but
Beaman et al. (1962) reported the same chromosome number in material from
Central America.
ACKNOWLEDGMENTS
lam indebted to the following persons for their assistance in supplying seeds
used in this research: L.C. Bliss, K.L. Chambers, WJ. Cody, R. Old, MJ. Oldham,
DJ. Pinkava, S.S. Talbot, E.K. Winterhalder and the late G.L. Stebbins.
REFERENCES
Beaman, J.H., D.C.D. De Jona, and W.P. Stoutamire. 1962. Chromosome studies in the alpine
and subalpine floras of Mexico and Guatemala. Amer. J. Bot. 49:41-50.
Bocuer, T.W. 1977. Cerastium alpinum and C. arcticum, a mature polyploid complex. Bot.
Notiser 130:305-309.
232 BRIT.ORG/SIDA 20(1)
BOOK REVIEW
SaLLy Wasowski. 2002. Gardening with Prairie Plants: How to Create Beautiful Native
Landscapes. Photography by Andy Wasowski. (ISBN 0-8166-3087-9, pbk.)
University of Minnesota Press. Ll] Third Avenue South, Suite 290, Minne-
apolis, MN 55401-2520. (Orders: www.upress.umn.edu, 612-627-1932, 612-
627-1980 fax). $29.95, 304 pp, 241 color photos, 335 figures, 81/2" « IL".
Most gé susie cass packs could be classified informative, educational and instructive. But rarely could
} ly “|
a ¢ be regarded asa page-t lening with Prairie Pl
ants was sucha pleasure
o read thal uthada hard time putting down. Wasowski wees her book by elaborating the on merits
of gardening with native prairie plants and remarks “a ‘ae landscape can also be eye-poppingly
gorgeous!” | would ¢
raracterize this book in a similar lig
Gardening with Prairie Plants is generally div ded ome main parts: L. Prairies and Prairie
Gardens, I. Design, Installation, and Maintenance, and HI. Plant Profiles. The first part defines a prai
rie, where prairies are located in North America and — prairies vary based on ge eas es The
ie: follows with many examples of home prairie gardens, which perhaps are described better as
testimonials to the advantages of native plant landscapes. Each one of these idimnonials more or
he homeowner installed and maintained their g eee with a color photo anda
figure of the landscape design to accompany the text. The next chapter, “Using Preserved and Repli-
le: s describe show t
cated Prairies as Models”, encourages the reader to determine the soil type for their proposed prairie
plot and locate a nearby prairie example to model their garden by. Lowland, mesic, upland and sand
prairies and savannas are all considered here
Putting your newfound knowledge of prairies is put to work in Part I, The chapter entitled,
“Designing Prairie Gardens’, gives advice on planning your garden for your lifestyle and yard. This
information is followed by clear and candid advice on garden installation, including preparing a
seedbed, tilling and enriching the soil, collecting native seed and planting transplants. As well, read-
ers will know exactly how to maintain their gardens with the authors advice on weed control, irri-
gation, prescribed burning and mowing. oe a safe controlled burn is considered in some
detail, wath an outline on the proper procedut
e author's labor shines through in IIL with the wealth of knowledge provided for each
species included in the plant profiles. This part is divided into four chapters, which deal first with
grasses, sedges and rushes, followec
by warm season forbs, then cool season forbs and finally trees
and shrubs. At the end of each chapter, a table is prov ided of all included species, which serves the
reader in quickly identifying species that would grow best in their garden habitat. For each plant
listed, information on common and scientific name, some diagnostic characters and habitat preter
ences are oe ided. Each specie ee dasa map of its geographic distribution and many species have
] +] 2
a lovely | y th scripti
say that the aeeerene sel photographs in this book are lovely would be a | ae
statement. Exquisite and inspiring are two terms that might do them justice. Hats off to Michael
Parkey and Andy Wasowski for their illustrations and
oo
see dhs, soni Their i is the
eye candy of this treatise. A short glossary of prairie terms, a list of gardening resources listed by
state and an index with common and scientific names ae this book
In sum, this book would be of interest to anyone interested in native landscape gardening—
homeowners, landscape architects, Master Gardeners and prairie plant enthusiasts. Those particu-
larly interested in prairie restoration should not be without a copy of this book. It is pure pleasure to
read.—Amy Trauth Nare, Botanical Research Institute of Texas, Fort Worth, EX, 76102-4000, U.S.A.,
amy trauth@yahoo.com
SIDA 20(1): 232. 2002
USOS DE MELASTOMATACEAE EN EL ECUADOR
*Alina Freire Fierro Diana Fernandez
Missouri Botanical Garden Herbario Nacional del Ecuador QCNE
0 17-21-1787
St. Louis, MO 63166-0799, U.S.A. Quito, ECUADOR
Email: alina.freire@mobot.org Universidad Central del Ecuador
“Autora para correspondencia Escuela de Biologia
uito, ECUADOR
Email: fernandezdiana@yahoo.com
Catalina Quintana
Herbario QCA
Pontificia Universidad Catélica del Ecuador
17-12-584, Quito, ECUADOR
E-mail: cquintana@puceuio.puce.edu.ec
RESUMEN
AS(\) bd ] past p
* Lf
Melastomataceae es una familia con los tropicos y
I L
subtropicos, deep epnens en América del Sur. En Ecuador se han registrado +3 géneros y 553
os de éstos, 24 géneros y 124 reportan alguna utilidad para el ser humano.
4
Segun registros de los herbarios QCNE y QCA y la literatura, las es
Meteiormaricsie son utilizadas principalmente para tratar heridas, eae de culebras y para
construcciones. De las 124 especies utiles, 27 (21,8%) son alimenticias, especialmente debido a los
frutos dulces y a los tallos agrios ieee ee calmar la sed; 32 (25,8 ‘ e son uilizaces para tablas y
lici 8 (6,4%) son usados pol l iviar algunas
construccion; 59 (47.6%) son
enfermedades fisicas ra ituales, solamente 4 (3,2%) son ornamentales y 50 (40, 3%) tienen otros
usos, tales I cobas, encofrados, forraje para animales, lena, tinciones, y
como cercas vivas.
ABSTRACT
il | inl the tropics, especis ally in
Melastomataceae | isa family with about 4500
South America. There are 43 genera and 553 ao oun for Fouader Based on the revision of
herbarium specimens housed at QCNE, QCA and the literature, 24 genera and 124 species are regis-
tered as useful for humans. Of these, 27 (21.8%) are used as food due to the sweetish fruits and acidic
stems. used against thirst; 32 (25.8 %) are used for construction; 59 (47.6%) are medicinal; 8 (6.4%) are
used by Shamans for treating fisical and spiritual ailments; only 4 (3.2%) are ornamentals; and 50
(40.3%) have various uses, such us for tools handles, brooms, fuelwood, charcoal, dyes, and for living
fences.
INTRODUCTION
El Ecuador incluye 553 especies (Renner et al. 1999) de las +500 existentes en el
mundo (Mabberley 1997). Registros de usos de Melastomataceae, junto con otras
familias de plantas se han concentrado principalmente a areas 0a grupos étnicos
determinados del pais. Asi, Ceron (1996) registra los usos de plantas del Parque
SIDA 20(1):; 233 - ~ 260. 2002
234 BRIT.ORG/SIDA 20(1)
Nacional Machalilla (prov. de Manabi), Kvist y Holm-Nielsen (1987), basados
en sus estudios en la comunidad indigena Cayapa (prov. de Esmeraldas) y en la
literatura, describen rituales curativos y usos de plantas por comunidades
indigenas de las tierras bajas. Para la region Andina, Kothari (1993) lista las
plantas usadas por pobladores de La Esperanza, comunidad indigena Zuleta
(prov. de Imbabura), Ceron (1996) reporta los usos de plantas de la Reserva
Geobotanica Pululahua (prov. de Pichincha), Rios 1993) lista las plantas de una
comunidad mestiza del noroccidente de Pichincha, Elleman (1991) publica los
usos maderables realizados por la comunidad Saraguro (prov. de Loja), y Cueva
(1997) y Van den Eynden (1997) listan las plantas alimenticias de la provincia
de Loja. Muchos otros trabajos etnobotanicos han sido realizados en
comunidades indigenas de la Amazonia ecuatoriana, entre ellos podemos citar
a Alarcon (1988), Ceron (1996), Ceron y Montalvo (1998), Lescure et al. (1987) y
Vickers y Plowman (1984). A una mayor escala geografica, Schultes y Raffaul
(1990) presentan usos etnobotanicos de Melastomataceae y demas familias de
angiospermas presentes en la Region Amazonica Nororiental.
la ausencia de publicaciones sobre los usos de la familia a nivel del Ecua-
dor nos motivo a la recopilacion de informacion sobre nombres vernaculares,
usos actuales y potenciales de especies de Melastomataceae a partir de las
etiquetas de especimenes depositados en los dos herbarios con mayor numero
de colecciones del pais (QCNE y QCA).
METODOLOGIA
La informacion sobre usos de plantas se la obtuvoa partir de los especimenes con
informacion etnobotanica depositados en el Herbario Nacional del Ecuador
(QCNE) y herbario de la Pontificia Universidad Catolica del Ecuador (QCA),
ambos en Quito. De cada especimen se recopil6 el nombre de la especie, el colector
y numero, el nombre vernacular e idioma (si presentes), y los usos. La
determinacion de los especimenes botanicos depositados en ambos herbarios fue
realizada principalmente por Elvira Cotton (actualmente en el Herbario AAU,
Aarhus, Dinamarca), especialistaen Melastomataceae ecuatorianas quien residio
en Ecuador por varios anos. Informacion publicada en la literatura, y aquella
producto de nuestra experiencia de campo también fueron incluidos en el presente
trabajo. Datos sobre el numero total de especies por género fueron tomados de
Renner (1993) y Mabberley (1997), e informacion sobre el numero de especies por
género y sudistribucion en el Ecuador se obtuvo de Wurdack (1980), Renner et al.
(1999) y Freire-Fierro (en prensa). E] material examinado esta listado en el Anexo,
enel cual se incluyen a los autores de los nombres cientificos
Aciotis D. Don
Este género herbaceo incluye 13 especies, y en Ecuador se registran cuatro
especies que se distribuyen principal en las faldas occidentales de la Cor-
FREIRE-FIERRO ET AL., MELASTOMATACEAE EN EL ECUADOR 235
dillera Occidental y en las faldas orientales de la Cordillera Oriental. Aciotis
rubricaulis, una especie distribuida a ambos lados de la cordillera es utilizada
para curar picaduras de culebras. Similar uso se registra para Aciotis ornata,
especie que en Ecuador se distribuye tnicamente en las tierras bajas al
noroccidente. Ceron (1994) registra el uso de Aciotis leoyara (probablemente
refiriéndose a A. levyana Cong,, un sindnimo de A. rubricaulis [Freire-Fierro
(en prensa)|) como antiofidica.
Arthrostemma Pavon ex D. Don
Este género arbustivo presenta siete especies, y solamente Arthrostemma ciliatum
esta registrada en el Ecuador, en donde se distribuye en la region costera, andina y
amazonica. Esta especie es utilizada para curar la diarrea en ninos y para curar
Los tallos suculentos son un tanto acidicos y son generalmente
utilizadosp para calmar la sed. Esta especie también podria ser utilizada como orna-
mental debido a su habito trepador y flores de un vistoso color rosado.
Axinaea Ruiz y Pavon
Este género arboreo o arbustivo consta de 30 especies, 10 de las cuales ocurren
en los Andes del Ecuador. De éstas, Axinaea affinis, A. macrophylla, A. quitensis,
y A. sclerophylla son utilizadas principalmente para la construccion y como
combustible en calidad de lena y carbon. Un potencial uso del género seria como
ornamental ya que las flores son bastante vistosas.
I de Axinaea son consideradas maderables, A. macrophylla por
Ceron (1994) y A. sclerophylla (como A. schlerophylla) por Elleman (1991).
Bellucia Neck. ex Raf.
Este género arboreo consta de siete especies, y solamente Bellucia pentamera
esta registrada en el Ecuador, en donde se distribuye en la Costa, Andes y
Amazonia. La madera de esta especie es utilizada para la construccion. Los
frutos, que nacen directamente de los troncos, pueden alcanzar hasta 10 cm de
diametro, son comestibles y de sabor ligeramente dulce.
El fruto maduro de Bellucia pentamera (como B. weberbaueri Cogn.) es
comestible para los Kichwas (como Quichuas) de la Amazonia Ecuatoriana
(Alarcon 1988, Lescure et al. 1987) y por colonos mestizos de la comunidad
Ahuano (provincia del Napo) (Rios y Caballero 1997). El nombre Bellucia
axinthera (probablemente refiriéndose a B. axinanthera Triana, un sinonimo
de B. pentamera) es listado como alimenticio por Cer6on (1994).
Blakea P. Browne
Este género arbustivo incluye 100 especies, 31 de las cuales ocurren en el Ecuador,
en donde se distribuyen en la Costa, Region Andina y Amazonia. Blakea bracteata,
B. glandulosa, B. involvens, B. punctulata, B. rosea, y B. subconnata son utilizadas
para tratar heridas, picaduras de culebras, y prevenir ampollas. También son
utilizadas para curar el “enduendado” [hechizo] y para la construcci6n de casas.
BRIT.ORG/SIDA 20(1
=
Especies de este género también tienen gran potencial ornamental, especialmente
idas debidoa que sus flores son bastante llamativas,
para reforestar parques y
puesto que existe un contraste entre los pétalos, generalmente rosados, con los
estambres y estaminoides que son de un amarillo dorado.
espécimen Vickers y Plowman 255 (QCA) listado como “Blakea atf.
ciliata Mgf. or B. rosea” por Vickers y Plowman (1984), y como B. rosea por
Lescure et al. (1987), es conocido por los Siona-Secoyas como “Yayurua” y es
utilizado para tratar quemaduras. B. sawadae es utilizada por los Cofanes como
medicinal (Cerén 1994). B. portentosa Wurdack es conocida por los Huaoranis
como “Namuntaqui” y tiene frutos que son consumidos por aves pequenas
(Ceron y Montalvo 1998).
EF
—
Brachyotum (A, DC.) Triana
Brachyotumes un género arbustivo de 58 especies, 21 de las cuales ocurren ene
Ecuador, en donde se distribuyen unicamente en los Andes. Cinco especies,
Brachyotum alpinum, B. azuayense, B. ledifolium, B. rostratum y B. trichocalyx
son utilizadas para curar el catarro en los pollos, tenir, construir, asi como para
lena y escobas. Este género puede ser también utilizado como cercas vivas.
Algunas especies de Brachyotum tienen potencial como ornamental ya que sus
flores presentan generalmente calices de un color rojo vivo y corolas morado
oscuras (e.g. Brachyotum andreanum Cogn.) o amarillas (e.g. B. ledifolium); sin
embargo, las plantas tendrian que ser sembradas especialmente en lugares frios
ya que el género es tipicamente de paramo.
Brachyotum ledifolium es utilizada medicinalmente por los Kichwas
andinos (Cerén 1994) y como combustible y para escobas en la zona del
Pululahua (Ceron 1996).
Clidemia D. Don
Este género arbustivo incluye 120 species, 38 de las cuales ocurren en el Ecua-
dor, en donde se distribuyen tanto en la Costa, Region Andina como Amazonia.
Las siguientes 14 especies son utilizadas: Clidemia acostae, C. allardii, C.
capitellata,C.crenulata, C.dentata, C.dimorphica, C. epiphytica, C. heterophylla,
C. hirta, C.ostrina, C. purpurea, C. septuplinervia, C. sprucei y C. variifolia. Estas
especies son utilizadas para tratar heridas, ulceras, diarrea con sangre, asi como
para cicatrizar, calmar el mareo, y bajar la fiebre. Ademas, una de estas especies
es utilizada por los shamanes para “asustar a los espiritus”, especialmente los
“malos” espiritus [entidades sobrenaturales que pueden hacer dano|. Debido a
que las bayas son agri-dulces, éstas son utilizadas para calmar la sed. Muchas
especies de Clidemia presentan formicarios en la base de las hojas o en los tallos
y tienen asociacion con hormigas, las cuales utililizan estas estructuras como
sus nidos. Los formicarios tienen un sabor acido debido a esta situacion y son
consumidos como alimento por algunos grupos étnicos de la Amazonia.
Clidemia dentata, conocida como “Paitzi”, es utilizada por los Kichwas de
FREIRE-FIERRO ET AL., MELASTOMATACEAE EN EL ECUADOR 237
la Amazonia ecuatoriana para bajar la fiebre (como Clidemia sp., Alarcon 1988,
Lescure et al. 1987). Los frutos de C. dimorphica y C. sprucei son comestibles
para los indigenas Huaoranis (Lescure et al. 1987), y es también conocida como
medicinal por los Cofanes (Ceron 1994). Los indigenas Kichwas y mestizos de
la comunidad Ahuano (provincia del Napo), también consumen los frutos de
una especie de Clidemia no determinada (Rios y Caballero 1997). Clidemia
heterophylla es listada como medicinal por Ceron (1994).
Conostegia D. Don
Conostegia es un género arbustivo de 43 especies, 16 de las cuales ocurren en
Ecuador, en donde se distribuyen principalmente en la Costa, aunque también
se han registrado especies para la Region Andina y la Amazonia. Conostegia
attenuata, C. centronioides y C. dentata son utilizadas para curar enfermedades
[no especificadas en las etiquetas].
Diplarpea Triana
Este género herbaceo 0 arbustivo tiene solamente una especie, la cual esta
presente en el Ecuador. Diplarpea paleacea ocurre en los flancos de la cordi-
llera norte andina y es utilizada para tratar dolores de estomago.
Graffenrieda DC.
Este género arbustivo o arboreo consta de 44 species, 1] de las cuales existen en
Ecuador, y estan distribuidas principalmente en la region andina. Dos especies
del género, G. aff. intermedia y Grafferieda sp. son utilizadas en la construccion
y como lena por los Huaoranis de Quehueiri-Ono (provincia del Napo) (Ceron
y Montalvo 1998).
Henriettella Naudin
Este género arbustivo o arboreo incluye 51 species, siete de las cuales se
encuentran en Ecuador, tanto en las tierras bajas comoen los flancos occidental
y oriental de los Andes. De acuerdo a Rios y Caballero (1997), el fruto de una
especie no identificada es conocidac comestible por los pobladores Kichwas
y mestizos de la comunidad de Ahuano (provincia del Napo).
Leandra Raddi
Este género arbustivo incluye 175 especies, 19 de las cuales ocurren en el Ecua-
dor, en donde se distribuyen principalmente en la Amazonia, aunque también
hay registros de la Region Andina y de la Costa. Leandra caquetana, L.
caquetensis, L. granatensis, L. secunda y L. subseriata, son utilizadas como co-
mestibles por sus frutos, y para curar ulceraciones de la piel. Los pequenios frutos
carnosos son comidos por las aves.
Loreya DC.
Loreya es un género arbustivo que comprende 14 especies. De las cuatro especies
ecuatorianas, una se distribuye en la Costa y las tres restantes en la Amazonia.
De acuerdoa Lescure et al. (1987), los frutos de Lore ya spruceana (bajo L. collatata
238 BRIT.ORG/SIDA 20(1)
Wurdack) son comestibles. Estos frutos también son comidos por tapires y
puercos salvajes (-bajo L. collatata -Schultes y Raffaut 1990).
Maieta Aubl.
Este género arbustivo incluye tnicamente dos especies, y ambas se distribuyen
en la Amazonia ecuatoriana. De éstas, Maieta guidanensis es utilizada para tratar
dolores estomacales, y de cabeza, mareos, y para cicatrizar heridas. Esta especie
también tiene frutos carnosos que son comidos por las aves.
Meriania Sw.
Este género arbustivo y arboreo consta de 74 especies, 27 de las cuales ocurren
en el Ecuador. Una sola especie esta registrada para la Costa y las restantes se
distribuyen en los Andes. Los troncos de Meridania tomentosa son utilizados para
la elaboracion de carbon (Cer6n 1996). Este género también tendria gran
potencial para forestacion de parques y avenidas por las vistosas flores. Aunque
este ultimo uso no se da en el Ecuador, en Colombia especies del género son
utilizadas como arboles ornamentales de avenidas (D. Neill, com. pers. 2001).
La madera del “Quihle o Quishle blanco,” una especie indeterminada de
Meriania, es utilizada en la construccién por los Saraguros (Elleman 1991).
Miconia Ruiz y Pav.
Miconia es el género mas grande de la familia, incluye aproximadamente 1000
especies, 240 de las cuales estan registradas para el Ecuador. La mayoria de
especies son andinas, aunque también ocurren en la Costa y en la Amazonia.
La alta diversidad de especies es acompanada por la diversidad de usos de
especies de Miconia en el Ecuador. En los herbarios revisados se tienen
registradas como medicinales a las siguientes especies: Miconia affinis, M.
barbinervis, M. bracteolata, M. bubalina, M.crocea, M. erioclada, M. goniostigma,
M. gracilis, M. lugonis, M. nervosa, M. paleacea, M. papillosa, M. procumbens, M.
theaezans, M. trinervia, M. triplinervis, M. venulosa, y M. zubenetana, las que
son utilizadas como vermifugo, para estimular las dilataciones durante el parto,
curar dolores de garganta y del cuello, tratar la tuberculosis, dolores de muelas,
infecciones bucales, micosis, sarnas y para curar diarrea de ninos recién nacidos,
e inclusive pueden ser utilizadas como antidoto de picaduras de la hormiga
“Conga” (Paraponera spp.) (ver Ceron 1994, 1996, Ceron y Montalvo 1998:
Schultes y Raffauf 1990). Miconia bubalina (como Miconia sp.) es utilizada por
los Kichwas de la amazonia para tratar pequenos abcesos (Alarcon 1988; Lescure
et al. 1987).
La madera de Miconia aeruginosa, M. astroplacama, M. dureoides, M.
brachycalyx, M. bracteolata, M. bubalina, M. caelata, M. corymbiformis, M. crocea,
M. decurrens, M. floribunda, M. lutescens, M. matthaei, M. napoana, M. obscura,
M. ochracea, M. paleacea, M. papillosa, M. pustulata, M. salicifolia, M. splendens,
M. subspicata, M. theaezans, y M. tinifolia, es utilizada principalmente para
construccion, para fabricacion de herramientas, produccién de carb6n y como
—
FREIRE-FIERRO ET AL., MELASTOMATACEAE EN EL ECUADOR 239
combustible (Ver Ceron 1994, 1996; Cerén y Montalvo 1998; Cueva 1997;
Elleman 1991; Van den Eynden 1997; Lescure et al. 1987; Schultes y Raffauf 1990;
Vickers y Plowman 1984). De acuerdo a Ceron y Montalvo (1998), otra especie
utilizada en la construccion y lena es Miconia ternifolia Triana, sin embargo,
este nombre no esta registrado en el Catalogo de Plantas Vasculares del Ecua-
dor (Renner et al. 1999) ni en la base de datos TROPICOS (Missouri Botanical
Garden 2002).
Si bien los frutos bacaceos de Miconia alcanzan hasta 1 cm de didmetro,
algunos son dulces y comestibles, tal es el caso por ejemplo de Miconia
asclepiadea, M. aspergillaris, M. crocea, M. latifolia, M. ledifolia, M. lutescens, M.
cf. splendens y M. theaezans (Ver Ceron 1994, 1996; Cerén y Montalvo 1998;
Cueva 1997; Van den Eynden 1997).
Los frutos de Miconia bubalina, M. calvescens, M. cazaletii, M. centrodesma,
M. cercophora, M. decurrens, M. nervosa y M. oraria son comidos por aves (Ver
Ceron y Montalvo 1998).
Algunas especies tendrian también potencial ornamental, por ejemplo,
Miconia crocea tiene paniculas de cerca de 20 cm de longitud con abundantes
flores con pétalos blancos y estambres amarillos. Otra especie con potencial
ornamental podria ser Miconia salicifolia, la cual se distribuye en la Region
Andina y presenta arbustos con apariencia de candelabros debido a la densa y
erecta disposicion de los tallos y a la posicion adpresa de las hojas, las cuales
muestran la densa pubescencia ferruginosa-amarilla de su envés.
Los indigenas Kichwas de la Amazonia ecuatoriana utilizan una especie
no determinada de Miconia conocida como “Chiquitapayansu” para tratar
granos infectados (Alarcon 1988). La madera de M. astroplacama es utilizada
por los Siona-Secoyas para pilares en construccion de casas (Lescure et al. 1987;
Vickers y Plowman 1984). Los frutos de otra especie no identificada de Miconia,
y conocida por los Kichwas como “Payanzo”, son comestibles (Lescure et al. 1987).
Los frutos de M. ledifolia, M. lutescens y dos especies no determinadas son co-
mestibles en la provincia de Loja (Cueva 1997; Van den Eynden 1997). Ceron
(1994) registra el uso de Miconia crocea en la medicina y M. palacea, M. pustulata
y M. theaezans en la construccion. Elleman (1991) lista como maderables para
los Saraguros a M. caelata, M. corymbiformis, M. obscura, M. cf. tinifolia y dos
especies no identificadas.
Monochaetum (DC.) Naudin
Este género arbustivo incluye 45 especies, y solamente tres estan registradas
para el Ecuador, en donde se distribuyen en la region andina. Monochaetum
hartwegianum es utilizado como ornamental y M. lineatum para lena. Esta
altima especie es conocida como maderable por los Saraguros (Elleman (1991).
Monolena Iriana
Monolenaes un género herbaceo de 10 especies, y Unicamente M. primulaeflora
240 BRIT.ORG/SIDA 20(1)
esta registrada tanto en la Costa y Region Andina como en la Amazonia
ecuatoriana. Esta especie es utilizada para tratar la gripe, dolores de estomago,
parasitos intestinales, conjuntivitis, siendo ademas usada como antiolidica.
Debido al sabor acido de los peciolos suculentos, esta especie también es utilizada
para calmar la sed. Monolena pee podria ser explotada como planta
de interior debido a que las hojas acaulescentes tienen peciolos rojizos y
suculentos, laminas variegadas y flores bastante llamativas.
Kvist y Holm-Nielsen (1987) mencionan que esta especie es usada por los
Coaiqueres como vermifuga y que es fuertemente hemeética, y Cerdn (1994) la
listacomo laxante. Schultes y Raffauf (1990) registran el uso de lamisma especie
para tratamientos de mordeduras de serpientes en la provincia de Pastaza.
Mouriri Aubl.
Este género arboreo consta de 81 especies, 10 de las cuales se han registrado para
el Ecuador, en donde ocurren principalmente en la Amazonia. De las tres
especies registradas como utiles, M. oligantha es utilizada como anticonceptivo
y esterilizante; y los frutos de Mouriri grandiflora y M. nigra son comestibles.
Su uso como comestible esta registrado también por Cer6on (1994).
x
)
Ossaea AP. DC.
Ossded es un genero arbustivo con 91 especies, 20 de las cuales ocurren tanto en
la Costa y Amazonia ecuatoriana como en los Andes. El Ecuador tiene cinco
especies nee bracteata, O. laxivenula, O. micrantha, O. quadrisulca, y
O. robusta. Los herbarios estudiados registran que estas especies han sido
utilizadas para bajar la fiebre y tratar la epilepsia. Los frutos estan registrados
Nn
como comestibles.
Una especie identificada como Ossaea sp.[muestra Alarcon 34] y conocida
por los Kichwas de la Amazonia ecuatoriana como “Hagapania/Payanzo” es
utilizada para tratar granos infectados y sarna (Alarcon 1988; Lescure et al. 1987).
En realidad, este espécimen fue mas tarde identificado como Miconia affinis.
Tessmanianthus Marker.
Este género arboreo neotropical incluye siete especies, de las cuales dos estan
registradas para la Amazonia ecuatoriana. Aunque no se estudio material de
herbario de este género, Ceron y Montalvo (1998) indican que el fuste de T.
heterostemon Marker. es utilizado por los Huaoranis de Quehueri-Ono para la
construccion y para lena.
et
Tibouchina Aubl.
Este género arbustivo incluye 240 especies. Solamente I+ especies estan
registradas para el Ecuador, las cuales se distribuyen principalmente en los
Andes, aunque también existen especies en la Costa y Amazonia. Siete especies,
Tibouchina gleasoniana, I. grossa, T. laxa, [.lepidota, T. longifolia, T. mollis, y T
oroensis son utilizadas para tratar cataratas, extraer sarcoma de los ojos, tratar
dolor de muelas, para forraje de los animales, lena o uso ornamental.
FREIRE-FIERRO ET AL., MELASTOMATACEAE EN EL ECUADOR 241
Tibouchina lepidota es muy comutn en la region andina nor-oriental, y
podria ser utilizada no solo como bustible, sino también como ornamental.
Las inflorescencias estan compuestas de grandes flores con colores blancos,
rosados y fucsias y con estambres amarillos vistosos. Con la excepcion de
Tibouchina oroensis, las otras especies de Tibouchina ornamentales observadas
en el Ecuador son importadas del Brasil (D. Neill, com. pers. 2001).
El uso como combustible de Tibouchina mollis esta registrado en Cer6on
(1994),
Tococa Aubl.
Tococa comprende cerca de 50 especies de plantas arbustivas usualmente
mirmecofilas. Ecuador posee 12 especies distribuidas principalmente en las
tierras bajas de la Amazonia, existiendo menos en la Regidn Andina y la Costa.
Tococa guianensis es utilizada para tratar los granos infectados de la piel. Los
especimenes examinados también registran que los frutos de T. spadiciflora y
de T. symphandra son comidos por las aves. Aunque las etiquetas de los
especimenes herborizados no lo registren, es posible que las pequenas y vistosas
bayas de otras especies de Tococa sean también comidas por aves.
Tococa guianensis (-erroneamente escrita “Cococa”) es utilizada por los
Kichwas de la Amazonia ecuatoriana para curar granos infectados y es conocida
como “Payanshu’ (Alarcon 1988), o como “Jatunpayanshu’ (Lescure et al. 1987).
Los frutos maduros de esta misma especie, bajo el sindnimo Tococa chuivensis
Wurdack var. ecuadorensis Wurdack, son comidos por personas y por aves
(Ceron y Montalvo 1998).
Triolena Naudin
Triolena es un género herbaceo de 22 especies, 11 de las cuales se encuentran en
Ecuador, especialmente en las tierras bajas tanto de la Costa como de la
Amazonia. Triolena hirsuta, T. obliqua, T. pluvialis, T. pedemontana, y T.
pustulata son utilizadas en Ecuador como medicinales. Estas especies son
utilizadas principalmente para tratar picaduras de culebras, reducir las
subsiguientes hinchazones, y para tratar tuberculosis y tos.
Los Cayapas y Coaiqueres utilizan Triolena para tratar mordeduras de
culebras, y los Siona-Secoyas usan I. pluvialis para tratar dolores de muelas
(Kvist y Holm-Nielsen 1987; Vickers y Plowman 1984). Similar uso es dado para
esta especie por los Kichwas de la Amazonia ecuatoriana (Alarcon 1988; Lescure
et al. 1987) y por los Cofanes (Schultes y Raffauf 1990). Cerdén (1994) también
registra el uso medicinal de esta especie.
CONCLUSIONES Y DISCUSION
El presente trabajo se baso primordialmente en la compilacion de la informacion
de las etiquetas de dos herbarios ecuatorianos. Contrariamente a la opinion
generalizada de que Melastomataceae no es una familia etnobotanicamente
242 BRIT.ORG/SIDA 20(1)
Tasta 1.Porcentajes de especies por tipos de usos de las 124 especies de Melastomataceae utilizadas
en Ecuador. Nota: El numero total de especies (177) aparece mayor que el real (124) debido a que
46 especies tienen al menos dos usos.
Usos Numero spp. utiles Porcentaje
Medicina 59 47,6
Otros 50 40,3
Construccion 32 25,8
Alimenticio 2/ 21,8
Magico Religioso 8 64
Ornamental 4 3,2
importante (e.g. Schultes y Raffauf 1990), al menos una de cada cinco especies
que ocurre en Ecuador tiene alguna utilidad.
Mas de la mitad de los géneros registrados para el Ecuador son utiles, sin
embargo, a pesar de que algunos géneros estan presentes en el pais y de ser
citados en la literatura como utiles (e.g. Adelobotrys DC. y Centronia D. Don
[Schultes y Raffauf 1990)), no registran uso alguno en Ecuador,
La mayoria de especies de Melastomataceae (Tabla 1) es utilizada
medicinalmente, en segundo lugar esta el uso para lena, carbon, alimentos de
animales, tinciones, etc. [Otros], una de cada cuatro especies utiles de la familia
es usada en la construccion, menos del 22% de especies es utilizado en la
alimentacion, muy pocas especies son utilizadas en rituales magico religiosos y
solamente cuatro de las 124 especies es utilizada como ornamental. Seguin
Schultes y Raffauf (1990), sin embargo, la unica importancia econdomica que la
familia tiene es por las especies ornamentales. Probablemente esta discrepancia
se deba a que si bien en Ecuador Melastomataceae no es muy utilizada a nivel
ornamental, este uso es comtin en otros paises.
Debido a la diversidad de especies, Miconia es el género que mayor numero
de especies utiles presenta, principalmente en la construccion, medicina y usos
varios. Clidemia es el segundo género mas importante (14 especies utiles) y la
mayoria de sus especies es utilizada en la medicina, aunque C. crenulata es
también utilizada en ritos magico religiosos. En tercer lugar estan Blakea y
Tibouchina, generos con 7 especies utiles cada uno. Los otros géneros tienen de
una a cinco especies utiles. A nivel de especies, Miconia theaezans, Blakea
subconnata, Miconia bracteolata, Miconia bubalina, Miconia crocea, y Ossaea
micrantha registraron el mayor numero de usos, tanto en la alimentacion, como
en medicina y construccion.
A nivel de uso, Arthrostema y Monolend son los mas utilizados para calmar
la sed debido a sus habitos suculentos y de sabor acido; y Bellucia, Henriettella,
Mouriri, y Ossadea son predominantemente usados como alimenticios debido a
que sus frutos son relativamente grandes y bastante dulces. Las especies de
FREIRE-FIERRO ET AL., MELASTOMATACEAE EN EL ECUADOR 243
Axinaea, Bellucia, Graffenrieda, y Tessmanianthus son las mas utilizadas en la
construccion, especialmente debido al habito arboreo de las mismas. Mas del
60% de las especies de Aciotis, Arthrostema, Blakea, Clidemia, Conostegia,
Diplarpea, Maieta, Monolena, Ossaea, y Triolena, son utilizadas
medicinalmente. A nivel magico religioso, Aciotis es el mas utilizado y menos
del 50% de las especies de Blakea, Clidemia, Conostegia, Miconia, y Ossaea es
también usado para este fin; ninguno de los otros géneros registrados para Ec-
uador es utilizado para el mismo proposito. De los 24 géneros usados en Ecua-
dor, unicamente Monochaetum (M. hartwegianum) y Tibouchina (T. gleasoniana
y T. grossa) son en la actualidad utilizados como ornamentales. Mas del 55% de
las especies de los géneros arboéreos y arbustivos Axinaea, Brachyotum,
Graffenrieda, Loreya, Meriania, y Tibouchina es utilizado primordialmente
para carbon, lena o por la fauna local como alimento.
De las 124 especies Utiles, 13 de ellas (cerca del 10%) fueron citadas en la
literatura pero no estaban representadas en los dos herbarios (Ceron 1994, 1996,
Ceron y Montalvo 1998; Elleman 1991; Rios y Caballero 1997), y de las 111 especies
restantes, solamente 36 (32%) fueron citadas en estudios etnobotanicos
anteriores (Alarcon 1988; Cerén 1994, 1996; Cerén y Montalvo 1998; Cueva 1997;
Elleman 1991; Freire-Fierro en prensa; Kvist y Holm-Nielsen 1987; Lescure et al.
1987; Rios y Caballero 1997; Schultes y Raffauf 1990; Van den Eynden 1997;
Vickers y Plowman 1984). Esta compilacion da una pauta sobre la necesidad de
publicar informacion que de otra manera permanece restringida Wnicamente a
los investigadores locales, quienes podrian darla a conocer a la comunidad
internacional por medio de publicaciones similares en otras familias de plantas.
La informacion sistematica sobre usos de Melastomaceae podria servir de
base por ejemplo, para la busqueda de principios activos en las especies listadas
como medicinales; o para la seleccion de especies para la construccién y ornato
que mejor podrian crecer en determinada region geografica; 0 para estudios
ecoldgicos de las especies utiles endémicas, especialmente de aquellas
amenazadas de extincion Cotton 2000); para futuros planes de manejo
sustentable y conservacion | y de campo,
enfatizados en la familia, podrian verificar los usos de Melastomataceae
registrados aqui y en la literatura.
AA
estudioset
ANEXO
Material de Melastomataceae examinado en los herbarios QCA y QCNE. Usos:
A = Alimenticio; C = Construcccién; M = Medicinal; MR = Magico-Religioso, O =
ornamental; R = Otros. Distribucion Jorgensen y Leon 1999, Renner et al. 1999,
Freire-Fierro en prensa): AZU = Azuay, BOL = Bolivar, CAN = Canar, CAR =
Carchi, CHI = Chimborazo, COT = Cotopaxi, ORO = El Oro, ESM = Esmeraldas,
GUA = Guayas, IMB = Imbabura, LOJ = Loja, RIO = Los Rios, MAN = Manabi,
MOR = Morona Santiago, NAP = Napo, PAS = Pastaza, PIC = Pichincha, SUC =
Especie
Provincias
MR
Nombre vernaculo
Especimenes
Referencias
Aciotis ornata (Miq.) Gleason
Aciotis rubricaulis oe
y Mart. ex DC.) Trian
Arthrostemma ciliatum Pav.
ex D.Don
Axinaea affinis (Naudin)
Cogn.
Axinaea macrophylla (Naudin)
Triana
Axinaea quitensis Benoist
Axinaea sclerophylla Triana
Bellucia pentamera Naudin
ESM
CAR, ORO, ESM,
IMB, MOR, NAP,
PAS, PIC, TUN,
ZAM
BOL, CHI, COT,
ORO, ESM, GUA,
LOJ, RIO, MAN,
MOR, NAP, PAS,
PIC, ZAM
CAN, CAR, IMB,
AZU, CAN, CAR,
CHI, IMB, LOJ,
MOR, PIC, SUC,
ZAM
BOL, CAN, CHI,
COT, IMB, LOJ,
NAP, PIC, TUN
AZU, LOJ, MOR,
ZAM
CAR, ESM, IMB,
MOR, NAP, PAS,
SUC, ZAM
x
Chulco, Pin tape (Y)
Agro Viro, Shimujcu
Schji-moh-co (Y)
Colca
Amarillo
Guihle (S)
Amarillo, Palo
blanco (S)
KA dol cil +
ia (K),
muyu yura (K),
Qui
ee guayaba (K),
Barfod 41016 (QCA)
Barfod 41478 (QCA),
Kvist y Asanza 40745
(QCA)
Barfod 48053 (QCA),
Ydnez et al.
Moran y Paisano 104
(QCA, QCNE)
Tipaz 98 (QCA, QCNE)
Ellemann 9876 (QCNE)
Cuamacas etal. 158
(QCNE), Ellemann
9862 (QCNE)
Alarcon 120 (QCA),
Baker 6154 (QCA),
Balslev y Alarcon 3030
=
(QCA, OCNE), Irvine 417
01 (QCNE)
Freire-Fierro
(en prensa)
Ceron (1994)*,
Freire-Fierro
(en prensa)
Ceron (1994)
Elleman (1991
Alarcon (1988)*,
ro
Ww
n (1994)*,
Lescure et al.
(1987)*, Rios y
bbe
(L)OZ VOIS/DYO'LINE
Especie Provincias MR Nombre vernaculo Especimenes Referencias
Tamia muyu (k), (QCA), Palacios 558 Caballero ed
Tunguia (U) (QCNE), Rios 358 (QCA) = Schulte
Ratfaut (1 oo
Blakea bracteata Gleason MOR, NAP, PAS, Huasca payatsi (K) Freire y Cerda
10 (QCA, QCNE)
Blakea glandulosa Gleason MOR, NAP. PAS, Uchan panga (kK) Hurtado y Alvarado 296
TUN (QCA, QCNE)
Blakea involvens Markar. BOL, RIO, PIC Quirdo (QO) Kvist 40049 (QCA)
Blakea punctulata (Triana) CAR, ESM Pilmo Barfod 41613 (QCA)
Wurdack
Blakea cf. rosea (Ruiz y MOR, NAP, PAS Hirt khaki (F),Shashapi Lewis et al. 14045 Lescure et al.
Pav.) D.Don (K), Shashaqui (A), (QCNE), Rios 343 (QCA), (1987),
Yayurua (I) Vickers 255 (QCA) Schultes
Raffauf (1990)
Vickers y
Plowman (1984)
Blakea sawadae J.F. Macbr. MOR, NAP. PAS Ttembaccdpicho (F) Cerdn (1994)
Blakea subconnata O. Berg AZU, BOL, CAR, X Cacha micite, Kin Barfod 48336 (QCA),
ex Triana COT, ORO, ESM, mullu-chua (Y) Kvist 40297 (QCA),
GUA, IMB, RIO, Pei-ca (Y), Quinlulo Kvist 40497 (QCA),
NAP. PIC, TUN tape (Y) Tipaz et al. 1228 (QCA,
QCNE)
Brachyotum alpinum Cogn. AZU, BOL, CHI, Urco fichana (K) Ceron etal. 11785
Brachyotum azuayense
IMB, MOR, PIC,
TUN
AZU,LOSJ
Guillo yuyo (S), Quillo
(QCA, QCNE)
Ellemann 9904 (QCNE),
YOQVNDI 13 NJ IVIDVIVWOLSYT3W “Ty 13 OWH3Id-3HIINd
She
Especie Provincias A MR Nombre vernaculo Especimenes Referencias
Wurdack yuyo Ellemann 66933 (QCA),
Ellemann 75440 (QCA)
Brachyotum ledifolium BOL, CAR, CHI, Lengua de gat Ceron et al. 6883 (QCA, — Ceron (1994,
(Desr.) Triana COT, IMB, NAP, Puca-chaglla ( ; Quijol QCNE), Nunez 75 (QCA, —- 1996)
PIC, TUN QCNE), Tinajero 127 (QCA)
Brachyotum rostratum AZU,LOJ, MOR, Quilloyuyu Ellemann 66862 (QCA)
(Naudin) Triana
Brachyotum trichocalyx Triana AZU,LOJ Escoba colorada Ellemann 66575 (QCA)
Clidemia acostae Wurdack IMB, RIO, PIC x Jaramillo 7012 (QCA)
Clidemia allardii Wurdack CAR,MOR,NAP x Chuegimo, Oa.-/‘do- Barfod 41060 (QCA),
PAS, SUC tape (Y), Mendoza 85 (QCA)
Clidemia capitellata ESM, IMB, NAP Aewaeniwae Oldeman 84 (QCA)
(Bonpl.) D.Don PAS, ZAM
Clidemia crenulata Gleason ESM x Oa-do-tape (Y), Barfod 41061 (QCNE),
Pin-remedio-tapé (Y) — Barfod 48077 (QCA)
Clidemia dentata Pav. ex D.Don CAR, CHI, COT, Paitzi (K) Alarcon 19523 (QCA) Alarcon (1988)*,
ESM, GUA, IMB, Lescure et al.
LOJ, RIO, MAN, (1987)*
MOR, NAP. PAS,
PIC, SUC, TUN,
ZAM
Clidemia dimorphica
J.F.Macbr
Clidemia epiphytica
MOR, NAP, PAS,
SUC, ZAM
CAR, COT, ESM,
><
=>
Aewaeniwaeé (H '
IcSa nanai yura,
Sambiri’si
Pishunguy tape (Y),
anil
Oldeman y Arévalo
79 (QCA), Ponce 236
(QCA, QCNE)
Kvist y Asanza 40371
Ceron (1994),
Lescure et al.
(1987)
9bZ
(LOZ VaIS/D¥O"LINa
Especie Provincias MR Nombre vernaculo Especimenes Referencias
(Triana) Cogn. RIO, MOR, NAP, Sacha-uchan-panga (QCA, QCNE)
PAS, PIC, SUC
Clidemia heterophylla NAP, PAS Situli panga (K), Asanza y Neill 79 Cerdén (1994),
(Desr.) Gleason Uchan (K), (QCA, QCNE) Schultes y
Uchanpanca (k) Raffauf (1990)
Clidemia hirta (L.) D. Don ORO, ESM, LOJ, Rios 434 (QCA) Schultes y
NAP, PAS, ZAM Raffauf (1990)
Clidemia ostrina Gleason CAR, ESM, MOR So-é (Y) Kvist 40307 (QCA)
Clidemia purpurea Pav. ESM, GUA, RIO, x Un ga lala jé tape (Y) Kvist y Asanza 40896
ex D.Don MAN, PIC (QCA, QCNE)
Clidemia septuplinervia Cogn. ESM, RIO, MOR, Henda-pe-chi (Y) Kvist 40510 (QCA),
NAP PAS, PIC Rios 357 (QCA)
Clidemia sprucei Gleason MOR, NAP, PAS Amarun uchu (K), Asdall 8228 (QCA), Lescure et al.
Ampar (U), Eweyiwe (H), Kohn 1162 (QCA, (1987)
Jemper (U), Sutulli (K), = QCNE), Lewis et al.
Teempe jari (U), 13805 (QCA,
Teempe-jari-Macuma, QCNE), Lewis et al.
Uchan (K),Walinumba 74090 (QCA, QCNE),
Palacios 10254 (QCA,
QCNE), Ponce 226
(QCA), Van Asdall
82-28 (QCA, QCNE)
Clidemia variifolia Wurdack MOR, NAP, PAS Pirana panga (K)
Neill et al. 7723 (QCA,
E
QCNE)
YOOVNDI 13. NF IVIIVIVWOLSY T3W “TY 13 OWN51S-341FNs
Lv?
Especie Provincias A MR Nombre vernaculo Especimenes Referencias
Conostegia attenuata Triana CAR, ESM Picha quare (Q) Tipaz et al. 1354 (QCA,
QCNE)
Conostegia centronioides COT, ESM, RIO, x Jendape mirogulachi, — Kvist y Asanza 40779
Markgr. MOR, PAS, PIC Ka-gen-tape (Y) (QCA, QCNE),
Kvist, 40528 QCA
Conostegia dentata Triana CHI, ESM Onia-tapé-ban-vino Barfod 48100 (QCA)
-tapé-micharo- kola
Diplarpea paleacea CAR, ESM Hierba buena Kvist 48723 (QCA)
Triana ex Cogn.
Graffenrieda aff. intermedia ZAM Ebenibe, Gonamonoyi, Ceron y
Triana Evenyive, Tahualagi Montalvo
(todas H) (1998)
Henriettella sp. ESM,CAR,SUC, x Rios y
PAS, NAP MOR Caballero
(1997)
Leandra caquetana Sprague MOR, NAP, PAS Obata médyabo Rios 496 (QCA)
Leandra caquetensis Gleason LOJ, MOR, NAP, x Aéwaéniwae Oldeman 81 (QCA)
PAS, ZAM
Leandra granatensis Gleason CAR, ESM, IMB, Quianchi (Y) Kvist 40326 (QCA)
PIC
Leandra secunda (D.Don) MOR, NAP. PAS, x Chanchak (A), Jaramillo 10912 (QCA),
Cogn. Urkupayas (K) Lewis et al. 13906 (QCA,
QCNE)
Leandra subseriata BOL, CAN, CAR, Colca Ceron et al. 10384 (QCA,
8bZ
(L)0Z vdls/D4O'LIYa
Especie
Provincias
Nombre vernaculo
Especimenes
Referencias
(Naudin) Cogn.
Loreya spruceana
Benth. ex Triana
Maieta guianensis Aubl.
Meriania tomentosa
(Cogn.) Wurdack
Miconia del ugit 105d Naudin
Miconia affinis DC.
Miconia asclepiadea
Triana
CHI, COT, IMB,
PIC, TUN
MOR, NAP
MOR, NAP PAS,
AZU, BOL, CAN,
CAR, CHI, COT,
IMB, LOJ, MOR,
NAP, PIC, TUN,
BOL, CAR, CHI,
ORO, PIC
IMB, NAP SUC
AZU, BOL, CAR,
MOR, NAP PIC,
ZAM
<
Chanchak (A),
Payas (Kk),
Quisa muyu yura (K),
Titemenca (H)
Chanchak chichi (A),
Egim6 (H), Payas (kK) ,
Putus panga (K),
Shallipu (K), Uchan (K)
Hualicon cari
Colca, sagalita
<<
Hagapania, Payanzo (K
Evenibe (H)
'
QCNE)
Irvine 128 (QCA),
Lewis et al. 13856
(QCA, QCNE)
Balslev 3016 (QCA),
Kohn 1211 (QCNE),
Lewis et al. 13899 (QCA,
QCNE), Neill et al. 8675
(QCA, QCNE), Rios 477
(QCA)
Nunez et al. 23 (QCA,
NE)
Ceron 12990 (QCA,
E
Alarcon 34 (QCA)
5.c. 485 (QCA)
Lescure et al.
(1988)*,
Schultes y
Raffauf (1990)*
Cerdon (1994,
1996)
Alarcon (1988)*,
Montalvo (1998),
Lescure et al.
(1987)
YOOWNDI 13 NF IVIDVIVWOLSVTIW “TV 13 O8YI1S-IYIFYS
6b
Especie
Provincias
Nombre vernaculo
Especimenes
Referencias
Miconia aspergillaris
(Bonpl.) Naudin
Miconia astroplocama
Donn.Sm
Miconia aureoides Cogn.
Miconia barbinervis
(Benth,) Triana
Miconia brachycalyx Triana
Miconia oo
(B
pl.) )D
Miconia bubalina
(D.Don) Naudin
AZU, BOL, CAN,
LOJ, MOR, PIC
ORO, RIO, MOR,
NAP, PAS, PIC
MOR, NAP, PAS,
SUC
ESM, GUA, LOJ,
RIO, MOR, NAP,
PAS, PIC, SUC,
TUN, ZAM
CAR, ESM, PIC
AZU, CAN, CAR,
CHI, COT, IMB,
LOJ, MOR, NAP
PIC
GUA, MOR, NAP.
PAS, SUC, ZAM
—
Ao nee au (|
Evenyibe (H)
Hén-tapé (Y)
Chicharron
Chilloyudo, Quijol
Ayamohue (H),
Chiquitapayanzo,
Ebenibe (H),
Pishcu micuna muyu
yura (Kk),
Puca payas (K)
Jaramillo 3829 (QCNE)
Vickers 207 (QCA)
Barfod et al. 48431
(QCA, OCNE)
Palacios 5178 (QCA,
QCNE)
Eriksen 59104 (QCNE),
Moran et al. 15 (QCA,
QCNE), Moran et al. 84
(QCA, OCNE)
Alarcon 35a (QCA),
i ia y Nenquerei
cn QCNE), Hurtado
a QCNE),
Irvine 126a (QCA)
Lescure et al.
(1987)
Vickers y
Plowman (1984)
Ceron y
Montalvo (1998
ete
Alarcon (1988)*,
-eron y
Montalvo (1998),
Lescure
et al. (1987)*
057
(L)0Z VaIS/9¥O'LINa
Especie
Provincias
Nombre vernaculo
Especimenes
Referencias
Miconia caelata (Bonpl.) DC.
Miconia calvescens Shrank
y Mart. ex
rae
Miconia centrodesma Naudin
nee } Relat
Miconia cercophora Wurdack
Miconia crocea (Desr.) Naud.
Miconia decurrens Cogn.
LOJ, ZAM
RIO, MOR, NAP,
PAS, SUC, TUN,
ZAM
MOR, NAP
ESM, MOR, NAP,
PAS, ZAM
MOR, NAP, PAS,
ZAM
AZU, BOL, CAN,
CAR, COT, IMB,
LOJ, NAP, PIC,
SUC, TUN
AZU, BOL, CAN,
CAR, CHI, COT,
IMB, LOJ, NAP
PIC, TUN
MOR, NAP, ZAM
x
Cerrag (S)
Pishcu micuna muyu
yura
(K)
Yana muyo (kK)
Payachic muyo (kK)
Payachic (K)
Cerrag (S), Colca,
Colca de
hoja grande, Taruguita
Amarillo, Colca
Pischu-micuna-
muyu-yura (K)
a
rvine 102a (QCA)
Cerdn 7850 (QCA, QCNE)
Hurtado et al. 2076
(QCA, OCNE)
Hurtado y Alvarado 2128
(QCA, QCNE)
Ceron y Alarcon 12303
(QCA, OCNE), Moran
etal. 7 (QCA, QCNE),
Moran et al. 56
(QCA, QCNE)
OCNE), Cerén et al. 7134
(QCA, QCNE), Cerdn et a
10436 (QCA, QCNE),
Cerén et al. 10718 (QCA
QCNE), Cerdn et al, 10841
(QCA, QCNE), Ceron et al.
10997 (OCA, OCNE),
Cerdnet al. 11
Irvine 127 (QCNE),
Irvine 127a (QCA)
Alvarez et al. 1198 (QCA,
1
Elleman (199
]
)
Elleman (1991)
Cerdn (1994,
1996
)
YOQVND9 1] NJ IVIDVIVWOLSVTAW “TY 13 OWNI14-IN1Ia4
L&Z
Especie Provincias A MR Nombre vernaculo Especimenes Referencias
Miconia erioclada Triana ESM, MOR, NAP Jedape nicha rocola (Y)
jubiquichono tape (Y) — Kvist 40569 (QCA, QCNE)
Miconia floribunda NAP Ebenyibe (H) Cerédn y
(Bonpl.) DC. Montalvo (1998)
Miconia goniostigma Triana CAR, COT, ORO, Monte hierba Barfod 41500 (QCA,
ESM, RIO, MOR, QCNE)
NAP. PAS
Miconia gracilis Triana CAR, ESM, PIC Chiknul (Q), Hé.n- Barfod 41033 (QCA),
tapé (Y) Beck 1777 (QCA)
Miconia latifolia CAN, CAR, CHI, x Yamuchuchu (K) Ceron et al. 11777 (QCA,
(D.Don) Naudin IMB, LOJ, MOR, OCNE)
NAP. PAS, PIC,
TUN
Miconia ledifolia AZU, LOS x Sierra (S) Eynden & Cueva 327 Cueva (1997),
(DC.) Naudin (QCA, QCNE) Eynden (1997)
Miconia ligustrina AZU, CAR, COT, Moran et al. 58 (QCA,
(Sm.) Triana IMB, LOJ, MOR, QCNE)
NAP. PIC, SUC,
TUN
Miconia lugonis Wurdack ESM, MOR, NAP, x A len tape (Y) Kvist 40769 (QCA)
Miconia lutescens (Bonpl.) DC. CAN, LOJ, ZAM x Taruma (S) Eynden 184a (QCA) Cueva (1997),
Eynden (1997)
Miconia matthaei Naudin NAP SUC, ZAM Gonomoroyi (H), Ceron y
Evenhibe (H)
Montalvo (1998)
787
(L)0@ vals/94¥0'LINd
Especie
Provincias
Nombre vernaculo
Especimenes
Referencias
Miconia napoana Wurdack
Miconia nervosa (Sm.)
iana
Miconia nutans Donn. Sm.
Miconia obscura (Bonpl.
Naudin
Miconia ochracea Triana
Miconia oraria Wurdack
Miconia pnaleacea Cog n.
Miconia nanillosa (Desr.)
Naudin
Miconia pilgeriana Ule
NAP, PAS
ESM, IMB, MOR,
NAP, PAS, PIC,
SUC
CAR, IMB, MOR,
NAP, PAS, TUN
AZU,LOJ, MOR,
TUN, ZAM
BOL, CAR, IMB,
NAP
CAR, ESM
MOR, NAP, PAS
CAR, IMB, LOJ,
NAP, PIC, TUN,
ZAM
NAP, PAS, SUC,
ZAM
Evengimo (H)
Ewengimo, Hein-
tapé (Y), Mif-cha-
rro-kola-hambi-
kéno-tape
Equive (H)
Cerrag (S)
Amarillo
Ma-sju-ke-mu-chi (Y),
Mashu-kimu-chi
Caracha panga (k),
Guala, Paijatsigi (kK),
Puca-paya-tsic-
panga (kK),
Tacupayas (K)
Colca, Tostado de
pajarito
Evenibe (H)
Barfod 41014 (QCA),
Barfod 48099 (QCA),
Miller 102 (QCA)
Elleman 66671 (QCNE)
Tipaz 63 (QCA, QCNE)
Barfod 48216 (QCA),
Barfod 48339 (QCA)
Bravo 42 (QCA), Kohn
1090 (QCA, QCNE),
Kohn & Alvarado 1263
QCA), Neill 7730
QCNE), Palacios 10347
(QCA, QCNE)
=
=
Moran & Paisano 114
(QCA, QCNE), Tinajero
183, 184 (QCA)
QCA,
Ceron y
Montalvo (1998)
Cerédn y
Montalvo (1998)
Elleman (1991)
Cerdn (1994)
Ceron y
Montalvo (1998)
YOOVNII 13 NI IVIDVIVWOLSV TSW “Tv 13 O8N514-341dNd
£57
Especie Provincias A MR Nombre vernaculo Especimenes Referencias
Miconia procumbe
(Gleason) Wa MOR, NAP Shishak (U) Warush RBAE 84
(QCA, QCNE)
Miconia pustulata Naudin AZU, BOL, CHI, Amarillo, Angoterio Bdez 35 (QCA), Cerdn (1994)
COT, IMB, PIC Ceron et al.6978
(QCA, QCNE),
Cuamacds & Gudifio
245 (QCA, OCNE)
Miconia salicifolia (Bonpl. AZU, CAN, CAR, Chacla negra, Espinilla Cerdn et al. 10867
ex Naudin) Naudin CHI, COT, IMB, (QCA, OCNE), Cer6n
LOJ, NAP. PIC, etal. 11069(QCA,
TUN QCNE
Miconia splendens (Sw,) NAP PAS, ZAM x Olla, Evenhibe (H) Cerén & Ayala 9995 Ceron y
Griseb (OCA, OCNE) Montalvo (1998)
Miconia subspicata Wurdack MOR, NAP, PAS Manzano blanco, Bravo 237 (QCA)
San Juan
Miconia theaezans AZU,BOL,CAN, — x Amarillo, Cerrac (S), Barfod 41495 (QCA, Ceron (1994,
(Bonpl.) Cong CAR, CHI, COT, Colca, Laichi, Sacha QCNE), Cerdn & Al i 1996), Elleman
oie LOJ, NAP. Colca 12236 (QCA, QCN (1991)
PAS, PIC, TUN, Cuamacds et al. ve
ZAM (QCA, OCNE), Ellemann
66506 (QCA), Moran
etal. 26 (QCA, QCNE),
Tipaz 7 (QCA, QCNE)
Miconia tinifolia Naudin BOL, CAR, | Amarillo, Cerrag Ellemann 66541 (QCA), — Elleman (1991)
LOJ, NAP, a ZAM
fino (S)
Tipaz 33 (QCA, QCNE)
pS
(L)0Z VdIS/9¥O'LINS
Especie
Provincias
Nombre vernaculo
Especimenes
Referencias
Miconia trinervia (Sw.)
.Don ex Loudon
Miconia triplinervis
Ruiz & Pav.
A
Viiconia venulosa Wurdack
Miconia versicolor Naudin
Miconia zubenetana
J.F. Macbr.
Monochaetum hartwegianum
Naudin
Monochaetum lineatum
(D.Don) Naudin
Monolena primuliflora
Hook.
CHI, ORO, ESM,
MOR, NAP, PAS,
SUC
ESM, MOR, NAP,
PAS, ZAM
COT, ESM, MOR,
NAP, PAS, PIC
CHI, IMB, PIC
NAP, PAS
AZU, BOL, CAR,
CHI, COT, IMB,
AZU, CAN, CAR,
CHI, COT, IMB, LOJ,
MOR, NAP, PAS,
PIC, TUN, ZAM
CAR, COT, ESM,
LOJ, MOR, NAP,
PAS, PIC, TUN,
ZAM
X
x
Jedape mincharogola
tape (Y)
Evengui (H),
Llagaspanga,
Uchan panga (K
Pengomo (Y)
Morochillo
Biasek (K), Palo de agua
Salapilla (S)
Apanguira, Aurutullo
(K), Auru-tulla, Corazon
panga, Jeen shiimujcu,
Lyofasufsapa‘cco,
Lyunk (U), Nimon
onabo, Quenca
)
Kvist 40489 (QCA, QCNE)
Balslev 2933 (QCA),
Gudino 1730 (QCA,
QCNE)
Kvist 40324 (QCA)
Miller et al. 2370
(QCA, QCNE)
Ceron et al. 11002
Elleman 75407 (QCNE
Ww
Aulestia & Aulestia 1321
(QCA, OCNE), Aulestia
et al. 1456 (QCA, QCNE
Barfod 41582 (QCA),
Cerdn 186 (QCA),
Kohn 1156 (QCA, QCNE),
i
4
Ceron y
Montalvo (1998)
Cerén (1994,
1996)
Elleman (1991)
Ceron (1994),
Kvist y Holm-
Nielsen (1987),
Schultes y
Raffauf (1990)
YOOVNDI 13 NJ IVIIVIVWOLSY TW “Ty 13 O8YI1S-IY1INd
$6?
Especie
Provincias
Nombre vernaculo
Especimenes
Referencias
Mouriri grandiflora DC.
Mouriri nigra (DC.) Morley
Mouriri oligantha Pilg.
Ossaea bracteata Triana
Ossaea laxivenula Wurdack
Ossaea micrantha
(Sw.) Macfad. ex Cogn.
Ossaea quadrisulca
(Naudin) Wurdack
Ossaea robusta (Triana)
Cogn.
MOR, NAP, PAS,
MOR, NAP
MOR
CAR, PIC
CAR, ESM, IMB,
MOR, PAS, PIC
CAR, ORO, ESM,
GUA, IMB, LOJ,
RIO, MAN, MOR,
NAP PAS, PIC
MOR, NAP PAS,
ZAM
CAR, ESM, MOR,
NAP, PAS, PIC
x
x<
querentebin (H), Urti
tullu (K)
Sharima (U), Sharimiat
Pagaramoncamo (H)
Madera de pambil
Hein-tapé-chi (Y),
O/.n-gala-hen-tapé (Y)
Isah tape (O),
Pichi ri chide (O),
Tajasanene
Cana agria, Fruto
de conga, Yuturi
muyu (K)
Pengomo (Y
as
Lewis et al. 14198 (QCA,
QCNE), Macias & Asanza
19 (QCA, QCNE), Neill
7728 (QCA), Bligaard
§7359 (QCA), Palacios
10291 (QCA)
Enynden et al. 914
(QCNE), Palacios 6653
(QCA, QCNE)
Mendoza 201 (QCA)
Miller et al. 2299
( CNE)
Buitron 654 (QCA)
Barfod 41013 (QCA,
QCNE), Barfod 48173
(QCA), Kvist 40364
—_,
Kvist 40048 (QCA),
Kvist 40162 (QCA)
Rios 389, 390 (QCA)
Kvist & Asanza 40324
(QCA, QCNE)
QCA)
Hernandez 111 (QCA),
Cerdn (1994),
Schultes y
Raffauf (1990)
Schultes y
Raffauf (1990)
967
(L)OZ VaIS/O¥O"LINa
Especie Provincias MR Nombre vernaculo Especimenes Referencias
Tessmannianthus NAP, PAS Evennive (H), Ceron y
heterostemon Markagr. Evennimo (H) Montalvo (1998)
Tibouchina gleasoniana BOL, CAR, COT, Flor de mayo Cuamacds & Gudinrio
Wurdack IMB, PIC 181 (QCA, QCNE)
Tibouchina grossa (L.f.) Cogn. CAN, CAR, SUC Onore 628 (QCA)
Tibouchina laxa (Desr.) Cogn. AZU, BOL, CAN, Dom brin Cerén 13473 (QCA,
CHI,ORO, LOJ, QCNE)
PIC, ZAM
Tibouchina lepidota AZU, CAR, CHI, Dumarin Ellemann 66543 (QCA)
(Bonpl.) Baill. IMB, LOJ, MOR,
NAP, PIC, TUN,
ZAM
Tibouchina longifolia BOL, CAR, CHI, Kukuj
(Vahl) Baill.
Tibouchina mollis
(Bonpl.) Cogn.
Tibouchina oroensis Gleason
Tococa guianensis Aubl.
COT, ORO, ESM,
GUA, IMB, LOJ,
RIO, MAN, MOR,
NAP, PAS, PIC,
TUN, ZAM
BOL, CAR, CHI,
COT, ORO, IMB,
LOJ, NAP, PIC,
ORO, LOJ
ESM, MOR, NAP.
PAS,SUC
Flor de mayo
Hallilumba, Payanshu
(K) Jatunpayanshu (K)
Moran & Paisano 79
(QCNE)
Tinajero 190-195, Cerdn (1994,
197 (QCA) 1996)
Palacios & Tirado 13013
(QCA, QCNE)
Alarcon 27 (QCA), Alarcon (1988),
Blane 96107 (QCA) Ceron y Montalvo
(1998)*, Lescure
et al. (1987
YOQVNDI 13 NI IVIDVLVWOLSW AW “Ty 13 OWNSIS-3u1ded
£82
Especie
Provincias
MR
Nombre vernaculo
Especimenes
Referencias
Tococa spadiciflora Triana
Tococa symphyandra
(Triana) Cogn.
Triolena hirsuta (Benth.)
Triana
Triolena obliqua (Triana)
Wurdack
Triolena pedemontana
Wurdack
Triolena pluvialis
(Wurdack) Wurdack
Triolena pustulata Triana
RIO, MOR, PIC
CAR, NAP. SUC
CAR, COT, ESM,
MOR, NAP, PAS,
TUN
AZU, ORO, ESM,
GUA, PIC
CAR, NAP, PAS,
PIC
COT, ESM, PIC
MOR, NAP, PAS,
CHI, COT, GUA,
Ouish (Q)
Allipas payas (K),
Cunua (K), Ho-horo-
tape (Y), Pini tape
(O), Raca pies
Maipe
Contra
Cu’a shocongo,
Cunua (K), Koshashi
(F), Shashaki panga
(K), Sutulli (K)
Maape tape,
Yahuar pallas (K)
Kvist 48701 (QCA)
Kvist 48725 (QCA)
Alarcon 19508 (QCA),
Barfod 41084 (QCA),
Barfod 48032 (QCA),
Blane 96553 (QCA),
Kvist 40094 (QCA)
Barfod 41418 (QCA),
Barfod 41642 (QCA),
Barfod 48807 (QCA),
Kvist 48714 (QCA)
Beck 1793 (QCA, QCNE)
Hurtado 2827 (QCNE),
Palacios 10344 (QCNE)
Blanc 9637
Yanez et al. 8 (QCNE)
Schultes y
Raffauf (1990
Alarcon (1988),
Ceron (1994),
Kvist y Holm-
Nielsen (1987),
(1987), Schultes
y Raffauf (1990),
Vickers y Plow-
man (1
Schultes y
Raffauf (1990)
857
(L)OZ VdIS/DHO'LIYa
FREIRE-FIERRO ET AL., MELASTOMATACEAE EN EL ECUADOR 259
Sucumbios, TUN = Tungurahua, ZAM = Zamora Chinchipe. A = Achuar, F =
Cofan, H = Huaorani, I = Siona-Secoya, K = Kichwa, O = Colorado, Q = Coaiquer,
S = Saraguro, U = Shuar, Y = Cayapa. * = Especie citada bajo un sindénimo, mal
escrita o no identificada.
AGRADECIMIENTOS
Agradecemos a todas aquellas comunidades que desinteresadamente
compartieron sus conocimientos con los colectores; a todos los colectores
listados en el apéndice quienes registraron los usos de las plantas en sus etiquetas,
especialmente a A. Barfod, C.E. Ceron, L. Eemann, J. Jaramillo, L.P Kvist y M.
Rios; al personal de los herbarios QCNE y QCA, quienes nos permitieron revisar
las colecciones de Melastomataceae, especialmente a Mercedes Asanza, David
Neill, Katya Romoleroux y Renato Valencia. David Neill y Rosa Ortiz-Gentry
(Missouri Botanical Garden), Kevin Janni y un revisor anonimo, contribuyeron
mucho para el mejoramiento del articulo gracias a sus comentarios en versiones
anteriores. La primera autora agradece al Plutarco Naranjo de la Universidad
Andina Simon Bolivar, por la oportunidad de presentar este trabajo en las II
Jornadas Ecuatorianas de Etnomedicina realizadas en 1999,a FUNDACYT yal
Missouri Botanical Garden por todo el apoyo logistico, y a Joseph C. Welling por
toda su ayuda durante la recopilacion de la informacion.
REFERENCIAS
ALARCON, R. 1988. Etnobotanica de los Quichua de la amazonia ecuatoriana. Misc. Antrop.
Ecuat. Ser. Monogr. 7:1-183.
Ceron, C.E. 1994. Manual de botanica ecuatoriana: Sistematica y métodos de estudio. Uni-
versidad Central del Ecuador, Quito.
Ceron, C.E. 1996. Plantas utiles de la Reserva Geobotdnica del Pululahua, Provincia de
Pichincha, Ecuador. Hombre y Ambiente 25:9-72.
CerOon, C.E. y C.G. Montatvo. 1998. Etnobotanica de los Huaorani de Quehueiri-Ono, Napo,
Ecuador. Abya-Yala, Quito.
Corton, E.2000.Melast taceae.En:R.Valencia,N. Pitman, S.Ledn-Yanez, y RM.J@rgensen,
eds. Libro Rojo de las plantas endémicas del Ecuador 2000. Publicaciones del Herbario
OCA, Pontificia Universidad Catélica del Ecuador, Quito. Pp. 230-250.
Cueva, E. 1997. Arboles y arbustos productores de frutos comestibles en la provincia de
Loja. En: Rios, M. y H.B. Pedersen, eds. Uso y manejo de recursos vegetales. Memorias
del Il Simposio Ecuatoriano de Etnobotanica y Botanica Econémica.Ed. Abya-Yala, Quito.
ELtemaN, L.1991.El uso de la madera del bosque montano por los Saraguro. En: Rios, M.y H.
Borgtoft Pedersen (eds.), Las Plantas y el hombre. Herbario QCA y Abya-Yala, Quito.
Freine-Fierro, A.En prensa. Monograph of Aciotis (Mel eae). Monog. Syst. Bot. 62
JORGENSEN, PM. y S. Leon-YAnez (eds.). 1999. Catalogue of the vascular plants of Ecuador,
Monogr. Syst. Bot. Missouri Bot. Gard. 75:42.
260 BRIT.ORG/SIDA 20(1
KorHaai, B. 1993. Nucanchic Panpa Janpicuna: plantas medicinales del campo. Ed. Abya-
Yala, Quito.
Kvist, L.P.y L.B. Houm-Nievsen. 1987. Ethnobotanical aspects of lowland Ecuador. Opera Bot.
92:83-107,
Lescure, J.P, H. Batstev, y R. Atarcon. 1987. Plantas utiles de la Amazonia ecuatoriana: Un
inventario critico de los datos disponibles en Quito. ORSTOM-P.U.CE.-ILN.C.R.A.E.
Programa Nacional de Regionalizacién Agraria PRONAREG, Quito.
Maseertey, D.J. 1997. The plant book: a portable dictionary of the vascular plants. 2nd ed.
Cambridge University Press.
Renner, S.S. 1993. Phylogeny and classification of the Melastomataceae and Memecylaceae
Nord. J. Bot. 13:519-540.
Renner, S.S., F. Aumeba, y E. Corton. 1999. Melastomataceae En: Jorgensen, P.M. y S. Leon-
Yanez, eds. Catalogue of the vascular plants of Ecuador, Monogr. Syst. Bot. Missouri Bot.
Gard. 75:561-585.
Rios, M. 1993. Plantas utiles en el noroccidente de la provincia de Pichincha. Hombre y
Ambiente 26:1-185.
Rios, M. y J. Casattero. 1997. Las plantas en la alimentacién de la comunidad Ahuano,
Amazonia ecuatoriana. En: Rios, M. y H.B. Pedersen, eds. Uso y manejo de recursos
vegetales. Memorias del Il Simposio Ecuatoriano de Etnobotanica y Botanica
Econdmica. Ed. Abya-Yala, Quito.
ScHULTES, R.E.y R.F. Rarraur. 1990. The healing forest: medicinal and toxic plants of the North-
west Amazonia. Dioscorides Press, Portland, OR.
Van den Eynden, V. 1997. Plantas comestibles en la provincia de Loja. En: Rios, M. y H.B.
Pedersen, eds. Uso y manejo de recursos vegetales. Memorias del II Simposio
Ecuatoriano de Etnobotdnica y Botanica Econdmica. Ed. Abya-Yala, Quito.
Vickers, W.T. y T. Plowman. 1984. Useful plants of the Siona and Secoya indians of eastern
Ecuador. Fieldiana, Bot. 15:1-63.
Wurback, J. 1980. Melastomataceae. En: Harling, G. y B. Sparre, eds. Flora of Ecuador 13:
1-406.
THE IDENTITY OF EAGLEWOOD (GYRINOPS,
THYMELAEACEAE), ANEW ECONOMIC RESOURCE
FOR PAPUA NEW GUINEA
W. Takeuchi M.Golman
C/o PNG Forest Research Institute PNG National Forest Service
OC). PO. Box 5055
Lae, PAPUA NEW GUINEA Boroko, NCD, PAPUA NEW GUINEA
ABSTRACT
] : ] at feeeeoaeres F | ] Sev-
ls aerate
eral t being confused with G. ledermannii but the true eipleqood: is readily distin-
guished by its floral characters.
TOK IGO PAS (MELANESIAN TOK PISIN)
Diwai ol i kolim Gyrinops ledermanni ae ene aa long tok latin na igalwud or gaharu long
tok malaia na nau em tu nem mipela i save long Papua New Guinea. Gyrinops, em nau i kamap
wanpela diwai insait long maket bilong salim na ae dispela diwai. Pastaim sampela luksave long
lif na flawa bilong dispela diwai ino bin kamap klia tumas. Tasol nau i gat wei bilong luksave long ol
flawa na lif bilong trupela diwai igalwud.
INTRODUCTION
In recent years, a lucrative trade has emerged in the Hunstein subdistrict of
East Sepik Province, involving a natural product commonly knownas ‘eaglewood’
(Fig. l). The newly established market is based on an arborescent species which
accumulates aromatic oils in older wood. Demand for the fragrant wood can be
likened to that for sandalwood, Santalum spp. (Santalaceae). However unlike
sandalwood, the eaglewood essence appears to be a contingent response to in-
jury or trauma, rather than a normal outcome of maturation. In its most desir-
able form, eaglewood becomes blackened by concentrated deposits of aromatic
compounds, attracting prices of K800-1,000 per kg for the highest grades. Fol-
lowing separation of the oils, the fragrant extracts are currently used in the
production of expensive incense and perfumes, for eventual purchase by Asian
consumers.
Due to its recent appearance in commerce and unusually high value,
eaglewood has become the focus of intense interest. Although the product is
known primarily from lowland and low montane forest, there has been con-
siderable uncertainty over the identity and number of species involved in the
current trade. In an attempt to resolve the taxonomic issues, the eaglewood
source areas near Ambunti township were visited by the authors in August 200],
SIDA 20(1): 261 — 267. 2002
BRIT.ORG/SIDA 20(1)
262
S
the
Gray
ieee
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a
e
By 909'LZg= Bolg
Vina ouinyes Wsdy
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Pr ee
&
3
- a
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1
Tor eagiewood.
ne ap
te ‘as
Se
fut
i
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ions are in
Most of the known populat
J
dictrihuth
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PproDabDly Nes Turner West
bw Ed
Fic. 1.H
Hunstein
shading indicates areas of closed canopy forest.
TAKEUCHI AND GOLMAN, GYRINOPS IN PAPUA NEW GUINEA 263
as part of a general floristic study of the upper Sepik. The following account
presents the results of these investigations!
TAXONOMY OF EAGLEWOOD
During the 1989 National Geographic Society expedition to Mt. Hunstein (cf.
Bakker 1994; Sohmer et al. 1991), over 1,200 botanical specimens were btained
from areas where eaglewood is now harvested. Among the survey vouchers was
Takeuchi 4848 (A!, LAE), collected from a tree described by village guides as
having a fragrant black wood which could burn even when wet. This specimen
was identified as Gyrinops ledermannii Domke on the original distribution
(also confirmed in 1995, cf. annotation on the A duplicate), a species considered
endemic to the upper Sepik and previously known with certainty only from
the type gathering (Ledermann 7401; Pentecost Mt along the May River). The
identification of the 1989 collection was facilitated by comparisons against two
additional specimens obtained within the last 50 years, but the other reference
materials have since disappeared from the Lae National Herbarium.
The Ledermann locality lies ca. 100 km WNW of the April River site des-
ignated as ‘Kamelsrucken’ (Samsai Mt) by early German explorers. Samsai vil-
lagers report that eaglewood is very common on the west-facing slopes of
Kamelsrtcken, in the direction of Pentecost Mt. Because the original specimen
was taken from the May River, the local testimony implies that the distribu-
tional center for the species lies further to the west of existing production areas.
The Hunstein populations are possibly only secondary occurrences.
Ledermann’s type was lost in the 1943 destruction of Berlin Her-
barium, the National Geographic survey voucher was unambiguously keyed
to the given species using Ding Hou (1960). After eaglewood became a resource
deserving of scrutiny, suspicion was thus directed to Gyrinops as the likely
source for the new product.* However, in the absence of herbarium material
specifically identified as eaglewood by local producers, it was difficult to be
sure of the presumed correspondence. Especially as eaglewood was recently
discovered in Jayapura (West Papua or Papua Barat, the former Irian Jaya), where
itis there identified as Aquilaria filaria (Oken) Merr.(Thymelaeaceae), consid-
erable confusion existed over the plant's identity.
A fertile collection taken during our Ambunti reconnaissance (Towati &
Golman 159) was pointed out by village suppliers as an example of the tree har-
vested for the eaglewood trade. The Ambunti voucher includes both flowers
and fruits, and its completeness leaves no room for doubt that Gyrinops
ledermannii is the Hunstein eaglewood. In the Ambunti specimen, the pen-
'The original submission of this manuscript was made to en ee Meee but subsequently withdrawn.
2A similar conclusion and t i d f | repo ted by Zich and
Compton (2001) while this paper was in submission.
264 BRIT.ORG/SIDA 20(1)
tamerous flowers have the same number of stamens as calyx lobes, each sta-
men also alternating with a fimbriate petaloid appendage. This clearly elimi-
nates Aquilaria from consideration, because in that genus the number of sta-
mens is twice that of the calyx lobes (Ding Hou 1960). Aquilaria filaria is
otherwise very similar to Gyrinops ledermannii, particularly in their shared
characteristic of loculicidal capsules that develop by rupturing through one
side of the corolla tube. It is easy to get the two species conlused unless the flow-
ers are carefully examined. Reports of eaglewood being present in West Papua,
and its identification as Aquilaria filaria, need to be reevaluated. Aquilaria has
thus far been recorded in New Guinea only from the Vogelkop region, so its
alleged presence at other West Papuan localities such as Jayapura is not unrea-
sonable. Due to the similarities between the respective taxa, it is entirely plau-
sible that Aquilaria represents another source lor eaglewood. However photo-
graphs of the Jayapura provenance taken by M. Golman are more suggestive of
Gyrinops ledermannii than of Aquilaria filaria (Figs. 2, 3).
Uncertainty over the identification of eaglewood is also evident among lo-
cal villagers searching lor the plant. Thymelaeaceae is a prominent family in
the Hunstein flora, with very close similarities among its representatives
Within the existing source areas, Phaleria coccinea (Gaud.) FvM.
(Thymelaeaceae) is the species most often mistaken for eee In superti-
cial aspect its inflorescence is comparable to Gyrinops, and the leaves are also
of similar appearance, particularly with respect to venation. As with other
members of the family, Phaleria coccinea has an extremely strong inner bark
which is often used in the construction of white bilums (ie., local netbags used
as carry-alls) noted for their soft texture.
Due to the black color of the branchlets, Diospyros papuana Val. (Ebenaceae)
is also mistaken by villagers for eaglewood, even though the differences in mor-
phological aspect are rather obvious. For example, unlike eaglewood the leat
venation in D. papuana is bifacially prominulous, and the fruits are indehis-
cent and large. The flowers are also unisexual and dioecious, rather than bi-
sexual. Table | summarizes some of the salient distinctions between Gyrinops
ledermannii and the plants with which it is presently confused.
In addition to the problems caused by similarities with other taxa, G.
ledermannii is highly variable and can exhibit considerable difference in leaf
size between individuals. The collection from the 1989 survey has lanceolate
blades averaging 5.0 x 2.5cm but the more recent material has elliptic leaves ca.
19.5 x 65cm. Presumably these distinctions are at least partly under environ-
mental control, especially in view of the wide range of substrates comprising
eaglewood habitat (cf. Takeuchi @ Golman 2002:63-64: this issue). The nonde-
script nature of the vegetative characters also discourages effective identifica-
tion of sterile plants, thus adding to the confusion.
TAKEUCHI AND GOLMAN, GYRINOPS IN PAPUA NEW GUINEA 265
1 ab | Iwai L Sey Cee of eR BY ee lod
Fic. 2. Eaql f J TI
and Aquilaria filaria. Photo by M. Golman, August, 2001.
LI £ i |
Fic. 3. Flowering | yapura population. The i | f the flower is critical to the proper
identification of eaglewood. Photo by M. Golman, August, 2001.
266 BRIT.ORG/SIDA 20(1)
{
Tape 1. Comparative list of reproductive characters for Gyri fi and the taxa with which
it is confused in the field. Tokples names are provided wie the village locality where the name
originates. Voucher source: H&C = R. Hoogland and L.Craven, CSIRO survey from 1966;T =W. Takeuchi,
Hunstein surveys from 1989, 199
Scientific name/Tokples name Flower Fruit
Gyrinops ledermannii Domke
(Thymelaeaceae), may-hasei bisexual small loculicidal capsule
(Bugabugi, J 15531) stamens same no. as
calyx lobes
y i
stamens included
Aquilaria filaria (Oken) Merr.
sais sp. not recorded bisexual small loculicidal capsule
from stamens 2 x no. of calyx
lobes; stamens included
Phal
(Thymelaeaceae), wanyip or yaru bisexual small drupe
(Wagu; H&C 10377); wonyip stamens 2 x no.of
(Wagu; H&C 10543); winyap calyx lobes; stamens
(Waskuk; T 10793) exserted from floral tube
Diospyros papuana Val. (Ebenaceae)
manukway (Ambunti; H&C 10214); unisexual, dioecious large berry
mankall (Waskuk; 7 10227) stamens 3-4 x no. of
calyx lobes; stamens
included
DISCUSSION
Although the economic value of eaglewood affords the promise of substantial
returns for stakeholders, high market prices clearly create incentives for
overexploitation and eradication of natural populations. In order to ensure long-
term viability of existing stocks, appropriate measures in resource management
will eventually require enactment. However the present dearth of information
on Gyrinops ledermannii will undoubtedly constrain its sustainable develop-
ment. Even 90 years after its initial discovery, the plant remains poorly under-
stood by botanical and forestry science. The demography, distribution, and ecol-
ogy of G. ledermanniiare unknown. This situation will require correction before
eaglewood can be properly developed as an economic asset. There is also an
obvious need for natural products research into the chemical principles which
are responsible for eaglewood’s desirable qualities. Depending on the nature of
the resource, the species could conceivably serve as a valuable subject for silvi-
cultural and natural products development.
TAKEUCHI AND GOLMAN, GYRINOPS IN PAPUA NEW GUINEA 267
ACKNOWLEDGMENTS
The aut] tl ion to parataxonomist Ali Towati of Waskuk,
for his assistance on the Ambunti reconnaissance. We also thank the Kokomo
clan of Bugabugi for their goodwill and hospitality during our visits to the April
River. Reviewers John Pipoly ILI (Fairchild Tropical Garden) and Kevin Janni
(University of Hawaii) provided helpful comments on the manuscript.
REFERENCES
Bakker, E. 1994. Return to Hunstein forest. Natl. Geogr. Mag. Feb., 40-63.
Dina Hou. 1960. Thymelaeaceae. Flora Malesiana ser. |, 6(1):1-48.
SOHMER, S.H., R. KIAPRANIS, A. ALLISON, and W. TakeucHi. 1991 (unpublished). Report on the
Hunstein River expedition—1989. Report to the National Geographic Society, Wash-
ington D.C, U.S.A.
TAKEUCHI, W. and M.Gotman. 2002.The present status of Ledermann’s April River localities in
Papua New Guinea. Sida 20:55-70.
Zick, F.and J.Compton. 2001.The final frontier. Towards sustainable management of Papua
New Guinea's agarwood resource. Traffic Oceania and WWF South Pacific Programme:
http://www.traffic.org/news/agar2.pdf.
268 BRIT.ORG/SIDA 20(1)
BOOK REVIEW
TM. Antonio and S. Masi. 2001. The Sunflower Family in the Upper Midwest: a
Photographic Guide to the Asteraceae in Ilinois, Indiana, Lowa, Michigan,
Minnesota, and Wisconsin. (ISBN |-883362-|1-3, hbk.). Indiana Academy
of Science, Indianapolis, in collaboration with Chicago Botanic Garden,
Glencoe, IL, U.S.A. $50.00, 419 pp, numerous color photos, maps, 7" « 10".
| love this book for the superlative photos and the detailed distribution maps—others may be more
ion
interested for the wealth of liberally referenced, genera- and species-specific information included
in the species discussions. 150 species are treated, approximé a one-half of the family in the Upper
Midwest (the region including, by definition, Illinois, Indiana, lowa, Michigan, Minnesota, and Wis-
consin). A full-page color photo for each species shows habit and habitat and smaller photos on the
facing page provide details. A distribution map for each species gives county-level distribution for
fe} “ i
the entire region (Appendix VI give references from which dic map data are derived). All Asteraceae
species of the region are included ina key at the back of the book adapted largely from Swink and
Wilhelm 1994°—Appendix IL), but here is an example of photos good enough to serve for eflective
ene identification, relegating the key to adjunct value (for example, compare among species
of Aster, Erigeron, E es Lactuca, and Lidtris). The Introduction includes echo mane infor
{ the family, general principles of nomenclat ture and classification, and habi-
tat and conservation, as well as “Comments on Distribution Maps” and “Comments on Photography”
(technical and otherwise: “Waiting for the wind to die down ina midwestern prair : totake
aneternity”
i context of general ees for this book, my ‘technical’ side can at least provide a
wish- . of features toward a second edition
rview. Synoptic list of all ad the | distri
(1) Ov maps forall species ~ these
could be added without accompanying photos and discussions
) Nomenclature. How did the authors decide what generic name to use? Readers are referred
to other floristic publications “for varying nomenclatural ei ken * but some generic con-
cepts used in the book have been modified within the last decade or longer with varying degrees ol
pes ance: Aster (= Doellingeria, Eurybia, lonactis, Symphyotric ae pane (= Arnoglossum),
Chr
rysopsis | recd). Eupdtorium (= Ageratind and Eupatorium sensu stricto), Gnaphalium (=
Ps ] ]
book. For other
end | . Senecio (= Packera) ~ not even synonyms are provided in the
genera, listed synonyms give notice of potential taxonomic variance: Chrysanthemum (AKA
Leucanthemum), Kuhnia (AKA Brickellia), Hymenoxys(AKA Actined. but not letraneuris not listed),
Microseris (AKA Nothocalais).
(3) Arrangement of species. The species are first arranged within four color classes—White or
Green, White with Yellow Centers, Yellow or Yellow with Brown Centers, and Blue, Violet, Rose or
Pink—then alphabetically by genus and species. | believe folks interested in Asteraceae, at almost all
-
levels of botanical sophistication, could appreciate the value of placing all species of one genus to-
gether and arranging the genera by relationship rather than eae Guy L. Nesom, Botani-
cal Research Institute of Texas, Fort Worth, TX, 76102-4060, U.
SIDA 20(1): 268. 2002
VASCULAR PLANTS OF KONZA PRAIRIE BIOLOGICAL
STATION: AN ANNOTATED CHECKLIST OF SPECIES IN A
KANSAS TALLGRASS PRAIRIE
E.Gene Towne
n of Biology
ee ssid See es ity
Trios KS 6 -4901, U.S.A.
oir
ABSTRACT
The native and naturalized vascular plants of Konza Prairie Biological Station, a 34+87-ha tallgrass
prairie research site located in the Flint Hills of northeastern Kansas, are listed along with their
primary habitat, relative abundance, flowering period, and life span. After more than 25 years of
ar flora of Konza Prairie consists of 576 species, representing 336 gen-
collections, the known vascul
era and 96 families. Families with the most species are Poaceae (84), Asteraceae Se Fabaceae ie
Cyperaceae (33) Non-native specie (n =96) co mpris se 16.7% of the total flora. W £
by life span, 64.6% of all species are perennials, 31.3% are annuals, and 4.2% are biennials. Species
I
are characte peueney) associated with a prairie habitat comprise 40.6% of the flora, followed by
4%), woodland (22.2%), and wetland (14.8%) habitats. In the prairie habitat,
uthway oc-
16.2% of the species are grasses andl 76.9% are herbaceous forbs. The C, photosynthetic pa
curs in 59.5% of all eu sais and in 70.8% of the annual grasses. The diversified habitats created
jeafl richness thatis likely greater
L
DY Konz a Prairie provide rflora
than ok of any other area of comparable size in the Great Plains region.
Key Worps: Prairie Flora, Vascular Plant Survey, Biodiversity, Species Richness
RESUMEN
| indi li das de la Estacion Biologica de la Pradera Konza, un sitio
e fais alto de 3487 hectareas en el noreste de Kansas, se listan
i. su habitat principal, abundancia relativa, periodo de floracion y su longevidad. Tras mas
de 25 anos de colectar, se han es 576 especies de flora vascular de la Pradera Konza, que
representan 336 géneros y 96 familias. Las familias con mayor cantidad de especies son Poaceae (84),
eae (79), Fabaceae (49) y Cyperaceae (33). Las especies no indigenas (n = 96) componen el
6.7% de la flora total. Cuando son clasificadas por su — 64.6% del total de especies son
cas de un habitat de praderas
L aS panies
vestigacion en una ade ra d
jen
perennes, 31.3% anuales y 4.2% bienales. Las especies caracteri
flora total, seguidas por las de bavieee aac (22.4%), forestal (22.2%)
componen el 40.6% de la
C, aparece en el 59.5% de las especies de pastos y en el 70.8%
y ea (14.8%). La ruta fotosintética
e los pastos anuales. Los habitats diversificados que se han producido a través de actividades de
} | ] lof] |
administracion e investigacion en la Pradera Kc [Ue |
sobrepasa cualquier otra area de t parable en esta region.
SITE OVERVIEW
Konza Prairie Biological Station is a 3487-ha tallgrass prairie located in the Flint
Hills of northeastern Kansas (39° 05' N, 96° 35' W, Fig. lL). The Nature Conser-
SIDA 20(1): 269 — 294. 2002
BRIT.ORG/SIDA 20(1
Konza Prairie
vancy initially purchased 371 ha of the site in 1971, followed by the remaining
procurement in 1977. The preserve is managed for ecological research by the
Division of Biology at Kansas State University. To study how fire and grazing
affect the ecosystem, Konza Prairie is parceled into 52 watershed units that are
burned at different frequencies and in different seasons, and are either ungrazed
or grazed by large herbivores. Mowed fireguards or gravel roads separate the
watersheds. Approximately 200 ha of agricultural land and abandoned brome
fields are scattered throughout the site. The headquarters area consists of old
homesteads with renovated buildings, a corral, and cultivated fields.
Topographically, the prairie landscape comprises upland plateaus, lime-
stone hillsides, and fertile lowlands. The upland soils are relatively shallow, silty
clay loams overlying limestone and shale layers (Udic Argiustolls and Lithic
Haplustolls), whereas the lowland soils are deeper colluvial and alluvial de-
posits (Pachic Argiustolls). Two major creeks meander through the lowlands,
but water flow is intermittent and dependent upon precipitation patterns. Per-
manent pools in the lower creek channels, localized springs, seeps, and several
—
—
TOWNE, CHECKLIST OF KONZA PRAIRIE 271
small stock ponds constitute the wetland habitat. Gallery forests (bands of de-
ciduous trees bordering the creeks and ravines) cover about 8.5% of the site.
Prairie vegetation is dominated by warm-season perennial grasses, with
numerous interstitial forb species. Although the grassland flora consists pri-
marily of tallgrass prairie plants, species that are characteristic of midgrass
prairies thrive on the uplands and hillsides, and many shortgrass prairie spe-
cies occur on the shallow, droughty sites.
For nearly 100 years, the site was a cattle ranch with both year-long and
season-long grazing operations. After acquisition by The Nature Conservancy,
Konza Prairie was not grazed by large herbivores until 1987, when a bison (Bos
bison) herd was established in a 992-ha internal enclosure. Seasonal grazing by
domestic cattle (Bos taurus) resumed in 1992 in 6 adjacent pastures (510 ha),
although not all are grazed every year. The remainder of Konza Prairie is
ungrazed by ungulates, except for a prolific white-tailed deer (Odocoileus
virginianus) population.
Konza Prairie has a temperate continental climate, characterized by hot
summers, cold winters, and moderately strong surface winds. Average annual
precipitation is 835 mm, with 75% of this occurring in the April through Sep-
tember growing season. The average frost-free season lasts 180 days. Mean an-
nual temperature is 13°C, with average minimum and maximum temperatures
ranging from -3°C in January to 27’ C in July. Elevations on Konza Prairie range
from 320 to 444 m.
FLORISTIC EFFORTS
After a decade of collecting, 436 species of vascular plants (plus 5 varieties)
were initially reported to occur on Konza Prairie (Freeman & Hulbert 1985),
with another 27 species subsequently added (Freeman & Gibson 1987). Six years
later, Kazmaier (unpublished pamphlet) listed 524 vascular plant species for
Konza Prairie. Because of Konza Prairie’s role as a benchmark tallgrass prairie,
it is essential that the flora inventory and pertinent ecological synopsis of the
species be current and readily accessible. With more than 70 new species dis-
covered since Kazmaier’s unpublished checklist, questionable validity of some
previously listed species, and extensive nomenclatural revisions, an updated
inventory of the Konza Prairie vegetation is necessary. Format for this flora fol-
lows standards proposed by Palmer et al. (1995). Voucher specimens are stored
in the Kansas State University Herbarium (KSC).
ORGANIZATION OF THE CHECKLIST
The vascular plants known to occur spontaneously on Konza Prairie are ar-
ranged alphabetically by family, genus, and species within four groups:
Pteridophyta (ferns and fern allies), Pinophyta (gymnosperms), Magnoliopsida
272 BRIT.ORG/SIDA 20(1)
(dicots), and Liliopsida (monocots). Each entry includes scientific name, com-
mon name, relative abundance, primary habitat, flowering period, and life span
(Appendix I). In addition, non-native species and the photosynthetic pathway
—
of grass species are listed.
Nomenclature follows Kartesz (1999), although authority abbreviations
conform to Brummitt and Powell (1992). A synonym is enclosed in brackets for
those species that have been annotated recently or if the scientific name changed
from that listed by Freeman and Hulbert (1985). Infraspecific taxa (subspecies
and varieties) are not delineated, although naturally occurring hybrids are
included.
The common name for each species was selected from regional sources
(Anderson 1961; Bare 1979; Great Plains Flora Association 1986) to reflect cus-
tomary usage for this area. Most species are identified with a single colloquial
name, but alternatives are presented ina few instances. Structure and format of
common names generally follow guidelines recommended by Kartesz and
Thieret 1991).
The listed habitat indicates where each species is most likely found on
Konza Prairie; however, it does not imply that the plant is restricted to that site.
Many species occur at different densities in heterogeneous locations. Designated
j=)
habitat categories for the Konza Prairie flora are:
wg?
Prairie—Open grassland that is subdivided by topographic Boe uplands
(clayey, cherty, or shallow soils on top of the hills), lowlands (deep bot-
tom land soils), and slopes (rocky hillsides). Additionally, some species
are — as occurring predominantly in grazed or infrequently burned
site
Woodland—Galler y forest and sites that are shaded by trees.
Wetland—Wet sites that include ponds, creeks, seeps, stock tanks, and mesic
lowlands.
Disturbed—Sites where the soil has been disrupted, include trails, wallows,
ditches, roadsides, homesteads, lawns, and ape fields.
(en,
me
a
Richness among the different plant habitats was estimated by assigning each
species to one habitat. Because many species occur ina variety of habitats, as-
signment to a specific category may be debatable. However, the criterion for
assigning species was based on which habitat the species is most characteristi-
cally associated with on Konza Prairie. Species that are found predominantly
in disturbed sites were not considered prairie plants even if they appear spo-
radically in grazed or unburned prairie. Likewise, woodland species that mi-
grate into unburned grassland were not grouped in the prairie habitat.
Relative abundance is an evaluation of how common and widespread a
species is for the particular habitat, and is based on extensive field experience
of the author. However, abundance of most annuals and some perennial species
may fluctuate yearly in response to precipitation ae and grazing or burn-
ing treatments. Descriptions of the five abundance classes are:
TOWNE, CHECKLIST OF KONZA PRAIRIE 273
Abundant—Species is dominant or co-dominant in the habitat.
Common—Species is found easily in the particular habitat.
Occasional—Species occurs sporadically but is not difficult to find.
Infrequent—Species is relatively scarce and not easily found.
Rare—Species is seldom encountered or found only in specific locations.
The flowering period is when peak blooming normally occurs on Konza Prairie
and often indicates when the species is most likely found. For vascular crypto-
gams, the dates represent spore formation. Flowering periods are based on field
observations, herbarium specimens, and phenology records for this area.
Each species is classified by life span as either annual, biennial, or peren-
nial. Non-herbaceous perennial plants were additionally subdivided into tree,
shrub, and woody vine categories. Although some biennials and short-lived
perennials may persist for only one year, designations were based on the ex-
pected longevity of the species for this area.
Grasses (Poaceae) are designated as either C3 or C4 species (Waller &@ Lewis
1979). The C3 grasses are cool-season plants that initially produce a 3-carbon com-
pound during photosynthesis and characteristically flower in May and June. The
C4 grasses are warm-season plants that initially produce a 4-carbon compound
in the photosynthetic pathway and characteristically flower in July and August.
Species that were not present in this region before European settlement are
considered introduced (Brooks 1986). Cultivated crops from the agricultural
fields (e.g., wheat, oats, sorghum, corn, and soybean), and other non-natural-
ized annual species are not included in this checklist. However, exotic horticul-
tural and ornamental species that were planted decades ago near the headquar-
ters area and have subsequently persisted, are listed in Appendix II.
RESULTS AND DISCUSSION
The known vascular flora on Konza Prairie consists of 576 species, represent-
ing 336 genera and 96 families (Table 1; Appendix I). Families with the most
species are Poaceae (84), Asteraceae (79), Fabaceae (49), and Cyperaceae (33).
Forty families are represented by a single species. The most species-rich genera
are Carex (17 species), Polygonum (12 species), Symphyotrichum (9 species), and
Ascle pias (9 species).
Twenty-three species from the last inventory (Kazmaier, unpublished pam-
phlet) are deleted from the present list because the specimens were misidentified
(n =8), had been reduced to synonymy (n = 4), or because there are no vouchers
and no bona fide evidence of their presence on Konza Prairie. Most of the un-
substantiated species appeared in preliminary inventories (see Freeman &
Hulbert 1985), and their unverified presence was subsequently perpetuated.
Taxonomic revisions of Kazmaier’s unpublished checklist also transferred 55
species into different genera and changed the specific epithet of 19 species.
274 BRIT.ORG/SIDA 20(1)
Taste 1.Summary of vascular plant families, genera, and species on Konza Prairie.
Species
Taxonomic group Families Genera Native Introduced Total
Pteridophyta 4 6 6 0) 6
Pinophyta | | 1 0 1
Magnoliophyta:
Magnoliopsida /6 254 349 73 422
Liliopsida 15 75 124 23 147
Total 96 336 480 96 576
When species are categorized to one predominant habitat, 40.6% of Konza
Prairie plants are characteristically associated with prairie grassland, followed
by disturbed (22.4%), woodland (22.2%), and wetland (14.8%) habitats (Table
2). In the prairie habitat, 16.2% of the species are grasses and 76.9% are herba-
ceous forbs. Only seven woody species are characteristically associated with
the prairie habitat; but without frequent fire, many woodland species invade
and proliferate. Species richness is lowest in wetland sites, but these areas are
the smallest and most fragmented habitats on Konza Prairie.
When partitioned into life span categories, 316 species (54.9%) are herba-
ceous perennials, 56 species (9.7%) are woody perennials, 180 species (31.3%)
are annuals, and 24 species (4.2%) are biennials. Within the prairie habitat, 76.1%
of the species are perennials, 20.9% are annuals, and 3.0% are biennials. In com-
parison, 23.3% of the species in disturbed sites are perennials, 69.0% are annu-
als, and 7.8% are biennials.
Non-native species (n = 96) account for 16.7% of the total flora. Families
with the most introduced species are Poaceae (21), Brassicaceae (13), Fabaceae
(9), and Asteraceae (8). Annuals and biennials comprise 59.4% of all introduced
species, compared to 30.8% of the native flora (Table 3). The highest density of
exotics is in the headquarters area where many species were planted intention-
ally or are cosmopolitan weeds. Most introduced species are either restricted to
disturbed soils or are weak competitors in native prairie. However, some inva-
sive species are capable of displacing native vegetation. Because the aggressive
ness of four exotic species (Bothriochloa bladhii, Bothriochloa ischaemum,
Carduus nutans, and Lespedeza cuneata) potentially threatens the integrity of
tallgrass prairie, they have been specifically targeted for eradication.
The C4 photosynthetic pathway occurs in 59.5% for all grass species and in
65.1% of the native grasses. Of the 24 annual grasses that occur on Konza Prai-
rie, 17 species (70.8%) use the C4 photosynthetic pathway. Most dicotyledons
use the C3 photosynthetic pathway, but 14 species representing five genera
(Amaranthus, Bassia, Chamaesyce, Portulaca, and Tribulus) have C4 photosyn-
a
a
TOWNE, CHECKLIST OF KONZA PRAIRIE 275
Taste 2. The number of vascular plant species within taxonomic groups that are characteristically
associated with different habitats on Konza Prairie.
Taxonomic group Prairie Disturbance Woodland Wetland
cea 0 0 5 1
Pin ) 0 1 0
we
Magnoliopsida 178 98 102 45
Liliopsida 56 31 20 39
Total numbers 234 129 128 85
Percent 40.6% 224% 22.2% 14.8%
Taste 3. The number and percent of native and introduced species within different life form catego-
ries on Konza Prair
Native Species Introduced Species
Life form Number % Number %
Total annuals 134 279 47 49.0
nual grasses 16 33 8 8.3
a alee forbs 111 23.) 39 40.6
Annual monocots 7 1.5 0 0
Biennial dicots 14 2.9 10 10.4
Total ee 332 69.2 39 40.6
Perennial g 47 9.8 13 135
an ae ot 176 36.8 18 18.8
Perennial monocots 53 11.0 2 ia
Woody 50 10.4 6 6.3
Vascular cryptogams 6 3 0 0
Totals 480 96
thesis (Downton 1975). All of the Konza Prairie forb species that utilize the C4
photosynthetic pathway are annuals
Although field collections of Konza Prairie flora have been ongoing for more
than 25 years, the compilation of vascular plants will continue to be dynamic.
New i ienugrants epee al annuals that fluctuate with precipitation patterns,
and super ficially similar species W ill undoubtedly be found in subsequent years.
Species also will disappear. Some species have not been observed on Konza Prai-
rie for more than 10 years and may be extirpated. Other species occur ina single
locale or consist of only a few plants. Few areas of tallgrass prairie have been
surveyed as intensively as Konza Prairie, and the diversified habitats created by
management and research activities provide a flora richness that is likely
greater than any other area of comparable size in the Great Plains region.
BRIT.ORG/SIDA 20(1)
APPENDIX |
Annotated Ch
PTERIDOPHYTA - Vascular Cryptogams
DRYOPTERIDACEAE — Wood fern famil
Woodsia obtusa (Spreng,) Torr; blunt-lobed cliff
srn-rare on shaded limestone outcrops; Jul;
ao
eel
EQUISETACEAE - Horsetail family
f quise etum laevigatum Braun; smoot
infrequent in wet sites; May-Jun;
) SCOUTING g
rush
perennial.
OPHIOGLOSSACEAE - Adder's-tongue family
m(L.) Sw. rattlesnake fern;
perennial.
Botrychium virginianu
occasional in woodlands; May-Jun
Ophioglossum engelmannii Pranth adder’s
re in woody lowlands; Apr—Jun;
tone
Ue; fe
perennial
PTERIDACEAE - Maidenhair fern family
albata (Pursh) Windham;
false cloak fern; rare
Argyrochosma d
Ne sa iena dea alas tal;
ellaea glabella Mett.ex Kuhn;smooth cliff brake;
rare on shaded limestone outcrops; May
Jun; perennial.
DIVISION PINOPHYTA - Gymnosperms
CUPRESSACEAE - ee oe
Jun wane virginial
al al
oodlanc me ie buried prairie; ‘Meh oe
=
a
ex
In
evergreen tree.
DIVISION MAGNOLIOPHYTA -—
Angiosperms
Class Magnoliopsida — Dicots
ACANTHACEAE - Acanthus famil
Ruellia humilis Nutt, fringe-leaf ruellia; common
on all prairie sites; Jun—Aug; perennial.
Ruellia strepens L.; limestone ruellia; occasional
in woodlands; May—Sep; perennial.
ACERACEAE - Maple family
Acer negundo L.; boxelder; occasional in
lands; Apr; tree.
AMARANTHACEAE - Pigweed family
nlitoides Watson; [Amaranthus
non in
wood
Amaranthus
graecizans]; prostrate pigweed; comr
disturbed sites and along roads; Jun—Sep;
annual.
Amaranthus pc
ecklist of Species for Konza Prairie
Matson; Palmer's piqweed;
infrequent in disturbed sites; Jul-Oct; annual.
Amaranthus retroflexus L.,rough pigweed; infre
entin cultivated fields and disturbed low
lands; Jul-Oct; introduced annual.
Amaranthus rudis Sauer; water-hemp;rare in wel
sites; Jul-Oct; annual.
ANACARDIACEAE - Cashew family
tica Aiton; aromatic sumac; infre
mal
que
Rhus aroma
quent on uplands and lowlands; Apr-May;
shrub.
Rhus glabra L..smooth sumac;common on shal
and slopes; May—Jun; shrub.
ntze; polison-ivy;
- woody
low uplands
Toxicodendron radicans (L.) Ku
common in woodlands; May-Jun;
vine.
NACEAE - Custard-apple family
jee ina triloba (L.) Dunal; pawpaw; rare in wood-
lands and along creeks; Apr-May; tree.
APIACEAE - Parsley family
Chaerophyllum procumbens (L.) Crantz; spread
ing chervil; infrequent at woodland edges
nd in unburned prairie; May—Jun; annual.
Chaerophyllum tainturieri Hook.; erect chervil;
infrequent at woodland edges; May—Jur
annual.
onium maculatum L.; poison-hemlock; infre
quent at woodland edges and in disturbed
sites; May-Jun; introduced biennial
Lomatium foeniculaceum (Nutt.) Coult. & Rose;
carrot-leaf lomatium; common on shallow
iplands and slopes; Apr-May; perennial.
Osmorhiza long! istylis (Torr.) DC.; long-style
rare in woodlands; May;
eee)
Sanicula canadensis L.; Canadian sanicle; occ
sional in woodlands; Jul-Sep; biennial.
Spermolepis inermis (Nutt.ex DC.) Mathias & Con
| | : | |-
stance;sc and
slopes; A ie
( 5.) Link; hedge-parsley;infre
s and in unburned
Torilis arvensis (Ht
aod| nd daec
quent at
prairie; Jun—Jul; introduc ed annual.
y, golden alexanders; occa
un; perennial.
Zizia aurea (L.) Koc
sional in WOO ae May
TOWNE, CHECKLIST OF KONZA PRAIRIE
APOCYNACEAE - Dogbane family
Apocynum cannabinum L.; hemp dogbane; oc-
asional on lowlands; Jun—Jul; perennial.
ASCLEPIADACEAE - Milkweed family
Asclepias lanuginosa Nutt.; woolly milkweed;rare
razed uplands; May; perennial.
Asclepias speciosa Torr, showy milkweed;rare on
plands and lowlands; Jun; perennial
Asclepias stenophylla A. Gray; narrow-leaf milk-
weed; occasional on uplands and lowlands;
Jun-—Jul; perennial.
Asclepias sullivantii Engelm. ex A. Gray; smooth
milkweed; occasional on lowlands; Jun;
nnial.
sae syriaca L.; common milkweed; occa-
nal on lowlands; Jun; perennial.
sce ias tuberosa L.; butterfly milkweed; occa-
on uplands and lowlands; Jun—Jul;
scene
Asclepias verticillata L.; whorled milkweed;com-
mon on all prairie sites; Jun—Sep; perennial.
Asclepias viridiflora Raf; green milkweed; occa-
nal on eee and slopes; Jun—Jul;
oS
fap)
ASC
perennial.
Asclepias ie Walter; spider milkweed; com-
mon on all prairie sites; May—Jun; perennial.
inanenti laeve wa ree Seng milk-
eed; i es and in
ee eniencs: Jul-Sep; ae
E - Sunflower family
Achillea millefolium L; western yarrow; common
on all prairie sites; May—Jun; perennial.
Ageratina altissima (L.) King & Rob, [Fupatorium
r oa mt snakeroot; Common in
nds; Aug—Oct; perennial.
Ambrosia arternisitalia L.; annual ragweed; oc-
nal in disturbed sites; Jul-Oct; annual.
penne psilostachya DC.; western ragweed;
on all prairie sites; Jul-Oct;
eren
aes ne L.; giant ragweed; Secasione) in
disturbed sites and at dl dges;Jul-
Oct; annual.
Amphiachyris dracunculoides (DC.) Nutt;
(Gutierrezia annua
broomweed; infrequent in disturbed sites;
ug—Oct; annual.
Antennaria neglecta Greene; field pussytoes;
common on uplands; Apr-May; perennial.
| / heen g 7
GFadCcuricuiOldesy],
277
Arctium minus Bernh.; burdock; infrequent in
woodlands and shaded disturbed sites; Jul-
Sep; introduced biennial.
Arnoglossum plantagineum Raf.; [Cacalia
plantaginea]; tuberous indian-plantain; oc-
casional on uplands and lowlands; Jun—Jul;
perennial.
Artemisia ludoviciana Nutt; Louisiana sagewort;
n all prairie sites; Aug-Sep;
perennial.
Bidens aristosa (Michx.) Britton; [Bidens polylepis];
a a aces ticks; infrequent in wet
5; Sep-Oct; annual.
innata L.; nee needles; rare in wet
ites; Sep-Oct; annual.
Bidens cernua L.; nodding beggar-ticks; infre-
wet sites; Sep—Oct; annual.
Bidens ae L.;devil’s a ee od
in wet sites; Sep-Oct; annual.
Brickellia eupatorioides (L.) Shinners; [Kuhnia
eupatorioides]; false-boneset; common on
lands and slopes; Aug—Sep; perennial.
Carduus nutans L.; musk-thistle; infrequent on
lowlands and in disturbed sites; May—Jun;
introduced biennial.
Cirsium altissimum (L.) Spreng,; tall thistle; com-
mon in unburned lowlands; Aug-Sep;
biennial.
Cirsium undulatum (Nutt.) Spreng.; wavy-leaf
thistle; Common on uplands and slopes;
Jun—Jul; perennial.
Cirsium vulgare (Savi) Ten.
turbed sites; Jun—Sep; introduced biennial.
Conyza canadensis (L.) Cronquist; Canadian
horseweed; common in disturbed sites; Jul-
Sep; annual.
Conyza ramosissima Cronquist; spreading
_ rare in disturbed sites; Jul-Sep;
ual.
Dpto eles (Vent.) Hitche.;fetid-marigold,
dogweed; occasional along roads;
u emcee
Echinacea angustifolia DC.; black-sampson
echinacea; common on ones and rocky
prairie; Jun: perennial
sa ne ae L.; yerba-de-tajo; rare along
Jul-Sep; annual.
ee tes hic (L.) Raf. ex DC; American
d; infrequent at woodland edges
ad me creeks; Aug—Sep; annual.
ae bip
bull th ul ist ale in dis
<7
+O
278
Erigeron annuus (L.) Pers.; annual fleabane; oc-
casional in disturbed lowlands; May-Jun;
nual.
Frigeron philadelphicus .;Philadelphia fleabane;
infrequent in mesic lowlands; Apr—Jun;
biennial.
Erigeron strigosus Muhl.e
C non uplands and in disturbed sites;
mes un; ania
Eupatorit ticciIm
VAL]
Willd. : daisy fleabat le;
L.; tall joe-pye-weed; oc-
casional on eee Aug-—Sep; perennial.
Euthamia gymnospermoides Greene; sticky
euthamia; infrequent on lowlands; Sep-Oct;
perennial.
Grindelia squarrosa (Pursh) Dunal; curly-cup
ed; common in disturbed sites and
along roads; Aug—Oct; biennial.
Helianthus annuus L.; common sunflower; com-
mon in ditches and disturbed sites; Jul-Oct;
nual.
i hirsutus Raf; hairy sunflower; com-
oodland edges; Jul-Oct; perennial.
Heli ants maximi il fani ee Maximilian’s
BRIT.ORG/SIDA 20(1)
Lactuca ee L. cou lettuce; infre-
rent at edges; Jul-Sep; biennial.
Lactuca ludovicia Nat) idl rouelie let-
tuce; oc¢
I DED,
biennial.
Lactuca saligna L.; willow-leaf lettuce;infrequent
odland edges and in disturbed sites;
Jul-Sep; introduced annual.
Lactuca serriola L; prickly lettuce; occasional in
unburned prairie and disturbed sites; Jul-
Sep; introduced annua
Liatris aspera Michx.; button gayfeather; infre-
quent on uplands and rocky prairie; Aug-
Sep; perennial.
Liatris mucronata DC.;eastern dotted gayfeather;
infrequent on uplands; Aug-Sep; perennial.
Liatris punctata Hook.; western dotted
gayfeather; common on shallow uplands
and rocky prairie; Aug-Sep; perennial.
ee lee nee ae cei
cuspidata]; wavy-leaf false-dandelion; infre
quent on uplands and slopes; Apr-May;
nial.
Aug-Oct; perennial.
ianthus pauciflorus Nutt, pice idus];
stiff sunfl uplands; Jul-
Soqeennal
Helianthus petiolaris Nutt; plains sunflower; rare
in disturbed sites; Jul-Sep; annual.
elianthus tuberosus L.; Jerusalem artichoke;
common at woodland edges; Aug-Oct;
perennial.
Heliopsis helianthoides (L.) Sweet; rough
heliopsis; infrequent on lowlands; Jul-Sep;
perennial.
Heterotheca canescens (DC.) Shinners;[(CArysopsis
canescens]; golden-aster; rare on uplands;
ug—Sep; perennial.
Hieracium longipilum Torr; long-beard hawk-
;occasional on uplands; Jul; perennial.
latest scabiosaeus (Her; white-bract
hymenopap old plainsman; occasional
on shallow ade and slopes; May—Jun;
biennial.
lva annua Ly annual sumpweed; rare in ditches
mesic disturbed sites; Aug—Oct; annual.
Krigia cespitosa (Raf.) Chambers; dwarf-dande-
lion; infrequent in mesic uplands and low
lands; May—Jun; annual.
He
—
f-
a
Oligoneuron rigidum (L.) Small; [Solidago rigidal;
stiff goldenrod; occasional on lowlands;
Aug—Oct; perennial.
Packera ene: (Nutt.) Weber & Love; [Senecio
plattensis]; prairie groundsel; common on
uplands; Apr-May; biennial.
Pseudognaphalium obtusifolium (L.) Hilliard &
Burtt; [Gnaphalium obtusifolium]; fragrant
cudweed; infrequent on uplands and low-
lands; Seo—Oct; annual.
atibida columnifera (Nutt.) Wooton & Standl.;
upright prairie-coneflower; common on
uplands and along roads; Jun—Jul; perennial.
Rudbeckia hirta L.; black-eyed susan; infrequent
n lowlands; Jun—Jul; biennial.
phium int he cas Michx.; whole-leaf rosin-
requent on lowlands; Jul-Aug;
Si
=
perennial
eye | | : |
on siden Jul Bie perennia al.
tall goldenrod, Canadian
Senne common on lowlands; Aug-Oct;
perennial.
Solidago gigantea Aiton; late goldenrod; infre-
quent in mesic lowlands; Sep—Oct; perennial.
Solidago missouriensis Nutt; Missourl goldenrod;
common on all prairie sites; Aug—Oct; perennial.
TOWNE, CHECKLIST OF KONZA PRAIRIE
Solidago mollis Bartl.; ashy goldenrod; rare on
uplands; Aug-Sep; perennial.
| granny 4
Bb / yg
quent on rocky prairie; Sep—Oct; perennial.
olidago speciosa Nutt, showy-wand goldenrod;
infrequent on uplands; Sep-Oct; perennial.
Sonchus asper (L.) Hill; prickly sow-thistle; infre-
quent in disturbed sites; Jun—Oct; intro-
uced annual.
sa ic serea ee ae Nesom;
aster-com-
mon in woodlands; ean Oct; say
Symphyotrichum ericoides (L.) Nesom; [Aster
ericoides]; heath aster; abundant on all prai-
rie sites; Sep-Oct; perenn
Symphyotrichum laeve (L.) ie [Aster laevis]:
smooth aster; occasional on uplands and
slopes; Sep—Oct; perennial.
Symphyotrichum lanceolatum (Willd.) Nesom;
[Aster si Moab panicled aster; occasional on
lowlands; Sep-Oct; perennial.
Symphyotri a oblongifolium (Nutt.) Nesom;
on uplands; Sep-Oct; perennial.
Symphyotrichum oolentangiense (Riddell)
Nesom [Aster oolentangiensis]; azure aster;
rare on lowlands; Sep-Oct; perennial.
Symphyotrichum pilosum (Willd.) Nesom; [Aster
pilosus]; hairy aster; infrequent on lowlands
and in abandoned brome fields; Sep-Oct
perennial.
Symphyotrichum sericeum (Vent.) Nesom; [Aster
sericeus]; silky aster; occasional on uplands
d slopes; Aug-Oct; perennial.
Symphyotchum subulatum (Michx.) Nesom; [As-
subulatus]; saltmarsh aster; infrequent in
peas any wet sites; Sep-Oct
LARISA VAI Als ae Fy
VVCVWCI
on
occasional around headquarters; Mar—Oct;
niente Derchila
lejfi
t’s-
peat) infrequent in n ditches and aioe
sites; May-Jun; introduced bien
Verbesina alternifolia (L.) Britton ex Dae
; common in woodlands; Aug-
Oct; p nial.
Vernonia ae inii Torr, western ironweed;com-
nall prairie sites; Jul-Sep; perennial.
i, strumarium L.; cocklebur; infrequent
in ditches and disturbed sites; Jul—Oct;annual.
279
BETULACEAE - Birch family
Ostrya virginiana (Mill.) Koch; hop-hornbeam;
rare on wooded slopes; Apr; tree.
BIGNONIACEAE - Bignonia famil
Campsis radicans (L.) Seem. ex Bureau; trumpet-
creeper; infrequent in woodlands; Jun—Aug;
oody vine
BORAGINACEAE - Borage family
Buglossoides arvensis (L.) Johnst.; [Lithospermum
a ; corn gromwell; infrequent in
ditches and disturbed sites; Apr-May; intro-
ced annual.
Cynoglossum officinale L.; hound's-tonque; rare
in woodlands; Jun—Jul; introduced biennial.
Hackelia virginiana (L.) Johnst.;Virginia stickseed;
occasional in woodlands; Jul-Aug; biennial.
Lithospermum canescens (Michx.) Lehm.; hoary
gromwell; rare on uplands and slopes; May;
perennial.
—
f Lehm.; leaf grom
well, fringed puccoon; occasional on up-
lands and slopes; Apr-May; perennial.
Myosotis verna Nutt.; Virginia forget-me-not; in-
frequent on uplands; May; annual.
KI
infrequent on uplands and slopes; Jun—Jul;
perennial.
BRASSICACEAE - Mustard family
Alliaria petiolata (Bieb.) Cavara & Grande; garlic-
mustard; Common in woodlands; Apr-May;
introduced biennial.
Arabis hirsuta (L.) a hairy rockcress; rare in
May—Jun; biennial.
Barbarea vul ce Br, wintercress; rare in ditches
nd disturbed sites; Apr-May; introduced
pere a
Camelina microcarpa DC.; little-pod false-flax;
infrequent along roads and in disturbed
sites; Apr-May; introduced annua
Capsella bursa-pastoris (L.) Medik.; shepherd’s-
purse; occasional in ae sites; Apr—
May; introduced a
Cardamine concatenata oe Schwarz;
[Dentaria laciniata]; cut- a rare
in woodlands; Apr-May; pere
ctr .) Desv.; hoary cress; rare in dis-
urbed sites; May; introduced perennial.
oo ae (Pallas) DC.; blue-mustard;
infrequent in disturbed sites; Apr-May;
introduced annual.
Descurainia ce, Waly Britton; pinnate
vustard; occ lin disturbed sites;
ene nual.
Descurainia sophia (L.) Webb ex Prantl; flixweed
nustard; rare in disturbed sites; May—
os voduced annual.
7 Nutt. ex Torr. & A. Gray; wedge-
a draba; occasional on shallow and rocky
o
lr, Ij
=)
Cc
1ov)
Draba reptans (Lam.) Fernald; white whitlow
wort; occasional on shallow uplands and
slopes; Apr-May; annual.
Erysimum repandum L.; bushy wallflower; infre-
quent in ditches and disturbed sites; Apr—
May; introduced annua
Hesperis matronalts L.,;dame’s rocket; rare around
fae eer eee ee acai aa
i Tie U 2 h
Lepidium densiflorum Schrad.; common
oe dues common on upianes and in
annual
Lepidium obl po Seal branched per ee
weed; occasional in disturbed sites an
along nore an Jun; annual.
Lepidium virginicum L.; Virginia pepperweed;
on in disturbed sites and along roads;
Apr-Jun; annual.
Microthlaspi perfoliatum (L.) M
iatum]; perfoliate pennycress; rare around
neadquarters; Apr; introduced annual.
Rorippa Dan egumn moss . .) Hayek; {Nas-
turtium officinale]; water s; occasional in
s; May-J cceeee perennial,
Pee) ustris (L.) Besser; bog yellowcress; rare
in wet sites; Le pea
Rorippa sessiliflora (Nutt.) Hitchc;stalkless yellow-
cress; rare in wet s ae me lal.
na sinuata (Nutt) Hitchc.;
infrequent along creeks; May Jun;
nial,
Tlaspaven nse L, nent peninyels ress;occasional in
|
-[Thlacni nerf
\ ag
oe Sets
>} dreadir IW ye lI IV
anntial
Ap
CACTACEAE - Cactus family
Escobaria missouriensis (Sweet)
[Coryphantha missouriensis]; nipple cactus;
dey on shallow uplands; Apr-May;
perennial.
a macrorhiza Engelm.;
Hunt.;
plains pricklypear;
BRIT.ORG/SIDA 20(1)
infrequent on shallow uplands; May-Jun
perennial.
CAMPANULACEAE - Bell-flower family
( ampanulastrum americanum (L.) Small; [Cam
panula americana]; American bellflower; in-
frequent in wet sites; Jul-Sep; annual.
delia cardinalis L.; red cardinal-flower; rare in
wet sites; Aug—Oct; perennia
Lobelia siphilitica L; blue cardinal-flower; infre-
quent in wet sites; Aug—Oct; perennial.
ae tocarpa (Nutt.) Nieuwl.,;slimpod Ve
ooking-glass; infrequent on uplands
wlands; May-Jun; annual.
eee perfoliata (L.) Nieuwl,; clasping Venus”
»oking-glass; occasional on uplands and
lowlands; May—Jun; annual.
CANNABACEAE - Hemp family
Cannabis sativa L.; marijuana, hemp; occasional
in ditches, at woodland edges, and around
headquarters; Jul-Sep; introduced annual.
Lo
ogans
CAPPARACEAE - Caper family
Polanisid dodec andra (L.) DC; clammyweed;
requent in disturbed sites and on slopes;
u ten seal
CAPRIFOLIACEAE — Honeysuckle family
Lonicera maackii (Rupr.) Maxim.; Amur honey
suckle; rare in shaded sites near headquar-
ters; May; introduced shru
Sambucus nigra L.; Se eies canadensis]:
rican aiderbertw: infrequent in wooc
lands; Jun—Jul; shrub.
. by | Pang |
n
Oo.
! M h;buckbrush;
common in woodlands and unburned low
lands; Jul; shrub
Iriosteum perfoliatum L.; horse-gentian; infre-
quent at woodland edges; May-Jun;
perennial,
CARYOPHYLLACEAE - Pink family
Arenaria serpyllifolia L; thyme-leaf sandwort; in-
t in disturbed sites; May—Jun; intro-
annual.
Cerastium brachypodum (Engelm. ex A. Gray)
ia emenee eed:rare in disturbed
sites and along roads; Apr; annual.
Dianthus armeria ae deptford pink; rare in dis-
dyed sites and along roads; Jun—Jul; intro-
ieee annual.
acd
OU
TOWNE, CHECKLIST OF KONZA PRAIRIE
Holosteum umbellatum L.; jagged chickweed;
occasional in disturbed sites; Apr-May; in-
troduced annual.
Saponaria officinalis L.; soapwort; rare around
| | | A . i J ee |
Silene antirrhina Li sleepy catchfly; occasional on
uplands; May—Jun; annual.
Silene stellata (L.) Aiton; starry silene; occasional
in woodlands; Jun—Jul; perennial.
Stellaria eee (Dumort.) Crep.; lesser chick-
ed; occasional around headquarters;
i ioe introduced annual.
CELASTRACEAE - Staff-tree family
Celastrus scandens L.; American bittersweet: in
requent in woodlands and on creek banks;
Ma vine.
Euonymus guoputnice Jacq.; eastern wahoo;
and on creek banks; May—
Tdi il
Jun; pais
CHENOPODIACEAE - Goosefoot famil
Bassia scoparia (L.) Scott; [Kochia scoparia];
ochia, fireweed; infrequent in disturbed
sites; Jul-Sep; introduced annual.
Chenopodium album L.; lamb’s-quarters; rare in
_ eee sites; Jun— SEP; introduced annual.
landi it-seed goose-
feet occasional in disturbed sites; Jun—Oct;
einual
henog I Standl goosefoot;
rare on rocky prairie; Jul- bab: annual.
Chenopodium simplex (Torr.) Raf; (Chenopodium
gigantospermum]; maple-leaf goosefoot; oc
casional at woodland edges and in dis-
turbed lowlands; Jul-Sep; annual.
Chenopodium standleyanum Aellen; Standley’s
ae sefoot; occasional in woodlands; Jul-Oct;
ual.
eee nuttalliana (Schult.) Greene; poverty-
weed; infrequent in ditches and disturbed
sites; Apr-May; annual.
CLUSIACEAE - St. John’s-wort one
Hypericum perforatum L.; common St. John’'s-
wort; infrequent on uplands ae lowlands;
ie Jul; introduced perennial.
CONVOLVULACEAE - Morning-glory family
Calystegia macounil (Greene) Brummitt; hedge
bindweed;infrequent in disturbed lowlands;
ay-Sep; perennial.
281
Convolvulus arvensis L.; field bindweed; infre-
quent in disturbed sites; May—Sep; intro-
duced perennial.
Evolvulus nuttallianus Schult.; Nuttall’s evolvulus;
infrequent on shallow and rocky uplands;
May-Jun; perennial.
Ipomoea hederacea Jacq.ivy-leaf morning- ee
infrequent in ditches and disturbed |
lands; Jun—Sep; introduced annual.
Ipomoea lacunosa L.;white morning-glory;infre
quent in disturbed recA nas
Ipomoea pandurata (L.) Mey.; big-root morning-
glory;infrequentin ae lowlands; Jun—
Sep; perennial.
CORNACEAE - Dogwood family
Cornus amomum Mill.; pale dogwood; rare in ra-
vines and on wooded slopes; May-Jun;
shrub.
Cornus drummondii Mey.; rough-leaf dogwood;
common in unburned prairie and wood-
lands; May-Jun; shrub.
CRASSULACEAE - Stonecrop family
Penthorum sedoides L.; ditch stonecrop; infre-
quent along creeks; Jul-Sep; perennial.
CUCURBITACEAE - Cucumber famil
Cucurbita foetidissima Kunth; buffalo gourd; in
ic wala in aie and along fence lines;
hin ug renn)
Sicyos engulotst L.; bur- eeedeaga in
ep; annual.
CUSCUTACEAE - Dodder family
casei igi gucialll cluster dodder, infre-
n lowlands; Jul-Sep; ann ual forb
arasitic on various Aoteyacee spec
Cuscuta indecora Choisy; showy doce ne
quent on lowlands; Aug-Sep; annual forb
parasitic on numerous dicot species.
ELAEAGNACEAE - Oleaster family
sacle pa aL.Russian-olive;rare along
cree ay—Jun; introduced t
EUPHORBIACEAE - Spurge family
Acalypha ostryifolia Riddell; hop-hornbeam
copperleaf; occasional in shaded lowlands
and disturbed sites; Jul-Oct; annua
Acalypha rhomboidea Raf.; rhombic copperleaf;
infrequent in shaded lowlands and dis-
turbed sites; Jul-Oct; annual.
=
282
Aca paar virginica L.; Virginia copperleaf; occa
yal in shaded lowlands and disturbed
Sees
Chamaesyce glyptosperma (Engelm.) Small; [Eu-
phorbia sence. ridge-seed mat-
spurge; infrequent on shallow uplands and
in disturbed sites; Jul-Sep; annual.
Chamaesyce maculata (L.) Small; [Euphorbia
maculata]; spotted mat-spurge; occasional
on grazed uplands and in disturbed sites;
Jul-Oct; annual.
CH : : Raf) cl Beall |
ly
missurica]; Missouri mat-spurge; infrequent
on rocky uplands; Jul-Sep; annua
Chamaesyce nutans (Lag.) Small; [Euphorbia
nutans]; eyebane; occasional in unburned
uplands and lowlands; Jul-Oct; annual.
Chamaesyce prostrata (Aiton) Small; eee
prostrata]; prostrate mat-spurge; occasional
along roads and in disturbed sites; Jul-Sep;
introduced annual.
Chamaesyce serpens Le Small: snptabi
serpens]; serpent mat-spurge,; infreque
grazed uplands He in disturbed sie
Oct.annual.
CH So
bia stic oinor slim- seed mat-spurge; infre-
quent in disturbed sites; Jul-Oct; annual.
Croton ieee Michx.; woolly croton; rare on
cky uplands; Jul-Sep; annu
Croton monanthogynus Michx.;one apnea croton;
sienerners on shallow uplands and slopes;
nual.
ae ie L.; flowering spurge; infre-
nae on uplands and lowlands; Jul-Sep;
nial.
bie: cyathophora Murray; painted spurge;
infrequent in disturbed sites; Jul-Sep;annual.
a acicls davidii Subils; western toothed
urge; infrequent in disturbed and un
burned sites; Jul-Sep; annual.
Euphorbia dentata Michx.; eastern toothed
urge; infrequent in disturbed and un-
ourned sites; Jul-Sep; annual.
BUP ROI neRagONG Do ex XO PIeng six-angle
A Sep;ar
Paphos! ia marginata Pursh; snow-on-the
ountain; occasional on uplands and in dis
turbed sites; Jul-Sep; annual.
) Small; [Euphor
nial
nual.
BRIT.ORG/SIDA 20(1)
aren spathulata Lam.; warty spurge; occa
yal on uplands and rocky sites; May—Jun;
rae
Tragia betonicifolia Nutt; nettle-leaf noseburn;
occasional on shallow uplands and rocky
sites; Jun— Aug; Perennial
Tragia ramosa J
on shallow delends ae rocky sites; Jun—Aug;
perennial.
FABACEAE - Bean family
Amorpha canescens Pursh; aap lant common
irie sites; Jun—Jul; shru
—_ 1a ae L; indigobush stereo oc-
ional in wet sites and along creeks; Jun—
ik ‘ rub.
Amphicarpaea bracteata (L.) Fernald; hog-pea-
nut; occasional on shaded creek banks; Aug
Sep; perennial.
Astragalus canadensis L.; Canadian milk-vetch;
rare on shaded slopes; Jul; perennial.
Astragalus crassicarpus Nutt; ground-plum milk-
vetch; occasional on uplands; Apr-May;
perennial.
Astragalus lotiflorus Hook.; lotus milk-vetch; in-
frequent on shallow uplands and slopes;
Apr-May; perennial.
Astragalus missouriensis Nutt.; Missouri milk
vetch; infrequent on shallow uplands and
slo eal
Astragalus plattensis Nutt; Platte River milk-vetch;
rare on rocky uplands; May-Jun; perennial.
Baptisia australis (L.) Br.ex Aiton; blue wild-indigo;
mmon on wae Reka ci
Raptis ia x bicolor Greenm. & Larisey; [B. australis
x B. bracteata]; ae wild-indigo; rare on
uplands; May; perennial.
Fyn ae ere ny ee
Casior al
o >
=,
+
wild-in
digo; common on all prairie sites; Apr-May;
perennial.
Cercis — : eastern redbud; occasional
in i q
C TIBMTACGING ial ita (Michx) Greene; [Cassia
oe ¢ Pa
on lowlands sae in ditches; Jul -Sep; annual.
Coronilla varia L.; crown-vetch; occasional on
airie adjacent to highway embankments;
Sep; introduced perennial.
C a Me sagittalis L.; rattlebox; rare on shaded
slopes; Jul-Sep; annual.
TOWNE, CHECKLIST OF KONZA PRAIRIE
Dalea aurea Nutt.ex Pursh;silk-top prairie-clover;
rare on shallow uplands; Jul-Aug; perennial.
Dalea candida Michx.ex Willd; white prairie-clo-
ver; comma
nial.
a hee ina (Aiton) Bullock; fox-tail prairie-clo-
er;rare in mesic lowlands; Aug—Sep; annual.
See multiflora (Nutt.) Shinners; round-head
rairie-clover; occasional on uplands and
slopes; Jul; perennial
Dalea purpurea Vent.; purple prairie-clover;
common on uplands and slopes; Jun—Jul;
erennial.
Desmanthus illinoensis (Michx.) MacMill. ex Rob.
& Fernald; Illinois bundle-flower; ara
on uplands and lowlands; Jul; perennial
Desmodium canescens (L.) DC.;hoary a ane
infrequent on lowlands; Aug—Sep; perennial.
Desmodium glutinosum (Muhl. ex Willd.) Wood;
large-flower tick-clover; occasional in wood
lands; Jul-Aug; perennial.
Desmodium illinoense A. Gray; Illinois tick-clover;
occasional on uplands and lowlands; Jul;
ou ial,
odium paniculatum (L.) DC.; panicled tick
ee ent at woodland edges; Aug—
Sep; perennial.
Desmodium sessilifolium (Torr.) Torr. & A. Gray;
sessile tick-clover; infrequent on lowlands
and slopes; Aug—Sep; perennial.
Gleditsia triacanthos L.; honey-locust; occasional
tree.
Glycyrrhiza lepidota Pursh; American licorice; in-
frequent on uplands and lowlands; Jun—Jul;
perennial
Gymnocladus dioicus (L.) Koch; Kentucky coffee-
tree; infrequent in woodlands; May; tree.
Kummerowia stipulacea (Maxim.) Makino; {Lespe-
eza stipulacea]; Korean lespedeza; occa-
sional in disturbed sites and along roads;
Au t; introduced annual.
ae capitata Michx,; ne head lespedeza;
common on lowlands; Aug-Sep; perennial.
Lespedeza cuneata (Dum. Cours.) Don; sericea
lespedeza; rare on uplands and lowlands;
Aug-Oct; introduced perennial
Lespedeza stuevei Nutt. tall bush-clover; rare on
lowlands; Aug—Sep; perennial.
non all prairie sites; Jun—Jul;
Lespedeza violacea (L.) Pers.; prairie lespedeza;
occasional on lowlands and slopes; Aug-
nial.
Medicago lputna
WN
laa medic; occasional in
Cc ; J ] |
Medicago sativa L.; alfalfa; rare in abandoned
brome fields and ditches; Jun—Sep; intro-
duced perennial.
Melilotus officinalis (L.) Lam.; yellow sweet-clov
ls wert good pec iall in a and
Mimosa nuttallii (OC) Turner: lat ad i];
t-claw sensitive-brier; occasional on up
and ares eee
Pediomelum argophylium (Pursh)
[Psoralea argophylla], silver-leaf scurfpea;
ccasional on lowlands; Jun—Jul; perennial.
ie esculentum (Pursh) Rydb.; [Psoralea
¢
uplands and slopes; May; perennial.
Psoralidium tenuiflorum (Pursh) Rydb.; [Psoralea
tenuiflora]; many-flower scurfpea; common
on all prairie sites; Jun; perennial.
Robinia pseudoacacia L.; black locust; rare in
odlands; May; tree
Senna marilandica (L.) Link; [Cassia marilandica];
wild senna; infrequent at woodland edges
and on lowlands; Jul-Aug; perennial.
Sei aet Nena wildbean;
shaded creek banks; Aug—Oct; annual.
Sophos eiosperma (Torr. & A. Gray) Piper;
th-seed besa: infrequent on low-
a Aug-Sep; annu
Trifolium pratense L.; a Cee infrequent
around headquarters; May—Sep; introduced
perennial
Trifolium repens L.; white clover; infrequent
ound headquarters; May—Sep; introduced
nnial.
Vicia americana Muhl.ex Willd; American vetch;
infrequent on lowlands and slopes; May-Jun;
perennial.
Vicia villosa Roth; hairy vetch; infrequent in
ditches and disturbed sites; May—Jun; intro-
duced annual.
FAGACEAE - Oak family
satay Macca Michx.; bur oak; common
Mavy-tree
May; LIOCe,.
Quercus muehlenbergli Engelm,chinquapin oak;
common in woodlands; May; tree
FUMARIACEAE - Fumitory family
Corydalis micrantha (Engelm.ex A.Gray) A. Gray;
slender fumewort; cl al on disturbed
sites; Apr-May; annual.
ee
rare in woodlands; Apr-May; perennial.
GENTIANACEAE - Gentian famil
Gentiana puberulenta Pringle; downy gentian
rare on lowlands; Sep—Oct; perennial.
RANIACEAE - Geranium family
Geranium carolinianum L.; Carolina crane’s-bill:
infrequent in disturbed sites and along
roads; May—Jun; annual.
GROSSULARIACEAE - Currant family
Ribes missouriense Nutt.; Missouri erent
-asional in woodlands; Apr-May; shru
nn Buckeye family
lus glabra Willd.; Ohio buckeye; infrequent
in woodlanes and along creeks; Apr-May;
shrub.
YDROPHYLLACEAE - Waterleaf famil
Ellisia nyctelea (L.) L.; waterpod; occasional in
woodlands and along creeks; May—Jun
annual.
JUGLANDACEAE - Walnut family
Carya cordiformis (Wangenh.) Koch; bitter-nut
ory; infrequent in woodlands; May; tree.
at walnut; occasional in
Sa
a ans nigra a
dland
LAMIACEAE - Mint family
ens) nepet toides (L.) Kuntze; catnip giant-
infrequent in woodlands; Aug—Oct;
perennial.
Hedeoma hispida Pursh; rough false-penny-royal;
occasional on grazed uplands and s
sites; May-Jun; annual.
lsanthus brachiatus (L.) Britton et al.;[7richostema
brachiatum]; rane rare on slopes and
rocky uplands; Jul-Sep; annual.
Lamium am ae L.; hen-
Mare
eit
around
yallow
bit; common
i} i J
annual
Lycopus americanus Muhl. ex Barton; American
bugleweed; infrequent in wet sites; Jul-Sep;
perennial.
Dicentra cucullaria (L.) Gene een meee
BRIT.ORG/SIDA 20(1)
Marrubium vulgare L.; common horehound; in-
frequent in shaded and disturbed sites; Jun—
Oct; introduced perenni
Ment
creeks; Jul-Sep; perennial.
Monarda fistulosa L.; mint-leaf bee-balm, wild
bergamot; occasional on lowlands and
Slopes; Jun—Jul; perennial
Nepeta cataria L., catnip; occasional in shaded dis-
turbed sites; Jun—Sep; introduced perennial.
Prunella vulgaris L.;selfheal; infrequent in wood-
lands and along creeks; Jun—Sep; perennial.
Salvia azurea Michx. ex Lam.; [Salvia pitcheri]:
Pitcher's sage, blue sage; common on up-
lands; Jul-Sep; perennial.
Salvia refleca Hornem.,; lance-leaf sage; occa-
sional in disturbed sites and along roads;
Jun—Sep; annual.
Scutellaria parvula Michx.; small skullcap; infre
quent on uplands and rocky sites; May—Jun
perennial.
Teucrium canadense L.; A abd germander;
ommon on lowlands; Jun—Jul; perennial.
LINACEAE - Flax family
Linum sulcatum Riddell; grooved flax; common
on uplands; Jun—Jul; annua
LOASACEAE - Stick-leaf family
Mentzelia oligosperma Nutt. ex Sims; stick-leaf;
infrequent on slopes and rocky uplands;
Jun—Sep; perennial.
LYTHRACEAE - Loosestrife famil
Ammannia coccinea Rottb.; purple toothcup;in-
frequent in wet sites; Jul-Sep; annual.
Lythrum alatum Pursh; winged loosestrife; occa-
sional in wet sites; Jun—Aug; perennial.
MALVACEAE - Mallow famil
Abutilon theophrasti Medik. velvetleaf; occa-
sional in disturbed lowlands and cultivated
fields; Jun—Oct; introduced ual.
iieeale aeoides (Michx.) A. ae pale poppy-
vallow; occasional on uplands and low
me s; May-Jun; perennial.
Callirhoe involucrata (Torr. & A. Gray) A. Gray;
purple poppy-mallow; occasional on up-
ands and lowlands; M
va arvensis L.; field mint; cae quent along
O
—
Ca
ay—Jul; perennial.
Hibiscus trionum L.;flower-of-an-hour infrequent
in disturbed lowlands and cultivated fields;
Jun—Sep; introduced annual.
TOWNE, CHECKLIST OF KONZA PRAIRIE
Malva neglecta Wallr; common mallow; infre
quent in disturbed lowlands; Jun—Oct;intro-
duced annual.
Malvastrum hispidum (Pursh) Hochr.; rough false-
mallow; infrequent in disturbed and un-
burned sites; Jun—Oct; annual.
Sida spinosa L.; prickly sida; ieaeeh in dis-
turbed lowlands and cultivated fields; Jul-
Oct; introduced annual.
MENISPERMACEAE - Moonseed family
Meni ispermum canadense L.; moonseed; com
mon in woodlands; May-Jun; woody vine.
MOLLUGINACEAE - Carpetweed family
Mollugo verticillata L.. carpetweed; infrequent in
disturbed sites; Jul-Sep; introduced annual.
CEAE - Mulberry family
Maclura a (Raf.) Schneid.; osage-orange,
apple; infrequent in unburned sites
and ine May; introduced tr
Morus alba L.; white mulberry; sean in
woodlands; May; introduced tr
NYCTAGINACEAE - Four-o’clock family
Mirabilis albida (Walt.) Heimerl;white four-o'clock;
infrequent on uplands and lowlands; Jun—
Sep; perennial.
Mirabilis (Pursh) Heimerl; narrow-leaf
four- o'clock, infrequent on uplands; Jun-Sep;
perennial.
Mirabilis eeaines (Michx.) MacMill. wild four
o'clock; occasional at woodland edges and
in disturbed sites; May—Sep; perennial.
OEENGERE DINE tami
ash:com-
Fall d “Apr; r
mon in prt
ONAGRACEAE - Evening-primrose family
Calylophus serrulatus (Nutt.) Raven; serrate-leaf
evening-primrose; infrequent on uplands
and slopes; May—Jul; perennial.
Gaura mollis James; [Gaura parviflora]; small-
flower gaura, velvet gaura; infrequent in
ditches and disturbed sites; Jul-Sep; annual.
Oenothera biennis L common evening-prim-
rose; infrequent in disturbed sites; Jul-Sep;
biennial.
Oenothera laciniata Hill; cut-leaf evening-prim-
rose:rare in disturbed sites; May—Jul; annual.
Relic cee
Oenothera macrocarpa Nutt.; Missouri evening-
285
primrose; occasional on shallow uplands
and slopes; May—Jun; perennial.
poe speciosa Nutt.; showy evening-prim-
occasional in mesic lowlands and
ditches; Jun—Jul; perennial.
Stenosiphon linifolius (Nutt. ex James) Heynh.;
flax-leaf stenosiphon; occasional on uplands
and slopes; Aug—Sep; biennial.
OROBANCHACEAE - Broomrape family
Orobanche uniflora L.; one-flower broomrape;
n lowlands; May; annual forb parasitic
on roots of various dicots.
OXALIDACEAE - Wood-sorrel family
Oxalis dillenii Jacq.; gray-green wood-sorrel; in-
{in woodlands; Apr-
ue ent on upla nds
un; perenni |
Oxalis stricta L., yellow wood-sorrel; occasional
on oa and in woodlands; Apr-Jun;
perennial.
Oxalis ieee L. violet wood-sorrel; occasional
on lowlands; Apr-May; perennial.
PAPAVERACEAE - Poppy family
Argemone eee (Fedde) Ownbey;white
prickly-poppy; infrequent on slopes and
shallow ater es Sep; annual.
iis lee iaenieal jenna family
tolacca americana L.; pokeweed; occasional
a roodlands and disturbed lowlands; Jun-
Sep; perennial.
PLANTAGINACEAE - Plantain family
Plantago aristata Michx.; bottle-brush plantain;
i | | qd Aj 1; } ej
p
Jun—Jul; annual.
Plantago lanceolata L.; buck-horn plantain; rare
in disturbed sites; Jun—Aug; introduced pe-
rennial.
Plantago patagonica Jacq.; woolly plantain; oc-
casional on uplands and slopes; Jun—Jul;
annual.
Plantago rhodosperma Decne.; red-seed plan-
tain; common on uplands and in disturbed
May-Jun; annual.
noe Beir i Decne.; black-seed Spite
mmon in woodlands and shaded d
ae 1 sites; Jun—Sep; perennial.
ee virginica L.; pale-seed plantain; occa-
sional on uplands and in disturbed sites;
May-Jun; annual.
286
seetisdileensi am Sycamore famil
nus occidentalis L.; sycamore; infrequent in
\ diaper Ap MAytieS
V IANS, API—-Midy, (ree.
POLEMONIACEAE - Polemonium family
Phlox divaricata L.; blue phlox; occasional in
woodlands; Apr-May; perennial.
OLYGALACEAE - Milkwort family
Polygala verticillata L.; whorled milkwort; infre
quent on uplands and rocky sites; Jun—Sep;
annual.
POLYGONACEAE - Buckwheat family
Polygonum lial cus L; water smartweed rare
in ditches and wet sites; Jul-Sep; perennial.
Polygonum aviculare \.; prostrate knotweed;
common on trails and in heavily disturbed
sites; May—Oct; introduced annual.
Pol sete bared L; i water-pepper,;
are in wet sites; Jul-Oct
eee ee ise mild water
pepper; rare in wet sites; Jul-Oct; perennial.
Polygonum lapathifolium ies smartweed: in
frequent in wet sites; Jul-Oct; annual.
Polygonum pensylvanicum ve Pennsylvania smart-
ed;infrequent in wet sites; Jul-Oct;annual
Polygonum persicaria L.; lady’s-thumb smart
weed; infrequent in wet sites; Jul-Sep; intro-
ausen annual.
ip) cH a Hl
toes,
infre eaten in wet sites; ae Oct; perennial
Polygonum ramosissimum Michx.; bushy knot-
weed; infrequent in disturbed sites and
along roads; Aug—Oct; annual.
Polygonum scandens L.; false buckwheat; infre
quent at woodland edges and in disturbed
sites; Aug-Sep; perennial.
olygonum tenue Michx.; slim knotweed; infre
quent on uplands; Jul-Oct; annual.
Polygonum ne ee ee 2illg smartweed;
occasional in woodlands; Jul-Sep; per
Rumex altissimus Wood; pale dock; infrequent in
ditches and disturbed lowlands; Apr-Jun;
perennial.
Rumex an L..curly dock;infrequent in ditches
and disturbed sites; May—Sep; introduced
perennial.
Aa li hacseilicat Purslane family
mon purslane; rare in
disvarbed: sites; ee introduced annual.
BRIT.ORG/SIDA 20(1)
PRIMULACEAE - Primrose family
Androsace occidentalis Pursh; western rock-jas-
mine; infrequent on shallow uplands and
rocky sites; Apr-May; annual.
sites and along creeks; Jun—Jul; perennial.
RANUNCULACEAE - Buttercup famil
Anemone caroliniana Walter; Carolina anemone;
infrequent on uplands; Apr-May; perennial.
Anemone virginiana L.; tall anemone; rare in
lands; Jun—Jul; perennial.
Clematis pitcheri Torr. & A. Gray; Pitcher's clema-
>in woodlands; Jun—Jul; perennial.
ascent carolinianum Walter; [Delphinium
scens]; plains larkspur; infrequent on Uup-
a eae perennial.
Myosurus tiny mousetail; rare in wet
sites; Apr May: annual.
Ranunculus abortivus L.little-leaf buttercup; rare
in wet sites; Apr-May; biennial.
ictrum dasycarpum Fisch. & Ave-Lall.; purple
meadow-rue; rare in woodlands; May-Jun
perennial.
RHAMNACEAE - Buckthorn family
Raf; New J
a
tis; rare
Tha
ante
Ceanothus herbaceus
rsey tea, inland
ceanothus; common in unburned uplands
and slopes; Apr-May; shrub.
ROSACEAE - Rose famil
Agrimonia pubescens Wallr; downy agrimony;
at woodland
Jul-Aug; perennial.
Crategus m dllis Scheele: downy hawthorn, sum-
mer haw; rare on wooded slopes and along
creeks; Apr-May; shrub.
faci) Ganadenseage cq.;white
e avens;common in
dlands and unburned lowlands; Jun—Jul;
nial,
Potenti aoe Pursh; tall cinquefoil; infrequent
on lowlands; Jun—Jul; perennial.
Potentilla recta L.; sulphur cinquefoil; rare in dis
turbed sites; May—Jun;introduced perennial.
Prunus americana Marshall; wild plum; occa-
sional along creeks and in unburned sites;
~Apr; shrub.
Prunus angustifolia Marshall; chickasaw plum;
infrequent along creeks and in unburned
sites; Apr; b.
Prunus virginiana L.; choke cherry; infrequent
along creeks and in unburned sites; Apr-
May; shrub.
:
-
TOWNE, CHECKLIST OF KONZA PRAIRIE
R h Porter; Arkansas rose; Common
in unburned pein May—Jun; shrub.
Rosa multiflora Thunb.; multiflora rose; rare in
woodlands and unburned lowlands; May-
at Iniioeaceg shrub.
blac k raspberry OEE asional
Jun; shrub.
Rubus ostryifolius Rydb.; high-bush blackberry;
common in woodlands and unburned low-
lands; May-Jun; s
RUBIACEAE - Madder family
Galium aparine L.; catch-weed bedstraw; occa-
sional in woodlands and unburned low-
lands; May-Jun; annual.
Galium circaezans Michx.; woods bedstraw; oc-
casional in woodlands; May—Jun; perennial.
Galium an OER MICOS sneak Saati ecco
Hedyot ee (Lam.) ae coreanast
bluets occasional on uplands and slopes;
Jun—Ju ie ial.
Houstonia i ean [Hedyotis crassifolia];
small bluets; rare on shallow uplands; Apr—
May; annual.
RUTACEAE - Citrus family
Zanthoxylum americanum Mill; prickly-ash; infre-
quent in woodlands and unburned low-
lands; Apr; shrub.
ALICACEAE - Willow family
Populus deltoides Bartram ex Marshall; cotton-
wood; occasional in wet sites; Apr
ow; infrequent in wet sites; May; tree.
Salix caroliniana Michx.; Carolina willow; infre-
quent in wet sites; Apr; tr
Salix exigua Nutt.; en wlcd: occasional in
wet sites; Apr; shrub.
Salix nigra Marshall; black willow; infrequent in
wet sites; Apr; tree
SANTALACEAE - Sandalwood family
omandra umbellata (L.) Nutt.; bastard toad-flax;
common on shallow uplands and slopes;
Apr-Jun; perennial.
Se OR OE A RIRCEAE Figwort family
galir Dougl. ex Benth.) Britton;rough
paar een on uplands and slopes;
Aug—Sep; annual.
Agalinis densiflora (Benth.) Blake; [Tomanthera
densiflora]; a ear nally: ORDIONG rare on
y Sep; annual.
ane rotundifolia (Michx) Wettst: water-hys-
sop;rare in shallow water and mud; Jul-Aug;
perennial.
Leucospora multifida (Michx.) Nutt.; paleseed;
UniSueniey in wel ales Aan Sep; annual.
=
Ah
=
infre equent in wet sites; May-—Aug; perennial.
Penstemon cobaea Note; eodars ies ono a
t Jun;
perennial.
Penstemon grandiflorus Nutt; shell-leaf beard-
tongue, large beardtongue; infrequent on
uplands and slopes; May—Jun; perennial.
lata ve Nh ett tr ites |
on uplands and slopes; May—Jun; perennial.
es marilandica L.; Maryland figwort;
ial in woodlands; Jul-Sep; perennial.
on banana L.:moth mullein; occasional
in ditches and disturbed sites; May—Sep; in-
troduced bie
Verbascum ee ‘ are mullein;infrequent
in ditches and oo sites; Jun—Sep; in-
duced biennia
vee age LE els Ae seen Oecdslorial
Apr- Jun;introduced annual.
agallis-aquatica L.; water speedwell;
occas onal in wet sites; May—Sep; perennial.
Veronica arvensis L., corn speedwell; infrequent
Dee | PE | i} A NA * J {
annual.
Veronica peregrina L.; purslane speedwell; occa-
sional in wet sites; Apr-May; annual.
SOLANACEAE - Nightshade family
wae stramonium L.; jimsonweed; rare in dis-
ded sites; Jul-Oct; introduced annual.
Psa rede Moe A. ‘Oley: neal leat grou und-
ee heterophylla Nees; clammy und-
herry; infrequent on lowlands; ie we
nial.
bie pubescens L, CNY oodles a rare in
Dh ye ear a
c
on all atti sites; iba —Sep; ea
Bie virginiana Mill; Virginia ground-cherry;
occasional on lowlands and in disturbed
sites; Jun—Sep; perennial.
Solanum ee - is horse-nettle;
occasional on lowla ea and in disturbed
sites; Jun—Sep; f nnia
Solanum eee Du a black nightshade;
rare in wet sites and ditches; Jun—Oct; annual.
Solanum r foie poral ue bur nightshade;
TILIACEAE —- Linden famil
Tilia americana L., basswood: rare in woodlands;
Jun; tree.
ULMACEAE - Elm family
Celtis occi Sea alis L.; hackberry; common in
Ayr M a
AO, PN
Lf limy ea) ric al
ay; tree.
Peas elm;common in
woddands Mar; tree.
Jimus pumila Lz Siberian elm; occasional in
dlands; Mar-Apr; introduced tr
Ulmus ee Muhl,; red elm, slippery aes ane
pes,
quen in oodlands and along creeks; Mar—
aera
URTICACEAE -— Nettle family
Laportea canadensis (L.) Wedd.; Canadian wood
nettle; infrequent in woodlands; Jul-Sep;
perennial.
Parietaria pensylvanica Muhl. ex Willd.; Pennsy!|
a pellitory;infrequentin woodlands and
ae rned lowlands; Jun—Sep; annual.
Pilea coe (L.) A. Gray; clearweed; occasional
wet sites; Aug—Oct; annual.
ca dioica L.; stinging nettle; common in
woodlands; Jul-Sep; perennial.
RBENACEAE - Verbena family
Glandularia Ane fida (Nutt.) Nutt,
bipinr WC
Urtic
- [Verbena
tific kota verbena;occasional on
slopes and a sites; May—Jul; perennial.
Glandularia canadensis (L.) Nutt.; [Verbena
Feber rose verbena; infrequent on
es; Apr-Jun; perennial.
Phoyme leptostachya L.; eed saan in
od
Jul ug; p
lands; Jul-Aug; p
ae lanceolata (Michx) Greene; [Lippia
lanceolata]; northern fogfruit; occasional in
we lune Co
—11d,
bigbract verbené na; occasional in csturbe
sites and along roads; Jun—Aug; perenni
we
BRIT.ORG/SIDA 20(1)
Verbena hastata L.; blue verbena; infrequent in
mesic lowlands and along creeks; Jul-Sep;
perennial.
Verbena simplex Le
oe
ym.;narrow-leaf verbena; rare
opes and rocky sites; Jun—Aug; perennial.
verbena stricta Vent.; woolly verbena; occasional
on uplands and along roads; Jun—Sep; perennial.
Verbena urticifolia L; white ial nettle-leaf
erbena; occasional |
creeks; Jul-Aug; perer cick
VIOLACEAE - Violet family
Hybanthus verticillatus (Ortega) Baill; nodding
saat et; i infreq uent on uplands and
slopes; May; pere
tola : icolor Pursh; i ee rafinesquii]; johnny
a 1p, field pansy; occasional on uplands
n disturbed sites; Apr- -May; annual.
a) ie ifida Don prairie viol
all prairie sites; Apr-May; perennial
/iola sana al Greene; Viola pratincolal;
common violet; common in woodlands;
Apr-Jun; perennial.
Nilld. ;downy blue | inf
in woodlands; Apr-Jun; perennial.
VITACEAE - Grape family
Parthenocissus quinquefolia (L.) Planch.; Virginia
creeper; common in woodlands; Jun—Jul:
woody vine.
Vitis riparia Michx.; riverbank
in woodlands; May; ey vine.
ZYGOPHYLLACEAE - Caltrop famil
Tr
=.
-as
along
Vio
—
et O -casional on
<
\ fola ororia
r
re
bulus terrestris L.; puncturevine; infrequent in
disturbed sites and along trails; Jul-Sep; in-
troduced annual.
Class LILIOPSIDA — Monocots
AGAVACEAE - Agave family
Yucca glauca Nutt; small soapweed, yucca; in
requent on uplands and slopes; May—Jun;
perennial.
ALISMATACEAE - Water-plantain family
Alisma subcordatum Raf; subcordate water-plan
tain;infrequent in wet sites; Jul-Sep; perennial.
Echinodorus berteroi (Spreng.) Fassett;
[E -hinodorus
in wet sites; Jun—Aug; annual.
ee a latifolia Willd; common arrowhead;
nfrequent in wet sites; Jul-Aug; perennial.
TOWNE, CHECKLIST OF KONZA PRAIRIE
ARACEAE - Arum family
Arisaema dracontium eee dragonroot;rare
oodlands; May; perennial.
COMMELINACEAE - y
— erecta L,; ie dayflower; wa
woodland edges; Jun—Oct; perennial.
a ae bracteata Small ex es bracted
spiderwort; occasional in mesic lowlands;
ae Jun; oan
far nil
IUCTWOTL
Raf.; Ohio spiderwort; rare
in mesic lowlands; May—Jun; perennial.
CYPERACEAE - Sedge family
Bolboschoenus fluviatilis (Torr.) Sojak; [Scirpus
uviatilis]; river bulrush; rare around ponds;
Jun-Jul; perennial.
Carex albicans Willd. ex Spreng.; [Carex artitecta];
white-tinge sedge; infrequent in woodlands;
May—Jun; perennial.
=>
C
on ipland
Ju
Carex blanda Dewey; wosdiena an occa-
sional in woodlands and along creeks; Apr—
Jun; perennial.
Carex brachyglossa Mack.; [Carex annectens|:yel-
low-fruit sedge; occasional in wet sites; May—
June; perennial.
Carex brevior (Dewey) Mack. ex Lunell; straw
-common on uplands and lowlands;
May-Jun; perennial.
Carex davisii Schwein. & Torr; Davis’ sedge; rare
oodlands; May-Jun; perennial.
Carex mee anes lal s sedge; rare in wet
s;Ma nial.
bs a eunthi ai sedge; infrequent in
wet sites; May—Jun; perennial.
cGiex glace Paley Healy Senge common on
| June rennia a|
ULI Crelit tal,
Carex grisea Wahlenb.; [Carex amen bola]; nar-
w-leaf sedge;common in woodlands and
: Apr-Jun; perennial.
Carex hyalinolepis Steud.; shoreline sedge; rare
in wet sites; May—Jun; perennial.
p ip NA ih Vid chart]
common in wet sites; May—Jun; perennial.
; a4 r Pate Ea ea oe {
abundant on uplands; Apr-May; perennial.
Carex meadii Dewey; Mead's sedge; abundant on
all prairie sites; Apr-May; peren
Carex molesta Mack. ex Bright; pest ihe occa-
sional on a and in wet sites; May—
June; per
Carex pellita ioe ex Willd.; [Carex lanuginosa];
woolly sedge; occasional in wet sites; May—
Jun; perennial.
Carex vulpinoidea Michx.; fox sedge; occasional
in wet sites; May—Jun; perennial.
Cyperus acuminatus Torr. & Hook. ex Torr; tape-
leaf flat-sedge; occasional in wet sites; Jul-
ep;annua
Cyperus bipartitus Torr; [Cyperus rivularis]; brook
lei infrequent in wet sites; Jul-Sep;
an
ee i. L; yellow nut-sedge; rare in
esic lowlands and ditches; Aug-—Oct;
perennial.
lef F
Cyperus x Resoenon Us Ge ise; [C. qs x C,
oe intermediate flat-sedge; infre
que uplands; Jun—Jul; perennial.
Cyperus od L.; rusty flat- fice occasional
in wet sites; Jul-Sep; annual.
Cyperus squarrosus L.; cae aristatus]; awned
lat-sedge; rare in wet sites; Jul-Aug; annual.
Cyperus ie L. straw flat-sedge; infrequent
in wet sites; Aug—Sep; perennial.
Eleocharis compressa Sull.,; flat-stem spike-rush;
nin wet sites; Apr—Jun; perennial.
Eleocharis erythropoda Steud,;red-foot spike-rush;
common in wet sites; May—Jun; perennial.
Fleocharis macrostachya Britton; long-stem spike-
rush;common in wet sites; POS Jun; pabily
iF
mbrella |
Fuirena simplex Vahl; u
in wet sites; Jul-Sep; annua
Schoenoplectus tabernaemontani (Gmel.) Palla;
[Scirpus validus]; soft-stem bulrush; occa-
sional in wet sites; Jun— ek, ; perennial.
eich
VAL ITA
76 wet sites; Jun—Jul; perennial.
Scirpus pendulus Muhl.; drooping bulrush; com-
mon in wet sites; May—Jul; perennial.
IRIDACEAE - Iris family
Sisyrinchium campestre Bicknell; prairie blue-
eyed-grass; common on uplands; Apr-May;
perennial.
JUNCACEAE - Rush family
Juncus dudleyi Wiegard; Dudley's rush; occa-
sional in wet sites; Jun—Aug; perennial.
290
Juncus interior Wiegard; inland rush; common
in c uplands and lowlands; May—Aug;
perennial.
Juncus torreyi Coville; Torrey's rush; common in
wet sites; Jun—Aug; perennial.
MNACEAE - Duckweed family
Lemna minor L.; lesser duckweed; rare in shal-
low water; Jul-Sep; perennial.
LILIACEAE - Lily family
Allium canadense L.; wild onion; occasional on
uplands; Apr—Jun; perennial.
Allium stellatum Nutt. ex Ker Gawl.; pink onion;
infrequent on uplands; Aug-Sep; perennial.
ei officinalis L.; asparagus; rare at wood-
lan ges; May-Jun; introduced perennial.
een um es white fawn- vty infre-
: Apr, perennial.
Ornithogalum umbellatum L,star-of Bet hlehem;
rare around headquarters; Apr-May;
duced perennial.
pal Ry \AJ4|
intro-
seal; rare in woodlands; Jun; perennial.
Zigadenus nuttallii (A. Gray) Watson; Nuttall’s
as; occasional on uplands and
lowlands; May-Jun; perennial.
NAJADACEAE - Naiad famil
Najas guadalupensis (Spreng.) Magnus;common
naiad; occasional in ponds; Aug—Sep; annual.
ORCHIDACEAE - Orchid family
Spiranthes cernua (L.) Rich. nodding ladies’
sses;rare on lowlands; Sep-Oct; perennial.
Spiranthes ee. Engelm. & A. Gray; spring
ladies’-tresses; rare on lowlands; Jul-Aug;
perennial
POACEAE - Grass family
Aegilops cylindrica Host; jointed goat grass; rare in
isturbed sites; May—Jun; introduced annual C,.
Agrostis hyemalis (Walter) Britton et al,; tickle
gra inter bent grass; occasional on up-
lands; May —Jun; perennial C
Agrostis ae L.; ie in eauent along
creeks; Jun—Jul; introduced perennial C,.
Alopecurus hee Walter, Carolina Foscail
infrequent in wallows and mesic uplands;
Apr—Jun; annual C
Andropogon ie vaeeer big bluestem;
abundant on all prairie sites; Jul-Oct; peren-
nial C,
=
a &
BRIT.ORG/SIDA 20(1)
Aristida oligantha Michx.; prairie threeawn,; infre
ent on disturbed sites; Aug—Sep; annual C,.
Aristi 2" purpurea Nutt; purple threeawn; iafie:
ane onu Peas and rocky sites; Jun—Aug;
a <a - se (Retz.) Blake; [Andropogon
bladhi i Coleen ln all common on
|
ueoce introduced perennial C,,.
Bothriochloa ischaemum (L.) Keng; [Andropogon
DENG a pee common
ir il-Oct; introduced perennial C
we ae loa laguroides (DC.) Herter; aingienbaon
saccharoides]; silver bluestem; rare in ditches
and disturbed sites; Jul-Sep; perennial C,,
Bouteloua curtipendula (Michx.) Torr; side-oats
grama; abundant on uplands and slopes;
Jul-Sep; perennial C,.
a gracilis (Willd. ex Kunth) Lag. e
Griffiths; blue grama; occasional on icc
slopes; Jul
et
—_
4
Bouteloua hirsuta ae hairy grama; infrequent
| | and slopes; ha Sep; pe-
DTT S| vallow
nnial C,
Bromus inermis Leyss.;smooth brome; common
in fields near headquarters, ditches, and at
woodland edges; May—Jun; introduced pe-
ennial C,.
ae icus Thunb.;J brome;com
n in disturbed sites and unburned pral-
n; introduced annual C
on ci Muhl. ex wild. woodland
ome; common in woodlands; May—Jun;
perennia
Bromus acai ai downy brome; infrequent in
iS urbe d sites and unburned prairie; May—
ntroduced annual C,,.
Peay ides gee we tial ess
common on clay u uplands and shall
sites;
|
May
Cenchrus eng asi NUS (Hack) Fernald; sandbur;
in a entin disturbed sites and along trails;
Jul-Sep; annual C
Chloris vertic eke Nutt; ; windmill grass; infre-
sr on shallow uplands; Jun—Sep; peren-
ialC
are aa fon (L.) Pers.; Bermuda grass; oc-
casional around headquarters; Jun—Sep; in-
troduced perennial C,
TOWNE, CHECKLIST OF KONZA PRAIRIE
=
Dactylis fei L., orchard grass; infrequent
around headquarters; May—Jun; introduced
perennial C,,.
Diarrhena obovata (Gleason) Brandenburg;
(Diarrhena americana var. obovata]; Ameri-
can beakgrain; occasional in woodlands; Jul-
Aug; perennial C,.
Dichanthelium acumi Een (Sw.) Gould 2 oa
woolly dict p
rie sites; May-Jun; perennial C,.
Dichanthelium oligosanthes (Nash) Gould;
Scribner's dichanthelium; eres on all
prairie sites; May—Jun; perennial C,
Digitaria cognata (Schult.) ae "eae
cognatum|; fall witch grass; infrequent on
uplands and slopes; Jul-Sep; perennial C,.
sl sanguinalis (L.) Scop.; hairy crab grass;
common around headquarters; Jul-Oct; in-
toduced annual C
Ee |
grass; seen in » ditches and disturbed
sites; Jun—Sep; an
Eleusine indica (L.) ae ; goose grass; occa-
sional in disturbed sites and along roads; Jul-
Oct; introduced annual
Elymus canadensis L.; Canadian wild-rye; occa-
sional on all prairie a ri eee ie io
Elymus villosus Muhl. ex Willd.; hairy wild-rye;
occasional in Vesna Jun—Jul;perennial C,.
Elymus virginicus L.;Virginia wild-rye; occasional
in woodlands; Jun—Jul; perennial C,.
ae capillaris (L.) Nees; lace grass: rare on
slopes; Jul-Sep; annual C,.
Eragrostis cilianensis (All) Vignolo ex Janch.;stink
ional in disturbed si talona
roads; Jun—Oct; introduced annual C,
— 5 ice aes (Michx.) Nees ex Steud.;
Carolina love grass; occasional in disturbed
ites an a along roads; Jul-Oct; ual C,,.
Eragrostis spectabilis (Pursh) en sine love
r
ee -
J~3€p,
perennial C,.
Eriochloa contracta Hitchc.; prairie cup grass;
occasional in disturbed sites and along
roads; Jul-Sep; annual C.
Festuca subverticillata (Pers.) Alexeev; [Festuca
sa]; nodding fescue; infrequent in
an ds; May— ne palin
aa astriata dena ) Hitche.; ft g
eee in wet sites and along creeks;
n; perennial C
Mts eee spartea Trin) Barkworth; [Stipa
spartea]; porcupine hie infrequent on up-
lands; May—Jun; perennial
ec ace ca {iy i a barley; infrequent
itches and mesic lowlands; Jun; peren-
ni al C,,
Hordeum pusillum Nutt, little barley; occasional
in disturbed sites and along roads; May; an-
nual C,.
eria macrantha (Ledeb.) Schult.; [Koeleria
pyramidata]; prairie June grass; common on
uplands; May—Jun; perennial C
Leersi ine ides (L.) Sw.;rice ee infrequent
wet sites and along creeks; Aug—Sep; pe-
rennial C
aa
Koe
VAGITA
Leersia virginica Willd.; white grass; occasional in
wet sites and along creeks; Jul-Sep; peren-
nial C,.
Leptochloa fusca (L.) Kunth; [Leptochloa
fascicularis]; bearded sprangletop; infre-
quent in disturbed sites; Jul-Sep; annual C,.
Leptochloa panicea (Retz.) Ohwi; [Leptochloa
filiformis]; red sprangletop; rare in disturbed
sites; J : ual C
Lolium arundinaceum (Schreb.) Darbyshire;
[Fest di J; tall fescue; occasional
=:
around headquarters; May—Jun; introduced
erennial C,.
ium perenne L.; rye grass; infrequent in dis-
turbed sites and ditches; May-Jun; intro-
duced perennial C
Lolium pratense dds) Darbyshire; [Festuca
pratensis]; meadow fescue; infrequent
around headquarters; May—Jun; introduced
oe
Lo
AAs ist n iL Deere «sm Pe | }y: bec]
sional in woodlands; Aug—Sep; perennial C,,.
stat esa (Torr. ex Hook.) Rydb.;
plains mu 7 ; occasional on uplands and
slope Sep; perennial C
ees on he eerrialel wire-stem
muhly; occasional in woodlands and along
creeks; Aug—Oct; perennial C
Muhlenbergia mexicana (L.) Trin.; leafy muhly;
eo in woodlands and along creeks;
p-Oct; perennial C,.
ieee racemosa (Michx) Britton et al.;
marsh muhly, green muhly; occasional at
ee edges and in mesic lowlands;
ct; perennial C
Pease la ee Emel nimblewill; occa-
sional in woodlands; Aug—Oct; perennial C,,.
Panicum capillare L., common witch grass; nae
onal in disturbed sites and along roads; Jul
nual C,
Panicum dichotomiflorum Michx.; fall panicum;
occasional in disturbed sites and along
roads; pace Soe ee. ar
2}
lowlands ann in mesic ‘euislande. Jul- sep pe-
nial C,,.
Panicum nea L.; switch grass; Common on
all prairie sites; Jul-Oct; perennial C
ae smithil (Rydb.) Love; ie eayen
smithif]; western wheat grass; occasional on
a uplands and in ditches; Jun; perennial C,.
Paspalum pubiflorum Rupr.ex Fourn,; hairy-seed
paspalum; rare along roads; Jul-Oct; peren
nial C,.
Paspalum setaceum Michx.; sand paspalum; rare
on clay uplands and shallow sites; Jul-Aug;
perennial C
Phi iclowlands
aad ditches; Jun; introduced perennial C,.
Poa arida eae plains blue grass; infrequent on
I wlands; May—Jun; perennial C,.
Poa oe Scribn.; Chapman's blue grass;
rare in disturbed sites; May-Jun; annual C,.
Poa compressa L.; Canadian blue grass; infre
quent in unburned uplands and disturbed
sites; May—Jun; introduced perennial C
Poa ee L.: Kentucky blue grass; abundant
in unburned prairie and woodlands; Apr—
Jun;intro aie ed perennial C
Schedonnardus paniculatus (Nutt. ) Trel.; tumble
> occasional on uplands and shallow
soils; J un—Aug; perennial C,.
Schizachyrium scoparium (Michx.) Nash;
[Andropogon scoparius]; little bluestem;
abundant on all prairie sites; Aug- Sep; pe
ial C
re pum ie (Poir.) Roem. & Schult.; [Setaria
glauca]; yellow bristle grass; occasional in
disturbed sites and along roads; Jul-Sep; in
troduced annual C,
Setaria viridis (L.) Beauv.; green bristle grass; in
frequent in disturbed sites and along roads;
Jun-Sep; introduced ann
=
4
BRIT.ORG/SIDA 20(1)
ae qutans (I ) Nash, Indian grass;abun
ennial C,,.
nall prairie sites; Aug—Oct; per
Sorghum halepense (L.) Pers.;
near cultivated fields; Jul-Oct; peecueee
perennial
ieee pecti inata Bosc ex Link; prairie cord
grass; common in mesic lowlands and
sane Jul-Sep; perennial C,.
sent is obtusata Mic 1x.) Scribn.; prairie
escale;Occe on uplands and low-
nies —Jun; perennial C,
Sporobolus compositus (Poir,) Merr; [Sporobolus
asper);tall dropseed; abundant on all prairie
sites and in abandoned brome fields; Aug-
ct; nial C
ea a ik (Torr.) A. Gray; sand
dropseed; infrequent on clay uplands and
shallow soils; Jul-Aug; perennial C,,.
Sporobolus heterolepis (A. Gray) A. Gray; prairie
dropseed; occasional on all prairie sites;
Aug-Sep; perennial C,,
Sporobolus neglectus Nash;
- Int
puff-sheath
1 ry
dQs, AUG ~SCp,
annual C
Soarebonasy vagini iflorus (Torr. ex A. Gray) Wood;
2 verty dropseed; infrequent along roads;
Aug—Sep; annual C
Tridens flavus (L.) Hitches: purpletop; occasional
on lowlands and in ditches; Jul-Sep; peren
nial C,
Tripsacum dactyloides (L.) L.; eastern gamma
grass; occasional on lowlands around head
quarters; Jun—Jul; perennial C,,.
Vulpia octoflora (Walter) Rydb.; [Festuca
octoflora]; six-weeks fescue; occasional on
grazed u ai and disturbed sites; Apr-
May; ann
POTAMOGETONACEAE - Pondweed family
tamogeton foliosus Raf; leafy pondweed; rare
H “dun wo:nerenni |.
In ) un Q; Pp
ats nodosus Poir;long-leaf pondweed,
nal in ponds; Jun—Aug; perennial.
Potamogeton pusillus L, baby pondweed; rare in
ponds; Jun—Aug;
SMILACACEAE - Catbrier famil
Smilax herbacea L.; carrion-flower greenbrier;
in woodlands; May-Jun; perennial.
Smilax tamnoides L.; [Smilax hispida]; bristly
greenbrier; common in woodlands; May
Jun; woody vine
erennial
r
rt
TOWNE, CHECKLIST OF KONZA PRAIRIE 293
Typha latifolia L.; broad-leaf cat-tail; occasional
in wet sites; Jun—Jul; perennial.
TNE HOCERE Cat-tail family
ph tifolia L. narrow-leaf cat-tail; rare in
wet sites. Jun—Jul; perennial.
APPENDIX II
Non-naturalized ornamental and horticultural species growing around the
headquarters area on Konza Prairie.
Family
Species
Common name
Araliaceae
ae
Caprifoliaceae
Hedra helix
Catalpa speciosa
Lonicera tatarica
hininer, hb ; trol}
English ivy
Northern catalpa
Tatarian honeysuckle
Cupressaceae D ae juniper
lridaceae Iris germanica
Liliaceae Amaryllis bella- donna ie ladies
Liliaceae Narcissus poeticu Poet's narcissus
Liliaceae Narcissus foun affodil
Oleacea Syringa vulgaris Lila
Paeoniaceae Paeonia lactiflora Pe
Pinaceae Pinus nigra Austrian pine
Rosaceae Malus sylvestris Apple
Rosaceae Pyrus communis Pear
Sapindaceae Koelreuteria paniculata Golden rain-tree
ACKNOWLEDGMENTS
Numerous individuals have contributed to compiling the Konza Prairie flora,
but the most prominent predecessors were Craig Freeman, Richard Kazmaier, Lloyd
Hulbert, and Linda Marple. Ted Barkley and Iralee Barnard were instrumental
in persuading me to collect formally and update the Konza Prairie flora. Earlier
drafts of this manuscript were improved by comments from Carolyn Ferguson,
Mark Mayfield, and Iralee Barnard. 1 also appreciate the meticulous review and
insightful comments from Robert B. Kaul and Craig C. Freeman. The National
Science Foundation Long-Term Ecological Research Program, Konza Prairie
Biological Station, Kansas State University Herbarium, and the Kansas Agri-
cultural Experiment Station provided support for this project. This report is
contribution no. 02-122-J of the Kansas Agricultural Experiment Station.
REFERENCES
ANDERSON, K.L. 1961. Common names of a selected list of plants. Kansas State Agric. Exp.
Stat. Tech. Bull. No. 117, Manhattan.
Bare, J.E.1979.Wildflowers and weeds of Kansas. The Regents Press of Kansas, Lawrence.
Brooks, R.E. 1986.Vascular plants of Kansas:a checklist. Kansas Biological Survey, Lawrence.
Brummitt, R.K.and C.E. PoweLt. 1992. Authors of plant names. Royal Botanic Gardens, Kew.
294 BRIT.ORG/SIDA 20(1
Downton, W.J.S. 1975. The occurrence of Cy photosynthesis among plants. Photosynthetica
9:96-105.
Freeman, C.C. and L.C. Huteert. 1985. An annotated list of the vascular flora of Konza Prairie
Research Natural Area, Kansas. Trans. Kansas Acad. Sci. 88:84-115.
Freeman, C.C.and D.J.Gigson. 1987. Additions to the vascular flora of Konza Prairie Research
Natural Area, Kansas. Trans. Kansas Acad. Sci. 90:81-84.
Great PLains Flora Association. 1986. Flora of the Great Plains. University Press of Kansas,
Lawrence, KS.
Kartesz, J.T. 1999.A synonymized checklist and atlas with biological attributes for the vas-
cular flora of the United States, Canada, and Greenland. First Edition. In: J.T. Kartesz and
C.A. Meacham. Synthesis of the North American flora, Version 1.0. North Carolina Bo-
tanical Garden, Chapel Hill.
Kartesz, J.T. and J.W. THierer. 1991. Common names for vascular plants: Guidelines for use
and applications. Sida 14:421—434.
Parmer, M.W., G.L. Wabe, and P. Neat. 1995. Standards for the writing of floras. BioScience
45:339-345,
Water, S.S.and J.K. Lewis. 1979.Occurrence of C3 and C4 photosynthetic pathways in North
American grasses. J. Range Manage. 32:1 2-28.
THE VASCULAR FLORA OF THE CERRADO IN
EMAS NATIONAL PARK (GOIAS, CENTRAL BRAZIL)
Marco Antonio Batalha Fernando Roberto Martins
Department of Botany Departmen Forsotny
Federal University of f Sdo Carlos Institute of Biology, S State University o fCam
PO Box 676, Sdo Carlos, SP 13565-905, BRAZIL PO Box 6109, Campinas, SP. 13083-970, en
marcobat@uol.com.br fmartins@unicamp.br
ABSTRACT
The cerrado, a savanna-like ecosystem, is the second largest a type in Brazil, originally cov-
P
ering about two million km? (or 23%) of the Brazilian territory. ihe Emas National Park (ENP), com-
prising about 133,000 ha, is one of the most important reserves within the cerrado. From November
1998 to October 1999, we carried out a floristic survey of all the cerrado physiognomies of the ENP
and found 601 species, belonging to 303 genera and 80 families. Among the collected species, 12 were
weeds and seven were new to science. The herbaceous to woody species ratio was 3.03:1. The richest
amilies were Asteraceae (88 species), Fabaceae (87), Poaceae (51), Myrtaceae (39), and Lamiaceae
(24); these five families alas 48% of oe total number of species. The results obtained show the
importance of the ENP t servation. since from 8 to 20% of the species recorded for this
vegetation type occur within mutts reserve. We emphasize the need for more floristic surveys in which
the frequently overlooked hert
p t should also be sampled.
Key words: cerrado, savanna, floristics, Emas National Park, central Brazil
RESUMEN
El cerrado, un tipo de sabana, es el segundo mayor tipo de vegetacion de Brasil. Una de las reservas
si Hee Sees a proteger el cerrado es el “P. oe Nacional das Emas” (PNE), con un area
aproximada de 133.000 ha. En el periodo de noviembre de 1998 a octubre de 1999, fue realizado
i ween en las fisionomias de cerrado del ne oo 601 especies
pertenecientes a 303 géneros y 80 familias. De entre las es] is, fueron encontradas 12
invasoras ie siete nuevas para la ciencia. La proporcion entre especies eee subarbus tivas yi
arbustivo-ar sf 3,03:1. Las familias mas ricas en especies fueron: A (88 especies),
Fabaceae (87), Poaceae (51), Myrtaceae (39) e Lamiaceae (24), compreendiendo 48% del el de
especies. Los resultados mostraron la importancia del PNE para la conservacion del cerrado, ya que
en esta reserva aparecen de 8a 20 %de las especies ee para dicha formacion vegetal. Resaltamos
aqui la necesidad de realizar levantamientos floristicos contemplando también el componente
herbaceo-subarbustivo, frecuentemente olvidado.
INTRODUCTION
The Cerrado Domain, the second largest Brazilian phytogeographic province,
once covered about 2 million km2, or 23% of the Brazilian territory, with its
core area in central Brazil (Ratter et al. 1997). As its name implies, cerrado veg-
etation prevails in the Cerrado Domain. The cerrado vegetation is not uniform
in physiognomy (Coutinho 1990), ranging from grassland to tall woodland, but
most of its physiognomies within the range defined as tropical savanna
SIDA 20(1): 295 — 311. 2002
296 BRIT.ORG/SIDA 20(1)
(Sarmiento 1983). In the Cerrado Domain, interspersed with the prevailing
cerrado vegetation, there are other vegetation types, such as seasonal forest, ri-
parian forest, rocky campo, and wet campo.
Although frequently neglected in the past, the cerrado vegetation stands
out in its high floristic richness (Ratter et al. 1997). After comparing a large
number of floristic and phytosociological surveys carried out in cerrado sites
from all over Brazil, Castro et al. (1999) estimated that 3,000 to 7,000 vascular
plant species occur in this vegetation type. In addition to its high floristic rich-
ness, the cerrado presents a high degree of endemism. Lenthall et al. 1999), for
instance, listed 234 woody species occurring in 10 cerrado sites and verified
that 80% of them were restricted to this vegetation type. Owing to its high rich-
ness, high degree of endemism, and present conservation status, Fonseca et al.
(2000) included the cerrado among the biodiversity hotspots for highest prior-
ity conservation in the world.
According to Rizzini (1963), the cerrado flora consists of an herbaceous
and a woody component, which compete because both are heliophilous, 1.e.,
there is no shade-adapted ground layer. Following Coutinho’s (1990) concept of
cerrado, the importance of the herbaceous component decreases from open to
{4
closed physiognomies, whereas the importance of the woody component in-
creases. In most forms of cerrado, the herbaceous component is much richer in
species, as demonstrated by Mantovani and Martins (1993), who found a ratio
between herbaceous and woody species ranging from 2:1 to 3:1 in the compari-
son of some southern cerrado sites. However, despite its richness, Castro et al.
(1999) pointed out the almost complete absence of existing surveys on the her-
baceous component of the cerrado. These authors also highlighted the uneven
distribution of surveys, which tend to be concentrated on few well studied ar-
eas (Castro et al. 1999).
The Emas National Park (ENP) is the largest and one of the most impor-
tant reserves among those in the Cerrado Domain (Conservation International
1999). Although some papers about its fauna and wildfires exist (e.g., Ramos-
Neto & Pivello 2000; Rodrigues & Monteiro 2000), the ENP’s vegetation remains
poorly studied. The present work was designed as an intense and systematic
floristic survey to provide a better knowledge of the ENP’s flora in particular
and of the cerrado flora in general, especially concentrating on the still poorly
known herbaceous component. Furthermore, this survey is intended to pro-
vide a basis for other studies to be carried out in the ENP and for phytogeo-
graphical studies on the cerrado flora.
Toachieve these aims, we have tried to answer the following questions: What
L
isthe floristic composition of the cerrado physiognomies in ENP? Which fami-
lies are the richest ones in its flora? Does the herbaceous to woody species ratio
lie within the range described by Mantovani and Martins (1993), that is, be-
BATALHA AND MARTINS, CERRADO FLORA OF EMAS NATION AL PARK 297
tween 2:1 and 3:1? Are there woody species that should be included in the check-
list elaborated by Castro et al. 1999)?
SITE SUMMARY
Created in 1961, the ENP is located in the Brazilian Central Plateau, southwest-
ern Goias State (17°49'-18°28'S, 52°39'-53°10'W), in the cerrado core region, and
comprises 132,941 ha. Regional climate is humid tropical with wet summer and
dry winter, classified as Aw following Képpen’s (1948) system. Annual rainfall
varies from 1,200 to 2,000 mm, concentrated from October to March, and mean
annual temperature lies around 24,60'C (Ramos-Neto & Pivello 2000). Three
quarters of the ENP consist of flat tableland, 820-888 m high, and the remain-
ing area consists of hilly terrain, 720-820 m high (Ramos-Neto & Pivello 2000).
Recently, ENP was included by UNESCO (2001) in the World Natural Heritage
List as one of the sites containing flora, fauna, and key habitats that character-
ize the cerrado.
The cerrado in ENP exhibits almost all physiognomies found in this veg-
etation type. Following Coutinho’s (1990) classification and Sarmiento’s (1984)
translation, the cerrado in ENP ranges from campo limpo (a grassland) to cerrado
sensu stricto (a woodland). In the reserve, open cerrado physiognomies—campo
limpo, campo sujo (a shrub savanna), and campo cerrado (a savanna woodland)—
prevail, covering 68.1% of the total area, especially on the flat tableland (Ra-
mos-Neto @ Pivello 2000). The more closed cerrado sensu stricto covers 25.1%
of the reserve, mainly on the hilly terrain. Other vegetation types, such as wet
campo (4.9% of the total area) and riparian and seasonal semideciduous forests
(1.2%), also exist within the park.
METHODS
We surveyed all cerrado physiognomies occurring within the reserve from
November 1998 to October 1999, in monthly field trips, each one with a 50-60
hr. sampling effort in the field. We established routes through the firebreaks
that cross the reserve (Fig. 1) and covered one of them each day by driving a
vehicle and stopping whenever vegetation of interest was seen. The routes com-
prised the following reference points (with approximate distances):i) U2, UL, Q,
R,S, T, U2 (51.5 km); ii) U2, V, Pl, O, U1 (52.5 km), iii) O, M, N, P1 (45 km); iv) V, P2,
W, Y, V (47.5 km); v) X, Z1, Z2, Z3, Z2, ZI, Y, W, X (47.5 km); vi) A, B,J, G, D, C, K2,
KI, A (48 km).
We collected fertile botanical material along the pre-established routes.
During the last two field trips, we also collected sterile material from species
previously not found in reproductive stages. The collected material was identi-
fied to species level by comparing it with lodged vouchers and consulting taxo-
nomic references. The specimens were then sent to taxonomists for confirma-
298 BRIT.ORG/SIDA 20(1)
P| ™ firebreak
Se river
contour lines
(20 m internal
Fic. 1. Firebreaks, with ref ints, in E National Park (17° 49'-18'28'S, 52° 39'-53° 10'W), Goias State, central
Brazil.
tion. The voucher material was stored in the herbaria of the Sao Paulo Botani-
cal Institute (SP) and State University of Campinas (UEC).
We classified the species in families according to the system proposed by
Judd et al. (1999) and in life forms following Raunkiaer’s (1934) system adapted
by Mueller-Dombois and Ellenberg (1974). We considered the chamaephytes,
BATALHA AND MARTINS, CERRADO FLORA OF EMAS NATIONAL PARK 299
epiphytes, hemicryptophytes, geophytes, lianas, vascular parasites, and
therophytes as belonging to the herbaceous component and the phanerophytes
as belonging to the woody component. We applied the chi-square test (Zar 1999)
to verify whether the herbaceous to woody species ratio was significantly dif-
ferent from the expected by Mantovani and Martins (1993). The results found
were also compared to the patterns obtained by Castro et al. (1999).
RESULTS
We collected 2,123 voucher specimens, representing 601 species, 303 genera, and
80 families (Appendix I). Out of these 601 species, 571 were identified to species
level, including one new to science, Piriqueta emasensis Arbo (Turneraceae).
Six out of the remaining 30 species were also new to science and are currently
being described by taxonomists: Annona sp. nov. (Annonaceae), Gyrostelma sp.
nov. (Apocynaceae), Dimmerostema sp. nov. (Asteraceae), Ipomoea sp. nov.
(Convolvulaceae), Hybanthus sp. nov. (Violaceae), and another Convolvulaceae
that belongs toa new genus. There were 22 species identified to genus level, one
identified to family level (Fabaceae sp.), and one that we could not identify even
to family level.
Twelve species (2.0% of the total number of species) were considered by
Mendonca etal. (1998) as weeds that do not occur spontaneously in the cerrado.
Of the 601 species, 149 were phanerophytes and thus included in the woody
component, while +52 belonged to other life forms and were included in the
herbaceous component. The herbaceous to woody species ratio was 3.03:1, a
value not significantly different (y? = 0.014, P = 0.906) from the highest ratio
(3:1) set by Mantovani and Martins (1993).
The richest families were Asteraceae (88 species), Fabaceae (87), Poaceae
(51), Myrtaceae (39), Lamiaceae (24), Malpighiaceae (23) Euphorbiaceae (20),
Apocynaceae (19), Malvaceae (16), Rubiaceae (16), and Convolvulaceae (15),
which together summed up 66.2% of the total number of species. In the herba-
ceous component, the richest families were Asteraceae (85 species), Fabaceae
(59), Poaceae (48), Lamiaceae (23), Euphorbiaceae (19), Malpighiaceae (16),
Myrtaceae (16), Convolvulaceae (15), and Apocynaceae (14), which comprised
65.27% of the herbaceous species. In the woody component, the richest families
were Fabaceae (28), Myrtaceae (23), Malpighiaceae (7), Melastomataceae (7),
Annonaceae (6), Apocynaceae (5), Vochysiaceae (5), Bignoniaceae (4),
Nyctaginaceae (4), and Rubiaceae (4), which together accounted for 62.42% of
the woody species.
Seventeen out of the 149 woody species (11.4%) did not appear on the check-
list of the cerrado woody flora established by Castro et al. (1999): Annona sp.
nov., Aiouea trinervis Meisn., Apoclada arenicola McClure, Banisteriopsis acerosa
(Nied.) B. Gates, Calliandra macrocalyx Harms, Dalbergia cuiabensis Benth.,
Mimosa amnis-atri Barneby, M. gemmulata Barneby, M. hebecarpa Benth.,
300 BRIT.ORG/SIDA 20(1)
Myrcia bracteata O. Berg, M. camapuanensis N.FE. Silveira, M. crassifolia (O.
Berg) Kiaersk. M. fall ax(Rich.) A. DC. M. linguaeformis Kiaersk., M. ener
Kiaersk., Olyra taquara Sw, and Psidium laruotteanum Cambess
DISCUSSION
If we assume that the number of species in the cerrado ranges from 3,000 to
7,000 (Castro et al. 1999), then the ENP contains approximately 8.5 to 20.0% of
the cerrado flora. These figures show the importance of this reserve for the con-
servation of the cerrado vegetation. The number of species in ENP might be
increased by species not found in our survey. Floristic surveys certainly miss a
number of species in a given area, especially those that are not at reproductive
stage at the time of the visit, flower sporadicaly, are ephemeral, or are incon-
spicuous, problems that particularly affect the herbaceous component of the
vegetation (Mantovani & Martins 1993; Castro et al. 1999).
Among the 601 species found in the ENP, seven are new to science. Although
the cerrado is one of the most studied vegetation types in Brazil (Castro et al.
1999), the fact that undescribed species keep on appearing in floristic surveys—
for example, also in Brasilia, Federal District (Pereira et al. 1993)—shows that
firmation that the
tropical flora remains undercollected (Prance et al. 2000) seems to apply also
to the cerrado vegetation. Some of the species not identified to species level in
our inventory may be new to science as well.
-_
the cerrado has not yet been satisfactorily sampled. The a
Although the proportion of weedy species in the ENP’s cerrado flora was
lower than the ca. 5% found by Mendonga et al. (1998) for the whole Cerrado
Domain, the invasion of ruderal plants in ENP is alarming, notably the African
grasses Brachiaria decumbens Stapf and Melinis minutiflora P. Beauv. Plant in-
vasion has become a great pro lem in virtually all cerrado fragments (Pivello
et al. 1999) and will grow into a serious problem in the ENP if precautions are
not taken.
The richest families in the ENP were also the richest ones in other cerrado
sites (Mantovani & Martins 1993; Batalha et al. 1997; Batalha & Mantovani 2000).
Exceptions were Convolvulaceae and Lamiaceae, well represented only in the
ENP. The herbaceous to woody species ratio, although not significantly differ-
ent from the maximum set by Mantovani and Martins (1993), was higher than
the ratios found in other surveys (Mantovani & Martins 1993; Batalha et al. 1997:
Batalha & Mantovani 2000). This probably was a consequence of the preva-
lence of open physiognomies (campo limpo and campo sujo) in ENP.
According to Castro et al. (1999), surveys in poorly sampled regions should
—_—
—
augment the cerrado woody species checklist. Indeed, the 17 species found in
our survey that should be included in their list represent a high percentage of
the species collected in ENP and indicate that even the woody component of
the cerrado vegetation remains undercollected. As for the herbaceous compo-
a
BATALHA AND MARTINS, EMAS NATIONAL PARK 301
nent, this undercollection might be even greater. In the visits we paid to her-
baria and in our search for references, the absence of surveys in this component
became evident.
Species inventories, even at the most basic level, are available for only about
1% of the tropical regions (Hammond 1992). Our floristic survey of the domi-
nant cerrado vegetation in the ENP represents a first step, and it will be neces-
sary to survey the other vegetation types existing within the reserve, such as
seasonal semideciduous forest, riparian forest, wet campo, and the aquatic veg-
etation of streams, rivers, and lakes.
APPENDIX I
Species collected in the floristic survey of the cerrado vegetation in Emas Na-
tional Park (17°49'-18°28'S, 52°39'-53°10'W), Goias State, central Brazil. Taxa were
listed alphabetically by family, genus, and species. The format used was species
name, authority, collection number, and life form. Authorities were abbreviated
according to Brummit and Powell (1992). Species designated by an asterisk (*)
were considered non-native weeds by Mendoncaet al. (1998). All specimens were
collected by M.A. Batalha (B). Life form was assigned according to Raunkiaer’s
(1934) system adapted by Muller-Dombois and Ellenberg (1974). Life form
classes were abbreviated as: Ch = chamaephyte, Ep = epiphyte, Geo = geophyte,
H = hemicryptophyte, Li = Liana, Ph = phanerophyte, Th = therophyte, and Vp
= vascular parasite.
APPENDIX |
The vascular flora of the cerrado in Emas National Park (central Brazil).
ACANTHACEAE Annona la Mart., B 3620,H
Hygrophila brasiliensis (Spreng.) Lindau, B 2306,H = Annona tomentosa R.E. Fries, B 2347,Ch
Ruellia gemini iflora Kunth, B 2169, H Annona warmingiana Mello-Silva & Pirani, B 3763,
Ruellia pta (Nees) Lindau, B 3592, Ch
AUEHIC HICOFT!
ALSTROEMERIACEAE
Alstroemeria gardneri Baker, B 2715,H
AMARANTHACEAE
Froelichia procera (Seub.) paren B 2123,H
a b > L.£,B 5,H
phalc ey B 2214,H
Gomphieta pohlii Woe: B 2091,
Pfaffia helichrysoides eeu ine B2152,H
Pfaffia jubata Mart., B 2
ANACARDIACEAE
Anacardium humile
A. St-Hil., B 2050, Ch
Tapirira guianensis Aubl., B 3804, Ph
oabidinsateg
\ \
Vid
5,Ph
Annona crassiflora en : es Ph
Annona sp. nov.,B 2621,
Bocageopsis anooronenke s (R.E. Fries) R.E. Fries,
Duguetia furfuracea (A. St-Hil.) Benth. & Hook. f,,
Duguetia glabriuscula (R.E. Fries) R.E.Fries,B 1955,
Ch
Ylopia a tica (Lam.) Mart.,B 2515, Ph
APIACEAE
Didymopanax macrocarpum (Cham. & Schltdl.)
Seem.,B 2107, Ph
os idmepanay a malmei Harms, B 2733, Ph
Cham. & Schitdl., B 3890, H
Eryngium junceum Cham. & Schitdl., B 2566, H
APOCYNACEAE
Asclepias mellodora A. St-Hil., B 2967, H
iliatr
Ery
302
i eas macrocarpon Martt., : ae Ph
pidosperma nobile Mull. Arg., B 3661, Ph
Aspidosperma tomentosum ae a ron Ph
sd FOUrN,, Zo,
K. a an 182,H
tee
mae:
<<
erecta (Vell,
Sennen bicuspidatur
Sacral nov., B 2081,
: 2106,Ph
ee B 2083,H
Himatanthus obovatus (Mull. Arg.) Woods., B
2 i
of
Heminogon aceros
[meet
Macrosiphonia longiflora Mall. Arg., B 2218, H
Macrosiphonia velame (A. St-Hil.) K. Schum., B
10,H
Mandevilla coccinea (Hook. & Arn.) Woods., B
2577
as
Mandevilla pohliana (Standelm.) A. Gentry, B
2099, H
Markgr., B 2645, Li
»Fourn, /B 2241, H
Pau eee lia ldhna a Mill. Arg., Bd 962, H
Rhodocalyx rotundifolius Mull. Arg., B 2281,H
EG uted Da )
ypetalun f
ARACEAE
Scaphispatha gracilis Brongn.ex Schott, B 2075,H
ARECACEAE
Acrocomia aculeata Jacq.) Lodd.ex Mart.,B 2498,
Ph
Acrocomia hassleri (Barb. Rodr.) Hahn, B 3828,Geo
Allagoptera campestris (Mart.) Kuntze, B 2006,
ae
eo
Allagoptera leucocalyx (Mart.) Kuntze,B 2215,Geo
Attalea geraensis Barb. Rodr., B 2509, Geo
Syagrus flexuosa (Mart.) Becc., B 2249, Geo
ARISTOLOCHIACEAE
Aristolochia gibertii Hook., B 2852, Li
Aristolochia gracilis Duch.,, B 4029, Li
ASTERACEAE
i a australe (Loefl.) Kuntze,B 2527,
Soe
ae
fain te satureoides (Lam.) A.DC., B 3400, Th
Apo opyros warm we (Baker) Nesom, B 2176, H
Aspilia foliacea (Spreng.) Baker, B 3377,H
Aspilia laevissima pe B ae :
Aspilia leucoglossa Malme, B 2
Aspilia platyphylla (Baker) ne B 2848, H
Ayapana amygdalina (Lam.) King & H. Rob., B
3458,H
RBaccharis camnportur
ae
nN A.DC.,B 3313,Ch
BRIT.ORG/SIDA 20(1)
Baccharis humilis Sch. Bip.,B 2172,H
Bidens gardneri Baker, B 3031, Th
Calea clausseniana Baker, B 3771,H
Calea cuneifolia A.DC.,B 4011,H
lepis Baker, B 2886, H
Calea platylepis Sch. Bip. ex Baker, B 3204,H
Campuloclinium chlorolepis Baker, B 3120,H
oclinium megacephalum (Mart.) King &
ob.,B
Calea hymen
pai
Campu
H.R
Chaptalia integerrima (Vell.) Burkart, B 2089, Th
Chromolaena chaeseae H.Rob., B 3314, Ch
Chromolaena leucocephala Gardner, B 3409, Ch
Chromolaena squalida (A. DC.) King & H. Rob., B
2 Ch
ee stachyophylla (Spr) King & H.Rob,,
B - 6,4
. B 2497,TH
passin eee ar (Gardner) Cabrera,
bi merostema asperatum Blake, B ies de
brasilianum C 316
Di ifoli Sch ap oe i
Dimmerostema sp. nov.,B 2010, H
Elephantopus biflorus Less.,B 3159,H
Flephantopus mollis L.,B 3128, H
ee racemosus Gardner, B 3412, H
(> Sims.) Sweet * iB 2364, Th
-htitesh
Eremanthus Sch. Bip., B 2213,Ph
Eremanthus glomerulatus Less.,B 3333, Ph
Eremanthus sphaerocephalus Baker, B 2273,H
a us patori ium betonicaeforme Baker, - _ H
ampestre A.DC.,B 19
¢ Hook. & hee g a H
Eupatorium myriocephalum Gardner, B 3406, Ch
Eu ses torium auadon ens ape a . 3093,H
L
Finatoritim |
a yaa
Fupator um L.f,,
Bapatonth /UM SP., B 3309, H
a fe loa Cabrera, B 3484, Ch
Cabrera, B 3014, Ch
eee trixoides pee Cabrera, B
596,
ct Lae sp.1,B 212
Sonica pean cum Scherff, B 2149, H
sost oe Pa se H
M | (L.) Willd. B 3156, Li
BATALHA AND MARTINS, CERRADO FLORA OF EMAS NATIONAL PARK
Orthopappus angustifolius (Sw.) Gleason *, B
2402,H
Piptocarpha rotundifolia (Less.) Baker, B 1957,Ph
Porophyllum angustissimum Gardner, B 2513, Th
ase pinifolia (Spr.) King & H. Rob., B
perocvion Mga (L.) A. DC., Bes liaacA
lia Gardner, B 2331,H
urtia Ul ngif TCA
n
Riencourtia tenuifolia Gardner, B 2256, H
SE ilanthec nenjncg Chod.,
Gant: glomerulatus a B 2877,H
Stilpnopappus speciosus Baker, B 2504, H
ag ee oe (A. — King & H.
Rob., B
bo lonia agente ms B 2388,H
5.,B 2514, .
Vernonia Siena ot Bip.,B 3189, H
Vernonia buddleiifolia Sch. Bip. ex ee B 2338, H
ene eae Mart. ex Baker, B 2867, H
! um Mart.ex A.DC.,B 3567,H
Weinenie erythrophyla Mart., 2094,H
Vernonia ferruginea Less.,B 3510, H
Vernonia grandiflora Less., B 2487,H
Vernonia herbacea (Vell.) Rusby, B 1992, H
Vernonia ignobilis Less., B 2168,H
Vernonia polyanthes ee Less., 7 2336,Ch
vernon psi lop nye ‘ DC.,B 2832,
nia rubricaull eee B 2375,H
ein! ia rubriramea ie B 3191,Ch
Vernonia simplex Less.,B 4014, H
Vernonia tomentella Mart.ex A.DC.,B 2063, H
Vernonia tragiaefolia A.DC.,B - ),
Vernonia VC rroniifolia A. DC,, B3 4,H
Vernonia venosissima Sch. Bip. ex . ker,B 2001,H
Vernonia virgulata Mart., B 2033,H
Viguiera bakeriana Blake, B 2414, H
ae sp. 1,B 2084, H
Viguiera sp. 2,B 1960, H
Wedelia macedoi H. Rob.,B 3920, H
BALANOPHORACEA
Langsdorffia hypogea ak B4
BIGNONIACEAE
Anemopaegma arvense (Vell.) Stellfeld ex de
Souza, B 2883,
Anemopaegma glaucum Mart.ex A.DC.,,B 3035,H
Anemopaegma scabriusculum Mart. ex A. DC., B
ee a
Ay chypoda (A. DC.) Bur, B 2272,Ch
Boab tee Bete (Cham.) Sandw., B 2460, Li
303
rae psa ica Mart., B 3246, Ph
ba (Vell.) A.DC.,B 2279, Ch
saldaia
Cham., B 3799, Ch
ieeas ae Silva Manso, B 2267,Ch
Memora pedunculata (Vell) Miers, B 2063, Ch
Tabebuia aurea (Silva Manso) S. Moore, B 3557,
Tabebuia ochracea (Cha ees B 3659, Ph
Zeyheria montana vars B 266
BORAGINACEA
Cordia villicaulis oe B 381
BROMELIACEAE
Aechmea bromeliifolia (Rudge) Baker, B 2474, Ep
Ananas ananassoides L.B.Sm., B 2878,
Bilbergia magnifica Mez, B 3755, Ep
Bromelia balansae Mez, B 2013,H
Dickia tuberosa (Vell. ) Beer, B 2839, H
BURSERACEAE
|
CNgI., B 1930, Ph
CACTACEAE
Fninhvilum phvilanthus (L.) Haw., B 3878, Ep
CARYOCARACEAE
Caryocar brasiliense Cambess., B 1989, Ph
CARYOPHYLLACEAE
Dnv!
(L.) Lam.,B 3359,Th
CELASTRACEAE
la a aun Reissek, B 2277, Ph
(Mart.) A.C.Sm.,B 1934, Ph
CHRYSOBALANACEAE
Couepia oe (Mart. & Zucc.) Benth. ex
Hook. f.,B
icania pana & Schitdl., B 3034, Ph
si ee excelsa Sabine, B 1978, Ch
Peaiaaie
qa abdita oe B 2095,Ch
21
rt
dl
Kieh never ice (wera) Saddi, . 2061,Ph
Kielmeyera rubriflora C ess.,B 2.
Kielmeyera trichophora Mart., B 2035, Ch
} ielmeyera variabilis Mart., B 191 Ds Ch
COCHLOSPERMACEAE
Cochlospermum regium (Mart.) Pilg., B 2428, H
COMBRETACEAE
Buc ia avid CUAPTICPI COs Eich, B 4031, Ph
eee hilarianum D. Dietr., BR?) 33, @
—
COMMELINACEAE
C lina obliqua Vahl, B 2060, Th
CONNARACEAE
Connarus suberosus Planch., B 2374, Ph
Rourea induta Planch.,B 1931,Ph
CONVOLVULACEAE
Evolvulus cressoides Mart., 7,H
Evolvulus fuscus Meisn., B 3974, H
Evolvulus ae ibe va : 7 H
m sn.,B 26
tris Meisn., B .
pening: breton nbens Mart. ex ae B 2613,H
{pomoea procurrens Meisn.,B 2295, Li
Ipomoea sp. Nov
ane vi gat a Meisn, B 3415,H
mitica Hassl.,B 2636,H
Jacquemontia cohucroceonala Meisn.,B 2363,H
(Choisy) f.,B 2600, Li
Merremia ohare Meisn., B 214
Turbina abutiloides (Kunth) O' al B 2512,H
Convolvulaceae sp. 1, gen.et sp.nov. B 1967, H
ieatienioas
li ogn.,B 2118,Li
pi qtOS nna hilarian 1Cogn., B 3645, Li
Melancium campestre Naud., B 2608, H
—
PERACEAE
Bulbostylis junciformis (Kunth) C.B.Clarke, B 2300,
Bulbostvlis
(Spreng.) Lindm.,B 3565, H
Bulbostylis sp fea (Nees) C.B. Clarke, B
H
2046,
Bulbostylis truncata (Nees) M.T. Strong, B 3971,H
pei aggregat Us en see B 2023,H
Cyr nth 7,H
Kyll inga pdoratNa | : 2931, tia
Rhynchosphora diamantina (C.B.Clarke) Kukenth,
2316,H
SUA sani emaciata Boeck., B 3087,H
Rhynchosphora exaltata Kunth, B 2466, H
irae rugosa (Vahl) Gale, B 3534,H
Scleria scabra Willd., B 2310, H
DILLENIACEAE
Davilla elliptica A. St-Hil., B 2080, Ph
Davilla nitida (Vahl) Kubitzki, B 3988, Li
DIOSCOREACEAE
sagan amaranthoides Presl, B 3124, Li
i Uline, B 3044, Li
BRIT.ORG/SIDA 20(1)
EBENACEAE
Diospyros hispida A.DC.,B 2031, Ph
en ROR ER Ene
nm campestre A. St-Hil.,B 2228, Ph
decid A. St-Hil., B 3716, Ph
ei suberosum A. St-Hil., B 1965, Ph
EUPHORBIACEAE
Chamaesyce caecorum (Mart. ex Boiss.) Croizat,
2137,H
Cnidosculus gt s Pohl, B 1946, H
shia a ins Mull Arg., B 2036,H
Mart, B 2017,H
Coton INCLUS 5 Mill Arg., B 3109, H
losus Mall. Arg., B 2389,H
Croton goyazensis Mall. Arg.,B 2222,H
Croton lundianus Mull. Arg.,B 2205, H
Croton p dhlianus Mull. Arg., B 2196,H
Croton sclerocalyx Mull. Arg., B 2885, H
Croton sp.,B 1964, H
Dalechampia humilis Mull. Arg., B 1981, H
Dalechampia linearis Baill, B 2076, H
lulocroton humilis Didr., B 2207,H
Manihot caerulescens Pohl, B 2459,H
ie Mull. Arg.,B 1908,H
ounea guianensis Aubl., B 3930, Ph
ai orbiculatus Mull. Arg., B 2445, Th
Sapium glandulatum (Vell) Pax, B 2301,Ch
Sebastiania bidentata (Mart.) Pax, B 2430,H
FABACEAE
Acosmium subelegans (Mohl.) Yakovlev, B 3815
Ph
Aeschynomene marginata Benth.
Aeschynomene oroboides Benth., B 5,H
Anadenanthera falcata (Benth.) Den : 2650,Ph
Andira cuiabensis Benth., B 2045, Ph
Andira laurifolia Benth., B 2342, Ch
Andira vermifuga (Mart.) Benth.,B 3718, Ph
Arachis tuberosa Bong. ex Benth., B 2275,H
Bauhinia rufa Steud., B 1907, Ph
Bowdichia virgilioides Kunth, B 3350, Ph
ot maid ee tha Benth.,B 1979, Ch
crocalyx Harms, B 2230, Ph
‘hodat ex
pie 6,Th
cements sericeum (Benth.) C
Hassl.,B 2569, Li
Camptosema ellipticum (Desv.) Burkart, B 3232, Li
Centrosema venosum Mart. ex Benth., B 1924, Li
Chamaecrista basifolia (Vogel) Irwin & Barneby,
B 2710,Ch
BATALHA AND MARTINS, CERRADO FLORA OF EMAS NATIONAL PARK
Chamaecrista campestris Irwin & Barneby,B 2723,
Ch
Chamaecrista cotonifolia (G. Don) Killip, B 4001,
Ch
Chamaecrista desvauxii (Collad.) Killip, 2457, Ch
Chamaecrista filicifolia (Benth.) Irwin & Barneby,
B 2325,
Chamaecrista flexuosa (L.) Greene, B 2290, Ch
Chamaecrista lundii (Benth.) Irwin & Barneby, B
3435,H
Chamaecrista nictitans (L.) Moench., B 2726,Ch
hamaecrista planaltoana (Harms) Irwin &
rneby, : 2208,
Chamaecrista rotundifolia (Pers.) Greene, B 1974,
Chamaecrista setosa (Vogel) Irwin & Barneby, B
2
Clitoria densi (Presl) Benth., B 2433, H
Copaifera langsdorffii Desf, B 2127, Ph
Crotalaria maypurensis aoe B 2597,Th
Crotalaria nitens Benth., B 2724,H
rotalaria velutina 81,H
Dalbergia cuiabensis Benth., B 3214, Ph
papegi a miscolobium Benn B 3504, Ph
arbatum (L.) Benth., B 3024, H
Desnodiinn incanum sw) A.DC.,B 2523,Th
Desmodium platycarpum Benth., B 3944, H
Dimorphandra mollis Benth., B 2047, Ph
Dioclea bicolor Benth., B 2042, Li
a i aurantiaca Tul., B 3056, Ph
tum (Kunth) Gardner, B 2483, H
CUTTT H Idiitts, B 2] 20, H
Evosenia marian Mart. ex Benth.,B 2251,H
Eriosema heterophyllum Benth., B 3992,H
Eriosema longifolium Benth., B 2194,H
Eriosema rufum Kunth, B 2444, H
Galactia decumbens (Benth.) Chodat & Hassl., B
2371,H
alactia dimorpha Burk., B 2130,H
Galactia martii A.DC.,B 3512,H
Harpalyce brasiliana Benth., B 2548, Ph
Hymenaea stigonocarpa Mart., B 2326, Ph
Indigofera gracilis Bong.,B 2057,H
ps nus subsessilis Benth.,B 2320,H
lium Vogel, B 2115, Ph
Mimosa amnis-atri Barneby, B 3589, Ph
Mimosa distans Benth., B 2070, Ch
Mimosa foliolosa Benth., B 2367, Ph
Mimosa gemmulata Barneby, B 2012, Ph
Benth., B 28
haeriiim
305
imosa gracilis Benth., B 2395, ee
Mimosa hebecarpa Benth., B 1911,Ph
Mimosa nuda Humb. & Bonpl., B 2074, H
Mimosa polycephala Benth., B 2391,Ch
Mimosa radula Benth., B 2452,Ch
pla ea thocentra Mart.,B 2114,Ch
eriandra mediterranea ee B 2671,Ch
eee firmulus Mart., B 3488,
Plathymenia reticulata Benth., am Ph
Poiretia al ia ea B 2386,H
Poiretia latifolia Vogel, B 3007,H
Aaa, hs oe Harms, B 2599, H
pubescens Benth., B 3833, Ph
Sonne platyphylla Benth., B 2653,H
Senna rugosa (G. Don.) Irwin & Barneby, B 2329,
Ph
Senna silvestris (Vell.) Irwin & Barneby 4,Ph
Aw
Senna velutina (Vogel) Irwin & Lach = ee
Ph
si ee a adstringens (Mart.) Coville, B
Reem obovatum B 2087,Ph
Share bracteata Vogel, B 2155,H
Stylosanthes gracilis Kunth, B se
Stylosanthes guianensis Sw., B 2423, Th
" ee scabra Vogel, B oe -
\/Atniron mracr,
Vigna linearis Kunth, B 2838, if
Zornia latifolia Sm., B 2676,H
Zornia reticulata Sm., B 2481,H
Fabaceae sp. 1,B 2796, Ph
FLACOURTIACEAE
Casearia grandiflora Cambess., B 2078, Ph
Casearia sylvestris A B 2473,Ph
Casearia sp.,B 2034
Se tahoe
nervosa Cham. & Schitdl., B 3457, H
i alata (Aubl.) Maas, B 2879,H
Irlbachia speciosa (Cham.& Schltdl.) Maas, B 3438,
H
GESNERIACEAE
Sinningia elatior (Kunth) Chautems, B 3989, H
HYPOXIDACEAE
Curculigo sp.,B 4019, Geo
oe CEAE
trim nitens (Renth ) Miers, B 3244, Ph
306
IRIDACEAE
iM vaginatum Spreng. B 2253,Geo
isyrinchi U
Trimezia juncifolia (Kl) Kunth, B 2062, Geo
Eriope crassipes Benth., B 2054, H
Hypenia macrantha (A. St-Hil. ex Benth.) Harley,
B 3704, H
Hyptidodendron canum (Pohl ex Benth.) Harley,
93,
Hyptis adpressa A. St-Hil.ex Benth., B 4013, H
Hyptis capriariifolia Pohl ex Benth., B 3006, H
Hyptis caudat a pling & Sativa, B 3643,H
it7 R th
Hy ry ti
nnitad benMN,, B
ane desertorum Pohl ex Benth, B 4015,H
Hyptis eriophylla Pohl, B 2849,
Hyptis ferruginosa Pohl ex seen
/B 3979,H
3076,H
Hyptis lythroides Pohl ex ae 8 3708, H
Hyptis multiflora Pohl ex Benth., B 2670, H
Hyptis recurvata Poit.B 3535,H
Hyptis saxatilis A. St-Hil.ex Benth., B 311
Hyptis villosa Pohl ex Benth., B 2255, H
Hyptis virgata Benth., B 1925,H
Hyptis s LH
Maleveant thes chamaedrys (Vahl) Kuntze, B 2531,
H
1,ch
7 Benth., B 3108, H
NM irsypianthes
Ocimum sp.,B 2151,H
Peltodon pusillus Pohl, B 3271,H
Pohl, B 3065, H
Dp|tr
/ eltodon tome Ntosus
Salvia sp.,B 2
LAURACEAE
Aiouea trinervis Meisn., B 3931, Ph
Cassytha filiformis L., B 2422,VP
LECYTHIDAC
Eschweilera nana oe Berg) Miers, B 2825, Ph
LOGANIACEA
ae ne St-Hil., B 3943, Ph
LYTHRACEAE
pais eOHAg ene nsIs 5 Jacq.) Macbr.,B 2179,H
Cuphe Koehne, B 2136, |
nee pacari A. St-Hil.,B 3782, Ph
MALPIGHIACEAE
eg Barium aie B. Gates, B 2809, Ph
ates, B 2429, Li
Bani eee Rance (A. i‘ ss.) Little, B 2346, Li
Banisteriopsis gardneriana (A. Juss.) W. Anderson
attl., B 3756, Li
BRIT.ORG/SIDA 20(1)
Banisteriopsis laevifolia (A. Juss.) B.Gates, B 2558,
Li
Banisteriopsis schizoptera (A. Juss.)
3049, Li
Banist ter lopsis stellaris (Griseb.) B. Gates, B 2479, Li
Banisteriopsis variabilis B. Gates, B 2304, Li
Byrsonima bastloba A. Juss.,B 2133,Ph
Byrsonima coccolobifolia A. Juss., B 2233, Ph
Byrsonima crassa Nied., B 1898, Ph
Byrsonima gaultherioides Griseb., B 2898, Ch
Byrsonima quilleminiana A. Juss.,B 2044, Ch
Byrsonima intermedia A. Juss.,B 2467, Ph
Byrsonima rigida A. Juss.,B 2135,Ch
(Griseb.) B.Gates, B 2643,
B. Gates, B
Byrsonima verbascifolia
Ph
nies affinis A. St-Hil., B 2417
5 tera A. JUSS., B 3149, Li
Het ne hveonietol ia A. Juss.,B 2476, Ph
Juss.,B 2172, Li
t pt
camnestris A
v
eterop fervs
-
He A SS., : 3089, Li
Pei ea ret Cla B 2079, Li
Tetrapteris ambi (A. Juss.) Nied., B 3279, Li
MALVACEAE
Byttneria oblongata Pohl, B 21
Eriotheca gracilipes (K.Schum. " in eobyne B 1936,
Ph
Eriotheca pubescens (Mart. & Zucc.) A. Robyns, B
36, Ph
Helicteres sacarolha A. St-Hil., B 2468, H
Krapovichasia macrodon (A.DC.) Fryxell,B 2175,H
Melochia villosa (Mill) Fawc. & Rendle, B 2499, H
Pavonia rosa-campestris A. St-Hil.,B 1962, H
Peltaea vial (Hochr.) Krapov. & Cristobal, B
1940,
rca eure (A.St-Hil.)
H
Krapov.& Cristobal,
B 2528,
ae ce ee longiflorum (Mart. & Zucc.) A.
ns,B 3507, Ph
Sida Bea kee Krapov., B 2960, H
Sida cordifolia 61,H
inearifolia A St-Hil., B 3051,H
* B 2840, H
A. St-Hil.B 2028, H
Waltheria indica L.*,B 2274,H
MELASTOMATACEAE
Miconia albicans Triana, B
ba iconta fallax A DC, B 1986
>
Sida
Sida rhombifolia L.
\A Ith i 4 ,
1987,Ph
5, Ph
Jil B 1988, Ph
Micon algo an (A. DC.) Naud, B 3465, Ph
BATALHA AND MARTINS,
CIAO WALI AL PARK
Miconia (Bon ipl.) : - B 1956, Ph
Mouriri elliptica Mart., B 213
Rhynchanthera ursina nn : 3536, Ch
Tibouchina gracilis (Bonpl.) Cogn., B 2635, Ch
Tibouchina stenocarpa (A.DC.) Cogn., B 2800, Ph
MENISPERMACEAE
Cissampelos ovalifolia Ruiz & Pav.,B 1954,H
MORACEAE
Brosimum gaudichaudii Trecul, B 2109, Ph
MYRISTICACEAE
Virola sebifera Aubl.,B 3219,Ph
MYRSINACEAE
Myrsine leuconeura Mart., B 2820, Ph
MYRTACEAE
Campomanesia adamantium (Cambess.) O. Berg,
Campomanesia pubescens (A. DC.) O. Berg, B
2226, Ph
Eugenia angustissima O. Berg, B 2413,H
Eugen a aurata 0. Berg, B 2699, Ph
Euget varginata A.DC.,B 3502, Ph
Eug ae alycit eens B 2618,Ch
Ei j pli Berg, B 3583,Ch
eoenta aisaenst is a Bae) B 2231,
Eugenia piauhiensis O. Berg, B 2025, Ph
Eugenia piloesis Cambess.,B 2511, Ph
Eugenia punicifolia (Kunth) A.DC.,B 2265, Ph
Eugenia sp.1,B 4017,Ch
Eugenia sp. 2,B 3601,Ph
Eugenia sp. 3,B H
Eugenia sp.4,B 2
Mice og: Combes, B2127,Ph
1a UTE
Myrcia bracteat erg, B 2673, Ph
My rcia camapuanensis N.F.E. Silveira, B 1 i Ph
aes crassifolia (O. Berg) Kiaersk.,,B 2228, Ph
ia O. Berg, B 1994, Ph
lanes fallax (Rich.) A. DC., B 3689, Ph
Myrcia guianensis A.DC.,B 3691, Ph
Myrcia laruotteana Cambess., B 3927,Ph
Myrcia lasiopus O. Berg, B 2689, H
Myrcia linguaeformis Kiaersk., B 3555, Ph
Myrcia rhodeosepala peas B 3817,Ph
MENS Ne A. DC,,B 23,Ch
oe B 3561, Ph
Wvrcia variabilis Mart. ex A.DC.,B 2041, Ph
Myrcia sp. 1,B 2238,Ch
Myrcia sp. 2,B 3260, Ph
Myrcia sp. 3,B 3442,H
Myrciaria delicatula (A.DC.) O. Berg, B 2995, H
Psidium australe Cambess. B 2777,Ch
Psidium cinereum Mart.,B 2212, Ch
sched ae O. ee B 3848, Ch
arl Cambess.,B 2250, Ph
multiflorum Cambess., B 2547,Ch
Psidium rufum Mart. ex A.DC.,B 2199, Ph
NYCTAGINACEAE
Guapira campestris (Netto) Lund., B 3969, Ch
Guapira gracilifora (Mart. ex J.A. Schmidt) Lund.,
dies laa)
voxia (Netto) Lund.,B 3552, Ph
Neca aeonedl a Poepp. & Endl.,B 3080, Ph
Neea theifera Oerst.,B 2110, Ph
OCHNACEAE
Pas acuminata (A.DC.) Engl., B 2463, Ph
lia (A.DC.) Engl, B 3577, Ph
Ouratea floribunda (A. St-Hil.) Engl, B 2219, Ch
Ouratea nana (A. St-Hil.) Engl., B 2462, Ch
Ouratea spectabilis (Mart.) Engl., B 2045, Ph
ORCHIDACEAE
istephium sclerophyllum Lindl., aes Geo
Gc Heanala montana Barb. Rodr.,, B 2727, Geo
H aria brevidens Lindl., a ]
Habenaria nasuta Rchb. f.& Warm., B 2751, Geo
Habenaria obtusa Lindl. B 2633, Geo
OXALIDACEA
Oxalis sel ovis B1910,H
PASSIFLORA
Mitostemma Aa F 5 Gontsch., B 3562, Ch
Passiflora mansoi (Mart.) Mast., B 3066, Li
OACEAE
Actinocladum verticillatum (Nees) McClure ex
Soderstrom, B 3837, Ph
Andropogon bicornis L., B 2866, H
Andropogon fastigiatus Sw.,B 3157,H
Andropogon leucostachys Kunth, B 1922, H
Andropogon selloanus (Hack.) Hack., B 2614, H
anna perforata (Nees) Parodi, B 2344,
| icola McClure, B 3855, Ph
Austide Runa Trin., B 3233
Aristida riparia Trin., B 2560, H
aureus P. Beauv., B 3017,H
arbigerus (Kunth) Hitchc., B 2763, H
Axonopus brasiliensis (Spr.) Kuhim., B 2890, H
Cane Ciao eins Black, B 2291,H
Brachia Stapf *,B 2311,H
308
Cteniun ne idense ( Trin.) one ; 3420,H
Eehinnipenn jalove (Poir,) Chas 2382,H
Flionurniys latiflorys Nees, B 377
ragrostis articulata (Schrank) B 2556, Th
Eragrostis hae s (Kunth) Steud.,B 2170, Th
Gymnopogon foliosus (Willd.) Nees, B 3083, H
Hyparrhenia eee (Humb. & Bonpl.) Stapf, B
3833,H
Hy rrhenia rufa (Nees) Stapf *,B 3181 H
ie natn procurrens (Nees) Sw.,B 2561,H
Leptocoryphium lanatum (Kunth) Nees, B 3700, H
Loudetiopsis chrysothryx (Nees) Conert, B 2845,H
Aelinis minutiflor 1P. Beau or, B 2278, H
Olyra taquara Sw.,B 3949, Ph
Panicum o eee Kunth, B 1928, H
Panicum em.& Shult., B 2491, H
Panicum s
a H
aupeiuns a carinatum Humb.& Bonpl. ex Fleug.,B
2002, H
Paspalum convexum Humb.& Bonpl. ex Fleug.,B
2972,H
Paspalum erianthum Nees, B 2405, H
ie gardnerianum Nees, B 2354, H
iff Steud.,B 3250,H
lum matacophyl um Trin.,B 3050, H
lt B 3168,H
Ppl a ae B 2343,H
Paspalum sp. 1, 0,H
Sane sp. 2,B 2938,H
Pennisetum setosum (Sw.) L.C. Rich. *, 3363, H
Rhynchelytrum repens (Nees) C.E. Hubb.*,B 2572,
Th
Pasa
wim lt jule Poi
—
Schizachyrium condensatum (Kunth) Nees, B
30,H
Setaria geniculata (L.) P. Beauv. *, B 2495, Th
Sporobolus acuminatus Boechat & Longhi-Wag-
ner,B 218
ciliatus (Trin.) Hack., B 2986, H
Sporobolus indicus (L.) R. Brown, B 2519, H
Sporobolus tenuissimus (Schrank) Kuntze, B 2178,
|
Snorohol)s«
Sp 5
Thrasya petrosa Nees, B 2630,H
Tristachya leiostachya Nees, B 2269, H
POLYGALACEAE
Polygala angulata A.DC.,B 2425,H
Polygala aphylla AW. Benn., B 3910, H
Polygala opit 1a Wurdack, B 2223, H
Polygala violacea Aubl., B 2492,H
Securidaca tomentosa A. St-Hil., B 3547, Li
Kea)
BRIT.ORG/SIDA 20(1)
POLYGONACEAE
C loba densifions Mart. ex Meisn.,B 2111, Ph
POLYPODIACEAE
Adiantum serratodentatum Humb. & Bonpl. ex
Willd.,B 3403, H
PROTEACEAE
Roupala montana Aubl.,B 2144,Ph
RHAMNACEAE
Crumenaria polygaloides Reissek, B 2920, H
RUBIACEAE
Alibertia sessilis (Vell.) K. soe 7 ia ch
20/eNS Mey ers BI
reria SUaVE
Bor
Chomelia Neale Benth. ex re ao B 3928, Ph
[; VATA
aU 1.
H
Dec lieu IXiQ a oenanth des Schult & Schult, B 2040,H
Paes Verte Hata } Arg., B 2245, H
i Standl., BI 972,Th
ene teres Walt. B 2524,Th
Ga janthe nega idifolia ELL. rae B 2944,H
7L.,B 2449, Ph
—
Palicoureacoracea Cham.) x ae B 1903,Ch
alicourea rigida | Kunth, B ]
Richardia humistrata (Cham. ‘ Schitdl. ) Steud.,B
1909, H
Richardia stellaris (
Cham. & Schltdl.) Steud., B
3036, H
Sipanea hispida Benth., B 2485, H
Tocoyena formosa (Cham. & Schltdl.) K. Schum.,
2124,Ph
RUTACEAE
ene pido abies ex - DC.,B 2765, Ph
St t-Hil.,B 2073, Ch
SAPINDACEAE
Matayba guianen be ae sa Ph
Serjania ci i4 3, Li
sey ese a se
nia ret Cambess., B 3040, Li
fais gost Radlk.,B 1935,Ch
cia tomentosa Radlk., B 3088, Ch
SAPOTACEAE
Pouteria ramiflora (Mart.) Radlk.,B 1984, Ph
Pouteria subcaerulea Pierre ex Dubard, B 1899,Ch
Pouteria torta (Mart.) Radlk., B 1918, Ph
Pradosia brevipes (Pierre) Penn.,B 3772,H
SCROPHULARIACEAE
Buchnera lavandulacea Cham.& Schltdl, B 2437,
H
BATALHA AND MARTINS, CERRADO FLORA OF EMAS NATIONAL PARK 309
Esterhazya petiolata Barr.,B 3358,H Lippia hirta Schauer, B 2415,Ch
Scoparia dulcis L.*, B 2486, Ch ae hoehnei Mold., B 3690, Ch
Lippia lupuli ina ena B 3642,Ch
SIMAROUBACEAE
Simaba suffruticosa Engl. B 3730, Ch
Simarouba amara Aubl.,B 3802, Ph
Lippia uer, B 2254, Ch
Lippia primulina . oe B 3966, H
Lippia stachyoides Cham., B 2257,H
SMILACACEA Lippia turnerifolia Cham.,B 2505,H
Smilax cissoi ne Mart. ex Griseb., B 2330, Li Stachytarpheta maximilliani Schauer, B 2401,4
SOLANACEAE Stachytarpheta simplex Hayek, B 2369, H
Solanum lycocarpum A. St-Hil.,B 2020, Ph VIOLACEAE
Solanum subumbellatum Vell, B 4005, Ch Hybanthus poaya (A. St-Hil) Baill. B 3947,H
STYRACACEAE Hybanthus sp. nov.,B 3513
Styrax ferrugineum Nees & Mart., B 3227,Ph VITACEAE
TURNERACEAE Cissus erosa L.C. Rich, B 2458, Li
iriqueta emasensis Arbo, sp.nov., B 1968, H VOCHYSIACEAE
Pir ioaeia sidifolia (Cambess.) a : 3059,Ch Qualea grandiflora Mart., B 2065, Ph
Turnera purpurascens Arbo, B Qualea multiflora Mart., B 2434, Ph
VERBENACEAE Qualea parviflora Mart.,B 3952, Ph
ene scar hme 2303, Ch Vochysia thyrsoidea Pohl, B 2892, Ph
_B 23] 5, Ph Vochysia ftucanorum Mart., B 2496, Ph
Arnasonia hirta Benth., ah 385,H Unknown
Casselia chamaedryfolia Cham., B 2066, H Unknown sp. 1, B 2305, Ch
ACKNOWLEDGMENTS
We are grateful to Fapesp, for financial support; to Ibama, for granting the re-
search permit; to the Emas National Park staff, for their help in field; to M.B.
Ramos-Neto, for logistical assistance; to M.ER. Siles, for translating the Abstract
into Spanish; to W. Mantovani, for useful discussions and assistance in species
identification; to the taxonomists K. Althott, M. Alves, M.M. Arbo, M.C. Assis, C.
Bestetti, R.S. Bianchini, V. Bittrich, A.C. Cervi, I. Cordeiro, C.L. Crist6bal, T.S.C.
Dorow, S.I. Elias, G. Esteves, M.A. Farinaccio, M.S. Ferrucci, P. Fiaschi, E. Forero,
A. Furlan, E. Goncalves, M.L. Kawasaki, M. Kirizawa, A. Krapovickas, J.A.
Lombardi, M. Magenta, A.C. Marcato, R. Mello-Silva, E. Melo, M. Moraes, R.C.
Oliveira, M.B. Paciencia, J.R. Pirani, A.P Prata, C. Proenca, N.A.T. Ranga, ER.
Salimena, J.U.M. dos Santos, J. Semir, M.F Simon, E.R.V. Silva, M.M. da Silva, J.P.
Souza, A. Spina, J.Y. Tamashiro, W. Thomas, R.B. Torres, M.G. Wanderley, and A.
Zanin, for their assistance in the identification of species; and to J. Ratter and
W. Thomas, for their useful comments on the text.
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BOOK REVIEW
IRWIN M. Bropo, SyivIA DuRAN SHARNOFF, and STEPHEN SHARNOEFF. 2001. Lichens of
North America. (ISBN 0-300-08249-5, hbk.). Yale University Press, 302 Temple
Street, PO. Box 209040, New Haven, CT 06520-9040, U.S.A. (Orders: 203-
432-0960, 203-432-0948 fax). $69.95, 795 pp, 924 color photos, distribu-
tion maps, 91/2" x 11".
This recently published magnum opus Lichens of North America is a beautiful volume that treats an
often overlooked eroup of oa And this book has everything! The book is divided into two
parts. The first part begins y material including a wonderful description of the sym-
biotic relationships that lichens form and moves on to discuss lichen morphology, anatomy, repro-
duction, physiology, chemistry, ecology and biogeography. Two chapters describe how humans have
found uses for lichens and how we may use them as environmental indicators. There are also two
chapters offering very practical information on how to describe and classify lichens and how to col-
lect and preserve them for future study. The final chapter in the first part of the book aged useful
information about how to use the maps, keys and descriptions in the second half of the boo
The second part of this volume provides easy to follow keys, we ‘ crafted descripti Cron’
ful photographs, and very useful range maps for 805 species of lichens. The photogr: a0 phs and range
maps should make oe aaa extre — useful anyone — in lichens whether they be
s A
sat
professionals
he back of the book provides
a most oe guide to the peteen oe enti a group a organisms about which many
people know very little.
ne introductory text in this book is extremely well written and immensely readable. And the
eee . simply wi a of art. This nes ou ace book should be on
fiil diver
the shelf o f every one wl , anc appreciate tt to be found even
a
—_
in one of the least well-known ned eee groups of organisms. It will aively inspire many of us
totake a closer look the next time we see a lichen.—Debra Trock, Botanical Research Institute of Texas,
509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A.
SIDA 20(1): 312. 2002
RUST FUNGI (UREDINALES) OF NORTHWEST ARGENTINA
José R. Hernandez Joe F.Hennen
Systematic Botany and Mycology Laboratory Botanical Research Institute of Texas
USDAZARS, Rm. 304, BOTTA €
Beltsville, MD 20705-2350, U.S.A. Fort Worth, TX 76102-4060, U.S.A
ABSTRACT
This paper presents the results of a study of rust fungi (Uredinales) on wild and culated plans in
the provinces of Catamarca, Jujuy, Salta and Tucuman that belong to the region
Argentina. The rust fungi are represented by 635 collections on 257 host plant species belonging to 51
vascular plant families. These rusts constitute 182 species in 30 genera of which 61 are new records
10
for Argentinaan
for Argentina, and 50 of these are new host records for rusts
Fundacion Miguel Lillo (LIL) or the US. National Fungus Collection (BPD.
are new species of rusts. Eighty-four of the collecuons tepresent new host records
|
S ted in either
ts
RESUMEN
En el presente trabajo se stan los resultados de la prospeccion de royas (Uredinales) realizada
sobre la flora silvestre y cultivada de las Provincias de Catamarca, Jujuy, Salta, y Tucuman,
pertenecientes a la region ener como Noroeste Argentino. Como resultado de este trabajo se
eliootaron 635 GOlecclgnes cortespelidientes: a 51 familias de hospedantes (257 especies). Ochenta y
| Argentina, de las cuales 50 son nuevas citas
I Se identifi 309 gies 182 especies de royas, de las cuales 61 son
nuevos ee para el pais y 10 nuevas especies. Las colecciones fueron depositadas en Fundacion
Miguel Lillo (LIL) y U.S. National Fungus Collection (BPD.
The fungi of Argentina have been studied by a number of mycologists, most
notably Carlos Luigi Spegazzini (1858-1926) whose taxa were compiled into a
comprehensive account by Farr (1973). Spegazzini’s collections of rust fungi
(Uredinales) as well as collections made by later mycologists were organized
and studied by Lindquist (1982). Lindquist’s studies resulted in numerous pub-
lications on rust fungi, culminating in his “Royas de la Republica Argentina y
Zonas Limitrofes” (Lindquist 1982). Although the fungi of Argentina are rela-
tively well known, the rust fungi of Northwest Argentina are less well studied
because few mycologists collected rusts in that region.
In this paper we present a study of rust fungi collected on wild and culti-
vated plants in the provinces of Catamarca, Jujuy, Salta and Tucuman in North-
west Argentina. This subtropical region is characterized by climatic and bio-
logical diversity and consists of a plain in the east and the Cordillera de los
Andes in the west where valleys closely alternate with mountains. The grada-
tions in latitude and altitude in Northwest Argentina result in a mosaic of
bioecosystems with vegetation types varying from desert to rain forest. The di-
versity of flora is reflected in the range of phytogeographical regions described
SIDA 20(1): 313 — 338. 2002
314 BRIT.ORG/SIDA 20(1)
for this area by Cabrera (1994). For these reasons, Northwest Argentina is an
interesting and rich area for collecting rust fungi.
Lindquist (1982) documented 450 species representing 28 genera of rust
fungi from throughout Argentina of which 124 species were reported from
Northwest Argentina. Between 1993 and 1999 the authors collected and identi-
fied or re-examined 635 specimens of rust fungi representing 182 species in 30
genera. These rust specimens were collected on 257 host plant species from 51
plant families. Fighty-four of these host plants are new host records for Argen-
tina, and 50 of these are new host records for rusts worldwide. Sixty-one rust
species are reported for the first time from Argentina and 10 of these are new
species. One of those, Baeodromus ranunculi J.R. Hern. & J.E Hennen, has been
described (Hernandez 2000) and the remainder will be described elsewhere.
RESULTS
The rust species reported here are presented according to host family. This is
the most useful form of organization because of the high level of host specific-
ity of the rust fungi. Collections for each rust species are then listed alphabeti-
cally by host species followed by the province, details of locality, date, collector(s)
and collector’s number, herbarium number and rust stage present. For the rust
stages symbols are used as follows: 0 = spermogonial stage, I = aecial stage, II =
uredinial stage, and II] = telial stage. The abbreviation “An” refers to the pres-
ence of an anamorph stage for which we are uncertain whether it is function-
ally an aecial or uredinial stage. Symbols representing spore types are X =
amphispores, M = mesospores, and lV = basidiospores.
New rust records are indicated in superscript following the species name
as follows: new rust species (N), new rust record for Argentina (A), Catamarca
(C), Jujuy QD, Salta (S), and Tucuman (T). Superscript Helauons following host
names indicate: new host plant genus for this rust worldwide (G), new host plant
species for this rust worldwide (S), new host plant species for any rust in Argen-
tina (A), and new host plant species for which another rust was previously
known from Argentina (AR).
Abbreviations of authorities for plant host and fungal names follow
Brummit and Powell (1992) and Kirk and Ansell (1992) respectively. Plant host
names were verified using W3TROPICOS at the Missouri Botanical Garden.
ACANTHACEAE Puccinia justiciae Puttemans
Accidium sp.* on Chaetothylax umbrosus Nees’, Tucuman:
on Dyschoriste sp.°; Salta: Quebrada de San San Pedro de Colalao, 2 Apr 1994,
Lorenzo, 6 Dec 1997, |. R.Herndndez 97- j.FHennen & M.M.Hennen 94-069 (LIL
162 (BPI 841258) [I]. Tucuman: 22 km W 54706, BPI 841031) [I].
of San Miguel de Tucuman, mtn, 27 Mar Puccinia lateripes Berk. Ravenel"
1993, J.RHennen, L.D.Ploper & on Ruellia ciliatiflora Hook.; Tucuman
J.R.Herndndez 93-019 (LIL or BPI Dept. Trancas, Vipos, 30 Mar 1995,
840996) III. J.R.Herndndez 95-043 (BPI 841086) [H-
HERNANDEZ AND HENNEN, RUST FUNGI OF NORTHWEST ARGENTINA
III], Las Salas, 5 May 1995, ].R.Herndndez
95-015 (BPI 841079) [I-II-III}; San Javier,
Pa rque Sierras de San Javier, 23 Jul 1994,
|. R.Herndndez 94-150 (LIL 54704, BPD [II-
Ill]; San Pedro de Colalao, 2 Apr 1994,
J.FHennen & M.M.Hennen 94-071 (LIL
54703, BPD [0].
Puccinia sp.
on Ruellia er ‘ythrop oni eee Catamarca:
E of La Vina, 24 Ser 7, .FHennen &
J. R-Herndndez 97-061 is 197) II]
Puccinia thunbergiae Cooke*
on hana sp., Tucuman: San Javier, Parque
rras de San Javier, 23 Jul 1994,
fe erndndez 94-144 (LIL 54705, BPI
841067) [IL-HIL.
Uromyces sp."
on undtn. Jujuy: Palma Sola, 12 Apr 1994,
J.FHennen & L.D.Ploper 94-127 (LIL
54707, BPI 841056) [II].
AMARANTHACEAE
Uromyces bonaerensis Speg
on Gomph rena elegans Mart. Tucuman: Cruz
Alta, 28 Jul 1913, leg. ? [II].
Holw.*
on Chamissoa altissima Jacq.)
Uromyces celosiae Dietel &
Kunth
Tucuman: San Javier, Parque Sierras de
San Javier, 23 Jul 1994, J.R.Herndndez 94-
165 (LIL 54708, BPI 841074) [II]
Uromyces iresines Lagerh. ex Syd. & P. Syd.
on Iresine diffusa Humb. & Bonpl. ex Willd.
var. diffusa*; Tucuman: San Javier. Parque
Sierras de San Javier, 23 Jul 1994,
).R.Herndndez 94-140 (LIL 54709, BPI
841064) [III]
ANNONACEAE
Phakopsora neocherimoliae (Cummins) Buritica
& J.F Hennen*
h
on Annona cherimol
ia Mill; Tucuman: Dept.
se 15 Sep 1995, J.R.Herndndez 95-
088 (BPI 841111) (II; Dept. Famailla, Sauce
Huascho, 23 Apr 1993, N.E.V.de Ramallo
s.n. (BPD [IIL
APIACEAE
Puccinia hydrocotyles Cooke’ >
on oe rocotyle bonariensis Comm. ex Lam.”;
nos Aires: La Plata, 29 Sep 1997,
J. 7 nnen & J.R.Herndndez 97-072 (BPD)
[II]. Catamarea: N of Catamarca, Hwy. 38,
24 Sep 1997, J. FHennen & J.R.Herndndez
oo road to El Rodeo, camp-
area, 29 Nov 1997, ie nee 97-
ca ) [UI W of Catamarca, 23S p 1997
J FHennen & J.R.Herndndez 97- ae 1)
[Il]; W side of Catamarca, 23 Sep 1997,
J.FHennen & J.R.Herndndez 97-035 (BPI
841184) [II]. Salta: Dept. Santa Victoria, La
Mision, 30 Jun 1996, J.R.Herndandez 96-
059 (BPD [I]. Tucuman: Lules, Aug 1994,
J. R.Herndndez 94-149 (LIL 54943, BPD [II]
on Hydrocotyle sp., Catamarca: El Rodeo, 29
Nov 1997, |.R. Hernandss aie a :
Tucuman: Amberes,
L.D.Ploper & G. eae ere $n. ae
Dept. Trancas, Rt. 9, km 1364, 26 Oct 1995,
J. R-Herndndez 95-052 (BPD [LH] San Javier,
— Sierras de San Javier, La Cascada.,
9 Mar 1996, |. R.Herndndez 96-014 (BPI
sai I].
ASCLEPIADACEAE
Puccinia sp.
on Morrenia ae d ee T. me
Tu : Alta Gracia, 997,
ee se oe a ne a
ASTERACEAE
Aecidium amplia HLS. Jacks
on Eupatorium ante
_& Holw.
hthalmum Griseb.;
Catamarca: near Balcozna, 28 Nov 1997,
J.R.Herndndez a 113 (BPI 841226) [0-I].
B. Alberdi, road to Escaba,
bank of creek, 28 Nov 1997,
J.R.Herndndez 97-103 oH 841221) [0-1].
CIs
Tucuman: J.
Aecidium niederleinii Henn.
n Conyza a ; Catamarca: Dept. Ambatos, 1
Ro 14 Nov 1999, |. R.Herndndez 99-
007 yes #1276) [O-I]. Salta: Dept.
Gtiemes, La Calderilla, 7 Dec 1997,
UR. Hemaandez oT- ne (BPI) [I]. Tucuman:
Alta Gracia, 19 Nov 1997, ].R. Herndndez
97-0932 (BPI pene ed Alta Gracia, 19
Nov 1997, ].R.Herndndez 97-087 (BPI) [0-
I], El Siambon, river next to monastery, 3
Nov 1997, J.R.Herndndez 97-145 (BPI
841243) [0-I],
Aecidium sp.
on Senecio cremeiflorus Matth; Catamarca:
Rio Las Juntas, 23 Sep 1997, ].FHennen &
J.R.Herndndez 97-051 (BPI 841192) [I].
316
Cionothrix praclonga (G. Winter) Arthur®
on Eupatorium sp.*“; Salta: Dept. Santa Vic-
toria, Los Toldos, El Nogalal, 29 Jun 1996,
J.R-Herndndez 96-040 (BPI 841135) [HII
Tucuman: San Javier, Parque Sierras de
San Javier, 27 Mar 1993, J. Hennen,
L.D.Ploper & J.R.Herndndez 93-033 (LIL
54734, BPD [III San Javier Parque Sierras
de San Javier, 27 Mar 1993, |. FHennen,
L.D.Ploper & E.R. Pee 93-035 (LIL
54735, BPI) [III].
Coleosporium tussilaginis (Pers.) Lév.
on Senecio bravensis Cabrera’; Buenos Aires:
Partido de Gral. Pueyrredon, 2 Nov 1944,
A.L. Cabrera s.n. [Hl].
on Senecio cremeiflorus Mattf.; Catamarca:
Dept. Ambatos, El Rodeo, 14 Nov 1999,
J.R.Herndndez 99-004 (BPI 841273) [II], N
of Catamarca, Hwy. 38, 24 Sep 1997,
J.RHennen & J.R.Herndndez 97-055 (BPI
841194) [II].
on Senecio deferens Griseb. Tucuman: Sier-
ras de San Javier, 17 Nov 1949, Singer &
ae io S.n. na
on S$ 5G », Catamarca: E|
Rede: 39 | ie - 1907. IR Herndndez 97-
126 (BPI841231) [II].
on Senecio rudbeckiifolius Meyen & Walp
Tucuman: Tali del Valle, 6 Apr 1995,
|.R-Herndndez 95-077 (BPD [I]; Tafi del
Valle, Apr 1993, j.RHennen &
J.R.Herndndez 93-120 (LIL 54736, BPDIIIL.
on Senecio sp.; Salta: road to Parque del Rey,
6 Dec 1997, |. R.Herndndez 97-158 (BPI
841254) [HI].
vernoniae Ber!
[oy
Sy
Coleosporium kK. &@ M.A. Curtis’
on Elephantopus mollis Kunth**; Tucuman:
San ae Parque Sierras de San Javier, 27
Mar 1993, J.kHennen, L.D.
J. R.He a 93-029 (LIL 54737, BPD [Uf
San Miguel de Tucuman, 20 Jun 1994,
J.R.Herndndez 94-168 (BPI 841075) [II].
Puccinia baccharidis Dietel @ Holw<*
on Baccharis lilloi Heering*®; Salta: Cafayate,
9 Apr 1994, |. RHennen, M.M.Henn
J.R.Herndndez 94-11] (LIL 54754, B
841046) [I-III].
Ploper &
on Baccharis salicifolia (Ruiz & Pav) Pers.:
Sep 1997,
J.EHennen & |.R.Herndndez 97-023 (BPI)
Balcozna, 22
en &
BRIT.ORG/SIDA 20(1)
II-III]; N of Catamarca, 22 Sep 1997,
jJ.EHennen & J.R.Herndndez 97-019 (BPI
841173) [II road to El Rodeo, ane:
area, 29 Nov 1997, | R Herndndez
(BP1 841228) [II-IL; W side of €
23 Sep 1997, |. FHennen & | R. Hernandez
97-028 (BPI 841178) [FIL].
Puccinia calcitrapae DC. var. centaureae (DC.)
ND
S
Catamarca,
Cummins“
on Carthamus tinctorius L.*; Sa
Tartagal, General Ballivian, 25 Sep
A.C.Ramallo s.n. (BPI 841201) [II-HI.
Puccinia cnici- oleracei Pers.
on Vernonia remotiflora Rich.
‘ancas, Rio de la Cuesta, elev. 1700
m, 19 Apr 1926, S. Venturi 4138 [II].
on Vernonia saltensis Hieron: Salta: Metan,
l ene 1933, AL Cabrera 3024 [II].
eg.!
Ita: Dept.
997,
Tucuman:
om
Dept.
Puccinia colos ¢
on Coa ar Lem. Tucuman: Tali
del Valle, 6 May 1995, ].R.-Herndndez 95-
078 (BPI 841106) [III]; Tafi del Valle, 6 Apr
1993, |. FHennen & [.R.Herndndez 93-123
(LIL 54738, BPI 841019) [III].
Puccinia conoclinii Seym. in Burrill
on ea aie ium macrocephalum Less.**;
nNWo AE Concenc de la
Sierra, I] ie 1969, A. Krapovickas et al.
s.n. {I}; Dept. Burruyacu,
Campo, elev, 1500 m, 29 Mar
iis
on Ita: Dept. Rosario de la
Cerro del
1930,
Horeones 6 Dec 1997,
ez 97-154 (BPI 841251) [II].
Tucuman: Cevil Pozo, 12-15 km E of San
a de Tucuman, 30 Mar 1993,
J. EHennen & |. R.Herndndez 93-055 (LIL
54761, a 841003) [I]
Puccinia enceliae Dictel & Hala var. aemulans
(Syd. & P. Syd.) Parmelee®
on Simsiad dombeyana DC» Salta: Cafayate, 9
Apr 1994, |. FHennen, M.M.Hennen &
J.R.Herndndez 94-112 (LIL 54740, BPI
841047) [II-III].
Puccinia eupatorii Dietel>'
on Eupdtorium sp., Salta: Paes 8 Apr
1994, |. FHennen, M.M.Hennen & |.R.
Hernandez 94-101 (LIL a BP]
841042) [ILI]. Tucuman: Dept. Trancas.
Vipos, 30 Mar 1995, |. R.Herndndez 95-048
Frontera,
J.R.Herndne
Q.
HERNANDEZ AND HENNEN, RUST FUNGI OF NORTHWEST ARGENTINA
9) [II-III]; San Pedro de Colalao,
r 1994, |} FHennen & M.M.Hennen 94-
- (LIL 54741, BPI ae
Puccinia exornata Arthur
on Baccharis sp. (A)**; Salta: Quebrada de San
Lorenzo, 6 Dec 1997, J.R.Herndndez 97-
163 (BPI 841259) [I-II-III].
on Baccharis sp. (B)**; Catamarcea: naer
Balcozna, 28 Nov 1997, ].R.Herndndez 97-
112 (BPI 841225) [II
Puccinia helianthi Schwein.'
on Helianthus annuus L.; Tucuman: Dept
Capital, 17 Dec 1995, ene z 95-
095 (BPI) [II anne Molle, Parque Sierras
de San Javier, 29 Mar 1996, .R.Herndndez
a ay 841129) [II-III]; Monte
ondo, 1996, ].R. Hernandez 96-
7 eerie II].
Puccinia ee aes .) H. Mart.
on 1 m officinale Weber, Catamarca:
a Ambatos, El Rodeo, 14 Nov 1999,
J.R.Herndndez 99-003 (BPI 841272) [II].
Puccinia investita Schwein?
\DPlLOTIVOY
on Ac eae sp. Salta: Dept. Santa Victo-
ria, La Mision, 30 Jun 1996, |. R. Herndndez
96- = (BPI 841140) [II].
dq.
Puccinia lilloana J.C. Lin
Baccharis lilloi Heering; Tucuman: Tafi, 16
Apr 1904, leg. ? [III].
Puccinia lorentzii Henn.)'
on Vernonia scorpioides (Lam.) Pers.; Jujuy:
Quinta prop. Laguna de la Brea, 31 May
1901, R.E.Frias s.n. [II], Tweuman: Horco
Molle, 17 Sep 1997, J.R.Herndndez 97-075
(BPI 841205) [I].
Puccinia noccae Arthur'
on ete mollis Cav; Tucuman: Cruz Alta,
stin, 29 Mar 1994, J. FHennen,
oa ne & J.R.Herndndez 94-039
(BPDII- 1}, Dept. Cruz Alta, San Agustin,
Monte Redondo, 29 Mar 1994, |. FHennen,
M.M.Hennen & |.R.Herndndez 94-025
(LIL 54743, BPI) [II-III]; San Pedro de
Colalao, 2 Apr 1994, J. FHennen &
M.M.Hennen 94-070 (LIL 54744, BPI
841032) [II-III].
Puccinia ocellifera Cummins“
on Pluchea sp.*®; Tucuman: J.B. Alberdi, road
a, 28 Nov 1997, ].R.Herndndez 97-
097 (BPI 841218) [HI.
317
Puccinia petrakii J.C. Lindq.
on Baccharis tucumanensis Hook. & Arn.
Salta: Dept. Santa Victoria, Los Toldos, El
1, 29 Jun 1996, J.R.Herndndez 96-
048 (BPI 841137) [I-III].
ks. &@ Holw.*
on Senecio peregrinus Griseb.*, Catamarca: E
of La Vina, 24 Sep 1997, J. EHennen &
J. R.Herndndez 97-057 (BPI 841196) [II], W
of Catamarca, 23 Sep 1997, |. kKHennen &
J.R.Herndndez 97-045 (BPI 841189) [II].
Tucuman: 15 Sep 1949, R.Singer & Digilio
s.n. [I-III], Dept. Trancas, 27 Oct 1995,
J. R.Herndndez 95-059 (BPI 841094) [II]
J.B. Alberdi, S of Escaba Dike, 22 Sep 1997,
J.FHennen & J.R.Herndndez 97-010 (BPI
841168) [II]; San Javier, Parque Sierras de
San Javier, 23 Jul 1994, J.R.Herndndez 94-
162 (LIL 54746, BPD [II]; San Javier, Parque
Sierras de cee ey: 27 Mar 1993,
Ploper & J.R.Herndndez
03- 028 (LIL ae BPI) [II-Ill].
Puccinia rata HS. Jacks. @ Holw.*
on Vernonia saltensis Hieron. Salta: Capital,
Cerro San Bernardo, 27 May 1933,
A.L.Cabrera 3021 [II-III]. Tucuman: Dept.
Trancas, Vipos, elev. 780 m, 19 Mar 1928.
Screiter 47843 [I-II-Illl, San Pedro de
‘olalao, 1 Apr 1994, J. Hennen &
.M.Hennen 94-058 (LIL 54747, BPI
841028) [II-III].
Puccinia schileana Speg. var. partheniicola (HS.
Jacks. & Holw.) J.C. Lin
on Parthenium hiyslerophar
Cruz Alta, San Agustin, Ma oot i
Hennen, M.M.Hennen & J.R.
94-031 (LIL 54748, BPI e022) ae D
Dept. Trancas, Vip
J.R.Herndndez 96- oD a ae 7
San Pedro de Colalao, 2 Apr 1994,
J.FHennen & M.M.Hennen 94-066 (LIL
54749, BPI) IIL.
a Speg. var. schileana
Puccinia proluviosa H.S. Jac
Puccinia schilean
on Verbesina lilloi S.F. Blake; Jujuy: Dept.
Tumbaya, volcano, on slope to abra de
Tiraxi, 31 Dec 1952, H. Sleumer s.n. [O-I-
I].
Puccinia singeri J.C. oe
on Verbesina sp, Tucuma s de San
Javier, 15 Nov oo a aoai Sn. aE TH].
Blake:
Tucuman: J.B. Alberdi, S of Escaba Dike,
22 Sep 1997, |. FHennen & | R. Hernandez
97-OLL (BPI 841169) [II].
Puccinia sp.
on Verbesina suncho (Griseb.) S.F
on Baccharis sp. Salta: Dept. Santa Victoria,
La Mision, 30 Jun 1996, |.R-Herndndez 96
062B (BPD [I-11]. Tucuman: Tafi del Valle,
6 Apr 1993, |. FHennen & |. R-Herndndez
93-122 (LIL 54757, BPI841018) [I-H-II)
Puccinia sp. A°
on Vernonia fulta Griseb.; Tucuman: Dept.
Trancas, Rt. 9, km 1364, 26 Oct 1995,
J.R.Herndndez 95-055 (BPI 841093) [ILE
san Javier, Parque Sierras de San Javier, 23
Jul 1994, |. R.Herndndez 94-141 (LIL
54758, BP] 841065) [III]
on Vernonia sp., Tueuman: Horco Molle,
Parque Sierras de San Javier, 6 Apr 1994,
JFHennen, M.M.Hennen& LR. Herndndez
94-O91A (BPI) [I-III].
Puccinia sp. B’
on Vernonia squamosad Gardner, Catamarea:
19907, J. F.Hennen &
J.R.-Herndandez 97-022 (BPI 841175) [I-HI].
Salta: Oran, Jul-Aug 1873, PG.Lorentz &
G.Hieronymus s.n.[H-H1. Tucuman: De
Capital, elev. 480 m, 15 Aug 1909, Litto
3792 (I-III.
Puccinia sp. C*
Balcozna, 22 Sep
on Vernonia sp.; Salta: road to Parque del Rey,
1997, |.R.Herndndez 97-161 (BPI
841257) [H-lI]. Tuweuman: El Siambon.
river next to monastery, 3 Nov 1997,
J. R.Herndndez 97-147 (BP1841245) {II J.B.
Alberdi, Escaba Dike, 28 Nov 1997,
J.R.Herndndez 97-106 (BPI 841223) [If
Yerba Buena, 29 Mar 1993, |. R.Herndndez
93-047 (LIL 54755, BPD [II].
Puccinia sp.(A)
on undtn., Salta: Dept. Santa Victoria, La
Mision, 30 Jun 1996, J.R.Herndndez 96-
056 (BPI 841142) [II].
Puccinia sp.(B)
on undtn., Tucuman: NW of San Miguel de
Tucuman, 31 Mar 1993, | FHennen &
J.R.Herndndez 93-088 (LIL 54759, BPI
841009) [II-III].
Puccinia spegazzinii De Toni
on Mikania micrantha Kunth*: Tucuman:
BRIT.ORG/SIDA 20(1
Horco Molle, 15 May 1995, |. R.-Herndndez
95-079 (BPD [III Horco Molle, Parque Si-
erras de San Javier, 6 Apr 1994, |. FHennen,
M.M.Hennen & |.R.Herndndez 94-088
(LIL 54752, BPD III; San Javier, Parque Si-
Javier, ul 1994,
J. R.Hernandez 94-160 (LIL 54753, BPD [IH]
on Mikania periplocifolia Hook. & Arn.;
Los Vaqueros, 7 Dec 1997,
|. R.Herndndez 97-165 (BPI 841261) [III
Metan, 2 Apr 1993, ].FHennen ©
L.D.Ploper 93-113 (LIL 54751, BPD) [III].
Tucuman: 19.5 km W of ne Miguel de
Tucuman, mtn, 27 Mar 1993, | FHennen,
L.D.Ploper & [.R. oats 93-010 (LIL
54750, BPD [III Dept. Trancas, Vipo
Mar 1996, ].R.Herndndez 96- er
).B. Alberdi, road to Escaba, 28 Nov 1997,
R.Herndndez 97-098 (BPI) [III].
on Mikania sp. Tucuman: 15 Jan 1895, C.
erras de San
a
=
Rey
Spegazzini s.n. [I], Dept. Capital, 15 Jun
1992, |. R.Herndndez 92-005 (BPI [III].
Puccinia tanaceti DC. var tanaceti®
on Chrysanthemum sp.*®; Tucuman: San
Miguel de Tucuman, 15 Dee 1994,
J.R.Herndndez 94-170 (BP1 841076) [II
Puccinia unicolor Arthur*
on Baccharis sp.**; Tueuman: Alta Gracia, 19
Nov 1997, ].R.Herndndez 97-0931 (BPI
841214) [II],
Uredo sp.
on Baccharis sp, Tucuman: Tafi del Valle, 6
Apr 1993, ]. Fk} Hennen & |.R.Herndndez 93-
118 (LIL 54756, BP] 841017) [I].
on Eupatorium sp. Tucuman: Valle de Las
Salas, bank of Rio Las Salas, 3 Nov 1997,
J.R.Hernandez 97-151 (BPI ay [II].
Uromyces gue E LS: : icks. & Holw
on Aspilia sp.."; Tu co > Molle: 14
Nov 1997, |.R. ae 97-130 (BPI
841234) [0-11-11].
on Wedelia saltensis Cabrera’. Tucuman
Horco Molle, 21 Dec 1992, |. R.Herndndez
92-004 (BPI 840991) [II-III].
Uromyces bidenticola Arthur
on Bidens pilosa |; Tucuman: Horco Molle,
17 Sep 1997, ].R.Herndndez 97-078 (BPI
841207) [II Horco Molle, Parque Sierras
de San Javier, 29 Mar 1996, |. R. Herndndez
96-022 (BPI 841128) [I]; Horco Molle,
HERNANDEZ AND HENNEN, RUST FUNGI OF NORTHWEST ARGENTINA
Javier, 6 Apr 1994,
gercinen: M.M.Hennen&J|.R. Herndndez
94-087 (LIL 54762, BPI) [II-III]; San Javier,
Parque Sierras de San Javier, 23 Jul 1994,
J.R.Herndndez 94-151 (LIL 54763, BPD III].
on Bidens sp; Tucuman: 19.5 km W of San
Miguel de Tucuman, mtn, 27 Mar 1993,
J.FHennen, L.D.Ploper & J.R-Herndndez
93- canvas 54764, BPI) [II-III]; J.B. Alberdi,
road to Escaba, 28 Nov 1997, ].R.
ez a 096 (BPI 841217) [II].
Uromyces megalospermus Speg.
on Tessaria absinthioides (Hook. & Arn.) DC:
a: N of Catamarca, 22 Sep 1997,
ae nen nee ez 97-017 (BPI
841171) [H-III], N of Catamarca, 22 Sep
1997, |. FHennen & J.R.Herndndez 97-018
(BPI 841172) [Il-HI road to El Rodeo,
camping 29 Nov 1997, JR.
Herndndez 97-115 (BPI 841227) [II W
side of Catamarca, 23 Sep 1997,
J.EHennen & J.R.Herndndez 97-024 (BPI
841176) [Il]; W side of Catamarca, 23 Sep
1997, |. FHennen & |.R.Herndndez 97-027
(BPD [II].
Uromyces polymniae Dietel & Holw.
on dao connata (Spreng.) S.F Blake;
Tucuman:
Q,
Herndnc
area,
W of San Miguel de Tucuman,
31 Mar cee nen &J.R.Herndne
93-090 (LIL 54765, BPI 841010) [II-H1L.
aa
ez
BERBERIDACEAE
Edythea quitensis (Lagerh.) H.S. Jacks. @ Holw.*
on pee Tucuman: Tafi del Valle,6 Apr
993, J FHennen & [.R. Herndndez 93-116
aa 54711, BPI 841016) [-tM)
BETULACEAE
Mel idi } S. Ito ex Hirats. f.4
on Alniis acuminata Kunth; Salta: Dept
Santa Victoria, La Mision, 30 Jun 1996,
J.R.Herndndez 96-058 (BPI 841143) [I];
Dept. Santa Victoria, Los Toldos, 28 Jun
1996, J-R.Herndndez 96-033 (BPI 841132)
HI]. Tucuman: road to Villa Nougués, 5
May 1995, ].R.Herndndez 95-035 (BPI
841082) [II]; San Javier, 27 Mar 1993,
J.EHennen, L.D.Ploper & J.R.Herndndez
93-039 (LIL 54712, BPD [II].
BIGNONIACEAE
Prospodium appendiculatum (G. Winter) Arthur
on Tecoma stans (L.) Juss. ex Kunth in H.B.K.;
marca: E of La Vina, 24 Sep 1997,
J FHennen&J.R Herndndez 97-060 (BPI)
[I]. Jujuy: Caimancito, 2 Apr 1993,
jJ.FHennen & L.D.Ploper 93-110 (LIL
54714, BPI) [I-Il]; Parque Nacional
Calilegua, 8 Dec 1997, J.R.Herndndez 97-
180 (BPI 841268) [I]; Santa Clara, 12 Apr
1994, |. FHennen & L.D.Ploper 94-I31(LIL
54716, BPD [II]. Salta: Dept. Santa Victo-
ria, La Mision, 30 Jun 1996, J.R.Herndndez
96-055 (BPI 841141) [II-III]; General San
Martin, 7 Dec 1997, |. R.Herndndez 97-172
(BPI 841262) [I]. Tucuman: Horco Molle,
15 Nov 1995, J.R. Hernandez 95-090 (BPI)
[I]; Horco Molle, 14 Nov 1997, J :
Herndndez 97-132 (BPI II J.B. Alberdi,
Sep 1997, ].FHennen pee esas
002 (BPI ‘84 LI6L) [Ik J.B. Alberdi, road to
Escaba, 28 Nov 1997, J.R.Herndndez 97-
099 (BPI 841219) [I road to Villa Nougués,
5 May 1995, J.R.Herndndez 95-036 (BPI
841083) [I-H1]; San Javier, Aiea Sierras
San Javier, 27 Mar 1993, J.RHennen,
L.D.Ploper & J.R.Hernd se 93-027 (LIL
54713, BPD [I-I; San Pedro de Colalao, |
Apr 1994, J. EHennen & M.M.Hennen 94-
061 (LIL 54715, BPI 841030) [II-IH}.
gans (J. Schrét) Cummins”
on Tecoma garrocha Hieron.; Salta: Cafayate,
9 Apr 1994, |. EHennen, M.M.Hennen &
J.R.Herndndez 94-115 (LIL 54717, BPI
841048) [III].
Prospodium perornatum Syd.!°
Prospodium ele
n Tabebuia avellanedae L
Jujuy: Parque Nacional Calilegua, 8 Bee
1997, J.R.Herndndez 97-181 (BPI 841269)
[O-I]. Salta: Dept. Santa Victoria, La
Mision, 30 Jun 1996, se ae 96-
057 (BPI) [III]. fucuman: Cev 15
km E of San Miguel de ee - Mar
1993, J.FHennen & J.R.Herndndez 93-056
(LIL 54739, BPI) [I-III]; Dept. Capel, |
Nov 1995, |.R.Herndndez 95-062 (BPI
841095) [O-I} Dept. Capital, 1 Nov 1995,
J] R.Herndndez 95-061 (BPI) [0-1]; Dept.
Capital, 4 Apr 1994, J.RHennen ©
J. R-Herndndez 94-080 (LIL 54718, BPD [LI-
Ul]; Dept. Capital, Quinta Agronomica, 3]
Oct 1997, J.R.Herndndez 97-070 (BPD [0-
I]; Dept. Monteros, 29 Oct 1996,
J.R.Herndndez 96-077 (BPI) [0-I}; El Corte,
15 Feb 1995, |.R. Hernandez 95-067 (BPI
841099) [0-1 El Manantial, 27 Sep 1997,
J.EFHennen & |. R.Herndndez 97-064 (BP]
841198) [0|, Horco Molle, Parque Sierras
de San Javier, 6 Apr 1994, |. FHennen,
M.M.Hennen & J.R.Herndndez 94-092
LIL 54719, BPI) [H-IHI; San Javier, 21 Mar
i J.R.Herndndez 95-072 (BPI hee
HHI-I]; Yerba Buena, 5 7,
J.R.Herndndez oO
Prospodium pithecoctenii (Pazschke) Cummins*
on Pithecoctenium sp. Salta: Las Lajitas, 12
EHennen & L.D.Ploper 94-130
(LIL 54722. BP1841057) [II-IL]. Tucuman:
Las Cejas, 30 Mar 1993, J. FHennen &
J). R.Herndndez 93-061 (LIL 54721, BPD [II-
Ul]; San Pedro de Colalao, | Apr 1994,
J.EHennen & M.M.Hennen 94-055 (LII
54720, BPI 841026) [II-III].
Prospodium singeri Petr."
°F T]act
on Clyt ;(Cham.) Bureau
ex Griseb.**; Tucuman: San Miguel, Jardin
Botanico Inst. Miguel Lillo, 30 Aug 1951,
R.Singer s.n. [I-II].
Uropyxis rickiana Magnus”!
on Macfadyena unguis-cati (L.) A.H. Gentry;
Salta: Dept. Rosario de la Frontera,
nap ees 6 Dec 1997, |.R.Herndndez 97-
53 (BPI 841250) [II-I]; Posta a pees
ste J.R Hernandez 97-155 (BPD III.
Tucuman: 19.5 km W of San an de
3, J.hHennen,
L.D.Ploper & J.R.Herndndez 93-013 (LIL
54723, BPD [II-IIk Horco Molle, 15 Mar
1992, |. R.Herndndez 92-002 (BPI) [H-HI].
Tucuman, mtn, 27 Mar 199
Q.
BORAGINACEAE
Accidium tournefortiae Henn.’
on side il sity srauan ;Tucuman:
aco .R.Herndndez 97-14
(BPI eet ets
Puccinia cordiae Arthur’
on aati sp. Jujuy: Parque Nacional
calilegua, 8 Dec 1997, |. R-Herndndez 97-
ns 841266) [I]. Salta: Dept. Giiemes, 7
Dec ee roms 97-171 (BPD) [IIL
Uromyces dolichosporus Dietel & Holw.*
on Tournefortia rubicunda DC.:; Tucuman:
San Pedro de Colalao, 2 on 1994,
BRIT.ORG/SIDA 20(1)
jJ-FHennen & M.M.Hennen 94-072
54724, BPI 841033) [II-HII.
(LIL
CACTACEAE
Uromyces pereskiae Dietel!
on Pereskia aculeata Mill; Jujuy: Santa Clara,
12 Apr 1994, |. FHennen & L.D.Ploper 94-
125 (LIL 54725, BPI 841055) [IH].
CANNACEAE
Puccinia thaliae Dietel!
on Canna ee Roscoe*, Tucuman: Yerba
Bueba, 15 Nov 1995, |. R.Herndndez 95-089
(BPI 841112) (1I-HIL.
on Canna sp.; Tueuman: Dept. Capital, 12 Aug
1994, [. R.Herndndez 94-153 (LIL 54726,
BPI 841070) [II-III
CARYOPHYLLACEAE
Uromyces dianthi (Pers.) Niessl
on Dianthus sp; Tucuman: Lules, 17 Aug 1995,
).R.Herndndez 95-086 (BPI) [I].
CHENOPODIACEAE
Puccinia subnitens Dietel>
on Sudeda divaricata Mogq.; Salta: Cafayate,
pr 1994, J. FHennen, M.M.Hennen &
J.R.Herndndez 94-110 (LIL 54727, BPI
841045) [I]
COMMELINACEAE
ecta HS. Jacks. & Holw.
on Commelina sp. Tucuman: Dept. Trancas,
Vipos, 28 Mar 1996, ].R.Herndndez 96-006
(BPI 841118) [II Horco Molle, Parque Sier-
ras de San Javier, 6 Apr 1994, |. FHennen,
M.M.Hennen & J.R.Herndndez 94-095
(LIL 54728, BPD[H} San Javier, 23 Jun 1995,
J. R-Herndndez 295 OS4 (DFLOTLLOY) [II].
Phakopsora t
Puccinia commelinae Hol w.’
on she aaties sp.“ Tucuman: Horco Molle,
v 1990, ).R.Herndndez 90-
Jy Poa (oe
on Ir
Y
Salta: Dept. sania a Victoria, Los Toldos, 28
Jun 1996, [.R.Herndndez 96-032 (BPI) [II].
Tucuman: Horco Molle, Parque Sierras de
San Javier, 6 Apr 1994, ].FHennen,
M.M.Hennen & J.R.Herndndez 94-082
(LIL 54729, BPD [I].
on Tripogandra sp.*; Tacuman: 19.5 kin W of
San a de 1
1993, |.F
Herndnc
Tucuman, mtn, 27 Mar
snnen, L.D.Ploper & ].R
ez 93-009 (LIL 54730, BPD) i
HERNANDEZ AND HENNEN, RUST FUNGI OF NORTHWEST ARGENTINA
orco Molle, 17 Sep 1997, |. R.Herndndez
a 079 (BPD [Il; Horco Molle, 14 Nov 1997,
J.R.Herndndez 97-134 (BPI 841237) III;
Yerba Buena, 29 Mar 1993, |. R._ Hernandez
93-046 (LIL 54731, BPI] 841000) [I-H1].
Uromyces commelinae Cooke
on Commelina sp., Tucuman: Dept. Capital,
17 Dec 1995, J.R.Herndndez 95-093 (BPI
841114) [Il]; San Javier, Parque Sierras de
San Javier, 23 Jul 1994, |. R.Herndndez 94-
155 (LIL 54732, BPI 841071 [II-HI].
CONVOLVULACEAE
Puccinia crassipes Berk. & Curt. var. pone
on Ipomoea cairica (L.) Sweet; Tucum
ept. Capital, 27 Mar 1993, /FHennen,
L.D.Ploper & J.R.Herndndez 93-042 (LIL
54767, BPI) [IIk J.B. Alberdi, 22 Sep 1997,
J.FHennen & J.R.Herndndez 97-001 (BPI
sae ee J.B. Alberdi, road to Escaba,
eee -095 (BPI)
a ce de Tucuman, 10 Jun 1992,
J. R.Herndéndez 92-001 (BPI) [II-Hl.
hieronymi (Kuntze) O’Donell;
Jujuy: Santa Clara, 12 Apr 1994, J.F
Hennen & L.D.Ploper s.n. (LIL 54768, BPI
841058) [II-III].
on [pomoea sp.; Jujuy: Pampa Blanca, | Apr
1993, |. FHennen & L.D.Ploper 93-099 (LIL
54769, BPI 841013) [II]. fucuman: Amberes,
May 1989, L.D.Ploper & G.Wiirschmidt
s.n. (BPD [II-HI.
Puccinia dichondrae Mont.
—
on Ipomoea
on Dichondra sericea Sw.*;, Buenos Aires: La
Plata, 29 Sep 1997, |.RHennen & J.R
Hernandez 97-071 (BPI 841203) [III].
Catamarca: Dept. Ambatos, El Rodeo, I4
Nov 1999, J.R.Herndndez 99-005 (BPI
841274) [IH]. Tucuman: El Siambon, river
next to monastery, 3 Nov 1997,
Herndndez 97-149 (BPI 841246) [III].
on Dichondra sp; Tucuman: San Javier, Parque
Sierras de San Javier, 23 Jul 1994, /.R.
Herndndez 94-158 (LIL 54766, BPI) [III].
CUCURBITACEAE
Uromyces novissimus Speg.
on Cayaponia citrullifolia (Griseb.) Cogn.;
Tucuman: Dept. Capital, 12 Aug 1994,
J.R.Herndndez 94-139 (LIL 54770, BPD
321
[Ill]; Las Salas, 5 May 1995, J.R.Herndndez
95-021 (BPI 841080) [II]
on Cucurbitella cucumifolia (Griseb.) Cogn."
Tucuman: San Javier, Parque Sierras de
San Javier, 23 Jul 1994, ].R. Hernandez 94-
148 (LIL 54771, BPI 841069) [III].
on undtn.; Tacuman: Horco Molle, Parque Si-
erras de San Javier, 6 Apr 1994, J. FHennen,
M.M.Hennen & J.R.Herndndez 94-094
(LIL 54772, BP] 841037) [II-HI].
CYPERACEAE
Puccinia cyperi Arthur
on Cyperus sp., Tucuman: San Pedro de
Colalao, 2 Apr 1994, J.RHennen &
M.M.Hennen 94-075 (LIL 54773, BPI
841035) [LI-IIL].
DRYOPTERIDACEAE
Desmella anemiae Syd. & P. Syd.4
lilloi CHicken) Alston’;
Tucuman: Las Lenguas, 22 Oct 1950,
R.Singer s.n. [I]; Las Lenguas, 22 Oct 1950,
R.Singer s.n. [I].
on Athyrium
ERYTHROXYLACEAE
Phakopsora coca Buritica & J.-F Hennen
on Erythroxylum coca Lam. Jujuy: 2 Apr 1993,
J.FHennen & L.DPloper 93-115 (LIL 54774,
BPI) [Il]. Salta: Cafayate, 10 Apr 1994,
J FHennen, M.M.Hennen &J.R. Hernandez
94-135A (LIL54775, BP] 841061) [IIL
EUPHORBIACEAE
Uromyces cisneroanus Speg.“!
haematospern > Mal.Arg.;
Catamarca, 23 Sep
0: 3 3
on Sapium
Catamarca: W side of
97, J. FHenne n GJ.R.H Hernandez 97
ae II]. Tucuman: Dept. Capi-
tal, 14 May 1995, |. R.Herndndez 95-051
(BPI 841092) [II-II; El Manantial, 5 Mar
1996, J.R.Herna hee 96-076 (BPI ae
[II-III]; Los Gutierrez, 15 Aug 1
J.R.Herndndez ae «roth San e
de Colalao, 1 Apr 1994, J.F.
M.M.Hennen 94-062 fn Beis an {Il-
Il]; San Pedro de Colalao, 31 Mar 1994,
J.EHennen & ea ee (LIL
54778, BPI) [IIIl.
Uromyces euphorbiae Cooke & Peck
on Euphorbia hirta L; Tucuman: Dept. Cruz
Alta, San Agustin, Monte Redondo, 29
322
Mar 1994, |. FHennen, M.M.Hennen &
J.R.Herndndez 94-028 (LIL 54776, BPD [I-
II, San Pedro de Colalao, 31 Mar 1994,
J.RHennen & M.M.Hennen 94-046 (LII
54777, BP] 841024) [II].
FABACEAE
Phakopsora meibomiae (Arthur) Arthur’
on Cologar lifolia Kunth®; Tueuman: 22
km W of San Miguel de Tucuman, mtn,
27 Mar 1993, |. EF Hennen, L.D.P| & LR.
Herndndez 93-018 (LIL a: . BPI
840995) [II].
Phragmopyxis deglubens (Ber
k. & M.A. Curtis)
Yietel in Engl. & Prantl’
on Coursetia brachyrhachis Harms”, Jujuy:
Pampa Blanca, | Apr 1993, J.E Hennen
& L.D.Ploper 93-097 (LIL 54820, BPI
841012) [I-III
Puccinia bergii Speg.’
on Potretia tetraphylla (Prior) Burkart’:
Corrientes: Dept. Concepcion, Rincon de
Luna, Fstancia Santillana, 26 Febl957,
TM. Pedersen 4483 [IIL
Ravenelia argentinensis Speg“~
on Acacia droma Gillies in Hook. & Arn.
Catamarca: N of Catamarca, Hwy. 38, 24
Sep 1997, ].FHennen & |.R.Herndndez 97-
052 (BPL 841193) [I]. Jujuy: Parque Nacio-
nal Calilegua, 8 Decl1997, |. R.Herndndez
97-179 (BP1841267) [I]. Salta: Metan, 2 Apr
1993). Hennen & L.D.Ploper 93-103 (LIL
54823, BPI 841014) [1-I-HI: road to
Parque del Rey, 6 Dec 1997, ].R.-Herndndez
97-160 (BP1 841256) [I]. Tucuman: Cevil
30 Mar 1993, |. F.Hennen & LR
Herndndez 93-051 (LIL 54824, BPD [II
Cruz Alta, San Agustin, 30 Mar 1993,
J.E:Hennen & L.R.Herndndez 93-059 (LIL
54825, BPD [I-TH; Cruz Alta, San Agustin,
30 Mar 1993, J EHennen & |. R.Herndndez
93-060A (LIL54845, BPD
30 Mar 1993, |. F}Hennen & |. R-Herndndez
93-076 (LIL 54821, BPD[L-H} Las Cejas, 30
Mar 1993, . FHennen & |. R.Herndndez
93-072 (LIL 54848, BPD [I-Ik Las Cejas, 30
Mar 1993, J. hHennen & |. R.Herndndez
93-078 (LIL 54822, BPD [E-IFIII| Las Cejas,
30 Mar 1993, |. FHennen & |.R.Herndndez
93-065 (LIL 54826, BPD [II-III Las Cejas,
30 Mar 1993.) FHennen& |. R.Herndndez
Pozo,
(HI; Las Cejas.
BRIT.ORG/SIDA 20(1)
93-069 (LIL 54847, BPD I-I-IIf Las Cejas,
30 Mar 1993, |. FHennen& |.R.Herndndez
93-074 (LIL 54827, BPD [II-III Raco, 3
Nov 1997, |. R.Herndndez 97-143 (BPD [Lk
Raco, 3 Nov 1997, |. R.Herndndez 97-142
(BPI841241 [I] San Javier, ae Sierras
de San Javier, 27 Mar 1993, |. FHennen,
L.D.Ploper ea 93-032 (LIL
54844, BPD [I-II-HI]
Ravenelia cebil Speg.!
on Anadenanthera sp“: Jujuy: Caimancito, 2
Apr 1993, |. KHennen & L.D.Ploper 93-111
(LIL ae BPI) [II-Il; Pampa Blanca, 1
J-Hennen & L.D.Ploper 93-100
(LIL 54820, BPI) [I-H}, San Pedro, 2 Apr
1993, J. -}Hennen & L.D.Ploper 93-107 (LIL
54830, BPD) [I-I]. Tucuman: San Pedro de
Colalao, | Apr 1994, JI
1.M.Hennen 94-059 (LIL
841029) [I-HI].
Ravenelia cohniana Henn.
“Hennen &
54828, BPI
on Legum. undetn; Jujuy: Chalican, 2 Apr
1993, .}Hennen & L.D.Ploper 93-105 (LIL
54832, BPL841015) [I- HI,
Ravenelia echinata ica & Dietel var. ectypa
(Arthur & Holw.)
on Calliandra formosa Kom Benth.: Salta:
Pichanal, 2 Apr 1993, |. FHennen &
L.D.Ploper 93- Cd 54835, BPL) [M-UT],
Tucuman: San Pe e Colalao, | Apr 1994,
J-FHennen & M. M. Hennen 94-054 (LIL
548306, BPD [II-TI; San Pedro de Colalao, 2
Apr 1904, |. FHennen & M.M. Hennen 94-
073 (LIL 54837, BI 1841 O34) (I-III).
Ravenclia hieronymi Speg.
)Cummin
on Acacia caven (Molina) Molina: Catamarca:
Rio Las Juntas,c amping area, 23 Sep 1997,
J E-Hennen & |. R.Herndndez 97 en
[An W of Catamarca, 23 Sep 1997,
J. FHennen & |. R. Hernandez 97 044 (BPI
841188) [I]. Tucuman: J.B. /
Iberdi, S
Escaba Dike, 28 Nov 1997, ].R. sane
97-108 (BPI 841224) [An-lIk SW of J.B.
Alberdi, 22 Sep 1997, J FHennen &
J.R.Herndndez 97-007 (BPI 841166) [An-
HI; SW of J.B. Alberdi, 22 Sep 1997,
j.EPHennen & [.R.Herndndez 97-006 (BPI
841165) [An-lll].
Ravenelia lindquistii ].F. Hennen @ Cummins‘
on Acacia praecox Griseb. Catamarea: bank
HERNANDEZ AND HENNEN, RUST FUNGI OF NORTHWEST ARGENTINA 323
of Rio Las Juntas 23 Sep 1997, J. FHennen J.R.Her ndndez 93-073 (LIL 54840, BPI)
& J.R.Herndndez 97-041 (BPI 841187) [II- (11-1111.
II]; Rio Las Juntas, camping area,23Sep —_ Ravenelia papillosa Speg.‘
1997, J. FHennen & J.R.Herndndez 97-039 on Acacia visco Lorentz ex Griseb.; Catamarca:
i 841185) [II- III]. Salta: Pichanal, 2 Apr Dept. Ambatos, El Rodeo, El Nogalal, 14
93, J-FHennen & L.D.Ploper 93-109 (LIL Nov 1999, J.R.Herndndez 99-009 (BPI
are BPI) [II-III]; Pichanal, 2 Apr 1993, ere Dept Ambatos. FI Rodeo, El
jJ.FHennen & L.D.Ploper 93-108 (LIL lal, 14 Nov 1999, J.R. Hernandez 99-
54850, BPD [II]. Tucuman: San Pedro de ce Tucuman: |_as Salas
Colalao, 2 Apr 1994, J-EHennen & M.M. 5 May 1995, J.R.Herndndez 95-019(B PDI
Hennen 94-067 (LIL 54833, BPD) [II-III]. Ill; Dept. Trancas, Vipos, 28 Mar 1996,
Ravenelia lonchocarpicola var. lonchocarpicola J] R.-Herndndez 96-007 (BPI 841119) [II-III].
Speg.! on pi nia excelsa (Griseb.) Burkart’
on Lonchocarpus lilloi (Hassl.) Burkart’: atamarca: Dept. Ambatos, El Rodeo, El
Jujuy: Ledesma, between Valle Grande cane = Sep 1997, |.FHennen & JR.
and Rio Agua Negra, 8 Apr 1971, Legname Herndndez 97-049 (BPI 841191) [An-Illl.
& Cuezzo 7970c [0-11]; Ledesma, from Rio Ravenelia platensis Speg.“
Agua Negra to Alba de Canas, 29 May on Erythrina crista-galli L.; Buenos Aires: La
1977, Legname & Cuezzo 8247c [Ill Plata, 29 Sep 1997, J.RHennen & J.R.
Ravenelia macrocarpa Syd. & P. Syd.“ Herndndez 97-073 (BPI 841204) [0],
on Senna birostris var. hookeriana(Gilliesex Ravenelia sp.
Hook. & Arn.) HS. Irwin & Barneby*; on undtn.; Tacuman: Cevil Pozo, 12-15 km E
Tucuman: Dept. Trancas, Cuesta Grande, of Sat ~~ de Tucuman, 30 Mar 1993,
22 Apr 1926, S.Venturi s.n. [III]; Dept. J.FHennen & J.R.Herndndez 93-052 (LIL
Trancas, Cuesta Grande, 22 Apr 1926, 54849, “ 841002) [II-II.
_ se s.n. [III]; Tafi del Valle, 6 Apr Ravenelia spegazziniana J.C. Lindq.”'
93, |. EHennen & J.R.Herndndez 93-117 on Acacia aroma Gillies ex Hook. & Arn.
7: 54838, BPD [I-II]. Salta: Metan, 2 Apr 1993, J.EHennen ©
on Senna hookeriana Batke*; Tucuman: Tali, L.D.Ploper 93-103A (LIL54823, BPI) [III].
19 Mar 1944, A.Courteig 516[LI-HUl. Tucuman: Cruz Alta, San Agustin, 30 Mar
on Senna subulata (Griseb.) H.S. Irwin & 1993, | FHennen & J.R.Herndndez 93-
Barneby*; Catamarca: E of La Vina, 24 Sep O59A (LIL54825, BPI) = Cruz Alta, San
1997, J. FHennen ai Herndndez 97-056 Agustin, 30 Mar 1993, J.F.Hennen &
ee sega: [II]. Tucuman: Dept. Tali, 7 J.R.Herndndez 93- 060(LIL 54845, BPI) [H-
vate S. Gane $.N. a San Javier, Il}; Cruz Alta, San Agustin, 30 Mar 1994,
P s de San Javier, 23 Jul 1994, J). FHennen,M.M.Hennen © J.R. Hernandez
Terenas: ae BPD[H]. 94-038 ce 54846, BPI) [III]; Dept.
Ravenelia mimosace-sensitivae He Trancas, Vipos, 30 Mar 1995, J.R.Herndndez
on Mimosa debilis Humb. ae ex Willd. 95-044 (BPI 841087) [III]; Las Cejas, 30
var. debilis®, Salta: Between Rosario de la Mar 1993, |. FHennen & J.R.Herndndez
Frontera and Horcones, ll Apr 1994, 93-076A (LIL54821, BPI 841005) [II];
J.FHennen & L.D.Ploper 94-120 (LIL Monte Redondo, 31 Mar 1996, J.R.
54843, BPI 841052) [II-LI[]. Tuacuman: Cruz Herndndez 96-089 (BPI) [II-III].
Alta, San Agustin, 30 Mar 1994, J.F. | Uromyces appendiculatus (Pers.) Unger
Hennen, M.M.Hennen & J.R.Herndndez on Phaseolus vulgaris L.; Salta: Between
94-035(LIL 54842, BPD [I-I], Dept. Cruz Rosario de la Frontera and Horcones, 11
Alta, San cone Monte Redondo, 29 Apr 1994, J.FHennen & L.D.Ploper 94- "121
Mar 1994, J.RHennen, M.M.Hennen & (LIL 54852, BPD [II]; Cafayate, 9 Apr 1994
J.R.Herndndez peptic 54841, BPD [II- J.FHennen, M.M.Hennen & J.R.Herndndez
Ill]; Las Cejas, 30 Mar 1993, |. FHennen & 94-107 (LIL 54851, BPD [II-HI]. Tucuman:
Dept. Trancas, 17 Mar 1995, ,.R. Herndndez
95-070 (BPI 841102) [I]; Las Criollas, 2
Nov 1986, L.D.Ploper & G.Wurschmidt
sn. (BPD [H-HH]
P. Syd. & Syd"
on Desmodium purpureum Hook, & Arn.
Uromyces castancus |
Salta: Dept. Santa Victoria, Los Toldos, F]
Nogalal, 29 Jun 1996, J.R.Herndndez 96-
041 (BPI 841136) [H-1].
Uromyces dietelianus Pazsche*
on Bauhinia oni Link*; Buenos Aires:
castelar, r 1996, |. R-Herndndez 96-
064 (BPI a {11-111]. Tucuman: F
Corte, 15 Nov 1997, |. R. Hernandez 97-085
(BPI 841211) {I-HIl.
Uromyces hedysari-paniculati (Schwein) Farl
apud Ellis
on Desmodium sp. Tucuman: NW of San
Miguel de Tucuman, 31 Mar 1993, J.-F.
Hennen & J.R.Herndndez 93-092 (LII
54860, SEO EIEN NW of San Miguel de
Tucuman, 31 Mar 1993, J.RHennen &
J.R-Herndndez 93-091 (LIL 54855, BPI
841011) [H-If; San Javier, 27 Mar 1993,
J.-FHennen, L.D.Ploper & J.R.Herndndez
93-022 (LIL 54856, BPD) [I-II-HI].
Uromyces orbicularis Dietel*
on Desmodium purpureum Hook. & Arn.*;
Tucuman: San Javier, Parque Sierras de
San Javier, 27 Mar 1993, |. FHennen, LD.
Ploper & J.R.Herndndez 93-034 (LI
BPI) [II-Ill]; San Miguel de
Tucuman, 23 Jun 1995, .R.Herndandez 95
082 (BPI 841108) [I-HI]
on Desmodium sp. Tucuman: NW ol San
Miguel de Tucuman, 31 Mar 1993, [LF
Hennen & J|.R.Herndndez 93-091A
(11154855, BPI) [I-III]; San Pedro de
Colalao, 31 Mar 1994, }FHennen & MM.
Hennen 94-048 (LIL 54857, BPD [I-UIT.
Malme
San Javier, Parque Sierras de
San Javier, 23 Jul 1994, ].R.Herndndez 94
159 (LIL 54854, BP) (H-IUL.
Uromyces striatus Schrot
ay
on Desmodium subsericeum
edicago sdliva L; Salta: Calayate, 9 Apr
1994, J RHennen, M.M.Hennen & JR.
ez 94-108 (LIL 54859, BPD) [I]
Tucuman: Dept. Trancas, Vipos, 28 Mar
1996, ]. R. Hernandez %6
Hernadne
=\ ft)
UUZVDELOTILIY J {LY
BRIT.ORG/SIDA 20(1)
Uromyces trifolii-repentis Liro!
on Trifolium repens L.; Tweuman: Cruz Alta,
15 Mar 1996, |.R.Herndndez 96-092 (BPI
841158) [Il]; NW of San Miguel de
Tucuman, 31 Mar 1993, J FHennen & JR.
Herndndez 93-093 (LIL 54858, BPD [II].
Uromyces viciae-fabae (Pers.) Schrot.!
n: Tafi Viejo, 9 Nov
on Vicia faba L., Tucu
07 OBE 841199)
19907, | R.Herndndez
[-HI]
Ypsilospora sp.*
on Inga edulis Mart.“*; Tucuman: Dept. Capi-
tal, Quinta Agronomica, 28 Mar 1994,
J-RHennen & M.M.Hennen 94-024 (LIL
54817, BPD [I]; Dept. Capital, Quinta
Agronomica, 30 Mar 1993, |. FHennen &
J.R.Herndndez 93-049 (LIL 54816, BPI
841001) [Ik Capital, Quinta
Agronomica, 6 Apr 1994, ].F.Hennen,
M.M.Hennen & J.R.Herndndez 94-096
(LIL 54818, BP] 841038) [I-1H].
Dept.
GERANIACEAE
Puccinia pelargonii-zonalis Doidge*
n Pelargonium hortorum L.H. Bailey *;
Tucuman: Dept. Capital, 29 Jul 1994,
|. R-Herndndez 94-146 (LIL 54781, BPD III;
Dept. Capital, Quinta Agronomica, 28
Mar 1904, |. }Hennen & M.M.Hennen 94-
023 (LIL 54780, BPD [I Dept. Capital,
Quinta Agronomica, 15 Apr 1995, [.R.
Hernandez 95-075 (BPI) [1-H San
Miguel de Tucuman, 15 Apr 199], a
Herndndez 91-001 (BPI 84.0990) [II Tat
iejo, 15 Aug 1994, [.Zarsoza s.n. (LIL
54782, BPD {Ik Yerba Buena, 19 Jul 1996,
J. R.Herndndez 96-072 (BPI 841148) [I].
US
IRIDACEAE
Uromyces transversalis (Thtim.) G. Winter
on Gladiolus sp.’; San Luis: Villa Mercedes,
1995, leg. ? (BPI 841104) [1-H].
Tucuman: Dept. Capital, 27 Jun 1996,
JR Hernandez 96-028 (BPI 841131) [If
Dept. Capital, lo Mar 1995, ].R.Herndndez
95-069 (BPL 841101) {H-1U].
22 Apr
LAMIACEAE
Puccinia farinacea |_ong!
on Salvia rypara Brig; Tucuman: San Pedro
de Colalao, 2 Apr 1994, J. hHennen &
M.M.Hennen 94-068 (LIL 54807, BPD [II-
HERNANDEZ AND HENNEN, RUST FUNGI OF NORTHWEST ARGENTINA
III]; San Pedro de Colalao, | Apr 1994, J.E
Hennen & M.M.Hennen 94-063 (LIL
54805, BPD [II-II|; San Pedro de Colalao,
Apr 1994, J.FHennen & M.M.Hennen 94-
074 (LIL 54806, BPD [II-III].
on Salvia sp. wk Palma Sola, 12 Apr 1994,
jJ.FHennen & L.D.Ploper 94-134 (LIL
54808, BPI ies [I-11].
Puccinia giberti Speg.
on Hyptis sp. Tucuman: Tafi del Valle, 6 Apr
1993, |. FHennen & |.R.Herndndez 93-125
(LIL 54760, BPI 841020) [II-III].
Puccinia hyptidis-mutabilis Mayor
on Hyptis mutabilis (Rich.) Brig.*"; Salta:
Cafayate, 8 Apr 1994, J.FHennen, M.M.
Hennen & J.R.Herndndez 94-102 (LIL
54810, BPD [I-X], Cafayate, 9 Apr 1994,
J.-FHennen,M.M.Hennen@].R. Hernandez
94-113 (LIL 54811, BPI) [II-X|. Tucuman:
Cruz Alta, San oe 30 Mar 1994,
J.-FHennen, M.M.Hennen& ].R.Herndndez
94-037 (LIL ey BPI) [II-X], El
Manantial, 15 May 1996, J.R.Herndndez
96-093 (BPI 841159) [H-X]; San pas
Parque Sierras de San Javier, La Cascada.,
29 Mar 1996, J.R.Herndndez 96- ae BPI
eel Xl.
on Hyptissp.**; Tucuman: Alta Gracia, 19 Nov
R.Herndndez 97-088 (BPI) [II-X];
Las Cejas, 30 Mar 1993, J.-Hennen &
J.R.Herndndez 93-077 (LIL 54813, BPI) [I-
X]; Las Cejas, 30 Mar 1993, J. Hennen &
J.R.Herndndez 93-066 (LIL 54812, BPD [H-
X-HI; NW of San Miguel de Tucuman, 31
Mar 1993, |. FHennen & J.R.Herndndez
93-085 (LIL 54814, BPD [II-X].
e Pers.!
Puccinia mentha
on Mentha pulegium L.: Tucuman: Valle de
Las Salas, bank of Rio Las Salas, 3 Nov 1997,
J.R.Herndndez sence ee 247) (U].
saan sp., Tucuma Capital, 15 Sep
995, J.R. Hernandez 95- ee HII].
Puccinia ae dissima Speg. |
on Stachys are Benth»; Tacuman: Dept.
Capital, Ouinta Agron omica. 15 Apr 1995.
J.-R.Herndndez 95-074 (BPI 841105) [III].
on Stachys sp., Tucuman: Yerba Buena, 29 Mar
1993, J.R.Herndndez 93-044 (LIL 54815,
BPD [II].
LILIACEAE
Puccinia allii F Rudolfi!
on Allium sativum L., Tucuman: Leales, 15
Aug 1992, L.D.Ploper & N.V.de Ramallo
s.n. (BPI 840992) [I-III].
MALPIGHIACEAE
Puccinia heteropteridis T htm.
on Janusia guaranitica (H. St-Hil.) A. Juss.
oad to El Rodeo, camping
area, 29 Nov 1997, ].R.Herndndez 97-116
(BPD [II]; W side of Catamarca, 23 Sep
1997, |. FHennen & J.R.Herndndez 97-034
(BP] 841183) [H-IU]. Tucuman: San Pedro
de Colalao, 1 Apr 1994, J. FHennen &
M.M.Hennen 94-056 (LIL 54861, BPI
841027) [II
I
MALVACEAE
Catenulopsora praelonga (Speg.) Buritica’
on Pavonia malvacea (Vell.) Krapov. &
Cristoval®; Tucuman: San Javier, Parque
Sierras de San Javier, 23 Jul 1994,
).R.Herndndez 94-143 (LIL 54863, BPD [II
San Javier, Parque Sierras de San Javier, 27
Mar 1993, J.FHennen, L.D.Ploper &
J.R.Herndndez 93-025 (LIL 54862, BPI
840997) [II-IL].
Puccinia heterospora Berk. & M.A. Curtis!
on Abutilon sp. Tucuman: 19.5 km W of San
Miguel de Tucuman, mtn, 27 Mar 1993,
J.RHennen, L.D.Ploper & J.R.Herndndez
93-007 (LIL 54864, BPI) [III
on Anoda cristata (L.) Schltdl; Jujuy: Capi-
tal, 15 Apr 1994, . R.Herndndez 94-167
(BPD [II]. Tucuman: San Javier, Parque Si-
erras de San Javier, 23 Jul 1994,
|. R.Herndndez 94-161 (LIL 54867, BPI)
{HW} San Miguel de Tucuman, 23 Jun 1995
2 95-08] oo pas San ee
de Colalao, 2 Apr 199
M.M.Hennen 94-076 (LIL een aes
San Pedro de Colalao, 31 Mar 1994,
Jw Hennen 94-042 (LIL
=
J.R.Herndnde
FHennen & M.A
54865, aus
1 A. St.-Hil: Tucuman:
Cruz aie San Agustin, 30 Mar 1994,
J. EHennen,M.M.Hennen &|.R.Herndndez
94-036 (LIL 54868, BPD [III Cruz Alta,
San Agustin, 30 Mar 1994, J.FHennen,
M.M.Hennen & J.R.Herndndez 94-040
(LIL 54869, BPD IIIT; Dept. Trancas, Vipos,
28 Mar 1996, ].R.Herndndez 96-003 (BPD)
[IU]
on Gaya sp., Tucuman: Las Cejas, 30 Mar 1993,
J. FHennen & |. R. Herndndez 93-068 (LI
54871, BPD [HI].
on si av. Tucuman: J.B. Alberdi,
scaba Dike, 22 Sep 1997, J.FHennen
- : eae 3PT 841167) [HII
on Sida spinosa L.; Tacuman: Las Cejas, 30
Mar 1993, |. FHennen & J.R.Herndndez
93-064 (LIL 54870, BPI 841004) [III
Puccinia malvacearum Bertero ex Mont.
on Malvastrum coromandelianum (L.)
Garcke; Salta: Cafayate, 8 Apr 1994, J.F
Hennen, M.M.Hennen & J.R.Herndndez
94-104 (LIL 54873, BPD [II]. Tucuman:
Dept. Trancas, Vipos, 28 Mar 1996,
|. R-Herndndez 96-008 (BPI 841120) [III
Dept. Trancas, Vipos, 30 Mar 1995, J.R
Herndndez 95-046 (BPD [II i. aes
road to Escaba, 28 Nov YR.
Herndndez 97-094 (BPI oe : San
Javier, Parque Sierras de San Javier, 23 Jul
1994, |.R.Herndndez 94-152 (LIL 54874,
BPD [III]; San Javier, Parque Sierras de San
Javier, La Cascada., 29 Mar 1996, [_R.
Herndndez 96-019 (BPD [IN| San Pedro de
Colalao, 31 Mar 1994, ]|.FHennen &
1.M.Hennen 94-041 (LIL 54872, iene
on Malvastrum sp. Tucuman: NW of San
Miguel de Tucuman, 31 Mar 1993, J.P
Hennen & J.R.Herndndez 93-082 (LIL
54875, BPD [III].
on undtin., Tacuman: Horco Molle, 20 Apr
1992, | R Herndndez 92-007 (BPI) [III|;
Horco Molle, 17 Sep 1997, J.R.- Hernandez
97-080 (BPI 841208) [III].
Puccinia platyspora (Speg.) HS. Jacks. @ Holw
)
Q
=
aa
Alcea rosea L.*; Jujuy: Tilcara, 3 Feb 1996,
J. R-Herndndez 96-001 (BPI 841116) [HI].
on Alcea sp.*; Tucuman: San Miguel de
Tucuman, 15 Apr 1991, ].R-Herndndez 9
003 (BPD [I].
on Modiolastrum malvifolium (Griseb.) kK
Schmann; Catamarca: Balcozna, 22 Sep
1997, |. FF} Hennen & |. R Hernandez 97-021
(BPI) [HE near Balcozna, 28 Nov 1997,
J.R.Herndndez 97-114 (BPD [III]; W side ol
Catamarca, 23 Sep 1997, |.;Hennen &
BRIT.ORG/SIDA 20(1)
J.R.Herndndez 97-025 (BPD [III]. Salta:
Posta de Yatasto,6 Dec 1997, ].R. Herndndez
97-156 (BPI 841252) [HI]. Tucuman: Cruz
Alta, 15 Mar 1996, [.R.Herndndez 96-090
(BPD [U1]; El Siambén, river next to mon-
astery, 3 Nov 1997, ].R.Herndndez sok
any
(BPI 841242) [IH Horco Molle, 15 \
1995, |. R.Herndndez 95-080 (BPI 841 i
[IIT]; Lules, 16 Nov 1996, |.R.Herndndez 96-
065 (BPD {III} San Pedro de Colalao, 31 Mar
1994, J. }Hennen & M.M.Hennen 94-043
(LIL 54876, BPD [IU].
on Modiolastrum sp. Tucuman: NW of San
Miguel de Tucuman, 31 Mar 1993, [.F
Hennen & |. R.Herndndez 93-089 (LIL
54879, BPD [IIIf San Javier, Parque Sierras
de San Javier, 27 Mar 1993, ]. FHennen, LD.
Ploper & J. R.Herndndez 93-024 (LI
54877, BPD III; Yerba Buena, 29 Mar 1993,
|. R.Herndndez 93-045 (LIL 54878, BPD
in I
ral atamarea: Dept. Ambatos
E | Rodeo, 14 Nov 1999, ].R. PernaindexOo-
006 (BPI 841275) [II]; Dept. Capital, 14
Nov 1999, |. R.Herndndez 99-002 (BPI
841271) [Il E of La Vina, 24 Sep 1997,
| FHennen & |. R-Herndndez 97-062 (BPD
[III]; Rio Las Juntas, 23 Sep 1997, J.F.
Hennen & |.R.Herndndez 97-050 (BPD
[III]. Salta: Cafayate, 8 Apr 1994, ].F
Hennen, M.M.Hennen & |.R.Herndndez
94-103 (LIL 54880, BPI 841043) [III].
Tucuman: Dept. Trancas, Vipos, 28 Mar
1996, |.R. Herndndez 96-004 (BPI) [III].
on a jorgensenii (.M. Johnst.) Krapov-;
Calayate, 9 Apr 1994, J. KHennen,
ae e nnen & [.R.Herndndez 94-114
(LIL 54881, BPD [II]
on pe : San Luis: Villa Mercedes, 15 Sep
92, leg. ? (BPI [IH].
a
MORACEAE
Phakopsora nishidana S. Ito"
on Ficus carica L.; Salta: Dept. Santa Victo-
ria, La Mision, 30 Jun 1996, | R. Hernandez
96-051 (BPI 841139) [I], Pichanal, 2 Apr
1993, J. }Hennen & L.D.Ploper 93-101(LIL
54883, BPD [I]. Tucuman: Dept. Capital,
30 Mar 1993, |. R.Herndndez 93-048 (LIL
54882, BPD [II]; San Miguel de Tucuman,
12 May 1991, [.R.Herndndez 91-004 (BPD
HERNANDEZ AND HENNEN, RUST FUNGI OF NORTHWEST ARGENTINA
[Il]; San Pedro de Colalao, 2 Apr 1994,
jJ.RHennen & M.M.Hennen 94-077 (LIL
54884, BPD [II].
MYRTACEAE
Puccinia psidii G. Winter
on Psidium guajava L., Tucuman: El Corte, 15
Feb 1995, J.R.Herndndez 95-065 (BPI) [III
El Corte, 15 Feb 1995, [R.Herndndez 95-
066 (BPI 841098) [I].
NYCTAGINACEAE
Puccinia ee ss oe :
1]
tamarca:
El Rodeo, 29 Nov 1997 J.R.Herndndez 97-
127 (BPI 841232) [I]. Jujuy: San Pedro, 2
Apr 1993, J. FHennen & L.D.Ploper 93-106
(LIL 54885, BPI) [0-I-II-II]. tucuman: San
Javier, Parque Sierras de San Javier, 23 Jul
1994, |.R.Herndndez 94-164 (LIL 54887,
es (1-II-II; San Pedro de Colalao, 16 Jul
2, J.R.Herndndez & A.Wiirschmid
fe 008 (BPI) [I; San Pedro de Colalao, |
Apr 1994, J. FHennen & M.M.Hennen 94-
050 (LIL 54886, BPI 841025) [I-II-III].
ONAGRACEAE
Endophyllum sp.*
on Fuchsia boliviana Carriére; Tucuman: F]
Corte, 14 Nov 1997, |. R.Herndndez 97-129
(BPI 841233) [III]; Lules, 27 Oct 1996,
JR. ieee 96-069 (BPI 841147) [IIIf
t. 367, 13 Sep 1986, A.Ganem s.n. (BPI)
‘ Tafidel vie 13 Sep 1986, C.Boero s.n.
(BPD [III
Puccinia jussiaeae Speg.'
on Ludwigia sp., Tucuman: NW of San Miguel
de Tucuman, 31 Mar 1993, . FHennen &
J.R.Herndndez 93-086 (LIL 54710, BPI
841008) [II-III].
Pucciniastrum epilobii (Chaillet) G.H. Otth
on Epilobium brasiliese Hausskn.; Salta:
rancas, Cuesta del Arca, Jan 1897,
leg.? (1-111)
OXALIDACEAE
Puccinia oxalidis Dietel & Ellis
on Oxalis sp. Jujuy: San Salvador, 12 Apr
. nnen® L ince ioe 7 IL
ie BP eee Tucuman: Dept. Trancas,
Vipos, 28 Mar 1996, ].R.Hernd a 96-005
(BPI) [II]; El Corte, 20 Feb 1995, .R.
Herndndez 95-063 (BPI 841096) [II San
Javier, 27 Mar 1993, J.EHennen, L.D.Ploper
& J.R.Herndndez 93-021A (LIL54888,
BPI) [II]; San Pedro de Colalao, 16 Jul 1992,
J.R.Herndndez & A.Wiirschmidt 92-009
(BPD [II].
PIPERACEAE
Puccinia peperomiae J.C. Lind
on Peperomia sp., Salta: camino de corniza
between Salta and Jujuy, 7 Dec 1997, J.R.
Herndndez 97-175 (BPI 841263) [II]; Dept.
Santa Victoria, Los Toldos, El Nogalal, 29
Jun 1996, J.R.Herndndez 96-049 (BPI
841138) [I]. Tucuman: E slope of Aconquija,
road to Tafi del Valle, 11 Nov 1952,
A.T.Hunziker s.n. [Il-If]|; Horco Mo
Parque Sierras de San Javier, 29 Mar 1996,
ede a -020 (BPI 841126) [III J.B.
Albe caba Dike, 28 Nov 1997
ey ee Sit Il].
le,
POACEAE
Physopella compressa (Mains) Cummins &
Ramachar*
on Paspalum sp.**; Tucuman: Yerba Buena, 24
Mar 1993, J. R.Herndndez 93-043 (LIL
54783, nae
Puccinia abnorm
on Echinoch a c eae Schult’;
Ita: Calayate, 9 Apr 1994, J. EHennen,
M.M.Hennen & J.R.Herndndez 94-117
(LIL 54784, sae dlaaige
on Echinoch a sp. : Amberes, 5
May 19 ae eee oe >.Wiirschmid
s.n. (BPI ){n- III-IV].
Puccinia aristidae [racy var. chaetaria Cumm.@
Husain*
on Aristida sp.**, Catamarca: bank of Rio Las
Juntas, 23 Sep 1997, J. EFHennen & JR.
Hernandez 97 ne (BPI 841186) [II-III].
Puccinia brach ii G.H. Otth var. poae-
nemoralis (G.H. ai Cummins & H.C. Greene
on Poa sp.; Tierra del Fuego: Est. Fique, 16 Feb
953, Ruiz Leal & Roig s.n. . PS ee
on Poa a ape rodi; Buen
talina, Oct
= ee $n. me Soot TL
Puccinia cacabata Arthur & Holw!
on Chloris gayana Kunth**; Salta: Las Lajitas,
12 Apr 1994, J. EHennen & L.D.Ploper 94-
128 (LIL 54785, BPD [II-III].
on Chloris polyd actyla (L.) Tucuman:
icas, Vipo ) Mar 1995, [.R.
Herndndez 95-04 een ace {1I-111.
Puccinia chaetochloae Arthur’
on Setaria lachnea (Nees) Kunth”: Salta: Las
Lajitas, 12 hei 1994, J. FHennen & LD.
Plope 94-129 (LIL 54802, BPI) [II].
on Setaria parviflora (Poir.) Kerguélen®;
Tucuman: Cruz Alta, San Agustin, 30 Mar
F-Hennen, MM.Hennen & JR.
Hernandex oe: 034 (LIL 54803, BPD [II].
side of
L994
a
on Setaria sp.**; Catamarca: W
Catamarca, 23 Sep 1997, J. FHennen &
J.R.Herndndez 97-026 (BPI 841177) [Il
Tucuman: Cruz Alta, 23 Jun 1995, |.R.
Herndndez 95-083 (BPD [IL Las Cejas, 30
Mar 1993, | FHennen & |.R-Herndndez
93-075 (LIL 54804, BPI) [II]; Monte
Redondo, 31 Mar 1996, |. R.- Hernandez 96-
O86 (BPI III.
Puccinia chloridis Speg.
n Chloris virgata Sw. Salta: Cafayate, 9 Apr
94. J.Hennen, M.M.Hennen & |.R.
Herndndez 94-118 (LIL 54786, BPI
841050) [LI].
Puccinia coronata Corda var. rangiferina (S
to)
Cummins*
on Avena sativa L.A“ i-
tal, Quinta Agronémica, 20 Oct 1994,
J.R.Herndndez noice il
Puccinia inane Lacroix ex Dest
onc ylon L; Salta: ae vyate, 9 Apr
1904, J. Hennen, M.M.Hennen & |.R
Herndndez 94-109 (LIL 54787, BPI
841044) [I-HI.
Puccinia inclita Arthur’
* Tucuman: Dept. Ca
aaa
a
on Ichnanthus minarum (Nees) Doéll*:
Tucuman: NW olf San Miguel de Tucuman,
31 Mar 1993, f FHennen & |.R. Hernandez
93-083 (LIL 54799, BPI841007)[I-II; San
Javier, Parque Sierras de San Javier, 23 Jul
1994, |. R.Herndndez 94-138A (LIL54801,
BPD [H-IH; San es Parque Sierras de
San Javier, La Cascada., 29 Mar 1996,
| R-Herndndez 96-015 (BPD [I-III Tafi del
Herndndez 93-126A (LIL54780, BPI)|III |
Puccinia leptochloae Arthur!
BRIT.ORG/SIDA 20(1)
on Leptochloa mucronata (Michaux) Kunth*;
Pa
Tucuman: Dept. Cruz Alta, San Agustin,
Monte Redondo, 29 Mar 1994, | FHennen,
M.M.Hennen & |. R-Herndndez 94-026
(LIL 54788, BP] 841021) {H-HI1.
Puccinia levis (Sacc.W Bizz.) Magnus var. panici-
sanguinalis (Rangel) Ramachar @ Cummins*
on Ichnanthus minarum (Nees) Doll;
Tucuman: San Javier, Parque Sierras de
San Javier, 23 Jul 1994, |.R. Hernandez 94-
138 (LIL 54801, BP] 841063) ties Tali del
Valle, 6 Apr 1993, J].F.He 1& ER.
Page 293-126(LIL op erie I],
on Panicum sp.“*; Catamarca: Rio Las Juntas,
a
amping area, 23 Sep 1997, |. hKHennen &
J.R-Herndndez 97-037 (BPD [I-UT; W side
of Catamarca, 23 Sep 1997, J. FHennen &
J. R.Herndndez 97-032 (BPI84118D [IE HL;
W side of Catamarca, 23 Sep 1997, JF:
Hennen & |. R.Herndndez 97-031 (BPI
841180) [II-HIL.
Puccinia melanocephala Syd. & P. Syd.
Qu
on Saccharum officinarum L. Jujuy: San
. 12 Apr 1994, J. RHennen & L.D.
24 (LIL ee BPD) [II-HIL.
Tucuman: Dept. Capit 3 Jun 1996,
JR. ae 2 96- a 53) (II-U|;:
Santa Lucia, 15 Mar 1988, L.D.Ploper &
G.Wiirschmidt s.n. (BPD) [I].
Ellis & Everh>!
on Digitaria insularis (L.) Fedde
Pampa Blanca, | Apr 1993, J.}Hennen &
L.D.Ploper 93-098 (LIL 54791, BPI) [I-HIL.
Tucuman: Dept. Cruz Alta, San Agustin,
Monte Redondo, 29 Mar 1994, |. FHennen,
M.M.Hennen & J|.R.Herndndez 94-029
(LIL 54792, BPD [I-II, Las Cejas, 30 Mar
1993, FHennen & [.R.Herndndez 93-071
(LIL 54790, BPD [Il; Monte Redondo, 31
Mar 1996, |.R. Hernandez 96-087 (BPD [I-
Il]; San Pedro de Colalao, 31 Mar 1994,
j-RHennen & M.M.Hennen 94-044 (LIL
54793, BPI) [IIL
on Digitaria sanguindlis(L.) Scop. Salta: be-
tween Rosario de la Frontera and
Horcones, Il Apr 1994, J. -Hennen &
L.D.Ploper 94-122 (LIL 54794, BPI
841053) [I-Hl].
zp!
Puccinia purpurea Cooke
Pedro
a 942]
Puccinia oahuensis
ie Jujuy:
on Sorghum cafrorum (Thunb.) P. Beauvy.*:
HERNANDEZ AND HENNEN, RUST FUNGI OF NORTHWEST ARGENTINA
Tucuman: Horco Molle, Parque Sierras de
San Javier, 29 Mar 1996, J.R.Herndndez
96-025 (BPD [II Monte Redondo, 23 Oct
1996, J.R.Herndndez 96-080 (BPD [II].
on Sorghum halepense (L.) Pers.; Tacuman
Alta Gracia, 19 Nov 1997, .R. Teenie
97-089 (BPI1841212) [II-III]; Cevil Pozo, 12-
15 km E of San Miguel de Tucuman, 30
Mar 1993, |.RHennen & |.R.Herndndez
93-054 (LIL 54795, BPD [I]; Dept. Trancas,
Vipos, 28 Mar 1990, J.R.Herndndez 96-010
(BPD [II-III]; San Pedro de Colalao, 31 Mar
1994, J EHennen & M.M.Hennen 94-049
(LIL 54796, BPD [II].
Puccinia recondita Roberge ex Desm.
on Bromus sp.; Tucuman: Tafi del Valle, 15 Jul
94, |.R.Herndndez 94-169 (BPD [Ik Tati
del Valle, 6 Apr 1993, J.FHennen & J.B.
Hernandez 93-119 (LIL 54797, BPI) [I-HIL.
on Bromus unioloides Kunth; Tucuman: El
Manantial, 8 Oct 1995, |. R.Herndndez 95-
041 (BPI 841084) [II]
on Triticum aestivum L.. Tucuman:
urruyacu, Loc. La Ramada, Finca Lo
Lapachos, 19 Sep 1999, |.R.Herndndez 56.
O11 (BPI841279) [II-III].
Puccinia sorghi Schwein.
on Zea mays L.; Salta: Between Rosario de la
Frontera and Horcones, ll Apr 1994, ].F
Hennen & L.D.Ploper 94-123 (LIL 54798,
BPI 841054) [HI]. Tucuman: Dept. Capital,
14 May 1995, J.R.Herndndez 95-050 (BPI
841091) [IIL]; Dept. Capital, 8 Mar 1988,
cea &G.Wiirsc es s.n. (BPD) [IT
El Colmenar, 14 Jun 1989, L.D.Ploper &
G. Mee aa 89-003 (BPD [II-III].
Puccinia s
on ae nthus minarum (Nees) Dall‘;
Tucuman: San Javier, Parque Sierras de
San Javier, 23 Jul 1994, |. R:Herndndez 94-
138B (LIL54801, BPD [H-HI1.
Puccinia substriata El]. & Barth.
on Digitaria sp, Tacuman: Dept. Capital, 10
Mar 1996, J.R.Herndndez 96-079 (BPI
841151) [TI-I1.
Uromyces setariae-italicae Yosh.!
on Setaria sp., Tucuman: San eee a
Sierras de San Javier, La Cascada.,
1996, J.R.Herndndez 96- ae (BPI ees
[11-111].
POLYGONACEAE
Cerotelium sp.*
on Ruprechtia apetala Wedd"; Tucuman: San
Pedro de Colalao, | Apr 1994, J.RHennen
& M.M.Hennen 94-051 (LIL 54890, BPI)
[II-III].
on Sli laxiflora Meisn.°; Tucuman:
Dept. Capital, 15 Aug 1994, .R. Hernandez
O4- 135 (LIL 54893, BPI 841060) [II].
on Ruprechtia sp.°; Tucuman: SW of J.B.
Alberdi, 22 Sep 1997, |. FHennen & J.R
Herndndez 97- ee I].
Puccinia polygoni-amphibii Pers.:Pers.‘
on Polygonum dcuminatum Kunth; Salta:
Cafayate, 9 Apr 1994, J.-F Hennen, M.M.
Hennen & J.R.Herndndez 94-116 (LIL
54892. BPD [II]. Tucuman: San Javier, 27
Mar 1993, J.FHennen, L.D.Ploper & J.R.
Herndndez 93-040 (LIL 54891, BPI) [II].
on Polygonum hydropiperoides Michx.:
Catamarca: FE] Rodeo, 29 Nov 1997, [.R.
Herndndez 97-123 (BPI 841229) [II] Salta:
Posta de Yatasto, 6 Dec 1997, ].R.Herndndez
7-157 (BPI 841253) [II]. Tucuman: road to
Villa Nougués, 5 May 1995, J.R. Herndndez
95-034 (BPI 841081) [II].
on Polygonum sp.; Catamarea: E of La Vina,
24 Sep 1997, J. FHennenG@J.R Herndndez
97-059 (BPI) [II W side of Catamarca, 23
Sep 1997, ].FHennen & J.R.Herndndez 97-
030 (BPI 841179) [II]. Tacuman: Road to
Balcozna, 22 Sep 1997, J.RHennen &
J.R.Herndndez 97-012 (BPD [I].
Uromyces crassipes Dietel & Neger'
a ees Manantial, Fac.
999, |. R.Herndndez
99-010 (BPI 841278) “i
PORTULA
Puccinia leptochloae Arthur & Fromme?!
on Talinum paniculatum Jack.) Gaertn. Jujuy:
Nacional Calilegua, 8 Dec
1997, |. R.Herndndez 97-177 (BPI aes
1]. Tucuman: Dept. Capital, 12 Mar 199
M.de la Vega s.n. (BPI 841249) [O-I].
PTERIDACEAE
Desmella anemiae Syd. & P. Syd.‘
on Pteris deflexa Link*; Tucuman: SW of J.B.
Alberdi, 22 Sep 1997, J.FHennen & J.R.
Herndndez 97-005 (BPI 841164) [II].
on Pteris sp.“"; Tucuman: rdi, S ol
J.B. Albe
Escaba Dike, 28 Nov 1997, |. R.Herndndez
97-110 (BPD [II].
RANUNCULACEAE
Bacodromus seman JR is t ae me FE Hennen
onkR at
Dept. Armibatos: El Rode
j.EHennen
841190) [III]; Dept. Ambatos, El Rodeo, 14
Nov 1999, |.R.Herndndez 99-008 (BPI
746516) [III]
Puccinia recondita Roberge ex Desm.
amarca:
0, 23 Sep 1997
yn Clematis
montevidensis Spreng.
odeo, 29 Nov 1997,
/ R.Herndndez > 124 (BPI) [0]. Tucuman:
ta Gracia, 19 Nov 1997, |. R-Herndndez
07-091 (BPI) [0-1 El Corte, 15 Feb 1995
|. R.Herndndez 95-068 (BPI 841100) [0-1
El Siambon, river next to monastery, 3
Nov 1997, ].R.Herndndez 97-148 (BPI) lO}:
|.B. Alberdi, road to Escaba, bank of creek,
28 Nov 1997, |. R-Herndndez 97-101 (BPI)
[0-1], NW of San Miguel de Tucuman, 31
Mar 1993, | FHennen & |.R.Herndndez
93-081 (LIL 54895, BPI) [O-I NW of San
Miguel de Tucuman, 31 Mar 1993, ].F.
Hennen & ].R.Herndndez 93-087 (LI
54894, BPI) [0-I].
on Clematis sericea Michx.; Tucuman: San
Catamarca
>
we
ne Parque Sierras de San Javier, La
ada., 29 Mar 1996, |. R.Herndndez 96-
016 (BPI 841123) [0-1].
RHAMNACEAE
Phakopsora zizyphi-vulgaris Dietel“
on Zizyphus mistol Griseb.; Tucuman: Las
Cejas, 30 Mar 1993, J. kK Hennen & J.R
Herndndez 93-079 (LIL 54896, BPI
841006) [II].
—
ROSACEAE
Frommeella mexicana var indicae |. W. McCain &
J.F Henn
on on o— indica (Andrews) Focke*; Bue-
s: Ensanada, date ?, G.A.Costa &
ee s.n. [I]. Salta: Dept. Santa Vic-
Los Toldos, 28 Jun 1996, JR.
nee >2 96-035 (BPI 841133) : 1-111].
uman: Dept. Tra ‘1 Potre
a 1996, J.R.He ie 06- nen ih
Dept. Trancas, Fl Potrero, 28 Oct 1996,
nos Air
toria,
& J.R.-Herndndez 97-046 (BPI
BRIT.ORG/SIDA 20(1)
~
].R.Herndndez 96-067 (BPI 841146) [III]:
W of San Miguel de Tucuman, 31 Mar
1993, |. FHennen&j.R.Herna ndez 93-095
(LIL 54898, BPI) [II]; Raco, 3 Nov 1997,
| R Hernandez 97-140 (BPI) [II Road to
Balcozna, 22 Sep 1997, J.RHennen &
TR. ae 97-014 (BPI 841170) [If
San Javier, 27 Mar 1993, |.AHennen, L.D.
Ploper & J.R.Herndndez 93-038 (LIL
54897, BPD [II San Javier, Parque Sierras
de San Javier, 23 Jul 1994, | R.Herndndez
94-157 (LIL 34899, BP] 841072) [II].
is (Speg.) J.
on Rubus boliviensis Focke*; Tucuman: Horco
Gerwasia imperial C. Linda.
Molle, Parque Sierras de San Javier, 6 Apr
1994, J. FHennen, M.M.Hennen & J.R.
Hernandez 94-085 (LIL 54902, BPD [I-IIII;
San Javier, Parque Sierras de San Javier, 27
Mar 1993, |. FHennen, L.D.Ploper & J.R.
Herndndez 93-030A (LIL54903, BPI
840998B) [0-1]; San Javier, Parque Sierras de
San Javier, 27 Mar 1993, J]. Hennen,
L.D.Ploper & J.R-Herndndez 93-026 (BPD
[O-I-ILl|; San Javier, Parque Sierras de San
Javier, 27 Mar 1993, J.FHennen, L.D.Ploper
& JR. Hernandez 93-037 (LIL 54901, BPD [I]
on Rubus imperialis Cham. & Schltdl. Salta:
camino de corniza between Salta and
Jujuy, 7 Dec 1997, ].R.-Herndndez 97-174
(BPD [0-I, Dept. Santa Victoria, Los Toldos,
El Nogalal, 29 Jun 1996, J.R. Hernandez
96-045 (BPD [O-I-H-II Dept. Santa Vic-
toria, Los Toldos, El Nogalal, 29 Jun 1996,
J.R.Herndndez 96-046 (BPI) [0-1-1].
Tucuman: F
Siambon, river next to mon-
astery, 3 Nov 1997, |. R.Herndndez 97-146
(BPI 841244) [O-I]; Horco Molle, 14 Nov
1997, |. R. Hernandez 97-138 (BPI) [O-I-II].
Kuchneola loeseneriana (Henn.) HS. Jacks. & Holw.
on Rubus boliviensis Focke*; Tucuman: San
Javier, Parque Sierras de San Javier, 27 Mar
1993, J.FHennen, L.D.Ploper & JR.
Herndndez 93-030 (LIL 54900, BPI
840998 A ) [O-I-H-III].
Phragmidium mucronatum (Pers.) Schlecht.
on Rosa sp.; Salta: Dept. Santa Victoria, Los
Toldos, 28 Jun 1996, J.R.Herndndez 96-
036 (BPI 841134) [I-11].
Phragmidium tuberculatum J. Miller!
on Rosd sp., Tucuman: Dept. Capital, 17 Dec
HERNANDEZ AND HENNEN, RUST FUNGI OF NORTHWEST ARGENTINA
1995, J.R.Herndndez 95-091 (BPI 841113)
(II).
Tranzschelia discolor (Fuckel) Tranzschel & Litv>
on Prunus armeniaca L.; Salta
Victoria, Los Toldo
He rndndez 96-029(BPDIII- II. Tucuman:
Raco, 23 Apr 1996, J.R.Herndndez 96-026
(BPI 841130) [II-II].
on Prunus domestica L.; Salta: Dept. Santa
Victoria, Los Toldos, 28 Jun 1996, |.R.
Herndndez 96-034 (BPI) [II]. Tucuman
Dept. Capital, 10 Apr 996, [.R. ndndez
96-083 (BPI 841154) [II San Javier, Parque
Sierras de San Javier, 23 Jul 1994, |.R.
Hernandez 94-147 (LIL 54904, BPD [II-III].
on Prunus persica (L.) Batsch; Salta: Dept.
Santa Victoria, La Misi6n, 30 Jun 1996,
J.R.Herndndez 96-060 (BPI) [II-III]; Dept.
Santa Victoria, Los Toldos, 28 Jun 1996
J.R.Her ps 96-030 (BPI) [II]. Tucuman:
El 5 Nov 1997, J.R.Herndndez 97-
083 ae 841209)|II} Horco Molle, Parque
94- Legare 54906, BPI )[II- III}; San Javier,
27 Mar 1 ‘Hennen, L.D.Ploper &
J.R.Her naniieg 9s: 021 (LIL 54905, BPI) [II]
=
RUBIACEAE
Uredo sp.*
on Psychotria carthagenensis Jacq.; Tacuman:
El Corte, 15 Nov 1997, J.R.Herndndez 97-
oe ae 841210) [II]; El Corte, 27 Mar
3, J.FHennen, L.D.Ploper & JR.
e 002 (LIL 54907, BPD [Lk
Horco Molle, 14 Nov 1997, .R.Herndndez
97-133 a e410 [1]; Horco Molle
Parque Sierras de San Javier, 29 Mar 1996,
ene 96-021 (BPI 841127) [II];
Horco Molle, Parque Sierras de San Javier,
6 Apr 1994, J. FHennen, M.M.Hennen&.R.
Hernandez 94-089 (LIL 54908, BPD) [II]
Uromyces holmbergii Speg.'
on Diodia aaa Cham. = _
ucuman: 95, JR.
Herndndez oa (BPI veces ) oa III};
San Pedro, 16 Mar 1996, O.Baino s.n. (BPI
841150) [I-III Tati del Valle, 6 Apr 1993,
J.EHennen & J.R.Herndndez 93-124 (LIL
ve ee IU.
SALICACEAE
Mcelampsora epitea (Kunze & J.C. Schmidt) Thtim.*
on Salix babylonica L.\*; Salta — 8
Apr 1994, J].RHennen, M.M.Hennen &
J.R.Herndndez 94-097 a ao. BPI
841039) [II]; Metan, 2 Apr 1993, |. FHennen
& L.D.Ploper 93-112 (LIL ee BPI) [II].
Tucuman: me Capital, 27 Mar 1993,
J.EHennen, L.D.Ploper & J.R.Herndndez
93-041A L910 BPI) - ce Capital,
4 Apr 19 FHennen & J.R. Herndnde
94-081 a eae BPI) ‘ San Pedro de
Colalao, 3 Apr 1994, J. FHennen & M.M.
Hennen 94-079 (LIL 54912, BPD) [II].
on Salix caprea L.4®; Tucuman: Dept. Capital,
17 Dec 1995, ].R.Herndndez 95-096 (BPI
841115) [I].
on Salix humboldtiana Willd.*®, Tucuman:
San Javier, 27 Mar 1993, | FHennen, i
Ploper & J.R.Herndndez 93-020 re
ie BPI) [11| San Pedro de Colalao, 2 Apr
994, J. FHennen & M.M.Hennen - 078
i 54916, BPD [II].
Melampsora larici-populina Kleb~
on ces nigra L; Salta: Cafayate, 8 Apr
4, J]. Hennen, M.M.Hennen & JR.
an ndez 94-100 (LIL 54917, BPI
841041) [II]
on Populus sp.; Santa Fe: 1941, J.R. Weir s.n. (II,
Melampsora medusae Thtim.
on Populus sp., Buenos Aires: La Plata, Dec
1923, J.R.Weir s.n. [II]; Buenos Aires, S.A.,
25 Dec 1923, J.R. Weir s.n. [II].
Melampsora populnea (Pers.) P. Karst.
on Populus alba L.; Buenos Aires: La Plata,
Dec 1923, |.R. Weir s.n. [II], Cordoba: Capi-
tal, 9 Nov 1997, ].R.Herndndez 97-069 (BPI
841202) [II Santa Fe, 1941, ESchiels.n.[H]
SAPINDACEAE
Puccinia arechavaletae Speg.'
on Serjania marginata Casar®; Tucuman: FE]
27 Mar J. FHennen,1 D.Ploper
R.Herndndez 93- 005 (LIL 54918, BPI
3) ([M-III].
& JR
ai
Skierka sp.*
on Cupania vernalis Cambess.*;
Dept. Trancas, Rt. 9, km 1364, 26 Oct 1995,
J.R.Herndndez 95-056 (BPI) [II-III]; El
Corte, 27 Mar 1993, |. FHennen, L.D.Ploper
& J.R.Herndndez 93-004 (LIL 54919, BPD
Tucuman:
ptt
332
[II]; Horco Molle, 14 Nov 1997, J.B.
Herndndez 97-135 (BPI 841238) [HI].
SCROPHULARIACEAE
Uredo sibthorpiae J.C. Lindq
on Sibthorpia conspicuda Diels; Tucuman:
Yerba Buena, 15 Feb 1996, |. R.Herndndez
96-084 (BPI 841155) [II].
SOLANACEAE
Aecidium
on on riparium Pers. Salta: Quebrada
de San Lorenzo, 6 Dec 1997, |. R. Herndndez
97-164 (BPI 841260) [0-1]. Tucuman: J.B
Alberdi, road to Escaba, shrine, 28 Nov
1997, |. R.Herndndez 97-105 (BPI 841222)
[0-1].
on Solanum tucumanense Poe » Salta: road
R
to Parque del Rey, 6 Dec 1997,
Herndndez 97-159 on 841255) [0- IL
on Mace ee (Sendtner) Hunz.;
ca: El Portezuelo, 30 Nov 1997,
J ae ez 97-128 (BPD [Ik El Rodeo,
29 Nov 1997, [.R.Herndndez 97-125 (BP
841230) [0-1].
Aecidium sp.
on Cestrum parqui UHér; Tucuman: San Miguel
deTucuman, 15 Aug 1995, [.R.Herndndez
95-085 (BPI 841110) [II].
Chrysocyclus cestri (Dietel & Henn.) Syd.
on Cestrum strigilatum Ruiz & Pav; Tucuman:
19.5 km W of San Miguel de Tucuman,
mtn, 27 Mar 1993, J. FHennen, L.D.Ploper
& J.R.Herndndez 93-012 (LIL 54922, BPD)
III; Dept. Trancas, Rt. 9, km 1364, 26 Oct
1995, |. R.Herndndez 95-057 (BPD [Ik El
Corte, 27 Mar 1993, J. hHennen, L.D.Ploper
& |. R.Herndndez 93-003 (LIL 54921, BPI)
HIII-IV]; Horco Molle, |4 Nov 1997,
J.R.Herndndez 97-131 (BPI 841235) [IIT
Horco Molle, 17 Sep 1997, J.R.Herndndez
97-081 (BPD [II], Horco Molle, Parque Si-
erras de San Javier, 6 Apr 1994, ).FHennen,
M.M.Hennen & ].R.Herndndez 94-090
(LIL 54923, BPD [III] San Javier, Parque
23 Jul 1994,
54924, BPI
ae
Sierras de San Javier,
J.-R.Herndndez 94-145 (LIL
841068) [III].
Endophyllum pampeanum (Speg.) J.C. Linda.
on Salpichroa origanifolia (Lam.) Baill.
Catamarca: Dept. Ambatos, Fl Rodeo, 23
Sy
BRIT.ORG/SIDA 20(1)
Sep 1997, J. FHennen & [.R.Herndndez 97-
047 (BPD [III Dept. Ambatos, El Rodeo,
997, J. FHennen &
15 Dec 1975, A.Villegas s.n. [1H]. Tucuman;
Dept. Trancas, Rt. 9, km 1364, 26 Oct 1995
|. R.Herndndez 95-054 (BPD [IH].
Puccinia araucana Dietel W Neger
on Cesirum sp. Salta: camino de corniza be-
tween Salta and Jujuy, 7 Dec 1997, J.B.
Herndndez 97-I76A (BPI 841264) [I].
Tucuman: San Pedro de Colalao, | Apr
1994, |. FHennen & M.M.Hennen 94-052
(LIL 54920, BPD II].
Puccinia incondita Arthur’
on Solanum abutiloides (Griseb.) Bitter &
ee Tucuman: cae Javier, Parque Sier-
ul 1994, | R. Herndndez
04-142 (LIL 54928, BPL 841066) [III
Puccinia negeriana Diete
de n lavie
on Lycopersicon esculentum Mill; Tweuman:
Jun 1987, L.D.Ploper s.n. (LIL 54925, BPI
840989) [M-IIT].
Puccinia pampeana Spe;
on Salpichroa origanifota (Lam.) Baill;
|. R.Herndndez 94-166 (BPD [II-I; Horco
Molle, 13 Sep 1997, .R.Herndndez sae
oe em 1-H; Horco Molle, 17 Sep
1997, |. R.Herndndez 97-077 (BI P1841206)
a |, NW of San Miguel de Tucuman,
31 Mar 1993, |. FHennen & |.R.Herndndez
93-094 (LIL 54926, BPI) [HI- HI].
Puccinia paradoxapoda Speg
on Grabowskia schizocdlyx Dammer® Salta:
Calayate, 9 Apr 1994, J. Hennen, M.M.
Hennen & J.R.Herndndez 94-119 (LIL
54927, BPI 841051) (I-III.
Uromyces cestri Mont.
on Cestrum lorentzianum Griseb, Catamarca:
El Rodeo, 29 Nov 1997, ].R.Herndndez 97-
122 (BPDII]; N of Catamarca, 22 Sep 1997,
J. RHennen & J.R.Herndandez 97-020 (BPI
841174) I Gracia, 19 Nov
1997, |. R.Herndndez 97-090 (BPI) [II
Dept. Traneas, 27 Oct 1995, ].R.Herndndez
95-060 (BPD [II San Pedro de Colalao, 31
Mar 1994, yas nnen & MM. Hennen 94-
045 (LIL 54937, BPI) [II].
on Cestrum parqui LHer: S
Tucuman: Alta ¢
alta: Cafayate, 8
HERNANDEZ AND HENNEN, RUST FUNG! OF NORTHWEST ARGENTINA
Apr 1994, J.RHennen & J.R.Herndndez
94-099 (LIL 54929, BPI) [LI-III|; Cafayate,
8 Apr 1994, J.EHennen & J.R.Herndndez
94- oe (LIL54931, BPD [II; Cafayate, 8
Apr 1994, J.FHenne -Herndndez
oa ae eae BPI 841040) [II-III].
Tucu . Capital, 15 Dec 1990,
paeniare ey _ 002 (BPD [II].
on Cestrum sp.; Salta: camino de corniza oe
tween Salta and Jujuy, 7
).R.Herndndez 97- en Tucuman:
Cevil Pozo, 12-15 km E of San Miguel de
Tucuman, 30 Mar 1993, |.FHennen &
J.R.Herndndez 93-053 (LIL 54934, BPI)
(Il; Cruz Alta, San Agustin, 30 Mar 1994,
J.-FHennen, M.M.Hennen& ].R. Hernandez
94-033 (LIL 54936, BPD [II Horco Molle,
Parque Sierras de San Javier, 6 Apr 1994,
J.-FHennen,M.M.Hennen& ].R.Herndndez
94-091B (LIL54733, BPD [II] Las Cejas, 30
Mar 1993, J.EHennen & J.R.Herndndez
93-067 (LIL 54935, BPI) [I]; San Javier,
Parque Sierras de San Javier, 27 Mar 1993,
J.EHennen, L.D.Ploper & J.R.Herndndez
93-023 (LIL 54932, BPI [II S
Parque Sierras de San Javier, 27 Mar 1993.
j.FHennen, L.D.Ploper & J.R.Herndndez
93-041 (LIL 54933, ae II]; SW of J.B
Alberdi, 22 Sep 1997, J. KHennen &
J
J.R.Herndndez 97 UUSUBELOTLIO IH HI].
San Javier,
THELYPTERIDACEAE
Desmella anemiae Syd. & P. Syd.*
Thelypteris sae el alg (Fée)
Schelpe*; Tucuman: Fl Corte, 27 Mar 1993.
j.EHennen, ae oper oe ndndez
93-001 (LIL 54938, BPD [II Horco Molle,
17 Sep 1997, J.R.Herndndez 97-076 (BPI)
[Il]; Horco Molle, Parque Sierras de San
Javier, 6 Apr 1994, J.RHennen, MM.
Hennen & J.R.Herndndez 94-084 (LIL
54940, BPD [II], San Javier, Parque Sierras
de San Javier, 27 Mar 1993, | FHennen,
L.D.Ploper & J.R.Herndndez 93-031 (LIL
54939, BPI) III.
on ae oe sp.*; Tucuman: Horco Molle, 14
7, J. R.Herndndez 97-137 (BPI
es ce I.
on
TILIACEAE
Puccinia triumfettae Dietel @ Holw.*
333
on Triumphetta sp.*, Tacuman: Horco Molle,
Parque Sierras de San Javier, 6 Apr 1994,
J.FHennen,M.M.Hennen & |.R.Herndndez
94-083 (LIL 54942, BPI 841036) [IIIh Tafi
del Valle, 6 Apr 1993, |. FHennen & .R.
Herndndez 93-121 (LIL 54941, BPD [III].
VALERIANACEAE
Accidium sp.
on Valeriana sp.*; Tucuman: J.B. eeu road
to Escaba, bank of creek, 2 v 1997,
J. R.Herndndez 97-102 (BPI a [0].
VERBENACEAE
Prospodium lippiae (Speg.) Arthur”!
on Aloysia ee (Gillies & Hook.) Trone:
Tucuman: Monte Redondo, 31 Mar 1996,
J. R.Herndndez oe 088 (BPI 841157) [II].
on oe polystachia (Griseb.) Moldenke;
ta: Cafayate, 8 Apr 1994, |. FHennen,
- M.Hennen & J.R.Herndndez 94-098
(LIL 54945, BPD [I-11] Tacuman: Ald
30 Mar 1993, [.R.Herndndez 93-070 (LIL
oe BPI) [II]; Dept. Capital, 17 Aug
994, ].R.Herndndez 94-136 (LIL 54946,
r 841062) [II-III]; Dept. Capital, 15 Dec
996, N.Cantén s.n. (BPI) [II]
on ee sp; Tucuman: Cruz Alta,San Agustin
30 Mar 1993, |. FHennen & J.R.Herndndez
93-058 (LIL 54950, BPD [II], Las Cejas, 30
Mar 1993, J]. FHennen & |.R.Herndndez
ee . IL sta BPD [II Las vee 30
Mar oes & J.R.Herndndez
ee i ie ue IIIk 1 as Cejas,
30 Mar 1993, J FHennen& |.R.Herndndez
93-062 (LIL 54951, BPI) [Il NW of S
Miguel de Tucuman, Mar 1993,
J.EHennen & J.R.Herndndez 93-084 (LIL
54953, BPD [II].
on Aloysia virgata (Ruiz & Pav.) Juss.*";
Metan, 2 Apr 1993, J. FKHennen & L.D.
Ploper 93-104 (LIL 54948, BPI) [II].
Tucuman: Cruz Alta, San Agustin, 30 Mar
1994. J.FHennen, M.M.Hennen &
J. R.Herndndez 94-032 (LIL 54949, BPI
841023) [I-III].
Puccinia lantanae Farl.
san
cal
Jadalla.
on Lantana sp., Tucuman: 22 km W of San
iguel de Tucuman, mtn, 27 Mar 1993,
J-FHennen, L.D.Ploper & J.R.Herndndez
93-017 (LIL 54955, “aa Mi40500) {111}; El
Manantial, 15 May 1996, J.R.Herndndez
96-094 (BPI) [Ik San ees Parque Sier-
ra ee ey 23 Jul 1994, |. R. Hernandez
94-163 (LIL 54956, a 841073) [II].
on - rid ae (Mill.) N.E.BrS; Tucuman: Dept.
Capital, 14 May 1995, |.R.Herndndez 95-
oan 841 090) i |.
on Li} 1 Cham.°; Tucuman
Denk Trancas, 5 oie 1905, Herndndez
95-002 (BPI 841078) [III].
on ss grisebachiana Moldenke*, Salta:
Dept. Santa Victoria, La Mision, 30 Jur
1996, |.R.Herndndez 96-062C (on oe
(H.W an: San Pedro de Colalao, | Apr
BRIT.ORG/SIDA 20(1)
Alberdi, road to Escaba, 28 Nov 1997,
J.R.Herndndez 97-100 (BPD) [I].
on Lippia sp. Salta: Dept. Santa Victoria, La
Mision, 30 Jun 1996, j.R.Her Lents 96-
062A (BPI) [III]. Tucuman: J.B erdi,
road to Escaba, shrine, 28 Nov een
Herndndez 97-104 (BPD [II|.
on Lippia turbinata Griseb. Tucuman: Dept.
Trancas, Vipos, 30 Mar 1995, ].R. Herndndez
95-045 (BPI 841088) [III].
VITACEAE
Endophyllum sieuaaLiaae Whetzel & Olive
var. catamarcensis J.C Linda. !
Horco
on Cissus sicyoides L.*, Tucuman:
1994, |} Hennen & M.M.Hennen 94-057
(LIL 54954, BPD [IIIL.
on Lippia modesta Briq.*:
Molle, Parque Sierras de San Javier, 6 Apr
1994, J FRHennen, M.M.Hennen & J.B.
ez 94-086 (LIL 54957, BPD [IH.
T an: |.b. 3
ucuman: J.B Herndnd
NEW RECORDS
New rust species (N):
Aecidium sp. (on Dyschoriste sp.), Aecidium sp. (on Solanum riparium Pers.,
Solanum tucumanense Griseb., Vassobia breviflora (Sendtner) Hunz.),
Cerotelium sp. (on Ruprechtia apetala Wedd., Ruprechtia laxiflora Meisn.,
Ruprechtia sp.), Endophyllum sp. (on Fuchsia boliviana Carriére), Puccinia sp.
(on Vernonia fulta Griseb., Vernonia sp.), Puccinia sp. (on Vernonia squamosa
Gardner), Puccinia sp. (on Ichnanthus minarum (Nees) Doll), Skierka sp. (on
Cupania vernalis Cambess.), Uredo sp. (on Psychotria carthagenensis Jacq.),
Ypsilospora sp. (on Inga edulis Mart.).
New rust records for Argentina (A):
Aecidium tournefortiae Henn., Catenulopsora praclonga (Speg.) Buritica,
Cionothrix praelonga (G. Winter) Arthur, Coleosporium vernoniae Berk. & M.A.
Curtis, Desmella anemiae Syd. & P.Syd., Edythea quitensis (Lagerh.) HSJackson
& Holw., Frommeella mexicana var. indicae J.W. McCain & J.E Hennen,
Melampsora epitea (Kunze & J.C. Schmidt) Thtim., Melampsoridium
hiratsukanum S. Ito ex Hirats. f., Phakopsora meibomiae (Arthur) Arthur,
Phakopsora neocherimoliae (Cummins) Buritica & J.F Hennen, Phakopsora
nishidana S. Ito, Phakopsora zizyphi-vulgaris Dietel, Phragmopyxis deglubens
(Berk. @ M.A. Curtis) Dietel in Engl. & Prantl, Physopella compressa (Mains)
Cummins & Ramachar, Prospodium pithecoctenii (Pazschke) Cummins,
Prospodium singeri Petr., Puccinia aristidae Tracy var. chaetaria Cumm. &
Husain, Puccinia bergii Speg., Puccinia calcitrapae DC. var. centaureae (DC.)
Cummins, Puccinia chaetochloae Arthur, Puccinia cnici-oleracei Pers., Puccinia
commelinae Holw., Puccinia cordiae Arthur, Puccinia coronata Corda var.
rangiferina (S. Ito) Cummins, Puccinia exornata Arthur, Puccinia hyptidis-
~—
HERNANDEZ AND HENNEN, RUST FUNGI OF NORTHWEST ARGENTINA 335
mutabilis Mayor, Puccinia inclita Arthur, Puccinia incondita Arthur, Puccinia
lateripes Berk. &@ Ravenel, Puccinia levis (Sacc.& Bizz.) Magnus var. panici-
sanguinalis (Rangel) Ramachar & Cummins, Puccinia ocellifera Cummins,
Puccinia pelargonii-zonalis Doidge, Puccinia proluviosa HS. Jacks. & Holw,
Puccinia rata HS. Jacks. & Holw, Puccinia tanaceti DC. var tanaceti, Puccinia
thunbergiae Cooke, Puccinia triumfettae Dietel & Holw., Puccinia unicolor
Arthur, Ravenelia cohniana Henn., Ravenelia echinata Lagerh. & Dietel var.
ectypa (Arthur & Holw.) Cummins, Ravenelia macrocarpa Syd. & P. Syd.,
Uromyces aspiliae H.S. Jacks. @ Holw., Uromyces castaneus P. Syd. & Syd.,
Uromyces celosiae Dietel & Holw., Uromyces dietelianus Pazsche, Uromyces
dolichosporus Dietel & Holw., Uromyces iresines Lagerh. ex Syd. & P. Syd.,
Uromyces orbicularis Dietel, Uromyces sp., Uromyces transversalis (Thtim.) G.
Winter.
New host plant genus for this rust worldwide (G):
Anadenanthera sp. (Ravenelia cebil Speg.), Chaetothylax umbrosus Nees
(Puccinia justiciae Puttemans), Dyschoriste sp. (Aecidium sp.), Ruprechtia
apetala Wedd. (Cerotelium sp.), Ruprechtia laxiflora Meisn. (Cerotelium sp.),
Ruprechtia sp. (Cerotelium sp
New host plant species for this rust worldwide (S):
Athyrium lilloi (Hicken) Alston (Desmella anemiae Syd. & P. Syd.), Baccharis
latifolia Lem. (Puccinia colossea Speg.), Calliandra formosa (Kunth) Benth.
(Ravenelia echinata Lagerh. & Dietel var. ectypa (Arthur & Holw.) Cummins),
Canna compacta Roscoe (Puccinia thaliae Dietel), Chamissoa altissima Jacq.)
Kunth (Uromyces celosiae Dietel & Holw.), Clematis montevidensis Spreng.
(Puccinia recondita Roberge ex Desm.), Clematis sericea Michx. (Puccinia
recondita Roberge ex Desm.), Cologania ovalifolia Kunth (Phakopsora
meibomiae (Arthur) Arthur), Coursetia brachyrhachis Harms (Phragmopyxis
deglubens (Berk. & M.A. Curtis) Dietel in Engl. & Prantl), Desmodium
subsericeum Malme (Uromyces orbicularis Dietel), Echinochloa cruspavonis
(Kunth) Schult. (Puccinia abnormis Henn.), Grabowskia schizocalyx Dammer
(Puccinia paradoxapoda Speg.), Iresine diffusa Humb. & Bonpl. ex Willd. var.
diffusa (Uromyces iresines Lagerh. ex Syd. & P. Syd_), Janusia guaranitica (H. St.-
Hil.) A. Juss. (Puccinia heteropteridis Thtim.), Lippia alba (Mill.) N.E.Br.
(Puccinia lantanae Farl.), Lippia cfr. turnerifolia Cham. (Puccinia lantanae
Farl.), Lippia grisebachiana Moldenke Wai lantanae Farl.), Lippia modesta
Brig. (Puccinia lantanae Farl.), Lonchocarpus lilloi (Hassl.) Burkart (Ravenelia
lonchocarpiicola Speg.), Mimosa debilis Humb. & Bonpl. ex Willd. var. debilis
(Ravenelia mimosae-sensitivae Henn.), Parapiptadenia excelsa (Griseb.) Burkart
oe nent papillosa Speg.), Pavonia malvacea (Vell.) Krapov. & Cristoval
(Cat ‘a praelonga (Speg.) Buritica), Ruellia ciliatiflora Hook. (Puccinia
ee Berk. & Ravenel), Ruellia erythropus (Nees) Lindau (Puccinia sp.),
ay
336 BRIT.ORG/SIDA 20(1)
Salvia rypara Brig. (Puccinia farinacea Long), Senecio bravensis Cabrera
(Coleosporium tussilaginis (Pers.) Lév.), Senecio cremeiflorus Matt.
(Coleosporium tussilaginis Pers.) Lév), Senecio otopterus Griseb. (Coleosporium
tussilaginis (Pers.) Lév.), Senecio peregrinus Griseb. (Puccinia proluviosa H.S.
Jacks. &@ Holw.), Senna subulata (Griseb.) H.S. lrwin & Barneby (Ravenelia
macrocarpa Syd. & P. Syd.), Serjania marginata Casar. (Puccinia arechavaletae
Speg.), Simsia dombeyana DC (Puccinia enceliae Dietel & Holw. var. aemulans
(Syd. & P Syd.) Parmelee), Solanum abutiloides(Griseb.) Bitter & Lillo (Puccinia
incondita Arthur), Solanum riparium Pers. (Aecidium sp.), Solanum
tucumanense Griseb. (Aecidium sp.), Stachys gilliesii Benth. (Puccinia
anime le Speg.), Iarasad jorgensenii (.M.Johnst.) Krapov.(Puccinia platyspora
eae S. Jacks. @ Holw.), Tournefortia rubicunda DC. (Uromyces
dolic ae us Dietel @ Holw.), Tripogandra elongata (G. Mey.) Woodson
(Puccinia commelinde Holw.), Vassobia breviflora (Sendtner) Hunz. (Aecidium
sp.), Vernonia fulta Griseb. (Puccinia sp. A), Vernonia remotiflora Rich.
(Puccinia cnici-oleracei Pers.), Vernonia saltensis Hieron. (Puccinid cnici-
oleracei Pers., Puccinia rata H.S. Jacks. @ Holw.), Wedelia saltensis Cabrera
(Uromyces aspilide HS. Jacks. & Holw.), Zizyphus mistol Griseb. (Phakopsora
zizyphi-vulgaris Dietel).
—
i
New host plant species for any rust in Argentina (A):
Alcea rosea L. (Puccinia platyspora (Speg.) H.S. Jacks. @ Holw.), Alcea sp.
(Puccinia platyspora (Speg.) HS. Jacks. @ Holw.), Alnus acuminata Kunth
(Melampsor idium hiratsukanum S. Ito ex Hirats. f.), Bauhinia forficata Link
(Uromyces dietelianus Pazsche), Carthamus tinctorius L. (Puccinia calcitrapae
DC. var. centaurede (DC.) Cummins), Cissus sicyoides L. (Endophyllum
circumscriptum Whetzel & Olive var. catamarcensis J.C. Lindq.), Cucurbitella
cucumifolia (Griseb.) Cogn. (Uromyces novissimus Speg.), Cupania vernalis
Cambess. (Skierka sp.), Desmodium purpureum Hook. & Arn. Uromyces orbicu-
laris Dietel), Desmodium purpureum Hook. & Arn. (Uromyces castaneus P. Syd.
& Syd.), Dichondra sericea Sw. (Puccinia dichondrae Mont.), Duchesnea indica
(Andrews) Focke (Frommeella mexicana var. indicae J.W. McCain & J.F
Hennen), Gladiolus sp. Uromyces transversalis(Thtm.) G. Winter), Hydrocotyle
bonariensis Comm. ex Lam. (Puccinia hydrocotyles Cooke), Ichnanthus
minarum (Nees) Déll (Puccinia sp. B, Puccinia inclita Arthur, Puccinia levis
(Sace.& Bizz.) Magnus var. panici-sdnguinalis (Rangel) Ramachar @ Cummins),
Leptochlod mucronata (Michaux) Kunth (Puccinia leptochlode Arthur),
Mikania micrantha Kunth (Puccinia spegazzinii De Toni), Pelargonium
hortorum L.H. Bailey (Puccinia pelargonii-zonalis Doidge), Poiretia tetraphylla
(Prior) Burkart (Puccinia bergii Speg.), Pteris deflexa Link (Desmella anemiae
Syd. & P.Syd.), Rubus boliviensis Focke (Gerwasia imperialis (Speg.) J.C. Lindq.,
Jacks. & Holw.), Senna birostris var.
—
Kuehneola loeseneriana (Henn.) H.S
HERNANDEZ AND HENNEN, RUST FUNGI OF NORTHWEST ARGENTINA 337
hookeriana (Gillies ex Hook. & Arn.) H.S. Irwin & Barneby (Ravenelia
macrocarpa Syd. & P. Syd.), Senna hookeriana Batke (Ravenelia macrocarpa Syd.
& P. Syd.), Setaria lachnea (Nees) Kunth (Puccinia chaetochloae Arthur), Se-
taria parviflora (Poir.) Kerguélen (Puccinia chaetochloae Arthur), Sorghum
cafrorum (Thunb.) P. Beauv. (Puccinia purpurea Cooke), Thelypteris
quadrangularis (Fée) Schelpe (Desmella anemiae Syd. & P. Syd.), Thelypteris
sp.(Desmella anemiae Syd. & P. Syd.), lournefortia paniculata Cham. (Aecidium
tournefortiae Henn.), Tripogandra sp. (Puccinia commelinae Holw.),
Triumphetta sp. (Puccinia triumfettae Dietel @ Holw.), Valeriana sp.(Aecidium
sp.).
ak
New host plant species for which another rust was previously known from
Argentina (AR):
Aloysia virgata (Ruiz & Pav.) Juss. (Prospodium lippiae (Speg.) Arthur), Aristida
sp. (Puccinia aristidae Tracy var. chaetaria Cumm. & Husain), Aspilia sp.
(Uromyces aspiliae H.S. Jacks. & Holw.), Avena sativa L.(Puccinia coronata Corda
var. rangiferina (S. Ito) Cummins), Baccharis sp.(A) (Puccinia exornata Arthur),
Baccharis sp.(B) (Puccinia exornata Arthur), Baccharis lilloi Heering (Puccinia
baccharidis Dietel & Holw.), Baccharis sp. (Puccinia unicolor Arthur), Chloris
gayana Kunth (Puccinia cacabata Arthur & Holw.), Chrysanthemum sp.
(Puccinia tanaceti DC. var tanaceti), Clytostoma callistegioides (Cham.) Bureau
ex Griseb. (Prospodium singeri Petr), Digitaria insularis (L.) Fedde (Puccinia
odahuensis Ellis & Everh.), Elephantopus mollis Kunth (Coleosporium vernoniae
erk. @ M.A. Curtis), Eupatorium macrocephalum Less. (Puccinia conoclinii
Seym. in Burrill), Eupatorium sp. (Cionothrix praelonga (G. Winter) Arthur),
Hyptis mutabilis (Rich.) Brig. (Puccinia hyptidis-mutabilis Mayor), Hyptis sp.
(Puccinia hyptidis-mutabilis Mayor), Inga edulis Mart. (Ypsilospora sp.), Pani-
cum sp. (Puccinia levis (Sacc.& Bizz.) Magnus var. panici-sanguinalis (Rangel)
Ramachar & Cummins), Paspalum sp. (Physopella compressa (Mains) Cummins
& Ramachar), Pluchea sp. (Puccinia ocellifera Cummins), Pteris sp. Desmella
anemiae Syd. & P. Syd.), Salix babylonica L.(Melampsora epitea (Kunze & J.C.
Schmidt) Thtim.), Salix caprea L. (Melampsora epitea (Kunze & J.C. Schmidt)
Thum.), Salix humboldtiana Willd. (Melampsora epitea (Kunze & J.C. Schmidt)
Thtim.), Setaria sp. (Puccinia chaetochloae Arthur), Tradescantia sp. (Puccinia
commelinae Holw,).
—
ACKNOWLEDGMENTS
] ] ]
The first author gratefully his advisor and friend, L. Daniel Ploper,
National University of Tucuman, who provided support in completing his doc-
toral thesis. Many thanks to Maria Magdalena Schiavone and her staff at the
herbarium of the Fundacion Miguel Lillo (LIL), San Miguel de Tucuman,
Tucuman, who organized and accessioned many specimens, and to Alberto
338 BRIT.ORG/SIDA 20(1)
Slanis for host plant identifications. The efforts of Erin McCray and David F-
Farr in providing accession numbers and assistance in the final publication,
respectively, are appreciated as are the encouragement and efforts of Amy Y.
Rossman that contributed to the completion of this paper. Finally, | would like
to thank my wife, Mary E. Palm, for her invaluable motivation and support.
REFERENCES
Brummit, R.K. and C.E. Powett (Eds.). 1992. Authors of plant names. Royal Botanic Gardens,
Kew, UK.
Caprera, A.L. 1994. Regiones fitogeograficas argentinas. Enciclopedia Argentina de
Agricultura y Jardineria (Ira reimpresion) 1. ll. Buenos Aires, Argentina.
Farr, M.L.1973.An annotated list of Soegazzini’s fungus taxa. Vol. 1:1-823,Vol.2:824-1661.
J.Cramer, Lehre, Germany.
Heananoez, J.R. 2000. Baeodromus ranunculi,a new rust on Ranunculus from Argentina and
a synopsis of Baeodromus. Mycotaxon 76:329-336.
Kirk, BM. and A.E. Ansett. 1992. The author of fungal names. CAB International.Wallingford,
United Kingdom.
Linoquist, J.C. 1982. Royas de la Republica Argentina y Zonas Limitrofes. Instituto Nacional
de Tecnologia Agropecuaria, Buenos Aires, Argentina.
ADDITIONS TO THE CYPERACEAE AND JUNCACEAE
FLORA OF THE RIO MAYO REGION, SONORA, MEXICO
Eric H.Roalson Thomas R.Van Devender and
School of Biological Sciences Ana Lilia Reina G.
Washington State University eae see Desert Museum
Pullman, WA 99164-4236, U.S.A. 2021 N. Kinney Rd.
Tucson, 7 85743, USA,
ABSTRACT
Intensive floristic surveys of the Rio Mayo region of the Mexican state of Sonora, particularly the
Municipio de Yécora, have uncovered more species of o peraceae not previously re reported from the
co | Te
region, the state of Sonora northern Me es of Cy peraceae
t 14 spec
and two species of Juncaceae not previously cas to occur in the Rio Mayo oe ‘art the pres-
ence of three varieties he one lane us (Cy]
extension involved witl y. This aes the total number of Cen aceae taxa known
{ the range
from the Rio Mayo region to 99 sade ae taxa to |4.
RESUMEN
I } 1 . : s) eee i : giond RinM ] J+A ; 1:6
particularmente en el Municipio de Yécora, han descubierto mas ae de Cyperaceae que no
habian sido citadas previamente de esta region, del est o, en algunos casos, el norte de
México. Presentamos qui 14 especies de Cyperaceae y dos especies de ee que no se conocian
previamente de la region de Rio Mayo, clarificamos la presencia de tres variedades y una especie de
Cyperus (Cyperaceae), y se discute el significado del rango de extension implicado en cada
1 brimi sto lleva el numero total de ta idosd de la region Rio Mayo
a
a 99 y el taxones de Juncaceae a 14.
Gentry’s 1942 Rio Mayo Plants was a major contribution to the knowledge of
the flora and vegetation of northwestern Mexico. The Rio Mayo region includes
the area in southern Sonora and adjacent Chihuahua between the Rio Fuerte
and Rio Yaqui drainages. Beginning in the 1970s, a new network of roads, espe-
cially Mexico Federal Highway 16 (MEX 16) between Hermosillo, Sonora, and
La Junta, Chihuahua, was completed in 1992 and provided access to many ar-
eas of the Sierra Madre Occidental not previously explored floristically (Burquez
et al. 1992). A revision of the Rio Mayo flora (Martin et al. 1998) incorporated
recent floras from the Cascada de Basaseachi (Spellenberg et al. 1996) and
Nabogame (Laferriére 1994), Chihuahua. The total of 2825 taxa reported for the
Rio Mayo region included 1] genera and 77 species of Cyperaceae and two gen-
era and 12 species of Juncaceae.
An intensive floristic survey of the Municipio de Yécora, an area of 3,300
km? along MEX 16 in the Sierra Madre Occidental in eastern Sonora, was be-
gun in 1995 (Reina et al. 1999; Van Devender et al. in press). The current total for
the Municipio is 1648 taxa, raising the total for the Rio Mayo region to over 3000
SIDA 20(1): 339 — 347. 2002
340 BRIT.ORG/SIDA 20(1)
taxa. The families with the most taxa in the Municipio de Yécora flora are
Compositae (248 taxa), Gramineae (186 taxa), Leguminosae (168 taxa),
Cyperaceae (59 taxa), and Euphorbiaceae (56 taxa). The genus Cyperus with 29
species is especially diverse. This ongoing floristic work in the Rio Mayo region
has uncovered more species of Cyperaceae not previously reported from the
region, the state of Sonora, or, in some cases, northern Mexico. Here we present
14 species of Cyperaceae and two species of Juncaceae not previously known to
occur in the Rio Mayo region, clarify the presence of three varieties of one
Cyperus (Cyperaceae) species, and discuss the significance of the range exten-
sion involved with each new discovery. Specimens were determined by Eric H.
Roalson, unless noted otherwise. Most of the noteworthy records are in cienegas
or other moist areas from 1200 to 2100 m elevation in oak woodland and pine-
oak forest in the Municipio de Yécora, or the grasslands and mudflow barrens
within them (Reina G. et al. 1999).
Bulbostylis barbata (Rottb.) C.B.Clarke
This species is previously known from the southeast United States, Martinique
Guatemala, and the Old World tropics (Kral 1971). Bulbostylis barbata appears
to be quite weedy and expanding its range in North America (Kral 1971). This is
the first report of the species in Mexico.
Voucher specimens. MEXICO. Sonora. Municipio de Yécora: 14.7 kim E of Maycoba, 7.7 km Eof El Kipor
on MEX 16(KM 342), pine-oak forest, 28°20'04'N 108°33'36"W, 1460 m elevation, locally common a
in soil pocket in bare area within forest, 15 Sep 1999, Van Devender 99-647 (WS), with A.L. Rein
Bulbostylis pubescens (J. @ C.Presl) Svenson
This species is widespread in the lowlands to the south along the Pacific slope
of the Sierra Madre including the Mexican states of Sinaloa, Nayarit, Jalisco,
Guerrero, Mexico, Oaxaca, and Chiapas, and south through Central America
and northern South America (Kral 1971; McVaugh 1993). This is the first report
of the species from the Rio Mayo region and the state of Sonora.
Voucher specimens. MEXICO. Sonora. Munici cipio de Yécora: NW yf the cemetery in Yécora, spare
oak woodland on barren volcanic hilltops, 28°22'40"N 108°50'W, 1540 m elevation, locally common
on moist rock surfaces, 23 Sep 1997, Reina G. 97-1172 (ARIZ, CAS, MEXU, NY, WS), with TR. Van
Devender and W. Trauba; 1.9 km SSW of Las Viboras on MEX 16 on road to Trigo Moreno, bare mud-
stone area in open pine-oak forest, 28'21'50"N 108'49'34"W, 1620 m elevation, locally common annual,
17 Aug 1998, Van Devender 98-992 (ARIZ, WS), with A.L. Reina G., M.E. Fishbein, and G.M. Ferguson;
tributary of Arroyo Los Pilares near bridge, a on W ai WE ee MEX 16, Gak eee with
scattered pines in shady 3'4+0"N 108°47'35"W, 1300 locally com
mon annual in moist depressions on “ bare sur cies 13 Sep 1999, Reina G. 99-515 (WS), with T.R. Van
Devender
Carex planostachys Kunze
This species is previously known [rom Texas to Guatemala, and in Mexico, it is
known from as far northwest as Chihuahua (Hermann 1974). This is the first
report of this species in Sonora and the Rio Mayo region.
ROALSON ET AL., ( 341
Voucher specimens. MEXICO. Sonora. Municipio de Yécora: 5.2 km W of Yécora on MEX 16, pine-
oak forest, 28°21'48"N 108°59'12"W, 1720 m elevation, 11 Mar 1996, Van Devender 96-87 (ARIZ, MEXU,
RSA), with A.L. Reina G., S.L. Friedman, and W. Trauba, determined by Merriam C. Fritts.
Carex thurberi Dewey
This species is known from Arizona and localities across Mexico (Sonora, Chi-
huahua, Jalisco, Veracruz, and Chiapas), as well as Guatemala (Hermann 1974;
although see Mc Vaugh 1993; Espejo &@ Lopez F 1997). This is the first report of
this species for the Rio Mayo region.
Voucher specimen. MEXICO. Sonora. Municipio de Yécora: ca. 2 km (by air) W of Yécora, pine-oak
forest, 28°22'15"N 108°57'30"W, 1580 m elevation, uncommon herbaceous perennial in moist soil, 21
Jul 1998, Trauba 425-98 (WS).
Cyperus arsenei O'Neil & Ben. Ayers
This species is previously known from the western Mexican states of Baja Cali-
fornia Sur, Nayarit, and Michoacan and the Districto Federal (Tucker 1994). This
is the first record of this species in the state of Sonora and the Rio Mayo region.
Voucher specimens. MEXICO. Sonora. Municipio de Yécora: | km W of Yécora on volcanic ridges/
outcrops, W of cemetery, E facing slope with occas | Arctostaphylos, Bouteloua, other Cyperus sp.,
Hackelochloa, Juniperus, Muhlenbergia, occasional Pinus, Quercus, Schizachyrium, Scleria, and Tra-
descantia, 28°22'12"N 108'56'23"W, ca. 1600 m elevation, 5 Sep 1996, Roalson 1364 (WS).
Cyperus manimae HBK
In Gentry’s Rio Mayo Plants (Martin et al. 1998), varieties of C. manimae were
not distinguished. Tucker (1994) recognized three varieties of C. manimae, with
only one of these (C. manimae var. asperrimus) known from Sonora. Cyperus
manimae var. divergens is known from Chihuahua and Coahuila south to
Chiapas in Mexico and south to Costa Rica and C. manimae var. manimae is
known from Baja California, Sonora, and Durango to Chiapas in Mexico, south
to Guatemala and northern South America (Tucker 1994; Espejo &@ Lopez F
1997). All three varieties are present in the Rio Mayo region. This increases the
range of var. asperrimus and var. manimae into the Rio Mayo region, and the
range of var. divergens west from Chihuahua into Sonora.
var. asperrimus (Liebm.) Kitk.
Voucher specimens. MEXICO. Sonora. Municipio de Yécora: Rio Yepachic near junction with Arroyo
Hondo, ca. 2 km (by air) W of Chihuahua border, . woodland with canyon riparian forest with
cer, Alnus, Cupressus, and Prunus gentry, 28°27' 108°32'15"W, 1380 m elevation, locally com-
mon perennial on shady slope, 27 Sep 1998, Van ea 98-1783 ae with A.L. Reina G.
var. divergens (HBK) Ktik
Voucher tpguieg MEXICO. Sonora. uv da de Yécora: Canada La Ventana (Arroyo El Otro Lado),
2.5 km (by air) ESE of pela rocky st pine-oak forest, 28°21'38"N 108°53'55"W, 1520 m
elevation. solitary *ky sl ope 18 Sep 1998, Van Devender 98-1340 (WS), with A.L. Reina
G.and Trauba; near Rancho El Bede egoso, 15.6 km N of Yécora on road to Agua Blanca, oak wood-
land wi assland, 26°30'04'N 108°55'38"W, 1460 m el locally common perennial in mud at
edge of oS 24 ae 1998, Van Devender 98- 659 W S), with A.L. Reina G. and W. Trauba.
342 BRIT.ORG/SIDA 20(1)
var. manimae
Voucher specimens. MEXICO. Sonora. Municipio de Yécora: Pueblo Viejo on Rio Mayo in Navojoa,
27°06'N 109°25'40"W, ca. 35 m elevation, ee common herbaceous perennial, 2 Oct 1995, Van
Devender 95-1127 (ARIZ, WS), with AL. 1G: 2 km Eof Yécora, northeast of Las Cabanas Los
Champinones, disturbed pine-oak forest, sik 108°54'53"W, Lo00 m elevation, common 60 cm
tall herbaceous perennial in moist places, 3 Oct 1996, Flores M. 4969 (USON, WS), with J. Sanchez; ca.
2km NW ol Yécora on old road to Santa Rosa, 28'22'33"N 108°56'24"W., 1560 m elevation, uncommon
perennial in moist soil, 17 Aug 1998, Van Devender 98-1009 (EIU, WS), with A.L. Reina G., M-E. Fishbein,
and G.M. Ferguson; Canada La Ventana (Arroyo El Otro Lado), 2.5 kin (by air) ESE of Yécora, rocky
stream canyon in pine-oak forest, 28°21'38"N 108°53'55"W, 1520 m elevation, uncommon perennial on
rocky slope, 18 Sep 1998, Van De vender 98-1342 (WS), with A.L. Reina G. and W. Trauba; Rio Maycoba
at MEX 16 (20.5 km W of Maycoba, 28.6 km EF of Yécora), 28°22'15'N 108 45) 30" W, 1220 m elevation,
rare sedge at base of cliff, 26 Sep 1998, Reina G. 98-1711 (WS), with T.R. Van Devender, road to Rio
Yepachic from MEX 16 near Arroyo Hondo (11.5 km Eot El Kiporon MEX 16), oak woodland, 28'26'50"N
108°32'20"W, 1480 m elevation, very common on disturbed roadside, 27 Sep 1998, Van Devender 98-
ISO8 (WS), with A.L. Reina G.
a
ou
Cyperus sphaerolepis Boeck.
This species is known from New Mexico, Arizona, and west Texas south to
Puebla and the Districto Federal in Mexico, and has been documented previ-
ously from Sonora (Mc Vaugh 1993; Tucker 1994; Espejo & Lopez FE 1997). This
species was not listed, however, for the Rio Mayo region (Martin et al. 1998). These
specimens verily its presence there.
Voucher specimens. MEXICO. Sonora. Municipio de Yécora: | kim W of Yécora on mudflow ridges/
outcrops, W of cemetery, F facing slope with occasional Arciosta phylos, Boule tou, other Cyperus sp.,
Hackelochloa, Juniperus, Muhlenbergia, occasional Pinus, Quercus, Sch cleria,and Tra-
descantia, 28°22'12"N 108°56'23"W, ca. 1600 m ae ration, 5 Sep 1996, Roalson 1360. 1362. and 1363 (all
WS): NW of the cemetery in Veeors apes oak woodland on barren mudflow hilltops, 28°22'40"N
108'56'W, 1540 m elevation ge in wet areaon bare mudtlow | hilltop, 23Sep 1997, Reina
G. 97-1162 (WS) and 97-1183 (ARIZ, MEXU, WS), with T.R. Van Devender and W. Trauba; Ciénega de
Camilo, 6.3 km E of El Kipor, 11.2 km W of Chihuahua border on MEX 16, sphagnum bog in clearing
in shady riparian ae pcan 28'46'43'N 108°31'50"W, 1580 m elevation, solitary in marsh, 25 Sep
1997, Reind G. 97-1412 (WS), with T.R. Van Devender (Van Devender et al. in press); El Encinal, ca. 3.5
km E of Maycoba, 28° atte 'N 108°37'W, L600 m elevation, locally common perennial in moist soil in
open rl woodland, 20 Nov 1997, Van Devender 97-1538 (ARIZ, WS), with ALL. Reina G., A.M. Rea, C
Cassa, and A.E. Gondor.
Eleocharis macrostachya Britton
This is a widespread species known from much of the United States, south-
western Canada, and north-central Mexico. In Mexico it is previously known
from Chihuahua, Durango, Aguascalientes, Jalisco, Michoacan, Mexico,
Querétaro, the Districto Federal, Oaxaca, Puebla, Hidalgo, and San Luis Potosi
(McVaugh 1993). This is the first report for its presence in the Rio Mayo region
and the first report for Sonora.
Voucher specimen. MEXICO. Sonora. Municipio de Yécora: Yécora, freshwater marsh, 28°22'02"N
108°55'47"W. 1505 m elevation, occasional herbaceous perennial in marsh, 16 Jul 1997, Van Devender
97-816 (ARIZ, MEXU, USON, WS), with A.L. Reina G., D. Larson, P Merlin, MJ. Martinez C., G. Fergu-
ROALSON ET AL., 343
son, M. Kaib, and R.L. Bellsey; below El Llano on Mesa del Campanero (W of Yécora), 28°20'30"N
109°01'55"W, 2100 m elevation, locally common sedge in moist soil around well, 1 Jun 1999, Reina G.
99-176 (WS), with T.R. Van Devender, K. Baker, P. West, and R. Scarborough.
Eleocharis svensoniana S.Gonzalez F.
This species was described in 1985 from the state of Durango, Mexico (Gonzalez
E.1985) and is also known from the states of Jalisco (Mc Vaugh 1993) and Sinaloa
(Espejo & Lopez F 1997). These collections represent the first record of this spe-
cies in the Rio Mayo region and Sonora and are a significant northward expan-
sion of the species distribution from Durango and Sinaloa.
Voucher specimen. MEXICO. Sonora. Municipio de Yécora: NW of the cemetery in Yécora, sparse
oak woodland on barren nea hilltops, 28°22'40"N 108°56'W, 1540 m elevation, uncommon on
moist rock surfaces, 23 Sep 1997, Reina G. 97-1171 (WS), with TR. Van Devender and W. Trauba; Ciénega
de Camilo, 6.3 km E of El Kipor, 11.2 km W of Chihuahua border on MEX 16, sphagnum bog in clear-
ing in shady riparian pine-oak forest, 28°46'43"N 108°31'50"W, 1520 m elevation, abundant sedge in
mud at edge of ciénega, 27 Sep 1998, Van De vender 98-1827 (WS), with A.L. Reina G. (Van Devender et
al. in press).
oe
Eleocharis yecorensis E.H.Roalson
This species was described in 1999 from material collected in a freshwater
marsh on the outskirts of Yécora (Roalson 1999). It now appears that this spe-
cies is quite widespread, with documented localities as far south as Nicaragua
(Stevens et al. 2001)
Voucher specimens. MEXICO. Sonora. Municipio de Yécora: 2.0 km S of MEX 16 at Yécora, fresh
water marsh in grassland, 28°21'42"N 108°55'48" W, 1540 m elevation, 7 Sep 1996, Reina G. 96-486 (ARIZ,
WS), with TR. Van Devender and W. Trauba; SSW side of Yécora, 28°22'09"N 108'56'04'"'W, ca. 1500 m
elevation, 8 Sep. 1996, Roalson 1385 (Type collection, MEXU, MICH, MO, RSA, UC, WS); Yécora,
28'22'02"N 108°55'47"W, 1,505 m, 16 Jul 1997, Van Devender 97-810 (ARIZ, WS).
Fimbristylis complanata (Retz.) Link
This species is previously known from the New and Old World tropics (Kral
1971; McVaugh 1993), and prior to Martin et al’s (1998) discovery of it in Chi-
huahua, the northernmost record of this species in the New World was from
Durango. This is the first report of this species from Sonora. This species has
also recently been collected by Richard S. Felger and collegues N of Guaymas,
onora (Municipio de Guaymas, Canon La Balandrona, N side of Sierra El Aguaje,
28°04.27'N, 111'04.44'W, 825 ft. elevation, 19 Dec 2001, Felger 01-659 (USON,
ARIZ, MEXU), with J. Sanchez E. and PA. West; RS. Felger, pers. comm.).
Voucher eo MEXICO. Sonora. Municipio de Yécora: Arroyo La Cueva, 8 km N of Yécora on
the road to Agua Blanca, rocky stream canyon in pine-oak forest, 28°2627"N 108°55'29"W, 1520 m
elevation, uncommon ag, in moist soil at edge of stream, 24 Sep 1998, Van Devender 98-1633
CWS), with A.L. Reina G. and W Trauba.
Fimbristylis decipiens Kral
This species was described by Kral in 1971“... to represent an intermediate mor-
phology between F dichotoma and F annua.” It is difficult to tell F decipiens
344 BRIT.ORG/SIDA 20(1)
—
from F dichotoma and F- annua, but the two specimens listed below seem to fal
within Kral’s circumscription. Kral recognized this species in the southeast
United States, but suggested it was likely in Central and South America and the
Old World (Kral 1971). This is the first report for Sonora and the Rio Mayo region.
Voucher specimen. MEXICO. Sonora. Municipio de Yécora: 2.1 km W of Arroyo Hondo, 6.1 km W ol
Chihuahua border on MEX 16, open Se sleclae ees woodland on steep pink and white volcanic
ash, 28°26'24"N 108°33'20"W, 1400 m elevation, 25 Sep 1997, Reina G. 97-1374 (ARIZ, MEXU, NY, WS),
h T.R. Van Devender; Arroyo El Toro on NE adge of Yécora, 28'22'45'N 108°55'15"W, 1520 m eleva-
ocally common sedge in mud near stream, 2 Oct 1998, Van Devender 98-1997 (WS), with ALL.
wit
—
tion,
Reina G.
Fimbristylis pentastachya Boeck.
This species has been traditionally known from southern Mexico (Kral 1971).
McVaugh (1993) listed FE pentastachya from southern Sonora and Martin et al.
(1998) listed it for the Rio Mayo region in Chihuahua. This collection expands
its range into the Sonoran portion of the Rio Mayo region.
Voucher specimen. MEXICO. Sonora. Municipio de Yécora: 1.0 km SSW of Las Viboras on MEX 16 on
road to Trigo Moreno, bare mudstone area in open pine-oak forest, 28'22'18'N 108°49'42"W, 1620 m
ee vation, uncommon herbaceous perennial in moist soil, 17 Aug 1998, Van Devender 98-998 (WS),
1 A.L. Reina G., M.E. Fishbein, and G.M. Ferguson.
=.
Lipocarpha maculata (Michx.) Torr.
This species is previously known from the southeastern United States, in Mexico
the states of Nayarit, Jalisco, Veracruz, and Chiapas, Central and South America,
and the West Indies, although this species is thought to be uncommon in Mexico
(McVaugh 1993). This is the first report of this species from the Rio Mayo region
and the state of Sonora, with the closest documented locality to the south in
Jalisco and Nayarit (Espejo & Lépez F 1997).
Voucher specimen. MEXICO. Sonora. Municipio de Yécora: SSW side of Yécora, 28°22'09"N
108°56'04'"W, approximately 1500 m elevation, 8 oe 1996, en 1389 gos! 5), Canada La Ventana
38"N
(Arroyo Fl Otro La do y), 251 m (by air) ESE of Yecc ora Yr cl 5 Oak forest, 28°21
108°53'55"W, 1520 m elevation, locally abundant dwarl a on shallow soil on mud flow slope, |
Oct 1998, Reina G. 98-1967 (ARIZ, WS), with T.R. Van Devender and W. Traub
Scleria interrupta Rich.
Two collections from the Rio Mayo region apparently fall within the circum-
scription of Scleria interrupta (Konraed Camelbeke, pers. comm.). This is an
extreme range extension with the closest localities of this species in Chiapas
(Espejo & Lopez F. 1997). The two collections vary somewhat in the character-
istics of pubescence and extent of the tubercles on the achene. Reina G. 97-1373
has achenes with a large number of tubercles covering the surface which at
times approach a reticulate pattern and most of the bracts subtending the spikes
are covered in bristly hairs on the margins and midrib. This appears to fit wel
within the circumscription of S. interrupta, somewhat tending towards the
sometimes-segregate S. pinetorum Britton which is included in synonymy of S.
—
ROALSON ET AL., ( 345
interrupta here (Core 1936; Raynal 1976; Konraed Camelbeke, pers. comm.). The
other collection, Reina G. 97-1174, includes plants with achenes that are mostly
smooth with a few tubercles at the achene apex and the bracts subtending the
spikes are often near glabrous with few, scattered, bristly hairs. These plants do
not fitas well in the circumscription of S.interrupta,and approach S. verticillata
Muhl. (due to the lack of bristly hairs), or maybe S. distans Poir. (due to the
smooth achenes; Core 1936). For now these two collections are treated as S.
interrupta, but more detailed studies of the variation in achene ornamentation
and pubescence in these annual Scleria species is necessary.
Voucher specimens. MEXICO. Sonora. Municipio de Yécora: NW of the cemetery in Yécora, spare
oak woodland on barren volcanic hilltops, 28°22'40"N 108°56'W, 1540 m elevation, locally common
on moist rock surface, 23 Sep 1997, Reina G. 97-1174 (ARIZ, NY, WS), with T.R. Van Devender and W.
auba; 2.1 km W of Arroyo Hondo, 6.1 km W of Chihuahua border on MEX 16, open Sao aie
_ woodland on steep pink and w ce mottled volcanic ash, 28°26'24"N 108°33'20"W, 14 eleva-
tion, locally common in moist soil depression on rocky slope, 25 Sep 1997, Reina G. - as (WS),
ith T.R. Van Devender
Juncus dichotomus Eliott
This species is quite widespread, occurring from Alaska, across the United States,
Mexico, Central America, and South America (Balslev 1996; Espejo & Lopez F
1996; Brooks &@ Clemants 2000). This is the first report of J. dichotomus for the
Rio Mayo region.
Voucher seein MEXICO. Sonora. Municipio de Yécora: Fl Divisidero . Bordo), | km SE of El
oe road to Bermudez, Mesa tae Campanero, pine-oak forest, 28'1648"N 109°02'31"W, 1800 m
aad common in moist soil, 26 May 1996, Reina G. 96- Se enna WS), with TR.
Van Devender, A. Burquez M., and L. Varela; Mesa del Campanero, Arroyo Largo, upper tributary or
Barranca El Salto, pine-oak forest, 28 21'18"N 109'01'48" W, 2000 m elevation, locally common, 14 Jul
1997, Van Devender 97-698 (ARIZ, WS), with A.L. Reina G., P. Merlin, A. Burquez M., G. Ferguson, D.
Larson, M. Kaib, and MJ. Martinez C; Arroyo Hondo, 11.5 km E of El sie a oe W of Chihuahua
border on MEX 16, pine-oak forest, 28°26'30"N 108'32'30"W, 1460 m el n herba-
ceous perennial in water in rocks, 25 Sep 1997, Reina G. 97-1392 (WS), with TR. 2 Van Devences 2.3km
above eter Puerto de la Cruz on road to microwave tower, Mesa del Campanero, ie oak for-
est, 28'22'118"N 109°01'54"W, ca. 2150 m elevation, uncommon herbaceous perennial on moist road-
side, 16 Aug — Reina G. 98-939 (WS), with T.R. Van Devender, mM . ee . Role and
G.M. Ferguson; Canada La Ventana (Arroyo El Otro Lado), ra, rocky stre
canyon in pine-oak forest, 28°21'38"N 108°53'55"W, 1520 m Aeasiion, solitary ne in moist see in
ow side canyon, 18 Sep 1998, Van Devender 98-1303 (ARIZ, WS), with A ee and W. Trauba;
énega de Camilo, 6.3 km E of El Kipor, 11.2 km W of oe nua border on MEX 16, ile ac bog
in eet in shady riparian pine-oak ae 28°46'43'N 108°31'50"W, 1520 mel olitary rush
in moist soil, 27 Sep 1998, Van Devender 98-1850 (WS), with re Reina G. (Van Devender et al. in
press).
Oo.
Juncus liebmanii J.-Macbr. var. polycephalus Balslev
This species is known from Chihuahua in Mexico south to Ecuador. The northern
Mexican collections are considered part of J. liebmanii var. liebmanii or var. poly-
cephalus Balslev (Balslev 1996). The Sonoran collection belongs to var. polycephalus.
This is the first report of this species for Sonora and the Rio Mayo region.
346 BRIT.ORG/SIDA 20(1)
Voucher specimens. MEXICO. Sonora. Municipio de Yécora: Arroyo La Cueva, 8 km N of Yécora on
the road to Agua Blanca, rocky stream canyon in pine-oak forest, 28'26'27'N 108°55'29"W, 1520 m
elevation, uncommon perennial in moist soil at edge of stream, 24 Sep 1998, Van Devender 98-1634
(WS), with A.L. Reina G. and W. Trauba.
ACKNOWLEDGMENTS
We thank Father Bill Trauba for help and companionship in the field and pro-
viding shelter in Yécora; George Ferguson, Mark Fishbein, Sam Friedman, and
Richard Spellenberg for help in the field and for sharing their collections; Miriam
C. Fritts for help with Carex identifications; and Richard S. Felger and M. Socorro
Gonzalez E. for helpful comments on a previous version of the manuscript.
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Agric. Agric. Handbook No. 46/7.
Krat, R. 1971. A treatment of Abildgaardia, Bulbostylis and Fimbristylis (Cyperaceae) for
North America. Sida 4:57-227.
Larerriere, J.E. 1994. Vegetation and flora of the Mountain Pima village of Nabogame, Chi-
huahua, Mexico. Phytologia 77:102-140.
Martin, P.S.,D. YETMAN, M. FisHBein, P. Jenkins, T.R. VAN Devenber, and R.K. Wilson. 1998.Gentry’s Rio
Mayo Plants. The University of Arizona Press, Tucson.
ROALSON ET AL ( 347
McVauch, R. 1993. Flora Novo-Galiciana. Volume 13. Limnocharitaceae to Typhaceae. The
University of Michigan Herbarium, Ann Arbor, Michigan.
Raynal,J.1976.Notes Cyperologiques: 27. Identification de deux Scleria de Poiret. Adanso-
nia, ser. 2,16:211-217.
Reina G., A.L., T.R. VAN Devenoer, W. Trausa, and A. Burquez M. 1999. Caminos de Yécora. Notes
on the vegetation and flora of Yécora, Sonora. In: D. Vasquez del Castillo, M. Ortega N.,
CA.Yocupicio C.,eds. Symposium Internacional sobre la Utilizacidn y Aprovechamiento
de la Flora Silvestre de Zonas Aridas, Universidad de Sonora, Hermosillo. Pp. 137-144.
Roatson, E.H. 1999. Eleocharis yecorensis (Cyperaceae),a new species of spike-sedge from
Mexico. Aliso 18:57-60.
SPELLENBERG, R.S., T. Leaaue, and R. Corrat D. 1996. A specimen-based, annotated checklist of
the plants of Parque Nacional “Cascada de Basaseachi" and immediately adjacent ar-
eas, southwestern Chihuahua, Mexico. Listados Floristicos de México, Instituto de
Biologia, Universidad Nacional Aut6noma de México, México, D.F
Stevens, W.D.,C. U. ULLoa, A. Poot, and O.M. Montiet (eds.). 2001. Flora de Nicaragua. Missouri
Botanical Garden Press, St. Louis.
Tucker, G.C. 1994. Revision of the Mexican species of Cyperus (Cyperaceae). Syst. Bot. Monogr.
43:1-213.
Van Devenoer, T.R., A.L. Reina G., M.C. Peniatea G., and C.l. Orteca R. In press. The Ciénega de
Camilo:a threatened habitat in the Sierra Madre Occidental of eastern Sonora, Mexico
Madrofno.
348 BRIT.ORG/SIDA 20(1)
Books RECEIVED/ NOTICES
Annotations and comments by the Editor
Texas Floras/Natural History/Wildlife
FREDERICK R. GEHLBACK. 2002. Messages from the Wild: An Almanac of Suburban
Natural and Unnatural History. (ISBN 0-292-72837-9, hbk.; 0-292-72838-
7, pbk.). University of Texas Press, PO. Box 7819, Austin, TX 78713-7819, 800-
252-3206, 800-687-6046 fax). $45.00 (hbk), $24.95 (pbk), 280 pp., 24 color
photos, 12 line drawings, 51/2" x 9"
animals migrations of birds,
Key Worps: Natural history, Texas, central Texas
plants, ecology, life cycles.
ALFRED RICHARDSON. 2002. Wildflowers and Other Plants of Texas Beaches and
Islands. (ISBN 0-292-77115-0, hbk; 0-292-7716-9, pbk.). University of Texas
Press, PO. Box 7819, Austin, TX 78713-7819, U.S.A. (Orders: 800-252-3206
fax; 512-471-4032). $65.00 (hbk), $29.95 (pbk), 271 pp., 316 color photos, 1
Map. t+ eo.
Key Worps: Texas, beaches, natural history, field guides, wildflowers, identification
Youll wish you had this book the next time you visit Padre Island for any reason. Richi srdson
provides an interesting guide to some 275 common and/or noteworthy flowering plants from the
hor |
Rio Grande to Louisiana.
RAYMOND C. TELFAIR IE (ed.). 1999. Texas Wildlife Resources and Land Uses. (ISBN
0-292-78159-8, pbk.). University of Texas Press, PO. Box 7819, Austin, TX
78713-7819, U.S.A. (Orders: 800-252-3206 fax; 512-471-4032). $26.95, 416
pp., 20 b/w photos, 39 tables, 6" x 9",
Key Worps: Texas, wildlife resources, land uses, environmental history, conservation, agricultural
w
sae and sustainability, water resources, ecological regions, plants, anima
If you need information at your fingertips on Texas Parks, National Parks, National Wildlife
deve
eee or National Forests and Grasslands, then you should have this book on your library shelf.
The handy book is divided into five major sections: Part 1) Perspectives on Texas Wildlife Resources;
Part 2 Future Expectations in Land Use; Part 3) The Public and Future Demands for Wildlife; Part
ildlite M 1 Research; and Part 3) Wildlife M Public Lands. Four great
appendices are aio ided: 1) Conservation Organizations in Texas; 2) Plants Cited in the text; 3) Ani-
mals Cited in the Text; and 4) Texas Threatened and Endangered Species, November oo.
SIDA 20(1): 348. 2002
LISTADO PRELIMINAR DE LAS CACTACEAS Y AGAVACEAS
DE LA SIERRA “EL VIEJO,” SONORA, REGION PRIORITARIA
PARA LA CONSERVACION EN MEXICO
Carlos G.Velazco Macias Glafiro J. Alanis Flores
Organizacion Vida Silvestre A.C. Facultad de Ciencias Bioldgicas
San Pedro Garza Garcia UA NL, San I Nicolds de los Garza
“XICO N.L,, MEXICO
RESUMEN
La ae 2 Migjo" se else en ae mata de Euneuite y Caborca, Sonora, en la porcion Centr
|
‘El Alamo”-Sierra “El Viejo” esta eee
oO
como una Region Prioritaria Te siesue pata la conservacion en México. Este trabajo tiene por objetivo
contribuir al conocimiento floristico de las familias Cactaceae y Agavaceae principalmente en la
Sierra “El Viejo.” Se hizo una revision BInoeran pane establecer las i bane que, ra se
encontrarian en el 4
r
localidades. Se registraron 2 | 4 Jictribuid 10
4 doc jecd
género | Agavaces as
en 2 géneros, lo que hace un total de 25 especies d teDinelas en = oe familias Las especies de
Cacrsceas ie ia un 21 % del total de |
I | ce hae dae
Lal nal |
—. son nendemicasa la Sierra “El Viejo" Agave ye A. cae Se sugiere continuar con aT
j ‘ ]
variedades y también estudios ecoldégicos en
}
gas especies, para establecer programas de manejo y proteccion de las poblaciones y el area en
conjun
ABSTRACT
Sierra “El Viejo” is located in the municipality of Hee ae Caborca, Sonora, in the west central
portion of the Sonoran Desert, the aes Sierra “El Alamo”-Sierra “El Viejo” is catalogued as a
terrestrial high-priority region for n ition in Mexico. The objective for this study is to
contribute to the floristic knowle sig af we ‘emilee Cactaceae and Agavaceae within this Se
mainly in the Sierra “El Viejo.” oe research was carried out and species reported w
corroborated in the field by hing
10 genera, and 4 species, and 2 genera of the Agavaceae family were recorded. A total of 25 species
are recorded in the two families. The total of species in the Cactus family represents 21 % of the total
ocalities in the area. Twenty-one species of cacti, in
species present in the state of Sonora. One species of cactus is protected by federal law, Ferocactus
cylindraceus. Two species of agaves, Agave pelona and A. zebra are endemic to the region. Further
floristic and ecological work is needed in the region that will support conservation, management
and protection of the natural resources of this particular region.
INTRODUCCION
Importancia
Las familias Cactaceae y Agavaceae cuentan con diversos estudios taxonomicos
y de distribucién en algunos estados y municipios de la republica mexicana.
No obstante existen estados en donde se tiene un pobre conocimiento en estas
SIDA 20(1): 349 — 354. 2002
350 BRIT.ORG/SIDA 20
1)
familias en lo que a taxonomia y distribucion se refiere. Dada la importancia
que estas familias tienen en el Ambito comercial (como plantas de ornato
altamente cotizadas, medicinales y de alimento), es necesario promover estudios
de estos grupos taxondmicos.
La sierra “El Viejo” esta catalogada como una region terrestre prioritaria
para la conservacion en México (Arriaga et al. 2000; RPT Clave 16). Bajo este
criterio, consideramos que para la region, existe un conocimiento de escaso a
regular en cuanto a inventarios de especies, por lo que el desarrollo de listados
floristicos y faunisticos es de suma importancia para la correcta planeacion en
acciones de conservacion y manejo del area. De igual forma, el mismo autor,
considera que la region se encuentra en un excelente estado de conservacion y
con una alta integridad ecoldégica, lo que permite una buena oportunidad para
levar con éxito planes de conservacion y de manejo.
—
Objetivos
l) Ampliar el conocimiento floristico y de distribucion para las especies de la
familia Cactaceae y Agavaceae en la Sierra “El Viejo.”
2) Aportar algunos datos sobre su etnobotanica e importancia ecologica.
Area de estudio
La sierra “El Viejo” se encuentra ubicada en los municipios de Caborca y
Pitiquito, Sonora, en el noroeste de la republica Mexicana (Fig. 1). Ocupa una
extension aproximada de 1,128 km?, constituida principalmente por lomerios
y elevaciones montanosas que van desde los 200 a los 1,000 msnm. El clima es
desértico o muy seco semicalido BWhw (x’)(e’). El ciclo de Iuvias es bianual y
se presenta en verano y otono e invierno. La vegetacion de la planicie
corresponde al matorral desértico microfilo y en la sierra la vegetacion
corresponde al matorral microfilo sarcocaule.
METODO
Durante los meses de enero a diciembre de 1999 se realizaron recorridos por
diferentes localidades de la Sierra “EL Viejo” y sus planicies aledanas. Se
corroboraron los es existentes en la literatura y la taxonomia de las
t ‘uentran en esta area. La identificacion
iascenies se hizo nate los criterios de Bravo-Hollis (1978), Bravo-Hollis
y Sanchez-Mejorada (1991) y algunas modificaciones segtin Anderson (2001).
RESULTADOS
Se encontraron un total de 10 géneros y 21 especies para la familia Cactaceae y
dos géneros con cuatro especies para la familia Agavaceae (Tabla 1 y 2). Las
especies de la familia Cactaceae representan aproximadamente el 21 % del total
registrado para el estado de Sonora.
VELAZCO AND ALANIS, CACTACEAS Y AGAVACEAS 351
Estados Unidos de América
Area de
estudio Mn
be %@ chihuahua
Océano Pacifico
. “~*
4°
Sinaloa
i,
P| Areas Peerage & en Ne
noroeste de MEex
(9) compiejo Sierra El Alamo-el Viejo
WW ats
eee er ree ere ere 7 : re F ee
1 AAdyi (M dite Ind CONABIO, 2000)
DISCUSIONES Y CONCLUSIONES
Si bien la sierra “El Viejo” ha sido explorada por diversos investigadores (Gen-
try 1951, 1966; Turner 1980, 1983; Dimmitt 1983; Yatskievych y Fischer 1981 y
1982) es considera por Arraiga (2000), como un area que no ha sido revisada de
una manera detallada. El conocimiento botanico en esta area podria aumentar
enormemente, teniendo como ejemplo que Yatskievych y Fischer (1984)
encontraron los siguientes nuevos registros de la familia Cactaceae:
Ancisirocatus uncinatus, Echinocactus horizontalonius, var. nicholii,
Echinocereus scopulorum y Mammillaria lasiacantha, de estas especies se ha
podido corroborar la presencia de todas ellas excepto de E. horizontalonius var.
nicholii, esta especie fue encontrada originalmente en la porcion norte de la
sierra, consistiendo su colecta en un solo ejemplar. Al examinar ejemplares de
Mammilaria lasiacntha, se tiene cierta reserva al definir esta especie, debido
352
TABLA 1.Especies de Cactaceas en la Sierra “El Viejo.”
BRIT.ORG/SIDA 20(1)
Nombre Cientifico
Nombre Comun
Carnegiea gigantea (Engelm.) Britton & Rose 1908
Echinocereus fendleri (Engelm.) F. Seitz 1870
Echinocereus nicholii (L.D. Benson) B.D. Parfitt 1987
Echinocereus scopulorum Britton & Rose 1922
Ferocactus covillei Britton & Rose 1922
Ferocactus decd indraceus (Engelm.) Orcutt 1926
Hamatocactus uncinatus (Galeotti ) Orcutt 1926
a cee schottil (Engelm.) Britton & Rose 1909
Sahuaro
Cholla
Cholla barbona
Cabecita de viejo
Biznaga
Biznaga
Biznaga de anzuelos
Cinita, cina, tuna barbona
Mammillaria grahamii Engelm. 1856
Mammillaria thornberi Orcutt 1902
Mammillaria sp. (aff. lashiacantha)
Cabecita de viejo
Cabecita de viejo
Cabecita de viejo
Opuntia acanthocarpa Engelm. & Bigelow 1856 Sibiris
Opuntia arbuscula Engelm. 1856 Cholla
Opuntia bigelovii Engelm. 1856
Cholla brincadora, velas de coyote
Opuntia engelmannii Salm-Dyck ex Engelm. 1850 Nopal
Opuntia fulgida Engelm. 1856 Cholla plateada
Opuntia leptocaulis A. DC. 1828 Tasajillo o sibirito
Opuntia macrocentra Engelm. 18
Pachycereus pringlei (S.Watson) Britton & Rose 1909
Peniocereus striatus (T. Brandegee) Buxbaum 1975 Sacamatraca
Stenocereus thurberi (Engelm.) Buxbaum 1961 Pitahaya
TABLA 2.Especies de Agavaceas en la Sierra “El Viejo
Nombre Cientifico Nombre Comun Usos Tradicionales
Extraccion de fibras
Usos medicinales y cosméticos
Fabricacion de
Uso alimenticio
gdve pelona Gentry Lechuguilla
fear schotii Engelm.
Agave zebra Gentr aguey
Yucca arizonica McKelvey Jobichi, palma
bebidas alcohdlicas
existen diferencias en el color de la flor y la espinacion, por lo cual actualmente
se encuentran en revision para determinar su categoria taxondémica de una
manera correcta.
Comparativamente, la Sierra El Viejo se puede considerar rica en especies
de cactaceas, ya que Felger (1992), al realizar una sinopsis de las plantas
vasculares del noroeste de Sonora, aproximadamente 15,000 km2 (incluyendo
la reserva de la biosfera “El Pinacate”), registra 10 géneros, 26 especies y 7
variedades (reconocidas por Anderson 2001). En la Sierra “El Viejo,” area 13 veces
menor, se senalan igual numero de géneros y cinco especies menos; para la fa-
milia Agavaceae solo existen 2 géneros con una especie cada uno.
VELAZCO AND ALANIS, CACTACEAS ¥ AGAVACEAS 353
Hernandez (1998) caracterizo la vegetacion en el area de la Sierra “El Viejo,”
y registro nueve cactaceas, ademas de Agave sp. y Yucca sp. sin profundizar en
su categoria taxondmica. El presente estudio arroja mas del doble de especies
de cactaceas y determina las especies de agaves mediante la revision de literatura
ya existente. Cabe destacar que dos especies (Agave pelona y A. zebra) son
endémicas de la Sierra «El Viejo» y por lo menos A. pelona es parte importante
de la dieta y fuente de agua para el borrego cimarron (Ovis canadensis mexicana)
que habita la region. Precisamente es con el borrego cimarrén, con quien se
observ6 un fenomeno interesante de interaccion entre fauna y flora, durante la
época de floracion de A. pelona, se da un alto consumo de los escapos florales de
esta especie por parte del borrego cimarron y a lo largo del ano se consumen
plantas enteras para satisfacer las necesidades tanto alimenticias como hidricas.
Cabe destacar que del total de las especies de cactaceas solo una, Ferocactus
cylindraceus, esta dentro de la norma NOM-059-ECOL-1994 (Diario Oficial de
la Federacion 1994), bajo la categoria de rara y que ninguna de las especies de
agavaceas esta en alguna categoria de dicha norma, a pesar de que dos de ellas
son endémicas del area.
Historicamente y de acuerdo con los pobladores locales, se ha realizado
extraccion de Agave pelona y A. zebra. El primero para la extraccion de fibras
vegetales y el segundo para la elaboracion, a pequena escala, de una bebida
alcohélica. A pesar de dicho aprovechamiento (realizado en la primera mitad
del siglo XX), las poblaciones se encuentran en un buen estado, ya que se
observan individuos de Agave de todos los tamanos y clases de edades.
La Sierra “El Viejo,” area prioritaria para la conservacion en México,
mantiene dentro de sus limites un total de 21 especies de cactaceas distribuidas
en diez géneros, siendo Opuntia el género (7 spp.) mas diverso, seguido por
Echinocereus y Mammillaria G spp.). Esto representa aproximadamente un 21
% del total de especies registradas para el estado de Sonora (Paredes et al. 2000).
Las especies de la familia Agavaceae estan representadas por el género Agave (3
spp.) y el genero Yucca (1 sp.).
AGRADECIMIENTOS
Nuestro mas sincero agradecimiento a la Biol. Liliana Ramirez Freire y Erika
Gerez Hernandez por sus atinados comentarios y correcciones al manuscrito.
A Reinaldo Gutiérrez, Jess Gutiérrez y Martin Armenta que cooperaron en los
trabajos de campo. Organizacion Vida Silvestre A.C. por las facilidades
brindadas.
REFERENCIAS
Anoerson, E.F.2001.The cactus family. Timber Press, Portland, OR.
ARrRIAGA, L., J.M. Espinoza, C. AGUILAR, E. MARTINEZ, L.GOmez, y E.Loa (coordinadores). 2000. Regiones
BRIT.ORG/SIDA 20(1)
terrestres prioritarias de México. Comisi6n Nacional para el Conocimiento y uso de la
Biodiversidad, México.
Bravo, H.H.1978.Las cactaceas de México. Vol.1.Universidad Nacional Aut6énoma de México,
México.
Bravo, H.H. y H. SANCHEZ-Mesoraba. 1991. Las cactaceas de México. Vols. 2 y 3. Universidad
Nacional Aut6noma de México, México
DIARIO OFICIAL DE LA FEDERACION. 1994. Norma Oficial Mexicana NOM-ECOL-059. Secretaria de
Desarrollo Social. Tomo CDLXXXVIII. No. 10. Mayo 16 de 1994, México.
FeLGER, R.S.1992. Synopsis of the vascular plants of north tern Sonora, México. Ecologica
2(2):11-44.
HERNANDEZ, C.M.C. 1998. Caracterizacion de la vegetacion del predio”El Plomito,” municipio
de Pitiquito, Sonora, México. Tesis inédita. Facultad de Ciencias Bioldgicas, Universidad
Autonoma de Nuevo Ledn, México
Parepes A, T.R. VAN Devenver, y R.S. Fetcer. 2000. Cactaceas de Sonora, México: su diversidad,
uso y conservacion. IMADES-ASDM Press, Canada.
Turner, R.M., J.E. Bowers y T.L. Buraess. 1995. Sonoran Desert plants: an ecological atlas. The
University of Arizona Press, Tucson.
YATSKIEVYCH, G.y P.-C. FiscHer. 1984. New plant records from the Sonoran Desert. Desert Plants
5:180-190.
VASCULAR FLORA OF
BIG LAKE BOTTOM WILDLIFE MANAGEMENT AREA,
ANDERSON COUNTY, TEXAS
Kay M. Fleming
Wildlife Division
Texas Parks and Wildlife Department
9 E. Clinton
Athens, Texas 75751-2908, U.S.A.
eming@tvec.net
Jason R. Singhurst Walter C. Holmes
Wildlife Diversity Program Department of Biology
Texas Parks and Wildlife De epdariment Baylor University
3000 South IH- re 100 Waco, TX 76798-7388, U.S.A.
Austin, 1X 78704, U.S.A. walter_holmes@baylor.edu
Jason.si ae state. tx.us
ABSTRACT
A floristic pene! of Big Lake Bottom Wildlife Management Area, a bottomland hardwood forest
]
located on the Trinity River approximately 16 km SW of Palestine in Anderson Co., Texas, was con-
ducted from ee 1997 ‘july 2001. The study area is characterized by eight distinct alliances: LD)
— Oak-Water Hickory; 2) Willow Oak; 3) Planer Tree; 4) Cottonwood; 5) Sugarberry-Cedar
Elm; 6) Bur Oak-Shumard Oak: 7) Post Oak-Blackjack Oak; and 8) Sand Post Oak-Bluejack Oak. The
annotated list of taxa includes 99 families, 298 genera, and 459 species.
RESUMEN
Se realizo un inventario floristico del Big Lake Bottom Wildlife Management Area, un bosque de
madera dura localizado en el rio Trinity unos 16 km al SW de Palestine en el Anderson Co., Texas,
desde octubre . 1997 a julio de 2001. El area de estudio se caracteriza por ocho re aie distintas: 1)
Overcup Oak-Water Hickory; 2) Willow Oak; 3) Planer Tree; 4) Cottonwood; 5) Sugarberry-Cedar
Elm; a Ole cnet Oak; 7) Post Oak-Blackjack Oak; y 8) Sand Post Oak- Bluejack Oak. El
catalogo foaie de taxa incluye 99 familias, 298 géneros, y 459 especies
INTRODUCTION
The Texas Parks and Wildlife Department began purchasing bottomland in 1991
in an effort to preserve this unique and rapidly disappearing hardwood habitat.
Among the first acquisitions was the area now known as Big Lake Bottom Wild-
life Management Area (BLBWMA), one of the largest remaining bottomland
hardwood tracts of its type in the central Trinity River basin. The area consists
of approximately 1684.8 ha that lie within the transition zone between the
Pineywoods and the Post Oak Savannah vegetational areas of eastern Texas
(Gould 1962), about 16 km SW of Palestine. This alluvial bottomland was formed
SIDA 20(1): 355 — 371. 2002
356 BRIT.ORG/SIDA 20(1)
by the periodic erosion and sedimentation of the river as it flowed through the
region. The poorly draining soils and uniform topography produced a forest
dominated by hydrophytes.
Because of its unsuitability for most agricultural pursuits, the management
area has remained relatively free from outside disturbances. Two of the past
major disturbances were selective harvesting of timber in the late 1930s and
early 1940s and the clearing of about 65 ha for cultivation in the 1950s. Current
use of the property is limited to human recreational activities. Past and present
water control and agricultural practices have generally widened the Trinity
River, resulting in severe erosion problems along sloughs and creeks. This ero-
sion problem has been compounded by the use of all-terrain vehicles on the
management area.
Since the management area was purchased to preserve the bottomland
habitat along the Trinity River, this systematic inventory of the vascular flora
is imperative for proper management. Results of the study will be used by re-
source managers in decisions concerning activities that might affect the plant
communities and ecosystems of the area. These activities include future land
purchases or trades, mineral exploration, granting of rights-of-way and ease-
ments, and public use.
Field work was conducted primarily from October 1997 to October 1999, with
occasional visits made until July 2001 to search for taxa expected to occur but
not previously collected. The survey included more than 50 field visits that pro-
duced 532 field collections including ferns, gymnosperms, and [lowering plants.
Voucher specimens are deposited in the Baylor University Herbarium (BAYLU).
DESCRIPTION OF STUDY AREA
BLBWMaA is located within the humid subtropical zone. Temperatures rarely
go below -12°C in winter or above 38°C in summer, with the mean temperature
being near 20° C (Hatch et al. 1990). The soil of the study area is predominately
composed of alluvium and fluviatile terrace deposits and a small area of car-
bonaceous clay, silt, and ironstone on the Recklaw Formation (University of
Texas Bureau of Economic Geology-Palestine Sheet 1993). The deep clay bot-
tomland soils are classified as part of the Kaufman-Trinity Association while
the more loamy and sandy uplands belong to the Axtell-Lufkin Association
(Coffee 1970). Elevation varies from 57.9-82.3 m above sea level.
Nixon and Willett (1974), in the only previous study of the site located,
described 5 bottomland hardwoods types. These were described as coalitions
of species and not afforded names as such. Many species were included in more
than one type, apparently based upon abundance. These types actually over-
lapped to some extent, thus we find it advantageous to use the alliance method
of community description.
FLEMING ET AL., FLORA OF BIG LAKE BOTTOM WILDLIFE MANAGEMENT AREA 357
The BLBWMA vegetation areas can be segregated into eight alliances (Na-
tional Vegetation Classification System (NVCS) 1997) based on dominant spe-
cies, landscape position, and soil water content. The location, characteristic
species, and a short description of each alliance follow.
Overcup Oak (Water Hickory) Seasonally Flooded Forest Alliance
The Overcup Oak (Water Hickory) Alliance is characterized by temporary
standing water during and after flood events and consists of approximately
818.3 ha. This alliance is found throughout the site on heavy clay soils (Fig. LD.
Dominant trees are Quercus lyrata, Carya aquatica, Celtis laevigata var. laevigata
and Ulmus americana. Other common woody species in this area include
Fraxinus pennsylvanica, Ilex decidua, and Crataegus viridis. Common herba-
ceous species include Iva annua, Xanthium strumarium, Justicia ovata, Leersia
lenticularis, Saururus cernuus, Lobelia cardinalis, Diodia virginiana, Gratiola
virginiana, and Boehmeria cylindrica.
Willow Oak Seasonally Flooded Forest Alliance
Willow Oak Alliance consists of seasonally flooded areas and is the smallest
wet bottomland feature on the area at 50.2 ha. These are depressed areas of pe-
riodic inundation in which hardwood trees dominate and form a dense canopy.
This alliance is located north of the drainage channel of Big Lake into Keechie
Creek, an area that has been carved out by flooding events (Fig. 1). Common
tree species include Quercus phellos, Q. nigra, Celtis laevigata var. laevigata,
Carya aquatica, Nyssa sylvatica, Ilex decidua, and Liquidambar styraciflua.
Characteristic herbs of this forest include Carex frankii, Juncus coriaceus, and
Trachelospermum difforme.
—
Planer Tree Seasonally Flooded Forest Alliance
Planer Tree Alliance comprises 85.8 ha and consists of mid-story forests in
backswamps and sloughs that are produced in soils created by rapid siltation.
This alliance occurs along the corridor of Cedar Creek Slough and along Keechie
Creek Slough just south of where Keechie Creek joins the Trinity River, as well
as in a small section of the Johnson Tract (Fig. 1). Common woody plants in-
clude Planera aquatica, Gleditsia aquatica, Forestiera acuminata, Carya
aquatica, Cephalanthus occidentalis, Fraxinus pennsylvanica, Salix nigra, and
Berchemia scandens. Herbaceous plants include Saururus cernuus, Boehmeria
cylindica, Sesbania drummondii, and S. exaltata.
Cottonwood Temporarily Flooded Forest Alliance
Cottonwood Alliance consists of 10.9 ha of periodically flooded forests on sandy
alluvium primarily on developed river fronts. It is restricted to a narrow strip
adjacent to the Trinity River bank (Fig. 1). Common tree and shrub species in-
clude Populus deltoides, Salix nigra, Acer negundo, Gleditsia aquatica, Celtis
laevigata var. laevigata, Betula nigra, Ulmus americana, Morus rubra, Cornus
358 BRIT.ORG/SIDA 20(1)
ANDERSON COUNTY
Sand Post Oak-Bluejack Oak alliance
290"
Cedar Creek Slough
oP
Johnson Tract
Trinity River aay
F eZ Keechie Creek
Slough
ilk By dé
FREESTONE COUNTY
1 0 1 2 3 4 Kilometers
Vegetation Alliances
Overcup Oak - Water Hickory re Bur Oak-Shumard Oak
Willow Oak = - Post Oak-Blackjack Oak
Planer Tree —_ Sand Post Oak-Bluejack Oak
Fi A H Rin labo Ratt \HITAN 4, AA tA yy
1.A vicinity getation Alliances, Trinity River, and Big
Lake (ESRI Inc. ArcView ver. 3. 1).
drummondii, Symphoricarpos orbiculatus, and Forestiera acuminata. Herba-
ceous and woody vines are well represented. Common species include Mikania
scandens, Vitis riparia, V. palmata, Smilax bona-nox, S. tamnoides, Ampelopsis
arborea, loxicodendron radicans, Brunnichia ovata, Ipomoea cordatotriloba,and
I. lacunosa.
Sugarberry-Cedar Elm Temporarily Flooded Forest Alliance
Sugarberry-Cedar Elm Alliance consists of periodically flooded, forested, flat
areas with rich base soils. This alliance occupies 412.4 ha along Cedar Creek
Slough and along drainages of Keechie Creek (Fig. 1). Common trees include
Celtis laevigata var. laevigata, Ulmus crassifolia, U. alata, Gleditsia triacanthos,
Sapindus saponaria var. drummondii, Quercus macrocarpa, Fraxinus
FLEMING ET AL., FLORA OF BIG LAKE BOTTOM WILDLIFE MANAGEMENT AREA 359
pennsylvanica, F americana, and Crataegus mollis. Herbaceous plants which
typify the area include Iva annua, Carex cherokeensis, C. crus-corvi, Panicum
anceps, and Leersia oryzoides.
Bur Oak-Shumard Oak Forest Alliance
Bur Oak-Shumard Oak Alliance occurs on bluffs and terraces with clay soils
and rich humus layers. This alliance also occurs on two bluff areas that origi-
nated asa result of severe changes in river direction (Fig. 1). The area occupies
178.9 ha. This alliance was not formally described as occurring in Texas by the
NVCS (1997) or as a series level community by Diamond et al. (1987) and conse-
quently is mentioned here as occurring in Texas for the first time. Common
trees and shrubs include Quercus macrocarpa var. macrocarpa, Q. shumardii, Q.
stellata, Sideroxylon lanuginosum, Sophora affinis, Juniperus virginiana,
Crataegus viridis var. viridis, Celtis laevigata var. laevigata, Ilex decidua, and
Cercis canadensis. Herbaceous plants which typify the area include Carex
cherokeensis, Scleria triglomerata, Chasmanthium latifolium, Poa autumnalis,
Tradescantia ohiensis, Arisaema dracontium, Podophyllum peltatum, Dioscorea
quaternata, Clematis pitcheri, Urtica chamaedryoides, Stachys crenata, S.
tenuifolia, Ruellia nudiflora var. runyonii, Elephantopus carolinianus, Matelea
gonocarpos, Myosotis macrosperma, Scutellaria cardiophylla, Euphorbia
spathulata, Galium circaezans, Oxalis violacea, and Parietaria pensylvanica.
—
Post Oak-Blackjack Oak Woodland Alliance
Post Oak-Blackjack Oak Woodland Alliance comprises 21.5 ha and consists of
upland wooded areas on gentle sandy-loam slopes and abrupt deep sand ter-
races. This alliance is very species rich and found on the small Johnson Tract
and near entrances at each of the area gates (Fig. 1). Common trees and shrubs
include Quercus stellata, Q. marilandica, Q. incana, Q. falcata, Q. nigra, Q. phellos,
Carya texana, Juniperus virginiana, Diospyros virginiana, Vaccinium arboreum,
Cercis canadensis, Ulmus alata, Fraxinus americana, Callicarpa americana,
Crataegus crus-galli, C. marshalli, Prunus angustifolia var. angustifolia, P.
mexicana, P. serotina var. serotina, Rubus trivialis, Rhus copallina, and
Symphoricarpos orbiculatus. Herbaceous species include Chasmanthium
sessiliflorum, Elymus virginicus, Eragrostis secundiflora, Commelina erecta, Tra-
descantia hirsutiflora, Vitis mustangensis, V. rotundifolia, Sanicula canadensis,
Symphyotrichum patens, Podophyllum peltatum, Hypericum hypericoides, Hy-
pericum drummondii, Passiflora incarnata,and Geum canadense.
Sand Post Oak-Bluejack Oak Woodland Alliance
This alliance occurs in deep sandy soils ona ridge above the floodplain. It con-
sists of less than 5 ha (Fig. 1) located on the east side of Big Lake. Common tree
species include Quercus margarettiae, Q. marilandica, Q. incana, and Carya
texana. Other woody plants are Opuntia stricta, Yucca louisianensis, Rhus
copallina var. latifolia, Cornus florida, Vaccinium arboreum, Sassafras albidum,
360 BRIT.ORG/SIDA 20(1)
and Prunus gracilis. Herbaceous plants include Tradescantia hirsutiflora,
Matelea cynanchoides, Eragrotis secundiflora, Loeflingia squarrosa,
Helianthemum rosmarinifolium, Lechea tenuifolia, Cnidoscolus texanus, and
Dalea villosa.
FLORISTICS
The BLBWMA flora includes 459 species in 298 genera in 99 families. Average
number of species per genus is approximately 1.5. Families with the largest
number of species (in parentheses) are Poaceae (55), Asteraceae (54), Fabaceae
(32) and Cyperaceae (25). Other families with a relatively large number of spe-
cies are Lamiaceae (14), Apiaceae (12), Euphorbiaceae (12), Rosaceae (12), and
Scrophulariaceae (10). Genera with the most species include Carex (12), Quercus
(9), Panicum (8), Cyperus (7), Smilax (6), and Juncus (5). The number of non-
native species is 23, which is about five percent of the total flora.
Three plants collected on BLBWMA are considered threatened by the Texas
Parks and Wildlife Department and The Nature Conservancy of Texas (Carr
2001). Brazoria truncata var. pulcherrima (B. pulcherrima Lundell) is consid-
ered sensitive and ranked G3 (globally) and S3 (state). Both it and Yucca
louisianensis are also notable for being included in Estill and Cruzan’s (2001)
list of narrow endemics of southeastern United States. Curcurbita texana is
also ranked G3 (globally) and S3 (state). The species was historically widely
distributed along major rivers in the eastern half of Texas but much of its habi-
tat has been lost to impoundments, agriculture, and urban development. An
accurate assessment of its current status is difficult because of its ephemeral
nature. Cyperus grayoides is ranked G3 (globally) and $3 (state).
Other noteworthy plants were new to Anderson County. Correll and
Johnston (1970) report Dioscorea quaternata as mainly occurring in more north-
eastern Texas. The BLBWMaA record is near the most southwestern limit of the
species distribution. Symphyotrichum eulae, a Texas endemic, also occurs in
the area. The distribution of this species is primarily in east-central Texas, where
it is considered rare or uncommon. Rhynchosia minima var. minima is known
from the coastal plain and northward to Newton, Houston, Travis, Bexar, Rob-
inson, and Grimes counties. The present record is the northernmost record for
the species within the state. Heliotropium procumbens is mostly a coastal (and
western) species known to occur northward to Liberty, Waller,and Brazos coun-
ties. The new record extends the known distribution within the state farther
northward. Finally, Phyllanthus pudens has been recorded in Texas as far north
as Brazos, Madison, and Robinson counties. The Anderson County record ex-
tends the distribution farther within the state and provides a new county record.
The area contains the only known occurrence of Bur Oak-Shumard Oak
Forest alliance within the state (see above).
FLEMING ET AL., FLORA OF BIG LAKE BOTTOM WILDLIFE MANAGEMENT AREA 361
ANNOTATED LIST OF VASCULAR PLANT TAXA
BIG LAKE BOTTOM WILDLIFE MANAGEMENT AREA, ANDERSON COUNTY,
TEXAS
The annotated checklist is divided into pteridophytes, gymnosperms, and flow-
ering plants, which are subdivided into monocots and dicots. Family, genus,
and species are arranged alphabetically beneath each major heading. Nomen-
clature follows that of Correll and Johnston (1970), Correll and Correll (1972),
and Gould (1975). As needed, names were updated using Jones et al. (1997). Com-
mon names generally follow those of Hatch et al. (1990). Collector and collec-
tion numbers are given as: KF = Kay Fleming or JS = Jason Singhurst followed by
a number. An asterisk (*) denotes an introduced species.
PTERIDOPHYTA
ASPLENIACEAE
Asplenium platyneuron (L.) B.S.P,, Ebony spleen-
wort, JS 6167
DRYOPTERIDACEAE
Woodsia obtusa (Spreng.) Torr. Blunt-lobed cliff
fern, JS 6166
MARSILEACEAE
Hook. & Grev., Hooked water clo-
ver, 1 KF 369.
OPHIOGLOSSACEAE
Botrychium biternatum era Underw.,
Cutleaved grapefern, KF 3
Botrychium dissectum Spreng,., ae fern,
KF 456.
Ophioglossum vulgatum L., Common adder's
tongue, KF 82
POLYPODIACEAE
Pleopeltis polypodioides (L.) Andr
var. michauxiana (Weath.) Andrews &
Windham, Resurrection fern, JS 6320.
GYMNOSPERMAE
CUPRESSACEAE
Juniperus virginiana L., Eastern red cedar, KF 548.
MONOCOTYLEDONEAE
AGAVACEAE
Yucca louisianensis Trel., Louisiana yucca, KF 341.
ALISMATACEAE
Echinodorus berteroi (Spreng.) Fassett, Beaked
burwee !
Echinodorus cordifolius (L.) Griseb., Heartleaf bur-
wee
< & Windham
Sagittaria latifolia Willd., Duck potato, KF 478.
ARACEAE
Arisa dracontium (L.) Schott, Green dragon,
JS 6693.
ARECACEAE
Sabal minor (Jacq.) Pers., Dwarf palmetto, KF 486.
BROMILIACEAE
landsia usneoides (L.) L., Spanish moss, JS 6681,
COMMEINATERE
a Burm., Spreading dayflower,
KE 633.
Commelina erecta L., Erect dayflower, KF 335.
Tradescantia hirsutiflora Bush, Hairy spiderwort,
DD
Tradescantia ohiensis Raf., Ohio spiderwort, KF
239
CYPERACEAE
Carex amphibola Steud., Amphibious sedge, KF
473.
Carex ae ophora Muhl. ex Willd., Woodbank
sedge, KF 454.
Carex che
hwein., Cherokee sedge, JS
164.
Carex conan TOK & Hook., Sedge, KF 457.
| Kunze, Crowfoot
Care as Gh U 5 ¥
sedge, KF 271,
Carex flaccosperma Dewey, Thinfruit sedge, KF
472.
Carex frankii Kunth, Frank’s sedge, KF 452.
Carex lurida Wahl., Shallow sedge, KF 470.
Carex oxylepis Torr.& Hook., Sharpscale sedge, KF
46/7.
Carex retroflexa Muhl. ex Willd., Reflexed sedge,
4606,
362
Carex tetrastachya Scheele, Britton’s sedge, KF
46]
Carex texensis (Torr) Bailey, Texas sedge, KF 286.
Cyperus croceus Vahl, Baldwin flatsedge, KF 457.
Cyperus erythrorhizos Muhl., Redroot flatsedge,
KF 636
C\ neriic ordyi
lg =
ides Mohlenb., flatsedge, KF 675a.
Cyperus retrorsus Chapm., Backward flatsedge, KF
662
Cyperus strigosus L. var. strigosus, False nutsedge,
KF 678
Cyperus surinamensis Rottb., Tropical flatsedge,
Cyperus virens Michx. var. virens, Green flatgrass,
KF 386,
cars microcarpa Torr., Smallseed spikerush,
KF 317.
Eleocharis obtusa (Willd.) Schult. Blunt spikerush
KF 228.
Eleocharis palustris (L.) Roem.& Schult. Spikerush
Rhynchospora ea (Lam.) Gray, Horned
beakru
Scirpus we Muhl. small bulrush, JS 6774.
Scleria triglomerata Michx., Tall nutrush, KF 266.
DIOSCOREACEAE
Dioscorea quaternata (Walt.) Gmel., Atlantic yam,
IRIDACEAE
Iris brevicaulis Raf, Shortstem iris, KF 112
Sisyrinchium albidum Raf, White blue-eyed grass,
al
Sisyrinchium atlanticum Bicknell, Blue-eyed
Sisyrinchium langloisii Greene, Pale blue-eyed
JUNCACEAE
Juncus coriaceus Mack., Leather rush, KF 302.
Juncus effusus L. var. solutus Fern. & Weig., Com
rush, KF 305,
Juncus marginatus Rostk. var. setosus Cov., Bor-
der rush, KF 437.
Juncus scirpoides Lam., Strang rush, KF 407.
Juncus tenuis Willd., Slender rush, KF 474.
LILIACEAE
Allium canadense L. var. canadense, Canada gar
lic, KF 289.
Allium drummondii Regel, Drummond's anion, KF
2/5
BRIT.ORG/SIDA 20(1)
Cooperia drummondii Herb., Rain lily, KF 709.
Hymenocallis eulae Shinners, Late spider lily, KF 529
Hymenocallis liriosme (Raf.) Shinners, Spider lily,
—
—
Hypoxis hirsuta (L.) Cov., Common goldstar, KF 337.
Nothoscordum bivalve (L.) Britt, Crow poison, KF 224.
HIDACEAE
Spiranthes lacera (Raf.) Raf. var. gracilis
ae, ues ieee tresses, KF 212.
Spira 1(Small) Ames, Spring ladies’
tresses, KF 399.
POACEAE
Agrostis hyemalis (Walt.) B.S.P.,,Winter bentgrass,
KF 429,
(Bigel.)
Agrostis perennans (Walt.) Tuckerman, Autumn
Aira elegans Willd. ex Kunth, Annual hairgrass, KF
B53.
Alopecurus carolinianus Walt., Carolina foxtail, KF
461.
Andropogon ternarius Michx. var. ternarius,
Splitbeard bluestem, KF 653
virginicus L., Broomsedge bluestem,
A
WIChOPOGOr!
KF 425,
Arist oligantha Michx., Oldfield threeawn, KF
a purpurea Nutt., Red threeawn, KF 625.
Bothriochloa ischaemum (L.) Keng. var. songarica
(Fisch. & Mey.) Celerier & Harlan, King Ranch
bluestem, KF 691.
Briza minor L., Little Suis grass, KF 397.*
Bromus secalinus L., Rye brome, KF 380. *
Bromus catharticus Vahl ae KF 264.
Buchloe dactyloides (Nutt) ees
KF 390
Cenchrus spinifex A.Cav., Coastal sandbur, KF 705.
Chasmanthium latifolium (Michx.) Yates,
Longleaf uniola, KF 3617.
Chasmanthium sessiliflorum deh a Sessile
flowered nthiu
Cynodon ae ) pie var. a on, Ber-
muda grass, KF 537.*
Digitaria ciliaris (Retz.) Koel. Southern crabgrass,
KF 645.
Echinochloa walteri (Pursh) Heller,Walter’s barn-
yard grass, KF 686
Elymus canadensis L., Canads wilatye: AP 680.
Elymus oo us |
KF 6
eiual
VITO YE,
var. virginicus, \
FLEMING ET AL., FLORA OF BIG LAKE BOTTOM WILDLIFE MANAGEMENT AREA 363
Eragrostis hypnoides (Lam.) B.S.P, Teal lovegrass,
KF 639.
— sccowa (Michx.) Nees ex Steud.,
Carolina lovegrass, KF 6
eee eae Presl.subsp. oxylepis (Torr)
Red lovegrass, KF 398.
Eragrostis spec (Pursh) Steud., Purple
lovegrass, KF
oa. name (Michx.) B.S.P, Bearded
Snail 4 65].
Hordeu utt., Little barley, KF 269.
Leersia lenticularis ie cutgrass, KF 644.
Leersia oryzoides (L.) Sw. var. oryzoides, Rice
cutgrass, KF 649.
Leersia virginica Willd.,White cutgrass, KF 673.
Lolium perenne L., Perennial ryegrass, JS 6670. *
Panicum acuminatum Sw. var. acuminatum,
Woolly rosettegrass, KF 316.
Panicum nodatum & Chase, Sarita
rosettegrass, JS 536
Panicum oligosanthes i ‘Shult Rosettegrass, KF
424
Panicum anceps Michx.,Beaked panicum, KF 664.
Panicum dichotomum L.,Forked panicum, KF 659.
Panicum hemitomon Schult., Maiden cane, KF
Pani ii Scribn.& Merrill, Ravenel’s pani-
cum, KF 375.
Panicum gaan lee, eis KF 643.
|
Poir., Dallisgrass, KF 389.
Basal um floridanum ae Florida paspalum,
Paspalum notatum Flugge, Bahiagrass, KF oe be
Really urville ada _Vaseygrass, KF 687.
P, liniana Walt., Carolina canarygrass,
JS 6671.
Piptochaetium avenaceum (L.) Parodi, Blackseed
needlegrass
Poa annua L., Annual bluegrass, KF 505.*
Poa autumnalis Muhl. ex Ell, Autumn bluegrass,
KF 280.
Setaria parviflora (Poir.) Kergeulen, Knotroot
bristlegrass, KF 637
Sorghum halepense (L.) Pers., Jonnstongrass, KF
Tridens albescens (Vasey) Woot. & Standl., White
tridens, KF 642.
Tridens flavus (L.) Hitchc., Purpletop, KF 672.
Tridens strictus (Nutt.) Nash, Longspike tridens,
Tripsacum oe aa (L.) L., Eastern gamma-
9 rass, KF 958.
Triticum aes eat, KF 363.*
Vulpia octoflora (Walt, Rydb.,Common sixweeks
grass, KF 4
sind ania
ilax bona-nox L., Sawtooth greenbriar, KF 284.
a lax glauca Walt., Cat greenbriar, KF 663.
Smilax rotundifolia L., Common greenbriar, JS
6679
Smilax smallii Morong, Small's greenbriar, KF 6317.
Smilax tamnoides L., Bristly greenbriar, KF 283.
TYPHACEAE
Typha latifolia L., Common cattail, KF 478.
DICOTYLEDONEAE
ACANTHACEAE
Dicliptera brachiata (Pursh) Spreng., False mint,
Justicia ovata (Walt.) Lindau var. lanceolata
m.) Long, Lance-leaved water willow,
KF 209.
Ruellia nudiflora (Englem. & Gray) Urban var.
ii (Tharp. & Barkl.) B.L. Turner, Violet
ane 6315.
Ruellia pedunculata Torr. ex Gray, Low ruellia, KF
ACERACEAE
Acer negundo L. var. negundo, Box elder, KF 208.
MARANTHACEAE
Amaranthus albus L.,White amaranth, KF 606.
Amaranthus blitum L., Green amaranth, JS 631]
Amaranthus rudis Sauer, Amaranth, KF 607.
Froelichia drummondii Moq., Drummond's snake
542
ANACARDIACEAE
Rhus copallina L. var. latifolia H. Engler, Wing-rib
sumac, KF 276
Toxicodendron radicans (L.) O. Ktze. var. radicans,
Poison ivy, KF 310,
APIACEAE (UMBELLIFERAE)
Chaerophyllum tainturieri Hook., Chervil, JS 6169.
Cyclospermum leptophyllum (Pers.) al ex
Britt. & PWils., Slimlobe celery, KF 4
Cynosciadium digitatum DC., Finger ia
=
KF 372.
Daucus pusillus Michx., Wild carrot, KF 3817.
Eryngium hookeri Walp., Simple leaf eryngo, KF
48/
I lydh ocotyle verticillata Thun b., Whorled penny-
Limnosciadium ales) pe Math. & Const.,
as dogshade, 0.
Po ioe anuttallii DC., ine parsley, KF 272.
Sanicula canadens Canada sanicle, KF 412.
ee is giana: nee Raf. ex Ser., Forked
scaleseed, KF 355
Spermo
Seal
epis inermis (Natt ex DC) Math.& Const,
eseed, KF 2?
Trepocarpus oe Nutt.ex DC. Trepocarpus,
KF 460
APOCYNACEAE
Amsonia tllustris Woods., Bluestar, KF 247.
Trachelospermum difforme (Walt.) Gray, Climbing
dogbane, KF 420
AQUIFOLIACEAE
Ilex decidua Walt., Deciduous holly, JS 6136
llex opaca Ait. var.opaca, American holly, KF 519.
Ilex vomitoria Ait, Yaupon holly, KF 404.
ASCLEPIADACEAE
Asclepias amplexicaulis Smith, Bluntleaf milk-
ed, KF 37
Asclepias tuberosa L. subsp. interior Woods., But-
terfly milkweed, KF 349.
Asclepias viridis Walt., Antelopehorn milkweed,
KF 319.
Matelea Feo (Engelm.) Woods., Milk-
eed vine, KF 6
Matelea decipiens i Woods., Milkweed vine,
KF 667
Matelea gonocarpos (Walt.) Shinners, Milkweed
vine, KF 306
ASTERACEAE
Acmella oppositifolia (Lam.) Jansen var. repens
(Walt.) Jansen, Creeping spot flower, JS 6305.
i ia psilostachya DC.,Western ragweed, KF
een trifida L., elait ragweed, KF 533.
accharis halimifolia L., Eastern baccharis, KF 516
Bidens bipinnata L. var. 2 oe ides E. Seri
anish needles,
Boltonia diffusa ae var. | aes Small head
boltonia, KF 596
Brickellia eupatorioides (L.) Shinners, False bonset,
KF 668
BRIT.ORG/SIDA 20(1
Chrysopst is pl ilosa Nutt., Soft gold aster, KF 444.
Cirsium horridulum Michx. var. elliotti 7. & G., Bull
thistle, KF oe
Conyza canadensis (L.) Cronq., Horseweed
n 6/6
Conoclinium coelestinum (L.) DC., Blue mist
flower, KF 205
Coreopsis basalis (Dietr.) Blake, Golden coreop-
sis, KF 346
Coreopsis oe Nutt. var. tinctoria, Plains core-
opsis,
Croptilon divaric
KF 539
Dracopis amplexicaulis (Vahl) Cass., Dracopis, KF
366.
Echinacea pallida (Nutt.) Nutt., Purple cone
flower, KF 405
Eclipta prostrata (L.) L., Yerba de tago, KF 594.
Elephantopus Sri Raeusch., Leafy el-
ephan t foo ie
Erigeron ten Slender fleabane, KF 248.
Eupatorium ee ium Walt., Yankeeweed,
KF 623.
atum (Nutt.) Raf, Scratch daisy,
Eupatorium serotinum Michx., Late eupatorium,
KF 545
Facelis retusa (Lam.) Sch. Bip., Facelis, KF 307.
Fleischmannia incarnatum . King & Robin-
ink eupatorium, KF 6
bee purpurea (L. a Greenleaf
cudweed, KF 298.
Grindelia papposa G. Nesom & Y. Suh, saw-leaf
daisy, KF 1105
LS
Helenium amarum (Raf.) H. Rock, Bitter sneeze-
weed, KF 483.
Helianthus annuus L.,cCommon sunflower, KF 1002.
Helianthus debilis Nutt., Cucumber leaf sun-
flower, KF 378.
Heterotheca subaxillaris (Lam.) Britt.& Rusby, Yel-
low camphorweed, KF 234.
Hieracium gronovii L. Gronovius’ hawkweed, KF
411.
se aabenl del aaa L'Her., Flattop
oollywhite, KF 3
ec nnua L. Siiiewee an
Krigia cespi itosa (Raf.) ca a dwarf dan-
6177.
Lactuca fordenall (L.) Gaertn, Florida wild lettuce,
KF 628.
Liatris squarrosa (L.) Michx., Variable gayfeather,
F 523.
FLEMING ET AL., FLORA OF BIG LAKE BOTTOM WILDLIFE MANAGEMENT AREA 365
Mikania scandens (L.) Willd., Climbing hemp vine,
KF 495,
Packera tampicana (DC.) C. Jeffrey, Golden
groundsel, KF 743.
Pluchea camphorata (L.) DC., Camphor weed, KF
515
Pluchea purpurascens (Sw.) DC., Purple camphor
weed, KF 647
Pyrrhopappus carolinianus (Walt.) DC., Carolina
false dandelion, KF 468.
Rudbeckia hirta L., Late coneflower, KF 383.
Senecio ampullaceus Hook., Texas groundsel, JS
6180
Colid lavatal
is L., Tall goldenrod, KF 632.
a ieconean: Ait., Downy goldenrod, KF 665.
Solidago radula Nutt., Rough goldenrod, KF 627.
Solidago ulmifolia Muhl. ex Willd., Noble golden-
rod, KF 658.
liva pterosperma (Juss.) Less., Lawn burweed,
KF 501,
Symphyotrichum dumosum (L.) G.Nesom, Bushy
aster, KF 213.
Symphyotrichum eulae (Shinners) G. Nesom,
Simple aster, JS 6308.
ae ichum patens (Ait.) G.Nesom, Skydrop
KF 657.
tree ina encelioides (Cav.) Benth. & Hook. ex
ar. encelioides, Cawpen daisy, KF 203.
ee virginica L. var. virginica, Frostweed, KF
583.
Vernonia baldwinii Torr, Baldwin's ironweed, JS
6319.
Vernonia texana (Gray) Small, Texas ironweed, KF
463,
Xanthium strumarium L.,cCommon cocklebur, KF
ceuaaimabecine
eltatum L., May apple, KF 39.
BETULACEAE
Betula nigra L., River birch, KF 540.
BIGNONIACEAE
Campsis radicans (L.) Seeman ex Bureau, Trum-
pet creeper, KF 484
BORAGINACEAE
Heliotropium indicum L., Turnsole heliotrope, KF
200
umbens Mill. var. procumbens
Fourspike heliotrope, KF 688.
Heliotronium pro
b
Myosotis macrosperma Engelm., Spring forget-
me-not, KF 296
BRASSICACEAE
Cardamine parviflora L. var. arenicola (Britt.)
Schulz., Sand bittercress, KF 463.
Cardamine pensylvanica ly ex Willd., Pennsyl-
vania bittercress, JS 615
Lepedium virgincum L.,, on man's peppergrass,
KF 253
orippa palustris (L.) Besser, Bog bittercress, KF 417.
Rorippa sessiliflora (Nutt. ex T.& G.) Hitche., Stalk-
less yellowcress, JS 630
BUDDLEJACEAE
Polypremum procumbens L., Juniper leaf, KF 352.
CACTACEAE
Opuntia stricta (Haw.) Haw., Southern spineless
actus, KF 393
CALLITRICHACEAE
Callitriche ee Pursh oe heterophylla
water starwort, JS 617
ete Ae caE
Lobelia appendiculata var. appendiculata,
Earlobe lobelia, KF 3
Lobelia cardinalis L. ad flower, JS 6314.
bd is ass iata (L.) Nieuw. var. ee
ing Venus looking glass, KF
CAPRIFOLIACEAE
Lonicera japonica Thunb., Jaf
KF 329.*
=>
WIESE honey uckle,
Lonicera sempervirens L. var. sempervirens, Trum-
pe HERE YSUENE: KE 25/.
,Coral berr
KF 364.
Viburnum rufidulum Raf., Rusty blackhaw, KF 241.
CA
Arenaria serpyllifolia L.subsp. serpyllifolia, Thyme-
leaved sandwort, KF 502.
Cerastium brachypodum (Engelm. ex Gray) Rob-
ins., Shortstalk chickweed, JS 6162.
Cerastium elo Clustered flower
chickweed, JS 6
Loeflingia squarrosa ae subsp. squarrosa,
Spreading loeflingia, KF 435.
Minuartia drummondii (Shinners) McNeill,
Drummond's sandwort, KF 244
Minuartia ae (Michx.) Mattf, var. patula,
Pitcher sandwort, KF 312
366
Silene antirrhina L., Seedy catchfly, KF 382.
Stellaria media (L.) Vill. subsp. media, Common
chickweed, JS 6160.*
Sdsearad ceanscirdes
ides L. var. ambrosioides,
Woameeeds KF 54].
CISTACEAE
Helianthemum rosmarinifolium Pursh, Rosemary's
rockrose, KF 351
-hea mucronata Raf., Hairy pinweed, KF 402.
Lechea tenuifolia Michx., Narrowleaf pinweed, KF
CLUSIACEAE
Hypericum drummondii (Grev. & Hook.) T. & G
rummond's St. John's wort, KF 449.
Hypericum hypericoides (L.) Crantz, St. Andrew's
cross, KF 24
CONVOLVULACEAE
Dichondra carolinensis Michx., Ponyfoot, JS 6170.
Ipomoea lacunosa L., Pitted morning glory, KF
~~
pomoea cordatotriloba A. Dennstaedt, Sharp
pod morning glory, KF 605.
Cornus drummondii C. Mey., Roughleaf dog-
wood, KF 256.
Cornus florida L., Flowering dogwood, JS 6179.
Nyssa sylvatica Marsh., Black gum, KF 707.
CUCURBITACEAE
Cucumis melo L. subsp. melo, Muskmelon, KF
o92a
b
CCH COACH IC
Gray, Texas gourd, KF 589.
His pendula L., Drooping melonette, KF
964
CUSCUTACEAE
Cuscuta pentagona Engelm., Dodder, KF 543.
EBEN
Diospyros virginiana L., Common persimmon, KF
ERICACEAE
Marsh., Farkleberry, KF 406.
EUPHORBIACEAE
Acalypha gracilens Gray, Slender copperleaf, KF
661.
Cnidoscolus texanus (Mull. Arg.) Small, Texas bull
nettle, KF 318
BRIT.ORG/SIDA 20(1)
Croton ales ee , Woolly croton, KF ee
C. hern croton, KF 53
Croton nigh 1aUxil res ie
KF 555.
Croton monanthogynus Michx.,One seed croton,
KF 635.
es fa dentata Michx., Toothed spurge, KF
ea ia nutans Lag., Evebane spurge, KF 597.
Euphorbia prostrata Ait., Prostrate euphorbia, KF
690.
a ne a Spat ata alain, Warty spurge, KF 469.
Birdseed leaf flower,
Ds
Tragia brevispica Engelm. & Gray, Heartleaf
noseburn, JS 6685
FABACEAE
Amorpha fruticosa L., Indigo bush amorpha, KF
44],
Astragalus crassicarpus Nutt., Ground plum
milkvetch, KF 235.
Baptisia nuttalliana Small, Nuttall’s wild indigo,
KF 342.
Centrosema virginianum (L.) Benth., Butterfly pea,
Cercl canader ; L.
ar. canadensis, Eastern red-
Chamaecrista fasciculata (Michx.) Greene, Par-
tridge pea, KF 477.
Clitoria mariana L., Atlantic pigeon wings, KF 967.
Dalea villosa (Nutt.) Spreng., Prairie clover, KF 350.
Desmanthus illinoensis (Michx.) MacM. ex Rob-
ins & Fern., Illinois bundle flower, KF 675.
Bere eae a (Nutt.) DC., Smooth
's ticks, KF 217.
Desmodium seg (L.
beqagar’s t
Desmodium sessi lifol ium DC, Sessileleaf beqgar’s
ticks, KF 1033
Galactia regularis (L.) B.S.P, Downy milkpea, KF
656,
) DC., Panicled
ST aquatica Marsh.,Water honey locust, KF
517.
Gleditsia triacanthos L., Common honey locust,
FIOI/.
Glottidium vesicaria Jacq.) R.M. Harper, Bagpod,
KF 604.
Kummerowia striata (Thunb.) Hook. & Arn., Japa-
nese bush clover, KF 626.
FLEMING ET AL., FLORA OF BIG LAKE BOTTOM WILDLIFE MANAGEMENT AREA 367
Lathyrus pusillus Ell., Low peavine, KF 262.
Lespedeza stuevei Nutt., Tall bush clover, KF 666.
edicago minima (L.) L., Small bur clover, KF 238.
Mimosa nuttallii (DC.) B.L. Turner, Sensitive briar,
KF 332
Neptunia lutea (Leavenw.) Benth., Yellow
unia, KF 105:
Pediomelum digitatum (T. & G,) Isely, Palmleaf
scurfpea, KF 408.
Rhynchosia minima (L.) DC. var. minima, Least
Senna occidentalis (L.) Link, Coffee senna, KF 1022.
Sesbania drummondii (Rydb.) Cory, Rattlebush,
KF 526.
Sesbania exaltata (Raf.) Rydb. ex A. Hill, Hemp
sesbania, KF 585
Sophora affinis T. & G., Texas sophora, KF 309.
Strophostyles helvula (L.) Ell. Trailing wild bean,
Trifolium carolinianum Michx.,Carolina clover, KF 23/7.
Trifolium vesiculosum Savi var. vesiculosum,
rowleaf clover, KF 340
Vicia ludoviciana Nutt., Deer pea, KF 297.
FAGACEAE
Oue Siaila hx.,S uth rn red oak, KF 5173.
Quercus incana Bartr., Bluejack oak, KF 376.
Quercus lyrata Walt., Overcup oak, KF 507
Quercus macrocarpa Michx. var.macrocarpa, Bur
oak, KF 308
Quercus margarettiae Ashe ex small, Sand post
oak, KF 11175.
Quercus nigra L., Water oak, KF ft
Quercus phellos L., Willow oak, KF 323.
Quercus shumardii Buckl., nies oak, JS 6689.
Quercus stellata Wang., Post oak, KF 374.
SENTANACERE
tris Nutt., Prairie gentian, KF 394.
GERANIACEAE
Geranium carolinianum L.var.carolinianum, Caro-
lina geranium, KF 255.
HALORAGRACEAE
Myriophyllum heterophyllum Michx., Changeleaf
parrot’s feather, KF 432
Liquidambar styraciflua L., Sweetgum, KF 511
HYDROPHYLLACEAE
Hydrolea ovata Nutt. ex Choisy, Hairy hydrolea,
52 Is
Nama hispidum Gray, Rough nama, KF 43
Nemophila ae ides Nutt., Large flowered
nemophila, KF
Phacelia hirsuta ae Hairy phacelia, KF 256.
JUGLAN AE
Carya aquatica (Michx.) Nutt., Water hickory, KF
371.
Carya texana Buckl., Black hickory, KF 373.
Juglans nigra L., Black walnut, KF 268.
LAMIACEAE
Brazoria truncata (Benth.) Engelm. & Gray var.
pulcherrima (Lundell) M.W.Turner, Centerville
brazos mint, KF 379.
Hedeoma hispida Pursh, Marsh mermaidweed, KF
famnsm ample
v
icaule L., Henbit, KF 719. *
lycopus rubellus Moench, Water horehound, KF
Monarda fistulosa L., Wild horsemint, KF 480.
Monarda punctata L., Spotted horsemint, KF 357.
Physostegia intermedia (Nutt.) En al & Gray,
Intermediate dragonhead, KF 4
Physostegia virginiana (L.) Benth. via drag-
onhead, JS 6697
Salvia lyrata L., Lyreleaf sage, KF
ae aria nel laEngelm. : as Heartleaf
skullca
Pe ean hx., Small skullcap, KF 279.
Stachys crenata Raf., Shade betony, JS 6682.
Stachys tenuifolia Willd. subsp. tenu/folia,
Slenderleaf ed KF 629.
ree canadense L., g
—
der, KF
LAURACEAE
Sassafras albidum (Nutt.) Nees, Sassafras, KF 263.
LINACEAE
Linum sulcatum Riddell var. sulcatum, Grooved
flax, KF 395
LYTHRACEAE
nann Rottb., Purple ammania, KF
479
Lythrum alatum Pursh, Lanceleaf loosestrife, KF
1086
Hibiscus laevis All., Scarlet rosemallow, KF 359.
Hibiscus moscheutos L. subsp. lasiocarpos (Cav.)
Blach., Swamp rosemallow, KF 359.
—
br caroliniana (L.) G.Don, Carolina modiola,
Jd7.
me aan L., Diamond leaf oo KF 618.*
Sida spinosa L., Prickly mallow, KF 603.
MENISPERMACEAE
Cocculus carolinus (L.) DC., Carolina snailseed, /S
6684
MOLLUGINACEAE
Mollugo verticillata L., Thread carpetweed, KF 652.
MORACEAE
Maclura da (Raf.) Schneid., Osage orange,
KFS
Morus om Red mulberry, KF 370.
MYRICACEAE
Myrica cerifera L., Southern waxmyrtle, KF 258.
NYCTAGINACEAE
Mirabilis albida (Walt.) Heimerl, Four o'clock, Kf
464
OLEACEAE
Forestiera acuminata (Michx.) Poir,, Swamp privet,
225,
Fraxinus americana L., White ash, KF 609.
Red ash, JS 6683.
Ligustrum sinense Lour., Chinese privet, KF 717. *
ONAGRACEAE
Gaura longiflora Spach., Tall guara, KF 492.
Gaura parviflora Dougl. ex Lehm., Small fawies
gaura, KF 952.
[ ralt ifolia L
Fraxinus penns) Ivanica Mar sh.
L., Bushy seedbox, KF 4
Lud id aecunan 5 Walt., Winged pee -
ie iecgienail osa Walt., Creeping seedbox, KF
44
idvaole peploides (Kunth) Raven pale
glabrescens Raven, Smooth seed box, KF 422.
Oenothera heterophylla = Spach aise
ee Varileaf evening primrose, KF
3/4,
Srey laciniata Hill, Cutleaf evening prim-
503.
Pree erst speciosa Nutt., Showy evening prim-
OXALIDACEAE
Oxalis dillenii Jacq. subsp. dillenii, Dillen’s wood
orrel, KF 246
=~ un
Oxalis violacea L., Violet wood sorrel, KF 297
BRIT.ORG/SIDA 20(1)
PAPAVERACEAE
Argemone albiflora Hornem. var. texanad (G.
Owenby) Shinners, White prickly poppy, KF
PASSIFLORACEAE
Passiflora incarnata L., Maypop passion flower,
617.
Passiflora lutea L., Yellow passion flower, KF 634.
SAA han eoinids
Phytolocca americana L.
mon pokeweec - KF!
PLANTAGINACEAE
var. americana, Com-
14.
hil tage igi aL.,Paleseeded plantain, KF 497.
Dene., Wright's plantain, KF
384
POLYGALACEAE
Polygala polygama L. var. obtusa Chodat, Bitter
milkwort, KF 403
OLYGONACEAE
Brunnichia ovata (Walt.)
Shinners, Eardrop vine,
Polygonum 1 aviculare L., Prostrate knotweed, KF
514,
set hydropiper L., Water smartweed, JS
06.
ue 1 pensylvanicum (L.) Small, Pennsylva
nia smartweed, KF 5
Polygonum persicaria L. ladys tame: KF 692.
Rumex chrysocarpus Moris, Interupted dock, KF
388.
Rumex crispus L., Curly dock, KF 414
Rumex hastatulus Baldw., Meer WiniG dock, JS
618].
PORTULACACEAE
Ch yt ) ae |
ia virginica L.var. virginica, Spring beauty
231.
Portulaca umbraticola Kunth,Wingpod purslane,
eo
610
PRIMULACEAE
Samolus valerandi L. subsp. parviflorus
Hulten, Thinleaf brookweed, KF 361a
RANUNCULACEAE
. ee tcheri T.&G.,Pitcher’'s clematis, KF 348.
us L., Tiny mouse tail, KF 230.
Ranunc fie dus Michx. var. nitidus (Chapm.)
Duncan, Seaside buttercup, KF 233
a
Raf.)
Surus MIP TY
FLEMING ET AL., FLORA OF BIG LAKE BOTTOM WILDLIFE MANAGEMENT AREA 369
RHAMNACEAE
Berchemia scandens (Hill) K. Koch, Alabama
pplejack, KF 285.
Pet: caroliniana (Walt.) Gray, Carolina buck
thorn, KF 281
Crataegus crus-galli L., Cockspur hawthorn, KF 295.
Crateagus marshallii Eggl., Parsley hawthorn, KF
Crataegus mollis Scheele, Downy hawthorn, KF 475.
— viridis L.subsp. viridis, Green hawthorn,
(EZ moana
q.,White avens, KF 290
Prunus ener Marsh. var. Sauna
Chickasaw plum, JS 6133.
Prunus gracilis Engelm. & Gray, Sand plum, KF
1060
Prunus mexi icana 5. Wats., Mexican plum, JS 6734.
Else
P runus serotina Ehrhart var. serotina Black cherry
Rubus peste Rydb., Persistent dewberry, KF 693.
bus riograndis Bailey, Dewberry, JS 61
ie trivialis Michx., Southern dewberry, KF
1092
RUBIACEAE
Cephalanthus occidentalis var. californicus
Benth., Buttonbush, JS 6
Diodia teres Walt. var. teres, ae buttonweed,
KF 547
Diodia vi ccna L, Virginia buttonweed, KF 530.
Galium aparine L.,Catchweed bedstraw, KF 498.
Galium circaezans Michx., Woods bedstraw, KF
Galium tinctorium (L.) J. Scopoli, Dye bedstraw,
KF 294,
Houstonia pusilla J. Schopf, Small bluets, KF 229.
Oldenlandi boscii (DC.) Chapm., Bluets, KF 228.
UTACEAE
Zanthoxylum clava-herculis L., Prickly ash, KF 482.
SALICACEAE
t.ex Marsh si ibsp deltonides
Cottonwood, KF 4
alix nigra Marsh., iii willow, JS 6696.
SAPINDACEAE
a al halicacabum L., Balloon vine, KF
ae me enna L. var. drummondii (Hook. &
n.) Benson, Western soapberry, KF 273.
SAPOTACEAE
Sideroxylon lanuginosum Michx., Woolly bucket
bumelia, JS 6690.
SAURURACEAE
Saururus cernuus L., Lizard’s tail, KF 442.
SCROPHULARIACEAE
a tetas oe Small ex Britt., Prai-
agalinis,
a indivisa ie Texas Indian paint-
brush, KF 232.
Gratiola brevifolia Raf., Sticky hedge hyssop, KF
Gratiola virginiana L. subsp. virginiana, Virginia
hedge hyssop, KF 373.
Leucospora multifida (Michx.) Nutt., Narrowleaf
leucospora, KF 587.
Nuttallanthus canadensis (L.) Sutton, Old field
oadflax, KF 236.
Penstemon laxiflorus Penn.
LOOSE PE terilOrl,
Verbascum thapsus L.,Common mullein, KF 720.
Veronica agrestis L., Wayside purslane, JS 6173.
Veronica peregrina L.,Purslane speedwell, KF 304.
SOLANACEAE
see angulata L., Cutleaf ground cherry, KF
Phe mollis Nutt., Field ground cherry, KF 670.
Solanum ptycanthum Dun., American night-
hade, KF 677.
Solanum a L. var. carolinense, Carolina
horse nettle, KF 33/7.
Solanum ee ea Cav., Silverleaf night-
shade, KF 333.
TILIACEAE
Tilia americana L. var. caroliniana (Mill.)
Castiglioni, American basswood, KF 527.
ULMACEAE
Celtis eal oe Willd. var. laevigata, Texas sugar-
berry, JS 66
Planera saa (Walt.) Gmel., Water elm, JS
6676.
Ulmus alata Michx. ,Winged elm, KF 512.
Ulmus aL.,American elm, KF 509.
Ulmus aun Nutt., Cedar elm, KF 508.
URTICACEAE
Boehmeria cylindrica (L.) Sw.,False-nettle, KF 392.
Parietaria pensylvanica Muhl. Willd.,
Hammerwort, KF 299.
370 BRIT.ORG/SIDA 20(1)
=
Urtica chamaedryoides Pursh, Heartleaf nettle, JS Viola missouriensis Greene, Missouri violet, KF 222.
668 Viola palmata L., Trilobe violet, 92.
/j QO
VALERIANACEAE Viola villosa Walt., mains violet, KF 759.
Valerianella radiata (L.) Dufr., Beaked corn salad,
Phoradendron tomentosum (DC.) Engelm. ex
VERBENACEAE Gray, Christmas mistletoe, KF 7718
Callicarpa americana L., American beautyberry, VITACEAE
KF 443. Ampelopsis arborea (L.) Koehne, Peppervine, KF
Glandularia canadensis (L.) Nutt. Rose vervain, 385,
Parthenocissus quinquefolia (L.) Planch. var.
Phyla lanceolat ata (Michx.) Greene, Northern frog quinquefolia, Virginia creeper, JS 6694.
fruit, KF 207. Vitis mustangensis Buckl., Mustang sbi KF 328.
Verbena fas liensis Vell, Brazilian verbena, KF 525.* Vitis palmata Vahl, Catbird grape, KF -
Verbena halei Small, Texas vervain, KF 261. Vitis riparia Michx., Riverbank grape, ie 367.
VIO Vitis rotundifolia oe ar. rotundifolia, Musca-
LACEAE oe
Viola bicolor Pursh, Field pansy, JS 6163 eae senae
ACKNOWLEDGMENTS
We want to thank Ron George and Gary Graham of the Texas Parks and Wild-
lite Department and Wildlife Division Region 3 staff for their support on this
baseline inventory cooperative project. We are also grateful to Tom Wendt of
TEX/LL for his assistance. Also, special thanks to Changxiang Liu, Kim Ludeke,
Vivian Ackerson, Michelle Valek, and Duane German of the Texas Parks and
Wildlife Department Geographic Information Systems Laboratory for their as-
sistance in the vegetation alliance mapping. Thanks to David Riskind, Coordi-
nator of the Natural Resource Program, Texas Parks and Wildlife Department,
and Heather White of GREE who read and commented on preliminary ver-
sions of the manuscript. We are also grateful to Guy Nesom and Monique Reed
lor their review of the manuscript. Their comments were exceedingly helpful.
REFERENCES
Carr, W.R.2001.(unpublished) An annotated list of the G3/T3 and rarer plant taxa of Texas.
Texas Conservation Data Center, The Nature Conservancy of Texas, Austin.
Corre, D.R. 1970. Soil survey of Anderson County, Texas. Natural Resource Conservation
Service, Temple.
Corrett, D.S. and M.C. JoHNston. 1970. Manual of the vascular plants of Texas. Texas Re-
search Foundation, Renner
Corrett, D.S. and H.B. Correit. 1972. Aquatic and wetland plants of southwestern United
States. U.S.Government Printing Office, Washington, D.C
Diamonp, D.D., D.H. Riskino, and S.L.Orzett. 1987. A framework for plant community classifi-
cation and conservation in Texas. Texas J. Sci. 39: 203-221.
Estiut, J.C. and M.B. Cruzan. 2001. Phytogeography of rare plant species endemic to the
southeastern United States. Castanea 66. 3-23.
FLEMING ET AL., FLORA OF BIG LAKE BOTTOM WILDLIFE MANAGEMENT AREA 371
GouLp, FW. 1962. Texas plants — a checklist and ecological summary. Texas Agric. Exp. Stat.
Publ. MP-585, Texas A&M University, College Station.
GouLp, FW. 1975. The grasses of Texas. Texas A&M University, College Station.
Harcn, S.L., K.N. GANDHI, and L.E. Brown. 1990. Checklist of the vascular plants of Texas. Texas
Agric. Exp. Stat. Publ. MP-1655, Texas A&M University, College Station.
Jones, S.D., J.K. Wirrr, and PM. Montcomery. 1997. Vascular plants of Texas: a comprehensive
checklist including synonymy, bibliography, and index. University of Texas Press, Austin.
NATIONAL VEGETATION CLASSIFICATION SYSTEM-OKLAHOMA- Texas SusseT. 1997.The Nature Conservancy,
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Nixon, E.S. and R.L. Witter. 1974. (unpublished) A vegetative analysis of the floodplain of
the Trinity River. Stephen F. Austin State University, Nacogdoches, Texas.
UNIVERSITY OF TEXAS BUREAU OF ECONOMIC GEOLOGY-PALESTINE SHEET. 1993.Map of sedimentary rocks.
University of Texas, Austin.
372 BRIT.ORG/SIDA 20(1)
Books RECEIVED/ NOTICES
Annotations and comments by the Editor
Texas Floras/Natural History/Wildlife
Matt WHITE (FOREWORD BY GREG W. Las_ey). 2002. Birds of Northeast Texas. (ISBN
0-58544-192-9, hbk.; 0-58544-193-7, pbk.). Texas A@M University Press,
John H. Lindsey bldg, Lewis Street, 4354 TAMU, College Station, TX 77843-
4354, US.A.(Orders:800-826-8911, 979-847-8752 fax, WJL@tampress.tamuedu,
www.tamuedu/upress). $44.95 (hbk), $19.95 (pbk), 160 pp., 30 color pho-
tos, map, 6" x 9".
Key Worpbs: Texas, northeast Texas, birds, identification, field guide, zoology.
is at a nice ia the aes has g ene nus. The author summarizes the status of 390 species
1s -|
that | in
—
ecorded from t
he 22 counties of northeast Texas. “The brief
species s explain each species’ status, distribution, and period of occurrence in the region.”
Contents andlide an introduction followed by chapters on Geography of the Region, Birdwatching
Areas, and Species Accounts. The color photos are very nice.
Joun W. TUNNELL, JR.and Frank W. Jupp (FOREWORD BY RICHARD C. BARTLETT). 2002.
The Laguna Madre of Texas and Tamaulipas. (ISBN 0-58544-133-3, hbk).
Texas AGM University Press, John H. Lindsey bldg, Lewis Street, 4354 TAMU,
College Station, TX 77843-4354, U.S.A. (Orders: 800-826-8911, 979-847-8752
fax, WJL@tampress.tamuedu, wwwtamu.edu/upress). $60.00, 372 pp., 83
color photos, 87 figures, +5 tables, 8 1/2" « 11’
Key Worps: Texas natural history, Gulf Coast, Laguna Madre Region, natural resources, environment
seagrass meadows, shorebirds and aces birds, sea turtles, fish and invertebrate fisheries organ-
isms, red and brown tides, conservation issues.
the authors: “This een is intended as an overview of the Laguna Madre for an
re of both scientists and nonscientists. It is firmly based in science but also for use by others
who want to
earn about and take care of the Laguna Madre now and in the future.” There is a nice
appendix of Laguna Madre researchers and interested parties. The literature cited is 30 pages of rel -
erences.
GUNNAR Brune (Introduction by Helen C. Bese). 2002. Springs of Texas. Second
Editon. (ISBN 0-58544-196-1, hbk.). Texas AGM University Press, John H.
Lindsey bldg, Lewis Street, 4354 TAMU, College Station, TX 77843-4354,
U.S.A. (Orders: 800-826-8911, 979-847-8752 fax, WJ_L@tampress.tamu.edu,
www.tamuedu/upress). $75.00 (hbk), 608 pp., 272 b/w photos, 38 line
drawings, 9 color maps, 8 1/2" x IL".
Kry Worps: Springs, Texas natural history, human, population explosion, 2900 springs in 183 Texas
counties, decline and disappearance of springs, any sical chari janes of springs, lore, flora and
fauna of springs, endangered species, Texas water law.
You
‘ve never heard of so many springs. Yes, there is Rattlesnake Springs, Prairie Dog Springs,
Walnut Springs, Onion Springs, Comanche Springs, and the list goes on.
SIDA 20(1): 372. 2002
FLORISTICS OF XERIC SANDYLANDS IN THE
POST OAK SAVANNA REGION OF EAST TEXAS
Barbara R. MacRoberts
and Michael H.MacRoberts James C. Cathey
Bog Research, 740 Columbia Gus Engeling Wildlife Management Area
Shreveport, LA, 71104, U.S.A. Texas Parks and Wildlife Department
Herbarium, Museum of Life Sciences Tennessee Colony, TX 75861, U.S.A.
Louisiana State University-Shreveport
Shreveport, LA 71115, U.S.A.
ABSTRACT
This study describes the floristics, soils, and small-scale species richness of xeric sandylands of the
post oak savanna region of east central Texas and maps the se of xeric sandylands through-
out the West Gulf Coastal Plain. The interrelation of vetlands and their interest
for regional conservation assessment are discussed.
Kry We i lylands, Post Oak Savanna, flora, Texas, Gus Engeling Wildlife Management Area.
RESUMEN
En este est uate se eccecuye a a Hieusie a, suelos, y riqueza de especies a pequena escala - los lugares
k del st Texas y se cartogre adistribucién
s Se discuen la eels de los
oF ; ]
de los lugares arenosos xéricos de la licen costars del West Gu
lugares arenosos xéricos con los humedales y su interés para |
INTRODUCTION
During the past few years, we have been studying wetlands (muck bogs, upland
marshes, baygalls, and seeps) in the post oak savanna region of east central Texas
and southeastern Oklahoma (Nesom et al. 1997; MacRoberts & MacRoberts
1998b). These wetlands appear to be associated with xeric sandylands (vari-
ously referred to as eee aa sandylands [Ajilvsgi 1979], Post Oak-Black
Hickory Series [Diamond et al. 1987], xeric sandylands [MacRoberts &
MacRoberts 1994, 1995, 1996], Gea: Dry Uplands[Turner et al. 1999], Sand
Post Oak - Bluejack Oak Alliance [Singhurst et al. 2000], and Quercus incana
woodland alliance [Hoagland 2000]). These deep sands act as a reservoir or
sponge holding water that feeds adjacent seeps and springs that are the head-
waters for the area’s wetlands and ultimately the streams and rivers. These
upslope soils are porous and drain readily; rainwater percolates through the
sand and moves down a gradient created by underlying impermeable or slowly
permeable clays. Eventually, water seeps laterally out of the hillside (Martin &
Smith 1991; Jones & Carpenter 1995; Drewa 1999; Summer 1999).
As part of our study of muck bogs, upland marshes, and their flora, we
SIDA 20(1): 373 — 386. 2002
374 BRIT.ORG/SIDA 20(1)
studied these adjacent xeric sandylands since they are clearly the water source
for these wetlands.
The primary objectives of this paper are to describe: 1) the distribution of
xeric sandylands throughout the West Gulf Coastal Plain, 2) the floristics of
this community in the post oak savanna region, 3) the small-scale species rich-
ness of this community, and 4) the soils upon which this community occurs.
In addition to our primary objectives for studying this community, we also
are stimulated by a recent spate of national and regional conservation assess-
ments (e.g., Diamond et al. 1997; Ricketts et al. 1999a, 1999b; Myers et al. 2000)
that propose to pinpoint ecological “hotspots.” These assessments are based on
diversity, species richness, endemism, endangered species, and unique habitat
availability. But, as is so often the case with such large-scale efforts, while some
regions are well known, others are not. The post oak savanna region of east Texas
is one of the poorest known. This is evident upon examining recent regional
descriptions of the area (e.g, McNab G Avers 1994; Keys et al. 1995; Ricketts et al
1999a), where numerous factual errors lead to inaccurate assessments of the
region's ecological “temperature.”
THE SETTING
In Texas, the post oak savanna region (Fig. 1), consisting of about 30,000 sq. km,
is gently rolling and hilly with elevations from 90 to 250 meters. Rainfall ranges
from 75 to 115 centimeters. The Carrizo Sands are virtually coterminous with
the region (McBryde 1933). Prairies are scattered throughout, notably in the
south where some prairies are large (Smeins @ Diamond 1983). The predomi-
nant floristic character of the region is southeastern without pines (Smeins &
Diamond 1986). The area contains a diversity of plant communities, from hill-
side pitcher plant bogs, peat bogs, and upland marshes to open xeric sandylands
and oak-hickory forests and woodlands. Characteristic communities within the
post oak savanna region also occur to the east within the piney woods (Marietta
& Nixon 1983; Ward & Nixon 1992). How the post oak savanna region relates
biotically to other regions in the area has yet to be studied in detail (Monk et al.
1990; Bryant et al. 1993; Skeen et. al. 1993), however, it appears to be floristically
similar to adjoining regions (MacRoberts & MacRoberts pers. obs.) and has no
vertebrate endemism (Telfair 1999).
McBryde (1933) conducted the only major floristic/edaphic study of the
region. Subsequently, very little research has been done on the post oak savanna
except for the inclusion prairies in its southern portion (Smeins @ Diamond
1986).
Xeric sandylandsare (or at least once were) very common in the Texas post
oak savanna region, but they also occur in the pineywoods regions of south-
eastern Oklahoma, southwestern Arkansas, western Louisiana, eastern Texas,
MACROBERTS ET AL., FLORISTICS OF XERIC SANDYLANDS IN EAST TEXAS 375
Fic. 1. Post oak savanna region of Texas.
and into the Coastal Bend of Texas (McBryde 1933; Jones 1977; Drawe et al. 1978;
Taylor & Taylor 1978; Ajilvsgi 1979; Marks & Harcombe 1981; Louisiana Natu-
ral Heritage 1988; Bridges & Orzell 1989; Harcombe et al. 1993; Foti et al. 1994,
MacRoberts & MacRoberts 1994, 1995, 1996; Jones & Carpenter 1995; Texas Natu-
376 BRIT.ORG/SIDA 20(1)
ral Heritage Program 1995; Turner et al. 1999; Hoagland 2000). Homologues of
xeric sandylands occur east of the Mississippi River (Stout @ Marion 1993). It is
generally assumed that oak savannas and xeric sandylands, like prairies, are
kept open largely by fire (Smeins & Diamond 1986; Cutter & Guyette 1994).
While encroachment of woody vegetation is ubiquitous in the absence of fire,
there are many areas that remain open even when fire is suppressed, suggest-
ing that edaphic conditions play an integral role. Characteristic tree species of
xeric sandylands include Quercus incana W. Bartram, Q. margarettiae Ashe ex
Small, Q. stellata Wangenh., and Carya texana Buckley, and a variety of fidel
herbaceous species (see Methods for list). The area appears to be rich in plant
endemism (Sorrie @ Weakley 2001), and a large number of West Gulf Coastal
Plain endemics are associated with this community, for example, Brazoria
truncata (Benth.) Engelm. & A. Gray, Palafoxia reverchonii (Bush) Cory, Parony-
chia drummondii Torr. & A. Gray, Pediomelum hypogaeum (Nutt. ex Torr. & A.
Gray) var. subulatum (Bush) J. W. Grimes, Penstemon murrayanus Hook.,
Polanisia erosa (Nutt.) H.H. Iltis, Rhododon ciliatus (Benth.) Epling,
Tetragonotheca ludoviciana (Torr. & A. Gray) A. Gray ex Hall, Tradescantia
reverchonii Bush, and I. subacaulis Bush.
METHODS
In order to develop an objective idea of the distribution of xeric sandylands, we
selected 42 fidel species from the total list of species occurring in this commu-
nity. We mapped these by county and parish over their ranges (Fig. 2). In the
map, the number of fidels per county or parish is indicated in the legend. Par-
ishes or counties with fewer than 10 species are left blank.
The 42 species chosen were Astragalus leptocarpus Torr. & A. Gray, A.
soxmaniorum Lundell, Berlandiera pumila (Michx.) Nutt., Brazoria truncata,
Clematis reticulata Walter, Cnidoscolus texanus (Muell.-Arg.) Small, Coreopsis
intermedia Sherff, Crataegus uniflora Muenchh., Croton argyranthemus Michx,,
Cyperus grayioides Mohlenbrock, Dalea phleoides (Torr. &@ A. Gray) Shinners,
D. villosa (Nutt.) Spreng. Eriogonum longifolium Nutt., E. multiflorum Benth.,
Froelichia floridana (Nutt.) Mog. (not distinguished from F. gracilis [C(Hook.)
Moq.|), Hymenopappus artemisiifolius DC. Lithospermum caroliniense (Walter
ex J.-F Gmel.) MacMill., Loeflingia squarrosa Nutt., Matelea cynanchoides
(Engelm.) Woodson, Minuartia drummondii (Shinners) McNeill, Palafoxia
hookeriana Torr. & A. Gray, P. reverchonti, Paronychia drummondii, Pediomelum
digitatum (Nutt. ex Torr. & A. Gray) Isely, P hypogaeum var. subulatum, Penste-
mon murrayanus, Phacelia strictiflora (Engelm. & A. Gray) A. Gray, Phlox
drummondii Hook., Physalis mollis Nutt., Polanisia erosa (Nutt.) H.H. Htis,
Polygonella americana (Fisch. & C.A. Mey.) Small, Prunus gracilis Engelm.& A.
Gray, Rhododon ciliatus, Selaginella arenicola Underwood ssp. riddelliit (Van
Eselt.) R. M. Tryon, Scutellaria cardiophylla Engelm. & A. Gray, Streptanthus
MACROBERTS ET AL., FLORISTICS OF XERIC SANDYLANDS IN EAST TEXAS 377
Fic. 2. Xeri Ie] J . Al etyiheti f tovt ¢, key).
hyacynthoides Hook., Stylisma pickeringii (Torr.ex M.A. Curtis) A. Gray, Talinum
rugospermum Holz., Tetragonotheca ludoviciana, Tradescantia reverchonii,
Yucca louisianensis Trel.,and Zornia bracteata J.E Gmel. Sources for this infor-
mation consisted of extensive herbarium searches, notably LSU, TAMU (both
on line), ASTC, BRIT, Corpus Christi Museum of Science and History, LSUS,
378 BRIT.ORG/SIDA 20(1)
SBSC, SHST, TEX, VDB, W WE atlases (e.g., Smith 1988; Thomas & Allen 1993-
1998; Turner in press), and a variety of literature (e.g., Jones 1977; Taylor & Tay-
lor 1978; Singhurst 1996; Turner 1996; Nesom & Brown 1998) and unpublished
sources (Billie Turner pers. comm.; Bruce Hoagland pers. comm.). We also made
field searches throughout the West Gulf Coastal Plain to look for the species
and the community in situ.
We surveyed the flora of xeric sandylands at Gus Engeling Wildlife Man-
agement Area (GEWMA), Anderson County, Texas, in the center of the post
oak savanna region. The GEWMA occurs in the central part of the Trinity River
drainage along Catfish Creek (Telfair 1988). This property consists of approxi-
mately 44 sq. km, of which 18 are the sand post oak-bluejack oak community
(Singhurst et al. 2000). We established a single study plot measuring 50 m x
100 m (0.5 ha), within which was another single plot measuring 20 m x 50m
(0.1 ha) with two nested 3.16 m x 3.16 m plots (0.001 ha) and 6 nested 1m x 1m
plots (0.0001 ha) in an opening in this community (see Peet et al. 1998 for plot
design). The larger plot ran parallel to the topographic gradient and the adja-
cent downslope bog and was about 100 meters north of the edge of the bog and
a few meters higher than the bog (see MacRoberts & MacRoberts 1998b, 1999;
Singhurst et al. 2000 for a description of Andrew’s Bog, GEWMA). We surveyed
this area monthly from March to October 2000 and listed all plant species in
each plot. This gave us information on both total floristics and species/area
measures for comparison with similar measures from other plant communi-
ties. We estimated ground cover and measured vegetation height in the 0.0001
ha plots. We also surveyed several other xeric sandylands in other parts of the
GEWMaA. Plant nomenclature throughout this paper follows Kartesz and
Meacham (1999). Voucher specimens are deposited at TEX.
We collected soil samples from the upper 15 cm of the two 0.001 ha plots
and from the 0.1 ha plot for comparison with this community elsewhere in its
range. These were analyzed by A & L Laboratories, Memphis, Tennessee. The
soil at the study site is described as deep, well-drained acidic sandy Pleistocene
terraces (Arenosa series, Typic Quartzipsammeents, Entisols) (Coffee 1975).
—_—
RESULTS
Figure | shows the location of the Post Oak Savanna region of East Texas in
which xeric sandylandsare located. Figure 2 shows the frequency of occurrence
of the 42 xeric sandyland fidel species in parishes and counties in the West
Gulf Coastal Plain. Counties and parishes with fewer than 10 species are left
blank. Table 1 lists the species found within the 0.1 ha plot. Table 2 gives addi-
tional species in the 0.5 ha plot. Table 3 shows the number of species occurring
in the 0.0001 ha, 0.001 ha, 0.1 ha, and 0.5 ha plots. Table 4 gives the ground cover
and vegetation height in the six 0.0001 ha plotsin May and August. Table 5 gives
the soil information from the study plots.
—_—
MACROBERTS ET AL., FLORISTICS OF XERIC SANDYLANDS IN EAST TEXAS
TABLE
(Anderson County, Texas).
1. Species present in the 0.1 ha study plot at the Gus Engeling Wildlife Management Area
AGAVACEAE: Yucca louisianensis Trel.
AMARANTHACEAE: Froelichia floridana (Nutt.)
Moa.
ANACARDIACEAE: Rhus aromatica Aiton, loxico-
dendron radicans (L.) Kuntze
APIACEAE: Spermolepis divaricata (Walter) Raf.
x Ser., S. inermis (Nutt. ex DC.) Mathias &
Constance
AQUIFOLIACEAE: Ilex vomitoria Sol. in Aiton
ASCLEPIADACEAE: Matelea cynanchoides
(Engelm.) Woods
ASTERACEAE: Croptilon divaricatum (Nutt.) Raf,
Evax candida (Torr. & A. Gray) A. Gray,
Helianthus debilis Nutt. ssp. cucumerifolia
(Torr. & A. Gray) Heiser, Krigia virginica (L.)
Willd., Palafoxia reverchonii ae h) Cory,
Pyrrhopappus carolinianus (Walte ene
cio ampullaceus Hook., oo eo filifol lium
(Hook.) A.Gra
i aepetecian
a
ge deo virginicum L.,
5 Hook.
CACTACEAE Lea humifusa (Raf.) Raf.
CA a (Nutt.) H.H. Itis
CARYOPHYLLACEAE Loeflingia squarrosa Nutt.,
d i Torr. & A. Gray
CISTACEAE. Helianthemum georgianum Chapm,
Lechea mucronata Raf, L. tenuifolia Michx.
COMMELINACEAE: Commelina erecta L., Trades
ntia reverchonii Bush, I. subacaulis Bush
Pe otne CEAE: Stylisma pickeringil (Torr. ex
M.A. Curtis) A. Gray
CRASSULACEAE: Sedum nuttallianum Raf.
CUPRESSACEAE: J
CYPERACEAE: Carex cephalophora Muhl.ex Willd,
C.retroflexa Muhl.ex Willd, Cyperus grayioides
uniperus virg TETTCARIGA L.
Mohlenbrock
EUPHORBIACEAE: Cnidoscolus texanus (Muell.-
Arg.) Small, Croton argyranthemus Michx., C.
capitatus Michx., C. glandulosus L., C.
michauxii G.L.Webster
FABACEAE: Astragalus leptocarpus Torr.& A.Gray,
Baptisia nuttalliana Small, Centrosema
virginiana (L.) Benth, Chamaecrista fasciculata
(Michx.) Greene, Dalea phleoides (Torr. &
Gray) Shinners, Indigofera miniata regs,
Mimosa nuttallii (DC) B.L. Turner, Pediomelum
digitatum (Nutt. ex Torr. & A. Gray) Isely
FAGACEAE: Quercus incana W. Bartram, Q.
margarettiae Ashe ex Small
HYDROPHYLLACEAE: Phacelia. strictiflora
(Engelm.& A. Gray) A. Gra
KRAMERIACEAE: Krameria lanceolata Torr.
a Brazoria truncata (Benth.) Engelm.
Gray, Monarda punctata L., Rhododon
a atus (Benth) ering
LILIACE L.
MOLLUGINACEAE Mollugo verticillata L.
ONAGRACEAE: Oenothera laciniata Hil
PANTGNACEAE Plantago hookeriana Fisch.&
A. Mey
Paet ee ternarius Michx., Aristida
desnantne Ts & Rupr, Al anosa a ex a
liott
e CA. ‘Claliy D.oli gesan ines) one
hx
(Walter Chapm,, Vulpia elliotea (Raf) Fernald
POLYGONACEAE: oe multiflorum Benth.,
Rumex hastatulus Baldw
PORTULACACEAE: Talinum rugospermum Holz.
ROSACEAE: Rubus argutus Link
RUBIACEAE: Diodia
SCROPHULARIAC
L.) D.A. Sutton
SOLANACEAE: Physalis heterophylla Nees
VISCACEAE: Phoradendron tomentosum (DC.)
Engelm. & A. Gray
Gourd
Lt
teres Walter
FAF: Nuttallanthus canadensis
DISCUSSION
Xeric sandylands occur from southwest Arkansas and southeastern Oklahoma
to central Louisiana and the Coastal Bend of Texas. The distribution of species
numbers shown in Figure 2 results in part from differential collecting: Caddo
Parish and Anderson County have been well collected while most of east Texas
380 BRIT.ORG/SIDA 20(1)
Taste 2.Species in the 0.5 ha plot not found in the 0.1 ha plot. Both plots at the Gus Engeling Wildlife
Management Area (Anderson County, Texas)
Bulbostylis ciliatifolia (Elliott) Fernald Schizachyrium ni lum bl Nash
Carya texana Buckley Scleria lel
salad us ede x Cav Sisyrinchium albidum a
Co lis micranth ene elm. ex A. Gray) A. Gray Trichostema dichotomum L.
Glandularia canadensis (L.) Nutt. Vaccinium arboreum Marshall
Mirabilis albida (Walter) Heimer| Verbascum thapsus L.
Oenothera heterophylla Spach Vitis aestivalis Michx.
Penstemon murrayanus Hook. Vitis rotundifolia Michx.
-
TABLE 3. Species richness in xeric sandylands at the Gus E1 igelir \Q Wildlife Mar l\aQGeMe! \t Area (Ander-
son County, Texas
Plot size (ha) No. of plots Av. no. species (range)
0.0001 6 24.5 (19-28)
0.001 2 38.5 (35-42)
0.1 1 74
0.5 ] 90
—
has not. Better sampling would undoubtedly fill in the picture, but the outlines
are clear. We have observed the community in situ in Miller County, Arkansas;
Bienville, Caddo, Natchitoches, Vernon, and Winn parishes, Louisiana; Atoka,
Choctaw, and Pushmataha counties, Oklahoma; and Anderson, Angelina,
Aransas, Bastrop, Caldwell, Cass, Cherokee, Colorado, Franklin, Gonzales,
Guadulape, Hardin, Henderson, Jasper, Lee, Leon, Marion, Milam, Nacogdoches,
Panola, Rusk, San Augustine, San Patricio, Shelby, Smith, Tyler, Upshur, Van
Zandt, Wilson, and Wood counties, Texas. We have little experience with the
Coastal Bend xeric sandylands (Drawe et al. 1978), but this community appears
to have affinities with the more northern and eastern xeric sandylands and
needs further study. Xeric sandlylands also occur in Hopkins, Navarro, Rains,
and Williamson counties and appear to have once occurred on the boundary
of Fannin and Grayson counties, Texas Jason Singhurst, pers. comm.).
There were 74 species in the 0.1 ha plot and 90 species in the 0.5 ha plot.
Other taxa in xeric sandylands at GEWMA that did not occur in our plots in-
Chie Apu anoste piilssieurtn obasis(DC.) Trel,, Apocynum L., Asclepias
amplexicaulis Sm., Berlandiera pumila, Bouteloua hirsuta L ag. Delphinium
carolinianum Walter, Descurained pinnata (Walter) Britton, Eragrostis secundiflora
J. Presl., Eriogonum longifolium, Hymenopappus artemistifolius, Liatris elegans
(Walter) Michx., Mirabilis albida, Oxalis priceae Small, Pediomelum hypogaeum
ve subulatum, Physalis turbinata Medik., Polygonella americana, Scutellaria
hylla, Selaginella icola ssp. riddellii, Sideroxylon lanuginosum Michx.,
one
MACROBERTS ET AL., FLORISTICS OF XERIC SANDYLANDS IN EAST TEXAS 381
Taste. 4. Ground cover and vegetation height in six 0.0001 ha plots at the Gus Engeling Wildlife
Management Area (Anderson County, Texas
Plot (month) Average height (cm) Tallest height (cm) Percent cover
1 (August) 20 40 20
( May) 10 55 20
2 (August) 20 30 20
(May) 10 25 30
3 (August) 20 40 20
(May) 15 35 30
4 (August) 15 50 15
(May) 10 20 20
5 (August) 20 40 20
(May) 15 30 40
6 (August) 20 45 25
(May) 15 30 40
Taste. 5. Soil characteristics of sample plots at the Gus Engeling Wildlife Management Area (Ander-
son County, Texas).
Exchangeable ions (ppm)
Sample pH P K Ca Mg Organic Matter%
] 4.9 26 27 172 18 1.0
2 48 21 20 97 14 0.8
3 44 24 18 80 10 0.8
Stillingia sylvatica Garden ex L., Tetragonotheca ludoviciana, Tragia urticifolia
Michx., Triodanis perfoliata (L.) Nieuwl, and Vicia ludoviciana Nutt. (see also
Singhurst et al. 2000).
The ground cover varied from about 15 percent to 40 percent throughout
the growing season. Non-vegetated areas always prominently showed and veg-
etation was never tall. In general, biomass was low and sunlight was directly
on the ground.
Species richness can be measured at many scales. At scales of 0.01 ha and
larger, tropical rainforests are the most species rich. However, at scales below
0.001 ha and often 0.01 ha, temperate grasslands and open savannas of the south-
eastern United States are the most species rich. Values of between 20 and 40
species per 0.0001 ha occur but are very uncommon (Peet et al. 1983; Walker &
Peet 1983; Peet & Allard 1993; Brewer 1998; Platt 1999). Peet et al. 1983) found
that for a broad range of forest and woodland types, no community type ex-
ceeded 17 species per 0.0001 ha and none averaged over 13. Even tallgrass prai-
ries, which were the highest, averaged only 18.
While very little information exists on species richness of West Gulf
382 BRIT.ORG/SIDA 20(1)
Coastal Plain plant communties, the scant small-scale information available
indicates that wetland pine savannas, bogs, and upland pine savannas average
around 20 species in 0.0001 ha plots (Allen et al. 1988; MacRoberts &
MacRoberts 1991, 1998a; Carr 2000).
In the light of these figures, it is interesting that the GEWMA xeric
sandyland plots show higher species richness at small scales than virtually all
other plant communities so far measured in the southeastern United States.
These xeric sandylands also show a low “z” value (about 0.19), meaning that
there is basically a species doubling for every 100-fold increase in area. In the
present case, a 0.0001 ha plot contains roughly one-half of the species found in
a 0.01 ha plot, and one-quarter the species found ina 1.0 ha plot (see MacArthur
& Wilson 1967; Harris 1984 for discussion of “z” value).
The reason for such high species counts in these small-scale plots is not clear
except that, in this case, most species are relatively diminutive witha variety of
growth forms (about 60 Sneae of the species are perennials), and many have
very brief above-ground life histories. Species packing is therefore no problem.
The role of fire in maintaining plant communities is well understood (Platt
1999). In the absence of fire, many plant communities (e.g., prairies) are rapidly
invaded by shrubs and trees (Packard & Mutel 1997). Oak-hickory savannas
and xeric sandylands appear to require fire for natural maintenance (Cutter &
Guyette 1994). We noticed in this study that, because of the long drought that
the West Gulf Coastal Plain has been experiencing, there has beena significant
die-off of woody vegetation, notably in the dryer areas. Fire, therefore, may not
be the only important force preventing woody invasion. Periodic droughts may
be another factor keeping xeric sandylands, as well as other communities, open.
Soils at GEWMA are virtually identical to soils tested from xeric sandyland
sites in Louisiana and east Texas (MacRoberts & MacRoberts 1994, 1995, 1996).
They are nutrient-poor and acidic. Soil conditions per se may also preclude
woody invasion.
—
We have emphasized the floristics of xeric sandylands largely to develop
baseline information, and we note that the species richness of this community
is considerably greater than generally recognized. The hydrologic and geomor-
phologic properties of these xeric sandhills also are significant, as the water
they supply underlies the existence of adjacent wetland communities—bogs
marshes, baygalls, and seeps.
)
ACKNOWLEDGMENTS
Bruce Hoagland, Connie Taylor, Robert Kalinsky, Suzanne Walker, and the staff
of the Gus Engeling Wildlife Management Area and all herbaria where we worked
aided with this research. Special thanks to Billie Turner for allowing us to use his
unpublished distribution maps for Texas plants. The work was supported by a
Texas Parks and Wildlife Section 6, Wildlife Diversity Program Grant: “Status sur-
MACROBERTS ET AL., FLORISTICS OF XERIC SANDYLANDS IN EAST TEXAS 383
vey of Dwarf Pipewort (Eriocaulon kornickianum) in Texas.” Guy Nesom, Ray
Telfair, and Jason Singhust made many helpful comments on the paper.
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EFFECTS OF FIRE ON TWO PITCHER PLANT BOGS
WITH COMMENTS ON
SEVERAL RARE AND INTERESTING PLANTS
Eric L. Keith and N. Ross Carrie
Raven Environmental Services Inc.
PO.B y
Huntsville TX 77342, U.S.A.
ABSTRACT
Pitcher plant bogs require frequent fires to prevent encroachment of woody vegetation. Little infor-
growing-season burns for maintaining plant com-
mation is available on the Mine effectiveness o
munities in bogs and nove wood y encioachment In this study, the effects of growing-season and
dormant-season burns on plant communities were assessed in two pitcher-plant bogs on the Peason
Ridge Military Reservation in western Louisiana. Measurements were collected every month from
March through November 1996 and from March through October 1997. Floristic composition and
forb and woody plant density were deter oe in both bogs. Both bogs had been burned on 3-year
bog, hereafter Coneflower bog, was
ane in the dormant season for approximately 20 years. One
ned during the dormant season in ee ber 1995 and is approximately 0.8 hectare (ha). The
ae bog, Woodpecker bog, was burned during the growing season in August 1996 and is approxi-
mately 0.7 ha. Over both years, we found 98 plant species in both bogs together with 94 of the 98
species in Coneflower bog and 90 of the 98 species in Woodpecker bog. No species were eliminated
from either bog after the fires. In addition, all species that were present after the growing season fire
in Woodpecker bog had been present before the fire. However, 1, several ie date were much more abun-
dant after the fire than before the fire. The vo | ts on the occurrence of se
eral rare and interesting species 1 during ats a including Sabine een
beckia scabrifolia), Texas seiapiass (Eriocaulon texense), Red-t Panicum rigidulum var.
combsii), Slender panicum (Panicum tenerum), Baia, Coonalosaey chalarocephala),
Drummond's lie grass (Xyris drummondii), and Rough-leaved yellow-eyed grass (Xyris
scabrifolia).
RESUMEN
reg ¢.
ainvasion de vegetacion lenosa. Esta disponible
if 5 Lf
f ion sobre la relativa efectividad de los fuegos en la época de crecimiento para mantener
ies comunidades vegetales en turberas y reducir la invasion de vegetacion lefosa. En este estudio, se
evaluaron los efectos de las quemas en la estacion de crecimiento y en la época de parada de
comunidades vegetales en dos turberas en la Reserva Peason Ridge Military en el oeste de Louisiana.
Las medidas se tomaron todos los meses de marzo a noviembre de 1996 y de marzo a octubre de 1997.
Se determino la composici6n floristica y densidad de hierbas y plantas lenosas en ambas turberas.
Ambas turberas han sido quemadas en rotaciones de tres anos en la estacion de parada durante unos
20 anos. Una turbera, concretamente la Coneflower, se quemo en diciembre de 1995 y tiene
aproximadamente 0.8 hectareas (ha). La otra, Woodpecker, se quemo durante la estacion de
crecimiento en ae ee 1996 p tiene Dea 0.7 ha. Durante ambos anos, encontramos
94 de las 98 especies en la turbera Coneflower y 90 de las 98
foe en la turbera Woodpecker Ninguna especie fue eliminada de las turberas después de los
«
4
fay
C
SIDA 20(1): 387 — 395, 2002
388 BRIT.ORG/SIDA 20(1)
fuegos. Ademas, todas las especies que estaban presentes después del fuego en la estacion de
crecimiento en laturbera Woodpecker estaban Seca eens HG eyoaibice varias especies
eran mucho mas abunds untes des spués del fuego que antes. El estudio también contien comentarios
sobe la presencia de
Rudbeckia sca Bal eee E riocarulon lexense, aus oe um var. combsii, Pate fenerum,
2b wncho ith]
sp | ; umm iy X abrifolic
INTRODUCTION
Pitcher plant bogs are unique ecosystems found in the southeastern United
States. In western Louisiana, they occur in Beauregard, Natchitoches, Rapides,
Sabine and Vernon parishes (Allen et al. 1988; Correll & Correll 1972.) Similar
pitcher plant bogs can be found in adjacent counties in East Texas as well as
areas in southeast Louisiana and eastward to Georgia and Florida (Folkerts 1982:
Nixon & Ward 1986). Pitcher plant bogs typically contain saturated sandy soils
underlain by an impervious sandstone or clay layer. Water that percolates
through the sandy soil and reaches the impervious layer seeps to the surface.
Mudstone from the Catahoula Formation usually forms the impermeable layer
in bogs in the Peason Ridge area (Hart & Lester 1993).
Pitcher plant bogs are declining in quantity and quality throughout their
range. Approximately 97% of pitcher plant bogs have been destroyed or severely
altered since European settlement (Folkerts 1982). Public lands are virtually
ner plant
bogs require frequent fires to prevent woody encroachment (Folkerts 1982). The
absence of fire results in eventual elimination of bog species (Folkerts 1982).
Until recently, the flora of pitcher aon bogs was relatively unknown and little
information is available on the relative effectiveness of fire for maintaining plant
communities in bogs and reducing woody encroachment (Barker & William-
son 1988). Kral (1955), MacRoberts and MacRoberts (1988, 1990, 1991, 1992, 1993).
and Nixon and Ward (1986) have recently conducted valuable research on the
floristics of bogs in this region, but information is scarce on the effects of grow-
ing-season burns on pitcher plant bogs in the West Gulf Coast Plain. This project
assesses the effects on vegetation of a growing season fire ona pitcher plant bog
compared toa bog burned during the dormant season. We will also comment
on several rare and interesting plants encountered during our study.
—
the only areas that this ecosystem can be managed effectively. Pite
STUDY SITE/METHODS
The study area was located on Peason Ridge of the Fort Polk Military Reserva-
tion at 31° 20’ 05"N 93° 17'00"W. The area surrounding the study sites is an open,
sandy longleaf pine forest with gently rolling hills and is dominated by herba-
ceous vegetation in the under story. The two bogs, hereafter Woodpecker bog
and Coneflower bog, are located approximately 100 m apart and are separated
py a narrow upland longleaf ridge. They are both open and relatively flat (2%
slope) witha few scattered old growth longleaf pines and are composed mainly
—
KEITH AND CARRIE, EFFECTS OF FIRE ON PITCHER PLANT BOGS 389
of herbaceous vegetation with a few scattered small trees, shrubs, and vines. A
small creek runs along the eastern side of both of the bogs. Both bogs were vis-
ited monthly from March through November 1996 and from March through
October of 1997. The two bogs were chosen because of their close proximity,
similar slope and aspect, and because no bog floristic studies have been con-
ducted on Peason Ridge. The nearest study conducted on bogs was 20 km to
the northeast of the study site in the Kisatchie District of the Kisatchie Na-
tional Forest (MacRoberts & MacRoberts 1988.) Woodpecker bog is approxi-
mately 0.7 ha, and Coneflower bog is approximately 0.8 ha. Ten | x 1 m plots
were spaced in a random pattern in each of the bogs (Fig. 1). Herbaceous and
woody plant measurements were collected in each of the ten plots. Measure-
ments included species frequency of occurrence and density.
Five 5 x 5m randomly selected quadrates were also used to measure woody
stem density in Woodpecker bog (Fig. 1). Measurements of woody stems were
collected in May of 1996 before the summer burnand again in May of 1998 after
the burn to allow fora full growing season. Woody stems over | m tall were also
counted to record the number of mature plants. The growing-season burn in
Woodpecker bog was conducted on 20 August 1996 in the afternoon with south-
erly winds and was effective in completely eliminating most herbaceous and
leafy material. Coneflower bog was burned in December 1995.
We follow MacRoberts (1989) and Allen (1992) for scientific nomenclature.
Voucher specimens for some species were deposited at the University of Louisi-
ana-Monroe and the Botanical Research Institute of Texas (BRIT).
RESULTS AND DISCUSSION
Burn Affects
We recorded a total of 98 species in both bogs, 94 species in Coneflower bog
and 90 species in Woodpecker bog. In Table 1, we list the species present in both
bogs; “C” indicates species found only in Coneflower bog, “W” indicates species
found only in Woodpecker bog, no letter indicates a species that was found in
both bogs. White-topped sedge (Dichromena latifolia), Red milkweed (Ascl-
epias hess and Longleaf milkweed (Asclepias longifolia) were the only spe-
cies found exclusively in Coneflower bog. Species more abundant in Cone lower
bog were “ buttons (Lachnocaulon anceps), Drum heads (Polygala cruciata),
and Sabine coneflower (Rudbeckia scabrifolia). Texas pipewort (Eriocaulon
texense), Narrow leaved-sunflower (Helianthus angustifolius), and Rough-
leaved yellow-eyed grass (Xyris scabrifolia), and red-top panicum (Panicum
rigidulum var. combsii) were the only species found exclusively in Woodpecker
bog. In addition, Rose pogonia (Pogonia ophioglossiodes) and Meadow beauty
(Rhexia petiolata) were much more abundant in Woodpecker bog (Table 2).
Several species numbers were significantly affected by each of the burn
treatments (Table 2). Pitcher plant (Sarracenia alata) numbers were similar
390 BRIT.ORG/SIDA 20(1)
Woodpecker Bog Coneflower Bog
Cl 1X 1m plots
[| 5 X 5 m plots
Fic. 1. Plot locations in both bogs.
before and after the growing-season burn. Before the fire, a total of 392 phyllo-
dia were present in the ten one meter plots. One month after the fire, 383 phyl-
lodia were counted, and in the next growing season, 395 phyllodia were present.
This burn maintained the number of individuals present, contrary to the find-
ings of Barker and Williamson 1988. They indicated a 24% increase in the num-
ber of phyllodia present in the growing season after a winter fire. A 3% increase
in the number of phyllodia occurred in Woodpecker bog after this burn. How-
ever, in Coneflower bog, 463 phyllodia were present in the ten plots immedi-
ately after the fire and only 321 phyllodia were present the next growing sea-
son representing a 31% decrease in the number of phyllodia present between
subsequent growing seasons. This decrease in phyllodia appears to be due to
increased competition from oune Nee tation that returned after the fire.
Our study and previ frequent fires are necessary to main-
tain the density of certain species over time. Barker and Williamson (1988)
indicated an annual decay rate of 30% in Sarracenia alata phyllodia in an un-
burned bog in St. Tammany Parish, Louisiana. At this rate of decay, pitcher plants
would lose 95% of their foliage in only 6 years (Barker & Williamson 1988). Bog
buttons (Eriocaulon decangulare) displayed similar results in our study. We
found 41 plants before the growing-season burn and 40 plants after the burn.
However, in Coneflower bog, this species declined 59%, from 51 to 21, in just one
je
fades
KEITH AND CARRIE, EFFECTS OF FIRE ON PITCHER PLANT BOGS
Taste 1. Taxa present in both bogs.
Lycopodiaceace—lycopodium appressum
(Chapm.) Lloyd & Underw.; Lycopodium
carolinianum L.
Osmundaceae—Osmunda cinnamomea L.
Pinaceae—Pinus palustris P. Mill ; Pinus taeda L.
tdi dese a a z
Burman ae— ae ie aie +)
art.
Cyperaceae—Carex glaucescens Ell ;Dichromena
latifolia Baldw. ex. Ell., (C); Eleocharis
uberculosa (Michx.) Roem chult.; Fuirena
squarrosa Michx.; Raynchospora chalaro
cephala Fern. & Gale; Rhyncospora gracilenta
A. Gray; Rhynchospora oligantha A. Gra
Rhynchospora plumosa Ell; RAynchospora
rariflora (Michx.) Ell; Scleria reticularis Michx.
Eriocaulaceae—Friocaulon decangulare L.
Pry | (\A/\ | | } - i]
ah
=
anceps (Walt.) Morong.
Juncaceae— Juncus scirpoides Lam.;
trigonocarpus Steud.
Liliaceae—Aletris aurea Walt.; Smilax laurifolia L.
Orchidaceae—Calpogon tuberos
Pogonia ophioglossoides (L cto Spiranthes
alas ) LC. Rich
see iepadon gyrans ae a
en (Ell.) Schult. Anthaena! il
Michx.) Beauv.; Aristida palustris (Chapm)
ve ae nga (Muhl.) Scribn.;
is (Lam.) Trin.; Panicum
dchoterior Le Banu ensifolium Baldw
Ell.; Panicum rigidulum Nees. var. see
we & Ball), (W); Panicum scabriusculum
nicum tenerum Beyr.; Panicum
verrucosum Muhl.; Panicum virgatum L.;
Paspalum flori Hae Michx.; Schizachyrium
scoparium (Michx.) Nash; Schizachyriu
tenerum Nees; Tridens ambiguus (Ell.)
hultes.
Xyridaceae—Xyris ambigua Bey.ex. Kunth
sans ions Schultes; Xyris drummondi
me; Xyris scabrifolia Harper, (W).
eee umL
nacardia se en vernix (L.)
Kuntze.
JUNCUS
—
mm
XYris
Apiaceae—Eryngium integrifolium Walt.;
Oxypolis rigidior (L.) Raf; Ptlimnium costatum
(Ell) Raf.
Aquifoliaceace—/lex coriacea (Pursh.) Chapm.
Asclepiaceae—Asclepias longifolia Michx.,(C).;
Asclepias rubra L., (C).
Asteraceae—Aster dumosus L.; Cacalia ovata
Walt.; Chaptalia tomentosa Vent.; Coreopsis
aifolia Nutt.; Eupatorium leucolepsis (DC.)
a: : eye eee um rotundi oe ium L;
VAI).
rictifoliy isl. (\ ); t
graminifolia (Michx.) Shinners; Liatris
pynchostachya Michx.; Sena tenuifolia
Raf.; Rudbeckia scabrifolia Brown
Pee che ne tear ee heal B.L.
Turner
Caprifoliaceae— Viburnum nudum L.
Clusiaceae—Hypericum galioides Lam.; Hyperi
cum hypericoides (L.) Cra ee stans
(Michx.) Adams & Robso
Droseraceae—Dyosera capi ie is Poir.
Ericaceae—Vaccinium corymbosa L.
Euphorbiaceae—Crotonopsis elliptica Willd.
Gentianaceae—Bartonia paniculata (Michx.)
Muhl.; Sabatia gentianoides Ell.
Lamiaceae—Scutellaria integrifolia L.
Lentibulariaceae—Pinguicula pumila Michx.;
Utricu ag cornuta Michx.; Utricularia
subulat
Loganiacene Cynoctonum sessilifolium (Walt.)
Hil.
ea aceae—Ma gnc lia virginiana ie
Melastomataceae—Rhexia lutea Walt.; Rhexia
mariana L.; Rhexia petiolata Walt.
a
Myricaceae— ee cerifera L.; Myrica
heterophylla
pies ceae— | Sylvdtied Marsh.
raceae—Ludwigia hirtella
Pies Polygala cruciata am Polygala
ramosa Ell.
Rosaceae—Aronia arbutifolia (L.) Pers.
aceae— Sarracenia alata Wood.
nis purpurea (L.) Penn.;
Violaceae—Viola primu hohe fe
392 BRIT.ORG/SIDA 20(1)
Taste 2. Species significantly affected by fire treatments. * Coneflower bog was burned in Decem
ber 19
Number of stems
Woodpecker bog Coneflower bog”
Species Pre-burn (96) Post-burn (97) % Change 1996 1997 % Change
Sarracenia alata 392 395 <I 463 321 -31
Aletris aurea 2 13 550 3 12 300
Coreopsis linifolia | 18 1700 21 1 95
Drosera capillaris 2 35 1650 19 6 68
Friocaulon decangul 4] 40 <1 5] 21 59
Lycopodium carolinianum — 9 42 367 26 179 588
Marshallia tenuifolia 4 132 3200 104 1] 9
Mitreola sessilifolia O 2 1 6 500
Pogonia ophioglossiodes 6 54 800 | 5 400
Polygala cruciata | 27 2600 1 7 36
Polygala ramosa 2 37 1750 0 16
Rhexia petiolata 14 5 507 2 6 200
Viola primulifolia 0 14 23 12 48
Xyris ambigua 7 21 200 3 5 67
Xyris baldwiniana 30 151 403 118 97 3
Xyris drummondii 2 20 900 0 4
Xyris scabrifolia 9 800 0 0)
—
growing season. Pitcher plants and bog buttons appear to require frequent fires
to maintain their presence.
Other species that also increased immediately after fire and declined after
one growing season were Colic root (Aletris aurea), Cynoctonum sessifolium,
milkworts (Polygala cruciata and Polygala ramosa), Rose pogonia (Pogonia
ophioglossiodes), Meadow beauty (Rhexia petiolata), and Primrose-leave violet
(Viola primulifolia) (Table 2). Unlike these species, Club moss (Lycopodium
caroliniana) continued to increase a year after the dormant-season burn. The
number of stems continued to increase, from 26 to 179, a year after the dor-
mant-season burn in Coneflower bog. Perhaps this moss-like prostrate species
requires more than one growing season to maximize the number of reproduc-
tive shoots that it produces.
The total number of woody stems increased after the growing-season burn,
however, woody stems greater than one meter tall decreased. (See Table 3 for
woody stem counts before and after the growing-season fire.) This increase in
stems after the fire can be attributed to the large number of root sprouts from
the base of the small hardwood trees, shrubs, and vines. Several small stems
replaced one large stem that had been top killed by the fire. However, many
shrubs were completely killed by the fire with no root sprouting. Sweet bay
KEITH AND CARRIE, EFFECTS OF FIRE ON PITCHER PLANT BOGS 393
Taste 3. Woody stem totals in the five 5 x 5 m plots in Woodpecker Bog.
Total number of stems Stems over 1 meter tall
Species Pre-burn Post-burn %Change Pre-burn Post-burn % Change
Aronia arbutifolia 6 4] 583 1 2 100
Mag! lolia virginiar nes 20 ] 7 -| 5 ] ] 7 -36
Myrica cerifera 1] 14 27 1 1 0)
Myrica heterophylla 327 359 10 83 25 -70
Nyssa sylvatica 5 iS 160 2 3 50
Pinus palustris ] 1 0 1 1 0
Pinus tae 7 6 -14 6 6 0
Rhus vernix 0 6 se 0 3 :
Smilax laurifolia 7 39 457 5 39 680
Vaccinitiim rumbocg O 6 os 0) 4
Total stems 384 502 31 110 91 17
(Magnolia virginiana) and Loblolly pine (Pinus taeda) were the only species
that decreased after the fire. In addition, Poison sumac (Rhus vernix) and Blue-
berry (Vaccinium corymbosum) were not present in the plots before the fire, but
six plants of each species were found in the plots after the fire.
Species Encountered
Rudbeckhia scabrifolia was common in the wetter areas of both bogs. This site is
the northernmost record for this species. MacRoberts and MacRoberts (1993)
did not record this species in Natchitoches Parish and did not find it in their
study sites in the Kisatchie National Forest (MacRoberts & MacRoberts 1988,
1990, 1991, 1992). However, our study site is only 2.5 kilometers south of the
Natchitoches Parish line suggesting Rudbeckia scabrifolia could be found in
this parish and elsewhere on Peason Ridge.
Eriocaulon texense was frequent in Woodpecker bog, but not present in
Coneflower bog. This plant was the only species that was found commonly in
one bog and was absent in the other. This occurrence could also be the north-
ernmost location for this species. MacRoberts and MacRoberts (1993) noted that
they did not find this species in the Kisatchie District of the Kisatchie National
Forest. The location of this species is approximately 20 kilometers north of any
other known location.
Panicum rigidulum var. combsii was very uncommon in Woodpecker bog
with only two small clumps found, and it was completely absent from Cone-
flower bog. This occurrence is only the second record for this distinctive vari-
ety in Louisiana (Allen 1992).
Panicum tenerum was common in both bogs. This plant was not found on
Peason Ridge in a floristic survey (Hart & Lester 1993) and is rare in bogs and
flatwoods in the southern part of the state (Allen 1992).
394 BRIT.ORG/SIDA 20(1)
Rhynchospora chalarocephala was abundant in both bogs. MacRoberts and
MacRoberts (1993) found this species in two bogs in the Kisatchie District and
one bog in the Vernon District. They suggested that this species could be more
common than previously believed. We have seen this species quite commonly
in many bogs on Fort Polk and on Peason Ridge.
Xyris drummondii was common in wet, bare ground in both bogs as well
as many other bogs on Peason Ridge and Fort Polk. This species seems to be
more common than previously documented.
Xyris scabrifolia increased drastically after the growing season burn. We
found | stem in our plots and three stems outside the plots before the burn. Af-
ter the burn, we found 9 stems in our plots and 50 stems outside the plots.
Godfrey and Wooten (1979) suggest that this is the rarest entity of Xyrisin the
southeastern United States, known only from Florida and Georgia. It is pos-
sible that this species could be more common than previously thought. In ad-
dition, it has been found recently in several locations on Fort Polk (Hart & Lester
1993).
The goal of this study was to collect observational data before and after
prescribed fires in two pitcher plant bogs. Although it would have been opti-
mal to collect data in many bogs over several years burned in different growing
seasons, logistica
mee
Saari and other duties prevented such a comprehensive
study. Based on data collected through observations over two growing seasons,
we conclude numerous bog species respond drastically to fire. The life history
and habits of those species suggest frequent burns may be necessary to main-
tain the unique bog ecosystem. We predict future fire ecology studies will fur-
ther support the necessary role of fire in the bog ecosystem.
ACKNOWLEDGMENTS
This research was supported in part by appointments to the postgraduate en-
vironmental management participation program administered by the Oak
Ridge Institute for Science and Education. We would like to thank Charles Stagg,
Stephen Parris, Stephanie Stephens, Kenneth Moore, Chris Melder, and Kevin
Mundorff of the Fort Polk Environmental and Natural Resources Division and
Michael and Barbara MacRoberts of Bog Research for all their help and advice
on this project. We would also like to thank Charles Allen at the University of
Louisiana-Monroe, Larry Brown at Spring Branch Science Center in Spring
Branch, Texas and Guy Nesom at the Botanical Research Institute of Texas for
their assistance in identifying specimens; and William Keith of Lufkin, Texas
for Spanish translation of the abstract.
—
O
REFERENCES
Auten, C.M. 1992. The grasses of Louisiana.Caj
Louisiana.
un Prairie Habitat Preservation Society, Eunice,
—
KEITH AND CARRIE, EFFECTS OF FIRE ON PITCHER PLANT BOGS 395
Auten, C.M.,C.H. Stace, and S.D. Parris. 1988. Analysis of the vegetation in pitcher plant bogs
in two baygalls at Fort Polk in west central Louisiana. Proc. Louisiana Acad. Sci. 50:1-6.
Barker, N.G. and G.B. Wituamson. 1988. Effects of a winter fire on Sarracenia alata and S.
psittacina. Amer. J. Bot. 75: 138-143.
Corrett, D.S.and H.B. Corrett. 1972. Aquatic and wetland plants of the southwestern United
States. Environmental Protection Agency, Washington, D.C
Fotkerts, G.W. 1982. The Gulf Coast pitcher plant bogs. Amer. Sci. 70:260-267,
Goorrey, R.and J.W. Wooten. 1979. Aquatic and wetland plants of the southeastern United
States. Volume |, Monocotyledons. University of Georgia Press, Athens.
Hart, B.L. and G.D. Lester. 1993. Natural community and sensitive species assessment on
Fort Polk Military Reservation, Louisiana. Louisiana Dept. of Wildlife and Fisheries, Ba-
ton Rouge, LA in cooperation with Nature Conservancy, Arlington, VA.
Krat, R. 1955.A floristic comparison of two hillside bog localities in Northeast Texas. Field
& Lab. 23:47-69.
MacRoseets, B.R. and M.H. MacRoseets. 1988. Floristic composition of two west Louisiana
pitcher plant bogs. Phytologia 65:184-190.
MacRoserts, B.R. and M.H. MacRoserts. 1990. Vascular flora of two west Louisiana pitcher
plant bogs. Phytologia 68:271-275.
MacRoberts, B.R.and M.H. MacRoserts. 1991.Floristics of three bogs in western Louisiana.
Phytologia 70:135-141.
MacRoserts, B.R. and M.H. MacRoserts. 1992. Floristics of four small bogs in western Louisi-
ana with observations on species/area relationships. Phytologia 73:49-56.
MacRoserts, B.R.and M.H. MacRoseets. 1993. Floristics of a bog in Vernon Parish, Louisiana,
with comments on noteworthy bog plants in western Louisiana. Phytologia 75:247—
MacRoserts, D.T.1989.A documented checklist and atlas of the vascular flora of Louisiana.
Bull. Mus. Life Sciences, Nos. 7-9, Louisiana State University-Shreveport.
Nixon, E.S.and J.R.Warb. 1986. Floristic composition and management of east Texas pitcher
plant bogs. In Wilderness and Natural Areas in the Eastern United States: A Manage-
ment Challenge, eds. D.L. Kulhavy and R.W. Conner. Center for Applied Studies, School
of Forestry, Stephen F. Austin State University, Nacogdoches, IX. Pp. 283-287.
396 BRIT.ORG/SIDA 20(1)
Books RECEIVED/ NOTICES
Annotations and comments by the Editor
Texas Floras/Natural History/Wildlife
RoLanp H. Wauer and Cart M. FLEMING. 2002. Naturalist’s Big Bend: An Intro-
duction to the Trees and Shrubs, Wildflowers, Cacti, Mammals, Birds, Reptiles
and Amphibians, Fish, and Insects. (ISBN 0-58544-1506-2, pbk.; 0-58544-
155-4, hbk.). Texas AGM University Press, John H. Lindsey bldg, Lewis Street,
4354 TAMU, College Station, TX 77843-4354, U.S.A. (Orders: 800-826-8911,
979-847-8752 fax, WJL@tampress.tamuedu, wwwtamuedu/upress). $29.95
(hbk), $15.95 (pbk), 208 pp., 15 color photos, 28 b/w photos, 90 line draw-
ings, 6" x 9"
ion
Key Worps: Texas natural history, Big Bend, identification, field guide, zoology, botany, trees and
shrubs, wildflowers, cacti, mammals, birds, reptiles, amphibians, fish, butterflies, invertebrates.
This is quite an impressive little guide to Big Bend. The 90 line drawings are of the more com-
mon trees, shrubs, and wildflowers. Good job. The +1-page bibliography is roughly 20% of the book.
Although not intended to be a complete list of publications, it sure is a nice contribution in one
place.
Davip J. Scumipty (forewords by Andrew Sansom and Robrt J.Potts, afterword by
Clyde Jones). 2002. Texas Natural History: A Century of Change. (ISBN 0-
8967 2-469-7, hbk.). Texas Tech Press, Box 41037, Lubbock, TX 79409-1037,
U.S.A. (Orders: 800-832-4042, pena ttup@ttu.edu). $39.95 (hbk),
576 pp., 144 b/w photos, 43 maps, 6" x
Key Worns: Science, natural history, Texas, U.S. biological survey, diane fe heritage, mammals, plants,
natural environment, changes in landscapes, land use, early wildlife
This is quite an accomplishment on Texas history. The author, Dr. Dav 7d J. Se ici is presi-
dent of Texas Tech University. Congratulations on a tremendous resource for Texas!
Cc t ra INA 1 lA we | Ta a |
Ss t Uru
Tom S. COOPpERRIDER, ALLISON W. Cusick, and Joun T. Kartesz (eds). 2002. Seventh
Catalog of the Vascular Plants of Ohio. (ISBN 0-8142-0858-4, hbk; 0-8142-
5061-0, pbk.). The Ohio State University Press, Columbus, OH U.S.A.,
wwwohiostatepress.org. (Orders: University of Chicago Distribution Center,
110305 Landley Ave., Chicago, IL 60628, U.S.A; 800-621-2736). $29.95 (pbk),
io pp. 22
Key Worps: Botany, Ohio, classification, vascular plants, catalog, checklist, systematics
SIDA 20(1): 396. 2002
NATIVE FLOWERING PLANT SPECIES NEW OR
OTHERWISE SIGNIFICANT IN KENTUCKY
Robert F.C. Naczi Ronald L. Jones,
Aude PiillinsBeibe F. Joseph Metzmeier,
Department ae cilleate and Natural Resources Mark A.Gorton,
Delaware 2a ae and Timothy J.Weckman
Dover, DE 1 2277, USA,
Rens ea a tment ornibIag ical Sciences
Fas University
a KY 40475, U.S.A.
ron. jones@eku.edu
ABSTRACT
Twelve species or varieties of vascular plants, all native to the southeastern U.S.A, are reported as
new or noteworthy for Kentucky. Ten of these are the first documented records for the state: Bromus
eile ali sale corrugata, C. oklahomensis, C. ouachitana, C.oxyle epis var. pubescens, C. rodnensis,
ocharis indre peas erythrocarpum, and Viburnum lantanoides.
daaiional coum: are pee for two rare species: Carex buxbaumii and C. seorsa.
RESUMEN
Se presentan 12 especies o variedades de plantas vasculares, todas nativas de la plas sudeste de los
Prados Unidos y se citan como nuevas 0 de interés para el estado de Kentucky | as plantas
n las primeras citas documentadas del estado que son: Bromus nott conan Cues eee C.
E| Gentiana
oklahomensis, C. ouachitana, C. oxylepis var. pubescens, C. roanensis
andrewsii, Vaccinium erythrocarpum, y Viburnum lantanoides. Al nivel del condado, se presentan
dos especies raras: Carex buxbaumii y Carex seorsa.
INTRODUCTION
Recent field and herbarium work have resulted in the discovery of several spe-
cies of native flowering plants new to Kentucky or very rare in the state. These
findings will update Kentucky distribution records as listed by Beal and Thieret
(1986), Browne and Athey (1992), and Medley (1993). Kentucky rarity status—
Special Concern, Threatened, Endangered, or Historical, is based on KSNPC
(2000, 2001). Additional information on Clinton County records is provided
by Gorton (2000). Terminology for physiographic regions is based on Browne
and Athey (1992), and herbarium abbreviations follow Holmgren et al. 1990).
TAXA NEW OR OTHERWISE SIGNIFICANT FOR KENTUCKY
Bromus nottowayanus Fernald (POACEAE). Though Kentucky is well within
the geographic range of this species as provided by Pavlick (1995), this record is
apparently the first published for the state. Almost certainly, earlier Kentucky
collections exist for this woodland grass.
SIDA 20(1): 397 — 402. 2002
398 BRIT.ORG/SIDA 20(1)
Voucher specimen: Kenton Co.: mesic deciduous forest, Independence, 10 Aug 1995, R. Naczi 4911 &
B. Dalton (XNK).
Carex buxbaumii Wahlenb. (CYPERACEAE). This is the second county in Ken-
tucky documented for this species. Earlier records for this species are from
Ballard County, in the westernmost portion of Kentucky, in the Coastal Plain
region (Medley 1993). These records are over 20 years old, and the species is cur-
rently listed in Kentucky as Historical. This collection therefore provides a
redocumentation of the species for the state, and is also the first record from the
Mississippian Plateau region of the state. Kentucky is near the southern limit
of this species; east of the Mississippi River, it ranges south to North Carolina
and Tennessee (Radford et al. 1968; Gleason & Cronquist 1991; Chester et al. 1993),
Voucher — imen: Clinton Co.: hardwood swamp NW of Albany, 20 May 1999, R. Naczi 8008 (DOV,
EKY, KN
Carex corrugata Fernald (CYPERACEAE). These specimens represent the first
record of the species from the state. Kentucky is the northern limit of this sedge,
which is most common on river floodplains in the Coastal Plain of the south-
eastern U.S.A. (Naczi 1992). Most of the collections cited here are from sites along
the Ohio or Mississippi rivers.
Voucher specimens: Butler Co.: Mudd River/Green River junction at Butler/Muhlenberg County line,
3 May 1995, L. McKinney 6373 (MICH). Campbell Co.: Silver Grove, forested floodplain of Ohio River,
12 May 1995, KR. Naczi 4581 & J. C. Ganss (DOV, MICH); Melbourne, forested floodplain of Ohio River,
17 May 1999, R. Naczi 7874 (DOV). Fulton Co.: SW of Hickman, 5 May 1990, L. McKinney 4124 (EKY).
Hardin Co.: Prewitt’s Lake, 29 May 1977, R. Cranfill 239a (MICH). McCracken Co.: Reidland, 26 May
1969, R. Athey 641 (MUR). Union Co.: near Caney Mound School, 6 Jul 1936, H. Shacklette 401 (GH).
Carex oklahomensis Mack. (CYPERACEAE). These specimens represent the first
record of the species from the state. This species also occurs in the midwestern
S.A., from Missouri and Kansas south to Texas, and in the eastern U.S.A. in
Mississippi, North Carolina, Tennessee, and Virginia Jones & Reznicek 1995;
Bryson et al. 1996; Wieboldt et al. 1998).
—
Voucher specimens: Calloway Co.: wet fallow field, Snipe Creek Embayment, 5 Jun 1989, ]. a
1501 (MUR). McCracken Co.: wet ditches on the Ilchneidman Road, 24 May 1970, R. Athey 932 (KN
MUR). Marshall Co.: wet woods, Sharp-Elva Road off US 68, LO May 1972, R. Athey 1666 (MUR).
Carex ouachitana Kral, Manhart & Bryson (CYPERACEAE). This recently de-
scribed sedge is known from sites in western Arkansas and eastern Oklahoma
(Kral et al. 1987; Naczi & Bryson 1990), anda single site in Clay County, Tennes-
see (McNeilus 1992). This collection from Clinton County, Kentucky, only afew
km from the Clay County site, represents a new record for Kentucky, and the
second record of the species from east of the Mississippi River
Voucher specimen: Clinton Co.: dry-mesic slope N of Albany, 11 Apr 1998, M. Gorton 98-147 (EK Y); 21
Apr 1999, M. Gorton 99-109 (EK Y).
NACZI ET AL., NEW KENTUCKY RECORDS 399
Carex oxylepis Torr. @ Hook. var. pubescens J.K.Underwood (CYPERACEAE).
Though C. oxylepis var. oxylepis is known from Kentucky (Medley 1993), this
collection appears to be the first of C. oxylepis var. pubescens from the state.
Additionally, the population documented by this record represents a portion
of the northern range limit of this variety. This taxon appears to be rare; litera-
ture reports are from only one county in Alabama (Bryson et al. 1992), ten coun-
ties of Arkansas (Hyatt 1998), one county of Illinois (Basinger & Shimp 1999),
one county of Mississippi (Bryson et al. 1992), and four counties of Tennessee
(Chester et al. 1993).
Voucher specimen: Clinton Co.: mesic deciduous-juniper forest NE of Albany, 20 May 1999, R. Naczi
7982 (DOV, EKY, KNK
Carex roanensis FJ. Herm. (CYPERACEAE). This is the first record of this sedge
from Kentucky. It was previously known from only a few populations in Geor-
gia, North Carolina, Tennessee, Virginia, and West Virginia, mostly from high
elevation sites (Wieboldt et.al 1998). It was found at several sites on Black Moun-
tain, at elevations between 1100-1150 m, but these sites are threatened by strip
mining activities. It has recently been added to the Kentucky list of rare plants
as Endangered.
Voucher specimen: Harlan Co.: Black Mountain, — S-facing slopes above Razor Fork. 19 Aug
1998, R. Jones and J. Metzmeier 8737, 8757 (EKY
Carex seorsa Howe (CYPERACEAE). These records add two counties to the two
previously reported, Bath and Graves (Ousley & Risk 1998; McKinney et a
2000). This species is most common on the Coastal Plain, and populations out-
side of that physiographic province are very few (Reznicek & Ball 1980). It is
currently listed as a Special Concern taxon in Kentucky.
pace
Voucher specimens: Calloway Co.: Blood River Embayment, 23 May 1989, J. Grubbs 1383 (MUR). Clin-
ton Co.: ei swamp NW ol Albany, 22 May 1998, Naczi 7175 & Ford (DOV): 20 May 1999, R.
Naczi 7986 (KNK).
Eleocharis tuberculosa (Michx.) Roemer & Schultes (CYPERACEAE). This col-
lection represents a state record for Kentucky. It ranges mostly on the Coastal
Plain, and it is quite rare outside of that province (Svenson 1937), though it does
occur nearby in Fentress County, Tennessee (Chester et al. 1993).
Voucher specimens: Clinton Co.: openings in hard ds NW of Albany, 20 May 1999, R. Nacz
I
8002 (DOV, EKY, KNK); 3 Aug 2000, R. Naczi 8598 & M. Wessel (DOV, EKY, KNK)
Gentiana andrewsii Griseb. (GENTIANACEAE). This collection is apparently
the first documented record of this species for Kentucky. Browne and Athey
(1992) cited specimens from the Mississippian Plateau and the Bluegrass re-
gions, but these citations were apparently based ona specimen from Mammoth
Cave (J. Bruna s.n., 23 Oct 1965, EK Y!), which has now been annotated by R. Jones
400 BRIT.ORG/SIDA 20(1)
as Gentiana saponaria L., and on a specimen once housed at the University of
Kentucky Agricultural Herbarium—this specimen was searched for but could
not be located. The collection cited by Medley (1993) asa voucher for Kentucky,
(Short s.n., 1837, MICH!) lacks locality data. The “Lexington, Kentucky” pre-
printed on the label was Short’s residence at the time (Stafleu @ Cowan 1985),
and not the place where the plant was collected. Short’s habit was to write the
collection locality on the label, in the space above pre-printed information. Since
his label lacks such an inscription, it must be regarded as of unknown prov-
enance. Pringle (1967) cited no Kentucky specimens of G. andre wsii, and states
that this Campbell County collection is the first he has seen for the state QJ. Prin-
gle, pers.comm.). The site of the collection is at or near the southern range limit
for G.andrewsii, which is known from nearby southern Indiana and southern
Ohio (Pringle 1967). At the Campbell County locality, the gentians were locally
common in an open, wet-mesic, deciduous floodplain forest dominated by
young Acer rubrum and Quercus palustris.
—_—
Voucher specimen: Campbell Co.: Silver Grove, floodplain of Ohio River, 13 Oct 1997, R. Naczi 7059
(KNK), dupl. det. by James S. Pringle; 29 Sep 1998, R. Naczi 7735 (KNK).
Vaccinium erythrocarpum Michx. (ERICACEAE). The mountain cranberry is
restricted to the southern Appalachians, from West Virginia to Georgia, mostly
above LOOO m. Medley (1993) reported a possible sterile specimen of this spe-
cies collected from Harlan County (L. Pounds 85-138) deposited at the Univer-
sity of Tennessee Herbarium, but a recent search failed to locate any Kentucky
collection (B. E. Wofford, pers. comm.). This fruiting specimen substantiates the
presence of the species in Kentucky. The site, however, is likely to be destroyed
by strip mining activities. It has recently been added to our state list of rare
plants as Endangered.
—
Voucher specimen: Harlan Co.: S slope of Black Mountain, woodland edge along mining road toward
Left Fork of Fugitte Creek, elevation 900 m, 14 Aug 1998, R. L. Jones & J. Metzmeier 8639 (ERY).
Viburnum lantanoides Michx. (CAPRIFOLIACEAE). This is the first record of
Viburnum lantanoides for Kentucky. Earlier reports were unsubstantiated or
based on misidentifications (Weckman 1999). The species is common in some
high elevation areas of the southern Appalachians, usually above 1000 m. It
has been documented from several western Virginia counties, including con-
tiguous Wise County (Harvill et al. 1986), but is not yet known from Tennessee
counties adjacent to the Kentucky border (Chester et al. 1997). A single popula-
tion of about 50 individuals was discovered in the Razor Fork streamhead, grow-
ing with Betula alleghaniensis and Magnolia fraseri. Persistence of this Ken-
tucky population is threatened by potential valley fill associated with proposed
strip mining on Black Mountain. It has recently been added to the Kentucky
list of rare plants as Endangered.
rH
NACZI ET AL., NEW KENTUCKY RECORDS 401
Voucher specimen: Harlan Co.: Black Mountain, at head of Razor Fork, elevation 1050 m, 19 Aug
1998, R. L. Jones & J. Met zmeier 8804 (EKY): 5 Sep 1998, T. Weckman 4349 (EKY).
REFERENCES
Basincer, M.and J. SHime. 1999. Noteworthy collections from Illinois. Castanea 64:272-275.
Beat. E.O. and J.W. THieret. 1986. Aquatic and wetland plants of Kentucky. Kentucky Nature
Preserves Commission, Scientific and Technical Series, Number 5, Frankfort.
Browne, E.T., Jk. and R. AtHey. 1992. Vascular plants of Kentucky: an annotated checklist.
University Press of Kentucky, Lexington.
Bryson, C.T., J.R. Macponato, R. Carter, and S.D. Jones. 1996. Noteworthy Carex, Cyperus,
Eleocharis, Kyllinga, and Oxycaryum (Cyperaceae) from Alabama, Arkansas, Georgia,
Louisiana, Mississippi, North Carolina, Tennessee, and Texas. Sida 17:501-518.
Bryson, C.T., R.EC. Naczi, and S. McDanieL. 1992. Notes on noteworthy records of Carex
(Cyperaceae) from the southeastern United States. Sida 15:125-135.
Cuester, EW., B.E. Worrorp, R. Krat, H.R. DeSeim, and A.M. Evans. 1993. Atlas of Tennessee vascu-
lar plants. Vol. 1.Misc. Publ.No.9. The Center for Field Biology Austin Peay State Univer-
sity, Clarksville, Tennessee.
Cuester, E.W., B.E. Worrorp, and R. Kral. 1997. Atlas of Tennessee vascular plants. Vol. 2. Misc.
Publ. No. 13. The Center for Field Biology, Austin Peay State University, Clarksville, Ten-
nessee.
Gteason, H.A. and A. Cronauist. 1991. Manual of vascular plants of northeastern United
States and adjacent Canada, 2" ed. The New York Botanical Garden, Bronx.
Gorton, M. 2000. The vascular flora of Clinton County, Kentucky. M.S. thesis, Eastern Ken-
tucky University, Richmond.
Harvict A.M., Jr., T.R. Branvey, C.E. Stevens, T.F.Wieboldt, D.M.E.Ware, and D.W. Ogle. 1986. Atlas
of the Virginia flora. 2"4 ed. Virginia Botanical Associates, Farmville.
Houmoren, PK., N.H. Houmeren, and L.C. Barnerr (eds.). 1990. Index herbariorum part |. The
herbaria of the world, 8th ed. New York Botanical Garden, Bronx.
Hyatt, PE. 1998. Arkansas Carex (Cyperaceae): a briefly annotated list. Sida 18:535-554.
Jones, S.D. and A.A. Reznicek. 1995. Carex conjucta (Cyperaceae) verified for Arkansas, and
notes on the range of Carex oklahomensis. Sida 16:772-774.
Krat, R., J.R. MANHaART, and C.T. Bryson. 1987. A new Carex sect. Oligocarpae (Cyperaceae)
from western Arkansas and eastern Oklahoma. Ann. Missouri Bot. Gard 74:440-442
[KSNPC] Kentucky State Nature Preserves Commission. 2000. Rare and extirpated biota of Ken-
tucky. J. Kentucky Acad. Sci.61:115-132.
[KSNPC] Kentucky State Nature Preserves Commission. 2001. Rare and extirpated biota of Ken-
tucky: 2001 update. J. Kentucky Acad. Sci.62:145-146.
McNetus, V.B. 1992. Carex ouachitana (Cyperaceae) new to Tennessee. Sida 15:154—155.
McKinney, L.E.,G. Ligsy, and R.Mears. 2000. New and noteworthy records of Carex (Cyperaceae)
from Kentucky. Castanea 65:221-224.
Meotey, M. 1993. An annotated catalog of the known or reported vascular flora of Ken-
tucky. Ph.D. dissertation, University of Louisville, Louisville, Kentucky
402 BRIT.ORG/SIDA 20(1)
Naczi, R.F.C. 1992. Systematics of Carex section Griseae (Cyperaceae). Ph.D. dissertation.
University of Michigan, Ann Arbor.
Naczi, REC. and C.T. Bryson. 1990. Noteworthy records of Carex (Cyperaceae) from the
southeastern United States. Bartonia 56:49-58.
Oustey, J.R.and A.C. Risk. 1998. Preliminary bryophyte and vascular flora of the Hog Hollow
Seeps, Bath County, Kentucky. J. Kentucky Acad. Sci. 59:97.
Pavuck, L.E. 1995. Bromus L. of North America. Royal British Coumbia Museum, Victoria.
Painate,J.S. 1967. Taxonomy of Gentiana, section Pheumonanthae, in eastern North America.
Brittonia 19:1-32.
Raproro, A.E., H.E. AHies, and C.R. Beit. 1968. Manual of the vascular flora of the Carolinas.
The University of North Carolina, Chapel Hill.
Reznices, A.A.and PW.Ball. 1980. The taxonomy of Carex section Stellulatae in North America
north of Mexico. Contr. Univ. Michigan Herb, 14:153-203.
Starteu, FA. and R.S. Cowan. 1985. Taxonomic literature, 2"? ed., Volume V: Sal-Ste. Bohn,
Scheltema, & Holkema, Utrecht.
SveNnson, H.K. 1937.Monographic studies in the genus Eleocharis. IV.Rhodora 39:210-231,
236-273.
Weckman, T.J. 1999. A distributional and taxonomic study of the genus Viburnum
(Caprifoliaceae) in Kentucky. MS. thesis, Eastern Kentucky University, Richmond.
Wiesotor T.F, G.P. FLemine, J.C. Lupwic, and F.C. Huser. 1998. Noteworthy collections from Vir-
ginia. Castanea 63:82-91.
NEW RECORDS IN ASTERACEAE
FOR ALABAMA AND ARKANSAS
Guy L.Nesom
Botanical Research Institute of Texas
509 Pecan Street
Fort Worth, 1X 76102-4060, U.S.A.
Continuing study of Asteraceae brings to light additions to the Arkansas and
Alabama floras.
Pluchea yucatanensis Nesom. When first described (Nesom 1989), P.
yucatanensis was known in the USA from only one collection in Hancock Co.,
Mississippi. It is now recorded from four contiguous counties in Alabama (first
report here) and Mississippi and appears to be an established element of the
flora in that area of the Gulf Coast. Its larger range is in Central America (Belize)
and Mexico (Veracruz, — and Quintana Roo).
Voucher specimens. ALABA Mobile Co.: just N of jct.Co. Rd 59and Ala. Hwy 188, sandy edge of salt
marsh in slash pine, 15 Jul ee Kral 35624 (VDB). MISSISSIPPI. Hancock Co.: S of Kiln, sandy area
along Jordan River, coastal flatwoods, 26 Aug 1966, Jones 9575 (NCU); S of Kiln, sandy, low area along
Jordan River, edge of mixed woods with Serenoa, Sabal, Nyssa, Taxodium, = May 1967, ae 12656
(TEX); S of Kiln, low sandy soil ae ie River, 6 Jul 1967, Jones 14155 (NCU). Harrison Co.: SW of
Wool Market, low sandy soil along Biloxi River, R LOW T 7S Sec 8, 23 Jul Cea (GH). Jackson
Co.: Ocean Springs PO., low wet areas, 2 Jul 1952, Demaree 32262 (BRIT-SMU, GH); E of Ocean Springs, near
borrow pit on Hwy 90, 25 Jul 1966, Deramus 548 (V DB); Ocean Springs, 30 Jul 1896, Pollard 1129 (GH).
Pluchea yucatanensis is similar in habit and general appearance to P. foetida
(L.) DC. and P rosea Godfrey and has been misidentified as both. The rose-tinted
phyllaries and florets are more similar to P. rosea, but the glabrous, slightly thick-
ened, shiny leaves and glabrous phyllaries are unambiguous recognition char-
acters for P yucatanensis.
Pseudognaphalium luteoalbum (L.) Hilliard & Burtt. A recent report expanded
the known range of this species into Texas and New Mexico (Nesom 2001). It
probably should be expected in Oklahoma and other states eastward. It is al-
ready documented in Florida by many collections.
Voucher specimen. ARKANSAS. Nevada Co.: flat clear-cut woods along Co. Rd 221 E of Co. Rd 37,N
of Missionary Grove Baptist Church, N of Prescott, 19 Aug 1999, Thomasand Young 162,394 (TEX).
Symphyotrichum puniceum (L.) A. & D. Love var. scabricaule (Shinners) Nesom.
Variety scabricaule previously has been recorded from localities in Texas, Loui-
siana, and Mississippi (Nesom 1997).
Voucher specimen. ALABAMA. Chambers Co.: 1.9 mi NW of Penton, seepage area in granite outcrop
area, 17 Oct 1969, Kral 37887 (BRIT, VDB-5 sheets).
SIDA 20(1): 403 — 404. 2002
404 BRIT.ORG/SIDA 20(1)
The occurrence of Symphyotrichum puniceum in Alabama was documented
by Mohr (1901), who noted that the only collection was by FS. Earle from Au-
burn in Lee County in the east-central section of the state (immediately adja-
cent to Chambers County). Mohr noted that the plants grew in “swampy bor-
ders of woods” in “metamorphic hills” of the “mountain region.” Both Alabama
localities are at the southwestern extreme of the southward Appalachian ex-
tension of the range of S. puniceum through Georgia. Some of the distinctly
montane Georgia populations are typical in morphology (var. puniceum) while
others (e.g., Haralson Co., Kral 77182-VDB) show features of var. scabricaule,
especially in reduction of the uppermost leaves. Prior to this report, var.
scabricaule was considered a Gulf coastal plain endemic (Nesom 1997). Cham-
bers and Lee counties are slightly north of the coastal plain (as marked by the
boundary between Cretaceous and older Paleozoic formations, e.g., see Sorrie &
Weakley 2001). Lee County, however, is included in four of the geographic pat-
terns of coastal plain endemism shown Sorrie and Weakley.
Kartesz (1999) includes Symphyotrichum puniceum for Alabama as var.
puniceum, but the morphology of the plants suggests that identification as var,
scabricaule is more appropriate. The lower leaf surfaces are about the same color
as the upper surfaces, without a reticulum of dark veins, the main veins of the
upper leaf surfaces are shallowly impressed, giving a slightly rugose appear-
ance, and the leaves of the capitulescence are markedly reduced in size com-
pared to the lower.
fo.
ACKNOWLEDGMENTS
lam grateful to the staffs at BRIT, GH, NCU, TEX, and VDB for help during
recent studies in those herbaria and to Bob Kral and Michael Woods for review
comments.
REFERENCES
Kartesz, J.T. 1999. A synonymized checklist and atlas with biological attributes for the
vascular flora of the United States, Canada, and Greenland (ed. 1). In: Kartesz, J.T., and
C.A. Meacham. Synthesis of the North American flora, Version 1.0. North Carolina Bo
tanical Garden, Chapel Hill.
Mone, C. 1901. Plant life of Alabama. Contr. U.S. Natl. Herb. 6:1-921.
Nesom, G.L. 1989. New species, new sections, and a taxonomic overview of American
Pluchea (Compositae: Inuleae). Phytologia 67:158-167.
Nesom, G.L. 1997.The status of Aster scabricaulis (Asteraceae: Astereae), an endemic of the
Gulf coastal plain. Phytologia 82:300-315.
Nesom, G.L. 2001. New records in Pseudognaphalium (Gnaphalieae: Asteraceae) for the
United States. Sida 19:1185-1190.
Sorrit, B.A.and A.S.Weakity.2001.Coastal plain vascular plant endemics: Phytogeographic
patterns. Castanea 66:50-82.
SIX NON-NATIVE SPECIES NEWLY DISCOVERED
IN THE IOWA VASCULAR FLORA
Allison W. Cusick
Division of Natural Areas and Preserves
Ohio Dep artment o
1889 Fountain Sq. Ct, F-1
Columbus, OH 43224, U.S.A.
With the continuing invasion of indigenous floras by non-native elements, it
isimportant to document early appearances of alien species. This paper focuses
on six species that have appeared in the lowa vascular flora in recent years. Two
species were first noted in 1976 and 1981, the remainder in 2000. None of these
species are included in the checklist of lowa vascular plants by Eilers and Roosa
(1994). Single populations of two species were first reported in Norris et al.
(2001). This paper documents additional populations. Although these species
are not widespread as yet in the state, there probably is no barrier to their spread.
Most are weeds of ornamental plantings and are spread primarily via horticul-
tural materials. Most previous studies of horticultural introductions have fo-
cused on the spread of woody landscape plants (Vincent &@ Cusick 1998). The
present work concerns the incidental introduction of herbaceous species. One
shrub discussed below, Tamarix parviflora DC., potentially is a serious pest spe-
cies, however. Careful searching should yield additional lowa populations of
all six species.
Specimen citations are abbreviated and full details are available from the
author,
Anethum graveolens L. (APIACEAE). Culinary dill is a sporadic waif in North
America. Apparently, this is the first report of this herb in the lowa flora.
Des Moines Co.: bank of Mississippi River, US Hwy. 34 bridge, Burlington, 15 Jul 2000, Cusick 35548
USC, MICH).
Fatoua villosa (Thunb.) Nakai (MORACEAE). Mulberry-weed isan annual spe-
cies from Asia which now is common throughout the southeastern United
States. It typically grows in nurseries and amid ornamental plantings. It also is
a common adventive in greenhouses. The lowa population is a northwestern
range extension from southern Missouri and central Ohio (Wunderlin 1997). In
the north, Fatoua villosa is restricted to protected situations, such as walls and
south-facing slopes. The climate at the Muscatine County site is moderated by
the nearby Mississippi River. Mulberry-weed likely will be found at other places
along the river in southern Iowa.
—
SIDA 20(1): 405 — 407. 2002
406 BRIT.ORG/SIDA 20(1)
Muscatine Co.: nursery on terrace of Mississippi River, St. Hwy. 22, Muscatine, 21 Jul 2000, Cusick
35602 (ISC, MICH, MU, OSH).
Oxalis corniculata L.(OXALIDACEAE). This species is adventive from tropical
America. In the southern U.S, it is widespread in ornamental plantings. It is
common in greenhouses as well (Lourteig 1979). As with Fatoua villosa, it ap-
parently spreads as a contaminant with potted ornamentals, based upon the
author's personal observations of nursery material.
Black Hawk Co.: lawn, Gilchrist Hall, University of Northern lowa campus, | Apr 1981, I: Lammers
4028 (ISC). Muscatine Co.: nursery on terrace of Mississippi River, St. Hwy. 22, Muscatine, 21 Jul
2000, Cusick 35603 (SC).
Sagina procumbens L. (CARYOPHYLLACEAE). European pearlwort was first
reported in lowa from Story County where it was discovered in 2000 (Norris et
al. 2001). It is common throughout the northeastern and northwestern U.S., but
sporadic in Midwestern states. This perennial species typically grows in crev-
ices in walls and sidewalks and as a weed in ornamental plantings. The similar
annual, Sagina decumbens (ELL) T.&G., grows in comparable situations and oc-
curs near lowa in northcentral Hlinois (Crow 1978).
Cerro Gordo Co.: garden store, 4th St. SW & S Polk Ave., Mason City, 19 Jul 2000, Cusick 35588 (SC):
Muscatine Co.: nursery on terrace of Mississippi River, St. Hwy. 22, Muscatine, 21 Jul 2000, Cusick
35601 (SC).
Tamarix parviflora DC. (TAMARICACEAE). Salt cedar is a widespread inva-
sive species of riparian corridors throughout the western United States. It is a
serious pest, often forming a monoculture that reduces natural biodiversity
(Weisenborn 1996). The plants are isolated at the Des Moines County station,
with no planted material nearby. Its presence in the lowa flora should be closely
monitored. Extirpation of the shrubs at the single known site probably is war-
ranted.
Des Moines Co.: t Il shrubs, edge of pond, Sullivan Slough Rd.,S of Burlington, 15Jul 2000, Cusick
35555 (ISC, MICH).
Viola arvensis Murray (VIOLACEAE). A single previous collection of European
field pansy from Story County, lowa in 1998 has been reported (Norris et al.
2001). Unlike most violets, this is an annual species that may bloom in any
month of the growing season.
k 35585 (SC). Des Moines Co.:
Cerro Gordo Co.: nursery, St. Hwy .122, Mason City, 17 Jul 2000, Cusic
flowerbed, 1326 Griswold St, Burlington, 4 Jul 1976, T. L s 876 Us
ACKNOWLEDGMENTS
| thank Diana H. Horton of the University of lowa and Deborah Q. Lewis of
lowa State University for their assistance with this article. My research was sup-
ported in part by the Division of Natural Areas and Preserves, Ohio Depart-
ment of Natural Resources.
Ls
CUSICK, NEW RECORDS FOR IOWA 407
REFERENCES
Crow, G.E. 1978. A taxonomic revision of Sagina (Caryophyllaceae) in North America.
Rhodora 80:1—91.
Eiters, L.J.and D.M. Roosa. 1994. The vascular plants of lowa. Univ. of lowa Press, lowa City.
Lourteic, A. 1979. Oxalidaceae extra-austroamericanae. Il. Oxalis section Corniculatae DC.
Phytologia 42:57-198.
Norris, W.R., D.Q. Lewis, M.P. WiortecHner, J.D. THomeson, and R.O. Pore. 2001. Lessons from an
inventory of the Ames, lowa, Flora (1859-2000). J. lowa Acad. Sci. 108:34-63.
Vincent, M.A.and A.W. Cusick. 1998. New records of alien species in the Ohio vascular flora.
Ohio J. Sci.98:10-17.
Weisensorn, W. 1996. Tamarisk. In: J.M.Randall and J. Marinelli,eds. Invasive plants. Weeds of
the Global Garden. Brooklyn Bot. Gard. Handbook 149, Brooklyn, NY. Pp. 43-44.
WuNbERLIN, R.P. 1997. Moraceae. In: Flora of North America Edit.Comm.(eds.).Flora of North
America. Vol. 3. Oxford Univ. Press, New York and Oxford. Pp. 388-399.
BRIT.ORG/SIDA 20(1)
Books RECEIVED/NOTICES
Annotations and comments by the Editor
Cc 7 IMA 1 (El ‘Tae -_j
WRI
BrucE G. GALDWIN, STEVE BOYD, BARBARA J. ERTTER, ROBERT W. PATTESON, THOMAS J. ROSATTI,
and Dieter H. WILKEN (eds), MARGRIET WETHERWAX (managing ed). 2002. The
Jepson Desert Manual. Vascular Plants of Southeastern Califonria. (ISBN
0-520-22775-1, pbk.). University of California Press, Berkeley, CA 94720,
U.S.A. (Orders: California Princeton Fulfillment Services, 1445 Lower
Ferry Road, Ewing, NJ 08618, U.S.A., 609-883-1759, 609-883-7413 fax,
www.ucpress.edu). $35.00, 624 pp., 128 color photos, many line drawings,
eT
ey Worps: Systematics, botany, identification, keys, natural history, California, southeastern Cali-
fornia, Mojave Desert, Sonoran Desert, southern Great Basin, White Mountains, desert plants, vascu-
lar plants.
a Bs
or photo section is quite a nice addition since its “mother, The Jepson Manual, did not
include color photos. California is big enough for good regional floras.
PETER Lesica, 2002. A Flora of Glacier National Park. (ISBN 0-87071-538-0, pbk.).
Oregon State University Press, Ol Waldo Hall, Corvallis, OR 97331-6407,
U.S.A. (Orders: 800-426-3797, 541-737-3170 fax, OSU.Press@orst.edu,
osuorst.edu/dept/press/osupress.htm). $32.95, 480 pp., 8-page color insert,
60 color photos, 351 line drawings, 6" « 9",
Key \ S ics, flora, field guide, keys, desc riptions, habitats hical taxonomy
ethnobotanic: “al uses, climate, geology, early botanical exploration, plant geogr seals introduced spe-
cies, Rocky Mountains.
Davip H. ENGEL and Suctiart PHUMMAI. 2002. A Field Guide to Tropical Plants of
Asia. (ISBN 0-88192-542-X, pbk.). Timber Press, Inc., 133 S.W. Second Avenue,
Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: wwwtimbe1 press.com,
800-327-5680, 503-227-2878, 503-227-3070 fax). $19.95, 280 pp., app.rox.
390 color photos, 8 figures, 6" 9"
Key Worps: Asia, tropical plants, field guide. flora, identification, descriptions, local names, over 300
species.
The flora is broken down by trees, shrubs, vines, groundcovers, and bedding plants. Under each
category, the plants are listed by lower, foliage and fruit
SIDA 20(1): 408. 2002
NOTEWORTHY VASCULAR PLANTS FROM ARKANSAS. II
Eric Sundell R. Dale Thomas
School of Matneinaieg! ang Natural Sciences Department of Biology
\rkans University of Louisiana at Monroe
iontieeles AR 71656, U. s A. Monroe, LA 71209, U.S.A.
Carl Amason Chris Doffitt
PO. Box 164 Department of Biolog
Calion, AR 71724, U.S.A. University of Louisiana at Monroe
Monroe, LA 71209, U.S.A.
ABSTRACT
The authors provide a list of 41 additions, reinstatements, and significant ass extensions for th
flora of Arkansas. aieroaniner pny hioides, ae incana, peeoenes sis, Hypoxis
,Nandi | UM % ee as new and
curtissit
persistent elements, while omen: cleucantha, Hae alapa, Nana: and Veronica polita are
]
€) ylanic iY 0 df Se Beg 8 usine Pecan
reinstated to the state flora. oe of alien meres —Cynoglossum 2
Euphorbia hirta, Scoparia dulci it _ Trifolium nigrescens, Verbena montevidensis—
are noticed and documented for Hie ae time, Rneeiel their persistence is not known.
RESUMEN
Los autores proporcionan una lista donde se enumeran 41 adiciones, restablecimientos, y extensiones
de area significativas para la flora de Arkansas. Alternanthera paronychioides, Bowlesia incana,
Eleocharis montevidensis, Hypoxis curtissii, Nandina domestica, Polygonum setaceum, y Ir ifolium
lappaceum son citadas como componentes nuevos y persistentes del area, mientras que aes xX
leucantha, Marah dalape, noe minor, ny oo son una rehabilitacion paraa la flora del
Cynoglossum zeylanicum, Eleusine tristachya,
~~
estado. U
Euphorbia hirta, Scoparia dulcis eee pratensis, Tifol ium nigrescens, Verbena montevidensis—
son mencionad as y Aeciienedes S por primera vez, sine mbargo su per sistencia no es conocida
The authors provide a list of 41 taxa representing additions and reinstatements
to the Arkansas flora as well as noteworthy range extensions within the state.
Herbarium abbreviations are taken from Holmgren et al. 1990
AMARANTHACEAE
Alternanthera paronychioides St. Hilaire. Thomas discovered large populations
of this prostrate chaff-flower around the edge of borrow pits inside the Missis-
sippi River levee, in Chicot and Lincoln counties, in August and September, 2000.
Duplicate specimens were confirmed by Kenneth R. Robertson of the Illinois
Natural History Survey in Champaign.
homas 166,893 & 167,194 (NLU, UAM). Lincoln Co.: Thomas & Sundell
Ww
—
Voucher specimens: Chicot Co.: |
167,417 (NLU, UAM)
SIDA 20(1): 4 409- a8. 2002
410 BRIT.ORG/SIDA 20(1)
APIACEAE
Bowlesia incana Ruiz and Pavon is a decumbent annual of low woods, clear-
ings, lawns, and other moist, weedy places, occurring sporadically in Louisiana
and eastern Texas (Thomas and Allen 1996, Diggs et al. 1999). We record its first
appearance in Arkansas from Chicot County in the southeast corner of the state,
along a road bank as well as in alluvial woods along Indian Creek south of
Eudora.
Voucher specimens: Chicot Co.: Thomas 164,552 (NLU), 164,589 (NLU, UAM).
Sanicula smallii Bickn. Smith (1988) documents this cryptic black snakeroot
from two counties in the Ozark Mountains. We add collections from the Oua-
chita Mountains and the Coastal Plain and suggest that the species might be
undercollected in Arkansas. It is distinguished in the field from the widespread
S.canadensis by rather subtle characters: sessile, slightly larger fruits and semi-
succulent leaves. (The tuberous, thickened roots are only helpful after the plant
has been recognized.) In addition, in at least one of our populations the two
woodland species were mixed, with S. canadensis much the more common.
Voucher pees Gen! Co.: Leslie 435B (NLU). Lafayette Co.: Sundell, Thomas, & Amason 12,300
UAM). Montgomer E. & J. Sundell 11,826 (UAM). Ouachita Co.: Thomas & Doffitt 163,219 (NLU).
Union Co.: T ee - 816 (NLU).
ASTERACEAE
Hypochoeris glabra L. Smith (1988) documents smooth cat's ear, a European
weed, from two counties in south Arkansas. We supply voucher specimens for
his two Atlas reports (*R” for Ashley and Bradley counties) and add eight addi-
tional counties to the record.
=>
icher Ashley Co.: Hooks 253 (NLU). Bradley Co.: Leslie 1756 (NLU). Cleveland Co.: Tho-
mas st 35; 389(NL U). Conway Co.: Edwin B. Smith 4032(UAM). Lafayette Co.: Thomas, Sundell & Amason
127,585 (NLU). Miller Co.: Thomas 160,883 (NLU). Nevada Co.: Thomas & Slaughter 104,341 (NLU,
UAM). Ouachita Co.: Thomas & Doffitt 169,040 (NLU) o.: Thomas, Sundell & Amason 166,304
(NLU). Union Co.: Sundell, Thomas, & Amason 8210 (UAM).
—
Marshallia caespitosa Nutt. In May, 1999, lavender-flowered Barbara’s buttons
was known in Arkansas from eleven occurrences recorded in the Arkansas
Natural Heritage Commission data base. During a field study commissioned
by the Heritage Commission, ten of those populations, scattered in glades and
open, riparian communities in the Ozark and Ouachita highlands, were relo-
cated and several voucher aie made.
—
Voucher specimens (M i it Carroll Co.: Sundell & Wallace 12,617 (UAM)
Madison Co.: Dalton & iy la IARK). Monte Co.: Sundell & Wallace 12,576 (UAM, UARK),
Sundell & Wallace 12,585 (UAM).
Voucher specimens (M var. signata Beadle & Boynt): Perry Co.: Thomas 139,690 (NLU)
Pulaski Co.: Sundell & Wallace 12,574 (u AM)
Tragopogon pratensis |. lacks the inflated peduncles of T. dubius, the common
SUNDELL ET AL., VASCULAR PLANTS FROM ARKANSAS 411
goat’s beard of the roadsides of north Arkansas. T. pratensis is widely estab-
lished in the northern U'S., extending south to Tennessee (Cronquist 1980), but
has not previously been reported from Arkansas.
Voucher specimen: Lawrence Co.: Sundell 10,417 (UAM).
BERBERIDACEAE
Nandina domestica Thunb. A candidate for the most popular ornamental shrub
in south Arkansas, “heavenly bamboo” is not documented as a naturalized spe-
cies of the state flora. It both persists after cultivation and escapes (presumably
dispersed by birds) to open areas and wood margins where it is sufficiently com-
petitive to reach reproductive maturity.
eee
| & McDonald 7619(UAM).
Voucher specimen: Ashley Co.: Sunde
BORAGINACEAE
Cynoglossum zeylanicum (Hornem.) Thunb. ex Lehm. This previously unre-
ported hound’s tongue grew as an aggressive pasture weed in Sevier County in
the foothills of the Ouachita Mountains. In addition, a second collection is re-
ported from sandy soil in Union County. The species is known in the Southeast
from Texas Jones et al. 1997; Diggs et al. 1999) and Louisiana (from Claiborne
and Union parishes along the Arkansas border) (Thomas and Allen 1996). The
burs (mericarps) are 3-4 mm long, smaller than those of C. virginianum and C.
officinale.
Voucher specimens: Sevier Co.: Tyler s.n.(NLU, UAM, UARK). Union Co.: Thomas 116,853 (NLU).
CAMPANULACEAE
Wahlenbergia marginata (Thunb.) DC. was first reported for Arkansas (Thomas
et al. 1991) from a road bank in Union County. The species was rediscovered in
2000 along railroad tracks on the south side of Camden.
Voucher specimen: Ouachita Co.: Thomas & Doffitt 167,860 (NLU).
COMMELINACEAE
Murdannia keisak (Hassk.) Hand.-Mazz. We add two more documented occur-
rences of this easily overlooked, rambling, succulent-stemmed herb, previously
known in Arkansas from two localities (Sundell et al. 1999).
Voucher specimens: Faulkner Co.: Thomas & Sundell 164,429 (NLU). Ouachita Co.: Thomas & Doffitt
63,274 (NLU).
CONVOLVULACEAE
Ipomea x leucantha Jacq. The pink bindweed treated by Fernald (1950) and
Smith (1988) as Ipomoea lacunosa L. forma purpurata Fern. is recognized by
Austin (1978) as a natural hybrid between I. lacunosa and L. trichocarpa EIL (1
cordatotriloba Dennst.). We adda third county record to the two cited in Smith,
all three in the Mississippi Delta of southeast Arkansas.
Voucher specimen: Drew Co.: Sundell & Wiley 8686 (UAM).
412 BRIT.ORG/SIDA 20(1)
Ipomea cordatotriloba Dennst. We add three county records, from agricultural
sites, to the documented Arkansas distribution (Lafayette County, Smith 1988)
of this weedy purple morning glory.
Voucher specimens: Crittenden Co.: Thomas 13,405(NLU, UAM) Jefferson Co.: Thomas 147,277 (NLU,
UAM). Ouachita Co.: Thomas & Doffitt 167,857 (NLU)
CUCURBITACEAE
Citrullus lanatus (Thunb.) Matsum. & Nikai var. lanatus. Six collections from
southeast Arkansas document the status of watermelonat least asa wail in the
state flora.
Voucher specimens: Bradley Co.: Thomas & Amason 141,965 (NLU, UAM),. Calhoun Co.: Sundell et al.
12.5360 UAM). Chicot Co.: Thomas 164,414 (NLU). Lincoln Co.: Thomas & Sundell 167,480 (NLU). Oua-
chita Co.: [homas & Doffitt 167,999 (NLU). Union Co.: Thomas L808 (NLU).
CYPERACEAE
Eleocharis montevidensis Kunth is known from northern Louisiana (Thomas
and Allen 1993) and included in Smith’s Keys to the flora of Arkansas(1994) asa
possible addition to the state flora. We document its occurrence with a collec-
tion from Lake June in Stamps, in southwest Arkansas.
Voucher specimens: Lafayette Co.: Sundell, Thomas, & Amason 12,273 (UAM, UARK).
Websteria confervoides (Poir.) Hooper (Scirpus confervoides Poir in Lam.). Walker
and Campbell (1997) first reported this submersed, aquatic sedge in Arkansas
from a Jefferson County collection. We add two collections from Union County.
The species is rather broadly distributed in Louisiana (twelve parishes, four
along the Arkansas border, Thomas & Allen 1993) and probably more wide-
spread in southern Arkansas than we know.
—
Voucher specimens: Jefferson Co.: Walker 1206961 (UAM). Union Co.: Thomas & Amason 143,312 (NLU,
UAM); Crossland s.n. (UAM).
ELATINACEAE
Bergia texana (Hook.) Seub. ex Walp. Collections from the banks of the Red
River and the Arkansas River represent the third and fourth confirmed occur-
rences of this species in Arkansas.
Vouchet imens: Lafayette Co.: Thomas 120,776(NLU). Lincoln Co.: Lhomas & Sundell 167,.421(NLU).
EUPHORBIACEAE
Caperonia palustris (1) St -Hil. isa wetland weed of tropical American origin
known to occur in the USS. in south Florida, Louisiana and southeast Texas
(Godfrey & Wooten 1981). Smith (1988) has seen specimens from three Missis-
sippi Delta counties in southeast Arkansas to which we add a fourth, where
SS
plants were collected from a rice field.
Voucher specimen: Jefferson Co.: Brady Harmon s.n. (UAM).
Euphorbia hirta L. was found growing as a weed in a [lower bed in southeast
SUNDELL ET AL., VASCULAR PLANTS FROM ARKANSAS 413
Arkansas, its stems ascending above a mat of E. maculata L. The species is pre-
viously unreported for Arkansas.
Voucher specimens: Bradley Co.: Sundell 15,130 (BRIFSMU, NLU, NY, UAM, UARK).
Phyllanthus tenellus Roxb. was reported (Sundell et al. 1999) as a tentative addi-
tion tothe Arkansas flora from weeds escaping their containers at a Drew County
nursery. We adda second locality, Ellis’s Nursery in Fountain Hill, Ashley County,
where again plants arriving as weeds in containers from Louisiana and Texas
have spread aggressively throughout the nursery’s ee: and greenhouses.
Voucher specimens: Ashley Co.: E.& M. Sundell 13,039 (BRIT-SMU, NLU, UAM, UARK).
tT
FABACEAE
Lathyrus aphaca L. Previously documented only from Miller County in south-
west Arkansas (Smith 1988), this unique Eurasian peavine (the stipules function-
ally replace the leaflets) was collected on a roadbank in central Arkansas. Estab-
lished on the West Coast (Isely 1998), the species is apparently taking hold in the
Southeast as well. It was first reported for the region from Tennessee in 1972
(Beardsley and Browne) and recently from Texas in 1997 Jones and Reznicek).
Thomas and Allen (1998) map collections from four Louisiana parishes.
Voucher specimens: Conway Co.: Thomas & Amason 135,545 (NLU, UAM).
Medicago minima (L.) Bartal. A fourth Arkansas county is added to the record
for little bur clover.
Voucher specimen: Washington Co.: Sundell, [homas & Amason 14,155 (UAM).
Trifolium lappaceum L. Lappa clover is a distinctive Old World clover natural-
ized on the Gulf Coast from Alabama to east Texas (Isely 1990). It is reported
here as new to Arkansas from Grandview Prairie, near Columbus, in Hempstead
f-
restol ing this
County. The Arkansas Game and Fish Commission i
blackland prairie site to its original vegetation. Plants were collected in a re-
stored area dominated by compass plant (Silphium laciniatum) and purple and
white prairie clovers (Dalea purpurea, D. candida)—a spring alien among sum-
mer natives.
—
Voucher specimen: Hempstead Co.: Sundell 12.371(UAM).
Trifolium nigrescens Viviani. Ball clover is a sprawling, weedy annual from
southern Europe and southwest Asia recently introduced into the southeast-
ern United States (Isely 1998). Two collections from ruderal sites document its
occurrence and extend its range as a naturalized species in Arkansas. Ball clo-
ver is easily mistaken for alsike clover (I. hybridum) el tet euonc by the
smaller umbels and scarious (rather than I lat tipules
Voucher specimens: Bradley Co.: Thomas & Leslie 92,208 & 974 (NLU, UAM). Drew Co.: Sundell 13
(NLU, UAM, UARK)
414 BRIT.ORG/SIDA 20(1
HYPERICACEAE
Hypericum nudiflorum Michx. Smith (1988) documents this shrubby St. John’s
wort from Drew County in southeast Arkansas. We here confirm his report (no
voucher specimen was seen) of the species’ presence in Ashley County with
two collections, and we document two additional counties.
Voucher specimens: Ashley Co.: Sundell, Hartrick, & Etheridge 7234 (UAM); Thomas 97,093 (NLU).
Poinsett Co.: Thomas 10,314 (NLU). Union Co.: Thomas 107,901 (NLU).
LAMIACEAE
Melissa officinalis L. Smith (1988) records lemon balm as an adventive from
several Ozark and Ouachita Mountain counties. We here make note of a col
tion from the Coastal Plain.
—_
ec-
Voucher speci -G Co.: Thomas & Sundell 164,430 (NLU)
LILIACEAE
Hypoxis curtissii J. Rose (H. leptocarpa (Engelm. & Gray) Small) grows in
swamps and floodplain forests of the Atlantic and Gulf coastal plains (Godfrey
and Wooten 1979). We record it as new to the state from Bodcaw Creek, in south-
west Arkansas, where it occurred with Styrax americana, Planera aquatica,
Fraxinuscaroliniana,and Crataegus opaca under an overstory of Quercus lyrata.
Voucher specimens: Lafayette Co.: Sundell, Thomas, & Amason 12,342 (UAM, UARK).
Lycoris radiata (l’Her.) Herb. With its leaves appearing in autumn after the flow-
ers, red spider lily is nicely adapted to the annual lawn mowing rhythms of
most homeowners and has become a common lawn weed in southeast Arkan-
sas as well as an occasional waif in other disturbed sites. We note its presence in
the Arkansas flora with three collections.
Voucher specimens: Chicot Co.: Thomas 163,666 (NLU). Drew Co.: Sundel! 9,239 (UAM). Ouachita Co.:
Thomas & Doffitt 167,947 (NLU)
NAJADACEAE
Najas minor Allioni. Northeast Arkansas is included by Haynes (1979) within
the range of this distinctive Old World naiad, but neither localities nor speci-
mens are cited. Thomforde’s collection from the shallow margin of a goldfish
pond in central Arkansas (21 August 2000) reconfirms the species’ occurrence
in the state. Recent north Louisiana collections unknown to Haynes (Thomas
S Allen 1993) suggest that Najas minor, which apparently invaded North
America around 1930, may still be extending its range to the west and is likely
more widespread in Arkansas than we know.
Voucher speci : Lonoke Co.: [homforde s.n.(UAM)
NYCTAGINACEAE
Mirabilis jalapa L. Reported by Leslie (1986) but excluded by Smith (1988) as a
naturalized element in Arkansas, four o’clock is reinstated with roadside col-
—_—
SUNDELL ET AL., VASCULAR PLANTS FROM ARKANSAS 415
lections from the Ozark Highlands and the Coastal Plain. At none of the sites
did plants appear to be merely persisting after cultivation. Four o’clock is
mapped by Thomas and Allen (1998) from six of eight of Louisiana’s northern
tier of parishes.
Voucher specimens: Bradley Co.: Thomas & Amason 142,066 (NLU, UAM). Newton Co.: Thompson
685 (NLU). Union Co.: Thomas & Amason 111373 (NLU, UAM).
POACEAE
Andropogon ischaemum L. var. songaricus Rupr. ex Fisch. & Mey. (Bothriochloa
ischaemum (L.) Keng var. songaricus (Rupr.) Celerier & Harlan), an intrusive,
pernicious Eurasian bluestem, is common to the roadsides and pastures of the
southern Great Plains (Gould 1975, Great Plains Flora Association 1986). It has
become widespread in Louisiana (Thomas & Allen 1993) and an occasional
weed east to Florida (Wunderlin 1998). We add three counties to its documented
occurrence (Franklin Co.) in Arkansas.
Voucher specimens: Drew Co.: Sundell 15,128(NLU, NY, UAM, UARK). Miller Co.: Miesners.n.CUAM).
Ouachita Co. Thomas & Doffitt 168.023 (NLU).
Eleusine tristachya (Lam.) Lam., a diminutive goosegrass of African origin, is
known as a waif in North America from a few scattered localities (Hitchcock
and Chase 1950, Kucera 1998) and is apparently naturalized in California (Smith
1993). We report it as a new record from northeast Arkansas, where it occurred
as a lawn weed.
Voucher specimen: Independence Co.: S.D. Carter s.n.(UAM).
POLYGONACEAE
Fagopyrum esculentum Moench. Buckwheat spreads easily from cultivation to
waste places but seldom persists (Great Plains Flora Association 1986,
Steyermark 1963). Collections from a small population at the edge of a campus
parking lot mark the species’ first documented occurrence as a waif in Arkan-
sas.
Voucher specimens: Drew Co.: Sundell 12,567 (UAM, UARK).
Polygonum setaceum Baldwin. Godfrey and Wooton (1981) include Arkansas
within the range of this widespread smartweed, and Smith (1988) made note of
the species as a possible addition to the state flora based on its occurrence in
several Louisiana parishes along Arkansas’ southern border (MacRoberts 1988).
We confirm its presence with voucher specimens from three counties.
Voucher specimens: Chicot Co.: Thomas 163,358 (NLU). Hot Spring Co.: Sundell & Crank 10,733 (UAM).
Union Co.: Thomas 112,374 (NLU).
RUBIACEAE
Diodia dasycephala C.MS. Especially in the field, “perennial poorjoe” so closely
resembles Spermacoce glabra (rather than either of the common, congeneric
416 BRIT.ORG/SIDA 20(1)
buttonweeds, D. teres and D. virginiana) that it has perhaps been overlooked
and misidentified as often as any taxon in southern Arkansas. The two can be
distinguished vegetatively. Leaves of the Diodia are shorter and dry greener;
those of the Spermacoce are more prominently veined beneath. Spermacoce
glabra bears numerous, whitish scalelike hairs at the stipular sheath summit
eae and between the bristles while D. dasycephala ee a (more or
less) prominent vein that traverses the broader summit of the glabrous stipular
sheath just below its bristle-bearing margin. Diodia dasyce phala is documented
from six parishes in north Louisiana (Thomas & Allen 1998) and two counties
in southeast Arkansas (Smith 1988). We add three more counties to the record,
one from central Arkansas.
=n
Voucher specimens: Chicot Co.: Thomas 164,170 (NLU). Faulkner Co.: Sundell & Thomas 15,954 (NY,
UAM, UARK). Lafayette Co.: Thomas, Sundell, & Amason 156,442 (NLU)
SCROPHULARIACEAE
Lindernia crustacea (L.) F von Muell. isa wetland species, originally from south-
ern Asia, that occurs sporadically in the coastal plain of the Southeast (Godfrey
S& Wooten 1981). Our collection represents the second documented occurrence
in Arkansas.
Voucher specimen: Pulaski Co.: Sundell 12,437 (0AM).
Scoparia dulcis |. Thomas and Amason collected this tropical American weed
in a clearcut. It is known regionally from the Gulf Coast states (Correll &
Johnston 1970) but unreported for Arkansas.
319 (NLU, UAM).
Qo
Voucher specimens: Cleveland Co.: Thomas & Amason 158
Veronica hederifolia L. is confirmed by Smith (1988) for two northwest Arkan-
sas counties. We add four more counties to the record of this relatively distinc-
tive speedwell.
Voucher specimens: Chicot Co.: [homas 164,620 (NLU). Mississippi Co.: Barbee 24 (UAM). Pulaski
Co.: Burgess & Bernstein OO] (UAM). Saline Co.: Thomas & Amason 148,563 (NLU).
Veronica polita Fries appears sufficiently distinct from V. agrestis to be recog-
nized as a separate species. Specimens seen by Smith (especially at UARK), who
merges the two in the Acleatisas Atlas 1988), should probably be reexamined
for additional material of the former species. We document the presence of V.
in Arkansas from four counties.
Vouche Ba « Hyatt 1063.03 (UAM). Drew Co.: Sundell 10,755 (UAM). Garland Co.:
Demarece 59, 005(NL U). ie oe Co.: Sundell 12,263 UAM).
VERBENACEAE
Verbena montevidensis Spreng. Thomas and Allen (1998) record this slender
vervain [rom three parishes in central and eastern Louisiana. Four collections
from south Arkansas suggest that it may have become established in this state
as well.
SUNDELL ET AL., VASCULAR PLANTS FROM ARKANSAS 417
Voucher specimens: Ashley Co.: Thomas and Hooks 92,761 (NLU, UAM). Bradley Co.: Thomas 90,519
(NLU). Chicot Co.: Thomas 166,906 (NLU). Union Co.: Thomas & Amason 111,853 (NLU).
ZANNICHELLIACEAE
Zannichellia palustris |. Horned pondweed is documented from three counties
in the Ozark Plateau of northeast Arkansas (Smith 1988). Two collections of the
submersed aquatic from central and southwest Arkansas extend its range to
the Mississippi River Alluvial Plain and West Gulf Coastal Plain.
Voucher specimens: Howard Co.: Lawson 2117 (NLU). Lonoke Co.: Thomforde sn. (UAM).
ACKNOWLEDGMENTS
—
The authors express their appreciation to Isabel Bacon of the School of Arts
and Humanities, University of Arkansas at Monticello, for translating the En-
glish abstract into Spanish. Thanks also to Ralph W. Tyler of the Sevier County
Extension Service for his efforts in collecting additional specimens of
Cynoglossum zeylanicum, to Hugh Thomforde of the Lonoke Agricultural Cen-
ter for his collections of Najas minor and Zannichellia palustris, and to Judy
Griffith of Ninestone Land Trust in Carroll County for leading us toa splendid
population of Marshallia caespitosa in full bloom.
REFERENCES
Austin, D.F. 1978. The Ipomoea batatas complex-l. Taxonomy. Bull. Torrey Bot. Club 105:
114-129.
Bearps.ey,R.L.and E.T. Browne, Jr. 1972. Lathyrus aphaca L.new to Tennessee and the South-
east. Rhodora 74:155.
Corrett, D.S. and M.C. JoHNsTon. 1970. Manual of the vascular plants of Texas. Texas Re-
search Foundation, Renner
Cronauist, A. 1980. Vascular flora of the southeastern United States. Vol. 1. Asteraceae. The
University of North Carolina Press, Chapel Hill.
Diccs, G.M., Jr., B.L. Lipscome, and R.J. O’Kennon. 1999. Shinners and Mahler's illustrated flora
of north central Texas. Botanical Research Institute of Texas, Fort Worth and Austin
College Center for Environmental Studies, Sherman, Texas.
FerNatp, M.L. 1950. Gray's manual of botany, ed.8. American Book Company, New York.
Goorrey,R.K.and J.W. Wooten. 1979. Aquatic and wetland plants of the southeastern United
States. Monocotyledons. The Univ. of Georgia Press, Athens.
Goorrey,R.K.and J.W.Wooren. 1981.Aquatic and wetland plants of the southeastern United
States. Dicotyledons. The Univ. of Georgia Press, Athens.
Goutb, FW. 1975. The grasses of Texas. Texas A & M University Press, College Station.
Great PLains Flora Association. 1986. Flora of the Great Plains. University Press of Kansas,
Lawrence.
Haynes, R.R. 1979. Revision of North and Central American Najas (Najadaceae). Sida 8:
34-56.
HitcHcock, A.S.and A.CHase. 1950.Manual of the grasses of the United States, ed. 2. (United
BRIT.ORG/SIDA 20(1
=
States Department of Agriculture, Miscellaneous Publication No. 200.) United States
Government Printing Office, Washington, D.C
HOLMGREN, P.K., N.H. Houmoren, and L.C. Barnett. 1990. Index herbariorum. Part |: The herbaria
of the world, ed. 8. New York Botanical Garden, Bronx.
IseLy, D. 1990. Vascular flora of the southeastern United States. Vol. 3, Part 2. Leguminosae
The University of North Carolina Press, Chapel Hill.
IseLy, D. 1998. Native and naturalized Leguminosae (Fabaceae) of the United States. Monte
L. Bean Life Science Museum, Provo, Utah.
Jones, S.D. and A.A. Reznicek. 1997. Lathyrus aphaca (Fabaceae), previously unreported for
Texas. Phytologia 82:1-2.
Jones, S.D., J.K.Wipff, and PM. Montcomery. 1997.Vascular plants of Texas. University of Texas
Press, Austin.
Kucera, C.L. 1998. The grasses of Missouri, rev. ed. University of Missouri Press, Columbia.
Lestit,S.A.1986.A preliminary survey of the vascular flora of Bradley County, Arkansas. M.S.
Thesis, Northeast Louisiana University [University of Louisiana at Monroe].
MacRoserts, D.T. 1988.A documented checklist and atlas of the vascular flora of Louisiana.
Part Ill. Dicotyledonae. Fagaceae to Zygophyllaceae. Louisiana State University in
Shreveport.
Smith, E.B. 1988. An atlas and annotated list of the vascular plants of Arkansas, ed. 2. Pub-
lished by the author, Fayetteville, Arkansas.
Smith, E.B. 1994. Keys to the flora of Arkansas. The University of Arkansas Press, Fayetteville.
SMITH, J.P., Jk. 1993. Poaceae. In: J.C. Hickman, ed. The Jepson manual: higher plants of Cali-
fornia. University of California Press, Berkeley.
STEYERMARK, J.A. 1963. Flora of Missouri. The lowa State University Press, Ames.
SUNDELL, E.,R.D. THomas, C. AMason, R.L. Stuckey,and J.Locan. 1999. Noteworthy vascular plants
from Arkansas. Sida 18:877-887.
THomas, R.D., E.B. Smity, E. Sunpett, RE. Hyatt and C. Amason. 1991. Additions to the flora of
Arkansas. Sida 14:483-491,
THomas, R.D. and C.M. Aten. 1993. Atlas of the vascular flora of Louisiana. Vol. |. Louisiana
Department of Wildlife and Fisheries, Baton Rouge.
THomas, R.D. and C.M. Aten. 1996. Atlas of the vascular flora of Louisiana. Vol. Il. Louisiana
Department of Wildlife and Fisheries, Baton Rouge.
THomas, R.D. and C.M. Atten. 1998. Atlas of the vascular flora of Louisiana. Vol. Ill. Louisiana
Department of Wildlife and Fisheries, Baton Rouge.
Wacker, S.A. and J. Campsett. 1997, A floristic survey and annotated checklist of the Pine
Bluff Arsenal. J. Arkansas Acad. Sci. 51:178-187,
Wunbertn, R.P. 1998. Guide to the vascular plants of Florida. University Press of Florida,
Gainesville.
COMMELINA BENGHALENSIS (COMMELINACEAE)
NEW TO NORTH CAROLINA AND AN UPDATED KEY
TO CAROLINA CONGENERS
Alexander Krings Michael G. Burton
Herbarium, Department of Botany Department of Crop Science
North Carolina State University North Carolina State University
Raleigh, NC 27695-7612, U.S.A. Raleigh, NC 27695-7620, U.S.A
Alexander_Krings@ncsu.edu Mike_Burton@ncsu.edu
Alan C. York
Department of Crop Science
pot Carolina State University
a NC 27695-7620, U.S.A.
n_York@ncsu.edu
ABSTRACT
A population of the federal noxious weed C lina benghalensis L.(C li was recently
fo) ie /
identified in a soybean field in North Carolina. Voucher sp land an updated key to
Commelina in the Carolinas is presented.
RESUMEN
[apa] on. Lh) F a le esge ees } 7 tee |
| Commelina beng halensis L. oe ae)
en un campo de setaill a de soja en Carolina del Norte. Ci
I ya
nueva para Commelina en las Carolinas
A common paleotropical weed, Commelina benghalensis L. (Tropical spiderwort,
Benghal dayflower) has been previously collected in the southeastern United
States only in Florida, Georgia, and Louisiana (Faden 1993; Thomas & Allen
1993). The taxon has not been reported for North Carolina by Radford et al.
(1968), Kral (1981), or Faden (1993, 2000). Federally listed as a noxious weed in
1983 (Lasseigne 1983), C. benghalensis has primarily been found in citrus plan-
tations, fields, yards, and other cultivated and disturbed sites in the southeast
as far north as Georgia, as well as in California (Faden 1993). During a field sur-
vey, a population of this weed was identified in a soybean field in Goldsboro,
Wayne County, in the Coastal Plain of North Carolina. The infestation occurs
over an area of less than 2 ha, north of SR 1008 and near the Southern Railway.
The density is quite high at the edges of the crop plots and lower in neighbor-
ing (intensively managed) cotton and soybean plots. The weed was first no-
ticed about three years ago, but was thought to bea different species (Commelina
caroliniana Walter occurs in a neighboring field). Representative material was
prepared for deposit at NCSC, US, and USCH in Oct 2001.
SIDA 20(1): 419 — 422. 2002
420 BRIT.ORG/SIDA 20(1)
a
pete
pone eee: a
Peoria ert ‘cai
mo |
on
\
\ I
NY Cathe Faequale’
Fic.1 li b hal is L.A, Whole plant, I i g I g | 1 bel g 1infl B Leaf sheath:
Po c phe ent a as;D, ies tea f r he, showi gk iti f fl E Imperfect
patfecehlowei 6G: | 1 fruit; H, Dissection of seed f k d fruit. A-B based on Tsiang 2498
(NY); C-H based on Seydel 203 (Mo). Illustration by Cathy Pasquale, courtesy of the U.S. Dept. Agric., Animal and Plant
Health Inspection Service.
Voucher specimens: U.S.A. NORTH CAROLINA. Wayne Co.: Goldsboro, in a soybean field, 35.397 deg
ut
N, 78.046 deg W, 27 Sep 2001, Burton, Hoyle, and Mendoza s.n. (NCSC, US, USCH).
Commelina benghalensis L. Fig. 1) can be distinguished from its southeastern
congenerics by the often clustered, infundibuliform spathes, the relatively broad
leaves, often bearing reddish hairs on the sheath apex, and the basally-borne,
KRINGS ET AL
421
sometimes subterranean cleistogamous flowers in addition to chasmogamous
flowers. An updated key, largely adapted from Faden (2000) follows:
KEY TO COMMELINA IN THE CAROLINAS
. Spathes with margins connate basally.
2. Roots stout; leaf sheaths auriculate apically; proximate petal white, minute
locules all 1-seeded Com meliaa erecta L.
2. Roots thin; leaf sheaths not eg proximate petal blue to lilac or lavender,
conspicuous; some locules 2-seeded.
3. Perennial with erect to ce stems; lamina lanceolate-elliptic to lan-
ceolate oblong, the apex acuminate; spathes 1.5-3.5 cm long; subterranean
cleistogamous flowers absent Co mmelina virginica L.
3. Annual with eecenalhig to decumbent or occasionally scrambling stems;
|
lamina ovate to lanceolate-elliptic, the apex rounded, obtuse, or acute; ue
0.5-2 cm long; subterranean, cleistogamous flowers usually presen
___ Commelina
benghalensis L.
1. Spathes with margins free to base.
4. Spathes typically discolored whitish or pale green near peduncle with contrast-
ing, darker green veins; proximal petal whitish or paler than the others; capsules
2-locular; seeds rugose pitted-reticulate Commelina communis |.
4. Spathes uniformly colored, without contrasting veins; proximal petal essentially
colored as the others; capsules 3-locular; seeds reticulate or smooth to some
what alveclate.
5. Spathes typically distinctly falcate; distal cyme in larger spathes usually well-
developed, 1-several-flowered; seeds reticulate melina diffusa Burman f.
5. Spathes not falcate to only slightly so; distal cyme in larger spathes usually
vestigial (rarely well-developed, 1-flowered); seeds smooth to somewhat
alv
eolate Commelina caroliniana Walter
ACKNOWLEDGMENTS
We thank: Jamie Hinton and Steve Hoyle (North Carolina State University) for
their attention to detail; Gene Cross and David Patterson (NC Dept. Agric. &
Consumer Services) for assistance in obtaining necessary transport permits;
Rodney Young (APHIS) and Robert Faden (US) for confirming our identifica-
tion; Robert Faden and an anonymous reviewer for thoughtful comments on
the manuscript.
REFERENCES
Fapen, R.B. 1993. The misconstrued and rare species of Commelina (Commelinaceae) in
the eastern United States. Ann. Missouri Bot. Gard. 80:208-218.
Fapen, R.B. 2000. Commelina. |n: Flora North America Editorial Committee, eds. Flora North
America, Vol. 22.Oxford University Press, New York. Pp. 192-197.
Kral, R. 1981.Some distributional reports of weedy or naturalized foreign species of vas-
cular plants for the southern states, particularly Alabama and middle Tennessee. Cas-
tanea 46:334—-339.
422 BRIT.ORG/SIDA 20(1)
Lasseicne, A. 1983. Noxious weeds of the Federal Noxious Weed Act,No.21:Commelina sp.
U.S. Dept. Agric., Washington, D.C
Raproro, A.E., H.E. AHies, and CR. Bett. 1968. Manual of the vascular flora of the Carolinas.
The University of North Carolina Press, Chapel Hill.
THomas, R.D. and C.M. Atten. 1993. Commelina benghalensis L. (Commelinaceae), Carex
hyalina Boott (Cyperaceae) and Chloris subdolichostachya C. Muell. (Poaceae): New to
Louisiana. Phytologia /4:336-338.
AGROSTIS AVENACEA (POACEAE: POOIDEAE):
FIRST RECORD FOR THE MEXICAN FLORA
Arely Nava-Rojo Manuel Gonzdlez-Ledesma
and Maricela Gomez- Sanchez! Centro de | ena ones IOI:
ni i ) Biologia | |rivere sid at A rl Hi 4.
LI nclatura ! gid lidalgo
Universidad Auténoma de Querétaro Apdo. Postal 1-69, Bana Judrez
Cerro de - Campanas s/n 42001 Pachuca, Hgo., MEXICO
Cent iversit 010 Querétaro, Qro., MEXICO
ee
ABSTRACT
Agrostis avenaced J.F Gmel. is reported from temporary and permanent ponds of the region of
Huimilpan and Amealco of the state of Queretaro, Mexico. Previously, it was only known [rom Aus-
tralia, south ue Argentina, ee U sia State of aie. the pawauay Islands and pean)
rom Costa Rica 1
and collecti : ta and Scachien epecumeigate cited.
io] I
RESUMEN
Agrostis avenaceaJ.F. Gmel.se cita de charcos temporales y permanentes ne la region de Huimilpan y
Amealcoenel estado de Querétaro, México ustralia, sur de Africa,
ig asec Chile, Estados . midds de América, ne as Hawa) y recientemente de Costa Rica. Se
HF
llustraciones y se citan los dat ALTOS
a a colectas y especimenes esenaadss
INTRODUCTION
Agrostis is a genus of grasses that was described by J.-F Gmelin in 1791. The ge-
nus is included in the Aveneae tribe and Alopecurinae subtribe (Pohl & Davidse
1994; Valdés-Reyna & Davila-Aranda 1995). As presently circumscribed, the
genus includes about 125-220 species (Acosta-Castellanos 1990; Beetle 1983;
Clayton & Renvoize 1986; Mc Vaugh 1983; Watson & Dallwitz 1992) which grow
in temperate and cold regions from both hemispheres. In the tropics, they are
mostly restricted to the high altitude mountains. Currently, the most recent
list of species for Mexico (Espejo-Serna et al. 2000) shows that 23 species are
registered for Mexico. While conducting an aquatic flora project of the Queretaro
state Mahinda Martinez made a collection of subaquatic grasses from the
Huimilpan and Amealco municipalities that proved to be Agrostis avenacea J.F
Gmel. This collection represents the first record of A. avenacea from Mexico
and increases the number of species to 24.
'Author for Correspondence
SIDA 20(1): 423 - 429. 2002
424 BRIT.ORG/SIDA 20(1
Agrostis avenacea J.F. Gmel., Syst. Nat. 2(1):171. 1791. Calamagrostis avenacea JE
Gmel.) Bech., ee 7519. 1938. Lachnagrostis avenacea J. Gmel.) Veldkamp,
Blumea 37:230. 1
Agrostis debilis Poir, Encycl. Suppl. i249. 1810. Vilfad
-( Dp,
ir.) P Beauv, Ess. Agrostogr. 16, 147,
Ag rostt: forsteri Roem. ex Schult Syst. Veg. 2:359. 1817
ee aaa Parodi, — Argent. Agron. 29(1-2):19. 1962.
Agrostis ligulata Steud., Syn. Pl. Glumac. 1:173. 1854.
Agrostis eens Willd. Enum. Pl. 1:94. 1809. Lachnagrostis Salaes (Willd.) Trin. Fund.
Agrost. 128, 1820. Vilfa retrofracta (Willd. P. Beauv. Ess. Agrostogr. 182.1812. Deyeuxia
a Sri (Willd.) Kunth, Rev. Gramin. 1:77. 1829. Calamagrostis ret hae (Willd.) Link,
Hortus Berol. 2:247.1
Avena filiformis G. Forst. F ‘ Ins. Aust. Prodr. 9. 1786. Agrostis filiformis(G. Forst.) Spreng., Prim. FI.
Hal. 32, 1807. AU el a a Forst.) Trin., Fund. Agrost. 128, t. 10, 1820.
Calamagrostis filiformis (G. Forst.) Rept. Bot. Surv. oe Natt. Park 35, 1908. Deyeuxia
filiformis (G. Forst.) Pe trie ot Isl. an 19
Deyeuxia oie ae Revis. Gramin. 1:77. 182¢
h ill ii Trin.,Gram. Unifl. en 217.1824.C grostis wi ii Trin.)
Send, Syn. Pl ¢ G jonas: 1:192. 1854
—
Plants annual, cespitose. Culms erect, thin, breakable, 20-60 cm tall, 1-2 mm
broad. Leaf blade flat, 5-20.cm long, 1-3 mm broad. Ligule 2-5 mm long. panicle
open, lax, 15-30 cm long; branches thin, slender, inferiors as longer as inflores-
cence axis, spikelets on the tips of the branches. Glumes acute, 2-5 mm long, l-
nerved, scabrous on the keel. Lemma membranous, pilose, 1.4-16 mm long, 5-
nerved, with a mid-dorsal once-geniculate awn. Palea membranous, |.1-1.3 mm
long, nearly as long as lemma, 2-nerved, tip bifid, callus pubescent. Chromo-
some number: 2n = 28. (Fig. [A-F)
Habitat.— Agrostis avendacea grows on dry to moist soils, with abundant clay
and from 15 to 2350 m in elevation. In Mexico, it is grows around small streams,
temporary and permanent ponds, and at sites at 2280-2350 m in elevation. it is
found as a member of the subaquatic vegetation (Fig 2B).
Distribution.—This species is native to Australia (Canberra, New South
Wales, Queensland and Tasmania) and New Zealand. In the Missouri Botanical
Garden's herbarium and VAST nomenclatural database (W* TROPICOS)
(www.mobot.org), there are records of its introduction in South Africa, Argen-
tina (Buenos Aires, depression of El Salado, Entre Rios and Santa Fé) and Chile
(Isla de Pascua) (Nicora & Rugolo 1987), as well as, some temperate regions of
the United States of America (California, Texas and Ohio), and the Hawaiian
Islands (Hawaii, Kauai, Molokai, Oahu) (Hitchcock 1950). Recently, it was re-
corded from San José, Costa Rica (Davidse 1998). In Australia, it isan abundant
weed of inundated places (Vickery 1941). Nicora and Rugolo (1987) and Zuloaga
et al. (1994) recognize this taxon in the segregate genus Lachnagrostis
In Mexico, Agrostis avenacea was collected from the municipalities of
Huimilpan and Amealco, in the southern portion of the state of Queretaro (Fig.
—
NAVA-ROJO ET Al 425
Fic. 1. Agrostis avenacea J.F. Gmel. (M. Martinez 2936, QMEX).A. Habit. B. Floret. C. Caryopsis. D. Spikelet. E. Inflorescence
branches. F. Indument of inflorescence branches. G-H. Leaf blade anatomy. G. Detail of transverse section (M. Martinez
2936, QMEX). H. Abaxial epidermi in surface view (M. Martinez 2953, QMEX). Stippling indi t tal
/ rr
gion in H.
426 BRIT.ORG/SIDA 20(1)
2A). This species has not been previously reported in any floristic study from
Mexico. This taxon was introduced to U.S.A (Hitchcock 1950; Davidse et al. 1998).
In California and texas, plants of A.avenacea act like tumble weeds and its dried
panicles are carried widely by wind and in the process dispersing seeds. It is
unknown whether the populations of A. avenaced from Mexico is the result of
long-range wind dispersion, or by some other manner, such as by vehicles pass-
ing through wetlands and picking up spikelets and panicles. It is probable that
this taxon is growing in other states of Northern Mexico, as well.
a
Specimens examined: MEXICO: Querétaro: Mpio. Huimilpan, arroyo La Beata, km 42 carretera
Querétaro-Amealco-La Beata, 20° 21'LN y 100° 13' LW, 2350 msnm, 15 May 1996, Martinez 2936. Mpio.
Huimilpan, “La Calle,” km 33 Carretera Querétaro-Amealco, hacia las alueras de Huimilpan, 20° 22.5'
LN, L00'16.5' LW, 2280 msnim, 15 May 1996, Martinez 2952, 2953, 2955; Mpio. Amealco, km 3l carretera
Amealco-San Juan del Rio, 2 km al F de Amealco, charco See a la orilla de la carretera, 20
2.84 LN, 100° 9.16' LW, 2490 msnm, 25 Jul 1996, Martinez 3331, 3333; Mpio. Amealco, km 5 carretera
Amealco-San Ildelfonso, charco temporal, 20° 10.80' LN, 100° 5.40' LW, 2040 msnm, 5 Nov 1996,
Martinez 3667 (QMEX, duplicates for distribution to [EB and MEXU)
Agrostis dvenaced is distinguished by its slender, thin and lax inflorescences
The spikelets of mature inflorescences are easily dispersed by the wind.
Leaf blade anatomy
The abaxial epidermis and inner structure of the leaf blade were studied. The
epidermis and transverse sections were obtained following Gomez-Sanchez and
Koch (1998).
Abaxial epidermis.—Differentiation of the costal and intercostal zones incon-
spicuous. Long cells, fusiform, longer than wide. Stomata abundant, 37-39 pm
long, parallel-sided subsidiary cells, guard cells hidden by interstomatal long
cells. Papillae absent. Prickles a few in the costal zone. Silica bodies, microhairs
and nee absent (Fig. 1H).
Transverse section.—Outline of the blade U-shaped, nodular. Abaxial and
adaxial surface with rounded ribs associated with all vascular bundles; abaxial
and adaxial furrows broad and deep. Vascular bundles nine; usually first order
vascular bundles. Outer vascular bundle sheaths absent. Sclerenchyma costal
strands present, the abaxial in 2-4 cells wide and 1-2 cells tall, the adaxial in 1-
3 cells wide and 1-2 cells tall or absent. Chlorenchyma nonradiate, its cells of
various sizes and shapes, continuous between adjacent vascular bundles. Col-
orless parenchyma cells absent. Bulliform cells in fan-shaped, adaxial groups
no conspicuous, central cell relatively small, or absent (Fig. 1G).
Agrostis avenaced is clearly a member of the Pooideae in its anatomical fea-
tures. Itisa Non-Kranz species, with C3 photosynthetic metabolism (Hattersley
S& Watson 1975). It has a typical “pooid” anatomy, characteristic of those spe-
cies that grow in temperate climates. The leaf blade is nodular with ribs and
deep furrows in the abaxial and adaxial surfaces. The limited development of
NAVA-ROJO ET AL
Huimilpan
HU Amealco
ERD AG L
Fic. 2. Regi
~s
ftho Ctat ££ A+ ; Maeyico. 2B. T
428 BRIT.ORG/SIDA 20(1)
abaxial and adaxial sclerenchyma explains the presence of the breakable and
lax culms and suggests that Agrostis avenacea is adapted to wet habitats.
Several taxonomic treatments of Agrostis exist (Acosta-Castellanos 1990;
Pohl 1980; Pohl & Davidse 1994; Davidse 1998), however these treatments are
partial and taxonomic affinities of the species are uncertain. The genus Agrostis
urgently needs a world revision (Pohl & Davidse 1994: Davidse 1998). In addi-
tion to the morphological, anatomical, micromorphological work, the develop-
ment of molecular studies will be a very important contribution to circum-
scription of Agrostis species.
ACKNOWLEDGMENTS
We are grateful to the curators of the ENCB, IEB, MEXU and QMEX herbaria
who so kindly accommodated our visit, to Mahinda Martinez who provided
the collections, to Clara Tinoco who prepared Figure |A-F and to Robert Jones
and an anonymous reviewer for helpful comments. Research was supported by
CONACYT through 29106-N project.
REFERENCES
AcosTa-CASTELLANOS, S, 2001. Agrostis. In: Rzedowski, J. and Graciela C. de Rzedowski, eds.
Flora Fanerogamica del Valle de México. Instituto de Ecologia, Centro Regional del
Bajo. Patzcuaro, Mich. México. Pp. 1014-1017.
Beettie, A.A. 1983. Las gramineas de México. Secretaria de Agricultura y Recursos
Hidraulicos- COTECOCA. México.
Crayton, W.D. and S.A. Renvoize. 1986. Genera graminum. Grasses of the World. Kew Bull,
Add. Ser. 13. London.
Davipse, G. 1998. Agrostis. In:G. Davidse, M. Sousa S.,and S. Knapp, eds. Flora Mesoamericana.
Version Internet (W?FM). Universidad Nacional Autonoma de México. México
EsPEJO-SERNA, A., A.R. Lopez-Ferrari, and J. VaLbes-Reyna. 2000. Poaceae. In: Espejo-Serna A.and
A.R.LOpez-Ferrari,eds.Las Monocotiledoéneas Mexicanas, una Sindpsis Floristica. |. Lista
de Refrencia. Parts IX—XI. Consejo Nacional de la Flora de México. Mexico. Pp. 8-14.
Gomez-SAncHez, M. and S.D. Koch. 1998. Estudio anatomico comparativo de la lamina foliar
de Eragrostis (Poaceae: Chloridoideae) de México. Acta Bot. Mex. 43:33-56.
Harterstey, PW. and L.Watson. 1975, Anatomical parameters for predicting photosynthetic
pathways in grass leaves. Phytomorphology 25:325-333.
HircHcock, A.S.and A. Case. 1951. Manual of the grasses of the United States. U.S. Dept. of
Agric., Misc. Publ. No. 200: 1051 pages (Second edition, revised by A. Chase)
McVaush, R. 1983. Flora Novo-Galiciana. 14. Gramineae. University of Michigan Press. Ann
Arbor. Missouri Botanical Garden: http://mobot.mobot.org/W3T/search/vast.htm|
Nicora, E.G.and Z.E.Rucoto be Acrasar. 1987.Los géneros de gramineas de América Austral.
Hemisferio Sur. Buenos Aires, Argentina.
Port, R.W. 1980. Flora Costaricensis. Family # 15. Gramineae. Fieldiana Bot. n.s. 14:1-607.
NAVA-ROJO ET AL 429
Pou, R.W.and G. Daviose. 1994. Agrostis. In: Davidse, G.,M. Sousa S.and A.O. Chater, eds. Flora
Mesoamericana. Vol. 6. Universidad Nacional Aut6noma de México. México. Pp.
237-240.
Vatoes-Reyna, J. and P. DAvita-Aranpa. 1995. Clasificacidn de los géneros de gramineas
Mexicanas. Acta Bot. Mex. 33:37-50.
Vickery, J.W.1941.A revision of the Australian species of Agrostis L.Contr.N.S.W. Natl. Herb.
1:101-119.
Watson, L. and M.J. Datiwitz. 1992. The grass genera of the World. C.A.C. International.
Wallingford.
ZULOAGA, F.O., E.G. Nicora, Z.E. Rucoto pe Acrasar, O. Morrone, J. Pensiero, and A.M, CIALDELLA.
1994. Catalogo de la Familia Poaceae en la Republica Argentina. Monogr. Syst. Bot.
Missouri Bot. Gard. 47:1-178.
430 BRIT.ORG/SIDA 20(1)
Books RECEIVED/ NOTICES
Annotations and comments by the Editor
ANCILLE GLOUDEN and Cicely Tosiscu. 1995, Orchids of Jamaica. (ISBN 976-640-
002-4, pbk.). The University of West Indies Press, la Aqueduct Flats, Kingston
7, Jamaica, WI (Orders: University of Oklahoma Press, 4100 28th Avenue,
Norman, OK 73069-8218; www.oupress.com, 405-307-9048 fax). $26.00,
b
222 pp. numerous color photos, 6" x 9",
Key Words: Orchids, Jamaica, flora, field guide, identification, descriptions, 125 species.
FRITZ HOCHSTATTER. 2000. Yucca I (Agavaceae). (Dehiscent-Fruited Species) in the
Southwest and Midwest of the USA and Canada with Seed Morphology by
Dr. Ginter Hentzschel. English Translation by Chris Holland. (ISBN 3-00-
005946-6, hbk.). Fritz Hochstatter, PO. Box 510201,D-68242 Mannheim,
Germany; Hamlyn Valley, Box 11, Modena, UT 84753, U.S.A. (Orders:
{hnavjo@aol.com; members.aol.com/fhnavajo; www.cactus-mall.com /navajo:
www.cactus-mall.com/cdrom/yucca.html; 0621-79-00332 fax). Price not
given, 256 pp. 189 figures, with many color photos, line drawings, and maps,
6 3/4" x 93/4".
Key Worps: Yucca, Agavaceae, flora, systematics, descriptions, illustrations, keys, morphology,
pals
anatomy of Yucca seeds, ecology, pollination, USA, Canada, reproductions of original descriptions.
At least two new names are proposed in the book, Yucca harrimaniae Trelease ssp. gilbertiana
(Trelease) Hochstatter stat. nov. (page 36) and Yucca arkansana Trelease ssp. freemanii (Shinners)
Hochstatter, comb. et. stat. nov. (page 61).
Horticulture/Gardening
BARBARA Perry LAWTON. 2002. Mints: A Family of Herbs and Ornamentals. (ISBN
0-88192-524-1, hbk.). Timber Press, Inc., 133 S.W. Second Avenue, Suite 450,
Portland, OR 97204-3527, US.A. (Orders: www.timberpress.com, 800-327-
5680, 503-227-2878, 503-227-3070 fax). $27.95, 272 pp., 61 color photos, 5
b/w photos, +6 line drawings, 6" x 9"
Kry Worbs: Lamiaceae, mints, herbs, ornamentals, gardening, cultivation, classification, catalog, 67
genera
SIDA 20(1): 430. 2002
AN UPDATE TO THE VASCULAR FLORA OF
CALCASIEU PARISH, LOUISIANA
Ray Neyland
Department of Biological & Environmental Sciences
McNeese State University
Lake Charles, LA 70609, USA
ABSTRACT
The publication of a vascular flora survey of Calcasieu Parish, Louisiana by Neyland et al. in 2000 is
updated and revised. Subsequent nelGs and Herban ium research has yielded the discovery of four new
that had been previously reported by other sources to
species for the pat ish. Additionally
t
occur the parish but were not discovered during Neyland et als original survey have been collected.
Two corrections from the original survey are also noted.
Key Worps: Vascular Plant Survey, Calcasieu Parish, Louisiana
RESUMEN
blicati | io de flora vascular de Calcasieu Parish, Louisiana por Neyland etal.en
2000 se pone al dia y se revisa. La see ane b le campo y herbario ha tenido como
5
consecuencia el cacaauamand de suatr s especies para par ogee Adicionalmente, se han
I | lo citadas previ I es pane perono
- ] ye ] Las tien a Viel CNT ] ] 1. Cat
al catalogo original.
This paper updates and revises the original vascular flora survey of Calcasieu Par-
ish, Louisiana of Neyland et al. (2000). Specifically, the purpose of this study is to
list new collections from the Parish and to correct errors from the original survey.
METHODS
Since the publication of the original survey by Neyland et al. (2000) until
present, plant collections from Calcasieu have continued. Voucher specimens
were prepared using standard herbarium practices and are housed at the
McNeese State University (MCN) Herbarium. Additionally, specimens from
other herbaria have been examined. Nomenclature follows Kartesz (1999).
RESULTS
The following entries are new additions to the vascular flora of Calcasieu Par-
ish. These taxa have not been reported previously as occurring in the parish.
Specimen references follow the same format as in the original survey (Neyland
et al. 2000). Authorities are abbreviated according to Brummitt and Powell
(1992). Following the collection data is an abbreviation for the vegetative region
SIDA 20(1): 431 — 433. 2002
432 BRIT.ORG/SIDA 20(1)
where each species typically occurs: prairie (Pr); longleaf pineland (Pi). Al-
though not a vegetation region, disturbed areas such as pastures, fallow fields,
ditches, urban areas and roadsides are collectively abbreviated (D).
AMARYLLIDACEA
‘phyranthes eer (Lind1.) Herbert (MCN) Neyland 1944 D
ASTERACEAE
Bidens alba (L.) DC. (MCN) Neyland 1940 D
LAMIACEAE
Nepeta cataria L.(MCN) Walker s.n. D
ORCHIDACEAE
Platanthera integra (Nutt.) Gray ex Beck (AMES) Cocks s.n. Pi
The following entries have been previously reported in the parish but were not
collected during the original survey of Neyland et al. (2000). Therefore, these
collections provide recent evidence for their presence in Calcasieu Parish.
ASTERACEAE
Coreopsis pubescens FIL. var. debilis (Sherff) E.B. Smith (MCN) Neyland 1951 D, Pi
olium Michx. (MCN) Neyland 1947 D, Pr
Sans
Silphium integri
EUPHORBIACEAE
Chamaesyce hirta (L.) Millsp. MCN) Neyland 1939 D
JUNCACEAE
Juncus bufonius L. Haning 8(MCN) D, Pi, Pr
LAMIACEAE
Scutellaria drummondii Benth. (MCN) Neyland 1927 D
OXALIDACEAE
Oxalis violacea L. (MCN) Neyland 1945 D, Pi, Pr
The following entries were misidentified in the original survey. Subsequent
annotations have resulted in the noted corrections.
1. The specimen originally reported as Asclepias variegata L. Neyland 1763
Piis Asclepias obovata Ell. Therefore, this species is removed from the survey.
2. The specimen reported as Juncus elliottii Chapm. Brooks 609 (MCN) D,
Pi, Pris Juncus acuminatus Michx. However, the species does occur in the par-
ish as evidenced by the specimen designated as Juncus elliottii Chapm. Giltner
37 (MCN) D, Pi, Pr.
DISCUSSION
The most notable addition to the flora of Calcasieu Parish is the discovery of a
specimen of Platanthera integra in the Oakes Ames Herbarium at Harvard.
This specimen was collected by R.S. Cocks in September, 1906; however, addi-
tional collection information is absent from the herbarium sheet. P. integra is
presently rated by The Natural Heritage Program of the Louisiana Department
of Wildlife and Fisheries as “S2” which is defined as rare and local throughout
NEYLAND, UPDATE TO THE FLORA OF CALCASIEU PARISH, LA 433
the state or found locally in a restricted region of the state. This apparently is
the only known specimen that has ever been collected in Calcasieu Parish. Be-
cause suitable habitat for this species largely has been destroyed in the parish
(Neyland et al. 1998), it may be extirpated.
As a result of these revisions to the original vascular flora survey of
Calcasieu Parish, the number of confirmed specific and subspecific entries for
the parish is increased to 1,153. The number of reported but unverified entries
for the parish is reduced to 131.
ACKNOWLEDGMENTS
I thank Sara Thames, University of Louisiana at Monroe, Monroe, LA, John
Miller, Lake Charles, LA, Bruce A. Sorrie, Longleaf Ecological, Whispering Pines,
NC and Lowell E. Urbatsch, Louisiana State University, Baton Rouge, LA for their
assistance.
REFERENCES
Brumitt, R.K.and C.E. Powett. 1992. Authors of plant names. Royal Botanic Gardens, Kew.
Kartesz,J.7.1999.A synonomized checklist of the vascular flora of the United States, Canada
and Greenland. 1st. ed. In: Kartesz, J.T. and C.A. Meacham, eds. Synthesis of the North
American flora, Version 1.0.North Carolina Botanical Garden, Chapel Hill.
NeyLanp, R., H.A. Meyer, and H. Harrincton. 1998. Woody vegetation of longleaf pine com-
munities in Calcasieu Parish, Louisiana. Phytologia 85:5 1-60.
NeYLAND, R., B.J. HOFFMAN, M. Mayrtetp, and L.E. Ursatscu. 2000.A vascular flora survey of Calcasieu
Parish, Louisiana. Sida 19:361-386
434 BRIT.ORG/SIDA 20(1)
Books RECEIVED/ NOTICES
Annotations and comments by the Editor
Horticulture/Gardening
LoRAN M. WuiTELock. 2002. The Cycads. (ISBN 0-88192-522-5, hbk.). Timber Press,
Inc., 133 S.W. Second Avenue, Suite 450, Portland, OR 97204-3527, U.S.A.
(Orders: wwwtimbet press.com, 800-327-5680, 503-227-2878, 503-227-3070
fax). $59.95, 532 pp., 505 color photos, 13 line drawings, 8 1/2" x 11"
Key Worps: Cyecads, horticulture, cultivation, ag distribution, classification, morphology,
reproduction, conservation, I] genera, ca. 300 moe
This is one encyclopedia of informati -ycads. The 505 color plates are exceptionally nice.
If you are into cycads then you will want this boas
JOHN Bryan. 2002. Bulbs, Revised Edition. (ISBN 0-88192-533-0, hbk.). Timber Press.
Inc., 133 S.W. Second Avenue, Suite 450, Portland, OR 97204-3527, USA.
(Orders: wwwtimberpress.com, 800-327-5680, 503-227-2878, 503-227-3070
lax). $89.95, 896 pp., L171 color photos, 10 line drawings, 2 charts, 81/2" x 11".
Key Worps: “ Bulbs, landscape, horticulture gardening, « catalog, worldwide.
This is an impressive revised edition. The book is worth the price in just the color plates alone.
There are many color plates reproduced from Curtis's Botanical Magazine. Just beautiful!
Jane McGary (editor). 2001. Bulbs of North America. (ISBN 0-88192-517-9, pbk.).
Timber Pressand North America Rock Garden Society (Orders: Timber Press,
133 SW Second Ave., Suite 450, Portland, OR 97204-3527, US.A.,
www.timberpress.com, 800-327-5680, 503-227-2878, 503-227-3070 fax).
$34.95, hbk., 308 pp., 101 color photos, 4 line drawings, 2 maps, 6" x 9"
Key Worbs: Bulbs, landscape, horticulture, gardening, catalog, North America.
Contents include: 1) The Genus Allium; 2) Amaryllidaceae of North America; 3) The Brodiaea
Alliance: Bloomeria, Brodiaea, Dichelostemma, and Triteleia, 4) The Genus Calochortus: 5) The Ge-
nus Erythronium; 6) The Genus Fritillaria; 7) lrids of the Southeast; 8) The Genus Lilium: 9) Bulbs
of the Northwest; 10) Bulbs of the Southwest; and LH) Bulbs of Eastern North America. Information is
provided on sources of bulbs and seeds.
—
SIDA 20(1): 434. 2002
NEW VASCULAR PLANT RECORDS FOR TEXAS
Guy L.Nesom and Robert J.O’Kennon
Botanical Research Institute of Texas
509 Pecan Street
Fort Worth, TX 76102-4060, U.S.A.
Study of previous collections brings to light records for two species (Asteraceae,
Caryophyllaceae) previously unreported for the state of Texas.
Anaphalis margaritacea (L.) Benth.
Anaphalis margaritacea occurs over most of the western and northern United
States and Canada—it has not been recorded from Texas, Oklahoma, and states
of the southeastern USA (Kartesz 1999). In Mexico it is known from localities in
northern Baja California (Wiggins 1980). It is reported from eight primarily
montane counties in New Mexico (Martin & Hutchins 1981), where it occurs at
7000-10,000 feet elevation, and from several prairie counties in Colorado (Great
Plains Flora Association 1976). The record from the Texas panhandle reported
here is a southeastward ‘outlyer from the main range but it isa relatively small
step eastward from the known localities in New Mexico and Colorado. The
Brewster Co. collection is disjunct southward about 350 kilometers from the
closest localities in Lincoln Co., New Mexico. Other UNM collections (fide Jane
Mygatt, UNM Collection Manager) confirm that Ora Clark was botanizing in
Brewster County in the first week of August 1931, adding a degree of confidence
that the Anaphalis collection did originate from there.
Voucher specimen: TEXAS. Brewster Co.: “arroyas, s. Terlingua, Texas,” 5 Aug 1931, Clark 4199 (UNM).
Hale Co.: Plainview, E of Co-op Cotton Compress on side of railroad, sandy soil, 3350 ft, 11 Sep 1967,
Hall s.n. WIS).
Gypsophila elegans M. Bieb.
Gypsophila elegans (var. elegans) is native to Russia (Black Sea region), Crimea,
Caucasus, eastern Turkey, and lran (Caspian Sea region) and is adventive in
western Europe and North America (Barkoudah 1962), where it is grown as a
garden plant. Gypsophila elegans var. latipetala Barkoudah is known only from
“Turkey and Transcaucasia” (fide Barkoudah 1962). Variety elegans is known
from scattered localities in Canada and the USA (including California, Utah,
Colorado, Kansas, Illinois, North Carolina (Kartesz 1999). The central and north-
central Texas localities reported here are the southernmost known stations in
North America.
A key in the treatment of Gypsophila for California (Hartman 1993) sepa-
rates G. elegans from G. paniculata L. and G. scorzonerifolia Ser. Gleason and
SIDA 20(1): 435 — 436. 2002
436 BRIT.ORG/SIDA 20(1)
Cronquist (1991, p. 127) included a brief description and comment regarding G.
elegans, following the description of G. muralis L., noting that it “occasionally
escapes from cult. in our range.” It is possible that the Texas plants are from
direct roadside seeding—the species is distributed as part of ‘Texas/Oklahoma,
‘Mountain,’ ‘Northeast, Gulf Coast/Caribbean, and ‘Southeast’ wildflower seed
mixes (see www.prairiefrontier.com/pages/pfmixpg/regional.html). Another
website touts the species as “One of the easiest seeds to grow. This plant literally
pops out of the ground in any soil, ready to bloom.” A search for the Tarrant
County plants at the Keller site in 2002 was unsuccessful, however, suggesting
that the species is not successfully reproductive in north central Texas.
The common name of Gypsophild elegans is Annual Baby’s Breath or Showy
Baby’s Breath. Many photos of the species are available on the internet, includ-
ing an excellent one of a herbarium specimen (linnaeus.nrm.se/flora/di/
caryophylla/gypso/gypsele.html).
Voucher specimen: TEXAS. Erath Co.: roadside of Hwy 377,ca.7 mi Sof Stephenville at intersection
of Co. Rds. 379 and 380, in ditch on NW side of hig
and bluebonnets, 15 Apr 2001, Kunselman 23 (TAC). Tarrant Co.: along railroad tracks in township
of Keller, NE of Fort Worth, Apr 2001, O’Kennon (sight record only).
hway, with ‘oats, wheat, bermudagrass, rescuegrass
ACKNOWLEDGMENTS
The new records reported here are from among specimens in loans to BRIT from
WIS, TAC, and UNM. The Tarrant County collection of Gypsophila was inde-
pendently identified by Richard Rabeler. Review comments of Lindsay Woo-
druff are greatly appreciated. Jane Mygatt provided helpful information on Ora
Clark and his collecting schedule.
REFERENCES
Barkoupad, Yl. 1962.A revision of Gypsophila, Bolanthus, Ankyropetalum and Phryna.Wentia
9:1-203.
Gieason, H.A. and A. Cronauist. 1991. Manual of vascular plants of northeastern United
States and adjacent Canada (ed. 2). New York Botanical Garden, Bronx.
GREAT PLAINS FLorA Association (R.L. McGregor, coord,; T.M. Barkley, ed.).1976.Atlas of the flora
of the Great Plains. lowa State University Press, Ames.
Hartman, R.L. 1993. Caryophyllaceae (except Silene). In: J. Hickman, ed. The Jepson manual:
higher plants of California. Univ. California Press, Berkeley. Pp. 475-497,
Kartesz, J.T. 1999. A synonymized checklist and atlas with biological attributes for the
vascular flora of the United States, Canada, and Greenland (Ed. 1). In: J.T. Kartesz and
CA. Meacham. Synthesis of the North American flora, Version 1.0. North Carolina Bo-
tanical Garden, Chapel Hill.
Martin, W.C. and C.R. HutcuHins. 1981. A flora of New Mexico. J.Cramer, Vaduz.
Wicains, LL. 1980. Flora of Baja California. Stanford Univ. Press, Stanford, California.
NOTES ON THE FLORA OF TEXAS WITH ADDITIONS
AND OTHER SIGNIFICANT RECORDS. II
Larry E. Brown |. Sandra Elsik
Houston Community College Dept. of Ecology and Evolutionary Biology, MS 170
1300 Holman, Houston, 1X 77007, U.S.A. P.O. Box 1892
and Spring Branch Science eal ter Herbarium Rice University
8856 Westview Dr Houston, TX 77251, U.S.A.
Houston, 1X 77055, USA elsik@rice.edu
larry@theplantman.net
ABSTRACT
The following ten taxa are documented new to Texas: Aster laevis var. purpuratus, Bidens alba var.
radiata, Tridax procumbens, Raphanus raphanistrum, Calycanthus floridus var. glaucus, Lyonia lu-
M Significant new
cida, Hyptis mutabilis, Lilium ee orum, pila ae and
collection records are provided for eight: Hyp 1, Cyperus -ephalanthus, Rhynchosia
tomentosa, Quercus arkansana, Allium stellatum, fore viscosissima, Denuthens elata subsp.
hirsutissima, and Leptochloa dubia. Taxa are listed alphabetically by family, genus, and species with
comments and citation of voucher specimens.
RESUMEN
Los siguientes diez taxa se citan nuevos para Texas: Aster laevis var. purpuratus, Bidens alba var.
radiata, Tridax procumbens, Raphanus raphanistrum, Calycanthus floridus var. glaucus, Lyonia lu-
ay is mutabilis, Lilium longiflorum, Lythrum lineare, y Myrtus communis. Se hacen citas
si ae s de otros ocho nuevas recolecciones: Hypochaeris glabra, Cyperus cephalanthus,
a aa tomentosa, Querct Allium stellatum fp Cunhea viscosissima, Oenothera elata
bh] Jarhar l
subsp. Hesse yl familia, género y especie
r
is
con a y cita de los especimenes testigo.
INTRODUCTION
Floristic work in the Houston area and other east Texas counties, coupled with
plants sent to the SBSC herbarium for identification, has revealed a number of
plants new to Texas and significant new records for others.
Unless otherwise indicated, all collections cited in this paper are in the
Spring Branch Science Center Herbarium (SBSC), Houston, Texas. Duplicates
of some collections are at the other indicated herbaria. Herbarium acronyms
follow Holmgren et al. (1990).
ASTERACEAE
Aster laevis L. var. purpuratus (Nees) A.G. Jones.—Aster laevis has been repre-
sented in Texas by the western variety ge yeri in the Guadalupe Mts of Culberson
Co. (Nesom 1993). Here we report variety purpuratus, the narrow leaved
southeastern entity, from two east Texas counties. Thomas and Allen (1996)
SIDA 20(1): 437 — 444. 2002
438 BRIT.ORG/SIDA 20(1
mapped it in six, mostly north central, Louisiana parishes. Kartesz (1999)
mapped it, as Symphyotrichum laeve (L.) A. & D. Love var purpuratum (Nees)
Nesom, in Georgia, Alabama, Mississippi, Louisiana, Arkansas, and Texas. At
the Polk County site, A. laevis var. purpuratus is distinguished from the similar
and more numerous plants of A. oolentangiensis Riddel by smooth leaves in
contrast to the harshly scabrous leaves of A. oole meee nsis.
Voucher specimens: Polk Co.: growing ina native cal ie surrounded by pine-oak woods
at the intersection of Lone wel Road and Wancinarn Road, N of Hwy 1988 between Goodrich and
Providence, 5 of Livingston, 30° 37'33"N and 94° 55'33"W, 28 Oct 2000, Brown 24913; same site, 23 Sep
1999, Brown 23563. Tyler Co.: acne inacalcareous prairie barren on U.S. Hwy 190, ca. 4.5 mi W of
intersection with FM 92, 12 Jun 2000, J. Liggio s.n.
Bidens alba (L.) A. DC. var. radiata (Sch. Bip.) Ballard ex T.E. Melchert—Ballard
(1986) divided the B. pilosa complex into three mostly tropical American spe-
cies, namely B. alba, B odorata Cav, and B. pilosa L. He mapped United States
records of B. alba var. radiata in Florida and B. pilosa in California. Wunderlin
(1998) reported B. alba var. radiata as common and distributed nearly through-
out Florida. Gandhi and Thomas (1989) reported B. pilosa. in Louisiana but in-
dicated that Ballard’s description of Bidens alba var. radiata was in concurrence
with their description of B. pilosa. Thomas and Allen (1996) maintained B. pilosa
in Louisiana and mapped it in thirteen parishes. We report B. alba var. radiata
(sensu Ballard) from four Texas counties. There is a photograph of the taxon in
Tveten and Tveten (1993).
Voucher specimens: Cameron Co.: common ina vacant lot in Brownsville, | Apr 1983, Brown 5954.
Chambers Co.: along [-L0 at the bridge over the Trinity River near the village of Wallisville, 29° 50
12"N and 94° 3908"W, 8 Oct 2000, Brown 24801 Harris Co.: weed along Old Katy Rd at Bunker Hillin
Houston, 19 Oct 1990, Tveten b-390; weedy plants at the intersection of Dell Dale and Brentwood Streets
in Channelview, 21 May 1992, Brown 16084. Montgomery Co.: edge of small woodland next to parking
lot near Woodlands Parkway, |/4 mi W of 145, 8 Dec 2001, FE. Keith 31.
Hypochaeris glabra |..—Diggs et al. (1997) reported this European weed new to
Texas from Williamson County. Since then additional collections have been
made in other regions of the state.
Voucher specimens: Harris Co.: weedy plants in a subdivision adjacent to a small native prairie at
intersection of Greenhouse Rd and FM 529, 29° 52'42"N and 95° 42'14"W, 12 Mar 1999, Brown 22866:
plants on ane soil in pasture on south side of the Harris County Katy Park, N of Katy, 29° 4834"N
and 95° 48'20"W, 14 Apr 2001, Brown 25083. Leon Co.: growing on sandy soil in the Boykin Cemetery
olf of Hwy - between Oakwood and Flo, 18 Apr 2001, Brown 25090. Waller Co.: on sandy soil at the
intersection of Holik Rd and Hwy 1887, west of the Monaville cemetery and between Hempstead
and Pattison, 29° 56'58"N and 960° 04'51"W, 8 May 2002, Brown 26702.
Tridax procumbens L.—Kartesz (1999)reported this exotic and federal noxious
weed in Florida and Hawaii. Wunderlin (1998) reported it frequent in the cen-
tral and southern peninsula of Florida. John Tveten recently sent a collection
of it from the Valley Nature Center in Hidalgo County for identification. Re-
garding the history of the plant in Texas, the following communication was
BROWN AND ELSIK, NOTES ON THE FLORA OF TEXAS 439
received from Ken King and Martin Hagne of the Valley Nature Center; “Seed
was collected in the Brownsville area by a native grower several years ago. That
grower is no longer in the area and we do not know the exact Brownsville loca-
tion. The plants at the Valley Nature Center were planted here from those seed-
lings. This comp is growing in empty lots throughout Weslaco and has been in
the area for at least fifteen years. We have seen several local establishments
around town. There are at least two south Texas counties that have communi-
ties of this plant; Hidalgo and Cameron.” Alfred Richardson (1995) did not in-
clude this taxon for the Rio Grande Delta area.
Voucl Hidalgo Co long trail tsi ite bush in the Weslaco
ey ner Center, 20 Now 2001 Gate S.A,
(=e
Val
BRASSICACEAE
Raphanus raphanistrum |.—Rollins (1993) distinguished the genus Raphanus
by a segmented fruit with the lower segment inconspicuous and seedless. The
petal colors vary from white, yellow, lilac, to violet. Rollins Cloc cit.) reported
two introduced weedy species in North America, R. raphanistrum and R. sa-
tiva. He indicated that in R. raphanistrum the fruits are linear, longitudinally
grooved, and strongly constricted between the seeds. The petals are yellow and
fade to whitish or white. In R. sativus the fruit body is widest at the base, smooth
or very slightly grooved, and is not or only poorly constricted between the seeds.
The petals are usually purple. The petal veins are prominent in yellow flow-
ered R. raphanistrum, which can distinguish it from yellow petaled plants of
Brassica and Sinapsis, if one does not notice the small lower fruit segment.
Kartesz (1999) mapped it in Texas fide Gould (1975a) but no voucher was pro-
vided. It is not listed in Correll and Johnston (1970) nor any of the recent check-
lists. We report voucher material with the traits of R. raphanistrum from three
Texas counties. Thomas & Allen (1996) mapped R. raphanistrum in numerous
northern and southern Louisiana parishes. Two of these parishes are adjacent
to Texas. In contrast, R. sativus is known from only seven parishes. Wunderlin
(1998) found R. raphanistrum more common in Florida than R. sativa. A reex-
amination of Texas material may reveal the same situation in Texas.
usa aie imens: Fort Bend Co.: yellow flowered weed 2 plant alonga ie road above a drainage
093
W edge of Barker Reservoir, ca. 1/2 mi E of Mason Rd at .1/2 miN of FM
10 Mar 1995, Brown 18435. Harris Co.: Brays Bayou at BE falo Speedway in Houston, 18 Mar 2000, D.
Johnson s.n. San Jacinto Co.: collected along roadside of Hwy 945 near the Evergreen Cemetery in the
village of Evergreen, 9 Mar 2001, E. Keith s.n.
CALYCANTHACEAE
Calycanthus floridus L. var. glaucus (Willd.) Torr. & A. Gray.—At least fifteen
plants of this shrub are present in woods adjacent to the David and Patricia
Lewis residence in Newton County. Ina personal communication, the Lewises
indicated that these plants were here when they established a residence on the
440 BRIT.ORG/SIDA 20(1)
site. Thomas and Allen (1996) were unsure of the native status of a collection
from West Feliciana Parish, Louisiana. Likewise we are uncertain of their na-
tive status here; however, they are well established and naturalized shrubs and
should be considered a member of the Texas flora.
Voucher specimens: Newton Co.: ic slope 1] | Screw Pin Branch adjacent
to CR 3062, W of Hwy 87 N of Bimal 22 Jul 2000, Brown & Elsik 24437 (SBSC, BRIT): same site,
13 Apr 1999, Lewis, Lewis & Elsik 6169 (SBSC, TEX).
—
CYPERACEAE
Cyperus cephalanthus Torr. &@ Hook.—Carter and McInnis (1993) reported the
type locality of this sedge from somewhere near Galveston Bay. The type col-
lection was made in 1835 and the species has not been recollected in Texas since.
We here report a recent collection from Harris County. Richard Carter (VSC)
has verified the identification.
Voucher specimen: Harris Co.: tall plants with thick stem in a moist prairie depression a few hun-
dred feet SE of the Texas ee College, on the S side of Spencer Blvd, a few hundred feet EF of
Beltway 8 in Pasadena, 29°39'39"N and 95'08'50"W, 29 May 1999, Brown 23076 (BRCH, SBSC, TEX).
This site is owned by the Harris County Flood Control District and is soon to be
formed into a detention basin for downstream flood control along Armand
Bayou. However, Ralph Taylor of the Flood Control Environmental Services has
removed all plants from the site and replanted them in their nursery near Lake
Houston. Plants have been given to the Mercer Arboretum and Botanic Gar-
dens and to the Armand Bayou Nature Center for establishment on their lands.
ERICACEAE
Lyonia lucida (Lam.) K. Koch.—Specimens of these shrubs, with flowers in March
and fruits in summer, have been recently collected in Newton County. The
shrubs were on the property of David and Patricia Lewis when they set up a
residence north of Bleakwood. There was no evidence of a prior human occu-
pation of the site. David has discovered six populations with well over a hun-
dred total plants (per.comm.) along Screw Pin Branch. Fetterbush is native and
common in the adjacent Sabine Parish of Louisiana (personal observation by
the second author, Thomas and Allen 1996). It is our opinion that this is also a
native population.
Voucher s ae Newton Co.: shrubs along Screw Pin Branch, ca. 300 m W of Hwy 87,1 mi N of
the intersection with Hwy 363i iy of peer 30° ace N ane 93'4934"W, 24 Mar 1999,
Lewis & Elsi a (SBSC, nine dupli ite, 22 Jul 2000
Brown & Elsik 24436. (SBSC, BRIT).
FABACEAE
Rhynchosia tomentosa (1.) Hook. @ Arn.—Turner (1959), Correll and Johnston
(1970), Johnston (1990), Hatch et al. 1990), Jones et al. (1997), and Isely 1998)
all reported this legume for Texas. Turner (1959) reported a collection by Charles
BROWN AND ELSIK, NOTES ON THE FLORA OF TEXAS 441
Wright that lacks locality data. Isely (1998) did not map it in the state. It is also
not mapped in the state by Turner et al. (in press). We here report an authentic
Texas collection from what is now the Little Rocky Preserve of The Nature Con-
servancy of Texas.
Voucher specimen: Jasper Co.: Little Rocky Nature Conservation area, 13 mi N of Jasper on Hwy 96
to Browndel, then ca. 3 mi E of Browndel on Hwy 1007, 15 Aug 1982, Peterson 631
FAGACEAE
Quercus arkansana Sarg —Hunt et al. (1995) first reported this oak as new to
Texas. They found two small populations in Cass County and an historical
record for Jasper County. We here report recent collections from Jasper and
Hardin counties. Jensen (1997) mapped it in the extreme northeast and south-
east portions of the state.
Voucher specimens: Jasper Co.: deep sandy hillside about 2 mi N of the end of FM 2800, NW of
Jasper in the headwaters of Hog Creek, 3 Jul 2001, E. Keith Ib.
ay
LAMIACEAE
Hyptis mutabilis (Rich.) Brig—Kartesz (1999) reported this exotic weed in seven
southeastern states from Virginia to Louisiana. Thomas and Allen (1998)
mapped it in three parishes east of the Mississippi. River. We here report a col-
lection for Texas.
Voucher specimen: Liberty Co.: plants growing in a fenced field on S side of the Davis Hill Salt Dome
adjacent to the Davis Hill Estates subdivision, east of Cleveland and off of Hwy 105, 30° 1832"N and
94° SL'OL"W, 5 Jul 1999, Brown 23169.
LILIACEAE
Allium stellatum Ker—Turner et al. (in press) mapped this onion in five, mostly
north central Texas counties. We here report collections far south of those
mapped.
Voucher specimens: Polk Co.: in a native prairie surrounded by pine-oak forest along Lone Wolf Rd
N of Hwy 1988 between Goodrich and Hwy 146 S of Livingston, 17 Nov 1996, Brown 19809. Brazos
Co.: population of red flowered plants on a gravel slope above Lick Creek, NE of the intersection of
Hwy 6 and Greens Prairie Rd, probably within the city limits of College Station, 2 Nov 2001, Brown
26401 (SBSC, TAMU).
Lilium longiflorum Thunb.—In the summer of 2000, we found this tall white
flowered plant to be common and conspicuous along roadsides near Newton. It
is also found as an escape from cultivation in Alabama, Florida, and Utah
(Kartesz 1999). We identified it to this species with the key in Wunderlin (1998).
Thomas and Allen (1993) reported L. formosum in Louisiana, but our plants have
the totally white flowers of L. longiflorum, in contrast to the red-purple outer
perianth of L. formosum.
Voucher specimens: Newton Co.: growing along the gravel CR 3062 between Screw Pin Branch and
442 BRIT.ORG/SIDA 20(1)
Thickety Creek, west of Hwy 87, N of Bleak wood, 22 ae 2000, Brown & Elsik 24438. Tyler Co.: 1.1 mi
N of jet Hwy 190/FM 256 W of Woodville, 3 Aug 1991, Schultz 061.
LYTHRACEAE
Cuphea viscosissima Jacq.—Correll and Johnston (1970) reported Blue Wax weed
in Texas but withouta specific locality.Graham (1975) eliminated it from Texas,
as did Johnston (1990), Hatch et al. 1990), Jones et al. 1997), Kartesz (1999), and
Turner et al. (in press). We here report a voucher to confirm its presence in Texas.
oucher specimen: San Augustine Co.: 8 mi W of San | Augustine on Hwy 21, then 1 mi S, Weches
iar 16 Jul 1976, George & Nixon 43 (SBSC, AS”
Lythrum lineare |.—Correll aria (1970), Graham (1975), Johnston (1990),
Hatch et al. (1990), Jones et al. (1997), and Turner et al. Gin press) did not report
this species in Texas. However, Kartesz (1999) listed Texas within its distribu-
lers to support its presence in the state. The identifi-
tion. We provide two vouc
cation was verified by S. Graham (KE).
Voucher 2 penis Chambers Co.: Anahuac National Wildlife Refuge, brackish marsh in the Jackson
Ditch Unit, 29°35'24'N and 942509" W, 3 Oct 2000, Brown 24774, Anahuac National Wildlife Refuge,
brackish m¢ a in the East Unit, 29°39'N and 94°28'W, 20 Jul 2000, Brown 24399 (SBSC, KE)
MYRTACEAE
Myrtus communis L.—Kartesz (1999) did not report this cultivated ornamental
plant as occurring wild in the United States. The first author found many well
developed shrubs scattered across a salt dome in coastal Texas. These plants
had mature fruits and were rather large, suggesting they have been here for some
time. The salt dome is isolated and surrounded by a coastal prairie and is for the
most part undeveloped except for some old machinery and buildings associ-
ated with prior oil extraction.
mn)
Voucher specimen: Brazoria Co.: shrubs with fruits along gravel road to the center of Hoskins Mound,
Galveston- ae oria Key Map 801 D and H, near 29°08'26"N and 95'13'26'W, 6 Aug 2000, Brown 24588
(SBSC, TEX, AS
ONAGRACEAE
Oenothera elata Kunth subsp. hirsutissima (A. Gray ex S. Wats.) W. Dietr.—
Dietrich et al.(1997) mapped this taxon largely in Trans-Pecos Texas with one
collection from the Panhandle. They also mapped three populations in east
Texas in Anderson, Brazos, and Leon counties. Plants matching the key traits
and description of this taxon have been collected in Harris and Liberty coun-
ties. They are more common on sandy soil in Liberty County. These are tall
plants with large yellow flowers whose stigmas are elevated above the stamens.
Voucher penne Harris Co.: tall herb along Hwy 59 at the San Jacinto River bridge, 28 Jul 1991,
Gregg s.n. Liberty Co.: plants oa ing on sands along the Trinity River at the Hwy 105 bridge west ol
Moss Hill, 30° ; 30" N and 94°47'57" W,1 Oct 2000, Brown 24758; along the Ace Hwy Sof the village of
Ace, across from the eee ae of the Menard Creek Unit of the Big Thicket Biological Pre-
BROWN AND ELSIK, NOTES ON THE FLORA OF TEXAS 443
serve, 30°28'59'N and 94°50'19" W, 25 Aug 2000, Brown 24680a; tall plants with rather | f]
deep sands along the Trinity River, Trinity River Nat Wildlife Refuge at cami oa on the Davis
Hill 7.Squad, 30 Aug 1996, Brown 19528; large flowered primrose on dry sandy soil along CR 2252
near the Davis Hill Baptist Church E of Cleveland and N of Hwy 105, 15 Sep 1996, Brown 19693; a
single tall plant at edge of Hwy 146, ca 3 mi N of Moss Hill, 21 Aug 1994, Brown 18124; tall weedy
plants with large yellow flowers, along dirt road in the Dayton Lake Estates E of Hwy 1008, adjacent
to the Trinity River, N of Dayton, 9 Sep 1994, Brown 18180.
POACEAE
Leptochloa dubia (H.B.K.) Nees. —Gould (1975b) reported this grass from all re-
gions of the state except for the east Texas pineywoods and the post oak savan-
nah area. We here report collections from two east Texas counties.
\ ! Harris Co.: plant lark cl il along a newly constructed road into a prairie
at the ieeectio of Onkesaed St. and Fairmont ee in Deer Park, 29 Sep 1984, Brown 8100.
Hardin Co.: fairly frequent tall grasses on verge of hwy between village of Honey Island and Hwy
69/287, N of Kountze, 6 Sep 1996, Brown 19565.
ACKNOWLEDGMENTS
We wish to thank Tom Wendt (TEX) and Leslie Landrum (ASU) for the identi-
fication of Myrtus communis, Richard Carter (VSC) for the verification of
Cyperus cephalanthus, and S. Graham (KE) for the verification of Lythrum
lineare. Michael MacRoberts provided a literature source and Guy Nesom (BRIT)
helped with some nomenclatural questions. Justin Williams (SHST) and Mo-
nique Reed (TAMU) reviewed the paper and provided many helpful suggestions.
The first author is thankful to the Houston Community College for a sabbati-
cal leave during the spring semester of 2002 that allowed him to complete this
project. Publication funds were provided by Rice University.
REFERENCES
BaLLaro, R. 1986. Bidens pilosa complex (Asteraceae) in North and Central America. Amer. J.
Bot. 73:1452-1465.
Carter, R.and N. Mclinnis. 1993. Final status report for Cyperus cephalanthus T. & G. Unpub-
lished report. U.S. Fish and Wildlife, endangered species office. Jackson, MS.
Corrett, D.S. and M.C. Jounston. 1970. Manual of the vascular plants of Texas. Texas Re-
search Foundation, Renner, TX.
Dietrich, W., W.L. Waaner, and PH. Raven. 1997. Systematics of Oenothera section Oenothera
subsection Oenothera (Onagraceae). Syst. Bot. Monogr. 50:1-234.
Diccs, G.M., R.J. O’Kennon, and B.L. Lirscome. 1997. Hypochaeris glabra, a new Asteraceae for
Texas. Sida 17:633-634.
GANoHI, K.N. and R.D. THomas. 1989. Asteraceae of Louisiana. Sida Bot. Misc. 4:1-202.
Goutp, EW. 1975a. Texas plants —a checklist and ecological summary. Revised edition.
Texas Agric. Exp. Sta. College Station.
Goutp, FW. 1975b. The grasses of Texas. Texas A&M University Press. College Station.
444 BRIT.ORG/SIDA 20(1)
GRAHAM, S. 1975. Taxonomy of the Lythraceae in the southeastern United States. Sida 6:
80-103.
Hatch, S.L, KN. GANDHI, and L.E. Brown. 1990. Checklist of the vascular plants of Texas, MP-
1655. Texas Agric. Exp. Sta., College Station.
Houmeren, P.K., N.H. Houmaren, and L.C. Barnett, 1990. Index herbariorum, 8"? ed. New York
Botanical Gardens. Bronx.
Hunt, D.M., M.H. MacRoserts, and B.R. MacRoserts. 1995. The status of Quercus arkansana
Sarg. (Fagaceae) in Texas. Phytologia 79:22-24.
Isety, D. 1998. Native and naturalized Leguminosae (Fabaceae) of the United States (exclu-
sive of Alaska and Hawaii). Monte L. Bean Life Sciences Museum, Brigham Young Uni-
versity. Provo, Utah.
Jensen, R.J. 1997. Quercus section Lobatae.|n Flora of North American, vol. 3.Oxford Univer-
sity Press. New York, NY.
Jounston, M.C.1990.The vascular plants of Texas, a list up-dating the manual of the vascu-
lar plants of Texas. 2" ed. Privately published by the author. Austin, TX.
Jones, S.D., J.K. Wire, and P.M. Montcomery. 1997. Vascular plants of Texas. A comprehensive
checklist including synonymy, bibliography, and index. University of Texas Press. Austin.
Kartesz, J.1.1999.A synonymized checklist and atlas with biological attributes for the vas-
cular flora of the United States, Canada, and Greenland. First edition. In J.T. Kartesz and
C.A.Meacham. Synthesis of the North American Flora, Version 1.0. North Carolina Bo-
tanical Garden, Chapel Hill.
Nesom, G.L. 1993. Three species of Aster (Asteraceae:Astereae) disjunct in northern Coahuila,
Mexico. Phytologia 74:296-304.
RicHarpson, A. 1995. Plants of the Rio Grande Delta. University of Texas Press. Austin.
Roiuins, R.C. 1993. The Cruciferae of continental North America. Stanford Univ. Press. Stan-
ford, CA.
THomas, R.D.and C.M. Aiten. 1993. Atlas of the vascular flora of Louisiana. Vol. |: ferns & fern
allies, conifers, & monocotyledons. Louisiana Department of Wildlife & Fisheries. Baton
Rouge.
THomas, R.D, and CM. Aten. 1996. Atlas of the vascular flora of Louisiana. Vol. Il: dicotyle-
dons, Acanthaceae — Euphorbiaceae. Louisiana Department of Wildlife & Fisheries. Baton
Rouge.
THomas, R.D.and CM. Atten. 1998. Atlas of the vascular flora of Louisiana. Vol. Ill: dicotyle-
dons, Fabaceae-Zygophyllaceae. Louisiana Department of Wildlife & Fisheries. Baton
Rouge.
Turner, B.L. 1959. The legumes of Texas. University of Texas Press. Austin.
Turner, B.L., H. NicHots, O. Doron, and G.C. Denny.In press. Atlas of the Texas flora Vol. 1 Dicots
and Vol. 2 Monocots. Sida Bot. Misc.
Tveten, J.L.and G.A. Tveten. 1993.Wildflowers of Houston and southeast Texas. University of
Texas Press. Austin.
Wunoertin, R.P. 1998. Guide to the vascular plants of Florida. University Press of Florida.
Tampa.
ERRATUM FOR SIDA 18(4):1227—1245. 1999.
Allan Nelson
Department of Biological Sciences, Box T-0100
Tarleton State University
— TX 76402, U.S.A.
elson@tarleton.edu
Negrete et al. wish to correct their paper, A checklist for the vascular plants of
Padre Island National Seashore published in the December 1999 issue of SIDA
(volume 18, number 4, pp. 1227-1245) by deleting Sesuvium trianthemoides from
the checklist, which will change the total number of species to 455.
SIDA 20(1): 445. 2002
446 BRIT.ORG/SIDA 20(1)
THE RUPERT BARNEBY AWARD
The New York Botanical Garden is pleased to announce that Aaron Liston, cur-
rently at the Department of Botany & Plant Pathology, Oregon State Univer-
sity, is the recipient of the Rupert Barneby Award for the year 2002. Dr. Liston
will be studying the phylogenetic systematics of Astragalus and Trifolium.
The New York Botanical Garden now invites applications for the Rupert Barneby
Award for the year 2003. The award of US $1,000.00 is to assist researchers to
visit The New York Botanical Garden to study the rich collection of
Leguminosae. Anyone interested in applying for the award should submit their
curriculum vitae, a detailed letter describing the project for which the award is
sought, and the names of 2-3 referees. Travel to the NYBG should be planned for
sometime in the year 2003. The application should be addressed to Dr. James L.
Luteyn, Institute of Systematic Botany, The New York Botanical Garden, 200th
Street and Kazimiroff Blvd., Bronx, NY 10458-5126 USA, and received no later
than December 1, 2002. Announcement of the recipient will be made by
December 15.
Anyone interested in making a contribution to THE RUPERT BARNEBY FUND
IN LEGUME SYSTEMATICS, which supports this award, may send their check,
payable to The New York Botanical Garden, to Dr. Luteyn.
SIDA 20(1): 446. 2002
BOOK REVIEWS 447
BOOK REVIEWS
Martco LAMBerTINI (Translated by John Venerella). 2000. A Naturalist’s Guide to
the Tropics. (ISBN 0-226-46828-3, pbk.). The University of Chicago Press,
58015 Ellis Ave., Chicago, IL 60637-1496, U.S.A. (Orders: 773-702-0279, 773-
702-7956 fax, da@press.uchicago.edu, wwwopress.uchicago.edu). $25.00, 338
pp, 57 color photos, 11 color plates, numerous b/w halftones, 51/2" x 71/2"
Though unknown to most American readers, the author apparently is professionally trained and
works as a conservationist with BirdLife International (UK) and as a freelance scientific journalist.
His locus is to educate the public about the importance of conserving the tropics. As he says in the
preface, “This is a work that is intended to establish among prospective world travelers a knowledge
of the natural life and ecology of the Tropics, to make them seem less distant and unreachable, less
unknown and mysterious.” In this, | would say he has achieved his goal.
=)
On the other hand, the publisher's flier highlights this book as “the first field guide in Englis
to cover all the world’s tropics in one volume” and the author states, “a guide to skim before the tripas
well as to take along.” I would say, “a reference to absorb thoroughly before the trip only.” If you are
looking for an all-in-one field guide (ie., birds, mam mele Eepiiles peacnell> trees, and lowers), you
LK } }
va
will be sorely disappointed. If, instead, you | land-
5
=
le that you will be seeing or want to pee stand why the tropics are so im pera nt, then this
s book is worth your sua! and ti
The first tl sh S a major physical factors—geologic history, climate and soils—
while the next two cover ener features of tropical plants and animals. There are five chapters d
tailing the structure and species of the major ecosystem types in the tropics, ie. rane ore ee
coral reefs, savannas, and deserts. The book concludes with an overview of eee conservation
and a series of precautions that travelers should observe to avoid elephant attacks and yellow feve
among others. The layout is very readable and includes many sidebars to extract more difficult or
specific information from the general flow of the text. The color photographs are beautiful, and the
drawings generally augment the writing.
The introductory chapters are well written and provide a good foundation for understanding
tropical environments. The soils chapter should have been longer to relate the greater diversity ol
tropical soils and the factors giving rise to them. Also, | was surprised that the term, mycorrhizae,
was not introduced, although the author did discuss “elevated recycling of nutrients”. These chap-
ters cover essential topics, such as, seasonality rel lee ted in the structure and behavior of organisms,
primary production, uh eatin similar different continents, species interactions,
and human use. Standard evolutionary explanations are given for the phenomena of symbiosis, de-
fense, population iene biodiversity niches, dis es rsal, and convergence. The colored plates of
birds given in this section are di 7 species each from Africa and Australasia, 8 from the
Americas. The sidebar on re was itaceraved by Seve of only seven species. Having studied
this group, I found at least EnIEe errors. eo) Uae drawing of Phytelephas macrocarpa, was incorrect. The
: . ]
different subfamily, probably a poorly drawn Sabal. 2) Licuala
c Lt ‘a
grandis is Asian not South American. 3) Coconut is pantropical, not just Indo-Pacific, although it
may have originated there
Th t I very informative, especially the one on mangroves, in terms of fea-
ts. For example, the one on forests distinguishes
r
primary and secondary forests and descril py structure, forest characteristics (e.g., root but-
tresses, lianas, and epiphytes), light gaps and pioneer species, life histories o forest plants, fig-wasp
b
symbiosis, growth of stranglers, ecology of forest animals (sidebars on cicadas, butterflies, ants, poi-
SIDA 20(1): 447. 2002
—=
448 BRIT.ORG/SIDA 20(1)
son dart frogs, treehoppers), adaptations of arboreal animals, and pollination and dispersal in the
forest
The last two chapters—conservation and traveling precautions—make this book especially use-
ful and timely.
My recommendation is: Buy this book and read it in depth, book a oe ecotour, then buy
some good field guides for that specific region actually to take with yo And, as the author says,
“Buon viaggio!” —Roger W.S eee: Associate Collections Manager, Bot il Research Institute of Texas,
Fort Worth, TX 76102-4060, U.S.A.
0
Davin Birk. 2000. Flora of Mount Rainier National Park. (ISBN 0-87071-470-8,
dk.) Oregon State University Press. 101 Waldo Hall, Corvallis, OR 97331-
6407. (Orders: http://osu.orstedu/dept/press/osupress.htm, 800-426-3797,
541-737-3166, 541-737-3170 fax). $29.95, 520 pp, 64 color photos, line drawings,
Cx
This accu marks the first comprehensive treatment ol the flora of Mount Rainier subsequent
to George Neville Jones’ work published in 1938. Mount Rainier was designated a National Park in1899
and since hes auc information has been gathered on the biota of the area. The Flora of Mount
Rainier National Park by David Biek seeks to update Jones’ earlier flora and covers the entire area
within the boundaries of the park. According to the author, the goal of this book is “to provide a
escriptions suffi-
Qo.
complete list of the native and introduced plants found in the Park, with keys and ¢
cient to distinguish each species from the others.”
“he volume begins with an introduction on how to use the book and clarifies its scope and
o
intent. The author provides information on how to use the keys to identify plants and forewarns his
readers of the inconstant nature of scientific names of plants. Next, facts on the Park, including loca-
tion, topography and climate are given. The author follows with a discussion of plant communities
and associations, citing 19 different types present in the Park and the groups of plants that can be
found growing in each zone or community. A brief discussion of plant geography, distribution, weeds
and rare plants allow the reader to become familiar with the flora of the Park. Last, a historical over-
view of explorations and studies, including ecological research on Mount Rainier, gives evidence ol
the long significance this area has had in our ee heritage.
course, a ae of the book is dedicated to plants included in the flora and the treatment
begins with a key to the major groups of vascular slant Alter its key, ane major groups are pre-
sented, lerna nd fe ernallies first, followed by gymnosperms, tl finally monocots. Within
each group, the families are listed alphabetically, followed by genera and species. There is a key to
accompany each family and genus description so the reader can ascertain the taxon at hand. Species
escription of diagnostic charac-
—
descriptions include scientific name and common name, a brief ¢
ters, elevation, habit and habitat, as well as locations where it has been collected in the Park. Hlustra-
tions, reprinted from Hitchcock’s Vascular Plants of the ae Northwest, are adjacent to most spe-
cies descriptions. Color photos olf selected species are found in the middle of the book. An appendix
includes 79 plants added to the flora, which were observed by the author in the field or from her-
barium specimens between 1996 and 1999. A brief glossary clarifies some essential terms and the
bibliography cites eutcall publications. The author provides a single index with accepted names,
synonyms, names misapplied and common names used in the book
Although it is very convenient to have illustrations directly adjacent to the species descrip-
tions, the limited margin results in a small space for inadequate illustration printing. The effect for
SIDA 20(1): 448. 2002
BOOK REVIEWS
some of the ill i i ia
to be useful. For example, the illustration for Pse ae
menziesii includes a cross section of both leaf and stem. Both of these figures are too small to be
eful toa reader. Likewise, an illustration of a branch of Tsuga heterophylla is so small it it is hard to
dent theo lat
Perhap woud have pect more useful Had theaubior provided theiiles:
]
The color
photos in the uiidle of the book woule Be more wlan e if scale bars or another unit of measure
were included. As the plates are presented, the reader has no frame of reference for how large or small
each plant is in the photo. MomOvel it would
of | Letter OF number
lave been uel to identify each photo with some sort
ly cross-referenced w e caption. The discussion on forest com-
munities of Mount Rainier proves a bit confusing. Following the forest classification of another au-
thor, Biek states that there are four different forest zones within the Park. These zones are alae
ided into 19 associations, seven of which have two or more phases. An attempt i
ante
plants commonly growing in each zone, but clearly the distribution and limits of these en are
een on iaaneenl e factors, including annual precipitation, disturbance, elevation and
tem-
perature. From reading this discussion, it seems likely that many of the communities listed herein
are intertwined and thus difficult to classify.
e book has some handy features that are rather palsies Most keys are written with a
minimal amount of technical terminology, making them accessible to those with limited botanical
knowledge. The author has provided a map of the Park on the an cover for quick reference to its
readers. In addition, a metric ruler provided on the back cover proves convenient when keying outa
specimen in the field
Without doubt, this book is an important update to the knowledge of the flora of Mount Rainier.
Those who enjoy the flora of this area would certainly benefit by purchasing a copy of this book,
which should prove a worthy and practical field tool. This is a must-have for amateur and profes-
sional botanists who enjoy the Northwest United States—Amy Trauth Nare, Botanical Research Insti-
tute of Texas, Fort Worth, TX 76102-4060, U.S.A.amy_ trauth@yahoo.com
DonaLD H. MANSFIELD. 2000. Flora of Steens Mountain. (ISBN 0-87071-471-6, pbk.)
Oregon State University Press. 10] Waldo Hall, Corvallis, OR 97331-6407.
(Orders: http://osu.orst.edu, dept/press/osupress.htm, 800-426-3797, 541-
737-3166, 541-737-3170 fax). $29.95, 424 pp, 64 color photos, line drawings,
6" x 9"
According to the Mansfield, the f aes of Steens Mountain, Oregon includes “all of Steens Mountain
from Long Hollow near Fields to Riddle M st
f Diamond, the entire Alvord Basin, the Pueblo
Valley, Catlow Rim and the eastern edge of Catlow Valley, Diamond Craters, and the Malheur Marsh.”
The author also claims this book would be useful for the surrounding Pueblo Mountains and Harney
Basin as well as most of southeastern Oregon, southwestern Idaho, and northern Nevada. Indeed,
this publication would provide a useful tool for those interested in the flora of the northwestern
ae States.
e book’s introduction begins with a peiscuesion on ies it is organize ‘ specifics on the geo-
gr mF c region covered and more detailed i
short geologic and peoneannnen history of Steens follows, then a floristic overview of Sreens Moun-
tain and an explanation of vegetation zones and plant communities. Mansfield provides an interest-
ing discussion on what makes a plant rare, followed by a listing of some rare plants of Steens. Next, a
brief history of botanical exploration on Steens is given, as well as information on how to use the
SIDA 20(1): 449, 2002
BRIT.ORG/SIDA 20(1)
keys in this book and how to make a Pope plant collection. Last, a brief discussion of scientific anc
common names and the benefits and detriments of each is prov
arge portion of the book is dedicated to plants anda in a flora, begins witha key to the
paihe and is followed by an alphabetical, rather than phylogenetic, listing of families. Within each
family contains the genera and species found on Steens. The author has provided recognition fea-
tures, or identifying characters, for each family; t yeas that would be especially useful oe an amateur
dea agit ach family, sucl 1om1
botanist. As well, he includes interesting tidbits of in
importance of Poaceae and the fruit-bearing cultivars of Rosaceae. The etymology ol a genus
helps the reader understand the origins of sometimes difficult scientific names. Within each genus
isa key to species, followed by a brief description of each species, common names, habitat and distri-
eunoR After Ee listing ct aS a lial Bea follows, which cle ny defines potentially con-
hat follow. These illustrations woul a gra
| y of an-
ete in teaching the amateur botanist sour the seemingly infinite
giosperms. The author provides three separate indices for families, gener ae common names, re-
spectively.
Although this book is in general well written, | have but a few minor criticisms. The author
claims that the keys are written hie nls motivatec e novice in mind. As with most useful keys, how-
{ \ forsomeone witha limited knowl-
ever, much technical verbias ge |
edge of botanical terminology or diagnostic n
the positive side, the glossary included at the e a ol ce book is get for clarifying any ‘confusing’
plants on Steens Mountain, Mansfield explains why plants are en-
demic, citing two types, oo a neoendemics. While he states that the endemic plants of
Steens are neoendemics, he | for this observation other than a passing comment
on alpine habitat (p.14) The discussion also includes a brief discussion of rare plants in the context
| ters of {lowering plant families. On
terms. In his discussion of rare
of geologic history. Few, if any, amateur botanists or outdoor enthusiasts would have any understand-
ing of geologic time periods. This discussion may be confusing or have little value for the novice
reader. By comparison, the information that follows on how to use a key and make proper plant col
lections are integral in training the reader in proper botanical exploration and collection. In the cen-
ter of the book, the author provides some high quality color plates, but these photos would be more
der has no
valuable had scale bars or another unit of measure was provided. As they are given, the re
frame of reference for how large or small each plant is in the photo. In addition, it would have been
ae to sine each photo with some sort of as or num as so as to be e ay cross- oe
|
othe author's credit
ness of ae Flore of Steens Mountain, pen that “the flora of Steens Tone is saedainly not
finished..there will undoubtedly be more collections of plants not now included in the flora.” Too
ften, this point is forgotten when a nee such as this is published. Knowledge of biodiversity is
certainly not complete and the dynamic nature of systematics ensures that we will continue learn-
ing ee the flora of our continen
s book provides a ia addition to the knowledge of the flora of the Northwest. Cer-
tainly, outdoor enthusiasts and naturalists of southwest Oregon and surrounding areas would ben-
efit by purchas copy ol Flora of Steens Mountain. [tis the perfect size for a field guide and would
prove a valuable tool for those who tool around eee: —Amy Trauth Nare, Botanic al Research Insti-
tute of Texas, Fort Worth, TX 76102-4060, U.S.A. amy ]
Yr th@
POULTRY VOAFLOO.COTIL
SIDA 20(1): 450. 2002
BOOK REVIEWS 451
Robert H. MOHLENBROCK. 2001. The Illustrated Flora of Illinois. Grasses: Pani-
cum to Danthonia, Second Edition. Illustrations by Miriam W. Meyer and
Paul W. Nelson. (ISBN 0-8093-2360-5, hbk.) Southern Illinois University
Press. PO. Box 3697, Carbondale, IL 62902-3697. (Orders: 618-459-6633, 618-
453-1221 fax). $50.00, 455 pp, 306 line drawings, 51/2" x 81/2"
ROBERT A. MOHLENBROCK. 2001. The Hlustrated Flora of Illinois. Flowering Plants:
Pokeweeds, Four-o’clocks, Carpetweeds, Cacti, Purslanes, Goosefoots,
Pigweeds, and Pinks. Illustrations by Paul W. Nelson. (ISBN 0-8093-2380-
X, hbk.) Southern Hlinois University Press. PRO. Box 3697, Carbondale, IL
62902-3697. (Orders: 618-459-6633, 618-453-1221 fax). $59.95, 277 pp, 139
line drawings, 6" x 9".
ROBERT H. MOHLENBROCK. 2001. The Illustrated Flora of Illinois. Sedges: Cyperus
to Scleria, Second Edition. Illustrations by Fredda Burton and Mark W.
Mohlenbrock. (ISBN 0-8093-2358-3, hbk.) Southern Illinois University Press.
PO. Box 3697, Carbondale, IL 62902-3697. (Orders: 618-459-6633, 618-453-
1221 fax). $39.95, 223 PP 128 line drawings, 51/2" x 81/2",
These three bool 1 and two revised editions to the flora of Illinois. Each book
begins swith a county ee I] ee and a brief introduction about the g BroupScoveree wa Taxo-
( | | ps for | The
books als tain a list of species excluded, a brief and basic oy il ndasingle
ne for both h common ae scientific names. The editions of Grasses and Sedges have aes
which follow the s] These appendices seek to update any changes in nomencla-
ture and classification since the et etticn In the appendices, only taxa new to the flora are illus-
trated and distribution maps are not provided.
The Grasses volume covers half of the 99 genera found in the Illinois flora, including those in
the following sublamilies: Panicoideae, Eragrostoideae, Bambusoideae, Oryzoideae, and
Arundinoideae. The ee to the Grasses volume includes a ae but nonetheless informa-
of grasses. The author also includes sion of the variety of
tive, discussion of the mot pholog gy
grass habitats in Hlinois. The key to the genera of Grasses is illustrated and would serve novice
aprosieldeiets a nee 1 subs rainy, ale ande eae has a brief description, which is followed by
]
: r
the most impor tant featur es for identili-
. Also included are common name, habi-
—
aioe and 6 written nom observations of Hlinois materia
tat d ertinent taxonomic information. Following each
=
species name is dite and 1 pl ace of public: ation, synonyms, if applicable, and is cross-referenced with
an gece eo, ing illustration.
ges volume covers all genera of Cyperaceae except Carex. The Sedges bears many simi-
larities to t the C rasses volume. Like Grasses, the poupeu on pies Sa enor and norte dis-
cussion on Pei morphology and distribution of sedges. relationship
of sedges to other monocot families and describes how a are classified. The taxonomic treat-
ment crass an Mstiatd key to genera and brief descriptions of tribes and genera. Species de-
aa
found in t] The appendix that follows clearly details
adéieens and latur to the ILlinois flora. Many species of Scirpus, for example, have
been transferred to segregate genera, iaindludiae Isolepis, Bulboschoenus and Schoenoplectus.
The new Flowering Plants edition includes taxa from the Chenopodiales: Phytolaccaceae,
SIDA 20(1): 451. 2002
452 BRIT.ORG/SIDA 20(1)
Nyctaginaceae, Molluginaceae, Cactaceae, Portulacaceae, Chenopodiaceae, Amaranthaceae and
Caryophyllaceae. Polygonaceae, a family included in this order, is treated in a previous volume. The
Flowering Plants volume does not include a key to the families at the ae of the taxonomic
treatment, but it would have been useful. Instead, readers are encouraged to use Mohlenbrock’s com-
ail aa Guide tothe Vascular Flora of Illinois. After a five-page introduction, eae ees
“henopodiales. Species descriptions are provided in an identical structure as
the he volumes. Moreover, the author provides other useful tidbits such as a se to sterile or im-
mature specimens of genera in the Chenopodiaceae and a table that details t ces between
Chenopodium lanceolatum and C.album. The illustrations in this volume i a and I found
myself admiring them as | paged through the text. Never have [seen such weedy species presented in
such a lovely fashion.
ormation included in these three texts isan invaluable addition to the knowledge of the
in the Midwest United States should consider
=
=)
—_
flora of North America. In particular, those interestec
making this series a part of their library.—Amy Trauth Nare, Botanical Research Institute of Texas,
Fort Worth, TX 76102-4060, U.S.A.amy_trauth@ yahoo.com
Joun A. Parrorta. 2002. Healing Plants of Peninsular India. (SBN 0-85199-501-
2, hbk.). CABI Publishing, LO East 40th Street, Suite 3203, New York, NY
10016, U.S.A. (Orders: 212-481-7018, 212-686-7993 fax, cabi-nao@cabi.org).
$140.00, 944 pp, numerous color photos, 7" x 91/2",
Healing Plants of Peninsula India is an important addition to the therapeutic plant knowledge base.
This work compiles valuable information on 545 species of pete plants eee in traditional
T
Indian medicine. This is NOT a flora or identification guide, as there are | s. There are
however, superb descriptions and ae photographs for all species. “Species are oe ane alphabeti-
cally, by family, genus, and species”, and all species are listed, under the appropriate family, in the
table of contents.
] p +h} ' eat | ai ] ] J
he introductory material consists of a general 1
servation issues, and the importance of this book in nodueine readers to the eel y useful
floristic wealth of central and southern India. Itis the author’s hope that “this work will contribute
to the recognition and appreciation 1 of the oe le dge of plants and their therapeutic uses accumu-
lated and pieservecoue th oe le sacred cone soa oe forts
CO CONServe the diversity of tural | li li | phar m ACY.” ’Thereisa short
section on the cultural and historical roots of traditional Indian medicine, hichighane g ek
and Siddha medical concepts, and the importance of traditional medicine to the cultures of the re
a
gion. There is an interesting section of only a few paragraphs called: Botanical identification of tr
ditional plant drug sources: current challenges. This section describes a major sieblen facing Indian
traditional medicine generated by a lack of precise botanical descriptions in the traditional texts
and general lack of e saruus in sc apo noeueniae oe the hese ity of traders, pharmacists, and
!
Ayul iti ‘India isthe ee section of the
icreeneciien and describes the region in ie ul, with s specific nies mation on vegetation zones. The
species ae start soon aitera brief section on preparatio of the book
Each of the 545 species descriptions contains at least one color nee Descriptions are
arranged as follows: The scientific name is followed by a listing of synonyms and authorities. Com
mon names are listed in the appropriate languages of the region. Some common languages seen are:
Bengali, Hindi, Sanskrit, Tamil, Kannada, and English. This feature makes this volume useful for In-
dian traditional healers and botanists alike, as the common names known by the locals can be used
SIDA 20(1): 452. 2002
BOOK REVIEWS 453
to help find a good botanical description, with correct scientific nomenclature. Botanical descrip-
tions are thorough, as is information on distribution and habitat. Of course the real importance of a
work like this is best revealed under the subheading medicinal properties and uses. Here, one can
find information on the part of the plant used in preparation of a traditional drug, historical infor-
mation on which cultural groups used the plant, and the chemically active components, if they are
identified. Each description also has a short list of references.
n extensive 186 pages of end matter follows the descriptions. A Bibliography is followed by a
as
Glossary of Medical Terms anda Glossary of Botanical Terms. A very well done and complete Common
Name Index comes next, separated into _ Mee of the region. Also included are a Scientific
Name Index and an Index of Medicinal U
This is certainly one of the most extensive auc well-designed medicinal plant books to date.
=
jen
Hopelully its vast information and intuit ation will set a standard for other regional me-
dicinal plant books. This volume should help pres readers to the vast unexplored areas of study
in medicinal plants. Many of the species ae ibed in this work have not been studied scientifically,
and could be of great pispea canis) value. Final this book makes one fully aware of the impor-
tance of conserving all biologically dive ireas, especially those containing useful and potentially
revolutionary healing ee —Cole Weathe rby, Dept. of Biology, Austin College, Sherman, 1X 75090,
and Botanical Research Institute of Texas, Fort Worth, TX 76102, U.S.A, wweatherby@austinc.ec
a
u
STEPHEN Foster and CHrisTOPHER HOBBS. 2002. Peterson Field Guide to Western
Medicinal Plants and Herbs. (ISBN 0-395-83806-1, Flexi, ISBN 0-395-83807-
X, hbk.). Houghton Mifflin Company, 222 Berkeley Street, Boston, MA 02116-
3764, US.A. (Orders: 617-351-3243, gracie_doyle@hmco.com, wwwhoughton
mifflinbooks.com). $22.00 (Flexi), $30.00 (hbk). 448 pp, 530 + color pho-
tos, 41/2" « 71/4".
This Peterson field guide to medicinal plants is the second by author Stephen Foster. The Peterson
Field Guide to Medicinal Plants and Herbs of Eastern and Central North America, released in 2000,
marked a sort of revolution in plant field guides. Not only did this guide allow for the identification
of many plant species, using the “Peterson System”, but it also provided important information o
oe contem pay and historical medicinal uses and toxicity. Having used field guides exte ee
d myself wanting more information than the descriptions had to offer. This usually
meant a trip home to look the species up in my i flora and, often, another trip to the library to
search for articles in the scientific literature. While I usually enjoy doing this kind of detective work,
I found it fascinating to have more information av nee right there in the field, with the plant star-
ing me in the face
nit ially thisd urably land covered be yok resembles any oth ler fie >| d- worthy guide. Pwas a
ee surprised to find entck oo yages, each ustally present ing at somes one color photograph. T
ly adjacent as othe ae
son pie have done. cre isa very nice feature, as it can often be frustrating f disping back and forth
between ste a and photographs. The identification system is decent. As in most Peterson bo-
tanical field guides into six flower color sections, followed by shrubs, trees,
woody vines ue on allies, snd grasses and grass-like plants. In the color-separated sections
things are also grouped by various key flower characteristics. This system requires one to thumb
through practically every page until they find the section they are looking for. A table of contents at
the beginning of each flower color section showing the sub-sections would be very useful, and speed
SIDA 20(1): 453.2002
454 BRIT.ORG/SIDA 20(1)
up identification. I'm sure that my copy will soon be cluttered with self-adhesive notes sticking out
of my favorite sections, so this problem is fixable with a little work.
The biggest surprise came when | started reading the excellent descriptions this book offers.
‘ach species hasa short botanical description, followed by range, uses, and related species and warn-
ings when applicable. The “uses” section is, of course, the most significant thing this book has to olfer.
This usually presents information on how the plant was used historically, and how the plant was
and is prepared. It is interesting to see so much preparation information. Although it states plainly in
the mired venion that this “..book is not intended to be rescriptive or totake the place ol a healthcare
5)
provider...”, there is certainly enough information here on preparation and indications for use to al-
low one to aeons use these plants. Co-author Christopher Hobbs isa line nsed acupuncturist, herb-
alist, and consultant to the dietary supplement industry wh
yse EXertise on preparing these plants
for actual use is apparent upon reading these sections. Dosage infor aon israrely listed in the Uses
section, which seems potentially hazardous. This work provides readers with the necessary infor-
mation to make use of these plants by containing detailed preparation instructions, and then ne-
glects to give any dosage information, not even ine about over-dose. It is likely that it would
have been difficult to compile this information, and hability issue
=
s may have also prevented the in-
clusion of dosages. It would also be help! ‘ul to provide references for the specific historical informa-
tion presented in the text. However, there isa nice list of references in the back, which seems to com-
pensate for this.
Icons are used throughout this book to indicate poisonous plants, plants that are known to
cause allergic reactions or contact dermatitis, and those used in modern medicine. The end matter
includes a checklist of plant species, with blanks to write observed [lowering or fruiting dates, and
two indices: an index of plant species by common and scientific name, and another of medicinal
topics. The medicinal topics index is excellent, providing ae numbers of plant descriptions orga-
nized by their usefulness i in beaune ae medical condition
This guide for anyone with an interest in medicinal plants.
5 | I y
The extensive color pl | | | iu ious that great amounts of time
were invested in Hadise some eal the best p iorescanis avi re e book’s identification material.
detailed information on plant uses, and brilliant color ak ae it especially deserving of a
spot in your backpack.—Cole Weatherby, Dept. of Biology, Austin College, Sherman, TX 75090, and Bo-
tanical Research Institute of Texas, Fort Worth, 1X 76102, U.S.A. wweatherby@austinc.edu
SIDA 20(1): 454. 2002
BOOK NOTICES 455
Books RECEIVED/ NOTICES
Annotations and comments by the Editor
Timber Press’ Gardener’s Guide to Growing
MicHacL JEFFERSON-BROWN and Harris HOWLAND. 2002. Gardener’s Guide to Growing
Lilies. (ISBN 0-88192-537-3, pbk.; 0-88192-315-X, hbk.). Timber Press, Inc.,
133 S.W Second Avenue, Suite 450, Portland, OR 97204-3527, U.S.A. (Or-
ders: wwwtimberpress.com, 800-327-5680, 503-227-2878, 503-22 723070
fax). $29.95 (hbk, 1995), $19.95 (pbk, 2002), 160 pp., 70 color photos, +0 b/w
line drawings, 7" x 91/2".
Key Words: Lilies, Lilium, related plants, gardening, identification.
Diana GRENFELL. 2002. The Gardener's guide to Growing Daylilies. (ISBN 0-88192-
550"5, boks 0-63102-45l- x, hbk.). Timber Press, Inc., 133 S.W. Second
Avenue, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: www.timber
press.com, 800-327-5680, 503-227-2878, 503-227-3070 fax). $29.95 (hbk,
1998), $19.95 (pbk, 2002), 160 pp., 74 color photos, 5 line drawings, 7" x 91/
>"
jen
Key Words: Daylilies, Hemerocallis, gardening, horticulture, identification.
Martin PAGE. 2002. The Gardener’s Guide to Growing Peonies. (ISBN 0-88192-
535-7, pbk; 0-88192-408-3, hbk). Timber Press, Inc.; 133 5.W. Second
Avenue, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: www.tim ber
press.com, 800-327-5680, 503-227-2878, 503-227-3070 fax). $29.95 (hbk,
1997), $19.95 (pbk, 2002), 160 pp., 76 color photos, 18 b/w line drawings, 7"
x 91/2".
Key Worps: Peonies, gardening, horticulture, identification, 600 species and cultivars.
GraHam Rice and ELizABeTH STRANGMAN. 2001. The Gardener’s Guide to Growing
Hellebores. (ISBN 0-88192-517-9, pbk.; 0-88192-266-8, hbk.). Timber Press,
Inc., 133 S.W. Second Avenue, Suite 450, Portland, OR 97204-3527, U.S.A.
(Orders: wwwtimberpress.com, 800-327-5680, 503-227-2878, 503-227-3070
fax). $29.95 (hbk, 1993), $19.95 (pbk, 2001), 160 pp., 74 color photos, 20 line
drawings, 7" x 91/2"
Key Worps: Hellebores, gardening, horticulture, identification.
Grorr STEBBINGS. 2001. The Gardener’s Guide to Growing Irises. (ISBN 0-88192-
519-5, pbk.; 0-88192-388-5, hbk.). Timber Press, Inc., 133 S.W. Second
Avenue, Suite 450, Portland, OR 97204-3527, U.S.A. Oonice: www.timber
SIDA 20(1): 455. 2002
456 BRIT.ORG/SIDA 20(1)
press.com, 800-327-5680, 503-227-2878, 503-227-3070 fax). $29.95 (hbk,
1997), $19.95 (pbk, 2001), 160 pp., 74 color photos, 3 line drawings, 7" x 91/2"
Kry Worbs: Irises, gardening, horticulture, identification, cultivar.
DIANA GRENFELL. 2001. The Gardener’s Guide to Growing Hostas. (ISBN 0-88192-
518-7, pbk; 0-88192-355-9, hbk.). Timber Press, Inc., 133 S.W. Second Avenue,
Suite +50, Portland, OR 97204-3527, U.S.A. (Orders: www.timberpress.com,
800-327-5680, 503-227-2878, 503-227-3070 fax). $29.95 (hbk, 1996), $19.95
(pbk, 2001), 160 pp., 74 color photos, 5 line drawings, 7" x 91/2"
Kry Worps: Hostas, gardening, horticulture, identification, cultivars, propagation, over 400 culti-
vars and forms
Ecology/Conservation/Landscape Ecology
Ropert H. Rosicuaux and Davin A. YetMan (eds.). 2000. The Tropical Deciduous
Forest of Alamos: Biodiversity of a Threatened Ecosystem in Mexico. (ISBN
0-8165-1922-6, hbk.). The University of Arizona Press, 355 S. Euclid, Ste. 103,
Tucson, AZ 85719, U.S.A. (Orders: 520-621-1441, 520-621-8899 fax). $50.00
(hbk), 259 pp., numerous b/w photos, 6" x 9",
Key Worps: Mexico, Sonora, Biological diversity, tropical decid vegetation, flora, checklist
of en uc ee plants, trees in couheen Sonor a, ee, the Mayo eae ethnobotanically useful
trees, columnar cacti, Mayo people, crop diversity, amphibians and reptiles, birds.
BARBARA TeLLMAN (ed.). 2002. Invasive Exotic Species in the Sonoran Region . (ISBN
0-8165-2178-6, hbk.). The University of Arizona Press, 355 S. Euclid, Ste. 103,
Tucson, AZ 85719, U.S.A. (Orders: 520-621-1441, 520-621-8899 fax). $75.00
(hbk), 460 pp., 37 illustrations, 6 1/8" x 91/4",
Kry Worbs: Invasive plants, exotics, amphibians, mammals, insects, fish, birds, 500 naturalized ex-
otic species, Sonoran Desert, adjacent grasslands, riparian areas, native landscapes, Mexico, U.S.A,
environmental concerns.
Ropnry W. Bovey. Woody Plants and Woody Plant Management: Ecology, Safety,
and Environmental Impact. (ISBN 0-8247-0438-, hbk.). Marcel Dekker, Inc.,
270 Madison Avenue, New York, NY 10016, U.S.A. (Orders: 212-696-9000,
212-685-4540 fax, wwwdeekercom). Marcel Dekker AG, Hutgasse 4, Postfach
812, CH-4001 Basel, Switzerland (Orders: 41-61-261-8482, 41-61-261-8896
fax) $195.00, 564 pp., 6" x 9"
Kry Worbs: Woody plants, woody plant management.
From the back cover: “Consolidating an extensive breadth of resources on woody plants, this
reference/text on the historical significance and botanical nature of woody plants details strategies
for managing woody plants and using research data to select the most : PRE ane eo
av
y
able
ods—analyzing the responses of over 370 North American woody |
herbicides.”
SIDA 20(1): 456. 2002
BOOK NOTICES 457
Ethnobotanical/E ic/Medicinal/Useful Plants/Toxic
Marta R. Emery and repecca J. McLain (eds.). 2001. Non-timber Forest Products:
Medicinal Herbs, Fungi, Edible Fruits and Nuts, and Other Natural Prod-
ucts from the Forest. (ISBN 1-56022-089-9, pbk.). Food Products Press, 10
Alice Street, Binghamton, NY 13904-1580 U.S.A. (Orders: 607-722-5857, 607-
722-1424 fax, getinfo@haworthpressinc.com). Price not listed, 176 pp., 6"
x 81/4".
Key Worps: Non-timber forest products, North America, review, history, sustainable uses.
NARAYAN P. MANANDHAR. 2002. Plants and People of Nepal. (ISBN 0-88192-527-
6). Timber Press, Inc., 133 S.W Second Avenue, Suite 450, Portland, OR 97204-
3527, U.S.A. (Orders: www.timberpress.com, 800-327-5680, 503-227-2878,
503-227-3070 fax). $69.95, 636 pp., 48 color photos, 3 tables, 834 line drawings,
2 maps, 81/2" « 11",
Key Words: Plants, Nepalese people, flora, ethnobotany, descriptions, common names.
This is quite a book with fantastic color photos of plants and Nepalese people. There are no
keys so the drawings, although not sophisticated, are excellent aids in identification. What a great
reference with several useful appendices.
Science/Society/Economy
DorotHyY SCHWIEDER, THOMAS Morain, and LYNN NicLsEN. 2002. lowa Past to Present:
The people and the Prairie. Third edition. (ISBN 0-8138-2998-4, hbk.). lowa
State Press, 2121 State Avenue, Ames, IA 50014, U.S.A. (Orders: 1-800-862-
6657, 1-515-292-3348 fax; ww [ ).$32.99, 312 pp., numerous
b/w photos and other illustrations, 7" x 10".
Key Worps: lowa, history, juvenile literature, American Indians, pioneers, prairies, settlers, frontier,
farms
Stuart McCook. 2002. States of Nature. Science, Agriculture, and Environment
in the Spanish Caribbean. (ISBN 0-292-75256-3, hbk.; 0-292-75257-1, pbk.).
University of Texas Press, P.O. Box 7819, Austin, TX 78713-7819, U.S.A. (Or-
ders: 800-252-3206 fax; 512-471-4032). $50.00 (hbk), $22.95 (pbk), 201 pp.,
6 photos, | map, 6" x 9"
Key Worps: Science, history Latin America, Caribbean, Venezuela, Puerto Rico, Colombia, Costa Rica,
environment, nature, economy, agriculture botanical science.
GERHARD SONNERT (with assistance of Gerald Holton). 2002. Ivory Bridges: Con-
necting Science and Society. (ISBN 0-262-19471-6, hbk.). The MIT Press, 5
Cambridge Center, Cambridge, MA 02142-1493, U.S.A. (Orders: 800-405-
1619, mitpress-orders@mit.edu, mitpress.mit edu). $30.00 (hbk), 227 pp.,
6" x 9
SIDA 20(1): 457. 2002
458 BRIT.ORG/SIDA 20(1)
Key Worps: Science, society, governmental science policy, scientists’ voluntary public-interest asso-
ciations, “pure” science, research in the national interest, activities of citizen scientists
Botany/Molecular/Physiology
Thomas J. GIVNisH and KENNETH J. SytsmaA. 1997. Molecular Evolution and Adap-
tive Radiation. (ISBN 1-521-57 329-7, hbk.). Cam bridge University Press, The
Edinburgh Building, Cambridge CB2 2RU, UK; 40 West 20" Street, New
York, NY 10011-4211, U.S.A. $39.95, 621 pp., b/w photos, figures, 7" x 10"
Key Worbs: Evolution, adaptive radiation, convergence, reproductive strategies, : lutionary
patterns.
There were forty authors who contributed to the book’s seven chapters: 1) Introduction; 2) In
tegrative Studies; 3) Convergence; 4) Rapid Radiations; 5) Reproductive Strategies; 6) Character Di-
vergence and Community Assembly; and 7) Macroevolutionary Patterns
R.S. SINGH. 2000. Diseases of Fruit Crops. (ISBN 1-57808-159-9, pbk; 1-57808-149-
|, hbk.). Science Publishers, Inc. PO. Box 699, Enfield, NH 03748, U.S.A. (Orders:
sales@scipub.net; wwwsscipub.net, or info@scipub.net). $49.50 (pbk), 310
pp., 31 b/w photos, 6" x 91
Kry Worbs: Fruit diseases, fruit pests, citrus, pome and stone fruits, mango, banana, grapevines and
grapes, guava, strawberry, coconut palm, pineapple, pomegranate, litchi, loquat, cashewnut, sapo-
dilla plum.
GEORGE G. KHACHATOURIANS, ALAN MCHucen, Rater Scorza, Wat-Kit Nip, and Y.H.
Hui(eds.). 2001. Transgenic Plants and Crops. (ISBN 0-8247-0545-9, hbk.).
Marcel Dekker, Inc., 270 Madison Avenue, New York, NY LOO16, U.S.A. (Or-
ders: 212-696-9000, 212-685-4540 fax, wwwdeekercom). Marcel Dekker
AG, Hutgasse 4, Postfach 812, CH-4001 Basel, Switzerland (Orders: 41-61-
261-8482, 41-61-261-8896 fax) $225.00, 876 pp., b/w photos, 7" 10".
<ty Worbs: Transgenic plants, crops, transgenic techniques, agriculture, molecular biology, food plant
biotechnology, genetics, social, political, legal aspects of genetic plant manipulation.
Yoav WalseL, AMRAM Eset, and Uzi KarKari (eds.). 2002. Plant Roots. The Hid-
den Half. Third Edition. (ISBN 0-8247-0631-5, hbk.). Marcel Dekker, Inc.
270 Madison Avenue, New York, NY 10016, U.S.A. (Orders: 212-696-9000.
212-685-4540 fax, wwwdeekercom). Marcel Dekker AG, H utgasse 4, Postfach
812, CH-4001 Basel, Switzerland (Orders: 41-61-261-8482, 41-61-261-8896
fax) $250.00, 1120 pp., color and b/w photos, line drawings, 8 1/2" x 11".
Kry Worps: Roots, characteristics of roots, root system, root structure, development, root genetics,
regulation of root growth, physiological aspects of root systems, roots under stress, roots of various
ecological groups, roots of economic value, roots with medicinal activity
SIDA 20(1): 458. 2002
BOOK NOTICES 459
This is quite a book on the subject of roots. There are some 99 individuals listed as contributing
to this incredible and massive compilation. The references follow each of the 59 chapters rather than
at the end. The editors are to be congratulatec
Anpre O. BareL, Marc Poaye, and Howarp L. Maisacu (eds). Handbook of Cosmetic
Science and Technology. (ISBN 0-8247-0292-1, hbk.). Marcel Dekker, Inc.,
270 Madison Avenue, New York, NY 10016, U.S.A. (Orders: 212-696-9000,
212-685-4540 fax, wwwdeekercom). Marcel Dekker AG, Hutgasse 4, Postfach
812, CH-4001 Basel, Switzerland (Orders: 41-61-261-8482, 41-61-261-8896
fax) $235.00, 886 pp., b/w photos, and other figures, 7" x 10"
Key Worbs: Cosmetics, cosmetic ingredients, finished products, target organs, delivery systems, cur-
rent technology in sacl), toxicology, dermatological testing.
TI
his appears up-to-date reference on the cosmetic industry. Part 1) Introduction, Part
2) Target Oueins for Cosmetic Products; Part 3) Safety Considerations; Part 4) Vehicles of Cosmetic
Products; Part 5) Cosmetic Ingredients; Part 6) Cosmetic Products; Part 7) Legislation and Regula-
tions Worldwide; Part 8) Testing Cosmetic Products; and Part 9) Cosmetic Claims. Contributions
from some 92 eapett scientists from acade mic dermatology and dermato-cosmetics, the cosmetics
industry, i |
toregulatory agencies, helped the ere toachieve
their goal.
MoHAMMAD PEssSARAKLI (ed). Handbook of Plant and Crop Physiology. (ISBN 0-
8247-0546-7, hbk.). Marcel Dekker, Inc., 270 Madison Avenue, New York,
NY LOO16, U.S.A. (Orders: 212-696-9000, 212-685-4540 fax, wwwdeekercom).
Marcel Dekker AG, Hutgasse 4, Postfach 812, CH-4001 Basel, Switzerland
(Orders: 41-61-261-8482, 41-61-261-8896 fax) $225.00, 973 pp., b/w pho-
tos, graphs, and other figures, 7" 10"
Key Worps: Plant crops, physiology, environmental factors, climatic changes, developmental states,
growth regulators.
This is a comprehensive, up-to-date reference on plant and crop physiology. Part 1) Plants/
Crops Growth Responses to Environmental Factors and Climactic Changes; Part 2) Physiology of
Plant/Crop Growth and Developmental Stages: Part 3) Cellular and Molecular Aspects of Plant/
Crop Physiology; Part 4) Plant/Crop Physiology and Physio
=
ogical Aspects of Plant/Crop Produc-
tion Processes; Part 5) Plant Growth Regulators: The Natural Hormones (Growth Promoters and In-
hibitors) and Plant Genes; Part 6) Physiological Responses of Plants/Crops Under Stressful (Salt,
rought, and Other Environmental Stresses) Conditions; Part 7) Physiological Responses of Plants/
Crops to Heavy Metal Concentration and Agrichemicals; Part 8) Physiological Relationships Between
Lower and Higher Plants; Part 9) Physiology of Lower-Plant Genetics and Development, Part 10)
Physiology of Higher-Plant/Crop Genetics and Development; Part 11) Using Computer Modeling in
Plant Physiology; and Part 12) Plant/C top a) siology ° nder Controlled Conditions, in Space, and
on Other Planets. There were some 76 spec to this volum
SIDA 20(1): 459. 2002
460 BRIT.ORG/SIDA 20(1)
Biographical
Witrrip Blunt (Introduction by William T. Stearn). 2001. Linnaeus: The Compleat
Naturalist. (ISBN 0-691-096 36-8, pbk.). Princeton University Press, 41 William
Street, Pinceton, NJ 08540, U.S.A. (Orders: California Princeton Fulfillment
Services, 1445 Lower Ferry Road, Ewing, NJ 08618, U.S.A., 609-883-1759,
609-883-7413 fax, www.pup.princeton.edu). $35.00, 264 pp., color photos,
73/4" x 93/4"
—_—
Key Worns: Linnaeus, biography, taxonomy, systematic botany, history.
Journals
Acta Botanica Hungarica. (ISSN 0236-6495). Akadémiai Kiado, Budapest, H-1519
Budapest, PO. Box 245, HUNGARY. (Subscriptions: 36-1-464-8221, email:
kiss.s@akkrt.hu). $192.00, 4 issues/year, 6 1/2" x 91/2"
CONTENTS FOR 4303-4) 2001 1) In honour of the seventy-year-old Gabor Fekete. 2) Jaraine Komlodi
Magda is seventy-years old. 3) Les componantes de la balance hydrique des deux type des mais
differents dans les deux périodes de végétation successives. 4) Anatomical investigation of 4,000-
years old Cedrus libani wood remains from Egypt. 5) Revision taxonomica del género Stevensia Poit.
6) Total soluble protein amount in vegetative buds and needles of Norway spruce aps - burst-
time. 7) Melitopalynological and antimicrobial properties of honeys from Elazi_ (E Turkey). 8)
effect of chemical and biological insecticides on growth, pigmentation, photosynthetic activity and
ey productivity of Vicia faba ee 8) Coenological data on temperate semidesert sandy grass-
lands in Hungary. 9) Soil seed bank and vegetation composition of two fen-meadow stands in the
Netherlands. 10) The elfects of low temperature on the growth and total protein amount of spa tae
(Cicer arietinum) seedlings. 11) Cambial activity and development of xylem in T
L. growing in different forests of Gujarat state. 12) Morpho-histogenic studies of the fruit wall of
Argemone mexicana L. (Pap idleness 13) Effect of industrial effluents polluting the river Nile or
growth, meta Pelion and productivity of Tritic st 1 Vicia faba plants. 14) Floral anatomy
of Act Ssq | Bidens biternata (Asteraceae).
SIDA 20(1): 460. 2002
—
“You don't have to be a botanist to use it! All the tools you need
to teach plant taxonomy, family characters, classification, etc
Excellent introductory materials on the history and geology of
North Central Texas. Great for identifying indigenous plants in
an ecosystem! Essential for student wildflower collection
projects! Use this one book instead of five, it has it all.”
—Sharon Foster, 7th grade teacher,
Fort Worth Country Day School
“The best available, user-friendly source of information about
our native and naturalized plants. This book should be in the
library of every gardener, landscape architect, horticulturist,
urban planner, farmer, rancher, and wildflower enthusiast.”
—Howard Garrett, The Dirt Doctor
“INCREDIBLE, FANTASTIC, WONDERFUL, BEAUTIFUL, ETC. WOW!
To be more specific: COMPREHENSIVE, ENCYCLOPEDIC,
INNOVATIVE. With every species illustrated - it’s a new
departure in Texas floras, a breath of fresh air.”
—Marshall Johnston, Co-author,
Manual of the Vascular Plants of Texas
“The Illustrated Flora on North Central Texas is a magnificent work.
This abundantly illustrated volume will not only be of use to
those people wanting to identify Texan plants, it will be a
model of how a local flora should be prepared.”
—Sir Ghillean Prance FRS
Director, Royal Botanic Gardens, Kew
neceitt & MAHLER’S
ILLUSTRATED
FLORA OF
NORTH CENTRAL
TEXAS
Now IN ITS SECOND PRINTING!
DETAILED BROCHURE WITH ORDER FORM
AVAILABLE BY REQUEST ON THIS GROUND
BREAKING NEW FLORA.
Catt BRIT Press at
817 332-4441 or
EMAIL: SIDA@BRIT.ORG
FOR HOW TO ORDER.
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SOURCE OF
SIDA CONTRIBUTIONS TO
BOTANY WAS FOUNDED BY
LuoyD H. SHINNERS (LEFT)
IN 1962. INHERITED BY
WILLIAM F. MAHLER
(RIGHT), DIRECTOR EMERITUS
oF BRIT In 1971, AND
SINCE 1993, IT HAS BEEN
PUBLISHED BY Brit PRESS.
aie
ey?
Now CELEBRATING FORTY YEARS...
23 Nov 1962—23 Nov 2002
BRIT
PRS
Barney L. Lipscome, EDITOR
Botanical Research Institute of Texas
509 Pecan Street
Fort Worth, Texas 76102-4060, USA
817 332-4441 / 817 332-4112 FAX
Electronic mail: sida@brit.org
Home page at the URL: http://www.brit.org/sida/
JOHN W. THIERET, ASSOCIATE EDITOR
Dept. of Biological Sciences
Northern Kentucky University
Highland Heights, KY 41099 USA
FEuIx LLAMAS, CONTRIBUTING SPANISH EDITOR
Dpto. de Botanica, Facultad de Biologia
Universidad de Leon
E-2471 Leon, SPAIN
Guidelines for contributors are available upon request
and on our SIDA home page as well as the last pac
of each issue.
Subscriptions for year 2003:
$85. Individual
sie) USA Institutions
55. Outside USA
numbers issued twice a year
OSIDA, CONTRIBUTIONS TO BOTANY
VOLUME 20, NUMBER 2, PAGES 461-876.
23 NovemBer 2002
CopyriGHT 2002
BOTANICAL RESEARCH INSTITUTE OF TEXAS
PRINTED IN THE UNITED STATES OF AMERICA
SSN 0036-1488
TABLE OF CONTENTS
SYSTEMATICS
NEW AND NOTEWORTHY ORCHIDS OF THE BISMARCK ARCHIPELAGO, PAPUA New GUINEA
N.H.S. Howcrort AND W. TAKEUCHI—461
UNA ESPECIE NUEVA DE PORTULACA (PORTULACACEAE) DEL NORTE DE MICHOACAN (MEXICO)
GILBERTO OCAMPO ACOSTA—487
GAVACEAE), UNA NUEVA ESPECIE DE MAGUEY DEL NORESTE DE MEXICO
GREG STARR AND JOSE A. VILLARREAL O.—495
A NEW SPECIES OF LITHOSPERMUM (BORAGINACEAE) FROM Nuevo LEON, Mexico
B.L. TURNER—501
A NEW SPECIES OF SCHISTOCARPHA (ASTERACEAE: HELIANTHEAE) FROM NORTHWESTERN OAXACA AND CLOSELY ADJACENT
GUERRERO, MEXICO
TURNER—505
SENECIO SPRIBILLEI (ASTERACEAE: SENECIONEAE), A NEW SPECIES FROM MONTANA, U.S.A.
WILLIAM A. WeeerR—511
EUCHITON (ASTERACEAE: GNAPHALIEAE) IN NORTH AMERICA AND HAWwall
Guy L.Nesom—515
NEW COMBINATION IN SALIX (SALICACEAE)
Guy L. NEsom—523
TAXONOMY OF THE MOST COMMON WEEDY EUROPEAN ECHINOCHLOA SPECIES (POACEAE: PANICOIDEAE) WITH SPECIAL EMPHASIS
ON CHARACTERS OF THE LEMMA AND CARYOPSIS
MIHAI COSTEA AND FRANGOIS J. TARDIF—525
SEED AND FRUIT CHARACTERS IN SELECTED SPERMACOCEAE AND COMPARISON WITH HEDYOTIDEAE
Epwarb E. TERRELL AND RICHARD P. WUNDERLIN—549
RUBIACEAE)
Notes ON CARPHEPHORUS ODORATISSIMUS (ASTERACEAE) IN PENINSULAR FLORIDA, U.S.A.
Steve L. ORZELL AND EDWIN L. Bribces—559
RESOURCE ALLOCATION PATTERNS AND PHENOTYPIC VARIATION IN THE ENDANGERED TEXAS WILDRICE (ZIZANIA TEXANA, POACEAE)
PAULA POWER—
NOMENCLATURE OF THE HEART-LEAVED HEDGE-NETTLE, STACHYS CORDATA (LAMIACEAE)
JAMES S. PRINGLE—583
A cypress (CUPRESSUS ARIZONICA, CUPRESSACEAE) IN Jerr Davis COUNTY, TEXAS?
M. Patrick GRIFFITH AND STEPHANIE C. BARTEL—
DAHLIA NEGLECTA (ASTERACEAE: COREOPSIDEAE), A NEW SPECIES FROM SIERRA MADRE ORIENTAL, MEXICO
Day_e E. SAAR—593
THE VARIETIES OF LIATRIS ELEGANS (ASTERACEAE)
Mark H. MAYFIELD —597
DOcUMENTED CHROMOSOME NUMBERS
DOCUMENTED CHROMOSOME NUMBERS 2002: 3. CHROMOSOME COUNTS FOR PACKERA PAUPERCULA VAR. APPALACHIANA
(ASTERACEAE: SENECIONEAE
ALISON McKENZIE MAHONEY, VIRGINIA W. KONG, AND RoBerT R. KowAL—605
ETHNOBOTANY
NORWEGIAN AND SAMI ETHNOBOTANY OF VERATRUM ALBUM (MELANTHIACEAE)
TORBJORN ALM—611
BOTANICAL HisTORY
Carto BERTERO (1789-1831) IN THE New Wor_b
PicRO G, DELPRETE, GIULIANA FORNERIS, AND ANNALAURA PISTARINC—62 1
—
ON THE DERIVATION OF THE GENERIC NAME ILIAMNA (MALVACEAE)
WituiAM A. WEBER AND PAUL A. FRYXELL—645
FLoRISTICS
WHAT |S THE WRITER OF A FLORA TO DO? EVOLUTIONARY TAXONOMY OR PHYLOGENETIC SYSTEMATICS ?
Georce M. Dicas, JR. AND BARNEY L. LiPSCOMB—647
THE VASCULAR FLORA OF THE LAGUNA DE ALEGRIA, A NATIONAL MONUMENT IN THE CRATER OF TECAPA VOLCANO, EL
SALVADOR
DENISE WILLIAMS, RuDyS W. HERRERA, AND JOSE L. LINARES—675
THE FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI
ANDREAS LEIDOLF, SIDNEY MCDANIEL, AND Tim NUTTLE—691
ENDEMISM IN THE WEST GULF COASTAL PLAIN: IMPORTANCE OF XERIC HABITATS
MicHaet H. MACRoseERTS, BARBARA R. MaAcRoserts, BRUCE A. SORRIE, AND Ropert E. EvANS—767
ANNOTATED CHECKLIST OF THE VASCULAR FLORA OF THE HICKORY CREEK UNIT OF THE BIG THICKET NATIONAL PRESERVE,
TYLER COUNTY, TEXAS
BARBARA R. MAcRoserts, MICHAEL H. MACROBERTS, AND LARRY E. BROWN—781
VASCULAR PLANTS OF THE MAPIM/ BIOSPHERE RESERVE, MEXICO: A CHECKLIST
ABEL GARCIA-AREVALO—797
A SIGNIFICANT NEW POPULATION OF THE RARE SEMAPHORE PRICKLYPEAR CACTUS, OPUNTIA CORALLICOLA (CACTACEAE)
KEITH A. BRADLEY AND STEVEN W. WooDMANSEE—809
PLANTS NEW TO THE UNITED STATES AND FLORIDA
RICHARD P. WUNDERLIN, BRUCE F. HANSEN, AND LorAN C. ANDERSON—-813
CRYPTOCORYNE BECKETTH COMPLEX (ARACEAE) INTRODUCED AT A FLORIDA SPRING
COLETTE C. JACONO—819
NEPHROLEPIS EXALTATA (DRYOPTERIDACEAE) NEW TO ARKANSAS
JAMES H. PECK—833
CYNOSURUS ECHINATUS (POACEAE) NEW TO TEXAS
R. DALE THOMAS—-837
ADDITIONS TO THE FLORA OF NAGS HEAD Woops (DARE COUNTY, NORTH CAROLINA) AND THE OUTER BANKS OF NORTH
AROLINA
ALEXANDER KRINGS—839
CIRSIUM NUTTALLII (ASTERACEAE: CYNAREAE) NEW TO NORTH CAROLINA AND AN ILLUSTRATED KEY
ALEXANDER KRINGS, RANDY WESTBROOKS, AND JANINE LLoyYo—845
REINSTATEMENT OF VIBURNUM OZARKENSE (CAPRIFOLIACEAE): AN ENDEMIC TAXON OF THE INTERIOR HIGHLANDS OF ARKANSAS,
MISSOURI AND OKLAHOMA
TIMOTHY J. WECKMAN—849
COMMEMORATIVE
CELEBRATION HONORS PIONEER FIELD BOTANIST ANDRE MICHAUX
WILLIAM R. BURK—861
Det WENIGER (1923-1999), REMARKABLE TEACHER AND NATURALIST
MARJORIE FOHN EPLER—865
BOOK REVIEWS AND NoTICES 494, 500, 504, 514, 522,558, 570,604, 620, 766, 796, 808, 812, 818, 836, 838, 844 867
ANNOUNCEMENT—-876
INDEX TO NEW NAMES AND NEW COMBINATIONS IN SIDA 20(2), 2002.
AGAVE OVATIFOLIA STARR & VILLARREAL, SP. NOV.— 495
DAHLIA NEGLECTA D.E. SAAR, SP. NOV.—
DENDROBIUM ARCHIPELAGENSE HOWCROFT & TAKEUCHI, SP. NOV,— 464
DENDROBIUM VEXILLARIUS J.J. SM. VAR. HANSMEYERENSE HOwcrcOFT & TAKEUCHI, VAR. NOV.—462
LIATRIS ELEGANS MICHX. VAR. BRIDGESI MAYFIELD, VAR. NOV.—598
LIATRIS ELEGANS MICHX. VAR. ELEGANS FORMA FLABELLATA (SMALL) MAYFIELD, COMB. NOV.—603
LIATRIS ELEGANS MICHX. VAR. KRALI MAYFIELD, VAR. NOV.—601
LITHOSPERMUM HINTONIORUM B.L. TURNER, SP. NOV.—501
PORTULACA RZEDOWSKIANA G. OCAMPO, SP. NOV.—487
SALIX EXIGUA VAR. SERICANS (NEES) NESOM, COMB, NOV.—523
SCHISTOCARPHA CALZADANA B.L. TURNER, SP. NOV.—505
SENECIO SPRIBILLEI WA. WEBER, SP. NOV.—511
¥
NEW AND NOTEWORTHY ORCHIDS OF THE BISMARCK
ARCHIPELAGO, PAPUA NEW GUINEA
N.H.S. Howcroft W. Takeuchi
Papua New Guinea Forest Service Harvard University Herbaria and Arnold Arboretum
-O.Box 406 c/o Papua New Guinea Forest Research Institute
Rabaul, East New Britain 614 Papua New Guinea Forest Service
GUINEA P.O.Box 314, Lae, Morobe Province 411
howcroft@daltron.com.pg PAPUA NEW GUINEA
ABSTRACT
A general account is provided of the orchid gatherings from the 1994 Conservation International
survey of southern New Ireland, an area identified by environmental assessment as one of Papua
(T) a era hi ] - >) |
New Guinea's major terrestrial unknown regions. One nev pelag
one new variety (Dendrobium vexillarius JJ. Sm. var. hansmeyerense) are described and illustrated.
Several distributional records are reported. Illustrations are provided for most of the cited taxa.
Kry Worns: Botanical survey, Dendrobium, New Ireland, Orchidaceae, Papua New Guinea
JAPANESE ABSTRACT
AFF =sa—-FATOPCORSLRHMEM BENT Sih e NS. =a
THI YEBRRBICKIT, 1994 (CI YE RK-Yartyvag—-tyatwles
SRE DNA, COMBICS|\CTREANAMICRT SHASTA.
Dendrobium archipelagense & D. vexillarius J.J. Sm. var. hansmeyerense 7) 2D
OPMOMORARICBL CitM eC MRMET. HOMODAMMICMLCORAT
®o
INTRODUCTION
Originally a part of the German Territory of New Guinea, the Bismarck Archi-
pelago consists primarily of New Britain, New Ireland, and Manus. These islands
are now separate provinces of the sovereign state of Papua New Guinea (PNG).
The Archipelago is collectively designated as the ‘Islands Region’ by national
administrators, and also includes several minor islets scattered outboard of New
Ireland, as for example, the St. Matthias group, Lihir, and Tabar (Fig. 1).
The Bismarck Archipelago represents one of PNG’s most poorly docu-
mented floristic regions (Takeuchi & Golman 2001). It is unfortunately the most
intensively logged territory in the country: 50% of the archipelagic land area is
currently under timber concessional development (ibid.). Because of the pau-
city of biodiversity information, the forested tracts in southern New Ireland are
ranked among PNG’s 16 terrestrial unknowns and are also included in the country
portfolio of high-priority sites for conservation action Johns 1993: 25; Sekhran
& Miller 1995; 113-15).
SIDA 20(2): 461 — 486. 2002
462 BRIT.ORG/SIDA 20(2)
St Matthias
%,
Manus :
La ae
Bismarck Archipelagc
New
mA, Australia 4
Fic. 1. Island of New Guinea, and the Bismarck Archipelago.
In order to assess the status of its environments, from January 14 to Febru-
ary 16, 1994, a 13-member team conducted an elevation-sequenced survey in
the Weitin Valley and Hans Meyer Range of southern New Ireland. Although
the survey's collective findings have been recently published (Beehler & Alonso
2001), the orchid vouchers were treated ina separate checklist (Howcroft 1994).
In the following discussion, the orchids from the 1994 survey are given further
consideration. Occurrence notes from other areas within the Archipelago have
also been added when appropriate.
DESCRIPTIONS OF NEW TAXA
Dendrobium vexillarius J.J. Sm. var. hansmeyerense Howcrolt & Takeuchi, var.
nov. (Fig. 2). Type: PAPUA NEW GUINEA. NEW IRELAND PROVINCE: Hans Meyer Range,
Angil Mountain, expedition camp 4 in pristine subcloud forest, 425.2’ S, 15256.8' E, 1800 m, 6
Feb 1994 (£1, fr, spirit), W. Takeuchi & J. Wiakabu 9453 (HOLOTYPE: LAE).
7 “Jetat Lire,
A var. vexillarius sepalis latioribusc baxial
ssep ue ab
Distribution and ecology —Known thus far only from the type locality, in mossy
montane forest at 1800 m.
Etymology.—The new variety is named after the type locality,
Dendrobium vexillarius was previously represented by six varieties sepa-
rated primarily by the color of the perianth (Reeve & Woods 1989: 250). A promi-
nent abaxial crest on the lateral sepals distinguishes var. hansmeyerense from
other infraspecific taxa.
The new variety was collected earlier during the 1975 Kew expedition to
New Ireland (ie., Sands et al. 1985, 2349, 2374, 2399). Reeve and Woods (1990)
had tentatively assigned these specimens to variety uncinatum. However that
variety is not prominently crested on the abaxial side of the lateral sepals as
with var. hansmeyerense. Keels are sometimes present among the varieties of D.
HOWCROFT AND TAKEUCHI, ORCHIDS OF THE BISMARCK ARCHIPELAGO 463
poor
Mie
Fic. 2. Dendrabi (Marius J.J.Sm. var. h H ft & Tak hi. A. Habit. B. Flower, side view. C. Flower,
from below. D. Fl f I E. Labellum, midsecti I ia ae apex with and (G) without anther.H.
Anther, front view. I. Antt ith pollen, obli ( ior) view.J. Ovary in cross -section. Scale bars: A, 4 cm; B—
E, 2 cm; F—-G, 3mm; H-I (enlarged 30x); J, 3mm. D f I y NHS, Howcroft.
464 BRIT.ORG/SIDA 20(2)
vexillarius but they are not prominent. The only other taxon in this section
with a prominent abaxial keel is D. brevicaule Rolfe ssp. pentagonum (Kraenzlin)
Reeve & Woods, with an otherwise very differently shaped ovary in cross-sec-
tional view,
Dendrobium archipelagense Howcrolt & Takeuchi, sp. nov. (Fig. 3). Typr PAPUA
NEW GUINEA, NEW IRELAND PROVINCE: junction of Niagara and Weitin rivers, expedition
camp 2, lowland rainforest, +30.210'S, 15290.242' FE, 240 m, Jan 1994 (f1, spirit), W. Ta
& J. Wiakabu 9979 (HOLOTYPE: LAE; ISOTYPE: A)
—
reuchi
l) ]
‘cei Dendrobio dntennato Lindley auiniesee ene ert aia ac icamplitudine
labelli, lobis sali 5 g oe ICIS G 1ON Gua idt rat I
r t
basim midlobi, et lamella triangulari carenti in parte Sista idlobe
Tufted epiphyte. Rhizomes short. Pseudobulbs swollen at the base, fusiform in
the lower part, distally more slender, subcylindrical, with persistent nodal
sheaths shorter than the internodes, leafy, to 50 by 1.0-2.0 cm. Leaves distichous,
lanceolate-ovate, 7.5-9.5 by 2.0-3.0 cm, asymmetrically emarginate, adaxially
slightly concave, abaxially convex, thick, coriaceous, green to yellow-green. In-
florescence racemose, one or several per pseudobulb, inserted at the apex or
slightly below, erect, rachis to 40 cm by 2-3 mm; peduncle to 9 cm long, pedun-
cular and floral bracts tubular; pedicel glabrous, ca. 2.2 cm long. Flowers 5-10
per raceme, usually wide-opening, 4. we 5 by 3.5cm, pale green to yellow-green.
Dorsal sepal recurved, twisted through one turn, oblong-lanceolate, 2.0-2.6 by
0.6-0.7 cm, acuminate. Lateral sepals recurved, oblong-lanceolate, 2.5-3.3 by 0.9
cm at the base, apex acute or acuminate. Mentum narrowly conical, 0.9-1.0 cm
long, apex straight or slightly decurved. Petals linear, acute, 3.0-4.5 by 0.25-0.4
cm, twisted through one or two turns, spreading, Labellum porrect, 3-lobed, 2.0-
2.3 by 1.0-15 cm, similar in color to sepals and petals but with some purple
venation; lateral lobes oblong-elliptic, 1.15 by 1.5 cm, over-lapping base of the
midlobe, apex rounded; midlobe ovate, 1.0 by 1.0 cm, acute to apiculate, not re-
curved; callus of 5 keels, the lateral ones not extending past the base of the
midlobe, the median keel extending to the apex of the midlobe, slightly raised,
not lamellate or crested. Column 0.5 cm long, apex laterally bilobed, foot at 10°
to column. Ovary glabrous, 0.9 cm long.
Distribution and ecology.—Widely distributed throughout the Bismarck
Archipelago, frequently on trees near the coast but also from inland areas. The
type locality is lowland rainforest at 240 m.
The new species seems to flower mainly during the first half of each year.
In addition to the type specimen (flowering in Jan), cultivated plants have pro-
duced flowers continuously from May to Jul. Seeds from the previous year's set,
are released simultaneously with the production of new flowers.
Etymology.—The epithet reflects the distributional restriction to the Bis-
marck Archipelago.
Within section Spatulata Lindl. the novelty’s closest relative appears to be
HOWCROFT AND TAKEUCHI, ORCHIDS OF THE BISMARCK ARCHIPELAGO 465
10 mm
, bh
jy fies!
Mya!
Fic. 3. Dendrobi hinel H ft &Tal i. A. Flower, aspect.B. Dorsal sepal.C. Lateral sepal.D. Labellum.E
hl 1 col F. Col from below.G. A her, frontal
view. Scale bars: A- E 10 mm; F, 4 mm; G-H, 2 mm. Drawn from the type by N.H.S. Howcroft.
466 BRIT.ORG/SIDA 20(2)
Dendrobium antennatum Lindl. In his treatment of this section, Cribb (1986:
618) recognized several species groups, among them the antennatum group with
9 species. The new species was formerly placed in D. strepsiceros J.J.Sm. by Cribb
(ibid: 637), O'Byrne (1994: 236), and Lavarack et al. (2000: 262).
Smith’s illustration of D. strepsiceros (1913: Fig. 4, Bl, B2) shows that its la-
bellum differs from the corresponding characters in the new species. In D.
strepsiceros the labellum midlobe is larger than the lateral lobes, and the apices
of the latter do not overlap the base of the midlobe as it does in live specimens
of D. archipelagense (Fig. 3). This distinction can be lost in the preparation of
dried specimens so the relationship between the lobes is best seen with spirit
material. Apart from the difference in shape and extension of the lateral lobes,
lagense hasa shorter labellum compared to D. strepsiceros and is more
similar to D.antennatum on that feature.
Compared to its closest congeners, the lateral lobes of the new species are
proportionately larger in relation to the length of the labellum. In D.
archipelagense, the lateral lobe is 70-80% of the labellum length, in D.
antennatum 55-70%, and in D. strepsiceros 50-58% (Fig. 4).
There are five ridges on the callus of the labellum in all of the preceding
species. The mid-callus ridge for D. archipelagense extends towards the apex of
the midlobe, but is not raised to a triangular lamella as described for D.
strepsiceros. This character may have been overlooked by other authors.
The new species has long petals which make the flower appear larger than
typical D. antennatum. As many as ten flowers can be produced by one inflo-
rescence. The color of the perianth is initially light green but turns yellow-green
with age. In most individuals, the ridges of the labellum are bordered with vio-
let, and there are also violet reticulations up to the median, and sometimes the
distal portion of the midlobe. The perianth is persistent on the fruit, a charac-
teristic shared by other species in the antennatum group.
No significant color differences have been seen between specimens from
West New Britain, East New Britain, and New Ireland. However in terms of
flower size, the type collection represents one of the largest and most attractive
forms seen by the senior author.
,:
D. archipe
£
DISTRIBUTIONAL RECORDS
Arachnis beccarii Reichb. f. var. imthurnii (Rolfe) Tan, Selbyana 1:1-15. 1975; 1:365-
373. 1976. (Fig. 5).
Specimen examined: PAPUA NEW GUINEA. New Ireland Province: junction of Niagara and Weitin
rivers, expedition camp 2, lowland rainforest, 430.210’ S, 15256.242' FE, 240 m, Jan 1994 (fl, spirit), W.
Takeuchi & J. Wiakabu 9968 (LAE).
The genus Arachnis is represented by two species in New Guinea, of which only
A. beccarii occurs in the Bismarck Archipelago.
Variety imthurnii had been previously recorded from Bougainville,
HOWCROFT AND TAKEUCHI, ORCHIDS OF THE BISMARCK ARCHIPELAGO 467
l
Mid lobe
length
Labellum length
S
3 C
|
di
Mere
wa alot
Fic. 4. A. Labell truct B1. Dendrobi + | g 1] Sm., from Smith (1913) RB?.) strepsiceros from Cribb
(1986). C. D. ant tum Lindl. D. D hipel H ft & Tak hi. All scale bars: 10 mm.
468 BRIT.ORG/SIDA 20(2)
f
F syle
4297
Fic 5 A L re L. tes eae Oe | € ara imthurnii (Rolfe) Tan A ri , f 7 R 1 7 ll j | side view
C. Labellum and column, frontal view. D, Labellum and column, cross-section. E. Anther, frontal view. F. Anther, from
above. G. Anther, fi back. showi N.H.S
wing pollen. Scale bars: A-G, 1 cm. Drawn from W. Takeuchi & J. Wiakabu 9968
7
Howcroft.
HOWCROFT AND TAKEUCHI, ORCHIDS OF THE BISMARCK ARCHIPELAGO 469
Guadalcanal, Malaita, and Santa Isabel (Lewis & Cribb 1991). It was also seen
by the senior author on the Toriu River side of the Kanak Range in East New
Britain. The New Ireland specimen is a distributional record for the island.
Although var. imthurnii is reported as having an entirely white perianth
(ibid.), the New Ireland provenance is marked by large brown spots. Cultivated
plants in the National Botanical Gardens at Lae have also produced flowers with
color markings different from mainland and archipelagic provenances. The dis-
tinctions can be used to justify eventual reinstatement of this variety to spe-
cific rank.
—_—
Corybas epiphyticus (J.J. Sm.) Schltr, Feddes Repert. Spec. Nov. Regni Veg. 19:21.
1923. (Fig. 6)
Specimen acoeeiees PAPUA NEW GUINEA. New Ireland Province: Hans Meyer Range, Angil ous
tain, ex p 4, pristine subcloud forest dominated by bryophytic and ferny growth, 425.2
152: 56.8 E, 1800 m, 6Feb 1 994 (fl, spirit), W. lakeuchi & J. ae 9461 (LAE).
ude known only from Indonesian Papua (van Royen 1983). The collec-
tion from New Ireland represents a significant eastward extension of the spe-
cies range. Corybas epiphyticus is still unrecorded from the PNG mainland.
Dendrobium gnomus Ames, J. Arnold Arbor. 14:106-107. 1933. (Fig. 7).
Specimen examined: PAPUA NEW GUINEA. New Ireland Province: Hans Meyer Range, ridge adja-
cent to the Weitin River, expedition camp 3, mossy montane forest, 427.205'S, 15256.489' EF, 1175 m,
27 Jan 1994 (fl, spirit), W. Takeuchi & J. Wiakabu 9597 (LAE).
This species was initially mistaken for Dendrobium erosum (BL) Lindl. because
of the spathulate shape of the labellum and its finely erose margins (Fig. 7F).
However the leaf apex is unequally bilobed as with D. gnomus (Fig. 7A), whereas
in D. erosum the apex is acute. The cucullate labellum apex on the New Ireland
specimen is probably an artefact of drying. The shape of the sepals and petals,
and the mentum being appressed to the ovary and pedicel rather than diverg-
ing, are also consistent with the redetermination (Figs. 7B-F).
The species had not been previously recorded for New Ireland.
Pedilochilus longipes Schltr, Feddes Repert. Spec. Nov. Regni Veg. Beih. 1:691.
Fig. 8)
1912. (Fi
Specimen examined: PAPUA NEW GUINEA. New Ireland Province: Hans Meyer Range, Angil Moun-
tain, expedition camp 4, pristine subcloud forest dominated by ee tic and ferny growth, 425.2’
S, 152568 E, 1800 m, 4 Feb 1994 (FD), W. Takeuchi & J. Wiakabu 9380 (LA
Previously known with certainty only from mainland New Guinea. The New
Ireland specimen represents a distributional record.
Phaius amboinensis Bl., Mus. Bot. Lugd.-Bat. 2:180. 1856. (Fig. 9).
Specimen examined: PAPUA NEW GUINEA. New Ireland Province: junction of Niagara and Weitin
rivers, expedition camp 2, lowland rainforest, 430.210! S, 15256.242' FE, 240-300 m, 15 F a 994 (1, fr,
spirit), W. Takeuchi & J. Wiakabu 9780 (A, LAE).
470 BRIT.ORG/SIDA 20(2)
5mm
= spe
ABs
Fic. 6. Corybas epip me (J.J. Sm.) Schitr. A. Flower and leaf, side view. B. Dorsal sepal. C. Labellum, frontal view. D.
Labellum, ith fl d out. E. Column, sh d F. Column with labellum re-
moved and shoving two lateral cae Scale bars: A-D, 1 cm; E-F, Sr mm. Drawn from W. Takeuchi & J. Wiakabu 9461 by
N.H.S.H
HOWCROFT AND TAKEUCHI, ORCHIDS OF THE BISMARCK ARCHIPELAGO 471
jblfonadl?
pe 3ioue
Fic.7. Dendrobium g A A. Leaf. B. Fl ide view. C. Dorsal sepal. D. Petal. E. Lateral sepal. F. Labellum. G
Column, obliq ques ide view Column ovary, and spur Antt J), Anther from side. Scale bars: A— B,1cm;C
H, 5 mm; IJ, 2 mm. Drawn from W. Takeuchi & J Wiakabu 9597 by N.H.S. Howcroft.
ce
BRIT.ORG/SIDA 20(2)
5mm
|
ts.B.—C. Flower, aspect. D. Dorsal sepal.E. Lateral sepal.F. Petal.G.
rom above. Scale bars: A, 4 cm; B-C, 1 cm;
side view H ditto oblique view. I. ditto
y N.H.S. Howcroft
wee
ahs
D-I,5 mm. Drawn from W. fakenchh &lb
HOWCROFT AND TAKEUCHI, ORCHIDS OF THE BISMARCK ARCHIPELAGO 473
Fic. 9. Phaius amboinensis Bl. A. Flower, side aspect. B. Dorsal sepal.C. Petal. D. Lateral sepal.E. Labellum, side view. F.
Labellum, flattened, view from above. G. Column, from side. H. Column, from below. Scale bars: A—H, 2 cm. Drawn from
W. Takeuchi & J. Wiakabu 9780 by N.H.S. Howcroft.
-
474 BRIT.ORG/SIDA 20(2)
A widespread orchid found on many islands in the Bismarck Archipelago, but
not previously recorded from New Ireland. The species distribution includes
Vanuatu, Fiji, Samoa, and Tahiti in the east. Also occurring in Java, the southern
Philippines (Sulu Archipelago), Sulawesi, and the Moluccas.
Phreatia loriae Schltr, Feddes Repert. Spec. Nov. Regni Veg. 3:318. 1907.
Specimen ex eee PAPUA NEW GUINEA. New Ireland Province: junction of Niagara and Weitin
rivers, eX] camp 2, lowland rainforest, 430.210’ S, 15256.242' E, 240 m, 21 Jan 1994 (f1, spirit),
W. Takeuc oi & J. Winkel 9885A (LA
The species was ue known only from the southern PNG mainland.
= montana (Schltr) Cretz. @ J.J.Sm., Acta Fauna Fl. Univ. Bucur. ser. 2,
, 14:4. 1934. (Fig. 10).
Specimen examined: PAPUA NEW GUINEA. New Ireland Province: Hans Meyer Range, Angil Moun-
tain, expedition +, pristine subcloud forest dominated by bryophytic and oy growth, 425.2’
S, 152568 E, 1800 m, “4 Feb 1994 (f1, spirit), W. Takeuchi & J. Wiakabu 9394 (1
The labellum of this specimen compares well with the illustration by Halle
(1977). The new occurrence represents a significant disjunction from the previ-
ously known stations in Vanuatu and New Caledonia.
OTHER ILLUSTRATED COLLECTIONS
pee Pt ane al Schltr, Feddes Repert. Spec. Nov. Regni Veg. 16:
125. 1920. (Fig. 11).
Specimen examined: PAPUA NEW GUINEA. New Ireland Province: Hans Meyer Range, Angil Moun-
tain, expedition camp 4, pristine subcloud forest dominated by bryophytic and ferny growth, 425.2’
S, 15256.8' E, 1800 m, 3 Feb 1994 ({l), W. Takeuchi & |. Wiakabu 9375 (Lé
Bulbophyllum pachyglossum isa member of section eer within which
3 species have been described for New Guinea. Of these, only B. pachyglossum
has been recorded from the Bismarck Archipelago. The species is also known
from the Solomon Islands (Lewis & Cribb 1991).
Cadetia echinocarpa Schltr, Feddes Repert. Spec. Nov. Regni Veg. Beih. 1:439.
1912. (Fig. 12).
Specimen examined: PAPUA NEW GUINEA. New Ireland Province: junction of Niagara and Weitin
rivers, expedition camp 2, low | rainforest, 430.210' S, 15256,242' E, 240 m, 21 Jan 1994 (£1), W.
Takeuchi & J. Wia it 9884 (LAE
Possibly synonymous with the later name C. homochroma (JJ. Sm.) Schltr
Calanthe triplicata (Willemet) Ames, Philip. J. Sci. Bot. 2:325. 1907. (Fig. 13).
Specimen oe PAPUA NEW GUINEA. New Ireland Province: junction of Niagara and Weitin
rivers, €X 2, lowla nee 430.210'S, 15256.242' E, 240 m, Jan 1994 (f1, spirit), W.
eh & J. Wiakabu 9976 (A, KL, LAF).
Widespread in New Guinea, the Bismarck Archipelago, and Southeast Asia.
HOWCROFT AND TAKEUCHI, ORCHIDS OF THE BISMARCK ARCHIPELAGO 475
4mm
ae
Ye
Fic. 10. Pristiglottis montana (Schitr.) Cretz. & J.J.Sm. A. Flowering habit. B. Labellum from cla . AN mith ate
i D. Labellum, from side. Scale bars: A, 10 mm; B—D, 4 mm. Drawn from W. Takeuchi & J. |
roft.
Fic. 11. Bult
BRIT.ORG/SIDA 20(2)
sepal. G. Labellum and column, side view. H. Labellum, side view. |
II hvali Schitr A Flowering habit. B. Flower. from side € Dorsal sepal D-E. oe : ca
ala
bars: A, 5 cm; B—G, 10 mm; H-I, «12. Drawn from W. Takeuchi & J. Wiakabu 9375 by N.H.S. Howcroft.
HOWCROFT AND TAKEUCHI, ORCHIDS OF THE BISMARCK ARCHIPELAGO 477
ayn
‘ail
Fic. 12. Cadetia echi Schitr. A. Stem and leaf. B. Flower, entavies G HTOMey and ovary, side view. D. Dorsal
sepal. E. Petal. F. Lateral eeual G. Eabellum, from: above. H. ditto: Le y fl iehoaang aneiet column,
£ hal
stigma, spur, an nd sheaths.J. Column
Scale 2 A, 5 cm; B-G, 5 mm; H, 2 mm; |, 5 mm; J-K, 2 mm. Drawn from W. Takeuchi & J. Wiakabu 9884 by N. H.S.
Howcro
BRIT.ORG/SIDA 20(2)
478
ee) a
Tem
F
Fic. 13. Calantt i fi (Will yA A. Fl “fi lvi B. Fl side vi C Dorsal sepal D. Lateral sepal
E. Petal. F. Labell it f I G. Column, side view. H. Floral bract. Scale bars: A-H, 1 cm. Drawn from W.
Takeuchi & J. Wiakabu 9976 by N.H.S. Howcroft.
HOWCROFT AND TAKEUCHI, ORCHIDS OF THE BISMARCK ARCHIPELAGO 479
Dendrobium cuthbertsonii F Muell., Trans. Roy. Soc. Victoria 24:175. 1888.
(Fig. 14)
Specimens examined: PAPUA NEW GUINEA. New Ireland Province: Hans Meyer Range, Angil Moun-
tain, expedition camp 4, pristine subcloud forest dominated by bryophytic and ferny growth, 425.2’
S, 15256.8' E, 1800 m, 3 Feb 1994 (f1, spirit), W. Takeuchi & J. Wiakabu 9341 (LAE); ibid., 4 Feb 1994
(LD), W. Takeuchi & J. Wiakabu 9400 (LAE): ibid., 6 Feb 1994 (f1, spirit), W. eae & J. Wiakabu
9452 (A, LAE)
The surface of the leaves is not verrucose, as in most mainland provenances.
Dendrobium masarangense Schltr. ssp. masarangense, Repert. Spec. Nov. Regni
Veg. 10:78. 1911. (Fig. 15).
Specimen examined: PAPUA NEW GUINEA. New Ireland Province: Hans Meyer Range, ridge adja-
cent to the Weitin River, expedition camp 3, mossy montane forest, 427.205' S, 15256.489 E, 1175 m,
26 Jan 1994 (fl, spirit), W. Takeuchi & J. Wiakabu 9534 (LAE).
The white-flowered ssp. masarangense, with a light yellow-tipped labellum, is
widespread and common on high islands. Lewis and Cribb (1991: 188) list its
distribution as Sulawesi to New Guinea, New Britain, Bougainville, Guadalcanal,
Vanuatu, Fiji, and New Caledonia. The senior author has examined this sub-
species in situ from the Lelet plateau of New Ireland. However much of the habi-
tat there has been recently destroyed by agricultural development and the or-
chid has now disappeared from that locality.
Dendrobium rhodostictum F Muell. & Kraenzl., Oest. Bot. Zeit. 44:300. 1894.
(Fig. 16)
Speci S ined: PAPUA NEW an New Ireland Province: Hans ae ee ridge adja-
cent to the Weitin Riv <pedition cam ssy montane forest, 427.205’ S, 15256.489' F, 1175 m,
25 Nee 1994 ({1, ae W. Takeitel hi & J. on 9514 (LAE); junction of oe a Weitin rivers,
p 2, lowland rainforest, 430.210’ S, 15256.242’ E, 240 m, Jan 1994 (£1), W. Takeuchi &
BE Wiakal bu 9974 (LAE)
Previously recorded from New Ireland. The pseudobulbs of these specimens
are less clavate than other collections from New Britain, Bougainville, and the
mainland, but the perianth segments and callus agree perfectly in shape and
size to the species.
Goodyera rubicunda (Bl.) LindL, Bot. Reg. 25: 61, misc. 92. 1839. (Fig. 17). Neottia
rubicunda BL, Bijdr. 408. 1825.
Specimen ined: PAPUA NEW GUINEA. New Ireland Province: junction of Niagara and Weitin
rivers, sieved at camp 2, lowland rainforest, ise 210' S, 15256.242' E, 240-300 m, 13 Feb 1994 (1,
spirit), W. Takeuchi & J. Wiakabu 9808 (A, K, L, LAE).
In the broad sense, Goodyera rubicunda is distributed throughout Malesia, from
peninsular Malaysia to Samoa.
Spathoglottis plicata Bl., Bijdr. Fl. Ned. Ind. 5: 401. 1825. (Fig. 18).
Specimen examined: PAPUA NEW GUINEA. New Ireland Province: junction of Niagara and Weitin
Fic. 14. Dend
robium cuthbertsonii F. Mu
bell , side view. E
ell. A. Flowering h
abit. B.
from ab
£.
BRIT.ORG/SIDA 20(2)
, aff
Mer
Flower, frontal view. C. Flower and spur, side view. D.
ove E cae j hd £1 £e° L . £. 1
Scale bars: AE, 10 mm. Drawn from W. Takeuchi & J.
ing. G. Leaf in cross-section. H. Enlarged view of ovary
52 by N.H.S Howc
Wiakabu 94
HOWCROFT AND TAKEUCHI, ORCHIDS OF THE BISMARCK ARCHIPELAGO 481
Fic. 15. Dendrobj. g Sch] ssp. masarangense A Flowering habit. B. Fl of lvi C. Flower and
spur, side view. D. Dorsal sepal. E. Petal. F. Lateral sepal. G. Labellum (basal part omitted). H. Flower with labellum
removed, showing column, anther, stigma, and spur. I. Leaf cross-section. Scale bars: A, 10 mm; B-I, 4 mm. Drawn from
W. Takeuchi & J. Wiakabu 9534 by N.H.S. Howcroft.
482 BRIT.ORG/SIDA 20(2)
Fic. 16. Dendrobium rhodostictum F. Muell. & Kraenzl. A. Flowering habit. B. Dorsal sepal. C. Lateral sepal. D. Petal. E.
Labellum (d ged). F. Labellum fi | i fl G. Callus f ture labellum (in subdiag E). H. Callus
from immature labellum (in subdiagram F). Scale bars: A, 5 cm; B—-F, 1 cm; G—-H, 5 mm. Drawn from W. Takeuchi & J.
Wiakabu 9574 by N.H.S. Howcroft.
HOWCROFT AND TAKEUCHI, ORCHIDS OF THE BISMARCK ARCHIPELAGO 483
NESS
5 mm wi
ee
Fic. 17. Good bicunda (BI.) Lindl. A. Fl f | view. B. Flower, side view. C. Dorsal sepal. D. Lateral sepal. E
7 A ff i
Petal. F. Labellum, frontal view. G. Labellum, from above. H
section.I. Column.J. Anther.
Scale bars: A-B, 1 cm; C-J,5 mm. Drawn from W. Takeuchi & J. Wiakabu 9808 by N.H.S. Howcroft.
484 BRIT.ORG/SIDA 20(2)
rivers, expedition camp 2, lowland rainforest, 430.210’ S, 15256.242' E, 240 m, Jan 1994 (f1, fr, spirit),
W. Takeuchi & J. Wiakabu 9972 (LAE).
The New Ireland specimen undoubtedly belongs to the S. plicata complex. How-
ever the ligulate lateral lobes on the labellum, with virtually no dilation of the
apices, indicate that this is not subspecies plicata. The lack of a distinctly arched
column is suggestive of cleistogamy.
Spathoglottis taxa are distinguished primarily by the labellum, especially
by the shape and angle of the lateral lobes, shape/size of the callus and its
vestiture, and by claw length and width. Plant habit and morphology is of value
in defining sections and subspecies (Howcroft 1986). Research by the senior
author indicates that six subspecies of S. plicata are present in the New Guinea
region, of which four have been recorded from the Bismarck Archipelago. A
formal revision of Malesian Spathoglottis will appear ina future publication by
the first author.
ACKNOWLEDGMENTS
Flowers of Dendrobium antennatum were provided by Wolf gang Bandisch, Gen-
eral Manager of the National Capital Botanical Gardens (NCBG), and by Judith
Raka, NCBG Administration Manageress and Scientific Officer. Specimens rep-
resenting New Ireland provenances were received from Oscar Melepia, of
Gaulim in East New Britain, and from Steven Kami of Port Moresby.
The 1994 New Ireland expedition was funded by Conservation Interna-
tional and the U.S. Agency for International Development (through the
Biodiversity Support Program). Bruce Beehler (Conservation International) and
the Papua New Guinea Department of Environment and Conservation were
the principal planners and organizers for the expedition. Hitofumi Abe (Eco-
system Research Group, University of Western Australia) provided the Japa-
nese translation. Guy Nesom (Botanical Research Institute of Texas) wrote the
Latin diagnosis for Dendrobium archipelagense, and John J. Pipoly HI (Fairchild
Tropical Garden) wrote the Latin diagnosis for D. vexillarius var. hansmeyerense.
Reviewers Paul Omerod and Andre Schuiteman made many helpful comments
on the manuscript.
REFERENCES
BeeHter, B. and L.E. ALonso (eds). 2001. Southern New Ireland, Papua New Guinea: a
biodiversity assessment. RAP Bulletin of Biological Assessment 21,Conservation Inter-
national, Washington, DC.
Crise, PJ. 1983. A revision of Dendrobium sect. Latouria. Kew Bull. 38:229-306.
Crips, PJ. 1986. A revision of Dendrobium sect. Spatulata. Kew Bull. 41:615-692.
Crips, PJ. and C.Z. Tana. 1982. Spathoglottis in Australia and the Pacific Islands. Kew Bull.
36:721-729.
HOWCROFT AND TAKEUCHI, ORCHIDS OF THE BISMARCK ARCHIPELAGO 485
F
5 mm dem
Aiipooore
ye Bee
Fic. 18. Spathogl li BI.A. Mature bud. B. Dorsal sepal.C. Lateral sepal. D. Petal. E. Col ith labell side
view. F. Labellum, from ad G-H. Bilobed callus. Scale bars: A-F, 1 cm;G—H, 5 mm. Drawn from W. Takeuchi & J. Wiakabu
9972 by N.H.S. Howcro
486 BRIT.ORG/SIDA 20(2)
Dockritt, A.W. 1992. Australian indigenous orchids, revised edition. Surrey Beatty & Sons
Pty Ltd, NSW Australia.
Hate, N. 1977. Flore de la Nouvelle-Caledonie et Depéndances 8: Orchidacées. Museum
National d'Histoire Naturelle, Paris.
Howcrort, N.H.S. 1986. The taxonomy of Spathoglottis Bl. Orchadian 8:139-150.
Howcrort, N.H.S. 1994. Orchid collections from New Ireland. Orchid Res. Bull. 3. Papua New
Guinea Forest Research Institute.
Jouns,R.J. 1993. Biodiversity and conservation of the native flora of Papua New Guinea. In:
B. Beehler, ed. Papua New Guinea conservation needs assessment report, vol. 2. PNG
Dept. of Environment and Conservation, Boroko. Pp 15-222.
Kores, PJ. 1989. A precursory study of Fijian orchids. Allertonia 5:1-222.
LAvARACK, P.S., W. Harris, and G. Stocker. 2000. Dendrobium and its relatives. Kangaroo Press,
NSW, Australia.
Lewis, B.A.and PJ. Criss. 1989. Orchids of Vanuatu. Royal Botanic Gardens Kew. Whitestable
Litho Printers Ltd.
Lewis,B.A.and PJ.Crigs. 1991.Orchids of the Solomon Islands and Bougainville. Royal Botanic
Gardens, Kew. Whitestable Litho Printers Ltd.
O'Byrne, P. 1994, Lowland orchids of Papua New Guinea. SNP Publishers Pty Ltd Singapore.
Reeve, .M.and PJ.B.Woops. 1990.A revision of Dendrobium sect. Oxyglossum (Orchidaceae).
Notes Royal Bot. Gard. Edinb. 46:1—305.
Royen, P. van. 1980. The orchids of the high mountains of New Guinea. J.Cramer, Germany.
Reprinted from:P. van Royen. 1979. The Alpine flora of New Guinea. Vol. 2. Orchidaceae.
J.Cramer, Germany. Pp 51-812.
Royen, P.vAN. 1983.The genus Corybas in its eastern areas. Phanerogamarum Monographiae
Tomus XVI. J.Cramer, Germany.
SCHLECHTER, R. 1911-14. Die Orchidaceen von Deutsch-Neu-Guinea. Feddes Repert. Spec.
Nov. Regni Veg. Beih. 1:1-1079. English translation 1982. D.F. Blaxell, ed. The Australian
Orchid Foundation, Melbourne
SeKHRAN, N. and S. Miiter (eds). 1995. Papua New Guinea country study on biological diver-
sity. Colorcraft Ltd, Hong Kong.
Smit, JJ. 1913. Orchidaceae. Nova Guinea 12:1-108, t.14, 39.
Takeuchi, W.and M.Goiman. 2001.Botanical documentation imperatives: some conclusions
from contemporary surveys in Papuasia. Sida 19:445-468.
Tan, K. 1975. Taxonomy of Arachnis, Armodorum, Esmeralda and Dimorphorchis
(Orchidaceae), Part |. Selbyana 1:1-15.
Tan, K. 1976. Taxonomy of Arachnis, Armodorum, Esmeralda and Dimorphorchis
(Orchidaceae), Part Il. Selbyana 1:365-373.
UNA ESPECIE NUEVA DE PORTULACA (PORTULACACEAE)
DEL NORTE DE MICHOACAN (MEXICO)!
Gilberto Ocampo Acosta
; do EF, if i
titut ga, AC
pS. Id RAZ
NEGlOrai 1 DG}l
Apdo. Postal 386, CP6I 1600
Pdtzcuaro, Michoacdn, MEXICO
ocampo@inecolbajio.edu.mx
RESUMEN
Se describe ie se pispone: a ponuulden neenavisntd nd como eee nueva ek la ciencia. De acuerdo
con el zal y matorral sub-
tropical eee de la parte norte del estado de Michoacan. Se caracteriza | ser
de habito poses con raiz enpetosa, por poseer hoj lanad Brollies laneceladencea jones
oblongo-| las y por el ppereals de la capsula que es de forma campanulada.
PALABRAS CLAVE: Portulaca, Portulacaceae, Michoacan, México.
ABSTRACT
Portulaca rzedowskiana is described and proposed as a new species. According to examined mate-
rial this taxon is found in grasslands and disturbed subtropical scrub in the north part of the Mexi-
can state of Michoacan. It is ee mainly by its ee : a with suberous root, by its flat
to subterete lanceolate leaves, s g-lanceolate t iceolate in shape, as well as by
the campanulated ee of the ae
Key worps: Portulaca, Portulacaceae, Michoacan, Mexico.
Durante la preparacion del fasciculo correspondiente a la familia Portulacaceae
para la Flora del Bajio y de regiones adyacentes, se encontraron algunos
ejemplares de herbario del género Portulaca, los cuales fueron colectados en la
parte norte del estado de Michoacan (México) y que corresponden a una entidad
que se puede ubicar en la subseccion Conocarpae D. Legrand (subgénero Portu-
laca, seccion Catoclasis D. Legrand), cuyo tnico miembro reportado para México
es P mexicana P. Wilson. Estos ejemplares presentan caracteristicas que no
corresponden a ninguna de las especies conocidas en la actualidad, por lo que
se propone como:
Portulaca rzedowskiana G. Ocampo, sp. nov. (Fig. 1). Tipo: MEXICO. MICHOACAN,
{unicipio de Morelia: Monte Rubio, pastizal, terreno plano, orilla de camino, por la via del
tren, alt. 1900 m, 1 me 1991, J.M. Escobedo 2035 (HOLOTIPO: IEB; ISOTIPOS: por distribuirse).
Herba is crassi adix incrassata, suberosa, 2-6 cm longa, 0.5-3.5 cm lat |
ad 26 cm longi, prostrati ae nee folia alterna, plana vel subteretia, fanenclata sliguands
'Trabaj izad smico del Instituto de Ecologia, A.C. (cuenta 902-03), del Consejo Nacional
de C Ciencia viene y de la Comision Nacional para el Conocimiento y Uso de la Biodiversidad.
SIDA 20(2): 487 — 493. 2002
488 BRIT.ORG/SIDA 20(2)
oblongo-lanceolata vel oblanceolata, 4-20 mm longa, 1-3(-4.5) mm lata, apice acuta; sepala ovata,
rubra, (3-)4.5-6.5 mm longa, 4-5.5(-6.5) mm lata; petala rosea vel purpurea, obovata, (5.5-)6.5-11 mm
longa, (3-)3.5-6(-7) mm lata; stamina 22-44; stylus 4.5-6(-8) mm longus, lobis stigmatosis (3-)4-5(-
6); capsula 3.5-6 mm longa, operculum campanulatum; semina coracina, 0.6-0.7(-0.8) mm longa,
tuberculata, tuberculis basi stellulatis ornata.
Planta herbacea perenne, carnosa. Raiz engrosada, suberosa, de 2-6 cm de
largo por 0.5-3.5 cm de ancho, en raras ocasiones surgen de ella raices
cilindraceas de hasta 15 cm de largo por 2-4 mm de ancho. Tallos varios
surgiendo de la raiz, de 4-26 cm de largo, postrados a decumbentes. Hojas
alternas, aplanadas a subrollizas, carnosas, pelos axilares de 3-5 mm de largo,
peciolo de 1-1.5(-2) mm de largo, lanceoladas, en ocasiones oblongo-lanceoladas
a oblanceoladas, de 4-20 mm de largo por 1-3(-4.5) mm de ancho, pice agudo.
Flores terminales, sésiles, agregadas en cabezuelas de (2-)3-8 flores, rodeadas
por un verticilo de (6-)8-LOG-13) hojas, pelos de hasta 5 mm de largo; sépalos
connados en su base, rojizos, ovados, de (3-)4.5-6.5 mm de largo por 4-5.5(-6.5)
mm de ancho, desiguales, apice agudo, margen escarioso, persistentes hasta la
madurez de la capsula; pétalos de color rosa a morado, connados en su base,
obovados, de (5.5-)6.5-11 mm de largo por (3-)3.5-6(-7) mm de ancho, obtusos
a emarginados, la mayoria de las veces con un pequeno mucro6n; estambres 22-
44, sus filamentos de (2.5-)3-4(-5) mm de largo, connados en su base formando
un anillo estaminal de +1 mm de largo, anteras de 0.6-0.8(-0.9) mm de largo;
estilo de 4.5-6(-8) mm de largo, ramas estigmaticas (3-)4-5(-6), de 1.5-2.5(-3)
mm de largo. Capsula sésil o sobre un pedicelo de hasta 1.5 mm de largo, la
capsula de 3.5-6 mm de largo por 3-4.5(-5) mm de diametro, con dehiscencia
en su tercio inferior o mas cercana a la base, opérculo campanulado a semi-
campanulado; semillas de color negro-azulado cuando maduras, brillantes,
reniformes, de 0.6-0.7(-0.8) mm de largo, tubérculos redondeados con estélulas
radiales en su base.
Mi eee adicional e sala México. Michoacan. Municipio de Acuitzio del Canje: Las Penas, | Jul
1986, H. Diaz 2372 (CHAPA, IEB). Pianteapyo de Charo: brecha rumbo a Las Mesas, 22 Jul 1987, J. San-
tos ere B), +15] bre el camino a Las Mesas, a un costado de la cortina del bordo, 22 Ago 2001,
G pineal 1128 (EB), 1129 ee Loaner de be sabimaads Oponguio, 5 Jul 1990, H. Diaz 6205
(CHAPA, EBUM, IEB, MEXU, Puerto de Los Copales, +8 km de Morelia sobre la
carretera a wn Cumbres, 22 es 2001, G. Ocampo 1124 (EB); alrededores de San José de las Torres, 14
Sep 1991, E. Garcia y E. Pérez 3930 EB); San José Coapa, 4 Oct 1989, J.M. Escobedo 1845 (IEB). Municipio
de Purépero: +3 km de Villa Mendoza, camino a San Antonio Carupo, 2 Sep 1999, E. Carranza et al.
5777 (EB). Municipio de Puruandiro: 9 km al SE de Puruandiro, carretera a Cuitzeo, 22 Jun 1985, H.
Diaz 1162 (EB). Municipio de Tlazazalca: 2 km al W de La Yerbabuena, 6 Jul 1990, E. Pérez y E. Garcia
ul 2001, G.
(IEB),; bordo en la desviacion a La Yerbabuena, sobre la carretera a Churintzio, 7.
Ocampo y A. Medellin 1053 (EB)
Portulaca rzedowskiana se ha encontrado en lugares con matorral subtropical
perturbado o con pastizal, en terrenos planos o con poca inclinacion, entre los
1800 y 2200 m de altitud (5920-7230 ft). Suabundancia local oscila entre escasa
OCAMPO ACOSTA, ESPECIE NUEVA DE PORTULACA 489
“RR
3em MES
sf
era
Ra,
ne
PA Sees
{sages ‘
os ee
3
“ead Sd
ee a af
ae
ot
ae
a3
‘F.
i
(i
Fic. 1. Portul dowski A.A t I de la planta. B. Capsula. C. Semilla
y francamente abundante, por lo que se puede considerar que carece de
problemas de supervivencia. Por el momento, tinicamente se conoce de la parte
norte de Michoacan, pero es de esperarse que esta especie se encuentre en zonas
colindantes de los estados de Jalisco y Guanajuato.
Las especies de la subseccion Conocarpae D. Legrand, muestran capsulas
] ] at ee |
con dehiscencia en sutercio inferior, el opére é
(raramente casi hemiesférico) y sus hojas son rollizas a aplanadas, dispuestas
en grupos de 6a |4 en las inflorescencias (Legrand 1962). P rzedowskiana tiene
mayor afinidad con P. mexicana P. Wilson, P minensis D. Legrand, P perennis R.
E. Fr. y con P. wedermannii Poelln., de las cuales se diferencia por una serie de
combinacion de caracteres (Cuadro 1), siendo los principales el tamaho mas
Cuapro 1. Comparacidn de Portulaca rzedowskiana con otras especies de la subseccion Conocarpae.
P. rzedowskiana
P. mexicana
P. minensis
P. perennis
P. werdermannii
Sépalos
Pétalos
Pétalos
Ralz
Tallo
Densidad de
pelos axilares
Forma
Tamano
Color
Tamano
Enarosada, suberosa
4-26 cm de largo
Regularmente
abundantes
Aplanadas a subrollizas,
lanceoladas, algunas
veces
|
(3-)4.5-6.5 mm de largo
por 4-5.5(-6.5) mm de
ancho
Rosa a morado
(5.5-)6.5-11 mm de
largo
Klana n
MIMNY 1 Pe
oblanceoladas
4-20 mm de largo por
1-3(-4.5) mm de ancho
Engrosada, no
suberosa
10 cm de largo
Regularmente
abundantes
Rollizas, lineales a
sublineales
4-10 mm de largo
por 1.5-2 mm
de ancho
4.5-5 mm de largo
por 1.5-2 mm de
ancho
Amarillo
45-6 mm de largo
2-6 cm de largo
Muy
abundantes
Aplanadas,
bh] lin
na
QUDIOTIQOT CHIL
oblongo-lanceoladas
6-12 mm de largo por
2-3 mm de ancho
7.5-8 mm de largo
por 6 mm de ancho
Pivotante, suberosa
2-9 cm de largo
Regularmente
abundantes
Rollizas,'
oblongas a sublineales
3-8 mm de largo por
1-2 mm de ancho
4-6 mm de largo por
4-6 mm de ancho
Purpureo, rojo-
anaranjado o amarillo
4-9 mm de largo
Suberosa
13 cm de largo
Muy abundantes
Aplanadas,
oblonga a ovado-
oblonga
10 mm de largo por
3 mm de ancho
10.5 mm de largo por
8-9 mm de ancho
Morado
15 mm de largo
06
(2)02@ VaIS/DYO'LIYA
P.rzedowskiana P. mexicana P. minensis P. perennis P. werdermannii
No. de estambres 22-44 14-21 25 8-32 >100
Tamano 4.5-6(-8) mm de largo ——-1.5-3 mm de largo 3-3.5 mm de largo 2-4mm De hasta 6.5 mm
Estilo de largo
No.deramas (3-)4-5(-6) 4-5 5-6 4-7 7
del estilo
Tamano 3.5-6 mm de largo 3-4.5 mm de largo 3-4 mm de largo (2-)3-4.5 mm de largo 3-4 mm de largo
Capsula Forma del Campanulado Campanulado Campanulado a Campanulado a Hemiesférico
Opérculo subtubuloso hemiesférico
Color Negro-azulado Negro Negro Negro anegro-rojizo— Gris
Semillas Tamano 0.6-0.7(-0.8) mm 0.5 mm de largo 0.8-0.9 mm 5-0.6 mm 0.7 mm
de largo de largo de largo de largo
Relieve Estelulado Diminutamente Tubérculos Granulado, Diminutamente
tuberculado subestelulados en subrugoso a tuberculado
su base diminutamente
tuberculado
Distribucion Parte N del Desde el centro de Brasil Desde el N de Brasil
estado de México al N de Argentina y Bolivia
Michoacan Colombia a Peru
VIVTNLUOd 30 VAINN 3193dS3 ‘VLSODV OdW¥I0
Lov
492 BRIT.ORG/SIDA 20(2)
grande que pueden alcanzar sus tallos, asi como las mayores dimensiones de
sus hojas y de sus capsulas (P. rzedowskiana presenta las capsulas mas grandes
de la subseccién (D. Legrand, op. cit.), las cuales se conocian de hasta 5 mm de
largo), asi como por el color negro-azulado de sus semillas. En particular, la
nueva entidad se diferencia de P mexicana y P. perennis por sus hojas aplanadas,
por sus pétalos de hasta 1.1 cm de largo, por presentar mayor numero de
estambres, asi como por tener mayores dimensiones en el tamano del estilo y el
tamano de la semilla; de P minensis y P wedermannii, especies de distribucion
sudamericana, se diferencia principalmente por la abundancia regular de la
pilosidad axilar y por el menor numero en las ramas del estilo.
Martin de Sessé y José Mariano Mocino, en suobra Flora Mexicana, registran
a Portulaca meridiana Lf. (p. 132, 1891-1897) de las cercanias de Puruandiro
(Michoacan), de donde también se ha colectado P. rzedowskiana. En la
actualidad, se considera que P. meridiana Lf. es sinonimo de P. quadrifida L.,
especie distribuida en Africa, Asia, Oceania y las Antillas. Por desgracia, de la
excursion efectuada por los autores mencionados, no se conocen ejemplares en
el herbario del Real Jardin Botanico de Madrid (MA) (Mc Vaugh, p. 446, 2000)
ni dibujos en la Coleccion Torner de la expedicion de Sessé y Mocino (Hunt In-
stitute for Botanical Documentation, Pittsburgh, E.U.A.) asociados con la especie
aludida. Al revisar la descripcion que Sessé y Mocino hacen del material
observado, ésta parece coincidir con las caracteristicas de P rzedowskiana. La
descripcion de P. meridiana que se encuentra en la Flora Mexicana se transcribe
—
a continuacion:
Portulaca foliis lineari-subulatis, carnosis, axillis pilosis; floribus
polyandris, terminalibus, sessilibus. Fl. Mex.
Portulaca foliis ellipticis, carnosis, planis, articulis pilosis, floribus
sessilibus, terminalibus. Herba dodrantalis, caule tereti, glabro, succulento. Fo-
lia alterna, lineari-subulata, carnosa, sessilia, glaberrima, pilis fasciculatis,
utrinque axillaribus ornata. Flores rubro-purpurei, polyandri, terminales, in
capitulum sessiles, verticillum foliorum suffulti. Capsulae ovatae, circumcisae.
Habitat prope Oppidium Puruandirum. Floret lulio.
Portulaca meridiana es una planta anual, con hojas opuestas, aplanadas,
con flores rodeadas por un verticilo de 4 hojas, con pétalos amarillos y con el
opérculo de la capsula de forma campanulado-tubulosa. La disposicion opuesta
de las hojas, entre otras caracteristicas, la ubican en el subgénero Enantiophylla
D. Legrand, por lo que la asignacion del nombre de P. meridiana que hicieron
Sessé y Mocino a los ejemplares que observaron cerca de Puruandiro es
incorrecta. La descripcion en Flora Mexicana concuerda con las caracteristicas
de la nueva entidad, por lo que muy probablemente se trate de la misma unidad
—
taxonomica.
Etimologia.—El epiteto especifico esta dedicado a la Maestra Graciela
OCAMPO ACOSTA, ESPECIE NUEVA DE PORTULACA 493
Calderon de Rzedowski y al Dr. Jerzy Rzedowski Rotter por su incansable e in-
valuable labor en pro del conocimiento botanico de México.
AGRADECIMIENTOS
Deseo expresar mi agradecimiento a Heike Betz (Field Museum) por su
apreciable ayuda en la consecucion de bibliografia. De igual forma, deseo
agradecer a Patricia Mayoral, a Emmanuel Pérez, a Sergio Zamudio (todos ellos
del Instituto de Ecologia, A.C., Centro Regional del Bajio), asi como a James F.
Matthews (University of North Carolina at Charlotte) por la revision critica del
manuscrito.
REFERENCIAS
Lecranp, D. 1962.Las especies americanas de Portulaca. Anales Mus. Hist. Nat. Montevideo
24 ser. 7(3):1-147.
Sesse, M. y J. M. Mocifio. 1891-1897. Flora Mexicana. La Naturaleza, 2? ser., 2 (apéndice):
1-263.
McVauch, R. 2000. Botanical results of the Sessé & Mocifo expedition (1787-1803). VILA
guide to relevant scientific names of plants. Hunt Institute for Botanical Documenta-
tion. Pittsburgh.
494 BRIT.ORG/SIDA 20(2)
Book NoTICc!
Garrett E. Crow and C. Barre Hettquist. 2000. Aquatic and Wetland Plants of
Northeastern North America: A Revised and Enlarged Edition of Norman
C. Fassett’s A Manual of Aquatic Plants, Volume 1. Pteridophytes, Gym-
nosperms, and Angiosperms: Dicotyledons. (ISBN 0-299-16330-X, hbk.).
$90.00, +80 pp, 338 line illus., 1 map, 81/2" « 11".
Garrett E. Crow and C. Barre He_tQuist. 2000. Aquatic and Wetland Plants of
Northeastern North America: A Revised and Enlarged Edition of Norman
—
C. Fassett’s A Manual of Aquatic Plants, Volume 2. Angiosperms: Mono-
cotyledons. (ISBN 0-299-16280-X, hbk.). $90.00, 400 pp, 268 line illus., |
map,ol/2 x LI.
The University of Wisconsin Press, 1930 Monroe Street, 3rd Floor, Madi-
son, W153711-2059, US.A. (Orders: Customer Service Department, The Uni-
versity of Wisconsin Press, c/o Chicago Distribution Center, 11030 S. Lan-
gley Ave., Chicago, IL 60628, U.S.A.; 773-568-1550; fax 800-621-8476,
(fa°000-2235).
This 2-volume set updates the original 1940 Fassett manual and its successor, the 1957 revision by
Eugene Ogden—it isa guide to identification of native or naturalized vascular plants growing in “the
region from Newfoundland west along the 50th parallel to southeastern Manitoba and Minnesota,
south to Virginia and Missouri. Thus, on the southern edge, our range abuts the range covered by Drs.
Godfrey and Wooten in their “Aquatic and Wetland Plants of the Southeastern United States.” aie
a
treats “Pteridophytes, Gymnosperms, and Angiosperms: Dicotyledons”, Volume 2 treats “An-
giosperms: Monocotyledons.” Species of “salt marshes and tidal waters” and those “which typically
occur in saturated peat” are included but “not terrestrial plants which may occur typically on flood
slains, in woodland springs, or along vern: al woodland or seasonal alpine brooks.” The keys, whie h
g g l
“utilize vegetative features ¢ ape ss ee ible,” “treat a total of 1139 species C118
1g
295 genera in LO9 families.” “The 606 aes include figures of 1087 taxa, with 92 percent ee inet tax
fully or partly illustrated, [and]... ref tl es are included in the keys.” Habitat ons 1
figu
tion and geographical distributions “have oa a n Naeeas rom Bae published literature ... and
from herbarium specimens documenting locat and from our own field
experience.” The nicely bound, large-size, easy-to-use format of the volumes, spacious plates with
black-and-white line drawings, effective keys, and wide taxonomic coverage promise that this con-
tribution will be useful in many contexts.—Guy L. Nesom, Botanical Research Institute of Texas, 509
Pecan Street, Fort Worth, TX, 76102-4060, U.S.A.
SIDA 20(2): 494. 2002
AGAVE OVATIFOLIA (AGAVACEAE), UNA NUEVA ESPECIE
DE MAGUEY DEL NORESTE DE MEXICO
Greg Starr José A. Villarreal Q.
Starr Nursery Depart to de Botdni
3340 W. Ruthann Rad. Universidad Auténoma Agraria “Antoni jo Narro"
Tucson, AZ 85745, U.S.A. Buenavista, Saltillo, Coahuila 25315, Méx
gstarrsprint@earthlink.net javillarrealo0O@hotmail.com
RESUMEN
Una especie de maguey, Agave ovatifolia Starr & Villarreal de la Sierra de Lampazos en el norte de
Nuevo Le6n es propuesta como nueva. Es ubicada en la seccion Parryanae de subgénero Agave y se
discute su relacion con A. havardiana Trel. y A.parrasana Berger. Se presenta una ilustracion de la
planta y una clave parcial y modificada de la seccion Parryanae (Gentry 1982:526).
ABSTRACT
A new species, Agave ovatifolia Starr & Villarreal from the Sierra de Lampazos in northern Nuevo
Leon is described and illustrated. It belongs in the section Parryanae of the subgenus Agave, and its
relationship to A. havardiana Trel. and A. pare sana Berger is discussed. A partial, modified key to
10@7).-&9
section Par ryande (Gentt y lyoz 6) is prov ided.
Key worps: Agavaceae, Agave, Flora of Mexico
La colecta de material proveniente del norte del estado de Nuevo Leon
proporciona informacion de una nueva especie de Agave.
Agave ovatifolia Starr & Villarreal, sp. nov. (Figs. 1 y 2). Tivo: MEXICO. NUEVO LEON:
Se de Bustamente, Sierra de Lampazos, rancho Minas Viejas, 26°39’ N, 100°24' W,
ue de Quercus canbyi, Quercus lace yi, Juniperus, Crataegus, Ribes, Dane, Vitis, entre
otras, L1OO-1200 m, aa oo. JA. Villarreal 9180, M.A. Carranza yJ.A. Encina (Hol
MEXU, tsoTipos: ANSM, T
| lti acaulis. rosula | | s, compactus, 80-120 cm diametro, 60-90 cm alta;
E
foliis 40-50 in 8-10 stratis, as a5: 45 cm longis, 20-24 cm latus, griseo-azureus, ae
acuminatus, spina terminalis 2 cm longa, margine 15-19 paribus dentes separatus 20-25 :
inflorescentia 3.5-4.0 m alta, en en 13-16 rami floribus 30-50 cm longis; cee
ellipticus-triangularis, inferiora 20-25 cm longis, 8-10 cm latis, superiora 12-15 cm longis, 7-9 cm
latis; pedicelli 4-8 mm longi; flores 67-74 mm longum, flavo-virens, ovarium cylindricus, 30-35 mm
longis, perianthii collo 7 mm longo, peranthii tubo 15-20 mm longo, 15 mm lato, tepala 20 mm longa,
5-7 mm lata, fila 50-60 mm longa insertus basis tepalas; capsula cylindricus, 50-60 mm longis, 15-
20 mm latis; semina 5-6 mm longa, 3-4 mm lata.
Plantas multianuales; rosetas hemisféricas, algo compactas, 80-120 cm
diametro, 60-90 cm alto; hojas arregladas en 8-10 hileras, 40-50 hojas por planta,
elipticas, ligeramente concavas a casi planas, 35-45 cm de largo, 20-24 cm de
ancho en la parte media, de color glauco-grisaceo, ocasionalmente con |-2
costillas dorsales, base ensanchada, apice acuminado con borde de color café,
SIDA 20(2): 495 — 499, 2002
496 BRIT.ORG/SIDA 20(2)
Fic. 1.Agave ovatifolia.A hoja B. bractea. C. flor, | itudinal. E. qlandul lext del tépalo. F. capsula
espina terminal de cerca de 2 cm de largo, margen recto, con 15-19 pares de
dientes ligeramente curvados, separados 20-25 mm, mas proximos cerca de la
base; inflorescencia 3.5-4 m alto, forma eliptica a largamente ovoide, escapo
floral 9-11 cm ancho en la base, 6-8 cm en la primer ramificacion, raquis algo
sigmoideo, 13-16 ramillas florales, de 30-50 cm de largo, ubicadas en la mitad
superior del escapo; bracteas eliptico-triangular, esparcidas, las basales 20-25
cm largo, 8-10 cm anchoen la base, las superiores 12-15 cm largo, 7-9 cm ancho
en la base, acuminadas con espina terminal, enteras a dentadas, delgadas
(papiraceas), glaucas, pronto se deshidratan y cambian a color café-claro, la
mayoria erectas; flores en 5-6 umbelas densas, 67-74 mm largo, verde
amarillentas; pedicelos 4-8 mm largo, ovario cilindrico, 30-35 mm largo, cuello
del perianto 7 mm largo, tubo del perianto 15-20 mm largo, 15 mm anchoen la
parte media, tépalos largamente triangulares, 15-17 mm largo, 5-7 mm ancho
STARR AND VILLARREAL, UNA NUEVA ESPECIE DE AGAVE 497
en la base, los tres mas externos con evidentes glandulas apicales, filamentos
50-60 mm largo, ligeramente aplanados insertos en la base de los tépalos, verde
amarillento, anteras 22-25 mm largo, verdes, estilo 8-9 cm largo; capsulas
cilindricas 50-60 mm largo, 15-20 mm ancho, color café oscuro; semillas
lacrimiformes, 5-6 mm largo, 3-4 mm ancho, color negro brillante.
Crece en laderas y mesetas rocosas entre los 1100-1500 m, tanto en bosque
de encino como matorral de Agave lechuguilla y arbustos espinosos. Su
distribucion es restringida en el area antes mencionada con la posibilidad de
que se encuentre también en otras sierras aledanas. Las poblaciones son
esparcidas, donde nose observan hijuelos ni la formacion de colonias. La planta
es reportada como buena forrajera para el ganado y las poblaciones son
menguadas por jabalies y otra fauna silvestre. Sdlo se observaron plantas
maduras y en floracion en laderas con fuerte pendiente, donde escapan a la
depredacion. Localmente se le conoce como “noga” y se le diferencia de las
plantas que carecen de hojas con costillas dorsales, las cuales son denominadas
“maguey noga.”
La especie se ubica en la seccion Parryanae del trabajo de Gentry (1982) y
al parecer esta relacionada morfoldgicamente con A. havardiana Trel. y A.
parrasana Berger, de las cuales se diferencia por sus hojas proporcionalmente
498 BRIT.ORG/SIDA 20(2)
mas anchas, de color gris-azuloso y flores con tamano intermedio, como se
muestra en la siguiente clave modificada. Aunque las dimensiones de las hoja
en A. havardiana y A. ovatifolia se sobreponen, la relacion entre el largo y ancho
es diferente, la de A. havardianaes de 2.0-2.8, mientras que la de A. ovatifoliaes
de 1.5-L8. La relacion entre largo y ancho de la hoja en A. parrasana es de 2.0-
2.5, pero con hojas consistentemente mas cortas. Agave havardiana se distribuye
en el suroeste de Texas y regiones adyacentes de Chihuahua y Coahuila, A
parrasana en sierras del centro y sur de Coahuila, mientras que A. ovatifolia en
la Sierra de Lampazos en el norte de Nuevo Leon.
Agave ovatifolia es ubicado en la seccion Parryanae, y la clave de Gentry
(1982:526) puede ser modificada como sigue:
2. Rosettes globose, suckering copiously with long rhizomes forming large, spread-
ing clones; panicles deep with 20-40 umbels; flowers 60-80 mm long 3
2. Rosettes not globose, more openly flat-topped, with few or no suckers; panicles
rather wide and open with 10-20 large umbels; flowers 67-90 mm lon 7
Ng
7. Leaves larger, 30-70 X 15-25 cm, long acuminate; teeth oe along middle
and lower leaf tie tepals about equaling to slightly longer than tube. Big
Bend Region and adjacent Mexico 8
7. Leaves smaller, 20-30 x os 12 cm, the apex abruptly acute; teeth not reflexed
along margins; tepals about twice as long as tube. Sierras of southern Coahuila.
Agave parrasana p.537
8. Leaves relatively long and narrow, 30-70 xX 15-25 cm, the length 2 or more
times the width, ovate-acuminate to long-acuminate, leaf tip acute, glaucous
gray to light green, spine stouter, 3—5(-10) cm long; flowers 68-88 mm long,
tepals 18-24 mm long; Big Bend, east-central Chihuahua and northwestern
oahuila, Mexico Agave havardiana p. 531
Leaves relatively short and broad, 35-45 x 20-24 cm, the length less than 2
8.
times the width, elliptic to ovate-elliptic, leaf tip obtuse, ee to grayish
sky blue, spine 2 cm long; flowers 67-74 mm long, tepals 15-17 mm long,
Nuevo Leon, Mexico Agave ovatifolia
Al parecer A. ovatifolia es la misma planta que Mrs. Anna B. Nickels colect6 y
distribuy6 como A. noah Nickels a finales del siglo antepasado. La especie de
Mrs. Nickels no fue descrita y tampoco fue apropiadamente publicada, a pesar
que Trelease (1911) la enlista como sinénimo de A. wislizeni Engelm. La
taxonomia de A. wislizeni ha sido ampliamente discutida por Gentry (1975) y
Ullrich (1992). El concepto de Trelease (1911) para A. wislizeni acomoda bien
con A. parrdsana pero no para A. ovatifolia. Lynn Lowery, renombrado
horticultor texano, colecté esta especie de Sierra de Lampazos, a principios de
los 1990's. El epiteto especifico hace referencia a las hojas relativamente anchas
en relacion a su largo.
AGRADECIMIENTOS
Nuestro agradecimiento a el senor Pedro Elizalde Cantu, por permitirnos
colectar material botanico en su rancho. A Ron Gass, Miguel A. Carranza y Juan
STARR AND VILLARREAL, UNA NUEVA ESPECIE DE AGAVE 499
A. Encina por acompanarnos en las visitas a campo. La ilustracion es obra de
Miguel A. Carranza.
REFERENCIAS
Gentry, H.S. 1975. The case of Agave scabra and Agave wislizenii. Cact. Succ. J.(Los Angeles)
47:102-104.
Gentry, H.S. 1982. Agaves of Continental North America. University of Arizona Press, Tuc-
son. 670 p.
Trevease, W. 1911 [1912]. Revision of the Agaves of the group Applanatae. Ann. Rep. Mis-
souri Bot. Gard. 22:85-97.
UtiricH, B. 1992. On the history of Agave asperrima and A. scabra (Agavaceae) as well as
some taxa of the Parryanae. Sida 15:241-262.
500 BRIT.ORG/SIDA 20(2)
Book NOTICE
LAURENCE J. Dorr, BASIL STERGIOS, ALAN R. SmitH, and Nipia L. Curio A. Chief Edi-
tor: LAURENCE J. Dorr. 2000. Catalogue of the Vascular Plants of Guaracamal
National Park, Portuguesa and Trujillo States, Venezuela. (ISSN 0097-1618,
pbk). Contributions form the United States National Herbarium, Volume
20:1-55. Department of Botany, National Museum of Natural History,
Smithsonian Institution, Washington, DC 20560-0166, U.S.A. Price not given,
155 pp, 3 figures, 7" « 10"
k, which lies southwest of the city of Bocon6, preserves lower montane
“Guaracamal National Par
forest, cloud forest, and paramo in the northeastern-most portion of the venezuelan Andes.” It covers
21.5 km2 and ranges between 1600-3100 meters in elevation. “The specimens examined for this cata-
logue are mostly our own collections. These have been deposited in NY, PORT, UC, and US, and tl
duplicates have been distributed widely.”
The catalogue treats a total of 147 families, 517 genera, and 1227 species—236 species Gn 62
genera) of ferns and fern allies, 237 species (in 92 genera) of monocots, 752 species (in 361 genera) of
dicots, and 2 species of pines (in 2 genera). The ten most important families in Guaracamal National
Park in terms of the number of species are (in order) Orchidaceae-115, Asteraceae-102, Rubiaceae-
Piperaceae-47, Dryopteridaceae-46, Melastomataceae-44, Solanaceae-35, Poaceae-33,
Grarnmnitidacede 0. and Lauraceae-27. Thirty-three vascular plant taxa have been described (as
new to science) from material collected in the park. A brief statement of collection localities,
elevational range, exsiccatae, common names and uses, endemic and near-endemic status, and ad-
ventive and cultivated species is given for each species, where applicable. Relevant taxonomic refer-
ences are provided at the end of each family treatment.—Guy L. Nesom, Botanical Research Institute
of Texas, 509 Pecan Street, Fort Worth, TX, 76102-4060, U.S.A.
JOURNAL Notict
Acta Botanica Hungarica. (ISSN 0236-60495). Akadémiai Kiad6, Budapest, H-1519
Budapest, PO. Box 245, HUNGARY. (Subscriptions: 36-1-464-8221, email:
kiss.s@akkrt.hu). $192.00, + epee Gly2 3 O72"
CONTEN NTS FOR ae ze cae " putes meer 7. al fungi Seoul ales) in see) pt. Il. An ecological
I land; 2) Ultrastructural stud-
view
ies on Vicia faba and its eathowen Botrytis fabae in ae to lithium ¢ hilor ide; 3) Taxonomy and
morphology of uncultivated hemp (Cannabis sativa L.) was weed in Hungary; 4) Los géneros
Ceuthorcarpus Aiello y - lice is (Rubiaceae 3) Salt tolerance variation in some common
gradient in the eastern Niger delta area of
trees, 6) A spectrum of ]
Nigeria; 7) Top-forests ‘Geratro nigri-Fraxine bum orni) of the Bakony iesouaay 8) The aida
characteristics of Fumana procumbens (Dunal) Gren. et Godron under different clim ati iti
9) Detection of phenoloids in some H iuear a Tata ane Centaurea at ee 10) Grbloereh ic acid
icia
effects on protein pattern, hydrolytic
faba in sea water, 11) 1 Disceibution and phy tosociological ‘ana two ecu ed plant species in
an open sand grassland area in the Great Hungarian Plain; 12) Cytotaxonomic studies on some Vicia
L. species growing in the eastern Mediterranean and southern Aegean regions II.
SIDA 20(2): 500. 2002
A NEW SPECIES OF LITHOSPERMUM (BORAGINACEAE)
FROM NUEVO LEON, MEXICO
B.L. Turner
Plant Resources Center
University of Texas
Austin, TX 78713, U.S.A.
ABSTRACT
ithospermum hintoniorum is newly described and illustrated. It reportedly occurs in marshy areas
of oak woodlands in southern Nuevo Leon, Mexico at elevations of about 1500 meters. It is seemingly
most closely related to the widespread L. calycosum but is distingished by a number of characters,
including both vegetative and floral features.
RESUMEN
Se describe ys se il I hi i Se ha | lizad , | ee 2 pee |
enel sur de Nuevo Leon, México en elevaciones de unos 1500 metros. Parece ser que su pariente mas
cercanoes el muy frecuente L. calycosum del que se diferencia por varios caracteres, tanto vegetativos
como florales
Routine identification of Hinton collections from northeastern Mexico has re-
vealed the following novelity:
Lithospermum hintoniorum B.L. Turner, sp. nov. (Fig. 1). Type: MEXICO. NUEVO LEON:
Mpio. Aramberri, San Juan de Aviles, “Marsh in oak woods,” ca. 1480 m, 13 May 1992, Hinton
et al. 21950 (HOLOTYPE: TEX; ISOTYPES: personal collections of the collectors).
Simile Lithospermo calycosa, lobos corollarum noncrenulatos, nuculasque majoris (ca. 3 mm altas
vice 2mm) a griseas.
Perennial herbs 12-17 cm high, the stems seemingly recumbent and arising from
woody roots which exude a violet-colored dye. Leaves basal at first, but soon
developing 20 or more overlapping cauline leaves, these gradually reduced up-
wards and grading into the floral bracts; blades linear-lanceolate to lanceolate,
essentially nonvenose, except for the midribs, the surfaces appressed-strigose,
the undersurfaces having finer hairs, the upper surfaces with coarser hairs hav-
ing attractive flattened basal cells 0.15-0.30 mm across. Inflorescence decid-
edly scorpioid, having numerous bracteate pale yellow flowers. Sepals 5, linear,
mostly 7-9 mm long, moderately pubescent without, glabrous within, weakly
glandular at the throat, markedly nodular-annulate at or near the base; lobes 5,
ovate, 1.4-1.6 mm long, 1.0-1.2 mm wide. Anthers 5, dorsifixed, ca. 1.7 mm long,
inserted in the throat but not exceeding the lobes. Styles ca. 11 mm long, gla-
brous, minutely bifid at apex. Nutlets 2-4, glossy-gray, not at all ornamented,
ca. 2mm high, 1.5 mm wide, basally attachment ca. 2 mm across.
The relationship of this taxon among Mexican species of Lithospermum is
SIDA 20(2): 501 - 503. 2002
502 BRIT.ORG/SIDA 20(2)
Fic. 1. Lithospermum hintoniorum. Freshly dug garden
J t vt 7
TURNER, A NEW SPECIES OF LITHOSPERMUM FROM MEXICO 503
moot. Assuming the plants to be consistently homostylous with weakly devel-
oped faucal appendages, it will begrudgingly key to L. calycosum (Macbride)
LM. Johnst. in Johnston’s (1952) treatment of Lithospermum, to which it is com-
pared in the above diagnosis. It differs in having much larger, broader, leaves,
noncrenulate corolla lobes, and larger (ca. 3 mm high vs 2 mm), glossy-gray
nutlets.
It isa pleasure to name this species for the remarkable Hinton family, who
first collected the taxon. When first sent to me for identification I recognized it
as possibly new but lacked fruiting material for descriptive purposes. George
Hinton (son of the living Jaime Hinton, and grandson of the legendary plant
collector, George B. Hinton) kindly sent me mature seeds nine years after its
initial collection, these obtained from garden grown plants collected at the type
locality. lam grateful to him for such diligence.
ACKNOWLEDGMENTS
lam grateful to my wife Gayle Turner for the Latin diagnosis and to Diane M.
Ferguson and James S. Miller for helpful comments.
REFERENCE
JOHNSTON, ILM. 1952. Studies in the Boraginaceae, XXIll. Survey of the genus Lithospermum.
J. Arnold Arbor. 33:299-363.
BRIT.ORG/SIDA 20(2)
Book NoTICcE
STANWYN G. SHETLER and SyiviA STONE Orit. 2000. Annotated Checklist of the Vas-
cular Plants of the Washington - Baltimore Area. Part I: Ferns, Fern Al-
lies, Gymnosperms, and Dicotyledons. (no ISSN or ISBN). Department of
Botany, National Museum of Natural History, Smithsonian Institution,
Washington, DC 20560-0166, U.S.A. Price not given, 186 pp [duplicated], 1
figure, 81/2" x 11",
STANWYN G. SHETLER and SYIVIA STONE Ori. 2000. Annotated Checklist of the Vas-
cular Plants of the Washington - Baltimore Area. Part Il: Monocotyledons.
(no ISSN or ISBN). Department of Botany, National Museum of Natural
History, Smithsonian Institution, Washington, DC 20560-0166, U.S.A. 95
pp [duplicated], 8 1/2" x 11"
The checklist in this pair of publications treats all native and patel ed species within the “Wash-
ington-Baltimore area,” which represents a radius of about 50 mil he Capitol in Washington,
D.C. and includes the District of Columbia as well as 23 counties ad 5 cities in Maryland and Vir-
ginia. “Species were accepted only on the basis of one or more voucher specimens in the D.C. Her-
barium or a reliable published record” (the District of Columbia herbarium is a subunit of the US
National H ane ium). Sih summarized in Part Il, the documented D.C. flora now includes 1801] native
species and 9 ced species
s He 8 h Vy ts/dcfle /) | | |
related v www.nmnhsiedu/| tany/ projects/ acyl
|
ongoing D.C. Flora Project oad much other ceaiiens information, alate a arene aphy, color
photographs, spring-flowering records, links to useful sites related to the project, and a searchable
Checklist database. The Checklist database returns full specimen data accessed by query to family,
genus, species, collector, locality, state, or county. For example, an otherwise unrestricted query for
Chenopodiaceae returned 148 records * a query for Asteraceae returned 7616 records; a query for
Apiaceae in D.C. returned 183 records.—Guy L. Nesom, Botanical Research Institute of Texas, 509 Pecan
Street, Fort Worth, TX, 76102-4060, U.S.A.
SIDA 20(2): 504. 2002
A NEW SPECIES OF SCHISTOCARPHA (ASTERACEAE:
HELIANTHEAE) FROM NORTHWESTERN OAXACA
AND CLOSELY ADJACENT GUERRERO, MEXICO
B.L. Turner
Plant Resources Center
The University of Texas
Austin, 1X 78713, U.S.A.
ABSTRACT
Schistocarpha calzadana B.L. Turner, sp. nov, is described from south-central Oaxaca (Mpio. San-
tiago Juxtlahuaca) and south-central Guerrero, Mexico. It is closely similar to S. bicolor but can be
distinguished by a number of features including those of the leaf, size and congestion of the
capitulescence, and number and size of ray florets. A re-evaluation of the Mexican taxa has resulted
in the submergence of S. longiligula into S. platyphylla. A revised key to the Mexican species is pro-
vided, along with maps showing their distribution.
Key Worps: Schistocarpha Asteraceae, Mexico, Guerrero, Oaxaca
RESUMEN
Se describe Schistocarpha calzadana B.L. Turner, sp. nov., del centro-sur de Oaxaca ae Santiago
Juxtlahuaca) y centro-sur de Guerrero, México. Fs mu y semejantea S. bicolor y nuede diferenciarse
L
por varios caracteres de la hoja, tamano y aglomeracion de la capitulescencia, asi como el numero y
tamano de las ligulas. Una re-evaluacion de los taxa mexicanos ha dado como resultado la inclusion
de S. longiligulaen S. platyphylla. Se ofrece una clave revisada par alas especies mexicanas, junto con
mapas de su distribucion.
Schistocarpha is a mostly Mexican genus of ten closely related species (sensu
Turner 1986 and the present paper). These have been variously interpreted (e.g,
Rydberg 1927; Robinson 1979; Turner 1996), the most recent appraisal of the
Mexican taxa, albeit for the state of Chiapas, being that of Strother (1999). With
the exception of S. eupatorioides (Fenzl) O. Kuntze and S. pedicellata Klatt,
Strother placed most of the Mexican species recognized by previous workers
under the fabric of S. bicolor Less. Because of Strother’s treatment I have looked
again at the Mexican taxa and have concluded that one can reasonably sink the
widespread, highly variable S. longiligula Rydb. into the earlier S. platyphylla
Greenm., but the remainder of Strother’s dispositions under a widespread S. bi-
color is moot. Indeed, having gone over this taxonomic landscape anew | would
still recognize six species as occurring in Mexico, including S. liebmannii Klatt,
S. matudinae H. Rob., and the newly described S. calzadana, proposed below.
Schistocarpha calzadana B.L. Turner, sp. nov. (Figs. 1, 2). Type: MEXICO. OAXACA:
Mpio. Santiago Juxtlahuaca, 17 km al SW de Santa Rosa sobre la carretera a San Miguel Cuevas
SIDA 20(2): 505 — 510. 2002
=
BRIT.ORG/SIDA 20(2)
UNIVERSITY OF
EXAS p=
HERBARI
The University of Texas Herbarium (LL, TEX)
Sch staph calzadana
. Turner, sp. nov.
Det: BL. Turner, 20 Jun 2001
| Isoryr: Sehistocarphn |
—_ |
« aland an Bbhitvere |
~ |
ve H
FLORA DE OAXACA
RACEAL
_- Schistocarpha bicolor Less
Fr SANTIAGO JUX CA: Santiago Juxtlahuaca, 17
kim al SW. se Santa Tio a paras la carretera a San Miguel
Cuevas y Manzanares , 17° 13’ 31.9", 98° 00" 24.4 W
Hierbas p perennes de 1.5-2.5 m de alto, Hgulas blancas, corolay
del disco amarillas. Bosque mesdfilo de montaha, 2150 m
r Panwro 5524 28 Feb 1995,
con Ismael Calzada
Tosa « Rutt JL. 10
Herbario Nacional de México (MEXU)
Michig, te University Herbarium (MSC)
east Aust, TO
Food
OM
Bacardi, Méwien,
EATS, MSTA
Fic. 1. Schistocarpha calzadana; isotype (TEX).
TURNER, A NEW SPECIES OF SCHISTOCARPHA FROM MEXICO 507
SCHISTOCARPHA Ni
YZ bicolor ne
o calzadana
» eupatorioides
Fid. 2. Distrit ti £ Mavi td £C-hi }
: Manzanares (17 13 31.9 N, 98 03 24.4 W), 2150 m, 28 Feb 1995, J.L. Panero 5524 (con Ismael
Calzada) (HOLOTYPE: MEXU; ISOTYPES: TEX, UC).
Similis S. bicolorised liis foli latis basi itul ia congesti itulis
é Sos
numerosis, et flosculis radii cum ligulis 1.5-2.5 mm longis (vice 3-6 mm).
Perennial, suffruticose herbs 1.5-2.5 m high. Stems moderately to densely pi-
lose, glabrate with age. Larger leaves 20-30 cm long, 14-16 cm wide; petioles 6-
8 cm long, winged throughout but much tapered below into a distinct petiole;
blades broadly subcordate, markedly serrate, their apices rounded or obtuse.
Primary capitulescences 14-20 cm across, ca. 10 cm high; ultimate peduncles
mostly 3-9 mm long, pubescent like the stems. Involucres 4-5 mm high and
about as wide, having imbricate 3-4 seriate bracts, the inner bracts scarious
with mostly rounded ciliate apices. Receptacle plane or somewhat convex,
paleate, the pales shorter than the subtended florets. Ray florets ca. 13 ina single
series; tubes ca. 3mm long; ligules white, 1.5-2.5(-3.0) mm long, ca.0.8 mm wide,
2-4 nervate. Disk florets 30-40; corollas ca. 4 mm long, tubes ca. 2 mm long,
glabrous or nearly so, the lobes 5, ca. 0.5 mm long, hispidulous without. Achenes
of ray and disk similar, ca. 1.5 mm long, glabrous; pappus of numerous white
bristles 3-4 mm long.
508 BRIT.ORG/SIDA 20(2)
—
Additional collections examined: MEXICO. GUERRERO: “Along road between El Paraiso and Puerto
del Gallo, 6.0-8.7 mi NE of El Paraiso. Moist tropical evergreen forest,” 3500 ft, 9 Mar 1987, Daniel &
Bartholomew 4936 (CAS, TEX). OAXACA: Mpio. Santiago Juxtlahuaca, Km 14 de San Miguel Cuavas a
om Manzal, desviacion en Santa Rosa (17 13 N, 98 03 W), 2300 m, 18 Feb 1995, Calzada 19761C (MEXU,
EX); 0.4 km del poblado El Manzanal, carretera a Infiernillo (17 13 N, 98 03 W), 2185 m, 8 Mar 1996,
an ada 20784 (MEXU, TEX).
In the account of Schistocarpha by Robinson (1979), the present taxon will key
to S. matudae, a species of southern Chiapas. In my treatment of the genus
(Turner 1986), it will key to or near S. ligbmannii, to which | originally referred
the collections from Guerrero. With the newly collected Oaxacan material in
hand, it seems likely that the several sheets relate better to S. bicolor, the latter
from the Gulf Coast sierras, the former from the Pacific sierras. Schistocarpha
calzadana differs from its more eastern cohort in characters as noted above.
It should be noted that Strother'’s (1999) treatment of Schistocarpha for the
Asteraceae of Chiapas would reduce er all of the Mexican taxa of that ge-
nus (other than S.e upatot ioidesand § S. pe dicellata)tobuta single widespr ead S.
bicolor, including the present novelty. His comments to justify such reduction,
“| have seen intermediate specimens that link all of the named extremes.” seem
exceedingly strained to me, especially since his examination of a wide range of
collections at LL-TEX resulted in no annotations to that effect. | had no great
difficulty in placing ‘segregate’ species names on the holdings at CAS, LL-TEX,
and UC. most of which served as the basis for Strother’s remarks. | have, how-
ever, accepted the likelihood that S. longiligula and S. platyphlla are synony-
mous, the two being largely sympatric and separated by relatively trivial fea-
tures (ligule length and floret number). | also freely admit that S. matudae, S.
licbmannii,and S.calzadana might be swept under S. platyphylla, as interpreted
here (the earliest name for the complex then being S. liebmannii), but such ti-
dying should take place only after more detailed field studies of the popula-
tions concerned. Regardless, S. bicolor (with its markedly winged petioles) does
not extend so far south in Mexico so as to include the state of Chiapas.
Etymology.—The species is named for J.T. Calzada, a Mexican botanist who
was the second worker to garner the taxon and who participated in all subse-
quent collections.
linclude below a revised key to the Mexican species of Schistocarpha, which
includes the present novelty. Figures two and three show the distributions of
these taxa. The revised key and up-to-date distribution maps should prove help-
ful to future workers interested in the group.
KEY TO MEXICAN TAXA OF SCHISTOCARPHA
. Pistillate florets (21—-)30—-80 in 2-4 series S. eupatorioides (Fig 2)
sais florets (3-)5-21 in a single series.
etioles conspicuously winged throughout, often expanding at the base and
ites across the node; northeastern Mexico S. bicolor (Fig 2)
TURNER, A NEW SPECIES OF SCHISTOCARPHA FROM MEXICO
SCHISTOCARPHA
4 liebmann
A matudae
2. Petioles not broadly winged throughout, nearly —_ tapering onto a distinct
a petiole; western and southern Mexic
. Cap
tulescence lax, containing 8-12 heads on ultimate peduncles a 5<Gm
long at maturit
pedicellata (Fig. 4)
3. Capitulescence strict, congested, containing 30 or more heads on fue
peduncles mostly 1.5-2.0(—2.5) cm long or les
4. Stems glabrate or nearly so at maturity; ae surface of blade (except for
veins) glabrous, the upper surface somewhat darker than lower; disk flo-
rets 10-12, i
2, their corollas sparsely pubescent; montane cloud forests of
north-central Oaxaca and adjacent Verac
ruz S. liebmannii (Fig. 3)
4. Plants without the above combination of characters and distribution.
5. Ray florets 5-12; ligules mostly 4- a mm lon . platyphylla (Fig. 4)
5. Ray florets 12-21; ligules mostly 1-4 mm long.
6. Involucral bracts with acute ero eeu Chiapas S. matudae (Fig. 3)
6. Involucral bracts with rounded ap Guerrero and Oaxaca
S.calzadana (Fig. 2)
With the exception of S. calzadana, all of the above-keyed taxa are treated in
more detail by Turner (1986).
509
510 BRIT.ORG/SIDA 20(2)
SCHISTOCARPHA
o pedicellata
@ platyphylla
Fig 4. Distributions of Mexican species of Schistocarpha.
ACKNOWLEDGMENTS
lam grateful to my wife, Gayle Turner, for the Latin diagnosis and to Jose Panero
for calling my attention to the Oaxacan plants concerned. John Strother re-
viewed an early draft of the manuscript, improving considerably its terseness
and readibility. The study is based upon materials from the following herbaria:
CAS, LL-TEX, UC, as follows: Schistocarpha bicolor CAS (22), LL-TEX (39), UC
(7). Schistocarpha liebmannii CAS (4); LL-TEX (10); UC (D). Schistocarpha
matudinae CAS (4); LL-TEX (3); UC (0). Schistocarpha pedicellata CAS (9); LL-
TEX (15); UC (1). Schistocarpha platyphylla CAS (68); LL-TEX (54); UC (8).
REFERENCES
Rosinson, H. 1979. A study of the genus Schistocarpha (Heliantheae: Asteraceae).
Smithsonian Contr. Bot. 42:1-20.
RypserG, P.A. 1927. Schistocarpha, in N. Amer. Fl. 34:303-306.
StrotHer, J.L. 1999. Schistocarpha. In: Flora of Chiapas, Pt. 5: Heliantheae s.l., Calif, Acad. Sci.,
San Francisco.
Turner, B.L. 1986. An underview of the genus Schistocarpha (Asteraceae-Heliantheae).
Phytologia 59: 269-286.
SENECIO SPRIBILLEI (ASTERACEAE: SENECIONEAE),
A NEW SPECIES FROM MONTANA, U.S.A.
William A. Weber
University of Colorado Museum
Boulder, CO 80309, U.S.A.
ABSTRACT
c : ibillei WA
. Weber, a new, putatively narrow endemic species of Asteraceae, is described
froma aunates in the Rocky Mountains of northwestern Montana. It is distinguished from its near-
est relative, Senecio dene Blake, ie foliage, proadey and shorter leaf ion wing-
less petioles, coarser g tati nd minute ray flowers.
RESUMEN
Se describe Senecio spribillei W.A. Weber, una nueva especie encenues de Ba eas ve un nunatak
n las Montanas Rocosas del noroeste de Montana. Se dif Senecio
neowebsteri Blake, por sus hojas glabras, foun folia rmasancha y me corta, Peolos sin alas, dientes
del margen con dientes mas juntos, capitulos racemosos mas pequenos y ligulas diminutas.
Senecio oer W.A. Weber, sp. nov. (Fig. 1). Type: U.S.A. MONTANA. Lincoln Co: Cabi-
et Mountains, E face of Snowshoe Peak, ca. 150 m below the summit, N4893'40" W11540'40";
on eee vegetated al pe er sae with Dryas octopetala and Carex albonigra, 2500 m
alt. 25 Aug 2001, Toby S} Ml. Arvidson 11264 (HOLOTYPE: MO; ISOTYPE: COLO).
1c Oey |
d 2.0 dm alta, tota glabra, caudices brevi-ramosa, folia caulis juvenilis basalibus
pngenetSlatis 0.5 dm longis, folia basalibus caulis floriferis longepetiolatis 0.5 dm longis induviis,
=e caulina 4-5, laminibus late triangulo-ovatis vel subrotundis 3-4 = 2.5-3.0 cm longis et latis,
truncatis vel ad basi brevissimo-cuneatis, grosse spinuloso-dentatis, petiolis non alatis, 4-5 cm longis,
purpureis; capituli +-6, nutantes, punnor renee peconewas involucris 8-10 mm ales. cisco 7
As
hyll floribus radii 5
mm diametris
de oOo Oo
Perennial from a well-developed, often branched caudex, up to 2 dm tall, to-
tally glabrous; first year shoots 0.5 dm long, consisting of a few basal leaves on
long petioles; flowering stem (second season) with withered basal leaves, cauline
leaves with lamina broadly triangular-ovate to subrotund, 3.0-4.0 x 2.5-3.0cm,
truncate or very shortly cuneate at the base, coarsely spinulose-dentate, peti-
oles scarcely winged, 4-5cm long, purple; cauline leaves 4-5, similar to the basal
leaves; heads 4-6, nodding, the lower on long, erect peduncles; involucre 8-10
mm high, the disk 5-7 mm diam, with a single row of phyllaries and a few
minute basal bracteoles; phyllaries lance-oblong, obtuse or broadly acute, with
broad hyaline margins, glabrous; ray flowers few, little exceeding the involucre,
the lamina 7 x 2 mm, the veins 4-5, simple, the tube 4 mm long; disk flowers
with limb 4 mm, tube 4 mm long; pappi 3 mm long; cypselae linear, strongly
ribbed, 3 mm long.
SIDA 20(2): 511 — 513. 2002
512 BRIT.ORG/SIDA 20(2)
Fic.1. Senecio spribillei, holotype.
This new species is most closely allied with S. neowebsteri Blake (S. websteri
Greenman, non Hook.), an endemic of the Olympic Mountains of Washington
(Table 1). lt appears to be a narrow endemic of northwestern Montana.
The ray flowers in the new species are so short and so narrow that the col-
lector believed that the plant was rayless. The structural anatomy of the ray
flowers differs from that of Senecio neowebsteri. In S. spribillei the nerves of
the ray flowers are simple, unbranched; in S. neowebsteri each nerve is forked
near the base, the branches continuing closely parallel distally. Baaghoe (1977,
1978) published on the taxonomic application of ligule micro-characters in the
WEBER, A NEW SPECIES OF SENECIO FROM MONTANA
Taste]. Comparison of Senecio spribillei and $
Senecio spribillei
Senecio neowebsteri
Foliage glabrous
Leaf lamina almost isodiametric,
coarsely dentate, the base truncate
Petioles not winged
Heads several,6—-8 mm tall, disk 5-7 mm diam
Ligules of ray flowers 7-12 mm
hardly longer than the involucre
Tubes of disk flowers 4 mm, limb 4 mm
Cypselae linear, strongly ribbed, 5 mm long
Pappi white, 7 mm lon
Foliage initially floccose, then glabrate
Leaf lamina usually longer than broad,
variably dentate, the base cuneate
Petioles winged
Heads solitary, 10-15 mm tall, disk 9-12 mm diam
Ligules of ray flowers 25-30 X 3-5 mm,
much longer than the involucre
Tubes of disk flowers 6 mm, limb 4 mm
Cypselae linear. strongly ribbed, 3 mm long
Pappi white, 3 mm long
Asteraceae but made no mention of venation or possible differences involving
venation. Her work dealt with light and electron microscope observations,
mostly on epidermal features. The existence of branched and non-branched
nerves in Senecio, s. lat. deserves attention.
Distribution.—Known only from the type collection. According to the col-
lector, the locality is likely a nunatak north of the southernmost limits of the
Pleistocene ice sheets.
Etymology.—Senecio spribillei is dedicated to the young taxonomist and
phytosociologist, Toby Spribille, with whom I have been associated for several
years. His keen eye and ability to deal with the lichen and bryophyte as well as
the phanerogamic flora, his knowledge of phytosociology, and his fine collec-
tions, marks him asan important figure in the recent history of the Rocky Moun-
tain flora in Montana.
REFERENCES
PO J; cece Taxonomical application of ligule microcharacters in Compositae. I.
leae, Heliantheae, and Tageteae. Bot. Tidsskr. 71:193-224.
Bree J. 1978. Taxonomical application of ligule microcharacters in Compositae. Il.
Arctotideae, Astereae, Calenduleae, Eremothamneae, Inulae, Liabeae, Mutiseae, and
Senecioneae. Bot. Tidsskr. 72:125-147.
514 BRIT.ORG/SIDA 20(2)
Book NOTICES
CHARLES M. ALLEN, DAWN ALLEN Newman, and Harry H. WINTERS. 2002. Trees, Shrubs,
and Woody Vines of Louisiana. (ISBN 0-9718625-0-8, pbk). Allen’s Native
Ventures, LLC, 5070 Hwy 399, Pitkin, LA 70656, U.S.A. (Orders: 337-328-
2252, native@camtel.net). Price not given, 333 pp, numerous b/w line draw-
ings, 3 maps, 51/2" x 81/2"
This book provides information on 414 species that are native and naturalized and, in a number of
cases, Cultivated and persisting. The entries are listed alphabetically (by scientific name, sans au-
thor) by genus and species. A long paragraph on each of the species gives common names, name
derivations, morphological features, oe medipinal, economic, pee wildlife uses. The last
sentence of each paragraph notes the distribution in Lousiana: e.g., “It is widely irene a Panouigle:
out the state but more commonly in sandy soil (FAC) along es coast with reports from 18 parishes”
(Cissus trifoliata); “In swamps, edges of sie ponds and enue water bodies Ade wath current re-
ports from 46 parishes” (Nyssa aquatica); “Widely culti | thick
ets; this species is a serious pest in many mae of the state (FAC)” (Ligustr inense): “The current
records, mostly from cultivated plants, include seven parishes” (Photinia serratifolia), “Planted
throughout the state (FACU) as an ornamental with some recent interest in its wood. The current
records include 14 parishes” (Paulownia tomentosa). Apart from the enumeration of which woody
species occur in the state eee their reproductive status, perhaps the most peel feature of the book is
the line drawing for cies, drawn froma range of previously published sources. Identification
ng
keys are not ae but 8 ‘tables’ in the introductory material list genera with distinctive charac-
ters such as strong odor, compound leaves, opposite or whorled leaves, etc. This compilation will
serve good use as a reference in company with an identification manual and range maps.—Guy L.
Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX, 76102-4060, U.S.A.
Joun Graves, Photos by Wyman MEINzeR. 2002. Texas Rivers. ISBN 1-885696-
38-8, hbk.) University of Texas Press, PO. Box 7819, Austin, TX 78713-7819,
US.A. (Orders 800252-3206 512-471-4032, 800-687-6046 fax)) $39.95, 144
pp.80 color a "x 1".
If Texas Rivers w ly k it would be an outstanding work. The photographs by Wyman
Meinzer are brebtakingy ee eae and close-ups alike are full of radiant color and
LE
telling detai of the particular area through which the river flow
text by Johny hn G is al plete in itself, a testament to the beauty and diversity and
total utility of Texas rivers. He touches on all aspects of riverine territories from their archeology,
otany, climate, economy through the alphabet of scientific studies to zoology. It is as if one is taking
a leisurely walking tour along the river witha very knowledgeable friend with time to stop to appre-
ciate the prea and small Hones rs of this multifarious land.
Together la very beautiful book,a welcome addition toan library.—
Ruth Ginsburg, Fort Worth, Texas.
SIDA 20(2): 514. 2002
EUCHITON (ASTERACEAE: GNAPHALIEAE) IN
NORTH AMERICA AND HAWAII
Guy L.Nesom
i
nstitute of Texas
—
509 Pecan Street
Fort Worth, TX 76102-4060, U.S.A.
ABSTRACT
Euchiton is treated apart from Gnaphalium primarily on the basis of its tendency for stolon produc-
tion and cypselar surfaces with paired, non-mucilaginous epidermal papillae. Three species of
Euchiton are recorded in North America: F. sphaericus (naturalized in California and Oregon), E.
gymnocephalus (naturalized in California and Oregon), and E. involucratus (apparent waif in Cali-
fornia and Massachusetts, the first published report of this species in North America). The first two
species also are naturalized in Hawaii. A nomenclatural and ecological summary is presented for
each species and a key clarifies their identifications, which have been confused in North America.
RESUMEN
ee eT en al ary Dali ‘ j } | i la produccion
x
7] en + ] |
LB i QC
] ; re uie 4
Se han
de ole ones y superficie Se ee are con}
Norte ae brica: FE
Glioma y Ores n),
-alifornia y y Oregon) - salaebidiaee aaa eG abandonada
en Cali ae y Massachusetts, es | sie en Norte América). Las dos Ss primeras
lL a ( i ]
especies estan también Pie en Hawaii. Se piece un resumen nomenclatural y ecoldgico
e cada especie y una clave clarifica sus identificaciones, que han sido confusas en Norte América.
Euchiton is treated apart from Gnaphalium primarily on the basis of its ten-
dency for stolon production and cypselar surfaces with paired, non-mucilagi-
nous epidermal papillae (Anderberg 1991). The plants are mostly perennial and
produce heads in terminal, capitate clusters immediately subtended by a whorl
of leafy bracts. The distinctiveness of the group was emphasized by Drury (1972)
and Holub (1974); nomenclatural transfers to Euchiton were made by Holub
(1974) and completed by Anderberg (1991) and Ward and Breitwieser (1998).
The species number about 22 (Drury 1972; Anderberg 1991), including recent
additions (Buchanan 1999, Walsh 1999) or 20-30 (Walsh 1993). They are native
to Australia and New Zealand and probably to New Guinea and eastern Asia;
some species have weedy tendencies and have become widely naturalized.
The following synopsis of Euchiton in North America and Hawaii is based
on surveys of collections at ARIZ, BRIFSMU, CAS-DS, GH, HSC, MO, NCU, TEX-
LL, UC-JEPS, and WIS.
Ichellus Cass.
ree Cass. in F Cuvier, Dict. Sci. Nat. ed. 2, 56:214. 1828. Type species: Euchiton
pu
= Euchiton involucratus (G. Forster) Holub
SIDA 20(2): 515 — 521. 2002
516 BRIT.ORG/SIDA 20(2)
Plants perennial or (less commonly) annual, herbaceous, stoloniferous in most
species (taprooted in 1), white-tomentose, eglandular. Leaves basal and cauline,
sometimes ina rosette, lanceolate to linear, entire, sessile or petiolate, bicolored,
usually witha close tomentum at least on the abaxial surfaces. Capitulescences:
heads usually ina terminal cluster subtended by a whorl of leafy bracts, some-
times with axillary clusters below. Capitula narrowly campanulate to short-
cylindric; phyllaries chartaceous, transparent, inner with larger stereomes. Pis-
tillate florets fertile, more numerous than the bisexual florets, corollas purple
or purple-tipped. Bisexual florets fertile, corollas purple or purple-tipped.
Cypselae oblong, 0.6-1.5 mm, epidermis minutely papillate (the papillae im-
bricate and paired, nonmyxogenic); pappus of caducous, separate or weakly
basally connate bristles separating in groups. Base chromosome number, x =
1. Annual, taprooted; leaves not at all clasping or ree capitula in a vine
cluster; bisexual florets 1 per capitulum; capitulescence bracts 4-8. uchiton sphaericus
. Perennial or biennial, fibrous-rooted; leaves subclasping or pe aelrtiaes th ina
Hevea cluster; bisexual florets 3-7 per capitulum; capitulescence bracts 2-5.
Stolons commonly present, plants forming colonies; basal leaves present in a
rosette at flowering; cauline leaves 2—4(-6), linear to oblanceolate, 0.5— a Ea
long, 1-3 mm wide; pistillate florets 40-60; capitulescence ar 2-3,1
ie g the capitulescence or only slightly so Euchiton cancel
2. nt usually absent, plants not evidently een: basal leaves usually with-
ered or absent at flowering; cauline leaves 6-10, mostly linear, 3-8 cm long, 2-3
mm wide; pistillate florets 80-150; capitulescence bracts 3-5, distinctly longer
than the capitulescence. Euchiton involucratus
1. Euchiton gymnocephalus (DC.) Holub, Folia Geobot. & Phytotax. 9:271. 1974
Inon (DC.) A. Anderberg 1991]. Gnaphalium gym nocephalum DC.,, Prodr. 6:235. 1838.
Type: AUSTRALIA
Gnaphaliumcollinum Labill., Nov. Holl. Pl. 2:44, t. 189.1806. Gnaphalium japonicum var.collinum
abill.) Maiden & Betche, Census New South Wales PI. 204. 1916. Type: AUSTRALIA
Flowering (late March-)May-October(-November). Grassy hills, margins and
openings in woods, roadsides, cutover areas; ca. L1OO-2500 ft (30-750 m); California
and Oregon, Hawaii. Apparently native to Australiaand New Zealand. Recorded
in Australia as “common and widespread, particularly in cooler parts ..., often
colonizing bare ground, stream and track margins etc.” (Walsh 1993, p.823) and
“woodland areas near Canberra and also extending to upper slopes of moun-
tains, widespread in many parts of Australia and Tasmania” (Burbidge & Gray
1970). In New Zealand, it occurs in a wide variety of habitats but is “typically
associated with forest and scrub communities between sea level and 1,000 (-
2,000) ft” (Drury 1972, p. 144).
A handwritten note by J.P. Tracy on the JEPS sheet of Tracy 14110 notes the
following: “My record of this plant [Euchiton gymnocephalus, presumably in
Humboldt County]extends back to 1900 when it was already well estab.” Cita-
NESOM, EUCHITON IN NORTH AMERICA AND HAWAII 517
tions below indicate that Tracy collected the species repeatedly in Humboldt
County from 1904 through 1946 and collections between 1960 and 1989 have
continued to document its occurrence in counties of northeastern California
and southeastern Oregon.
Collections examined. UNITED STATES. eae Del Norte Co.: 3 mi S of Klamath River, grassy
hills near coast, 300 ft, 12 May 1929, Tracy 8588 (CAS, UC). Humboldt Co.: near Azalea Reserve State
Park, cool woods in partial shade, 100 ft, 15 Jul 1960, yee: s.n. (HSC); along Mad River ca. 2 mi S of
Blue Lake, 28 Jun 1971, Anderson and Smith J-1230 (HSC); 8 mi W of Berry Summit along US 299,
2500 ft, roadcut on serpentine soil, 15 May 1965, Bos 254 (HSC); Lord Ellis road near Shingle Mill, 28
Mar 1926, Kildale 1670 (DS); Lord Ellis road near Blue Lake, 25 Nov 1926, Kildale 2830 (DS), Trinidad,
Camp Twenty One, 600 ft, 30 Jul 1932, Parks and Tracy 1133 (UC), above dirt road from old 101 to the
t the N bank of the Mad River, grassy hillside in half shade, 22 Jul 1967, Stevens 61 (HSC);
Garberville Quad., old skid road, shady edge of redwood forest and mixed evergreen forest, 850 ft,
moss-covered rocky subsoil, 13 Aug 1976, Sutherland 9.13.8.76 (ASC); Shubrick Peak Quad., Kinsey
Ridge above Oat Creek, logged doug fir forest, 24 Apr 1978, Sutherland 6.24.4.78 (HSC); immediate
vicinity of Eureka, small patches in open places in woods, 0-500 ft, 31 Jul 1904, Tracy 2134 (CAS, DS,
GH, UC); Boynton Prairie road, near Carroll place, 1000 ft, 23 Jun 1918, Tracy 4958 (NCU, UC); near
- Lake, in woods, spreading by stolons and forming patches, 1000 ft, 1 Apr 1923, Tracy 6175 JEPS,
near Boynton Prairie, in recently logged area, among “fireweeds,” 2000 ft, 28 Aug 1927, Tracy
nee ae logged-oft ands eee mats, 10-200 ft, 13 Aug 1932, Tracy 10349 (ARIZ,
CAS, UC); ne ue Lake, road ods on Trinity Hwy, spreading and forming mats, 900
ft, 20 Jul ee Pa 14110 (CAS, JEPS, UC-2 ee 4 mi SE of Korbel, near “Angels Ranch,” local in
grassy moist places, 1000 ft, 22 Mar 1936, Tracy 14784 (CAS, DS, JEPS, UC-2 sheets); Fickle Hill, 6 mi
ata, in recently cleared land, 2000 ft, 17 May 1936, Tracy 14811 (UC); 2 mi NE of Orick, on
Bald ale road, local by oe 500 ft, 29 May 1936, Tracy 14829 (CAS, DS, GH, JEPS, UC); hillsides
near Canyon Creek, 6 mi SE of Blue Lake, in logged-over land, 1200 ft, 1 Aug 1936, Tracy 15058 (UC);
Carlotta, in recently ee area, with ee haeta ustulata] and [Euchiton sphaericus], 100 ft, 23
Aug 1936, Tracy 15164 (UC); Fickle Hill, 6 mi SE of Arcata, in country recently logged, 2000 ft, 24
Aug 1936, Tracy 15175 (UC); vicinity of C ae in recently cleared land, ca. LOO ft, 15 Jun 1938, Tracy
15936 (UC); “Riverside Park” near old Strong’s Station District School House (burned down), 30 May
1946, Tracy 17548 (UC); Kneeland Prairie, Dan McBride's place (SE end of Kneeland), local, in partial
shade of open fir woods, 2500 ft, 23 Jun 1946, Tracy 17596 (UC). Mendocino Co.: 15 air mi SW of
Garberville, 60 air mi SSE of Eureka, 1/4 mi E of Bear Harbor on Lost Coast Trail, edge of trail bed,
California ua riparian forest, 120 ft, 8 Jul 1989, Bowcutt 1369 (HSC); 3.2 km N of jet with Rte 1, along
Usal Rd, openings in second growth redwoods, 305 m, I] Jun 1981, Smith 6632 (CAS, HSC). elk
Co.: Forest See Road 17NO4, Doe Flat, TI7N, R4E, Sec. 35, [no date], Van Deventer s.n. (HSC). H.
WAIL. Hawaii, Upper Waiakea Forest Reserve, along Disappointment Trail (Pwu Maka’ala access “a )
of Stainback Hwy., in Metrosideros rainforest, ca. 1100 m, 26 Jul 1983, Wagner et al. 4842 (US). OR-
Curry Co. gment of Hwy 101 that les ads as a dead-end from Harris Beach State Park,
just N of Brookings, 10 Jun 1964, Chambers 2241(DS, NCU); Brookings, 100 ft, | Aug 1937, Tracy 15606
(UC); moist slope 5 mi N of Brookings, 7 Jul 1939, Desk 20445 (CAS, UC). Klamath Co.: Winema Na-
a7
tional Forest, Cold Springs Road and State Route 140, T36S R6E, Sec. 18, 14 Oct 1978, Sawyer 3320
(HSC)
—
ct
Wagner et al.(1997) recorded Euchiton gymnocephalus, identified as E. japonicus
(Thunb.) Holub, as a new state record for Hawaii. In Australia it was identified
as Gnaphalium japonicum Thunb. by Burbidge and Gray (1970), but the syn-
onymy of G. japonicum with G.gymnocephalum was later rejected without spe-
cific comment by Cooke (1986) and Walsh (1993). In the present review, plants
518 BRIT.ORG/SIDA 20(2)
from Japan and China appear to be consistently distinct from related ones of
Australia and New Zealand, although differences are subtle, and until some-
one may provide a more detailed study of the complex, the morphological af-
finity of the North American plants with those of Australia is reinforced by the
nomenclature. Differences between the two taxa are outlined in the following
couplet.
Basal leaves usually narrowly oblanceolate, attenuate to a distinct petiolar portion,
the cauline abruptly differentiated in size and shape from the basal, smaller and lin-
ear-lanceolate to linear-oblanceolate; adaxial leaf surfaces quickly glabrescent and
shiny-green; stereome thickened and opaque, distinctly narrowed apically and sepa-
rated from the lamina by a short, completely translucent portion; cypselae ca
mm long. Euchiton ayminecephaliis
Basal a cauline leaves linear-lanceolate to linear-oblong, without a distinct petiolar
portion, the lower cauline ag htly reduced in size ene slmlet in also to the basal;
adaxial leaf surfaces | arachnoid, ap = thin-translucent,
not evidently narrowed apically, more or ice directly eonneees to the lamina;
cypselae ca. 1.0 mm long. Euchiton japonicus
Plants from Japan, China, and Taiwan (specimens examined), and probably from
Ryukyus and Korea are typical E. japonicus (see citation below). Collections re-
ferable to E. japonicus or E. gymnocephalus from New Guinea, New Caledonia,
and Java need to be studied in order to assess their relationship. Reports of E.
japonicus from the Philippines evidently refer, at least in part, to Gnaphalium
oblanceolatum Elmer, which apparently is more closely related to E.
involucratus (but probably not conspecific with it) than to E. japonicus.
Euchiton japonicus (Thunb.) Holub, Folia Geobot. & Phytotax. 9:271. 1974 [non
(Thunb.) Anderb. 1991]. Gnaphalium j japonicum Thunb., FL Jap. 311. 1784. Type: JAPAN
(“prope urbem Nagasaki”).
2. Euchiton involucratus (G. Forster) Holub, Folia Geobot. & Phytotax. 9:271.
1974 (non A. Anderberg 1991). Gnaphalium involucratum G. Forster, Fl. Ins. Austr,
55.1786. Type: NEW ZEALAND.
Flowering July-October. Grassy open places, often moist or wet; ca. 50-600 m:;
California and Massachusetts. Native to Australia and New Zealand; collections
also observed (present study) from New Guinea, New Caledonia, Java, and Tai-
wan. The species in Australia is recorded as a “weed of gardens and disturbed
ground” (Burbidge & Gray 1970); it also is common “particularly in swampy
sites, from near sea-level to the higher alps where usually in Sphagnum bogs”
(Walsh 1993).
Both of the North American collections cited below probably represent
waifs, because this species apparently has not been subsequently recorded in
floristic summaries from anywhere in the USA. This apparently is the first
eee report of E. involucratus from North America.
Collecti ined. UNITED STATES. California. Humboldt Co.: 5 end of Fickle Hill, 6 mi SE of
Arcata, ca. 2000 [t, moist ae among rushes and sedges, spreading by rootstocks, perennial, 26
NESOM, EUCHITON IN NORTH AMERICA AND HAWAII 519
Sep 1948, Tracy 18232 (ARIZ, NCU, UC-2 sheets, WIS). Massachusetts. Hampshire Co.: Northampton,
weed in garden, Sep 1900, Mrs. E.H. Terry s.n. (GH).
3. ees peers (Willd.) Holub, Folia Geobot. & Phytotax. 9:271. 1974
A. Anderberg 1991). Gnaphalium sphaericum Willd., Enum. Hort. Berol. 2:868.
ee Type: AUSTRALIA.
Gnaphalium involucratum G. Forster var. simplex DC., Prodr. 6:236. 1838. Based on G. sphaericum.
Gnaphalium involucratum G. Forster var. ramosum DC,, Prodr. 6:236. 1838. TyPE: AUSTRALIA.
Gnaphalium morii Hayata, Ic. Pl. Formos. 8:58. 1919. TyPE: Taiwan (“Formosa”).
?Leontopodium japonicum (Thunb.) H. Lév. var. sandwicense H. Lév, Fedde Repert. 10:121. 1911.
BE: All.
Gnaphalium japonicum sensu various authors, non Thunberg 1784.
Flowering (late June, Tracy 15974; July-)August-October. Grassy open places in
wooded areas, recent clearings and clearcuts, disturbed soil, especially along
roadsides; ca. 100-2000 ft [30-600 ml}; California and Oregon, Hawaii (Molokai,
Lanai, Maui, and Hawaii, fide Wagner et al. 1999). “Reported from San Joaquin
County,” California (Ferris 1960; voucher not seen in present study). Native to
Australia and New Zealand; also documented from New Guinea, New
Caledonia, Java, Philippines, Japan, and Taiwan (Drury 1972). In Australia re-
corded as “a very common and widespread species, occurring on a wide variety
of substrates, e.g., mallee and coastal sands, clayey floodplain area, and often
colonizing disturbed ground” (Walsh 1993).
Euchiton sphaericus has long been identified in California, apparently be-
ginning with Howell (1937), as Gnaphalium japonicum; it recently was correctly
identified by Wagner et al. (1997) and by Peter W. Michael (annotations on speci-
mens at GH). Correct nomenclature was earlier provided by Drury (1972).
Euchiton sphaericus was established in California at least by 1915, when appar-
ently first collected there. J.P. Tracy collected it repeatedly from that year through
1949; from one locality (Tracy 14752) he noted that it was “scarce now, abun-
dant 15 years ago.” Tracy also observed (fide label of Tracy 5097) that the species
was “probably introduced in grass seed sown on logged-over country.” Relatively
recent collections (1954, 1977, 1992) confirm its persistence in northwestern
California.
Euchiton sphaericus was collected in Hawaii, perhaps for the first time, in
1909 (on Maui, Faurie 928-BISH, as noted by Wagner et al. 1999). Rock (1914, p.
352) noted that in Hawaii E. sphaericus was among several species “imported
accidentally during the last 10 or 20 years, by the cattle estates with grass seeds.”
Collections ined. UNITED STATES. California. Del Norte Co.: Sutton Creek, 2 mi E of the Van
Deventer Ranch and 2.5 miair-line E of Fort Dick, 350 ft, 15 Oct 1954, Van Deventer 412 (UC). Humboldt
.: Eel River, 16 mi N of Garberville, 29 Aug 1936, Howell 12888 (GH, LL, WIS); logged redwood flat,
xa mi E of Carlotta, 15 Aug 1936, Jepson 17,898 JEPS); mouth of Laribee Creek, common as a weed but
local, 200 ft, 12 Sep 1915, Tracy 4677 (CAS, MO, UC); around Humboldt Bay, Newells Camp on old
Kneeland Prairie road, probably introduced in grass seed sown on logged over country, 1500 ft, | Sep
1918, Tracy 5097 (NCU, UC-2 sheets); South Fork of Eel River, 4 mi above the mouth, common road-
520 BRIT.ORG/SIDA 20(2)
side weed, 200 ft, 29 Sep 1918, Tracy 5103 (TEX, UC); at Eureka, introduced along old road in stump
land, 0-200 ft, 6 Oct 1918, Tracy SII8 (UC); North Fork of Mad River, along newly constructed road,
evidently i luced and not yet c here, LOOO ft, 19 Sep 1920, Tracy 5399 (UC); South Fork of
Rel River, near etme 300 ft, 9 Aug 1925, Tracy 7229 (UC); along Lord-Ellis Road, headwaters of
North Fork of Mad River, scarce, 2000 ft, 8 Aug 1931, Tracy 9608 (UC); near Dyerville, roadside weed,
scarce now, ne 15 ae ago, 300 ft, Sep 1935, Tracy 14752 (UC), Carlotta, locally abundant asa
“fireweed” in recently cleared land, 100 ft, 23 Aug 1936, Tracy 15162 (GH, HSC-2 sheets, MO, TEX, UC-
3 sheets, WIS); Fickle Hill, 6 mi SE of Arcata, restricted to land recently cleared and soil disturbed,
2000 ft, 24 Aug 1936, Tracy 15176 (GH, JEPS, UC); Trinity River Valley, at Willow Creek, weed in gar-
den, not common, 500 ft, 20 Jun 1938, Tracy 15974 (UC); Trinity River Valley, along new road to Hors
Linto Creek, disturbed soil, 800 [t, 4 Sep 1938, Tracy 16142 (UC); Trinity River Valley, near Raccoon
Creek, moist ground disturbed by road work, 500 ft, 1] Aug 1940, Tracy 16694 JEPS, LL-2 sheets, UC):
Valley of South Yager Creek, recently logged land as a ‘fireweed’ on the Redwood House Road at old
mill site, | mi N of Redwood House, 1500 ft, 26 Jul 1942, Tracy 17299 (UC); Park’s place, 10-200 ft, 28
Sep 1947, Tracy 17933 (UC); Fickle Hill, 6 mi SE of Arcata, recently logged area, scarce, ca. 2000 ft, 26
Sep 1948, Tracy 18230 (UC); Van Duzen River, near Grizzly Creek, frequent in disturbed soil of recent
logging operations, 300 ft, lo Oct 1949, Tracy 18591 (UC, WIS); Aikens Campground, TION R5E, Sec.
30, mixed evergreen forest, | Jul 1977, Sawyer 2990 (HSC). Lake Co.: road to Bartlett Mountain, Aug
1930, Mason 5732 (UC). Mendocino Co.: 10 air mi W of Leggett, plone and in dirt roadbed of Hotel
-]
Cc:
Gulch Road 5.3 road mi from Usal, S-facing slope, coast redw s fir forest logged about 15-
20 years ago, 21 Jul 1992, Boweutt 1571 CHSC); Mendocino Woodlands, 10 Jul | 938, Eastwood and How-
ell 6238 (UC). Trinity Co.: Sharber Slough on the Trinity River, 3 mi E of the South Fork, along trail,
sparsely introduced in moist ground, 6 Sep 1926, Tracy 7783 (CAS, UC). HAWAIL. Hawaii: Hawaii Na-
tional Park, Kilauea Crater, on dry lava, 18 Jul 1931, van Loben Sels 595 (UC); Waimea Mts., Jun 1910,
Rock 8433 (GH); Hawaii National Park, weed on xeric cinder flats in Kau Desert near Halemaumau,
3600 ft, 18 Jun 1948, Webster and Wilbur 1793 (TEX). Lanai: Mts., E end, Jun 1913, Forbes 276.2 (UC).
Maui: Koolau Gap, Haleakala, volcanic cinders, 10 Aug 1927, Degener 18451 (GH); Haleakala, 16 Aug
1927, Degener 18450 (GH); Olinda, Pipe-Line Trail, 30 Jul 1927, Topping s.n.(GH); Haleakala, Aug 1909,
Faurie 928 (BISH, as cited by Rock 1914 and Wagner et al. 1999; not seen). OREGON. Curry Co.:
ground along Chetco River, 3 mi above Harbor, 20 Jul 1919, Peck 8924 (GH).
ACKNOWLEDGMENTS
lam grateful to the staffs at GH, MO, NCU, and TEX-LL for help during recent
visits to those institutions. Loans were studied from ARIZ, CAS, DS, HSC, JEPS,
UC, and WIS. Comments by John Strother contributed to the consistency and
clarity of presentation.
EFERENCES
ANDERBERG, A.A. 1991. Taxonomy and phylogeny of the tribe Gnaphalieae (Asteraceae).
Opera Bot. 104:5-195.
BucHANAN, A.M. 1999. A new species of Euchiton (Gnaphalieae: Asteraceae) from southern
Tasmania, Australia. Pap. Proc. Roy. Soc. Tasmania 133:115-116
Bursioce, N.T.and M. Gray. 1970. Flora of the Australian Capital Territory. Australian National
Univ. Press, Canberra.
Cooke, D.A. 1986. Gnaphalium. In: Jessop, J.P.and H.R. Toelken, eds. Flora of South Australia.
Part Ill, Polemoniaceae—Compositae. South Australian Govt. Printing Division, Adelaide.
Pp. 1516-1519.
NESOM, EUCHITON IN NORTH AMERICA AND HAWAII 521
Drury, D.G. 1972. The cluster and solitary-headed cudweeds native to New Zealand:
(Gnaphalium section Euchiton - Compositae). New Zeal. J. Bot. 10:112-179.
Ferris, R.S. 1960. Vol.IV,Bignoniaceae to Compositae.In Abrams, L.and R.S. Ferris, Illustrated
flora of the Pacific States. Stanford Univ. Press, Stanford, California.
Hotus, J. 1974. New names in phanerogamae. Folia Geobot. & Phytotax. 9:261-275.
Howe, J.T. 1937. Three species of Gnaphalium adventive in California. Leafl. W. Bot. 2:
10-12.
Rock, J.F.1914.Revisio plantarum Hawaiiensium a Léveillé descriptarum. Repert. Spec. Nov.
Regni Veg. 13:352-361.
Wacener, W.L., R.K. SHANNON, and D.R. Heresr. 1997. Contributions to the flora of Hawai'i. VI.
Bishop Mus. Occ. Papers 48:51-65.
Wacner, W.L., D.R. Hersst, and S.H. Somer. 1999. Manual of the flowering plants of Hawai'i
(rev.ed.), Volume 1. Univ. of Hawai'i Press and Bishop Museum Press, Honolulu, Hawaii.
Watsh, N.G. 1993. Euchiton. In: N.G.Walsh and T.J. Entwhistle, eds. Flora of Victoria, Volume
4.Inkata Press, Melbourne, Australia. Pp. 820-825.
WALSH, N.G. 1999. New species in Asteraceae from the subalps of southeastern Australia.
Muelleria 12:223-228.
Warb, J.M. and |. Breitwieser. 1998. New combinations in Euchiton (Compositae -
Gnaphalieae) from New Zealand. New Zeal. J. Bot. 36:303-304.
522 BRIT.ORG/SIDA 20(2)
Book NotIcE
DENNIS BLAGG. 2002. Big Bend Landscapes. (ISBN 1-58544-202-X, hbk.). Texas AQM
University Press. John H. Lindsey Building, Lewis Street, 4354 TAMU, College
Station, TX 77843-4354, U.S.A. (Orders: wwwtamu.edu/upress, 979-458-
3982, 800-826-8911, 979-847-8752 fax). $40.00, 160 pp, 50 color paintings,
15 b/w drawings, 12 1/4" x 93/4".
Ron Tyler's introduction to Big Bend Landscapes is a succinct history of the region from Indian times
to the creation of a public park in a portion of this great wilderness. He also traces the rise of the
Romantic movement and the enthusiasm for landscape painting asa result of the awe-inspiring ex-
panses and mountains of the West. The Big Bend country is certainly a manifestation of the ‘untar-
nished handwork of the Creato
his essay is a fitting sete to the fifty paintings and nineteen drawings which are repro-
duced here in beautiful color The diversity of the country, the effect of light, and the astonishing
formations of rocks and clouds are very well portrayed. In the vast panoramic views, the details are
se r-
astonishing. The short explanations which accompany each picture are interesting and adda
sonal note. But the paintings speak for themselves. They co pie) a Seite and a passion for na-
ture. The artist isa wot thy successor of such g Sees American landsc< ape painting as Cole, Durand,
Church, Kensett, and Bierstadt.—Ruth Ginsburg, Fort Worth, Texas.
Witrrip BLUNT with an introduction by WiLLiAM T. STERN. 2001. Linnaeus. The
Compleat Naturalist. (ISBN 0-691-09636-8, hbk.). Princeton University Press,
41 William Street, Princeton, NJ 08540, U.S.A. (Orders: California Prine-
eton Fulfillment Services, 1445 Lower Ferry Road, Ewing, NJ 08618, US.A.,
609-883-1759, 609-883-7413 fax, www.pup.princeton.edu). $35.00, 264 pp.,
color photos, b/w photos, line drawings, maps, 7 5/8" x 97/8"
What a beautiful book and what a wonderful way to remember William T. Stearn (1911-2001). The
book, originally titled The Compleat Naturalist: A Life of Linnaeus, is published in his memory. The
following description of the book is an extract from the dust jacket. “This book gives a fascinating
and rounded portrait of Linnaeus the man, charting his rise from a poor student at Lund University
o Professor of Medicine at Uppsala and a founder of the Royal Academy of Sciences. Wilfrid Blunt's
engaging text is interspersed with vivid passages [rom his subject's own writings—from riveting de-
scriptions of adventures in the wilds of Lapl and toa ena account of Sjupp the raccoon.
Linnaeus’s family life and his relations witl |
1 pul I tee aoe gat mak ing scien
tific achievements. William Stearn’s appendix on Linnaean cl: mscilicaiaon provide oncise survey
of the basics necessary for understanding Linnaeus’s work.”
SIDA 20(2):: ):522. 2002
NEW COMBINATION IN SALIX (SALICACEAE)
Guy L.Nesom
Botanical Research Institute of Texas
509 Pecan Street
Fort Worth, TX 76102-4060, U.S.A.
Dorn (1998; following Cronquist 1964) has treated Salix interior Rowlee at infra-
specific rank within S. exigua Nutt., observing that a broad region of morpho-
logical intergradation exists between the two where their ranges are in contact
(see Dorn’s Fig. 2, p. 200). This observation is confirmed in field studies relating
toa floristic inventory of Lake Meredith National Recreation Area in Hutchin-
son, Moore, and Potter counties of the north-central Texas panhandle (Nesom
& O’Kennon in progress). In this area many populations exist of S. interior sensu
stricto (colonial shrubs with completely glabrous, linear-lanceolate leaves with
widely spaced and elongate teeth) as well as S. exigua (similar to S. interior but
with densely silky-sericeous lower leaf surfaces). Also common are populations
with foliar vestiture in varying degrees of intermediacy (e.g., Nesom & O’Kennon
LM359, LM198 - BRIT). Salix nigra Marsh and S. amygdaloides Anderss. also
commonly occur in the area, apparently without intergrades.
Argus (1986) treated Salix exigua as including S. interior without formal
rank, but he noted (p. 91) that the “eastern phase (S. interior) differs from the
western phase (S. exigua) in having leaves less densely sericeous, more distinctly
toothed, and more veiny.” In my observation (based on BRIT and VDB collec-
tions), variability in vestiture is greater in the western phase than in the east-
ern phase, where the green-glabrous leaves are consistently evident. Sericeous
undersurfaces are characteristic of the western phase.
Dorn recognized three varieties within Salix exigua subsp. exigua but
treated S. interior only at subspecific rank: S. exigua subsp. interior (Rowlee)
Cronquist. The following combination provides consistent nomenclatural rec-
ognition of all infraspecific taxa, following ICBN Article 4.1: “The secondary
ranks of taxa in descending sequence are tribe (tribus) between family and ge-
nus, section (sectio) and series (series) between genus and species, and variety
(varietas) and form (forma) below species” (see commentary in Turner & Nesom,
Sida 19:257-262. 2000).
Salix exigua var. sericans (Nees) Nesom, comb. nov. Basiony™: Salix longifolia var.
sericans Nees in Wied-Neuw., Reise Nord-America 2:448. 1841. TYPE: not seen.
This is the oldest available name at varietal rank for plants identified as Salix
interior. As noted by Dorn (1994, p. 92), the type locality apparently is “in or
near the present Cass Co., Nebraska” (eastern Nebraska, well within the range
of typical S. interior and east of the area of its co-occurrence with S. exigua var.
SIDA 20(2): 523 — 524. 2002
524 BRIT.ORG/SIDA 20(2)
exigua, although the epithet implies that the leaves probably were not com-
pletely glabrous.
ACKNOWLEDGMENTS
Comments by George Diggs and an anonymous reviewer are appreciated.
REFERENCES
Arcus, G.W. 1986. The genus Salix (Salicaceae) in the southeastern United States. Syst. Bot.
Monogr. 9:1—1 70.
Cronauist, A. 1964. Salix. Vasc. PI. Pacific Northw. 2:37-71.
Dorn, R.D. 1994. North American Salix (Salicaceae): typifications and notes. Phytologia
7789-95.
Dorn, R.D. 1998. A taxonomic study of Salix section Longifoliae (Salicaceae). Brittonia
50:193-210.
TAXONOMY OF THE MOST COMMON WEEDY EUROPEAN
ECHINOCHLOA SPECIES (POACEAE: PANICOIDEAE)
WITH SPECIAL EMPHASIS ON
CHARACTERS OF THE LEMMA AND CARYOPSIS
Mihai Costea! Francois J. Tardif
se ee ant Agriculture Department of Plant Agriculture
niversity of Guelph University of Guelph
Guelph, ae ee 2W1, CANADA Guelph, Ontario, NIG 2W1, CANADA
coste_amihai@hotmail.com ftardif@uoguelph.ca
ABSTRACT
The most common weedy Echinochloa species 1 in Europe E.colona (L.) Link, E. crus-galli (L.) Beauv,
E. hispidula (Retz.) Nees ex Royle, E. oryzoides (Arduino) Fritsch, E. oryzicola (Vasing.) Vasing. are
f
for the first time analyzed regarding the micromorphology of as ang caryopsis i anatomy o
the caryopsis and the size of the starch grains. New potentially val { the
caryopsis, the pattern of surface variation of the caryopsis coat, and the size of the starch grains. A
cautionary approach is suggested towards the taxonomic utility of the silica bodies shape. Based on
the new characters, the relationships between the species are analyzed. Some nomenclatural issues
involving E. oryzicola, E.oryzoides, and E. hispidula are also discussed. For the first time, E. colona is
recorded from Romania.
RESUMEN
Las especies mas comunes de Echinochloa en Europa que son E. colona (L.) Link, E. crus-galli (L.)
Beauv, E.hispidula (Retz.) Nees ex Royle, E.oryzoides (Arduino), Fritsch, E.oryzicola (Vasing.) Vasing.
han oe an anegas as pun vez mediante pulcromionologss del lemma y del caviopsids, la
> Jo] l
YI
considerar de gran alee son el tamano del caridpside, la variacion en el patron de la Cee de la
cubierta del pe ae y el tamano vee los Sian de almidon. No se sugiere como herramienta
taxonomica Basandose en los nuevos caracteres, se analiza la relacion
entre las especies. Algunos ponies de nomenclatura que nee a E. oryzicola, E.oryzoides y E
hispidula son también discutidos. Por primera vez E. colona se cita de Rumania.
INTRODUCTION
Echinochloa is an economically important genus because many of its species
are noxious weeds. In Europe, E. hispidula, E.oryzoidesand E.oryzicola are most
often found as weeds in rice fields while E. crus-galli and E. colona although
present in rice fields, are more often encountered in irrigated fields and amongst
vegetable crops (Kossenko 1947; Vasconcellos 1954; Pirola 1965; Chirila 1984;
Tzvelev 1976; Haflinger & Scholz 1980; Carretero 1981; Mosyakin 1996; Ciocarlan
2000). The taxonomy of the genus is controversial due to nomenclatural prob-
‘Corresponding author
SIDA 20(2): 525 — 548. 2002
526 BRIT.ORG/SIDA 20(2)
—
lems and the continuous morphological variation exhibited by the taxa. For
the practical purpose of plant identification, the species often lack conspicu-
ous identification characters. Previous treatments and taxonomic studies are
often contradictory and, consequently, the limits of the taxa are uncertain and
specimens in herbaria are often misidentified. Other important taxonomic treat-
ments of the genus (or part of the genus) are those of Stapf (1899, 1934), Hitch-
cock (1920), Wiegand (1921), Kossenko (1947), Ohwi (1942, 1962), Martinez
Crovetto (1942), Vasconcellos (1954), Bor (1960), Pirola (1965), Chirila (1967,
1984), Yabuno (1962, 1981), Ali (1968), Gould et al. (1972), Tzvelev (1976),
Haflinger & Scholz (1980), Clayton (1980), Carretero (1981), Michael (1983) and
Brussoni (1994). A general characterization of the genus can be found in Wat-
son and Dallwitz (1992-onwards). Carretero (1981) provided a summary of the
mor pnolagie characters for European weedy species of the genus Echinochloa.
The identification of new characters useful in determining taxonomic
boundaries have shed light on our understanding of difficult groups such as
grasses. The study of lemma micromorphology can provide characters valu-
able in elucidating the delimitation of taxa, and these characters may reflect
systematic relationships. Micromorphological features of glumes and bracts
(lemma and palea) have been studied in other genera from Poaceae by: Bjorkman
(1960), Hsu (1965), Baum (1971), Lucas (1979), Clark and Gould (1975), Thomas-
son (1978a; 1978b, 1980, 1981, 1986), Shaw and Smeins (1979), Terrel and Wergin
(1981), Terrel et al. 1983), Webster and Hatch (1983), Bark worth (1983), Peterson
(1989), Soderstrom and Zuloaga (1989), Kellogg (1990), Zuloaga and Judziewicz
(1991), Valdes-Reyna and Hatch (1991), Molina (1993), Naredo et al. (1993), Ball
et al. (1993, 1999) and Snow (1996, 1998).
Caryopsis morphology has received a lot of attention since the beginning
of the century; however, most studies have focused on the general morphology
and the structure of the embryo. There is comparatively less information avail-
able on the anatomy of the caryopsis coat in various other grasses (Netolitzky
1926; Anderson 1927, Avery 1930; Krauss 1933; Hayvard 1938; Akerberg 1943;
Bradbury et al. 1956; MacLeod & Palmer 1966; Kowal @ Rudnicka-Sternowa
1969; Rost 1973: Jones & Rost 1989; Rost et al. 1990; Ungurean & Costea 1994,
1997). According to our knowledge, micromorphology and anatomy of the cary-
opsis in Echinochloa have not been studied so far.
The purpose of this study is to evaluate the taxonomic usefulness of select
floral characters in Echinochloa, namely, micro- and macrocharacters of the
lemma and caryopsis, the caryopsis coat, and the size of starch grains. Further-
more, the nomenclature, the taxa limits and the interspecific relationships in
Echinochloa are also discussed.
MATERIAL AND METHODS
Samples were collected and identified from populations in Spain and Romania.
COSTEA | AND CARVNDCIC 527
The accessions from Spain were collected and identified by Professor José Luis
Carretero from Polytechnic University Valencia, Spain (Table 1). The samples
from Romania were collected and identified by the first author (Table 1). Mature
spikelets were collected from the median region of the panicles as the spikelets
from the upper parts of the inflorescence tend to be smaller. Micromorphologi-
cal characteristics were assessed on 20 specimens in each population. Voucher
specimens for all the taxa are preserved in the VALA, BUAG and BUC Herbaria
collections.
Micromorphology of sterile lemmas and caryopses.—Ten mature caryopses
for each specimen were examined. Only the adaxial (ventral) face of sterile lem-
mas and caryopses was observed. Sterile lemmas and caryopses were mounted
on aluminium stubs with Avery's spot-o-glue and then coated with 20 nm of
gold using Bio-Rad Sputter-Coatter SC-500. Samples were examined with 0°
tilt at 5-15 KV ona Hitachi S-4100 Scanning Electron Microscope.
Structure of the caryopsis coat.—Mature caryopses with the glumes and
bracts removed were soaked in warm water for 1 hour. Afterwards, they were
fixed in FAA (90 ml 95% ethanol, 5 ml formalin and 5 ml glacial acetic acid) for
48 h. Caryopses were transferred to 70% ethanol, dehydrated in tertiary butyl
alcohol (TBA), and embedded in Tissuemat. Ten mature caryopses for each speci-
men were serially sectioned to 5-10 ttm, stained with safranin and fast green,
mounted in Canada balsam, and examined with standard brightfield optics
and with polarized light. A small number of caryopses belonging to each ac-
cession were soaked, fixed, embedded and sectioned without removing the
glumes and bracts. The anatomy drawings were made using a Reichart camera
lucida. The lengths of the largest fifty starch grains located in the mealy en-
dosperm from each caryopsis, from each accession were measured as well.
SPIKELETS AND LEMMATAL MICROMORPHOLOGY
Depending on the environmental conditions, an Echinochloa spp. plant can
produce 1500-22000 spikelets distributed on 9-25 panicle bearing culms. The
spikelets have 2 flowers; the lower floret is sterile consisting only of lemma and
palea. Sometimes the lower floret is staminate especiany in E.colona). The ster-
ile lemma may be awned; but this cl
lets of E.colona are unawned, ia of E.oryzoides are always awned, and those
of the other species may be awned or not. An important character, emphasised
by all authors, is the size of the spikelets. The lower glume is about 1/2 the length
of the spikelets in E. colona;1/2-3/4 in E.crus-galli, E.oryzoidesand E. hispidula,
and 1/2-3/5in E. oryzicola. The lemma (in the species examined) is becoming
indurated and difficult to remove. Micromorphological characters of the lemma
for Echinochloa include: short cells (silica cells), long cells, bicellular microhairs,
papillae and microhairs. Stomata may occasionally occur in all species. Cork
cells were not observed on the Echinochloa sterile lemmas. Silica bodies are
is relatively insigi if icant. The spike-
528 BRIT.ORG/SIDA 20(2)
Taste 1.Description of sample populations.
Species Source Accesions
Echinochloa colona Spain (VALA’*) c1,c2,¢3
Romania (BUAG?, BUC‘) c11,¢c12
Echinochloa crus-galli Spain (VALA) cs5501, c5606, cs825, cs820
Romania (BUAG, BUC) cs21,c522, 523
Echinochloa hispidula Spain (VALA) h802, h803, h805, h811.
Echinochloa oryzoides Spain (VALA) 05808, 05819, os909
Romania (BUAG) 0531, 0535, 0540
Echinochloa oryzicola Spain (VALA) Oa oad
Romania (BUAG o0a64, 0a69, 0a/0, 0a71
aS
a—Department of Botany, Polytechnic Institute Valencia, Spain.
b-Department of Botany, University of Agronomical Sciences Bucharest, Romania
c-Botanical Garden, University of Bucharest, Romania
restricted to the intercostal regions. They can be cross-shaped or dumbbell-
shaped. Within each category, 2 types can be further recognised.
Cross-shaped type (ratio length:width = LL).
CL. Silica bodies with the two endings deeply divided (Fig. le).
C2. Endings only slightly bilobed (Fig. 2b).
Dum bbell-shaped type (ratio length:width = 1:2).
D1. The two endings rounded (Fig. La, £).
D2. The two endings emarginated or bilobed (Fig. Id).
Even if one or two types of silica bodies predominate in one species, one sterile
lemma may often contain sporadically the other types as well.
Long cells are easily recognizable by their length and their sinuous mar-
gins. Long cells are associated with papillae at their distal ends, especially near
the apex of lemmas. Bicellular hairs, 30-60 1m long, are present in all species
and belong to the panicoid type (Tateoka et al. 1959; Amarasinghe & Watson
1988, Watson & Dallwitz 1992-onwards). All the trichomes observed in
Echinochloa (bicellular microhairs excepted) are more or less macroscopic. If
shorter, they are only an earlier stage in the development of macrohairs. They
tend to increase in frequency and length toward the distal parts of the lemmas.
The marginal veins usually have the longest hairs. They are somewhat shorter
or even missing from the median and lateral veins. Macrohairs are present in
the intercostal regions but they are usually shorter compared to those of the
marginal veins. In E. oryzicola, sterile lemmas often appear glabrous in the in-
tercostal regions. The macrohairs are rigid, pointed and oriented toward the
apex of the lemma.
COSTEA AND TARDIF, TAXONOMY OF AND CARYOPSIS 929
PD OS cam
Lm Wa ee
C-hj, hI Pe Pare ln bk Cn b. £ hisnidula ( In |} cn
Fic.1.Mi hol f | General view.a L
een len eE icola (scale bar 25 im), f. E. hispidula (scale
um), cd. E
bar 25 um)
7 \ t by
530 BRIT.ORG/SIDA 20(2)
Fic 7) RAS L | {1 J 7
b. silica bodies in £. crus-galli (scale bar 50 um
hispidula, f. E. oryzicola (scale bar 50 tm).
rama? 1 + rope bl wee la |} cn
=
. €-f Micromorphology of caryopsis: ¢. £. oryzoides, d. E. crus-galli,e. E.
COSTEA AND TARDIF, TAXONOMY OF AND CARYOPSIS 531
CARYOPSIS
Morphology of caryopsis (Fig. 3, a-e). Caryopsis is the dry, monospermous, inde-
hiscent fruit of grasses in which the layers of pericarp are fused with the seed
coat. The adnation between the pericarp and the seed coat starts in the placento-
chalazal region and extends toward the rest of the caryopsis (Izaguire de Artucio
& Laguardia 1987).
The size of caryopsis varies proportionally with the size of the spikelet and
therefore isalso an important differential character. The caryopsis is round, ellip-
soid to ovate, with the ventral (adaxial) face rounded and the dorsal face (abaxial)
more or less flat. The ventral face shows the axis of the embryo. The embryo is
large, about 0.5 times the caryopsis length in E. colona; almost as long as the
caryopsis in E. oryzicola and 0.6-0.7 times the caryopsis length in the other
examined species. The scutellar region is visible around the embryo axis. The
dorsal face features the basal pointed coleorhiza, which is also encircled by a
rounded scutellar region. The hilum may or may not project on the outline of
the caryopsis. When plants shed their spikelets, they land with the convex face
down, in a position that brings the embryo in the most favourable position for
germination. The glumes and sterile lemma then absorb the water necessary
for germination. When they land in water, the spikelets float until imbibition
is completed after which they sink and fall to the bottom of the water (Costea,
unpublished). The mature caryopsis coat is shiny and whitish, yellowish or
brown. The scutellar region prolonged around the embryo (on the convex, ven-
tral face), is smooth or wrinkled, and may have black spots (E. oryzicola).
Micromorphology of caryopsis (Fig. 2,c-f) is not similar in the Echinochloa
species examined. The epicarp cells are elongated ranging from 60-120 pm in
length and 10-20 pm in width. The periclinal walls are flat, except for E.
oryzicola which has concave periclinal walls (Fig. 2f). The anticlinal walls are
protuberant and undulated with the amplitude, the width at the base and the
shape of the undulations having diagnostic value. For example, in E. crus-galli
(Fig. 2d) the anticlinal walls are Q-undulated (undulations rounded, wider to-
ward the apex and narrower at the base), in E. orzyoides and E. hispidula they
are S-undulated, (Fig. 2c and e) and in E.oryzicola they are Z-undulated (Fig, 2f).
Anatomy of the caryopsis coat (Fig. 4, a-c) in the examined species is simi-
lar. The differences observed are minute and quantitative. The caryopsis coat
consists of adnate layers of pericarp, seed coat and nucellus that surround the
endosperm and embryonic axis. If the lemma is not removed, cross-sections
reveal its connivence with the caryopsis. The structure of the lemma resembles
the structure of the leaf. A 1 genous mesophyll consisting of 2-4 cell layers
can be observed between the 2 epidermis. During the early stages of develop-
ment the mesophyll cells contain chloroplasts. These cells are larger than the
epidermis cells, with thickened, sclerified cell walls at maturity (Fig 4, l-c).
532 BRIT.ORG/SIDA 20(2)
|
Fic 3. Morphology of caryopsis.a. Echinochloa crus-galli, b. E. hispidula, c. E. oryzicola, d. E. oryzoides, e. £. colona. Scale bar
1mm.
(NCTA AND CARYNDCIC 533
aoe cedars
ea ‘
K OSU CRU ass- _lw
aires ep ee
Fic.4.A y of caryopsis.a. Echinochi icola,b. £. crus-galli, c. E. oryzoides. up—upper epidermis of lemma, m-
mesophyll of lemma, Ilw-lower epidermis of lemma, ep-epicarp, ms-mesocarp, sc-seed coat, al-alleurone layer, ce-cen-
tral endosperm, sg-starch grains. Scale bar 10 um.
The veins in young lemmas resemble the tertiary veins of the leaves. The cary-
opsis coat is thin (7-15 wm thick), with a simple structure. It consists of only 2-3
cell layers. The one-layered epicarp is the most representative component of the
pericarp. The epicarp cells have thickened walls. The mesocarp is also single-
layered with smaller and often crushed cells. At caryopsis maturity the endocarp
is usually no longer visible. In younger caryopses it consists of long thin-walled
cells (“tube-cells”), parallel to the epicarp cells. The seed coat is fused with the
pericarp and it is not structured, visible only as brown line. The nucellus per-
sistsasathin remnant. A single layered alleurone i ti d the entire
endosperm. The alleurone layer can be locally 2-layered in E. oryzoides (Fig.
4c). The central endosperm consists of large, irregularly shaped cells that con-
tain simple starch grains. The central endosperm does not have a homogenous
534 BRIT.ORG/SIDA 20(2)
appearance. Around the embryo the endosperm is “mealy” and soft while in
the rest of the caryopsis is hard and glassy. In most of the species examined, the
glassy endosperm predominates. However, in E.oryzoides the mealy endosperm
isequaly or better developed than the glassy endosperm. The mealy endosperm
contains rounded, isolated starch grains; only rarely they are aggregated (but
not compound) in small groups of 2-4 granules. The hard endosperm has po-
lygonal (rarely round) starch grains, closely packed together. This last type cor-
responds to the Panicum type described by Tateoka (1962). The average size of
the starch grains is significant and it varies between 5.5 tm and 10.5 um de-
pending on the species. The embryo has a scutellar tail, without an epiblast.
SPECIES DESCRIPTION AND NOMENCLATURE
Nomenclature.—Most of the nomenclatural problems of E.oryzicola are related
tothe binomial Panicum phyllopogon on which Echinochloa phyllopogon (Stapf)
Stapf ex Koss. is based. Stapf’s diagnosis of Panicum phyllopogon addresses only
the vegetative features of the plants, particularly the presence of hair tufts in
the collar region of the leaves. For a long time this feature was erroneously con-
sidered to be an exclusive characteristic of only one species—E. phyllopogon,
thus generating extensive nomenclatural confusion. The individuals of
Echinochloa oryzicola—a more recent name—are constantly showing this veg-
etative characteristic. Consequently E. oryzicola has almost generally been con-
sidered a synonym of E. phyllopogon (Pirola 1965; Morariu 1972; Kerguélen 1975,
1993, 2002; Czerepanov 1981; Michael 1983; Chirila 1984; Ciocarlan 2000).
Kossenko (1940) further contributed to this confusion when he subordinated
E.oryzicolaasa subspecies of E. phyllopogon (1940). Other authors erroneously
considered E. oryzicola to be asynonym of E. hispidula (Ohwi 1962; Gould et
al. 1972) or even of E. oryzoides (Clayton 1980).
Stapf’s collections of Panicum phyllopogon at Kew comprise both vegeta-
tive and fertile specimens. Tufts of hairs are obvious in most of the vegetative
specimens and in several plants bearing inflorescences. Carretero (1981) noted
that tufts of hair might also occur in the leaf collar of E. hispidula (1981). After
examined Stap!’s collection, Carretero (1981) synonymized E. phyllopogon with
E. hispidula. Michael initially regarded Stapf’s collection (1975 in herb, 1983) as
a mixture of E. phyllopogon (the plants showing the vegetative characteristics)
and E. oryzoides. Later, after examining populations from Spain, Italy and
France he observed that individuals of E. oryzoides may also have leaves with
hair tufts in the collar region and accordingly he synonymized E. phyllopogon
with E. oryzoides. The author selected as a lectotype for Panicum phyllopogon
the one sheet in which the inflorescence and tufts of hairs are evident on the
same specimen (1985, note in the Kew Herbarium on the specimen selected asa
lectotye, collected by Jacometti from “Novarra, in rice fields”). We support
—
COSTEA AND TARDIF, TAAVINGUIIE UT
ANN CARYVODCIC
Michael's view in that E. phyllopogon may be asynonym of E. oryzoides and that
Stapf’s collection isa mixture of E. oryzicola and E.oryzoides(=E. phyllopogon).
Many recent studies on the biology, ecology, herbicide resistance and physi-
ology have been also using the binomial “Echinochloa phyllopogon” referring
probably to E. oryzicola (Fox & Kennedy 1994; Fox et al. 1995; Mujer et al. 1995;
Gibson et al. 1999; Fischer et al. 2000; Fischer et al. 2000).
Identification key—The following key is modified from Carretero (1981)
and Michael (1983).
In order to improve chances of correctly identifying the species, a range of
individuals from the same population and a range of spikelets and caryopses
belonging tothe same plant should be collected and examined. Spikelets length
measurements do not include the awns. Caryopsis measurements refer to dry
caryopses. Starch grains should be observed in the mealy endosperm (around
the embryo).
é
Spikelets 2-3 mm long, regularly arranged on the racemes. Caryopses whitish, 0.7—
1.2 mm long with the embryo 0.4-0.5 of the caryopsis length
E.colona
Spikelets larger, irregularly arranged on the racemes. Caryopses larger, light-yellow
or in different shades of brown, with the embryo 0.6-0.9 of the caryopsis lengtt 2
2. Spikelets 3.9-5 mm long; caryopses 2-3 mm long; starch grains 7-10 Lum in
diameter 3
2: fiona 2.8-3.7 mm long; caryopses 1.4-1.8 mm long; starch grains 5—5.6 um
in diam 4
Se aes red-purplish. Leaves ae with a tuft of brown hairs in the collar
region. Lower glume 1/2-3/5 the length of the spikelets, with the 3 veins
visible only at the base. Sterile ae often glabrous and shiny, rarely with
stiff macrohairs, up to 0.8 mm. Caryopses brownish, 2-2.4 mm long, with the
mbryo 0.9 of the caryopsis length, and the glassy endosperm equal or more
developed than the mealy endosperm E. oryzicola
3. Coleoptyle green. Leaves only rarely with a tuft of hairs in the collar region of
the leaves. Lower glume 1/2-1/3 the length of the spikelets, with the 3
visible along their entire length. Sterile lemma never glabrous and shiny, pro-
vided with dense, stiff hairs up to 1mm long. Caryopsis light-yellow, 2.2-2
mm long, with the embryo 0.6-0.7 of the oss length and the mealy
endosperm more developed than
veins
th 1€ glassy 4 | \dosper Td
4. Weeds of rice fields, occasionally with a tuft of hairs in the collar region of
the leaves. The panicle is not pyramidal with branches often whorled. Spike-
lets ovate-elliptical, 3.3-3.8 mm long. Caryopsis 2—2.2 mm long with the
scutellar zone transversely wrinkled
E. oryzoides
E. hispidula
aw
. Weeds not obligate of rice, but sometimes occurring in rice fields espe-
cially on the margins of the ponds. Leaves never with hairs in the collar of
the leaves. Panicle pyramidal with the branches never obviously whorled
Spikelets ovate, 2.8-3.4 mm. Caryopsis ovate, 1.4-1.9 mm, with the scutel
lar Zone smooth
E. crus-galli
1. Echinochloa colons ML) ink, eas ie 2:209. 1833. Type: JAMAICA?’ Antilles,
IMs}; n yst
Browne (LINN). P Nat., ed. 10, 2:870. 1759. Echinochloa crus-galli (L.)
Beauv. subs p colona (L.) Honda, Bot. oven ae yo), 37:22.1923.
536 BRIT.ORG/SIDA 20(2)
= Panicum zonale Guss., Fl. Sic. Prod. 1:62. 1827.
= Echinochloa verticillata Berhaut, Bull. Soc.Bot. Fr, 1953:11. 1954.
There is still disagreement over how to write the specific epithet, as various
authors have used either “colonum” or “colona’” to designate the species The spe-
cific epithet is most likely derived from the medieval Latin adjective “colonus-
a-um,’ in which case “Echinochloa colona” is the correct spelling. Another hy-
pothesis (Carretero 1981) is that the specific epithet results from the contraction
of the word “colonorum, the plural genitive of “colonus-i,” although we believe
this to be less probable.
Echinochloa colona is the most easily recognised species. It is annual, often
rooting at the lower nodes. The leaves may have purplish transversal bands. The
inflorescence is usually erect, with short branches (the lower ones are shorter
to equalling 3 cm long). The spikelets are regularly arranged on the racemes.
Often the lower floret of the spikelets is staminate. The lower lemma is awn-
less, not exceeding 2 mm long. The spikelets are ovate, 2-3 x 1.2-1.8 mm. The
lower glume is 3-veined, about 1/2 as long as the spikelet. The upper glume is 5-
veined. The sterile lemma is 7-veined. The median vein may or may not have
macrohairs. Longest macrohairs, | mm long. Lateral veins are only distally con-
spicuous. The silica bodies are of type D2. The stigmas are dark-red.
The caryopsis (Fig. 3e) is round to almost so, whitish and translucent, (0.7-)
0.9-1.2 x 0.7-1.1 mm. The embryo is 0.4-0.5 of the caryopsis length. The epicarp
cells are 40-65 x 8-12 um. The periclinal walls are flat. The anticlinal walls are
moderately thick 2.1-2.7 wm), undulated; the amplitude of the undulations is
5-10 um, and their width is 6-12 um. The undulations are Q-shaped or S-shaped.
Starch grains, 5.6 (+ 0.24) um long. The chromosome number is 2n = 6x = 54
(Carretero 1981; Yabuno 1985; Koul & Gohil 1991; Devesa et al. 1991).
This species is a widespread weed in rice fields in tropical and subtropical
areas of the globe. In Europe it has been recorded in the warmest countries
(Spain, Portugal, Italy, France, Turkey, Greece), where it grows as a ruderal or
segetal weed in irrigated crops (and sometimes also in rice). It seems to be ex-
panding towards typical temperate countries where it usually grows as a rud-
eral. Asan example, we are recording the occurrence of this species for the first
time in Romania where we have encountered it since 1996 growing asa ruderal
in the Railroad Station in Bucharest.
2. Echinochloa oryzicola (Vasing.) Vasing. in Komaroy, Fl. U.R.S.S. 2:33. 1934. Type:
“Oriente Extremo inter segetes Oryzae sativae L.” Panicum oryzicola Vasing., Bull. Appl. Bot
Pl.-Breed. (Leningrad) 25(4):125. 1931. ae, sen A al el seca subsp. ced
(Vasing.) Koss., Not. Syst. Herb. Acad. U S.8C.2):210.19 i : galli(L.) Beau
var. oryzicola (Vasing.) Ohwi, Acta ae wee 1 - 1942.
= Echinochloa phyllopogon auct., non Stapf
Annuals; the European populations with a tuft of brownish hairs in the collar
COCSTEA ANT CARYVNDCIC 537
}
region of the leaves. The hairs are obvious from the two or four leaf stage. Yabuno
(1962, 1981) studying Asian populations found that different forms - notably
his Cand F forms - do not always exhibit the tuft of hairs in the collar region of
the leaves. The inflorescence is normally erect and spreading but occasionally
can be horizontal or pendent and may be green to red coloured. Spikelets are
ovate-elliptical, awned (awn not exceeding 2 cm in length) or not, measuring
3.9-4.8(-5) x 2.2-2.4 mm. The spikelets persist in the panicle longer than in E.
oryzoides. The lower glume is about 1/2-3/5 the length of the spikelet, with 3
veins only partially visible. The upper glume is 5-7 veined. The sterile lemma
is 4-5 veined. The median veins may or may not have short hairs (0.5-0.6 mm);
the laterals veins are conspicuous only toward the apex. The longest macrohairs
are up to 0.8 um long. The sterile lemma can be:a) glabrous and shiny, convex—
with the lateral veins visible only along their distal part. In the intercostal re-
gions the macrohairs are rare, short, 0.1-0.4 mm, with a swollen base (Fig. 1, c
and d) (C-form of Yabuno 1962, 1981). This is the most common type encoun-
tered in Europe. b) more or less flat and coarse, with dense long macrohairs, up
to 0.8 mm in the intercostal regions (F-form of Yabuno 1962, 1981). The silica
bodies may be type C2 (Fig. le) or D2 (Fig. 1d). The stigmas are red. The cary-
opsis is ellipsoidal to almost round measuring 2-2.4 x 1.8-2.1 mm (Fig. 3c). The
embryo is 0.75-0.9 of the caryopsis length. Often the stigmas are persistent. The
hilum is prominently visible on the outline of the caryopsis. The colour is
brownish-red, brownish-green or brownish-grey. The scutellar zone is irregu-
larly wrinkled often with black spots. The epicarp cells are 100-120 x 15 um.
The periclinal walls are concave (Fig. 4a). The anticlinal walls are moderately
thick (2.6-3.2 um) and undulated; the amplitude of the undulations is 8-10 um,
and their width is 6-7 um (Fig. 2f). The undulations are Z-shaped (acute and
narrow toward the apex) (Fig. 2f). The glassy endosperm predominates or it is
as developed as the mealy endosperm. Starch grains are 7.14 (+ 0.51) um in
length. The chromosome number is 2n = 4x = 36 (Carretero 1981; Yabuno 1985,
1996).
This species originated in China and SE Asia and is now a weed of rice in
Europe, Asia, North and South America. The biology and ecology of a culti-
vated form of E. oryzicola has been recently described by Hirosue et al. (2000).
3. Echinochloa crus-galli (L.) Beauv., Ess. Agrost.:53, 161, tab. 11, fig. 2. 1812. Tver:
“Europae, Virginiae cultis” (LINN). Panicum crus-galli L., Sp. PL, ed. 1:56. 1753. Milium crus-
galli (L.) Moench, Meth:202. 1794.
The herbarium sheet number 80.18 (LINN) contains a specimen collected by Kalm in
Canada, annotated by Linnaeus himself as Panicum crus- sa mas: al eo (Hitchcock
1920; Baum 1967; Carretero 1981; Michael 1983) observed that thi s not belong
to E. crus-galli. The same situation occurs with the other two Eieats li abelted: Panicum crus-
galli (80.19 and 80.20) by Linnaeus. According to Carretero, only the specimen 80.17 labelled
by Linnaeus as Panicum crus-cor ds to E. crus-galli, Ali 1968) and Gould et al.
(1972) designated a neotype. Michael (1983) designated as a lectotype the specimen |: 303
538 BRIT.ORG/SIDA 20(2)
from Burser’s Herbarium at the University of Uppsala lin order to retain the specific epithet,
Baum suggested the same solution but he did not select a lectotype (1967).
= Panicum crus-corvi L., Sp. PL. ed. 2, 84. 1762.
Annuals with erect to spreading or decumbent stems, up to 120 cm. The panicle
is often pyramidal with purplish nuances, erect, but sometimes nodding. The
inflorescence branches are longer than 3 cm, patent or erect-patent, never ob-
viously whorled. The lower branches are frequently twice branched. The spike-
lets are irregularly arranged on the racemes and the lower floret is rarely stami-
nate. Spikelets are ovate to broadly-ovate, 2.8-3.4 x 1.6-1.8 mm. The lemma of
the lower floret isawnless or awned, the awn not exceeding 5cm in length. The
lower glume is 3-veined and about 1/2-1/3 the length of the spikelet. The up-
per glume is 5-7 veined. The sterile lemma is 7-veined. The lateral veins are
visible along their entire length or only toward the apex. The longest macrohairs
are about | mm long. The silica bodies belong to types C2 or D2 (Fig. 2b). The
stigmas are white or red.
The caryopsis is ovate, 1.4-1.9 « 1.3-1.6 mm, brownish-red or brownish-grey
in colour. The embryo is 0.6-0.7 of the caryopsis length (Fig. 3a). The epicarp
cells are 60-80 x 10-15 um. The periclinal walls are flat. The anticlinal walls
are thick (2.6-3.2 um) and undulated; the amplitude of the undulations is 8-10
yum, and their width is 6-7 um. The undulations are Q-shaped (Fig. 2d). The
glassy endosperm predominates. The starch grains are 5.9 um (+ 0.37) in length
The chromosome number is 2n = 6x = 54 (Carretero 1981).
This species is probably the most widespread of the genus, being a com-
mon and noxious weed all over the world, especially in irrigated crops. It is less
limited to warm climate compared to E. colona and E. hispidula. In rice it usu-
ally grows on the pond margins, but it may also penetrate the interior of the
rice fields.
4. Echinochloa hispidula (Retz.) Nees ex Royle, Ill. Bot. Himal. 11:416, 420.1840.
Type: “India Orientali,” Konig (LD). Panicum hispidulum Retz., Obs. Bot. 5:18.1789. Echinochloa
crus-galli (L.) Beauv. var. eae = ) aeeree Bot. Mag. (Tokyo) 37. 12? 1923. Echinochloa
crus-galli (L.) Beauv. subsp. his} z.) Honda, J. Fac. Sci. Tokio Univ, Bot. 3:267. 1930.
= Echinochloa phylloryzoides Novelli, Giorn. Risic. 2:306 (1912).
= Echinochloa erecta (Pollacci) Pignatti, Arch. Bot. (Italy) 31, ser. HI, 15(1):2. 1955. Echinoch
galli (L.) Beauv subsp. erecta (Pollaci) Ciferri @ Giacomini Nomencl. FI. Ital. 1:20. 1950.
Annuals which may occasionally have tufts of hairs in the collar region of the
leaves. The panicle is rarely pyramidal, erect to pendent, green or with purplish
nuances. The branches are whorled and more or less erect, except for the lower-
most ones. Spikelets are ovate-elliptical of 3.3-3.6(-3.8) x L.7-L8 (-2) mm. The
lower glume is 3-veined, about 1/2-1/3 of the spikelet length, and the upper
glume is 5-7 veined. The sterile lemma is 7-veined resembling EF. crus-galli. The
longest macrohairs are about 0.8 mm long (Fig. 1b). The silica bodies are very
COSTEA AND TARDIF, TAXONO : AND CARYOPSIS 539
variable, depending on the population - Cl (h802) (Fig. 1b), Cl and D1 (h803,
h805), C2 and D2 (h805). The stigmas are white but may occasionally be red.
The caryopsis is larger that of E. crus-galli, ovoid to oblong, 2-2.2 x 1.5-18
mm and brownish-red or brownish-grey in colour (Fig. 3b). The embryo is 0.6-
0.7 the caryopsis length, and the scutellar zone is transversally wrinkled (Fig.
3b). The hilum is projecting on the outline of the caryopsis. The epicarp cells
are 65-90 x 10-15 um. The periclinal walls are flat. The anticlinal walls are thick
(2.6-3.2 um) and S-undulated; the amplitude of the undulations is 8-10 um and
their width is 6-6.5 pm (Fig. 2e). The glassy endosperm predominates. The starch
grains are in average 6.3 (+ 0.47) um in length. The chromosome numbers re-
ported are: 2n = 6x = 54 (Devesa et al. 1991) or 2n = 36 (Carretero 1981; Feng &
Zhang 1993).
This taxon probably originated in the SE Asia. It is widespread in tropical
and temperate areas of Asia, Africa, America and Australia. In Europe it grows
only in the warmest regions (Spain, Italy, Portugal and France) asa weed of rice
fields.
5. Echinochloa oryzoides (Arduino) ee Verh. Zool.-Bot. Ges. Wien 41: 742.
1891.
TYPE: “Semina hujus Panici inventa a me fuere inter Oryzam” Panicum oryzoides
Arduino, Animady. Bot. Spec. Alt. 16, tab tien Panicum crus-galliL. var. oryzoides (Arduino)
Fiori, Nuov. FL Anal. Ital. 1:79. 1923.
= Echinochloa crt te L.) Beauv. subsp. oryzoides Bolos & Masclans, Collect. Bot. 4:420. 1955.
= Panicum hostii Bieb., Fl. Taur. Cauc. 3:57. 1819. Echinochloa crus-galli (L.) Beauv. subsp. hostii
(M. Bieb.) K. Richter, al Eur. 1:26. 1890.
= Panicum phyllopogon Bane Hook, Ic. Plant., ser. 4, plate 2698. 1901. E ee phyllopogon
(Stapf) Stapf. ex Koss., Not. Syst. Herb. Acad. U.R.SS. 8 (12):208. 194
= Ec Pees macrocarpa Vasing. in Komaroy, Fl. U.R.S.S. 2:739. ou Ee hinochloa crus-galli L.
Beauv. var. macrocarpa (Vasing.) Morariu in Savulescu, Fl. Rom. 12:86. 1972
= Echinochloa coarctata Koss., Not. Syst. Herb. Acad. U.R.S.S. 9(1):28. 1941.
= Echinochloa commutata Schultes in Roemer & Schultes, Syst. Veg., ed. 15, Mant. 2:267. 1824.
Annuals with erect to spreading or decumbent stems, up to 130 cm tall. The
seedlings have a reddish coleoptyle. Tufts of hairs in the collar region may be
infrequently present. The inflorescence is green, pendent and at maturity hang-
ing almost horizontally, resembling rice in habit. The branches of the panicle
are often adpressed to the main rachis. The spikelets are rather persistent, usu-
ally awned (awn up to 5cm long), broadly-ovate to ovate and (3.6-)4.1-5 x 2.2-
2.6 um. The lower glume is about 1/2-1/3 the length of the spikelet, with 3 veins
visible across their entire length. The upper glume has 5 veins also entirely vis-
ible. The sterile lemma is 7-veined, with the lateral veins usually visible along
their entire course. The longest macrohairs are about | mm long (Fig. la). The
silica bodies belong to the type D1 (Fig. la). The stigmas are red.
The caryopsis is ovate to almost round, 2.2-2.8 x 1.9-2.3 mm (Fig. 3d) and
yellowish. The embryo is 0.6-0.7 the caryopsis length, with the scutellar region
540 BRIT.ORG/SIDA 20(2)
zone smooth (Fig. 3d). The epicarp cells are 100-120 x 10-15 um. The periclinal
walls are flat. The anticlinal walls are moderately thick (2-2.6 um) and weakly
S-undulated (Fig. 2c). The amplitude of the undulations is 1-3 ym, and their
width is 8-11 um. The mealy endosperm predominates. The starch grains are
9.5 um (+ 0.58) in length. The chromosome numbers reported are: 2n = 36
(Carretero 1981) and 2n = 6x = 54 (Yabuno 1984; Feng & Zhang 1993).
The species is a common weed in rice occurring in many countries from
Asia, Europe, North America, South America and Australia. It is probably one
of the most widespread species after E. crus-galli and E. colona.
DISCUSSION
Characters
The genus Echinochloa does not have qualitative characters which clearly de-
marcate its species. A good example is the presence of hair tufts in the collar
region of the leaves which has been considered an exclusive feature of E.
phyllopogon. This assumption has subsequently caused a widespread nomen-
clatural and taxonomic confusion because as we have previously indicated, three
species may show this feature (E. oryzicola commonly; E. hispidula and E.
oryzoides rarely). Even the most significant qualitative character states overlap
between species. When such characters are noted they are always accompa-
nied by the words “often,” “usually,” “normally,” etc. because exceptions are pos-
sible. The occurrence of a feature should be understood as a predilection and
not as an exclusive characteristic.
Lemmatal micromorphology.—The long cells, microhairs and papillae are
the same in all species examined. The pattern of papillae in Echinochloa spe-
cies is similar to the pattern described in the fertile lemmas of some Panicum
species (Clark & Gould 1975). The macrohairs are more or less similar in the
examined taxa. Shorter hairs that could be described as prickles (Metcalfe 1960;
Ellis 1979) were observed in all species. However, because these shorter hairs
represent a developmental stage, only the macrohairs are recognised in this
study as a distinct category (see Snow 1996, 1998 for considerations on homol-
ogy and ontogeny of hairs in Poaceae). Variations in length, distribution and
density of macrohairs are minute and there is considerable overlapping among
species. The only exception is the short, rare macrohairs, with swollen bases
from the intercostal regions of sterile lemmas in E. oryzicola (in the “form C” of
Yabuno 1962, 1981). These hairs are constant in their morphology and can be
considered intermediate to prickles (Metcalfe 1960; Ellis 1979). Silica bodies have
previously been considered structures of taxonomic significance in Poaceae
(Metcalfe 1960; Ellis 1979; Palmer & Tucker 1981; Evoli & Pirola 1971). In
Echinochloa, silica bodies vary in shape even within a small area of the same
lemma. However, in most species there is a tendency toward one or two types.
Echinochloa hispidula was the most variable of all species examined in this
—
COSTEA AND TARDIF, TAXONOMY OF AND CARYOPSIS 541
study. The silica bodies varied from population to population, with all the dif-
ferent types observed. The previous epidermis studies conducted on leaves
(Sanchez 1968; Pirola & Evoli 1971; Carretero 1981: Jin et al. 1986) are rather con-
tradictory but they all showed an even greater variation of this character. Even
if the results obtained in this study revealed some differences between species
(E. hispidula excepted), the shape characteristics of the silica bodies should be
regarded with caution. More populations need to be examined before a final
conclusion is reached. The same cautionary approach towards the taxonomic
utility of the silica body shape is suggested by other studies conducted on
Zizania (Terell @ Wergin 1981), Oryza (Whang & Kim 1994) and Leptochloa
(Snow 1996). Ball et al. 1999) showed that silica phytoliths examined individu-
ally are taxonomically irrelevant. The authors instead used detailed
morphometrics, computer-assisted imagery and statistical analysis to develop
a classification key of several Triticum and Hordeum species. The presence or
absence of silica bodies in lemmas can prove useful at higher ranks (genera
and above) since their presence is probably symplesiomorphic (Snow 1996).
Caryopsis.—The simple morphology of the caryopsis provides good char-
acters for species identification. The size of caryopsis is important and corre-
lated with the size of the spikelets. The other characters overlap between spe-
cies but may be useful diagnostics is some cases, eg: caryopsis whitish,
translucent (E. colona), yellowish (E. oryzoides), embryo 0.4-0.5 of the caryopsis
length (E. colona) or 0.75-0.9(E. oryzicola). Surprisingly, surface patterns of the
caryopsis proved to be significant in the differentiation between almost all
Echinochloa species. However, more populations should be examined before any
definite conclusions can be reached.
The anatomy of the caryopsis is of minimal systematic value. The overall
organization of the caryopsis coat resembles the structures described in Setaria
(Rost 1973) or Melica (Rost and Izaguire de Artucio 1990). The structural sim-
plicity of the pericarp can be a functional result of the indurated lemma that
protects the caryopsis. In those grasses where no coalescence between the cary-
opsis and lemma occurs, or when the lemma is thin, the pericarp is more com-
plex structurally (as for example in many Iriticoideae—Netolitzky 1926; Avery
1930; Krauss 1930; Bradbury et al. 1956; Ungurean & Costea 1994). The anatomy
of the sterile lemmas in Echinochloa species could be more important as a taxo-
nomic character than the structure of the caryopsis coat. The structural fea-
tures and surface pattern of the lemma indicate its ontogenetic connection with
the leaves.
The average size of the starch grains examined in a large number of en-
dosperm cells and caryopses is constant and significant among the species ex-
amined and therefore can be considered a diagnostic feature. This character is
as significant in the taxonomy of the genus as other quantitatively important
characters such as the size of spikelets, and size of caryopses.
542 BRIT.ORG/SIDA 20(2)
Relationships between species
The individuality of Echinochloa species as treated in the present study is ap-
parently reinforced by studies of their biology and ecology (Holm et al. 1977,
1997, Szilvassy 1976; Chirila 1967; Carretero 1981; Yabuno 1983; Norris 1996;
Honek & Martinova 1996). The appropriate understanding of E. oryzicola was
delayed by nomenclature problems since this binomial has been almost gener-
ally considered a synonym of E. phyllopogon (Pirola 1965; Morariu 1972,
Kerguélen 1975, 1993, 2002; Czerepanov 1981; Michael 1983; Chirila 1984,
Ciocarlan 2000). Apart from this nomenclatural confusion E. oryzicola is the
second species easiest to differentiate (after E. colona) even using classic mor-
phologic characters. It has the second largest (after E. oryzoides) spikelets, cary-
opses and starch grains; the micromorphology of the sterile lemma and cary-
opsis are quite peculiar as well.
Yet apart from E. colona and E. oryzicola, which are usually easily
recognisable, the eae 3 taxa comprise a cages oe Echinochloa
hispidula is extremel ly variable and shares clos ies with both E. oryzoides
and E. crus-galli. Several current treatments tend to view E. hispidula asa sub-
species or as a variety of E. crus-galli (Michael 1983; Mateo Sanz 1990; Devesa
1991; Janzein 1993; Mateo Sanz & Crespo Villalba 1995; Kerguélen 1993, 2002;
Asins et al. 1999). However, Gonzales- Andres et al. (1996) studying the izoenzyme
variation of these species, reached another conclusion: only 3 species—E. colona,
E. crus-galli and E. oryzicola—were clearly defined. Echinochloa hispidula and
E. oryzoides clustered together, and the variability between the populations of
E. hispidula was higher than the variability between both taxa. Therefore, the
authors suggested that the most appropriate classification would be E. hispidula
as a subspecies of E. oryzoides. Asins et al.(1999) studying the morphologic and
isozyme patterns of variation of the same species, found only FE. colona and E.
oryzicola to be distinct from both morphologic and molecular point of view.
Based on their morphology, the populations of E. crus-galli, E. hispidula and E,
oryzoides congruently clustered together within each species. However, based
on the isozyme variation, there was a considerable overlapping between spe-
cies suggesting a high degree of genetic variation. Consequently, Asins et al.
(1999) advocated that E. hispidula and E. oryzoides as infraspecific taxa of E.
crus-galli would be the most appropriate classification. We consider that in
order to reach a final conclusion more populations worldwide should be
analysed using combined molecular (such as RAPD, AFLP. ISSR and DNA fin-
gerprints) and morphologic methods, and the data resolved in a cladistic ap-
proach involving all characters. The enormous variation observed makes the
boundaries between these taxa rather uncertain and in order to avoid an arbi-
trary classification, we maintained each taxa at specific level. In Echinochloa
the amplitude of morphologic variation can serve as a basis for taxa differen-
COSTEA AND TARDIF, TAXONOMY OF AND CARYOPSIS 543
tiation. Each taxon has a distinct morphologic core and the variation—statisti-
cally understood—can serve to their delimitation.
ACKNOWLEDGMENTS
I owe many thanks to Jose Luis Carretero for the useful discussions and the
material from Spain. I am also very grateful to Julio Iranzo who kindly made
possible the SEM study. Our sincere thanks go also to Peter Michael and WD.
Clayton who reviewed an earlier version of the manuscript providing valuable
comments and suggestions. Many thanks goalso to Cheryl Corbett for the help-
ful criticism.
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SEEDAND FRUIT CHARACTERS IN SELECTED
SPERMACOCEAE AND COMPARISON
WITH HEDYOTIDEAE (RUBIACEAE)
Edward E. Terrell Richard P Wunderlin
Research Collaborator Institute for Systematic Botany
Depa ens oh Systematic Biology-Botany Department of Biology
Natural Hi , smithsonian Institution University of South Florida
Weshin han DC 20560, U.S.A Tampa, FL 33620-5200, U.S.A
ABSTRACT
Seed and fruit morphology of ten species of five American genera of Spermacoceae (Diodia, Ernodea,
Mitracarpus, Richardia, Spermacoce) are described, illustrated, and compared with that in the
Hedyotideae. These data support the taxonomic recognition of two distinct tribes.
ESUMEN
l -f >] 1 aT ARG | 7: . z $ peer
Tita
la
se décciibe; se ilustra, y se nce acon la de edyati deae. . stos datos apoyan el reconocimiento
taxonomico de dos tribus distintas.
INTRODUCTION
Seed morphology in Houstonia L. and related genera in the tribe Hedyotideae
Cham. & Schltdl.ex DC. (Rubiaceae) has contributed valuable information, es-
ecially when combined with data from chromosome number and pollen struc-
ture (Terrell et al. 1986; Terrell 1996). More recent taxonomic studies have con-
tinued to emphasize the importance of seed data (Terrell 2001la, 2001b, 2001c).
This study investigates the morphology of the seeds and fruits in selected
genera in the tribe Spermacoceae Bercht. & J. Presl. We contrast this informa-
tion with previously acquired seed data for the Hedyotideae. The two tribes in-
clude some superficially similar genera and species (e.g. Oldenlandia L. and
Spermacoce L.), but structure and development of their fruits and seeds are fun-
damentally different and deserve further study. The Hedyotideae have few to
many small seeds per locule, nearly always in subglobose biloculate capsules.
In contrast, the fruits of Spermacoceae have one large seed per locule and are
generally more structurally complex and vary considerably among the genera.
Bremer and Manen (2000) expanded the tribe Spermacoceae to include the
Hedyotideae; however, our data from seed and fruit morphology in five genera
do not appear to support this inclusion.
Verdcourt (1958) proposed a classification of the Rubiaceae in which he
recognized three subfamilies. The Rubioideae, the largest subfamily, contained
17 tribes including the Hedyotideae and Spermacoceae. Bremekamp (1966)
SIDA 20(2): 549 — 557, 2002
550 BRIT.ORG/SIDA 20(2)
—
modified Verdcourt’s system and recognized eight subfamilies in the family and
19 tribes in the Rubioideae (including the Hedyotideae and Spermacoceae).
Verdcourt (1976), still recognizing his original three subfamilies, reduced the
number of tribes in the Rubioideae to 12 and maintained Hedyotideae and
Spermacoceae. Robbrecht (1988, 1993) recognized four subfamilies and 19 tribes
in subfamily Rubioideae. The latter included Hedyotideae and Spermacoceae.
MATERIALS AND METHODS
Scanning electron microscopy was employed in this study of seeds and fruits.
The investigation was carried out at the Biology Department, University of
South Florida, using a JEOL JSM-35 microscope operating at 15 kilovolts. Ten
species in five genera were examined from herbarium material at the University
of South Florida (USF) (Table 1). The species examined were those most readily
available and representative of species native to tropical America, especially of
the Caribbean basin. Tables | and 2 provide nomenclatural author’s names.
RESULTS
The Spermacoceae is a pantropical tribe that may consist of as many as 19 gen-
era and 450 species. Its generic delimitations are a matter of dispute. Eleven
currently recognized genera native to the New World are summarized in Table
2.Morphological characters of the seeds and fruits of five genera of Spermacoceae
native to the Americas are described as follows:
Diodia: Ovary 2(3-4)-locular; ovules solitary in each locule, attached to
the middle of the septum. Fruit with 203-4) indehiscent mericarps, these fleshy
or dry capsules. Seed enclosed within the hardened mericarp (accessible only
by dissection), oblong, somewhat compressed, the dorsal surface convex, the
ventral surface with a slightly sunken, central, longitudinal groove (see com-
ments in Discussion), the testa finely reticulate (Fig. lA-D).
Ernodea: Ovary 2-locular; ovules solitary in each locule, attached to the
middle of the septum. Fruit fleshy, oblong, becoming hardened in drying, not
easily separated into indehiscent mericarps (accessible only by dissection).
Seeds oblong, somewhat compressed, the dorsal surface convex, the ventral sur-
face witha slightly sunken, central, longitudinal groove, the testa finely reticu-
late (Fig. 3C,D)
Mitracarpus: Ovary 2(3-4)-locular, ovules solitary in each locule, attached
near the middle of the septum. Fruit a thin-walled circumscissile capsule, the
distal portion falling away with the calyx limb, the septum usually persistent.
Seed oblong to globose, the dorsal surface convex, the ventral face divided into
4 distinct areas by an x-shaped groove, the testa finely reticulate (Fig. 3A,B).
Richardia: Ovary (2-)3-4(-6)-locular, ovules solitary in each locule, at-
tached near the middle of the septum. Fruit dehiscing into dry mericarps, the
ventral mericarp face with either a medial keel or narrow groove. Seed filling
TERRELL AND WUNDERLIN ey
Table 1. List of species and collections examined for this study.
Species Source/Voucher Information
Diodia teres Walter Florida. St. Johns Co.: Hansen & Hansen 9844 (USF)
Diodia virginiana L. Florida. Collier Co.: Lakela 27450 (USF)
Ernodea littoralis Sw. Florida. Monroe Co.:Gann & Bradley 449 (USF)
Mitracarpus hirtus (L.) DC. Florida. Lake Co.: Daubenmire & Daubenmire s.n., 18 Oct 92
(USF
Richardia brasiliensis Gomes Florida. ee a Rates )
Richardia humistrata (Cham. & Florida. Escambia rkhalter 7877 (USF)
Schltdl.) Schult. & Schult.
Richardia scabra L. Florida. Hillsborough Co.: Robinson s.n., 14 Jul 77 (USF)
Richardia tricocca (Torr. & A. Mexico. Hidalgo: Herndndez M. & Vdsquez 3466 (USF)
Gray) Stand.
Spermacoce assurgens Ruiz & Pav. Florida. Pinellas Co.: Fleming 3862 (USF)
Spe nGcore prostrata Aubl. Florida. Collier Co.: Lakela 31721 (USF)
Spermacoce tetraquetra A. Rich. Florida. Miami-Dade Co.: Bradley 1999 (USF)
Spermacoce verticillata L. Florida. Dade Co.: Avery 589a (USF)
Taste 2.Eleven currently recognized genera of Spermacoceae native to the Americas, the estimated
number of species, native distribution, and reference to a recent taxonomic treatment
Crusea Cham. & Schltdl. 13 spp. a hoea rn U.S., Mexico; Anderson 2.
Diodia L.5 spp. rth America, Mexico, Contel America, West Indies, and
su America; Bacigalupo & Cabral 1999.
Emmeorhiza Pohl ex Endl. 1 sp. azil.
Ernodea Sw.4 spp. oe West Indies, Belize, a and Guatemala;
egron-Ortiz and Hickey 19
Galianthe Griseb. ca. 50 spp. Central America and South en 1991,
Mitracarpus Zucc.ca.30 spp. Mexico, Central America, and South America.
Psyllocarpus Mart. & Zucc. 8 spp. Brazil; Kirkbride 1979
Richardia L. 15 spp.; Mexico, Central erie South America; Lewis and Oliver
1974.
Scandentia E.L. Cabral & Brazil; Cabral & Bacigalupo 2001.
Bacigalupo. 4 spp.
Spermacoce L.ca. 150 spp. North America, Mexico, Central America, West Indies, and
South America; [Including Borreria G. Mey.; Hemidiodia K.
hum]
Staelia Cham. & Schltdl. 15 spp. Brazil.
the mericarp tightly (sect. Richardia) or slightly smaller than the mericarp (sect.
Asterophyton (K.Schum.) W.H.Lewis & R.L.Oliv.), oblong or broadly ovate, the
dorsal surface convex, the ventral surface with a slightly sunken, central, lon-
gitudinal groove, the testa finely reticulate (Fig. 3E,F;4A-F)).
Spermacoce: Ovary usually 2-locular, ovule solitary in each locule. Fruit
BRIT.ORG/SIDA 20(2)
f seed 1 by SEM. Diodia teres (A-B), D. virginiana (C-D), Spermacoce prostrata
Sources of collections listed in Table 1
—
. f). Scale bars are 1000 microns (A-B), 100 aaiemne( F).
dehiscing into 2 mericarps, both opening to release the seed (sect. Borreria (G.
Mey.) Verdc.) or one mericarp opening to release the seed while the other re-
mainsclosed by a fragile, easily removable septum (sect. Spermacoce). Seeds are
) g ) [
ith
ellipsoidal, ovoid, or oblong, the dorsal surface convex, the ventral surface wit
ZTOOVe eet to the ends, this sometimes
a slightly sunken, longitudinal g
elaiosomes, the testa reticulate (Fig. 1E,F; 2A-F).
with smal
DISCUSSION
1e broad sense, while others
Some workers continue to treat Spermacoce in t
divide it into several genera. Borreria is treated either as a separate genus or as
a section of Spermacoce based on fruit dehiscence (capsule with both carpels
opening to release the seeds in Borreria or a capsule with | carpel opening to
release the seed and | remaining closed). Based on our study of seed morphol-
ogy, Borreria does not appear to warrant recognition as a genus. Recently
TERRELL AND WUNDERLIN, SEED 553
r hart £ J ° L ChA en
Fic. 2
(E-F
(A-RB) S tetraquetra (C—D),S. assurgens
. Scale bars are 100 microns. Sources of collections listed in Table 1.
Galianthe was resurrected as a segregate genus from Borreria (Cabral 1991) and
Scandentia was described (Cabral & Bacigalupo 2001). Both Galianthe and
Scandentia are distinguished from Borreria primarily in habit with more or
less lax, thyrsoid, apical inflorescences in contrast to the congested, apical and/
or axially glomeruliform inflorescences of Borreria s. str. Emmeorhiza, a mo-
notypic South American genus closely related to Borreria, Galianthe, and
Scandentia has an umbelliform inflorescence. The seeds of Spermacoce
assurgens, S. prostrata, and S. verticillata, which would be placed in Borreria
on the basis of fruit dehiscence, do not differ significantly from that of S.
tetraquetra traditionally placed in Spermacoce (Fig. LE,F; 2A-F). Further work
on the Spermacoce complex is needed and is underway by various other work-
ers (e.g., E.L. Cabral, N.M. Bacigalupo, and S. Dessein).
Diodia also presents similar problems in circumscription. Hemidiodia, a
monotypic genus with 2 indehiscent mericarps and long considered closely
related to Diodia, was transferred to Borreria subg. Dasycephala (DC.)
nd
m+
554 BRIT.ORG/SIDA 20(2)
Fic. 3D | J lug £ j 1 . . Ahi COAA AA; eg i-/A B), Ernodea littoralis
seeds (C—D), Richardia brasiliensis mericarps (E-F). Scale bars are 100 microns (A—B),1000 microns (C—F). Sources of
collections listed in Table 1.
Bacigalupo & E.L.Cabral by Bacigalupo and Cabral (1996). Another study by
Bacigalupo and Cabral (1999) defined Diodia as comprised of only five Ameri-
can species; the other species previously referred to the genus are transferred to
Galianthe or Borreria. These workers exclude Diodia teres and related species
from Diodia s. str. The placement of these species is still under study by
~—
_—
Bacigalupo and Cabral. The seed morphology is markedly different between
the two southeastern U.S. species, Diodia virginiana and Diodia teres, exam-
TERRELL AND WUNDERLIN, ne
ined in this study (Fig. |A-D). The seed of D. virginiana is oblong with an equally
rounded apex and base. The testa has more or less isodiametric cells. In con-
trast, that of D. teres is oblong with an apical projection and a truncate base. On
the ventral surface, the base and sides are somewhat enrolled onto the longitu-
dinal groove. The cells of the testa surface are elongate (2-3 times as long as
wide). Further study by other workers may prove D. teres and related species to
be distinct at the generic or subgeneric level.
The seeds of the four Richardia species examined are fairly uniform, differ
only slightly in size and shape, and the genus appears to be a natural assemblage.
Ernodea seeds and fruits are most similar to those of Diodia virginiana.
The seeds of Mitracarpus are unique among those examined in this study.
The distinct x-shaped groove and the circumscissile dehiscent capsule, both
features unique in the tribe, suggest that it may be a distinct subtribe.
e Hedyotideae is much simpler in seed morphology than is the
Spermacoceae, but has much more variation in seed shape and size. The seeds
are typically in subglobose capsules, each bearing few to many seeds borne on
complexly structured placentas. The variation shown by the seeds of the vari-
ous taxa is very great and each genus, subgenus, or section often has its own
particular seed shape. The following are some examples of seed diversity found
in the Hedyotideae: (1) Houstonia L. has crateriform seeds, each with a ventral
subglobose cavity (subg. Houstonia) or with a hilar ridge in a ventral shallow
depression (subg. Chamisme Raf. sect. Amphiotis (DC.) Terrell) or with a hilar
ridge in a ventral boat- or cup-shaped depression and with other complex struc-
ture (subg. Chamisme sect. Ericotis Terrell) Terrell) (Terrell et al. 1986, Terrell
1996); (2) Oldenlandia L. typically has very small trigonous seeds, 50-100 or
more per capsule (Terrell 1996); (3) the Hedyotis fruticosa L. group has dorsiven-
trally compressed seeds with a short to long raised hilar ridge (Terrell 1996),
and (4) Stenotis Terrell and Stenaria (Raf.) Terrell have ellipsoid seeds with a
central punctiform hilum (Terrell 2001a, 2001b).
The Spermacoceae, in contrast, have one seed per locule, and the locules
develop into varied mericarps. The complexity occurs in the mericarps as well
as in the seeds; the seeds examined here do not seem to exhibit much variation,
although the Mitracarpus seed is an exception. In the Spermacoceae, some
mericarps are hardened and indehiscent (e.g., Diodia), others are fleshy
(Ernodea), and some open widely (e.g., Spermacoce). Each genus often has its
characteristic kind of mericarp, and its structure is important in classification
(e.g., Richardia, Fig. 4A,B). The mericarps may be more important in classifica-
tion than the seeds (e.g., our figures show the similarity of seeds among the vari-
ous genera). The presence of such varied mericarps contrasts with the mor-
phology of the Hedyotideae, which have no such seed covering, just the bare
seeds attached to a multi-branched placenta.
An additional seed feature not found in the Hedyotideae is the ventral
556 BRIT.ORG/SIDA 20(2)
Fic.4 n | J lor £ J J . | H Yh,,CCRA Dj-b 4° } H I (A B),R. humistrata
seeds (C—D) and R. tricocca seeds (E-F). Scale bars are 1000 microns (A—B), 100 microns (C—F). Sources of collectior
listed in Table 1.
(adaxial) longitudinal groove which resemblesa raphe. This structure was des-
ignated as a strophiole by Kirkbride (1979), Bacigalupo and Cabral (1996), and
other South American botanists. The structure and terminology of this appar-
ent raphe needs further study. It occurs on all seeds of Spermacoceae that we
have examined. The Hedyotideae differ in having a so-called hilum, a puncti-
form scar or a more elongated ridge-top scar on the seed.
Although our evidence from five selected genera represents a limited sur-
vey of the possibly 19 genera of Spermacoceae, it is sufficient to lead us to seri-
ously question the union of the Spermacoceae and Hedyotideae.
ACKNOWLEDGMENTS
We thank Betty Loraamm, Electron Microscope Manager, and Clinton J. Dawes,
Biology Department, University of South Florida for providing SEM data and
TERRELL AND WUNDERLIN ey
advice on SEM matters. Joseph Kirkbride and John Wiersema contributed valu-
able reviews. Piero Delprete provided early ic advice. Paul Peterson and
Pedro Acevedo contributed the Spanish translation.
REFERENCES
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Bacicaturo, N.M. and E.L. Carat. 1996. Infrageneric classification of Borreria (Rubiaceae-
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Bacicaturo, N.M. and E.L. Casral. 1999. Revisidn de las especies Americanas del género
Diodia (Rubiaceae, Spermacoceae). Darwiniana 37:153-165.
Bremekame, C.E.B. 1966. Remarks on the position, the delimitation and the subdivision of
the Rubiaceae. Act. Bot. Neerl. 15:1-33.
Bremer, B. and J.-F. MANeN. 2000. Phylogeny and classification of the subfamily Rubioideae
(Rubiaceae). Pl. Syst. Evol. 225:43-72.
Casral, E.L. 1991. Rehabilitacidn del género Galianthe (Rubiaceae). Bol. Soc. Argent. Bot.
27:235-249,
Casrat, E.L. and N.M. Bacicaturo. 2001. Scandentia, nuevo género de Rubiaceae-
Spermacoceae. Darwiniana 39:29-41.
KirKeride, JH. 1979. Revision of the genus Psyllocarpus (Rubiaceae). Smithsonian Contrib.
Bot.,no.41.32p.
Lewis, W.H. and R.L. Ouver. 1974. Revision of Richardia (Rubiaceae). Brittonia 26:271-301.
Necron-Ortiz, V, and R.J. Hickey. 1997.The genus Ernodea (Rubiaceae) in the Caribbean Ba-
sin: 2. Morphological analyses and systematics. Syst. Bot. 21:445-458.
Ropsrecut, E. 1988. Tropical woody Rubiaceae. Appendix 4:235-250. Nationale Plantentuin
van Belgie, Meise.
RossrecHr, E. 1993. Supplement to the 1988 outline of the classification of the Rubiaceae.
Index to genera. Opera Bot. Belg. 6:1 73-196.
Terre, E.E. 1996. Revision of Houstonia (Rubiaceae-Hedyotideae). Syst. Bot. Monogr. 48:1-118.
Terrett, E.E.2001a. Taxonomy of Stenaria (Rubiaceae-Hedyotideae), a new genus includ-
ing Hedyotis nigricans. Sida 19:591-614.
Terrett, E.E.2001b.Stenotis (Rubiaceae),a new segregate genus from Baja California, Mexico
Sida 19:899-911.
Terrett, E.E. 2001. Taxonomic review of Houstonia acerosa and H. palmeri, with notes on
Hedyotis and Oldenlandia (Rubiaceae). Sida 19:913-922.
TerRELL, E.E., W.H. Lewis, H. Rosinson, and J.W. Nowicke. 1986. Phylogenetic implications of di-
verse seed types, chromosome numbers, and pollen morphology in Houstonia
(Rubiaceae). Amer. J. Bot. 73:103-115.
Verocourt, B. 1958.Remarks on the classification of the Rubiaceae. Bull. Jard. Bot. Etat 28:209-
KS
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BRIT.ORG/SIDA 20(2)
Book NOTICE
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P y
gardeners, se plant ee iasts, and commercial 1 ery eae Gary L. Jennings, Botani-
cal Research Institute nee Fort Worth, TX ie ee U.S
SIDA 20(2): 558. 2002
NOTES ON CARPHEPHORUS ODORATISSIMUS
(ASTERACEAE) IN PENINSULAR FLORIDA, U.S.A.
Steve L. Orzell Edwin L. Bridges
Avon Park Air Force Range Botanical and Ecological Consultant
9 South Boulevard 52 folly Tree Place
Avon Park Air Force Range, FL 33825, U.S.A. Bremerton, WA 98312, U.S.A.
steve.orzell@avonpark.macdill.afmil ebridges@earthlink.net
ABSTRACT
Morphology, distribution, ecology and phenology for Carphephorus odoratissimus and C.
horus odoratissimus var.
=
odoratissimus var. subtropicanus are discussed and compared. Carphep
varietal status is quantified. It occurs in a fire-
subtropicanus is illustrated and evidence s
f] ] ]
maintained pine savanna-flatwoods/dry prairie Tandseape where it is a conspicuous, perennial,
rer
autumn flowering forb, endemic to south and c al peninsular Florida.
RESUMEN
Se discute y « 1 la morfologia, distribucion, ecologia y fenologia de Carphephorus odoratissimus
yc. adorauissims var. supeinpleanus Carphephor us odoratissimus var. subtropicanus se ilustra y se
cuantifican | varietal. Se da en una sabana/pradera seca de pinos
mantenida mediante fuego. es una hierba perenne, otonal, endé lel sur y centro de la peninsula
INTRODUCTION
Carphephorus sensu lat. (Asteraceae-Eupatorieae-Liatrinae) comprises seven
species (Correa & Wilbur 1969), and one recently recognized variety (Wunderlin
& Hansen 200) of Carphophorus odoratissimus, all of which are endemic to the
southeastern United States. Six of these taxa occur in Florida, of which five [C.
carnosus (Small) C.W James, C. corymbosus (Nutt.) Torr & A. Gray, C. paniculatus
(J.F Gmelin) H. Hebert, C. odoratissimus (J. Gmelin) H. Hebert, and C odoratissimus
var. subtropicanus (DeLaney, N. Bissett @ Weidenhamer) Wunderlin & BE
Hansen] are known from peninsular Florida (DeLaney et al. 1999; Wunderlin
1982; Wunderlin et al. 1996; Wunderlin 1998; Wunderlin @ Hansen 2001).
he iption of Carphephorus has been much debated (Hebert 1968;
Correa & Wilbur 1969; Cronquist 1980; Cox 1998). Some authors (Hebert 1968;
Correa & Wilbur 1969) include Trilisa and Litrisa within Carphephorus, while
others recognize them as distinct genera (Cassini 1823, 1828; Robinson 1913;
Gaiser 1954; James 1958; King & Robinson 1987; Bremer 1994). Evidence pre-
sented by both Hebert (1968) and Correa & Wilbur (1969) support the inclu-
sion of Trilisa within Carphephorus. Most recent floristic works have adopted
this broad definition of Carphephorus (Cronquist 1980; Godfrey & Wooten 1981;
Long & Lakela 1971; Wunderlin 1982; Wunderlin et al. 1996; Wunderlin 1998).
—~
SIDA 20(2): 559 — 569. 2002
560 BRIT.ORG/SIDA 20(2)
In the most recent taxonomic study of Carphephorus sensu lato (Correa &
Wilbur 1969), Trilisa and Litrisa are included within Carphephorus. Since
Correa and Wilbur (1969), no new taxa in Carphephorus were recognized until
DeLaney et al. (1999), described C. subtropicanus DeLaney, N. Bissett, &
Weidenhamer. Following DeLaney et al. (1999), Wunderlin and Hansen (2001)
treated C. subtropicanus at varietal rank within C. odoratissimus.
Based upon field observations in the fall of 1995, there seemed to be two
entities of C.odoratissimus within peninsular Florida. Carphephorus specimens
from flatwoods and prairie habitats of south-central peninsular Florida seemed
somewhat morphologically different from C. odoratissimus, as known to us from
previous fieldwork in southern Alabama, southern Mississippi, southern Geor-
gia, and the Florida panhandle. Subsequent field investigations and critical taxo-
nomic study of Carphephorus conducted from 1995-1997 led us to conclude
that the Carphephorus entity in south-central peninsular Florida was worthy
of recognition. Since that time, our on-going (1998-2001) field investigations
and taxonomic studies of Florida Carphephorus, particularly in the region of
range overlap for C. odoratissimus and C. subtropicanus in central Florida, lead
us to conclude that C. subtropicanus should be recognized as a variety of C.
odoratissimus. In this paper we present ecological, geographical, morphologi-
cal, and Pacnolegica! evidence to corroborate Wunderlin & Hansen’s (2001)
recognition of C. subtropicanus at varietal rank. All our data on morphological
characters are derived from field sampling of randomly chosen individuals in
each of ten populations of these varieties scattered in central Florida. Both en-
tities have mainly separate geographic ranges but intergrade where they are
contiguous or overlapping; both exhibit slightly divergent but apping flow-
ering phenology; and despite quantitative differences in mean values af veg-
etative morphology and numbers of flowers per head, they lack differences in
achene and floral morphology; all these features are indicative of varietal rather
than species level recognition. Phenotypically, C. odoratissimus var.
subtropicanus is distinguished from C. odoratissimus var. odoratissimus by its
lack of a coumarin janice like odor), shorter and narrower basal leaves,
strongly clasping and entire stem leaves, broader and more diffuse inflorescence,
and more flowers per head. An illustration of Carphephorus odoratissimus var.
subtropicanus is provided in Figure |
TAXONOMIC RELATIONSHIPS
Carphephorus odoratissimus var. subtropicanus differs from var. odoratissimus
most notably in leaf character and some inflorescence characters. Small
(1933:1337) noted that “extreme forms” of Trilisa odoratissima (= C
odoratissimus) might represent two entities, “one with a strong coumarin odor,
broad, clasping, coarsely toothed upper leaf -blades and slightly viscid involu-
cres[var.odoratissimus], the other with only a faint coumarin odor, narrow entire
=“
561
\ MI (
ORZELL AND BRIDGES
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AN. AN
25,261.A. Habit of base of plant; B.
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Inflorescence: C. Si
,
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562 BRIT.ORG/SIDA 20(2)
upper leaf blades and very viscid involucres. Small never published the latter
as a distinct taxon, and subsequent floristic workers have treated C.
odoratissimusasa single entity. A revision of the genus (Correa & Wilbur 1969)
also failed to account for the two as separate entities. According to Correa and
Wilbur (1969), the type of C.odoratissimus is “presumably from the outer Coastal
Plain of South Carolina ... which was the site of most of the species mentioned
in Thomas Walter's Flora Caroliniana.” Although this type has not been seen
by the authors, the collection location is outside the range of C. odoratissimus
var. subtropicanus.
There is considerable character overlap between the two C. odoratissimus
varieties (Table 1). The presence of numerous quantitative differences in inflores-
cence and leaf morphology are the most divergent character states. However,
the lack of major differences in achene and floral morphology indicates differ-
entiation at the varietal level. One useful character is the average number of
inflorescences per clump in well-developed plants. Most plants of C.odoratissimus
var. subtropicanus have one or two flowering stems per clump (mean=L.6,
std.=0.8), whereas in central Florida var. odoratissimus averages over twice as
many inflorescences per clump (mean=4.l, std.=1.9). Several characters of the
basal and stem leaves have different mean character states. Although there is
wide variation in leaf size and shape between plants within a population in
either entity and between lower and upper leaves on individuals, comparison
of leaves in the same relative position on plants of the two varieties reveals con-
sistent differences. The basal leaves of C. odoratissimus var. subtropicanus are
both shorter (mean=l1 cm) and narrower (mean=3.0 cm) than those of var.
odoratissimus (mean length=19.8 cm; mean width=5.6 cm). The stem leaves of
var. odoratissimus are broadly elliptic, and although the base of the leaf clasps
the stem, the apex of the leaf is often divergent or flared away from the stem.
The margins of the stem leaves in var. odoratissimus are often coarsely shal-
lowly toothed. In contrast, the stem leaves of var. subtropicanus are narrowly
elliptic, clasp the stem for their entire length, and typically have entire margins.
Other characters that differentiate the two varieties involve the size and
branching pattern of the inflorescence. The inflorescence of var. subtropicanus
is much broader than that of var. odoratissimus, and on well-developed mature
plants it is almost always as broad as tall or broader than tall (height:width
ratio mostly from 0.5:1 to 1:1). This difference is mostly attributable to the angle
of branching of the primary inflorescence branches from the main axis. The
inflorescence branches of var. subtropicanus diverge from the main axis at an
angle of 30°to +5°and are often arcuate, resulting in a rather open and diffuse
inflorescence. In contrast, the main inflorescence branches of var. odoratissimus
diverge from the main axis at an angle of 10°to 20° resulting in a narrower,
taller, inflorescence that is typically taller than broad (height:width ratio from
L5:1 to 3:1). This open, diffuse inflorescence, more diffuse than any other species
ORZELL AND BRIDGES 563
Taste 1: Ranges of character states in selected populations of Carphephorus odoratissimus.
Character var. odoratissimus var. subtropicanus
Stem height 140-180 cm 50-150 cm
Stems per clump 1-9 1-5
Basal leaf length 17-24 cm 9-14. cm
Basal leaf width 4.5-6.8cm 2.5-4.0 cm
Inflorescence height:width ratio 1.5:1 to 3:1 0.5:1 to 1:1
wer stem leaf length 10-15 cm 4-11. cm
Lower stem leaf width 5-7.cm 1.7-4.0 cm
Midstem leaf length 2.2-4.5 cm 1.6-6.0 cm
Midstem leaf width 1.8-3.5 cm 0.8-2.4 cm
Flowers per head (4-)7-10 (7-)10-14
Involucre height 4-6 mm 7-9 mm
Involucre width 2-3 mm 4-6mm
Achene length 2-2.5mm 1.9-2.5(-2.8) mm
of Carphephorus, is a reliable field character but is often difficult to discern in
dried herbarium specimens or plants with an immature inflorescence.
Differences in inflorescence, floral, and achene characters between the two
varieties are minor. There is considerable variation even within an individual
in the number of florets per head, size of heads, and size of floral parts.
Carphephorus odoratissimus var. subtropicanus tends to have more florets per
head (10-13) than is typical for var. odoratissimus (7-10). However, this could be
attributable to the fact that most var. subtropicanus inflorescences consist of
only 50-75 heads, in contrast to the hundreds of heads present in some inflo-
rescences of var. odoratissimus; therefore a larger proportion of heads fully de-
velop all of their potential florets. We can not find any consistent differences in
achene length (1.9-2.5 mm) (-2.8 mm) for C. odoratissimus var. subtropicanus in
comparison to var. odoratissimus (2.0-2.5 mm) or other floral characters be-
tween the two varieties. DeLaney et al. (1999) also noted overlap in achene length
and number of florets per capitulum.
The following key can be used to distinguish the two varieties:
Plants with a strong odor of coumarin or vanilla; basal leaves usually more than 15 cm
long and 5 cm wide; midstem leaves broadly elliptic, the apex flared away from the
stem, the margins often shallowly toothed; inflorescence taller than broad
(height:width ratio of 1.5:1 to 3:1), the primary inflorescence branches diverging from
the main axis at a 10° to 20° angle; heads mostly with 7-10 pss Carphephorus
ratissimus var. Po li
Plants with very slight or no odor of coumarin or vanilla; basal Bee less than
long and 4 cm wide; midstem leaves narrowly elliptic, tightly clasping the stem, ae
margins entire; inflorescence broader than tall (height:width ratio of 0.5:1 to 1:1 . the
primary inflorescence branches diverging from the main axis at a 30° to 45°
heads mostly with 10-14 flowers er ae odoraticsimus
var. subtropicanus
a:
a
564 BRIT.ORG/SIDA 20(2)
DISTRIBUTION AND ECOLOGY
Carphephorus odoratissimus var. subtropicanus is endemic to central and south-
ern peninsular Florida, where it has been collected from 21 counties (Fig. 2),an
area encompassing approximately 50,000 sq. km (20,000 sq. mi). It is a con-
spicuous, autumn-flowering forb of burned pine savanna-flatwoods/dry prai-
rie landscape throughout south-central Florida and is quite frequent in High-
lands, Polk, and Okeechobee counties. In the western part of the peninsula, its
range extends somewhat further north as compared to the eastern part of its
peninsular range. In south Florida, C. odoratissimus var. subtropicanus is rather
infrequent in Collier County, having been collected most frequently in scrubby
flatwoods near Immokalee and historically from scrubby flatwoods at Marco
Island, the only offshore island location presently known. In southeastern
Florida it has been collected from flatwoods and prairies in St. Lucie, Martin,
and northern Palm Beach counties. It is not known from the Everglades region
of south Florida, presumably due to the absence of acidic pine flatwoods and
prairies. There is an historical collection from a sandy low pineland in north-
ern Miami-Dade County, presumably from sandy flatwoods that once occurred
on the Atlantic Coastal Ridge.
Carphephorus odoratissimus var. odoratissimus, in contrast, is found
throughout most of the southeastern United States coastal plain, from south-
eastern North Carolina south and west to southeastern Louisiana (Correa &
Wilburl969). It is quite common in northern Florida and in southern Georgia
and Alabama, where it is typically found in upland longleaf pine (Pinus
palustris Mill.) dominated woodlands and savannas. Throughout its range var.
odoratissimus tends to be associated with sandy surface soils that are better
drained than those for var. subtropicanus. Carphephorus odoratissimus var.
odoratissimus does not occur in South Florida slash pine (Pinus elliottii Engelm.
var. densa Little & K.W. Dorman) dominated flatwoods. However, ranges of both
varieties overlap in central Florida. Here Carphephorus odoratissimus var.
odoratissimus seems to reach its southern limit in ecotonal habitats on the lower
sideslopes of sandy ridges, both associated with the Central Ridges in extreme
northern Polk and Osceola counties and with coastal sand ridges in Brevard,
Volusia, Citrus, and Hillsborough counties.
Carphephorus odoratissimus var. subtropicanus is characteristic of the pine-
savanna flatwoods/dry prairie landscape of the Okeechobee, Osceola, and
Desoto Plains (as defined by Cooke 1939; Schmidt 1997) in south-central penin-
sular Florida. Pine savanna-flatwoods and dry prairie occupy nearly level, in-
ter-drainage flatlands on acidic, low nutrient, poorly drained sandy to sandy
clay alfisol or spodosol soils (Abrahamson & Hartnett 1990; Bridges 1997; Orzell
& Bridges 1997). Pine savanna-flatwoods are fire-maintained, open-canopied
pine stands dominated in south-central Florida either by Pinus palustris or P.
peel
ORZELL AND BRIDGES,
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location within county i 1 SMOWT
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I g few ions i ty. For Clit COUN:
in center of county when species is widely distributed throughout county. Distribution of C. odoratissimus var.
elliottii var. densa, with Aristida beyrichiana Trin. & Rupr, Serenoa repens (W.
Bartram) Small, and low-growing Querc (Sarg.) Small typically domi-
nant in the ground cover as well as numerous regionally endemic species that
vary floristically between geographic regions (Orzell & Bridges 1997). Dry prai-
ries are similar in ground cover composition to pine-savanna flatwoods but are
naturally treeless. Historically, dry prairies had annual or biennial naturally
occurring fires (Harper 1921, 1927; Orzell & Bridges 1999) and are the highest
fire-frequented community type in central Florida (Orzell & Bridges 1999).
Associated species that are endemic or near-endemic to peninsular Florida in-
clude Asimina reticulata Shuttlew.ex Chapm., Polygala setacea Michx., Liatris
tenuifolia Nutt. var. quadriflora Chapm., Rhexia nuttallii C.W. James, Bejaria
racemosa Vent., Andropogon brachystachyus Chapm., Andropogon ternarius
Michx. var. cabanisii (Hack.) Fern. & Griscom, Gymnopogon chapmanianus
Hitche., Phoebanthus grandiflorus (Torr. &@ A. Gray) S.F Blake, and Polygala
rugelii Shuttlew. ex Chapm.
Carphephorus odoratissimus var. subtropicanus can be sympatric with any
566 BRIT.ORG/SIDA 20(2)
of the four other Carphephorus species occurring in central Florida. It most of -
ten occurs with C. corymbosus at drier sites, C. paniculatus at mesic sites, and in
very rare cases with C. carnosus at wet-mesic sites. Carphephorus odoratissimus
var. subtropicanus is sympatric with var. odoratissimus at its northern (Polk
and Osceola counties) and northwestern (Citrus and Hillsborough counties)
range limits. Plants intermediate between the two are occasionally found in
ecotonal and disturbed sites in central Florida where the boundaries of the two
overlap. Where the two varieties are sympatric at a site in Osceola County, var.
subtropicanus gr ows in dry-mesic poorly drained pine flatwoods, whereas var.
odorat is found ona somewhat poorly drained slight sandy rise between
pee
the pine flatwoods and a wetland depression.
PHENOLOGY
Ata disturbed site in Orange County, specimens intermediate in characteris-
tics deaf morphology, inflorescence, and coumarin odor) between var.
odoratissimus and var. subtropicanus were observed with both flowering in late
September 1999. The co-habitation and overlapping flowering at the site of both
varieties indicates that ecological separation of the two varieties is incomplete.
In 1997 at an Osceola County site, C.odoratissimus var. odoratissimus flow-
ered from early September till late October, and C. odoratissimus var.
subtropicanus flowered from late October into November. Elsewhere in central
Florida, var. odoratissimus generally flowers from early to mid September into
late-October and even earlier in north-central Florida (August thru mid-Sep-
tember), whereas var. subtropicanus flowers primarily from early to mid Octo-
ber thru early to mid November, with flowering having been noted in Septem-
ber at several locations. Plants flowering in early or late September were also
noted by DeLaney et al. 1999). Furthermore, atypical flowering dates for var.
subtropicanus (usually the result of burning followed by available soil mois-
ture) have been observed. Flowering plants were collected on 2 April 1998 fol-
lowing a 19 August 1997 burn in Osceola County.
APPENDIX |
Citations here are abbreviated, however full label data is available upon request
from the authors, including detailed habitat and location information for the
Orzell & Bridges collections.
aon specimens of Carphe} loratissimus var. subtropicanus: FLORIDA: Charlotte
:9miN of Placida, off SR 775 & sib 45, 20 Ge 1977, Fulton 35 (USF). Citrus Co.: SR 480,
ee ka, 7 Oct 1972, Genelle & Fleming 1606 (FLAS, USF). Collier Co.: off Hwy 82 Wo
Immokalee, 25 ann Lakela 27406 (USF). Miami-Dade Co.:7'" Ave., N of Miami, 30 Nov 1946, Ledin
s.n. (FLAS). Desoto Co.: oak scrub, ca. | mi W of Arcadiz von FL 70, 29 Oct 1978, Wunderlin, Arcuri &
Hansen 6386 (USF). Hardee Co.: ca. 1 air mi S of Fort Green Springs, 9 Nov 1993, Hansen & Wunde rlin
12435 (USF). Hendry Co.: ca. 12.5 air mi SE of | ‘aBelle, 21 Sep 1904. Orzell & Bridges 23233 (FTG).
Hernando Co.: ca. | mi W of Weekiwachee Springs, 15 Nov 1983, 8. Godley s.n. (USF). Highlands Co.:
ORZELL AND BRIDGES,
Sebring, 24 Oct 1945, Brass 15613 (ARCH); dry-mesic, sandy, treeless flatwoods/dry prairie, Avon Park
Air Force Range, 12 Nov 1997, Orzell & Bridges 25261 (ARCH, BRIT, FLAS, FSU, MO, NY, TEX, USF).
Hillsborough Co.: Flatwoods Park, ca.7 mi E of Lutz, 2 Oct 1994, Wunderlin 10558 (USF). Lee Co.: Six
Mile Cypress Preserve, 17 Oct 1997, Bradley 791 (FTO). Manatee Co.: cal mi N of Myakka River State
1978, Dodson 4798 (USF). Martin Co.: W of canal, S of Salerno ject. off US 1, 29 Sep1962,
OFOLATILATL
Park, 17 Se
Lakela 25383 (FLAS, pie Okeechobee Co.: US 68, 15 mi N of Okeechobee, 19 Oct 1969, McCart 11141
(FLAS, USF). Osceola Co.: ca. 12 air mi S of Kissimmee, ll Oct 1997, Orzell & Bridges 25237 (FLAS,
rzell & Gold 25263 (USF). Palm Beach Co.:
USF), Three L ee Wi ldlife ee Area, 2 Apr 1998, Orzel
Loxahatchee River Corridor, | Oct 1997, ness & Woodmansee 559 (FTG). Paseo Co.: Upper
*t 1974, Rochow s.n. (USF). Pinellas Co.: Brooker
Park Air
Hillsborough Flood Detention Area, Zephyrhills
Creek Preserve, 17 Nov 1993, Hansen 12533 ee Polk Co.: N of Tick Island Slough, Avon I
Force Range, 9 Nov 1994, Orzell & Ae 23431(FTG). Sarasota Co.: ca. 5 mi S of Venice, off US 41,
20 Sep 1961, Lakela 24653 (FLAS, USF). St. Lucie Co.: Savanna State Reserve, along E-W track to W
side of Savannas, 24 Sep 1992, an 971(FLAS).
] RIDA: Alachua
Representative specimens of Carpheph
Co.: NW of ee 14 Sep 1978, oe I71 (FL AS). Bradford Co.: 19 Oct 1977, Conde s.n. (FLAS).
Brevard Co.: 16 972, Shuey M0345 (USF): N of Cocoa.ll Oct 1963, Lakela 26595 (USF). Calhoun
Co.: Sof ee 28 Sept | : Inverness,10 Nov 1982, Mawhinney
40 (USF). Crystal River, 6 Jun 1941, Murrills.n. (FLAS). Dixie Co.: N of Oldtown, 18 Aug 1937, Arnold
s.n. (FLAS). Duval Co.: Fort Caroline Club Estates, 18 Aug 1963, Creager 269 (FLAS). Escambia Co.: .
of Pensacola, 20 Nov 1983, Wilhelm 11852 (USF). Flagler Co.: Palm Coast, 4 Oct 1978, Tabb s.n. (UST
SE of oes 12 Oct 1979, Hansen, Wunderlin, Sauleda, & Richardson 6719 (USF). N of canes
18 Apr 1940, West & Arnold s.n. (FLAS). Gilchrist Co.: E of Spent 10 ven 1940, Martin, De Vall &
rate s.n. (FLAS). Hamilton Co.: N of White Springs, 8 Oct 1966, D’Arcy 1219 (FLAS). Hillsborough
Co.: 21 Sep 1967, Lakela 31111 (USF). Holmes Co.: W of Ponce oo eon, 28 Sep 1981, Hansen 906 (USF).
e Co.: Ocala National Forest, 3 Sep
SF
983, Nelson 2988 (FLAS). Citrus Co
E of Walton Co line on US 90, 23 Sep 1967, Blum 2745 (USF). L
1976, Daubenmire s.n. (USF). Leon Co.: Apalachicola National a 29 Oct 1983, Parker 1790 (USF).
Levy Co.: vicinity of Cedar Key, 15 Sep 1971, Carlton, ies Lakela, & Long 3351; (USF). Ce-
erve, 19 Oct 1990, Amoroso 521 ( ees Liberty Co.: Apalachicola National
er River State Park, 10 Sep 1990, Buckner
of Chri
dar Key Scrub State Res
Forest, 9 Dec 1988, Gholson & Hill 20133 (USF). Marion Co.:
I11 (FLAS). Okaloosa Co.: N of Niceville, 26 Sept 1967, cae 1196 (FLAS). Orange Co.:
mas, 1] Sep 1979, Hansen & Richardson 6240 (USF). 10 mi ce Oallanc 7 fan: 1958. Kral 7745 (USF)
JSF). Polk Co ca Haines
elaka,
Osceola Co.: W of Intercession City, 14 Aug L981, Hans¢
City, 2 Sep 1968, Shape n. ae ae Lake Wales, 1 Oct 1964, C a S.A. (FL AS). P
16 Jun 1939, De Vall s.n.(F anta Rosa Co.: N of Hernandez Pt., 28 Nov 1980, Wi ea, en
NE of Holley, 26 Sep 1967, coe 2003 (FLAS). Seminole Co.: Oviedo, 7 Sep 1947, Schallert 6762 (USF)
St. Johns Co.: Faver Dykes State Park, 29 Sep 1983, Hansen 9857 (USF). W of Cresent Beach, 14 Oct
1941, West & Arnold s.n. (FLAS). Sumter Co.: Wildwood, 20 Sep 1981, Correll & Correll 52625 (USF).
Taylor Co.: US 27, W of Lafayette Taylor Co. line, 12 Sep 1960, Ward & Myint 2175 (USE, FLAS). E of
Perry, 12 Aug 1956, Knobloch 1429 (FLAS). Union Co.: S of Raiford, 2 ie ae 1942, West & Arnold s.n.
Funiak Spr ing
(FLAS). Volusia Co.: Near Ormond, 5 Aug 1943, Butts s.n. (USF). W: aSof D
27 Sep 1967, Smith 2006 (FLAS). Washington Co.: N of Chipley, sae 1954, Ford 3682 (FLAS). me
bama: Covington Co: F of Wing, 20 Oct 1969, Kral 38120 (USF). Escambia Co: Riverview, 27 Nov
1980, Wilhem 8180 co GEORGIA: Thomas Co: N of Boston, 3 Oct 1967, eee 2668 (USF). MeIn-
tosh Co: NE of Sapelo Island, 17 Sep 1956, Duncan 20544 (USF). LOUISIANA: Washington Par: N of
Hackley, 16 Sep 1983, Taylor & Dutton 5618 (USF). MISSISSIPPI: Harrison Co: Pass Christian, 22 Oct
1954, Demaree 36238 (USF). Jackson Co: Hurley, 27 Sep 1953, Demaree 34398 (USF). Pearl River Co: I-
59 exit 4, 22 Oct 1984, Hermann 483 (USF). SOUTH CAROLINA: Beaufort Co: SE of Yemassee, 6 Sep
1956, Bell & Ahles 17999 (USF). Berkley Co: S of Cross, 23 Sep 1966, Bradley & Sears 3551 (USF
568 BRIT.ORG/SIDA 20(2)
ACKNOWLEDGMENTS
We thank Paul Ebersbach, Chief of the Environmental Flight at Avon Park Air
Force Range (APAFR), for his support of scientific research on the military in-
stallation; Rebecca Yahr for the illustration; Scott Penfield (APAFR) for provid-
ing funding from the Rangelands Program at APAFR for the illustration; and J.
Douglas Ripley formerly of the Environmental Planning Division of the U.S.
Air Force in Washington DC., who secured funding for both the illustration
and manuscript preparation. Fieldwork was facilitated by Sam VanHook, who
conducted prescribed burns of sites for C. odoratissimus var. subtropicanus at
APAFR, and Chris-Ann Kosel formerly of the TNC Disney Wilderness Preserve,
who provided access to the preserve. We especially thank Guy Nesom of BRIT,
Billie Turner of TEX, and Richard Wunderlin of USF for providing stimulating
views and critical edits to earlier drafts of the manuscript.
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Bot. Missouri Bot. Gard. 22:277-280.
Lona, R.W. and O.Laketa.1971.A flora of tropical Florida. University of Miami, Coral Gables.
Orzett, S.L. and E.L. Brioces. 1997. Regional floristic diversity in peninsular Florida pine
flatwoods and savannas. Abstr. of the Southeastern Flatwoods Ecosystem. First Ann.
Conf. Soc. Ecol. Rest. Coastal Plain Chapter. May 16-17, 1997. Gainesville, FL.
Oprzett, S.L. and E.L. Brioces. 1999. Dry prairie. U.S. Fish and Wildlife Service. eds. In: South
Florida multi-species recovery plan, a species plan, an ecosystem approach. South-
eastern Region, Atlanta. Pp. 3-279 thru 3-346.
Rosinson, B.L. 1913. A key to the genera of the Compositae — Eupatorieae. Proc. Amer.
Acad. Arts. 49:429-437,
Scumipt, W. 1997. Geomorphology and physiography of Florida. A.F. Randazzo and DS.
Jones. eds. In: The geology of Florida. University Press of Florida, Gainesville. Pp. 1-12.
Smatt, J.K. 1933. Manual of the southeastern Flora. Published by author, New York.
WunNbeRrLin, R.P. 1982. Guide to the vascular plants of central Florida. University Press of
Florida, Tampa.
WUNDERLIN, R.P. and B.F. Hansen. 2001.Seven new combinations in the Florida flora. Novon
11:366-369.
WUNDERLIN, R.P.,, B.F. Hansen, and E.L. Brioces. 1996. Atlas of the flora of Florida. Florida De-
partment of State, CD-ROM version of publication. (Also electronically published at:
http://www.plantatlas.usfedu/default.asp)
WUNDERLIN, R.P. 1998. A guide to the vascular plants of Florida. University Presses of Florida,
Gainesville.
—=
BRIT.ORG/SIDA 20(2)
Book REVIEW
Davip G. Fropin. 2001. Guide to Standard Floras of the World. Second Edition.
(ISBN 0-521-9077-0, hbk). Cambridge University Press, Cambridge, U.K.
$240.00. 1100 pp, 7 1/2" x 10".
The entire history of botanical exploration of the world can be gleaned from this new edition of
Frodin’s 1100-page bibliography, which is nearly twice as long as edition one (1984). It is a source of
an incredible—almost unbelievable—amount of information. We are here dealing with a landmark
iterature on plants. The a ral ‘monumental,” mouse hackneyed comes to mind.
Floras exist for every part of the world, no matter how remote. In North America north of Mexico,
they are available from county ae to state-, country-, or even continent level, e.g., Flora of North
America. Almost every Bate has its own aes canada has a four- volume ora of Canada plus floras
work on Flora Brasiliensis,
in
for many
for example, extended over 66 years. A few never are ae beans: of lack of funds or botanists.
Frodin includes not only full-fledged floras klists and floristic atlases.
Areas covered by florasare of all sizesand oS oe are some exam Sane ar renee rom small
to large, with the number of species in each—A
an island group: the Falklands, 256; part of a country: Michigan, 2465; a country: Canada, 4100; part
of a continent: North American Great Plains (extreme southern Prairie Provinces of Canada to the
Texas pele 3000; and, finally, continents: Australia, 20,000, and Antarctica, 2 native and |
introduced spec
‘rodin’s book is eee by major geographic area, ie, North America, Asia, Europe, Africa,
ithi states, islands, etc.). Coverage is strongly 20th century,
he book is good reading, including
ai ample representation from the 19th and even earlier. All of t
the 1424 footnotes.
Among the tidbits in the Guide are these about Easter Island. The first floristic account of Eas-
ter appeared in 1914, with others in 1920, 1922, 1951, 1958, and 1991. The island has 195 species, aie!
introduced (only 46 are eate Badly treated by its inhabitants, it is now virtually treeless. One o
the trees, the legume So} is extinct except in cultivation off the island. The last ne
of ae tree from the island was by the Thor ete 1955-1956 expedition. The island's sole palm is
— too, known only from fossil remains.
ore tidbits? Which USS. state has the most plants? Leading the pack are California, 5800, fol-
lowed i Texas, 4800, and Florida, 3800. North Dakota is last with about 1000
The compiler of all this wealth, David Frodin, is a botanist at Kew Gardens in England. He
spent many years br inging together in one place an abundance of floristic literature; the user is thus
spared this aes effort. The myriad titles he lists are testimony to the thousands of dedicated
botanical explorers who have worked over the centuries to discover and make known the world’s
plants.—John W. ee Northern Kentucky University.
SIDA 20(2): 570. 2002
RESOURCE ALLOCATION PATTERNS AND PHENOTYPIC
VARIATION IN THE ENDANGERED TEXAS WILDRICE
(ZIZANIA TEXANA, POACEAE)
Paula Power
San Marcos National — ees and Technology Center
McCarty Lane
San ee TX 78666, U.S.A.
ABSTRACT
|
Zizania texana Hitche., an re endemic to the upper 4.9 km of the San Marcos
River, Hays County, Texas was Hederally listed in 1978 after the species experienced a population
decline between about 1940 and 1967. Recent interest in restoration has focused attention on the
need for a better understanding of the species’ response to a variety of microhabitats. This study
documents biomass allocation patterns in three ei over 7 months. Current velocities
ranged from 0.0-0.010 m/s in the slow site, 0.038-0.142 m/s in the moderate site, and 0.250-0.369
m/s in the fast site. ieee were Par vested eo eh site, on seven occasions, washed, Separateg into
roots, submersed leaves.
and weighed. At the end of the study, mean total biomass of plants grown in the fast site was 28.42
gdw vs 4.24 gdw for plants grown in the slow site. Resource allocation patterns differed among sites
and two distinct phenotypes were apparent. One phenotype, associated with relatively higher cur-
rent velocities, had higher net productivity, a well-developed root system, and allocated proportion-
ally more biomass to non-reproductive organs (49.2% gdw root biomass in fast site vs 24.8% gdw
root biomass in slow site). A second phenotype, associated with relatively slower flowing water, had
lower net plore aly and allocated proportionally more ene to reproductive organs (22.1%
w lm in fast site vs. 65.0% gdw reprodt Im in i site). Because of lower
productivity and the potenti for loss from herbivory toa sipfican proportion of the panei in
with relati vely low CULTEILIL velocity would MO
for eoeetion purposes.
RESUMEN
as Be ce Ey: } Af: 1 AK A | 49] . Ba be [ier dea
TCOS.
Hays Co., Texas se incluy6 en la lista federal en 1978 después que la ae experimentase una
se see poblacional entre 1940 y 267. a interés reciente en _ restauracion ane pr res pigs atencion
oh
ee Este estudio document a los modelos de situacion mn de ope en tres microhabitats du
rante 7 meses. Las | l to), 0.038-0.142 m/
s (en el sitio pases y 0.250-0.369 m/s (en el sitio wk Las ee se cosecharon de cada
pacrehabite se lavaron, separaron - raices, las hojas sumergidas, y los culmenes
las hoj inf] y pesaron. A final del estudio,
la isiouas total media a plantas del sitio rapido fue 28. 42 gdw contra 4.24 gdw para plantas del
sitio lento. Los modelos de recursos difirieron significativamente entre los diferentes sitios y fueron
aparentes dos fenotipos claros eran. Un fenotipo asociado con velocidades actuales relativamente
mas altas, ale una piodncoviea neta mas alta, un sistema radical bien desarrollado, y coloc6é
vegetativos (49.2% de peso seco de biomasa de raiz en el
sitio rapido contra 24.8% de peso seco eae biomasa de raiz en el sitio lento). Un segundo fenotipo,
SIDA 20(2): 571 - 582. 2002
572 BRIT.ORG/SIDA 20(2)
asociado con agua de corriente relativamente mas lenta, tuvo una productividad neta mas baja y
coloco proporcionalmente mas biomasa en los organos de reproductores (22.1% de peso seco de
biomasa en calmenes reproductores en el sitio rapido contra. 65.0% de peso seco de biomasa en
culmenes reproductores en el sitio lento). A causa de tasas bajas de productividad y la pérdida
ances debida a los herbivoros en una proporcion significativa en el agua de corriente mas lenta,
este tipo de microhabitat no se deberia recomendar para las restauraciones.
INTRODUCTION
Trade-offs between reproductive allocation and vegetative allocation with
changes in environmental conditions have been reported by Harper and Ogden
(1970); Hickman (1975); van Baalen, et al. (1990); Dunn and Sharitz (1991);
Madsen (1991) and Neill (1993). Environmental factors associated with alloca-
tion trade-offs include substrate, nutrient availability, and water depth
(Idestam-Almquist & Kautsky 1995; Blanch et al. 1999; Lorenzen et al. 2001;
Vretare et al. 2001). Little information is available on current velocity and its
affect on allocation patterns in macrophytes although numerous studies have
identified current velocity as an important factor influencing macrophyte dis-
tribution, photosynthesis and growth (Nilssen 1987; Chambers et al. 1991;
Madsen & Sondergaard 1983; Power 1996a). The work reported here was under-
taken to document resource allocation patterns in the endangered Zizania
texana Hitche. over seven months in three different habitat types differing in
current velocity and pH.
Zizania texana is an endangered macrophyte endemic to the San Marcos
River, Hays County Texas (USS. Fish & Wildlife Service 1995). It commonly oc-
curs midchannel in swiftly flowing water. Terrell et al. (1978) and Poole and
Bowles (1999) provide thorough descriptions of the habitat for Z. texana. Threats
to the species include reduced spring flow from the source aquifer due to
overpumping of ground water for human use, competition and herbivory by
nonnative plant and animal species, and absence of sexual reproduction in the
wild, along with other human impacts including alteration of historic hydro-
logic patterns from dams located along the river and within the watershed, and
recreational use (U.S. Fish & Wildlife Service 1995).
Under flowing water conditions, Z. texana produces long ribbon-like, sub-
mersed leaves and emergent culms, each with a terminal, wind pollinated in-
florescence; however, sexual reproduction is nearly absent in the wild (U.S. Fish
& Wildlife Service 1995). Historical accounts suggest this was not always the
case. Photographs dep g fully developed inflorescences and verbal accounts
suggest recruitment by seed occurred (Silveus 1933). More recent descriptions
of floral structures and floral development by Emery and Guy (1979), and Power
(1996b; 1997) indicate that sexual reproduction is most likely limited by envi-
ronmental factors rather than cytological factors. It is not well understood why
fully developed, emergent inflorescences are rare in the wild, although drifting
mats of floating vegetation and herbivory play a role (Power 1996b, c). The spe-
POWER, RESOURCE ALLOCATION IN ZIZANIA TEXANA 573
cies produces asexual clones (tillers) which form at the nodes of reproductive
culms and it is assumed that tiller production is the primary mechanism for
recruitment of new individuals in the wild (U'S. Fish & Wildlife Service 1995).
Zizania texana has two distinct phenotypes under wild and cultured con-
ditions. When grown in the wild, Z. texana is primarily submersed and tends
to be a long lived perennial plant. Under cultivated conditions, Z. texand is pri-
marily emergent, reproductive, and short lived (Terrell et al. 1978 and pers.
observ.). Information on plant response to environmental conditions, gains im-
portance given the endangered status of the species and the need for restora-
tion of the habitat as outlined in the Recovery Plan for the species (US. Fish &
Wildlife Service 1995). Adequate restoration protocols cannot be developed with-
out a thorough understanding of the response of Z. texana toa variety of con-
ditions found in its historic range.
METHODS AND MATERIALS
This research project was carried out in Spring Lake on the Southwest Texas
State University (SWT) campus. Spring Lake is an impoundment formed by a
dam and spillway originally constructed across the San Marcos River in 1849.
The dam and spillway are approximately 750 m downstream from the San
Marcos Springs.
Study plants were obtained by germinating captive grown seed in the lab.
Seedlings were then transplanted to 15cm plastic pots, lined with small plastic
bags and filled with sediments collected from one location in the study site in
Spring Lake. Pots were placed in an outdoor cement raceway on the SWT cam-
pus and seedlings were allowed to grow for about 6 weeks. Water was supplied
by an artesian well from the source aquifer for Spring Lake and the San Marcos
River. In March 1995, potted plants were transplanted into three sites in Spring
Lake and one site in the outdoor raceway on the SWT campus. This wasa nested
design with plants nested in plots and plots nested in sites. There were three
replicate plots at each site. Each replicate plot had 36 potted plants for a total of
432 plants. All plants were protected from herbivores with 1 m?, floating
exclosures ructed of polyvinyl chloride (PVC) pipe and 2.5 cm wire mesh.
Initially, four plants were harvested to obtain baseline biomass values for the
newly transplanted individuals. The study design called for harvesting four
plants from each replicate plot on nine separate occasions. However, during the
study period some plants were lost to herbivory and some plants were washed
away by the current. Asa result, plants were harvested from each replicate plot
on six dates, May 3, June 1, June 30, July 27, September 1, and October 16. At
each harvest, plants were selected at random. Sediment was washed from plant
roots, then plants were divided into vegetative parts, reproductive parts, and
roots. Vegetative parts were defined as submersed leaves and reproductive parts
were defined as reproductive culms with associated leaves, inflorescences, seeds,
574 BRIT.ORG/SIDA 20(2)
and tillers. Plants were then dried at 65°C for at least 48 hours and weighed to
the nearest 0.01 g. Data are reported as means plus-minus standard error.
On nine occasions between May 1995 and October 1995 pH, water depth,
and current velocity were recorded in each replicate plot, in each site. Current
velocity was measured with a Marsh McBirney Model 201 portable water cur-
rent meter and calculated as the average velocity at 20%, 60%, and 80% depth.
The four sites differed in mean current velocity and were identified as follows:
slow (raceway), no flow, moderate and fast.
RESULTS
Plants in the no flow site in Spring Lake were lost to herbivory, probably by
crawfish, prior to the second harvest and this site was dropped from the study.
Water depth, current velocity, and pH at the remaining sites were recorded
at the study sites eight times between May and October (Table 1). In the slow
site, mean water depth was 0.71 m (+0.03) mean current velocity was 0.001 m/
s (+0.004), and pH ranged from 7.50 to 7.68. In the moderate site, mean water
depth was 0.88 m (+0.06), mean current velocity was 0.090 m/s (+0.048), and
pH ranged from 7.23-7.26. In the fast site, mean water depth was 0.85 m (+0.07),
mean current velocity was 0.290 m/s (+0.076) and pH ranged from 7.16-7.28.
Plants in all study sites increased in size during the study period (Fig. L). At
each harvest, fast site plants had the greatest root, submersed leaf, and total net
biomass, furthermore, data from the September harvest showed mean net total
biomass of fast site plants was an order of magnitude greater than mean net
total biomass of slow site plants (34.05 g vs. 3.66 g).
Number of submersed leaves increased in all sites during the study period,
however fast site plants produced six times as many submersed leaves as slow
site plants. At the end of the study period, mean number of submersed leaves in
fast site plants was 18.0 (+ 1.414) while mean number of submersed leaves in
slow site plants was 3.0 (+ 0.211).
Reproductive culms were present in every harvest between 1 June 1995 and
16 October 1995. Culm number was greatest during the September harvest in
plants grown in the fast site with 2.9 (+ 0.380) culms/plant. For plants grown
in the slow site, number of culms was greatest in the October harvests with 1.6
(+ 0.159) culms/plant. (Fig. 2).
Biomass allocation to plant roots and reproductive culms varied through
time and among sites (Fig. 3). The proportion of biomass allocated to roots in-
creased from 27% in March at the beginning of the study to 49% in October in
plants grown in the fast site. In contrast, the proportion of biomass allocated to
roots decreased from 27% at the beginning for the study to 25% at the end of the
study in the slow site.
Plants in all sites produced similar numbers of culms, however biomass
allocation varied greatly among sites. In October, culm biomass constituted over
POWER, RESOURCE ALLOCATION IN ZIZANIA TEXANA 575
Taste 1.Means and ranges for environmental factors in two sites (Moderate and Fast) in Spring Lake
and one site (Slow) on the SWT campus. Data were collected on nine separate days between May
1995 and October 1995.
Slow Moderate Fast
Depth (m)
mean (sd) 0.71 (£0.03) 0.88 (£0.06) 0.85 (+0.07)
range 0.69-0.75 0.87-0.91 0.80-0.87
Current velocity (m/s)
ean (sd) 0.001 (+0.004) 0.090 (40.048) 0.290 (£0.076)
range 0-0.010 0.038-0.142 0.250-0.369
pH
range 7.50-7.68 7.23-7.26 7.16-7.28
half of total plant biomass in slow site plants (65%; culm number = 1.640.159),
in moderate site plants, 48% of total net biomass was culm biomass (culm num-
ber = 1.740.129 and fast site plants allocated only 22% of total net biomass to
reproductive culms (culm number = 2.25+1.591
The proportion of biomass allocated to submersed leaves was similar
among sites even though the number of leaves varied among sites. In October,
submersed leaf biomass was 29% of total biomass in the fast site (leaf number
= 18+1.414), 15% of total biomass in the moderate site (leaf number = 5.341.862),
and 28% of total biomass in the slow site (leaf number = 3+0.211).
DISCUSSION
In this study, plants exhibited markedly different growth patterns among study
sites. Net total biomass accumulated over the study period was an order of
magnitude greater in plants grown in water flowing between 0.146-0.442 m/s
compared with plants grown in water flowing between 0-0.01 m/s. Net biom-
ass accumulation in individual plant organs (roots, submersed leaves, and re-
productive culms) also was greater in fast flowing water compared with slow
flowing water. Submersed leaf biomass was 18 times greater in fast site plants
compared with slow site plants and 4.5 times greater compared with moderate
site plants. Zizania texana exhibited a similar response to flowing water in
other studies (Power 1996a, b).
Flowing water has been shown to influence macrophyte photosynthetic
rates (Westlake 1967; Smith @ Walker 1980; Madsen & Sondergaard 1983), dis-
tribution (Fonseca & Kenworthy 1987; Nilssen 1987); and growth (Chambers
et al. 1991). Plants occur in a range of current velocities and there is consider-
able variability in optimum flow rates for macrophytes. Chambers et al. 1991)
found an inverse relationship between biomass and current velocity between
0.01-1.0 m/s. Nilssen (1987) found species richness reached a peak at about 0.3
576 BRIT.ORG/SIDA 20(
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577
RESOURCE ALLOCATION IN ZIZANIA TEXANA
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POWER, RESOURCE ALLOCATION IN ZIZANIA TEXANA 579
m/s along a current gradient from 0.04-1.23 m/s, with some species growing
in current velocities greater than 1 m/s. Many species occur in slower current
velocities, while fewer species are specialized to withstand the forces of veloci-
ties over 0.3 m/s. Apparently, Z. texana with smooth, ribbon-like leaves is one
of few species able to withstand velocities over 0.3 m/s. Poole and Bowles (1999)
found wild Z. texana stands primarily in current velocities > 0.46 m/s and the
current velocity tolerance range for Z. texana exceeds 1.0 m/s; from this study
however, it was not possible to identify a maximum or an optimum current
velocity for growth.
Zizania texana exhibits phenotypic variation in response to current ve-
locity. One phenotype, associated with relatively higher current velocities, has
higher net productivity, a well-developed root system, and allocates propor-
tionally more biomass to nonreproductive organs. A second phenotype, associ-
ated with relatively slower flowing water, has lower net productivity and allo-
cates proportionally more biomass to reproductive organs. Plants face a tradeoff
between the need for a well-developed root system to anchor plants in poten-
tially unstable sediments, the need for submersed leaves with which to photo-
synthesize, and the need for emergent stems for reproduction. In this study, the
proportional allocation t |
decreased with decreasing current velocity as the proportional allocation to
reproductive organs increased. Apparently there is a trade-off in favor of sub-
mersed organs in faster flowing water.
Other factors directly and indirectly influenced by current velocity may
play a role in net biomass accumulation in plant organs. They include herbivory,
deposition of debris and sediments on leaves interfering with metabolic pro-
cesses, colonization of leaves with epiphytes, and the plant's inability to utilize
O3 and its dependence on CO? as an inorganic carbon source (unpublished
data). Ribbon-like submersed leaves of Z. texana are adapted to withstand the
forces of flowing water and can reduce carbon limitation by exploiting flow-
ing water habitat where boundary layer surrounding leaves and diffusion dis-
tances for CO2 are reduced, and leaves are continually bathed with carbon rich
water. In contrast, in slower flowing water, photosynthesis by submersed leaves
of Z. texana is carbon limited and few submersed leaves are produced. Emer-
gent reproductive culms with associated emergent leaves most likely are not
carbon limited because culms obtain CO2 from the atmosphere where CO? is
more readily available owing to the higher diffusion rate and current velocity
in air relative to water (Madsen & Sand-Jensen, 1991; Denny 1993).
Increased proportional allocation to emergent reproductive organs in rela-
tively slower flowing water concurs with observations of captive grown Z.
texana when grown in current velocity =0.015 m/s in which plants allocate a
greater proportion of biomass to reproductive organs and typically set seed and
senesce after one growing season. This is similar to Z. palustris and Z. aquatica,
tive organs (roots and submersed leaves)
580 BRIT.ORG/SIDA 20(2)
annual species which commonly occur in shallow water along the margins of
lakes and streams (Ferren & Good 1977; Weir and Dale 1960). The importance
of having leaves and flowers above the surface of the water may be due to CO?
limitation in submersed leaves. In relatively slower flowing water, gas exchange
and photosynthesis may be insufficient to support vegetative organs and re-
sources shift toemergent organs where CO} is plentiful.
Herbivory isa factor contributing to sexual reproducti Z.texana
(US. Fish & Wildlife Service 1995; Power 1996c). Plants growing in microhabi-
tats with relatively slow flowing water and potentially over 60% of biomass
allocated to reproductive parts, are especially vulnerable to herbivory by wa-
terfowl. Microhabitats with conditions which would trigger a low productiv-
ity/high reproductive phenotype in the wild include back eddies and protected
stream edges, emergent yte beds, impoundments upstream from dams,
and potential y, reduced springflows due to drought and overpumping of the
source aquifer (the Edwards Aquifer) for human use. These microhabitats would
not be recommended as possible Z. texana restoration sites.
C2)
ACKNOWLEDGMENTS
This research was supported by a Section 6 grant from USS. Fish and Wildlife
Service and Texas Parks and Wildlife. 1 would like to thank Kathryn Kennedy,
Center for Plant Conservation, Robert Doyle, Baylor University for their sup-
port and Francis Rose, Southwest Texas State University, for permission to use
outdoor raceways and Spring Lake.
REFERENCES
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flooding in the emergent sedge Bolboschoenus medianus. Aquatic Bot. 63:145—160.
CHameers, PA, E.E. Prepas, H.R. Hamitton, and M.L. BotHwelt. 1991. Current velocity and its ef-
fect on aquatic macrophytes in flowing waters. Ecol. Applications 1:249-257
Denny, M.W. 1993. Air and water: The biology and physics of life's media. Princeton Univer-
sity Press, New Jersey.
Dunn, C.P.and R.R. SHaritz. 1991, Population structure, biomass allocation, and phenotypic
plasticity in Murdannia keisak (Commelinaceae). Amer. J. Bot. 78:1712-1723.
Emery, W.H.P. and M.N. Guy. 1979. Reproduction and embryo development in Texas
wildrice(Zizania texana Hitchc.). Bull. Torrey Bot. Club 106:29-31.
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tion of seagrasses. Aquatic Bot. 27:59-78.
Harper, J.L.and J.Oapen. 1970. The reproductive strategy of higher plants.|. The concept of
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fected by phosphorus and oxygen availability. Aquatic Bot. 70:117-133.
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582 BRIT.ORG/SIDA 20(2)
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NOMENCLATURE OF THE HEARFLEAVED HEDGE-NETTLE,
STACHYS CORDATA (LAMIACEAE)!
James S. Pringle
Royal Botanical Gardens
Hamilton, Ontario, CANADA L8N 3H8
Jpringle@rbg.ca
ABSTRACT
Stachys cordata Riddell is the correct name for the species usually called S. nuttallii Shuttlew. ex
Benth.
RESUMEN
Stachys cordata Riddell es el nombre correcto para la especie usualmente conocida como S. nuttallii
Shuttlew. ex Benth.
Stachys cordata was described from Ohio by Riddell in 1836. The identity of the
species so named was never in question, and the use of this name prevailed dur-
ing the following 85 years. Riddell’s type specimen is extant at US and micro-
fiches are widely accessible.
In 1921, House published the name Stachys riddellii in direct substitution
for S.cordata Riddell, although he inappropriately used the designation “n. sp.”
Because this was merely a new name for the same species, no new description
was provided and no new type was cited. House considered the name S. cordata
Riddell to be illegitimate, in that it wasa homonym of S.cordata Gilib. Whether
the new plant names in Gilibert’s Flora Lituanica Inchoata should be consid-
ered validly published under the provisions of the International Code of Bo-
tanical Nomenclature was formerly a matter of doubt. This matter is now settled.
Flora Lituanica Inchoata and Gilibert’s other publications are listed in the Code
among the “opera utique oppressa,” and new names at the rank of species are
decreed not to have been validly published in those works.
Immediately upon its being published, the name Stachys cordata Gilib. was
widely recognized as being taxonomically synonymous with S. sylvatica L., the
name of a species native to Europe. Consequently, it was not inadvertently vali-
dated by subsequently being accepted by any other author. Bentham, in his com-
prehensive treatment of the genus in 1848, did not note the use of this name by
any other authors in the interim, nor did he even cite it in synonymy. Since the
provisions of the International Code of Botanical Nomenclature pertaining to
homonyms apply only to validly published names, the use of the combination
a
‘Contribution No. 101 from the Royal B | } Hamilton, Ontario, Canada.
SIDA 20(2): 583 — 584. 2002
584 BRIT.ORG/SIDA 20(2)
by Gilibert no longer constitutes a reason for rejecting the name S. cordata
Riddell.
Following House’s publication of the name S. riddellii, that name came into
general use for the species, although as late as 1933 the name S. cordata was
used in Small’s Manual of the Southeastern Flora. In 1979, Nelson and Fairey
noted that the long-misapplied name S. nuttallii Shuttlew. ex Benth. (1848),
which antedated S. riddellii House, was correctly applicable to the same spe-
cies. The species has usually been called S. nuttallii since their paper was pub-
lished. They rejected the name S. cordata Riddell only for the reason given by
House. In his monograph on Stachys in the southeastern United States, Nelson
(1981) included S. cordata Riddell in the synonymy of S. nuttallii but did not
address the question of the legitimacy of the earlier name.
The only post-1981 flora in which the name S.cordata Riddell was accepted
is that by Gleason and Cronquist (1991). The names S. riddellii and S. nuttallii
were cited in synonymy, without comment. In other standard references, both
printed and electronic and including those published since 1991, the name S.
nuttallii has continued to prevail.
Iconclude that there is no basis for the rejection of the name Stachys cordata
Riddell, and that it is the correct name for the species that has been called S.
riddellii House and S. nuttallii Shuttlew. ex Benth.
REFERENCES
BeNTHAN, G. 1848. Labiatae. In: Candolle, A.P. de, ed. Prodromus systematis naturalis regni
vegetabilis. Paris: Victor Masson. 12:27-603.
Epuina, C.C. 1934. Preliminary revision of American Stachys. Repert. Spec. Nov. Regni Veg.
Beih. 80:1-73.
Gteason, H.A. and A. Cronauist. 1991, Manual of vascular plants of northeastern United
States and adjacent Canada, ed. 2. Bronx: The New York Botanical Garden.
House, H.D. 1921. Necessary changes of certain plant names. New York State Mus. Bull.
233-234:60-69.
Newson, J.B. 1981. Stachys (Labiatae) in southeastern United States. Sida 9:104-123.
Newson, J.B.and J.E. Fairey Ill. 1979. Misapplication of the name Stachys nuttallii (Lamiaceae
to a new southeastern species. Brittonia 31:491-494.
Riopett, J.L. 1836. A supplementary catalogue of Ohio plants. W. J. Med. Phys. Sci. 9:567-
a
Smatt, J.K. 1933. Manual of the southeastern flora. Chapel Hill: The University of North
Carolina Press.
A CYPRESS (CUPRESSUS ARIZONICA, CUPRESSACEAE)
IN JEFF DAVIS COUNTY, TEXAS?
M. Patrick Griffith Stephanie C. Bartel
Rancho Santa Ana Botanic Garden Department of Biolo
1500 N. College Ave. Sul Ross State University
Claremont, CA 91711, U.S.A. Alpine, TX 79832, U.S.A.
ABSTRACT
(Cc ) iT
We investigated a recent report of a disjunct individual of Cup
lected in the Davis Mountains of Jeff Davis County, Texas. Cileropleet DNA nee were obtained
for the novel disjunct specimen and putative related taxa. Phylogenetic analysis suggests that the
Davis Mountains C. arizonica specimen consists of ana tissue from Juniperus deppeana,acom-
mon species of the Davis Mountains. Based on this evidence, the present known distribution of C.
arizonica in Texas remains restricted to the Chisos ene of southern Brewster County.
RESUMEN
Hemos aeatciou ga un reciente registro de un individuo disyunto de Cupressus arizonica
Cur as montanas de Davis del condado de Jeff Davis, Texas. Se obtuvieron
secuencias de ADN aca del nuevo registro y los supuestos taxa relacionados. El analisis
que el especimen de C. arizonica de las montanias de Davis consiste en tejidos
veretativos de Juniperus deppeana, una especie comun de las montanas de Davis. En base a esta
evidencia, la actual distribucion de C. arizonica en Texas, permanece restringida a las montanas
Chisos del condado de Brewster.
INTRODUCTION
Cupressus arizonica, as broadly circumscribed (Wolf @ Wagener 1948,
Eckenwalder 1993) includes a number of variable, isolated populations in south-
western North America. The recent report of a single individual of Cupressus
arizonica at Bridge Gap Spring in the Davis Mountains of Jeff Davis County,
Texas (Karges & Zech 2001) suggests a more continuous distribution in suit-
able habitats from California, through the southwestern United States, into
Trans-Pecos Texas, and south into Mexico. Previously, C. arizonica was known
from northern Mexico (Standley 1920; Wolf and Wagener 1948; Little 1978
Correll & Johnston 1979; Rehfeldt 1997) southern Brewster County, Texas (Pow-
ell 1988; 1998), and isolated localities in Arizona, California, and New Mexico
(Sudworth 1927; Vines 1960; Little 1971; Minnich & Everett 2001). The Davis
Mountains specimen led Karges and Zech (2001) to postulate a greater Pleis-
tocene range for C. arizonica, represented by an occurrence in this additional
Madrean sky island habitat.
The specimen upon which this phytogeographic postulate is based (Karges
& Hedges 2480, SRSC) may not represent Cupressus arizonica, however. This
SIDA 20(2): 585 - 592. 2002
586 BRIT.ORG/SIDA 20(2)
sheet consists of several short (< 12 cm), separate, yellowed terminal branches,
and three small (<8 mm), detached woody scales. Another voucher of the Bridge
Gap putative Cupressus arizonica (Karges s.n., SRSC) consists of a single branch
with green leaves, but no visible reproductive structures. Common members of
the Cupressaceae that occur in the Davis Mountains are Juniperus deppeana
var. deppeana and J. pinchottii (Powell 1998). Also present in the Davis Moun-
tains, J. deppeana var. sperryi is an extremely rare endemic known from three
individuals (Adams 1973; 1993; Watson & Eckenwalder 1993; Powell 1998). The
Bridge Gap specimens could represent one of the above taxa, as concluded by
B.L. Turner in an annotation (17 Jan 2001) of Karges and Hedges 2480 as Juniperus
deppeana. The distal vegetative growth of Cupressus arizonica and the above
taxa of Juniperus are nearly indistinguishable. This may reflect on the sister
relationship between these genera (Brunsfeld et al. 1994; Gadek et al. 2000).
The putative occurrence of Cupressus arizonica in the Davis Mountains is
of biogeographic importance, and we attempt to address the veracity of the re-
cent report in Jeff Davis Co, Texas, through an alternative means. Cupressaceous
taxa that are difficult to identify by conventional means have been accurately
identified through chemical analysis (Gough & Welch 1978; Adams 1993; Hsiang
& Huang 2000). In order to verify the determination of the two Davis Moun-
tains putative Cupressus arizonica specimens, we have gathered and compared
chloroplast DNA sequence data from these specimens and specimens of puta-
tive related taxa.
METHODS AND MATERIALS
The Davis Mountains specimens were compared to 4 individuals representing
Cupressus arizonica, 2 other Cupressus species, 1] individuals of Juniperus, and
1 specimen each of the outgroups Calocedrus, Chamaecyperis, and Thuja (Table
1). This sampling was based on the generic relationships within Cupressaceae
suggested by recent studies (Brunsfeld et al. 1994; Gadek et al. 2000). Specimens
used for DNA sequencing were either collected during fieldwork in Mexico and
the United States in 2001, previously vouchered herbarium specimens, or
vouchered live plantings growing at Rancho Santa Ana Botanic Garden
(RSABG).
For all specimens collected in 2001, DNA was extracted using 2X CTAB,
followed by precipitation in cold isopropanol (Friar et al. 1996). One gram of
terminal shoot tissue (including leaves) was used for these extractions. For speci-
mens collected before 2001, a modified small prep for dried leaf tissue was used
for extractions (Doyle & Doyle 1987). Amplification of cpDNA templates of the
trnL intron and trnL-trnF intergenic spacer follows the methods outlined by
Porter et al. (2000). Purified template amplifications were sequenced directly
with four primers, trnLc, trnLd, trnLe, and trnLf (Taberlet et al. 1991), using “big
GRIFFITH AND BARTEL, PHYLOGENETIC ANALYSIS OF CUPRESSUS ARIZONICA IN TEXAS
Taste 1. Specimens used in molecular analysis.
Species
Location
Voucher
Calocedrus decurrens Torr.
Chamaecyparis lawsoniana(A. Murr.) Parl.
Cupressus arizonica Greene
cf Cupressus arizonica Greene
Cupressus lindleyi Klotzsch ex Endl.
Cupressus macnabiana A.Murr.
Juniperus coahuilensis (Martinez)
Gaussen ex Adams
Juniperus deppeana Steud.
Juniperus flaccida Schlecht.
Juniperus jaliscana Martinez
Juniperus monticola Martinez
Juniperus pinchottii Sudw.
Juniperus virginiana L.
Thuja plicata Donn ex D.Don
Cauirornia. Los Angeles Co.:
Catirornia. Del Norte Co.:
CAurorNia. San Diego Co.:
Texas. Jeff Davis Co.:
Texas. Jeff Davis Co.:
Mexico. D.F::
Mexico. Michoacan:
Catirornia. Shasta Co.:
Texas. Brewster Co.:
exas. Brewster Co.:
Mexico. Puebla:
Texas. Jeff Davis Co.:
Texas. Jeff Davis Co.:
Texas. Brewster Co.:
Mexico. Coahuila:
Texas. Jeff Davis Co.:
Texas. Bastrop Co.:
Catirornia. Humboldt Co.:
RSABG 13148 (RSA)
RSABG 14672 (RSA)
Bartel 571 (SRSC)
Bartel 572 (SRSC)
Warnock 7138 (SRSC)
arg
(SRSC)
Karges s.n. (SRSC)
Griffith 325 (RSA)
Griffith 287 (RSA)
RSABG 11434 (RSA)
Bartel 575 (SRSC)
Powell 5146 (SRSC)
Griffith 324 (RSA)
Powell 5186 (SRSC)
Keough 205 (RSA)
Griffith 352 (RSA)
RSABG 13452 (RSA)
RSABG 11746 (RSA)
rges and Hedges 248
dye” chemistry from Applied Biosystems Incorporated, according to the
manufacturer's specifications. All sequences were gathered using an Applied
Biosystems Incorporated 3100 automated DNA sequencer.
Chromatograms derived from sequencing were assembled into contigs, and
edited using Sequencher v4.1 (Gene Codes Corporation, Inc.). Consensus se-
quences were manually aligned using Se-Al v2.0a72 (Rambaut 1996). The
aligned DNA data matrix is available from the first author upon request. The
phylogenetic relationships among these specimens were estimated using Fitch
parsimony, in PAUP* v4.088 (Swofford 1998). Estimations of confidence in the
clades were obtained through bootstrap analysis (Felsenstein 1985) with 10,000
pseudoreplicates, and through jackknifing (Farris et al. 1996), also with 10,000
pseudoreplicates (63% deletion) as implemented in PAUP™.
RESULTS AND DISCUSSION
The specimens of Cupressus arizonica form a well-supported (94% bootstrap,
80% jackknife) monophyletic group with three other Cupr pecimens, an
94
BRIT.ORG/SIDA 20(2)
C. arizonica Warnock 7138
C. arizonica Bartel S71
a C. arizonica Bartel 572
—
—
poems (CO indieyi Griffith 287
C. lindleyi Griffith 325
80
C. arizonica RSABG 11746
J. deppeana Powell S186
J. deppeana Griffith 324
J. deppeana Kalle S71
J. coahutlensis Powell 5146
J. coahuilensis Bartel 575
89 |
J. pinchottii Keogh 205
J. jaliscana Griffith 279
J. monticola Griffith 304
J. flaccida Bartel 574
J. flaccida Griffith 253
J. virginiana Griffith 352
Calocedrus
Chamaecyperis
Fic. 1. A phylogeneti lysis of th
Thuja
a al
iene (CC napDIGNA RSABG 11434
cf C. arizonica Karges s.n.
cf C. arizonica Karges and Hedges 2480
jE AEN
osaiimammaiiiaiiaeaiiel
ARN dK, \ " | 1 Ctr
f
£ICQIVAN . . £
A Ub.
J
bound analysis of the te ‘ Wealon With ey ntoGnans Cisiactels excluded, Jengsh— a d= — leds a
0.6
rescaled consistency =
and iac
. 8854;
port above 50% is ol below the branches: = Cupressus, J.= Juniperus.
GRIFFITH AND BARTEL, PHYLOGENETIC ANALYSIS OF 589
this clade excludes the two Bridge Gap putative C. arizonica specimens (Fig. 1).
The three specimens of Juniperus deppeana plus the two Bridge Gap specimens
form a clade, though lacking strong support (63% bootstrap). Juniperus
deppeana is a species present at Bridge Gap (Karges & Zech 2001). In context
with the known flora of the Davis Mountains, chloroplast DNA data clearly
suggest that the two Bridge Gap specimens are not Cupressus arizonica, and are
most likely Juniperus deppeana.
The first author examined the Bridge Gap cypress closely in July 1999 and
made the following observations: The specimen’s trunk bark is divided into
rough squarish plates near the base, characteristic of Juniperus deppeana var.
deppeana (Powell 1988). Ata height of about 0.5 m, the gray-brown bark is sepa-
rated into longitudinal ridges, a key character for Juniperus deppeana var.
sperryi (Adams 1973; Powell 1988). In contrast, the inner bark of Cupressus
arizonica is often described as reddish (Wolf & Wagener 1948; Powell 1998),
maturing often into thin fibrous strips on large trees (Correll & Johnston 1979;
Bartel 1993), although the bark of Cupressus arizonica sensu lato can be quite
variable. Martinez (1963) notes that J. deppeana var. patoniana forma obscura
may have the lower bark checkered and the upper bark furrowed.
Although other characters may be of use, in most taxonomic keys the ma-
jor feature diagnostic between Juniperusand Cupressus is the presence of fleshy,
fused cones (Juniperus), versus woody, dehiscent cones (Cupressus) (Correll &
Johnston 1979; Powell 1988: Bartel 1993, Watson & Eckenwalder 1993). Of the
two specimens of the Bridge Gap putative Cupressus, one (Karges s.n.) is devoid
of reproductive features and the other (Karges & Hedges 2480) has three sepa-
rate orbicular-spathulate scales that are 5-7 mm long and 6-8 mm wide. These
scales appear to have been sessile at the proximal end rather than peltate.
Cupressus arizonica has 4-5 partially peltate proximal pentagonal scales ap-
proximately 13-15 mm long, 12-14 mm wide, and 2-3(-4) distal oblong, trun-
cate scales 5-8 mm wide and 9-11 mm long, completely peltate and valvate
(Table 2). Morphologically, the scales present on Karges & Hedges 2480 do not
appear to be produced by C. arizonica.
One of the two putative Cupressus arizonica specimens (Karges s.n.) may
be misdetermined because of a lack of reproductive characters, while the other
(Karges & Hedges 2480) may have been determined as C. arizonica based on the
three woody scales collected. Although the location of discovery of these three
scales is faithfully recorded, we cannot be certain of their identity. Although
unfortunate, this is not the first instance of a mixed collection being mistaken
for something more significant (Thomson 1991). Given the above evidence, the
present known distribution of Cupressus arizonica in Texas remains limited to
the Chisos Mountains of southern Brewster County. This may change if a speci-
men exhibiting clear characters of C. arizonica is collected elsewhere in Texas.
590 BRIT.ORG/SIDA 20(2)
Tape 2. Specimens of Cupressus arizonica measured for morphological discussion.
Location Voucher
California: San Diego Co. Everett 33264 (RSA)
Mexico: Sonora. Wolf 2564 (RSA)
Texas: Brewster Co. Bartel 571 (SRSC)
Texas: Brewster Co. Bartel 572 (SRSC)
Texas: Brewster Co. Warnock 7138 (SRSC)
ACKNOWLEDGMENTS
The authors wish to wholeheartedly thank Sharon Yarborough of SRSC and
Steve Boyd of RSA for making herbarium specimens available; Bart O’Brien of
Rancho Santa Ana Botanic Garden for making live material available; Travis
Columbus, Naomi Fraga, Clem Hamilton, Mark Porter, Mike Powell, Linda
Prince, Smita Sanbui, and Gary Wallace for providing very useful comments
ona draft of this manuscript; and we are especially indebted to Nancy Refulio
for providing the resumen. Two anonymous reviewers provided excellent feed-
back. We are grateful for a Mellon Foundation grant through Rancho Santa Ana
Botanic Garden, which supported this work.
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non-coding regions of chloroplast DNA. PI. Mol. Biol. 17:1105-1109.
THOMSON, K.S. 1991. Piltdown Man:The great English mystery story. Amer. Sci. 79:194—201.
Vines, R.A. 1960. Trees, shrubs,and woody vines of the Southwest. University of Texas Press,
Austin.
592 BRIT.ORG/SIDA 20(2)
Watson, F.D.and J.E. ECkeNwALbeR. 1993.Cupresseaceae Bartlett: Redwood or Cypress Family.
In Flora of North America Editorial Committee, ed. Flora of North America Volume 2:399-
422.Oxford University Press, New York.
Wotr, C.B.and W.E. Waconer. 1948. The New World Cypresses. Aliso 1:1-144,
DAHLIA NEGLECTA (ASTERACEAE: COREOPSIDEAE), A
NEW SPECIES FROM SIERRA MADRE ORIENTAL, MEXICO
Dayle E. Saar
Department of Biological Sciences
Northern Illinois University
DeKalb, IL 60115-2861, U.S.A.
ABSTRACT
Dahlia neglecta, a new species from the state of Hidalgo, Mexico, is described and illustrated. This
species is distinguished by the combination of glaucous stems, tufts of light-colored hairs at the
rachis nodes, strongly oblique blade bases, and mucronate ligules.
RESUMEN
Sed | l da Dahl l le] do de Hidalgo, México. Esta especie
oO [
se distingue por la combinacién de tallos glaucos y un copete de pelos de colores suaves en los nudos
del raquis. Ademas, las bases de las hojas pequenas son dominantemente oblicuas, y con ligulas
mucronadas. Se ofrece una clave del género.
While collecting material from wild populations of Dahlia to expand natural
history information and for cytological and molecular analyses of the genus,
several undescribed species from Mexico were encountered, one of which is
described here.
Dahlia neglecta D.E. Saar, sp. nov. (Fig. 1). Type: MEXICO. HipaALco: ca. 7 km NE of
Tulancingo, just below the first pass along Mex 130 to the gulf coast, zone of Quercus and
arborescent Opuntia; SE-facing slopes, among rocks and recent road fill, sunny to mostly
sunny locations, elev. 2320 m, 11 Sep 1995, J.P. Hjerting, D.E. Saar, & PD. Sorensen 95-86 (HOLO-
TYPE: MEXU; ISOTYPES: C, DEK, F IEB, MO, TEX).
Herba perennis 1.1-1.5 m alti. Folia pinnata vel bipinnata. Multiflorus, capitula 10-12.5 cm diam.,
corollae radii ligulatae lavendulus, squammis exterioribus involucri reflexis sub anthesi
Distinguibilis a caules glaucedine, fasciculati densi ad rachis nodis, obliqus basis laminis, ligulae
mucronatusque necnon.
Plants to 1.5 m (type locality) or 1.2 m (paratype population) with 1-3(-5) her-
baceous stalks arising from perennial tuberous rootstock; internodes glaucous,
ollow, nodes solid. Median leaves 19-24 cm long including petiole, bipinnate
with stipels frequently at the second (occasionally second and third) rachis
node, becoming pinnate without stipels to simple at the base of the flowering
portion; petioles solid; rachis with prominent tufts of light-colored hairs at
nodes, pinnules opposite on the rachilla, primary pinnae 5-7; leaflets dentate
and ciliolate, blade bases strongly oblique, veins lighter-colored, sometimes cre-
ating a dusty appearance at first glance. Capitula to 12.5cm in diameter includ-
ing rays, borne on unbranched peduncles 6-19 cm long and projecting above
SIDA 20(2): 593 — 596. 2002
594 BRIT.ORG/SIDA 20(2)
the foliage; involucral bracts reflexed at anthesis. Ray florets sterile; ligules
lavender with a spot of yellow at the base, veins darker below, tips mucronate.
Disc florets hermaphroditic, yellow. Cypselae elliptic-oblong, 8.6 mm wide x
1.2-1.6 mm long; black.
Etymology and distribution. —Flowering commences in August (in 1995).
Plants from the type locality grow along and in full view of a busy highway
(Mex 105) near the large city of Tulancingo. It is surprising, therefore, that this
showy species has not been described previously, hence the specific epithet,
neglecta. The known distribution for this species is presently limited to the
holotype and paratype localities.
Paratype: MEXICO. Hipatco: at K-8, 2 km SW of Mineral del Monte, along route Mex 105, zone of
Cupressus and scattered shrubby Quercus; road cut with dense vegetation on SE-facing slope, elev.
2800 m, Ll Sep 1995, J.P Hjerting, D.E. Saar, & PD. Sorensen 95-97 (DEK, MEXU).
The combination of glaucous stems, tufts of light-colored hairs at the rachis
nodes, strongly oblique blade bases, and mucronate ligules sets this species
unmistakably apart from all others. Sorensen’s (1969) key would place Dahlia
neglecta in section Dahlia. Within the key for this section, D. neglecta pairs with
D. australis and D. sherffii at couplet “N” (page 325) and incorporates some
morphological characters from each species. Dahlia neglecta is similar to D.
australis in that the lower leaf surfaces are conspicuously lighter green or sil-
very green, but the former differs by having flat (not revolute or obscurely so)
margins, a smooth (not rugose) upper leaf surface and glaucous (not glabrous
or pubescent) canes. The fruit is of similar length but only 1.2-1.6 mm wide
versus 1.5-2.6 mm for D. australis. The smooth upper leaf surfaces of D. neglecta
are similar to D. sherffii. Dahlia neglecta differs from D. sherffii by its glaucous
(not glabrous) canes, bicolored leaf surfaces, and narrower fruit (D. sherffii =
2.2-3 mm). The unique combination of characters mentioned earlier also dis-
tinguishes D. neglecta. None of these species are sympatric in their distribu-
tions.
A molecular study of the genus which utilizes combined molecular se-
quences from the internal and external transcribed spacer regions ITS and ETS,
respectively) of nuclear ribosomal DNA, shows good support (86% bootstrap
value) for placing Dahlia neglecta in a pectinate clade (Variable Root Clade”)
that includes the “tree dahlias” (sect. Pseudodendron: D. imperialis, D. excelsa,
and D. tenuicaulis), sect. Epiphytum (D. macdougallii), two species from sect.
Dahlia (D. rudis and D. apiculata), and an unpublished species (Saar et al. in
press). Within this clade, D. neglecta is most closely allied with D. apiculata and
D. tenuicaulis, but support is minimal (57% bootstrap value). Dahlia neglecta
is not sympatric with these species and both D. apiculata and D. tenuicaulis
have lignified perennial canes, whereas D. neglecta is herbaceous.
The description of Dahlia neglecta brings the number of “wild” or natu-
rally-occurring species of Dahlia to 35, but does not include the cultivated forms
595
SAAR, A NEW SPECIES OF DAHLIA FROM MEXICO
Fic. 1. Dahlia neglecta: A. leaf bases: B. flowering head: C. median leaf: D. general habit. D
a
photographs.
596 BRIT.ORG/SIDA 20(2)
often called D. variabilis or occasionally D. pinnata (see Hansen & Hjerting
1996 for clarification of the latter Latin binomial). This total does, however, in-
clude three previously undescribed species currently “in review” or “in press:”
D. hjertingii (Hansen and Sorensen in press), D.campanulata, and D. cuspidata.
ACKNOWLEDGMENTS
| thank Nathan A. Saar for preparing the illustration and Portia Gallegos for
translating the abstract into Spanish. Paul Sorensen and Jens Peter Hjerting were
invaluable throughout the field studies in Mexico. Guy Neson and an anonymous
reviewer provided helpful comments to an earlier version of the manuscript.
REFERENCES
Hansen, H.V. and J.P. Huertinc. 1996, Observations on chromosome numbers and biosys-
tematics in the genus Dahlia (Asteraceae, Heliantheae) with an account on the iden-
tity of D. pinnata, D.rosea and D.coccinea. Nordic J. of Bot. 16:445-455.
Hansen, H.V.and PD. Sorensen. (In press).A new species of Dahlia (Asteraceae, C ideae)
from Hidalgo state, Mexico. Rhodora
Saar D.E., N.O. Potans, and P.D. Sorensen. (In press). A phylogenetic analysis of the genus
Dahlia (Asteraceae) based on internal and external transcribed spacer regions of
nuclear ribosomal DNA. Syst. Bot.
Sorensen, P.D. 1969. Revision of the genus Dahlia (Compositae, Heliantheae—Corepsidinae).
Rhodora 71:309-365, 367-416.
THE VARIETIES OF LIATRIS ELEGANS (ASTERACEAE)
Mark H. Mayfield
Herbarium-Div. of Biology
Kansas State University
Manhattan, KS 66502, U.S.A.
markherb@ksu.edu
ABSTRACT
A desc cription of two new inf I 1, Liatris elegans var. bridgesii,
var. nov. occurs on the Carrizo and adjacent EOCERS sand fi rmations in central ae Liatris elegans
Var. kralii, var. nov, occurs on the Fall | stal plain of Alab to the
7
northern panhandle of nee The newly described taxa are similar to each other in their cream-
white to yellow (vs. anthocyanin-colored) phyllary apices but they are widely separated g geographi-
cally and each is distinct in other morphol four they are viewed here as having indepen-
dent origins. Liatris elegans var. flabellata is reduced toa form of L. elegans var. elegans, while Liatris
elegans var. carrizana is maintained for southern-central Texas populations. Liatris elegans var.
elegans is widespread on the coastal plain from South Carolina to eastern Texas.
Key Worps: Liatris, Asteraceae, Texas
RESUMEN
Se presenta la descripcion de dos nuevos taxa infraspecificos en Liatris elegans. Liatris elegans var.
bri meses var. nov, se da en el Carrizo y formaciones arenosas adyacentes del Eoceno en el centro de
Texas. Li gans Var. kralii, var. nov. oe enla - Line > Sand ey llanura costera de Alabama
y Georgia hasta el estrecho de Florida ntre si por sus apices de
los filarios de blanco-crema a amarillo (vs. color = antonio) pero estan au anens sane
geograficamente y se diferencian
independientes. Liatris elegans var. = flabel ata se ha reducido a una a foams de L. lesan var, Sex ns,
mientras que Liatris elegans var. carrizanad se mantiene para las poblaciones del sur-centro de Texas.
Liatris elegans var. elegans es comtn en la Ilanura costera desde Carolina del Sur al este de Texas.
Liatris elegans Michx. is an obligate psammophile, occurring in well-drained,
loose to somewhat compacted sandy soils in open to sparsely forested habitats
across much of the Gulf and Atlantic coastal plains and in the southern Inte-
rior Highlands province in Arkansas. Its colored, elongated, conspicuous phyl-
lary apices in combination with white to yellowish-cream corollas distinguish
it as one of the most recognizable species in the genus (cf. King @ Robinson
1987). All other species have more or less herbaceous, relatively short, incon-
spicuous phyllaries with exserted, lavender to pink corollas (Gaiser 1946). Thus,
in L. elegans the phyllaries rather than the flowers provide primary visible col-
oration of the columnar inflorescences. Throughout most of the range of L.
elegans (herein, L. elegans var. elegans and L. elegans var. carrizana L.O. Gaiser)
populations have phyllaries that are predominantly shades of lavender, pink,
bluish, or magenta, colors that are presumably generated by anthocyanin
SIDA 20(2): 597 — 603. 2001
598 BRIT.ORG/SIDA 20(2)
pigments. Two sets of populations, one on the western margin of the range in
Texas, another in the eastern Gulf Coastal Plain in Alabama, Georgia, and
Florida, mostly lack classic anthocyanin coloration at anthesis. Instead, these
have primarily yellowish, cream, or nearly white phyllaries with some popula-
tions variably cream to faintly lavender. The Texas populations were discussed
briefly by Gaiser (1946, p. 174, as L. elegans forma fisheri Steyermark), but accu-
rate description of the variation is lacking. In addition, the well-defined pat-
tern of distribution of the eastern coastal plain pale-phyllaried populations has
never been addressed. While phyllary color alone may not be taken asa reliable
indicator of genetic cohesion, the geographic integrity and populational uni-
formity of these unique populations suggests that selection for phyllary color
has taken place within these populations. A recognizable suite of other mor-
phological features supports their recognition but the presence of intergrada-
tion of both sets of populations with var. elegans further suggests that recogni-
tion at an infraspecific rather than specific rank is warranted. | have chosen
variety to indicate that intermediate populations occur along the margins of
these novel taxa.
Liatris elegans Michx. var. bridgesii Mayfield, var. nov. (Fig. 1). Type: US.A. TEXAS.
M Co.: 2.6 mi W of [FM 487 in] Rockdale on US 79; S side of railroad track, Simsboro Sand
Formation, 505 ft, N 30°38°34" W 97°02’52", 6 Sep 1992, Mark H. Mayfield 1549 (HOLOTYPE:
TEX; ISOTYPES: F FLAS, GA, GH, MO, NCU, NY, US).
Liatris elegans forma fisheri Steyermark, Field Mus. Nat. Hist., Bot. Ser. 11: 275. 1936. TYPE: TEXAS:
CORYELL Co. near and on the highway, 4 mi S of Copperas Cove, 26 Aug 1934, George L. Fisher
s.n.CHOLOTYPE: F #761244!)
Differt a L. eleganti var. eleganti apicibus phyllariis petaloideis flavis aut eburneis vel pallide roseis
aut lavandulis; aient a Sa eleganti var. eens apicibus phyllariis valde recurvatis ac capitulis
CUAaACCI v
Corms eae stems ay to densely tomentose; lateral heads usually
densely crowded, divergent-ascending t ding at right angles, sessile; phyl-
laries parallel and appressed below the petaloid portion, the colored apices re-
curved, abaxially pubescent to tomentose, initially pale lemon yellow, then fad-
ing to cream or very pale lavender, rarely rosescent at fruiting; corollas
cream-white, tubular, the tube scarcely broadened to the top of the limb, the
lobes scarcely flared apically; stigmas cream to light yellow or nearly white.
ParATYPES: U.S.A. TEXAS. Anderson Co.: George Engling Wildife Manag Area, Upper North Pas-
ture, W Side, 1954 Burn, 18 Sep 1958, Marsh 55-8 (TEX). Bastrop Co.: 1.8 mi SE of McDade ane on
highway by railroad face 30 Jul 1992, Mayfield & Westlund 1486 (NCU, NY, TEX). Freestone Co.
1848, 5.8 mi N of Buffalo at jet. US 79, lower Queen City formation, N 31°32’46" W 96°03'22", 6 -~
1992, “Mayfield 1557 (F, FLAS, GA, GH, NCU, NY, TEX, US); 14 mi $ of Fairfield, 15 Aug 1955, Turner
aa eal ay Grimes Co.: along main highway E of Brazos county line, 4 Sep 1993, Mayjieid etal.
(TEX). Hardin Co.: 2.5 mi W of Silsbee, 2 Oct 1945, Cory 49880 (BRIT). Henderson Co.:
“ oe grassy open oak woods in sandy soil, flowers whitish, 20 Oct 1963, Correll 28558 (LL).
MAYFIELD, VARIETIES OF LIATRIS ELEGANS (ASTERACEAE) 599
Fic. 1. Liatris elegans var. bridgesii. A. Habit. B. Close-up of
heads showing yellow to cream phyllary apices.
Fic. 3. Liatris elegans var. carrizana with consistently Fic. 2. Liatris elegans var. elegans with phyl-
colored blue, magenta, lavender, or pink, rather than cream _laries that are predominantly shades of lav-
phyllaries. ender, pink, bluish, or magenta.
Fic. 4. Liatris elegans var. kralii.
600 BRIT.ORG/SIDA 20(2)
Houston Co.: Davy Crockett N. Forest, N of SR 7 froma ee 2.6 mi W of the junction of FM 357 and
0.9 mi E of FM 1733, 21 Sep 1991, Mayfield et al. 1123 (NCU, NY, TEX); 2 mi S of Grapeland, 12 Oct 1937,
Cory 26136 (F). Lee Co.: NW of Lexington, on county ieee to a ears Cemetary, by small pond, Carrizo
loxmation, 500 ft, N 30°27°30" W 97°03'15", 15 Sep 1991, Mayfield et al. 1117 (GH, NCU, NY, TEX). Leon
NE of Buffalo along hwy 79, 11 Sep 1958, Turner 4433 (TEX). Limestone Co.: 5.5 mi E of
ee 9 eee 1964, Shinners 30579 (BRIT). Nacogdoches Co.: 4 mi W by N of Garrison, 7 Sep 1948,
Cory 54734 (LL). Robertson Co.: county road 330 ca 2.0 mi E of its junction with FM 2246, 500 ft, N
31°0134" W 96°2159", 6 Sep 1992, Mayfield 1559(GH, NCU, NY, TEX). San Augustine Co.: sandy open
forest ca. 1.5 mi N of San Augustine, flowers whitish, 14 Oct 1962, Correll 26282 (LL). Shelby Co.: ca
0.8 mi SE of SR 7 ona parallel unpaved road, ca 0.1 mi W of a small lake on Upper Iron Ore Cr, 1.2 mi
E of Mt Hermon, 21 Sep 1991, Mayfield et al. 1130 (NCU, NY, TEX). Van Zandt Co.: Ca. 1.5 mi NW of Ben
Wheeler on SR 64, 0.2 mi S of SR 64 opposite a cemetary on the N side of the road, 22 Sep 1991,
Mayfield et al. 1138 (F GH, NCU, NY, TEX). Walker Co.: ca. 3 blocks E of Sam Houston St Un, 27 Sep
1980, Kessler 3679 (FLAS). Williamson Co.: 0.7 road mi N of FM 112 on county road, ca 0.8 mi W of
Milam Co. line and 9 miS of Thorndale, Simsboro Formation, 530 ft, 30°29°24" N, W 97°L0°40", 15 Sep
1991, Mayfield et al. 1120 (F GH, NCU, NY, TEX). Wood Co.: Lake Ellis, 5 mi NE of Crow, 2 Sep 1942
Lundell 11730 (LL)
Forma asa category fails to represent the clear geographic pattern that occurs
in var. bridgesii. | have chosen to provide a novel epithet for the present taxon
because the type locality of forma fisheri is apparently inaccurate. No popula-
tions of L. elegans var. elegans (Fig. 2) occur anywhere close to the designated
location “on the highway 4 miles south of Copperas Cove” in Coryell County.
While there are some thin, Cretaceous sand-bearing strata in this area as well
as some river terrace sands, searches around Copperas Cove have not yielded
populations or habitat suitable for L. elegans. Furthermore, no other collections
from the area were located in searches of three major herbaria in central Texas,
among others. The establishment of a new type and name will serve to solidify
the concept of the taxon.
Etymology.—This variety is named for Edwin L. Bridges, prolific plant col-
lector and knowledgeable floristician/taxonomist. Bridges’ tenacity in the field
and his incredible plant recognition skills are truly inspiring. Our memorable
field trips into the sand-bearing strata east of Austin, Texas, were crucial to my
understanding of this and many other sand-loving species.
Distribution, variation and ecology—Liatris elegans var. bridgesii is best
represented in the southwesternmost part of its range from north of the Colo-
rado River in Bastrop and Lee Counties northeast to Henderson County in Texas.
Here, populations consist entirely of individuals with phyllaries that are creamy
white to very pale lavender, and possess long-tubed corollas with relatively short
weakly flared lobes. In the Post Oak savannas of Williamson, Milam, Lee, and
Robertson counties, all populations I found had consistently cream-colored
phyllaries. The latter begin the flowering season with a very striking, nearly
lemon yellow color in the phyllaries, a color that is otherwise not present in the
phyllaries of any Liatrinae. Later in the season, well into fruiting, the color fades
to creamy white, sometimes infused with a rose or lavender tinting. Further
MAYFIELD, VARIETIES OF LIATRIS ELEGANS (ASTERACEAE) 601
northeast, populations seem to intergrade in terms of phyllary color but other
features are consistently more like var. bridgesii than var. elegans. The phyllar-
ies of var. bridgesii tend to be longer and more strongly recurved with longer,
more narrowly tubular, pale cream corollas than in var. elegans. The var. bridgesii
is similar to the next newly described variety in the pale coloration of the phyl-
laries, but it has a more compact inflorescence with more strongly recurving
phyllary tips and sessile heads (see further discussion below regarding pedun-
culate heads).
The var. bridgesii occurs almost exclusively on Eocene sand-bearing strata
running from southwest to northwest from the north side of the Colorado River
in Bastrop County to Wood County in the northeast. Its influence is also appar-
ent in population eastward through Cherokee and Houston, to Nacogdoches,
Shelby, and San Augustine Counties where the Carrizo formation is exposed
around the western margin of the Sabine uplift. Post-oak savanna is the pri-
mary habitat where this variety is found. Mixed pine forests of eastern Texas
are more likely to have populations of L. elegans var. elegans (Fig. 2) but both
occur therein and some intergradation is observed.
Liatris elegans var. carrizana [_.O. Gaiser (Fig. 3) occurs south of the Colo-
rado River in Texas on deep, sandy soils of xeric sandhills (cf. Gaiser 1946). This
variety is characterized by its elongated, napiform corm suited to the drier cli-
mate and deep sands that prevail in this region. Its phyllaries are structurally
similar to those of the var. bridgesii, with elongated strongly recurving apices,
but they are consistently colored blue, magenta, lavender, or pink, rather than
cream. Populations of var. carrizana just to the south of the Colorado River in
Bastrop and Caldwell Counties, Texas, may have more or less globular corms
but are otherwise more similar to the var. carizzana. In all other areas, L. elegans
has depressed globose corms as is true of many species of the genus.
Liatris elegans Michx. var. kralii Mayfield, var. nov. (Fig. 4).Type: U.S.A. ALABAMA.
LEE Co. Locally abundant in open pine woods 6 mi S of Auburn, 23 Sep 1899, ES. Earle and
E.S. Earle 94 (HOLOTYPE: KSC} tsoTyPes: F!GH!ND NY! US).
pa
] .. - ed peer ca : lhic
(* 3 LT I >
Similisa L. eleganti var. bri | pallide lavandulis sed capitulis
| +1 ] | soe
] ‘ cee sf } 1]
oO Le c
petaloideum z leniter recur Jat ad asieas
Corms globose; stems sparsely to densely lanose-tomentose; lateral heads (at
anthesis) usually loosely arranged, ascending to divergent, pedunculate; phyl-
laries abaxially tomentose to densely lanose, divergent along their length, the
petaloid apices flaring only slightly or not at all, initially light yellow or cream,
then fading to cream; less often pale lavender to rose, corollas pure-white, tu-
bular, the tube flaring gradually to the top of the limb, the lobes strongly flared;
stigmas white to very pale lavender.
ParAtyPes: U.S.A. ALABAMA. Baldwin Co.: Gateswood, 30 Oct 1903, Tracy 8568 (FGH). Barbour Co.:
602 BRIT.ORG/SIDA 20(2)
6.4 mi N of the Barbour/Dale County line along Alabama route 51, ca. 4.5 mi N of Doster, Turkey
Oak /Bluejack Oak/Hickory sandhill area, 495 ft, 31°40°55" N, 85°38°10" W, 12 Aug 1994, Mayfield &
Fuertes 201 (TEX). Covington Co.: Dozier, 31 Oct 1941, Reed 2053 (TEX). Escambia Co.: dry, sandy
area above low hillside seepage bog re powerline, on N side of western extension of Co Rd 6, 2.4
mi W of Co Rd 27 and 2.6 mi W of AL 113 ata point 3.8 mi N of int US 31 E of Flomaton, just W of
Pond Cr, SH SWQ SEQ of sec 1, TIN RTE, sees 7.5' quad, Soils-Plummer (Grossarenic paleaquults),
elev 130 {t, 31°04'14" N, 87°19°OL" W, LL Sep 1989, Orzell and Bridges 11815 (TEX); 3 mis of Canoe, 17
Oct 1929, O'Neill 6171 (FLAS). Geneva Co.: Sandy west banks of Choctawhatchee River between
Geneva and Eunola by Al 52,18 _ 1971, Kral 44342 (NY). Lee Co.: 2.1 mi NW of Phenix City, 17 bel
1971, Kral 44247 (NY). Mobile USA Property, 17 Oct 1968, me 879(NCU). Russell Co.: 1 n
of Glenville, 26 Sep 1976, ee (NCU). FLORIDA. Okaloosa Co.: Crestview, 29 Sep 1946, ae
sn. (NCU). Santa Rosa Co.: pine woods on Juniper Creek, Rd. i 6 Oct 1949, Hood 3226 (FLAS). Wash-
ington Co.: vicinity of Falling Waters Sink, longleaf pine deciduous scrub oak forest, 17 Sep 1956,
Kral 3574 (GH). GEORGIA. Ben Hill Co.: 7 mi NNW of Fitzgerald, 26 Oct 1968, Faircloth 5691 (NCU).
Coffee Co.: 7.2 mi E of ridge on outcrop of Altamaha Grit, 17 Oct 1970, Faircloth 6936 (NCU). Grady
Co.: Sandy clay ridge, open, 2 mi W of Wigham, 26 Oct 1963, Godfrey 63218 (LL). Macon Co.: Near
Marshallville, 28 Sep 1926, Dunegan 15 (LL). Meriwether Co.: Warm Springs, | Oct 1945, Totten etal.
s.n. (NCU). Muscogee Co.: Columbus, 17 Sep 1902, Biltmore Herbarium [no further collector informa-
tion on label] (NY). Taylor Co.: Sandy scrub oak habitat of deep sands of Fall Line Sand Hills, ca. | mi
NE of Charing and 11-12 mi SW of Butler on GA 137, 10 Sep 1974, Jones et al. 22598 (NCU); 3.8-4.0
mi N of Butler on high wand ridge along HWY 19, xeric longleaf pine/turkey oak sandhill wood-
land, Fall-Line Sandhills, 570 ft, 32°36'34" N, 84°14'15" W, Ll Aug 1994, Mayfield & Fuertes 1998 (TEX).
Etymology.—This variety is named to honor Dr. Robert R. Kral, well known as
one of the most knowledgeable botanists on the southeastern US flora. Kral’s
personal collections amassed over four decades from the coastal plain have con-
tributed greatly to the understanding of thisand many other species. His classes
| took as an undergraduate in Plant Taxonomy and Dendrology stimulated my
interest in botany.
Distribution, variation, and ecology.—Liatris elegans var. kralii occurs in
two major population centers. Populations to the north, in the Fall-Line
Sandhills of western-central Georgia to eastern-central Alabama, including the
type, consistently bear phyllary apices that are white to light tan in color. A
second, somewhat separated area of occurrence is further to the south in coun-
ties along the southern tier of Georgia, northern tier of Florida, and west to
Mobile County in Alabama. In this area scattered populations of var. kralii oc-
cur sympatrically (but not apparently in mixed populations) with var. elegans.
The var. elegans is much more common in this area but relatively pure popula-
tions of var. kralii do occur, albeit usually with both white and pale lavender
forms (e.g., Kral 37333, Mobile Co., AL; Kral 44342, Geneva Co., AL; Faircloth
5691, Ben Hill Co., GA; Kral 3574, Washington Co., FL). All populations refer-
able to var. kralii have short peduncles bearing the heads and the heads are
well-spaced and ascending. Although pedunculate heads may otherwise occur
throughout the range of the species in shaded or apically injured plants, these
are rare and unusual. Also, the phyllaries of var. kralii are longer and narrower,
with the petaloid portion especially elongate in proportion to the basal herba-
MAYFIELD, VARIETIES OF LIATRIS ELEGANS (ASTERACEAE) 603
ceous portion. Another recognizable trait of var. kraliiis the divergent phyllar-
ies with the tips ascending, rather than recurved. All other varieties have more
strongly recurved phyllary apices, with the lower herbaceous portions closely
appressed.
Variety flabellata.—The type specimen of var. flabellata apparently con-
sists of a single plant divided into two separate sheets. The only two sheets re-
ferable to this taxon are at NY, one has a single plant with root and two stems
attached and a third stem severed at the base. The second sheet is a single stem
that appears to have come from the severed base of the plant on the other sheet.
The only feature separating this plant from var. elegans is its emarginate to trun-
cate, relatively broadened (flabellate) phyllary apices. From the same locality
(St. Helena Island, S.C.) numerous other sp collected by Cuthbert at NY
and FLAS have phyllary morphology as is ae ee for the var. elegans. Thus, |
view the type specimen as consisting of a single plant from a locality with plants
otherwise identical to the var. elegans. Thus it is best treated as an aberrant or
freak, admittedly distinctive, but not persisting as a viable population. It is re-
duced here toa form of var. elegans.
Liatris elegans Michx. var. elegans forma flabellata (Small) Mayfield, comb. nov.
Laciniaria flabellata Small, Bull. Torrey Bot. Club 25:472. 1898. Liatris flabellata (Small) K.
Schum., Just’s Bot. Jahresber. 26:378. 1900. Liatris elegans (Walter) Michx, var. flabellata (Small)
Gaiser, Rhodora 48:345. 1946. TYPE: SOUTH CAROLINA. BEAUFORT Co. St. Helena Island, dry
barrens, Sept 1894, A. Cuthbert s.n. (HOLOTYPE: NY! ISOTYPE: NY).
ACKNOWLEDGMENTS
The curators of herbaria cited are acknowledged for loaning specimens and the
many field trips were in part supported by my graduate institution-Plant Re-
sources Center, Dept. of Botany, at the University of Texas at Austin. Thanks are
also due to my several field companions who so graciously gave their time to
allow me to collect Liatris elegans. Guy Nesom greatly improved an earlier draft
of the manuscript and provided the Latin translations for the diagnoses.
REFERENCES
Gaiser, L.O. 1946. Series VII. Elegantes. In: The genus Liatris.Rhodora 48:340-345.
Kina, R.M.and H. Rosinson. 1987.The genera of Eupatorieae (Asteraceae). Monogr. Syst. Bot.,
Missouri Bot. Gard. 22:1-581
604 BRIT.ORG/SIDA 20(2)
Book REVIEW
RONALD A. COLEMAN. 2002. The Wild Orchids of Arizona and New Mexico. (ISBN
0-8014-3950-7, hbk.) Cornell University Press. Sage House, 512 E. State Street,
Ithaca, NY 14850. (Orders: wwwcornellpress.cornell.edu). $39.95, 272 pp,
35 line drawings, 36 maps, 160 color plates, 6" « 9 1/4"
This volume is just one of many regional orchid floras published for North America within the last
15 years. Though the Native Orchids of the United States and Canada was published in 1975 by Car-
lyle Luer, regional floras, such as this one, update nomenclature and more clearly define geographic
ranges. Since Orchidaceae is one of the largest flowering plant families, has a cosmopolitan distribu-
tion, and occupies many cliverse
roe
jabitats, small regional floras, like this one, provide a baseline of
knowledge. Coleman presents to his readers a rather varied orchid flora in Arizona and New Mexico,
composed of 35 species and 14 genera. The most diverse region in both states for spotting orchids is
the Mogollon Rim. The combination of high elevation and oe ipitation bee habitat for orchids,
in comparison to the rest of the two states d high plains dominate. For those interested
in collecting orchids, you might think twice. All of the orchids of Arizona te afew in New Mexico
are protected by law, so all you should take away from your orchid-hunting adventures are photo-
graphs and memories.
The book begins with a lucid discussion of floral morphology, habitats and geographic distri
bution in Arizona and New Mexico, as well as information on blooming seasons ne conservation.
|
Next, a key to orchid genera is provided and could not be more straightforward. The terminology is
basic and would aes useful to anyone, even those isos only ra ies understanding of botanical termi-
iF 1]
| Genera and species are listed
]
N0
I
in eon For each genus enue: provi sitation of effective publication, etymol-
ogy and a rather informal description or discussion of the group. For example, in the discussion on
Cypripedium, the reader learns that the lip of the flower serves as a trap to the pollinator, who once
inside finds it hard to esc tl
>
—
discussion on Corallorhiza helps enthusiasts clear ae dis-
tinguish the members of . genus from those of Hexalectris. If there is more than one species pres
a key is provided. Following the generic discussions are species descriptions, which also ier a
citation of publication and etymology of the epithet. In addition, synonymy, a short non-technical
description, distribution, habitat, blooming season, conservation issues and threats to each species
are included. Because the author has taken an informal approach to describing each species, it be-
comes much more interesting to the reader. A case in pointis the interesting story on nomenclature
of Schiedeella arizonica and how misapplied names can lead to taxonomic contusion. In the margin
next to each species description is a line drawing of the flower. These illustrations provide a practi-
cal identification tool. A dot map for each species shows presence of that species in Arizona and New
Mexico counties.
Several apy 5 concl the book: 1) Excluded species, 2) Watch list of species not yet re-
ported from Arizona or New Mexico, 3) Herbarium collections studied, and 4) Distribution of or-
chids in Arizona and New Mexico listed by county. Color photographs seem strategically placed at
the beginning of the book, where they can whet the reader's appetite. Most of the photos are close-
ups of flowers or fruits, > but also include (> Sains ol Eee ants in napa
This isa wonderful pul | f information useful to
anyone with any affinity to orchids. een writing style is clear, concise, and unpretentious. It
would make a lovely addition to the library of a native plant lover, botanist, or orchid enthusiast.—
Amy lrauth Nare, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX, 76102 U.S.A,,
L@ [
amy trauth@yahoo.com.
SIDA 20(2): 604. 2002
DOCUMENTED CHROMOSOME NUMBERS 2002:3.
CHROMOSOME COUNTS FOR PACKERA PAUPERCULA VAR.
APPALACHIANA (ASTERACEAE: SENECIONEAE)
Alison McKenzie Mahoney Virginia W. Kong
Department of Biological Sciences
Minnesota State University-Mankato
Mankato, MN 560017, U.S.A.
Robert R. Kowal
Department of Botany, Birge Hall
0 Lincoln Drive
University of Wisconsin- he son
Madison, WI 53706, U.S
Cornell University
Ithaca, NY 14853, U.S.A.
ABSTRACT
Certain populations of more or less tomentose Packera (Asteraceae: Senecioneae) in southeastern
North America have recently | ated as P paupercula var. appalachiana based on evidence from
| ; ldi Hf ] t pa ] : Tal Ae] . |
8g ) & 8
a preliminary RAPDs analysis, and crosses. Previously they had been treated as disjuncts of P.
plattensis, a species of the Great Plains, which may have pans into the mid Appalachians enune
the post-glacial Hypsithermic Period and survived there as relicts. Howeve
with n= 46 and x
= 23. Our two chromosome counts of - new variety from North Carolina and
from Virginia are n = 22 and n= ca. 44, res e tively, a diploid and a coos both with x = 22.
Statistical analyses of p pollen dis evidence that the other individuals in the two © popu-
lations are diploid and mostly tetraploid, oa Hybridization with P aurea may be involved
n the origin of the tetraploid. Thus, cytological evidence supports other data showing that these
ane may be more closely related to P. paupercula.
ie)
a
RESUMEN
Ciertas Coan ae Packera (Asteraceae: Senecioneae) mas 0 menos tomentosas del sureste de
Norte
yaaa reciente meat como P. Be has var. apna lachiana pavandess en
oy F ]
y datos florals, un gnats preliminar de RA oe y cruces. piven panlap yee tratados c como
lant lra
Apala
medios Eat el eri hipstermico a Blac y sobrevivid alli como relict. Sin embargo E
plattensises tetraploide co
t
de Carolina del Norte y = vi irginie asonn=22 yn-~ca.44, respectivamente, un diploide y un fuaploue
ambos con x= 22 lel dia
lel pole evidencian que los otros individuos
de las dos poblaciones son diploidies,s pret La hibridacion
con P aurea puede ser la causa del origen del tetraploide. Asi, la ev idencia citolégica wcla otros
datos que muestran que estas p yblaciones pueden estar mas emparentadas con P. paupercu
INTRODUCTION
Packera plattensis (Nutt.) Weber and A Love is distributed in dry grasslands
from the eastern flanks of the southern Rocky Mountains to the Mississippi
SIDA 20(2): 605-610. 2002
606 BRIT.ORG/SIDA 20
p<
2)
River with scattered populations eastward in Wisconsin and Missouri (Barkley
1962, 1978) (Fig. 1). It is tetraploid with a base chromosome number of x = 23
(Kowal 1975; Kowal et al. unpubl. ms.).
Apparently disjunct populations occur in Pennsylvania, Virginia, Tennes-
see, North Carolina and South Carolina (Barkley1962, 1978, 1988; Uttal 1982;
Rhoads & Klein 1993) (Fig. 1). Barkley hypothesized that these are remnants of
the widespread eastern migration of prairie-adapted plants during the
Hypsithermic Period of about 8000 years ago. Such populations in Montgom-
ery Co., Virginia commonly hybridize with P aurea, P anonyma, and P obovata
(Uttal 1982). Because the latter three species are usually diploid with x = 22
(Kowal et al. unpubl. ms.), widespread hybridization with P plattensis would
be unexpected (Mahoney & Kowal ms. subm.; Kowal et al. unpubl. ms.). More-
over, herbarium specimens from the entire range of the eastern disjunct popu-
lations are morphologically different from P. plattensis of the Great Plains in
their rosette and cauline leaves and in their smaller heads.
Individuals from North Carolinian and Virginian populations of putative
P. plattensis were included in a larger study of the P paupercula complex and
allied species (Mahoney & Kowal ms. subm.). Replicates were grown in a stan-
dard environment transplant garden, and data on morphology and flowering
time were analyzed by canonical discriminant analyses (CDA) and analyses of
variance (ANOVA). Herbarium specimens of the complex, emphasizing its dis-
tribution east of the Mississippi River, were studied, anda preliminary analysis
of genetic similarity using random amplified polymorphic DNAs (RAPDs) was
made. One conclusion is that these central Appalachian populations are best
included in the polymorphic P paupercula complex as P. paupercula var.
appalachiana A.M. Mahoney var nov. ined. Chromosome numbers provide criti-
cal evidence supporting this conclusion.
MATERIALS AND METHODS
Individuals of putative eastern disjunct Packera plattensis (P paupercula var.
appalachiana) were collected from two sites. The Paint Rock Road population
(NC, Madison Co. near Hot Springs, 3.2 mi northwest along Paint Rock Road
from Hwy 25-70, Kowal & Mahoney 3120[WIS]) grows in loose, gravelly soil on
a steep, wooded, southwest-facing slope and road cut overlooking the French
Broad River. The Lover’s Leap population (VA, Montgomery Co. 2 mi west from
Price’s Fork on McCoy Road [rt. 652], Kowal & Mahoney 3123 [WIS)) consists of
scattered clones on exposed and wooded rocky outcrops ona south-facing bluff
overlooking the New River and along the south side of McCoy road on a grav-
elly bank. Replicated individuals were randomly positioned in a standard-en-
vironment transplant garden in Madison, WI.
Heads from test garden individuals were fixed in Jackson’s solution (4
ethanol: 2 methanol: 2 chloroform: | propionic acid: | acetone, v/v), transferred
MAHONEY ET AL APPALACHIANA 607
aw i fe. Ye”
3 = e / as % s
D.
Fic. 1 ion of Packera pl is and hyt
Carolina (P paupercula var. appalachiana).®@ P. pl is; @ Disj
annalachiang\
Ale Pr
after about a day to 70% ethanol, and stored in a freezer. Counts were made on
meiocytes of young anthers from florets 2.5-3.0 mm in length. Using a squash
technique, described in Beeks (1955) as modified by Kowal (1975) and Kong
(1998), anthers were prestained in aceto-carmine and squashed between a slide
and a cover slip. Each slide was scanned under the 40 x objective of a Zeiss
Standard WL phase-contrast microscope, and cells with promising meiotic fig-
ures were resquashed until their chromosomes were well-separated and in one
plane. Coverslips were sealed with “Zut.” Counts were made with the 100 x ob-
jective under oil immersion, and cells with well-separated chromosomes were
photographed with Kodak Ektachrome 160 and Technical Pan (exposed at an
ASA of 100) films using a Nikon Microflex Model AFM photomicrographic at-
tachment (Kong 1998). Results are in Table 1 and substantiating photographs
are in Figures 2 and 3.
RESULTS
Individual 9 from Paint Rock Road (NC, Madison Co.) is diploid with n = 22
(Kong 1998; Figs. 2 & 3). Individual 9 from Lover’s Leap (VA, Montgomery Co.)
608 BRIT.ORG/SIDA 20(2)
Table 1.Chromosome counts for Packera paupercula var. appalachiana
Taxon Haploid number Cells counted Phase
NC, Paint Rock Rd, #9 22 34 Metaphase | & II
VA, Lover's Leap, #9 44 | Anaphase |
is tetraploid with n = 44, based ona single, but fairly good preparation (Figs. 4
& 5). A poor anaphase I from individual 10 from Lover's Leap gave an extremely
rough count of 2n =ca.67, superficially, an approximately triploid count. How-
ever, the regular meioses, terminating in normal tetrads of nuclei without
micronuclei, in this individual prompts the interpretation that individual 10 is
tetraploid as well.
Pollen diameters from the transplant garden can be used to extend the
counts of the two individuals to their respective populations, because pollen
diameter is correlated with polyploid level (kowal 1975; Mahoney & Kowal ms.
subm.). A nested ANOVA indicates that the difference in mean pollen diameter
between the North Carolinian and Virginian populations, 27.9 vs. 31.2 wm, is
real (F).g = 24.16, p = 0.0018) and that individuals within each population do
not differ (Fg7 = 2.19, p = 0.16). Taken together, this implies that the populations,
the former containing an individual with n = 22 and the latter with n = ca. 44,
consist of all diploids and all tetraploids, respectively. ANOVAs testing for dif-
ferences among individuals within the two populations separately give abso-
lutely no evidence (F57 = 0.46, p = 0.80) of differences in pollen diameter (and,
implicitly chromosome number) in the “diploid” population but moderate evi-
dence (F37 = 5.11, p = 0.035) in the “tetraploid” population. However, the one in-
dividual with unusually small pollen diameter (29.0 um) is, in fact, individual
10, which is approximately tetraploid, so its aberrancy has other causes.
Uttal (1982) notes that among the specimens from Madison Co., NC (pre-
sumably the Paint Rock Road population), he saw hybrids with P. anonyma.
This species does occur ca. 0.5 miles downstream from the var. appalachiana
population, and 3 of the 11 individuals collected in 1995 showed morphological
similarities to P anonyma. Introgression here is likely, but none of the individu-
als in the transplant garden provided any obvious evidence of it.
Uttal (1982) reports that P aurea and P. paupercula var. appalachiana (as
“P. plattensis”) “thoroughly introgress” along McCoy Road (rt. 652). Individual
10 grew on the road cut and individual 9 grew nearby, on the bluff’s edge; the
other individuals were from less disturbed bluff sites. Some individuals, in par-
ticular #9, have deeply pinnatifid cauline leaves with wide, distally dilated lobes
and broad, undivided terminal lobes like P aurea, which is rampant in an ap-
parent sink-hole in the woodland above the bluff. A garden-grown individual
of P. aurea from this site is lighter green than is typical for the species and
MAHONEY ET Al APPALACHIANA 609
Fic. 2. NC, Paint Rock Road, #9. Metaphase |, 2n =
22 Il.
Fic. 3. NC, Paint Rock Road, #9. Metaphase Il, 2n =
22+22
Fic.4. VA, Lover’s Leap, #9. Late anaphase |,2n =22+ ca, _ Fic. 5. Figure 3 retouched using Corel Photo-Paint 8.
88
610 BRIT.ORG/SIDA 20(2)
reproduces abundantly and invasively by adventitious shoots on roots, unlike
normal P. aurea. Therefore, it is likely that at least a portion of the Lover's Leap
population is introgressed with P. aurea.
Chromosome counts indicate that Packera plattensis does not occur in the
central Appalachians. This Great Plains species appears to be entirely tetraploid
with x = 23 and the North Carolinian population is diploid with x = 22. At least
some members of the Lover’s Leap population in Virginia are tetraploid but
probably with x = 22. This evidence supports the morphological and RAPDs
evidence that these central Appalachian populations are best treated as P.
paupercula var. appalachiana (Mahoney & Kowal ms. subm.).
ACKNOWLEDGMENTS
We would like to thank GH, MO, NCU, NY, PHIL, and UNCC for specimen loans,
Kiefer Kuah and Justin Lo for assistance with photo scanning and retouching
(Fig. 5), Kandis Elliot for the distribution map (Fig. 1), Dale Johnson, the staff at
WIS, and especially Ted Barkley for his bountiful good advice.
REFERENCES
Barktey, T.M. 1962.A revision of Senecio aureus Linn. and allied species. Trans. Kans. Acad.
Sci. 65:318-408.
Barktey, T.M. 1978. Senecio. In Rogerson,C.T.,ed.North American flora, series Il, part 10. New
York Botanical Garden, pp. 50-139.
Bark ey, T.M.1988.Variation among the aureoid senecios of North America: A geohistorical
interpretation. Bot. Rev. 54:82-106.
Beeks, R.M. 1955. Improvements in the squash technique for plant chromosomes. Aliso
3131-1233.
Kona, V.W. 1998. Putative Packera p! is (Asteraceae: Senecioneae) in North Carolina and
Virginia. (Honors senior thesis, eee af Wlisconsi: Madison
Kowal, R.R. 1975.Systematics of Senecio aureus and allied species on he Gaspé Peninsula,
Quebec. Mem. Torrey Bot. Club 23:1-113.
Kowal, R.R., C.-C. Freeman, and T.M. Barktey. Unpubl.ms.cChromosome numbers of the genus
Packera (Asteraceae: Senecioneae): a summary.
poe
Matoney, A.M.and R.R. Kowat. Ms. submitted to Sida. Races of the Packera paupercula com-
plex (Asteraceae: Senecioneae) in the upper Midwest and southeastern United States.
RHoaps, A.F.and W.M. Kcein, Jk. 1993. The vascular flora of Pennsylvania: Annotated checklist
and atlas. Philadelphia, PA: American Philosophical Society.
Urtat, LJ. 1982. Promiscuity of Senecio plattensis Nutt. in a Virginia county. Castanea 47:
344-346.
NORWEGIAN AND SAMI ETHNOBOTANY OF
VERATRUM ALBUM (MELANTHIACEAE)
Torbjorn Alm
Department of botany
University of Troms
N-9037 Troms, NORWAY
ABSTRACT
In Fennoscandia, Veratrum album L. (Melanthiaceae) is restricted to th rt of Nor-
way, Finland and ee Itis locally abundant in coastal areas of oe Finnmark (Norw ce and
well known in both Nor | Sami folk tradition as a toxic plant, frequently causing li
ves
poisoning. Until tealated By tobacco, the roots were used as snuff by the Sami, who had also ae
restricted use of the plant for medicinal purposes, e.g. as a remedy for rheumatism.
RESUMEN
n Fennoscandia, Veratrum album L. Haan aceae) esta euigide a la parte norte de Noruega,
Finlandia y Rusia. Es localmente abundante en | noreste de Finnmark (Norway), y
bien conocida tanto la tradicion noruega como en la finlandesa como planta toxica, que causa
frecuentemente envenenamiento del ganado. Hasta que fue reemplazado por el tabaco, las raices
fueron usadas como rape por los finlandeses, quienes también encontraron un uso restringido de la
planta con fines medicinales, por ejemplo como remedio para el reumatismo.
NORWEGIAN SAMMENDRAG
I ee finnes Veratrum album L. (Melanthiaceae) bare i den nordligste delen av Norge, Fin-
land o land. Arten er lokalt vanlig i kysttraktene i NO-Finnmark (Norge), og velkjent bade i
norsk a ee tradisjon som en an oe som ofte er opphav til forgiftning av husdyr. Den
ble tidligere brukt som snus av samene, en rolle tobakk na helt har overtatt. I samisk tradisjon har
arten ogsa spilt en viss rolle som medisinplante, bl.a. som en kur for reumatisme.
INTRODUCTION
Veratrum album L.(Melanthiaceae) is a rare species in Fennoscandia, and con-
fined to the area’s northeastern part—the Kola peninsula in Russia (Pojarkova
1966), Finnmark in Norway (Alm 1991a), and a single, somewhat disjunct sta-
tion in northeast Finland (Hamet-Ahtiet al. 1998). The plants found in Fennoscandia
have yellow-green flowers, and are sometimes referred to V.album var. virescens
Gaudin, although their relationship to middle European plants, where both
white- and yellow-flowered forms occur, has not been investigated.
Within its Norwegian distribution area, V.album is common locally, and often
predominant on meadows, especially along brooks and rivers. It is also found
in tree-less subarctic areas along the outermost coast, and grows vigorously even in
Empetrum heath and Salix herbacea-type snowbeds (Alm 199]a, 1991b, 1991c).
~~
SIDA 20(2): 611-619. 2002
612 BRIT.ORG/SIDA 20(2)
Veratrum album is easily recognized, due to its large size and characteris-
tic habit. Within the species’ distribution area in Finnmark, both Norwegian
and Sami inhabitants have noted and named the species, for which a one-to-
one relationship exists between the ethnotaxonomical unit and the scientific
species (sensu Berlin 1992).
Ina previous paper (Alm 1991b), dealing mainly with the species distribu-
tion and ecology in Norway, I have included some data on the ethnobotany of
Veratrum album, compiled from various literature sources. The present paper
presents an expanded and revised compilation of ethnobotanical data, and adds
recent ethnobotanical information from interviews with local inhabitants of
both Norwegian and Sami descent in Finnmark. Transcripts and recordings of
the original interviews are stored at the Department of botany, Tromse@ mu-
seum (TROM).
VERNACULAR NAMES
The vernacular names used for Veratrum album in Finnmark are all related to
its toxicity. A characteristic feature of the chemical components of V. album is
their ability to cause sneezing (Frohne @ Pfander 1983). The only Norwegian
vernacular name recorded for V.album, nysrot or nyserot Csneeze root”), reflects
this property. The latter form is now also well established as an “official” (e.g. in
floras) Norwegian name for the species. As such, it was first used by Gunnerus
(1772).
Gunnerus mentioned two Norwegian names for V. album, Hviit Nysegras
(hvitt nysegras in modern Norwegian), i.e. “white sneeze-grass,” and Hviit
Nyseroot (= hvit nyserot), ie. “white sneeze-root.” Although his flora includes
much ethnobotanical information, vernacular names provided by Gunnerus
should be treated with some caution. Gunnerus coined Norwegian names for
numerous species, including e.g. many fungi that had little or no chance of be-
ing recognized by the layman (Eckblad & Heiland 1985). In the case of Ver-
atrum, at least the preceding hvit Cwhite”) must have been added by Gunnerus,
probably by translating the Latin epithet of V. album; there is no factual basis
for connecting this term with the yellow-flowered specimens found in Norway.
This comment also applies to the notion of Hvid Nyserot as a Norwegian ver-
nacular name in Wahlenberg (1833).
Gunnerus original record had no indication of colour, and was a straight-
forward Nyseroot or Nysroot, as revealed by his letters to Carl von Linné
(Linnaeus) dated 13 February and | April 1768 (printed in Amundsen 1976; origi-
nals in the Linnaean Society, London). Furthermore, nyserot has now been con-
firmed as a genuine and still extant vernacular name in Finnmark by inter-
views e.g. at the Nordkinn peninsula.
The most frequent Sami name for Veratrum album is gastinrdssi (in mod-
ALM, ETHNOBOTANY OF VERATRUM ALBUM 613
ern North Sami, gastemrasse in older spelling), of similar meaning (“sneeze
plant”); from gastin, to sneeze, and rdssi, “grass” or “plant’—the latter being a
preferable translation. It was first recorded by K. Leem and J.E. Gunnerus in the
mid-18th century (Alm 1991b; Gunnerus 1772; Nielsen & Nesheim 1979;
Qvigstad 1901), and was also noted by Wahlenberg (1812) during his 1801 visit
to Finnmark. It is still widely known in Finnmark (interviews 1996).
A deviant Sami name, mirkkordssi “poisonous plant”) has been recorded
in the Deatnu/Tana area. It was first mentioned by Qvigstad (1901), but in eth-
nobotanical context asa nomen nudum, as he did not identify the species. Alm
(1991b) suggested that the name might refer to Veratrum album; this has now
been confirmed by interviews with Sami inhabitants in the Sieidda area of
Deatnu/Tana.
HUMAN USE
Literature sources reveal past use of Veratrum album by the Finnmark Sami as
a kind of snuff, perhaps with intoxicating abilities. Such use was first suggested
by Hans Lilienskiold, district governor of Finnmark in the late 17th century,
and author of a large, hand-written description of the area:
“If the tobacco-hungry knew the roots strength, little would be left standing
(cited from the printed version in Solberg 1942-43; translated from Norwegian).
Gunnerus (1772) noted that the Sami used the pulverized root as a kind of
sneezing powder, ie. as snuff. A slightly more detailed description was given
by Jens Rathke, in an account of his 1801-1802 journey in Finnmark:
“The use of the roots of this plant as snuf{-tobacco has formerly been very
frequent among the Sami, but is now increasingly substituted by [ordinary] to-
bacco (..)” (Rathke 1899; translated from Norwegian).
Use of Veratrum as snuff was also noted by the Swedish botanist Géran
Wahlenberg during his 1801 visit in Finnmark (Qvigstad 1932). A single 20th
century record of this practice is found in Lillevik (1970), who noted that a Sami
walking companion at the Nordkinn peninsula was eagerly interested in some
Veratrum stands they passed, showing visible pleasure. Lillevik (op.cit.) also
noted that only full-grown plants were selected for human use.
The effects of using Veratrum snuff are not described in any Norwegian
sources. The roots have a strong and bitter taste (Hoppe 1975). According to
Frohne and Pfander (1983), 1-2 grams of the dried root may be lethal for hu-
mans. However, since the poisonous constituents provoke violent sneezing, it is
hardly possible to consume such amounts, except in alcoholic extracts (Hruby
et al. 1981). Used as snuff, the Veratrum powder was supposedly spit out in a
short while, certainly after causing some sort of sensation. There are no records
of any kind of intoxication, although V.album is known to possess psychoactive
abilities (Ratsch 1998). In fact, sneezing may have been just the desired effect; in
bs
614 BRIT.ORG/SIDA 20(2)
Sami folk medicine, sneezing was considered healthy, probably asa way of “get-
ting rid of” diseases; ordinary tobacco has also been much used to provoke sneez-
ing (Qvigstad 1932).
Apart from this, V.album has found little use in folk medicine. According
to Qvigstad (1932), Wahlenberg had noted that Veratrum was used to treat rheu-
matism in Finnmark. No further details are known, e.g. on ethnicity of those
who used it, but it is likely to be a Sami tradition. According to Steen (19601),
Sami inhabitants of the Lyngen area of Troms, North Norway, used V. album to
treat rheumatism. If so, the cure must have been based on plant material im-
ported from Finnmark, as the Lyngen area is situated far outside the present
distribution of V.album. Steen suggests that the cure was carried out mainly by
drinking a decoction of Veratrum; surely a dangerous practice. It could also be
applied as a compress on the painful area.
Gunnerus (1772) listed a whole range of diseases that could be treated with
Veratrum, but his account seems to be based on literary sources related to Central
Europe, including Gouan (1765). Hiorthay (1785) mentioned Veratrumasa cure
for psychic disorders in Gudbrandsdalen, southeast Norway, again probably
based on a learned tradition, and certainly on imported plant material—the
nearest stations with indigenous V. album are situated in Central Europe; the
plant used may have been Helleborus niger (Reichborn-Kjennerud 1922).
LIVESTOCK POISONING
Veratrum album is strongly poisonous (Bertrand et al. 1990; Frohne & Pfander
1983; Hoppe 1975; Jaspersen-Schib et al. 1996; Keeler et al. 1978; Quatrehomme
et al. 1993). In Finnmark, this is well known to anyone living in areas with large
stands (Alm 1991b; Hoeg 1974; Lillevik 1970). However, since the plants sprout
in early spring, and unfold considerable leafage before almost any other herbs,
they are frequently consumed by animals. The resulting poisoning was first
commented on by Lilienskiold in the late 17th century:
“When the creatures eat its leaves in abundance towards spring, it affects
their hairs, which are felled” (cited from the printed version in Solberg 1942-
43; translated from Norwegian).
Lethal poisoning of a small herd of cattle was reported from the Deatnu/
Tana area in Finnmark by Ruud (1938). Dissection showed haemorrhages of
the heart, stomach and intestines, liver damage, and lungs with a bloody foam.
The incident occurred in June, in circumstances that were typical of past pov-
erty. Both in Norwegian and Sami husbandry, the animals were customarily
hunger-fed during the latter part of winter, and would eagerly eat anything
available when released in spring—including the large, but poisonous shoots of
Veratrum album.
Nowadays, the remaining cattle in Finnmark are usually fed on cultivated
pastures, and poisoning is unlikely to occur. Sheep, on the other hand, frequently
—
ALM, ETHNOBOTANY OF VERATRUM ALBUM 615
eat shoots of Veratrum album. According to people at the Nordkinn peninsula,
lambs would often consume at least some leaves of Veratrum. Poisoning is rarely
lethal (though death may occur, as suggested by Lillevik 1970), but causes se-
vere sickness, characterized by a “foam” issuing from the mouth. A recent de-
scription may be included:
“Nysrota ... itis the worst weed we've got. Almost every lamb will eat it, and
then they get this foam at the mouth.”... “It becomes all foam. It’s really awful, it
even gets into the eyes” (interview 1994; translated from Norwegian).
Vomiting is frequent (Hoeg 1974). pecording to people at Veernes (Nordkinn
peninsula), the only remedy that would help was giving the affected sheep milk.
When symptoms subdued, the lambs seemed to suffer no lasting conse-
quences—and would subsequently avoid Veratrum, for which reason grown-
up sheep are rarely affected (interview 1994).
CONTROL MEASURES
Asaconsequence of livestock poisoning, farmers in Finnmark at least formerly
tried to decimate or eradicate V. album—a difficult task, given that the species
has very effective vegetative propagation by its thick rhizomes. The Norwegian
botanist J.M. Norman made a characteristic observation at Bonjakas in Deatnu/
Tana in the late 19th century:
[Veratrum album occurs] “in large quantity, somet g into sites,
where is was not previously noted, and threatening during the [18]80's to be-
come a real pest, in particular in spring, when the livestock is released. One
tries to curb its further spread by scything it in early spring” (Norman 1900;
translated from Norwegian).
The Swedish botanist Nikolaus Svensson noted a similar practice during
an 1892 visit to Deatnu/Tana:
[Veratrum album] “is very frequent around the Tana fjord and ascends high
up into the mountains, but only in the immediate vicinity of the fjord. In Tana,
a veritable war of extermination is fought against it due to its toxicity. However,
it seemed to manage well, and often grew as high as a man on the small pas-
tures along the river” (Svensson 1894; translated from Swedish).
Conscious eradication may explain the present absence of V. album from
several sites in Finnmark where literature records suggest that it did occur in
the 18th and 19th century, e.g. at Nordkapp and Porsanger (Gunnerus 1772;
Wahlenberg 1812) and Varda (Solberg 1942-43), well outside the present range
(see map in Alm 1991b).
Stands of V. album with all stems cut at base level—obviously as a control
measure—were observed in Deatnu/Tana as late as in 1975 (Alm 1991b). Such
practices are probably rare by now.
At present, V.album is frequent on meadows e.g. in Deatnu/Tana, and little
effort seems to be directed at controlling it. This is partly a consequence of
]:
616 BRIT.ORG/SIDA 20(2)
altered agricultural practices; numerous farms have been deserted, and those
remaining have largely adopted modern practices of intensive grass cultiva-
tion; the former extensive use of both in- and outfield areas is hardly practised.
Like many other large herbs, V. album thrives on abandoned meadows, and is
probably more common nowadays than in the past.
VERATRUM IN OTHER AREAS
Veratrum album is widely distributed in Eurasia, from Spain to Beringia (Hulten
& Fries 1986), with a close relative (V. viride Ait.) in North America. The latter
is also strongly poisonous (Prince & Stork 2000). Throughout the area,
Veratrum’s characteristic ability to cause sneezing has been well known.
Veratrum album is found in the mountains of mainland Greece, and may
have been known to the ancient Greeks, though to what extent it was used is
disputed. Theophrastus (book IX, X) noted that “The white and the black helle-
bore appear to have nothing in common except the name” (Hort 1916). Accord-
ing to Baumann (1993), V. album was not used by the Greeks, wheras Ratsch
(1998) suggested that it was considered a medicinal plant of prime importance.
Raven (2000) rejected previous ident ifications of the white hellebore of the
ancient Greeks, and suggested Drimia maritima (L.) Stearn, syn. Urgineda mar-
itima(L.) Baker, asa more likely candidate, whereas the “black hellebore” prob-
ably refer to Helleborus species.
Just as in Sami tradition, the Greeks (as recorded by Pliny the elder) con-
sidered sneezing as a way to promote health, by forcing the demons of disease
to leave the body; its main use was for psychic diseases (e.g. depression), epi-
lepsy, etc. (Ratsch 1998). Similar use is mentioned in early printed European
herbals, e.g. Fuchs (1543).
In Italy, an extract of Veratrum album roots in alcohol has served as an
antirheumatic remedy (Cappelletti et al. 1982). In Iranian folk tradition, a paste
of the fresh root of Veratrum is used externally to relieve headache and neural-
gic pains (Hooper 1937).
In North America, numerous Indian tribes have used Veratrum species for
medicinal and other purposes (Moerman 1998). Antirheumatic use of V. viride
is rather widespread, and known e.g.among the Bella Coola, Cherokee, Gitksan,
Haisla, Hanaksiala, Iroquois, Kitasoo, Okanagan-Colville, Oweekeno, Quinault,
Salish Thompson, and Tsimishian tribes (Gottesfield & Anderson 1988; Gunther
1988: Moerman 1998; Smith 1929; Turner et al. 1980, 1990). The Blackfoot Indians
used powdered Veratrum viride to provoke sneezing and thus clear the respira-
tion tract (Hart 1979). V.californicum Dur. has been used as a remedy for rheu-
matism, wounds and sores, and as a contraceptive (Kelly 1932; Train et al. 1941).
Veratrum species feature in a variety of other folk beliefs and traditions
among the Indians of North America. The Gitksan and Haisla Indians made
ceremonial use of V. viride to drive off evil spirits, and could wear a piece of the
ALM, ETHNOBOTANY OF VERATRUM ALBUM 617
root to ensure good luck, ie. as an apotropaic. It was also used in purification
rituals (Gottesfield @ Anderson 1988). Haisla Indian shamans applied the plant
to arrows to shoot at and drive off “disease spirits” (Moerman 1998). Among the
Okangan-Colville Indians, V. viride featured in witchcraft (Turner et al. 1980).
Gunther (1988:24) recorded that the Cowlitz indians used a chewed piece of
Veratrum to scare off sea-monsters.
In summary, the use of Veratrum species as an anti-rheumatic remedy is
common to the Old (Norway, Italy, Greece) and New World (American Indi-
ans). Their ability to provoke sneezing is well known in both areas, and was
considered attractive either as a way to expel evil sprits, get rid of disease, or
merely regarded “healthy” as in Sami tradition.
ACKNOWLEDGMENTS
The author expresses his gratitude to people in Finnmark who have offered in-
formation on vernacular names and other traditions related to Veratrum al-
bum, in particular Karstein Adriansen and Eva Mikalsen (Norwegian tradition)
and Ole Larsen, Terje Pedersen and Johannes Persen (Sami).
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620 BRIT.ORG/SIDA 20(2)
Book REVIEW
DonaLp A. Cox. 2002. A Naturalist’s Guide to Wetland Plants: An Ecology for
Eastern North America. [|]ustrations by Shirley A. Peron. ISBN 0-8156-0740-
7, pbk.) Syracuse University Press. 621 Skytop Road, Suite 110, Syracuse,
NY 13244-5290. (Orders: 315-443-2597, Fax: 315-443-5545, supress@syredu)
$17.95, 203 pp, 123 line drawings, 6" x 9".
Once viewed as wastelands, wetlands are now better appreciated by the public for the value they
bring to human life. By some estimates, wetlands once covered more than 200 million acres in the
United States. Yet, oo half of these eas ecosystems were destroyed by the 1950's for use in
agriculture and t. Recent leg I wetlands and associated biota,
yet these ecosystems continue to be destroyed, albeit at a slower rate, and wetland species become
threatened or endangered. This book focuses on palustrine wetlands, ie. marshes, bogs, and swamps
and within the first few pages raises the reader's awareness of the value of wetlands.
Beginning, appropriately, with a definition of wetlands as ecosystems, Cox enumerates in the
first chapter dhe eine Bi Werlandeds Habitat for el noode protection, recreation eae oa ater
tae: ge, oe oe Ss. 7
d global climate chan lu cothe ee of fossil fuels. . latter
5
half of the first cl landsin North A ibed—the Atchafalaya Swamp,
Cranberry Glades ai West vee the Everglades, the Great Dismal Swamp, Okefenokee, the Pocosins
and prairie pothole marshes. Information about each wetland details geography, dominant vegetation,
and historical and current ranges. The next five chapters emphasize the plants that grow in wet-
lands. Chapter two, entitled Types of Plants, explores the different forms of plant life in wetlands or
for that matter, any ecosystem on the planet. Details on the life history and general characteristics of
algae, fungi, slime molds, lichens, bryophytes, ferns and allies, gymnosperms and angiosperms are
prowiee: In the ReRrChApes ie ares Survival, the author | pollinati | seed
ft | ] |
iter in wetland Sine C
I
sed bogs and peatlands focus or imarily on i vascular plants growing there. Though illustrations
are scattered throughout the book, more are presented here to enhance the plant descriptions
In the chapter, Plants of Special Interest, the amount Gea on poisonous plants far out-
weighs the details on hallucinogenic and edible plants. In fact, only one hallucinogenic plant is men-
tioned, sweetflag or Acorus calamus, and | found myself wondering about the merit of listing this
plant in its own heading because of its psychoactive properties. Perhaps it would have been better served
in the medicinal plants section since Native Americans used it to treat some physical ailments
Transitions in the physical characteristics of wetlands due to seasonal climate change are dis-
cussed in chapter seven, Through the Year. Noxious aquatic weeds are also considered in some detail
here. The author provides tips on collecting, preserving and identifying plants, including sound ad-
vice on where and what not to collect, obtaining permission from land owners prior to collecting
and how to collect good, representative specimens. In the final chapter, suggested activities on wet-
ife history of a plant or investigating the cl a
lands, such as making a study on the
local wetland, encourage readers to learn more.
his book raises awareness regarding the importance of wetlands. The dynamic attributes of
ecosystems and interconnected nature of wetland biota is stressed throughout. Clarity and brevity
of Cox’s writing style makes this book relevant to nearly all groups of people— naturalists, educators
college students, amateurs interested in birds, amphibians, reptiles or plants. Emphasis on conserva-
tion and habitat preservation make this book especially valuable.—Amy Trauth Nare, Botanical Re-
search Institute of Texas, 509 Pecan Street, Fort Worth, 1X 76102, U.S.A, amy_trauth@ yahoo.com.
SIDA 20(2): 620. 2002
CARLO BERTERO (1789-1831) IN THE NEW WORLD
Piero G. Delprete Giuliana Forneris
Institute of Systematic Botany Erbario
The New York Botanical Garden ee es di Biologia vegetale
Bronx, NY 10458-5126, U.S.A. niversita degli Studi di Torino
pdelprete@nybg.org Viale sh Mattioli 25, 1-10125, Torino, ITALY
giuliana.forneris@bioveg.unito.it
Annalaura Pistarino
Museo Regionale di eae Naturali
Via Giolitti 36, |-10123, Torino, ITALY
annalaura.pistarino@regione. piemonte.it
—
ABSTRACT
Carlo Giuseppe Bertero (1789-1831) is, among the AMON Italian botanical collectors that contrib-
uted toward the knowledge of New World flora, the one Ll dand lled tl Bertero
gathered a oe number of anes which were in turn distributed to sae of a ann
dents, and are now present in many herbaria worldwide. During his travels
1
which he sent to numerous ee contributing toward the collections of private and ane gar-
dens. Several hundreds of his botanical collections turned out to be plants unknown to science, which
were in turn described by m of his con sae ae oe botanists. The botanical work of
Carlo Bertero in Italy has ieee ly treated b ny specialists. Nevertheless, his bo-
tanical expeditions and eee in ee New World have never ae completely studied, although
ae reports have ened peen Sea Such studies aa further vests: as new ae
ve been poccney obtained g CHeSpeneence field Boe sand herbar
]
o's first aie to the Antilles
and northern Colombia (1816-1821), and his secoull eredinen to cre (1827-1830), Juan Fernandez
Islands (1829) and Tahiti (1830-1831). This preliminary study is part of an ongoing project that will
produce a complete biography of Carlo Bertero and a comprehensive account of his botanical work
in the New World and the South Pacific, that will be published in the near future (Delprete, Forneris
& Pistarino, in progress)
Key worps: Carlo Bertero (1789-1831), Giovanni Battista Balbis (1765-1831), Luigi Colla 1766-1848);
Eee seems botanical collections, botanical explorations, itineraries; Greater Antilles,
Guadalupe, Puerto Rico, Santo Domingo, Jamaica, South America, Colombia, Chile, Juan Fernandez
Islands, ae Society ae
RESUMEN
Carlo aes Bertone O789- 1831) es, entre los Taeee fon Polaco: italianos que
t Jo la fl INu levo 5A BR
Mundo viajo y ertero acopio
| C f Jjctribniid
5 b 1
h t4 =| } } } : ] =
sala oe de sus oe y que
]
rante sus viajes recolect6 también
semillas, que mand6 a muchos botanicos, contribuyendo a las Penere de ee publicos y
peiadonr Varios centenares de s sus pspecumencs se revelaron como Pele er scenueias para la
arlo
ClemCla yf if t t
SIDA 20(2): 621-644. 2002
622 BRIT.ORG/SIDA 20(2)
liat | | Sin embargo
Berteroen It )
colecciones en el Nuevo Mundo nunca es Sanne estudiadas, aunque se Soaileaten
ee reportes oa Estos esiudton necesitan de oe: inv estipacion, puesto que fueron
|
etiqu
de herbario, en el intento por reconstruir los itinerarios - la secuencia cronologica re su primera
paras a Las Antillas y norte de Colombia (1816-1821) y su segunda expedicion a Chile (1827-
1830), Islas de Juan Fernandez (1829) y Tahiti (1830-1831). Este estudio preliminar es parte de un
proyecto en curso que producira una biografia completa = Carlo Ber ey una rejacion comp ronan a
ifj 4
L £ I
n
de su trabajo botanicoen el Nuevo Mundo y enel P
(Delprete, Forneris & Pistarino, en curso).
PALABRAS CLAVE: Carlo Bertero (1789-1831), Giovanni Battista Balbis (1765-1831), Luigi Colla (1766-
1848); ore bibliografia, colecciones botanicas, exploraciones botanicas, itinerarios; Antillas
Mayores, Guadalupe, Puerto Rico, Santo Domingo, Jamaica, América del Sur, Colombia, Chile, Islas
de Juan Fernandez, Tahiti, Islas de la Sociedad.
Among the many Italian naturalists who explored the New World, Carlo
Giuseppe Bertero (1789-1831) stands out as one of the early collectors who vis-
ited little-known or unexplored areas, and for the impressive amount of mate-
rial that he collected in these regions. The historical, floristic and ecological
importance of Bertero’s collections is also that they were made in areas that are
nowadays almost entirely destroyed, particularly in the Greater Antilles, Chile,
and in the Juan Fernandez Islands. He engaged two maior trips, covering his
travel expenses as a medical doctor: the first in the Antilles and the Caribbean
coast of Colombia (1816-1821), and the second in central Chile 1827-1830), Juan
Fernandez Islands (1829) and Tahiti (1830-1831). The bulk of Bertero’s collec-
tions was shipped to Giovanni Balbis (1765-1831) and Luigi Colla (1766-1848;
cf. Delprete 2000), both resident at Torino, and were in turn distributed to many
major European herbaria (and now partially present in many North and South
American institutions), and his seeds and living material were sent to several
European botanical gardens (Spin 1823; Parlatore 1841; Visiani 1855).
A considerable portion of his collections turned out to be genera and spe-
cies unknown to science, and were described by many contemporary botanists.
Most of Bertero’s new taxa from the Caribbean region were primarily described
by: A.P de Candolle (823-73) and Kunth (1833-50) and those from Chile by
Moris (1831, 1832, 1833, 1834, 1835), A. de Jussieu (1832), and Colla 1824, 1834,
1835, 1837) among many others (ie., Balbis, Decaisne, Delessert, Delile,
Guillemin, Hooker & Arnott, Montagne, Nees, Philippi, Sprengel, Trinius, Ur-
ban and Vignolo-Lutati). Four generic names (Berteroa DC., Berteroella O.E.
Schultz, Berteromyces Ciferri and Terobera Steud.) and more than 300 specific
epithets were dedicated to Bertero (Mattirolo 1932a, 1932b). The correct orthog-
raphy of specific epithets honoring Bertero has been discussed in Stuessy and
Marticorena (1990) and Kartesz and Gandhi (1993). At last, a reef discovered a
few years after Bertero’s death, was named after him (Bertero’s Reef, also Récif
Ebrill, Tuamotu Archipelago, 22°40! South, 133°30' West, Pacific Ocean) by J.A.
DELPRETE ET AL., CARLO BERTERO IN THE NEW WORLD 623
Morenhout, the Consul of the United States in the Pacific Islands and his last
travel companion in the Society Islands.
Bertero also contributed greatly to the knowledge of the Italian flora
through his collections before and in-between his two American expeditions.
Many authors have focused their work on Bertero’s botanical collections in Pied-
mont (northwestern Italy), which are preserved at TO, in the Herbarium
Pedemontanum (1O-HP: Re 1827: Bertoloni 1833-54; Colla 1833-37; Burnat 1883:
Ferraris & Ferro 1904; Mattirolo 1929; Castiglioni 1931; Steinberg 1971; Vignolo-
Lutati 1926, 1927, 1928, 1929, 1932, 1950, 1960; Forneris & Pistarino 1987-88,
Forneris & Pistarino 1991; Forneris et al. 1992).
Several authors have dedicated their attention to Bertero’s life and work in
the Americas (e.g., Vignolo-Lutati 1955, 1956), but the subject has never been
treated extensively. Therefore, it has become necessary to start a major project
on Bertero’s biography and botanical expeditions in the New World (Delprete
et al. 2000; Delprete, Forneris & Pistarino, in progress), with the main goals to
catalogue Bertero’s itineraries and collections, and to produce a study on his
life, dedicated to botany, until his premature death caused by a shipwreck in
the high seas of the Pacific Ocean. The goal of the present article is to introduce
botanists and historians who are interested in the floras of the Americas (and
the Society Islands) to Bertero’s major accomplishments, and to his dedication
and passion for botanical research that lasted until the last day of his short life.
BIBLIOGRAPHIC SOURCES
Bibliographic information has been compiled from the works of many authors
who in various occasions have mentioned Bertero’s work and life in the New
World (Guillemin 1830; Anonymous 1831, 1833, 1834; Hooker & Arnott 1833;
Colla 1832, 1839; Astegiano 1857; Candolle 1862; Fournier 1876; Molinari 1880;
Urban 1902; Mattirolo 1930, 1932a, 1932b; Gunckel 1931; Looser 1931; Mussa 1931;
Vignolo-Lutati 193la, 1931b, 1951, 1955, 1956; Pichi Sermolli 1951, 1952, 1988;
Lanjouw & Stafleu 1954; Perez-Arbelaez 1970; Piovano 1970; Stafleu 1974; Stafleu
& Cowan 1976; Del Vitto 1986; Montacchini et al. 1986; Brondino 1989; Forneris
et al. 1989: Molinaro 1989; Pistarino @ Clemente 1989; Pistarino et al. 1989; Stafleu
& Mennega 1993).
Among the works that focused, at least in part, on Bertero’s American ex-
peditions, should be mentioned the Elogio storico dell’Accademico Dottore Carlo
Bertero published by Colla (1839), Bertero’s friend and colleague, and the works
of Vignolo-Lutati (1955, 1956) and Pistarino et al. 1989). Colla’s Elogio is par-
ticularly important in this biographic project, because he was one of Bertero’s
dearest friends, and also because it is based on the many letters, with many
biographical notes, that were written to him by Bertero. Vignolo-Lutati (1955)
studied Bertero’s American collections preserved at TO-HG, from the Balbis and
624 BRIT.ORG/SIDA 20(2)
Colla herbaria (for a total of 2,248 taxa), with scholarly work of analysis and
organisation [the taxa are arranged according to the Durand (1888) classifica-
tion, as the specimens still are at TO-HG]. Pistarino et al. 1989) represents the
first step of the Bertero Project, of which this article is the logical continuation.
Another important source of information is contained in the letters exchanged
among Bertero, Colla and Balbis, which are united into 22 volumes of
Corr ispondenza botanica ed micale [Botanical and personal COrrTe sponde nce]
and is preserved at the Biblioteca Reale di Torino[Royal Library of Turin]. These
letters represent an extremely important documentation in the recontruction
of the stages of life and the scientific activities of Bertero. A few letters are also
preserved at the Accademia delle Scienze di Torino [Academy of Sciences of
Turin] and at the Biblioteca Civica di Torino [Civic Library of Turin]. Neverthe-
less, Turin is not the only city were Bertero’s letters can be found, mostly be-
cause he kept close epistolary relationships with many botanists of his time
(Anonymous 1831).
Finally, the most important document examined in this study was Bertero’s
field book of his first trip to the Antilles and northern Colombia (Fig. 2). This
precious document, currently preserved at the Library of the Department of
Plant Biology of the University of Turin, is a result of 14 field books that were
later bound together, to form a book of 1,095 pages, where he described 1,746
collections. The manuscript is in Latin, in Bertero’s minute handwriting, and
with exhaustive descriptions (sometimes more than one page long), habitats,
localities, common names, local medicinal uses, and sometimes augmented by
accurate discussions on systematics, taxonomy and nomenclature. In addition,
many pages of the manuscript are enriched by Bertero’s pencil drawings of
morphological details that are often fundamental for the determination of the
species collected. This manuscript was donated by Balbis to Augustin Pyramus
de Candolle [who used (and sometimes copied) Bertero’s descriptions of the
Caribbean taxa and published them in the Prodromus, 1823-73], and was later
returned by Alphonse de Candolle (Augustin’s son) to Turin in 1857 (Vignolo-
Lutati 1931a; Mattirolo 1932a). Unfortunately, Bertero’s field books of his trips
in Chile, Juan Fernandez Islands and Tahiti have been lost.
aa
BERTERO'S LIFE, TRAVELS, AND BOTANICAL COLLECTIONS
Bertero’s Early Years In Europe
Carlo Giuseppe Bertero (Fig. Ll) was born on 14 October 1789, in Santa Vittoria
d’Alba,a small town in the hilly region of Langhe, about 60 km South of Turin,
Piedmont region, northwestern Italy. He studied medicine and natural history
at the University of Turin, and later studied botany under the famous botanist
Giovanni Battista Balbis, and finally received his Laurea in Medicine in 1811.
His doctoral dissertation clearly shows the first signs of his botanical inclina-
625
DELPRETE ET AL., CARLO BERTERO IN THE NEW WORLD
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Fic. mili of petieta® field book, compiled By viding is igendion ue fascicles with collections numbers,
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loca drawings, etc.) p (1816-1821)
tion (Bertero, 1811). While studying for his medical degree, he was a frequent
visitor to the Orto Botanico, and his botanical interests are shown in a letter
[hereafter, between brackets are indicated the locality and date of the letters
sent by Bertero] (Alba [Piedmont], 21 August 1809) where he asked his botani-
cal master Balbis to find hima copy of the Flora Pedemontana of Allioni (1785).
In these early years of formation, Bertero strengthened his own botanical knowl-
edge and built the foundation of a strong friendship with Balbis, as is demon-
strated in their frequent letters.
In the following years, Bertero proved to be quite successful in the medical
sciences, but he refused to continue his medical studies at the Collegio di
Medicina [College of Medicine], and started eagerly botanizing in the hills and
valleys of Piedmont instead. The epistolary relationship between Bertero and
Balbis continued strongly through this period, even when the latter moved to
Pavia (Torino, 24 May 1815). In these letters, Berter icated his frequent
botanical collections (Alba, | and 7 January 1816, and 6 March 1816) and put his
own herbarium at the complete disposition of Balbis (Alba, 7 June 1816) and
offered (Alba, 18 March 1816) to help him in the recompilation of the indexes of
Willdenow’s Enumeratio plantarum (1806-1816) and Species plantarum (1797-
my
—
DELPRETE ET AL., CARLO BERTERO IN THE NEW WORLD 627
1825), which were recompiled by Balbis because a great number of taxa were
missing in both works.
Before moving to France, Bertero was living in his family home in the small
town of Santa Vittoria d’Alba. The frustration caused by the increasing isola-
tion and his desire to be part of the academic world stimulated him to search
for objectives outside of Italy. He condensed these feelings of dissatisfaction in
a sentence included ina letter that he sent to Balbis: “... Ce qui me console encore
cest que touls] mes parents sont persuadés qu’Alba n’est [pas] faite pour moi,
nayant aucune ressource dans ce pays” [what consoles me is that all my rela-
tives are persuaded that Alba is not made for me, there are no resources in this
town (Alba, 7 June 1816)]. In July 1816, with the help of Balbis, who had connec-
tions with many European scientific personalities, Bertero moved to Paris. In
this city, he had the occasion to meet many famous contemporary botanists as,
for example, René Louiche Desfontaines (Paris, 20 July 1816), Jean Louis
Loiseleur-Deslongchamps, and Christiaan Hendrik Persoon (Le Havre, 5 Au-
gust 1816). The Parisian cultural and scientific environment, the study of her-
baria containing tropical collections and the frequent visits to the Jardin des
Plantes contributed to Bertero’s rapidly increasing knowledge. It was in this
exciting cultural atmosphere, and with the help of Christiaan Persoon (1761-
1836), that Bertero decided to the study the flora of the Antilles.
Bertero’s Expedition to the Antilles and Colombia (1816-1821)
General Foujaz de Saint Fond, captain of the ship Guadalupe, offered an invita-
tion, extended through Persoon, to Bertero to travel on his boat as ship doctor;
an offer that he gladly accepted. On August 1816, Bertero departed, on the
Guadalupe, from the port of Le Havre, with destination to the West Indies. Dur-
ing the trip, the entire crew was infected with yellow fever, and Bertero suc-
ceeded in saving most of them, so that at the arrival in Martinique he was wel-
comed with extraordinary honors.
In Martinique, Bertero was much appreciated as a medical doctor and as
naturalist, to the point that he was offered the direction of the local museum of
natural history and of the botanical garden. This position Bertero did not ac-
cept because he wanted to be free for his botanical explorations. The earliest
letter examined documenting his stay in Guadeloupe was dated 31 December
1816, where he affirmed to have collected more than 350 species and to have
prepared LOO packages of seeds to be sent to Europe (Pistarino & Forneris 1992).
During this expedition that lasted nearly five years, Bertero visited
Guadeloupe in 1816-1818, Saint Thomas in 1818, Puerto Rico in 1818-1819, Santo
Domingo (Rio Yaqui, etc.), Haiti Jacmel, Les Cayes, Port-au-Prince, etc.) in 1819-
1820, and Colombia (Santa Marta, Barranquilla, Mompos, and the lower Rio
Magdalena) in 1820-1821. Because of ongoing revolution in Colombia, he re-
turned to Europe, with a short stay in Jamaica, in 1821.
628 BRIT.ORG/SIDA 20(2)
During his stay in the Caribbean region, the correspondence of Bertero with
Balbis and Colla is occasional; this may be related to the difficulties of deliver-
ing the mail from such remote places to Europe. The letters that we have stud-
ied were in chronological order as follows: France: Paris 20 July 1816), Le Havre
(5 August 1816), Guadeloupe: Pointe-a-Pitre (31 December 1816), Grande Terre
(30 January 1817), Petit Canal (30 May 1817, 15 August 1817, 1 September 1817, 13
May 1818), Pointe-a-Pitre (lO July 1818); Saint Thomas (30 July 1818); Puerto Rico:
Patillas (15 January 1819); Hispaniola: Santo Domingo (4 November 1819); Ja-
maica: Kingston 2 June 1821); France: Paris (3 October 1821).
In several letters (Petit Canal, 1 September 1817; Pointe-a-Pitre, 10 July 1818;
Santo Domingo, + November 1819), Bertero mentioned several trips that he
planned but never realised. Colla (1839), based on the letter of 15 August 1817
from Petit Canal, referred to an itinerary that Bertero planned (and Colla as-
sumed the he realised) from Guadeloupe to Martinique, Trinidad, Tobago and
to the mouth of the Orinoco River (Venezuela), to which some other authors
have referred in the reconstruction of the itinerary of Bertero’s first trip
(Astegiano 1857; Molinari 1880). Most likely this trip never took place, and no
collections, field books, nor correspondence from any of these areas has ever
been found. Urban (1902), studying Bertero’s herbarium specimens, was able to
make some corrections to the inexact information supplied by Colla about
Bertero’s trip to the Antilles and northern South America. Urban’s corrections
and additions were later followed by Mattirolo (1932a, 1932b) and Spada
Sermonti (1967). Both specimen labels and correspondence enable us to con-
struct a fairly accurate itinerary of Bertero’s first trip (Fig. 3). Also, Table |
summarises the localities and the periods in which Bertero made botanical col-
lections in the Antilles and in Colombia. This table has been constructed based
on data obtained from the above-cited letters, Bertero’s field book, and the refer-
ences published by Vignolo-Lutati (1955), who in turn extracted this informa-
tion from TO-HG and Bertero’s field book.
The botanical research conducted by Bertero in the Antilles took place with
many difficulties. The various problems that he referred to in several letters
were mostly the amount of time that he had to dedicate to his medical practice,
necessary to cover his travel and living expenses, and by the few references avail-
able to him for the identification of his own collections (Pointe-a-Pitre, 31 De-
cember 1816; Petit Canal, 30 May 1817,and 1 September 1817). In those days Curt
Sprengel’s (1827) and A.P. de Candolle’s (1823-1873) treatments were not yet
available; in fact, these texts were published later and were based on many of
Bertero’s neotropical collections.
He stayed in the West Indies Jamaica) until the end of July 1821, extremely
saddened to have to abandon those beautiful lands but, at the same time, anx-
ious to meet again his relatives after his trip of nearly five years. But what made
Bertero even more anxious about his return to Piedmont was the possibility of
DELPRETE ET AL., CARLO BERTERO IN THE NEW WORLD 629
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meeting Balbis, at Turin, in order to study and determine his new collections,
discuss his botanical observations, and update himself on the new publications
which had come out during his absence (Kingston, 2 June 1821).
Bertero’s collections from the West Indies and Colombia were received by
Balbis, who, after a rapid determination by Curt Sprengel (at Halle, Germany),
distributed them to various botanists that specialised on Caribbean flora and
to several European herbaria. The most substantial set of Bertero’s Caribbean
collections is preserved at TO-HG, at the University of Turin, which arrived
here from Balbis’ and Colla’s personal herbaria, which was carefully studied by
Vignolo-Lutati (1955) and is of cardinal importance in systematic, taxonomic,
floristicand monographic studies of the genera occurring in the Greater Antilles
(e.g, Anderson 1986; Delprete 1995; Delprete et al. 2000). Duplicate sets of
Bertero’s Caribbean collections were sent to A.P. de Candolle (G-DC), K. Kunth
(B, destroyed), K. Krug (B, destroyed), K. Sprengel (transferred to Urban, B, de-
stroyed), and A. Bertoloni (BOLO; Forneris et al. 1992), to the institutional her-
baria of Berlin (B, destroyed), Paris (P), Munich (M), Wurzburg (WB), Florence
(FD, and additional Bertero’s Caribbean specimens are also found at B, FI,
BRIT.ORG/SIDA 20(2)
Taste 1. Summary of Bertero’s field book of his trip to the Antilles and northern Colombia (1816—
1821), organized by localities, fascicle numbers (notebooks
cessively numbered by Bertero;fas-
cicle numbers followed by * were pencilled by Vignolo- er aes collection numbers, and page
numbers of the manuscript.
Fascicles numbers Collecting dates Collection numbers Book pages
In plantas Guadalu; imad j [Guadalupe]
primus 1816-17 {e212 1-92
secundus 1817 213-412 93-184
ertius 1817 413-587 85-26
quartus 1817-18 588-786 265-360
quintus 1818 787-1079 361-448
sextus 1818 1080-1171 449-529
Stirpium portoricensium historia [Puerto Rico]
septimus 1818 1172-1232 530-597
octavus 1818 1233-1348 598-685
Plantae domingenses (Santo Domingo, Dominican ey blic]
Nonus* & decimus”* 181 1 61 686-905
Stirpi d fl Maqdal Amer. merid. lect / ip [Rio Magdalena,
Colombia]
undecimus 1820 1562-1591 906-937
duodecimu
Stirpes in die S. ae oo Australi — pi Marta Province, seen
0-21 592- 938-9
anneen i i 1 a
[Plants collected in Jamaica, without title in front page]
82]
decimus quartus* | 1724-1747
994-] -
1070-1082
*The numbers of these fascicles were pencilled by Vignolo-Lutati.
HAL, L, M, MEDEL, MO, MPU, NY, P, P-JU, S, WB and probably some other
institutions.
At last, Bertero also collected many samples of seeds in the Greater Antilles
and sent them to Balbis, who had active exchange with several European bo-
tanical gardens (e.g., Delprete 1995). In many letters preserved at the Biblioteca
Reale di Torino it is possible to ascertain the frequent requests [rom various
botanical gardens of the precious seeds collected by Bertero in the West Indies.
Bertero’s Return to Europe (1821-1827)
In 1821 Bertero arrived in Paris, and from there he returned to Italy, where he
resumed his floristic explorations in Piedmont. In 1825 he was assigned to as-
sist Giuseppe Giacinto Moris (1796-1869) in botanical collections in Sardinia
(Moris 1837: Colla 1839; Mussa 1911; Mattirolo 1929; Pistarino et al. 1989), but
this collaboration was interrupted. He returned to Santa Vittoria d’Alba and
DELPRETE ET AL., CARLO BERTERO IN THE NEW WORLD 631
for some time he continued botanical collections in Piedmont, as documented
by his collection in the Gesso Valley of 1826.
Returning to Santa Vittoria d’Alba, Bertero’s feelings of frustration, caused
by the isolation from cultural centers and the monotony of small town life, were
eventually expressed ina letter to Balbis (Alba, 3 March 1826): “.. Il faudrait étre
au courant de la science, avoir des livres en quantité et des correspondants; je
suis loin de tout cela, et je me trouve, malgré moi, forcé de rester dans cette inac-
tion” [I need to be up to date with science, and to have books and correspon-
dents; | am far from all that, and | found myself, against my will, forced into
this inaction (Alba, 3 March 1826)|. The emptiness left by the death of his much
beloved mother, the only bond that could have kept him in Piedmont (Torino,
18 April 1827), made Bertero’s desire for new travels irrepressible. In 1827 he
returned to Paris, and followed the suggestions made by A.P. de Candolle and B.
Delessert (Paris, 25 July 1827) to choose Chile as his next goal, a land that was
botanically poorly known at that time. In preparation for this expedition, he
consulted the Joseph Dombey herbarium (P), constituted by material collected
in southwestern South America with Hippolito Ruiz and José Pavon (Steele 1964).
In several letters from Paris (e.g., 18 August 1827), Bertero communicated
to Balbis to be literally harassed by scientists, who masked their interests be-
hind apparent manifestations of friendship in order to gain the favor of receiv-
ing aset of his future South American collections. Candolle himself offered him
money in exchange fora set of his future Chilean collections, an offer that Bertero
refused with disdain. Nevertheless, in his correspondence, Bertero declared his
interest in sharing his collections not only with his colleagues Balbis and Colla,
but also with A.P. de Candolle and several other botanists. At the same time,
Bertero was conscious of the ruthless attitude of some botanists to appropriate
materials and results from their colleagues. As a consequence, he considered
suitable the offer of Candolle to include the descriptions of his newly found
Caribbean taxa in the Prodromus, before they would be published by others.
Candolle published several hundreds new species based on Bertero’s detailed
decriptions (sometimes copying them literally) of the Caribbean plants he ob-
served and collected. The field book was returned to the Turin Herbarium 16
years after Bertero’s death, by Alphonse de Candolle (A.P. de Candolle’s son),
with the following inscription on the front page (translated from French):
“Bertero, botanical manuscripts, previously donated by Balbis to Aug. Pyramus
de Candolle, with the agreement of Bertero, transmitted in 1857 to the Botani-
cal Museum of Turin by Alphonse de Candolle, as a token of recognition for a
present that he gave me of dry plants from Brazil. Geneva, 27 October 1857, Alph.
de Candolle” (Fig. 4).
In addition, Bertero planned to publish the new taxa that would result from
his future South American collections, before they would be supplanted by con-
BRIT.ORG/SIDA 20(2)
Z oitad 4s) |
ped ee
ee
Fig. 4, Pr i £+h fs} £[D 4 |e Oe Pe 7 tel L Lr gt y Alpt A Ct i|] “Bertero, Manuscrits
botaniques, Donnés anf par Balblse Aude h ie le Candolle, I’ i le Bertero, transmis en 1857
doll i d i t le don quiluia
au Musée botan que de Turin par A
été fait de ae séches du Bresil. Genie 27 Octobre 1857, Alph.de Grdolle®
DELPRETE ET AL., CARLO BERTERO IN THE NEW WORLD 633
temporary European botanists. These feelings were expre ed in several letters,
just before his departure for Chile: “... puisque je me suis de nouveau exposé a des
dangers pour faire des découvertes je veux que rien ne parait ce qu’en mon nom...”
[.. because I have again exposed myself to dangers in order to make discoveries,
I want them to appear under my name..] (Paris, 10 August 1827), and “.. mon
intention était de ramasser des matériaux les quels étant réputés de
quelqu’interét je voulais les publier moi méme et payer par ce moyen un juste
tribut 4 mon amour propre...” [... my intention was to accumulate the materials
that [regarded of some interest, and wanted to publish them myself and through
these means pay a tribute to my self-respect..] (Paris, 18 August 1827).
Bertero’s expedition to Chile, Juan Fernandez Islands, and Tahiti (1827-1831)
For the second time, Bertero travelled from Paris to Le Havre, and from there, in
October 1827, he embarked on a ship with destination Chile, again as a ship
doctor. That would be the last time that he saw the European continent. After
112 days of sea voyage, Bertero arrived at Valparaiso, and shortly after he re-
ceived permission for medical practice in the country, which allowed him to
pay his travel and living expenses.
Bertero’s explorations in South America turned out to be even more diffi-
cult than those he undertook in the Antilles. In fact, in his correspondence he
mentioned the many obstacles that he had to overcome: endless rains, impen-
etrable forests, unusable roads, earthquakes, and frequent dangers of all sorts.
In addition, he had to dedicate a considerable part of his time to his medical
practice, which he alternated with his botanical work, and had to postpone the
shipments of his specimens to Europe several times because of the obstacles
encountered in several Chilean ports. To make things more complicated, Bertero
travelled in Chile while the country was in revolution (Rancagua, 30 June 1828).
The Chilean government contacted Bertero in order to teach a course in Botany
and Agriculture, and even planned the creation of an acclimation garden, but
these projects were never realized (Valparaiso, 24 June 1829).
Bertero’s correspondence with Balbis and Colla includes a wealth of infor-
mation related to the flora of the regions explored, and documents his travels
from Europe to South America, and his life in Chile: France: Paris (25 July 1827,
8 August 1827, 10 August 1827, 18 August 1827, 26 August 1827, 14 September
1827), Le Havre (16 September 1827, 10 October 1827); Chile: Santiago de Chile
(11 March 1828, 11 March 1828), Rancagua (17 May 1828, 30 June 1828), Valparaiso
(21 June 1829, 24 June 1829, 6 July 1829, 28 November 1829, 28 November 1829,
6 February 1830, 6 February 1830, 6 July 1830, 26 July 1830, 7 August 1830, 10
September 1830).
Contrary to the first expedition, the information that can be extracted from
herbarium labels from the second expedition cannot be easily integrated with
634 BRIT.ORG/SIDA 20(2)
the data obtained from his correspondence. In fact, the collection dates of her-
barium specimens are limited to the year, reducing the possibility to reconstruct
the exact chronological sequence of Bertero’s movements (Fig. 5). The only sure
data obtained from the material preserved at TO-HG are those correlated with
his collections in Valparaiso June 1829, June-August 1830), Quillota July 1829),
and Juan Fernandez Islands July-November 1829) (Table 2). No herbarium
material was found at TO-HG from the trips that he mentioned in various let-
ters (Santiago, ll March 1828; Valparaiso, 24 June 1829, 6 July 1829, 26 July 1830),
in the surroundings of Concepcién, Petorca and Coquimbo. Nevertheless, in a
letter (preserved at the Academy of Sciences of Turin) he mentioned the send-
ing of a sample of native silver, extracted from the mines of Coquimbo, to the
Academy of Sciences of Turin.
In Chile, he collected mostly in the central region. First he botanized in the
surroundings of Valparaiso, and then in Vina del Mar and Quillota, on his way
to Santiago; from there he travelled south to Rancaguaa. It is still doubtful if he
travelled to Concepcion, Petorca and Coquimbo, which he mentions in one let-
ter to Colla (see below). Some references related to Bertero’s work in the region
were published by the Royal Society of London (1867), Pritzel (871), Saccardo
(1895-1901), Vignolo-Lutati (1956), Stafleu @ Cowan (1976), and Del Vitto 1986).
Bertero published an account on the Chilean plants known to him in sev-
eral articles in El Mercurio Chileno (Bertero 1828-1829), where taxa were listed
alphabetically, with notes on common names, their uses and peculiarities. This
work remained incomplete, arriving only to the letter Q, and was ended with
“(Se concluira)” [to be concluded], because the publication of the journal was
stopped, due to political turmoil. Bertero’s articles were translated in Italian
(Colla 1829a, 1829b, 1830), English (Bertero 1831, 1833) and German (Bertero
1832) immediately after their publication in Chile, and reprinted with notes by
Looser (1931), proving the importance of his work. Unfortunately, Bertero’s new
taxa were not supplied witha description and should be treated as nomina nuda
(invalid names according to the code of botanical nomenclature - Greuter et al.
2000). The logical continuation to Bertero’s work published in the Mercurio
Chileno is to be found in Colla’s Plantae rariores in regionibus chilensi pes
1834, 1835, 1837) and Moris’ Plantae chilenses novae minusque cognitae 183
1835). Also, a few of Bertero’s new taxa were described by Moris (1831, 1832, re
from plants grown in the Royal Botanical Gardens of Turin (from seeds sent by
Bertero from Chile). In addition, Bertero’s new taxa from his Chilean collections
were published in the works of many botanists from all over Europe, sometimes
validating his nomina nuda, or disregarding his proposed binomials.
In his botanical observations in Chile, Bertero was also the first to realize
that Cyttaria (Ascomycetes, described by Berkeley, 1841) was a fungus, and not
the strange “fruits” of Nothofagus (Minter et al. 1987).
Because of the ongoing war in Chile, he had to discontinue his medical
—
—
DELPRETE ET AL., CARLO BERTERO IN THE NEW WORLD 635
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ia rats
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7
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Fic. 5. Map of both Bertero’s two itineraries in the Antilles and Colombia (1816-1821), and in Chile, Juan Fernandez
Islands, and Tahiti (1827-1831). Rep ; wi : 9)
Taste 2. Bertero’s main localities and dates of his trips to Chile, Juan Fernandez Islands, and Tahiti
(1827-1831). Localities and periods were extracted from the letters exchanged among Bertero,
Colla and Balbis, and from labels of herbarium specimens preserved at the Turin Herbarium (TO-
HG).
Localities Collection periods
Valparaiso February 1828
antiago March 1828
Rancagua March 1828-February 1829
ntiago Herne 1829
Valparaiso Ju
Juan Fernandez Islands ih y- Ce cans
Valparaiso December 1829- ee 1830
Quillota March—May 1830
Tahiti November 1830-April 1831
practice. Therefore, from March to May 1830, he visited the Isla Masa Tierra, of
the Juan Fernandez Islands, accompanied by A. Caldeleugh, and returned to
Chile with plant specimens accounting for about 300 species. Guillemin (1830)
636 BRIT.ORG/SIDA 20(2)
published an extract of one letter written by Bertero, dated Valparaiso 7 July
1830, on the natural history of the Juan Fernandez Islands. This account, for
which Bertero should be rightly considered the author, is the first scientific ob-
servations of this archipelago. Among the plants collected by Bertero in Juan
Fernandez Islands, should be mentioned the genus Rea, a name that he pro-
posed in a letter to Colla (February-March 1830), and that was later published
by MJ. Decaisne (1833). Unfortunately, this genus was described one year ear-
lier by David Don (1832) as Dendroseris, and Rea should be treated as synony-
mous to it. Regarding the cryptogams collected by Bertero in Juan Fernandez
Islands, the vast majority of the new taxa were described by Camille Montagne
(1835a, 1835b).
On 28 September 1830, Bertero sailed from Chile, with the General Consul
of North America J.A. Morenhout, to the Society Islands and arrived at Tahiti
on 4 November 1830. Detailed information on Bertero’s trip to Tahiti was sup-
plied in a letter written by Morenhout (25 March 1832; Vignolo-Lutati 1956).
According to Morenhout, as Bertero arrived in Tahiti, he immediately started
actively botanizing and made substantial collections. A few months later, as he
received news about the ongoing revolution in France, Bertero sailed on a small
boat (of Morenhout’s property) directed to Valparaiso on 2 April 1831, leaving
his Tahitian collections to Morenhout, and arriving on the same day at the small
island of Raiatea (Society Islands), where he made his last botanical collections.
From Raiatea, Bertero wrote two letters (2 April 1831 to Colla, and 9 April 1831
to Morenhout); these were his last communications, at the young age of 42. In
his last letter to Colla, probably feeling a premonition of his imminent death,
Bertero wrote: “Adieu, bonne Santé, un Pater et un Ave pour mon ame dans le
cas quelle soit submergée!” [Good bye, and good health, recite a Pater Noster
and an Ave Maria for my soul in case that it will be sunk].
Almost one year later, after loosing any hope of Bertero’s survival,
Morenhout wrote a long letter (25 March 1832) to the Academy of Sciences of
Turin, where he informed them of Bertero’s disappearance, and of his grand
future plans to explore Peru, return to Colombia, and start his explorations in
North America. Morenhout concluded his letter with the following sad notes
(free translation from French): “Destiny has not permitted that he [Bertero] re-
alized his grand projects, as he never stopped loving his interesting research!
He deserves the sorrow of all his friends in science, and most of all from You,
Sirs [the members of the Academy of Sciences of Turin], and from his homeland
that he always fondly remembered. Honoured by his friendship, | have known
the most intense sentiments of his soul, and I know that he had no other passion
than Botany nor other ambition than being proud of You and his Homeland!”
While still in Chile, Bertero sent his collections to Baron Delessert at Paris,
who took the responsibility of distributing duplicate sets to Balbis (now at TO-
HG), Colla (now at TO-HG) and A-P. de Candolle (G-DC), keeping one for him-
DELPRETE ET AL., CARLO BERTERO IN THE NEW WORLD 637
self (G-DEL), and conserving the rest for the return of Bertero, as he explained
in several letters from Paris (10, 18, and 26 August 1827) and from Valparaiso
(24 June and 6 July 1829). Fearing that some of the boxes might become lost,
Bertero also planned duplicate sets to be sent directly to Balbis (Paris, 26 August
1827). Such a plan was not followed, however, because Bertero did not receive
any communications from Balbis for more than a year anda half. The last let-
ters sent from Balbis were dated 3 August 1828 and 31 January 1829, which were
delivered contemporarily to Bertero on June 1829 (Valparaiso, 24 June 1829).
Bertero’s entire collection from Chile—approximately 15,000 specimens—
was sold by Delessert’s heirs at an auction, several years after Bertero was de-
clared lost at sea. The collection was bought for 1,200 Francs by the travel com-
pany Unio Itineraria (“Reise Verein”), based at Esslingen (Germany) and owned
by E. Steudel and C.F Hochstetter. Unio Itineraria dismantled Bertero’s Chilean
collection, and sold the duplicate sets to the institutional herbaria of Berlin (B),
the British Museum of London (BM), Kew (K), Leiden (L), Nancy (NCY) and
Paris (P), and to the personal herbaria of E. Cosson (now P), B. Delessert (G-DEL),
A.P. de Candolle (G-DC), J. Cambessédes (now at MPU), A. Wigand (now at MB),
and probably some others. According to Turrill (1920), 5,000 specimens are pre-
served in the Paris herbarium (P), with labels reporting localities of collection
and accurate descriptions. Other than those indicated above, additional Bertero’s
Chilean collections are found at A, CGE, CN, E, E Fl, G, GH, KIEL, LE, M, MEL,
MO, NY, OXE PC, SGO, TCD; and those from Juan Fernandez Islands at B, FE G,
GH, K, L, MEL, MPU, NY, P P-JU, PC, SGO (Lanjouw & Stafleu 1954, and pers. obs.).
In 1834, Morenhout sent Bertero’s Tahitian collections to Alcide Dessalines
d’Orbigny (1802-1857) who travelled in South America in 1826-1834 (Orbigny
1834-1847), at Paris in a large crate. As the crate arrived at his destination, it
was noticed that it had been opened, and part of its contents was missing. A
detailed account on the Tahitian collection made by Bertero and Morenhout
sent to d’Orbigny, as well as descriptions of many new species, was published
by J.A. Guillemin (1836, 1837). In the first article of the series, Guillemin stated
that duplicates of these collections were sent to the Museum of Natural History
of Paris (P), to the Royal Academy of Turin (which material has been entirely
transferred to the herbarium of the University of Turin, TO), and to the per-
sonal herbaria of A.T. Brongniart (now at P) and B. Delessert (now at G-DEL),
and other non specified botanists. More research on the distribution of dupli-
cates of Bertero’s Tahitian collections is necessary (none cited by Lanjouw &
Stafleu 1954). Nevertheless, specimens of his Tahitian incomplete sets were
found in Paris (P), Berlin (B, destroyed), Kew (K), and at the Delessert herbarium
(G-DEL). Only four specimens of Bertero’s Tahitian collections were found at
the Turin herbarium (TO-HG), and no reference to them could be found in the
correspondence.
Additional references about the distribution of Bertero’s collections can be
i
638 BRIT.ORG/SIDA 20(2)
found in: Anonymous (1833), Laségue (1845), Caruel (1874), Parlatore (1841,
1874), Saccardo (1895-1901), Urban (1902), Mattirolo (1907), Ciferri (1931),
Martelli (1931), Vignolo-Lutati (1931b), Lanjouw & Stafleu (1954), Del Vitto
(1986), Matthei (1986), Rodriguez Rios (1986), and Del Vitto et al. 1993).
—
ACKNOWLEDGMENTS
We are grateful to the Library of the Department of Plant Biology of the Uni-
versity of Turin, the Royal Library of Turin (Biblioteca Reale di Torino), the Acad-
emy of Sciences of Turin (Accademia delle Science di Torino), the Civic Library
of Turin (Biblioteca Civica di Torino), the Regional Museum of Natural Sciences
at Turin (Museo Regionale di Scienze Naturali), and The New York Botanical
Garden for their support toward this project. We are also thankful to Jim Grimes
for information about Bertero’s Chilean collection at the Herbarium of the Royal
Botanic Gardens, Victoria (Australia), to Lucia Cortez for proofreading the Span-
ish version of the abstract, and to Mike Nee, Alain Liogier, Richard Howard and
Tod Stuessy for reviewing the manuscript and for their valuable comments.
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ON THE DERIVATION OF THE GENERIC NAME
ILLIAMNA (MALVACEAE)
William A.Weber Paul A. Fryxell
University of Colorado Museum Herbarium, Plant Resources Center
Boulder CO 80309, U.S.A. University of Te
Austin, TX 78712, a
Edward Lee Greene is well knownasa classics scholar anda botanist with strong
opinions. His prowess as a classics scholar is well attested to by his historical
work (Greene 1909). His idiosyncratic ideas are too well known to need docu-
mentation. But this combination of characteristics leads to some interesting
speculations with regard to certain generic names that he proposed. In particu-
lar, we deal here with the name Iliamna, which he proposed for a genus of west-
ern American Malvaceae (Wiggins 1936).
In publishing the name, Greene (1906) did not state how he chose the name
or offer any indication of its derivation. Other sources indicate its origin to be
unknown (e.g. Gleason 1952; Genaust 1983) or at least mysterious (“Greek, sig-
nificance obscure”: Hitchcock & Cronquist 1973).
Our first considerations of the question took into account Greene's classi-
cal learning and led us to the roots ilyos (Greek: mud) and amnis (Latin: per-
taining toa river), which makes a certain amount of sense, since the plants con-
cerned occur in wet habitats. However, it seemed unlikely that Greene would
have mixed Greek and Latin roots, so this derivation left us unsatisfied.
We next serendipitously discovered that there is a place name in Alaska,
Iliamna, applied to a bay, a lake, a point, a village, and a volcano. The native
name of the lake was reported as Ozle|ro Bollschi] Ilyamna on the 1852 Russian
Hydrog. Dept. Chart 1455. This feature was earlier called Ozyero Shelekl
an 1802 Russian map. According to G.G. Martin, USGS, Hiamna is said to be “the
name of a mythical great blackfish, supposed to inhabit this lake, which bites
holes in the bidarkas [boats] of bad natives” (Orth 1967). The volcano’s name
was published by the Russians (Orth 1967) as Sopka Ilymna (Capt. M.D.
Tebenkov, 1852, map # 5). It has therefore been suggested that the Russian name
Ilym is the root, but this is speculative.
We presume that Greene encountered the name, liked the sound of it, and
found an occasion to apply it botanically. But did he apply it because of its sound
or because of its meaning? Is the origin of the name Russian or Amerindian?
The genus Iliamna does not occur in Alaska; its species have their northern-
most limit in British Columbia.
We have now learned that what appears to be the “true” derivation of the
SIDA 20(2): 645 — 646. 2002
646 BRIT.ORG/SIDA 20(2)
geographical name, which Greene presumably appropriated for botanical
purposes. The Dena’ina are an Athabascan people of Cook Inlet and the Alaska
Peninsula where the geographical name was used. Their name Nilamna (pho-
netically) was modified by the early Russian colonists into Iliamna. The name
derives from two words, nila (islands) and vena (lake), that is, “islands in the
lake.” We are grateful to Lydia Black of the Anthropology Department, University
of Alaska, Fairbanks, for this information.
Thus, although we now know the derivation of the name Iliamna, we re-
main in the dark as to why Greene chose it as a botanical name. He may have
simply liked the sound of it, but it is also possible that he chose it to confound
his botanical critics into seeking a classical derivation of a name that did not
have classical roots.
REFERENCES
Genaust, H. 1983. Etymologisches Wérterbuch der botanischen Pflanzennamen. ed. 2.
Birkhauser Verlag, Basel.
Gteason, H.A.1952. The new Britton and Brown illustrated flora of the northeastern United
States and adjacent Canada. Hafner, New York. 3 vols. [vol. 2, p. 532].
Greene, E.L. 1906. Certain Malvaceous types. Leafl. Bot. Observ. Crit. 1:205-209.
Greene, E.L. 1909. Landmarks of botanical history, Part |. Smithsonian Institution, Washing-
ton, D.C.
HircHcock, C.L.and A. Cronouist. 1973. Flora of the Pacific Northwest: an illustrated manual.
Univ.Washington Press, Seattle.
Ort, D.J. 1967, Dictionary of Alaska place names. Geol. Survey Prof. Pap. 567.
Weber, W.A. and P. Fryxeit. 1992.What's in a name? Derivation of /liamna. Aquilegia 9(2):9.
Wicains, ILL. 1936. A resurrection and revision of the genus /liamna Greene. Contr. Dudley
Herb. Stanford Univ. 1:213-231.
WHAT IS THE WRITER OF A FLORA TO DO?
EVOLUTIONARY TAXONOMY OR PHYLOGENETIC
SVo LEMATICS:
George M. Diggs, Jr. Barney L. Lipscomb
ee of Biology Botanical Research Institute of Texas
n College 509 Pecan Street
U.S Ft. Worth, TX 76102, U.S.A.
& Botanical Research Institute of Texas barney@brit.org
iggs@austincollege.edu.
ABSTRACT
Plant clas sication angim momenclatire are in a continuing state of flux and heated debate between
PL g “evolutionary” or “Linnaean taxonomy”,
and 2) cladists SUpEOrenG: ype sy stematics” or ue While it isa multifaceted con-
troversy that has spanned several decades, relatively little attention has been focused specifically on
the Be for ae ‘The two goals of this paper are 1) to give special emphasis to the
een of the eee for a writers loras, ne 2) to provide an overview of some of the
fe
ormat
So
otanists interested in floristics. We exam-
ae some of the issues we have confronted in our floristic work, discuss how we are attempting to
balance the strengths and weaknesses of both approaches, and indicate what we believe is the best
albeit imperfect, approach to the writing of floras at the present time. We argue that, for both practi-
cal and theo dca reasons, a modit Hed taitional system pein: jempenenrat ranked hierar-
chies) be used in f1 grouy iminating Il po Dpayene groups despite
some nomenclatural instabil ie) Fuster hese fl hould i newly dis-
covere ‘ Eby SERSUE teleuonships (even if too 20 Pre aINary; eaierye or DpappropHats for nomen-
O facilitate
as oe an nandérstanding of plant Ganon as Soible,
RESUMEN
] j,] ] rad ey ea - a ot fi : pa ]
] rier ce
La LIASTLINACIUIL Ds
1 ‘eo ] ]
j
op Ss “taxonomia Linneana’o la “evolutiva”™ y2) cladistas
ae apoyan la “sistematica ilogenética’ fe) “cladonomia’ Mientras esta controversia se ha extendido
ay. ] er 8s ‘ ae (ents [meen pene
a ee ]
durante VALIAS UCLCAaUaAS
c
Las dos metas de este artieio son 1) dar un énfasis seSPealg a ba implicaciones de la controversia para
los autores de floras, y 2) ofrecer una visio 1 ry “17
J; : a ]
te)
1 iad F a
fm |
Pa ee E i | le | ltad
que hemos confrontado en bajo floristi 1 i | ilil as
bondades y debilidades de ambos mierodoe e indicamos lo que creemos que es el igor aunque
imperfecto, método de escribir floras e en la ee pac ncntanlos que, tanto aa razones tedricas
como pt acticas, se use e fl sistema
1 ooh Bare le * J ] ] Paces
Oo ae r A vg oO L
| 1). Ademas, estas f] deberian incorporar inf acion sobre las
relaciones filogenéticas recién descubiertas (incluso si son demasiado preliminares, tentativas, o
causen iniestabilidad
inapropiadas para cambios nonienclating: es) y aan éstas en la familia apropiada ei tratamientos
genéricos para facilit a compr I posible de | |
SIDA 20(2): 647 — 674. 2002
648 BRIT.ORG/SIDA 20(2)
INTRODUCTION
Plant taxonomy is the science that deals with the identification, nomenclature,
and classification of plants. The term plant systematics (or systematic botany)
is often used synonymously with plant taxonomy (as is done here), but some-
times has the connotation of mainly using recently developed techniques or
approaches such as chromosomal studies, electron microscopy, molecular biol-
ogy, or cladistics to answer questions about plant relationships. From the defi-
nition of plant taxonomy it follows that the primary goals of the discipline are
to:
1) identify and describe all the various kinds of plants;
2) develop a system of naming plants le.g., International Code of Botanical No-
menclature (Greuter et al. 2000) or potentially a future version of the
PhyloCode (2002);
3) arrange plants with common characteristics into groups that reflect their
evolutionary relationships (Lawrence 1951; Porter 1967; Radford et al. 1974;
Jones & Luchsinger 1986; Judd et al. 1999, 2002).
In terms of nomenclature, the goal of plant taxonomy has been to develop a
uniform, practical, and stable system of naming plants—one that can be used
by both plant taxonomists and others needing a way to communicate precisely
and retrieve information about plants. In the words of Stevens (2002), “The value
of any naming system is how effectively it establishes conventions that allow
people to communicate and to develop their ideas. ...”
Regarding classification, the goal has been to arrange plants with common
characteristics into groups that reflect their relationships—in other words, to
develop a scheme of classification that is useful—that conveys maximum in-
formation and has predictive value. Since the time of Darwin,a primary goal of
plant taxonomists has been to reflect phylogeny or evolutionary history in their
systems of classification. There are several reasons for this. One is that taxono-
mists want their classification system to reflect the reality of the evolutionary
history of life on earth. Second, a system that reflects evolution should have
maximum predictive value and usefulness (since related species should share
similarities due to common descent). While this basic evolutionary approach
is agreed on by virtually all botanists, in recent years there has been heated
debate between two main schools of taxonomists:
=
1) traditional taxonomists practicing what is sometimes referred to as “Linnaean
classification” (Brummitt 1997) or “evolutionary taxonomy.” Traditional or evo-
lutionary taxonomists, while attempting to have a classification system based
on evolutionary relationships, also try to reflect the amount of evolutionary
change undergone by groups. In addition, they try to incorporate other goals,
including practicality and stability, into the classification system (see Brummitt
FLORAS TAXONOMY SYSTEMATICS? 649
1997 for a detailed discussion of traditional classification). The names Linnaean
classification or Linnaean taxonomy (Stuessy 2000; Forey 2001, 2002; Nicolson
2002), are perhaps inappropriate since the system is very different from that
established by Linnaeus. In its more recent version it is perhaps better called
“evolutionary taxonomy” (Sanders & Judd 2000) or “evolutionary systematics”
(Grant 2001b) signifying the attempt to reflect evolutionary relationships. Grant
(2001b) uses the econyan TTES “to include the two subschools of traditional
taxonomy y systematics. ” When considered from the standpoint
of nomenclature, this isa system that incorporates binomial nomenclature (two-
part scientific names consisting of a genus name and specific epithet) and a
hierarchy of formal ranks (e.g., family, genus, etc.). The nomenclatural applica-
tion of this viewpoint has been referred to as the “L-code” and its principles are
embodied in the International Code of Botanical Nomenclature (Greuter et al.
2) cladists, whose method of constructing phylogenies is derived from the ideas
of the German entomologist Willi Hennig, practicing “phylogenetic
systematics’ or “cladistic classification” (referred toas “cladonomy” by Brummitt
1997) based explicitly and solely on phylogenetic relationships. In other words,
the overriding goal is that classification should reflect the branching patterns
of evolution. It should be noted that in a clade-based classification and result-
ing nomenclature system, there are no formal ranks, including family or genus,
and no formal binomial nomenclature (de Queiroz & Gauthier 1992; Lidén et
al. 1997; Cantino 1998; Brummitt 2002). The nomenclatural application of this
viewpoint has sometimes been referred to as the “P-code’” and its ideas are em-
bodied in the PhyloCode (2002).
Currently, classification and
| lsystemsare in astate of flux between
these two main opposing camps—both of which attempt to reflect evolution-
ary relationships. Those practicing cladistic systematics have made major con-
tributions to our understanding of plant evolution, and have brought about some
long overdue changes. In fact, some of their methodology has contributed toa
well-recognized revival in taxonomy/systematics. Some of the most evident
examples of this are the incredible breakthroughs in knowledge of plant rela-
tionships resulting from molecular phylogenetics. It should also be noted that
there is significant variation in the extent to which various “phylogenetic” sys-
tematists follow all of the implications of the cladistic approach—most, for ex-
ample, still use binomial nomenclature even though they may disagree with it
on theoretical grounds. At the same time, the system of nomenclature (bino-
mial, etc.) and hierarchical classification that has developed over the past 250
years has served and continues to serve the botanical and broader communi-
ties well. There are thus positive aspects to both of these approaches.
Because of major differences in philosophy and methodology, the classifi-
650 BRIT.ORG/SIDA 20(2)
cation systems produced by proponents of evolutionary taxonomy and cladis-
tics are often quite different. It is not unexpected then that the two conflicting
viewpoints have produced a vigorous and heated debate (e.g., Nixon & Carpen-
ter 2000), which has even been referred to as a “maelstrom” (Benton 2000). The
proposed approach of phylogenetic classification has certainly not “mostly been
politely accepted by the systematic community” as stated by Schander (1998).
In fact, the tone of a few of the articles and discussions (on both sides of the
argument) has been surprisingly impolite by the standards of modern scien-
tific discourse, with Webster (2002) referring to the arguments as “an ideologi-
cal cacophony of bombast and invective.” There is a voluminous literature on
the subject, including numerous recent articles about the different taxonomic
and nomenclatural approaches (e.g., Moore 1998; Stevens 1998; Diggs et al. 1999
(Appendix 6); Mishler 1999; Benton 2000; Cantino 2000; de Queiroz 2000;
McNeill 2000; Nixon & Carpenter 2000; Sanders & Judd 2000; Stevens 2000;
Stuessy 2000, 2001; Withgott 2000; de Queiroz & Cantino 2001; Grant 200la,
2001b; Langer 2001; Lee 2001; Pennisi 2001; Berry 2002; Brummitt 2002; Forey
2002; Kress & DePriest 2002: Nicolson 2002; Stevens 2002; Webster 2002). Sym-
posia and workshops have also been held (XVI International Botanical Con-
gress— August 1999; Smithsonian’s National Museum of Natural History—March
2001; Hunt Institute for Botanical Documentation—June 2002), and a new sys-
tem of nomenclature has been proposed (PhyloCode 2002). However, few au-
thors or discussions have specifically addressed the special problems faced by
writers of floras (but see Stevens 1998, Sanders & Judd 2000, and Berry 2002).
This controversy is actually multifaceted, with many quite different aspects
(e.g.,are taxon names defined?—Stuessy 2000, 2001, de Queiroz 2000, de Queiroz
& Cantino 2001; which system will ultimately be more stable?—Forey 2002)
that are beyond the scope of the discussion here. In this paper we are focusing
on the implications for floristics.
IMPLICATIONS FOR FLORISTICS
As writers of a flora (the Illustrated Flora of East Texas project—Diggs et al. in
prep, wwweasttexast lora.org), we and our co-authors are faced with the ques-
tion of what type of classification and nomenclatural concepts should be followed
in a large regional flora (ca. 3,300 species or roughly 1/6 the species in North
America north of Mexico). We are acutely interested in this question, because
as floristicians we must translate and synthesize a variety of types of botanical
research, both practical and theoretical, into a form usable by a very diverse
audience—one that ranges from professional taxonomists and other scientists
to lay botanists, students, and interested amateurs, many of whom are unac-
quainted with taxonomic methods. In fact, we believe that addressing the needs
of diverse users is one of the most important tasks of floristicians. Further, de-
veloping a “general-purpose system” that effectively addresses the needs of
TAD Re SYSTEMATICS? 651
multiple users is often considered to be the “historical and continuing func-
tion” of taxonomy as a whole (Cronquist 1987). The answer to the question of
what type of concepts should be followed varies greatly depending on who one
asks. The most conservative voices would say that the traditional system of
nomenclature, a ranked hierarchical system of classification, traditional fami-
lies, etc., should be used due to both theoretical and practical considerations.
Some would even argue that clearly polyphyletic traditional families (e.g.,
Liliaceae in the broad sense) should continue to be used in floras since this is a
very useful and practical approach. The most extreme voices on the other side
(ie.extreme cladists) would say that no set categories should be recognized (e.g.,
no families, no genera; instead, only supportable clades), only monophyletic
groups (=a common ancestor and all its descendants; Fig. 1) should be given
taxonomic recognition (ie., no paraphyletic groups should be allowed—cur-
rently many genera and families are paraphyletic; paraphyletic groups are de-
fined as those containing a common ancestor and some, but not all, of its de-
scendants). Further, the extreme cladists argue that binomial nomenclature
should be replaced (since genera have no objective reality, there can be no ge-
neric names and hence no binomials; only clade-based names should be used).
Interestingly, our previous flora (Shinners & Mahler’s Illustrated Flora of North
Central Texas—Diggs et al. 1999) was criticized by individuals from both ex-
tremes. Despite having more information on cladistics (lengthy appendix dis-
cussing the issue, discussions in numerous family synopses) than any other
regional or state flora we know of (e.g., Hickman 1993; Yatskievych 1999; Rhodes
& Block 2000; Wunderlin & Hansen 2000), we were criticized for not applying
the cladistic approach throughout the flora. Likewise, some conservative bota-
nists were disturbed by decisions such as lumping some groups (e.g., Najas into
the Hydrocharitaceae), splitting others (e.g., Senecio into Senecio and Packera),
and following an alphabetical rather than a traditional sequence (showing sup-
posed relationships) of families and genera (a logical impossibility since a writ-
ten flora is linear and evolution is a branching process). One thing to keep in
mind when discussing this clash of viewpoints is the realization that any sys-
tem of classification, nomenclature, and written presentation will be an im-
perfect reflection of the complexity represented by the evolutionary history of
life on earth. In the words of Benton (2000), ‘phylogeny is real, classification is
not.” While we now | ly sophisticated and diverse sources
of data, factors such as extinction, an incomplete fossil record, and the com-
plexity of evolutionary processes (e.g., reticulate evolution—Wagner 1954; see
discussion below) will prevent us from producing completely accurate phylo-
genetic reconstructions. Despite these limitations, as floristicians attempting
to produce a useful flora, we have to use approaches to classification and no-
menclature that best reflect a diverse and complex set of needs.
Itis generally agreed that the primary goal of a flora is to allow identifica-
—
652 BRIT.ORG/SIDA 20(2)
POLYPHYLETIC GROUPS
MONOPHYLETIC GROUPS
PARAPHYLETIC GROUPS
Fic. 1 ni : H £ | hyl +t} monophyletic I | ee . (f. Dig aes | 1999).
tion of the plants treated. However, there are a number of secondary goals. Ac-
cording to Sanders and Judd (2000), these are: 1) to provide entry into the sys-
tematic literature; 2) to provide thumbnail summaries of the current state of
FLORAS TAX MY SYSTEMATICS? 653
9 systematic, ecological, ethnobotanical, etc.); 3) to serve
our knowledge (including g
as a reference for other professionals; and 4) to fix the concepts of taxa, espe-
cially families and genera, in the minds of users. Generally we agree with these
secondary goals, and expended considerable thought and effort in applying
them in our previous floristic effort (Diggs et al. 1999). We also agree with Sand-
ers and Judd (2000) that there isa critical need for the collaboration (and prob-
ably more importantly cross-training) of floristic, monographic, and phyloge-
netic researchers. However, we disagree with Sanders and Judd (2000) in how to
accomplish the fourth of their stated goals. They argue that the methods of
phylogenetic classification should be applied consistently in floras (e.g., only
monophyletic groups allowed, hence precluding paraphyletic families). We
believe that this approach, if inflexibly applied, would hinder the primary goal
(of a flora) of allowing effective identification. Further, if all aspects of the cla-
distic approach are followed (e.g., elimination of ranks and binomial nomen-
clature) in a flora, we envision significant erosion of this primary goal. The loss
of many morphologically coherent and easily recognizable paraphyletic fami-
lies, the discontinuity in information retrieval due to a radical change in no-
menclature, the lack of effective mnemonic devices to replace such widely rec-
ognized and practically important ranks as family and genus, and instability
in nomenclature (and hence identification) associated with rapidly changing
cladograms are a few of the reasons for this concern.
In some cases, floristicians, for practical or historical reasons, are unable to
apply even the most important recent phylogenetic discoveries in their floras.
For example, the critically important Flora of North America Project, because
of the long time span necessary for such a massive multi-volume work, had to
adopt a standard years ago (Cronquist system of families)—hence, the Liliaceae
(in the broad sense), now known to be clearly polyphyletic, is still being recog-
nized in a forthcoming volume (with an extensive discussion of phylogeny).
While we strongly agree with cladists that polyphyletic groups should be elimi-
nated (whenever possible), we disagree with the advisability of eliminating the
numerous useful and meaningful paraphyletic groups, particularly at the lev-
els of family and genus (see further discussion below). We would add three other
goals to the four (Sanders & Judd 2000) enumerated above: 5) to address spe-
cifically the needs of diverse users (discussed above); 6) to connect the work of
monographers and other researchers to the “consumers of botanical informa-
tion” (T. Barkley, pers. comm.; Barkley 2000) who need to use these discoveries;
and 7) to use systems of classification and nomenclature that allow meaning-
ful comparisons with other floristic works. In other words, for conservation,
biogeographical, ecological, etc. purposes, it should be easy to compare data
such as the total number of species, the number of endemics, or the number of
introduced species from flora to flora (with the realization that the compari-
sons will be far from perfect, but useful nonetheless). Ultimately, we somehow
654 BRIT.ORG/SIDA 20(2)
hope to combine several important but not necessarily compatible approaches.
We not only want to produce a useful, informative, and user-friendly flora, but
also one that accurately reflects evolutionary history (ie., be phylogenetically
informative) and incorporates recent discoveries in botany.
APPROACH TAKEN IN THE ILLUSTRATED FLORA OF EAST TEXAS
After considerable thought, discussion with a variety of individuals, and a re-
view of the pertinent literature, we are taking what we hope is an intermediate,
albeit somewhat conservative, approach in the Illustrated Flora of East Texas.
Our goal is to provide maximum information while retaining a practical and
utilitarian framework.
Cladistic side
On the cladistic/phylogenetic systematics side of the argument, a number of our
decisions have been influenced by the desire to increase information content
and accuracy:
1) We are attempting to provide detailed information on the known evolution-
ary relationships of various plant groups. A tremendous amount of new infor-
mation has become available recently (primarily, but not exclusively, as a re-
sult of the application of Hennigian principles to molecular data), and as much
of this as possible is being included and references provided. For example, in
the draft family synopsis of the Lemnaceae (duckweed family), we (Diggs et al.
in prep.) have included the following statement.
Lemnaceae are tiny and extremely reduced morphologically making it difficult in the past to deter-
mine the phylogenetic relationships of the family. Kvacek (1995) suggested that the fossil genus
Limnobiophyllum is a fossil link between Araceae and Lemnaceae, and Stockey et al. (1997), using a
cladistic approach and mater i o Lim pao hyllum, soleus ae ane (a free-floating member
of the Araceae) plus Limnobio} Indeed, this link-
age of Lemnaceae to Araceae goes back over 175 seat (Hooker and Brown in Smith 1824; see discus-
sion in Leset al. 2002). In addition ies | linked Lemna with Pistia (Araceae) (Duvall
etal. she raorererenty 9 faeere subtan yahreldese(Sienen etal. 1995). In fact, the Lemnaceae
is considered by many within Araceae jACK-IN-THE-PULPIT family) by
extreme ae tion, and it has been suggested that Lemnaceae be reduced to a subgroup within a
more inclusive Araceae (Mayo et al. 1995, 1997, 1998; Stockey et al. 1997). From a cladistic standpoint,
——
Araceae (without Lemnaceae) are paraphyletic and inappropriate for formal recognition
Even where very preliminary information is available, we have included and
referenced it in an attempt to foster a better understanding of evolutionary re-
lationships.
2) Also on the cladistic side (and on that of most other plant taxonomists), we
are rejecting all clearly polyphyletic groups, even when these are practical and
of long-term or wide usage. The best example of this is the Liliaceae (lily fam-
ily) sensu lato (in the broad sense). Extensive morphological and molecular data
nna SYSTEMATICS? 655
now clearly indicate that as broadly conceived, this family is a heterogeneous
mixture based on superficial similarities in flower structure (e.g., Fay et al. 2000;
Rudall et al. 2000b). In fact, recent molecular studies (e.g., Chase et al. 2000)
have shown that species traditionally treated in the Liliaceae should be placed
in at least four different orders. Asa result, we are recognizing 14 separate fami-
lies (all previously treated in the Liliaceae) for East Texas. However, from the
standpoint of usability, we are incorporating a table in the Liliaceae (narrow
sense) family treatment that clearly indicates in what family the genera for-
merly included in the Liliaceae (broad sense) are now placed. Furthermore, in
the main key to families, as many as possible of the liliaceous (broad sense)
families will be clustered together and clearly indicated. Likewise, the genus
Nolina (bear-grass) and its relatives, which have often been included in the
Agavaceae (agave family), are now known to not be closely related to that fam-
ily and we are excluding them. In a draft family synopsis we (Diggs et al. in
prep.) say,
In the past, taxa included here in the Nolinaceae were sometimes included in a apreadly conceived
Liliaceae (e.g., Kartez 1999) or often in the Agavaceae (e.g rell & Johns 1.1999;
Verhoek & Hess 2002 following Cronquist 1988) based on certain earn eanilacnice How-
ever, recent evidence suggests that the Agavaceae and Nolinaceae are not closely related and should
be recognized separately (Dahlgren et al. 1985; Eguiarte et al. 1994; Bogler & Simpson 1995, 1996,
Kubitzki et al. 1998; Chase et al. 2000). Molecular euldetice indicates that Nolinaceae is closely re-
lated to Convallariaceae and Ruscaceae, anc es (e.g., Chase et al. 1995a; Chase et al. 2000,
inaceae arouse pe meade in the Convallanaccac Fol-
Fay et al. 2000) have suggested that the Noli
lowing such preliminary studies, Judd et al. (1999), lumped
On the other hand, Rudall et al. (2000a) and Judd et al. (2002) included the Nolinaceae in a broadly
ire anelad USSR EE HOWENEE there has been disagreement in molecular analyses of the family
Rudall et al. 2000a; Yamashita & Tamura 2000). S he Nolinaceae
sepeaee to be a well-defined ne letic group (Bogler & Simpson 1995, 1996), and until the phy-
logeny of this complex is clarified and the nomenclature more stable, we are recognizing it as a dis-
tinct family.
Hopefully, such explanations will allow users to see that the understanding of
plant relationships is still changing and improving. With such insights, we hope
that non-taxonomists will be less resistant to needed nomenclatural changes.
3) Again on the cladistic side, when established useful family concepts are not
excessively distorted, we are lumping small groups whose relationships have
now become clear. For example, the monogeneric family Najadaceae (the ge-
nus Najas) is now known to be derived from within the Hydrocharitaceae (R.
Haynes, pers. comm.; Shaffer-Fehre 1991; Les et al. 1993; Les & Haynes 1995;
Hayneset al. 1998; Haynes 2000). Including Najas in the Hydrocharitaceae more
accurately reflects evolutionary history, yet does not substantially modify the
concept of the Hydrocharitaceae nor distort it beyond the bounds of usability.
We are therefore following several recent floristic treatments (e.g., Thorne 1993;
Diggs et al. 1999) in lumping Najas into the Hydrocharitaceae.
656 BRIT.ORG/SIDA 20(2)
4) Our families, genera, and species are arranged alphabetically. Some very tra-
ditional taxonomists want “related” families placed together in the linear se-
quence physically required of a book. However, the complex branching pattern
of evolution does not follow such a linear form and thus any linear sequence is
highly arbitrary and distorts actual evolutionary relationships. An easy to use
alphabetical sequence, while not reflecting relationships, at least does not dis-
tort them. In addition, an alphabetical arrangement allows quick and easy ac-
cess to the material so arranged.
Evolutionary taxonomy side
On the evolutionary taxonomy side of the argument, a number of our decisions
have been influenced by both practical and/or theoretical considerations:
1) We are continuing to use the system of nomenclature that has developed
over the past 250 years (International Code of Botanical Nomenclature—Greuter
et al. 2000). This (particularly the use of binomials) is an eminently useable
system that addresses the needs of an audience far broader than the taxonomic
community (“the consumers of botanical information”—T. Barkley, pers.comm:;
Barkley 2000). We believe that eliminating it would cause great confusion
among the many non-taxonomists who use plant names. In fact, it is likely that
if plant taxonomy went toa specialized non-binomial, clade-based system, some
separate static system of “accepted plant names” would be developed by the
horticultural community or other user groups (e. 8 agricultural, eee) con-
servation). Such a move would both marginalize plant taxonomy and ultimately
result in a nomenclatural system with much less information content than at
present. This practical consideration may well be one of the most important
reasons for maintaining our current system of classification and nomenclature
In fact, even those developing the PhyloCode (2002), the nomenclatural system
being produced by phylogenetic systematists, have not yet come to grips with
what to do regarding the naming of “species.” It is interesting to note that Stevens
(2002) has argued that binomials have been used so long and so widely (across
many cultures and in many contexts) because they are inherent in human per-
ception—in other words, having such a two word nomenclature system may be
built into the organization of our nervous systems. Nixon and Carpenter (2000)
in a similar vein suggested that, “Our natural form of communication (even as
evidenced by the common human binomial system of naming ourselves) is
clarified by the use of ranks and binomials.” Likewise, anthropologist Brent
Berlin (1992) noted that there are widespread cross-cultural regularities in the
classification and naming of living organisms by people in traditional, nonlit-
erate societies—these systems more closely approximate Linnaean binomials
than clade-based nomenclature. Further, when the diverse users of a flora are
considered, a radical shift in the system of nomenclature used seems particu-
larly ill-advised, especially at a time when the public needs to be brought closer
FLORAS | MY SYSTEMATICS? 657
to, rather than pushed further away from, an appreciation and understanding
of botany. Indeed, these ideas raise questions about a “dominant” code (T. Bark-
ley, pers. comm.) of nomenclature. Undoubtedly, a P-code will be used, but it
remains to be seen how widely such a system will be accepted by the diverse
users of botanical information—the L-code may continue to be used as the pri-
mary or dominant code by the user community long after a functioning P-code
(presumably the PhyloCode) is finalized.
2) While many taxonomists have long realized that the traditional ranked cat-
egories (e.g., family, genus) are not used consistently and are simply human con-
structs (unlike species which have some biological reality), they do, however,
provide important mental pegboards or mnemonic devices to allow a practical
way to arrange our thinking. As Stevens (2002) has said, “Hierarchical naming
systems pervade our whole language and thought.” We are thus retaining a tra-
ditional ranked hierarchical system of classification (in other words, groups of
organisms arranged in a hierarchy of categories—genus, family, etc.). While
there are evident problems with such a system, the “cornucopia of categories”
(Colless 1977) resulting from a cladistic approach does not seem to be an over-
all improvement in communication, and in fact seems less suited to human
mental abilities. In a clade-based system, a particular species is in dozens if not
hundreds of successively larger clades—how does one choose which of these
clades to use in real-world situations (e.g., floras, textbooks). This point seems
to be particularly crucial to writers of floras—unless all genera or even species
are to be arranged alphabetically, which would be extremely user unfriendly,
some higher level organization must be agreed upon. It seems clear that some
arbitrary convention to replace the convenience of currently used families
would have to be developed for use ina completely clade-based system—in other
words, someone would arbitrarily have to decide which of the innumerable
clades to recognize. Otherwise, there would be no practical way to group spe-
cies in floras (now grouped in families and then genera) and no groups to refer
consistently to when comparing different floras, etc. In fact, for practical rea-
sons many cladists still use traditional ranks such as family and genus (e.g.,
Judd et al. 2002), despite stating that “one logical step would be to eliminate
ranks altogether.” Currently, only three words (e.g., Fagaceae, Quercus alba) are
needed to communicate a great deal of information about a particular organ-
ism and these are used consistently by botanists. In our flora for example, gen-
era are being arranged alphabetically under alphabetically arranged families.
Does this mean that these families are viewed as somehow being equivalent?—
of course not. They are, however, effective means of conveying information. As
stressed by Stevens (2002), if we are going to be able to effectively communi-
cate, “..conventions will be needed. To paraphrase Linnaeus, without conven-
tion, all is chaos.” As Stevens (2002) so clearly pointed out, even early botanists
(e.g., Linnaeus, Bentham) were quite ned about effective communication.
658 BRIT.ORG/SIDA 20(2)
What would be the system of organization under a system of unranked clades
and how could consistency (and thus communication, information retrieval,
comparability, etc.) be assured in different floras, textbooks, etc.? Ultimately,
some arbitrary convention (not unlike our current family system) would seem
to be required. To be fair, it should be noted that the PhyloCode (2002) is not yet
complete and nomenclatural conventions will have to be worked out in the fu-
ture. Whatever system is ultimately settled upon, the conventions used should
take into consideration a variety of factors (e.g., accuracy of information con-
veyed, effectiveness of communication toa broad range of users, compatibility
with the organization of the human nervous system, etc.).
3) Furthermore, we are not rejecting paraphyletic groups (e.g., families). For ex-
ample, it now seems clear (as stated above) that the Lemnaceae (duckweed fam-
ily) arose from within the Araceae (arum family), which is thus paraphyletic.
It seems more reasonable to us to continue to recognize both easily distinguish-
able aa and Geamnigueu=)y state in discussions associated with both what
the y relationships between the twoare. Clearly the duckweeds have
undergone extraordinary morphological and genetic divergence in adapting
to an aquatic environment—so much so that the question of their ancestry was
only recently resolved. To lump the two families and have their genera mingled
in a taxonomic treatment accomplishes little and seems to result in a loss of
clarity and evolutionary information. As currently recognized, both families
have significant morphological coherence and thus recognition and predictive
value—together, they are a mixture of two very different types of morphology.
A similar example can be seen with the Cactaceae. That family has recently
been shown to have evolved from within the Portulacaceae (Hershkovitz &
Zimmer 1997; Applequist & Wallace 2001). Lumping the two families in a flo-
ristic treatment would only obscure the many differences between these useful
and easily recognizable groups. This reflects the view of evolutionary taxono-
mists, who while attempting to have a classification system based on evolu-
tionary relationships, also try to reflect the amount of evolutionary change
eaeoay by groups (Fig. 2). Evolutionary taxonomists argue that classifica-
tion is “.. more than just branching patterns of evolution” (Stuessy 1997).
Brummitt (2002) stressed Mayr’s (1995) observation that Darwin indicated evo-
lutionary classification depends on two factors, descent and modification, not
descent alone. To use a well known animal example, because birds are so differ-
ent from other vertebrates (e.g., fly, have feathers, etc.), they are treated as a dif-
ferent class of vertebrate even though they are known to have evolved from
within the paraphyletic class known as reptiles (Fig. 3). From the practical
standpoint, there are numerous other useful and evolutionary meaningful
paraphyletic groups. Examples include the Araceae, Capparaceae, Clusiaceae,
Moraceae, Portulacaceae, etc. Radically changing their circumscription (e.g,.,
FLORAS SYSTEMATICS? 659
A B C D E F
Fic. 2. Pt ic dist liag horizontal di I pecies indi pl ic diff Note that species
F hil ea | 1 ee ee ee * 1 to snecies F . ie bt «2 Pane 8 {nh Pi Rinne ot a]
y WHE THU Ot CIUocly PHYIUg y u , 4 v \ yy
1999).
adding Cactaceae to Portulacaceae, Podostemaceae to Clusiaceae, or Lemnaceae
to Araceae) results in confusion and a loss of information. This risk of confu-
sion is particularly problematic if the same name is variously used to include
quite different sets of species (e.g., Portulacaceae in the narrow sense or in the
broad sense including Cactaceae). The risk of a nomenclatural “train wreck”
(Stevens 2002) is a troubling possibility. Very different uses of the same names
“will be decidedly unsettling for society and perhaps damaging for our disci-
pline” (Stevens 2002). Unfortunately, taxonomists have long had the reputation
of changing names without regard for the implications, and great care needs to
be taken to avoid non-essential changes. Grant (2001b) stressed that splitting
up paraphyletic groups (e.g., genera such as Gilia or Linanthus) obscures rela-
tionships and multiplies generic names where one would be sufficient. Does
the splitting up of recognizable and well known genera really best serve the
broad constituency of users (keeping in mind that professional taxonomists
are a small fraction of the users of scientific names)? Changes in classification
at the generic level are particularly significant because of the resulting changes
in scientific names.
Further, Brummitt (1997, 2002) has argued that paraphyletic groups are
inevitable. He goes on to say that any genus “must have originated from a spe-
cies in another genus, which is thus paraphyletic.” The same logic applies at
other levels in the hierarchy, including the species level. Species, unlike other
categories in our hierarchical system of classification, have some objective bio-
logical beat (e & Rieseberg © Burke oy However, if a small subgroup of a
hto become a distinct entity, the origi-
aval species is thus tendered paraphyletic even though it may still be a repro-
ductively isolated or otherwise distinct group. Species can be excluded by defi-
660 BRIT.ORG/SIDA 20(2)
TURTLES
LIZARDS
SNAKES
CROCODILES
BIRDS
Fic. 3. Dendrogram showing relationships of some vertebrate groups. Note the paraphyletic nature of the “Reptilia”
(from Diggs et al. 1999).
nition as not being paraphyletic (Sanders & Judd 2000) as done by some cladists,
but in reality, by standard use of the term, they are often paraphyletic (see
Mishler 1999, 2000). As Brummitt (2002) noted, the abandonment of species
“seems to me to be a necessary logical extension to abandoning all other ranks”
(Brummitt 2002). Further, Rieseberg and Brouillet 1994) argued that based on
the modes of speciation known to occur in plants, paraphyletic species are com-
mon. In the words of Sosef (1997), “When a single diaspore [e.g., seed] acciden-
tally reaches an isolated habitat and its offspring gradually changes (a linear
process) and produces a new species, nothing ‘happens’ to the parent species.”
The new species thus renders the unchanged previous species paraphyletic.
Ultimately, paraphyly thus seems unavoidable. An excellent example of this
problem can be seen in the California tarweeds (genus Raillardiopsis) and the
Hawaiian silverswords (genera Argyroxiphium, Dubautia,and Wilkesia) in the
Asteraceae. It now appears (Baldwin et al. 1991) that Raillardiopsis (which had
previously been considered a “phenotypically conservative genus of two nearly
identical species”) was the “ancestral genetic source” for the cee ane ex
tremely morphologically divergent Hawaiian silversword alliance. R
—
L
TiApac SYSTEMATICS? 661
is thus clearly paraphyletic, even though its two species are “nearly identical.”
When taken to its logical extreme, the failure to recognize paraphyletic groups
would require the rejection of many useful groups (e.g. tarweeds). In animal tax-
onomy, this means the rejection not just of the group Reptilia (which gave rise
to birds), but also Amphibia (which gave rise to reptiles), and Pices (which gave
rise to Amphibia). Anyone understanding the basics of evolution realizes that
fishes gave rise to amphibians. Does that make the class Pices any less useful a
concept? Likewise, the capers (Capparaceae) gave rise to the mustards
(Brassicaceae)—both families are still useful and meaningful concepts that can
help us understand evolution and organize our thinking. Cronquist (1987)
stressed the need for paraphyletic groups, indicating that both evolutionary
relationships and the amount of evolutionary divergence among taxa is impor-
tant. He said that “the reasons for this belief are rooted in the historical and
continuing function of taxonomy asa general-purpose system of classification
that can be used by all who are concerned with similarities and differences
among organisms’—in other words, diverse users of botanical information.
While we believe it is critically important for paraphyletic groups to be clearly
distinguished from monophyletic groups, the usefulness of paraphyletic taxa
(e.g., effective communication, recognition of divergence, morphological simi-
larities, etc.) seems to be an important consideration.
4) Additionally, there are many cases where it is not yet clear what should be
done cladistically. Thus in these cases we are retaining traditional usage until
more information is available. For example, it is very likely that such families
as the Lamiaceae (mint family), Verbenaceae (vervain family), and
Scrophulariaceae (figwort family), as traditionally conceived, will have to be
changed substantially. However, if we had to finalize treatments of these fami-
lies today for our flora (which fortunately does not have to happen since these
dicot groups will be treated in Vol. 2 of the Illustrated Flora of East Texas to be
published in 2008), we would probably follow the traditional circumscriptions
and accompany them with substantial explanations. The reason is that adequate
research has not yet been done on these groups to provide answers that are de-
finitive enough to warrant major changes in classification and nomenclature.
In the words of Berry (2002), “it will be some time before our sampling of or-
ganisms at the molecular level will be good enough that we can get an adequate
idea of relationships across the entire span of biological diversity.” Further, it
does not seem desirable that classification (and nomenclature) should change
with every new cladistic discovery—do we really want nomenclature that
“depend[s] rigidly on the particular cladogram favoured at the moment” (Ben-
ton 2000)? Indeed, Sanders and Judd (2000) discuss the criteria for accepting
revised classifications. Before making major nomenclatural changes, there
should be substantial taxonomic evidence, to avoid more of the numerous ex-
662 BRIT.ORG/SIDA 20(2)
amples where initial cladistic hypotheses have proven to be wrong. For example,
in the past, some authorities have suggested major changes in family circum-
scription based on preliminary information (e.g., lumping the Apiaceae (car-
rot) and Araliaceae (aralia) families—Judd et al. 1994; Zomlefer 1994), only to
have more detailed work (Plunkett et al. 1996 [1997]) clarify the situation and
indicate that the families should be maintained in nearly their traditional cir-
cumscriptions. According to Plunkett et al. (1996 [1997]), the approach taken by
Judd et al. (1994) “hides rather than resolves the difficulties in Apiales.” Like-
wise, Downie et al. (2001) did not lump the Araliaceae. Another example is the
genus Trillium—it has often been treated in the Liliaceae in the broad sense
(e.g., Correll &@ Johnston 1970) or based on early phylogenetic analyses in the
Trilliaceae (e.g., Zomlefer 1996; Tamura 1998d; Judd et al. 1999). However, more
recent phylogenetic analyses (e.g., Rudall et al. 2000b) indicate that it belongs
in the Melanthiaceae. There are also instances where the best and most recent
evidence conflicts. A number of molecular studies have suggested that
Burmanniaceae (previously considered to be related to Orchidaceae) is in the
Dioscoreales and thus more closely related to such families as Dioscoreaceae
and Nartheciaceae than to Orchidaceae (which is in order Asparagales) (Chase
et al. 1995b; Caddick et al. 2000, 2002; Chase et al. 2000). In contrast, other re-
cent molecular research including more genera than previously sampled, indi-
cates that the family (minus the superficially similar but unrelated genus
Corsia) plus Thismiaceae is ina relatively isolated position “not closely aligned
with either the Dioscoreales or the Orchidaceae” (Neyland 2002). Writers of
floras are thus faced with having to judge which cladistic studies to accept and
when there is enough evidence to use the new discoveries in floristic treatments.
At the same time, writers of floras must avoid the instability and confusion
that would result from changing classification and nomenclature with every
new study Puen eee Furthermore, recent tests (Grant 2001a) of the accuracy
of cladograms when compared with known phylog
experimental plant groups with known pedigrees), raises questions about bas-
ing classification and nomenclature solely on cladistic methodology. Grant
(2001a) demonstrated that the “cladograms of the four plant groups [tested] all
differ in significant details from the known pedigrees.” Particularly important
is his following point: “It is also recognized by all evolutionary systematists and
most cladists that reconstructed phylogenies are unverified hypotheses. Some
cladists, however, seem to regard their cladograms as real phylogenies.” He (Grant
2001b) also indicated that “Molecular cladograms are very good indicators, but
we should not lose sight of the fact that the groupings they indicate are mo-
lecular clades, not taxa.” A recent paper by Rydin et al. (2002) on Gnetales is
particularly telling in this regard. Depending on which molecular analysis was
used, the phyogenetic position of Gnetales differed significantly. According to
Rydin et al. (2002), “It is becoming increasingly clear that the understanding of
omesticated and
FLORAS AXONOMY SYSTEMATICS? 663
molecular evolution and its impact on phylogenetic studies is poor. Nucleotide
data alone might not be able to solve phylogeny and evolution of this ancient,
once rapidly evolving group, and attempts to do so should include a compre-
hensive taxon sampling and several genes. Molecular data can definitely be
misleading, and by ignoring that, science will not progress.” From the stand-
point of a floristician, while recognizing the obviously valuable contributions
made by molecular systematists and cladists, careful thought must be given to
avoid accepting major, sometimes disruptive (and occasionally incorrect) clas-
sification and nomenclature changes prematurely.
5) Finally, we are not accepting some aspects of phylogenetic classification (par-
ticularly strict monophyly—sometimes and perhaps better referred to as
holophyly) since there are serious theoretical problems that make it at least par-
tially incompatible with the reality of the natural world (however, see
Freudenstein 1998 for an opposing viewpoint). Brummitt (2002, pers. comm.)
has stressed the importance of these theoretical considerations, and in terms of
the implications for floristics, these are perhaps as important as or even more
important than the purely practical considerations. One of the most serious
problems (referred to asa “fatal one” by Stuessy 1997) is that cladistics uses only
branching information in phylogeny. In contrast, evolution is a complex pro-
cess including such phenomena as asexual reproduction, progenitor-derivative
species pairs, lateral gene transfer, polyploidy, and reticulate evolution (the lat-
ter resulting from hybridization between species and subsequent speciation in
the offspring) (Rieseberg & Brouillet 1994; Sosef 1997; Stuessy 1997). These com-
plexities cannot be accommodated in a classification system requiring only
strict monophyletic groups and dichotomous branching. In the words of Stuessy
(1997), “.. simple dichotomous branching diagrams cannot do justice to the real
world of higher plant phylogeny.” An excellent example can be seen in the re-
ticulate evolution of the fern genus Asplenium as discussed in the classic paper
by Wagner (1954). In this case, the hybrids between two parental species be-
come reproductively isolated (and thus constitute a separate new species). An-
other example can be seen in the Triticeae (Elymus, Hordeum, Secale, Triticum,
and their relatives; Poaceae). This tribe has an extremely complex evolutionary
history involving hybridization, polyploidy, and reticulate evolution (Bark worth
2000; Mason-Gamer & Kellogg 2000). Both of these examples emphasize that
some evolutionary relationships are simply impossible to reflect accurately in
a system requiring strict monophyly and only dichotomous branching. As
Brummitt (1997) indicated, “No matter how much we may long for all our taxa
to be monophyletic, if we are considering the whole evolutionary process, it isa
logical impossibility.” Another way of stating the problem is that simple branch-
ing patterns are unable to reveal all significant dimensions of phylogeny
(Stuessy 1997).
A second theoretical problem with strict monophyly is that “only the tips
664 BRIT.ORG/SIDA 20(2)
of evolutionary branches can be classified” (Meacham & Duncan 1987) in such
a system, and “Species at the interior nodes of the tree must remain unclassi-
fied.” Not including ancestral species, some of which may well have survived to
the present, seems untenable. However, including them leads cladists insisting
on strict monophyly (holophyly) down the path of the “telescoping” or “snow-
ball” effect (also known as the “taxonomic black hole”) where more and more
organisms have to be included in a futile attempt to reach the mythical (and
impossible to reach) strictly monophyletic group (Sosef 1997; Brummitt 2002;
but see Stevens 1998 and Sanders and Judd 2000 for a different viewpoint). Ul-
timately, the whole Tree of Life (2002) would have to be included in one giant
monophyletic group. In the words of Brummitt (2002), “If we are classifying all
the products of evolution, ie. the wh lutionary tree of life, every taxon we
recognise must make another taxon paraphyletic. That is a simple logical fact.
It is obvious to most people that if you cannot have aes haa taxa, you can-
not have a classification showing anything beyond one original species, genus,
family, etc.” He points out that if ranks are to be used (which seems essential from
the standpoint of practicality), paraphyly is unavoidable (Webster 2002 referred
to this as “Brummitt’s Paradox”). Thus, there is a “fundamental incompatibility
between Linnaean classification and a system of monophyletic taxa, or clades”
(Brummitt 2002). In other words, including all extinct species would mean that
no monophyletic groups (or only one huge one) could be recognized. From this
standpoint, “extinction [and our lack of knowledge about extinct species] is the
saving grace of phylogenetic systematics” (T. Barkley, pers. comm.). Importantly,
“Brummitt’s Paradox” means that simply converting cladograms into Linnaean
nomenclature is impossible. Ultimately, this means that in order to have strict
monophyly, phylogenetic systematists must develop arankless system such as
the PhyloCode. While some modern phylogenetic systematists continue to
recognize families for practical/pedagogical reasons (e.g,, Judd et al. 2002), if carried
to its logical conclusions, cladistics would mean that these families (particu-
larly if extinct taxa are considered) would be telescoped into ever larger groups.
A third related theoretical problem demonstrates that a strictly monophyl-
etic system using Linnaean ranks is incompatible with evolution (Brummitt,
pers. comm.). As Brummitt (2002) indicated, *..1 think we all understand that
evolution has continually been throwing up greater and greater diversification
of plants and animals, and yet the cladistic idea of classification requires that
all successive descendant groups have to have lower and lower taxonomic rank.
Something which has evolved from within one genus must have a rank lower
than genus. Is this view of taxonomy really a sensible idea? How can we apply
lower and lower ranks when evolution is producing wider and wider variation?
Inaclade-based classification you can go on extending your clades as evolution
progresses ad infinitum, but if youare using a traditional taxon-based classifi-
cation you can't keep on giving them lower ranks.” Without paraphyly, the use
FLORAS SYSTEMATICS? 665
of ranks ina monophyletic system therefore is simply not workable (hence the
advocacy of a rankless Phylocode by some cladists). This problem seems to
clearly indicate the logical impossibility of a rank-based, strictly monophyl-
etic system.
A fourth theoretical problem is that our current methodology of obtaining
phylogenetic trees is based on a series of assumptions and indirect evidence
(e.g., character polarity, choice of outgroups, etc.) (Stuessy 1997), as well as math-
ematical algorithms (e.g, parsimony) that result in trees that “are almost al-
ways inappropriate as phylogenetic hypotheses in any but the most general
sense” (Zander 1998). Zander (1999) further indicated that, “Selecting one phy-
logenetic hypothesis of several or many reasonable alternatives as ‘best’ and
presenting it as a reconstruction cannot provide a probabilistic or dependable
basis for action.” Is the pursuit of the shortest tree the ultimate goal of system-
atics? The answer seems clearly to be no, since there are well-documented non-
parsimonious pathways of evolution (Stussey 1997). Thus, totally tying classi-
fication and nomenclature to sucha system seems problematic. A final concern
that follows from this is that the current classification and nomenclature sys-
tem is independent of a particular approach. As stated by Jorgensen (2000), “A
problem inherent with the system proposed by Cantino & de Queiroz (2000)
[PhyloCode] is that their nomenclature depends upon a specific way of taxo-
nomic thinking, i.e. nomenclature is ruled by the taxonomic system. This prob-
ably reflects that they are as convinced of their taxonomic system as Linnaeus
was of his, but, please, at least consider the possibility that new taxonomies
may evolve. Should we then change the nomenclatural system each time?”
FURTHER IDEAS AND DISCUSSION
The following ideas and quotes seem to provide substantial insight into the con-
troversy. Because of both philosophical and practical implementation problems,
Brummitt (1997) pointed out that while the controversy should be debated, it
seems unlikely that “Linnaean classification” will soon be abandoned. Brummitt
(1997) suggested that both a “Linnaean classification” system and a clade-based
phylogeny are desirable because they have different functions. He argued that
both be allowed to exist side by side and that the nomenclature of the two should
be easily recognized as different (Brummitt 1997). In summarizing his ideas he
stated, “.. we should not follow traditional practices just because they are tradi-
tional, but neither should we adopt new ideas just because they are new. We
need to understand the possibilities and appreciate the different objectives and
functions of the different options. In the meantime, it seems to me and to many
others that the compromise of maintaining Linnaean classification but trying
to eliminate paraphyletic taxa is nonsensical and should be abandoned before
any more damage is done to existing classifications and nomenclature.” Lidén
et al. (1997) indicated, “If applied consistently, Phyllis [= Lidén et al. term for
666 BRIT.ORG/SIDA 20(2
nomenclatural application of cladistics] will cause confusion and loss of infor-
mation content and mnemonic devices, without any substantial scientific or
practical advantage. ...any attempts to make Phyllis formal would be disastrous.
We can find no conclusive, valid arguments against keeping the body of our
current system intact.” Sosef (1997) stressed the same idea saying, “The quest
for monophyletic taxa and the epeang x of former paraphyletic ones should halt
immediately, as they unnecessarily deteriorate classifications which will often
prove to be valid.”
An interesting point was also made by Stuessy (1997) when he said, “... in
this urgent climate of seeking to inventory the world’s biota (Anonymous 1994),
and requesting funds from the rest of society to do so, it would be highly coun-
terproductive to simultaneously recommend whole-scale change of names of
organisms for any reason.” In the words of Paul Ehrlich (2002), who was stress-
ing the need for taxonomists to focus on conservation activities, “Others spend
their time trying to replace the functional Linnaean system for general com-
munication about organisms with one based on estimates SOL times of phyloge-
netic divergence; a sillier enterprise is hard to imagine...
While recognizing it is not perfect, Wheeler (2001, and quoted in Forey
2002) noted that Linnaean nomenclature “is stable enough to say what we know,
flexible enough toaccommodate what we learn; independent of specific theory,
yet reflective of known empirical data; compatible with phylogenetic theory,
but not a slave to it; particular enough for precise communication, general
enough to reflect refuted hypotheses.”
While strongly supporting a cladistic system, Welzen (1998) also noted that
a compromise between the two types of classification is impossible. He also
understood that because of practical reasons it is impossible to abandon Lin-
naean classification “.. because too few cladograms are available to replace the
existing system with a complete phylogenetic classification. Moreover, quite a
few cladograms will not be that trustworthy due to the many homoplasies |re-
sult of convergent evolution] that have evolved; they will therefore, provide an
unstable classification at best.” Welzen (1997) went on to say, “I think, therefore,
that the best solution is to choose the second option that Brummitt (1997) pro-
vides in his paper, namely, ‘retaining Linnaean classification, with paraphyletic
taxa, but developing alongside it an independent clade-based dichotomous sys-
tem with its own separate nomenclature. ” Recently, Brummitt (2002) indicated
that, “If people insist on monophyly, the clade-based PhyloCode will provide a
logical solution. If they want to use the traditional ranks, the answer is very
simple: recognise paraphyletic taxa.” Brummitt (1997) made what seems to bea
very reasonable suggestion: “Our task is to produce an optimally practical clas-
sification, and indicate which genera have evolved from which other genera,
which families from which other families, and so on.”
Indeed, recently it seems to have become clearer that two separate systems
FLORAS SYSTEMATICS? 667
will be necessary (Cantino 2000; Brummitt 2002). Years ago, Woodger (1952,
and quoted by Brummitt 2002) concluded that, “The taxonomic system and
the evolutionary phylogenetic scheme are quite different things doing quite
different jobs, and only confusion will arise from identifying or mixing them.”
Similarly, it has been argued more recently that to attempt to apply cladistic
rules (e.g., elimination of all paraphyletic groups) on the traditional Linnaean
system isa logical impossibility (Brummitt 2002) or at minimum highly detri-
mental (see discussion above). According to Brummitt (2002), “I believe there is
no middle way which will combine universal monophyly with formal Linnaean
ranks, and this is now increasingly being realized.” However, while it is becom-
ing more widely recognized that the two systems are incompatible (but see
Stevens 2002), there is clearly value in having a strictly phylogenetic system of
classification. It would thus seem that the most reasonable course for the near
term would be toallow the concurrent existence of two separate systems—Lin-
naean and phylogenetic. The Linnaean system would retain hierarchical ranks,
binomial nomenclature, and paraphyletic taxa, while the phylogenetic system
(e.g., PhyloCode) would recognize monophyletic clades (but have neither ranks,
binomial nomenclature, nor paraphyletic taxa, and possibly not even species).
Because of practical considerations, the Linnaean system will probably con-
tinue to be the “dominant” system used (nearly exclusively) to communicate
information about plants by scientists outside of geen: (e.g., ecologists,
horticulturalists, conservationists, etc.) and by the general Bune The De
genetic system will make contributions among
ing to further understand the evolutionary history of life on earth.
SUMMARY
Where does all this leave the writers of floras? We would argue that a modified
traditional system (binomial nomenclature, ranked hierarchies—L-code) be
used in floras (allowing paraphyletic groups but eliminating all polyphyletic
groups despite some nomenclatural instability), and that these floras (and this
is a critical point) should also incorporate newly discovered information on
phylogenetic relationships. In order to facilitate as complete an understanding
of plant evolution as possible, this new information should be briefly discussed /
summarized in appropriate family and generic treatments even if too prelimi-
nary, tentative, or inappropriate for nomenclatural change (e.g., that would re-
sult in the loss of meaningful paraphyletic taxa).
Insummary, in the Illustrated Flora of East Texa, we are thus attempting to
reflect some of the many contributions and insights from “phylogenetic sys-
tematics,” while retaining the practical benefits of an “evolutionary taxonomy”
framework. As indicated above, no system will be a totally accurate represen-
tation of the complexity of the evolutionary history of life on earth. We believe,
however, that by reflecting both evolutionary relationships and the amount of
668 BRIT.ORG/SIDA 20(2)
evolutionary change, while maintaining a flexible approach punctuated with
practicality, that a classification and nomenclature system useful to a broad
audience, including non-taxonomists, can be achieved. We agree fully with
Berry (2002) who said, “there are many users of scientific names—myself in-
cluded—who are interested in both floristic inventories and evolutionary rela-
tionships, and nomenclatural stability as well.” Ultimately, writers of floras need
to present information accurately (ie. reflect evolution) and in ways that can
be used—that allow effective communication and identification, promote in-
formation retrieval, and are useful comparatively (e.g., in conservation assess-
ments, to evaluate levels of endemism, to determine levels of introduced exot-
ics, to form the basis of biogeographical studies, to do ecological surveys). All of
these uses depend on having a unit (species) that can be meaningfully com-
pared and classification and nomenclatural systems that allows effective com-
munication. Taxonomic botanists are thus faced with the challenge of working
toward systems that make such communication and comparisons possible
(Berry 2002). Writers of floras, in particular, as the translators of botanical in-
formation to a wider audience, are uniquely faced with a difficult task—to ap-
ply the evolving concepts of modern systematic botany to floristics in a way
that allows modern floras to be both accurate and useful. It is hoped that this
article will stimulate discussion among those involved in or interested in the
writing of floras.
—
ACKNOWLEDGMENTS
We would like to thank Ted Barkley for helpful discussions and comments. We
also thank Richard Brummitt, Verne Grant, Roger Sanders, and Rudolf Schmid
for their extremely constructive reviews which improved this paper.
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—
ience.ora/
THE VASCULAR FLORA OF THE LAGUNA DE ALEGRIA,
A NATIONAL MONUMENT IN THE CRATER OF
TECAPA VOLCANO, EL SALVADOR
Denise Williams and Rudys W. Herrera José L. Linares
118 Fox Crest Drive pela Pay) G Standley
Lynchburg, VA 24502, USA. Escuela Panamericana
Pat eos, PO. Box 93
Tegucigalpa, HONDURAS
ABSTRACT
Extreme deforestation and overuse of resources have devastated El Salvador’s natural environment
while continued population pressure underscores the need for the few remaining natural areas to be
studied and preserved. The 150-ha Monumento Nacional Laguna de Alegria is located inside the cra-
ter of Tecapa Volcano in southeastern El Salvador and includes both primary and disturbed sub-
BOP cits and rocky crater walls, and a sulfuric lake in the basin. The flora of the Laguna
de Alegria wa yed from December 1998 to November 1999 and results of this survey were com-
bined ae reports aoe ae collectors and published reports to compile a list of 447 specific and 2
hybrid taxa representing 106 families. The taxa are cataloged in an annotated list that includes col-
lection number and herbarium of deposit. See species are designated. Trigonospermum annum
McVaugh & Lask. is a new report for El Salvador.
RESUMEN
La deforestacion extrema y la sol | 5n de rec i | lol | le El Salva-
dor, mientras la incesante presion Hemogniten subraya la necesidad de estudiar y preservar ce
restantes areas naturales. El Monumento Nacional Laguna de Alegria mide 150 hectareas y es
Jo]
ater del Volcan Tecapa en el sur-este de El Salvador. El Bae incluye aa
su eo primario y ee, paredes volcanicas rocosas y empinadas, y una laguna sulftrica
en la cuenca. Se identificé la flora de la Laguna de Alegria desde diciembre de 1998 hasta noviembre
de 1999 y se combinaron los resultados del estudio con los reportes de otros colectores y reportes
publicados para cereus un oe de 447 taxa especificos y 2 hibridos. Estas representan 106
familias. Los taxa fueron los en un listado anotado que incluye el ntmero de la coleccion y
E
WY
del herbario, y se reer las especies e>
Mc Vaugh & Lask. en El Salvador.
c primer reporte de Trigonospermum dnnum
INTRODUCTION
The Republic of El Salvador, witha land area of only 21,000 km? anda popula-
tion of more than 6 million, is the smallest and most densely crowded of the
Central American countries. Extreme population pressure has devastated El
Salvador’s environment, causing the loss of 97% of the country’s original forest
(Benitez 1996 unpublished, cited from Berendsohn 1991) and forcing farmers to
convert ever more forest and marginal land to crop production. Recent estimates
of forest cover indicate that only 130,000 ha, or 6.1% of the national territory, is
SIDA 20(2): 675 — 690. 2002
676 BRIT.ORG/SIDA 20(2)
now covered by natural forest (DIPRAT 1997), most of which is composed of
small, relictual parcels.
The government of El Salvador recognizes the urgency of protecting the
few remaining forests and forest fragments in the country and has identified
125 natural areas to form the “Sistema de Areas Naturales Protegidas” (http:/
www.marn.gobsv/sia/anp.htm). These natural areas are all relatively small,
but are of supreme importance as the country’s last refuges for wildlife and gene
pools that could provide the primary materials for reforestation and regenera-
tion of other areas. However, there is no infrastructure to administer most of
these areas; only three areas have legal protection by legislative or executive
decree; and nearly all of them are still exploited for firewood, grazing lands,
and for squatter’s homesteads. Moreover, there is an almost complete lack of
basic biological information about these areas and few in-country resources to
gather this information. Apart from El Imposible National Park, none of El
Salvador'’s protected areas has previously had a complete floral survey. While
biologists from developed countries tend to disregard El Salvador in favor of
studying other countries with better preserved and more extensive wild areas,
the very precarious ecological situation in El Salvador should make it a high
priority for study so that the natural resources still intact can be managed and
preserved in the most effective manner possible.
THE LAGUNA DE ALEGRIA
The Laguna de Alegria isa tiny but spectacular natural area in the crater of the
extinct Tecapa Volcano in the department of Usulutan, and is the only national
monument proposed within the Salvadoran system of protected areas. Tecapa
is located at 1329'40'N latitude and 8829'43" W longitude (Fig. 1) and rises toa
height of 1599 m. It is the second largest peak in the Chinameca chain of volca-
noes that dominates much of the southeastern corner of El Salvador and forms
part of the Central American chain that extends from southwest Mexico,
through Guatemala, El Salvador, Nicaragua, Costa Rica and Panama.
The interior of the Tecapa crater is 150 ha and is comprised of an emerald-
green, sulfuric lake in the caldera surrounded by natural forest. The ecological
value of this small forest is immense in this heavily deforested country and the
natural beauty of the lake and sheer crater walls make for a striking setting.
There are no historical records of volcanic activity, though there are emis-
sions of steam mixed with volcanic gases on the southwest side of the lake. The
gaseous emissions, which have a high concentration of sulfur compounds, have
contributed to the low pH of soils in much of the basin of the caldera and to the
extreme acidity of the lake. The sulfur compounds give the lake a characteris-
tic emerald green color and have bleached the beach and lake substrate white.
It has been suggested that only algae and diatoms are able to live permanently
in the extreme conditions of the water (SACDEL 1997), but exoskeletons of drag-
WILLIAMS ET AL., FLORA OF THE LAGUNA DE ALEGRIA, EL SALVADOR
677
al
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Fic. 1. Map of C tral A howing the approxi | i f the Lag de Alegria, El Salvador.
onfly nymphs have been observed (Williams, pers. obs.) and a species of
Eleocharis (Cyperaceae) has been reported growing in the margins of
(Fassett 1952).
Holdridge (1975) classifies the area in and around the summit of Tecapa
Volcano as very humid subtropical forest and much of the forest is primary
growth, especially those parts located on the steep inner slopes. Local residents
report that some of the valuable timber species have been removed from the
more accessible areas of the forest in the basin, and three areas of the forest were
cleared and planted in coffee: one in a small area of the basin, now abandoned,
and two others, on the northeast and northwest slopes, both still in cultivation.
The inner walls of three sides of the crater are a nearly vertical 300 m tall mak-
ing them inaccessible without specialized climbing equipment and only there
has the forest been left completely undisturbed by humans.
The outer slopes of the volcano are not officially included in the protected
area, and what parts have not been planted in coffee have been deforested for
firewood. The coffee grown in plantations on the outer slopes is mixed with
the lake
678 BRIT.ORG/SIDA 20(2)
shade trees, thereby offering some protection for the buffer zone. Unfortunately,
the coffee plantations do not have the density of vegetation in natural forest
and some recently replanted areas have come so close to the edge of the steep
inner slope that severe erosion and landslides have resulted from the loss of
a
cover (Williams, pers. obs.).
The Laguna de Alegria is currently being managed by the Asociacion de
Desarrollo Comunal Alberto Masferrer (ADESCAM), a local organization cre-
ated in 1995 and given charge of the area for development for ecotourism and
recreation by the municipal government. The area has become popular asa tour-
ist destination due to promotion and development by ADESCAM ona national
scale. However, the balance between conservation and tourism is often diffi-
cult, especially in the case of the Laguna de Alegria because the amphitheater-
like setting of the interior of the crater amplifies the presence of humans in the
natural area. This study was undertaken to provide ADESCAM with part of
the basic biological information necessary to write a management plan for sus-
tainable use of the area.
METHODS
The majority of the species listed in this survey are documented by herbarium
vouchers we collected in the Laguna de Alegria between 9 Dec 1998 and 20 Nov
1999. Most of the vouchers were taken from the basin of the crater, e.g., the
beaches of the lake, the edges of the forest along the sides of the road, and those
parts of the forest accessible from the basin. Some collections were made from
the top of the rim, but most of the vegetation of the cliff areas of the crater is
not accounted for in this survey because we did not attempt to scale the steep
slopes.
The samples were pressed and dried and sent to the herbarium at the Jardin
Botanico La Laguna (LAGU) in San Salvador where we later identified them.
Major references used to identify specimens include Davidse et al. (1995) and
Standley and Steyermark (1960).
At LAGU we encountered specimens collected in the 1960s and 1970s in
the Laguna de Alegria by the German botanist Hans Winkler. We revised those
reports and added them to ours.
We also added a list of species collected by Alex Monro that are vouchered
at The Natural History Museum in London (BM). Unfortunately, we were not
able to visit BM and not all of Monro’s collections had been identified to species
at the time of this report. More information may be available when he com-
pletes his work.
In addition to reporting herbarium collections, we included reports of
plants from the Laguna de Alegria in two previous publications: Fritz Hamer’s
book Las Orquideas de El Salvador (1974) and a descriptive survey of tree spe-
cies of the Laguna de Alegria made by the Salvadoran dendrologist Maria Luisa
WILLIAMS ET AL., FLORA OF THE LAGUNA DE ALEGRIA, EL SALVADOR 679
Reyna de Aguilar (SACDEL 1997). Unfortunately, Hamer did not publish the
sources of his reports and Reyna de Aguilar took only four voucher specimens
during her survey, so it is not possible to verify their identifications
Edy Montalvo (pers. com.), former curator of the herbarium of the Univer-
sidad de El Salvador (ITIC), made several collecting trips to the Laguna de
Alegria in the 1970s. We had hoped to peruse ITIC’s collections and add
Montalvo’s reports and any other specimens from the Laguna de Alegria that
might be housed there, but due to a shortage of staffing, we were never able to
gain permission to ITIC. It would be valuable to revise ITIC’s collections and
add them to this list.
After the identification phase was completed, we returned to the Laguna
de Alegria and noted another 17 species that had not been collected in the pre-
vious year nor noted by other authors. We also verified an older collection of
100 woody species made in 1998 for ADESCAM.
It should be noted that we also collected non-vascular plants, i.e., mosses
and lichens, during the 1999 survey. Those vouchers were deposited in LAGU
as well, but since materials and expertise are not yet available to identify them,
they are not included in this list.
RESULTS AND DISCUSSION
We compiled a list of 821 voucher specimens and visual reports to identify a
total of 447 species and 2 hybrids of vascular plants from 319 genera and 106
families from the Laguna de Alegria (Table 1). Of the two hybrids reported in
our list, one isa natural hybrid orchid, the other, the artificial hybrid that cre-
ates the many varieties of bananas and plantains. Neither of the parent species
for either of these two hybrids are known from the Laguna de Alegria.
The most important families in terms of numbers of species present are
Asteraceae (54 species), Fabaceae (41 species), Poaceae (34 species), and
Orchidaceae (29 species and | hybrid). The families Asteraceae and the Poaceae
both tend to be composed of pioneer and secondary vegetation specialist spe-
cies and most of the species from these families found in the Laguna de Alegria
were found along the roadsides, in landslide zones, and on the beaches of the
laguna. Many of the species of Poaceae collected in the Laguna de Alegria are
naturalized exotics. The high numbers of species in these families indicates the
large amount of disturbance in many parts of the protected area.
The orchid species found in the Laguna de Alegria, on the other hand, tend
to be primary forest species, and none are exotic. All of the orchids we found
were in the forest or undisturbed areas of the cliffs with the exception of just
two terrestrial species found in open areas. The large number of species of or-
chids, therefore, supports the notion that the less accessible parts of the forest
have been intact and undisturbed for a long time.
The hybrid orchid, Mormodes x salvadorensis Hamer & Garay, is a natural
680 BRIT.ORG/SIDA 20(2)
Taste 1. Statistical summary by taxa of the flora of the Laguna de Alegria.
Families Genera Species Hybrids
Ferns and fern allies 13 22 34 0
Gymnosperms 3 3 4 0
Monocots 12 69 99 2
Di 78 225 310 0
Total 106 319 447 2
hybrid first reported by Hamer (1974) from two locations in El Salvador. One
of these locations was the Laguna de Alegria, but M. x salvadorensis was not
found in our 1999 survey. It is possible that it has become extirpated in the 25
intervening years. However, neither of the two parent species was found either
by Hamer or by us, though they most likely exist or once existed within the
bounds of the park. This fact suggests that there probably are more orchid spe-
cies present which have not been recorded, a fact which is very likely consider-
ing the difficultly of encountering epiphytic orchids in the dense forest and
the steep cliffs of the crater,
We found 36 exotic species in the Laguna de Alegria, making up just 8.2
of the total number of species. In comparison with a national figure of 28%, the
percentage of exotic taxa in the Laguna de Alegria is very small,a fact that serves
as another indicator that the area is still in a very natural state. Many of the
exotics found are plants cultivated in the coffee plantations and around the
margins of the lake, but nine of the species have been introduced intentionally
by ADESCAM for ornamental purposes in recreational areas or for “reforesta-
tion” in areas where natural growth would have occurred. Apparently only eight
species have invaded under their own power: seven grasses and “higuero” or “cas-
tor bean” (Ricinus communis L. Euphorbiaceae ). These eight plants are all ag-
gressive colonizers and have taken advantage of man-made disturbances such
as roads and trails and other clearings to spread to new locations. One species,
“frijol de palo” or “pigeon pea” (Cajanus cajan (L.) Millsp. Fabaceae), is of uncer-
tain invasion status, being a plant that is widely cultivated for its edible beans,
but also widely naturalized and an aggressive colonizer.
The distribution status of the majority of El Salvador’s flora is poorly
known, and there is no official list of threatened or endangered species for the
country. At least one species collected in this survey, Cibotium regale Verschatf.
& Lem. Dicksoniaceae, isendangered throughout its habitat (Davidse et al. 1995).
The only individual of Bixa orellana L. Bixaceae in the Laguna de Alegria was
destroyed by a landslide in 1998. However, B. orellana is common throughout
most of El Salvador.
Despite the poor knowledge of El Salvador’s flora, we found relatively few
O/
/0
WILLIAMS ET AL., FLORA OF THE LAGUNA DE ALEGRIA, EL SALVADOR 681
new reports in our survey. [rigonospet annum McVaugh & Lask. Asteraceae
is the only verified new report for El Salvador. Peperomia cf. liebmanii C. DC.
Piperaceae and Tillandsia cf. kammi Rauh Bromeliaceae, if confirmed as such,
will also be new reports.
The total species count is relatively large for such a small area, especially
when one considers that more extensive studies would most likely provide new
reports and that the cliffs of the crater have never been surveyed. The total num-
ber of species registered for El Salvador is 4670 (Reyna de Aguilar 1995), mean-
ing that the Laguna de Alegria, with an areal extent of less than 0.005% of the
land area of the country, harbors 9.4% of the country’s plant species. It is evi-
dent that the continued preservation of the tiny Laguna de Alegria is crucial to
maintaining El Salvador’s plant diversity.
APPENDIX
Annotated list of the vascular species of the Laguna de Alegria
Collectors/authors of reports: DW: Denise Williams and Rudys Herrera, AM:
Alex Monro, FH: Fritz Hamer, JBL/Author: specimen housed at the Jardin
Botanico La Laguna, MLRA: Maria Luisa Reyna de Aguilar, ADESCAM: collec-
tion of the Asociacion de Desarrollo Comunal Alberto Masferrer.
* refers to introduced exotic species, ** refers to naturalized exotic species,
T extinct,
Note.—At the end of each entry with multiple collections follows a list of
herbaria where a duplicate of at least one of the collections was sent. This does
not mean toimply that each herbarium hasa duplicate of every collection listed
for that entry. LAGU was the main repository for the specimens we collected
and it retained one duplicate of all our collections.
DIVISION LYCOPODIOPHYTA ASPLENIACEAE
Asplenium auriculatum Sw., DW53, DW413 (LAGU,
SELAGINELLACEAE Mo)
Selaginelia sp., DW Asplenium cuspidatum Lam., DW52, DW137,
299 (B, LAGU, MO)
Dyn SDE Asplenium formosum Willd., DW315, DW503
ADIANTACEAE LAGU, M
Adiantum princeps 1. Moore, DW136 (B, LAGU, — Asplenium sp., DW139, DW427 (LAGU)
Me) . BLECHNACEAE
Anogramma leptophylla (L.) Link, DW424 (LAGU) Bleshindnnck occidentale ., DWS5 (LAGU. MIO)
Boe Og ne A eile Blechnum glandulosum Kaulf. ex Link, DW54
MO) aa
L
Cheilanthes kaulfussii Kunze, DW348 (LAGU)
Mildella intramarginalis (Kaulf. ex Link) Trevis., DAVALIACEAE
DW57, DW60 (B, EAP. LAGU, MO) eee $s ae flora (Roxb.) F.M. Jarrett ex CV.
Pityrogramma calomelanes (L.) Link, DW14, I (B, EAP, LAGU, MO)
DW297 (LAGU) appa aan Seni
m (L.) Maxon, DW527 (LAGU)
682
DICKSONIACEAE
Cibotium regale Verschaff. & Lem., DW471 (B,
LAGU, MEXU, MO)
DRYOPTERIDACEAE
Polystichum muricatum (L.) Fee, DW58 (LAGU, MO)
Tectaria mexicana (Fée) C.V. Morton, DW132
(LAGU, MO)
GLECHENIACEAE
Gleichenia bifidus (Willd.) Spreng.,DW141 (LAGU,
MO)
HYMENOPHYLLACEAE
Trichomanes hymenophylloides Bosch, DW247 (B,
LAGU, MO)
Trichomanes krausti Hook. & Grev., DW380 (LAGU)
POLY PODIACEAE
Campyloneurum angustifolium (Sw.
(LAGU)
Pecluma alfred
(LAGU, MO)
Phlebodium pseudoaureum (Cav.) Lellinger,
? (LAGU)
Pleopeltis angusta Humb. & Bonpl. ex Willd.,
DW129 (B, LAGU, MO)
Fée, DW455
ii (Rosenst.) M.G. Price, DW56
Polypodium furfuraceum Schltdl.& Cham., W300
(LAGU)
Polypodium plumula Humb. & Bonpl. ex Willd.,
DW298 (LAGU)
Polypodium polypodioides (L.) Watt var.
polypodioides, DW50, DW243, DW436 (B,
GU
~
MO)
SCHIZAEACEAE
Anemia hirsuta (L.) Sw., DW217, DW354 (B, LAGU,
MO)
A ia oblongifolia (Cav.) Sw., DW140 (B, LAGU,
MC
Anemia cf. phyllitidis (L.) Sw., DW127 (LAGU)
Anemia speciosa C. Pres|., DW131, DW302 (LAGU,
MO)
THELYPTERIDACEAE
Thelypteris oligocarpa (Humb. & Bonpl. ex Willd.)
Ching, DW59, DW237 (LAGU, MO)
Thelypteris puberula (Baker) C.V. Morton. var.
puberula, DW261 (LAGU)
DIVISION CONIFEROPHYTA
ARAUCACEAE
*Araucaria heter IL hylla (Salisb.) Franco, DW
BRIT.ORG/SIDA 20(2
=
CUPRESACEAE
*Cupressus lusitanica Mill., DW425 (LAGU)
PINACEAE
*Pinus caribaea Morelet
Pinus oocarpa Schiedue ex aide ADESCAM
DIVISION MAGNOLIOPHYTA
MONOCOTYLEDONS
AGAVACEAE
*Yucca guatemalensis Baker, ADESCAM
flies
rophyllum Eng., DW631 (LAGU)
nzantedeschi a deb jopica (L.) Spreng., DW626
(LAGU)
ARECACEAE
Chamaedora sp., DW579 (LAGU)
mina
na rr
Diane hee awla (Lindl.) Beer, DW118,
DW528 (LAGU)
Pitcairnia imbricata (Brongn.) Regel, DW337
ies
z, DW100 (B, LAGU, MO)
Namaste anne m aoa E. Morren, DW530
(LAGU)
illandsia fasciculata Sw., DW338 (LAGU)
landsia cf. kammi Rauh, DW412, DW529 (LAGU)
Tillandsia vicentina Standl., DW254 (LAGU)
COMMELINACEAE
Commelina diffusa Burm. f., DW151, DW190
DW327, DW331 (LAGU, MO)
Commelina erecta L., DW328 (LAGU, MO)
Commelina leineaipe ee sere (LAGU)
Commelina tubero Uu Le 82 (L LAGU U)
Gibasis pellucida (M. ree & Galeotti) DR.
nt, DW180 (LAGU)
Tinantia longipedunculata Standl. & Steyerm.,
AGU)
Tradescantia zanonia (L.) Sw., DW155, DW519
(LAGU)
Tripogandra montana Handlos, DW322 (LAGU,
MO)
CYPERACEAE
Bulbostylis tenuifolia (Rudge) J.F.Macbr., DW206,
DW390 (B, LAGU, MO)
aie
Pas
WILLIAMS ET AL., FLORA OF THE LAGUNA DE ALEGRIA, EL SALVADOR
Cyperus ee (Willd.) Endl, 0W221,DW335
(B, LAGU, MO)
Cyperus Bae Vahl, DW222? (LAGU)
Cyperus hermaphroditus Jacq.) Standl., DW12
DW134, DW149 (LAGU, MO)
Cyperus laxus Lam., DW195 (LAGU, MO)
ae tenerrimus J. Presl. & C. Presl., DW496
(LAGU)
Eleocharis filiculmis Kunth, DW274,AM3025 (BM
O)
Rhynchospora contracta (Nees) J.Raynal,OW318
(LAGU)
Torulinium odoratum (L.) S.S. Hooper, DW334 (B,
LAGU, MO)
DIOSCOREACEAE
Di icana Scheidw., DW381, DW520 (B,
LAGU, MO)
HYPOXIDACEAE
Hypoxis decumbens L., DW377 (LAGU)
IRIDACEAE
Sisyrinchium tinctorium Kunth, DW623 (LAGU)
LILIACEAE
Bomeria sp., DW619 (LAGU)
Echeandea skinnerii (Baker) Cruden, DW290 (B,
LAGU, MO)
MUSACEAE
*Musa x paradisiaca L.(pro.sp.), DW
ORCHIDACEAE
Bletia purpurea (Lam.) A. DC., DW244, DW500,
DW516 (LAGU, MO)
Cattleya qurantiaca (Batem.) P.N. Don, DW150
GU)
Cranichis apiculata Lindl., FH
Cycnoches egertonianum Batem., JBL 142 (LAGU)
Deiregyne hemichrea (Lindl.) Schltr., DW43,
DW527, FH (LAGU, MO)
Encyclia baculus (Rchb.f.) Dressler & G.E. Pollard,
DW487 (LAGU)
Paeveld eae ae : zene DW536
MO)
Gotenia ciliilabia ee : ie JBL/68/Win-
kler (LAGU
—
LAGU)
)
Govenia liliacea (La Llave & Lex.) Lindl., JBL386/
Hamer, FH (LAGU)
Govenia mutica Rchb. f., DW391, FH (LAGU)
Govenia utriculata (Sw.) Lindl., FH
Habenaria alata Hook., DW347 (LAGU)
683
Habenaria floribunda Lindl., JBL793/Winkler, FH
(LAG
eee crurigera Batem. ex Lindl., DW248,
W404, DW501 (B, LAGU, MO)
Pea: micrantha Lindl., DW414 (B, LAGU,
MO)
Isochilus amparoanus Schltr., DW148, DW346,
DWS510 (LAGU)
Laelia rubescens Lindl., DW537 (LAGU)
Lycaste sulfurea Rchb.f, DW5 18, FH, JBL394, JBL842
(B, K, LAGU, MO, SEL)
Maxillaria variabilis Batem. ex Lindl., DW246,
DW403 (B, LAGU, MO)
Mormodes x salvadorensis Hamer & Garay, FH
ie quadrifida (Lex.) Lindl., DW267,
FH (LAGU, MO)
rota cf. triloba Schitr., DW340
LAGU)
Sobral ia macrantha Lind|., DW384 (LAGU)
Spiranthes sp. |, DW252 (LAGU)
Spiranthes sp. Il, DW379 (LAGU, MO)
Stelis vestita Ames, DW535 (LAGU)
Trichopilia maculata Rchb. f., FH
POACEAE
Aegopogon cenchroides Humb.& Bonpl.ex Willd,
DW356 (B, LAGU, MO)
Andropogon angustatus (J. Pres!) Steud., DW292
(LAGU)
Andropogon leucostachyus Kunth, DW126 (B,
LAGU, MO)
Aristida capillacea Lam., DW173 (B, K, LAGU, MO)
** Arthraxon hispidus (Thunb.) Makino var.
hispidus, DW5, DW178 (B, LAGU, MO)
Axonopus capillaris (Lam.) Chase, DW314, DW523
(LAGU)
Chloris radiata (L.) ae DW17 (LAGU, MO)
**Cynodon dactylon (L.) Pers., DW423 (LAGU)
Digitaria ciliaris (Retz) Koeler, DW281, DW287,
DW288, DW289 (B, LAGU, MO)
**Fleusine indica (L.) Gaertn., DW71, DW179,
DW309 (B, LAGU, MO)
Eragrostis hypnoides (Lam.) Britton et al., DW512
(B, EAP, K, LAGU, MO)
Eragrostis mexicana Davidse subsp. mexicana,
DW220, DW294 (B, LAGU, MO)
** Hyparrhenia rufa (Nees) Stapf, DW70, DW108
(LAGU, MO)
Lasiacis procerrima (Hack.) Hitchc., DW321
(LAGU)
**Melinis minutifolia P. Beauv., DW76, DW123
DW?12 (LAGU, MO)
beg ey tenella (Kunth) Trin., DW341,
7, DW361 (B, LAGU, MO)
ae nus burmanii ee McVaugh var.
aE DW133, 52 (B, LAGU, MO)
Oplismenus hirtellus (L.) Se (LAGU)
Panicum arundinariae Trin. ex E. Fourn., DW176
(LAGU
¥¥D Inicum maximum Ja acq
DW293 (LAGU)
Panicum tri oe Sw., DW199 (LAGU)
Panicum sp., DW39 (K, LAGU, MO)
Paspalum can an (Humb.& Bonpl. ex Flugge)
Kunth, DW264, DW364 (LAGU, MO
Paspalum ocuiads C.Wright, DW312 (LAGU)
Paspalum um Humb.& Bonpl. ex Fluggé,
185, DW21 Sees U)
Paspalum costaricense Mez, DW28?, DW283
a=
(LAGU
Paspalum notatum Fluggé, DW286, DW310
(LAGU, MO)
mM naniciulatiym
W284 (B, LAGU, MO
Penniseturn aomolanaiin (Nees) Hemsl., DW5 14
=
(LAGU, MO)
**RAynchelytrum repens (Willd.) C.E. Hubb., DW6 1,
DW311, DW479 (LAGU, MO)
Setaria parviflora (Poir.) Kerguelen, DW291,
DW360 (B,L O
Sporobolus indicus (L (L .) R.Br, DW4, DW285 (B, K,
LAGU, MO)
cf. Urochloa sp., DW462 (LAGU, MO)
undetermined sp., DW/363, DW504 (LAGU, MO)
DIVISION MAGNOLIOPHYTA
DICOTYLEDONS
ACANTHACEAE
Blechum browneii Juss., DW19 (LAGU)
Elytraria imbricata (Vahl) Pers., DW65, DW120 (B,
LAGU, MO, US)
Henrya insularis Nees, DW153, DW459 (B, LAGU,
MO)
Tetramerium nervosum Nees, DW146 (B, LAGU,
MO)
ACTINIDACEAE
Saurauia kegeliana Schitdl., DW565 (LAGU)
AMARANTHACEAE
Achyranthes indica (L.) Mill, DW71, DW109 (B,
LAGU, MO)
BRIT.ORG/SIDA 20(2)
Amaranthus spinosus L., DW230 (LAGU)
Chamissoa altissima Jacq.) Kunth, DW256 (B, F,
U, MO)
lresine calea (Ibanez) Standl., DW26 (B, F LAGU,
MO)
lresine diffusa Humb. & Bonpl. ex Willd., DW20,
DW42, DW46, DW69, DW355 (B, F, LAGU, MO)
Pleuropetalum sprucei (Hook. f.) Standl., DW471
(LAGU)
ANACARDIACEAE
*Mangifera indica L., DW513 (LAGU)
Rhus terebinthifolia Schltdl. & Cham., DW453
(LAGU)
Spondias purpurea L., ADESCAM
ANNONACEAE
Annona cherimola Mill, DW434, AM3019 (BM,
AGU)
a diversifolia Saff., DW629 (LAGU)
Annona holosericea Saff., MLRA
APIACEAE
Spananthe aie Jacq., DW345 (LAGU, MO)
AeceeaNin
onia longifolia on DC.) Pichon, ADESCAM
*Thevetia peruviana (Pers.) K. Schum., DW551
(LAGU)
ARALIACEAE
Oreopanax xalapensis (Kunth) Decne. & Planch.,
DW8 G
ARISTOLOCHIACEAE
Aristolochia sp., DW
ASCLEPIADACEAE
Gonolobus lasiostemma (Hemsl.) Woodson,
DW428 (LAGU, MO)
ASTERACEAE
Acmella alba (UHer.) R.K. Jansen, DW117, DW231
( U, MO)
Pratum conyzoides L., DW125 (B, LAGU, MO)
Baccharis nents Pers., DW78 (LAGU, MO)
Baccharis monoi 1G.L.Nesom, DW116 (LAGU)
Bidens ilosa L. “DW200, DW303 (LAGU)
Bidens squarrosa Kunth, DW25, DW27, DW228 (K,
LAGU, MO)
Calyptocarpus wenlandii Sch. Bip., DW415 (LAGU)
Chaptalia nutans (L.) Pol., DW386 (LAGU, MO)
Chromolaena odorata (L.) R.M. King & H. Rob.,
DW86 (LAGU)
oak
WILLIAMS ET AL., FLORA OF THE LAGUNA DE ALEGRIA, EL SALVADOR
L.) Cronquist, DW410, DW431
haa
Conyza bonariensis (
(LAGU)
Coreopsis mutica D.J.Crawford var. microcephala,
1 (LAGU)
Critonia daleoides DC., DW112, DW250 (LAGU,
MO)
Critonia morifolia (Mill.) R.M. King & H. Rob.,
DW277, DW463 (LAGU, MO)
*Dahlia imperialis Roezl ex Ortgies, DW/4 (B,
LAGU, MO)
Eclipta prostrata (L.) L., DW326 (LAGU, MO)
Elephantopus mollis Kunth, DW68, DW207 (B,
LAGU, MO
ee spicatus B. Juss. ex Aubl., OW78 (B,
K, LAGU, MO)
Erechtites valerianaefolius (Link ex Spreng.) DC.,
DW373 (LAGU, MO)
Erigeron karvinskianus DC., DW422, DW488
(LAGU)
Fleischmanniopsis leucocephala
King & H. Rob., DW226 (LAGU)
Galinsoga caracasana (DC.) Sch. Bip., DW103
(LAGU, MO
Galinsoga urticifolia (Kunth) Benth., DW419,
DW421 (B, LAGU, MO)
Jaegeria hirta (Lag.) Less., DW191 (LAGU)
anophyllon cf. standleyi (Robinson) R.M. King
& H. Rob., DW165 (B, LAGU, MO)
Lasianthaea fruticosa (L.) K.M. Becker, DW209
(LAGU)
Melampodium divaricatum (Rich. ex Pers.) DC.,
DW306, DW308, DW401 (B, LAGU, MO)
Melampodium paniculatum Gardner, DW115,
W305, DW307, DW323 (B, K, LAGU, MO)
Melanthera nivea (L.) Small, DW105 (B, K, LAGU,
MO
(Benth.) R.M
Onoseris onoseroides (Kunth) B.L. Rob., DW271
(LAGU)
sa glabratus (J.M. Coult.) S.F. Blake,
DW164, DW402 (LAGU,
Perymenium grande Hemsl. var. grande, DW88
(LAGU, MO)
Peteravenia schultzii R.M. King & H. Rob., DW6,
DW113, W395 (B, K, LAGU, MO)
oldana petasioides (Greenm.) H. Rob., DW85 (B,
Sinclaria discolor Hook. & Arn., DW12 (LAGU)
Sinclaria sublobatum H. Rob., DW75 (LAGU, MO)
Sinclaria sp., DW510 (LAGU)
685
Sonchus oleraceus L.,DW229, DW295 (LAGU, MO)
Stevia ovata Willd., DW84, DW389 (B, K, LAGU, MO)
Stevia tephrophylia S.F. Blake, DW351 (LAGU)
Synedrella nodiflora (L.) Gaertn., DW90 (B, K, LAGU,
MO)
Tagetes filifolia Lag., DW397 (LAGU)
Tagetes tenuifolia Cav., DW21, DW83 (B, LAGU,
MO)
Telanthophora arborescens (Steetz) H. Rob.& Bret-
tell, DW238 (B, LAGU, MO)
Tithonia rotundifolia (Mill.) S.F. Blake, DW187
(LAGU, MO)
Tridax procumbens L., DW216, DW430 (B, LAGU,
MO)
Trigonospermum annuum McVaugh & Lask.,
DW203 (LAGU, MO)
Verbesina cf. punctata B.L.Rob.& Greenm., DW82
(LAGU)
Verbesina sp., DW367 (B, K, LAGU, MEXU, MO)
ell ia cf. deppeana Less., DW263 (B, LAGU, MO)
arpa DC., DW9 (B, K, LAGU, MO)
4 (B, K, LAGU, MO)
VONIA leio
Vernonia patens Kunth, DW114
cf. Vernonia sp., DW517 (LAGU)
Wedelia parviceps S.F. Blake, DW23, DW157
DW249, DW420 (LAGU)
** Zinnia elegans Jacq., DW251, DW399 (B, kK,
LAGU, MO
BEGONIACEAE
Begonia plebeja Liebm., DW79 (LAGU)
BIGNONIACEAE
* Jacaranda mimosifolia D. Don, DW550 (LAGU)
Macfadyena unguis-cati (L.) A.H. Gentry, DW435
GU)
Tabebuia cf. chrysantha (Jacq.) G. Nicholson,
SCAM
ADE
Tabebuia rosea (Bertol.) A. DC., DW260 (B, EAP,
LAGU, MEXU, MO)
Tecoma stans (L.) Juss. ex Kunth, DW28, DW106
(LAGU, MO)
BIXACEAE
+Bixa orellana L., ADESCAM
BOMBACACEAE
Ceiba aesculifolia (Kunth) Britt.& Baker f, MLRA
Ceiba ee (L.) Gaertn., DW576 (LAGU)
BORAG AE
Cordia eh (Ruiz & Pav.) Oken, ADESCAM
Cordia bullata (L.) Roem. & Schult., DW296,
DW400 (B, LAGU, MO)
—
Cordia inermis (Mill.) LM. Johnst., DW426 (LAGU)
Heliotropium rufipilum (Benth.) LM. Johnst.,
DW259 (B, LAGU, MO)
Heliotropium sp., DW124 (B, EAP, LAGU, MO)
BRASSICACEAE
Lepidium virginicum L., DW268, DW417 (LAGU,
MOQ)
BURSERACEAE
Bursera simaruba (L.) Sarg., DW445, AM3023 (BM,
LAGU)
CAMPANULACEAE
Diastatea micrantha (Kunth) McVaugh, DW8,
DW272, DW330 (B, LAGU, MO)
Lobelia laxiflora Kunth, DW67, DW152 (LAGU)
CARICACEAE
Carica cauliflora Jacg., DW121 (LAGU)
CARYOPHYLLACEAE
Arenaria lanuginosa (Michx.) Rohrb., DW407
(LAGU, MO)
Drymaria cordata (L.) Willd. ex Schult., DW213,
DW38 MO)
Stellaria ovata Willd. ex Schltdl., DW408 (LAGU,
MO)
CASUARINACEAE
*Casuarina equisetifolia L., DW621 (LAGU)
CECROPIACEAE
Cecropia peltata L., DW577 (LAGU)
CLETHRACEAE
Clethra lanata M. Martens & Galeotti, MLRA
Clethra mexicana DC., DW143, DW438 (B, EAP.
DW524 (LAGU, MO)
cf. Garcinia sp., DW452 (LAGU)
COCHLOSPERMACEAE
Cochlospermum vitifolium (Willd.) Spreng.,
ADESCAM
CONVOLVULACEAE
[pomoea cholulensis Kunth,
DW241 (LAGU, MO)
Ipomoea cf. mairetii Choisy, DW208 (LAGU)
Ipomoea santae-rosae Standl.& Steyerm., DW145
(LAGU)
CLUSIACEAE
Calophyll l 1.) Stand. var. rekoi
DW154, DW158
Ipomoea squamosa Choisy, DW177, DW350 (B,
LAGU, MO)
BRIT.ORG/SIDA 20(2)
Ipomoea cf. trifida (Kunth) G. Don, DW47 (LAGU,
MO
Ipomoea umbraticola House, DW44, DW227,
DW278 (B, LAGU, MO)
CUCURBITACEAE
Cayaponia sp., DW620 (LAGU)
Microsechium sp., DW188 (LAGU, MO)
Rytidostylis ciliata (Cogn) Kuntze, DW62, DW98
(LAGU)
Sechium edule Jacq.) Sw., DW
Sycios sertuliferus Cogn., DW142 (LAGU, MO)
ERICACEAE
Gaultheria erecta Vent., DW437 (LAGU, MO)
EUPHORBIACEAE
Acalypha macrostachya Jacq., DW255 (B, LAGU,
MO
Chamaesyce hirta (L.) Millsp., DW324, DW511
(LAGU)
Croton xalapensis Kunth, DW89 (B, F, LAGU, MO)
sare graminea Jacq., DW172, DW498
(LAGU)
Euphorbia heterophylla L., DW66 (B, K, LAGU, MO)
*Fuphorbia pulcherrima Willd. ex Klotzsch,
ADESCAM
¥RICI is L., DW122 (LAGU)
FABACEAE
Acacia hindsii Benth., DW443 (LAGU)
*Caesalpinia pulcherrima (L.) Sw., DW80 (LAGU,
MO)
*Cajanus cajan (L.) Millsp., DW175 (LAGU)
Calliandra sp., DW627 (LAGU)
Sl ae DW38 (K, LAGU, MO)
ea Lam., DW575 (LAGU)
eee virginianum (L.) Benth, DW210
Chamaecrista nictitans (L.) Moench, DW239,
DW359 (LAGU)
hyviloides (Rri & Rose) H.S.
Irwin & Barneby var. deamii, DW444 (LAGU)
Clitoria mexicana Link, DW204 (LAGU)
Cologania cf. rufescens Rose, DW18] (LAGU)
Coursetia caribaea (Jacq.) Lavin, DW730 (LAGU,
MO)
Crotalaria mollicula Kunth, DW138 (B, LAGU, MO)
Dalea cliffortiana Willd.,, DW349 (B, LAGU, MO)
peas sp., DW240 (LAGU)
esmodium barbatum (L.) Benth. DW342 (LAGU)
pale cajanifolium (Kunth) DC., DW447
(LAGU)
=
WILLIAMS ET AL., FLORA OF THE LAGUNA DE ALEGRIA, EL SALVADOR
Desmodium incanum DC., DW406, DW414
(LAGU)
Desmodium procumbens (Schindl.) B.G. Schub.
var, nS eee (B, K, LAGU, MO)
Diphysa rob 5 Benth., DW520 (LAGU, MO)
eahiing berteroana Urb., ADESCAM
Hymenaea courbaril L., a
Inga paterno Harms, D
Inga pavoniana G.Don, ane (LAGU)
Inga punctata Willd., DW523 (LAGU)
Lonchocarpus rugosus (Lundell) M.Sousa subsp.
apricus, DW448, DW454 (LAGU, MO)
Lysiloma cf. acapulcense (Kunth) Benth., DW439
(B, LAGU, MEXU, MO)
Macroptilium atropurpureum (Moc. & Sessé ex
DC.) Urb., DW273 (LAGU)
Mimosa albida Humb.& Bonpl. ex Willd., DW170
(B, K, LAGU, MO)
Mimosa pudica L., DW378 (LAGU)
Phaseolus leptostachyus Benth., DW169, DW214
(LAGU, MO)
Phaseolus lunatus L., DW3 (B, LAGU)
Piscidia grandifolia (Donn. Sm.) 1.M. Johnst.,
DW469 (LAGU)
Poiretia punctata (Willd.) Desv., DW174 (LAGU,
MO)
Senna hirsuta H.S. Irwin & Barneby var. hirta,
DW372 (B, LAGU, MO)
Spathodia campanulata Bea
Stylosanthes guianensis ne ‘ [ an DW96
(B, K, LAGU, MO)
Te inatus (L.) Sw., DW182 (LAGU)
Vigna sp., DW183 (LAGU)
Zapoteca tetragona (Willd.) H.M. Hern., DW135
(LAGU, MO
Zornia diphylla (L.) Pers. DW171
FAGACEAE
epee Liebm., DW144,,MLRA1506 (B,
K, LAGU,M
(LAGU)
Quercus Sp., ae DW269 (LAGU)
FLACOURTIACEA
Casearia sylvestris ae DW467, DW451,MLRA1504
(B, LAGU, MEXU, MO, )
Prockia crucis ‘ Br. ex L., DW375, AM3012 (B, BM,
LAGU, M
Xylosma oe (Kunth) Hemsl., 0W339 (LAGU
GESNERIACEAE
Achimenes erecta (Lam.) H.P. Fuchs, DW394,
DW492 (LAGU, MO)
So,
687
oe menes teen DC., DW329 (B, LAGU, MO)
indl., DW344 (LAGU, MO)
Achimenes ae Benth., DW63, DW365,
DW374 (B, LAGU, MO)
Phinaea repens (Donn. Sm.) Soler., DW316 (B,
GU, MO)
HERNANDACEAE
Gyrocarpus americanus Jacq., DW569 (LAGU)
HYDROPHYLLACEAE
Wigandia urens (Ruiz & Pav.) Kunth, DW223
(LAGU)
himenec micerq
JUGLANDACEAE
Juglans olanchana Standl.& L.O. Williams, DW
LAMIACEAE
Hyptis pectinata (L.) Poit., DW45 (B, LAGU, MO)
Hyptis suaveolens (L.) Poit., DW94 (LAGU)
Hyptis urticoides Kunth, DW101, DW205 (B, LAGU,
MO)
—
=
AA : +h L f 5 an ee
(B, LAGU, MO)
via occidentalis Sw., DW2, DW111, DW353 (B,
K LAGU, wee
_DW370
poe
Sa
Ortega, DW110 (LAGU, MO)
Salvia ais pea Cav., DW270 (B, LAGU, MO)
*Salvia splendens Sellow ex Roem. & Schult.,
DW40 (B, K, LAGU, MO)
LAURACEA
nestaner sf inuata Mez, DW538 (LAGU)
Mill, ADESCAM
LOASACEAE
Klaprothia fasciculata (C. Presl.) Poston, DW332,
DW524 (B, LAGU, MO)
Metzelia sp., DW358, DW376 (LAGU)
LOGANIACEAE
ddleja americana L., DW266 (LAGU)
LORANTHACEAE
Oryctanthus cordifolius (C. Pres.) Urban, DW522
(LAGU)
MAnRISHIACESE
lia (L.) Kunth, ADESCAM
Goud ie albida Cham. & Schitdl., DW93,
8 (B, LA )
ee, schiedeana Cham. & Schlitdl., DW37,
DW3 W494, DW499 (LAGU)
lon ellipticum (Kunth) A. Juss., DW33
MO)
IN
a
Stigmaphy:
(LAGU
MALVACEAE
*Hibiscus rosa-sinensis L., DW236 (B, LAGU, MO
Malvaviscus arboreus Cav., DW75 (B, EAP LAGU,
, US)
Sida acuta Burm. f., DW92 (B, LAGU, MO)
Sida glabra Mill., DW34 (LAGU, MO)
Sida urens L., DW72 (LAGU, MO)
MELASTOMATACEAE
sa tices xalapensis (Bonpl.) D. Don ex DC.,,
W446, AM3001 (B, BM, LAGU, MO)
hihi et hondurense Gleason, DW95 (LAGU,
MO)
Whiffin, DW
Miconia in nee DC., MLRA
Miconia guatemalensis Cogn., DW369, DW468,
Aine? (8, BM, a LAGU, MEXU, MO)
Aj is Naudin, DW196 (LAGU)
Monochaetum flori ne (Schitdl.) Naudin,
W35 (B, CAS, LAGU, MO)
Pterolepis pumila (Bonpl.) Cogn., DW102, DW325
(B, LAGU, MO)
Schwackaea cupheoides (Benth.) Cogn., DW32
(LAGU)
Tibouchina longifolia (Vahl) Baill., DW91, DW99
(LAGU)
Heterocentron 7 landulosum var. longicalyx
7 (LAGU)
MELIACEAE
Cedrela odorata L., DW392, DW474 (LAGU)
Cedrela tonduzii C. —
*Melia azedarach L., D
Trichilia havanensis i DWo4, DW371,AM3020
ae
=<
Trichilia hirta L., DW257, AM3009 (BM, LAGU)
Trichilia tomentosa Kunth, MLRA
MENISPERMACEAE
Cissampelos pareira L., DW509 (LAGU, MO)
ORACEAE
on a ae ii Standl., DW2717 (LAGU)
Fic tusifolia | unth, DW? 15, DW539 (LAGU)
poet s(Liebm.) Miq., DW163, DW166, DW405,
D U, MO)
Ficus pertusa L.f,, DW186 (B, F, LAGU, MO)
MYRICACEAE
Myrica cerifera L., DW3 1, DW440 (B, LAGU, MO)
MYRSINACEAE
Ardisia compressa Kunth, DW119, DW162 (LAGU,
MO)
BRIT.ORG/SIDA 20(2)
Rapanea myricoides (Schltdl.) Lundell, DW477
(LAGU)
MYRTACEAE
Eugenia cf. acapulcensis Steud., AM3003 (BM,
LAGU
Psidium guajava LRA
Psidium guineense a. DW16, DW77, DW533
(LAGU, MO)
Psidi tedianum O. Berg, MLRA
NYCTAGINACEAE
*Bougainvillea glabra Choisy, DW235 (LAGU)
pe DW161, DW253 (LAGU)
cf. Mirabilis sp., DW388 (LAGU)
ve aaculeata L., DW265, DW336 (LAGU)
OXALIDACEAE
Oxalis corniculata L., DW505 (LAGU)
Oxalis oe (DC.) Lourteig var. corymbosa,
DW622 (LAGU)
Oxa infuse L., DWI, DW159 DW497 DW517
(B, U, MO)
PAPAVERACEAE
Bocconia frutescens lis, DW234 (LAGU)
PASSIFLORACEAE
Passiflora ornithoura Mast., DW49 (LAGU)
PHY TOLACCACEAE
Phytolacca icosandra L., DW160, DW418 (LAGU,
MO
Rivina humilis L., DW13, DW107 (B, LAGU, MO)
PIPERACEA
Peperomia ae aC. DC., DW280 (LAGU, MO)
Peperomia granulosa Trel., DW317 (LAGU, MO)
Peperomia deppeana Schltdl. & Cham., DW279
AGU, MO)
Peperomia lanceolato-peltata (C. DC.), DW531
LAGU, MO
Peperomia cf. liebmannii C.DC., DW515 (F, LAGU,
MO
Peperomia sp., DW184 (LAGU)
Piper amalago L.,DW456, DW458, DW462, DW466
(LAGU, MO
Piper hispidum Sw., DW473, DW464 (LAGU, MO)
Piper jacquemontianum Kunth, DW304, DW485
(LAGU)
Piper marginatum Jacq., 0W475, DW486 (LAGU)
Piper umbellatum L., DW461, DW474 (LAGU)
WILLIAMS ET AL., FLORA OF THE LAGUNA DE ALEGRIA, EL SALVADOR
POLEMONIACEAE
Loeselia glandulosa (Cav.) G. Don, DW7, DW73,
DW507 (B, LAGU, MO)
POLYGALA
Polygala Gane Chodat, DW48, DW320
(LAGU)
Polygala longicaulis Kunth, DW393, DW525
(LAGU, MO)
Polygala paniculata L., DW225, DW398, DW493
(LAGU)
Polygala sp., DW189 (LAGU)
PORTULACACEAE
Talinum paniculatum (Uacqg.) Gaertn., DW6718
(LAGU)
PROTEACEA
Roupala ae ma Pittier, DW470 (LAGU)
RHAMNACEAE
Colubrina reclinata (L'Her.) Brongn., DW5/0
(LAGU)
ROSACEAE
*Eriobotrya japonica (Thunb.) Lindl., DW
Prunus brachybotrya Zucc., MLRA1505, AM3013
(BM, LAGU)
Rubus irasuensis Liebm., DW449 DW476 (LAGU,
MO)
*Rubus niveus Thunb., DW4817 (B, LAGU, MO)
RUBIACEAE
Borreria cf. latifolia (Aubl.) K. Schum., DW362
(LAGU)
Borreria ocymoides (Burm.f) DC., DW201, DW218
(B, F, LAGU, MO)
Bouvardia leiantha Benth., DW276, DW480,
DW484 (B, LAGU, MO)
*Coffea arabica L., DW460 (LAGU)
*Coffea liberica W. Bull ex Hiern, DW
Crusea longiflora (Willd. ex Roem. & Schult.) W
Anderson, DW495 (LAGU)
Hamelia patens Jacq., DW233, DW464 (B, LAGU,
MOQ)
Hoffmannia angustifolia Standl., AM3002 (BM
LAGU)
Manettia reclinata L., DW192 (LAGU)
Mitracarpus villosus (Sw.) Cham. & Schltdl., DW22,
D W534 (LAGU, MO)
Psychotria pubescens Sw., DW366 (LAGU, MEXU,
MO)
Richardia scabra L., DW30 (LAGU, MO)
689
RUTACEAE
Casimiroa edulis La Llave & Lex.,DW197 (B, LAGU,
MQ)
*Citrus limetta Risso, heat
*Citrus sinensis (L.) Osbeck, D
Zanthoxyllum cf. culantrillo oi ADESCAM
SAPINDACEAE
apindus saponaria L., DW457 (LAGU)
SAPOTACEA
— on nei (Hemsl.) T.D. Penn,, DW242,
AM3018 (BM, LAGU, MO)
SCROPHULARIACEAE
Castilleja arvensis Schitdl.& Cham., DW104 (B, BKL,
F, LAGU, MO)
Russelia sarmentosa Jacq., DW275, DW506 (LAGU)
SOLANACEAE
Brachistus stramoniifolius (Kunth) Miers, DW528
Cestrum lanatum M. Martens & Galeotti, DW432
(LAGU)
Physalis pubescens L., DW529, DW530 (B, LAGU,
MO)
So
anum lanceifolium Jacq., DW513, DW521
(LAGU, MO)
anum ni cia var. americanum (Mill) O.E.
chulz, DW LAGU)
Solanum torvum an DW473 (LAGU)
=
So
Aa IACEAE
altheria glomerosa C. Presl., DW198 (LAGU)
toe indica L., DW36, DW198 (LAGU, MO)
STYRACACEAE
Styrax argenteus C. Presl., DW41
ienetin
mexicanus (Turcz.) Sprague, DW479
6, EAP, LAGU, NEN MOQ)
a5,
1 (B, LAGU, MO)
W202 (LAGU)
DC., DW478 (LAGU)
Tri mfetta b norenc
J
LMACEAE
Trema micrantha (L.) Blume, ADESCAM
mus mexicana (Leibm.) Planch, DW
URTICACEAE
Boehmeria caudata Sw., DW24, DW482, AM3011
(B, a LAGU, MO)
noine
S
—
a)
=
8
Liebm., DW483 (LAGU, MO)
(LAGU)
Pilea hyali na pene: DW319
690
Urera cf. baccifera (L.) Gaudich., AM3016 (BM,
LAGU)
Urera caracasana (Jacq.) Grisebach., DW396,
W.
DW490 (LAGU)
VALERIANACEAE
Valeriana scandens (Gardn.) K.A.E. Muell. var.
BRIT.ORG/SIDA 20(2)
Citharexylum donnell-smithii hole DWIO,
DW29, DW429 (B, EAP, LAGU, MO)
Lantana camara L., DW156 (B, oe LAGU, MO)
Lippia myriocephala Schltdl. & Cham., DW472 (B,
EAP, LAGU, MEXU,M
VITACEAE
candolleana, DW224, DW245 (LAGU, MO)
VERBENACEAE
Cornutia pyramidata L., DW368, DW416
MO)
Cissus erosa Rich., DW624 (LAGU)
(LAGU,
REFERENCES
BeRENDSOHN, W.G. 1991. The arboreal vegetation of the Laderas de La Laguna, a Neotropical
Forest Fragment in El Salvador, C.A. Dissertationes Botanicae 165. Cramer, Stuttgart.
Daviose, G., M. Sousa, and S. Knapr. (eds.). 1995. Flora Mesoamericana. Vol. 1. Psilotaceae a
Salviniaceae. IBUNAM, Missouri Botanical Garden y The Natural History Museum
(London).
DIPRAT (DirECCION DE PLANEAMIENTO, PROGRAMACION, PROYECTOS, Y AUDITORIA TECNICA). 1997. El Salva-
dor: Degradacién y Perspectivas de Manejo de los Recursos Naturales Renovables. In-
stituto Interamericano de Co-operacién para la Agricultura, San José, Costa Rica.
Fassett, N.C. 1952.La Lagunita de Alegria. San Salvador, Comunicaciones: Tomo II (Anuario)
p. 24.
Hovorinee, L.R. 1975. Zonas de vida ecoldgica de El Salvador. FAO y Direccién General de
Recursos Naturales Renovables del Ministerio de Agricultura y Ganaderia, Soyapango,
El Salvador.
Hamer, F. 1974.Las orquideas de El Salvador, Tomo I-III. Ministeri
dor, El Salvador.
SACDEL (Sistema De ASESORIA Y CAPACITACION PARA EL DESARROLLO Local). 1997. Base conceptual
para el Parque Turistico Municipal en el Area Natural de la Laguna de Alegria. El
Salvador.
STANDLEY, PC. and J.A. StevermarK. 1960. The flora of Guatemala. Fieldiana: Botany. Chicago
Natural History Museum, Chica
Reyna be Acuitar, M.L. 1995. Flora. ane VIl de Historia Natural y Ecologia de El Salvador,
Tomo |. Ministerio de Educacion, El Salvador.
de Educacion, San Salva-
THE FLORA OP OK TIBBEHA COUNTY, MISSISSIPPI
Andreas Leidolf!, Sidney McDaniel, and Tim Nuttle?
Institute for Botanical oe on (IBE)
O Drawer EN
Mississippi State, MS oe 5668, U.S.A.
ABSTRACT
We surveyed the flora of Oktibbeha County, Mississippi, U.S.A., from February 1994 to 1996. Occupy-
ing 118 square kilometers in east-central Mississippi, Oktibbeha County lies among 3 physiographic
sees ee include, from west to east, Interior Flatwoods, Pontotoc Ridge, and Black Prairie. Ac-
co unty harbors a diverse flora. Based on field work, as well as an extensive review of
ete hone and herbarium records at IBE and MISSA, we recorded a total of 1,148 taxa (1,125
species, 7 hybrids, 16 infraspecific taxa) belonging to 514 genera in 160 families; over 85% of all taxa
ocumented pared to 3 other counties in east-central Mississippi, Oktibbeha County
has the second vaieest Recorded Nora ane number of state-listed kendaneoeG threatened, or of spe-
far exceeds that I any o other county en
\
rriére,
the region. Three introduced species, Ilex cornuta Lind]. & Paxton, Mahonia
ane Ne pala Gemesiiea Thunb., are reported i first time from Mississippi.
| ies belongi 5 broad habitat categories: bottomland
fo)
pee popiane forests ay ae aquatic Habitats meepane areas, and human-influenced habitats.
detai ription of the vegetation associated with each of these communities is provide
RESUMEN
H igado la flora del condado de Oktibbeha, Mississippi, 7 ea nawedianaas
ha ectaentn
Con 118 kilometros cuadrados en el centro-este de Mississippi,
e oeste aeste, Interior Blacwoods Pontotoc oe y Black Prairie. Con-
o Oo t
el lado alberg: flora diversa BaSaaos en el apale es an asi como en
do] j PR sy FE tz li 1; ISSA, a
un total d L148 t taxa a, 125 pe 7 hi nbedos 16 taxa sates tone Late ea
514 géneros de 160 familias; mas del 85% de los taxa sea eron autoctonos. ee rado
oa a | L mas
con otros 3 Gauls del conto» este oe Mississippi, el con
numerosa | I stado ( pl amend eace ee atencion especial)
} bY } ] ] nyt
| gion. Tres especies
introducidas, Ilex cornuta a & Paxton, Mahonia bealei (F ) Carriére, y Nandina domestica
Mi i. También se describen 16 comunidades
: Desi |
Thun Aississi
- B Ay
] 5 Hope ty DS 1 ]
cae diferentes pertenecientesa5 g | jas, bosq
y praderas, hal le filtracion, y habitats influenciados por el hombre. Se aporta
una des ae on detallada de la vegetacion asociada a cada una de estas comunidades.
ZUSAMMENFASSUNG
I ks Oktibbeha im Bund Mi i, Vereinigte
Wir untersuchten die Flora des Ver
Staaten von Amerika, im Zeitraum eee 1994 bis 1996. Der Vewaltunesbeanc: umfafst 118
‘Present address: USGS Utah Cooperative Fish and Wildlife Research Unit, Department of Forest, Range, and
Wildlife Sciences, Utah State University, Logan, UT 84322-5230, U.S.A.
Present address: Department of Biology, Utah State University, Logan, UT 84322-5305, U.S.A.
SIDA 20(2): 691 — 765. 2002
692 BRIT.ORG/SIDA 20(2)
Quadratkilometer im Mittelosten Mississippis und gehort, von West nach Ost, zu 3 physiographischen
Regionen (Interior Flatwoods, Pontotoc Ridge und Black Prairie), weshalb die Flora sehr artenreich
ist. Aufgrund unserer Freilandarbeit, sowie eines umfangreichen Studiums der Fachliteratur und
zahlreicher | oe aes in IBE und MISSA, waren wir in der Lage |. ee Taxa (1.125 Arten, 7
Hybriden und 16 Unterarten oder Varietaten) aus 514 Gattungen und 160 F katalogisieren
85% aller Taxa waren in Mississippi beheimatet. Der Verwaltungsbezirk Oktibbeha hat im Vergleich
zu 3 anderen Verwaltungsbezirken im Mittelosten Mississippis die zweitgrofte Flora. Die Anzahl
der Arten auf f der Roten - iste Mississippis (vom Aussterben bedroht oder von besonderem Belang)
ubersteigt mit it di randeren Verwaltungsbezirke in der Region. Drei eingeschleppte Arten,
lex cornuta L indl. & Paseo Mahonia bealei (Fortune) Carriére und Nandina domestica Thunb.
werden hier zum ersten Mal von Mississippi erwahnt. Im weiteren Verlauf beschreiben wir aufSerdem
im Detail 16 verschiedene Pflanzengesellschaften in 5 Lebensraumkategorien: Auenwalder,
Hochlandwalder und Prairien, Aquatische Lebensraume, Sickerstellen und vom Menschen
beeinflufSte Lebensraume.
INTRODUCTION
Unlike other areas in the southeastern United States, Mississippi is largely unex-
plored botanically (Duncan 1953; Pullen 1966; Bryson & Carter 1992: Bryson et
al. 1996; Sorrie & Leonard 1999; Alford 2001). To date, there is no comprehen-
sive flora for the state, although there are 8 public herbaria (DSC, HGCRL, IBE,
MISS, MISSA, MSCW, SWSL, University of Southern Mississippi) with combined
holdings of more than 400,000 specimens (Holmgren et al. 1990). In addition,
the Mississippi Museum of Natural Science (MMNS) maintains a herbarium of
ca. 8,000 specimens, as well as a computer database of plant records, both of
which are accessible to the public.
Historically, botanical explorations of Mississippi have been sporadic, and
largely concentrated on the southern part of the state (Flint 1882a; Halsted 1891).
The naturalist William Bartram traveled through parts of Mississippi in 1777
(Van Doren 1928), and the renowned botanist Thomas Nuttall reportedly col-
lected plants around Natchez, Mississippi, in 1811 and 1820 (Graustein 1967). In
1840, John Claiborne began his exploration of the pine forests of southern Mis-
sissippi (McDaniel 1986); twenty years later, Hilgard (1860) published comments
on the phytogeography of Mississippi. In the early twentieth century, Harper
(1906, 1914) gave a superficial account of the pine barren vegetation of south-
ern Mississippi based on his travels through that region. Lowe (1921) published
the first checklist of vascular plants for the state.
In 1964, the Flora of Mississippi Project, funded by the National Science
Foundation, was initiated by Samuel B. Jones, Jr, Thomas M. Pullen, and Ray
Watson. A number of publications (Pullen et al. 1968a. 1968b; Jones et al. 1969;
Watson 1969, 1970a, 1970b; Jones 1974a, 1974b, 1975a, 1975b, 1976a, 1976b) re-
sulted, but the goal of a comprehensive floristic treatment was never achieved.
A definitive treatment of the flora of Oktibbeha County has also been lacking
so far, although floristic surveys have been performed for several Mississippi
counties (Flint 1882b; Ferrari 1970; Morgan 1979: Meeks 1984; Morris 1987;
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI 693
Winstead 1990; MacDonald 1996; Alford 2001), as well as other selected areas
(Lassetter 1968; Massey 1974; Carter 1978; McCook 1982; Moore 1993). Hilgard
(1860) stated: “With the Flatwoods of Octibbeha [sic] county, | am not person-
ally acquainted,” and the floristic literature for Oktibbeha County remains
sparse. In the first published checklist of Mississippi plants, Lowe (1921) cites
several vascular plant collections from Oktibbeha County, among them many
species considered rare today by the Mississippi Natural Heritage Program
(MNHP). However, the first botanical work dealing explicitly with the flora of
Oktibbeha County was a survey of liver- and hornworts (Woods 1964). In re-
cent years, some of the rare plant communities found in Oktibbeha County have
been studied (Morris et al. 1993; Leidolf & McDaniel 1998).
The objectives of this study were to develop an annotated checklist of vas-
cular plants and bryophytes (excluding Bryopsida) for Oktibbeha County. In
addition, we wanted to provide information on habitat association and abun-
dance of the plant species included in this checklist. In doing so, we hope to
establish a botanical baseline for ecological research in this area, while at the
same time making an important contribution to the recently revived Flora of
Mississippi Project (e.g., Bryson & Carter 1992).
STUDY AREA
Location and Description of Oktibbeha County
Oktibbeha County is located in east-central Mississippi, approximately 200 km
northeast of the state capital Jackson, and ca. 60 km west of the Mississippi-
Alabama state line. It covers an area of approximately 118 km2, and is bordered
by 6 other counties: Lowndes County to the east, Winston and Noxubee Coun-
ties to the south, Webster and Choctaw Counties to the west, and Clay County
to the north (Fig. 1). The county seat of Starkville is located 10 km northeast of
the geographic center of Oktibbeha County, at the junction of State Highway
25 and US. Highway 82. Mississippi State University (MSU) and the Mississippi
Agricultural and Forestry Experiment Station are located 2.5 km east of
Starkville (Fig. 1). Starkville is the largest town in Oktibbeha County, followed
by Maben and Sturgis (Brent 1973).
Climate
The climate of Oktibbeha County is warm and humid, and influenced by the
subtropical latitude, the extensive landmass to the north, and the warm tem-
peratures of the Gulf of Mexico (Brent 1973). Annual precipitation averages
141.86 cm, with monthly precipitation ranging from 8.23 cm in October to 15.24
cm in March (Table 1). Winter and spring are the wettest seasons of the year,
with fall being the driest. Snow is rare and remains on the ground for short du-
rations only (Brent 1973). Temperatures in Oktibbeha County range from an
average low of 5.2°C in January to an average high of 27.0°C in July (Table D.
The average annual number of frost-free days (above 0°C) is 226 (Brent 1973).
oma
694 BRIT.ORG/SIDA 20(2)
4 TENNESSEE
ARKANSAS oe
= Taio aber OKTIBBEHA COUNTY
‘ Trim Cane Creek
1 tin ge
=“ Hwy 82
Starkville
i. Mississippi
Ps ‘“ State
il
oa ™, at
Jackson 0 10 20 30 40
Sh JASPER CLARKE
CNBR
COPLAH SIMPSON
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Topography, Physiography, Geology, and Soils
Oktibbeha County is located within the Hilly Coastal Plain physiographic prov-
ince (Fenneman 1938), a predominantly pine (Pinus spp.)-covered region in the
southeastern United States. Elevation in the county ranges from 50 m as.l. in
the Tibbe Creek floodplain in the northeastern corner of the county to 180 m
a.s.l.in the southwestern corner of the county. The county is mapped on the 7.5
series U.S. Geological Survey topographic maps Artesia, Bluff Lake, Bradley,
Cedar Bluff, Crawford West, Double Springs, Longview, Maben, Pheba, Starkville,
Sturgis, and West Point. Oktibbeha county is drained into the Tombigbee River
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI 695
Taste 1. Climate data for Oktibbeha County, Mississippi, based on data from the U.S. Department of
Agriculture, Natural Resources Conservation Service, National Water and Climate Center, for West
station, 1961-1990
—
Temperature (°C)
Avg. daily Avg. daily Average Avg. rainfall
Month maximum minimum daily (cm)
January 10.7 -0.5 5.2 1331
February 13.4 1.6 74 12.40
March 18.4 6.2 12:3 15.24
April 23.6 11.0 17.3 14.43
May 27.6 15.4 21.5 11.76
June 31.3 19.5 254 9.52
ly 32.8 21.3 27.0 1255
August 32.6 20.7 26.6 8.58
September 29.6 17.6 23.6 9.78
O 24.2 10.7 17.4 8.23
November 18.4 6.2 1253 10.80
December 12.8 1.5 72 15.06
Total: 22.9 10.9 16.9 141.86
by two major drainage systems: the Tibbe Creek system, which includes Sun
Creek, Trim Cane Creek, Sand Creek, and Catalpa Creek, drains the northern
half of the county; in the southern part of the county, the Noxubee River and
its tributaries drain southeastward into the Tombigbee River (Fig. 1, Brent 1973).
Oktibbeha County is situated among three physiographic regions (Fig. 2).
The eastern part of the county extends into the Black or Northeastern Prairie
Region. Vegetation in this region is influenced by alkaline soils derived from
underlying chalk of Upper Cretaceous age, including Demopolis, a bedded chalk
consisting primarily of calcite, with small additions of montmorillonitic clay,
mica, and sand; and Prairie Bluff, a mixture of bedded chalk and calcareous
sandstone (Brent 1973). To the west, the Black Prairie is bordered by the Pontotoc
Ridge; here, the Prairie Bluff chalk is, on the highest points, overlain with red
clays derived from the Ripley formation, also of Upper Cretaceous age and made
up of calcareous sands and clays (Brent 1973). The western portion of the county
lies within the Interior Flatwoods, a 5 to 20 km wide band of level to slightly
hilly lands (Hilgard 1860). This region was formed from geologic formations of
Tertiary age, which include Clayton, a heterogenous formation consisting of
sand, calcareous sandstone, chalk conglomerate, clay, or marl; Porters Creek,
made up of montmorillonitic clays; Wilcox, a formation consisting of quartz
sand, silty clay, bauxitic clay, and clay-ball conglomerates; and Naheola. In ad-
dition to these substrates, recent alluvial deposits, as well as older terrace de-
posits, may be found in the stream valleys and flood plains (Brent 1973).
BRIT.ORG/SIDA 20(2)
.
3
W- -E
l
| 6
S
4
7
| ale
SCALE vn)
0 50 100 150
Fic. 2.M ray + 4 . = 7 1 - ee ie fall kL L os y J AAC J ippi physiog | gi 4
Tannecsee River Hills 2 Black Pr irie 3 =P toc Ri 4 ior Fl North Central Plateau 6 = Loess
Bluff
Hills 7 = Mississippi Alluvial Valley vaio: Mississippi Delta) i — Sacksoi Prairie 9 = Longleaf Pine Belt 10 =
Coastal Pine Meadows (after Lowe 1921).
The soils found in Oktibbeha County have been grouped into 10 soil asso-
ciations (Brent 1973). Three associations consisting of nearly level, somewhat
poorly to moderately well-drained soils, occur in floodplains in the Black Prai-
rie (Leeper-Marietta-Catalpa) and the Interior Flatwoods (Mathiston-Urbo,
Mantachie-Mathiston-Ochlockonee) physiographic regions. The remaining 7
associations are found on uplands. The most prominent upland soil association
in the Black Prairie is the Kipling-Savannah-Oktibbeha Association, which
covers 21% of the county and consists of somewhat poorly to moderately well
drained soils with clayey subsoils or moderately well-drained soils with loamy
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI 697
subsoils and a fragipan. The Kipling-Sumter-Gullied Land Association, made
up of somewhat poorly drained to well-drained soils with clayey subsoils de-
veloped from chalk that are severely gullied in some areas, is another interest-
ing soil association of the Black Prairie: although it covers only 4% of the county,
it is of botanical interest because of the chalk outcrops it includes. Prominent
upland soil associations of the Interior Flatwoods physiographic province in-
clude the Longview-Falkner-Prentiss Association, which consists of somewhat
poorly to moderately well-drained soils with loamy to clayey subsoils and cov-
ers 25% of the county, and the Stough-Prentiss-Myatt Association, which is made
up of poorly to moderately well-drained soils with loamy subsoils and covers
8% of the county. The narrow ridgetops and sidelopes in the extreme western
portion of Oktibbeha County are dominated by the Maben-Ruston-Savannah
Association, consisting of moderately well- to well-drained soils with loamy
subsoils (Brent 1973).
Presettlement Vegetation
The first white settlers did not arrive in the area of present-day Oktibbeha
County until 1820, when Presbyterian missionaries built a mission in Mayhew.
Oktibbeha County was created through an act of the state legislature on 23
December 1833, after the land had been acquired from the Choctaw Indians in
the Treaty of Dancing Rabbit Creek on 27 September 1830 (Brent 1973). By the
1840 census, 4,276 people lived in Oktibbeha County, of which 1,702 were en-
gaged in agriculture (U.S. Census Bureau 1840). The best available information
on pre-and early settlement vegetaion in the area comes from Hilgard’s (1860)
seminal report on the phytogeography of Mississippi, which was based on his
travels through the state in the early to mid-1850's. Although Hilgard (1860)
did not visit Oktibbeha County, his comments on the Prairie Belt of neighbor-
ing Lowndes County, as well as on the Flatwoods of neighboring Chickasaw,
Calhoun, and Choctaw Counties (which, at the time, included present-day Clay
and Webster Counties) provide a reasonable estimate of Oktibbeha County's
early settlement vegetation. At the time, Oktibbeha County had a population
of 9,171, with 20% of its land area (23,655 ha) in improved farmland, distrib-
uted among 560 farms (U.S. Census Bureau 1850). However, it is likely that most
of this farmland, and therefore, most of the population, was concentrated in
the Black Prairie part of the county, with the Interior Flatwoods “.. thus far but
thinly settled ...” (Hilgard 1860:281).
Based on Hilgard’s observations, it is apparent that presettlement vegeta-
tion of the Black Prairie was dominated by grasslands, which were “... level, or
very gently undulating tracts, possessing a deep black, heavy soil, on which
timber is very much scattered or altogether wanting...” (Hilgard 1860:261). Chalk
outcops were already an important component of the landscape at the time, “..
forming ‘bald prairies and ‘bald hilltops—in which the limestone is too close
—
698 BRIT.ORG/SIDA 20(2
to the surface to allow of the growth of trees or other deep rooted plants, and
not unfrequently forms white areas many acres across in extent, strewn with
ossils (especially oysters) washed out of the mass, and only here and there a
patch of Verbena [probably V. bipinnatifida and V. simplex], or Cassia (C.
obtusifolia, occidentalis, marilandica).” (Hilgard 1860:77). Interspersed with
these tracts of open prairie and chalk outcrops were “.. tracts of a more rolling
surface, mostly with shallow, pale, light soil, timbered with the common up-
land Oaks—Spanish (“Red”) [Quercus falcata], Post [Q. stellata], Black Jack [Q.
marilandica], and sometimes Red and Black (“Black”) [Q. velutina] and Scarlet
(“Spanish”) Oak [Q. coccinea)...” (Hilgard 1860:261). Hilgard’s 1860) statements
regarding Black Prairie vegetation are echoed by Lowe (1921:32): “The region
has a gently rolling surface, and was originally prairies, having only here and
there scattered patches of trees, except on the stream bottoms, which supported
heavy growths of timber. This Prairie Belt is now largely in cultivation, but some
timber remains in the bottoms.”
Little can be said regarding the presettlement vegetation of the Pontotoc
Ridge in Oktibbeha County. Hilgard (1860) crossed the Pontotoc Ridge far to
the north of Oktibbeha County, so his observations may not be entirely repre-
sentative. However, speaking of the Pontotoc Ridge in general, he referred to it
asa“... narrow belt of fertile land underlaid [sic] by marl beds” (Hilgard 1860:266),
and further commented on the variable nature of the Pontotoc Ridge vegeta-
tion, which ranges from open prairie-like communities in calcareous areas to
ae
mesophytic upland hardwood forests on moist, north-facing exposures and xe-
ric upland hardwood forests on sandstone ridges. Lowe (1921:35) noted that the
red soils of the Pontotoc Ridge “... support a rich growth...” and that “[t]he plants
of the Pontotoc Ridge present decided differences from those of the prairies ly-
ing to the east, and the tertiary flatwoods on the west.”
=
rh
Presettlement vegetation of the Interior Flatwoods in the western part o
Oktibbeha County likely consisted of upland hardwood forest dominated by
Quercus stellata Wangenh., Q. marilandica Muenchh., and Pinus spp., with
Liriodendron tulipifera L. common on sideslopes and in hollows. Vaccinium
spp. was abundant in the understory; herbacious ground cover was sparse. Mov-
ing east and south into the Interior Flatwoods, these upland species were gradu-
ally replaced by Quercus alba L., Q. falcata Michx., Q. rubra L., and Carya spp.,
with the ground “... covered witha fine growth of grass.” (Hilgard 1860:280-282).
Land Use, Present-day Vegetation, and Disturbance
Prior to 1923, most (> 75%) of Oktibbeha County had been farmed at one time
or another, with the main crop being cotton (Gossypt umsp.; U.S. Census Bureau
1920, Brent 1973). As recent as 1964, 67% of the county’s land area was farm-
land (Brent 1973); today, in part due to such government programs as the Con-
servation Reserve Program (U.S. Department of Agriculture [USDA] Farm Ser-
camel
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI 699
vice Agency) and the Stewardship Incentives Program (USDA Forest Service),
forestry and recreation have supplanted agriculture as the primary land uses
in the county. As of 1997, there were 329 farms covering 35,000 ha, or 30% of
Oktibbeha County (USDA National Agricultural Statistics Service [NASS], http:/
/www.nass.usda.gov). On the other hand, forest land now makes up 64% of the
county, with 28% pine, 58% hardwood, and 14% mixed forest (Mississippi For-
estry Commission, http://www.mfc.state.ms.edu). The John W. Starr Memorial
Forest and Noxubee National Wildlife Refuge (NWR), which represent exten-
sive areas of forested land and provide a variety of recreational opportunities,
are partially located in Oktibbeha County. The remainder of the county is ur-
ban land.
With the exception of the John W. Starr Memorial Forest and Noxubee
NWR, most areas of the county have been heavily impacted by anthropogenic
disturbance, including agriculture and forestry, and the condition of the veg-
etation is correspondingly poor. Disturbance from agriculture is most evident
in the Black Prairie physiographic region, with much of the native prairie veg-
etation having been replaced by crop- and pasture land. Despite the fact that a
sizeable percentage of land has been taken out of cultivation over the last de-
cades, these lands have shown a tendency to grow up in trees rather than re-
turning to their original grassland vegetation (Lowe 1921, Leidolf @ McDaniel
1998). Furthermore, the few remnant prairies that remain today are subject toa
combination of anthropogenic and natural disturbances, including exotic spe-
cies invasions, recreation, and erosion.
Disturbance from forest operations is a major factor affecting the vegeta-
tion of the Interior Flatwoods. This is especially true for privately owned forest
land, where oe appears to be driven primarily by economic consid-
erations t t ften severe, with minimal rotation lengths.
On ee aes W. Start vee ial Forest and a NWR, forest sanagemen!
—
Management Practices that attempt to minimize the effect of ees operations
on land condition.
METHODS
We made vascular plant and bryophyte collections from February 1994 to 1996.
Collecting trips were conducted ona regular basis, at an average rate of one trip
per week, covering all parts of Oktibbeha County. We processed all collections
using standard field and herbarium techniques (Blake 1932; Smith 1971; Jones
& Luchsinger 1986). Field notes included information on habitat, species asso-
ciation, relative abundance, and natural history/ecology. Whenever possible,
we collected duplicate vouchers, except in the case of rare species. A complete set
of voucher specimens has been deposited at IBE, with partial sets distributed to
700 BRIT.ORG/SIDA 20(2)
BRIT, FR, UTC, and the Mississippi State University College of Forest Resources
Herbarium (MSUSFR).
After completion of field work, we conducted a complete herbarium re-
view of existing records at IBE and MISSA. All other herbaria in the state were
judged to have too few specimens from Oktibbeha County to warrant close at-
tention. Specimens from Oktibbeha County were verified and annotated. Where
available, we incorporated label information on physiographic region, habitat
of occurrence, and relative abundance into our annotated checklist. We exam-
ined 710 specimens in this manner. We also reviewed pertinent literature and
published checklists for records from Oktibbeha County and, after critical
evaluation, incorporated them into the checklist. Since the specimens upon
which the reports are based were usually not seen, only the reference is provided.
Sources used for identification and verification of specimens, as well as for
nomenclatural reference, included Allen (1992), Bailey and Bailey (1976),
Barkworth et al. (In preparation), Britton and Brown (1970), Burkhart (1969),
Correll and Johnston (1970), Cronquist (1980), Davenport (1988), Godfrey and
Wooten (1979, 1981), Great Plains Flora Association (1986), Hitchcock (1951),
Isley (1990), Krtissmann (1977), Lellinger (985), Morris (1989), Radford et al.
(1968), Shinners (1962), and Steyermark (1963). Where used, herbarium acro-
nyms follow Holmgren et al. (1990). Abbreviations of botanical journals follow
Bridson and Smith (1991).
RESULTS AND DISCUSSION
General
We documented 1,148 plant taxa (1,125 species, 7 hybrid taxa, and 16 infraspe-
cific taxa) from Oktibbeha County, belonging to L60 families and 514 genera.
Of these taxa, 976 (85%) are native (Table 2). Seventeen families are represented
by 5 genera or more (Table 3); 46 genera are represented by 5 taxa or more (Table
4+). Noteworthy collections include 3 introduced species documented ina natu-
ralized state from Mississippi for the first time: Ilex cornuta Lindl. & Paxton,
Mahonia bealei (Fortune) Carriére, and Nandina domestica Thunb. [The last
species has since been reported from Amite County, Mississippi (Alford 2001).
All 3 species are used frequently for landscaping by MSU and were encoun-
tered in a mesophytic upland hardwood forest in the Pontotoc Ridge physi-
ographic region adjacent to campus. Given the common characteristics of their
seeds (all 3 species produce fleshy berries readily consumed by birds|Dirr 1990),
animals, specifically birds, represent the most likely disperal vector.
Compared to 3 other counties in east-central Mississippi, Oktibbeha County
has the second largest documented vascular flora. It has 17% and 11% more vas-
cular plant taxa than Attala (Winstead 1990) and Grenada County (Morris 1987),
respectively, and 14% less vascular plant taxa than Monroe County (MacDonald
1996). Compared to these 3 counties, Oktibbeha County has the highest species
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI 701
Table 2. Taxonomic summary for the checklist of plants of Oktibbeha County, Mississippi
Species and lesser taxa‘
Division/Class? Families? Genera? Native Introduced ota
Bryophyta® 12 14 22 0) 22
Hepaticopsida 11 13 21 0 2]
Anthocerotopsida 1 1 1 0 1
Equisetophyta 0) 0 0) 0 0
Lycopodiophyta 2 3 5 ) 5
Polypodiophyta 10 12 14 ] 15
Pinophyta 3 3 4 0 a
Magnoliophyta 133 482 931 171 1102
Magnoliopsida 105 368 624 114 738
Liliopsida 28 114 307 57 364
Total 160 514 976 172 1148
® Nomenclature follows Flora of North America Editorial Committee (1993).
» Counts are based on families and genera as recognized by Brummitt (1992).
“Counts are based on species and lesser taxa as recognized by Kartesz (1994) and Barkworth et al.
(In preparation).
“ Bryopsida was not included in this study.
Taste 3. Plant families? in Oktibbeha County, Mississippi, represented by 25 genera? (in decreasing
order of importance; followed by number of species and lesser taxaS in parentheses).
seas 57 (140) Lamiaceae-14 (24) Cyperaceae-8 (100) Euphorbiaceae—5 (19)
Poacea a 59) Brassicaceae-13 (20) Boraginaceae—6 (10) Malvaceae-5 (7)
oe (75) Rosaceae-13 (28) Caryophyllaceae-6 (8)
Se ein en. (34) Rubiaceae-10 (22) Ranunculaceae-7 (21)
Apiaceae-19 (24) Orchidaceae-9 (17) — Caprifoliaceae—5 (6)
* Nomenclature follows Brummitt (1992).
> As recognized by Brummitt (1992).
‘As recognized by Kartesz (1994) and Barkworth et al. (In preparation).
richness in 5 out of 20 representative flowering plant families; for an additional
ll families, Oktibbeha couny ranks second in species richness (Table 5). The
1,103 species of vascular plants documented from Oktibbeha County comprise
38.7% of all vascular rere species known to occur in Mississippi (USDA, Natural
Resources Conservation Service INRCS] Plants database, http://plants.usda.gov).
Rare/Protected Species
Oktibbeha County features a large number of rare and protected species. Sixty-
seven species are currently listed as sensitive (MNHP 2002a, 2002b; Table 6),
far exceeding the number of state-listed species reported from 3 other counties
in east-central Mississippi (Table 5). Statewide, Oktibbeha County ranks 4th in
the number of sensitive species, after Jackson, Tishomingo, and Perry Counties
702 BRIT.ORG/SIDA 20(2)
Taste 4. Plant genera? in Oktibbeha County, Mississippi, that are represented by =5 taxa? (in decreas
ing order of importance).
Carex-57 Carya-10 Silphium—7 Lactuca-5
Dichanthelium-24 Hypericum—10 Smilax-7 a-
Juncus—17 Polygonum-9 Spiranthes—7 Rhus—5
Quercus—17 Trifolium—9 Sporobolus—7 Rhynchospora-5
Cyperus-15 Asclepias-8 Verbena-7 Rubus-5
Paspalum-15 Galium-8 Agalinis—6 Rumex-5
ster-14 Helianthus-8 Fimbristylis—6 Scirpus—5
Solidago-12 Panicum-8 Ludwigia-6 Sisyrinchium—5
Desmodium-11 Crataegus—/ /icia—6 Vaccinium-5
Eragrostis—11 Eleocharis—7 Amaranthus—5 Viola—5
Eupatorium-11 Lespedeza—/ Chamaesyce—5
Ranunculus-11 Oenothera-7 Euphorbia-5
*Nomenclature follows Brummitt (1992).
» As recognized by Kartesz (1994) and Barkworth et al. (In preparation).
(MMNS database, June 2002). Four species are also listed as either threatened
or rare by the USDA Forest Service (Kral 1983, Table 6); of these, Apios priceana
B.L. Rob. is currently the only species listed as federally threatened (5 January
1990, 55 FR 433) under the Endangered Species Act (16 USCS §1531 et seq.). Nine-
teen species in the families Araliaceae, Cactaceae, and Orchidaceae are also af-
forded special protection as Appendix II species under the Convention on In-
ternational Trade in Endangered Species of Wild Flora and Fauna (27 UST 1087;
TIAS No. 8249; Table 6). The most significant discovery in the category of rare
and protected species is that of asmall population of Alisma subcordatum Raf.
at Oktibbeha County Lake, a plant species previously considered to be of only
historical occurrence in Mississippi.
Plant Communities
We recognized 16 different plant communities that can be grouped into 5 broad
categories: bottomland forests, upland forests and prairies, aquatic communi-
ties, seepage areas, and human-influenced communities.
BOTTOMLAND FORESTS
Bottomland forests are formed by alluvial processes of erosion and sediment
deposition that result in diverse topographical formations such as ridges,
sloughs, flats, and terraces. Topographical variation results ina variety of flood-
ing regimes; thus, small differences in topography effect large differences in
edaphic conditions, and the resulting differences in plant associations are ob-
vious. The two main types of bottomland forest we recognized, bottomland
hardwood forest and swamp forest, owe their respective plant communities
primarily to differences in flooding regime. Because Oktibbeha County lacks
major streams, the extent of bottomland sites is limited.
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI 703
Taste 5. Species richness of selected taxonomic groups in 4 counties of east-central and north-
central Mississippi.
Taxon? Attala Co.” Grenada Co.‘ Monroe Co.! Oktibbeha Co.
Bryophyta 91 N/A‘ 5! N/A\
Anthocerotopsida ] N/A® ] ]
Hepaticopsida 24 N/A® 12 2]
Bryopsid 66 N/A‘ 38 N/A\
Equisetophyta ] 2 1 0
Lycopodiophyta 4 2 4 5
Polypodiophyta 19 19 19 15
Pino a 5 4 4 4
Magnoliophyta 916 969 1253 1079
Magnoliopsida 643 661 871 731
Apiaceae 17 22 32 2
Asteraceae 107 116 156 137
Brassicaceae 9 10 21 20
Ericaceae 9 7 12 7
Euphorbiaceae 12 1] 2] 19
Fabaceae 67 70 85 74
Fagaceae 23 21 24 18
Hypericaceae 10 9 13 12
Juglandaceae 10 12 12 11
Lamiaceae 22 24 28 23
Polygonaceae 14 15 is 1
Ranunculaceae 13 12 25 20
osaceae 34 27 40 28
Rubiaceae 16 15 22 22
Scrophulariaceae 24 26 29 34
Lilliopsida 273 308 382 348
Cyperaceae 79 90 112 98
Juncaceae 12 14 18 19
Liliaceae (sensu lato?) 27 23 32 36
Orchidaceae 16 19 17 17
Poaceae 98 12] 146 146
Total: 1036 N/A® 1332 N/AT
Total (vascular only): 945 996 1281 1103
State-listed species?: 13 42 50 67
Nomenclature follows Brummitt (1992), Flora of North America Editorial Committee(1993), and
Barkworth et al. (In preparation).—»Winstead (1990).—‘Marris (1987).—*MacDonald ae
not assess Bryophyta; therefore, a comparison can not be made with other counties—'Did no
assess Bryopsida; therefore, a comparison can not be made with other counties. ee
Alliaceae, Se es oes Convallariaceae, Hemerocallidaceae, Hyacinthaceae,
Hypoxidaceae, Liliac milacaceae, and Trilliaceae —" Mississippi Natural Heritage
Program (2002a, ae Mississippi ae of Natural Science database June 2002)
704 BRIT.ORG/SIDA 20(2)
Taste 6.Rank/status of protected plant species known to occur in Oktibbeha County, Mississippi
MNHP Rank?
Species* State Global — ESA‘ CITES? ~—sUSDA®
Aesculus glabra Willd. $2? G5
Agalinis pseudaphylla (Pennell) Shinners S2 G1G2 7
Alisma subcordatum Raf. SH G4G5
Amphiachyris dracunculoides (DC.) Nutt. S] G4G5
ae be ceana B. ba Rob. S] G2 qT R
is Gray) Al-Shehbaz &Bat S182 G4?
Asarum canad $253 G5
Asclepias hirtella Pennell) Woodson $2 G5
Asclepias purpurascens L. S| G4G5
Aster ericoides $2 G5
Astragalus canadensis L. var. canadensis 52 G5
Boutelou a curtipendl Aer ) Torr.in Marcy S3S4 G5
scilloides (Raf.) Cory $2S$3 G4G5
ea aeen: Steud. $283 G5?
Carex jamesii Schwein. $182 G5
Carex laxiflora Lam. S1 G5?
Carex meadii Dewey 5354 G4G5
Carex microdonta Torr. & Hook. $2? G4
Carex stricta Lat 52 G5
Carya laciniosa (F. Michx.) G. Don S283 G5
Carya leiodermis Sarg. $253 G5
Coelorachis cyli a icd (Michx.) Nash 5] G4G5
Corallorhiza wisteriana Conrad I|
Crataegus calpodendron (Ehrh.) Medik. S? G5
Dasistoma maeiopinila (Nutt.) Raf. S354 G4
Delphinium tricorne Mic hx S2 G5
Dodecatheon m di iL. S2 G5
ou elas dalle (L.) Moench S354 G4
m albidum Nutt. S2 G5
Euan prolifera Nutt. ex DC, S1 5
nymus atropurpureus Jacq. $253 G5
ee liniensis Walter $283 G5
Fraxinus quadrangulata Michx. $2 G5
federlea sa om oo Barnhart S2 G5 lI
F.Forst.) Spreng. $2 G5
Nine peo us Buckley S354 G5
Lilium superbum L S354 G5
Linum sulcatum Riddell $354 G5
Listera australis Lindl, II
Lobelia appendiculata A.DC. $253 G4G5
Malaxis unifolia Michx II
ermum ¢ scence L. $354 G5
unl eee slabrifora Scribn. S? G4?
Nemastylis geminiflora Nutt. $2 G4
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI
Species?
MNHP Rank?
State Global — ESAS CITES’ ~—sUSDA®
Neptunia lutea (Leavenw.) Benth.
Oenothera triloba Nutt.
Onosmodium molle Michx. ssp.
Pela imum (Mack.) B. Boivin
nelmoanni |
Pran
eee humi fia (Raf.) Raf.
Osmorhiza longistylis (Torr.) DC.
Panax quinquefo fius L
Penstemon tenuiflorus Pennell
id,
Perideridia americana (Nutt.ex DC.) Reichenb.
Platanth liaris (L.) Lindl.
Platanthera clavellata (Michx.) Luer
latanthera cristata (Michx.) Lindl.
Polytaenia nuttallii DC.
Ponthieva racemosa (Walt.) C.Mohr
Prenanthes aspera Michx
Ptelea trifoliata L.
ercus Dao cipg Michx.
lalaal ic la lata
rsh
Sedum nie Michx.
Spiranthes cernua (L.) Rich.
Spiranthes lacera (Raf.) Raf.
Spiranthes magnicamporum Sheviak
Spiranthes ovalis Lindl.
Spiranthes praecox (Walt.) S.\Watson
oy! iranthes tuberosa Raf.
Spiranthes vernalis Engelm. &A. Gray
Staphylea trifolia
Taenidia integerrima (L.) Drude
Thalictrum debile Buckley
Tipularia discolor (Pursh) Nutt.
ese a ol (Michx.) Raf.
ml.
=
Ulmus serotina Sar
Uvularia floridana Chapm.
Liga Sea (Sw.) Rydb.
arg.
S354 G5
SU G4
R
S] G5
II
S3 G5
G364 II
$283 G3?
S182 G4
II
II
$3 G5 II
S2 G5
$2? G4G5 II
S2 G4?
$354 G5
S2 G5
$2 G5
S1 G5
II
$354 G5 II
$283 G4 II
S283 G5? II
II
II
II
53 G5
51 G5
S182 G2 R
II
S] G3
*#Nomenclature follows Kartesz (1994) and Barkworth et al. (In preparation).— Mississippi Natural
Heritage Program (2002a, 2002b
oe
rank: Smee) imperiled cman
a
1), imperiled cain (G2),
rare and local throughout range, found locally ina
restricted FalNQe,O
ulnerable t LOLS tit n (G3),
apparently secure globally, but may be rare in parts of its range (G4), paar secure global
(G5), with?” indicating uncertainty about or inexactness of a rank. State (S) ranks are
ranks, except in that “state” is cee for “global” with SH, SU, and S? ea nicionealo occur-
ly.— e ne
rence but suspected to be e
fig el per ae
ant, uncertain rank, and unranked, respective
cies Act, 16 USCS §1531 et seq. i threatened species
Lea species likely to become endanger
within the foreseeable future throughout all or a significant portion of its range—4Convention on
706 BRIT.ORG/SIDA 20(2)
1. Bottomland hardwood forests.—Bottomland hardwood forests in
Oktibbeha County are found primarily in the Interior Flatwoods physiographic
region, where they occur on seasonally flooded sites associated with creeks and
minor streams. The high foliage volume of these forests gives credence to their
reputation for high productivity. Depending on mode of establishment and dis-
turbance history (e.g., logging), bottomland hardwood forests in Oktibbeha
County may have multiple layers, including overstory, midstory (subcanopy),
and woody and herbaceous understory layers (the latter may be weakly devel-
oped or vacking in forests subject to recent logging); however, due to variations
in tree height and extensive growth of such lianas as Rhus radicans L., Bignonia
capreolata L., Ampelopsis spp., Parthenocissus quinquefolia (L.) Planch., Vitis
spp., and Smilax spp., stratification may not be readily apparent at any given
site. In addition to the vegetation layers already mentioned, unique and locally
important features include palmetto [Sabal minor Jacq.) Pers] thickets and dense
cane [Arundinaria gigantea (Walter) Muh] brakes.
The overstory of bottomland hardwood forests is generally dominated by
oaks (Quercus spp.) with species composition depending on moisture. From wet
to dry, dominants include Q. lyrata Walter, Q. michauxii Nutt. Q. nigra L., Q.
phellos L.,and Q. pagoda Raf. Co-dominants include Q. alba L., Q. falcata Michx.,
OQ. laurifolia Michx., Q. rubra L., Q. shumardii Buckley, Q. stellata Wangenh., Q.
texana Buckley, Fagus grandifolia Ehrh., Liquidambar styraciflua L., Carya
carolinae-septentrionalis (Ashe) Engl. & Graebn., C. cordiformis (Wangenh.) K.
Koch, C. glabra (Mill) Sweet, C. ovalis Wangenh.) Sarg., C. ovata (Mill) K. Koch,
C. tomentosa (Lam. ex Poir.) Nutt.,and Liriodendron tulipifera L., noteworthy is
Carya leiodermis Sarg,, which is listed as imperiled to rare and local in the state
(S283, MNHP 2002a). Common intermediate/midstory trees include Acer
rubrum L., Nyssa sylvatica Marshall, Celtis laevigata Willd., Ulmus americana
Land U. alata Michx. Examples of small trees/large shrubs include Asimina
triloba Dunal, Ilex decidua Walter, Carpinus caroliniana Walter, Ostrya
virginiana (Mill.) K. Koch, Rhododendron canescens (Michx.) Sweet, Aesculus
pavia L., Hamamelis virginiana L., Lindera benzoin (L.) Blume, Sassafras
albidum (Nutt.) Nees, Amelanchier arborea (F. Michx.) Fernald, Crataegus
viridis L., and Planera aquatica J.- Gmel; noteworthy records include Ptelea
trifoliata L. and Staphylea trifolia L., which are listed as rare and local to ap-
arently secure (S2S3) and rare and local (S3) in the state, respectively (MNHP
2002a, 2002b). The understory consists of a mixture of small shrubs, includ-
i
en
Table 6 legend (continued)
International Trade in Endangered Species of Wild Fauna and Flora, 27 UST 1087; TIAS No.8249. 1] =
Appendix Il species, .e., (a) a species which, although not necessarily threatened with extinction,
may become so unless trade is subject to strict regulations, or (b) a species which must be sHIBIEEE
to regulation in order that trade in certain specimens of species referred to under (a) may be brought
under effective control, i.e., species similar in a *USDA Forest Service Technical Publi-
cation R8-TP2 (Kral 1983). T = threatened, R =
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI 707
ing Cornus stricta Lam., Vaccinium spp., Hydrangea quercifolia W. Bartram, and
Itea virginica L; and herbs, including such ee as Botrychiu m biternatum (Sav.)
Underw. and Onoclea sensibilis L., such forbs as Thaspium trifoliatum (L.) A.
Gray, Dicliptera brachiata (Pursh) Spreng., Amsonia tabernaemontana Walter,
Aristolochia tomentosa Sims, Pluchea camphorata (L.) DC., Solidago caesia L.,
Lobelia cardinalis L., Specularia biflora (Ruiz & Pav.) Fisch. @ C.A. Mey.,
Desmodium canescens (L.) DC., D. cuspidatum (Muhl. ex Willd.) DC. ex Loudon,
Vicia caroliniana Walter, Triadenum spp., Collinsonia tuberosa Michx., Lycopus
virginicus L., Spigelia marilandica L., Ludwigia spp., Polygonum virginianum
L., Epifagus virginiana (L.) Baton, Gratiola spp., Arisaema dracontium (L.)
Schott, Commelina virginica L., Tradescantia virginiana L., Maianthemum
racemosum (L.) Link, and Dioscorea villosa L., and such graminoids as Carex
abscondita Mack., C. annectens (E.P. Bicknell) E.P. Bicknell, C. caroliniana
Schwein., C. corrugata Fernald, C. flaccosperma Dewey, C. leavenworthii Dewey,
C. longii Mack., C. louisianica L.H. Bailey, C. reniformis (L.H. Bailey) Small, C.
typhina Michx., Cinna arundinacea L., Dichanthelium commutatum (Schult.)
Gould, Leersia oryzoides (L.) Sw, L. virginica Willd., Panicum rigidulum Bosc
ex Nees in Mart., and Paspalum repens PJ. Bergius. Pleopeltis polypodioides (L.)
E.G. Andrews & Windham var. michauxiana (Weath.) E.G. Andrews &
Windham in Windham isa common epiphyte. Noteworthy understory species
listed as sensitive (MNHP 2002a, 2002b) include Dasistoma macrophylla Nutt.)
Raf. (rare and local to apparently secure-S3S4), Lilium superbum L. (S3S4),
Muhlenbergia glabriflora Scribn. (uncertain status-S?), Spiranthes ovalis Lindl.
(imperiled to rare and local-S2S3), and Triphora trianthophora (Sw.) Rydb.
(S283),
In the Black Prairie physiographic region, intensive agriculture has reduced
once extensive bottomland hardwood forests to narrow gallery forests along
small streams. Here, noteworthy species include Quercus nigra L., Q. durandii
Buckley, Carya tomentosa (Lam. ex Poir.) Nutt., and C. laciniosa (EF Michx.) G.
Don, which is listed as imperiled to rare and local in the state (S253, MNHP
2002a), in the overstory, with Acer negundo L., Morus rubra L.,and Ulmus rubra
Muhl. as intermediates. Shrubs include Asimina triloba (L.) Dunal, Aesculus
glabra Willd., which is listed as possibly imperiled in the state (S2?, MNHP
2002a), A. pavia L.,and Prunus americana Marshall; Zizia aurea (L.) WDJ. Koch
and Eupatorium rugosum are common herbaceous species.
2. Swamp Forests.—Swamp forests are permanently /semi-permanently in-
undated, and occur along the shallow margins of existing stream channels and
old oxbow lakes that have been cut off from the main channel and have been
partially silted in. Because of the small size of streams in Oktibbeha County,
swamp forests are fairly limited in extent, with shallow water depths. Further-
more, they are restricted to the Interior Flatwoods physiographic region; the
Black Prairie lacks streams of sufficient size. Frequency and duration of flood-
—
_—
708 BRIT.ORG/SIDA 20(2)
ing limits the species that can occur in swamp forests to extremely water-toler-
ant species. Germination for most species can only occur during periods of low
water when the soil is exposed. Thus, in contrast to bottomland hardwood for-
ests, swamp forests have a fairly sparse foliage profile. The overstory is domi-
nated by Quercus lyrata Walter and, occasionally, Taxodium distichum (L.) LC.
Rich. Where present, water-tolerant shrubs and small trees (e.g., tea virginica
L. and Vaccinium arboreum Marshall) grow in small patches or singly and
sparsely distributed. Because of the presence of standing water, ground vegeta-
tion is often absent, or may consist of small patches or scattered individuals of
Justicia ovata (Walter) Lindau var. lanceolata (Chapm.) R.W. Long, Bidens
discoidea (Torr. & A. Gray) Britton, Polygonum densiflorum Meisn., Saururus
cernuus L., Gratiola virginiana L., Pilea pumila (L.) A. Gray, Carex intumescens
Rudge, C. joorii L.H. Bailey, and Leersia lenticularis Michx.
UPLAND FORESTS AND PRAIRIES
L Mesophytic upland hardwood forests.-Mesophytic upland hardwood forests
are typically found in the Pontotoc Ridge physiographic region (see Morris et
al. 1993); in the Interior Flatwoods, they rarely occur in the transition zone be-
tween upland pine/pine-mixed hardwood forests and bottomland forests; in
the Black Prairie, they rarely occur in association with streams. Mesophytic
upland hardwood forests are among the most diverse plant communities in
Oktibbeha County; they are also home to some of the rarest plants in the county,
featuring 18 species listed by the MNHP (2002a, 2002b), with many of them
occurring only in this community. Mesophytic upland hardwood forests are
characterized by an overstory of mixed deciduous hardwoods, including Acer
barbatum Michx., Fagus grandifolia Ehrh., Quercus alba L., Q. muehlenbergii
Engelm., Q. shumardii Buckley, Q. stellata Wangenh., Carya carolinde-
septentrionalis (Ashe) Engl. & Graebn., C. myristiciformis (F Michx.) Nutt,
Juglans nigra L., Liriodendron tulipifera L., and Platanus occidentalis L., note-
worthy overstory species are Fraxinus quadrangulata Michx. and Ulmus
serotina Sarg., which are listed as imperiled (S2) and possibly rare and local
(S3?) in the state, respectively (MNHP 2002a). Conifers are absent from this com-
munity. Common under- and midstory trees and shrub include Asimina triloba
(L.) Dunal, Viburnum rufidulum Raf., Ostrya virginiana (Mill) K. Koch, Cornus
florida L., Rhododendron canescens (Michx.) Sweet, Aesculus pavia L., Tilia
americana L.,and Ulmus americana L. noteworthy mid- and understory asso-
ciates are Evonymus atropurpureus Jacq. and Crataegus calpodendron (Ehrh.)
Medik., which are listed as imperiled to rare and local (S2S3) and being of un-
certain status (S?) in Mississippi, respectively (MNHP 2002a). Alsocommon are
lianas, including Smilax bona-nox L., S. ldsioneura Hook., and S. pulverulenta
Michx. The herbaceous plant layer is characterized by such forbs as Sanicula
odorata (Raf.) K.M. Pryer & LR. Phillippe, Thaspium barbinode (Michx.) Nutt.
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI 709
Heliopsis gracilis Nutt., Podophyllum peltatum L., Cynoglossum virginianum L.,
Lithospermum latifolium Michx., Myosotis macrosperma Engelm., Cudamnine
concatenata (Michx.) Sw. Monotropa hypopithys L., Desmodium glutinosum (L.)
DC,, D. rotundifolium DC,, Vicia caroliniana Walter, Spigelia marilandica L.,
Oxalis violacea L., Sanguinaria canadensis L., Anemone virginiana L.,
Thalictrum thalictroides (L.) Eames & B. Boivin, Agrimonia rostellata Wallr,
Galium uniflorum Michx., Pedicularis canadensis L., Scrophularia marilandica
L., Physalis heterophylla Nees, Viola spp., Arisaema spp., Lilium michauxii Poir,
Tipularia discolor (Pursh) Nutt., and Trillium recurvatum L.C. Beck, including
such spring ephemerals as Claytonia virginica L., Allium canadense L.,
Nothoscordum bivalve (L.) Britton, Uvularia sessilifolia L.,and Hypoxis hirsuta
(L.) Coville; such graminoids as Carex albicans Willd. ex Spreng. var. albicans,
C.lupulina Muhl. ex Willd. C. striatula Michx., C. vulpinoidea Michx., Luzula
echinata (Small) FJ. Herm., Agrostis perennans (Walter) Tuck., Chasmanthium
latifolium (Michx.) H.O. Yates, C. sessiliflorum (Poir) H.O. Yates, Dichanthelium
boscii (Poir.) Gould & C.A. Clark, D. polyanthes (Schult.) Mohlenbr.,, Melica
mutica Walter, Panium anceps Michx., and Poa dutumnalis Muhl. ex Elliott; as
well as a diverse array of ferns and fern allies, including Selaginella apoda (L.)
Spring, Woodwardia areolata (L.) T.Moore, Polystichum acrostichoides (Michx.)
Schott, Botrychium virginianum (L.) Sw. Osmunda cinnamomeda L.,, O. regalis
L., Pleopeltis polypodioides (L.) E.G. Andrews & Windham var. michauxiana
(Weath.) E.G. Andrews & Windham in Windham, Athyrium filix-femina (L.)
Roth ex Mert. ssp. asplenioides (Michx.) Farw,, and Onoclea sensibilis L; note-
worthy species from this association listed by MNHP (2002a, 2002b) are Apios
priceana B.L. Rob. (critically imperiled-Sl, federally threatened), Asarum
canadense L. (imperiled to rare and local-S2S3), Carex gracilescens Steud. (S2S3),
C. jamesii Schwein. (critically imperiled to imperiled-S1S2), C. laxiflora Lam.
(SL), Dasistoma macrophylla (Nutt.) Raf. (rare and local to apparently secure-
S384), Frasera caroliniensis Walter (S283), Hexalectris spicata (Walter) Barnhart
(imperiled-S2), Hybanthus concolor (T.F. Forst.) Spreng. (S2), Menispermum
canadense L. (S354), Panax quinquefolius L. (rare and local-S3), Taenidia
intergerrima (L.) Drude (SD), Thalictrum debile Buckley (SIS2), and Triosteum
angustifolium L. (S3).
2. Xeric upland hardwood forests.—In Oktibbeha County, xeric upland hard-
wood forests occur on dry secondary sandstone ridges in all 3 physiographic
regions: on isolated hills scattered throughout the Black Prairie, on the Pontotoc
Ridge, and in the transition zone from the Interior Flatwoods to the North-cen-
tral Plateau physiographic region. Xeric upland hardwood forests in Oktibbeha
County feature a unique complement of floristic elements, many of which are
found in no other plant community in the county. In the Black Prairie, domi-
nant overstory species include Quercus marilandica Muenchh., Q.
muehlenbergii Engelm., Q. velutina Lam., Liquidambar styraciflua L., Carya
710 BRIT.ORG/SIDA 20(2)
myristiciformis (F Michx.) Nutt. and C. tomentosa (Lam. ex Poir.) Nutt; note-
worthy overstory species include Quercus macrocarpa Michx., which is listed
as imperiled in the state (S2, MNHP 2002a). Common trees, shrubs, and woody
vines of the mid- and understory are Campsis radicans (L.) Seem. ex Bureau,
Symphoricarposorbiculatus Moench, Cornus drummondiiC.A. Mey., Vaccinium
arboreum Marshall, Amorpha fruticosa L., Sassafras albidum (Nutt.) Nees,
Maclura pomifera (Raf.) C.K. Schneider, Crataegus engelmannii Sarg.,and Zan-
thoxylum clava-herculis L. Examples of characteristic herbaceous species are
Matelea gonocarpos (Walter) Shinners, Rudbeckia hirta L., Sagina decumbens
(Elliott) Torr. & A. Gray, Desmodium ciliare (Muhl. ex Willd.) DC., Anagallis
minima (L.) Krause, Ranunculus micranthus Nutt., Agrimonia rostellata Wallr.
Fragaria virginiana Duchesne, Galium pilosum Aiton, Carex blanda Dewey, C.
bushii Mack., C. umbellata Schkuhr ex Willd., Juncus tenuis Willd.,and Luzula
bulbosa (Alph.Wood) Smyth & Smyth; noteworthy associates are Erythronium
albidum Nutt. and Spiranthes lacera (Rat.) Raf., which are listed as imperiled
(S2) and rare and local to apparently seure (S354) in the state, respectively
(MNHP 2002a, 2002b). On the Pontotoc Ridge, characteristic woody species
include Campsis radicans (L.) Seem. ex Bureau, Lonicera sempervirens L.,
Quercus shumardii Buckey, Liquidambar styraciflua L., Carya myristiciformis
(F Michx.) Nutt., Sassafras albidum (Nutt.) Nees, Celtis laevigata Willd., and
Smilax lasioneura Hook., with Antennaria plantaginifolia (L.) Richardson, Eri-
geron st ks Muhl. ex Willd., Vicia minutiflora FDietr, Oxalis violacea L.,
Passiflora lutea L., Carex blanda Dewey, C.cephalophora Muhl. ex Willd., C. tri-
angularis Boeck, and Juncus tenuis Willd. being typical herbaceous species;
Echinacea purpurea (L.) Moench, listed as rare and al to apparently secure
in the state (S354, MNHP 2002a), is a noteworthy record from this association.
The overstory of xeric upland hardwood forests in the Interior Flatwoods physi-
ographic region is dominated by such species as Quercus alba L., Q. falcata
Michx.,Q. nigra L., 9. pagoda Raf.,Q. shumardii Buckley, Liquidambar styraciflua
L.,and Carya glabra (Mill.) Sweet; Campsis radicans (L.) Seem. ex Bureau, Sassa-
frasalbidum Nutt.) Nees, Crataegus spathulata Michx.,and Ampelopsis arborea
(L.) Koehne are common shrubs and vines of the mid- and understory. Ascl-
epias amplexicaulis Small, Matelea gonocarpos (Walter) Shinners, Euphorbia
corollata L., Lespedeza virginica (L.) Britton, Pycnanthemum tenuifolium
Schrad., Trichostema brachiatum L., Rhexia mariana L. var. mariana, Oxalis
violacea L., Passiflora lutea L., Aureolaria flava (L.) Farw,, Juncus tenuis Willd.,
Luzula bulbosa (Alph.Wood) Smyth & Smyth, Chasmanthium sessiliflorum
(Poir.) H.O. Yates, and Dichanthelium boscii (Poir.) Gould & C.A. Clark are typi-
cal herbaceous species.
3. Pine Forests and Pine-Mixed Hardwood Forests.—By and large, pine for-
ests and pine-mixed hardwood forests in Oktibbeha County are restricted to
the Interior Flatwoods physiographic region. The vast majority are intensively
~—
—
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI 711
managed pine plantations; naturally regenerated, unmanaged pine forests are
rare. Management activities associated with timber production, including pre-
scribed burns on variable rotations, pre-commer« ialand commercial thinning,
and herbicide injection of undesirable hardwoods, result in reduced vegetation
stratification and woody species richness in managed pine forests, and prevent
naturally occurring plant succession. As a result, species composition differs
significantly between managed and natural stands (Table 7). For example, natu-
ral pine or pine-hardwood forests, in addition to having a more diverse over-
story, usually feature a well-developed midstory consisting of a diverse array of
ardwoods, including Acer rubrum L., Carya tomentosa (Lam. ex Poir.) Nutt.,
Cornus florida L., Liquidambar styraciflua L., Nyssa sylvatica Marshall,
Diospyros virginiana L., Quercus spp., Sassafras albidum (Nutt.) Nees, Prunus
serotina Ehrh., and Ulmus alata Michx. (Table 7). Managed pine forests, on the
other hand, wile development of midstory vegetation is actively prevented,
have only two vegetation layers: an overstory composed almost exclusively of
Pinus taeda L. (and some P echinata Mill), and an understory that, much like
that of pine-mixed hardwood forests, is characterized by a diverse assemblage
of plants, including such graminoids as Dichanthelium acuminatum (Sw.)
Gould & CA. Clark ssp. lindheimeri (Nash) Freckmann & Lelong in ed., D.
laxiflorum (Lam.) Gould, Saccharum brevibarbe (Michx.) Pers. ssp. contortum
(Elliott) R.D. Webster, S.giganteum (Walter) Pers.,and Schizachyrium scoparium
(Michx.) Nash in Small, as well as the fire-adapted Andropogon gerardii Vitman
and Sorghastrum nutans (L.) Nash in Small; a variety of forbs, including Apo-
cynum cannabinum L.,, Asclepias tuberosa L., Aster solidagineus Michx., Bidens
aristosa (Michx.) Britton, Eupatorium spp., Euthamia tenuifolia (Pursh) Nutt.,
Liatris spicata (L.) Willd., Solidago spp., Chamaecrista fasciculata (Michx.)
Greene, Lespedeza spp., lephrosia virginiana (L.) Pers., Seymeria cassioides J.F
Gmel.) S.F. Blake, Aletris aurea Walter, and Spiranthes tuberosa Raf. such li-
anas as Rhus radicans L., Lonicera japonica Thunb., Gelsemium sempervirens
(L.) W.T. Aiton, Vitis rotundifolia Michx., Smilax glauca Walter, and S.
rotundifolia L. shrubs and small trees, including Aralia spinosa L., Vaccinium
spp., Hyperium hypericoides (L.) Crantz, Crataegus spp., and Rubus spp., and
tree seedlings, inluding Carya glabra (Mill.) Sweet and Quercus spp. (Table 7).
Noteworthy pine and pine-mixed hardwood forest associates include Asclepias
hirtella (Pennell) Woodson and Lobelia appendiculata A.DC,, which are listed
as imperiled (S2) and imperiled to rare and local (S253) in the state, respec-
tively (MNHP 2002a).
4, Prairies.—Betz (1977) defines a prairie as a vegetative community com-
posed of native perennial grasses and other herbaceous plants, and in which
grasses contribute much of the vegetative cover. Prairies in Oktibbeha County
are located largely in the Black Prairie physiographic region, and are of the
blackland type, so named after the deep mantle of fertile black soil high in organic
eh
712
BRIT.ORG/SIDA 20(2)
Taste 7. Importance value’ of under-, mid-, and overstory plant species in representative managed
and unmanaged pine forests and pine-mixed hardwood forests, Oktibbeha County, Mississippi, Fall
1993 (data from Theriot et al. 1993
)
b
Pine forest Pine forest Pine-mixed
Species (managed) (unmanaged) hardwood forest
Understory
Carya glabra (Mill) Sweet 38.98
Lonicera japonica Thunb. 61.71 72.19 58.34
Rubus spp. 9.23 93.87 10.23
Smilax spp 15.67 22.6) 8.14
Solidago spp 9.12 22.33 16.91
Rhus radicans L. 11.67 6.13 17.03
Ulmus alata Michx. 2.83 14.4] 2.45
Misc. graminoids 17,73 16.19 2.58
Misc. forbs 17.69 15.13 47.51
Eupatorium spp. 37.14 3.62
Bidens spp. 11.13
L iOSpyt Os virgit var Ud ks ] 402 6.) 0
Saccharum giganteum (Walter) Pers. 24,21 18.79
Lespedeza 2.46 10.99
Panicum spp. (s.1.) 2.58 14.95
Pinus taeda L 11.45 5.82
Quercus sp 18.36 39.62
Vitis rotundifolia Michx. 28.60 11.09
Misc. tree seedlings 14.00 - 7.14
Total: 300.00 300.00 300.00
Midstory no midstory present
Diospyros virginiana L. 15.14
Nyssa sylvatica Marshall 53.64
Acer rubrum L. 32.70 9.36
Carya tomentosa (Poir.) Nutt. 54.76 6.44
Cornus florida L. 3.94 7.90
Liquidambar styraciflua L 55.27 104.44
Pinus spp. 7 12.27 60,20
Quercus spp 61.61 28.74
Ulmus alata Michx. 10.65 44.98
Juniperus virginiana L. 24.58
Prunus serotina Ehrh. 13.36
Total 0.00 300.00 300.00
Overstory
Pinus spp. 75.8] 266.65 178.28
Quercus spp. 24.19 18.12 714
Liquidambar styraciflua L. - 6.99 87.00
Juniperus virginiana L. - 19.93
Ulmus alata Michx. - 7.65
Acer rubrum L. - 8.24
Total: 300.00 300.00 300.00
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI 713
matter, which occurs over a substrate of Cretaceous chalk or marl (Foti 1989).
Today, most native prairies in Oktibbeha County have been replaced by agri-
culture (Lowe 1921; Brent 1973), with only few relicts of this highly threatened
ecoregion remaining along highway and powerline rights-of-way. Prairies are
among the most diverse plant communities in Oktibbeha County, albeit the
condition of the prairie vegetation is generally quite poor due to the extent of
human disturbance. Prairies are also home to some of the rarest plants in the
county, featuring 19 species listed by the MNHP (2002a, 2002b), with many of
them occurring only in this community. Prairie habitats in Oktibbeha County
are dominated by a complex of graminoids, composites, and legumes. Charac-
teristic graminoids include Carex basiantha Steud., C. cherokeensis Schwein., C.
granularis Muhl. ex Willd. C. hirsutella Mack. C. muehlenbergii Schkuhr ex
Willd., Scleria oligantha Michx., Juncus torre yi Coville, Andropogon glomeratus
(Walter) Britton, Sterns & Poggenb., A. virginicus L., Aristida longespica Poir.in
Lam., A. oligantha Michx., Bothriochloa laguroides (DC.) Pilger in Engler ssp.
torreyana (Steud.) Allred & Gould, Dichanthelium depauperatum (MuhL.)
Gould, Panicum virgatum L., Phalaris caroliniana Walter, Schizachyrium
scoparium (Michx.) Nash in Small, Setaria parviflora (Poir.) Kerguélen, and
Sporobolus vaginiflorus (Torr.ex A.Gray) Alph.Wood.Common forbs are Ruellia
humilis Nutt., Apocynum cannabinum L., Asclepias spp., Aster spp., Cacalia
plantaginea (Raf.) Shinners, Dracopis amplexicaulis (Vahl) Cass., Heterotheca
camporum (Greene) Shinners, Liatris spp., Ratibida pinnata (Vent.) Barnh.,
Rudbeckia fulgida Aiton, Silphium spp., Solidago spp., Lithospermum canescens
(Michx.) Lehm., Lobelia spicata Lam., Cuscuta pentagona Engelm. var. pentagona,
Croton spp., Euphorbia spp., Tragia urticifolia Michx., Crotalaria sagittalis L.,
Dalea candida Willd., D. purpurea Vent., Sabatia angularis (L.) Pursh, Hyperi-
cum sphaerocarpum Michx., Monarda citriodora Cerv. ex. Lag., Anemone
caroliniana Walter, Delphinium carolinianum Walter ssp. carolinianum, Ra-
nunculus fascicularis Muhl. ex Bigelow, Potentilla simplex Michx., Hedyotis spp.,
Agalinis spp., Buchnera floridana Gandog., Penstemon digitalis Nutt. ex Sims,
P. laevigatus Aiton, Verbena bipinnatifida Nutt. V. simplex Lehm., Manfreda
virginica (L.) Salisb. ex Rose, Hypoxis hirsuta (L.) Coville, and Sisyrinchium
albidum Raf. In disturbed areas, such ruderal species as Ambrosia psilostachya
DC., A. trifida L., Iva annua L., Lactuca saligna L., Lepidium virginicum L.,
Medicago lupulina L., and Melilotus alba Medik. may be evident. Noteworthy
prairie species listed by MNHP (2002a, 2002b) include Perideridia americana
(Nutt.ex DC.) Reichenb. (critically imperiled to imperiled-S1S2), Amphiachyris
—
a
The importance value is the sum of relative frequency, relative density, and relative dominance
(Daubenmire 1968).—®Data for understory estimates based on 100 ft. of line transect per habitat
type using the intercept method (Canfield 1942); data for mid- and overstory estimates based on
10 sample points per habitat type using the point-centered-quarter (PCQ) method (Cottam & Cur-
tis 1956).
714 BRIT.ORG/SIDA 20(2)
dracunculoides (DC.) Nutt. (critically imperiled-Sl), Aster ericoides L. Gmper-
iled-S2), Evax prolifera Nutt. ex DC. (SL), Prenanthes aspera Michx. (S2),
Neptunia lutea (Leavenw.) Benth. (rare and local to apparently secure-S3S4),
Linum sulcatum Riddell (S354), Dodecatheon meadia L. (S2), Delphinium
tricorne Michx. (S2), Agalinis pseudaphylla (Pennell) Shinners (S2), Penstemon
tenuiflorus Pennell Gmperiled to rare and local-S2S3), Tomanthera auriculata
(Michx.) Raf. (SL), Carex jamesii Schwein. (SIS2), C. meadii Dewey (S354), C.
microdonta Torr. & Hook. (possibly imperiled-S2?), Nemastylis geminiflora
Nutt. (S2), Spiranthes magnicamporum Sheviak (S253), and Bouteloua
curtipendula (Michx.) Torr. in Marcy (S354); Onosmodium molle Michx. ssp.
hispidissimum (Mack.) B. Boivin is considered rare by USDA Forest Service (Kral
1983). Further discussion of Black Prairie floristics in Oktibbeha County can
be found in Leidolf and McDaniel (1998).
5. Prairie Cedar Woodlands.—Leidolf and McDaniel (1998) define a prairie
cedar woodland as *... Juniperus virginiana L.-dominated patches of woody veg-
etation restricted to thin, highly erodible, calcareous soils over chalk ... that are
often interspersed with areas of exposed chalk, giving them a glade-like ap-
pearance”. Prairie cedar woodlands are commonly found in the Black Prairie
physiographic region of Oktibbeha County. Leidolf and McDaniel (1998) sug-
gested that the apparent abundance of this plant community isa rather recent
feature, and linked it to the absence of natural disturbance. Lowe (1921) also
noted that “... lands formerly in cultivation and now thrown out, show a strong
tendency to grow up in trees, ...”
Characteristic tree and shrub species of this community include Juniperus
virginiana L. var. virginiana, Rhus aromatica Aiton, Ilex decidua Walter, Vibur-
num rufidulum Ral., Cornus drummondii C.A. Mey., Quercus marilandica
Muenchh., 9. muehlenbergii Engelm., Q. stellata Wangenh., Sassafras albidum
(Nutt.) Nees, Maclura pomifera (Raf.) CK. Schneid., Morus rubra L., Fraxinus
americana L., Ligustrum sinense Lour, Ceanothus americanus L., Rhamnus
caroliniana Walter, Crataegus crus-galli L., C. engelmannii Sarg., Rosa carolina
L., Rubus cuneifolius Pursh, Bumelia lycioides (L.) Pers., Celtis laevigata Willd.,
Ulmus alata Michx.,and U. rubra Muhl; Campsis radicans(L.) Seem. ex Bureau,
Lonicera japonica Thunb., Berchemia scandens (Hill) K. Koch, Ampelopsis
arbored (L.) Koehne, Parthenocissus quinquefolia (L.) Planch., Vitis aestivalis
Michx., and Smilax bona-nox L. are common lianas. Herbaceous vegetation is
sparse and includes Ruellia caroliniensis J.F Gmel.) Steud., Sanicula canadensis
L., Euphorbia spathulata Lam., Apios americana Medik., Cocculus carolinus (L.)
DC., Fragaria virginiana Duchesne, Carex bushii Mack., C. cherokeensis
Schwein.,and C.oxylepis Torr. & Hook var. pubescens J.K. Underw; occasionally
ferns, such as Asplenium platyneuron (L.) Oakes and Pleopeltis polypodioides
(LJ) E.G. Andrews & Windham var. michauxiana (Weath.) E.G. Andrews &
Windham in Windham can be found. A number of unique floristic elements
—
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI 715
found in no other habitat in Oktibbeha County were also found: Rhus
toxicodendron L., Helianthus divaricatus L.,and Panicum flexile (Gatt.) Scribn.
in Kearney. Noteworthy prairie cedar woodland species listed by MNHP (2002a,
2002b) include Ophioglossum engelmannii Prantl (critically imperiled-S]),
Neptunia lutea (Leavenw.) Benth. (rare and local to apparently secure-S3S4),
Dodecatheon meadia L. (imperiled-S2), Rhamnus lanceolata Pursh (S2), Pen-
stemon tenuiflorus Pennell (imperiled to rare and local-S2S3), and Ponthieva
racemosa (Walter) C.Mohr (possibly imperiled-S2?). Morris et al. 1993) and
Leidolf and McDaniel (1998) provide additional comments on prairie cedar
woodland floristics in Oktibbeha County.
6. Chalk Outcrops.—Chalk outcrops in Oktibbeha County are restricted to
the Black Prairie physiographic region. They were first recognized as being
floristically distinct by Hilgard 860), who referred to them as “bald prairies”.
Leidolf and McDaniel (1998) defined chalk outcrops as “...areas of exposed chalk,
with all of the overlying soil eroded away ...”, and suggested that aggravated
erosion during periods of intense cultivation may have been responsible for an
apparent decline in native prairie grasslands accompanied by a simultaneous
increase in the abundance of chalk outcrops. Because conditions for plant
growth in this unique habitat are severe, chalk outcrops do not support a di-
verse plant community; however, they harbor floristic elements not found else-
where in Oktibbeha County, including Senecio obovatus Muhl. ex Willd.,
Chamaesyce humistrata (Engelm.) Small, C. serpens (Kunth) Small, Lespedeza
procumbens Michx., and Mirabilis albida (Walter) Heimerl. Due to ample rain-
fall and low evaporation, the ground is usually moist during winter and spring
months, and colonies of the blue-green alga Nostoc sp. can occasionally be found;
in the summer, harsher, drier conditions prevail. Plant species characteristic of
this habitat type include both native prairie grassland remnants, and plants
that colonize chalk outcrops after erosion. Commonly found trees and shrubs
include Juniperus virginiana L. var. virginiana, Rhus aromatica Aiton, Cornus
drummondii CA. Mey, Hypericum sphaerocarpum Michx., Ceanothus americanus
L., Rhamnus caroliniana Walter, and Prunus angustifolia Marshall. Character-
istic herbs are Amaranthus rudis J.D. Sauer, Asclepias viridis Walter, Aster patens
Aiton, Brickelia eupatorioides (L.) Shinners var. eupatorioides, Coreopsis
lanceolata L., Heterotheca camporum (Greene) Shinners, Liatris squarrosa (L..)
Michx., Silphium laciniatum L., Lithospermum canescens (Michx.) Lehm.,
Brassica erucastrum L., Euphorbia corollata L., Dalea candida Willd., D.
purpurea Vent., Sabatia angularis (L.) Pursh, Agalinis tenuifolia (Vahl Raf.,
Verbena bipinnatifida Nutt., V. simplex Lehm., and Sisyrinchium albidum Raf.
Noteworthy members of this association listed by MNHP (2002a) are Sedum
pulchellum Michx. (critically imperiled-SD, Rhamnus lanceolata Pursh Gim-
periled-S2), Penstemon tenuiflorus Pennell Gmperiled to rare and local-S2S3),
and Nemastylis geminiflora Nutt. (S2). Further discussions of chalk outcrop
716 BRIT.ORG/SIDA 20(2)
floristics in Oktibbeha County are provided by Morris et al. 1993) and Leidolf
and McDaniel (1998).
AQUATIC COMMUNITIES
Aquatic communities in Oktibbeha County include both such lotic systems as
rivers, creeks, canals, and drainage ditches, and lentic systems, including lakes,
ponds, and artificial impoundments. Vegetation in these communities is mostly
restricted to narrow regions near the shore referred to as riparian areas. Ripar-
ian areas generally have higher species diversity than either the adjacent ter-
restrial or aquatic communities, because they serve as a transition between the
two systems. They are also generally more productive because of periodic in-
fluxes of nutrients from seasonal floods, and favorable moisture conditions year-
round. Species composition at any point ina riparian area is largely a function
of the degree or periodicity of inundation from the adjacent stream (Mitsch &
Gosselink 1986).
l. Rivers/Creeks.—We discuss here the flora associated with riparian areas
immediately adjacent to the stream bank, including those species growing on
banks, associated mud flats, and directly in the stream (emergents). Submergent
species are generally absent in local streams, because the turbid water effec-
tively prevents photosynthesis; where clear-water streams do exist (ie. in the
Black Prairie physiographic region) pH is likely too high for plant growth, and
consequently, no submergent species were observed. As mentioned previously,
there are no major alluvial systems in Oktibbeha County; thus extensive ripar-
ian swamp forests are not represented.
In the Interior Flatwoods physiographic region, streambank associations
include such small trees and shrubs as Quercus lyrata Walter, Planera aquatica
J.EGmel, Salix nigra Marshall, Acer negundo L., Betula nigra L., Prunus serotina
Ehrh., and Evonymus americanus L., as well as such woody vines as the intro-
duced Lonicera japonica Thunb,, and herbaceous plants, including Thaspium
trifoliatum (L.) A. Gray, Aristolochia tomentosa Sims, Leersia virginica Willd.,
and Paspalum repens PJ. Bergius; noteworthy members of this association in-
clude Ptelea trifoliata L., which is listed as rare and local to apparently secure
in the state (S354, MNHP 2002b). Due to their ephemeral nature, mudflat com-
munities in the Interior Flatwoods are generally limited to such herbaceous
plants as Lobelia cardinalis L., Lycopus virginicus L., Samolus valerandi L. ssp.
parviflorus (Raf.) Hulten, Lindernia dubia (L.) Pennell, Mimulus alatus Aiton,
Physalis virginiana Mill., Boehmeria cylindrica (L.) Sw., and Commelina
virginica L. Emergents are few, and include Ammania coccinea Rottb., Hydro
uniflora Raf.,and Penthorum sedoides L.
—
ed
Streambank associations in the Black Prairie physiographic region are char-
acterized by Aesculus pavia L., Tilid americana L., Lonicera japonica Thunb.,
Apios americana Medik., Mimulus alatus Aiton, and Silphium perfoliatum L.
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI 717
Noteworthy streambank associates include Aesculus glabra Willd., which is
listed as possibly imperiled in the state (S2?, MNHP 2002a), and was found ex-
clusively on Catalpa silty clay loam. Common emergents are Ludwigia peploides
(Kunth) Raven and Juncus acuminatus Michx.
2. Canals/Drainage Ditches.—Canals/drainage ditches include both arti-
ficial, human-made structures, such as irrigation canals and culverts, and semi-
natural features, such as unimproved roadside ditches. These habitats vary con-
siderably with respect to extent, seasonality, and speed of water flow; they
include habitats subject to year-round water flow as well as ephemeral habi-
tats. Like rivers and creeks, the vegetation associated with canals/drainage
ditches can be separated into riparian species (i.e., those species growing on the
banks) and emergent species; additionally, the flora of canals/drainage ditches
includes a small number of submergent species.
In the Interior Flatwoods physiographic region, plants commonly found
on canal and ditch banks include such small trees and shrubs as Sambucus
canadensis L. and Cephalanthus occidentalis L.,as well as such herbaceous spe-
cies as Justicia ovata (Walter) Lindau var. lanceolata (Chapm.) R.W. Long,
Ptilimnium capillaceum (Michx.) Raf., Rotala ramosior (L.) Koehne, Ranuncu-
lus pusillus Poir., Dichanthelium polyanthes (Schultes) Mohlenbr., D.
sphaerocarpon (Elliott) Gould, and Urochloa ramosa (L.) T.Q. Nguyen; notewor-
thy bank species include Muhlenbergia glabriflora Scribn., which is listed as
being of uncertain status in Mississippi (S?, MNHP 2002a). Dominant emer-
gent species primarily include such obligate and facultative aquatic forbs,
sedges, and rushes as Physostegia angustifolia Fern., Mollugo verticillata L.,
Ludwigia spp., Penthorum sedoides L., Polygonum spp., Gratiola spp. Commelina
virginica L., Carex flaccosperma Dewey, C. joorii L.H. Bailey, Cyperus odoratus
L., C. strigosus L., Eleocharis spp., Fimbristylis spp., Rhynchospora corniculata
(Lam.) A. Gray, Scirpus atrovirens Willd, Iris virginica L. Juncus spp.and Typha
latifolia L.: noteworthy emergents include Alisma subcordatum Raf., which is
listed as being of historical occurrence in Oktibbeha County (SH, MNHP 2002a).
Submergent species include Callitriche heterophylla Pursh and Najas
guadalupensis (Spreng.) Magnus.
In the Black Prairie physiographic region, canal and ditch banks are char-
acterized by Sambucus canadensis L., Salix exigua Nutt, Rorippa sessiliflora
(Nutt.) Hitche., Apios americana Medik., Ranunculus pusillus Poir, and
Dichanthelium ravenellii (Scribn. &@ Merr.) Gould. Characteristic emergent spe-
cies in the Black Prairie are Physostegia angustifolia Fern., Ludwigia peploides
(Kunth) Raven, Polygonum lapathifolium L., Lysimachia nummularia L.,
Commelina communis L., Carex frankii Kunth, Cyperus odoratus L., C.
acuminatus Torr. & Hook. ex Torr, Eleocharis obtusa (Willd.) Schult. Scirpus
pendulus Muhl., Juncus spp.,and Typha s
3.Lakes/Ponds/Impoundments.—All phe and ponds in Oktibbeha County
718 BRIT.ORG/SIDA 20(2)
>
are artificial impoundments formed by damming a river, creek, or other drain-
age. Only two sizable (>250 ha) examples are present, both in the Interior
Flatwoods physiographic region: Oktibbeha County Lake and Bluff Lake (most
of which is in neighboring Noxubee County). Most other lentic systems are less
than 10 ha in size and completely surrounded by private land; thus the flora of
these areas may be underrepresented in this document because of poor access.
eT
We were able to obtain a fair representation from one such smaller lake, Dorman
Lake, which is on public land and also located in the Interior Flatwoods.
The flora of lakes and ponds can be discussed in terms of those species
growing in the riparian zone, including the margin or bank, and associated
mudflats; those growing in the littoral zone, i.e., the shallow part of the lake
immediately adjacent to the shore, including emergents, submergents, rooted-
floating, and free-floating species; and those growing in the profundal zone, ie.,
the deeper or interior part of the lake, including rooted-floating and free-float-
ing species.
Common species along the margins of lakes and ponds include such small
trees and shrubs as Baccharis halimifolia L., Betula nigra L., Itea virginica L.,
Cephalanthus occidentalis L., which also occurs in standing water in mudflats
and in the littoral zone, and Salix humilis Marshall, as well as herbaceous spe-
cies, including Baptisia alba (L.) R.Br. var. macrophylla (Larisey) Isley and
Leptochloa panicoides QJ). Presl) Hitche. Mudflat associates include Eryngium
prostratum Nutt.ex DC., Pluchea camphorata (L.) DC., Lobelia cardinalis L., Po-
lygala mariana Mill., Polygonum aviculare L., Ranunculus abortivus L.,
Micranthemum umbrosum (J.E Gmel.) S.F Blake, Arisaema dracontium (L.)
Schott, Cyperus erythrorhizos Muhl., Cinna arundinacea L., Juncus elliottii
Chapm., Scirpus cyperinus (L.) Kunth, Echinodorus cordifolius (L.) Griseb., and
Sagittaria platyphylla (Engeli.) J.G. Sm. These last 3 species, along with
Hydrolea uniflora Ral., Ludwigia palustris (L.) Elliott, Polygonum spp.,
Eleocharis quadrangulata(Michx.) Roem. & Schult.,and Juncus nodatus Coville,
are also representative emergent species. A noteworthy collection from this as-
sociation is Alisma subcordatum Raf., which is listed as being of historical oc-
currence in Oktibbeha County (SH, MNHP 2002a).
As their name implies, rooted-floating plants are attached to the substrate,
with leaves floating at the lake surface. Rooted-floating and submergent veg-
etation occurs from the lake margin toa depth of ca. 2 min the littoral zone. In
contrast, free-floating species, including Azolla caroliniana Willd. and
Spirodela polyrrhiza (L.) Schleid., occur in both the littoral and the profundal
zone, but generally are blown into clumps on the lee side of water bodies.
Sul gent species of the littoral zone include Callitriche heterophylla Pursh,
nee bbaL., Najas minor All, and Zanichellia palustris L. Representa-
tive rooted-floating plants are Brasenia schreberiJ.- Gmel., Limnobium spongia
(Bosc) Rich. ex Steud., Nelumbo lutea Willd., Nymphaea odorata Aiton,
~~
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI 719
Potamogeton diversifolius Raf.,and P. nodosus Poir. Frequently, dense vegetation
mats formed by rooted-floating plants provide a substrate for other plants to
grow hydroponically. However, no evidence of this was found in Oktibbeha
County.
SEEPAGE AREAS
Seepage areas are highly localized habitats that occur on relatively thin soil
covering an underlying hardpan or rock formation and that, depending on lo-
cal soil conditions and vegetation, vary considerably with respect to the source
and extent of water flow, acidity, and degree and duration of stagnation. How-
ever, with the exception of a small number of natural springs, most seepage
areas in Oktibbeha County consist of small ground depressions along road-
sides in the Interior Flatwoods physiographic region. They are fed by sub-lat-
eral flow from associated pine forests and occur on sandy loams overlain by a
thin layer of clay. Consequently, water flow is acidic and highly unpredictable,
with very little stagnation occurring. Within the constraints imposed by the
local species pool, the vegetation of seepage areas most closely resembles that
of a southern pine bog. Characteristic plant species include ferns and fern al-
lies, such as Lycopodiella alopecuroides(L.) Cranfill, L. appressa (Chapm.) Cran-
fill, L. prostrata (R.M. Harper) Cranfill, Woodwardia areolata (L.) T.Moore, and
Pteridium aquilinum (L.) Kuhn var. pseudocaudatum (Clute) A.Heller;
graminoids, including Carex frankii Kunth, Cyperus polystachyos Rottb., C.
rotundus L., Kyllinga odorata Vahl, Rhynchospora caduca Elliott, R. inexpansa
(Michx.) Vahl, Panicum virgatum L., and Paspalum spp. as well as numerous
forbs, including Aster solidagineus Michx., Pityopsis graminifolia (Michx.) Nutt.
Buchnera floridana Gandog,, Ruellia caroliniensis J.F Gmel.) Steud., Drosera
brevifolia Pursh, Eryngium prostratum Nutt. ex DC,, E. yuccifolium Michx.,
Lysimachia ciliata L., Polygala nana (Michx.) DC., Aletris aurea Walter,
Spiranthescernua (L.) Rich., S. praecox (Walter) S.Watson, and Xyris torta Small;
noteworthy species from this association include Platanthera cristata (Michx.)
Lindl., which is listed as rare and local in the state ($3, MNHP 2002a). Occa-
sionally, the small shrub Salix humilis Marshall can be found. Where stagnant
pools of water occur, Callitriche heterophylla Pursh may be found submergent.
HUMAN-INFLUENCED COMMUNITIES
1 Cultivated Fields.—As of 1997, 17,500 ha, or 15%, of Oktibbeha County was
cropland (USDA NASS, http://www.nass.usda.gov). Due to its productive soils,
most cropland in the county is located in the Black Prairie physiographic re-
gion. Crops commonly planted include corn (Zea mays L..), cotton, and soy bean
(Glycine max [L.] Merr). A variety of weedy species, both native and exotic,
readily invades these cultivated fields. Common natives found in this habitat
include such forbs as Amaranthus spp., Ciclospermum leptophyllum (Pers.)
Sprague ex Britton & Wilson, Aster pilosus Willd. var. pilosus, Helenium
720 BRIT.ORG/SIDA 20(2)
autumnale L., Ipomoea lacunosa L., |. pandurata(L.) G.Mey., Myosotis verna Nutt.,
Rorippa sessiliflora (Nutt.) Hitche., Sibara virginica (L.) Rollins, Chamaesyce
spp., Oenothera laciniata Hill, Myosurus minimus L., Ranunculus hispidus
Michx. var. hispidus, and Linaria canadensis (L.) Chaz. as well as graminoids,
including Juncus bufonius L., J. torreyi Coville, Digitaria ciliaris (Retz.) Koel.
Panicum capillare L., Paspalum pubiflorum Rupr. ex E.Fourn., and Urochloa
platyphylla (Munro ex Wright) R.D. Webster. Frequently encountered exotic
forbs are Arctium minus Bernh., Ipomoea hederacea Jacq., Capsella bursa-
pastoris (L.) Medik., Cerastium glomeratum Thuill., Spergula arvense L.,
Medicago arabica (L.) Huds., Vicia grandiflora Scop., Abutilon theophrasti
Medik., Polygonum aviculare L., Rumex crispus L., Ranunculus arvensis L., and
R. muricatus L. representative exotic graminoids include Cyperus esculentus
L.,C. rotundus L., Cyonodon dactylon (L.) Pers., Echinochloa colona (..) Link, E.
muricata (PBeauv.) Fernald, Lolium multiflorum Lam., Paspalum dilatatum Poir.
in Lam., Poa annua L., Setaria faberi R.A.W. Herm., and Sorghum halepense (L.)
Pers.
2. Grass/Forb Meadows.—Plant species occurring in grass/forb meadows,
which include hay fields, pastures, and wildlife food plots, have to withstand or
escape the frequent grazing or mowing characteristic lor this community. Some
species escape this disturbance by growing close to the ground, such as Galium
pilosum Aiton, Hedyotis crassifolia Raf., Ranunculus pusillus Poir, Veronica
persica Poir, and Kyllinga odorata Vahl., whereas other species, such as Pani-
cum scoparium Lam., Trifolium arvense L., or T. pratense L., persist even under
heavy grazing. Another group of plants found in this community are early spring
ephemerals, such as Claytonia virginica L. and Nothoscordum bivalve (L..)
Britton.
Grass/forb meadow associations in the Interior Flatwoods physiographic
region are characterized by such species as Euthamia tenuifolia (Pursh) Nutt.,
Eupatorium rotundifolium L., Cardamine bulbosa (Schreb. ex Muhl.) Britton,
Sterns & Poggenb., Cuscuta pentagona Engelm. var. pentagona, Euphorbia
spathulata Lam., Hypericum punctatum Lam, Prunella vulgaris L., Trichostema
brachiatum L., Hedyotis crassifolia Raf, Glyceria declinata Bréb.,and Dichanthelium
scoparium (Lam.) Gould; moist depressions may hold such wetland species as
Ludwigia alternifolia L. L. hirtella Ral., and Gratiola neglecta Torr; noteworthy
records from this association include Asclepias hirtella (Pennell) Woodson
which is listed as imperiled in the state (S2, MNHP 2002a).
In the Black Prairie physiographic region, species representative of grass/
orb meadows include Ruellia humilis Nutt., Silphium laciniatum L., Vernonia
gigantea (Walter) Trel. var. gigantea, Lithospermum canescens (Michx.) Lehm.,
Opuntia humifusa (Raf.) Raf., Galium pilosum Aiton, Carex festucacea Schkuhr
ex Willd.,and Kyllinga odorata Vahl; noteworthy collections from this associa-
tion include Armoracia lacustris (A. Gray) Al-Shebaz & Bates and Camassia
~
—
’
—
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI 72)
scilloides Corey, which are listed as critically imperiled to imperiled (SIS2), and
imperiled to rare and local (S253) in the state, respectively (MNHP 2002a).
3. Roadsides.—Roadsides are among the most interesting plant communi-
ties in Oktibbeha County, as they represent a conglomerate of floristic elements
of diverse origins. Because roads are frequently associated with human habita-
tion, both past and present, roadside plant communities often feature species
that have escaped from or persist after cultivation. Examples in Oktibbeha
County include Catalpa bignonioides Walter, Wisteria sinensis (Sims) DC.,
Broussonetia papyrifera (L.) LHér. ex Vent., Populus alba L., Lycoris radiata
(LHér.) Herb., Narcissus spp., Hemerocallis fulva (L.) L., Muscari neglectum Guss.
ex Ten., and Belamcanda chinensis (L.) DC. The high frequency of disturbance,
e.g., mowing of rights-of-way, associated with roadsides also provides suitable
habitat for a diverse array of native herbs and exotic weeds (“ruderals”). New
introductions are likely to be seen first along roads that can provide far-reach-
ing connections between seed source and suitable dispersal habitat. Examples
of common native roadside species include Bidens aristosa (Michx.) Britton,
Cirsium horridulum Michx., Erigeron annuus (L.) Pers., Iva annua L., Senecio
anonymus Wood, Acalypha virginica L., Chamaesyce maculata (L.) Small, Cro-
ton capitatus Michx., Desmanthus illinoensis (Michx.) MacMill. ex B.L. Rob. &
Fernald, Lespedeza cuneata (Dum.Cours.) G. Don, Blephilia ciliata (L.) Benth.,
Hedeoma hispida Pursh, Prunella vulgaris L., Plantago rugelii Decne., Rumex
hastatulus Baldwin, Carex frankii Kunth, C. triangularis Boeck., Dichanthelium
laxiflorum(Lam.) Gould & C.A. Clark, D. scabriusculum (Elliott) Gould @ C.A.
Clark, Elymus virginicus L., and Tridens flavus (L.) Hitche. Exotics frequently
encountered include Daucus carota L., Torilis arvensis Huds.) Link, Taraxacum
officinale Weber ex FH. Wigg., Capsella bursa-pastoris (L.) Medik., Lonicera
japonica Thunb., Cerastium glomeratum Thuill., Medicago lupulina L., Trifo-
lium repens L., Lamium spp., Verbena brasiliensis Vell. Bothriochloa pertusa
(L.) ACamus, Setaria parviflora (Poir.) Kerguélen, and Sorghum halepense (L.)
Pers. Whereas the species mentioned thus far can be found along roads through-
out the county, roadside communities, to a large degree, also reflect the plant
communities that immediately surround them. Thus, roadsides in the Interior
Flatwoods often feature such species as Boltonia diffusa Elliott, Elephantopus
carolinianus Raeusch., Acalypha rhomboidea Raf., Desmodium paniculatum(L.)
DC., Lespedeza capitata Michx., Hypericum lobocarpum Gatt., Rhexia mariana
L. var. mariana, Phlox pilosa L., Polygala sanguinea L., Aletris aurea Walter,
Spiranthes vernalis Engelm. & A. Gray, Paspalum floridanum Michx., P. laeve
Michx., Tridens strictus (Nutt.) Nash in Small, and Tripsacum dactyloides (L.) L;
note wor thy collections from this association include Asclepids hirtella (Pennell)
Woodson and Lobelia appendiculata A.DC., which are listed as imperiled (S2)
and rare and local to apparently secure (S354) in the state, respectively (MNHP
2002a). Typical Black Prairie roadside species are Aster novae-angliae L., A.
722 BRIT.ORG/SIDA 20(2)
praealtus Poir. var. praealtus, A. undulatus L., Ratibida pinnata (Vent. Barnhart,
Rudbeckia fulgida Aiton, Silphium laciniatum L., Specularia perfoliata (L.)
A.DC., Oenothera speciosa Nutt., Anemone caroliniana Walter, Penstemon
laevigatus Aiton, Panicum capillare L.,and P.dichotomiflorum Michx.; notewor-
thy records include Dasistoma macrophylla (Nutt.) Ral. (rare and local to ap-
parently secure-S3S4), Penstemon tenuiflorus Pennell (imperiled to rare and
local-S2S3), Tomanthera auriculata (Michx.) Raf. (critically imperiled-SI), and
Spiranthes magnicamporum Sheviak (S283, MNHP 2002a, 2002b).
4. Urban Areas (including lawns).—The flora of urban areas is depauper-
ate, and consists mostly of plants that have escaped from, are spreading from,
or persist after cultivation; as well as weedy species that aggressively invade
lawns, flower beds, sidewalks, and roadsides. Examples of local escapes include
Vinca major L., Ratibida columnifera (Nutt.) Wooton & Standl., Evonymus
fortunei (Turez.) Hand.-Mazz., Elaeagnus umbellata Thunb., Wisteria sinensis
(Sims) DC, Populus alba L., Ailanthus altissima (Mill) Swingle, Lycoris radiata
(LHér.) Herb., Narcissus spp., Hemerocallis fulva (L.) L., and Hyacinthoides
nonscripta(L.) Chouard ex Rothm. Commonly found native species include such
forbs as Bowlesia incana Ruiz & Pav, Acmella oppositifolia (Lam.) R.K. Jansen
var. repens (Walter) R.K. Jansen, Ambrosia trifida L.. Hieracium gronovii L.,
Lactuca serriola L., Draba brachycarpa Nutt. ex Torr & A. Gray, Sagina
decumbens (Elliott) Torr & A. Gray, Chamaesyce spp., Plantago spp., Rumex
hastatulus Baldwin, and Ranunculus spp. as well as many graminoids, includ-
ing Cyperus spp., Fimbristylis annua (AIL) Roem. & Schult., Kyllinga spp.,
Scirpus koilolepis (Steud.) Gleason, Juncus tenuis Willd., Axonopus fissifolius
(Raddi) Kuhlm., Hordeum pusillum Nutt. Dichanthelium spp., Sphenopholis
obtusata (Michx.) Scribn., and Urochloa ramosa (L.) T.Q. Nguyen. Exotic weeds
characteristic of urban areas include Arctium minus Bernh., Centaurea cyanus
L., Lactuca saligna L., Taraxacum officinale Weber ex FH. Wigg., Youngia
japonica(L.) DC,, Brassica juncea (L.) Czern., Capsella bursa-pastoris(L.) Medik.,
Cardamine hirsuta L., Cerastium glomeratum Thuill, Stellaria media (L. Vill,
Medicago arabica (L.) Huds., Trifolium spp., Lamium spp., Sherardia arvensis L.,
Valerianella locusta (L.) Betcke, Verbena brasiliensis Vell., and Poa spp.
ANNOTATED CHECKLIST
Species and lesser taxa are grouped by division, class, and family. Plant divi-
sions and classes are arranged according to phylogeny following Flora of North
America Editorial Committee (1993); families, and genera and species within
families, are arranged alphabetically. Non-vascular plant family and genus con-
cepts follow Schuster (1966, 1969, 1974, 1980) for Anthocerotopsida and
Hepaticopsida. Vascular plant family and genus concepts follow Brummitt
(1992). Nativity status is based on the USDA NRCS Plants database (http://
_——
eae
—
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI 723
plants.usda.gov). Species names are followed by the authority, with abbrevia-
tions according to Brummitt and Powell (1992). In general, nomenclature fol-
lows Kartesz (1994), with the exception of Poaceae, which follow Barkworth et
al.(n preparation). Synonymsare given in brackets where pertinent. The physi-
ographic region/regions in which a species occurs is/are indicated by the ab-
breviations BP (Black Prairie), PR (Pontotoc Ridge), or IF (Interior Flatwoods),
followed by the plant community/communities in which that species is found.
Relative abundance within a plant community is listed as very abundant (AA),
abundant (A), common (C), occasional (O), uncommon (U), rare (R), very rare
(RR), or historical (H). A plant that is common, abundant, or very abundant,
but local in distribution, is indicated with (1) following the abundance designa-
tion. The status of a species is indicated in bold as federally threatened (FT) or
sensitive (S), which includes the designations critically imperiled ($1), imper-
iled ($2), rare and local ($3), apparently secure ($4), demonstrably secure ($5),
of historical occurrence (SH), and of uncertain status (SU; MNHP 2002a, 2002b).
This is followed by a literature citation or accession number, composed of col-
ector name, collection number (with “R” indicating a recollection), and source/
depository herbarium acronym (Holmgren et al. 1990). Collections of the se-
nior author are cited first (no collector name given), followed by those of others
(collector name given, ina list for first specimen only); absence of an herbarium
acronym indicates the specimen is referenced at IBE. Species preceded by an
asterisk (*) are non-native species. An ampersand (®) indicates the species is a
new record for Mississippi.
j—
BRYOPHYTA
HEPATICOPSIDA
CEPHALOZIACEAE
Cephalozia bicuspidata (L.) Dumort., IF, lakes/
ponds/impoundments, U; Woods (1964)
CEPHALOZIELLACEAE
Cephaloziella hampeana (Nees) Schiffn. ex
Loeske, IF, U; Woods (1964)
GEOCALYCACEAE
Chiloscyphus pallescens (Ehrh.
Dumort., IF rivers/
Lophocolea bidentata (L.) Dumort., IF, lakes/
ponds/impoundments, U; Woods (1964)
Lophocolea heterophylla (Schrad.) Durmort., IF,
lakes/ponds/impoundments, C; Woods
(1964)
Lophocolea minor Nees, IF, lal
poundments, C; Woods (1964)
Ay aan Jas:
2 POIIGo/ ITT
JUBULACEAE
Frullania sents Gottsche, PR, mesophytic
upland hardwood forests, C; Woods (1964
Frullania ae Gottsche, IF, lakes/ponds/im-
poundments, U; Woods (1964)
ullania kunzei Lehm. & Lindenb.,
onds/impoundments, U; Woods (1964)
ania squarrosa (Reinw., a & pe Dumort.,
+
IF, lakes/
S
Fru
PR, mesophytic upland | | forests, A;
Woods (1964)
JUNGERMANNIACEAE
Solenostoma pumilum (With.) K. MUull.
Jungermannia pumila With], wees
U; Woods (1964
LEJEUNEACEAE
igen esi minutissima (Smith) Schiffn., BP,
prairie cedar woodlands, O; Woads (1964)
Leucolejeunea clypeata (Schwein.) A. Evans, IF,
bottomland hardwood forests, C; Wood
(1964)
ell
Leucolejeunea unciloba (Lindenb.) A. Evans, IF,
pine forests, C; Woods (1964)
PALLAVACINIACEAE
Pallavacinia lyellii (Hook.) Carruth., IF, lakes/
ponds/impoundments, U; Woods (1964)
PORELLACEAE
Porella platyphylloidea (Schwein.) Lindenb., PR,
mesophytic upland hardwood forests, C;
Woods (1964)
RICCIACEAE
Riccia fluitans |
Woods (1964
Riccia hirta (Austin) Underw., IF, roadsides, U;
Woods (1964)
Riccla membranacea Gottsche & Lindenb, IF, riv-
ers/creeks, C; Woods (1964)
Riccia sullivantii Austin, IF, rivers/creeks, C; Woods
(1964)
cl
ponds/impoundments,
SCAPANIACEAE
et ianemorosa (L.) Dumort., IF, alla
oundments, C; Woods (1964)
SPHAEROCARPACEAE
Sphaerocarpus texanus Austin, IF, grass/forb
meadows, C; Woods (196
ANTHOCEROTOPSIDA
ANTHOCEROTACEAE
Anthoceros punctatus L., IF, lakes/ponds/im
poundments, U; Woods (1964)
LYCOPODIOPHYTA
LYCOPODIACEAE
Lycopodiella alopecuroides (L.) Cranfill, IF, seepage
areas, C;651
Lycopodiella aes (Chapm.) Cranfill, IF, seep
oadsides, O/C; 482, 1557, McDaniel
toes,
24335
Lycopodiella prostrata (R.M. Harper) Cranfill, IF,
e areas, C; 530
Lycopodium digitatum Dill. ex A. Braun, IF, pine
forests, RR; McDaniel pers. obs.
SELAGINELLACEAE
Selaginella apoda (L.) Spring, IF, mesophytic up-
— hardwood forests, R; 9/4, 1298 (R914)
BRIT.ORG/SIDA 20(2)
POLYPODIOPHYTA
ASPLENIACEAE
Asplenium daeueniane .) Oakes, BP, IF, ae for-
ests, prai cedar woodlands,0;619, 720, 879,
Evans (1 a Leidolf and McDaniel (1 s
AZOLLACEAE
Azolla caroliniana Willd., BP, lakes/ponds/im-
fete ments (littoral/profundal-free-float-
A(l); McDaniel pers. obs.
BLECHNACEAE
alien chon (L.) T. Moore, IF, mesophytic
upland hardwood forests, pine-mixed hard-
wood Bi canals/drainage ditches
ie banks), seepage areas, O; 907, 1556
978), Jones et al. (1969), McDaniel
ey Ray 6793 (MISSA)
DENNSTAEDTIACEAE
Pteridium aquilinum — (L. Kuhn — var.
persia (Clute) A. Heller, IF, pine for-
ests, seepage areas, O/A(I); 983, McDaniel
ae
<2
fev)
—
—
—
DRYOPTERIDACEAE
PO UEN a AIOe ticho Le ANG) Schott, PR, IF,
hy
Evans (1978), H.B.B. s.n. (MISSA), McDaniel
28886
OPHIOGLOSSACEAE
Botrychium biternatum (Sav.) Underw., PR, IF, bot-
mland hardwood forests, pine forests, O/
C; 878, 971, Bennett s.n., Brooks 562, Evans
(1978), Thetict et al. (1993)
otrychium virginianum (L ) Sw., a A meso-
phytic upland hard 0,1350
Jones et al. (1969), McDaniel 3 re Morris et
al. (1993), Tracy s.n. (MISSA
a ea crotalophoroides Walter; Evans
(1978)
—
Ophioglossum engelmannii Prantl, BP, prairie
cedar woodlands, A(l), $1; Evans (1978),
McDaniel 10583
OSMUNDACEAE
Osmunda cinnamomea L., IF, mesophytic upland
hardwood forests, O/C; 899, 1380 (R899),
Evans (1978), Jones et al. (1969)
Osmunda regalis L., IF, mesophytic upland hard-
wood forests, O/C; 900, 1379 (R900)
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI
POLY PODIACEAE
Pleopeltis polypodioides (L.) E.G. Andrews a
Windham var. michauxiana (Weath.) E
Andrews & Windham in Windham, BP. PR, i
bottomland hardwood forests, mesophytic
upland hardwood forests, prairie cedar
woodlands, O/C; 44, Beal s.n. (MISSA), Chan-
nell s.n. (MISSA), Evans (1978), Jones et al.
(1969), Leidolf and McDaniel (1998), Morris
et al. (1993), Ray 8005 (MISSA), Theriot et al.
(1993)
THELYPTERIDACEAE
*Macrothelypteris torresiana (Gaudich.) Ching [=
Thelypteris torresiana (Gaudich.) Alston], IF,
bottomland hardwood forests, R; Theriot et
al. (1993)
WOODSIACEAE
Athyrium filix-femina (L.) Roth ex Mert. ssp.
asplenioides (Michx.) Farw., IF, mesophytic
upland hardwood forests, C; 460,659 (R460),
H.B.B.s.n. (MISSA), Jones et al. (196
Onoclea sensibilis L., IF, bottomland hana
forests, mesophytic upland hardwood for
oe fo a eee
a
PINOPHYTA
CUPRESSACEAE
— virginiana L. var. virginiana, BP, PR, IF,
bottomland hardwood forests, mesophytic
ca hardwood eee pine forests, pine-
mixed hardwood forests, prairies, prairie ce-
dar woodlands, chalk outcrops, R/AA; 6
1605, Bryson (1980), Henry s.n.(MISSA), be
and McDaniel (1998), Morris et al. (1993),
Theriot et al. (1993)
PINACEAE
Pinus echinata Mill, IF, pine forests, pine-mixed
hardwood forests, O/C; 1377a
Pinus aoe L., BP IF, bottomland hardwood for-
ests, pine forests, pine-mixed hardwood for-
ests, prairie cedar woodlands, chalk out
O; Leidolf and McDaniel (1 a eae et ne
(1993)
TAXODIACEAE
Taxodium distichum (L.
ests, C; Leidolfs.n.
—
L.C. Rich, IF, swamp for-
MAGNOLIOPHYTA
MAGNOLIOPSIDA
ACANTHACEAE
a setae (Pursh) Spreng, IF, bottom-
and hardwood forests, A; Smith 946
Justicia ovata (Walter) sea var. lanceolata
(Chapm.) R.W. Long, IF, bottomland hard-
wood forests, swamp on canals/drain
age ditches (riparian—banks), seepage areas,
CU) 359, 336, . Theriot et ‘ i a
II F.Gmel.) Steud., BP. IF, bot-
tomland sa forests, prairie cedar
woodlands, seepage areas, roadsides, O/C;
496, 1561, Leidolf and McDaniel (1998),
Theriot et al. (1993)
Ruellia humilis Nutt., BP, prairies, grass/forb
ws, O/C; 366, Leidolf and McDaniel
98), McDaniel 13447, Morris et al. (1993)
Ruellia strepens L., BP, PR, mesophytic upland
hardwood forests, roadsides, O/C; 278,
McDaniel 31326, Morris et al. (1993)
ACERACEAE
Acer barbatum Michx., PR, mesophytic upland
hardwood forests, C(I); 1337, Morris et al.
(1993
Ac ered: undo L.,BPPR,IF bottomland hardwood
s,rivers/creeks (riparian—banks), O; 226,
Askew 25, Morris et al. (1993), Theriot et al.
(1993)
Acer rubrum L., PR, IF, bottomland hardwood for-
ests, mesophytic yes hardwood forests,
pine forests, pine-m hardwood forest
C;25,35, 129 (R25), en. etal. aoe a
et al. (1993)
MARANTHACEAE
Amaranthus albus L., BP. cultivated fields; Mac-
Donald 10885
Amaranthus hybridus L., BP, cultivated fields, C(I)/
A; 896, Bryson 34486, Gasparini 201, McDaniel
28083
Amaranthus rudis J.D. Sauer, BP. chalk outcrops,
cultivated fields, roadsides, C(I)/A; 68
ay on 3450, McDaniel 22196, 31332
er spinosus L., BP, IF, cultivated fields,
grass/forb meadows, C(1); 897, Clonts 1406
Amaranthus viridis L., BP, cultivated fields, C(I);894
i
ANACARDIACEAE
Rhus aromatica Aiton, BP, mesophytic upland
hardwood forests, prairies, prairie cedar
woodlands, chalk outcrops, A; 186, 389, 788
(R186), 1327 (R389), 1342 (R186), Leidolf and
McDaniel (1998), MacDonald 7528, McDaniel
25016,Pullen et al. (1 968b), Smith 1410, 1896,
1936
Rhus copallinum L., \F, bottomland hardwood
forests, mesophytic upland hardwood for-
ests, pine forests, O; 445, Theriot et al. (1993)
Rhus glabra L.,BP. PR, IF, pine forests, prairies, pral-
rie cedar woodlands, roadsides, A(|); 301, 644,
Leidolf and McDaniel (1998), McDaniel 13464,
31647,Morris et al. (1993)
Rhus radicans L. [= Toxicodendron radicans (L.)
Kuntze ssp. radicans], PR, IF, bottomland hard-
wood forests, pine forests, pine-mixed hard-
wood forests, C; 472, Leidolf and McDaniel
(1998), Morris et al. (1993), Theriot et al.(1993)
Rhus toxicodendron L. [= Toxicodendron
pubescens Mill.], BP. prairie cedar woodlands,
ANNONACEAE
Asimina triloba (L.) Dunal, BP, PR, IF, bottomland
hardwood f mesophytic upland hard-
wood forests, A(l) 120, 353 (R120), 666, Askew
31, Gordon 2059 (MMNS), Leidolf s.n., Morris
t al. (1993), Theriot et al. (1993)
APIACEAE (UMBELLIFERAE)
Bowlesia incana Ruiz & Pav., PR, urban areas, A(I);
1215, Maddux 1287
158, Lo
Peat ie (Pers.) Sprague ex
Britton & Wilson, BP cultivated fields;
yson 1349
Cicut tamac ulata L., IF, gees areas, A(l); 1543
is (L.) DC., BP PR, bottom-
ane hardweod ee mesophytic upland
hardwood forests, pine-mixed hardwood
forests, O; 308, 391, 1
Cynosciadium digitatum DC., BP, roadsides
Sundell 1542
—
Fy i 1oc yk BP. roadsides Cc:
“Daucus carota L., BRIF, prairies, roadsides, C; 739,
Leidolf and McDaniel (1998), Pullen et al.
(1968b)
Eryngium prostratum Nutt.ex DC.,IF, bottomland
mesophytic upland hard-
| \ardwood f west
BRIT.ORG/SIDA 20(2)
wood forests, pine forests, lakes/ponds/im-
poundments (riparian—mudflats), seepage
areas, U; 334,498, 561, 1585 (R334), Bryson 674,
Copeland s.n.
Eryngium yuccifolium ae var. yuccifolium, IF,
seepage areas, R; 566, 6
Osmorhiza longistylis (Torr) i PR,$3;Green s.n.
(MISSA)
Perideridia americana (Nutt.ex DC.) Reichenb., BP.
prairies $1/S2; Brooks 150
Polytaenia nuttallii DC., PR, 82; Baker 57 (MISSA),
Lott s.n. (MISSA
Ptilimnium capillaceum (Michx.) Raf., BP, IF, bot-
tomland hardwood forests, mesophytic up-
nd har yee forests, canals/drainage
ditches (riparian—banks), seepage areas, O/
2,464, Bryson 719, Clonts 818
imnium nuttallii (OC.) Britt., BP, IF, seepage ar-
ea s/forb meadows, urban areas, O; 500,
ond /289, McDaniel 13494
Sanicula canadensis L., BP. PR, IF, bottomland
hardwood forests, mesophytic upland hard-
wood forests, prairie cedar woodlands, O;
302, Bryson /20,Leidolf and McDaniel (1998),
Lowe (1921), Morris et al. (1993), Theriot et
al. (1993)
Sanicula odorata (Raf) K.M. Pryer & LR. Phillippe,
mesophytic a hardwood forests, O;
Morris et al. (19
Sanicula smallii EP. a ecel IF, ee up-
land hardwood forests, O; 3
ee is inermis (Nutt.ex DC. ; ae s&Con-
e, BP, prairies, O/C; 1512, MacDonald
2
Ptill
a
Taenidia integerrima (L.) Drude, PR, mesophytic
upland hardwood forests, $1; Gordon 2034
Thaspium barbinode (Michx.) Nutt., PR, meso-
phytic upland hardwood forests, C; Gordon
2037
Thaspium trifoliatum (L.) A. Gray, BP, PR, IF, bot-
tomland hardwood forests, rivers/creeks (ri-
parian—banks), O; 356, Lowe (1921), McDaniel
24966, Morris et al. (1993), Theriot et al.(1993)
. Torilis arvensis (Huds.) Link, BP, PR, prairies, road-
1502, Clonts 816, McDaniel 13461,
ae, et al. (1968b)
sia ae aethusae Nutt. ex DC., BP, canals/
e ditches ae banks), seepage
ees ee Lowe (1921)
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI
Zizia aurea (L.) W.D.J.Koch, BP, bottomland hard-
wood forests, O; 10
pEOOUNA EAE
McDaniel in ed. BP Is/d
age ditches wee ae peepedé areas,
A(l); 209, McDaniel 2
Amsonia se anc Walter, IF, bottom-
land hardwood forests, O; Leidolf pers. obs.,
Theriot et al. (1993)
Apocynum cannabinum L., BP IF, prairies, pine
forests, A(I); 1517, Brooks 130, McDaniel 13463
Trachelospermum difforme (Walter) A. Gray, IF,
bottomland hardwood forests, pine forests,
C(l) 1497, Brooks 397, Theriot et al. (1993)
*Vinca major L., BP urban areas, A(|); 1448
eidebeae angie
“*/lex cornuta Lindl. & Paxton, PR, mesophytic
upland hardwood aa 887
llex decidua Walter, BP, PR, IF, bouomiend hard-
wood forests, mesophytic upland hardwood
forests, pine forests, pine-mixed hardwood
forests, prairies, prairie cedar woodlands,
chalk outcrops, roadsides, C; 38, 40, 946, Askew
69a, Clonts s.n., Leidolf and McDaniel (1998),
Morris et al. (1993), Theriot et al. (1993)
llex opaca Aiton, BP, IF, bottomland hardwood
forests, mesophytic upland hardwood for-
ests,C; 951, Laird 22, Theriot et al. (1993)
ARALIACEAE
Aralia spinosa L., IF, bottomland hardwood for-
ests, pine forests, pine-mixed hardwood for-
ests, roadsides, C(I); 586, Theriot et al. (1993)
Panax quinquefolius L.,BP, PR, mesophytic upland
hardwood forests, R, $3; Bryson 8830, Morris
et al. (1 Sees ith 1415
ARISTOLOCHIACEAE
Aristolochia tomentosa Sims, IF, bottomland
hardwood forests, rivers/creeks (riparian—
banks), U; Smith 542a, Theriot et al. (1993),
Timme 5435
Asarum canadense L., PR, mesophytic upland
hardwood rests C, $253; Harrelson 21
(MISSA)
ASCLEPIADACEAE
Asclepias amplexicaulis Small, IF, xeric upland
hardwood forests, R; 1546
ne hil Penne) Woodson, IF, pine for-
roadsides, C(|), $2:
1563,Bryson 681,6001, 7000, Ray 4751 (MISSA),
Stauffer s.n. (MISSA)
Asclepias purpurascens L., $1; Easley s.n. (MISSA)
Asclepias tuberosa L., BP, IF, prairies, pine forests,
roadsides, C(|); 291, 853, 1536, Bryson 679, 703,
Leidolf and ee (1998), McDaniel 31093,
Morris et al.
ariega a roadsides; Copeland s.n.
cea ee L., IF, bottomland hardwood
sophytic upland hardwood for-
ae ie U; 919, Clonts 827,Gordon ne
Pee ye Raf., BP, prairies, R/C; /
1526 2, Leidolf and McDaniel Hes
McD ae ae 098, Morris et al. (1993)
ree viridis Walter, BP, IF, prairies, chalk out-
crops, roadsides, C/A; 204, 1457 (R204), 1525,
Brooks 148, Carter 3017,Leidolf and McDaniel
(1998), McDaniel 29757, 31035, 31067, Morris
et al. (1993), Sundell 1536
Matelea gonocarpos (Walter) Shinners, BP, PR, IF,
bottomland hardwood forests, pine-mixed
hardwood forests, xeric upland hardwood
ie eae fields, R; 1527, 1574, Bryson
2599, 0, McDaniel 31066, Theriot et al.
nec
ASTERACEAE (COMPOSITAE)
“Achillea millefolium L., BP, prairies, O; McDaniel
cle
13492
Acmella oppositifolia (Lam.) R.K. Jansen var. repens
(Walter) R.K. Jansen, PR, urban areas, RR;
McDaniel 12494
Ambrosia artemisiifolia L., IF, bottomland hard-
ood forests, roadsides, C/A(|); 984, Brooks
ae ent et al. (1993), Wigley 314085
a Michx., IF, roadsides, C; 975
Aaibiedtnde sachs DC., BPIF, pine forests, prai-
res, ee cedar woodlands, chalk outcrops
C/ 9, Leidolf and McDaniel (1998),
a 15463, 22929
ee ae L., BP prairies, urban areas, C/A;
ae oa 468, McDaniel 24964, Wigley
3:
>
Am onc is fede culoides ve ) Nutt., BP.
ries, AA(I), $1; McDaniel 22
ee aria plant ee (L.) aon BP IF,
xeric upland hardwood forests, roadsides, O;
1262, 1535
“Arctium minus Bernh., BP, PR, cultivated fields,
urban areas, O; 706, Bryson 3078
Aster attenuatus Lindl. ex Hook.,, BP, prairies, O/
C(I); 1015, MacDonald 8145
Aster aes L. var. sagittifolius (Wedem. ex
Willd. es, PR, bottomland hardwood
ee i ors et al. (1993)
PR, IF bottomland hardwood
6 rests, pine pai prairies, C; 961, 991, Co-
peland s.n., Leidolf and McDaniel (1998),
Morris et al.(1993), Theriot et al. (1993), Wigley
Aster ericoides L., BP, prairies, C(I), $2; 1004, 1009a,
1011,Leidolf and McDaniel (1998), MacDonald
8142, 8152
Aster laevis L., BP, prairies, O; 933, Leidolf and
McDaniel (1998)
Aster novae-angliae L., BP, prairies, cultivated
fields, roadsides, O/C; / 1007, 1019
Gasparini 208, Leidolf and McDaniel (1998),
McDaniel 1044
Aster paludosus Aiton ssp. hemisphericus (Alex-
ander) Cronquist, BP IF, pine forests, prairies
seepage areas, roadsides, O/A(I); 650, 654,
Brooks 410, Bryson 6986, Clonts 571,Copeland
$n.
Aster patens Aiton, BP, IF, pine forests, prairies,
chalk outcrops, O; 968, Leidolf and McDaniel
(1998), Morris et al. (1993)
Aster pilosus Willd. var. pilosus, BP IF, prairies, culti-
vated fields, O; 990, Leidolf ee oo
1998), Gasparini 207, Wigley 313
Aster praealtus Poir. var. ate a prairies,
C; 1009, 1012, Leidolf and
McDaniel (1998), Macl ld 814
Aster sericeus Vent. var. microphyllus Dc. BP. prai-
ries, R/C(I); 1073, Leidolf and McDaniel (1998),
MacDonald 8147
Aster pee cane Michx., IF, pine forests, pine-
d hardwood forests, seepage areas,
roadsides Cl );349, 529, 1494, 1540, Bruza 929,
Bryson 684, Clonts 831
Aster subulatus Michx. var. ligulatus Shinners, PR,
urban areas; oe 12528
Aster undulatus | prairies, prairie cedar wood-
lands, rte U/C(I); 1003, 1008, 1014,
Leidolf and McDaniel (1998), MacDonald
8143, 8148, Gis}
=
—
7.., BP. IF, bottomland hard-
wood forests, lakes/ponds/impoundments
(riparian—banks), roadsides, C(I); Clonts 472,
BRIT.ORG/SIDA 20(2)
McDaniel 31324, 31325, Theriot et al. (1993),
Wigley 313120
Bidens aristosa (Michx.) Britton, BP. IF, bottomland
hardwood forests, pine forests, roadsides, O;
873, Brooks 482, Copeland s.n., Theriot et al.
(1993), Wigley 314083
sea discoidea (Torr. & A. Gray) Britton, IF, bot-
mland hardwood forests, — forests;
Seat a eles et al. (1993)
Elliott, IF, pine forests, roadsides
664, Copeland s.n.
ie eupatorioides (L.) Shinners var.
eupatorioides, BP, IF, chalk outcrops, C(1);
McDaniel 2193, 31310
Brintonia discoidea (Elliott) Greene, IF, bottom-
land hardwood forests, pine forests, road-
sides, O; 922,Theriot et al. (1993)
Cacalia ovata Walter [= Arnoglossum ovatum
(Walter) H.Rob.], IF mesophytic upland hard-
wood forests, O; 902
Cacalia plantaginea (Raf.) Shinners [=
Arnoglossum plantagineum Raf], BP, prairies,
R/A; 375, 1445 (R375), Leidolf and McDaniel
(1998), McDaniel 13456, 31074, Morris et al.
(1993)
*Centaurea cyanus L., PR, urban areas, RR;
McDaniel 13442
Chrysopsis mariana (L.) Elliott, IF, mesophytic
upland hardwood forests, roadsides, O/C;
ake Mathies 867
intybus L., BP, roadsides, RR; 680
Cirsi er scolar ex Willd.) Spreng., BP, prai-
ries, chalk outcrops, lakes/ponds/impound-
ments (riparian—banks), O; 823, 832, Wigley
313092
Cirsium horridulum Michx., BP. IF, prairies, prairie
cedar woodlands, roadsides, O; 1438, 1455,
Eee tate bce ae = re
*
s,chalk outcrops
fae dee C(I); “ wel en Leidolf and
McDaniel (1998)
wy pia | Exe £
Dral-
ries, roadsides; 343, Brooks 122, 143, McDaniel
1089
Coreopsis tinctoria Nutt., IF, roadsides, urban ar-
s,C; 1573, Copeland s.n.
Coreopsis tripteris L., IF, roadsides, C(I); 658, Brooks
494
*Crepis pulchra L., BP, urban areas, O; 1450
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI
Dracopis amplexicaulis (Vahl) Cass., BP. IF, prairies
roadsides, C; 289, 396, Sa ae Leidolf
and McDaniel (1998), Sundell 15
pes purpurea (L.) Moench, PR, on upland
ardwood forests, $3$4; Ray 4715 (MISSA)
Eclipta Reece .) L, BP IF, bottomland hard-
WwW forests, pine forests, canals/drainage
ae (riparian—banks), lakes/ponds/im-
poundments (riparian—mudflats), seepage
areas, cultivated fields, C/A; 609, Brooks 481,
Bryson 3423, McDaniel 31333, Theriot et al.
(1993
Elephantopus carolinianus Raeusch., IF, bottom-
land hardwood forests, pine forests, road-
sides, O/A; 806, 877, Lowe (1921), Smith 945,
Theriot et al. (1
Elephantopus tomentosus L., IF, mesophytic up-
land hardwood forests, pine forests, pine-
mixed hardwood forests, lakes/ponds/im-
oundments (riparian—banks), C; 1582,
Brooks 488, Carraway 174, Meanie! 9797,
15726
Erechtites hieracifolia (L.) Raf. ex DC., IF, bottom-
land hardwood forests; Theriot et al. (1993)
Erigeron annuus (L.) Pers., BP, PR, roadsides, O/A;
ter 3
\O
Ne)
SS)
—_
Erigeron canadensis L.[= Conyza canadensis (L.
ronquist var Grades BP, PR, IF, oe
roadsides, C; 551, 1597, McDaniel 31625,
Wigley 313062
g L fe L.IF, see] O;138
Erigeron strigosus Muhl. ex Willd., BP, IF, bottom-
land hardwood forests, xeric upland hard-
wood forests, prairies, chalk outcrops, O; 203
470, Bryson 716, Clonts 828, Leidolf and
McDaniel (1998
Eupatorium altissimum L., BP, as chalk out-
crops, roadsides, O/ Leidolf and
McDaniel (1998), Pee 20934, 22928,
28077, 31649
eee capillifolium (Lam.) Small, lF, pine for-
s,urban areas; Copeland s.n., Snow 39
ae ae coelestinum L., BP, IF, bottomland
hardwood forests, pine forests, chalk out-
crops, lakes/ponds/impoundments (ripar-
ian—banks), roadsides, C/A(I); 528, 958, Bryson
706, Lowe (1921), McDaniel 31319, Wigley
313098
cuties
Eupatorium hyssopifolium LIF, pine forests,O; 700
Eupatorium incarnatum Walter, BP, C; McDaniel
31322
ee pee L., IF, pine forests, O/C;
11, Brooks 475, Copeland s.
oe ae L, IF, grass/forb
ws, roadsides, A; 471, Clonts 841,
Mathies 841
Eupatorium rugosum Houtt. [= Ageratina
altissima (L.) King & H. Rob. var. al/tissima], BP,
IF, bottomland hardwood forests, C; 972,
McDaniel 31328, Theriot et al. (1993)
Eupatorium scabridum Elliott [= £. rotundifolium
L.var.scabridum (Elliott) Gray], IF, pine forests,
roadsides, 0,655, Wigley 31408
BHO semen DC,, IF, gan
pine forests,O;55
Wigley 314087
Eupatorium serotinum Michx., BP. IF, pine forests,
chalk outcrops, seepage areas, C(I)/A; 683,
Copeland s.n Daniel 22192, 2293?
Euthamia leptocephala (Torr. & A. Gray) Greene,
IF, pine forests; Wigieyeaetl OS
Euthamia tenuifolia (Pursh) Nutt., PR, IF, pine for-
ass/forb meadows, O; 848, 875
Evax oa ae Nutt. ex DC., BP, prairies, AA(I), $1;
195, 1442 (R195), Leidolf and McDaniel (1998)
*Facelis retusa (Lam.) Sch. Bip., PR, IF, roadsides,
urban areas, C(|); 1317, 1504
Gnaphalium helleri Britton, IF, pine forests, O; 955
Gnaphalium obtusifolium L IF, bottomland
hardwood forests, pine forests, R/O; McDaniel
1560, 31644, Theriot et al. (1
Gnaphalium purpureum L. [= Gamochaeta
pulputed . .) Cabrera], PR, IF, pine-mixed
ts, seepage areas, roadsides
, 1290, McDaniel 15072,
eee areas, O/C: 1
Wigley 313126
Helenium amarum (Raf.) H. Rock, BP IF, roadsides,
394, ee eae
Heleniu ale L., BP.IF, bottomland hard-
wood Forests pine forests, oie esekls
woodlands, chalk outcrops
O; 830, Copeland s.n., pee 209, Leidolf
and McDaniel (1998), Hees 1435, 15731,
31315, Theriot et al. (19
Helenium flexuosum Raf,, IF, ane upland
hardwood forests, O; 444, McDaniel 2540
Helianthus ea a L., IF, roadsides, A(I);872,
Mathies 840
730
Helianthus annuus L., BP, roadsides, A(I); 397
Helianthus divaricatus L., BP, prairie cedar wood-
lands, O; 427
Helianthus giganteus L., IF, roadsides, O; 881
Helianthus hirsutus Raf, IF, pine forests, grass/forb
ws, O: 577, Clonts 840
Helianthus mollis Lam. IF, pine forests, O;589, 699
Helianthus silphioides Nutt., IF, pine-mixed hard-
od forests, O; 808, Brooks 489, McDaniel
Hr
He gees an BP prairies, roadsides, O
927, Leidolf and McDaniel (1998), McDaniel
15 a
Heliopsis gracilis Nutt., IF, mesophytic upland
hardwood forests, O; 336
Heterotheca camporum (Greene) aha _ BP,
prairies, chalk outcrops, ae 841, 1596,
Leidolf and McDaniel (1 se ends
8146, McDaniel 22925, Morris et al. (1993
Hieracium gronovii L., IF, pine forests, urban ar-
648, McDaniel 2195, 2196
Iva annua L., BP, IF, prairies, roadsides, A(l); 992,
Copeland s.n., McDaniel 31630
Krigia dandelion (L.) Nutt., PR, IF, roadsides, O; 1
Askew 48
Krigia cespitosa (Raf.) K.L.Chambers, BP PR, culti-
ated fields, roadsides, O; 179, Bryson 3499
Lactuca canadensis L., BP. IF, xeric pein nard-
wood forests, prairies, lak
poundments (riparian—banks), aes fe
Carraway 175,McDaniel 30273, Wigley 312049
Lactuca floridana (L.) Gaertn., BP, IF, bottomland
hardwood forests, prairie cedar woodlands,
roadsides, O; 838, Leidolf and McDaniel
(1 dau Theriot et al. (1993)
Lactu ita Muhl. ex Nutt., IF; Bryson 677
*Lactuca see L.,BP.PR, xeric upland hardwood
forests, prairies, urban areas, O; MacDonald
897, McDaniel 30271
*Lactuca serriola L., PR, urban areas, O; 1598
*Leucanthemum vulgare Lam., BP. urban areas,
C(I); 1447, 1463
Liatris aspera Michx., BP. IF, prairies, roadsides, A;
516, 615 (R516), Copeland s.n., Leidolf and
aniel (1998)
Liatris spicata (L.) Willd., BP, IF, pine forests, pine-
mixed hardwood forests, prairies, R/U; 515,
563,Clonts s.n., Leidolf and McDaniel (1998),
niel 2660
Liatris squarrosa (L.) Michx., BP, prairies, chalk out-
BRIT.ORG/SIDA 20(2
R/A; 370, Clonts s.n., Leidolf and
nea (1998), McDaniel 15505, Morris et
al. (1993)
Liatris squarrulosa Michx., BP, IF, prairies, chalk
outcrops, roadsides, R/O; 969, Leidolf and
aniel (1998), McDaniel 229274
Mikania scandens (L.) Willd., BP, IF, bottomland
hardwood forests, chalk outcrops, lakes/
ponds/impoundments (riparian—banks),
roadsides, C/A(I); 585, Brooks 479, McDaniel
2216/7, Theriot et al. (1993), Wigley 314084
Pityopsis graminifolia (Michx.) Nutt., IF, seepage
areas, C; 868
—
Pluchea peeing .) DC, BP, IF, bottomland
hardwood forests, See eas ainenes
eee Ceiey
ments (riparian- silences eens areas, 5C
849, Theriot et al. (1993), Wigley 313100
Prenanthes altissima L., PR, mesophytic upland
hardwood forests, O; Morris et al. —
Prenanthes aspera Michx., BP, 2;
Leidolf and McDaniel (1 998), pene 15622
Bee carolinianus (Walter) DC., IF, pine
for
pyopopp ay iflorus (D. Don) DC., BP, prai-
oadsides, O; McDaniel 31072, Sundell
*Ratibida columnifera (Nutt.) Wooton & Stand,
' é 5 6.
Ratibida pinnata 8 Barnhart, BP, prairies,
prairie cedar woodlands, chalk outcrops,
roadsides, O; a nee s.n., Leidolf and
McDaniel (1998), McDaniel 13445, Morris et
al. (1993)
Hier fulgida Aiton, BP. prairies, roadsides, O;
44, Leidolf and McDaniel (1998)
1 ¢
ests, xeric upland hardwood forests, prairies,
A; 282, Bryson 723, Clonts 823, eee and
ie (1998), Hier 619
Rudbec ie laciniata L.,|F, bottomland nen
forests; Brooks 564
a il Wood, BP IF, pine forests, road-
ides areas, C;255,1441,1449 McDaniel
13216, 29753, 31031
Senecio glabellus Poir., PR, IF, bottomland hard-
hase forests, mesophytic upland hardwood
fores ea ae C; 133, 1372, Smith 1932,
ae et al. (19
Senecio obovatus ae ex Willd., BP, chalk out-
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI
crops, O; McDaniel 3
et al. (1993)
Silphium integrifolium Michx., PR, roadsides, C;
Leidolf pers. obs. (with 1001)
Silphium laciniatum L., BP, chalk outcrops, grass/
forb meadows, roadsides, C; 392, Clonts s.n.,
Leidolf and McDaniel (1998), Morris et al.
(1993
Silphium laciniatum L. x 7 een
Jacq., BP, roadsides, RR; 8
Silphium perfoli atumL., BP IF, cae (ripar-
ian—banks), roadsides, O; 625, 687, McDaniel
25285, 28877
Silphium perfoliatum L. x S. trifoliatum L. var.
latifolium A. Gray, BP, roadsides, RR: 83
Silphium terebinthinaceum Jacq., BP, PR, bottom-
land hardwood forests, prairies, O;843, Leidolf
and McDaniel (1998), Morris et a
Silphium trifoliatum L. var. latifolium A Gray, BP.
prairies, chalk outcrops, C;517,622, McDaniel
15502
olidago caesia L.,|F, bottomland hardwood for-
ests, mesophytic upland hardwood forests,
ie 953, slener oe (1993)
lidag ar. scabra Torr.& A. Gray,
BP. IF, pine forests, nal outcrops, roadsides,
O/C; ee i s.n., McDaniel 28076,
Wigley 3
Solidago gi porta Aiton, BP, IF, pine forests, prai-
ries, chalk outcrops, O; 824, Brooks 316
McDaniel 313
Solidago hispida a ex Willd. IF, pine-mixed
hardwood forests, C; McDaniel 14665
Solidago juncea Aiton, IF; Wigley 312050
Solidago nemoralis Aiton var.haleana Fernald, IF;
Bryson 678
1032, Morris 3097, Morris
Soli adage pemonaits pun Y Var. nemoralis, BP. IF
s, chalk
outcrops, eo cae (ri-
parian—banks), roadsides, O/C; 792, Carraway
176, Leidolf “nd McDaniel (1998), Theriot et
al. (1993)
Solidago odora Aiton var. odora, IF, pine forests,
O/C; 962, Copeland s.n., McDaniel 2192
Solidago ho ale Aiton, IF, pine forests, C;
McDaniel
Solidago ie L, C prairies, C; 790, Leidolf and
McDaniel (19
Solidago rugosa Ait ssp. aspera (Aiton)
Cronquist, IF, pine-mixed hardwood forests,
roadsides, O/C; ee s.n., Mathies 844,
Mebane 9795, 1
Solida ifolia ates ex Willd. BP IF. bottom-
land bardwasd forests, pine forests, pine-
mixed hardwood forests, prairies, O/C; 596,
te Leidolf and McDaniel (1998), McDaniel
31631,Morris et al.(1993), Theriot et al. ee 3)
*Soli iva sessilis Ruiz & Pav.,PR, urban areas, — 214
*Sonchus asper (L.) Hill, BP roadsides, O; 1339, 1464
*Sonchus oleraceus L., BP roadsides, O; 1459
*Taraxacum officinale Weber ex FH.Wigg., BP, PR,
bottomland hardwood forests, prairie cedar
woodlands, roadsides, urban areas, O; 1268
Leidolf and McDaniel (1998), Morris et al.
(1993), Wigley 313111
Verbesina alternifolia (L.) Britton ex Kearney, IF,
Resell hardwood forests, roadsides, C;
73,1537, Brooks 563, Theriot et al. (1993)
ee ina helianthoides Michx., IF, pine forests,
roadsides, C(l); Brooks 277, Blea 936
Verbesina iothica L., BP. chalk outcrops, lakes/
ponds/impoundments (riparian—banks),
roadsides, O/A; 836, McDaniel 1491, 31321
Vernonia gi as (Walter) Trel. ssp. gigantea, BP,
PR, IF, bottomland hardwood forests, pine
or. meadows, roadsides, O/C;
578, 628, Bruza 1001, McDaniel 22930, Morris
et al. (1993)
*Xanthium strumarium L., BPIF, lakes/ponds/im-
ments (riparian—banks), cultivated
Dayar meadows,R/A(I);819,Clonts
404, McDaniel 31329, Wigley 313099
oe ica (L.) DC., BR. urban areas, O; 7520
BALSAMINACEAE
Impatiens capensis Meerb.; Jones (1976b)
BERBERIDACEAE
ate bealei (Fortune) Carriére, PR, meso-
sland hardwood forests, R;886, 1307
oa ina vane ica Thunb., PR, mesophytic
upland Seles forests, R; 889
Podophyllum pelta IF, mesophytic up-
land Bdwoord Hers AA(); 88, 99, 1308,
Askew 41, Morris et al. (1993)
BETULACEAE
Betula nigra |
rivers/creeks (riparian—banks), lakes/ponds/
impoundments (riparian—banks); Anderson
s.n. (MISSA), Hansbrough s.n. (MISSA)
IF | 1 f ic
732
BIGNONIACEAE
Bignonia capreolata L., BP, PR, IF, bottomland
od forests, mesophytic upland hard-
wood forests, O/C; 134, Askew 60, McDanie
2355,Morris et al. (1993), Theriot et al. (1993)
Campsis radicans (L.) Seem. ex Bureau, BP, PR, IF,
bottomland hardwood forests, pine-mixed
hardwood forests, xeric upland hardwood
forests, prairie cedar woodlands, C; 300, 345,
Leidolf and McDaniel (1998), Morris et al.
1993), Theriot et al. (1993)
Catalpa bignonioides Walter, IF, rivers/creeks (ri-
parian—banks), roadsides, O; 1492, Clants 748
BORAGINACEAE
*Buglossoides arvensis (L.) ILM. Johnst., BP, road-
sides, A(l); 144
Cynoglossum virginianum L., PR,mesophytic up-
land hardwood pier A(l); 1332, Jones
(1976b), Morris et al. (19
*Heliotropium indicum L., s bottomland hard-
wood forests, pine forests, urban areas; Clonts
1405, Copeland s.n., Jones (1976b), Theriot et
al. (1993)
Heliotropium tenellum (Nutt.) Torr, BP, prairies,
chalk outcrops, O/A; 368, Jones (1976b),
Leidolf and McDaniel (1998), MacDonald
7529, McDaniel 15507,31088,31313,Morris et
al. (1993), Pullen et al. (1968b)
mecadba canescens (Michx.) Lehm., BP, prai-
ries, chalk ou eee grass/forb meadows,
roadsides, O/C(l); 210, 1269, Bryson 414, Jones
(1976b), Leidolf oe McDaniel (1998),
McDaniel 31015, Morris 3096, Morris et al.
(1993)
Lithospermum latifolium Michx., PR, mesophytic
upland hardwood forests; Gordon 2064
Lithospermum tuberosum Rugel ex DC., PR, me
sophytic upland hardwood forests, O/C;
31, Jones (1976b), Smith 1933
Myosotis macrosperma Engelm., PR, mesophytic
upland hardwood forests, O; 1334, 1352,
Jones (1976b)
Myosotis verna Nutt., BP, cultivated fields; Bryson
3484, Jones (1976b)
Onosmodium molle Michx. ssp. Aispidissimum
(Mack.) B. Boivin, BP, prairies, O; 1603, Jones
(1976b), McDaniel 31064, Morris et al. (1993)
BRASSICACEAE (CRUCIFERA
*Arabidopsis thaliana (L.) ok Jones (
oe
—
wie
—
—_
975a)
BRIT.ORG/SIDA 20(2)
Armoracia lacustris (A.Gray) Al-Shehbaz & Bates,
BP, grass/forb meadows, O, $182; McDaniel
25
*Brassica erucastr
(W fe
um L. [= Erucastrum gallicum
O.E. Schulz], BP chalk outcrops, O/C;
1461, McDaniel 24994, Smith 1448
i ier ies (L.) Czern., PR, urban areas, RR;
*Brassica napu PR roadsides, RR: 1260
*Capsella ees pastoris (L.) Medik., BP, PR, culti-
vated fields, Rouen urban areas, C; 63,
Bryson 3488, Stew
Cardamine ie (ane ex Muhl.) Britton,
Sterns & Poggenb,, PR, IF, bottomland hard-
wood forests, mesophytic upland hardwood
See ey ee meadows, O; 90, 1333,1377,
w 45, Jones (1975a)
Eres ine concatenata (Michx.) Sw., PR, meso-
phytic upland hardwood forests, O; 157, Mor-
ris 3091, Morris et al. (1993)
*Cardamine hirsuta L.,PR, urban areas, A; 23, 1209,
1975a), Stewart 145
Cardamine parviflora L.; Jones (1975a)
*Coronopus didymus (L.) Small, PR, IF, pine forests,
irban areas,O; 1205, Copeland and Warren s.n.
Draba brachycarpa Nutt. nae & A. Gray, BP pral-
ries, urban areas, A; /
Draba cuneifolia Nutt. ex ot & A. Gray, BP, prai
5, C(I); McDaniel 10415
*lepi ae perfoliatum L., BP, urban areas, RR; Ray
s.n. (MISSA)
Lepidium ie inicum L., BP, ie cultivated
fields, roadsides, A; 199, 395, 1287
raphanistrum L., i roadsides, O; 76
Rorippa sessiliflora (Nutt.) Hitchc., BP PR, canals/
drainage ditches (riparian—banks), cultivated
fields, grass/forb meadows, urban areas
1305, Bryson 7420, McDaniel 24753, 25027
Qara virginica (L.) Rollins, BP, cultivated fields,
roadsides, A; 1272, 1279, Bryson 3490, Jones
sag 2d mph INUS
Si
pats
nh ri ffi -j [
—
mF
SiSY L.) Scop.; Jones (1975a)
*Thlaspi perfoliatum L., BP, roadsides, C; McDaniel
33025
BUDDLEJACACEAE
Polypremum procumbens L., BP. IF, xeric upland
hardwoad forests, canals/drainage ditches
(riparian—banks), seepage areas, cultivated
fields, roadsides, urban areas, C(|); 441, 1570,
Bryson 2612, McDaniel 1503
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI
CABOMBACEAE
Brasenia schreberi J.F. Gmel., IF, lakes/ponds/im-
poundments (littoral-rooted-floating), A(/)
580, 657
CACTACEAE
ee humifusa (Raf.) Raf., BP PR, grass/forb
roadsides, R; 873, 940
CALLITRICHACEAE
Callitriche oekae Pursh, IF, canals/drainage
s (submergent), lakes/ponds/im
Balnaaeee (littoral-submergent), seepage
areas, C(I); 126, McDaniel 10491
CAMPANULACEAE
ampanula americana L. [= Campanulastrum
americanum (L.) Small], BP PR, IF, bottomland
hardwood forests, mesophytic upland hard-
wood forests, xeric upland hardwood forests,
prairie cedar woodlands, roadsides, O/C(\);
624, 822, 1604, MacDonald 9720, McDaniel
28888, 31640, Morris et al. (1993), Smith 1411
Lobelia appendiculata A.DC.,IF, pine-mixed hard-
wood forests, roadsides, A(l), $253; 322, 342,
anaes
363, 364
Lobelia cardinalis L., IF, bottomland hardwood
forests, mesophytic upland hardwood for-
ests, rivers/creeks (riparian—mudflats), lakes/
ponds/impoundments (riparian—mudflats),
R/C; 812, 944, 980, Bruza 1332, Theriot et al.
Lobelia puberula Michx., BP, IF bottomland hard-
wood forests, mesaphytic upland hardwood
forests, xeric upland hardwood forests, pine
forests, pine-mixed hardwood forests, seep-
age areas roadsides, O; 524a,809, 957, Clonts
572, McDaniel 31621, Theriot et al. (1993)
Lobelia spicata Lam., BP, prairies, O; 295, Leidolf
Daniel (1998), McDaniel 15571, Mor-
ris et al. (1993)
Specularia biflora (Ruiz & Pav.) Fisch. & C.A. Mey.
= Triodanis perfoliata (L.) Nieuwl. var. biflora
(Ruiz & Pav.) Bradley], IF, bottomland hard-
ests, O; 230
Specularia perfoliata (L.) A. DC. [= Triodanis
perfoliata (L.) Nieuwl. var. perfoliata], BP, prai-
ries, roadsides, O; 1361, 1454, 1465, Brooks 155,
Lowe (1921), McDaniel 31013
CANNABACEAE
Cannabis sativa |
—
IF roadsides, RR: 80
733
CAPRIFOLIACEAE
*l onicera japonica Thunb., BP, PR, IF, bottomland
hardwood forest mesophytic upland hard-
wood forests, prairie cedar woodlands, pine
forests, pine-mixed hardwood forests, rivers/
creeks (riparian—banks), roadsides, C; 215,
Brooks 129, Leidolf and McDaniel (1998),
Morris et al. (1993), Theriot et al. (1993)
Lonicera sempervirens L., PR, xeric upland hard-
wood forests, R; 1366
Sambucus canadensis L., BP, PR, IF, bottomland
h oad forests, Lasii upland hard-
wood ie nage ditches (ripar-
eno ee et al.(1993), Theriot
et al.
Symphoricarpos orbiculatus Moench, BP, PR, me-
sophytic upland hardwood forests, xeric
upland hardwood forests, roadsides,
799, McDaniel 1501,31650,Morris et al.(1993)
Triosteum angustifolium L., PR, mesophytic up-
land hardwood forests, R/C, $3; Channell s.n.
(MISSA), Funchess s.n. (MISSA), Hull s.n.
(MISSA), Rainwater s.n. (MISSA), Ray 931
(MISSA), Smith 1934
ua oe Raf., BP. PR, bottomland hard-
est meSOe ye upland As eee
aa prairie cedar woodlands, O; /
160, Leidolf and eps ne! (1998), Hel et
al. (1993)
CARYOPHYLLACEAE
*Arenaria serpyllifolia L., BP, roadsides, A(I);
McDaniel 29754
*Cerastium brachypetalum Desportes, BP, road-
sides, C; 71a
*Cerastium glomeratum Thuill, BP, PR, es
fields, ae urban areas, C/A; 77, 1211,
pian
agina d a ns (Elliott) Torr. & A. Gray, BP, PR,
xeric upland hardwood forests, urban areas
COGNATE AB a7.
Silene antirrhina L., BP, IF grass/forb meadows,
dsides, O/A; 1485, Bryson 8862, Pullen et
al. (1968b)
Silene stellata (L.) W.T. Aiton, PR, IF, mesophytic
upland hardwood forests, O; 885, McDaniel
28885, Morris et al. (1993
pergu , BP, cultivated fields, A; 1286
*Stellaria media (L.) Vill, PR, urban areas, A; 70,
1272
¥CHprnys la arveonce |
734
CELASTRACEAE
Evonymus americanus L., BP, PR, IF, bottomland
hardwood forests, mesophytic upland hard-
wood forests, rivers/creeks (riparian—banks),
roadsides, O; 259, 945, Bryson 728, Morris et
al. (1993), Theriot et al. (1993)
Evonymus atropurpureus Jacq., PR, mesophytic
upland hardwood oa RR, $283; Morris
3086, Morris et al. (19
*Evonymus fortunei ae Hand. -Mazz., PR,
urban areas, RR; MacDonald 9994
CISTACEAE
Lechea mucronata Raf,, IF, grass/forb meadows,
O; 1586
Lechea tenuifolia Michx.,\F, pine forests, seepage
areas, roadsides, C(|), 485,573, 1534, McDaniel
13489
CONVOLVULACEAE
Cuscuta compacta Juss. ex Choisy; Anonymous
5.n. (MI
Cuscuta pentagona Engelm. var. pentagona, BP.
IF, pias leads meadows, A(|); 428, -
q., BP. cultivated fields
A; 821, Bryson 25 sop.
Ipomoea lacunosaL.,BP, cultivated fields, C/A;820,
ipomoea pandurata (L.)
fields, A; 519
*Ipomoea wrightii A. Gray, IF, cultivated fields, A(|);
711
Jacquemontia tamnifolia (L.) Griseb.,, IF, lakes/
/\ pourament: (riparian—mudflats),
oadsides, O: 598 Clonts
G, Mey., BP, cultivated
aos
CORNACEAE
Cornus drummondii C.A.Mey., BP PR, bottomland
hardwood forests, mesophytic upland hard-
wood forests, xeric upland hardwood forests,
prairie cedar woodlands, chalk outcrops,
roadsides, O; 274, 620, Bryson 8294, Carter
3005, Leidolf and McDaniel (1998), McDaniel
31068, Morris et al. (19
Cornus florida L., PR, IF, bottomland hardwood
forests, mesophytic upland hardwood for-
ests, pine forests, pine-mixed hardwood for-
ests, C; 124, 351, 964, Askew 39, Morris et al.
(1993), Theriot et al. (1993)
Cornus stricta Lam. [= C. foemina Mill.], IF, bottom-
BRIT.ORG/SIDA 20(2
land hardwood forests, O; 244, McDanie
Theriot et a 93)
Nyssa sylvatica Marshall, IF, bottomland hard-
Ww rests, pine forests, pine-mixed hard-
wood forests, C; 1482, Smith 2540, Theriot et
al. (19
CORYLACEAE
Carpi lini
11562,
\O
wood forests, C; 234
Ostrya virginiana (Mill.) K. Koch, PR, IF, bottom-
land hardwood forests, mesophytic upland
hardwood forests, O; Leidolfs.n., Morris et al.
(1993)
CRASSULACEAE
Sedum pulchellum Michx., BP, PR, chalk outcrops,
R, $1; Brent 181, Stauffer s.n. (MISSA)
CUCURBITACEAE
Cayaponia quinqueloba (Raf.) Shinners; Jones
(1976b), Pullen et al. (1968b
othria pendula L., BP, PR, i peel
oundments (riparian—banks), r
ae 724, Jones (1 Gee 31327
DROSERACEAE
Drosera brevifolia Pursh, IF, seepage areas, R; 1289,
1437 (R1289)
EBENACEAE
Diospyros virginiana L., BP, PR, IF, bottomland
hardwood forests, mesophytic upland hard-
wood forests, pine forests, pine-mixed hard-
wood forests, prairie cedar woodlands, chalk
outcrops, O; 1462, Leidolf and McDaniel
(1998), Morris et al.(1993), Theriot et al.(1993)
ELAEAGNACEAE
*Elaeagnus pungens Thunb., PR, mesophytic up-
land hardwood forests, O; 888
*Flaeagnus umbellata Thunb., PR, urban areas, RR;
Me
Q
ERICACEAE
Monotropa hypopithys L.,PR, mesophytic upland
es Moe : Morris et al. (1993)
Pa L F: Stewart 116
Rhododendron canescens (Michx.) Sweet, IF, bot-
tomland hardwood forests, mesoph
ytic
land eer anes forests, O/C; 125, 130, nee
40, pen Sh Tenet et eh (1993)
\ i BPIF bottomland
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI
hardwood forests, swamp forests, xeric up-
land hardwood forests, pine forests, O; 1359,
1550, Brooks 132, Carter 3020, Clonts 830,
McDaniel 1097, Theriot et al. (1993)
Vaccinium elliottii Chapm., IF, bottomland hard
wood forests, pine-mixed hardwood forests,
O; 36, Theriot et al. (1993)
Vaccinium elliottii Chapm. x V.fuscatum Aiton, IF,
ttomland hardwood forests, RR; Theriot et
al. (1993)
Vaccinium stamineum L., PR, IF, bottomland hard-
mesophytic upland hardwood
woo
ee,
wood forests
Brooks 120, Theriot et al. (19
Vaccinium virgatum Aiton, IF, bottomland hard-
wood forests; McDaniel 3000
ESCALLONIACEAE
[tea virginica L., IF, bottomland nareweed 1OFE
px} /
swamp for
(riparian— a O; 265, weDaniel 1828
EUPHORBIACEAE
Acalypha gracilens A. Gray, IF, seepage areas, O;
649
Acalypha ostryifolia Riddell, BP. IF, prairies, culti-
vated fields, roadsides, C _1544,Gasparini
203, Pullen et al. (1968b)
Acalypha rhomboidea Raf. [= A. virginica L. var.
rhomboidea (Raf.) A eee IF rivers/creeks
(riparian—banks), roadsides, O
Acalypha virginica L., BP, IF, chalk eutcroos lakes/
ponds/impoundments (riparian—banks),
roadsides, O; 600, 630, Brooks 485, McDaniel
22197, Pullen et al. (1968b)
CH eee Engel
outcrops, C; McDaniel 22201
Chamaesyce maculata (L.) Small, BP, PR, IF, chalk
outcrops, rivers/creeks (riparian—banks), cul-
tivated fields, roadsides, urban areas, C; 423,
544,690, McDaniel 2202, 22199
Chamaesyce nutans (Lag.) Small, BP, IF, rivers/
creeks (riparian—banks), canals/drainage
ditches (riparian—banks), cultivated fields, O
A; 504, | i) 469, Bryson 3443, MacDonald /,
McDanie
Chamaesyce Pe (Aiton) Small, BP PR, culti-
a fields, urban areas, C; Bryson 3445,
McDaniel 2703, Stewart 151
) Small, BP. chalk
735
Chamaesyce serpens (Kunth) Small, BP, chalk out-
op G McDaniel 22198
r Michx., BP. IF, pine-mixed hard-
wood forests, prairies, roadsides, C; 379, 565,
Brooks 323
Croton glandulosus L. var. septentrionalis Mull.
Arg., IF, pine-mixed hardwood forests, road-
sides, C(I); 538, Brooks 319
NAv-|
hx., BP. prairies, chalk
outcrops, cultivated fields, C/A(I); 367, Bryson
8315, Leidolf and McDaniel (1998), McDaniel
1492, 24447, 31317,31629, Morris et al.(1993),
Pullen et al. (1968b
Euphorbia nutata Engelm., PR, mesophytic
ane nardweod forests, O; Morris et al.
(1993)
oe a corollata L., BP, IF, xeric upland hard-
df sts, prairies, chalk out-
crops, roadsides, C; 281, 641, Brooks 144, 318,
Bryson 8296, Leidolf and uae (1998),
McDaniel 15622a, Morris et al. (19
Euphorbia dentata Michx., verse ae
ian—banks), O; 684, Pullen et al.(19
Euphorbia heterophylla L., BP, prairies, A; ae
15620
Euphorbia spathulata Lam., BP, IF, prairies, prairie
cedar woodlands, grass/forb meadows, O;
1346, 1511, Morris 3081, Morris et al. (1993)
a cra Michx., BP, PR, IF, xeric oo hard-
rests, R; 1528, McDaniel 2
ae ae Michx., BP, prairies, a 1519
cDaniel 31632, Morris et al. (1993), Pullen
et al. a 968b)
FABACEAE (LEGUMINOSAE)
*Albizia julibrissin Durazz., PR, roadsides, C; 462
uae: ace LS ee IF, pero lanig hard-
Ww ests, Xeric
es 643, McDaniel 31 026, Theriot
et al.
Amphicarpaea bracteata (L.) Fernald, IF, meso-
phytic upland hardwood forests, O; 904
cae americana Medik., BP, PR, prairie cedar
oodlands, rivers/creeks (riparian—banks),
canolsinage ditches (ipa banks),O/
A(l):677, 1606, McDanie
Apios priceana B.L. Rob.,BP., i oe hard-
wood forests, mesophytic upland hardwood
forests, C(I), FT, $1; Gordon 2050, 2051, Morris
3100, Morris et al. (1 993)
ts, pine fore
a
~—
Act / ot: Fe|
Gordon 2048
Baptisia alba (L.) R. Br. var. macrophylla (Larisey)
IF, eal eleee egal (ripar-
ian—banks), roadsides, O; ] 1568
Centrosema virginianum (L.) Nie BP PR, IF, bot-
tomland hardwood forests, mesophytic up-
land hardwood forests, rivers/creeks (ripar-
—banks), roadsides, 0; 533,674, Bruza 1007,
eecaal 28883
Cercis canadensis L., BP, PR, IF, bottomland hard-
wood forests, mesophytic upland hardwood
forests, xeric upland hardwood forests, prai-
ries, prairie cedar woodlands, rivers/creeks
(riparian—banks), C; 64, 74, 201, Leidolf and
McDaniel (1998), Morris et al. (1993), Theriot
et al. (1993)
Chamaecrista fasciculata (Michx.) Greene, BP. IF,
pine forests, prairies, urban areas, C; 365,
Brooks 315, Leidolf and McDaniel (1998)
Chamaecrista nictitans (L.) Moench, IF, pine for-
s4
var.canadensis, BP. U;, $2:
Clitoria mariana L., IF, pine forests, lakes/ponds/
impoundments ie banks), O; 637,
Brooks 317,Copeland s.r
*Coronilla varia L., BP. sone Brooks 210,Mac
Donald 10888
Crotalaria sagittalis L., BP. IF, prairies, seepage ar-
eas, O; 297, 701, Leidolf and McDaniel (1998)
Dalea candida Willd., BP, prairies, chalk outcrops,
C/A; 284, Bruza 938, Leidolf and McDaniel
(1998), McDaniel 13457, 31094, Morris et al.
(1993)
Dalea purpurea Vent., BP, prairies, chalk outcrops,
R/A; 283, Bruza 939, Leidolf and McDaniel
(1998), oi 13459, 31095, 31101, Morris
etal. (19
Prin ee illinoensis (Michx.) MacMill. ex B.L.
Rob. & Fernald, BP IF, xeric upland hardwood
forests, prairies, canals/drainage ditches (ri-
parian—banks), road O; 398, 642, Clonts
s.n., Leidolf and McDaniel (1998), Wigley
315064
sides,
Desmodium canescens (L.) DC., BR. IF, bottomland
hardwood forests, rivers/creeks (riparian—
me . he Theriot et al. (1993)
(Muhl. ex Willd.) DC., BP, xeric
upland hardwood forests, prairies, chalk out-
crops, O/C; 793, Leidolf and McDaniel (1998),
McDaniel 1502, 15504, Wigley 315067
BRIT.ORG/SIDA 20(2)
Desmodium cuspidatum (Muhl. ex Willd.) DC. ex
Loudon, PR, bottomland hardwood forests,
O; Morris et al. (1993)
Desmodium glabellum (Michx.) DC., IF, seepage
areas, roadsides, O; 986
Desmodium glutinosum (Muhl. ex Willd.) Alph.
Wood, PR, IF, mesophytic upland hardwood
forests, O/C; 560, McDaniel 28884
Desmodium laevigatum (Nutt.) DC., IF, pine for-
ests; Brooks 4/73
Desmodium nudiflorum (L.) DC., PR, IF, meso-
phytic upland hardwood forests, C; 559, Mor-
ris et al. (1993)
Desmodium paniculatum (L.) DC.,IF, pine forests,
roadsides, O; 858, Wigley 314113
Desmodium pauciflorum (Nutt.) DC., PR, meso-
phytic hardwood forests, O; Morris
et al. (1993)
Desmodi ium in Pel pies B.G. Schub., BP, PR, meso-
upland hardwood forests, rivers/
ae (riparian—banks), canals/drainage
ditches (riparian—banks), O; 670, 834
Desmodium rotundifolium DC, IF, mesophytic
upland hardwood forests, C(I); 907,
1581(R90/)
Dioclea multiflora (Torr. & A.Gray) C. Mohr, IF, bot-
tomland hardwood forests, mesophytic up-
land hardwood forests, O; 337, 557(R331),
Bryson 721,Clonts ileal et al. (1993)
Galactia ste (L.) Britton, BP IF, prairies, seep-
5,0; 518, 532, Bee 924
ne triacanthos L., BP, PR, IF, bottomland
hardwood forests, mesophytic upland hard-
roadsides, O;Leidolfs.n., Morris
et al. (1993), Theriot et al. (1993)
*Kummerowia striata (Thunb.) Schindl., IF, pine
ie ts, roadsides, A; 545, Brooks 409, Wigley
—
ae
wood forests
14023
ty hirsutus . BP. IF, prairies, lakes/ponds/
mpoundments (riparian—banks), canals/
Hee ies nae banks),roadsides,
O; 316, 1470, Brooks 152,Pullen et al.(1968b),
Sundell 1549
*Lespedeza bicolor Turcz., IF, mesophytic upland
aawcod jor O; 443
Michx., IF, pine — seep-
age areas, ode U; 867, 92
*lespedeza cuneata (Dum. Cours.) - Don, BP. IF,
pine forests, prairies, prairie cedar wood-
lands, rivers/creeks (riparian—banks
~~
cL a a
road-
ot
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI
sides, C; 569,676, Leidolf and McDaniel (1998),
MED ane ee?
l Michx.,BP,chalk outcrops
737
Strophostyles umbellata (Muhl. ex Willd.) Britton,
ne forests, O/C;698, Brooks 495, Copeland
5.N., McDaniel 2191
‘C(); Mebaneeiai6: 31648
Lespedeza repens (L.) Barton, IF, pine forests, road-
sides, O; 570, Brooks 496, Lowe (1921)
Lespedeza violacea e Pers., IF, pine forests; Co-
pe alan ds. se Low
/
gil n, BP IF. xeric upland
Tat dined forests ae i 702, Brooks
399, McDaniel 31624, Wigley 314
*Medicago arabica (L.) Huds., BP, in cultivated
fields, urban areas, U: 1206, 1288,Lowe (1921
*Medicago lupulina L., BP, PR, prairies ane.
o 75, MEP CET OZ Pullen et al. (1968b)
J t BP, prairies, chalk outcrops
roadsides, C/A; 287, Brooks 142, Leidolf and
McDaniel (1998), McDaniel 13451, 31096,
Morris et al. (1993)
*Melilotus officinalis (L.) Lam. [=
Medik.], B adsides, C; 2
Neptunia lutea (Leavenw.) Benth., BP, prairies,
prairie cedar woodlands, O/A(I), $3$4; 299,
399, 426, 466, Channel! 73 (MISSA), Davis s.n.
(MISSA), Leidolf and McDaniel (1998), McDaniel
460, Morris et al.(1993), Smith 1412
Psoralea psoralioides (Walter) Cory [= Orbexilum
pedunculatum (Mill.) Rydb. var. psoralioides
Vie lsely], BP, IF, pine forests, seepage ar-
eas, roadsides, U/C; 272, 1439, McDaniel 3147
ae tomentosa (L.) Hook. & Arn., BP IF, prai-
ries, grass/forb meadows, urban areas, O;
1589, 1600, McDaniel 31634
Robinia pseudoacacia L., PR, roadsides, urban ar-
as, O; 1321
Schrankia microphylla (Dryand.) J.F. Macbr. [=
Mimosa quadrivalvis L. var.angustata (Torr. &
A. Gray) Barneby], IF, pine forests, pine-mixed
hardwood forests, roadsides, O/C; 330, Bruza
928, McDaniel 13488
Senna Ail nee .) Link, IF seen hard-
woo ests; Theriot et al. (1993)
*Senna fe ifolia (L.) Irwin & vee IF, lakes/
ponds/impoundments (riparian—banks), C;
a
x A
Melilotus albus
1
29)
_
zo
=>
Sesbania exaltata (Raf.) Rydb. ex A.W. Hill, BP, IF,
rivers/creeks (riparian—banks), seepage areas
roadsides, C(I); 704, McDaniel 2528
Strophostyles helvula (L.) Elliott, IF, pine forests, O;
595
Stylosanthes biflora (L.) Britton, Sterns &
Poggenb., IF, mesophytic upland hardwood
forests, O; 339, Wigley 312
Tephrosia spicata (Walter) Torr. & A. Gray, IF, pine-
mixed hardwood forests, roadsides, C(I); 379,
1567, Clonts 820
sage irginiana (L.) Pers. IF, pine forests, pine-
xed hardwood forests, O; 346, 531, Brooks
ee
*Trifolium arvense L., BP, | meadows;
O'Brein s.n. (MISSA) A
*Trifolium campestre eae PR, : roadsides,
urban areas, C; 13717, 1315, 1323
Trifolium dubium cae : PR, prairie cedar
woodlands, O; 223, Pullen et al. (1968b)
*Trifolium incarnatum L., PR, roadsides, A(|); 700,
Pullen et al. (1968b)
*Trifolium lappaceum L.,BP,PR, prairies, roadsides
urban areas, C; 1514, 1533a, Brooks 145, Mac-
Donald 8721, Morris et al. (1993)
*Trifolium pratense L., BP, prairies, grass/fo 7
ws, roadsides, C(I); 164, Brooks 147,
Locke s.n. (
n
Trifolium reflexum L. IF, pine forests, C(I); McDaniel
21235
*Trifolium eae L., BP PR, IF, roadsides, urban ar-
405, Askew 37, Wigley 314080
area resupinat prairies, roadsides,
O; 145, Brooks 1] is ee et al. (1968b)
Vicia caroliniana Walter, PR, IF, bottomland hard-
forests, mesophytic upland hardwood
se ests, O; 248, 1296 (R248), Morris et al. (1993)
*Vicia grandiflora Scop., BP, IF, cultivated fields,
roadsides, O; 139, 1283, Pullen et al. (1968b)
Vicia minutiflora F. Dietr., PR, xeric upland hard-
wood forests, O; Ray 8006 (MISSA)
*Vicia sativa L.ssp.nigra (L.) Ehrh., IF, roadsides, O;
, Pullen et al. (1968b)
*Vicia tetrasperma (L.) Schreb., PR, canals/drain-
itches (riparian—banks); Fletcher s.n.
(MISSA
*Vicia villosa Roth ssp. varia (Host) Corb., BP, PR,
roadsides, O; 178, Brooks 209, Pullen et al.
(1968b)
Wisteria frutescens (L.) Poir., IF, oe all
poundments (riparian—banks), R
*Wisteria sinensis (Sims) DC, IF, roadsides, urban
areas, O/C; 91, Askew 38 (R Leidolf 91)
icaieke
ndifolia Ehrh., BP IF, bottomland hard-
arene forests acorn
forests, : 475,952, 1583 pee
et al. (19
Quercus i a PR, IF, bottomland hardwood
forests, mesophytic upland hardwood for-
ests, xeric upland hardwood forests, pine
forests, C; 306, 1554, Theriot et al. (1993)
asda ne Buckley [= Q. sinuata Walter var.
sinuata], BP bottomland hardwood forests;
ieoune 1067
Mx
land hardwood
aird 6, Theriot
ne xeric ee hardwood forests, pine
fore pine-mixed hardwood forests, C;
1551 ee et al. (1993)
Quercus laurifolia Michx., |F, bottomland hard-
wood forests, O; Leidolf pers. obs., Theriot et
al. (19
Quercus lyrata Walter, IF, bottomland hardwood
orests, swamp forests, rivers/creeks (ripar-
ian—banks), C; 355, Leidolf s.n., Theriot et al.
(1993)
Quercus macrocarpa Michx., BP, xeric upland
hardwood forests, R, $2; 1526a, Bryson 3123,
McDaniel 32330, Smith 364
Quercus marilandica Muenchh., BP, IF, xeric up-
land hardwood forests, prairie cedar wood-
lands, C; 387, 1553, pec and McDaniel
(1998), Morris et al. (1993)
Quercus michauxii Nutt, IF, bottomland hard-
wood forests, C; Leidolfs.n. Theriot et al.
(1993)
Quercus muehlenbergii Engelm., BP, PR, bottom-
land hardwood forests, mesophytic upland
hardwood forests, xeric upland hardwood
forests, prairie cedar woodlands, O; 435,
Bryson 3124, 7440, Leidolf and McDaniel
(1998), McDaniel 28151, 31646, Morris et al.
(1993)
Quercus nigra L., BP, IF, bottomland hardwood
Leidolf s.n.,
(1993)
Quercus pagoda Raf., BP, IF, bottomland hard-
wood forests, xeric upland hardwood forests,
McDaniel 31612a, Theriot et al.
BRIT.ORG/SIDA 20(2)
O;1 “e Leidolf s.n., McDaniel 3
et al. (1993)
Quercus ae L. BP IF bottomland hardwood
forests, pine forests, prairie cedar woodlands,
lakes/ponds/impoundments (riparian—
banks), C; 982, 1592, Leidolf s.n., Leidolf and
McDaniel (1998), Theriot et al. (1993)
Quercus rubra L., BP. IF, bottomland hardwood
forests, O; Laird 29, Leidolfs.n., Theriot et al.
(1993)
Quercus shumardii Buckley, BP, PR, IF, bottomland
hardwood forests, mesophytic upland hard-
wood forests, xeric upland hardwood forests,
pine forests, C; 354, 1555, Bryson 3126, Leidolf
s.n., Morris et al. (1993), Theriot
31613, Theriot
5.n., McDanie
et al. (1993)
Quercus stellata Wat gel 1h., BP. PR, IF, bottomland
| {\ 1£
1a s mesophytic upland hard-
ou forests, pine-mixed hardwood forests,
prairie cedar woodlands, C; 305, Leidolf and
McDaniel (1998), Morris et al. (1993), Theriot
93)
Quercus texana haan IF, bottomland hard
wood forests, O; Leido/fs.n., Theriot et al.
(1993)
Quercus velutina Lam.,BP PR, mesophytic upland
hardwood forests, xeric upland hardwood
ani R: 307, McDaniel 31614
GENTIANACEAE
Frasera caroliniensis Walter, PR, mesophytic u
7 nd hardwood forests, pains ip $3; ree
99, Morris et al. (1993), Ra
a angularis (L.) Pursh, BP, ae prairie
c woodlands, chalk outcrops, O; 372,
von oe
Sabatia brachiata Elliott, IF, pine-mixed hard-
ood cee roadsides, O/C; 347, 1539,
ean 25:
GERANIACEAE
Geranium carolinianum L., BP, lakes/ponds/im-
poundments (riparian—banks), roadsides, C;
89, McDaniel 31050
*Geranium dissectum L., BP, roadsides, C; 157
Geranium maculatum L., PR, Web upland
hardwood forests, O; Smith |
©
Ne)
QJ
HALORAGACEAE
*Myriophyllum aquaticum (Vell.) Verdc.; Jones
1975b
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI
HAMAMELIDACEAE
Hamamelis virginiana L., IF, bottomland hard-
d forests, O; 975, Theriot et al. (19
Liquidambar styraciflua L., BP. PR, IF, bottomland
hardwood forests, pine forests, see mixed
hardwood forests, xeric upland hardwood
forests, C; 333, Bryson (1980), McDani ne 1615,
Morris et al. (1993), Theriot et al. (1993)
HIPPOCASTANACEAE
Aesculus x bushiiSchneid. [= A. pavia L. x A.glabra
Willd.], BP. PR, poromlane naooe forests,
mesophytic
creeks (riparian—banks), O/C; 109, 172, ae
2 (UTC), 3 (UTC), aoe 5 (UTC), 6 (UTO), 7
(UTC), Morris et al. (1993)
Aesculus glabra Willd., BP. PR, bottomland hard-
wood forests, rivers/creeks (riparian—banks),
C(I), $2?; 72,108,111, 128,438, 1270, Askew 32,
Carpenter s.n. (MISSA), Channell s.n. (MISSA),
Funchess s.n. (MISSA), Graham s.n. (MISSA),
Lowe (1921), McDaniel 1066, Morris 3084,
Morris et al. (1993), Persons s.n. (MISSA), Read
$n. ea ae th 1895, Tracy s.n. (MISSA),
Webb 80 (MISSA)
Aesculus pavia a PR, IF, bottomland hardwood
orests, mesophytic upland hardwood for-
ests, rivers/creeks (riparian—banks), oe 10,
119, McDaniel 1069, 31027a, Morris et al.
(1993), Theriot et al. (1993)
HYDRANGEACEAE
Hydrangea quercifolia W. Bartram, IF, bottomland
dwood forests, pine-mixed hardwood
forests, R; 1498, Bryson 732, Laird 16
HYDROPHYLLACEAE
S93
lakes/ponds/impoundments (littoral-emer-
gent), A(); 576, 725, Jones (1975b), McDaniel
20935
HYPERICACEAE (GUTTIFERAE)
ericum crux-andreae (L.) Crantz, IF, pine-
mixed hardwood forests, seepage areas, O/
C; 647, Brooks 403, McDaniel 2659
Hypericum drummondii (Grev. & Hook.) Torr. & A.
aes IF, roadsides, O/C; 719, McDaniel 1568,
028
iypercum gentianoides (L.) Britton, Sterns &
Poggenb.,, IF, seepage areas, roadsides, A(|);
645, a 729
=
Hypericum gymnanthum Engelm. & A. Gray, IF,
canals/drainage ditches (riparian—banks), O;
03
Hypericum hypericoides (L.) Crantz,|F, bottomland
hardwood forests, pine forests, pine-mixed
hardwood forests, seepage oo 1481,
Brooks 404, Theriot et al. (19
Hypericum lobocarpum Gatt., c rivers/creeks
(riparian— panic) cata ersinaae ditches (ri-
parian—banks), ace eet
(riparian—banks), seepage areas, roadsides
O/C; 547, 588, 985, 1566, Clonts 832, McDa ne
1434
Hypericum mutilum L.,|F, bottomland hardwood
forests, pine-mixed hardwood forests, ca-
nals/drainage ditches (riparian—banks), C;
477, 562, Brooks 402, Theriot et al. (1993)
wood forests, canals/drainage ditches (ripar-
ian—banks), grass/forb meadows,O;715, 1588,
Theriot et al. (1993)
Hypericum sphaerocarpum Michx., BP, prairies,
halk outcrops, C(I); 374, Leidolf and
McDaniel (1998), MacDonald 8718, McDaniel
31049, Morris et al. (1993), Smith 1473
Hypericum stragulum W.P. Adams & E. Robson [=
H. hypericoides (L.) Crantz ssp. multicaule
Michx. ex Willd.) E. Robson], IF, grass/forb
—
meadows, R; 1
Triadenum tubulosum (Walter) Gleason, IF, bot-
tomland hardwood forests; Theriot et al.
(1993)
Triadenum walteri (J.G. Gmel.) Sue IF, bot-
| 1 | | 1 f 908, Theriot et
al. (1993)
JUGLANDACEAE
Carya carolinae-septentrionalis (Ashe) Engl. &
Graebn., PR, IF, bottomland hardwood forests,
mesophytic upland hardwood forests, O;
Morris et al. (1993), Pullen et al. (1968b),
Theriot et al.(1993)
Carya cordiformis (Wangenh.) K.Koch, IF, bottom
landh ee ood forests, O; Leido/fs.n., Theriot
et al. (19
Carya ae (Mill.) Sweet, IF, bottomland hard-
wood forests, xeric upland hardwood forests,
pine forests, O; 1552, Theriot et al. (1993)
Carya laciniosa (F.Michx.) G.Don, BP, bottomland
hardwood forests, RR, $283; 167, McDaniel
3490
740
Carya leiodermis Sarg. [= C. glabra (Mill.) Sweet
Lh; tafAach Ache] tad | {| |
wood forests, $283; Brooks 565, Theriot et al.
(1993)
Carya myristiciformis (F.Michx.) Nutt., BP, PR, bot-
tomland hardwood forests, mesophytic up-
land hardwood forests, xeric upland hard-
wood forests, O; McDaniel 4233, 4324, Morris
et al. (1993)
Carya ovalis (Wangenh.) Sarg., IF, bottomland
hardwood forests; Theriot et al. (1993)
Carya ovata (Mill. IF, bottomland
wood forests; Theriot et al. (1993)
aa (Ashe) Engl.& Graebn., BP roadsides
K. Koch Vard-
fs,
ae tomentosa (Lam. ex Poir.) Nutt. [= C. alba
(L.) Nutt. ex Elliott], BP, IF, bottomland hard-
wood forests, xeric upland hardwood forests,
pine forests, pine-mixed hardwood forests,
O; McDaniel 31612, Theriot et al. (1993)
Juglans nigra L., BP. PR, bottomland hardwood
forests, mesophytic upland hardwood for-
ests, U; 7573, Morris et al. (1993)
LAMIACEAE (LABIATAE)
Blephilia ciliata (L.) Benth., BP, PR, prairies, chalk
outcrops, roadsides, C; 269, 1444, Brooks 149
(MMNS), Carter 3013, Leidolf and McDaniel
(1998), ad la 31069, Morris et al.
(1993) sete
Colli Michx., IF I
wood forests, O; 906, ie DiaRial 26487
Hedeoma hispida Pursh, BP, PR, IF, prairies, road-
ee O/A(I); 1505, Leidolf and McD
98), McDaniel 1820, Pullen et al. ae
res eee aes BP PR , roadsides, ur-
ban areas, A; 9,80, Jones aa Stewart 247
*Lamium purpureum L., BP, PR, roadsides, urban
areas, C/A(l);59, 142, 1210, Askew 35,McDanie
mess
1588
Lycopus rubellus Moench, IF; McDaniel 1566
Lycopus virginicus L., IF, bottomland hardwood
forests, rivers/creeks (riparian-mudflats);
McDaniel 1437, Smith 943, Theriot et al.(1993)
*Mentha x rotundifolia (L.) Huds. (pro sp.), IF
grass/forb meadows, A(I); MacDonald 10902
M da citriodora Cerv. ex |
ag., BP, prairies, riv-
ers/creeks (riparian—banks), O; 369, Jones
a ee Bryson 3077, Leidolf nd McDaniel
(1998), Morris et al. (1993), Pullen et al.
ae
—
BRIT.ORG/SIDA 20(2)
Monarda fistulosa L., IF, roadsides, C; Bruza 931,
ones (1976a
Physostegi ia angustifolia Fern., BP IF, canals/drain-
a es (e Se ae seepage areas, road-
sides, A(I); 312, 154
Physostegi virginiana (L L.) Be nth., Jones (1976a)
Prunella vulgaris L., BR IF, pine forests, prairies, prai-
rie cedar woodlands, grass/forb meadows,
roadsides, C; 174, 184, 235, 1443, Brooks 133
Jones (1976a), Leidolf and McDaniel (1998),
Stewart 502
Pycnanthemum albescens Torr. & A. Gray, IF, pine
forests, pine-mixed hardwood forests, lakes/
ponds/impoundments (riparian—banks), O/
C; 810, Bruza 973, Jones (1976a), McDaniel
2197, 15729, 24534
Pycnanthemum tenuifolium Schrad., IF, meso-
phytic upland hardwood forests, xeric up
land hardwood forests, pine forests, canals/
drainage ditches (riparian—banks), roadsides,
O; 320,337, Brooks 278, Bruza 598, Bryson 298,
683, Clonts 825, Jones (1976a)
Salvia lyrata L., BP, PR, IF, bottomland hardwood
forests, prairies, prairie cedar woodlands,
roadsides, C; 136, 152, 187, Askew 70, Jones
(1976a), Leidolf and McDaniel (1998), Morris
eg
a)
+
Ge
Scutellaria elliptica Muhl. ex Spreng., IF, meso-
phytic upland hardwood forests, O; 457
Scutellaria incana Biehler, IF, pine forests, O; 594
Scutellaria integrifolia L., IF, bottomland hard-
wood forests, roadsides, O; 325, Bryson 718
Jones (1976a), McDaniel! 1092, Theriot et al.
(1993)
Scutellaria parvula Michx., BP, prairies, roadsides,
81, 212, Jones (1976a), Leidolf and
McDaniel (1998), Morris et al. (19
ate ium canadense L., BP, IF, canals/drainage
es (riparian—banks), O; 467, Bryson 724
Tr Pee brachiatum L., BP, IF, prairies, grass/
for ws, O/C; 791, Jones (1976a)
Leidolf a McDaniel (1998), MacDonald
7290, McDaniel 15506
Tr ichostena dichotomum L., BP. IF, xeric upland
tf ts,chalk outcrops, roadsides
R/O; 920, Brooks 486, McDaniel 31311, 31622
LAURACEAE
Lindera benzoin (L.) Blume, IF, bottomland hard-
wood forests, R/O; Leidolfs.n., Theriot et al.
(1993)
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI
Sassafras albidum (Nutt.) Nees, BP, PR, IF, bottom-
land hardwood forests, mesophytic upland
hardwood forests, xeric upland hardwood
forests, pine forests, prairie cedar woodlands,
roadsides, C; 140,376, Askew 50,63, Leidolf and
McDaniel (1998), Morris et al. (1993), Theriot
et al. (1993)
LENTIBULARIACEAE
Utricularia gibba L., IF, lakes/ponds/impound-
ments (littoral-submergent), A(l); 696a
LINACEAE
Linum medium eet Britton, BP, IF, prairies,
roadsides, C; 285,
Linum striatum es IF, mesophytic upland
hardwood forests, O; 556
Linum sulcatum Riddell, BP prairies, R, S3S4;Mac-
nalds.n
LOGANIACEAE
Gelsemium sempervirens (L.) W.T. or IF, pine-
mixed hardwood forests, C(|); /
Spigelia marilandica L., PR, IF, ne hard-
wood forests, mesophytic upland hardwood
forests, pine forests, O; 249, 1499, Morris et al.
(1993)
LYTHRACEAE
mmannia coccinea Rottb., BP, IF, rivers/creeks
(emergent), cultivated fields, A(I);697, Bryson
2607, Jones (1975b) 88
*Cuphea carthagenensis (Jacq.) J.F. Macbr., IF,
mesophytic upland hard I f 660
McDaniel 252
lythrum alatum Pursh var. lanceolatum (Elliott)
orr. & A. Gray ex Rothr, BP, prairies, rivers/
creeks (riparian—mudflats), A(I); 373, Jones
(1975b), Leidolf ee McDaniel (1998),
McDaniel 25290, 3162
Rotala ramosior (L.) ae IF, canals/drainage
ditches (riparian-mudflats), C; 537, Jones
(1975b)
MAGNOLIACEAE
Lirl pane tulipifera L.,PR,IF, bottomland hard-
mesophytic upland hardwood
ne pine forests, O/C; 974, Morris 3101,
Theriot et al. (1993)
MALVACEAE
—
fae oem pe
Bryson 3419
Anoda cristata (L.) Schitdl., BP roadsides, R; 798
Hibiscus laevis All. IF, rivers/creeks (riparian—
banks), grass/forb meadows, O; 712, 716
Hibiscus moscheutos L., BP roadsides, U; 938
Modiola caroliniana (L.) G. Don, PR, urban areas,
U; 1204
Sida rhombifolia L., BP, cultivated fields, A;
McDaniel 28085
Sida spinosa L., BP,« roadsides,O
A; 409, 800, Bryson 3422, Lowe (1921),
McDaniel 28084
MELASTOMATACEAE
Rhexia mariana L. var. mariana, IF, xeric upland
hardwood forests, pine forests, canals/drain-
age ditches (riparian—banks), roadsides, O/
C; 440, Bryson 714, Clonts 817
MELIACEAE
*Melia azedarach L., PR, roadsides, U; 224
I+; fy eee
MENISPERMACEAE
Calycocarpum lyonii (Pursh) A. Gray, BP, PR, IF, bot-
tomland hardwood forests, mesophytic up-
land hardwood forests, O; Lowe (1921),
McDaniel 24964, Morris et al. (1993), Theriot
et al. (1
occulus ee nus (L.) DC., BP, PR, IF, bottomland
hardwood forests, mesophytic upland hard-
wood peara pine forests, prairie cedar
woodlands, urban areas, U; 883, Brooks 582,
Leidolf and McDaniel (1998), Morris et al.
(1 993), Theriot el al. ey
. PR, mesophytic up-
land hardwood ier O/R, $384; Morris
3088, Morris et al. (19
MOLLUGINACEAE
Mollugo verticillata L., IF, canals/drainage ditches
(emergent), O; 507, Jones (1976b)
Cy
—
=
—
MORACEAE
*Broussonetia papyrifera (L.) L'Hér.ex Vent.; Pullen
et al. (1968b)
Maclura pomifera (Raf.) C.K. te, BP PR, bot-
tomland hardwood forests, mesophytic up-
land hardwood forests, xeric upland hard-
wood forests, prairie cedar woodlands, C; 202,
377, Brooks 467, Bryson 7439, Leidolf and
McDaniel (1998), McDaniel 11622, Morris et
al. (1993)
Morus rubra L., BP PR, IF, bottomland hardwood
forests, mesophytic upland hardwood for-
ests, prairie cedar woodlands, C; 156, Askew
58, Bryson 7438, Leidolf and McDaniel (1998),
Morris et al. (1993), Theriot et al. (1993)
MYRICACEAE
Myrica cerifera L.,\F, pine forests, O; 121
NELUMBONACEAE
Nelumbo lutea Willd,, IF, lakes/ponds/impound-
ments (littoral-rooted-floating), O; 855
NYCTAGINACEAE
Boerhavia erecta L.; Pullen et al. (1968b)
Mirabilis albida (Walter) Heimerl, BP. chalk out-
crops, R; McDaniel 13454
NYMPHAEACEAE
Nymphaea odorata Aiton, lakes/ponds/im
poundments (littoral-rooted- ocr):
Jones (1975b)
Fraxinus americana L., BP, PR, IF, PonOm lett nals
WOO od fo orests mesophytic
forests, prairie cedar woodlands O; Leidolf
and McDaniel (1998), Morris et al. (1993),
Theriot et al. (1993)
Fraxinus pennsylvanica Marshall, BP, PR, IF, bot
tomland hardwood forests, mesophytic up-
land hardwood forests, pine forests, lakes/
mpoundments (riparian—banks),
mess O/C; 612, 1490, McDaniel 31639,
Morris et al. (1993), T Gave et al. (1993)
Fraxinus gcane ia ta me - PR, meso
phytic uy 2: Bryson
10123,Gordon 2031,Lowe (1 eae 3083,
Morris et al. (1993), Smith 141
*Ligustrum sinense Lour.,, BP, PR, IF, bottomland
hardwood forests, mesophytic upland hard-
wood forests, xeric upland hardwood forests,
prairies, prairie cedar woodlands, C; 205,854
Leidolf and McDaniel (1998), Morris et al.
(1993), Theriot et al. (1993)
—
ONAGRACEAE
Gaura shale L., BP, IF, Suara ere ditches
(riparian—banks), O;679, McDaniel 1565
Gaura aon se Jones (1 ae
Gaura parviflor glas ex Lehm.; Jones
(1 ae le et a (1968b
Ludwigia alternifolia L, IF, pine forests, grass/forb
MESON: O; 502, 587, Brooks 328, Clonts 843
Ludwigia decurrens Walter, IF, bottomland har
wee forests, canals/drainage ditches
(emergent), O; 693, Theriot et al. (1993)
BRIT.ORG/SIDA 20(2)
pl Perea | Pare | \Afa] Ich | {| !
ee forests, pine-mixed hardwood forests,
canals/drainage ditches (emergent), O; 553,
McDaniel 22135, heriot et al. (1993)
Ludwigia hirtella Raf., IF, pine-mixed hardwood
sts, canals/drainage ditches (emergent),
grass/forb meadows, 479, Brooks 32/,
Clonts 842, McDaniel 2
Ludwigia palustris (L : sae - canals/drainage
ditches (emergent), lakes/ponds/impound-
ments (littoral-emergent), O; 452, 610
Ludwigia peploides (Kunth) Raven, BP, rivers/
are (emergent), canals/drainage ditches
(emergent), O/A; 407, McDaniel 25287
Oen ae biennis L., BP.IF, pine-mixed hardwood
roadsides, C; 850, Brooks 484, Jones
cay
Oenot! thera fruticosa L.ssp.glauca (Michx.) Straley,
, pine-mixed hardwood forests, seepage
5,0;495, McDaniel 2532
Oenothera laciniata Hill, BP, cultivated fields, A(|);
a) BLBOD he Jones (1974b)
t, IF roadsides, U; Copeland
5.n.
Oenothera pilosella Raf. IF, roadsides, U; Copeland
., Jones (1974b)
Oenothera speciosa Nutt., BP, prairies, roadsides
_ Leidolf and McDaniel (1998)
BP, SU: Donald s.n.
Ww
Oenothera triloba Nutt.,
MISSA)
OXALIDACEAE
Oxalis articulata Savigny ssp. rubra (St. -Hil.)
Lourteig; Pullen et al.(1 a )
Oxalis dillenti Jacq.,|F,t {hard 1 fi
ests, O; 263,635
Oxalis stricta L., BP, PR, IF, bottomland hardwood
forests, Sean (riparian—banks), urban
re 626, McDaniel 13440, Theriot et al.
(] a
Oxalis violacea L.,PR, IF, mesophytic upland hard-
wood forests, xeric upland hardwood forests,
O; 1314, Morris et al. (19
PAPAVERACEAE
Sanguinaria canadensis L., PR, mesophytic up-
land hardwood forests, A(l); Morris 3089, Mor-
ris et al. (1993)
PASSIFLORACEAE
Passiflora incarnata L., BP. IF, bottomland hard
wood forests, lakes/ponds/impoundments
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI
(riparian—banks), grass/forb meadows, road-
sides, O/C; 329, 1587, Bryson 709, Clonts 833,
Jones (1976b), McDaniel 31323
Passiflora lutea L., PR, IF, bottomland hardwood
forests, xeric upland hardwood forests, O;
Clonts 822, Jones (1976b), Ray 4734 (MISSA),
Theriot et al. (1993)
PENTHORACEAE
ie)
| ee | exe! | 1 | fl |
forests, mesophytic upland hardwood for-
ests, rivers/creeks (emergent), canals/drain-
age ditches (emergent), A(l); 478, 692, 1584,
Theriot et al. (1993)
PHRYMACEAE
Phryma leptostachya L., PR, mesophytic upland
hardwood forests, O;467, Morris et al.(1993),
Smith 1416
PHY TOLACCACEAE
Phytolacca americana L., IF, roadsides, C; 522
PLANTAGINACEAE
Plantago aristata Michx., IF roadsides, C; 327, 348
*Plantago lanceolata L., BP roadsides, C; 273, 400
Plantago rugelii Decne., PR, IF, roadsides, urban
areas, O; 344, Pullen et al.(1968b), Stewart 106
Plantago virginica L., BP, prairies, urban areas, C/
A\(l); 183, McDaniel 31037
PLATANACEAE
Platanus occidentalis L., PR, mesophytic upland
hardwood forests, O; Morris et al. (1993)
POLEMONIACEAE
Phlox carolina L. ssp. angusta Wherry, IF, pine-
mixed hardwood forests, seepage areas, O/
C; 497, Brooks 119, McDaniel 15073, Pullen et
al. (1968b)
Phlox divaricata L., PR, IF, bottomland hardwood
forests, Mevephytle sake hardwood for-
ests, O; 127, Morris et al. (1993)
Phlox punieilate L., IF, C(I); Lowe (1921), Mac-
Phlox pilosa LIF, pine forests, roadsides, O/A; 238,
Brooks 118, McDaniel! 1093, 314
POLYGALACEAE
ue incarnata L., IF, pine forests, R; 1565,
Bryson 713
Polygala mariana Mill, IF, mesophytic upland
hardwood forests, pine forests, pine-mixed
hardwood forests, canals/drainage ditches
743
(riparian—banks), lakes/ponds/impound-
ments (riparian—mudflats), O/C 447, Bennett
5.n., Brooks 325, Bryson 731, Daugherty 2,
McDaniel 2529, 2641, 13487
Polygala nana (Michx.) DC., IF, mesophytic up-
land hardwood forests, pine forests, pine-
mixed hardwood forests, seepage areas, O/
C; 243, 1483, 1495, 1564, Bryson 702, 708,
McDaniel 2535
Polygala sanguinea L.,|F, bottomland hardwood
forests, pine forests, grass/forb meadows,
roadsides, O; 323, 582, Bryson 680, Clonts 829,
POLYGONACEAE
Brunnichia ovata (Walter) HNIC BPIFE, bottom-
land hardwood forests
poundments (riparian— Ban) O/A(I); 599,
Jones (1975b), McDaniel 31616, Theriot et al.
(1993)
a aviculare L., BP, IF, lakes/ponds/im-
poundments (riparian— ee cultivated
a A(l); 404, 608, Jones )
acces cespitosum ie : lakes/ponds/
Quadmens Wipatlal melon os 979
©
Da |
A;McDan jel 24537, Stewart 124
Polygonum hirsutum ye PR, ae
itch Leidolf per
Polygonum hydropiperoides Michx., : bottom-
land hardwood forests, canals/drainage
ditches (emergent), lakes/ponds/impound-
ments (littoral-emergent), C/A; 450, 856,
Bruza 962, 987, Jones (1975b), Theriot et al.
es (emergent), R
Polygonum lapathifolium L., BP, canals/drainage
ditches (emergent), lakes/ponds/impound-
ments (littoral-emergent), . 893, Jones
(1975b), McDaniel 31330, 3164
Polygonum pensylvanicum L., BP.IF, aie
age ditches (emergent), lakes/ponds/im
Seurdene (ee ene ea aes
U/C; 410, 597, 882, Jones (1975b)
Polygonum punctatum Elliott; Jones (1975b)
sein oS ianum L., IF, bottomland hard-
od forests, pine-mixed hardwood forests,
ei (1975b), Leidolfs.n., Theriot et
o 721
*Rumex acetosella L.; Jones (1975b)
Rumex altissimum Alph.Wood; Jones (1975b
a
*Rumex crispus L., BP, IF, cultivated fields, road-
anes CG ee en aie ras (1975b)
adsides urban
p
BRIT.ORG/SIDA 20(2)
Myosurus minimus L., BP, cultivated fields, C(I
71,1284, Bryson 3500
aS
areas, C())/A(I); 468, 137 12 pa
(1975b), Pullen et al. (1968b)
*Rumex pulcher L.; Jones (1975b), Pullen et al.
(1968b)
PORTULACACEAE
Claytonia virginica L., PR, IF, bottomland hard-
wood Geer eee meadows, road-
sides, O/C; 34, e (1921), McDaniel 1680,
Morris ee
PRIMULACEAE
Anagallis minima (L.) Krause, BP, xeric upland
hardwood forests; McDaniel 29336a
Dodecatheon meadia L., BP, PR, prairies, prairie
woodlands, R, $2; Bryson 5400, Lowe
1921), Morris 3095, Morris et al. (199
Lysimachia ciliata L., IF, seepage areas, U; Bryson
712
Lysimachia lanceolata Walter; Jones (1976b)
*Lysimachia nummularia L., BP, canals/drainage
ditches a -bar 08
Samolus Sp.pan iflorus (Raf.) Hultén,
BP. IF, vee (riparian— mga) ca-
nals/drainage ditches (emergent),
1487 (R448), Bryson 3076, Jones (1976b)
RANUNCULACEA
Actaea sar Elliott, BP PR; Gordon 2062
\ks, emergent), R
Anemone caroliniana Walter, BP, prairies, chalk
crops, roadsides, O/C; 115, 143, 126
. . ico and Mebane (1998), Morris
3077, Morris et al. (19
Anemone virginiana L., 7 bottomland hard-
wood forests, mesophytic upland hardwood
forests, O; Lowe (1921), Morris et al. (19
Clematis crispa L., PR, IF, bottomland hardwood
forests, canals/drainage ditches (riparian—
banks), roadsides, O; 534, Morris et al. (1993)
*Clematis terniflora DC., BP, PR, prairie cedar
O nds, roadsides, O; 707,McDaniel 31617
Delphinium carolinianum Walter — ssp.
carolinianum, BP. prairies, O/A(I); 273, 1509
Maes Brooks 154,MacDonald 8716,McDaniel
—
rene tricorne Michx., BP, prairies, C(
McDaniel 31027
), $2;
Ranunculus abortivus L., BP PR, mesophytic up-
land hardwood forests, xeric upland hard-
wood forests, lakes/ponds/impoundments
(riparian-mudflats), urban areas, C; 1278,
1292, 1335, aes jel 1589
nunculus a L., BP, cultivated fields, A(I);
1273, 13430
Ranunculus fascicularis Muhl.ex Bigelow, BP, PR,
mesophytic upland hardwood forests, prai-
ries, O; 5, 67, 117(R5), Leidolf and McDaniel
(1998), McDaniel 2326
Ranu qunculus hispidus Michx. var. hispidus, BP, cul-
ted fields, A; McDaniel 1826
tee le Michx. var. nitidus (Chapm.)
P urban areas, A; McDaniel 24967
es micranthus Nutt., BP, xeric upland
hardwood forests, O; 13
*Ranunculus muricatus L., BP, PR, IF, bottomland
hardwood epee cultivated fields, urban
areas, C/A(l); 1275, 1282, Bryson 3497, Clonts
744,McDani oe
*Ranunculus parviflorus L., BP, cultivated fields;
Bryson 34
—
Ranunculus pusillus Poir., BP. IF, canals/drainage
itches (riparian-banks), grass/forb
ws, O; 1345, 1363
Ranunculus recurvatus Poir., BP, PR, mesophytic
nd hardwood forests, R/U; 1349,
pe ei 1023
eile ele sardous nie BP.PR, roadsides, C/
7,McDaniel 3105
hain trum Gin a Enea upland
hardwood forests, A, $182; McDaniel 31385
Thalictrum thalictroides (L.) Eames & B. Boivin, PR,
IF, mesophytic upland hardwood forests, O
89 Askew 47, Morris et al. (1993)
RHAMNACEAE
Berchemia scandens (Hill) K. Koch, BP, PR, IF, bot-
tomland hardwood forests,
land hardwood forests, prairies, prairie cedar
sone chalk outcrops, O/C; 227, Bryson
7441, Carter 3002, Leidolf and McDaniel
(1998), ee el 10581, Morris et al. (1993),
Theriot et al. (1993)
eras ied BP prairie cedar wood-
lands, chalk outcrops, O/A(I); 286, 432, Leidolf
and McDaniel (1998), McDaniel 13452,31065,
Morris et al. (1993)
mesoph ytic up-
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI
Rhamnus caroliniana Walter [= Frangula
caroliniana (Walter) A. Gray], BP, PR, bottom-
land hardwood forests, mesophytic upland
hardwood forests, prairie cedar woodlands,
chalk outcrops, O/C; 388, . . Leidolf and
hel iel (1998), MacDon 717, McDaniel
316 eee et ie
mnus lanceolata Pursh, BP. PR, prairie cedar
woodlands, chalk outcrops, C(I), $2; 837, 1326
(R831), 1340, - and McDaniel (1998),
MacDonald 8150, Pullen et al. (1968b), Ray
8008 (MISSA), Smith 1447, 1937
ROSACEAE
Agrimonia rostellata Wallr., BP, PR, mesophytic
upland hardwood fe ates upland hard-
wood forests, O/A(l);671, McDaniel 31611
Amelanchier arborea (F. Michx.) Fernald, IF, bot-
tomland hardwood forests, O; Leido/f s.n.,
Theriot et al. (1993)
*Aphanes microcarpa (Boiss.& ieee Rothm., PR,
roadsides, urban areas, O/C; 1213, 1506
Crataegus calpodendron (Ehrh.) nae PR, me-
sophytic upland hard | f O,$?;21
a»
=
S
orris 3087, Morris et al. (1993)
Crataegus crus-galli L., BP, IF, pine-mixed hard-
wood forests prairie cedar woodlands, road-
sides, O; 279,697, cairo McDaniel (1998)
1 engelmannii Sarg., BP, xeric upland
hardwood forests, prairie cedar woodlands,
O; 706, Leidolf and McDaniel (1998), McDaniel
11619
Crataegus marshallii Eggl., PR, IF, bottomland
hardwood forests, mesophytic upland hard-
wood forests, pine-mixed hardwood forests,
O; 135, 1370, Leidolfs.n seen 1687, Mor-
ris et al. (1993), ae et al. (19
Wendl.) K. a IF, bot-
ioniione mre forests, O; Leidolf s.n.,
Theriot et al. (19
Crataegus sata Michx., BP, IF, bottomland
hardwood forests, xeric upland hardwood
ee 0 Leidolfs.n., McDaniel 3493, Mor
ris 3102, Morris et al. (1993), Theriot et al.
(1993)
Crataegus viridis L., IF, bottomland hardwood for-
ests, R/O; Leidolfs.n., Theriot et al. (1993)
*Duchesnea indica (Andrz.) Focke, PR, meso-
hytic upland hardwood forests, O; 1306
Fragaria virginiana Duchesne, BP, xeric upland
hardwood forests, prairie cedar woodlands,
Yrijin a (
O/C; 168, 214, a Askew 27, Morris 3078,
Morris et al.19 -
Geum d Jacq., BP, PR, bottomland hard-
wood forests SN upland hardwood
forests, rivers/creeks ae banks), C; 304,
Bryson 3079, Morris et al. (1993)
Potentilla simplex Michx., BP. prairies, O; 169, 1365
Prunus americana Marshall, BP, IF, rivers/creeks
(riparian—banks), roadsides, U; 103, 141, 43
Prunus angustifolia Marshall, BP, PR, IF, ls
chalk outcrops, roadsides, C; 30, 37, 7
Prunus mexicana S. Watson, BP, IF, Sanaa
hardwood forests, pine-mixed hardwood
forests, O; 43, Morris et al. (1993), Pullen et al.
68b), Theriot et al. (1993)
Prunus serotina Ehrh., BP. PR, IF, bottomland hard-
wood forests, mesophytic upland hardwood
forests, pine forests, pine-mixed hardwood
forests, rivers/creeks (riparian—banks), . 73,
49 Morris et al. (1993), Theriot et al.
!
~—
nee
*p f, j M.Roem.,BP prairies, chalk
outcrops, ee 207, 621, Leidolf and
McDaniel (1998), MacDonald 7531, 81506,
McDaniel 15513, 29756
a ae Decne., BP IF, pine forests, prai-
5,828 Leidolf and
woodlands, U: 80
a eesipe 998)
Rosa carolina L., BP, prairies, prairie cedar wood-
lands, O; 842, 1453, Brooks 146, Leidolf and
McDaniel (19
*Rosa chinensis Jacq., BP. IF, mesophytic upland
ardwo 208, 1488
*Rosa multiflora Thunb.ex Murray, BP PR, bottom-
land hardwood forests, prairies, prairie cedar
oodlands, roadsides, C; 8, 177, Leid
od forests, roadsides, R/U;
idolf and
io niel (1998), Morris et al. (1993)
Rubus argutus Link, BP, PR, IF, bottomland hard-
forests, prairies, O; Leidolf and
McDaniel (1998), Morris et al.(1993), Theriot
tal. (1993
Rubus betulifolius Small [=
bottomland hardwo
upland hardwood forests, roadsides, C; 154
R. argutus Link], PR, IF,
od forests, mesophytic
[373
Rubus cuneifolius Pursh, BP, heer cedar
ands, O; McDaniel 311
Rubus eee Willd. IF, pine- ie hardwood
forests, O; 122
Rubus trivialis Michx., BP, IF, bottomland hard-
746
wood forests, C; McDaniel 2324, Theriot et al.
(1993)
RUBIACEAE
G 4 / L
wood forests
ian—banks), akes/ponds/impouncments (ri-
parian—banks), C; 584, 607, Jones (1976b),
Theriot et al. (19
*Cruciata pedemontana (Bellardi) Ehrend., BP,
roadsides, C(l); MacDonald 9305
Diodia teres Walter, IF, roadsides, O; 439, Jones
(1976b)
Diodia virginiana L., BP IF, bottomland hardwood
forests, la ae saree ae (ripar-
| Peeat el
ian—mudflats) roadsides, C;
416,611 heal al. (1993)
Galium aparine L., BP, PR, lakes/ponds/impound-
ments (riparian—banks), roadsides, C; 155
Jones (1976b), McDaniel 31049
Galium circaezans Michx., BP. PR, bottomland
hardwood forests, nesophytic upland hard-
wood forests, prairie cedar woodlands, C; 303,
Jones (1976b), Leidolf and McDaniel (1998),
Morris et al. (1993)
Galium obtusum Bigelow, IF, bottomland hard-
wood forests, A(l); 245, Jones (1976b)
*Galium parisiense L., BP roadsides, A(I); 374
Galium pilosum Aiton, BP, IF, xeric upland hard-
wood forests, grass/forb meadows, U; 1590,
Clonts 826, Jones (1976b)
1(L.) Scop.; Jones (1976b)
Gali ium uniflorum Michx., IF, mesophytic upland
naleniees er O; 446
Galium vil 1 Nutt., BP, prairies, O; 746, Leidolf
and McDaniel (1998)
= australis W.H. Lewis & D.M, Moo iE
tonia micrantha Canes ena
een areas, U/C(|); 1316, MacDonald 8 eo
Hedyotis crassifolia Raf. [= Houstonia pusilla
Schoepf], BP, PR, IF, prairies, grass/forb
meadows, urban areas, C; 42, 62, 132, Jones
1976b), Leidolf and McDaniel (1998)
Hedyotis nigricans (Lam.) Fosberg, BP, prairies,
chalk outcrops, O; 794, Jones (1976b), Leidolf
and McDaniel (1998), McDaniel 155
Hedyotis purpurea (L.) Torr.& A. Gray var.calycosa
(A osberg, BP, IF, bottomland hard
wood ae pranes, aie cedar wood
lands, grass/f oadsides, C; 165,
alium CHIClCOTIUET
—
NR
BRIT.ORG/SIDA 20(2)
185,257, 1341 (R185), 1371, Brooks 125,Bryson
57,715, Jones (1976b), Leidolf and McDaniel
(1998), McDaniel 10523, Morris et al. (1993),
Theriot et al. (1993)
Mitchella repens L.,|F, bottomland hardwood for-
ae cela upland hardwood forests,
C251, 1580, Brooks 474, Jones (1976b), Theriot
a (1993)
Oldenlandia boscii (DC.) Chapm. IF, pine forests,
pine-mixed hardwood forests, canals/drain-
age ditches (riparian—banks), C/A; 505, Brooks
322, Copeland s.n., McDaniel 2661
Oldenlandia uniflora L., IF, pine-mixed hardwood
forests; Brooks 324
Richardia scabra L.; Jones (1976b)
*Sherardia arvensis L., PR, roadsides, urban areas,
C/A; 102, Jones (1976b), McDaniel 1610
Pullen et al. (1968b)
Spermacoce glabra Michx., BP, canals/drainage
ditches (riparian—banks), roadsides, O; 631
RUTACEAE
*Poncirus trifoliata (L.) Raf., BP, PR, IF, bottomland
hardwood forests, mesophytic upland hard-
wood forests, roadsides, U:69,870, Theriot et
al. (1993)
Ptelea trifoliata L., IF, bottomland hardwood for-
ests, rivers/creeks (riparian—banks), U, $3$4;
717
—
Zanthoxylum clava-herculis L., BP PR, mesophytic
upland hardwood forests,xeric upland hard-
wood forests, U; 935, McDaniel 1
31042, Morris et al. (19
SALICACEAE
*Populus alba L.,IF, roadsides, urban areas, U; 1542,
1545
1988, 24446,
— deltoides W. Bartram ex Marshall, PR, bot-
mland hardwood forests, mesophytic up-
a hardwood forests, C; 1322, Morris et al.
(1993)
Salix exigua Nutt., BP, canals/drainage ditches (ri-
arian—banks), R; 673, Lowe (1921)
Salix humilis Marshall, |F, pine forests, pine-mixed
hardwood forests, lakes/ponds/impound-
ments (riparian—banks), seepage areas, road-
sides, U; 508, 1501 (R508), Edwards s.n., Mac-
Donald 6132, McDaniel 2840
J | | |
forests, rivers/creeks (riparian—banks/
mudflats), canals/drainage ditches (riparian—
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI
banks), C; 13
(1993)
SAPINDACEAE
Cardiospermum halicacabum L., IF, rivers/creeks
(riparian—banks), canals/drainage ditches (ri-
parian—banks), cultivated fields, O; 694, 710,
Pullen et al. (1968b), Smith 942
SAPOTACEAE
Bumelia lycioides (L.) Pers. [= Sideroxylon lycioides
LJ, BP PR, IF, bottomland hardwood forests,
mesophytic upland hardwood forests, prai-
rie cedar woodlands, rivers/creeks (riparian—
banks), C; 434, 688, 827,Leidolf and McDaniel
(1998), Morris et al. (1993)
SAURURACEAE
Saururus cernuus L., PR, IF, bottomland hardwood
forests, swamp forests, O; 341, Jones (1975b),
McDaniel 28882, Theriot et al. (1 he
CROPHULARIACEAE
Agalinis fasciculata (Elliott) Raf, IF, pine forests, O;
1, 1362, Askew 129, Theriot et al.
~~
Agalinis gattingeri (Small) Small ex Britton, BP,
pr anes O; 929, Leidalf and McDaniel (1998)
lla (Nutt.) Small ex Britton, BP
R; MeBoiniehs) 1653
Agalini Bae (Pennell) Shinners, BP, prai-
ries, R, 829, Leidolf and McDaniel (1998),
a 15617
Agalinis purpurea (L.) Pennell, BP, IF, pine forests,
rairies, roadsides, C(1); 877, 9317, 960, Leidolf
and McDaniel (1998), McDaniel 24535, 31308,
Stewart 117
AI uae (Vahl) Raf., BP. IF, pine-mixed
chalk outcrops, roadside
989, McDaniel! 14664, 31312
Aureolaria flava (L.) Farw., BP. IF, xeric upland hard-
wood forests, pine-mixed hardwood forests,
roadsides, U/A(I); 718, Clonts 1400, McDanie
Aureolaria pectinata (Nutt.) Pennell, IF, pine-
mixed hardwood forests, U/A(I); 1538,
McDaniel 2724
Buchnera floridana Gat dog. [= 8. americana La
BP, IF, prairies, ae areas, O/R; 296, 371
(R296), 539, Leido io McDaniel (1998),
McDaniel 13450, 3
Dasistoma ie on Raf., BP. PR, bot-
tomland hardwood forests, mesophytic up-
land hardwood forests, roadsides, O, $3S4;
1601, Morris et al. (1993)
Epifagus virginiana (L.) Barton, IF, bottomland
hardwood forests, A(|); 943
Gratiola neglecta Torr, IF, bottomland hardwood
forests, canals/drainage ditches (emergent),
uae meadows, C; 264, 1347, McDaniel
Grati re pilosa Michx., IF, seepage areas, O; 481
Gratiola virginiana L., 7 botoniand hardwood
forests, swamp forests, canals/drainage
ditches (emergent), O/A(I); 1294, 1440,
McDaniel 2451
Leucospora multifida (Michx.) Nutt., BP cultivated
ields, O;
Linaria canadensis (L.) Chaz. [=
canadensis (L.) D.A. Sutton],
fields, A(l); 12817
Lindernia dubia (L.) Pennell, BP, IF, bottomland
hardwood forests, rivers/creeks (riparian—
mudflats), O; McDaniel 25289, Theriot et al.
(1993)
Mecardonia acuminata (Walter) Small, IF, pine
forests, seepage areas, lakes Is/i
pela (riparian—banks),
Copelan
cn ee umabronin (J.F.Gmel.) S.F. Blake,
IF, lakes/ponds/impoundments (riparian—
mudflats), O/C; 1595, McDaniel 1570, 1570a
Mimulus alatus Aiton, BP. IF, bottomland hard-
wood forests, ver creeks (riparian—banks/
mudflats), canals/drainage ditches (riparian—
banks), O; 554,672, Theriot et al. (1993)
Caruel, PR, urban areas,
Nuttalanthus
BP, cultivated
—
ponds/im
O; 646, 652,
xD + be : /|
FUPCHlaceliGd VisCO5a (L.)
C()); MacDonald 9377
Pedicularis canadensis L.,PR, mesophytic upland
rdwoad forests, O; Smith 193
Penstemon digitalis Nutt. ex Sims, BP, PR, meso-
phytic upland hardwood forests, prairies,
prairie cedar woodlands, rivers/creeks (ripar-
ian—banks), O; 217, 1452, Morris 3098
Penstemon laevigatus Aiton, BP, prairies, road-
sides, O; 2717, 1
Penstemon tenuiflorus Pennell, BP, prairies, prai-
rie cedar woodlands, chalk outcrops, road-
sides, W/C(O, $2S3; 270, Bryson 129, 7442,
McDaniel 10584, 31020, Morris 3079, Morris et
al. (1993), Stewart 357
Penstemon tubiflorus Nutt., BP. PR, mesophytic
upland hardwood forests, lakes/ponds/im-
poundments alopecia, R/U; McDaniel
31051,Morris et al. (19
Scrophularia marilandica PR, mesophytic da
land hardwood forests, U/O; McDaniel 2888
Morris et al. (1993)
Seymeria cassioides (J.F.Gmel.) S.F. Blake, IF, pine
forests, pine-mixed hardwood forests, C/A(\);
959, McDaniel 22134, 24536
Tomanthera auriculata (Michx.) Raf. [= Agalinis
auriculata (Michx.) S.F. Blake], BP, prairies,
roadsides, RR, $1; 928, 930, Leidolf and
NeDanil 998), McDaniel 15621
blattaria L., BP, prairies, roadsides, O;
* /orha
318, Brooks 156
*Verbascum thapsus L., IF, roadsides, C(|); 521
*Veronica arvensis L., BP roadsides, C; 70, 159
Veronica peregrina L., BP PR, cultivated fields, ur-
ban areas, C; 1208, 1274, 1285
*Veronica persica Poir.,, PR, urban areas, O; 1207
SIMAROUBACEAE
*Ajlanthus altissima (Mill.
ea ith 363
—_
Swingle, PR, urban ar-
SOLANACEAE
BP urban areas: Carraway
490
Physalis angulata L., BP. IF, chalk outcrops, cullti-
vated
713,Clonts 1408, McDaniel 22195
Physalis de Nees, ag A ead Ne
upland hardwood fores
poun iia aio mile lane) ‘o
McDaniel 31047, Morris et al. (1 on
Physalis pubescens L.; Pullen et al. (19
Physalis virginiana Mill., BP. IF, ae rests rivers/
creeks (riparian-mudflats drainage
ditches seater |
McDaniel 21234
| li, LIF roadsides, 0;252, Clonts
=
O; a 835,
821
Solanum ptychanthum Dunal, PR, mesophytic
upland hardwood forests, U; 890
STAPHYLEACEAE
Staphylea trifolia L., IF, bottomland hardwood
forests, A(l), $3; 280, 352 (R280), 978 (R280),
1368 (R280), Blocker s.n. (MISSA), Channell s.n.
(MISSA), Theriot et al. (1993)
STYRACACEAE
Styrax americanus Lam.,
—
as
IF, bottomland hard-
BRIT.ORG/SIDA 20(2)
wood forests, lakes/ponds/impoundments
(riparian—banks), U/C; 949, Leidolf s.n.,
McDaniel 1820a, Iheriot et al. (1993)
SYMPLOCACEAE
Symplocos tinctoria (L.) L’Hér., IF, bottomland
hardwood forests, C; 903, 948, Askew 47,
Theriot et al. (1993)
TILIACEAE
‘ilia americana L., BP, PR, bottomland hardwood
forests, mesophytic upland hardwood for-
ests, rivers/creeks (riparian—banks), O/C(I);
Bryson 7430, Morris 3085, Morris et al. (1993)
ULMACEAE
Celti fee eel BP, PR, IF, bottomland hard-
ood forests, mesophytic upland hardwood
ee xeric upland hardwood forests, pine-
mixed hardwood forests, prairie cedar wood-
lands, C; 233, 936, Leidolf and McDaniel
(1998), Lowe (1921), Morris et al. (1993),
Theriot et al. (1993)
Planera aquatica J.F.Gmel., IF, bottomland hard-
wood forests, rivers/creeks eet eels :
R/O; Leidolfs.n., Theriot et al. (19
Ulmus alata Michx., BP, PR, IF, Ean hard-
wood forests, mesophytic upland hardwood
forests, pine forests, pine-mixed hardwoo
forests, lone Prairie cedar woodlands,
chalk outc
(riparian— ae iogeeides Cc 39, 350, 634,
1336, Leidolf and Bid : 998), Morris et
al. (1993), Theriot et al. (19
Ulmus americana L., BP, PR, IF, | eae hard-
Ww forests, mesophytic upland hardwood
eens pine tree pine-mixed hardwood
-24,37,Morris et al. (1993),
—
=
ee
es
forests, roadsid
Theriot et al. (1 -
Ulmus rubra Muh. a PR, bottomland hardwood
forests, mesophytic upland hardwood for-
ests, prairie cedar woodlands, me ate
(riparian—banks), O; 104, Leidolf and
McDaniel (1998), Morris et al. (1993)
Ulmus serotina Sarg., PR, mesophytic upland
hardwood forests, R, $3?; 884, Anderson s.n.
(MISSA), Crawford s.n. (MISSA), Henry s.n.
(MISSA), Howell s.n. (MISSA), — Sn.
(MISS 10) Medal: ioe Smith 1
URTICACEAE
Boehmeria cylindrica (L.) Sw., BP, IF, bottomland
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI
hardwood forests, rivers/creeks (riparian—
mudflats), A();673, Theriot et al. (19
Parietaria pensylvanica Muhl. ex Willd., BP, lakes/
ponds/impoundments (riparian—mudflats),
A(l); McDaniel 31056
Pilea pumila (L.) A.Gray, IF, bottomland hardwood
forests, swamp forests, rivers/creeks (ripar-
ian—banks), O/A; 905, Smith 944
VALERIANACEAE
*Valerianella locusta (L.) Betcke, PR, urban areas,
O; McDaniel 21186
Valerianella radiata (L.) Dufr., BP. PR, IF, bottom-
land hardwood forests, roadsides, urban ar-
eas, C/A; 101, Askew 66, Jones (1976b),
McDaniel 1666, 31036
ERR ENS ERE
Callicarpa americana L., BP IF, bottomland Pall
ene) forests mesophyt
ey grass/forb Ae C; 449, 857,
Clonts 844, Morris et al. (1993), Theriot et al.
a
Lippia lanceolata Michx. [= Phyla lanceolata
(Michx.) Greene], BP chalk outcrops, culti-
vated fields, O; 475, McDaniel 22926
Verbena bipinnatifida Nutt. [= Glandularia
bipi gees (Nutt.) Nutt. var. see pia
BP. IF, prairies, chalk outcrops, roadsides, R/C;
167, 275, 1329, Bruza 925, ae on /446,
McDaniel 13449, 31019, Morris 3082, Morris et
al. (1993), Pullen et al. (1968b)
a bonariensis L., IF, grass/forb meadows,
; 1655, Bruza 963
“ebeno brasiliensis Vell., BP PR, IF, grass/forb
oadsides, urban areas, C/A(I); 290,
463, i ee: 730, Scrug
Verbena canadensis (L.) es a Glandularia
candensis (L.) Nutt.J], BP; Morris et al. (1993)
*Verbena rigida Spreng., PR, IF, grass/forb
meadows, roadsides, U; 847, 1532?
—
A
Verbena simplex Lehm., BP IF, prairies, chalk out-
crops, roadsides, O; 267, 298, Clonts &15,
Leidolf and McDaniel (1998), Lowe (1921),
McDaniel 13448, 31040, Morris 3080, Morris et
al. (1993), Sundell 1538
Verbena urticifolia L., BP, canals/drainage ditches
(riparian—banks), roadsides, U;833, Lowe (1921)
VIOLACEAE
hanthtsc n
, lor (T.F. Forst.) Spreng., PR, me-
sophytic upland hardwood forests, C(I), $2;
Channell s.n. (MISSA), a s.n. (MISSA), Ray
8214 (MISSA), Smith 1
ola affinis Leconte, PR, fs aie hard-
wood forests, mesophytic upland hardwood
forests, pine forests, canals/drainage ditches
(riparian—banks), C; 47, 86, 98, McDaniel 2884,
Morris et al. (1993), Theriot et al. (1993)
Viola bicolor Pursh, PR, roadsides, C; 60
Viola palmata L.,PR, IF, mesophytic upland hard-
wood forests, O; 97, 123, 153(R123)
Viola primulifolia L., IF, mesophytic upland hard-
wood forests, R/O; 198, Bryson 963
Viola sagittata Aiton, BP, IF, bottomland hard-
wood forests, pine forests, pine-mixed hard-
wood forests, chalk outcrops, rivers/creeks
(riparian—banks), C; 87, 106, 118, Bryson 5392,
McDaniel 1590, 1608
SCACEAE
Phoradendron leucarpum (Raf.) Reveal & M.C.
Johnst., PR, urban areas, C; 1203
VITACEAE
Ampelopsis arborea (L.) Koehne, BP. IF, bottom-
land hardwood Loe st) xeric upland hard-
Leidolf and McDaniel (1 a McDaniel 31634,
31655, Theriot et al. (1993)
Ampelopsis cordata Michx., BP PR, IF, bottomland
hardwood forests, roadsides, urban areas, C;
614, 1491, 1533
Cissus incisa Des Moul.,, PR, urban areas;McDaniel
Parthenocissus quinquefolia (L.) Planch., BP. PR, IF,
mland hardwood forests, mesophytic
upland hardwood forests, prairie cedar
woodlands, O; Leidolf and McDaniel (1998
Morris et al. (1993), Theriot et al. (1993)
Vitis aestivalis Michx., BP, PR, IF, bottomland hard-
sts, mesophytic upland hardwood
forests, Rig cedar woodlands, chalk out-
crops, O; Leidolf and McDaniel (1998), Mor-
ris et al. 1983 Theriot et al. (1993)
Vitis cinerea (Engelm.) Millard, BP IF, bottomland
hardwood forests, pine-mixed hardwood
forests, urban areas, R/O; 1446, 1496, fa
Theriot et al. (19
Vitis rotundifolia Michx., PR, IF, bottomland hard-
wood forests, pine forests, pine-mixed hard-
wood forests, C; 1593, Morris et al. (1993),
Theriot et al. (1993)
—
wood fore
750
LILIOPSIDA
GAVACEAE
Manfreda virginica (L.) Salisb. ex Rose, BP IF, prai-
ries, roadsides, C425, 627,802,839, Bruza 967
Leidolf and McDaniel (1998), Locke s.n.
(MISSA), Ray 49 (MISSA)
ALISMATACEAE
Alisma subcordatum Raf., IF, canals/drainage
ditches (emergent), lakes/ponds/impound-
ee ao : SH; Frasier s.n.,
Leidolf p s., MacDonald 10884
Echi pa pene ius (L.) i IF, lakes/
onds/impoundments (riparian—mudflats,
littoral-emergent), O; 726, Easley s.n.(MISSA),
Jones (1974a)
Sagittaria calycina Engelm.; Hare s.n. (MISSA)
Sagittaria lancifolia L.; Blocker s.n. (MISSA), Jones
(1974a)
Sagittaria platyphylla (Engelm.) J.G.Sm.,|F, lakes/
ponds/impoundments (riparian—mudflats,
littoral-emergent), O; 727
ALLIACEAE
*Allium ampeloprasum L.,
meadows, cues O; 31
Allium canadense L., BP, PR, bottomland hard-
wood forests, nesophy aprane nareHOge
BP, grass/forb
]
forests, prairies
(riparian- ee none meadows,
urban areas, O/A; 276, Fisher 82 (MISSA),
Leidolf and peat (1998), McDaniel 31054,
Morris et al. (1993)
Allium mobilense Regel [= A. canadense L. var.
mobilense (Regel) Ownbey], PR, urban areas;
Prescott 5.n. (MISSA
*Allium vineale L., BP, prairies, roadsides, O; 310
idolf and McDaniel (1998
Nothoscordum bivalve (L.) Britton, BP. PR, meso-
phytic upland hardwood forests, prairies,
prairie cedar woodlands, C; 61, 173, Leidolf
and McDaniel (1998), Mitchell s.n. (MISSA),
ee Oe
water 5.n. (MISSA), Sistrunk s.n. (MISSA),
Stauffer s.n. (MISSA), Tringle s.n. (MISSA)
AMARYLLIDACEAE
a
forests, R; Askew 46
*Lycoris radiata (L’Hér.) Herb., PR, roadsides, ur-
ban areas, O; 996
BRIT.ORG/SIDA 20(2)
*Narcissus biflorus Curtis [= N. x medioluteus Mill.
R, roadsides, urban areas,
ies x meoaaal ilis Mill, BP, roadsides,
urban areas, O; 79
*Narcissus jonquilla L., PR, roadsides, urban areas
ARACEAE
Arisaema dracontium (L.) Schott, BP, PR, IF, bot-
tomland hardwood forests, mesophytic up-
land hardwood forests, lakes/ponds/im-
poundments (riparian—mudflats), A(l); 278,
258, Brown s.n. (MISSA), Jones (1974a), Lowe
(1921), McDaniel 31057, Morris et al. (1993),
Theriot et al. )
Arisaema quinatum (Buckley) Schott [= A.
triphyllum (L.) Schott ssp. quinatum (Buckley)
Huttleston], PR, mesophytic upland hard-
wood forests, R; Morris et al.(1993), Tracy s.n.
(MISSA)
Arisaema triphyllum (L.) Schott, PR, bottomland
hardwood forests, mesophytic upland hard-
wood forests, U; 147, Channell s.n. (MISSA),
Jones (1 )
ARECACEAE (PALMAE)
Sabal minor (Jacq.) Pers., IF, bottomland hard-
od forests, pine-mixed hardwood forests,
A(l); 714, Askew 61, McDaniel 2549
ASPARAGACEAE
*Asparagus Officinalis L.,BP, PR, prairies, roadsides
urban areas, U; 804, Channell s.n. (MISSA), 78
(MISSA), Christian s.n. (MISSA), Clonts 490,
Leidolf and McDaniel (1998)
COMMELINACEA
Commelina communis L., BP. canals/drainage
ditches (emergent), roadsides, C; 797, Jones
(1976b)
Commelina diffusa Burm.f. PR, ere
ditches (emergent), urban are
Ke)
Ne)
uw
74a), Moore s.n. (MI
—
be
Commelina virginica L., BP, IF, ee hard-
W rests, rivers/creeks (riparian—
mudflats), canals/drainage ditches (emer-
gent), C; 675, 695, 1374, Brooks 400, Jones
(1976b), Moore s.n. (MISSA), Stauffer s.n.
MISSA)
Tradescantia Hoge is Raf, IF, ae eae road-
sides, C; 256, 326, Brooks 124, Easley s.n.
ea Dea 5.n. (MISSA), pla: 976b),
Rainwater s.n. (MISSA), Stauffer s.n. (MISSA)
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI
Tradescantia virginiana L., BP. IF, bottomland hard-
od ts,mesophytic upland hardwood
ae xeric upland hardwood forests, O/C;
Persons 5.n. (MISSA), Presley s.n. (MISSA), Price
52 (MISSA), Ray 8008 (MISSA), Theriot et al.
(1993), Tracy s.n. (MISSA)
CONVALLARIACEAE
nthemum racemosum (L.) Link, IF, bottom-
gee hardwood forests; Locke s.n. (MISSA),
Shelley s.n. (MISSA), Stauffer s.n. (MISSA),
Theriot et al. (1993)
Polygonatum biflorum (Walter) Elliott, PR, meso-
phytic upland hardwood forests, O; 219, 1348
(R219), Bryson 8331, Channell s.n. (MISSA),
Donald s.n. (MISSA), Easley s.n. (MISSA), Ham-
mer s.n. (MISSA), Morris et al. (1993), Stauffer
s5.n. (MISSA)
Uvularia floridana Chapm., $1; Eckles s.n. (MISSA)
Uvularia grandiflora Sm.; Easley s.n. (MISSA)
Uvularia sessilifolia L., IF, bottomland hardwood
forests, mesophytic upland hardwood for-
ests, O; 1297, 1367, 1486(R1297)
CYPERACEAE
Carex abscondita Mack., IF, bottomland hard-
wood forests; Theriot et al. (1993)
Carex albicans Willd. ex Spreng. var. albicans, PR,
mesophytic te hardwood forests, O;
Morris et al. (19
VALIIIA
ex Spreng. var. australis
ic albicans ie H.
Bailey) ene PR, mesophytic upland hard-
wo O; 66
bee Schwein.,PR, IF, canals/drain-
age ditches (riparian—banks), roadsides, O;
176, Bryson 3138 (Charles T. Bryson persona
herbarium [ctb]), 3191 (ctb), 3833 (ctb)
aares aaimeetele ie P. Blesel) E.P. Bicknell, PR, IF,
ts, roadsides, O;
266, Bryson 149, Morris et al. (1993)
Carex atlantica Bailey ssp.atlantica, IF, pine-mixed
hardwood forests, seepage areas; Bryson
3841 (MMNS)
Carex austrina (Small) Mack., PR, IF; Bryson et al,
(1992)
Carex Sete Steud., BP PR, mesophytic up-
land hardwood forests, prairies, O/C(I); 1530,
Bryson is 10,3217, McDaniel 31053, Morris et
al. (1993)
=)
>
Carex blanda Dewey, BP, PR, bottomland hard-
wood forests, mesophytic upland hardwood
751
forests, xeric upland hardwood forests, O; 197
Bryson 416, 2880a, 2880b, 3215, Morris
et al. (1993
Carex brevior (Dewey) Mack. ex Lunell, BPIF, pine
forests, roadsides, O; 225, Bryson 5391, 8600,
Bryson et al. (1992)
Carex bushii Mack., BP xeric upland hardwood
ae prairie cedar woodlands, Cil); Bryson
ae {1 ie a
aa
Ick | 1 | fl
welin
wood forests, roadsides, A(|); 262, Bryson 92
Carex cephalophora Muhl. ex Willd., BP, PR, IF,
mesophytic upland hardwood forests, xeric
upland hardwood forests, pine forests, prai-
ries, rivers/creeks (riparian—banks), urban ar-
eas, O/C; 221, Bryson 19, 40, 42, 98, 139, 346,
350, 1337a, 2877, 3214, McDaniel i &, 2439,
Morris et al. (1993)
Carex cherokeensis Schwein., BP, PR, bottomland
hardwood forests, mesophytic upland hard-
wood forests, prairies, prairie cedar wood-
lands, AA(|); 5, 107, Bryson 5,6, 13, 16,20, 26,27,
126, 352, Bryson (1980), Leidolf and
McDaniel (1998), McDaniel 1816, Schuster 9,
Morris et al. (1993)
Carex complanata Torr.& Hook., PR, IF, mesophytic
upland hardwood forests, pine forests, ca-
nals/drainage ditches (riparian—banks), road-
sides, C; 237, 240, 1335a,1476,Bryson 148, 3141
Carex corrugata Fernald, BP, PR, IF, bottomland
hardwood ee mesophytic ae hard-
s, O; 190, 229, 1520a, Bryson 93
a on = oo ae 3827 fe 5631 (ctb),
5635 (ctb), 5641 (ct
Carex crebriflora vena IF, bottomland hard-
wood forests, pine-mixed hardwood forests;
Bryson 675, 3063, 3200, 3656, Rosenkranz s.n
Carex crus-corvi Shuttlew. ex Kuntze, IF; Bryson
Carex debilis Michx., IF, bottomland hardwood
Bryson 376, 3139, Theriot et al. (199
Carex digitalis Willd. va da F ld [=
digitalis Willd.], IF; et 373 (ctb), 377 a
Carex festucacea Schkuhr ex Willd., BP, IF, bottom-
land hardwood forests, pine forests, canals/
drainage ditches (riparian—banks), grass/forb
meadows, roadsides, O; Leidolf pers. obs.,
Brooks 131, Bryson 282, 366, 3191
Carex ainmecaie Dewey, BP, IF, bottomland
pee Whee wee forests, meso-
1 pine forests
a
Sie Maine: ditches (emergent), G aa
1295, 1299, 1309, 1478, 1523a, Brooks 127,
Bryson 17,100, 128,147, 155,309,429 (ctb), Hp
(ctb), 1853 (ctb), 3060 (ctb), 3071, 3209 (ctb),
3829 (cto), McDaniel 1675, 2438, Theriot et al.
(1993)
Carex frankii Kunth, BP. PR, IF, bottomland hard-
wood forests, lakes/ponds/impoundments
(littoral-emergent), canals/drainage ditches
g ; pee areas, roadsides, C; 384,
575, 1493, 1559, 1 1, Bryson 283, 286,
Morris or ee a 4017
Carex glaucodea Tuck. ex Olney [= C.
flaccosperma Dewey var.glaucodea (Tuck. ex
Olney) - BP, Bryson 11671 (ctb), Morris
et al. (199
Carex occ Steud., BP, PR, IF, ey
wood forests, A(l), $2S
McDaniel 31 pe et al. (1993)
Carex granularis Muhl.ex Willd., BP, prairies, O/C(1),
1529, Bryson 135, McDaniel 31432, Morris et
al. (1993)
Carex hirsutella Mack., BP. IF, prairies; Brooks 138,
McDaniel 11621
Carex hyalinolepis Steud.,, IF, ees
ditches Bryson 285,
21)
Carex intumescens Rudge, IF, bottomland hard-
wood forests, swamp forests, A(I); 473, 1376,
Bryson 90, Theriot et al. (1993)
ee ld Perel BP, PR, oe up-
prairies, R, $182; 7537,
McDaniel ce Morris et al. (1993), Need
and Bryson (1990)
Carex joorii L.H ee IF, bottomland hardwood
fo rests, swamp forests, canals/drainage
es (emergent), O; 912, Bryson 304, 3440
eae laevivaginata (KUk.) Mack., PR, seepage ar-
eas; Bryson 3839 (ctb)
Carex laxiflora Lam. var. serrulata FJ. Herm., PR,
mesophytic upland hardwood forests, R, $1;
334a, Morris et al. (1993)
Carex leavenworthii Dewey, BP. PR, IF, bottomland
hardwood — roadsides, C; 196,228, Mor-
riset al. (19
~~
4
(emergent);
ottomla hardwood for-
BRIT.ORG/SIDA 20(2)
2029 (ctb), 3069 (ctb), 3431 (ctb), 7013 (ctb),
8608 (ctb), 8610 (ctb)
Carex louisianica L.H. Bailey, IF, bottomland hard
wood forests, A(I);456, 970 (R456), 1578 (R456)
Carex lupulina Muhl.ex Willd. IF, mesophytic up-
land hardwood forests, O; 558, 1579 (R558)
Carex lurida Wahlenb., IF, seepage areas, road-
sides, O; 1479, Bryson 150
Carex meadii Dewey, BP. PR, prairies, O/C(I), 3$4;
524a, Bryson 44, 59,415, 1822 (ctb), 3143 (ctb),
3640 (ctb), 3644 (ctb), 3738 (ctb), 3825 (ctb),
oe (ctb), 5402 (ctb), 11668 (ctb), Bryson
1980), Morris et al. (1993), Tracy 37 (MISSA)
Carexm icrodonta lorr.& Hook., BP, prairies, O,S2?;
1522, Bryson 11667 (ctb), Bryson (1980),
Baon et al. (1992)
Carex molesta Mack. ex Bright, BP, PR; Bryson et
al. (1992)
Carex muehlenbergii Schkuhr ex Willd., BP. PR, IF,
bottomland hardwood forests, mesophytic
upland hardwood forests, prairies, canals/
drainage ditches (riparian—banks),O/C(|); 172,
61, Bryson 4409, McDaniel 11620, Morris et
al. (1993)
Carex ease is Torr. & Hook. var. oxylepis, BP, PR,
mland hardwood forests, mesophytic
a hardwood forests, prairie cedar
woodlands, O; Bryson 130,353, 354 (ctb), 2881
(ctb), 3830 (ctb), 5637 (ctb), 5643 (ctb), 5644
(ctb)
C
Carex oxylepis Torr. & Hook. var. pubescens J.K.
Underw., PR; Bryson 3218 (ctb), Bryson et al.
(1992)
Carex pigra Naczi, BP, PR; Bryson 11672 (ctb), 12348
Carex ae. Naczi, BP, PR, IF, mesophytic
land hardwood forests, ecetaees (ri-
in banks), roadsides, O; 222, 1356(R222),
on 24, 25,93, 133, 2879, 3516,
ae ae (L.H. Bailey) Small, IF bottomland
hardwood forests, O/C; 246, McDaniel 1824
Carex tote an ee willa, PR, IF, oe
pine forests,
ee 351
Carex rosea Schkuhr ex Willd., IF; Bryson 369 (ctb)
Carex socialis Mohlenbr.& Schwegman, BP, PR, IF,
ottomland hardwood forests; Bryson 966
(MISSA), 5394, Tracy s.n. (MISSA)
Carex longii Mac
ests; an 153 va 282 as 623 (ctb), 707,
Carex squarrosa L.,|F; Bryson 284
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI
Carex striatula Michx., PR, mesophytic upland
hardwood forests, O; 130
Carex stricta Lam., $2; Bryson (1980)
Carex oe Buckley, PR, IF, mesophytic up-
land hardwood forests, pine-mixed hard-
- c rests, O; 193, Bryson 3055, 3137
ae oe Boeck., BP, PR, IF, mesophytic
upland hardwood forests, xeric upland hard-
wood forests, pine forests, rivers/creeks (ri-
parian—banks), canals/drainage ditches (ri-
parian—banks), roadsides, O/C; 236, Bryson 23
38, ve ee ae 156, 1859, 2028, Copeland
niel 18
oe aa ae IF, bottomland hard-
ests, rivers/creeks (riparian—banks),
eae ditches (emergent), O; 358,
474, Bryson 676, 3072, Theriot et al. (1993)
Carex typhina Micky IF bottomland hardwood
forests,
Carex umbellata Schkuhr ex Willd., BP, PR, xeric
upland hardwood forests, roadsides, U; 1263,
Bryson 5164, Naczi and Bryson (1990)
Carex vulpinoidea Michx., PR, mesophytic vs
hardwood forests, O; Morris et al. (19
Cyperus acuminatus Torr. & Hook. ex ae ca-
nals/drainage ditches (emergent), C; 406,
10110, MacDonald 10887
ban areas, O; 7
Bryson Debs Copemnaeine McDani see
Cyperus croceus Vahl, IF, pine forests, U; 876a,
Bryson 698
=
—
Cyperus echinatus (L.) Wood, IF, See
meadows, roadsides, C; 483, 506, 546, 6
Potts 3536, Wigley 314 019
Cyperus erythrorhizos Muhl., IF, lakes/ponds/im-
ndments a mudflats), C; 987,
Bryson 3439a, 3439b
*Cyperus pes L., BP IF, cultivated fields, C;
Cyperus flavescens L., PR, urban areas, U; 785
*Cyperus iria L., IF, lakes/ponds/impoundments
(riparian—-mudflats), C; 606, Bryson 345, 3428,
Pullen et al.(1968b), Wigley 312059
Cyperus odoratus L., BP IF,canals/drainage ditches
ae, banks), urban areas, C; 623, 967,
Bryson 3441, Carraway 179, McDaniel 31331
see pola Rottb., IF, seepage areas,
oadsides, O;
Ce er eae Steud., IF, rivers/creeks (ri-
parian—banks), C; 357, Bruza 980, Bryson 689
Cyperus retroflexus Buckley, BP, cultivated fields;
and Carter (1992)
Cyperus retrorsus Chapm., IF, grass/forb meadows,
O; 662, Bryson 3437, Lowe (1921)
*Cyperus rotundus L., BP. IF, seepage areas, culti-
vated fields,C; 412,540, 708, Bruza 957, Bryson
2135, 3427, Wigley 313036
Cyperus strigosus L., IF, canals/drainage ditches
(riparian—banks), roadsides, C; 484, 550, Bruza
960,961, Bryson 701,3438,Lowe (1921), Wigley
315070
Eleocharis compressa Sull., BP; Bryson 3146 (ctb)
Eleocharis elliptica Kunth, BP; Bryson 1858 (ctb)
Eleocharis montevidensis Kunth, BP; Bryson 8920
ctb
Eleocharis obtusa (Willd.) Schult. BP, IF, canals/
drainage ditches (emergent), C;430,814, 866,
Bruza 959, Bryson 685, McDaniel 2539, Wigley
312060
Eleocharis quadrangulata (Michx.) Roem. &
Schult. IF, lakes/ponds/impoundments (lit-
toral-emergent), U; 607, Bruza 982, McDaniel
Britton, BP; McDaniel ne
(\A/i ill.) Sc chult UF
2550
oe. smallii
qrIic FONTS
e ditches (emergent), O; /
ee annua (All.) Roem. i Baile PR, IF,
cons ditches (emergent), urban
areas, O; 783, 862, McDaniel 272
pena autumnalis (L.) Roem. & Schult., IF
nals/drainage ditches (emergent), O; 860,
863, Bryson 2610, McDaniel 2728
Fimbristylis decipiens Kral, BP; MacDonald 10886
Fimbristylis littoralis Gaudich. [= F. miliacea (L.)
ahl], IF, canals/drainage ditches (emergent),
n i
O; 864
Fimbristylis p (Mi eo 151
Fimbristylis tomentosa Vahl; Bruza 96
Kyllinga brevifolia Rottb., PR, urban areas, O; 786,
McDaniel 7526
Kyllinga odorata Vahl, PR, IF, ane areas, grass/
orb m s, urban areas, C; 546, 846, 965,
1560, Bryson oy, Carraway a
Rhynchospora caduca Elliott, IF, seepage areas,
C(I); 491, McDaniel 2547
Rhynchospora corniculata (Lam.) A. Gray, IF, ca-
ai ditches (emergent), C; 442,
Bruza 9/9
erynchspo ea aris Sau ) Small; Brooks
490, Bru 35, McDaniel 2
Rhy merata (L.) Vahl, IF, CG; 1477
Res menoande (Michx) Vahl, IF, seep-
C eee
ey
Scirpus atrovirens Willd., IF, oe
ditches (emergent), roadsides, O; 5
Scirpus cyperinus (L.) Kunth, IF, in.
poundments (riparian-mudflats, littoral-
emergent), O; 583
Scirpus koilolepis (Steud.) Gleason, IF, urban ar-
A(l); 1313
Scirpus lineatus Michx.; Carter 3010
Scirpus pendulus Muhl., BP, canals/drainage
itches (emergent), O; 170
Scleria oligantha Michx., BP, prairies, R/O; 15217,
McDaniel 1087
DIOSCOREACEAE
Dioscorea villosa L., PR, IF, bottomland hardwood
forests, mesophytic upland hardwood for-
ests, U;668, Jones (1976b), Phares s.n. (MISSA),
Theriot et al. (1993)
HEMEROCALLIDACEAE
*Hemerocallis fulva (L.) L., BP roadsides, urban
areas, C; 317
HYACINTHACEAE
Camassia scilloides (Raf.) Cory, BP, grass/forb
meadows, $283; Bryson 58, McDaniel 31379
*Hyacinthoides nonscripta (L.) Chouard ex
R ,PR, urban areas, R; Askew 55
*Muscari botryoides (L.) Mill.; Easley s.n. (MISSA)
*Muscari neglectum Guss. ex Ten., PR, roadsides;
urphy s.n. (MISSA)
HYDROCHARITACEAE
Limnobium spongia (Bosc) Rich. ex Steud., IF,
lakes/ponds/impoundments (littoral—
rooted-floating); Stewart 123
Najas i mae (Spreng.) Magnus, PR, ca-
nals/dr e ditches (submergent), O; 924
wee minor at PR, lakes/ponds/impound-
ments (submergent); Nuttle pers. obs.
HYPOXIDACEAE
Hypoxis hirsuta (L.) Coville, BP, IF, bottomland
hardwood forests, mesophytic upland hard
wood forests, pine-mixed hardwood forests,
prairies, roadsides, O/A(I); 173, 242, Arnett 91
(MISSA), Channell s.n. (MISSA), Locke s.n.
(MISSA), McDani ey
fo
BRIT.ORG/SIDA 20(2)
snplineniea
*B hinensis (L.) DC., PR, roadsides, U;
Ray 104 (MISSA), oe 1444
ris virginica L., IF \s/drainage ditches (emer-
gent), O; 254
Nemastylis era hae Nutt., BP, prairies, chalk
outcrops, C(I)/R, $2; 1510, McDaniel 31002,
orris 3094, ae re al. (1993)
Sisyrinchium albidum Raf., BP, PR, prairies, chalk
outcrops, C; 116, 148, Leidolf and McDaniel
(1998), Morris et al. (1993)
Sisyrinchium angustifolium Mill, IF, bottomland
ardwood eee pine forests, O; 250,
McDaniel 24
Sisyrinchium ane E.P. Bicknell, BP. PR, me-
sophytic upland hardwood forests, prairies,
O; 166 op. McDaniel 31024
ue um mucronatum Michx., IF; Copeland
=
we un rosulatum E.P. Bicknell, IF, seepage
areas, cultivated fields, urban areas, O/C; 499,
JUNCACEAE
Juncus acuminatus Michx., BP, IF, rivers/creeks
(emergent), yoda aes pls (emer-
gent), O; Bryson 1854, Mc 028
Juncus biflorus Ell, IF, iacadares ae for-
s, canals/drainage ditches (emergent), C/
362, Bryson 687, McDaniel 2538, 3149
Juncus ie oe Engelm., IF; Webster 1539
Juncus bufonius L., BP, PR, cultivated fields, CG; 1469,
Bryson 3289, 3501
Juncus coriaceus Mack.,
IF, ea
ditches (emergent), C; 360, Br 56
Juncus debilis A. Gray, IF, oe ditches
(emergent), O; 865
Juncus dichotomus Elliott, IF, canals/drainage
— sleet Bryson 1855
ur ckley, IF, canals/drainage
ditches aa. a
Juncus effusus L., IF, canals/drainage ditches
(emergent), O; 241
Juncus elliottii Chapm,, IF, lakes/ponds/impound-
ts (riparian-mudflats); Bryson 686
~_ enue Buckley, BP, $384; Maddox
es eusnene Rostk,, IF, canals/drainage
ditches (emergent), O; 362, McDaniel 2538,
149
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI
Juncus nodatus Coville, IF, canals/drainage
ditches (emergent), lakes/ponds/impound-
ments (littoral-emergent), U; 510, 656
Juncus scirpoides Lam., IF, canals/drainage
ditches (emergent); Bruza 930
Juncus tenuis Willd., BP PR, IF, mesophytic upland
hardwood forests, xeric upland hardwood
forests, pine forests, prairies, rivers/creeks (ri-
parian—banks), canals/drainage ditches
(emergent), urban areas, O/C; 378,476, 1293,
Bryson 8, 15, 699, 1682, 2027, Copeland s.n.,
Leidolf and McDaniel (1998
Juncus torreyi Coville, BP IF, prairies, canals/drain-
age ditches (emergent), cultivated fields,
grass/forb meadows, C; 288, 383, Bryson 700,
2030, 3994, Leidolf and McDaniel (1998),
McDaniel 31082
Juncus validus Coville, IE bottomland hardwood
forests, canals/drainage ditches (emergent),
lakes/ponds/impoundments (riparian—
mudflats, littoral-emergent), C; 367, Bruza
926,972, Bryson 727
Luzula bulbosa (Alph.Wood) Smyth & Smyth, BP
xeric upland hardwood forests, O; 1264,
318
CS
aechinata (Small) F.J.Herm., PR, mesophytic
upland hardwood forests, O; Morris et al.
(1993)
LEMNACEAE
Spirodela polyrrhiza (L.) Schleid., PR, lakes/ponds/
impoundments (littoral/profundal—free-
floating), A(I); 925
LILIACEAE
Erythronium albidum Nutt., BP, xeric ale hard-
wood forests, C(I), $2; McDaniel 1
Lilium michauxii Poir. es co
hardwood forests, R ne oa Morris et al.
(1993)
Lilium superbum L.,PR, IF, bottomland hardwood
forests, RR, $3S4; Leidolf s.n., Phares s.n.,
Theriot et al. (1993)
MELANTHIACEAE
gan aurea WNaltel IF, ae forests, pine-mixed
age areas, roadsides
U/O; 501, 527 ene 564, nes (MISSA),
Jordan s.n.(MISSA), McDaniel 2638,McKay s.n.
(MISSA), Ray 4844 (MISSA), ae (MISSA)
Aletris farinosa L., IF, pine-mixed hardwood for-
ests; Ray 6248 (MISSA)
—
ORCHIDACEAE
Corallorrhiza wisteriana Conrad; Morris pers.obs.
— spicata (Walter) Barnhart, PR, meso-
upland hardwood forests, U; $2; Gor-
on ae
Listera australis Lindl.; McDaniel pers. obs. (speci-
Malaxis unifolia Michx.; McDaniel pers.
Platanthera ciliaris (L.) Lindl. IF, ae pine-
mixed hardwood forests, canals/drainage
ditches tens banks), R/A(I); 597, Bennett
s.n., McDaniel 2637, Ray 4756 (MISSA), 4845
(MISSA), pe a 1442
estes clavellata (Michx.) Luer, IF, pine-
xed hardwood forests; Ray 4818 (MISSA)
ene cristata (Michx.) Lindl. IF, seepage
areas, A(I), $3; 526, Ray 5258 (MISSA)
Mohr, BP. prairie
cedar woodlands, RR, $2?; 926, Leidolf and
aeDain| Mee
p (L.) Rich., IF, seepage areas, O;
966, Morris (1 986 )
Spiranthes lacera (Raf.) Raf, BP.xeric upland hard-
wood forests, RR, $3$4; 937
Spi f I Sheviak, BP, prairies
cee AA(l), $283; 1002, 1006, 1010, Hare
n. (MISSA), Leidolf and McDaniel (1998
“Mortis pee doe (MISSA )
nd, Ick 11 | |
wo
forests, RR, os 970
Spiranthes praecox (Walter) S. Watson, IF, lakes/
impoundments (riparian—banks),
sr areas, O; 1480, Morris (1989), Patts
&7 (MISSA), Tracy s.n. (MISSA)
Spiranthes tuberosa Raf. IF, pine forests, pine-
mixed hardwood forests, U; 876, Ewing s.n.
(MISSA), Morris (1989), Ray 4848a (Missa)
scien vernalis Engelm. & A. Gray, IF, pine-
d hardwood forests, lakes/ponds/im-
ie (riparian—banks), seepage ar-
eas, grass/forb meadows, roadsides, O/C; 321,
Hollimon s.n. (MISSA), Miles s.n. (MISSA), Mor-
ris (1989), Ray 6244 Ween
Tipularia discolor (Pursh) Nutt. PR, ee, ytic
upland hardwood ae e 667, 1216 (R667)
Triphora trianthophora (Sw.) Rydb., IF, tee
hardwood forests, O/C(I), $283; 976, Leidolf
s.n., Theriot et al. (1993)
756
POACEAE (GRAMINEAE)
Agrostis hiemalis (Walter) Britton, Sterns &
Poggenb., IF, seepage areas, cultivated fields,
C; 1467, 1500,Barkworth et al.(In preparation),
Brooks 134, Lowe (1921)
Agrostis perennans (Walter) Tuck., BP, IF, bottom-
land hardwood forests, mesophytic upland
hardwood forests, O; 941, 950, Lowe (1921),
Theriot et al. (1993)
*Aira caryophyllea L.var.capillaris (Host) Mutel {=
A. elegans Willd. ex Gaudin], IF, roadsides, O;
1484, McDaniel 3007
Alopecurus carolinianus Walter; Askew 33, Bryson
Ne)
Apahopaden gerardii Vitman, IF, ag forests, C;
18, Bryson (1980), Low 21)
Andropogon glomeratus aie Britton, Sterns
& rairies, ©: 995 Barkworth et
al. (IN preparation), Leidolf and McDaniel
(1998), Lowe (1921), Morris et al. (1993),
Wigley 314108
Andropogon ace icus L. var. virginicus, BP, prai-
ries, C; 1078, Barkworth et al. (In preparation),
Bryson | Leidolf and McDaniel (1998),
Lowe (1921), Morris et al. (1993
Aristida dichotoma Michx.; Barkworth et al. (In
reparation), McDaniel 22
Aristida longespica Poir.in Lam. var. longespica, BP,
prairies, 7 fe Leidolf and McDaniel (1998),
Lowe (19
Aristida o ha Michx., BP, prairies, roadsides,
: 851, 1599b, Barkworth et al. (In prepara-
ee rald 18
So ae (Walter) Muhl., PR, IF, bot-
tomland hardwood forests, mesophytic up-
land hardwood forests, A(l); 977, 1369 (R977),
Barkworth et al. (In preparation), Brooks 590,
Laird 21, Morris et al. (1993), Theriot et al.
(1993)
Axonopus fissifolius (Raddi) Kuhlm. [= A. affinis
Chase], IF, urban areas, C; 577, Barkworth et
al. (In preparation)
Bothriochloa barbinodis (Lag.) Herter; Barkworth
In preparation)
*Bothriochloa ischaemum (L.) Keng; Barkworth
et al. (In preparation
Bothriochloa laguroides (DC.) Pilger in Engler ssp.
torreyana (Steud.) Allred & Gould [=
Andropogon saccharoides Sw.], BP, prairies, C;
436, Barkworth et al. (In preparation), Leidolf
—
a
=
=~
BRIT.ORG/SIDA 20(2)
and McDaniel (1998), Morris et al. (1993),
Pullen et al. (1968b), Wigley 315065
*Bothriochloa pertusa (L.) A. Camus [=
Andropogon heres .) Willd], BP PR, road-
sides, O; 1001, Allison 8016, McDanie lee
Morris et al.(] | Stewart 167, Wigley 313
Bouteloua curtipendula (Michx.) Torr.in eee
prairies, A(l), $3S4 6/6, Barkworth et al. (In
preparation), Bryson (1980), Leidolf and
McDaniel (1998), Lowe (1921), Morris et al.
(1993)
*Briza minor L., BP. roadsides, O; 1460, Barkworth
In preparation
*Bromus japonicus Thunb.in Murray, BP, prairies
A(l); 1523, Barkworth et al. (In preparation),
Leidolf and McDaniel (1998)
=
iaae
—~
*Bromus tectorum L., BP, roadsides, A(l); 1338,
oo Et al. (In preparation)
asmanthium latifolium (Michx.) H.O. Yates, BP,
PR, IF, Goreomiand hardwood forests, meso-
phytic upland hardwood forests, C; 390,
Barkworth et al. (In preparation), Lowe (1921),
Morris et al. (1993), Theriot et al. (1993)
Chasmanthium ornythorhynchum Nees;
B al. (IN preparation)
Chasmanthium sessiliflorum (Poir.) H.O. Yates, IF,
t hardwood forests, mesophytic
upland hardwood forests,xeric upland hard-
wood forests, C; 453,640, 1547, Barkworth et
al. (IN preparation), Bruza ee Clonts 541,
Lowe (1921), Theriot et al. (19
*Chloris pectinata Benth.; aie et al. (In
eee
L,, IE bottomland hardwood
forests, ickes pone sipoumements (ripar-
ian—-mudflats), O; 633, Theriot et al. (1993)
Coelorachis cylindrica (Michx.) Nash, $1;
Barkworth et al. (In preparation)
*Cynodon dactylon (L.) Pers., BP, cultivated fields
O; 421, Barkworth et al. (In preparation),
Bryson 3421,Lowe (1921)
*Dactylis glomerata L., BP, prairies, A(
Barkworth et al. (In preparation)
bey ec aegyptium (L i Barkworth
al. (In preparation), Lowe (19
oxham acic aie (Desv. ex in Lam.)
Gould ark ssp. aciculare; Barkworth
); 1524,
et al. (IN preparation)
Dichanthelium aciculare la ex Poir. in Lam.)
Gould & CA. Clark ssp. istifolium (Elliott)
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI
Freckmann & Lelong in ed.;Barkworth et al.
(In preparation
Dichanthelium aciculare (Desv. ex Poir. in Lam.)
Gould & C.A.Clark ssp. neuranthum (Griseb.)
Freckmann & Lelong in ed., IF, lakes/ponds/
mpoundments (riparian— lene O; 703
Di ane ium acuminatum (Sw.) Gould & CA.
Clark ssp. fasciculatum (Torr.) Freckmann &
Lelong in ed.;Barkworth et al. (In preparation)
Dichanthelium acuminatum (Sw.) Gould & C.A
Clark ssp. implicatum (Scribn.) Freckmann &
Lelong in ed., IF, seepage areas, roadsides, 494
Dichanthelium acuminatum (Sw.) Gould & CA.
Clark ssp. lindheimeri (Nash) Freckmann &
Lelong in ed., IF, bottomland hardwood for-
ests, pine forests, urban areas, O; 514;
Barkworth et al. (In preparation), McDaniel
13491, Theriot et al.(1993
Dichanthelium boscii (Poir.) Gould & CA. Clark, PR,
IF, mesophytic upland hardwood forests, xe-
ric upland hardwood forests, C; 459, 639,
Barkworth et al.(In preparation), Morris et al.
1993)
Dichanthelium clandestinum (L.) Gould;
Bele wone) et al. (In ieee
Di ult.) Gould ssp.
commutatum; Barkworth et el (In prepara-
tion), Lowe (1921
Dichanthelium commutatum (Schult.) Gould. we
aa (Vasey) Freckmann & Lelong in ed.,P
F, bottomland hardwood ae ave
eek Apeliat= banks), O; nb?
C.A. Clark, IF, roadsides, O; 8
Dichanthelium depauperatum ane Gould, BP,
IF, xeric upland hardwood forests, pine for-
ests, prairies, O; 1358, 1522a, Barkworth et al.
(In preparation) McDaniel 1086, 13490
Dichanthelium dichotomum (L.) Gould ssp.
dichotomum; Barkworth et al. (IN prepara-
tion), Lowe (1921)
Dichanthelium dichotomum (L.) Gould ssp.
microcarpon (Muhl. ex Elliott) Freckmann &
Lelong in ed. [= P microcarpon Muhl. ex El-
liott], IF, bottomland hardwood forests, me-
sophytic upland hardwood forests, O; 455,
Barkworth et al. (In preparation), Theriot et
al. (1993)
Dichanthelium dichotomum (L.) Gould ssp.
nitidum (Lam.) Freckmann & Lelong in ed;
Barkworth et al.(In preparation), Lowe (1921)
Dichanthelium laxiflorum (Lam.) Gould, BP, PR, IF,
pine forests, pine-mixed hardwood forests,
prairies, roadsides, C; 429, 592, 1366a,
Barkworth et al. (In preparation),Lowe (1921),
McDaniel 979
Sienanins nae CigOTanies (Schult. one ue
tal. (|
Dichanthelium oligosanthes (Schult) Gould ssp.
scribnerianum (Nash) Freckmann & Lelong in
ed., BP, xeric upland hardwood forests, O;
Barkworth et al. (IN preparation), McDaniel!
—
1500
Dichanthelium ovale (Elliott) Gould & C.A. Clark
ssp. villosissimum (Nash) Freckmann &
Lelong in ed,;Barkworth et al. (In preparation)
a a (Schult.) Mohlenbr, IF,
phytic upland hardwood forests, ca-
sa canage eee (riparian—banks), O;
2, Barkworth et al. (In preparation
Dichanthelium ravenelii (Scribn. & Merr,) G Gould,
BP. canal litches ( banl
O; 171, 1, Barkworth et al. (In preparation),
Dichanthelium scabriusculum (Elliott) Gould &
C.A. Clark, BP, chalk outcrops, urban areas, R/
O; aie et al.(In preparation), McDaniel
227194, 31045
Dichanthelium scoparium (Lam.) Gould, IF,grass/
forb meadows, C; 493, Barkworth et al. (In
preparation), Lowe (1921)
Dichanthelium sphaerocarpon (Elliott) Gould, IF,
canals/drainage ditches (riparian—banks), O
71,Barkworth et al. (In preparation), Lowe
(1921)
Digitaria ciliaris (Retz.) Koeler var. ciliaris, BP cul-
tivated fields, A(|); 403, Barkworth et al. (In
preparation)
Digitaria fil fae (L.) Koeler; Barkworth et al. (In
preparation) ae a Lowe (1921)
( Muhl., BP prairies
A(|); 994, Barkworth et al. (In preparation),
Copeland s.n., Lowe (1921
*Digitaria violascens Link; Barkworth et al. (In
preparation)
*Echinochloa colona (L.) Link, BP, cultivated fields,
C;402,Barkworth et al.(In preparation), Lowe
(1921), MacDonald 1, Wigley 312053
*Echinochloa crusgalli(L.) PR Beauv.; Barkworth et
al. (In preparation), Wigley 312054
=
. [7
*Echinochloa esculenta (A. Braun) H. Scholtz;
Barkworth et al. (In preparation)
*Echinochloa muricata (P Beauv.) Fernald, BP, cul-
tivated fields, O; 413, Barkworth et al. (In
ation)
*Fleusine indica (L.) Gaertn., BP roadsides, C(I); 68 1,
Barkworth et al. (In ee 3425,
Lowe (1921), Wigley 31300
a site (Vasey ex L.H. Dewey) Scribn.
var.glabriflorus; Barkworth et al. (In
ae
Elymus villosus Muhl. var. villosus; Barkworth et al.
(In preparation)
Elymus vi co inicus L., BP PR, IF, ee ae
wood forests, roadsides, C(I); -
ene etal.(In dase ayn - ne
nald 24, Morris et al.19
Eragrostis capillaris (L.) Nees in Mart., BP, prairies
prairie cedar woodlands, C; 795, 825,
nee et al. (In preparation),Leidolf and
niel (1 ia aml 15510, 22168
eas frankii C.A teud.; Barkworth
ces Lowe (1921)
ee 4 rsuta (Michx.) Nees, BP chalk outrops,
R;Barkworth et al. (In preparation), McDaniel
2293] ANN)
Eragrostis hypnoides (Lam.) Britton, Sterns &
Poggenb.; Lowe (1921), McDaniel 33287
Eragrostis intermedia Hitchc.; Barkworth et al. (In
preparation)
Eragrostis japonica (Thunb.) Trin. [= E. glomerata
(Walter) L.H. Dewey]; Lowe (1921), McDaniel
33290, 33296
a
—
me
—
ah
R | |
AA; hx. KI
al. (In t sfacaration) Lowe (1921)
Eragrostis pilosa (L.) P. Beauv.; Barkworth et al. (In
preparation)
Eragrostis refracta (Muhl. ex Elliott) Scribn., IF,
lakes/ponds/impoundments (riparian-
banks), O; 696, Barkworth et al. (In prepara
ion), Lowe (1921), McDaniel 2730
ee 5 spectabilis eoren) eG IF, ce 101e
cae
meadows; 633
Eragrostis trichodes (Nutt.) Alph. Wood;
Barkworth et al. (In preparation)
Eriochloa acuminata (J. Presl) Kunth [= E. gracilis
(E.Fourn.) Hitchc.];Barkworth et al. (In prepa-
ration), McDaniel 220
Eriochloa aristata Vasey; Barkworth et al. (In
preparation) Lowe (1921)
BRIT.ORG/SIDA 20(2)
*Glyceria declinata Bréb.,|F,grass/forb meadows,
U; 1344, Barkworth et al. (In preparation)
Glyceria striata (Lam.) 15]
Barkworth et al. (In preparation)
Gymnopogon ambiguus (Michx.) Britton, Sterns
& Poggenb.; Barkworth et al. (In preparation),
Lowe (1921)
Gymnopogon brevifolius Trin. Barkworth et al. (In
preparation), Lowe (1921), McDaniel 26518
Hordeum pusillum Nutt., BP, urban areas, O; 1456,
Barkworth et al.(In preparation), Bryson 3487
*Imperata cylindrica (L.) Raeusch., BP, roadsides,
C(I); 1507, Barkworth et al. (In ue eee
eersia lenticularis Michx., IF, bottomland har
d forests, swamp forests, O
Leersia oryzoides (L.) Sw., IF, bottomland hard-
wood forests; Barkworth et al. (In prepara-
tion), Lowe (1921), McDaniel 33292, Theriot
tal. (1993)
Leersia virginica Willd. IF, bottomland hardwood
less rivers/creeks (riparian—banks), O; 689,
(1921), MacDonald 23, Theriot et al.
Hitchc.,BP, prairies, O; 7
—
ne 3)
Leptochloa panicea (Retz.) Ohwi ssp. brachiata
(Ste Snow, BP, canals/drainage ditches
(foaians banks), O; 807, Barkworth et al. (In
preparation)
Leptochloa panicoides (J. Presl) Hitchc., IF, lakes/
pondsimpoundments pata hes A\(l);
Barkworth et al. (In prepara
ie Itiflorum Lam., BP, ee fields, C;
414,Lowe (1921)
Melica mutica Walter, PR, IF, mesophytic upland
hardwood forests, O; 1300, 1310, Barkworth
et al. (In preparation), Lowe (1921)
*Miscanthus sinensis Hsaieee a et al.
(In ation), Mathies
Wee: glabriflora ae IF, bottomland
hardwood forests, canals/drainage ditches
(riparian—banks), U $?; Leidolfs.n., Theriot et
=
=
Nassella leucotricha (Trin. & Rupr.) R.W. Pohl;
Barkworth et al. (In preparation)
Nassel : viridula (Trin.) Barkworth; Barkworth et
al. (IN preparation)
eee anceps Michx. ssp. anceps, IF, meso-
phytic upland hardwood forests, O; 454,552
(R454), Barkworth et al. (In preparation)
Panicum capillare L. ssp. capillare, BP, cultivated
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI
fields, roadsides, C; 471,629, Barkworth et al.
(IN preparation)
Panicum dichotomiflorum Michx. ssp.
dichotomiflorum, BP, roadsides, O; 892,
Barkworth et al. (In preparation)
Panicum flexile ) Scribn
(In | preparation), Lowe (1921
——
Panicum rigidulum Bosc ex Nees in ae a
aed mann & le
a
in Kearney, BP. prai-
rie cedar woodlands, O; 826, Barkworth et al.
*Paspalum racemosum Lam.,; Barkworth et al. (In
picearationy
| Re IC lead teaced
‘wood forests rvers/ereek (riparian— banks),
Leidolfs.n
McDaniel 33289, 33099 Theriot et al. (1993)
Paspalum setaceum Michx. var. ciliatifolium
(Michx.) Vasey; Barkworth et al. (In prepara-
tion)
¥D | ree 1 ic
|
pubes gee
AC ):602
mpoundments eens banks),
pene rigidulum
Lowe (1921)
Panicum verrucosum Muhl.; Barkworth et al. (In
preparation), Lowe (1921), McDaniel 33288
Panicum virgatum L., BP IF, prairies, seepage ar
eas, C; 590, 617, Barkworth et al. (IN prepara-
tion), Leidolf and McDaniel (1998), Lowe
(1921)
Paspalum bifidum (A. Bertol.) Nash; Barkworth et
al. (IN preparation), Lowe (1921)
Paspalum boscianum ee Barkworth et al.(In
picpale ation), Lowe (1921), Wigley 312056
pal njugatum PJ. Bergius; Barkworth et
al. (IN preparation)
*Paspalum dilatatum Poir. in Lam., BP. prairies,
cultivated fields, C; 382, 47 is ea et al.
ie preparation) MacDo
Do
i
J
ments (riparian— audiae U. 632 Banwarth
et cr Us pepaiaton), Lowe (1971)
Fas! Nx
age areas, roadsides, C; 488, Barkworth et al.
(IN preparation), Carraway 177, Clonts 1401,
Leidolf and McDaniel (1998), Lowe (1921)
Paspalum laeve Michx., IF, seepage areas, road-
sides, C; ne Barkworth - . (In preparation),
Lowe (1921), McDaniel 2
*Paspalum notatum Fluggé, - Te C;401,
Barkworth et al. (In preparation), Wigley
314024
BPIF, prairies, seep-
ae
Paspalum pate L.; Barkworth et al. (In
preparatio
Paspalum praecox Walter; Barkworth et al. (In
preparation), Lowe (1921
Paspalum pubiflorum Rupr. ex E. Fourn., BP, culti-
vated fields, U;878, Barkworth et al. (In prepa-
ration), McDaniel 2265
ees in Mart. ssp.
rigidulum; ae et al. (IN preparation),
p | ,road-
sides, C; 486, Barkworth et al.(In preparation),
Brooks 476, Webster 277, Wigley 314026
*Pennisetum ciliare (L.) Link; Barkworth et al. (In
preparation)
*Pennisetum setigerum (Vahl) Wipff; Barkworth
al. (IN preparation)
ae caroliniana Walter, BP, prairies, O; 1364,
Brooks 126, Wigley 313006
KA
ot
lure: MacDonald
1368
“Phyllostachys flexuosa Riviere & C. Riviere, IF,
been ue (riparian-
banks), A(l); 869, MacDonald 13
*Phyllostachys meyeri McClure, ie IF, lakes/
s/impoundments (riparian—banks),
roadsides, A(I); 723,845, MacDonald 1371
*Poa annua be aa euigivete TEI, ubpania areas,
C: 1265
al_(|
Brys ae ee 921)
Poa arachnifera Torr; Barkworth et al. (In prepa-
ation), Lowe (1921
Poa en Muhl. ex Elliott, PR, mesophytic
upland hardwood forests, C; 1336a, 1354,
Barkworth et al. (In preparation), Lowe (1921)
Poa chapmaniana Scribn.; Barkworth et al. (In
preparation), Lowe (1921)
*Poa pratensis L., PR, urban areas, C; Leidolf pers.
obs., Lowe (19721)
Saccharum brevibarbe (Michx.) Pers. var.
brevibarbe; Barkworth et al. (In preparation),
Lowe (1921)
sisal erat (Michx.) Pers. var.
Webster [= Erianthus
contortus us Elliott], ae pine Lee C; 988, Lowe
(1921)
Saccharum giganteum (Walter) Pers. [= Erianthus
giganteus (Walter) F.T. Hubb], IF, pine forests,
C;916,Barkworth et al.(In preparation), Lowe
(1921)
*Schedonorus arundinaceus (Schreber) Dumort.
—
—
760
= Lolium arundinaceum (Schreber) Darbysh.,
Festuca elatior L.J]—BP, PR, bottomland hard-
wood forests, cultivated fields, O; 478, Morris
et al. (1993
sales ce scoparium (Michx.) Nash in Small
um, BP, IF, pine forests, prairies, C;
917, ees et al. (In preparation), Leidolf
and McDaniel (199
*Setaria faberi RAW. Herrm.in F.Rosen, BP culti-
vated fields, O; 898
Setaria parviflora (Poir.) Kerquélen [= S.geniculata
auct. non Willd], BP, IF, prairies, roadsides, C;
386, 543, Barkworth et al. (IN preparation),
Leidolf and McDaniel (1998
*Setaria pumila (Poir.) Roem.& Schult.ssp.pumila
[= S.glauca (L.) P. Beauv], IF, roadsides, C; 542,
Barkworth et al. (In preparation)
Sorghastrum nutans (L.) Nash in Small, IF, pine
forests, O; 987, Barkworth et al. (In prepara-
tion)
ete x almum Parodi, BP, anak fields,
U; 817, Barkworth et al. (In preparation)
*Sorghum halepense (L.) Pers., BP PR, aa ine
hardwood forests, prairies, cultivated fields,
eis A; 380, 420, Barkworth et al. (In
preparation), oo 9, Morris et al.
Wigley 31
Sphenophoti ae - ydb.) ios IF, bot-
| hardwood forests, O; /
ae obtusata (Michx.) mui om urban
O; 1324a, Lowe (1921), McDaniel 1830
Sporobolus clandestinus (Biehler) Hitchc.;
Barkworth et al. (In preparation)
Sporobolus compositus (Poir.) Merr.
compositus |= S. asper (Michx.) Kunth]; ewe
1921), McDaniel 22169
Sporobolus compositus (Poir.) Merr. var.
drummondii (Trin.) Kartesz & Gandhi;
Barkworth et al. (In preparation)
Sporobolus compositus (Poir.) Merr. var. macer
(Trin.) Kartesz & Gandhi; Barkworth et al. (In
preparation)
gabe cryptandrus (Torr.) A.Gray;Barkworth
al. (IN preparation), Lowe (1921)
an robolus indicus (L.) R. Br. [= S. poiretii eel
& Schult.) Hitche], BP. IF, xeric upland h
wood forests, prairies, C;618, 1548, ane
et al. (In preparation), Leidolf and McDaniel
(1998), Lowe (1921)
—
BRIT.ORG/SIDA 20(2)
Sporobolus vaginiflorus ee ex A. Gray) Alph.
Wood, BP prairies, O; 1 Barkworth et al.
(In ash enna ria ves (1998),
Lowe (1921), McDaniel 31318, 31320
Steinchisma hians (Elliott) Nash in Small, IF, ur-
ban areas, O; 513, Barkworth et al. (In prepa-
ation)
Tridens flavus (L.) Hitche. var. flavus, PR, IF, bottom-
land hardwood forests, roadsides, C; 5417,
Barkworth et al. (In preparation), Copeland
s.n., Lowe (1921), Morris et al. (1993), Wigley
314022
Tridens strictus (Nutt.) Nash in Small, IF, roadsides
C; 568, Barkworth et al. (In preparation), Co-
E icinered sae 1921), Wigley 312057
actyloides (L.) L., IF, roadsides, O; 548,
“Barkworh et al. (in preparation), Carraway
181, Lowe (1921), Snow 66
*Trisetum flavescens (L.) P. Beauv.; Barkworth et
al. (In preparation)
aera aes (Munro ex Wright) R.D
Webster [= Brachiaria platyphylla a Giiseb)
Nash in Celi BP, cultivated fields, A(I); 897,
oe et al. (IN preparation), Wigley
—
a
>
12052
eae oa ramosa (L.) T.Q. Nguyen [= Panicum
ramosum LJ, PR, IF, canals/drainage ditches
(riparian—banks), roadsides, urban areas, O/
A(l); 567, Barkworth et al. (In preparation),
McDaniel 2204
Urochloa texana oe R. D. Webster;
Barkworth et al. (In prepara
Vulpia octoflora (Walter) ra var. octoflora [=
Festuca octoflora Walter]; Brooks s.n., Lowe
(1921)
POTAMOGETONACEAE
Potamogeton diversifolius Raf., lakes/ponds/im-
(littoral-rooted-floating);
Channells.n. (MISSA), Jones (1974a), McDaniel
pers. obs. (specimen extant)
Potamogeton nodosus Poir,, IF, lakes/ponds/im-
poundments (littoral-rooted-floating), C(I);
Bruza 983
SMILACACEAE
Smilax bona-nox L., BP, PR, IF, bottomland hard-
wood forests, mesophytic upland hardwood
forests, prairie cedar woodlands, chalk out-
crops, O/C; Easley s.n. (MISSA), Leidol/f s.n.,
LEIDOLF ET AL., FLORA OF OKTIBBEHA COUNTY, MISSISSIPPI
Leidolf and McDaniel (1998), Lowe (1921),
Morris et al. (1993), Ray 100 (MISSA), 6749
(MISSA), Theriot et al.(1993), Tracy s.n. (MISSA
Smilax glauca Walter, IF, bottomland hardwood
forests, pine-mixed hardwood forests, C;
Lowe (1921), Ray 6781 (MISSA), Theriot et al.
(1993)
Smilax lasioneura Hook., BP, PR, mesophytic up-
land hardwood forests, xeric upland hard-
wood forests, C; Easley s.n. (MISSA), Morris
3092, Persons s.n. (MISSA), Ray 6722 (MISSA),
8059 (MISSA)
Smilax pulverulenta Michx., PR, mesophytic up-
Qa
PRIEPIAEENE
Ae Raf IF } | { | Hl |
forests; Blocker s.n. (MISSA)
Trillium recurvatum L.C. Beck, PR, IF, bottomland
hardwood forests, mesophytic upland hard-
wood forests, R/C; 96, 149 (R96), 1261 (R96),
Channel! s.n. (MISSA), Easley s.n. (MISSA),
Fletcher s.n. (MISSA), Morris et al. (1993), Per-
sons $.n. (MISSA), Smith 1935
TYPHACEAE
*Typha angustifolia L., BP, canals/drainage ditches
(emergent), A(l); McDaniel 31081
Typha latifolia L., BP. IF, canals/drainage ditches
=
land hardwood forests, O; 220, Easley s.n. (emergent), A(l); 315, 603, 1569
(MISSA), Lowe (1921), McDaniel 28154, Mor-
ris et al. (1993) XYRIDACEAE
Smilax rotundifolia L., BPPR,IF, bottomland hard- —_ Xyris torta Small, IF, pine forests, canals/drainage
wood forests, pine forests, pine-mixed hard- ditches (emergent), seepage areas, R/U; 480,
wood forests, prairies, C; 232, 1320, Askew 62, 525, 1558, McDaniel 2640, Smith 14
68, Leidolf and McDaniel (1998), Lowe (1921 ZANNICHELLIACEAE
McDaniel 1676, 31028, Morris et al. (1993 Zannichellia palustris L., IF,
Price s.n. (MISSA), Theriot et al. (1993) ents (littoral- eae: tones
Smilax smallii Morong,|F,bottomland hardwood (] ae Ray s.n. (MISSA)
forests; Leido/fs.n., Theriot et al. (1993)
roe, ee
Td KES
forests; Ray 100 (MISSA), Theriot et al. (1993)
ACKNOWLEDGMENTS
We gratefully acknowledge the MNHP for funding part of the field work for
this study through a 1994 Wildlife Heritage Research Grant. Appreciation is
also extended to John R. MacDonald and Randy C. Warren for their assistance
with field work. Mary E. Barkworth and Charles T. Bryson generously offered
their expertise in the identification of Poaceae and Cyperaceae, respectively.
Use of the facilities at UTC was crucial in the final stages of preparation of this
manuscript, and is hereby gratefully acknowledged. Finally, we would like to
thank Mary E. Barkworth, Scott C. Barras, C.T. Bryson, Barney Lipscomb, and
J.R. MacDonald for providing helpful comments on earlier versions of the manu-
script, as well as three anonymous referees for review of the final draft.
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=
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766 BRIT.ORG/SIDA 20(2)
Book REVIEW
RICHARD DwiGuT PorcHEr and DouGLas ALAN Rayner. 2001. A Guide to the Wild-
flowers of South Carolina. (ISBN 1-57003-437-0, hbk.). University of South
Carolina, 718 Devine Street, Columbia, SC 29208, U.S.A. (Orders: 800-768-
2500, 800-768-0740 fax). $49.95, 551 pp, 1] figures, numerous color pho-
tos, 7" x 10 1/4".
This book is near the top of the ‘must-have’ list for anyone interested in plants and plant communi-
ties of the eastern United States. The range of subject matter is far broader than implied by the title—
so much so that one has to wonder at the title selection. The wildflower photos and brief accompa-
nying descriptions (about 700 species) occupy a little more than half of the 551 total pages—the
remainder Bon a cies of information on acc of botany and plant communities in South
Carolina. The | ily serve as the primary basis for courses in plant tax xonomy, plant com-
munities, or related topics, especially as supplemented by “Natural Vegetation of South Carolina”
J.M. Barry 1980) and “Wildflowers of the Carolina Low Country and Lower Pee Dee” (Porcher 1995).
With this nearly unmitigated commendation in introduction, an outline of the chapters con-
é
vevs]
}
- Introduction
Natural History of Selected ene of Vascular Plants (including carnivorous plants, native or-
chids, poisonous plants, woody vines, epiphytic saproph medicinal plants
and folk remedies, edible wild ae natuvalived ae and ducleweeds).
: History of Field Botany in South Carolina
‘Selected Topics on Natural History and Ecology (including ‘Succession in Natural Communities,
Fire in the South Carolina Landscape, Carolina Bays of the Coastal Plain, ‘Reproductive Strategies
of Vascular Plants, ‘Economic and Cultural Notes, ‘Longleaf Pine and the Naval Stores Industry,
‘Agriculture and eas Effects on South Carolina’s Physical Landscape, and ‘Marshes, Swamps,
Seen Bogs, and Fens’
uth Carolina’s Natural Wildflower ae (The ona The neGnent The Fall-Line
ee The Coastal Plain, The Mariti
- Species Descriptions and Color Plates (divided into the same ‘natural communities’ and numerous
subdivivisions to match the preceding chapter, with the addition of ‘The Ruderal Communities’)
- Field Guide to Natural Plant Communities (also divided first by community type, then by county -
then, for those wanting to walk through the real thing, various natural area preserves, State Parks,
and National Forests are identified and discussed under individual counties)
- Indices and other “end” ee Sane keys to selected plant groups, a guide to South Carolina
herbaria, references ntifi
names, and a “general index”)
| register only a single objection—if one is trying to match a plant in hand witha photo in the book,
an inordinate amount of page-flipping may be required. The photographs and brief notes on indi-
vidual species are arranged first il eae: and then by some other criterion that is not clear to
n n within cies of a single family are not grouped together. This objection,
newer: must sow a personal ae of mine, poe pincers recent books (including the earlier S.C.
volume by P )are published the much- ee ssitid
(Porcher & Ray ner) explicitl y intend the book to be a field guic de |
dents find the arrangement useful.—Guy L. Nesom, Botanical Researc h heute of Te xas, 509 Pecan
Street, Fort Worth, TX, 76102-4060, USA.
SIDA 20(2): 766. 2002
ENDEMISM IN THE WEST GULF COASTAL PLAIN:
IMPORTANCE OF XERIC HABITATS
Michael H. MacRoberts Barbara R.MacRoberts
Bog Research Bog Research
740 Columbia 740 Columbia
ee LA 71104, U.S.A. Shreveport, LA 71104, U.S.A.
rbarium and Herbarium
rie um of Life Sciences Museum of Life Sciences
Louisiana State University in Shreveport Louisiana State University in Shreveport
Shreveport, LA 71115, U.S.A. Shreveport, LA 71115, U.S.A.
Bruce A. Sorrie Robert E.Evans
3076 Niagara-Carthage Road NatureServe
Whispering Pines, NC 28327, U.S.A 6114 Fayetteville Road
Durham, NC 27713, U.S.A.
ABSTRACT
We studied the relationship between West Gulf Coastal Plain plant endemics and habitat. Approxi-
uae half of all ae Gulf Coastal Plain endemics are associates of one habitat: xeric sandylands.
Ww m ti f these data to age and former extent of xeric sandylands in the re-
gion, oa cored of the West Gulf Coastal ve from the East Gulf Coastal Plain and a long
and distinctive biotic history.
Key worps: West Gulf Coastal Plain, endemic plants, gross ic dry uplands, xeric sandylands.
RESUMEN
Se han estudiado las relaciones entre las plantas endémicas y el habitat de la llanura costera del
Golfo Oeste. Aproximadamente la mitad de los enema: estan asociados a un habitat: terrenos
arenosos xéricos. Se oper algunas impli tos dat la edad y extension anterior
la region bl islami de la ll
Oeste de la llanura costera del Golfo Este y una historia biotica larga y diferente.
INTRODUCTION
The Atlantic and Gulf Coastal plains have been recognized as a floristic prov-
ince of distinction in part due to the high number of endemic and near-en-
demic species (Takhtajan 1986). In North America, the Atlantic and Gulf Coastal
floristic province is second only to the California floristic province in endemics,
approximately 27 percent of its native species are endemic or near endemic to
it while in the latter province 48% are endemic or near endemic (Sorrie
Weakley 2001). While important local centers of endemism, such as the
Apalachicola and Lake Wales regions of Florida, are well known, the signifi-
cance of the West Gulf Coastal Plain (WGCP) has received almost no attention
in the published literature or has been portrayed as largely depauperate in
SIDA 20(2): 767-780. 2002
768 BRIT.ORG/SIDA 20(2)
endemics (Ricketts et al. 1999). That the WGCP constitutes a regional center of
endemism has been only recently demonstrated (Estill @ Cruzan 2001; Sorrie
& Weakley 2001). In this paper, based on a list of endemics from the WGCP
and their habitat affiliation, we begin a dialogue about evolutionary origin, di-
vergence, and migration for the region.
The WGCP center of endemism encompasses most of eastern Texas and
western Louisiana, southern Arkansas, and southeastern Oklahoma (Fig. L). Its
western margin is not clearly defined, either geologically or botanically
(Fennemen 1938; Christensen 2000). Its eastern boundary is the Mississippi
River floodplain. Our definition is approximately the same as that of Sorrie and
Weakley (2001) except that we encompass slightly more of south-central Texas
because the east Texas flora clearly extends into those counties just east of Aus-
tin and San Antonio (Region 3 of Hatch et al. 1990). Although within the geo-
logical coastal plain, the gulf prairies and marshes (Region 2 of Hatch et al.
1990) are excluded because it appears that this region is a separate center of
endemism (Sorrie @ Weakley 2001), which itself needs detailed study.
METHODS
To develop a list of endemics, we obtained species distributional data from a
wide variety of sources, including regional, state and local floras, floristic at-
lases, published papers, and monographs (among which are Correll & Johnston
1970; Jones 1977; MacRoberts 1984, 1989; Poole & Riskind 1987; Smith 1988;
Bridges & Orzell 1989; Taylor & Taylor 1989; Hatch et al. 1990; Isely 1990; Nixon
S& Kell 1993; Texas Organization for Endangered Species 1993; Thomas & Allen
1993-1998; Walker 1993; Texas Natural Heritage Program 1995a; Turner 1996;
Nesom 1997; Diggs et al. 1999; Kartesz @ Meacham 1999; Louisiana Natural
Heritage Program 1999; Turner in press; and several web sites centering on the
Flora of Texas Consortium), Natural Heritage Program databases, our own field
work, and an examination of herbarium specimens, notably at ASTC, BRIT,
Corpus Christi Museum of Science and History, GH, LSU, LSUS, NCU, NLU,
SBSC, SHST, TAMU (on line), TEX, VDB, and WWE The list was begun inde-
pendently of Sorrie and Weakley (2001) and has continued to develop after their
list appeared.
We consider those species to be endemic whose occurrence is limited to
the WGCP; near-endemic refers to species that are at least 90 percent centered
on the WGCP. These species are listed in Appendix I.
We encountered two major problems in developing the list. First, distribu-
tional information is incomplete in many cases. Various state and regional dis-
tribution maps and descriptions are either incomplete or do not agree. Second,
taxonomic difficulties are frequent: for example, should Trillium texanum
Buckley be considered a species, a subspecies, or a distinct taxon at all? We sim-
ply have followed the predominant view where possible (e.g., Jones et al. 1997:
—
a
MACROBERTS ET AL., ENDEMISM IN THE WEST GULF COASTAL PLAIN 769
i
— ) eg
pe
Fic. 1. Map showing West Gulf Coastal Plain.
Kartesz @ Meacham 1999). We have developed a fairly conservative list, which
we will continue to update and refine in future.
After we had developed the endemic/near-endemic list (which, for the sake
of brevity, will be referred to hereafter as “endemics”), we attempted to deter-
mine for each taxon its community association (habitat) in the WGCP. We ex-
amined all major classifications (Diamond et al. 1987; Texas Organization for
Endangered Species 1992; Harcombe et al. 1993; Foti et al. 1994; Texas Natural
Heritage Program 1995b; Louisiana Natural Heritage Program 1998; Turner et.
770 BRIT.ORG/SIDA 20(2)
al. 1999; Van Kley 1999a, 1999b; Weakley et al. 1999; Hoagland et al. 2000). We
selected the Turner et al. (1999; see also Van Kley 1999a, 1999b) classification
because it is based on quantitative data and is manageable and applicable over
a wide geographic area. With a few modifications, it is presented in Appendix 2
with associated plant group in parentheses and with The Nature Conservancy
(TNC) “ecological group” designations in brackets (Weakley et al. 1999). We then
assigned each endemic to habitat of best fit on the basis of the literature, per-
sonal experience, and herbarium labels. It was not always possible to determine
community type, a few taxa were assigned to two types, anda few were left blank.
RESULTS
Table 1 presents number and percent of WGCP endemics associated with each
community. Of the 96 taxa in our endemic list, we were able to classify all but
three to one or more communities. Number I, xeric sandylands, proved to have
by far the most endemics. Of the 96 endemics, 53 percent are associated with
xeric sandylands. The communities with the next highest number of endemics
are barrens, glades and weches, and bogs and wet pine savannas, each with nine
percent of the total.
Taste 1.Plant communities and number and percent of WGCP endemics associated with each.
Community Number Percent
eal
WUAOAWA= -ONDODOSA =
1. Xeric sandylands
2 Upland pine savanna, mixed woods
3 Loamy dry-mesic uplands
4 Bogs, wet pine savannas
5 Calcareous forest
6
7
8
Sa
Ww
nN
Mayhaw pond
Barrens, glades, weches barrens
Beech slopes
9 Stream course
10 Baygall
11 Bottomland, floodplain
12 Ponds, marshes
13 Prairies
Unknown
Total
mW
Cal
WaAODWAN—-ON— UOC fA
\o
fon)
se
oO
(o)
DISCUSSION
Xeric sandylands have the most endemics, a fact recognized by Bridges and
Orzell (1987, 1989) some years ago. Of the three genera endemic to the WGCP,
Maclura, Brazoria, and Rhododon, two, Brazoria and Rhododon, are associated
with xeric sandylands (Turner 1995, 1996). Xeric sandylands are open to sparsely
a
MACROBERTS ET AL GULF COASTAL PLAIN 7
wooded areas that typically occur on terraces or ridges composed of deep sand,
generally of marine Tertiary origin (McBryde 1933; Bridges & Orzell 1989;
Harcombe et al. 1993; MacRoberts & MacRoberts 1994, 1995, 1996; MacRoberts
et al. 2002; Turner et al. 1999). Water and air move rapidly through these soils.
Soils, where undisturbed, are often cryptogamic. Foliose lichens (Cladonia spp.)
may be common. Stunted Quercus stellata Wangenh., Q. margarettiae Ashe ex
Small, Q. incana W. Bartram, and various Carya species are dominant trees in
xeric sandylands. Other characteristic species are Bulbostylis ciliatifolia (El-
liott) Fernald, Cnidoscolus texanus (Muell.-Arg.) Small, Croton argyranthemus
Michx., Cyperus grayioides Mohlenbrock, Eriogonum longifolium Nutt., E.
multiflorum Benth., Froelichia floridana (Nutt.) Moq., Hymenopappus
artemisiifolius DC., Matelea cynanchoides (Engelm.) Woodson, Mirabilis albida
(Walter) Heimrl, Opuntia humifusa (Raf.) Raf., Paronychia drummondii Torr. &
A. Gray, Pediomelum digitatum (Nutt. ex Torr. & A. Gray) Isely, P hypogaeum
(Nutt. ex Torr. @ A. Gray) Rydb. var. subulatum (Bush) J.W. Grimes, Phlox
drummondii Hook., Polygonella americana (Fisch. & C.A. Mey.) Small, P.
polygama (Vent.) Engelm. & A. Gray, Selaginella arenicola Underwood subsp.
riddellii (Van Eselt.) R.M. Tryon, Streptanthes hyacinthoides Hook., Talinum
rugospermum Holz, Tetragonotheca ludoviciana (Torr. & A. Gray) A. Gray ex
Hall, Thelesperma filifolium (Hook.) A. Gray, Tradescantia reverchonii Bush,
Tragia spp., Yucca louisianensis Trel., and Zornia bracteata J.F. Gmel.
Sorrie and Weakley (2001) identify the Carrizo Sands of Texas as one en-
demic center for the Atlantic and Gulf Coastal plains (see McBryde 1933 for a
description of this area). While the Carrizo Formation is a narrow band run-
ning from the Rio Grande to northeastern Texas and then southward to eastern
Texas and has xeric sandylands scattered throughout, xeric sandylands are not
confined to this formation but occur virtually throughout the WGCP in both
piney woods and post oak savanna in Oklahoma, Arkansas, Louisiana, and Texas
as shown by the distribution of species fidel to thi as discusse d by
MacRoberts et al. (2002). In that study, in order to develop an objective idea of
the distribution of xeric sandylands, we chose 42 fidel species from the total
list of species occurring in this community and mapped these by county and
parish over their ranges (Fig. 2) (see also McBryde 1933; Drawe et al. 1978; Ajilvsgi
1979; Marietta & Nixon 1983; Matos & Rudolph 1985; Bridges & Orzell 1989;
Ward & Nixon 1992: Harcombe et al. 1993; MacRoberts & MacRoberts 1994,
1995, 1996; Hoagland 2000).
While widespread, xeric sandylands are not a dominant community across
the WGCP but are generally found in relatively small patches. At the time of
European settlement it has been estimated that xeric sandylands accounted for
less than one percent of western Louisiana (Louisiana Natural Heritage Pro-
gram 1993) and, although there are no published figures, probably no more than
this in either Oklahoma or Arkansas. In Texas, especially along the Carrizo for-
772 BRIT.ORG/SIDA 20(2)
Fic. 2. Xeric sandyland species distribution. Key ber of species i le list of 42 species | fi h t
-
or literature (see MacRoberts et al. 2002 for further explanation).
mation (McBryde 1933), xeric sandylands were common, but for the whole of
the region this ity is unlikely to have amounted to more than five per-
cent of the area.
While there appears to be disagreement among palynologists concerning
MACROBERTS ET Al GULF COASTAL PLAIN 773
short-term climatic events in the southeast in general and the WGCP in par-
ticular during glacial and post-glacial times as to whether or not there were
south Texas and Mexican refugia (Martin & Harrell 1957; Ellis et al. 1995;
Schmidtling @ Hipkins 1998), the major long-term patterns are relatively clear
(Bryant 1977; Watts 1980; Bryant & Holloway 1985; Webb 1988; Webb et al. 1993;
Delcourt et al. 1993; Delcourt & Delcourt 1993, 2000; Bousman 1998). The pic-
ture that emerges supports the thesis presented here: xeric sandylands (open
post-oak savannas and woodlands on sandy soils) were present as a commu-
nity type within the WGCP since the last glaciation. We know that coarse sands
necessary for the physical development of xeric habitats were present since the
Eocene (McBryde 1933; Sorrie @ Weakley 2001 and references therein). Prone
to frequent water deficits and nutrient limitations, these soils undoubtedly have
always dramatically affected vegetation structure and composition, which
probably explains why percent sand present in surface soils of the Coastal Plain
isa key determinant of vegetation patterns throughout this area (McBryde 1933;
Harcombe et al. 1993; Peet & Allard 1993; Christensen 2000). We also know from
pollen counts from dozens of bogs along the western edge of the WGCP (unfor-
tunately the only area of east Texas with lengthy pollen records) that oaks and
grasses dominated after the last glaciation (Bryant 1977; Bryant & Holloway
1985; Holloway et al. 1987). The prehistoric landscape that is revealed is one
that is not dissimilar to that seen in the region today: oak-dominated sandy
“savannas” and hilltops that act as reservoirs for water that feeds the adjacent
seeps, marshes, streams, and peat bogs of the area (McBryde 1993; MacRoberts
& MacRoberts 1998, MacRoberts et al. 2002). Unfortunately, palynology is sel-
dom precise to species; our hypothesis would be that oaks were probably those
of dryer habitat: Quercus stellata and Q. incana, and that the herbaceous veg-
etation included, in part, the evolving endemics described here. After all, the
peat bog habitats that contain the continuous pollen profiles extending back
thousands of years still exist (see MacRoberts @ MacRoberts 1998; Potzger &
Tharp 1947; Graham & Heimsch 1960; Bryant 1977; Holloway et al. 1987 for
pollen analyses of east-central Texas bogs).
Additionally, the Mississippi River/Embayment has been a migration bar-
rier since the Eocene (Rock 1957; Blair 1958; Sorrie & Weakley 2001). Many of
the similarities between east and west have come about in the last 6000 years
as the climates have approximated one another. Sandylands of the WGCP re-
tain a distinctive flora rich in endemic and near-endemic taxa, which express
more clearly than other communities in the region a long and distinctive phy-
logenetic history. The flora of many WGCP xeric sandylands has been enriched
by the addition of taxa more widespread in the Coastal Plain. In contrast, xeric
sandylands along the western drier boundary of the WGCP appear to retain a
greater percentage of taxa locally characteristic of the region and with a rela-
tively longer affiliation with it (McBryde 1933).
774 BRIT.ORG/SIDA 20(2)
Our data on endemism coupled with palynological and climatic data
strongly suggest that communities analogous, if not homologous, to xeric
sandylands may have beena significant component of the WGCP since the last
glaciation and perhaps earlier. Open oak-hickory woodlands and savannas with
a rich herbaceous layer and many openings may have been the characteristic
landscape over broad areas; perhaps moving south during glacial advances and
north again with their retreat.
We suggest that the WGCP should no longer be overlooked as a distinct
and critical region of endemism within the Coastal Plain. It may be more mean-
ingful to look upon it asa dynamic entity with an independent and, toa certain
extent, isolated ecological and biological history.
APPENDIX 1
Endemics (E) and near-endemics (NE) of the West Gulf Coastal Plain. Nomen-
clature follows Kartesz and Meacham (1999) in most cases. Community (habi-
tat) types in brackets (see Appendix 2
AGAVACEAE fora hookeriana Torr. & A. Gray, (E) [1]
Yucca louisianensis Trel., (E) (1] alafoxia reverchonii (Bush) Cory, (E) [1]
ALISMA ie dbeckia maxima Nutt., (NE) [13]
Rudbeckia scabrifolia L.E. Brown, (E) [10]
Rudbeckia texana (Perdue) P Cox & Urbatsch, (NE)
[4]
ml
=
TACEAE
Sagittaria papillosa Buchenau, (NE) [6
APIACEAE
Eryngium hookeri Walp., (NE) [9 & 13]
Solidag liad,
(A. Gray) Small, (NE) [1]
Symphyotrichum puniceum (L.) A. & D. Love var.
scabricaule (St INT vers) GL. Nesom, (E) [4]
Tetragonotheca ludoviciana (Torr. & A. Gray) A
ASTERACEAE
Berlandiera pumila (Michx.) Nutt. var. scabrella GL.
& B.L. Turner, (E) [1]
Chrysopsis texana Nesom, (E) [1] Gray ex Hall, (E) [1]
Coreopsis intermedia Sherff, (E) (1] Thelesperma flavodiscum (Shinners) B.L. Turner,
Echinacea sanguinea Nutt., (E) [1] (E) [1]
Evax candida (Torr. & A. Gray) A. Gray, (NE) [1] Vernonia texana (A. Gray) Small, (NE) [1]
Gaillardia Sa. (Walter) H. Rock var. winkleri
Peds B.L. aes ae ! [1]
H. Rock, (E) [4]
Helianthus debilis Nutt.ssp. silvestris Heiser, (E) [1]
Helianthus occidentalis Riddell ssp. plantagineus
ray) Shinners, (NE
Hymenopappus artemisiifolius DC., (NE) [1]
Hymenopappus carrizoanus B.L. Turner, (NE) [1]
Krigia wrightii (A. Gray) K.L.Chambers ex KJ.Kim,
(NE) [ J
er hirsuta Muhl.ex Nutt. var. a/biflora (Torr.
oe Shinners, (NE) [1
a cymosa (H. Ness) K. Schumann, (E) [7]
Liatris tenuis eae (E) [2]
Oligoneuron nitidum (Torr. & A. Gray) Small, (NE)
[7]
—
iat
=
=>
ss
BRASSICACEAE
Leavenworthia aurea Torr. var. texana (Mahler)
Rollins, (E) [7]
Lesquerella angustifolia (Nutt. ex Torr. & A. Gray)
S.Watson, (E) [7
Lesquerella pallida (Torr. & A. Gray) S.Watson, (E)
(7]
CAMPANULACEAE
Lobelia puberula Michx. var. pauciflora Bush, (NE)
CAPPARACEAE
Polanisia erosa (Nutt.) H.H. Iltis, (E) [1]
CARYOPHYLLACEAE
Datanuchiacd
' y
dii Torr.& A. Gray, (E) [1]
MACROBERTS ET AL LF COASTAL PLAIN
Paronychia setacea Torr. & A. Gray, (NE) [1]
Silene subciliata B.L. Robins, (E) [2]
COMMELINACEAE
Tradescantia reverchonii Bush, (NE) [1]
Tradescantia subacaulis Bush, (E) [1]
CUSCUTACEAE
Cuscuta gronovii Willd. ex J.A. Schultes var.
calyptrata Engelm.., (E) [11]
CYPERACEAE
Fuirena bushii Kral, (NE) [4]
Rhynchospora glomerata (L.) Vahl. var. angusta
Gale, (NE) [4,10]
EPPHOREIACEAE
Engelm. & A. Gray, (NE) [13]
FABACEAE
Amorpha paniculata Torr.& A. Gray, (E) [4]
Astragalus distortus Torr. & A. Gray var.
engelmanniil (E. Sheld.) M.E. Jones, (E) [1]
Astragalus leptocarpus Torr. & A. Gray, (NE) [1]
Baptisia bracteata Muhl. ex Elliott var. [aevicaulis
(A. Gray ex W. Canby) Isely, (NE) [2,13]
Baptisia nuttalliana Small, (NE) (1]
Dalea phleoides (Torr. & A. Gray) Shinners var.
phleoides, (NE) (1]
Dalea villosa (Nutt.) Spreng var. grisea (Torr. & A.
Gray) Barneby, (E) [1]
Mimosa hystricina (Small) B.L. Turner, (NE) [1]
Pediomelum hypogaeum (Nutt. ex Torr. & A.Gray)
Rydb. var. subulatum (Bush) J.W.Grimes, (NE)
[1
Trifolium amphianthum Torr.& A. Gray, (NE) [2,13
Trifalipipmy bejari rn Moric., (
E) [1]
GENTIANACEAE
Bartonia texana Correll, (E) [10]
HYDROPHYLLACEAE
Nemophila phacelioides Nutt., (NE) [9]
Phacelia glabra Nutt., (NE) [13]
IRIDACEAE
Iris hexagona Walter var. flexicaulis (Small) R.Fos-
ter, (NE) [6]
Sisyrinchium sagittiferum Bicknell, (E) [4]
LAMIACEAE
Brazoria truncata (Benth.) Engelm. & A. Gray var.
ulcherrima (Lundell) M.W. Turner, (E) [1]
Brazoria truncata (Benth.) Engelm. & A. Gray var.
truncata, (NE) [1]
Monarda a Engelm.& A. Gray ex A.Gray,
(2) 5,13]
Monarda viridissima pest (E ) [1]
Physostegia digitalis Small, (E) [2]
Ngee a eee Cantino, (E) [4,13]
ee pulchella Lundell, (NE) [11]
iliatu Benth) Epling, (E) [1]
Jiophylla Engelm. & A. Gray, (E) [1]
Cc tell
LILIACEAE
Trillium gracile J.D. Freeman, (E) [8
Trillium texanum Buckley, (E) [10]
MALVACEAE
Hibiscus dasycalyx Blake & Shiller, (E) [11]
MORACEAE
Maclura pomifera (Raf.) C.K. Schneid., (E) {1 1]
NYCTAGINACEAE
Abronia macrocarpa L. Galloway, (E) [1]
ONAGRACEAE
Oenothera heterophy II 1 Spach SSP. heterophylla,
(NE) [1]
ORCHIDACEA
Spiranthes eae ii Correll, (E) [7
PAPAVERACEAE
Argemone albiflora Hornem. ssp. texana G.B.
Ownbey, (NE) [1]
POACEAE
Panicum brachyanthum Steud., (NE) [4]
Sporobolus silveanus Swallen, (E) [7]
POLEMONIACEAE
Phlox cuspidata Scheele, (NE) [ ]
Phlox dr'ummondii Hook. var.drummondii, (NE) [1]
Phlox nivalis Lodd. ex Sweet ssp. texensis Lundell,
(E) [1]
POLYGONACEAE
a longifolium Nutt. var. plantaginium
elm. & A. Gray, (E) [1]
Eri iogonum multiflorum Benth., (N
Polygonella parksti Cory, (E) [1]
RANUNCULACEAE
Delphinium carolinianum Walter ssp. vimineum
on) M.J.Warnock, (NE) [1]
Thalictrum arkansanum Boivin, (NE) [11]
Thalictrum texanum (A. Gray) Small, (E) (11]
ROSACEAE
Crataegus nananixonii Phipps & R.O’Kennon, (E)
[1]
E) (1]
776 BRIT.ORG/SIDA 20(2)
Crataegus warneri Sarg., (E) [1] Penstemon murrayanus Hook., (NE) [1]
SCROPHULARIACEAE SOLANACEAE
Agalinis caddoensis Pennell, (E) [1 Physalis mollis Nutt.var. variovestita (Waterfall)
Agalinis navasotensis M. Dubrule & J. Canne- Sullivan, (NE) [1]
Hilliker, (E) [7] VALERIANACEAE
Gratiola flava Leavenw., (NE) [7] Valerianella florifera Shinners, (NE) [1]
APPENDIX 2
Community classification of West Gulf Coastal Plain (from Turner et al. 1999;
The Nature Conservancy [TNC] designations in brackets [Weakley et al. 1999)).
1. Xeric sandylands. Grossarenic dry uplands (Tragia group). As a modification of this type we
emphasize the xeric conditions, notably areas referred to as xeric sandylands and Post Oak-Blue
: ack Oak Savanna[TNC 305 and 320 in partl.
Upland pine savanna, mixed woodlands. Arenic dry upland (Schizachyrium group). [TNC 305
and 307 in partl.
. Loamy dry-mesic uplands (Schizachyrium, - allic eld a Chasmanthium groups). We modify
this entry to exclude communities where | below) dominates and place them
with the next entry.[TNC 307 in part].
ed
Ww
} ] ] ] ‘ T)
Tt Pig
4. Bog et pine savannas. ee loz AMY wet |
a [TNC 330 and 3
. Calcareous forest. se ary mesic ae and (C hasmanthium group). Oe 375 in part].
. Mayhaw pond. Clayey wet uf sion (Justicia group).{TNC 340].
. Barrens, ae ae oe bar eee Callirhoe, Dalecovoups) Open, herb-dominated
areas on | s. [TNC 350].
a WV
“I
x
. Beech slopes. Mesic one ne and terraces (Callicarpa and Mitchella groups). [TNC 308].
Stream course. Loamy mesic stream bottom (Mitchella and Arisaema groups) and loamy wet-
mesic stream bottoms (Bignonia and Justicia groups). [TNC 365 in part]
. Baygall. Loamy wet forested seeps (Osmunda group). [TNC 360
11. Floodplain, bottomland hardwood/river bottom communities; includes ae (Callicarpa,
Bignonia, Justicia, and Ceratophyllum groups). This combines Turner et al.1999) ¢
types 12-15.[TNC 385]
ponds and emergent marshes (not in Turner et al. 1999). [TNC 345].
13. Prairies (not in Turner et al. 1999). [TNC 375 in part].
.
_
Co
TEEEEL unity
w
©
UO
m
ms]
ACKNOWLEDGMENTS
Billie Turner, Jason Singhurst, Vaughn Bryant, the curators of all herbaria where
we worked, and two anonymous reviewers are thanked for their help.
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ANNOTATED CHECKLIST OF THE VASCULAK FLORA OF
THE FICKORY- CREEK UNIO TP BIG Pekher
NATIONAL PRESERVE, TYLER COUNTY, TEXAS
Barbara R.MacRoberts and Larry E. Brown
Michael H. MacRoberts ire ee College
Bog Research, 00 Holman
40 Columbia ae TX 77004, U.S.A.
Shreveport, LA 71104, U.S.A. and Herbarium
and Herbarium spun Brann pacer Center
m of Life Sciences 8856 We
Louisiana State University in Shreveport Houston, 1x 77055, u. s A.
Shreveport, LA 71115, U.S.A.
ABSTRACT
Anannotated, vouchered checklist is provided of the vascular plant taxa of the 284 hectares Hickory
Creek Unit of the Big Thicket National Preserve, Tyler County, in southeast Texas. The Hickory Creek
Unit consists of four plant communities: wetland pine savanna, upland pine savanna, pond, an
baygall. Four hundred one taxa are documented and their community affiliations are given. Previ-
ous lists for this unit numbered over 500 taxa. We believe that the discrepancy is the result of the
larger lists being, in part, “conceptual” (what could be there, not what was actually there).
Key Worps: Big Thicket National Preserve, floristics, Hickory Creek Unit, Texas
RESUMEN
de | l le las 284 hectareas
de la ae eee Unit de la Big Thicket National Preserve, Condads de Tyler, en el sureste de Texas.
La Hickory Creek Unit consiste en idad vege sabana humeda de pings sabana
en tierras altas de pinos, estanque, y matorral de Ilex coriacea. S
y se da la filiacion a su comunidad. Los listados previos de esta unidad contenian mas de 500 taxa.
Creemos que la discrepancia es el resultado de que las grandes listas fueron en parte “conceptuales”
(lo que puede estar alli, no lo que esta actualmente alli).
INTRODUCTION
The Big Thicket region of southeast Texas has been characterized as floristi-
cally rich (Gunter 1971; Eisner 1973; Ajilvsgi 1979; Watson 1979, 1986; Peacock
1994; Official Guide 1997), a characterization that persists in spite of the fact
that botanically the entire area is understudied and underdocumented. As yet,
there is no vouchered and annotated plant list for the Big Thicket region, for the
Big Thicket National Preserve, or for any of the twelve Big Thicket National
Preserve units. The best plant lists from the entire area are the one for the Roy E.
Larsen Sandylands Sanctuary, Hardin County (Matos & Rudolph 1985) and the
one for the Little Thicket Nature Sanctuary, San Jacinto County (Peterson &
Brown 1983). Even at the county level, the only vouchered lists are those of
SIDA 20(2): 781-795. 2002
782 BRIT.ORG/SIDA 20(2)
Nesom and Brown (1998), which are considered by the authors to be “prelimi-
nary.” All other plant lists for the Big Thicket region, beginning with Parks &
Cory (1936) and continuing through Watson (1982), U.S. National Park Service
(1993), and Yu (n.d.) are unannotated and are not based on vouchered speci-
mens. Consequently there is no way to determine whether the lists represent
what the author found or whether they represent what Cozine (1993) calls a
“speculative checklist”; a conjectural compilation of what possibly might be
there, not what is actually known to be there.
The purpose of this study was to begin a baseline floristic list for the Big
Thicket region and the Big Thicket National Preserve by producing an anno-
tated, vouchered checklist of the vascular flora for one unit of the Big Thicket
National Preserve.
SITE AND METHODS
The Big Thicket is located mostly within the longleaf pine region of the West
Gulf Coastal Plain in southeast Texas (Parks & Cory 1936; McLeod 1971;
Harcombe & Marks 1979; Marks & Harcombe 1981; Harcombe et al. 1993). The
Big Thicket National Preserve consists of 12 units scattered over seven Texas
counties. The units range from 222 hectares to 10,100 hectares and total about
34,000 hectares (Peacock 1994). The unit we selected for a floristic inventory is
the 284 hectare Hickory Creek Unit, located in southern Tyler County (Fig. 1).
The Hickory Creek Unit, which consists of four plant communities (pond, wet-
land pine savanna, upland pine savanna, baygall), was chosen for inclusion in
the Big Thicket National Preserve because of its pine savannas (see Ajilvsgi [1979],
Watson [1979], Harcombe & Marks[1981|, and MacRoberts & MacRoberts [1998]
for descriptions of these communities). The natural characteristic of the unit
was relatively flat open “grasslands” with scattered longleaf pines. The unit has
little relief and ranges from 35 meters to 42 meters elevation. Aerial photographs
show the unit still partly open in 1974 when the Big Thicket National Preserve
acquired it, but most openings have subsequently been invaded by shrubs and
offsite pines due largely to fire suppression and dormant-season prescription
burns (Streng @ Harcombe 1982; McClung 1988; Bridges & Orzell 1989;
MacRoberts & MacRoberts 2000). Details regarding edaphic and climatic fac-
tors of the Hickory Creek Unit can be found in Deshotels (1978), Watson (1979),
Marks and Harcombe (1981), and Streng and Harcombe (1982). A map of the
unit is in Harcombe and Marks (1979) and in Streng and Harcombe (1982).
Like all the units of the Big Thicket National Preserve, the Hickory Creek
Unit has not been the subject of a floristic inventory although extensive but
desultory collecting and floristic sampling has occurred there (Streng 1979;
Streng & Harcombe 1982; Watson 1982). Nonetheless, the Hickory Creek Unit
is perhaps the best collected unit of the Big Thicket National Preserve. Streng
(1979 and Streng & Harcombe 1982) conducted a study of factors affecting tree
PRESERVE 783
MACROBERTS ET AL., FLORA OF
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density in the Hickory Creek Unit in the late 1970s and made extensive plant
collections. Geraldine Watson, former botanist with the Big Thicket National
Preserve, also collected there in the 1970s and 1980s. Their plant specimens have
been housed largely at Rice and Lamar universities along with a few collec-
tions by Paul Harcombe, Peter Marks, and C. Liu. These collections number about
300 sheets. There are also a few collections housed at SHST and SBSC.
The MacRoberts’ collected 430 specimens from the Hickory Creek Unit
during 13 field trips beginning 7 March 2000 and ending 13 September 2001.
Larry Brown collected about 179 specimens from the unit during four field trips
between 27 April and 5 October 2001 (he had previously collected about 30
specimens between 1979 and 1997).
784 BRIT.ORG/SIDA 20(2)
Since our purpose is to produce a list of taxa known to occur on the Hickory
Creek Unit, a vouchered specimen was considered to be the only evidence ac-
ceptable for inclusion in the list. In all, about 950 herbarium specimens form
the data for this report.
Rejected are three previous lists: Watson (1982) produced a list of about
530 taxa for the Hickory Creek Unit, the U.S. National Park Service (1993) pro-
duced a list of about 1200 taxa for the entire Big Thicket National Preserve, and
Yu (n.d) produced a list of 525 taxa, excluding trees, for the Hickory Creek Unit.
The Yuand National Park Service lists are based largely on the Watson list. Aside
from being unannotated and not vouchered, inclusion criteria are not explained
for any of these lists. Also, all three lists were found to have extensive
misidentifications and dubious entries (Brown & Brown 1996; Brown n.d.).
Nomenclature follows Kartesz (1994), Jones et al. (1997), and Kartesz and
Meacham (1999) in most cases.
The following abbreviations are used:
CL = C. Liu. His specimens are at Rice University.
DS = Donna Streng. Her collections are unnumbered and are at Rice University.
GW = Geraldine Watson. Unless ot
—
her wise indicated, her National Park Service specimens are at
Rice Universit
LB = Larry Brown. Unless eeaiilen indicated, his specimens are at t SBSC.
McL = C.A. McLeod. His unnumbered and are
MM = sae R. and Michael H. MacRoberts. Except where aout their collections are tempo-
rarily at Rice University.
MW = Michael Warnock. His specimens are at SHST.
PH = Paul Harcombe. His specimens are at Rice University.
PM = Peter Marks. His collections are at Rice University.
* = eXOLIC
=
+ =see note at end of list.
The MacRoberts, Streng, Watson, Harcombe, Marks, and Liu specimens have
been annotated by Larry Brown (Brown & Brown 1996; Brown n.d.). Barbara
MacRoberts and Justin Williams examined the specimens of McLeod and
Warnock at SHST. Where we had more than one voucher, up to four have been
listed (see annotated list under results).
In addition to the taxa listed, we designate the plant community or com-
munities with which we found each species most closely associated. Commu-
nity abbreviations are given below and are put in parentheses (see Ajilvsgi [1979]
for additional description of these communities).
(U) = Upland pine savanna (aka Longleaf-Bluestem Uplands).
(W) = Wetland pine savanna (aka Longleal-Blackgum Savannas).
(B) = Baygall/Stream course. A small part of this is Sweetgum-Oak Floodplain and Palmetto-
Oak Flats.
(P) = Ephemeral pond. (These s
—
nallow ponds, which may become dry in drought years, but hold
water longer than other areas, have not been adequately described on the Big Thicket.)
MACROBERTS ET AL., FLORA OF
PRESERVE 785
RESULTS
Annotated list of the vascular plants found in the Hickory Creek Unit taxa.
ACANTHACEAE
Justicia ovata (Walt.) Lindau var. lanceolata
(Chapman) R.W. Long, (P,W) MM 4416; LB
25539
Ruellia humilis Nutt.,(U) DS; MM 4245,4452,4527
ACERACEAE
Acer rubrum L. var. rubrum, (W) MM 4861
AGAVACEAE
eles virginica (L.) Salisb. ex Rose, (U) GW
; DS; MM
MM 4838
a
ee louisianensis Trel., (U
eae
gittaria papillosa Buch., (P ) GW 2298; DS; MM
414] 4160
ANACARDIACEAE
Rhus copallinum L., (VU) MW 6490
Toxicodendron pubescens P. Mill, (U) LB 19236,
26135
Toxicodendron radicans (L.) Kuntze, (U) LB 25164
ANNONACEAE
Asimina parviflora (Michx.) Dunal, (U) MM 4284,
444)
APIACEAE
Centella erecta (L.f.) Fern., (
Eryngium integrifolium Walt., (
4708, 4814; LB 26380
Eryngiu m yuccifolium Michx., (U) DS; MM 4497
age is oe (Walt. : Britt.ssp. filiformis, (W,P)
470
W) DS; MM 4172,4402
W) PM s.n.; MM
Pree nuttallii tee Britt., (W) McL s.n.
AQUIFOLIACEAE
llex coriacea (Pursh) Chapman, (W,B) MM 4862
llex opaca Ait.,(U) MM 4827
llex vomitoria Ait., (U) MM 4830
ARECACEAE
Sabal minor (Jacq.) Pers., (W,B) MM 4866
AUISTOFOCHIACERE
lata Jacq., (U) MM 4272
ASCLEPIADACEAE
Asclepias amplexicaulis Sm., 5
Asclepias longifolia Michx.ssp.longifolia,(W) MM
4
we
S
i)
Asclepias obovata Ell., (U) GW 2215; DS; LB 25550
Asclepias tuberosa L.,(U) GW 2214:DS
ASTE
Amorosia artemisiifolia L.,
Arnoglossum ovatum (Walt.) H
MM 4494, 4706
—
U) MM 4758
E.Robins., (W) DS:
Baccharis halimifolia L.,(U) MM 4912
Berlandiera pumila (Michx.) Nutt. var. pumila, (U)
DS; MM 4277; LB 25152, 26359
Bidens aristosa (Michx.) Britt. (U) DS; MM 4759
Boltonia diffusa Ell., (W) GW 3429; DS; MM 4533
Chrysopsis mariana (L.) Ell. (U) DS; MM 4449
sae ae pilosa Nutt., (U) MM 4697, 4768
m horridulum Michx., (U) MM 4834
a
eS
: noclinium coelestinum (L.) DC., (U) MM 4756
Conyza canadensis (L.) Cranq.,(U) MM 4762
Coreopsis linifolia Nutt., (W) PM s.n.; MM 4794
pa
Coreopsis tinctoria Nutt.,(U) DS
Echinacea sanguinea Nutt.,(U) DS; MM 4439, 4453,
4528
Flephantor linianus R h.,(U) LB 26347
Elephantopus tomentosus L.,(U) MM 4447
Erechtites hieraciifolia (L.) Raf.ex DC.,(U) LB 26161
Erigeron strigosus Muhl. ex Willd., (U) GW 2302;
4162, LB 25165
Eupatorium capillifolium (Lam.) Small, (U) MM
4913
Eupatorium compositifolium Walt, (U) PH & CL
1108
Eupatorium hyssopifolium L., (W) MM 4810; LB
25537, 26150, 26351
Eupatorium lancifolium (T. & G.) Small, (U) LB
26138, 26139
Eupatorium ae we )T.&G.,, (W) GW 2181;
DS; MM 4520; LB 2
Eupatorium mohrii eae: (U) GW 2182;LB 8150,
5545, 26379
Eupatorium rotundifolium L., (W) GW 404, 2282;
DS; MM 4539
Epa enue Sunset DC., aU seme
DS
iN
MM 4787
Euthamia leptocephala (T. & G.) Greene, (U) GW
*Facelis retusa (Lam.) Schultz-Bip., (U) GW 3348
Gamochaeta purpurea (L.) Cabrera, (U) MM 4248;
25171
786
Helenium amarum (Raf.) H. Rock, (U) DS
Helenium drummondii H. Rock, (W) DS; MM 4146
U) DS; MM 4237, 4458
Helianthus angustifolius L.,(W,U) PM s.n.; DS; MM
Helenium flexuosum Raf., (
Helianthus annuus L., (U) DS
Helianthus debilis Nutt., (WU) GW 2357
Helianthus mollis Lam.,(U) GW 2183;DS; MM 4700
Hieracium gronovil L.,(U) MM 4839
Hymenopappus artem istifolius DC., (U) MM 4300
Liatris acidota Enge Im & Gray, (W) PH 901105;
MM 4538;
alot
Liatris elegans (Walt.) Michx.,(U) DS; MM 4764
Liatris pycnostachya Michx., (U,W) DS; MM 4801
Marshallia graminifolia (Walt.) Small, (W) PM s.n.;
M 4507, 4702
Oligoneuron nitidum (T.&G.
7
Small, (U) GW 354:
S; MM 4532
Pityopsis graminifolia (Michx.) Nutt., (WU) D5; MM
4/66
Pluchea foetida (L.) DC., (W,P) DS; MW 6496; MM
4519,470
Pluchea rosea Godfrey, (W,P) MM 4511,4712,4714
Rudbeckia grandiflora (D.Don) J.£.Gmel ex DC. var.
alismifolia (T.& G.) Crong., (U) DS; MM 4526;
25561
HUB REG! a hirta L., (U) DS; MM 4448
n gracile Gray, (U) MM 4294, a 4698
Solidago canadensis L.,(U) LB 2636/7
saiiaoe ludoviciana (Gray) Small, (U)
26146, 26148, 26162
Solidago odora Ait., (U) MM 4767;LB 5574b [RICE
Solidago rugosa P. Mill. (B) GW 3461
*Soliva sessilis Ruiz & Pavon, (U) LB 25150
Symphyotrichum dumosum (L.) Nesom, (B,W,U)
GW 2278, MM 4457, 4534
DS; LB
1 4)
a hum patens (Ait.) Nesom, (U) GW
Symphyat pes ense (Raf.) Nesom, (U) GW
1,4531
8; DS;
ian texanad ater Small, (WU) PH & Liu 907170;
S; LB 26136
BIGNONIACEAE
Bignonia capreolata L.,(U
BLECHNACEAE
Woodwardia areolata (L.) 1. Moore, (B) MM 4886,
4950
MM 4254
>
Woodwardia virginica (L.) Sm., (W,B) MM 4755,
4304; 1B 19231
BRIT.ORG/SIDA 20(2)
BUDDLEJACEAE
Polypremum procumbens L., (U) DS; MM 4537
CAMPANULACEAE
Lobelia appendiculata A. DC.,(U) DS; MM 4236
Lobelia flaccidifolia Small, (W) MM 4399, 4900
Lobelia puberula Michx.,(W,U) GW 2184, 3454; DS;
MM 4754
+Lobelia reverchonii B.L. Turner, (W) GW 3453;DS;
Triodanis perfoliata (L.) Nieuwl. var. biflora (Ruiz
& Pavon) Bradley, (U) MM 4240
CAPRIFOLIACEA
*/ onicera japonica Thunb., (U) we
mea al eae L.,(U) LB 2
m L.,(U) MM 41 i on
CISTACEAE
Helianthemum carolinianum (Walt.) Michx., (U)
D
Lechea mucronata Raf., (U) DS; LB 26140
Lechea tenuifolia Michx., (U) LB 19510
Lee eene
) crux-andreae (
4509. 18261 37
Hypericum galioides Lam., (W) PM s.n.; DS; MM
4157, 4424
Hypericum gentianoides (L.) B.S.P., (U) DS; MM
4500; LB 25572
Hypericum gymnanthum Engelm. & Gray, (U) LB
25571
Hypericum hypericoides (L.) Crantz., (U) MM 4899
Hypericum mutilum L., (P) LB 26131
L.) Crantz, (W) DS; MM
COMMELINACEAE
Commelina virginica L., (U) D
Tradescantia hirsutiflora za (U) MM 4943
CONVOLVULACEAE
Ipomoea cordatotriloba Dennst.,(U) MM 4785
[pomoea pandurata (L.) G.F.W. Mey., (U) DS; LB
26127
a
Jacquemontia tamnifolia (L.) Griseb., (U) MM
4/788
CORNACEAE
Cornus florida L., (U) MM 4852
Nyssa biflora Walt., (P,B,W) GW 2253; MM 4493:
B 25166
Nyssa sylvatica Marsh., (W,B) MM 4311,4909, 4910;
we
he ia
MACROBERTS ET AL.,
CUSCUTACEAE
+Cuscuta sp.,(U) MM 4979
CYPERACEAE
Carex complanata Torr. & Hook., (B,U) MM 4888
Carex frankii Kunth., (P) DS
Carex glaucescens Ell. (P,B) DS; LB 25560
Carex intumescens Rudge, (P,B) MM 4885
Cyperus croceus Vahl, (U) MM 4774
Cyperus haspan L., (W) GW 2202 [SHST]; DS; LB
26363b
Cyperus retrorsus Chapman, (U) LB 26370
ee virens s Michx. (P) DS
arpa Torr. (P,W) MM 4394
Eleocharis tuberculosa (Michx.) Roemer & J.A.
Schultes, (RW) MM 4265, 4282, 4401, 4406
mbristylis dichotoma (L.) Vahl., (W) MM 4395
Fuirena breviseta (Coville) Coville, (W,P) MM 4709;
—
26152
Fuirena bushii Kral, (W,P) MM 4387,4536;LB 25548,
25551
Fuirena squarrosa Michx, (W) GW 2196 [SHST]
Isolepis carinata Hook. & Arn. ex Torr., (W,U) MM
ora colorata (L.) H. Pfeiffer, (PW) DS
Rinenosbord corniculata (Lam.) Gray, (P) LB
25548, 26151
Rte elliottii ‘i Dietr., (RW) MM 47517,
8; LB 2551
pyncropae filifolia ae (RW) MM 4461,4545b;
Rhy gl ta (L.) Vahl. (W) DS
aie Siacient Gray, (W) MM 4421; LB
25525, 25529, 25542a
Rhynchospora harveyi W. Boott, (W) LB 25565
Rhynchospora inexpansa (Michx.) Vahl., (W) LB
2553]
Rhynchospora latifolia (Baldw.ex Ell.) W.Thomas,
(PW) GW 2216; MM 4460
Rhynchospora oligantha Gray, (W) MM 4250,
4310, LB 25521a
Rhynchospora perplexa Britt. (W) MM 4513; LB
4483 [RICE], 25555
Rhynchospora plumosa Ell, (
4259, 1B 25172
Rhynchospora pusilla Chapman ex M.A. Curtis,
(W) DS; MM 4389; LB 25563, 25573
sa rariflora (Michx.) Ell, (W) MM 4252;
25521C
W) DS; MM 4249,
ryncospr recognita (S.Gale) Kral, (W
LB 25536, 25547
) DS; MM
PRESERVE 787
+Scleria be see a Steud, (PW) MM 4514;
LB?
Scleria te eae ) LB 25144
Scleria georgiana Core, (W) MM 4260, 4407; LB
Scleria pauciflora Muhl. ex Willd., (W) LB 25173
Scleria reticularis Michx., (P,W) LB 1606, 6606,
25524b,
Scleria triglomerata Michx., (W,U) MM 4916
CYRILLACEAE
Cyrilla racemiflora L., (B,W) MW 3978, 6483: MM
4869
DENNSTAEDTIACEAE
Pteridium aquilinum (L.) Kun, (WU) MM 4302,4865
DROSERACEAE
Drosera brevifolia Pursh, (W) MM 4163
Drosera capillaris Poir., (W) MM 4306, 4432
EBENACEAE
Diospyros virginiana L.,(U,W) MM 4901;LB 26360
ERICACEAE
Lyonia mariana (L.) D. Don, (W,B) MM 4871
Rhododendron viscosum (L.) Torr., (U,W,B) DS; MM
4450
AccINiTnM arp
um Marsh., (U) MM 4288
ee corymbosum L., (U) MM 4147, 4153,
B 26134
L.,(U) DS; MW 3291; McL
ERIOCAUEACEDS
gulare L., (W) GW 3365; DS; MM
4423, 1B ]
ete anceps (Walt.) Morong, (W) GW
3368; DS; MM 4239, 4434
EUPHORBIACEAE
Chamaesyce maculata (L.) Small, (U) LB 26345
Cnidoscolus texana (Muell.-Arg.) Small, (U) MM
4904
Croton argyranthemus Michx., (U) DS; MM 4292,
4775
Croton capitatus Michx., (U) MM 4757
ton willdenowii G.L. Webster, (U) LB 14891,
523. 25553
llata L.,(U) DS; MM 4777
*Phyllanthus urinaria L., (U) LB 26348
Stillingia sylvatica Garden ex L., (U) MM 4246
Tragia smallii Shinners, (U) MM 4441; LB 19230,
25148
| Bs ph hj
Tragia urticifolia Michx., (U) DS; MM 4238; LB
25149, 25163
*Triadica sebifera (L.) Small, (U,B,W) MM 4850
BACEAE
Aeschynomene indica L.,(U) LB 14893
*Albizia julibrissin Durz., (UV) LB 26349
Baptisia nuttalliana Small, (U) MM 4835, 4854,
4873
Centrosema virginianum (L.) Benth, (U) MW 6493
Chamaecrista fasciculata (Michx.) Greene, (U) DS;
LB 26 144
Cro otolaria sagittalis L _ (U) DS; MM 4256; LB 26] 42
Desmodium paniculatum (L.) DC., (U) LB 26350,
—a
26366
*Kummerowia striata (Thunb.) Schindl., (U) GW
Mimosa hystricina (Small) B.L. Turner, (U,W) DS;
M4289, 4440; LB 2616
Orbexilum simplex (Nutt. ex T.& G.) Rydb., (U) DS
MM 4915; LB 25157
Rhynchosia latifolia Nutt. ex T.& G.,(U) DS
Strophostyles umbellata (Muhl. ex Willd.) Britt., (U)
& Liu 901107
Stylosanthes biflora (L.) B.S.P,
Tephrosia onobrychoides Nutt., (
M :
—=
U) DS
W) DS; MM 4393;
W 6486
*Trifolium dubium Sibthorp, (U) MM 4893
FAGACEAE
Quercus falcata Michx., (U) GW 2281: MM 4840,
4848, 4859
Quercus hemisphaerica Bartr. ex Willd., (B) MM
3 :
Quercus incana Bartr., (U) MM 4
Quercus laurifolia Michx., (U in Ao
Quercus margarettiae Ashe ex Small,(U) GW 2276
Quercus marilandica Muenchh., (U) MM 4837
Quercus muehlenbergil Engelm.,(U) MM 4841
Quercus nigra L., (U,B) GW 2277; MM 4843; LB
26147
Quercus similis Ashe, (B) LB 25154
Quercus stellata Wangenh., (U) MM 4847
Quercus virginiana P. Mill, (U) DS; MM 49171
GENTIANACEAE
Sabatia gentianoides Ell., (
HALORAGACEAE
Prosperpinaca pectinata Lam., (P) MM 4540; LB
26160
HAMAMELIDACEAE
Liquidambar styraciflua L., (U,W) MM 4856
W) DS; MM 4426, 4508
BRIT.ORG/SIDA 20(2)
HYDROPHYLLACEAE
Hydrolea ovata Nutt. ex Choisy, (P) DS; MM 4713
IRIDACEAE
Alophia pe (Graham) R.C. Foster, (U) DS;
MM 4451,
Sisyrinchium a ee Bickn., (UW) MM 4144b,
4159, 4167, 4301[SBSC]
Sisyrinchium rosulatum Bickn.,(U) MM 4257,4914
Sisyrinchium sagittiferum Bickn., (U,W) GW 2304;
JUNCACEAE
Juncus brachycarpus Engelm.,(U) MM 4944,4948
Juncus bufonius L., (W) LB 25173
Juncus dichotomus Ell, (W) MM 4253, 4872, 4902,
LB 25143
luncus diffusissimus Buckl., (
Juncus effusus L., (W) DS
Juncus ellottii Chapman, (
5147, 25
W) MM 4883
W) DS; MM 4881; LB
Rostk., (W) DS; LB 25159
Juncus aoa Coville, (P) DS; MM 4939, 4940
Juncus repens Michx., (P) MM 4946
Juncus scirpoides Lam DS
Juncus tenuis Willd., (W) MM 4896; LB 25145
Juncus validus Coville, (P) MM 4945,4247, 4427
LAMIACEAE
Hedeoma hispida Pursh, (U) MM 4894
Hyptis alata (Raf.) Shinners, (W) DS; MM 4495,
4802
Monarda punctata L.,(U) DS
Physostegia digitalis Small, (W
4525;1B 19235
irl vulgaris L., (U) MM 4255
anthemum tenuifolium Schrad.,(U) MM 4454
ae laria elliptica Muhl.ex Spreng.,(U) MM 4908;
—
—
) GW 355; DS; MM
25161
Scutellaria integrifolia L., (W) DS; MM 4295, 4917
B 25161
t L., (U) MM 4763
LAURACEAE
Persea borbonia (L.) Spreng.,(U) LB 26352, 26377
Persea palustris (Raf.) Sarg.,(U,B,W) MM 4166;MW
a)
a)
>
m (Nutt.) Nees, (U) MM 4855: LB
26376
LENTIBULARIACEAE
Pinguicula pumila Michx., (W) MM 4876; MW
4157
PRESERVE 789
MACROBERTS ET AL.,
Utricularia radiata Small, (P) MM 4875
Utricularia subulata L., (W) GW 3384; DS
LILIACEAE
Aletris aurea Walt., (W) DS; MM 4242; MW 3977
Allium canadense L., (U,W) 4769
Hymenocallis liriosme (Raf) Shinners, (P) GW 2332;
MM 4145; 1B 25170
Hypoxis curtisii Rose, (P) LB 26384
Hypoxis rigida Chapman, (W) MM 4270, 4897; LB
5142
raed
2
Nothoscordum bivalve (L.) Britt, (U) MM 4143
Schoenolirion croceum (Michx.) Wood, (W) DS;
Tofieldia racemosa (Walt.) B.S.P, (
MM 4505; LB 19229
LINACEAE
Linum medium (Planch.) Britt., (U,W) PM 4; DS;
MM 4251, 44
LOGANIACEAE
| mpervirens A. St-Hil. (U) MM 4183
Mitreola sessilifolia (J.F. Gmel.) G.Don, (W) PM 1;
DS; MM 4544
W) GW 350; DS;
LYCOPODIACEAE
evens appressa (Chapman) Cranfill, (W)
P. M 4307, 4430, 4541
topos ella caroliniana (L.) Pichi Sermolli, (W)
LYGODIACEAE
Lygodium japonicum (Thunb.ex Murr.) Sw.,(U) L
6164
oo)
MAGNOLIACEAE
Magnolia ic ae L., (U) MM 4849
Magnolia virginica ins (W,B) MM 4305
MALVACEAE
Hibiscus moscheutos L.ssp.lasiocarpos (Cav.) O.J.
Blanchard, (W,B,P) GW 2185; MM 4403, 4545
MELASTOMATACEAE
Rhexia lutea Walt., (W) DS; MM 4425, 4523
Rhexia mariana L., (W) DS; MM 4428, 4522
Rhexia petiolata Walt., (W) DS
MYRICACEAE
Morella caroliniensis (P. Mill.) Small, (B,W) MM
4170; McL; MW 3292
Morella cerifera (L.) Small, (U,B,W
OLEACEAE
Chionat thus Vil gil CUS ly (U) LB 25151
~
MM 4828
SCS
*/igustrum sinense Lour., (U) MM 4154, 4868
ONAGRACEAE
Ludwigia hirtella Raf., (W,U) DS; MM 4391, 4415;
B 25522
rm
Ludwigia linearis Walt., (W) PM 13;DS; MM 4503,
4710
Ludwigia pilosa Walt., (P) MM 4790; LB 14895
Oenothera linifolia Nutt., (U) DS; MM 4235, 4244
OPHIOGLOSSACEAE
Botrychium biternatum (Sav.) Underwood, (U)
MM 4898
ORCHIDACEAE
Calopogon oklahomensis D.H. Goldman, (W,U)
MM 4864; GW 3358 [TAMU]
Calopogon tuberosus (L.) B.S.P, (W
4937
/sotria verticillata Raf. (B) GW 1475
Platanthera nivea (Nutt.) Luer, (W) GW 349; DS;
MW 6480; McL
) DS; MM 4429,
Pogonia ophioglossoides (L.) Ker-Gawl., (W) DS
Spiranthes brevilabris Lindl., (W) DS
Spiranthes praecox (Walt.) S. Wats., (U,W) GW
3380; MM 4271, 4279
Spiranthes tuberosa Raf. (U,W) LB 18627
daaoaaiin
Osmunda cinnamomed L., (W,B) MM 4863
Ssunda ea L., (W,B) DS; MM 4158, 4261
OXALIDACEAE
Oxalis dillenii Jacq., (U) MM 4142, 4178
Oxalis lyonii Pursh, (U) GW 2303;MM 4161
Oxalis violacea L, (U) DS; LB 26385
PASSIFLORACEAE
Passiflora lutea L.,(U) MM 4443
PINACEAE
Pinus echinata P. Mill, (U) MM 4836
Pinus palustris P. Mill. (W,U) MM 4857
Pinus taeda L., (W,U) MM 4858
PLANTAGINACEAE
lantago aristata Michx., (U) DS
Plantago virgit Ca Le (U) MM 4298
PLATANACEAE
Platanus occidentalis L.,(U) LB 26369
POACEAE
Andropogon gerardii Vitman, (U) DS
Andropogon ternaris Michx., (U) GW 2270; LB
26365
790
Andropogon virginicus L.,(W,P) GW 2272; DS; MM
Anthaenantia rufa (Nutt.) J.A. Schultes, (W) GW
2284; DS; MM 4797; LB 26381
Anthaenantia villosa (Michx.) Beauv., (W) GW
- DS; LB 26364
Aristida longespica Poirvar.geniculata (Raf.) Fern.,
(U) GW 2219; DS
Aristida palustris (Chapman) Vasey, (P,W) GW
2205; MM 4704; LB 26153
Aristida purpurascens Poir. var. purpurascens, (U)
MM 4783
Aristida purpurascens Poir. vat. oe ie All-
red, (U, Sail DS; ele 4800; LB 5. 6143
A di) Kuh! nae
4390, 4778
are laxum (L.) Yates var. laxum, (B)
5157b; LB 26344, 26368
Peli cy pie (Michx.) Nash, (U) MM 4942
Coelorachis rugosa es Nash, (RW) GW 2188;
me MM 4711; LB 261
Pere A pee ae Poir.) Gould &
Clark,(U) DS
Dichanthelium acuminatum (Sw.) Gould & Clark
var. fasciculatum (Torr.) Freckmann, (P.U,W
4262:1B25155,2
Sienaneieliin comm. el (J.A. Schultes
Gould, (W) DS; MM 4
WS
eer
fa) Ky lH 2
\
BRIT.ORG/SIDA 20(2)
NDT) if } Cc } Crt kh py Miihbl
(U) DS
Eragrostis elliottii S.Wats., (W) GW 2
Eragrostis refracta (Ell.) Scribn., (W
Eragrostis secundiflora J. Presl,(U) D
Eragrostis spectabilis (Pursh) Steud, (U) GW 2268,
2287, 1B 14894, 2614]
247
) DS; MM 4796
4
Gymnopogon ambiguus (Michx.) B.S.P., (U) GW
2312; DS; LB 6605
MM 4780
a
+Gymnopogon brevifolius Trin., (U
[SBSC]
Muhlenbergia capillaris (Lam.) Trin. var.capillaris,
W) DS; MM 4793; LB 26373
Panicum anceps Michx., (W) DS
Panicum brachyanthum Steud,., (U,
4770
a)
W) DS; MM
Panicum hemitomon J.A. Schultes, (P) LB 25543
pane cum rn um Bosc ex Nees, (P,U,W) MM
4715 2, LB 25558, 261
ae tenerum Beye x Trin, (RW) MM 4410,
502, 4543, 4815;, LB 6669, 14896, 75523a
Panicum verrucosum Muhl., (W) MM 4812; LB
6607
Panicum virgatum L., (P,W) DS
Paspalum bifidum (Bertol.) Nash, (U) LB 6666
Paspalum ada Michx., (U,W) GW 2190;DS;
MM 477 8
Paspalum fe Michx., (W) LB 26383
Clark, (U,W) MM 4263, 4280, 4412;LB 19232
Dichanthelium dichotomum (L.) Gould var. tenue
aMol I) eouln . sos (W) DS LB 25146a
h) Freckmann,
(W) MM 4234, 4462: LB 25168, 25175
Dichanthelium scabriusculum (Ell.) Gould & Clark,
Dichanthelium scoparium (Lam.) Gould, (U,W)
44] ]
Dichanthelium sphaerocarpon (Ell) Gould var.
sphaerocarpon, (U) MM 4290; LB 25554
Se enriea sphaerocarpon (Ell.) Gould var.
isophyllum (Scribn.) Gould & Clark, (U) LB
25162
—
Dichanthelium villosissimum (Nash) Freckmann,
(U,P,W) MM 4286, 4299, 4444: 1B 26354
Dichanthelium wrightianum (Scribn.) Freck-
mann, (P) LB 26126
Digitaria ciliaris (Retz.) Koel, (U) LB 26346
Digitaria cognata (Schult.) Pilger.ssp.cognata, (U)
MM 4779
=
Paspalum lividum Trin., (P) GW 2222
*Paspalt mm noratiym Flue
U) DS
Paspalum plicatulum Michx.,(U,W) DS; MM 4243,
Paspalum praecox Walt., (W) GW 2192; DS; MM
54]
Paspalum setaceum Michx.,(W) GW 2221;DS; MM
*D alu it :
ru ria
*Poa. annua L.,(U) MM 4831
Saccharum giganteum (Walt. Pers.,(P) CL 901176,
901177
*Sacciolepis indica (L.) Chase, (P) GW 2194
Schizachyrium scoparium (Michx.) Nash var.
divergens (Hack.,) oe (U,W) GW 2186; DS;
M 4765; LB 263
as ium tenerum ane (U) MM 4696, 4782.
ae ee (Poir.) Kerguelen, (U) DS
Sporobolus junceus (Beauv.) Kunth, (U) DS; MM
4/773, 1B 26356
Steinchisma hians (Ell.) Nash, (U) MM 4398, 494]
MACROBERTS ET AL., THE HICKORY
PRESERVE 791
Tridens ambiguus (Ell.) J.A. ai (U,W) GW
2208; DS; MM 4760; LB 2
Tridens strictus Ve Nash, ie 93:LB 5577
[RICE]; MM 4792, 4806
Tripsacum dactyloides (L.) L., (U) DS; MM 4459
POLEMONIACEAE
Phlox pilosa L., (WU) DS
POLYGALACEAE
Polygala cruciata L., (W) GW 353; DS
Polygala incarnata L., (U,W) GW 3385; DS; MM
4276; LB 19233
Polygala mariana P.Mill., (U,W) DS; MM 4278, 4530
Polygala polygama Walt., (U) D
Polygala ramosa Ell., (W) DS; MM 4405
POLYGONACEAE
Polygonum punctatum EIl., (P
PRIMULACEAE
Anagallis minima (L.) Krause, (U) MM 4895
RANUNCULACEAE
Nelnhiniry liny
LB 26372
hss
Walt., (U) MM 4496
RHAMNACEAE
pcre scandens (Hill) K. Koch, (U) MM 4829
gul li (Walt.) Gray, (U) MM 4826
ROSACEAE
Crataegus marshallii Egglest., (U) GW 2301
Crataegus opaca Hook. & Arn., (B,B.W) GW 2305;
MM 4135, 4136, 4890
Crataegus spathulata Michx., (B) MM 4949
Photinia pyrifolia (Lam.) Robertson & Phipps, (W)
DS; MM 4148
Prunus caroliniana (P. Mill.) Ait., (U) LB 26353
Prunus serotina Ehrh., (U) MM 4187
*Pyrus calleryana Dcne.,(U) MM 4184
Rubus argutus Link, (U) GW 2299; MM 4867
Rubus trivialis Michx., (U) GW 2300; MM 4176
RUBIACEAE
Cephalanthus occidentalis L., (W,P) MM 4417,
S15
Diodia teres Walt., (U) DS; LB 26145
Diodia virginiana L., (W) MM 4542;1B 26133
Galium pilosum Ait., (U) MM 4437,LB 25153
Hedyotis nigricans (Lam.) Fosberg, (U) DS; MM
4498
—=
Houstonia pusilla Schoepf, (U) MM 4177
Mitchella repens L, (U) MM 4853
Oldenlandia boscii (DC.) Chapman, (P) LB 19227,
5 70
'
SARRACENIACEAE
+Sarracenia alata Wood, (
SCROPHULARIACEAE
Agalinis fasciculata (Ell.) Raf., (U
Agalinis oligophylia Pennell, (U
M 4769; LB 6603
Agalinis purpurea (L.) Pennell, (U) LB 26343, 26378
Agalinis viridis (small) Pennell, (U) GW 3426
Gratiola brevifolia Raf., (U,W) DS; MM 4266
Gratiola pilosa Michx.,(U) MM 4499
Teiasinia crustacea (L) F.Muell., (P) LB 26363
uminata (Walt.) Small, (U,W) MM
W) MM 4175; McL
DS
GW 3448, 3455;
a FS
4776
Penstemon laxiflorus Pennell, (U) DS; MM 4274
SMILACACEAE
Smilax bona-nox L., (U) PM 15;DS
Smilax glauca Walt., (W,U) DS; MM 4185;LB 19246
Smilax laurifolia L., (W,B) MM 4870
Smilax pumila Walt., (U) MM 4
Smilax rotundifolia L., (U) MM 4903; LB 25160
Smilax smallii Morong, (U) DS; MM 4182,4851
SOLANACEAE
Solanum ptychanthum Dunal, (U) LB 26155
SYMPLOCACEAE
Symplocos tinctoria
VERBENACEAE
Callicarpa americana L., (U) MM 4445
VIOLACEAE
Viola lanceolata L, (W) MM 4174
Viola palmata L.,(U) MM 4138, 4164, 4186
Viola primulifolia L., (W) MM 4144a, 4918
Viola sagittata Ait.,(U) MM 416
VITACEAE
Parthen ;
(L.) Her. (U) MM 41717
—
folia (L.) Planch., (U) MM
4860
Vitis rotundifolia Michx., (U) MM 4833
XYRIDACEAE
Xyris ambigua Bey. ex Kunth, (W
Mcl; LB 26132
Sr
) DS; MM 4703;
Xyris baldwiniana J.A. Schultes, (W) MM 4156,
4397, 4419;/1B 25524a
Xyris stricta Chapman var. obscura Kral, (W) LB
25540, 26129
Xyris torta Sm.,(W) DS; MM 4517.LB 18626a,25527
792 BRIT.ORG/SIDA 20(2)
Notes:
Cuscuta sp. The specimen had no flowers or fruit.
Gymnopogon brevifolius. This is the third record for this species in Texas.
Lobelia reverchonii. We prefer to keep this a good species.
Sarracenia alata. This species was introduced to the Hickory Creek Unit by
Geraldine Watson in the early 1970's. It occurs naturally in wetland pine sa-
vannas nearby.
Scleria baldwinii. This is the first pineywoods region record.
DISCUSSION
Our plant list numbers 401 taxa (385 native) for the Hickory Creek Unit. Previ-
ous plant lists (Watson 1982; Yu n.d.) numbered well over 500 taxa. Why the
discrepancy? Three possibilities suggest themselves: 1) our list is only about 75
percent complete, 2) the Unit was much richer 25 years ago when the first lists
were developed, and 3) previous lists are at least partly conjectural, being “pad-
ded” with taxa that could possibly be there but which have not actually been
found there.
These possibilities are not mutually exclusive and, at the present time, it is
impossible: to fully assess them. However, some comments are apropos.
Before we began collecting, the documented plant list for the Hickory Creek
Unit numbered about 190 taxa (Brown n.d.). We doubled the list in 17 field trips
in 2000 and 2001 If the total number of taxa at Hickory Creek numbered 530—
ie., only about one third had previously been documented—one would expect
that, at least at first, approximately two out of every three of our collections
would have been an addition. This was not the case, and we actively avoided
recollecting in many cases. Additionally, on the basis of the specimens collected
from the Hickory Creek Unit in the 1970's and 1980's and reported by Watson
(1982), there is no evidence that there then existed plant communities addi-
tional to those that occur there now that would account for the additional taxa.
The same taxa collected by Streng and Watson in the 1970's were recollected in
2000-2001, and Watson’s listed species do not signal other communities. Fi-
nally, aerial photographs, scientific literature, historical accounts, and remi-
niscences by “old timers” clearly do not place additional plant communities
there. With only four communities represented, one would expect on the basis
of typical community counts (75 to 125 taxa per community type with some
overlap among communities) only 400 taxa. Consequently, while we have ob-
viously not found every taxon that exists at the Hickory Creek Unit, we feel
that our list of 401 taxa is over 90 percent complete.
There is some evidence to support the second hypothesis. The Hickory
Creek Unit is severely fire suppressed and has lost much of its herbaceous layer
(Streng & Harcombe 1982; McClung 1988; MacRoberts & MacRoberts 2000).
Today there remain only small patches of the extensive pine savannas that were
MACROBERTS ET AL., FLORA OF PRESERVE 793
previously there. Some taxa undoubtedly could have been lost or at least be-
come very rare in the ensuing years. For example, we did not find several of the
conspicuous orchids that had been collected in the 1970's. Nonetheless, it is
unlikely that 125 taxa have been lost in that period.
The third hypothesis probably explains a good deal of the discrepancy. The
Parks and Cory (1936) Big Thicket plant list was partly conjectural Ccould pos-
sibly be there”) (Cozine 1993) and there is no reason to suppose that the Watson
(1982) list was not also partly conjectural. Many species on Watson’s (1982) list
do not occur in the area and were probably added either because of
misidentification (both in the field and of herbarium specimens) or because of
conjecture. Cases in point include Eriocaulon koernickianum Van Heurck &
Muell.-Arg. (undoubtedly misidentified Lachnocaulon anceps) and E.
septangulare With. (probably misidentified E. decangulare) (see Brown & Brown
1996).
Thus, while it is the case that our list isincomplete and that a few taxa may
have been lost, most taxa that have grown in the Hickory Creek Unit during the
past quarter century have been collected (probably 90+ percent). We believe
that the Hickory Creek Unit has about 400 to 425 native taxa, far fewer than
have been attributed to it.
ACKNOWLEDGMENTS
This study was supported in part by a Southwest Parks and Monuments Asso-
ciation Grant # 99-2: “Distribution of bogs and wetland pine savannas.” Roy
Zipp, U.S. Park Service, aided with the work. Brooke Wheeler, Paul Harcombe,
and Sandi Elsik provided information about several specimens at Rice Univer-
sity. Monique Reed provided information about a specimen at TAMU. Justin
Williams provided information about four specimens at SHST. Thanks to Jason
Singhurst and Walter Holmes for their comments on the manuscript.
REFERENCES
Ajivscl, G. 1979.Wild flowers of the Big Thicket, east Texas, and western Louisiana. Texas A.
& M. Press, College Station.
Brioces, E.L. and S.L. Orzeit. 1989. Longleaf pine communities of the West Gulf Coastal
Plain. Natural Areas J. 9:246-263.
Brown, L.E. and R.L. Brown. 1996. Flora of the Big Thicket National Preserve: review of her-
barium collections and development of a computerized specimen database. Abstract:
Big Thicket Science Conference. Beaumont, Texas. P. 16.
Brown. L.E. n.d. Vouchered list of Big Thicket National Preserve plant collections at Rice
and Lamar universities. Unpublished report. Big Thicket National Preserve, Beaumont,
Texas.
Cozine, J. 1993. Defining the Big Thicket: prelude to preservation. East Texas Hist. Assoc.
32:57-71.
794 BRIT.ORG/SIDA 20(2)
DesHorets, J.D. 1978. Soil survey for the Big Thicket National Preserve, Texas. U.S.D.A. Soil
Conservation Service, College Station, Texas.
Eisner, 1. 1973. The Big Thicket National Park. Science 179:525.
Gunter, PAY. 1971. The Big Thicket, a challenge for conservation. Jenkins, New York.
Harcomet, P.A. and PL. Marks. 1979. Forest vegetation of the Big Thicket National Preserve.
Unpublished report: U.S. Park Service, Santa Fe, New Mexico.
Harcomet, P.A., J.S. GLITZENSTEIN, R.G. Knox, S.L. Orzett, and E.L. Brinces. 1993. Vegetation of the
longleaf pine region of the West Gulf Coastal Plain. Proc. Annual Tall Timbers Fire Ecol.
Conf. 18:83-103.
Jones, S.D., J.K. Wipre, and PM. Montaomery. 1997. Vascular plants of Texas. Univ. Texas Press,
Austin.
Karesz, J.T. 1994.A synonymized checklist of the vascular flora of the United States, Canada,
and Greenland. Timber Press, Portland, Oregon.
KarTesz, J.T. and C.A. Meacham. 1999. Synthesis of North American flora. Version 1.0. North
Carolina Botanical Garden. Chapel Hill.
MacRoserrs, B.R.and M.H. MacRoserts. 1998. Floristics of wetland pine savannas in the Big
Thicket National Preserve, southeast Texas. Phytologia 85:40—50.
MacRoserts, M.H. and B.R. MacRoserts. 2000. Maintaining the natural integrity of pine sa-
vannas on the Big Thicket National Preserve, Texas. Unpublished report. Big Thicket
National Preserve, Beaumont, Texas.
Marks, PL. and PA. Harcomee. 1981. Forest vegetation of the Big Thicket, southeast Texas.
Ecol. Monogr. 51:287-305.
Matos, J.A.and D.C. Rupotp. 1985. The vegetation of the Roy E.Larsen Sandylands Sanctu-
ary in the Big Thicket of Texas. Castanea 50:228-249,
McCune, M.A. 1988. Effects of fire exclusion on longleaf pine savannas in the Big Thicket
National Preserve. Thesis, Stephen F. Austin State University, Nacogdoches, Texas.
McLeop, C.A. 1971. The Big Thicket forest of east Texas. Texas J. Sci. 23:221-233.
Nesom,G.L. and L.E. Brown. 1998. Annotated checklist of the vascular plants of Walker, Mont-
gomery, and San Jacinto counties, east Texas. Phytologia 84:107-153.
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any
Parks, H.B. and V.L. Cory. 1936. Biological survey of the east Texas Big Thicket area. Texas
Agricultural Experiment Station, College Station.
Peacock, H.H. 1994. Nature lover's guide to the Big Thicket. Texas A. & M. Press, College
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Peterson, C.D. and L.E. Brown. 1983.Vascular flora of the Little Thicket Nature Sanctuary, San
Jacinto County, Texas. Brunswick Press, Houston.
STRENG, D.R. 1979. Edaphic and pyric influences on two contiguous savannas in east Texas.
Thesis, Rice University, Houston.
STRENG, D.R. and P.A. Harcomee. 1982. Why don't east Texas savannas grow up to forest?
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MACROBERTS ET AL., FLORA OF PRESERVE 795
U.S. NATIONAL PARK Service. 1993. Species in parks: flora and fauna databases. http://
www.ice.ucdavis.edu/nps/
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report, Biology Department, Rice University.
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796 BRIT.ORG/SIDA 20(2)
BOOKS RECEIVED/NOTICES
Annotations and comments by Editor
Horticulture/Gardening
MAGNUs JOHNSON. 200L. The Genus Clematis. Edited and Translated by Bengt
Sundstrom. (ISBN 91-631-1030-X, hbk.). Magnus Johnson Plantskola AB and
Bengt Sundstrom, Branningestrandsvagen 63, 151 39 SODERTALJE, Swe-
den. (Orders: 08-550 979 77, +468 550 970 77). $110.00, 896 pp, 24 gouaches,
about 100 illustrations from classical literature and ink drawings by the
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cluding many new names and new combinations. The third part addresses Descriptions of sections
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Perer FE Yeo. 2002. Hardy Geraniums. Second Edition. (ISBN 0-88192-541-1, hbk.).
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3527, U.S.A. (Orders: www.tim ber press.com, 800-327-5680, 503-227-2878,
503-227-3070 fax). $34.95, 218 pp, 57 color photos, 151 line illustrations, 7
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Fr. JOHN L. Fic a. 1988 (paper back edition 2002). Lilacs: The Genus Syringa. (ISBN
0-88192-530-6, pbk.). Timber Press, Inc., 133 S.W. Second Avenue, Suite 450,
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SIDA 20(2): 796. 2002
VASCULAR PLANTS OF THE
MAPIMI BIOSPHERE RESERVE, MEXICO: A CHECKLIST
Abel Garcia-Arévalo
HHA Pie Eel A.C
Centro Regi ional Duane
Apdo. Postal 632
Durango, i 34000, MEXICO
ABSTRACT
The Mapimi Biosphere Reserve is located at the boundaries of the Mexican states of Chihuahua,
Coahuila, and Durango in the part central of the Chihuahuan Desert. It extends over 342,387 ha.
within an endorreic basin locally known as the Bolson de Mapimi. Elevation varies between 1000
and 1480 m. Weather is very arid mostly with summer rains, the average annual precipitation is 264
mm. Vegetation types are mostly xerophytic scrub and halophytic grassland.
he vascular flora is composed of approximately 71 families, 242 genera, +03 species and 426
taxa approximately. Families with the greatest diversity are Asteraceae, Poaceae, Cactaceae, and
Fabaceae. Genera with the greatest number of species are Sporobolus, Opuntia, Aristida, and Acacia.
I recorded 31 species endemic to the Chihuahuan Desert.
RESUMEN
La Reserva de la Biosfera de Mapimi, se ubica en el limite de union de los estados de Chihuahua,
Coahuila y Durango en la parte central del Desierto Chihuahuense, conforma una extension
aproximada de 342,387 ha y constituye una cuenca endorreica, denominada Bolsén de Mapimi. La
altitud varia entre los 1000 y 1480 m, suclimaes muy ar ido c con na ias en verano y la PEecpracion
es de e aproxime adamente 264 mm anuales. Lo tipos de g
y pase halon:
| Pasatalaccl : |
71 familias, 242 géneros, 403 especies y 426 taxa.
s familias con mayor diversidad son Asteraceae, Poaceae, Cactaceae y Fabaceae. Los géneros
Sporobolus Opuntia, Aristida y Acacia son los que presentan el mayor numero de especies. Se registran
31 especies endémicas al Desierto Chihuahuense.
INTRODUCTION
The Mapimi Biosphere Reserve is representative of the arid ecosystems of the
Chihuahuan Desert (Martinez & Morello 1977; Halffter 1981). Geographically
it is located at the boundaries of the Mexican states of Chihuahua, Coahuila,
and Durango (Bartolino 1988), between 26° 29' and 26° 52'N latitude, and 103°
32' and 103° 58' W longitude (Fig. L). It has an area of 342,387 ha and constitutes
an endorreic basin where fresh water sediments, basalts, and rhyolites have
accumulated until the Miocene. The Mapimi Biosphere Reserve is part of the
Bolson de Mapimi, and it includes the basins of Laguna de Patos, Laguna de
Palomas, and part of Laguna del Rey. It is part of the Mapimi subprovince.
The climate is very arid with most rain falling mostly between July and
SIDA 20(2): 797-807. 2002
COAHUILA
~
rr dR
10206
oss
Fig. 1 Mapimi
Biosphere
Reserve, Mexico.
GRAPHIC SCALE
oO 3 5
—— ———————
KILOMETROS
(7)0@ vals/o¥o'LINa
862
GARCIA f 799
September. The average annual rainfall is 264 mm. The highest record is 513
mm and lowest is 81 mm, anda mean annual temperature of 20.8°. Elevation is
about 1000 m at the lowest zones or “playas” and 1480 m at the summit of the
Cerro San Ignacio, the highest point.
VEGETATION
Martinez and Morello (1977) described nine types of physionomic-floristic units
for this region of the Mapimi. Later, Gonzalez (1983) noted, on her description
of the vegetation of Durango, desert xerophytic scrub, grasslands and halo-
phytic vegetation as representative types for the region.
On the basis of the biological forms and dominant species, Montana (1988)
grouped vegetation into 76 associations. These associations were grouped into
14 categories of higher rank defined by combinations of dominant life forms.
Out of the 35 species, 12 were more widely distributed as dominants: Prosopis
glandulosa Torr, Larrea tridentata (DC) Cov., Hilaria mutica (Buckley) Benth.,
Fouqueria splendens Engelm., Acacia constricta Benth., Flourensia cernua DC,
Cordia parvifolia A. DC, Atriplex acanthocarpa (Torr) S. Watson, Acacia greggii
A. Gray, Opuntia rastrera FA.C. Weber, Dalea scoparia A. Gray y Yucca elata
Engelm.
In general, there are two dominant vegetation types, xerophytic scrub
(Chihuahuan desert scrub) and halophytic grasslands, with a great diversity
of associations.
Xerophytic scrub (Chihuahuan desert scrub).—This vegetation type is
widely distributed in the reserve and is characterized by the dominance of
shrubby species with a wide variety of plant associations. Species most com-
monly represented are: Larrea tridentata, Prosopis glandulosa, Opuntia rastrera,
Acacia constricta and Fouqueria splendens, among others.
Halophytic grasslands.—Grasses are dominant in this vegetation type, and
are found, mostly at places with high salt concentrations. Principle dominant
species in the associations are Sporobolus airoides and Pleuraphis mutica.
FLORA
For the vascular flora of the Mapimi Biosphere Reserve 71 families, 242 genera,
403 species and 426 infraspecific taxa are registered. Families and genera with
the greatest diversity are shown in Table L. Endemic species to the Chihuahuan
Desert are shown in Table 2. Previously, Ruiz de Esparza (1988) recorded 60 fami-
lies, 197 genera and 313 species.
There are isolated collections from the Bolson de Mapimi basin. In the 1980s
these collections increased as it was the interest of various studies from Insti-
tuto de Ecologia A.C. (Carlos Montana, Antoine Cornet and Rosario Ruiz de
Esparza). Recently, from 1994 on, Garcia-Arévalo collected most of the speci-
mens on which this catalogue is based.
800
Taste 1. Most diverse families and genera.
BRIT.ORG/SIDA 20(2)
Families Species Genera Species
Asteraceae 68 Sporobolus 10
Poaceae 62 Opuntia 8
Cactaceae 29 Aristida 7
Fabaceae 21 Acacia 7
Taste 2.Endemic species to the Chihuahuan Desert.
AMARANTHACEAE
ie oer eeratt (Torr) Stand.
ASTERACEAE
Nicolletia edwa sli A. Gray
Pectis incisifolia |.M.Johnst.
Pectis pringlei Fernald
Psathyrotes 5c ApOsa A. Gr ay
ileus edie Rydb.
A GPCOTIUTT.
Visuieaphien nax S.F. Blake
BORAGINACEAE
Tiquilia ne (Wooton & Standl.) A.T. Rich
ards¢
—
Tiquilia greggii
CACTACEAE
Ariocarpus fissuratus (Engelm.) K.Schum.
Coryphantha macromeris (Engelm.) Lem.
Echinocereus enneacanthus Engelm.. var.
Torr. & A. Gray) A.T. Richardson
enneacanthus
Echinomastus ungispinus (Engelm.) Britton &
ose
Mammillaria pottsii Scheer ex Salm-Dyck
Opuntia bradtiana (J.M.Coult.)
Salm-Dyc
K. Brandegee ex
CARYOPHYLLACEAE
Drymaria axillaris Brandegee
CONVOLVULACEAE
Ronamiq
DOLAI/TI
multicaulis (Brandegee) House
FABACEAE
Senna pilosior (Robinson ex J.F.Macbr.) H.S. Irwin
& Barneby
FOUQUERIACEAE
Fouqueria shrevei |.M. Johnst.
HYDROPHYLLACEAE
Nama stenop
Nam
ryllum A. Gray ex Hemstl.
torynophyllum weenmM.
LAMIACEAE
Salvia purpusii Brandegee
ACEAE
onix crenatus A. Gray ex S.Watson
a WN
Peta
eee
is pl IKOLISIQNYIS Heimer
POACEAE
Sp orobolus coahuilensis Valdés- Reyna
Sporobolus spiciformis Swallen
POLYGONACEAE
FEriogonum hemipterum
orr. & A Gray) S.G.
Stokes
RUBIACEAE
Randia pringlei (S.Watson) A. Gray
ZYGOPHYLLACEAE
Fagonia scoparia Brandegee
Kallstroemia perennans B.L. Turner
To complement the catalogue, | reviewed the plant collections hosted at
CHDIR and ANSM, and also an unregistered herbarium hosted at Instituto de
Ecologia A.C., Centro Regional Durango. Plant specimens will be deposited at
these collections.
GARCIA EF
CATALOGUE OF KNOWN VASCULAR PLANTS OF THE
MAPIMI BIOSPHERE RESERVE (*NON-NATIVE SPECIES)
ACANTHACEAE
Justicia pg Nee) R.A. Hilse
a) Hi. i-f, (A
AGAVACEAE
Agave lechuguilla Torr.
Agave scabra Salm-Dyck ssp. scabra
Manfreda variegata (Jacobi) Rose
ZOACEAE
Glinus lotoides L.
Sesuvium sessile Pers.
AMARANTHACEAE
Amaranthus arenicola |.M. Johnst.
Amaranthus palmeri S.Watson
Gomphrena decumbens Jacq.
Guilleminea lanuginosa (Poir.) Hook.f.
cee ada (Hook.f.) Hen. , S.D. Sundb.
—
Stand.
Nee
iz
ot
romia ge LM.John
eee suffruticosa (Torr.
AMARYLLIDACEAE
ephy ranthes lonajfoli a Hemsl.
ANACARDIACEAE
Rhus microphylla Engelm.
APOCYNACEAE
Telosiphonia macrosiphon (Torr.) Henr.
ARISTOLOCHIACEAE
Al istolochia WI ightii Seem.
ASCLEPIADACEAE
ASC sede rahe Engelm. ex Torr.
pias linaria
epia ee, es Cham. & Schltdl.
epias svirifora Rak
ie
Sarcostema es Decne. subsp.
hartwegil (Vail) R.W. Holm
ASTERACEAE= COMPOSITAE
Acourtia pay (A. Gray) Reveal & R.M. King
Acourtia ae (A. nee Reveal & R.M. King
- reel
Amorosia psi leche i
Artemisia filifolia Torr.
Aster spinosus Benth.
Aster subulatus Michx.
Baccharis salicifolia (Ruiz & Pavon.) Pers.
Bahia absinthifolia Benth.
—
nb.
Gray) Tharp & FA. Barkley
Bahia biternata A. Gray
Baileya multiradiata Harv. & A. Gra
Barroetea subuligera (S. Schauer) A. Gray
Brickellia laciniata A. Gray
Rric-lplli It iA Gray
Conyza coulteri A. Gra
Dicranocarpus parviflorus A. Gray
Eupatorium solidagnifolium A. Gray
Flaveri a palmer! J. . Johnston
Flaveria trinervia (S prer \g.) C. Mohr
Fi lorestina tripteris DC.
cl DC.
Gaillardia pinnatifida Torr.
Gnaphalium falcatum Lam.
Grindelia oxylepis Greene
Gymnosperma glutinosum (Spreng.) Less.
Helenium autumnale L
Helianthus ciliaris DC.
Helianthus petiolaris Nutt.
Iva ambrosiifolia (A. Gray) A. Gray
lva dealbata A.Gra
Jefea brevifolia (A. Gray) Strother
doaieaalaass australis (Greene) Shinners
era brevilingulata (Hemsl.)
Turner
Machaeranthera linearis Greene
Machaeranthera pinnatifida (Hook.) Shinners
Nicolletia edwardsii
Palafoxia sphacelata Ate ex Torr.) Cory
P th ; fort A Gray
thenitim (in canum H.B.K.
Pecti 5 engus” ifolia Torr.
lindrica (F € rnald) ) Rydb.
ae incisifolia \.M.Johnst.
Pectis papposa Harv. & A. Gray
Pectis pringlei Fernald
Pectis tenella
Porophyllum me um A. Gray
Psathyrotes 5c Gra
suas ee A: ey
DC
>
eee ia puberula Ry.
Sconuhie oe ) et
*Sonchus oleraceus L.
Thymophylla aurea Greene var. polychaeta (A.
Gray) J.L. Strother
hymophylla pentachaeta Small
Trixis californica Kellogg var. californica
Verbesina encelioides tees Benth & Hooker
ae, EON Greenm
ee
Viguier dentata (Cav.) Spreng.
Viguiera multiflora (Nutt.) S.F. Blake
Viguiera phenax S.F. Blake
Viguiera stenoloba S.F. Blake
Xanthium strumarium L.
Xylothamia triantha (S. F. Blake) G.L.Nesom
Zinnia acerosa A. Gray
BIGNONIACEAE
Chilopsis linearis (Cav.) Sweet subsp. linearis
Tecoma stans (L.) H.B.K. var. angustata Rehd.
BORAGINACEAE
Antiphytum heliotropioides D
Tiquilia es (Wooton : Standl) A.T. Rich-
ardso
—
Tiquilia greggii (Torr. & A. Gray) A.T. Richardson
Tiquilia hispidissima (Torr. & A.Gray) AT. Richardson
-ordia parvifolia DC
Cryptantha pusilla (Torr. & A. Gray) Greene
eu convolvulaceum (Nutt.) A. Gray
L. var. curassavicum
helowapiin a usculum (Torr) A.Gray
Heliotropium greg
He liotropium molle
=
ae ILM. Johnst.
BROMELIACEAE
Hechtia glomerata Zucc.
ieprinimaninedes:
HOH Benth.
pee scordioides H.B.K.
CACTACEAE
Ancistrocactus uncinatus (Galeotti) L.D. Benson
pial aaer| Suna, ac aapes K.Schum.
DC.
Con olantne macromeris on ) Lem,
ae scheeri (Muhlenpf.) L.D.Benson var,
robustispina (Schott ex Engelm.) L.D.Benson
ea horizonthalonius Lemaire
Echinocactus texensis Hopffer ex Regel
Echinocereus enneacanthus Engelm.. var.
Be
ereus pectinatus (Scheidw.) Engelm. var.
reais (Engelm.) Engelm. ex Rumpler
BRIT.ORG/SIDA 20(2)
Echinomastus durangensis (Ruenge) Britton &
Rose
Echinomastus ungispinus (Engelm.) Britton &
Rose
Epithelantha micromeris ExXEnge Ina) F.A.C.Weber
Hamatocactus hamatacanthus (Muhlenpf.) F.M.
aun
rophora williamsii (Lem.) J.M.Coult.
anal laria heyderi ee var.
gummifera (Engelm.) L.D. Ben
var aie Secs Muehen porate var. heyderi
PCP EA Et igelm.
Momenflana pott ssjischee
Opuntia bradtiana (J.M. eit ex Salm Dyck) K
Brandegee
Opuntia imbricata (Haw.) DC.
Opuntia leptocaulis DC. var. brevispina (Engelm.)
S.Watson
Opuntia leptocaulis DC. var. robustior A. Berger
pu untia macrocentra Eng
pur ea Pfeiff.
Opunti ia rastrera F.A.C.Weber
Opuntia schottii Engelm.
Peniocereus greggil (Engelm.) Britton & Rose var.
—
ae
greggii
Thelocactus bicolor (Galeotti ex Pfeiff.) Britton &
Rose
CAPPARIDACEAE
Polanisia dodecandra (L.) DC.
CARYOPHYLLACEAE
Drymaria axillaris Brandegee
Drymaria molluginea (Ser.ex DC.) Didr.
Drymaria pachyphylla Wooton & Stand.
COCHLOSPERMACEAE
Amoreuxia wrightil A. Gray
COMMELINACEAE
Commelina erecta L.
Commelina erecta L. var. angustifolia (Michx.)
Fernald
CONVOLVULACEAE
Bonamia multicaulis (Brandegee) House
Cressa truxillensis H.B.K. var. vallicola (A. Heller)
Munz
Cuscuta umbellata Kunth
Ipomoea costellata Torr.
Ipomoea cristulata Hallier f.
[pomoea purpurea (L.) Roth
GARCIA
803
CRUCIFERAE
Descurainia pinnata (Walter) Britt.
Dimorphocarpa wislizenii (Engelm.) Rollins
“Eruca versicaria (L.) Cav. subsp. sativa (Miller)
Thell
Nerisyrenia camporum (A. Gray) Greene
Nerisyrenia linearifolia (S.Watson) Greene
*Sisymbrium
Stanleya pinnata Gane Britton
CUCURBITACEAE
Apodanthera undulata A. Gra
lbervillea tenuisecta (A. Gray ) Small
CYPERACEAE
Cyperus aff. odoratus L.
Eleocharis caribaea (Rottb.) S.F. Blake
Eleocharis macrostachya Britton
Eleocharis parvula(R & Schult.) Link ex Bluff,
Nees & Schauer
n maritimes |
CHENOPODIACEAE
Allenrolfea occidentalis (S. Watson) O. Kuntze
Atriplex acanthocarpa (Torr.) S.Watson
Atriplex canescens (Pursh) Nutt.
Atriplex obovata Moq
Chenopodium glaucum L.
*Salsola tragus
Suaeda nigrescens ILM. Johnst.
Suaeda suffrutescens S.Watson
EPHEDRACEAE
Ephedra trifurca Torr.ex S.Watson
EUFHORBIACERE
av.
ae nati i (Klotch) Muell-A rg.
Ditaxis neomexicana (Muell.-Arg.) A. Heller
Euphorbia oe ZUCC,
Euphorbia den ichx,
Euphorbia ee ae
Euphorbia strictior Holz.
Jatropha di
Tetraccocus a eee (S.Watson) Croizat
Tragia nepetifolia Cav.
FABACEAE=LEGUMINOSAE
Acacia angustissima (Mill.) O. Kuntze
Acacia berlandieri Benth.
Acacia constricta Benth.
Acacia farnesiana (L.) Willd
Acacia greggii A.Gra
Acacia neovernicosa |sely
Acacia roemeriana Scheel
alea aurea Nutt. ex Fraser
alea lanata Spr
alea leporina Hee Bullock
alea neomexicana (A. Gray) Cory
lea hes ot A. a
alea scoparia A
modium neomexicanum A. Gra
Hoftmansegia glauca (Ort.) Eifert
*Parkinsonia aculeata L
Phaseolus polymorphus S.Watso
Prosopis glandulosa Torr. var. ee (L.D.Ben-
nst.
Senna bani ides (A.Gray) H.S.Irwin & Barneby
Senna covesii (A.Gray) H.S. Irwin & Barneby
Senna pilosior (Robinson & Macbride) H.S. Irwin
& Barneby
ale
peeetes
FOUQUERIACEAE
Fouqueria shrevei |.M. Johnst.
Fouqueria splendens Engelm.
GENTIANACEAE
Centaurium calycosum (Buckley) Fernald
DEORE CEne
n (Torr.) Greenm.
Af
Nama stenocarpum A. Gray
Nama ea A. Gray ex Hemsl.
c
OP hy lum MOTECriTt.
Noma sono oe & Standl.) CL. Hitche.
ok.
KOEBERLINIACEAE
Koeberlinia spinosa Zucc.
KRAMERIACEAE
Krameria glandulosa Rose & Painter
Krameria erecta Willd ex Schult.
LAMIACEAE=LABIATAE
Salazaria mexicana Torr.
Salvia purpusii Brandegee
age ee Hornem.
LILIACEAE
Dasylirion wheeleri S.Watson ex Rothr.
Milla biflora Cav.
nace elaia Engelm.
(Engelm.,) Trel.
ee ‘one Shafer
LOASACEAE
Cevallia sinuata Lag.
Eucnide bartonioides Zucc.
Mentzelia albicaulis (Hook.) Torr. & A. Gray
Mentzelia pumila (Nutt.) Torr. & A. Gray
Petalonix crenatus A. Gray ex S. Watson
ORANTHACEAE
Phoradendron tomentosum (DC.) Engelm. ex A.
Gray subsp. tomentosum
LYTHRACEAE
Ammannia coccinea Rottb.
MALPIGHIACEAE
Janusia gracilis A. Gray
MALVACEAE
Abutilon malacum S.Watson
anon pen rea sta A. Gray
ri Harv. ex A. Gray
Hihicctis dentidatts 5 Benth.
Malva parviflora L.
Malvastrum coromandelianum (L.) Garcke
Malvella lepidota (A. Gray) Fryxell
Malvella leprosa (Ortega) nee
see sagitiifolia (A. Gray) Fryxell
a abutifolia Miller
ralcea angustifolia (Cav.) G.Don
cate cea hastulata A. Gray
MARTYNIACEA
Proboscidea althaefolia Decne.
Proboscidea fragans (Lindl.) Decne.
MELIACEAE
“Melia azedarach L.
pililtnadadaite
d A. Gray
Alin’ iac choisy. stand
An, /, [i i |
Star
Boerhavia gracillima Heimer
ae ue M. E. Jones
Cho
Boerhavia wana A. ae a
Mirabilis rotata (Standl) IM. Johnst.
ee viSCOSa 1 Cav.
pitata Choisy
Se elocaious angustifolius Torr.
Selinocarpus purpusianus Heimer!
Forestiera angustifolia Torr.
Menodora scabra A. Gray
BRIT.ORG/SIDA 20(2)
ONAGRACEAE
Clarkia purpurea (Curt.) A. Nelson & LF. Macbr.
tbsp. quadrivulnera (Douglas) F.H. Lewis &
Wis
Oenothera aff. texensis PH. Raven & D.R. Parnell
Oenothera speciosa Nutt.
a)
ROBANCHACEAE
Orobanche cooperi (A. Gray) Heller
OXALIDACEAE
Oxalis aff. decaphy
Oxalis L.
=.
la Kunth
PALMAE
“Washingtonia filifera (Linden ex André) H
Wendl.
PAPAVARACEAE
Argemone ochroleuca Sweet
POACEAE= GRAMINEAE
; idscencionis L
Aristida flor)
Aristida pdansda Wooton & Standh
Aristida purpurea Nutt. var. nealleyi (Vasey) All
Aristida roemeriqna Scheele
Aristida ternipes Cav.
Aristida wrightii Nash
Bouteloua aristidoides (Kunth) Griseb.
Ray, teloi;q harhnvte |
Lad.
Bouteloua cri (Michx.) Torr.
ee ramosa Yea
' Scribn. & Merr.
Bouteloua uni ou Vas sey
Brachiaria arizonica (Scribner & Merrill) S.T. Blake
lata (Sw.) Parodi
Bra Ai aria fi
bId 1d
Chloris crinita a
Chloris virgata S
Cottea seen ne Kunth
*Cynodon dactylon (L.) Pers.
lees cali es Ske Henrard
n(L.) Link
Echinochloe Beauv.) Fernald
Gea desvauii P. Beauv.
Fragrostis barrelieri Daveau
a is cilianensis ait ) Vignolo ex Janch.
1} (Hornem.) Link
Er, agrostis pi inca (L.) eae
Friochloa inata (I. Presl) Kunth
Eriochloa contracta Hitche.
tis mexicar
GARCIA E 805
Erioneuron pulchellum (Kunth) Tateoka Talinum aurantiacum Engelm
Heteropogon contortus (L.) P. Beauv.ex Roem.& — Talinum paniculatum (Jacq.) Gaertn.
Ser POTAMOGETONACEAE
| pt hl dub (H.B.K.) Nees Pot nodosus Poir.
Leptochloa fascicularis (Lam.) Beauv.
uhlenbergia arenicola Buckley PRIMULACEAE
Muhlenbergia fragilis Swallen Samolus ebracteatus Kunth
Muhlenbergia porteri Scribn. ex Beal PTERIDACEAE
Panicum flexile (Gatt.) Scribn. Astrolepis cochisensis (Goodd.) Benham &
Pani icum hallii Vasey Windham
Panicum hirticaule J. Pres| Astrolepis sinuata (Sw.) Benham & Windham
Pani bt ae Notholaena standleyi Maxon
trami tchc. & Chase N
Pappophorur micronltur Ness BANE HOSEN EERE 7
e(L) Clematis drummondii Torr.& A. Gray
Basen australis (Cav.) Trin. ex Steud. RESEDACEAE
Pleuraphis mutica Buckley Oligomeris linifolia (Vahl) J.F. Macbr.
een nogen brevifolius Phil. RHAMNACEAE
(Forssk.) Chiov. oa warnockii M.C. Johnst.
Setaria grisebachii E.Fourn. iphus obtusifolia (Hook. ex Tort. & A. Gray.) A
ae scheelei (Steud.) Hitchc.
en
por MBO aNOIds (Torr.) Torr.
is Valdés-Reyna RUBIACEAE
ea ibaliis contractus Hitche. Randia pringlei (S.Watson) A. Gray
pp onoms coromandelianus (Retz.) Kunth RUTACEAE
lic cryntanari se
BP
Thamnosma texana (A. Gray) Torr.
SALICACEAE
Salix exigua Nutt.
r.) A. Gray
Sorel: flexuosus (Thurb. ex Vasey) Rydb.
Sporobolus patens Swallen
Sporobolus poiretii (Roem. & Schult.) Hitche.
Sporobolus spiciformis Swallen SCROPHULARIACEAE
jeden Us da ius R.M. Harper Casti hele integra A. Gray
is Schult. eucophyllum laevigatum Stand.
fidens albescens mee Wooton. & Stand. eee He minus A. Gray
Tridens muticus (Torr.) Nash Maurandya antirrhiniflora Humb. & Bonpl. ex
Willd
POLEMONIACEAE
pomopsis laxiflora (|.M.Coult.) V.E. Grant
[po mopsis lonaiflora (Torr.) V. E.Grant
—
Mecardonia procumbens (Miller) Small
Stemodia schottii Holz
lpomopsis polycladon (Torr.) V.E.Grant ee
POLYGALACEAE ella lep hylla (Hook. & Grev.) Spring
Polygala ob Benth. SIMAROUBACEAE
pie sent tl
POLYGONACEA Castela texana (Torr.& A. Gray) Rose
ee het (Torr.& A.Gray) S.Stokes eee
alibrachoa parviflora Juss.) D’Arcy
Pol Neonun vee L. Chamoesraco eomcide: (Dunal) Britton
Rumex verticillatus L. a Ortega
Daon inoxia Mill.
PORTULACACEAE |
Datura quercifolia Kunth
inum berlandier| M. Dunal
Se poet Ly
ra fineee IS eee A Gray Wiese Pica Graban
806 BRIT.ORG/SIDA 20(2)
Nicotiana obtusifolia Mart. & Gal Citha thum (A. Gray) A. Gray
Lee cinerascens (Dunal) Hitche. Giinduiata bininnad fida (Nutt.) Nutt.
alis hederifolia A. Gray Lantana achyranthifolia
Physalis aff. microphysa A. Gray Lippia pee Kunth
eenae lobata (Torr.) Raf. Phyla incisa Small
ifoli A.Braun Phyla eal (L.) Greene
Sol leagnifoli av. Phyla strigulosa (M. Martens & Galeotti)
Sola tratum M. Dunal Moldenke
Solanum triquetrum Cav. Verbena canescens Kunth
TAMARICACEAE Verbena gracilis Desf.
“Tamarix ramosissima Ledeb. VITACEAE
ULMACEAE Cissus trifoliata L.
Celtis pallida Torr. ZYGOPHYLLACEAE
Fagonia scoparia Brandegee
Kallstroemia grandiflora Torr. ex A. Gray
ee hirsutissima ie il
L. Turner
Larrea tridentata (Sessé g Moc. ex DC.) Coville
Peganum aff. mexicanum A. Gray
“Tribulus terrestris L.
VERBENACEAE
Aloysia gratissima (Gillies & Hook.) Tronc.
Aloysia gratissima (Gillies & Hook.) Trone. var.
schulzae (Standl.) Mold.
Aloysia wrightii (A. Gray) A. Heller ex Abrams
Bouchea prismatica (L.) K.E. O. Kuntze
ACKNOWLEDGMENTS
To the Comision Nacional para el Conocimiento y Uso de la Biodiversidad
(CONABIO ) for their support (Proyect L 035 ), to Juan Pablo Ramirez, Adalberto
Herrera and Francisco Herrera for their invaluable help with field work, to José
Villarreal, Socorro Gonzalez, Richard Spellenberg, Yolanda Herrera, Jests Valdés,
Miguel Carranza and José Panero for their help in the identification of some
plant specimens, and to Jorge Nocedal for his help in the translation of this
manuscript.
ol
REFERENCES
BarTOUNo, J.R. 1988.Cenozoic geology of the eastern half of the La Flor quadrangle, Durango
and Chihuahua, México. |n:C. Montana, ed. Estudio integrado de los recursos vegetacion,
suelo y agua en la reserva de la biosfera de Mapimi. | Ambiente natural y humano.
Publ. 23, Instituto de Ecologia México. Pp. 77-97.
GONZALEZ ,S.1983.La vegetacién de Durango. Cuad.|Inv.Tecnol. CIIDIR-IPN Unidad Durango
1(1):1-114
Hacrrter, G. 1981. The Mapimi Biosphere Reserve: local participation in conservation and
development. Ambio. 10 (2—3):93-96.
Martinez, E.and J. Moretto. 1977. El medio fisico y las unidades fisondémico-floristicas del
Bolson de Mapimi. Instituto de Ecologia, México, DF.
Montana, C. 1988. Las formaciones vegetales. In: C. Montana, ed. Estudio integrado de los
recursos vegetacion, suelo y agua en la reserva de la biosfera de Mapimi.| Ambiente
natural y humano. Publ. 23, Instituto de Ecologia , México. Pp. 167-197.
GARCIA fy, JASCII 807
Ruiz be Esparza, R. 1988. Lista de especies vasculares. In:C. Montana, ed. Estudio integrado
de los recursos vegetacion, suelo y agua en la reserva de la biosfera de Mapim. |
Ambiente natural y humano. Publ. 23, Instituto de Ecologia, México. Pp. 225-239.
808 BRIT.ORG/SIDA 20(2)
BOOKS RECEIVED/NOTICES
Annotations and comments by Editor
Horticulture/Gardening
HAROLD Koopowttz. 2002. Clivias. ISBN 0-88192-546-2, hbk.). Timber Press, Inc.,
133 S.W. Second Avenue, Suite 450, Portland, OR 97204-3527, U.S.A. (Or-
ders: www.timber press.com, 800-327-5680, 503-227-2878, 503-227-3070
fax). $34.95, 384 pp, 118 color illustrations, 3 charts, | bn/w photo, 2 keys,
| map, 6" x 9"
ay
Clivia is a small genus (four species) in Amaryllidaceae family from southern Africa. “They are all
evergreen herbs with leaves arranged in two rank borne on a thick rhizome.” The name honors Lady
Charlotte Florentia Clive. “Clivias is the first practical book on this fascinating genus. The result is a
comprehensive treatment that covers the history and biology of clivias, and provides profuse infor-
mation on cultivation, color, and breeding.” There are 118 great color photographs. Contents: Intro-
duction, What is ina Name? Clivia and Cryptostephanus: The Species; The Biology of a Clivia Plant;
Cultivating and Growing Clivias, Understanding Clivia Colors, Hybridizing and Growing Clivias
from Seed; Clivia Obsession; Standard Clivia miniata Hybrids: Orange and Red; Standard Clivia
Lady Clive?, Appendix II. Sources of Information; Appendix III. Sources of Plants; Glossary; Map of
South Africa; References and Additional Reading; Index of Plant Names.
Lewis J. MATTHEWS. 2002. The Protea Book: A Guide to Cultivated Proteaceae.
(ISBN 0-88192-553-5, pbk.). Timber Press, Inc., 133 S.W. Second Avenue, Suite
450, Portland, OR 97204-3527, U.S.A. (Orders: www.timberpress.com, 800-
327-5680, 503-227-2878, 503-227-3070 fax). $34.95, 184 pp, 286 color photos,
Cit x10 1/4,
“Proteaceae is a large and ancient family of incredible diversity in both foliage and flower form. In
recognition of this variety, Linnaeus named them after the legendary Greek sea god Proteus, who
could change his shape at will.” This is a must book for anyone interested in Proteas. Whether you are
a grower or the cut-flower industry you will like this book. Contents: Introduction; Cultivating pro-
a
teas; Proteas in the garden; Commercial cultivation; Identification; Glossary; References; Index.
Micuae A. Dire. 2002. Dirr’s Trees and Shrubs for Warm Climates: An Illustrated
Encyclopedia. (ISBN 0-88192-525-X, hbk.). Timber Press, Inc., 133 S.W. Second
Avenue, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders:
www.timberpress.com, 800-327-5680, 503-227-2878, 503-227-3070 fax).
$69.95, +48 pp, 1438 color photos, | map, 8 1/2" x 1"
This isa com panion volume to Dirr’s Hardy Trees anc
Qu
Shrubs: An Ilustrated Encyclopedia also pub-
lished by Timber Press. This is truly an encyclopedia of beauty and detail for more than 400 species
and nearly 500 additional cultivars and varieties, many of which are illustrated. Even if you are not
into woody plants this book will surely please you. The 1438 color photos are worth the price alone.
SIDA 20(2): 808. 2002
A SIGNIFICANT NEW POPULATION OF THE RARE
SEMAPHORE PRICKLYPEAR CACTUS, OPUNTIA
CORALLICOLA (CACTACEAE)
Keith A. Bradley and Steven W.Woodmansee
The Institute for Regional Conservation
22601 S.W. 152 Ave.
Miami, FL 33170, U.S.A.
bradley@regionalconservation.org
ABSTRACT
Anewly discovered population of Opuntia corallicola is reported. This new population is in Miami-Dade
County, Florida, 145 km north of what was believed to be the last remaining population in the wild.
RESUMEN
Se inf | bl ToC) : eal acalac LI
-e sg ae
esta en el Condado Miami-Dade, Florida, 145 km al norte de lo que se pensaba que era la ultima
poblacion que crecia silvestre en la naturaleza.
The semaphore pricklypear cactus, Opuntia corallicola (Small) Werderm.
(Cactaceae) is considered one of the most threatened plants in the United States
(Stiling et al. 2000). It is endemic to the Florida Keys, first discovered by John
Kunkel Small in 1919 (Small 1930). Previously, Florida plants were thought to
be conspecific with O. spinosissima P. Miller of Jamaica, but research has shown
Florida plants to be distinct (Austin et al. 1998; Gordon & Kubisiak 1998). The
Florida Keys are a chain of islands at the southern tip of Florida, extending in
an arc approximately 320 km from the Dry Tortugas in Monroe County north
to Soldier Key in Miami-Dade County.
Small’s discovery of O. corallicola was made on Big Pine Key, an island in
the lower Florida Keys. Plants were known [rom this island until the 1960s when
the last plants were eliminated by poaching and road construction. Sometime
during the 1960s a new population was discovered on Little Torch Key, an is-
land immediately west of Big Pine Key. Subsequently, the station was acquired
by The Nature Conservancy. Small (1930) also reported the species from Key
Largo, an island in the upper Florida Keys just over 80 km north of Big Pine
Key, although we have been unable to locate specimens or other reports from
this island.
The population on Little Torch Key presently has only nine adult plants
(C. Bergh, pers. comm.). Plants at this site are threatened, originally by poach-
ing, and subsequently by both an exotic moth and its inability to sexually re-
produce. The exotic moth, Cactoblastis cactorum Berg (Lepidoptera: Pyralidae),
SIDA 20(2): 809-811. 2002
810 BRIT.ORG/SIDA 20(2)
was discovered in the Florida Keys in 1989 (Habeck & Bennett 1990). This pest
infested two of the remaining wild plants, killing one of them (Stiling @ Moon
2001). While vegetative reproduction results in numerous recruits (C. Bergh,
pers.comm.), Negron-Ortiz (1998) found that fruit set in the species is rare. Less
than half of the seeds set were viable in laboratory culture, but no seedling re-
cruitment was found under natural conditions. The few seeds produced are
believed to result from agamospermy, asexual seed formation common in
Cactaceae (Negron-Ortiz 1998). Negron-Ortiz (1998) proposed that the species
is unable to reproduce sexually either because of meiotic problems resulting
from polyploidy, or because the existing plants are self-incompatible.
The authors and George D. Gann have been conducting extensive field work
in Biscayne National Park (BNP) in part to update the floristic inventory pub-
lished by Stalter et al. (1999). The park is comprised of 42 islands of the upper
Florida Keys in Miami-Dade County. On November 20, 2001, the authors dis-
covered O. corallicola on Swan Key. This island is within the southern bound-
ary of BNP and is approximately 140 km from Big Pine Key, the closest histori-
cally documented occurrence, and 145 km from Little Torch Key, the site of the
other known population. On January 31, 2002 a subsequent survey was con-
ducted and 570 plants with trunks were found. The population consisted of
plants of diverse sizes and ages, including flowering individuals.
The new Swan Key population occupied approximately 0.5 km by 10 m
along the edge of a rockland hammock. Plants were in partial sun to shade grow-
ing on Key Largo Limestone, sometimes in light leaf litter and duff. While plants
were found primarily along the edges of the hammock, some small plants were
found inside the shady interior of the hammock up to 50 m from the edge. Asso-
ciated plant species include Acanthocereus tetragonus, Agave decipiens, Borrichia
arborescens, Bursera simaruba, Canella winterana, Capparis flexuosa, Coccoloba
diversifolia, Conocarpus erectus, Eugenia foetida, Guapira discolor, Metopium
toxiferum, Pithecellobium keyense, and Sideroxylon celastrinum.
We believe that O. corallicola merits Federal Endangered Species status.
Even with the discovery of this new population, only two populations of this
species are known. Both populations are threatened by Cactoblastis cactorum.
Opuntia corallicola is being considered by the U.S. Fish and Wildlife Service as
a candidate for listing (Federal Register 1999; Bradley & Gann 1999). Endan-
gered Species status would enable the National Park Service to focus on the
management and protection of the species. We encourage the US. Fish and
Wildlife Service to continue with the listing process and we support the estab-
lishment of an agreement with the National Park Service to focus on the man-
agement and protection of the species.
=
d
Rhodes Channel and Broad Creek, S of Old Rhodes Key, just N of the Monroe Co. line, common along
an 2002, Bradley 2152 (FTG).
Voucher specimen: FLORIDA. Miami-Dade Co.: Biscayne National Park; Swan Key, between O
the edge of rockland hammock, 31.
i
ACKNOWLEDGMENTS
The BNP inventory was funded through the National Park Service (NPS) In-
ventory and Monitoring Program, as part of the South Florida / Caribbean Net-
work of Parks Inventory of Vertebrates and Vascular Plants. We would like to
thank NPS ecologist Matt Patterson and NPS biologist Toby Obenauer. Fairchild
Tropical Garden (FTG) provided funding for the follow up survey which was
conducted with FTG biologists Meghan Fellows and Jennifer Possley. Robert
Pemberton, entomologist for U.S. Department of Agriculture-Agricultural
Research Service, also assisted with the follow-up survey. George Gann, Doria
Gordon, Dawn Jennings, Colette Jacono, and an anonymous reviewer provided
helpful comments on the manuscript.
REFERENCES
Austin, D.F., D.M. Binnincer, and DJ. Pinkava. 1998. Uniqueness of the endangered Florida
semaphore cactus (Opuntia corallicola). Sida 18:527-534.
Brantey, K.A.and G.D. GANN. 1999. Status summaries of 12 rockland plant taxa in southern
Florida. Report submitted to the U.S. Fish and Wildlife Service, Vero Beach, Florida. Mi-
ami: The Institute for Regional Conservation.
FEDERAL REGISTER. 1999. Candidate notice of review. Vol.69(205):57533-57547.
Gorodon, D.R. and T.L. Kusisiak. 1998. RAPD analysis of the last population of a likely Florida
Keys endemic cactus. Florida Sci.61:203-210.
Haseck, D.H. and F.D. Bennett. 1990. Cactoblastis cactorum Berg (Lepidoptera: Pyralidae), a
phyticine new to Florida. Fla. Dep. Agric. Consum. Serv., Div. Plant Ind. Entom. Circ. 333.
NeGron-Ortiz, V. 1998. Reproductive biology of a rare cactus, Opuntia spinosissima
(Cactaceae), in the Florida Keys: why is seed set very low? Sexual Pl. Reprod. 11:208-
Smal, J.K. 1930. Consolea corallicola, Florida semaphore cactus. Addisonia 15:25-26, pl.
Stacter, R., J. Tamory, P. Lynch, and B. Lockwoop. 1999. The vascular flora of Biscayne National
Park, Florida. Sida 18:1207-1226.
STILING, P.and D.C. Moon. 2001. Protecting rare Florida cacti from attack by the exotic cac-
tus moth, Cactoblastis cactorum (Lepidoptera: Pyralidae). Florida Entomol. 84:506-509.
STILING, P., A. Rossi, and D. Gorpon. 2000. The difficulties of single factor thinking in restora-
tion: replanting a rare cactus in the Florida Keys. Biol. Conserv. 94:327-333.
812 BRIT.ORG/SIDA 20(2)
BOOKS RECEIVED/NOTICES
Annotations and comments by Editor
Horticulture/Gardening
Francis HALLE. Translated and with foreword by Davip Ler. 2002. In Praise of Plants.
(ISBN 0-88192-550-0, hbk.). Timber Press, 133 SW Second Ave., Suite 450,
Portland, OR 97204-3527, U.S.A. (Orders: www.tim berpress.com, 800-327-
5680, 503-227-2878, 503-227-3070 fax). $24.95, hbk,, 334 pp., 99 line drawings,
6" x 9"
Contents.—Chapter 1. Plants, Animals, and Humans. Chapter 2. A Visit to the Landscape of Form.
Chapter 3. The Cell. Chapter 4. Plant Biochemistry in a Nutshell. Chapter 5. Evolution. Chapter 6. Of
Other Living Beings. Chapter 7. Ecology. The book concludes with an Epilogue, References, and In-
dex. This is a book that invites you to look closer at the world of plants and animals around us. The
first chapter contains a section on “Comparing Plants and Animals” and the ee paragraph will
give youa taste of what is tocome. “Whatare plants? Despite the important
occu our
landscapes, they are often ignored or scorned by ae n biologists, by icgeeia ait the old eee
by people in general. Does that place us on a contrasting quest to understand plants? Plants please
most women, they pleased Christ and the Buddha, the last emperor of China, the scholars of earlier
centuries, and they have pleased innumerable artists and philosophers, including Goethe, Cioran,
Colette, Valéry, Mandela, Durer, Giono, Hugo, and Rilke. First what do plants resemble and how can
we comprehend the forms they have adopted.”
ALLAN M. ARMITAGE. 2001. Armitage’s Manual of Annuals, Biennals, and Half-Hardy
Perennials. (ISBN 0-88192-505-5, hbk.). Timber Press, 133 SW Second Ave.,
Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: www.timberpress.com,
800-327-5680, 503-227-2878, 503-227-3070 fax). $39.95, hbk., 604 pp., 125
color photos, 107 line drawings, | map, 75 tables, 6" x 9"
EmLy S. Sircrrist. 2001. Bulbophyllums and Their Allies: A Grower’s Guide. (ISBN
0-88192-506-3, hbk.). Timber Press, 133 SW Second Ave., Suite 450, Port-
land, OR 97204-3527, U.S.A. (Orders: www.timberpress.com, 800-327-5680,
503-227-2878, 503-227-3070 fax). $34.95, hbk., 284 pp., 77 color photos, 6"
x 9"
Jim Cootes. 2001. Orchids of the Philippines. (ISBN 0-88192-516-0, hbk.). Timber
Press, 133 SW Second Ave., Suite 450, Portland, OR 97204-3527, U.S.A.(Orders:
www.imberpress.com, 800-327-5680, 503-227-2878, 503-227-3070 fax).
$49.95, hbk., 232 pp., 339 color photos, | diagram, | map, 7 1/2" x 10".
SIDA 20(2): 812. 2002
PLANTS NEW TO THE UNITED STATES AND FLORIDA
Richard P. Wunderlin Bruce F. Hansen
Institute for Systematic Botany Institute for Systematic Botany
Department of Biology Department of Biology
University of South Florida University of South Florida
Tampa, FL 33620-5200, U.S.A. Tampa, FL 33620-5200, U.S.A.
Loran C. Anderson
Department of Biological Sciences
orida State University
Tallahassee, FL 32306-4370, U.S.A
ABSTRACT
Eight taxa are reported as new to continental United States and the state of Florida: Alstonia scholaris,
Bromelia pinguin, Costus pulverulentus, Enydra fluctuans, Eucalyptus torelliana, Hygrophila
coins Roe se aR NETSS and Oxalis ¢ ae lar is var. Raney The reports for the non-
2 ; ;
[pee eon eee
ror Florida.
The occurrence of Didiplis d a a in ee is verified
RESUMEN
Se citan ocho taxa como nuevos para los Estados Unidos continentales y el estado de Florida: Alsto-
nid scholaris, Bromelia pinguin, Costus pulverulentus, Enydra fluctuans, Eucalyptus torelliana,
Hygrop tle diffor mis, Khaya senegalensis, y Oxalis triangularis var. papilionacea. Las citas de las no
nativas Berberis bealei y Canna glauca a la autoctona R | riculata son las primeras
para Florida ‘ Se verifica la existencia de Didipl Ira en Florida.
Continuing study on the flora of Florida and for the Guide to the Vascular Plants
of Florida, Second Edition (Wunderlin & Hansen, in press) has resulted in 12
taxa new to the state. Fight of these, all introduced, also likely represent new
records for the continental United States. Three additional species, two intro-
duced and one native, are new records for Florida, and the occurrence of Didiplis
diandra is verified for the state. This paper emphasizes the potential for more
natives to be discovered in Florida, as well as the high probability for invasion
by exotics. Of the over 4,119 taxa of plants in Florida, 1,305 (32%) are non-na-
tive (Wunderlin & Hansen 2002). The criteria for including plants in the flora
follow that of Wunderlin and Hansen (2000, p. 4).
SPECIES NEW TO CONTINENTAL UNITED STATES AND FLORIDA
Alstonia scholaris (L.) R.Br—Palm Beach Co. Village of Wellington, wetland
preserve at Wellington Mall, T445, RIIE, Sec. 13, 22 Feb 2002, Miller s.n. (USF).
Native to the Indo-Pacific area and occasionally cultivated in Florida, “dita” (or
“white cheesewood”) can be expected as an escape. The seeds are wind dispersed,
SIDA 20(2): 813-817. 2002
814 BRIT.ORG/SIDA 20(2)
and the plant at Wellington Mall apparently became established from nearby
plantings. It now has been eradicated from the site.
Bromelia pinguin L.—Hillsborough Co. Riverside Park, Temple Terrace, 6
Dec 2001, Dickman s.n. (USF). Native to tropical America, “pinguin” is an es-
cape from cultivation in central Florida.
Costus pulverulentus C. Presl—Orange Co. Valencia Community College,
East Campus, 700 block of North Econlockhatchee Trail, ca. LO mi E of Orlando,
T225S, R31E, Sec. 19, edge of riverine hammock, | Nov 2000, Myers 2 (USF). Semi-
nole Co.: Sweetwater Park, 100 block of Magnolia Street, Oviedo, 28%0.668'N,
8112.352'W, oak hammock, 2 Sep 2001, Myers 290 (USF). Native to tropical
America and escaped from cultivation in Orange and Seminole Counties.
Enydra fluctuans Lour—Hillsborough Co.: Tampa, ditch along the E side of
Manhattan Avenue just S of the intersection of Manhattan Avenue and Rem-
brandt Drive on the Interbay Peninsula, T30S, RI8E, Sec. 16, 7 Apr 2000, Shuey
& Poyner s.n. (USF). “Buffalo Spinach” is a pantropical aquatic plant, sometimes
cultivated for use in Asiatic foods. It is potentially highly invasive, readily root-
ing from stem fragments.
Eucalyptus torelliana F Muell.—Palm Beach Co: E of the intersection of Al-
ternate AlA and Hood Road, Palm Beach Gardens, 14 Jun 2001, Miller s.n. (USF).
“Torell’s eucalyptus” or “cadaga’” is native to Australia and escaped from culti-
vation in southern Florida. Trees and seedlings occur in a designated preserve,
these apparently coming from a development north of the site (Frenchman’s
Creek) where this species is used in landscaping.
Hygrophila difformis (L.) Blume—Hillsborough Co. spreading on clayey
shelf of man-made pond on N side of Cherry Street, 0.7 mi W of park Road,
Plant City, T28S, R22E, Sec. 21, SE!/+of SW !/4,5 May 1999, Hansen & Wunderlin
12893 (USF). “Water wisteria,” native to southeastern Asia, is often grown as an
aquarium plant. The Hillsborough County plants probably originated asa dis-
card from an aquarium and have now formed a sizable population in the pond.
It easily propagates from stem fragments and has high invasive potential in
lakes, ponds, and rivers. When growing out of the water, it is erect and flowers
profusely. However, mature seeds were not observed by Hansen and Wunderlin.
Khaya senegalensis (Desr.) A. Juss.—Collier Co.: Collier Seminole State Park,
Campsite #7 of 1-19, T51S, R27E, Sec 27, SE!/4, SW 1/4, SE !/4, rockland ham-
mock, 17 Apr 1998, Hattaway CS0319 (USF). “African mahogany” is native to
Africa and widely grown asa street and timber tree in the tropics (Irvine 1961).
It has the potential to invade disturbed tropical forests. It is naturalized in West-
ern Australia (Csurhes & Edwards 1998). The population in Collier-Seminole
State Park consisted of several trees (one very large), but these have now been
eradicated.
Oxalis triangularis A. St.-Hil. subsp. papilionacea (Hoffmanns. ex Zucc.)
WUNDERLIN ET AL., PLANTS NEW TO THE UNITED STATES AND FLORIDA 815
Lourteig—Leon Co. several plants naturalized in deeply shaded loam along road-
side of Duparc Circle near its W junction with Lakeshore Drive, Tallahassee, 23
Feb 2001, Anderson 19691 (FSU). “Purpleleaf false shamrock,” native to South
America, is occasionally cultivated in North America for its purple leaves. The
Leon County site is the only location known where it is naturalized in North
America.
SPECIES NEW TO FLORIDA
Berberis bealei Fortune—Leon Co. naturalized single shrub (vegetative) in moist
loam of deeply shaded creek swamp NNE of marker 20 in Elinor Klapp-Phipps
Park, W of Meridian Rd, N of Tallahassee, T2N, R1W, Sec. 36, NW 1/4 of NW !/4,
30 May 1996, Anderson 16530 (FSU); flowering shrub in pine-oak woodland rem-
nant on S side of Duparc Circle just N of Lakeshore Drive, Tallahassee, 23 Feb
2001, Anderson 19688 (FSU). These two sites are about 1.6 miles apart. Natural-
ized in Delaware, Virginia, North Carolina, South Carolina, Georgia, and Ala-
bama (USDA, NRCS 2002), “Beale’s barberry” or “leatherleaf mahonia’” is now
reported from Florida. It is an evergreen shrub native to China and frequently
cultivated outside its native range.
Canna glauca L.—Madison Co. along US 90, Greenville, 29 July 1967, Beckner
1885 (FLAS). The species is native to tropical America and previously known in
the United States only from Louisiana and Texas where it is considered to be
native (Kress & Prince 2000). It is escaped from cultivation at one site in Madi-
son Co., Florida, where it occurs as frequent clumps around several burned-out
houses near a lake where it is spreading vegetatively and by seed.
Didiplis diandra (Nutt.ex DC.) A.W. Wood—Sarasota Co. Pinelands Reserve,
ca. 1 mi SW of Lower Myakka Lake, T385S, RI9E, Sec. 1, freshwater marsh, 6 Apr
1998, Morrison 98064-1 (USF). “Waterpurslane,” a submergent or emergent
aquatic is apparently native from Minnesota and Wisconsin south to Georgia,
Alabama, Mississippi, Louisiana, and Texas (USDA, NRCS 2002), and now docu-
mented in Florida. This species has been reported (sometimes as Peplis diandra
Nutt.ex DC.) for Florida in many floras, (e.g., Chapman 1860; Correll & Johnston
1970; Crow & Hellquist 2000; Fassett 1957; Fernald 1950; Gleason & Cronquist
1963, 1991; Radford et al. 1968; Small 1933). The basis for these reports is not
known. No Florida material was found at NY, the principal repository for A. W.
Chapman and J. K. Small collections (Stafleu 1967). A search of the herbaria at
FLAS and FSU also yielded no Florida material. It is not reported by Godfrey
and Wooten (1981), Clewell (1985), or Wunderlin (1982, 1998). The species is
apparently rare in much of its natural range, but is rather inconspicuous and
perhaps has been overlooked. It is sometimes grown as an aquarium plant, but
it is not believed that the Sarasota plants represent an introduction.
Rudbeckia auriculata (Perdue) Kral—Walton Co. along US 331, 0.5 mi N of
816 BRIT.ORG/SIDA 20(2)
FL 2 at Gordon, ca. 6.5 mi S of Paxton, roadside bog, 17 Aug 2000, Searcy s.n.
(USF). A rare species, “eared conef lower” is known only from Georgia, Alabama
(USDA, NRCS 2002), and now Florida. A second attempt to locate the plants
was unsuccessful.
ACKNOWLEDGMENTS
We thank Steve Dickman, Bobby Hattaway, Ray Miller, Melissa Morrison, Steven
Myers, Darryl Searcy, and Allen Shuey for bringing various of these species to
our attention and providing collections.
REFERENCES
CHapman, A.W. 1860. Flora of the southern United States. New York: Ivison, Phinney & Co.
Crewelt, AF. 1985. Guide to the vascular plants of the Florida panhandle. Gainesville: Uni-
versity presses of Florida/Florida State University Book.
Corrett, D.S.and M.C. Johnston. 1970.Manual of the vascular plants of Texas. Renner: Texas
Research Foundation.
Crow, G.E. and C.B. Hettquist. 2000. Aquatic and wetland plants of northeastern North
America. Volume 1. Pteridophytes, Gymnosperms, and Angiosperms: Dicotyledons.
Madison: The University of Wisconsin Press.
CsurHes, S. and R. Eowarps. 1998. Potential environmental weeds in Australia: Candidate
species for preventive control. Queensland Department of Natural Resources.
Fassett, N.C. 1957.A manual of aquatic plants.With revision appendix by E.C.Ogden.Madi-
son: University of Wisconsin Press.
Feanaco, M.L. 1950. Gray’s manual of botany. 8th ed. New York: American Book Co.
Gteason, H.A. and A. Cronauist. 1963. Manual of vascular plants of northeastern United
States and adjacent Canada. Princeton, NJ:D.Van Nostrand Co.
Gteason, H.A. and A. Cronauist. 1991. Manual of vascular plants of northeastern United
States and adjacent Canada. New York: The New York Botanical Garden.
Goorrey, R.K. and J.W. Wooten. 1981. Aquatic and wetland plants of southeastern United
States: Dicotyledons. Athens: University of Georgia Press.
InviNE, FR. 1961.Woody plants of Ghana. London: Oxford Press.
Kress, W. J. and L.M. Prince. 2000. Cannaceae. In: Flora of North America Editorial Commit-
tee. Flora of North America north of Mexico. 22: 310-314. New York/Oxford: Oxford
University Press.
Raproro, A.E., H.E. Adves, and C.R. Bett. 1968. Manual of the vascular flora of the Carolinas.
Chapel Hill: University of North Carolina Press.
Smatt, J.K. 1933. Manual of the southeastern flora. New York: Published by the Author.
StarFteu, FA, 1967. Taxonomic literature. Regnum Veg. 52:1-556
USDA, NRCS. 2002. The PLANTS database, version 3.5 (http://plants.usda.gov). National
Plant Data Center, Baton Rouge, LA. (Accessed September 12, 2002).
Wunpbertin, R.P. 1982. Guide to the vascular plants of central Florida. Tampa: University of
South Florida Book/University Presses of Florida.
WUNDERLIN ET AL., PLANTS NEW TO THE UNITED STATES AND FLORIDA 817
WUNDERLIN, R.P. 1998. Guide to the vascular plants of Florida. Gainesville: University Press of
Florida.
Wunpbeetn, R.P. and B.F. Hansen. 2000. Flora of Florida. Volume 1:Pteridophytes and gymno-
sperms. Gainesville: University Press of Florida.
WuNDERLIN, R.P.and B.F.HANseN.2002. Atlas of Florida vascular plants (
[S.M. Landry and K.N. Campbell (application cineca Florida Gants for Com-
munity Design and Research] Institute for Systematic Botany, University of South
Florida, Tampa, FL. (accessed September 12, 2002).
WHUNDERLIN, R.P. and B.F. Hansen. In Press. Guide to the vascular plants of Florida, second
edition. Gainesville: University Press of Florida.
/ / Yang |
818 BRIT.ORG/SIDA 20(2)
BOOKS REGEIVED/NOVICES
Annotations and comments by Editor
Taxonomic/Systematic
James D. Stuart and Joun O. Sawyer. Illustrarted by ANpREA J. PICKART. 2002. Trees
and Shrubs of California. (ISBN 0-520-22110-9, pbk.). University of Cali-
fornia Press, Berkeley, CA 94720, U.S.A. (Orders: California Princeton Ful-
fillment Services, 1445 Lower Ferry Road, Ewing, NJ 08618, U.S.A., 609-
883-1759, 609-883-7413 fax, www.ucpress.edu). Price not given, +67 pp., +0
color photos, over 200 line drawings, 300 range maps, 4 3/4" « 7 1/4",
Key Worps: Trees, Shrubs, Identification, California, Pictorial Works.
GRAHAM NICHOLLS, Rick Lupp (Consulting Editor), and Foreword by Bossy Warp.
2001. Alpine Plants of North America. An Encyclopedia of Mountain Flowers
from the Rockies to Alaska. (ISBN 0-88192-548-9, hbk.). Timber Press, 133
SW Second Ave., Suite 450, Portland, OR 97204-3527, U.S.A. (Orders:
www.timberpress.com, 800-327-5680, 503-227-2878, 503-227-3070 fax).
$49.95, hbk., 344 pp., 495 color photos, 2 maps, 7 3/8" x 10 3/8".
Key Worps: Alpine Plants, Mountain Plants, Wild Flowers, West (U.S., Canada), Pictorial Works
DONALD E. SCHNELL. 200L. Carnivorous Plants of the United States and Canada.
Second Editon. (ISBN 0-88192-540-3, hbk.). Timber Press, 133 SW Second
Ave., Suite 450, Portland, OR 97204-3527, U.S.A. (Orders:
www.timberpress.com, 800-327-5680, 503-227-2878, 503-227-3070 fax).
$39.95, hbk., 468 pp., 200 color photos, 7 line drawings, 27 maps, 6" « 9".
Key Worps: Carnivorous Plants, United States, Identification, Venus Flytrap, Pitcher Plants, Califor-
nia Pitcher Plant, Sundews, Butterworts, Bladderworts.
ARMANDO T. HUNZIKER. 2001. Genera Solanacearum: The Genera of Solanaceae
Illustrated, Arranged According to a New System. (ISBN 3-904144-77-4,
hbk.). A.R.G. Gantner Verlag, Ruggell / Liechtenstein. (Orders: Koeltz Sci-
entific Books, PO. Box 1360, D 61453 Koenigstein / Germany; +49 6174 937240
fax; koeltz@t-online.de, http://www.koeltz.com). $120.00, 500 pp, 136 line
drawings, 61/2" x 91/2".
an
Key W
, keys, descriptions, illustrations, 92 genera.
SHARON C. YARBOROUGH and A. MicHaAri PoweLL. 2002. Ferns and Fern Allies of
the Trans-Pecos and Adjacent Areas. (ISBN 0-096072-476-X, pbk.). Texas
Tech University Press, Box 41037, Lubbock, TX 79410, U.S.A. (Orders: 800-
832-4042, wwwttup.ttuedu/). $17.95, 128 pp., 37 line drawings, range maps,
6" x 9"
Kry Worps: Ferns, Pteriodophyta, Trans-Pecos, Texas, New Mexico, identification.
SIDA 20(2): 818. 2002
CRYPTOCORYNE BECKETTIITCOMPLEX (ARACEAE)
INTRODUCED AT A FLORIDA SPRING
Colette C. Jacono
United States Geological Survey
lorida Caribbean Science Center
7920 NW 71ST St.
Gainesville, Florida, 32653, U.S.A.
ABSTRACT
A vegetative population ot CK ee, yne (Araceae), introduced at a Florida spring, appeared to rep-
resent three closely rel ©. beckettii complex: C. beckettii Thw. ex Trimen, C. wendtii
de Wit and C. undulata We nd. Indiv igual sof C. undulata were true to type and seule be caine ated
at the site. Intergradation of diag
ering. While some transplants produced spathes characteristic of one er Tour ii or é bec
intermediates between the two species were common. Neither C. beckettii nor C. wendtii could be
delineated at the site. The seclusion of the stream and the integrity of native plant communities have
downstream
ae ee ee
likely p nt p
>
ze lll,
RESUMEN
Una poblacion vegetativa de Cr ypeee ya (Araceae), introducida en un manantial de Florida, al
parecer representa tres especies relacionadas del complejo C. beckettii: C. bechettii
Thw. ex Trimen, C. wendtii de Wit y C. undulata Wendt. En el lugar de estudio, fueron clasificados
como verdaderos tipos ejemplares de C. undulata. En otros ejemplares, después del transplante y
floracion, fue comun obser var caus _ caracteres diagnoésticos. Aun cuando algunos
transplantes produjeron Sues C. we ndtii o de C. beckettii, fue comun observar
ejemplares con caracteristicas int dias d . Las especies C. beckettii y C. wendtii
no fueron Geuneagee en pe sitio as estudio. | La condicion ie aislamiento del manant iia he la conde on
integra de p han impedido probablemente | |
ahaa
INTRODUCTION
Cryptocoryne Fisher ex myers are aly sought-after and commercially
marketed as decorative plants for tropical aquariums. The most commonly cul-
tivated species belong to the C. beckettii complex, a uniform and specialized
group endemic to Sri Lanka Jacobsen 1977; Jacobsen 1987; de Graaf & Arends
1986)
In their native range, members of the Cryptocoryne beckettii complex in-
habit pristine rivers and springs. In some areas, they form large submersed or
emergent stands Jacobsen 1987), although some populations have declined from
commercial collecting and habitat destruction (de Graaf & Arends 1986). Spe-
cies in this group commonly root in the sand and silt that overlay rocky, shal-
low streambeds (de Graaf & Arends 1986; Rataj @ Horeman 1977) and occupy
SIDA 20(2): 819-832. 2002
820 BRIT.ORG/SIDA 20(2)
shady, sheltered niches rarely inhabited by other plants Jacobsen 1977; Doyle
2001). They adapt readily to terrestrial conditions and flower with water draw
down, when leaves become emersed and the roots well aerated lacobsen 1977,
Clark 1991). Vegetative reproduction is important in population establishment
and expansion JJacobsen 1977). In cultivation, C. beckettii members are hardy
and vigorous, propagating abundantly by rhizomes and stolons, and enduring
hard, calcium rich water (Miller 1998; Rataj @ Horeman 1977).
In treating Cryptocoryne for the Flora of Ceylon, Jacobsen (1987) portrayed
four species as closely related within the Cryptocoryne beckettii complex:
Cryptocoryne walkeri Schott, C. beckettii Thw. ex Trimen, C. wendtii de Wit,
and C. undulata Wendt. Although mainly diploids, all four have been found as
triploid species and several combinations within the complex have resulted in
the formation of hybrids Jacobsen 1977; Arends et al. 1982; Jacobsen 1987). Be-
cause Cryptocoryne share many similarities in their vegetative features, an in-
florescence is requisite for species identification. Determination of the species
in the C. beckettii group rests on subtle differences in the morphology of the
spathe, which isa fleshy, ornate bract enclosing the inflorescence. Key features
of the spathe include the size and color of the collar and the shape and degree
of twist of the terminal limb. Since differences in these characters are essen-
tially quantitative, it has been recognized that the four taxa could prove to con-
stitute a single species lacobsen 1987).
Wunderlin’s (1998) inclusion of Cryptocoryne wendtii de Wit as an intro-
duced component of the Florida flora was the first recording of this Southeast
Asian genus for North America. Soon after, Rosen (2000) reported C. beckettii
Thw. ex Trimen as introduced to Texas, followed by Doyle (2001) who detailed
itsexpansion and spread in the upper spring-fed regions of the San Marcos River.
Rosen's description of C. beckettii from the San Marcos River, Texas, was based
on vegetative material (Rosen 2000). Doyle (2001) reported that C. beckettii had
been adequately confirmed in Texas after K. Sanders and D. Lemke identified
flowering transplants. Wunderlin‘s report of C. wendtii in Florida was simi-
larly based ona sterile specimen collected at Rainbow Springs Aquatic Preserve,
Florida, in 1989. The objective of this inquiry was to confirm the persistence of
Cryptocoryne at Rainbow Springs pavers Preserve and to collect material for
cultivation and subsequent species determination from flowering plants.
The Rainbow River is a karstic, spring-fed stream receiving artesian
groundwater that is high in clarity, rich in soluble carbonates and moderate in
temperature (USGS 1999). The run traverses 8 km before joining the
Withlacoochee River near the town of Dunnellon. A site of exceptional scenic
beauty anda popular destination for recreation, it was designated the Rainbow
Springs Aquatic Preserve in 1986 (Florida Statute 258.39 [32)).
JACONO, CRYPTOCORYNE BECKETTII COMPLEX IN FLORIDA 821
METHODS
In May 2000 and in June 2001, approximately 50 individuals were uprooted
from the Cryptocoryne colony found at Rainbow Springs Aquatic Preserve. Four
sided, open-ended aluminum units (0.25 m2) enclosed in double netting were
placed over each area during sampling to delineate the sampling area and to
trap stray plant fragments. Sampling attempted to localize variation observed
in leaf color and in total represented seven areas within the colony. Samples
were transplanted to 10 cm pots ina bark and peat-based potting medium and
cultivated out of water, outdoors, with shade and screening, during the warmer
months. Transplants established readily, losing submersed leaves and produc-
ing new terrestrial foliage. Pots were moved toa heated greenhouse when night-
time temperatures dropped below 10°C. Flowering began in December and con-
tinued through July. The inflorescences survived for several days and were
documented with photographs and voucher specimens before decline. On press-
ing and drying, the inflorescences lost color and the angle of terminal limbs
appeared more upright. Photography provided for better study than did pre-
pared specimens. Species identification was based on Jacobsen (1987).
To assess plant density, the extent of subsurface rhizomes, and the poten-
tial of hand removal as a control method, total vegetation was harvested from
two 0.25 m2 areas within the colony. Limestone gravel was removed with fin-
gers as necessary to extract plants in whole (ie. with basal parts and some ex-
tent of rhizomes intact). Roots were harvested to approximately 10 cm deep.
Top portions of plants were separated from subsurface material (roots). Weight
was taken before and after material was oven dried at 60T
Adjacent, sterile colonies of Myriophyllum and Cyperaceae were sampled
for subsequent transplanting, flowering and identification. Myriophyllum was tank
cultivated in 50 cm of water, the sedge was grown under terrestrial conditions.
RESULTS AND DISCUSSION
Cryptocoryne was found persisting and well established at Rainbow Springs
Aquatic Preserve, in a shaded cove of a spring fed stream. The plants grew sub-
mersed in an area of geen — m2. Water depth ranged from 28 to 75
cm, although outlying runner to crevices as deep as 150 cm. The low
growing plants reached an average ae ol cm: The vegetation consisted of
dense mats that grew to the exclusion of other macrophytes.
Transplants from the Cryptocoryne colony produced approximately 100
inflorescences. Voucher specimens and photographs documented 49 represen-
tative inflorescences. Twenty-five appeared characteristic of three species in the
C. beckettii complex: C. undulata, C. beckettii, and C. wendtii (Fig. 1-3). However,
20 were intermediate between C. beckettii and C. wendtii. Cryptocoryne
822 BRIT.ORG/SIDA 20(2)
Fic. 1. Cryptocoryne undulata Wendt.
undulata was the only species that could be delineated at the Preserve. The re-
maining, and predominant, vegetation at the site coulc
ther C. beckettii or C. wendtii.
{
not be delineated as ei-
—
JACONO, CRYPTOCORYNE BECKETTII COMPLEX IN FLORIDA 823
Fic. 2. Cryptocoryne wendtii de Wit.
Spathes of C. undulata were uniform in color and morphology. The collar
zone graded from yellow to cream and the terminal limb darkened from yellow
to golden brown upon maturing. The terminal limb was always nearly doubly
twisted and positioned diagonally to the axis of the spathe tube. Cryptocoryne
undulata was delineated within a small (approx. 4m), distinct region of the
colony. Submersed, unopened inflorescences found within this region were simi-
larly yellow at the collar zone and on the inside of the unfurled limb.
Submersed leaf blades of C. undulata were narrowly lanceolate. They
ranged in length from 4.0 to 9.0 cm (mean 6.1, SD L5) and in width from 0.7 to
1.8 cm (mean 1.3, SD 0.4). Bases were cordate to cuneate and the entire leaf mar-
gin was crisped. Submersed leaf color was dark brownish green on the adaxial
surface; abaxial sides and petioles were maroon. Terrestrial plants of C. undulata
produced smaller lanceolate leaves with acuminate bases and margins that were
crisped only along the leaf base. Terrestrial leaf color was deep green with veins
and petioles red.
While four transplants produced spathes characteristic of C. wendtii, and
17 characteristic of C. beckettii, more than 20 individuals appeared intermedi-
824 BRIT.ORG/SIDA 20
=
NS
)
ate and could not be determined to the species level. Jacobsen (1987) separated
C. beckettii from C. wendtii by the angle on the terminal limb of the spathe.
The limb of C. beckettii is more upright or sub-erect, and twisted to recurved
while that of C. wendtii is obliquely twisted, often extending intoa tail Jacobsen
1987). Jacobsen (1976) noted these characters as difficult to describe and vari-
able, pat tly in ad aes to environmental conditions. In the present study, the
key characters were found toi across a population grown under green-
house conditions. The angle of the terminal limb (as measured from the spathe
tube) ranged widely from 93°to 175°(Table 1). Limbs assumed different shapes,
as well as angles, and spathes frequently shared diagnostic features. For example,
spathes with sub-erect limbs (limbs not obliquely twisted) were found extend-
ing into well-defined tails (Fig. +b).
Jacobson (1976; 1987) further depicted the terminal limbs of C. beckettii as
yellowish to brownish and those of C. wendtii as yellowish to purplish brown.
In this study limbs varied in color from golden to green to greenish or purplish
brown. On occasion they darkened with maturity. Limb color did not always
correlate with limb position, for example, greenish brown limbs ranged from
more erect to nearly perpendicular (Fig. 4d) and true green limbs were found
twisted at an angle (Fig. 4a). Thus, in form and color many individuals in this
study appeared intermediate between C. beckettii and C. wendtii.
The identity of those individuals that could be determined according to
floral characteristics was often in question when apparently distinct individu-
als (C. beckettii and C. wendtii) were traced back to the same sampling block at
the Preserve (Fig. 5 and representative specimens Jacono 373 vs. Jacono 368). As
a result, clear clonal populations, although expected for long established, veg-
etatively reproducing plants, could not be delineated for either C. beckettii or C.
wendtii.
Submersed inflorescences found in sunny shallow areas had purple collar
zones and green inner limbs. However, with spathes tightly furled, they con-
tributed little in separating C. beckettii from C. wendtii, and merely confirmed
the ability of this type to flower in situ.
Plants of this type (C. beckettii, C. wendtii, and the intermediates) demon-
strated differences in leaf color while growing at the Preserve. Submersed leaf
color of the adaxial surfaces varied from green marbled with brown to pink
infused with green venation. Abaxial sides were violet to pink. This color varia-
tion was apparent in zones and may be attributed to factors of substrate and
sunlight at the site. Green-brown plants grew in organic sediment under con-
tinuous shade of the tree canopy; pink plants grew on limestone substrate in
zones of higher sunlight. After transplanting, the terrestrial leaves that replaced
the submersed vegetation were without variation and uniform in color. The
submersed leaves were lanceolate to ovate with cordate bases and sinuous mar-
gins. The length of their leaf blades ranged from 3.8 to 9.6 cm (mean 6.2, SD 15)
JACONO, CRYPTOCORYNE BECKETTII COMPLEX IN FLORIDA 825
Table 1. Distribution of limb angles (measured from the spathe tube) of 22 inflorescences
photographed.
No. spathes
i) i) -) © © oe) i) io)
S=- GW © Ts oe)
i tie) on ie
O- iO: WS renee: ) Sinn
eo) © - N oe) + ce) ™
Nar os Te be Sa be es » i ea
Limb angle in degrees
and their width measured 1.1 to 3.3 cm (mean 2.2, SD 0.6). Terrestrial leaves were
smaller and narrowly ovate with cordate bases, olive-green adaxial surfaces and
violet on the abaxial side.
Variability and intergradation of the features upon which these species are
based prevented delineation of C. beckettii from C. wendtii. While certain indi-
viduals seemed easy to place to the species level, they could not be traced back
to distinct areas at the site. Other individuals were intermediate and could not
be determined to species. Regardless of spathe morphology, terrestrially grown
transplants within this type were consistently uniform in vegetative features.
The predominating Cryptocoryne material at the Preserve can only be described
as C. beckettii sensu lato.
The Cryptocoryne subsurface biomass consisted of large amounts of roots,
rhizomes and stolons that were difficult to uproot by hand. Roots were well
entrenched in contrasting substrates: silty, t near the shoreline
and pebbled to rocky limestone extending into the stream.
Plant density values were 1480 plants / m? for Cryptocoryne beckettii sl.
and 1880 plants / m? for C. undulata. Dry roots of C. beckettiis.l. weighed nearly
four times more than dry top growth: dry roots of C. undulata weighed eight
times that of corresponding top growth (Table 2).
Like many aroids, Cryptocoryne employ contractile roots to adjust plant
level after heaving or flooding (Bown 1988). Tiny rootlets sent deep into the
rock use root pressure to contract in length and pull the roots farther into the
BRIT.ORG/SIDA 20(2)
Fic. 3. Cryptocoryne beckettii Thw. ex Trimen.
JACONO, CRYPTOCORYNE BECKETTII COMPLEX IN FLORIDA 827
d C. wendtii, a—d (left to right).
828 BRIT.ORG/SIDA 20(2)
Taste 2.Plant density and top to root growth, per m? of submersed plants.
; op fresh Top dry Roots fresh Roots dry Roots/
of plants wt. ing wt.ing t.ing wt.ing op
Cryptocoryne undulata 1880 349g 23.2 g 1007q 183.8g 79
Cryptocoryne beckettii s.1. 1480 251g 20g 426g 74g 3.7
substrate. This adaptation firmly anchors plants in swift currents and likely
accounts for the deep rooting in limestone substrate at the Preserve, comp
ing mechanical removal without destructive draglines or dredging
Shortened runners, basal shoots, were formed at the base of both submersed
and transplanted material of all three species. Jacobsen (1987) described these
“bulbil-like runners” on forms of C. wendtii and C. undulata, but did not report
them for C. beckettii. Small (< 5mm), abundant (up to 8-20 per individual), and
fragile, the shoots are loosely associated with the mother plant and readily break
off. They quickly develop an initial root and primary leaf to establish as new
plants. Kane et al. (1999) successfully employed basal shoots for the in vitro
propagation of C. wendtii. At the Preserve, basal shoots were abundant in spring-
time. In transplants, they provided the flush of new season growth after spring
flowering. Walking on submersed plants can dislodge basal shoots, which im-
mediately sink to the bottom. Conversely, dislodged rhizomes float on the wa-
ter surface. Because of their abundance and small size, the shoots are difficult
to contain and may be carried downstream with bottom currents; they may be
more effective than rhizome fragments as propagules for downstream spread.
Reproduction in Cryptocoryne, like other submersed macrophytes, is pri-
marily accomplished through vegetative structures. Rhizomes, runners (stolons)
and basal shoots provide multiple means for expansion of an apparently well-
suited population at Rainbow Springs Aquatic Preserve.
Submersed inflorescences were observed in April and June on both types
of Cryptocoryne at the Preserve. In their native range, Cryptocoryne commonly
grow submersed during rainy periods and emerge to flower when seasonal
water levels drop Jacobsen 1977). Submersed inflorescences, while not uncom-
mon in nature, must emerge to be insect ae before forming seeds. Typi-
cally short-lived, the seeds may be dispersed in water but are not vectored long
distances over land (Bown 1988; Jacobsen 1976; Bogner 1987). Constant water
level ensured by the spring flow should keep the inflorescences of submersed
plants underwater. However, species in the C. beckettii complex are well adapted
to terrestrial life Jacobsen 1977) and runners could establish amphibious plants
that might flower along the damp, shady shoreline at the Preserve.
Cryptocoryne was found encroaching on adjacent Sagittaria kurziana
Gliick, Myriophyllum heterophyllum Michx. anda continually inundated popu-
—
icat-
JACONO, CRYPTOCORYNE BECKETTII COMPLEX IN FLORIDA 829
lation of Rhynchospora colorata (L.) H. Pfeiff. Several meters downstream,
Zizania aquatica L., Rorippafloridana Al-Shehbaz & Rollins, Ludwigia repens
J.R. Forst., Sagittaria lancifolia L., Hydrocotyle umbellata L., Chara, Fontinalis,
Utricularia, and Leersia spp. contributed to one of the few intact aquatic com-
munities remaining at the Preserve. The nator! ae may curtail the
downstream spread of Cryptocoryne by t shoots
Extensive underwater monitoring conducted - in 1996 ae 2000 detected no
additional sites with Cryptocoryne at Rainbow Springs Aquatic Preserve (P.
Sleszynski, pers. comm.).
Assessing Risks
In North America, weedy infestations of Cryptocoryne beckettii sensu lato may
be limited to karstic spring environments of the southern states. Laboratory
experiments demonstrated that four Cryptocoryne species, C. affinis Hook f., C.
cordata Griffith, C. elliptica Hook.f. and C. minima Ridley, effectively used bi-
carbonate as the main carbon source when pH ranged between 7.5 and 9.1
(Cheng and Mansor 2000). In high pH (>7.0), hard water systems, relatively
large amounts of dissolved inorganic carbon exist as bicarbonate (Spencer and
Bowes 1990). At Rainbow Springs Aquatic Preserve the spring water is consis-
tently alkaline and carbonate-rich. Within the last decade pH ranged from 7.87
to 8.27 (USGS 1999). Total hardness value, last determined in 1980, was 58 mg/
L, measured as calcium carbonate, and noncarbonate hardness was 3 mg/L,
expressed as equivalents of calcium carbonate, indicating a high level of car-
bonate in the water (USGS 1999). Similar pH values (7.6 to 8.0) were recorded
near the headspring of the San Marcos River, although hardness values were
not determined (USGS 2000). The ability of Cryptocoryne to selectively use bi-
carbonate under alkaline conditions may contribute to its successful coloniza-
tion at karstic spring-fed streams in Florida and Texas.
The original Cryptocoryne “mother plants” were probably planted at Rain-
bow Springs Aquatic Preserve before 1986 when the system came under state
ownership. Now well established, its potential for dispersal there is consider-
able. In the San Marcos River, disturbance caused by wading and sporting ac-
tivities are suspected to contribute to the proliferation of Cryptocoryne beckettii
downstream (P. Power, U.S. Fish and Wildlife Service, San Marcos, Texas, pers.
comm.). Owens et al. 2OO1) reported a significant increase in dislodged plant
fragments below high recreation use areas on the San Marcos River. Likewise,
recreational activities were reported as significant in the uprooting of large
masses of vegetation in Rainbow River (Mummaetal. 1996). The secluded stream
harboring Cryptocoryne at Rainbow Springs Aquatic Preserve is currently re-
stricted, but under consideration for public use.
Original voucher specimen. Cryptocoryne wendtii de Wit, U.S.A. FLORIDA, Marion Co.: Dunnellon,
Rainbow Springs Aquatic Preserve, quiet spring runs along the Rainbow River, lvs pink/green, fre-
quent, 24 Apr 1989, Joe Hinkle s.n. (FLAS), Det: Walter Driggers.
830 BRIT.ORG/SIDA 20(2)
Fic. 5. Apparently distinct individual planted pling
Representative specimens from this study: U.S.A. Florida. Marion Co., Dunnellon,
Rainbow Springs Aquatic Preserve, spring run along the Rainbow River.
Cryptocoryne undulata Wendt—Submersed to 75cm at site, sterile: Ll Apr 2001, Jacono 253 (FLAS,
MO), 12 Jun 2001, Jacono 280 (FLAS). Submersed to 75cm at site, fertile with spathes closed: 11 Apr
2001, Jacono 254 (FLAS). Flowering in cultivation: 10 Apr 2001, Jacono 282 (MO), 31 July 2001, Jacono
296 (FLAS); Apr 2002, Jacono s.n. FLAS); 7 Apr 2002, Jacono 408 (FLAS).
Cryptocoryne beckettii [hw.ex Trimen—Flowering in cultivation: 17 Dec 2001, Jacona 368 (MO);
18 Feb 2001, Jacono 211 (FLAS) 7 Apr 2001, Jacono 231 (FLAS), 7 Apr 2001, Jacono 233 (MO), Lo Apr
2001, sant AS).
coryne wendtii de Wit—Flowering in oo 14 Feb 2001, Jacono 210 (FLAS); 29 Feb
2001, p60 AS); 27 Jan 2002, Jacono 373 (N
e beckettii sensu lato—Submerse : to 28cm at site, sterile: 11 Apr 2001, Jacono 249,
(FL ee eee au. Jacono 275 (MO). Submersed to 28 cm at site, fertile with spathe closed, 1] A
2001, Jacono 250 (MO); 12 June 2001, Jacono 276 (FLAS). Flowering in cultivation with spathes appar-
ently intermediate between C. beckettii and C. wendtii: 16 Apr 2001, Jacono 243 (MO); 16 Apr 2001,
Jacono 244 (FLAS); 21 Feb 2001, Jacono 247 (FLAS), 21 Feb 2001, Jacono 271 (FLAS); 20 Feb 2002,
Jacono 379 (FLAS); 26 Mar 2002, Jacono 402 (FLAS).
Myriophyllum heterophyllum Michx.—Submersed plants, rooted, sterile, 19 May 2000, Jacono
I40(FLAS);C ne plants from sterile collection Gacono 140) flowering and fruiting, 20 May 2001,
Jacono 270(F
Rhynchospora colorata (L.) H. Pfeiff.—Sterile plants submersed in ca. 70 cm of clear, flowing
spring water, 12 Jun 2001, Jacono 281 (FLAS); Cultivated plants from sterile collection Gacono 281)
flowering and fruiting, 8 Oct 2001, Jacono 360 (NY, FLAS)].
JACONO, CRYPTOCORYNE BECKETTII COMPLEX IN FLORIDA 831
ACKNOWLEDGMENTS
eee 1.f£
The author thanks the following i
Sleszynski granted site access and helped with sample collection; Michael FE
Kane, Nancy L. Philman and A.E. Dudeck provided greenhouses for over-win-
tering; Niels Jacobsen, confirmed species identification with the photographs
shown in Figures 1-3; Jan Bastmeijer kindly offered opinions on identification
of Figures 1-3; Maria S. Sepulveda translated the abstract; Nancy C. Coile, R-E.
Smith and David W. Hall reviewed versions of the manuscript; C.D.K. Cook pro-
vided insightful discussion; and W. Wayt Thomas confirmed Rhynchospora
colorata.
their unique contributions: Pete
REFERENCES
Arenbs, J.C., J.D. Bastmever and N. Jacossen. 1982. Chromosome numbers and taxonomy in
Cryptocoryne (Araceae). II. Nordic J. Bot. 2:453-463.
Boaner, J. 1987. Morphological variation in Aroids. Aroideana 10:4—-16.
Bown, D. 1988. Aroids: plants of the arum family. Timber Press, Portland.
CHENG, H.S.and M. Mansor. 2000. A comparative study on the acquisition of inorganic car-
bon by four native aquatic plants from the genus Cryptocoryne and an aquatic weed
Hydrilla verticillata. Malaysian Appl. Biol. 29:43-53.
Crark, R. 1991. Flower and seed production in Cryptocoryne. Aquatic Galcener 4(1):7-14.
be Grarr, A.and J.C. Arenos. 1986. The occurrence of Cryptocoryne and Lagenandra (Araceae)
on Sri Lanka. Nord. J. Bot. 6:757-764.
Doyte, R.D.2001. Expansion of the exotic aquatic plant Cryptocoryne beckettii (Araceae) in
the San Marcos River, Texas. Sida 19: 1027-1038.
Jacossen, N. 1976. Notes on Cryptocoryne of Sri Lanka (Ceylon). Bot. Not. 129:179-190.
Jacossen, N. 1977. Chromosome numbers and taxonomy in Cryptocoryne (Araceae). Bot.
Not. 130:71-87.
Jacossen, N. 1987. Cryptocoryne, in M.D. Dassanayake, ed. A revised handbook to the flora
of Ceylon. Amerind Publishing Co. Pvt. Ltd., New Delhi.
Kane, M.E, G.L. Davis, D.B. McConnett and J.A. GaraiuLo. 1999. In vitro propagation of
Cryptocoryne wendtii. Aquatic Bot. 63:197-202.
Miter, R. 1998. Caring for groups of Cryptocorynes or waylaying your wandering wendtii.
Aquatic Gardener 11:187—-189
Mumma, M.T., C.E. CicHra, and J.T. Sowarps. 1996. Effects of recreation on the submersed
aquatic plant community of Rainbow River, Florida. J. Aquatic Pl. Mangem. 34:53-56.
Owens, C.S., J.D. MADSEN, R.M. Smart, and R.M. Stewart. 2001. Dispersal of native and nonna-
tive aquatic plant species in the San Marcos River, Texas. J. Aquatic Pl. Mangem.39:
75-79,
Rataj, K.and T.J. Horeman. 1977. Aquarium Plants: their identification, cultivation and ecol-
ogy. I.F.H. Publications, Inc., Neptune, New Jersey.
832 BRIT.ORG/SIDA 20(2)
Rosen, D.J. 2000. Cryptocoryne beckettii (Araceae), a new aquatic plant in Texas. Sida 19:
399-401.
Spencer, W. and G. Bowes. 1990. Ecophysiology of the world’s most troublesome aquatic
weeds. In A.H.Pieterse and K.J.Murphy (Ed.). Aquatic weeds, the ecology and manage-
ment of nuisance aquatic vegetation. Oxford University Press, Oxford.
USGS. 1999. United States Geological Survey Water Quality Database (http://
water.usgs.gov/nwis/qwdata).
USGS. 2000. United States Geological Survey Water Quality Database (http://
waterdata.usgs.gov/tx/nwis).
Wunber.in, R.P. 1998. Guide to the vascular plants of Florida. University Press of Florida,
Gainesville.
NEPHROLEPIS EXALTATA (DRYOPTERIDACEAE)
NEW TO ARKANSAS
James H. Peck
Department of Biology
rsity of Arkansas at Little Rock
2801 S, University Ave.
Little Rock, AR 72204, U.S.A.
ABSTRACT
Nephrolepis exaltata (Dryopteridaceae) is reported as new to Arkansas.
RESUMEN
Nephrolepis exalt D id ita como nueva de Arkansas
[ = [
Nephrolepis Schott, the Sword Fern genus, consists of 25-30 species, mainly
tropical in range, and is sometimes considered to be a small, monomorphic fam-
ily (Kramer 1990). The genus name is from the Greek, meaning “kidney” and
“scale,” referring to the shape of the indusium. Species of this genus have given
rise to many common cultivars that are excellent plants for growing on porches
or other shaded locations outdoors in southeastern United States (Benedict 1916;
Sessions 1978; Hoshizaki &@ Moran 2001). Four species and one hybrid are known
to occur outside of cultivation in North America north of Mexico (Nauman
Native to southern one-half of peninsular Florida, Nephrolepis exaltata
(L.) Schott has been extensively horticulturally selected and marketed as “Bos-
ton Fern”. It is widely cultivated outdoors in the South. The specific epithet is
from Latin, meaning “raised on high” or “lofty,” referring tothe perched site typi-
cally occupied in swamps. Its native range includes Florida, the West Indies
and in scattered locations in the Pacific Islands, probably introduced, escaping,
and becoming naturalized in many places. All populations in North America
beyond Florida originated as escapes from cultivation from spores or abandoned
plants. The species has escaped from cultivation and naturalized in peninsular
Florida (Nauman 1981; Nauman etal. 2000), Orleans Parish, Louisiana (Thieret
1980), and Highland Park, Texas (pers. comm., G. Gibbs). In Arkansas it isa com-
mon horticultural or cultivated summer plant in every county in the state, with
some winter-hardiness evident in the southernmost tier of counties.
Nephrolepis exaltata was first found in the wild in Arkansas on 25 Sep-
tember 2001 in natural woods east of Pine Bluff near the Arkansas River in Jef-
ferson Co. One plant well mired ina mucky soil was found with 20 sterile fronds,
obviously more than one year in age. The plant may have originated as the result
SIDA 20(2): 833-835. 2002
834 BRIT.ORG/SIDA 20(2)
of escaped spores or more likely from plants abandoned in some manner. The
park area accumulates flotsam and jetsam from the Arkansas River at high
water levels. Stranded debris, plastic cemetery decorations, and other fluvially
deposited materials attest to the efficacy of this mechanism. The plant will be
monitored for persistence, production of spores, and reproduction of new plants.
This is the sixth non-native fern of the Arkansas pteridophyte flora of 97.
The other exotics are Lygodium japonicum, Macrothelypteris torressiana,
Ophioglossum petiolatum, Psilotum nudum (in part), Pteris multifida, and
Salvinia minima. Only the Lygodium shows any ecological activity to be con-
sidered as an invasive species. Until this single plant Ne phrolepis exaltata dem-
onstrates reproductive competence, it will best be regarded asa waif. Additional
populations are expected from time to time in the southern part of state follow-
ing a few years with mild winters. (It seems unlikely that they would survive
an extremely hard winter.)
This first report for this genus and species in Arkansas is surprising late in
coming in that this and other horticultural-cultivated sword ferns are grown
extensively across the state. Apparently, they have not escaped or become rou-
tinely established mainly because of the prolonged heat and drought of late
summer in Arkansas.
The Arkansas vernacular name is “Yankee Fern,” reputed to be a gentle re-
taliation for the renaming of Dryopteris Xaustralis or Southern Log Fern by
some Connecticut Yankees who placed that hybrid into the horticultural trade
through tissue culture under the market name of Dixie Fern.
—
Voucher Specimen: ARKANSAS: Jefferson Co.: Wet riparian woods SE of Trulock Campground, off
access road to Atkins Lake-Nobel Lake along Arkansas River, 10 mi E of Pine Bluff, TOS R7W $5, Peck
2001388 (LRU)
REFERENCES
Benevict, R.C. 1916. The origin of new varieties of Nephrolepis by orthogenetic saltation. |.
Progressive variations. Bull. Torrey Bot. Club 43:207-234,.
Hosuizaki, B.J.and R.C. Moran 2001.Ferns grower’s manual. Revised and Expanded Ed. Tim-
ber Press, Portland, OR.
Kramer, K.U. 1990. Nephrolepidaceae. In: K.U. Kramer and PS. Green. Vol. 1, Pteridophytes
and gymnosperms.!n:K. Kubitski,ed. The families and genera of vascular plants. Springer-
Verlag, NY. Pp. 188-190.
Nauman, C.E. 1981.The genus Nephrolepis in Florida. Amer. Fern J. 71:35-40.
Nauman, C.E. 1993. Genus Nephrolepis Schott. Boston Fern. In: Flora of North America Edi-
torial Committee, Flora of North America north of Mexico.Oxford University Press, NY.
Vol. 2:305-308.
Nauman, C.E., R.P. WuNbeRLIN, and B.F. Hansen. 2000. Pteridophytes. In: Wunderlin, R.P. and B.F.
Hansen. Flora of Florida. Vol. 1: Pteridophytes and Gymnosperms. University Press of
Florida, Gainesville. Pp. 101-300.
PECK, NEPHROLEPIS EXALTATA NEW TO ARKANSAS 835
Sessions, K. 1978. Key to the cultivars of Nephrolepis exaltata ‘Bostoniensis’ Privately pub-
lished.
Tuieret, J.W. 1980. Louisiana ferns and fern allies. Lafayette Natural History Museum,
Lafayette, LA.
836 BRIT.ORG/SIDA 20(2)
BOOKS RECEIVED/NOTICES
Annotations and comments by Editor
Taxonomic/Systematic
Watter S. Jupp, St. Louis, PETER FE STEVENS, and MICHAEL J. DONOGHUE. 2002. Plant
Systematics: A Phylogenetic Approach, Second Edition. (ISBN 0-87893-403-
O, hbk.). Sinaurer Associates, Inc., 23 Plumtree Road, Sunderland, MA 01375-
0407, U.S.A. (Orders: 413-549-1118 fax; orders@sin« com, publisher
@sinauer.com). $86.95, 576 pp, numerous line drawings, 8 1/2" x 11"
From Sinauer website ( i om): “Plant Systematics: A Phylogenetic Approach, Second Edi-
tion is an introductory text that incorporates phylogenetic principles and methods throughout—
from the careful explanation of phylogenetic methods and principles in the initial two chapters to
he taxonomic survey of vascular plant families in the last two chapters.”
“The text is s accompanied ey a much expanded CD-ROM, containing over 2, ou colon pe
illustr ating the diz g f (an \ ariability W ithin) the vascular plant {
in the text, midudiner many images een floral and fruit dissections.”
Joun M. Gittett and Norman L. Taytor. MICHAEL COLLINs (Editor). 2001. World of
Clovers. (ISBN 0-8138-2986-0, hbk.; 0-8138-0282,-2, CD-ROM). lowa State
Press, 2121 State Avenue, Ames, IA 50014, U.S.A. (Orders: 1-800-862-6657,
1-515-292-3348 fax; www.iowastatepress.com). $144.95 (hbk & CD-ROM),
$99.95 (CD-ROM only), 488 pp, numerous b/w photos, 6" x 9",
From the back cover: “Descriptions are presented alphabetically by species name and enhanced by
beautiful color photographs of the plants and their seeds on the fully searchable CD-ROM (black
=
and-white photos in the book).” The book and CD-ROM include the following information for each
of the Nero species: Botanical name, Common name, Detailed c
jan
description of stipules, leaf-
lets, heads, flowers, and pods, Lifespan, Flowering period, Habitat, and Distribution. Clovers are in
the genus Trifoli um and belong to the Fabaceae (bean family).
Plant Physiology/Pathology/Agriculture
LINCOLN Taizand Epuarbo ZEFIGER, With contributors. 2002. Plant Physiology, Third
Edition. (ISBN 0-87893-82 3-0, hbk). Sinaurer Associates, Inc., 23 Plumtree
Road, Sunderland, MA 01375-0407, U.S.A. (Orders: 413-549-1118 fax:
orders@sinauer.com, publisher@sinauer.com). $104.95, 690 pp, illustrated,
81/2" x 11".
From Sinauer website (wwwsinauercom): “.. the new edition incorporates all the important new de-
velopments in plant physiology, especially in cell, molecular, and developmental biology.”
SIDA 20(2): 836. 2002
CYNOSURUS ECHINATUS (POACEAE) NEW TO TEXAS
R. Dale Thomas
Department of Biology
University of Louisiana at Monroe
Monroe, LA 71209, U.S.A.
Cynosurus echinatus L.—TEXAS. Walker Co.: Ebenezer Cemetery beside Hwy
30, | mi E of Farm Rd 2550, W of Huntsville, 30 May 1992, R.D. Thomas 129,383
(BRIT, NLU).
The species has been reported from Oklahoma, Louisiana, Missouri, Arkan-
sas, and many states of the eastern United States (Kartesz 1999). It also occurs
in the Pacific coast states of California, Oregon, and Washington, and in British
Columbia. Its closest known locality tothe Walker County, Texas, site isin Caddo
Parish, Louisiana (Thomas & Allen 1993). No plants of this species were relo-
cated in a search of the Walker County site in June 1997 (pers. comm., Guy
Nesom).
Cynosurus echinatus is native to southern Europe (Tutin 1980) and Africa,
and Asia (Yatskievych 1999), but it is now established in North America, South
America, and Australia. It is annual in duration and a colonizer of open dis-
turbed sites, including fields, roadsides, and railways. Among various common
names are “Hedgehog Dogtail,” “Rough Dog's Tail,” and “Bristly Dog’s Tail Grass.”
A photograph is posted on the internet at <http://plants.montara.com /
ListPages/FamPages/ Poa3.html#cynech>. Detailed line drawings are found at
<http://biodiversity.uno.edu/ delta/grass/www/cynosuru.htm>.
REFERENCES
Kartesz, J.T. 1999. A synonymized checklist and atlas with biological attributes for the
vascular flora of the United States, Canada, and Greenland. First edition. In: Kartesz, J.T.,
and C.A.Meacham. 1999. Synthesis of the North American flora, Version 1.0.North Caro-
lina Botanical Garden, Chapel Hill, North Carolina.
THomas, R.D. and C.M. Aten. 1993. Atlas of the vascular flora of Louisiana. Volume |. Louisi-
ana Natural Heritage Program & The Nature Conservancy.
Y ATSKIEVYCH, G. 1999. Steyermark’s flora of Missouri.Vol.1 (rev.ed.). Missouri Dept. of Conser-
vation, Jefferson City, in cooperation with Missouri Botanical Garden Press, St. Louis.
Tutin, T.G. 1980. Cynosurus. Flora Europaea 5:171-172.
SIDA 20(2): 837. 2002
838 BRIT.ORG/SIDA 20(2)
BOOKS RECEIVED/NOTICES
Annotations and comments by Editor
Plant Physiology/Pathology/Agriculture
ZDENKO RENGEL (Editor). 2002. Handbook of Plant Growth pH as the Master Variable.
(ISBN 0-8247-0761-3, hbk.). Marcel Dekker, Inc., 270 Madison Avenue, New
York, NY 1LOO16, U.S.A. (Orders: 212-696-9000, 212-685-4540 fax,
www.dekkercom). Marcel Dekker AG, Hutgasse 4, Postfach 812, CH-4001
Basel, Switzerland (Orders: 41-61-261-8482, 41-61-261-8896 fax) $175.00,
+70 pp, 0 x,
From Marcel Dekker website GQwww.dekkercom): “Thi biochemical
functional, structural, and developmental mechanisms of pH in nak ant growth, as well as examining
the role of pH in plant symplasm, plant app ism, the nee here, the ecosystem, and in plant inter-
action with biotic and abiotic environments.
ALLEN R.OveRMAN and RicHarD V. ScHortz IIL. 2002. Mathematical Models of Crop
Growth and Yield. (ISBN 0-8247-0825-3, hbk.). Marcel Dekker, Inc., 270
Madison Avenue, New York, NY 10016, U.S.A. (Orders: 212-696-9000, 212-
685-4540 fax, www.dekker.com). Marcel Dekker AG, Hutgasse 4, Postfach
812, CH-4001 Basel, Switzerland (Orders: 41-61-261-8482, 41-61-261-8896
fax) $159.00, 328 pp, illustrated, 6" 9"
From Marcel Dekker website (www.dekkercom): “This insightful ref / lescril
tion of viable mathematical models in data anal i se crop growth and yields— boblighiee
ing effective, analytical functions that have been found useful lor the comparison of alternative man-
agement techniques to maximize water dnd nutrient resources
SAMUEL S. GNANAMANICKAM (Editor). 2002. Biological Control of Crop Diseases.
(ISBN 0-8247-0093-5, hbk.). Marcel Dekker, Inc., 270 Madison Avenue, New
York, NY 10016, U.S.A. (Orders: 212-696-9000, 212-685-4540 fax,
wwwdekkercom). Marcel Dekker AG, Hutgasse 4, Postfach 812, CH-4001
Basel, Switzerland (Orders: 41-61-261-8482, 41-61-261-8896 fax) $175.00,
468 pp, illustrated, 6" x 9".
From Marcel Dekker website ( lek}
). “This innovative reference combines theory with
current global practices involved in the biological control of diseases in 12 major crops—highlight
ing the day-to-day challenges of organic crop management for cost-elfective real-world application.”
SIDA 20(2): 838. 2002
ADDITIONS TO THE FLORA OF NAGS HEAD WOODS
(DARE COUNTY, NORTH CAROLINA)
AND THE OUTER BANKS OF NORTH CAROLINA
Alexander Krings
Herbarium, Department of Botany
North Carolina State University
Raleigh, NC 27695-7612
Alexander_Krings@ncsu.edu
ABSTRACT
Including an extensive system of dunes, interdunal ponds, and wetlands, the Nags Head Woods Eco-
logical Preserve Dae om North Carolina) comprises one of the best remaining examples of mid-
Atlantic t s forests. This study presents additions to the flora of the preserve and
the larger Outer Banks, based on recent collections. Thirty-four taxa have not previously been reported
for the preserve, five have not been reported for the Outer Banks, and two have been rediscovered.
RESUMEN
La Nags Head Woods Ecological Preserve (Dare Co., North Carolina), que incluye un sistema extensivo
de dunas, lagunas interdunares, y humedales, comprende uno de los mejores ejemplos restantes de
bosques caducifolios maritimos sulgavicos Este estudio presenta adiciones a a ore aie a reserva y los
mayores Outer Banks, basado en colecciones recientes. Treinta
viamente de la reserva, cinco no habian sido citados de los Gis Banks, y dos han sido redescubiertos.
The Nags Head Woods Ecological Preserve (Kill Devil Hills, Dare Co.) comprises
one of the best remaining examples of mid-Atlantic maritime deciduous forest
(Schafale @ Weakley 1990) and includes an extensive system of dunes,
interdunal ponds, and wetlands. Recent progress toward a flora of the preserve
provided a checklist of vouchered collections housed at the herbarium of the
National Park Service Cape Hatteras National Seashore Unit (CAHA), Manteo,
North Carolina (Krings 2002). Although not listed in either the Index
Herbariorum (Holmgren et al. 1990) or Aulbach-Smith et al. (1986), CAHA con-
stitutes a valuable resource of North Carolina barrier island collections. The
collections from the Nags Head Woods Ecological Preserve (NHWEP) housed
at CAHA include material collected by various members of the Dunes of Dare
Garden Club, McCaffrey, as well as Stalter and Lamont (1997). This study pre-
sents new additions to the flora of the preserve and the larger Outer Banks (sensu
Stalter @ Lamont 1997), resulting from recent collections by the author and
preserve staff.
METHODS
The list was compiled based on field collections by the author and preserve
SIDA 20(2): 839-843. 2002
840 BRIT.ORG/SIDA 20(2)
staff over the period of Mar 2001-Sept 2002. Specimens were critically studied
and identified at the herbarium of North Carolina State University (NCSC).
RESULTS
Species at the NHWEP listed below include new additions to the flora, as well
as species previously reported but not vouchered by Blonder and Stalter (1992)
or Stalterand Lamont (1997). Asterisks preceding a scientific name indicate that
taxon to be a new addition to the flora of NHWEP (.e,, not previously reported
by Blonder and Stalter (1992) or not specifically reported for the preserve by
Stalter and Lamont (1997)).
The list includes 89 species, 34 (38%) of which have not been previously
reported for the preserve, and 5 (~6%) of which have not been reported for the
Outer Banks (ie., not listed in either Burk (1961), Blonder and Stalter (1992), or
Stalter and Lamont (1997)).
Pteridophyta
ASPLENIACEAE
*Asplenium platyneuron (L.) BSP., Krings 450
ee
Krings 534 (NCSC)
sen nde egal i Var. teenie (Willd.) A.Gray,
Krings 538 (NCSC)
Gy mnospermae
CUPRESSACEAE
Juniperus virginiana L., Krings 373 (NCSC)
PINACEAE
Pinus palustris Mill, Krings 563 (NCSC)
Pinus taeda L., Krings 564 (NCSC)
TAXODIACEAE
(L.) Rich., McCall s.n. (NCSC)
Angiospermae
Monocotyledonae
CYPERACEAE
“Carex albolutescens Schwein., Krings 554 (NCSC):
Krings 555 (NCSC)
Sc oe tus americanus (Pers.) Valk ex
Keller, Krings 546 (NCSC)
JUNCACEAE
Juncus canadensis J. Gay, Krings 556 (NCSC);
Krings 560 (NCSC)
Scheele, Krings 540
Schinz.
lunes roemerianus
NCSC)
pa
LILIACEAE
*Hemerocallis fulva (L.) Le Krings 45] (NCSC)
ORCHIDACEAE
Cypripedium acaule Aiton, Krings 471 (NCSC)
POACEAE
Arundinaria tecta Walt., Krings 541 (NCSC)
“Erianthus giganteus (Walt.) Muhl., Krings 559
(NCSC)
Glyceria septentrionalis Hitchc,, Krings 561 (NCSC
Panicum amarum Elliott, Krings 558 (NCSC)
“Panicum scoparium Lam., Krings 557 (NCSC)
Phragmites australis (Cav.) Trin. ex Steud., Krings
ae
*Setaria geniculata P. Beauv., Krings 521 (NCSC)
*Setaria magna Griseb., Krings 544 (NCSC)
SMILACACEAE
Smilax glauca Walter, Krings 465 (NCSC);
(NCSC)
Krings
487 (NCSC
ii Morong, Krings 525 (NCSC)
*Smilax smal
TYPHACEAE
lypha angustifolia L., Krings 549 (NCSC)
Dicotyledonae
ACERACEAE
Acer rubrum L., Krings 377 (NCSC)
ANACARDIACEAE
Rhus copallina L., Krings 514 (NCSC)
APIACEAE
Centella erecta (Lf) Fern. Krings 562 (NCSC)
KRINGS, FLUKA UP
*Ptilimnium capillaceum (Michx.) Raf. ex Ser.,
rings 533 (NCSC)
AQUIFOLIACEAE
llex vomitoria Aiton, Krings 386 (NCSC)
ARALIACEAE
Aralia spinosa L., Krings 463 (NCSC)
ARISTOLOCHIACEAE
“Aristolochia serpentaria L., Krings 517
ASTERACEAE
*Achillea millefolium L., Krings 543 (NCSC)
Carduus spinosissimus Walter, Krings 466 (NCSC)
*Chrysanthemum leucanthemum L., Krings 452
(NCSC)
Eupatorium hyssopifolium L., Krings 522 (NCSC)
*Gaillardia pulchella Foug., Krings 456 (NCSC)
Iva frutescens L., Krings 551 (NCSC)
BETULACEAE
Ostrya virginiana (P. Mill.) K. Koch., Krings 374
(NCSC)
BRASSICACEAE
*Cardamine hirsuta L., Krings 389 (NCSC)
Lepidium virginicum less Krings 530 (NCSC)
BIGNONIACEAE
Campsis radicans (L.) Seem., Krings 523 (NCSC)
CAPRIFOLIACEAE
Lonicera japonica Thunb. ex Murray, Krings 472
SC)
Lonicera sempervirens L., Krings 535 (NCSC)
CELASTRACEAE
*Celastrus orbiculatus Thunb., Krings 532 (NCSC
CISTACEAE
*Helianthemum canadense (L.) Michx., Krings 550
(NCSC)
CONVOLVULACEAE
Calystegia sepium (L.) R.Br., Krings 527 (NCSC)
CORNACEAE
*Cornus stricta Lam., Krings 536 (NCSC)
Cornus florida L., Krings 486 (NCSC)
EUPHORBIACEAE
Cnidoscolus stimulosus (Michx.) Engelm.& A.Gray,
Krings NCSC)
*Tragia urens L., Krings 515 (NCSC)
FABACEAE
*Desmodium nuttallii
(NCSC)
=
~~
(Schindler) Schubert,
NORTH CAROLINA) 841
Krings 512 (NCSC)
*Pueraria lobata (Willd.) Ohwi — var.
esen, Krings 531 (NCSC)
(Elliott) DC., Krings 509
mo a(Lour.) Ma
*Rhynchosia difformis
(NCSC)
*Trifolium arvense L., Krings 458 (NCSC)
*Wisteria sinensis (Sims) Sweet, R. Paris s.n. (NCSC)
AGACEAE
Quercus falcata Michx., Krings 537 (NCSC)
*Quercus hemisphaerica W.Bartr.ex Willd. Krings
518 (NCSC)
Quercus stellata Wangenh., Krings 516 (NCSC)
GERANIACEAE.
HAMAMELIDACEAE
Liquidambar styraciflua L., Krings 464
cident
‘al
INAdeEN
VL. Krings 477 (NCSC)
—
NCSC)
L., Krings 526 (NCSC)
LAURACEAE
Sassafras albidum (Nutt.
LOGANIACEAE
Gelsemium sempervirens St. Hilaire, Krings 369
ae
Nees, Krings 380 (NCSC)
MAGNOLIACEAE
Magnolia grandiflora L., Krings 485 (NCSC)
MORACEAE
Morus rubra L., Krings 475 (NCSC)
MYRICACEAE
Myrica cerifera L., Krings 387
ONAGRACEAE
*Oenothera laciniata Hill, Krings 455 (NCSC);
Krings 479 (N
OXALIDACEAE
Oxalis dillenii Jacq., Krings 478 (NCSC)
PASSIFLORACEAE
*Passiflora lutea L., Krings 529 (NCSC)
PLANTAGINACEAE
*Plantago hookeriana Fisch.& C.A.Mey.var.nuda
(A. Gray) Poe, Krings 454 (NCSC)
*Plantago psyllium L., Krings 457 (NCSC)
POLYGONACEAE
Polygonum hydropi hes Michx., Krings 552
NCSC); Krings 553 (NCSC)
—>
NCSC)
S
842 BRIT.ORG/SIDA 20(2)
PYROLACEAE SAURURACEAE
Chimaphila maculata (L.) Pursh, Krings 372. Saururus cernuus L., Krings 405 (NCSC)
iNCeO) SCROPHULARIACEAE
*Bacopa monnieri (L.) Pennell, Krings 528 (NCSC)
Amelanchier canadensis (L.) Medic. Krings 375 *Linaria canadensis (L.) Dum. Cours., McCall s.n.
(NCSC); Krings 381 (NCSC) (NCSC)
Rosa eee ona a 406 (NCSC) SOLANACEAE
Rubus trivialis Michx,, Krings 476 (NCSC); R. Paris *Physalis walteri Nutt., Krings 524 (NCSC)
5.n. (NCSC); Krings - ee
VERBENACEAE
UBIACEAE Callic ricana L., Krings 411 (NCSC)
SU apolay is L., Krings 410 (NCSC)
ium hispidulum Michx., Krings 370 (NCSC) VIOLACEAE
“Gali ium Sie eecos Krings 519 (NCSC) *Viola sororia Willd., Paris 5.n. (NCSC)
Mitchella repens L., Krings 484 (NCSC) VITACEAE
SALICACEAE Ampelopsis arborea (L.) Koehne, Krings 407
Salix caroliniana Michx., Krings 539 (NCSC) (NCSC)
Parthenocissus quinquefolia (L.) Planch., Krings
468 (NCSC)
DISCUSSION
Table | lists species apparently discovered for the first time from the Outer Banks
or rediscovered after not having been seen by Stalter and Lamont (1997). The
find of Aristolochia serpentaria (Aristolochiaceae) is rather interesting, al-
though not altogether surprising as it is generally associated with mixed de-
ciduous forests (Radford et al. 1968). As few examples of maritime deciduous
forests remain in North Carolina, the species likely remains rare due to habitat
availability on the Outer Banks. Although relatively common in the North Caro-
lina Coastal Plain, Tragia urens has been previously reported for the Outer Banks
only by Burk (1961). Stalter and Lamont (1997) did not mention the taxon and
presumably did not observe it. It remains unknown whether the taxon persists
on Bogue Banks as reported by Burk (1961). Celastrus orbiculatus and Pueraria
lobata var. montana have been weeds of concern to the preserve staff and ef-
forts are underway to severely control, if not eradicate, them from the site.
ACKNOWLEDGMENTS
I thank the Nature Conservancy for access to the preserve and Aaron McCall,
Rhana Paris, and Jeff DeBlieu for thoughtful conversations and collection help.
REFERENCES
AULBACH-Smit, C., J. Massey, and J. MartHews. 1986. Guide to herbarium resources in the Caro-
linas. Friends of the UNC Herbarium 1:1-38.
Bionper, B.and R. Statter. 1992. Vascular plant species list: Nags Head Woods Preserve. The
Nature Conservancy, Kill Devil Hills, NC.
KRINGS, FLORA OF NORTH CAROLINA) 843
Taste 1. Newly discovered or rediscovered taxa for the Outer Banks.
Taxa Burk (1961) Stalter & Lamont (1997) Present study
Aristolochiaceae
Aristolochia serpentaria Not listed Not listed Vouchered
from NHWEP
Celastraceae
Celastrus orbiculatus Not listed Not listed Vouchered
from NHWEP
Euphorbiaceae
Tragia urens Reported from Not listed Vouchered
Bogue Banks from NHWEP
Fabaceae
Desmodium nuttallii Not listed Not listed Vouchered
fromm NHWEP
Pueraria lobata var. montana Reported from Listed as not Vouchered
Nagshead recently observed from NHWEP
Plantaginaceae
Plantago hookeriana var.nuda Not listed Not listed Vouchered
from NHWEP
Smilacaceae
Smilax smallii Not listed Not listed Vouchered
from NHWEP
Burk, C.J. 1961.A floristic study of the Outer Banks of North Carolina. Ph.D. dissertation,
University of North Carolina, Chapel Hill, NC.
Hotmoren, P.K., N.H. HoumoGren, and L.C. Barnett. 1990. Index herbariorum. Part |: The herbaria
of the world. 8th edition. NY Botanical Garden, New York, NY.
Kartesz, J.7.1994.A synonymized checklist of the vascular flora of the United States, Canada,
and Greenland. Timber Press, Portland, OR.
Krincs, A. 2002. The Nags Head Woods Collections of the National Park Service Cape
Hatteras National Seashore Unit Herbarium (CAHA).J.North Carolina Acad. Sci. 118:145-
ScHaFALe, M.P. and A.S. Weaktey. 1990. Classification of the natural communities of North
Carolina, third approximation. NC Natural Heritage Program, Raleigh, NC.
Stacter, R.and E.E. Lamont. 1997. Flora of North Carolina’s Outer Banks, Ocracoke Island to
Virginia. J. Torrey Bot. Soc. 124:71-88.
BRIT.ORG/SIDA 20(2)
844
BOOKS RECEIVED/NOTICES
Annotations and comments by Editor
Plant Physiology/Pathology/Agriculture
CHRISTOPHER P. DUNN (Editor). 2000. The Elms Breeding, Conservation, and Dis-
ease Management. (ISBN 0-7923-7724-9, hbk.). Kluwer Academic Publishers,
Customer Service Department, P.O. Box 358, Accord Station, Hingham, MA
02018-0358, U.S.A. (Orders: 781-871-6600, Toll Free 866-269-9527, fax 781-
681-9045, kluwer@wkap.com). $125.00, 361 pp, illustrated, 6" x 9"
lc ultivated, throughout much
From Kluwer website ale nl: “Elms occur, both naturally anc
W ide-
of the reli perate world. Because of their high tolerance g §
] Sauer J ] }
spread d distr ibution, elms have been W idely I
America and northern Europe. As such, their current demise due to several pandemics of Dutch elm
disease resistance, conservation and
disease has spurred a huge body of research on breeding for
systematics. The Elms: Breeding, Conservation and Disease Management provides the current state
of knowledge in these areas and is an important reference work for pathologists, breeders, taxono-
mists and arborists
GERALD C. Nrison (Editor). 2001. Genetically Modified Organisms in Agriculture,
Economics and Politics. (ISBN 0-12-515422-4, hbk.). Academic Press, 200
Wheeler Road, Burlington, MA 01803, U.S.A. (Orders: 888-677-7357, 781-
Pelee 72 ak (Gl-77 1615, http:/ wwwacademic press.com www/ap/
reps.htm). £46.95, 300 pp, illustrated, 6 1/2" x 91/2".
fa qT] AA ] fi ]() ; {f{
From Academic website(www.apcatalog.com): “In Part 1,
aaa look at the science, economics, and politics a the use c agric ultaral eM0s It provides
f the three most- ig) used GMOs:
resistant
pact on
ne
import/export patterns and the im
soe ans. Issues such as US and EU regule COTY concerns,
res with widely different
ennai trade are covered. Part 2 presents articles from leading figu
views in the debate, such as representatives from the American Corn Growers a, ation, the EU
Com chore The Foundation for Science, Technology and Natural Resource Policy, India, Center for
the Foeasena Industry Organization. Finally, Part 3
aistory of biotechnology inno-
Global Food iSSES, Consumers U nion soa
provides more topics, including the
vations, the acai: of lacs the latest research on the consequences of the Bt corn for
the monarch butterfly, and a list of European GMO field trial approvals.
CIRSIUM NUTTALLII (ASTERACEAE: CYNAREAE)
NEW TO NORTH CAROLINA AND
AN ILLUSTRATED KEY TO SOUTHEASTERN CONGENERS
Alexander Krings Randy Westbrooks
Herbarium, Department of Botany US Geological Survey
North Carolina State University 233 Border Belt Drive
Raleigh, NC 27695-7612, U.S.A. Whiteville, NC 28472, U.S.A.
Alexander_Krings@ncsu.edu
Janine Lloyd
US Geological Survey
233 Border Belt Drive
Whiteville NC 28472, U.S.A.
ABSTRACT
Cirsium nuttallii (Asteraceae) is documented for North Carolina. The species had previously been
known from Florida to South Carolina and from disjunct populations in Virginia. An illustrated key
is provided to aid others in the diagnosis of Cirsium in North Carolina and the southeast
RESUMEN
Se documenta Cirsium nuttallii (Asteraceae) para el estado de Carolina del Norte. Previamente se
2
conocia la alae oe — Florida a Carolina del Sur y de poblaciones disyuntas en Virginia. Se
enta a para ayudar los demas en las ae oe de Cirsium en el sudeste.
A southeastern United States endemic, Cirsium nuttallii DC. (Asteraceae:
Cynareae) has been previously known only from peninsular Florida to Louisi-
ana and South Carolina, and from apparently disjunct populations in south-
eastern Virginia (Radford et al. 1968, Cronquist 1980). However, recently Randy
Westbrooks found an individual in a pasture in Whiteville, Columbus Co.,, North
Carolina. When surveyed, the plant was about 2.4 m tall. Apparently, it had not
been seen in the pasture previously. Additional survey by Robert Eplee located
a population lightly scattered over 10 acres, north of Hallsboro (also Columbus
Co.). The species is known from at least 16 counties in South Carolina, includ-
ing nearby Marion Co. North Carolina now includes all 12 species of native and
naturalized Cirsium known from the southeast. Voucher specimens of C.
nuttallii were prepared for deposit at NCSC, OBI, and US.
Voucher specimen: U.S.A. NORTH CAROLINA. Columbus Co.: Whiteville, in a pasture, roughly 8 ft
tall, 24 Jun 2002, R. West brooks s.n. (NCSC, OBI
Cirsium nuttallii DC. (Fig. 1H) is one of the tallest Cirsium species in the south-
east, capable of attaining heights of 3.5 m (Cronquist 1980). Only Cirsium
altissimum (L.) Sprengel and C. discolor (Muhl.) Sprengel are known to grow to
SIDA 20(2): 845-848. 2002
846 BRIT.ORG/SIDA 20(2)
a similar maximum size (Cronquist 1980). These latter two can be distinguished
from C. nuttallii by the abaxial leaf surface densely and persistently white to-
mentose (arachnoid-tomentose to glabrate in age in C. nuttallii).
Cirsium nuttallii can be distinguished from its southeastern congeners by
the combination of the following traits: (1) plant biennial, 1.5-3.5 m tall, (2) stems
not winged, but branched and many-headed, (3) abaxial leaf surface arachnoid-
tomentose to glabrate in age, (4) involucres 1.5-2.5 cm long, and (5) middle and
outer involucral bracts with a glutinous adaxial ridge and tipped by spines |-
2(-3) mm long. We are providing a key, largely adapted from Cronquist (1980),
as well as illustrations in hopes they might prove of diagnostic use to others.
KEY TO CIRSIUM IN THE SOUTHEASTERN UNITED STATES
1. Stems saa winged by spiny, decurrent leaf bases (Fig. 1A, i), the wings
Ci
nearly or as long as the internodes irsium vulgare (Savi) Ten. (Fig. 1A, ii)
. Stems not ee by decurrent leaf bas
2. Heads conspicuously and Closely pended by a series of narrow, reduced,
=
strongly spiny leaves (Fig. 1 Cirsium horridulum Michx.
2. Heads not closely subtended by a series of reduced spiny leaves.
3. Plant colonial ie (by creeping roots); ee ‘ia the heads
nearly unisexual with the pappus of the pistillate head longer than the corre
sponding corollas and the apne s of the staminate heads shorter than the
corresponding corollas (Fig. 1D, i & ii) Cirsium arvense (L.) Scop.
3. Plant not colonial, biennial or perennial; heads all similar and perfect.
4, |Involucral bracts completely lacking spiny tips or only exhibiting a vesti-
gial spinule to 0.5 mm long (Fig. 1 Cirsium muticum Michx.
4. Involucral bracts b bearing spiny tips, the spines typically at least 1 mm lon
5. Abaxial leaf surface densely and persistently white tomentose.
6. Plants robust, 1—3(—4) m tall; largest lobed leaves generally > 5 cm
wide; involucres generally 2.5—3.5 cm long
7. Leaves deeply pinnatifid (Fig. 1F,i) _ sd Cirsium discolor (Muhl. ex os
Sprengel (Fig. 1F, ii)
)
K)
a.
—
7. Leaves toothed or shallowly lobed Cirsium altissimum (L
Sprengel (Fig. 1
6. Plants relatively slender, 0.5—1.5 m tall; Aer lobed leaves generally
wide; involucres generally 1.5-2.5
8. Cauline leaves mostly 30-70; Sune cere in Aug-O
savannahs, bogs, and wet pinelands Cirsium virginianum (L.)
Michx. (Fig. 1)
8. Cauline leaves mostly 10-25; plants flowering Apr—Jun; open
woods and dry, sandy soil Cirsium carolinianum (Walt.)
Fern. & Schub. (Fig. 1G)
5, Abaxial leaf surface arachnoid-villous (sometime sparsely so) to glabrate
parieviany in
9. Plants 1.5-3. : m tall, branched and many-headed; involucres 1.5-2.5
cm cag fic, 1H) Cirsium nuttallii D
9. Plants 0.2—1 m tall, unbranched or only sparingly branched; involu-
cres 2.5-5 cm long.
10. Young stems and abaxial leaf surfaces thinly and loosely white
YVDIN
Fic.1.5
\°A Grom vulgare
are . if
Oh\-D fF;
1 (Ushi.
inal (Vet 36, Cirsium Eel pistillate flower, note |
);E,
ce leaves; ii, involucre (Wilbur 4856); 16,0 Ci
involucre, note dorsal, el
h
; yD
g
1s.n.):C, Cirsium horridul , pr :
ii, st fl note pappus
g Pap} Papf
O CAGETAANVCELTISE TT EeaT .
y -F, Cirsium discolor — i, deep!
Pee
I /
iva
8): H. Cirsium nuttallii
Pratt ype
Cirsium repandum liwalucre (Fox 1693): K, Cirsi
cts (R. Westbrookss.n.):1, Ci
. . * 1 {Ali -kwl
(Fox 3687). All specimens from NCSC.
BRIT.ORG/SIDA 20(2)
tomentose, older stems and abaxial leaf surfaces glabrate; heads
strongly pedunculate Cirsium lecontei Torr. & A. Gray (Fig. 1!)
10. Young stems and abaxial leaf surfaces Sa villous, hirsute
or rarely arachnoid tomentose; heads scarcely or more or less
a pedunculate
_ Cauline leaves crowded, the internodes frequently < 1 cm
long; inner involucral bracts gradually tapering toa ee
Poets tip (Fig. 1J) Cirsium repandum Michx.
. Cauline leaves typically not crowded rae sometimes
so near the base); inner involucral bracts with expanded,
chartaceous, crisped, and erose tips (Fig. 1L)
Cirsium pumilum
(Nutt.) Spreng.
REFERENCES
Cronauist, A. 1980. Vascular flora of the southeastern United States: Asteraceae. University
of North Carolina Press, Chapel Hill.
Raprorb, A.E., H.E. AHLes, and C.R. Beit. 1968. Manual of the vascular flora of the Carolinas
University of North Carolina Press, Chapel Hill.
REINSTATEMENT OF VIBURNUM OZARKENSE
(CAPRIFOLIACEAE): AN ENDEMIC TAXON OF THE
INTERIOR HIGHLANDS OF
ARKANSAS, MISSOURI AND OKLAHOMA
Timothy J.Weckman
Department of Biological Sciences
astern I entucky University
Richmond, KY 40475, U.S.A.
ABSTRACT
Based on field work and phological analysis of herbarium specimens, Viburnum ozarkense Ashe
d distinct species. Recognition of the occurrence and distribution of V. ozarkense has
been eine by its Teduetion to synonymy under Game molle Michx. A se pesees
characterization of V.ozarkense and V. molle, a key to species, and a distribution map are pro
This research supports the recognition of V. ozarkense as an endemic shrub of the Interior an
lands of Arkansas, Missouri and Oklahoma.
RESUMEN
Rp a . | a ae ok SP A
Viburnumozarkense
gone se repone ¢ como una especie distinta. Elr eesgocimients de la ocurrencia y distribucion de V.
le Viburnum molle Michx. Se ofrecen
una caracterizacion monolegica de V.ozarkense y V. molle, una clave de especies, y un mapa de
distribucion. Esta investigacion apoya el reconocimiento de V.ozarkense como un arbusto endémico
de las montanas Ouachita y Ozark de Arkansas, Missouri y Oklahoma
INTRODUCTION
Viburnumozarkense was described from material W. W. Ashe collected in Stone
County, Arkansas with the type locality providing the only distributional ref-
erence (Ashe 1928). Ashe was a prolific collector, amassing some 30,000 speci-
mens by his death (Massey 2001). Ashe also published 510 species and varietal
names from these collections, but many have been reduced to synonymy (Coker
et al. 1932). Viburnum molle was described by Michaux from material he col-
lected in Kentucky in the late 1700s (Michaux 1803). Ashe was aware of V. molle
Michx., and provided characteristics to separate V. ozarkense from V. molle in
his species description. Original material from both of these collections has not
been readily accessible to American botanists: the Michaux collection of V. molle
was deposited in the Paris herbarium (P)(acronyms after Holmgren et al. 1990),
and most of Ashe’s collections were inaccessible in his private herbaria at the
time of his death (Coker et al. 1932).
The name V. ozarkense was retained in an early checklist for Arkansas
(Moore 1941) and in a generic treatment of Viburnum by McAtee (1956). How-
SIDA A20(2): 849- 860, 2002
850 BRIT.ORG/SIDA 20(2)
ever, it was reduced to synonymy under V. molle Michx. by Steyermark (1963)
and this placement has been followed by current treatments (Tucker 1976; Smith
1988; USDA, NRCS 2002). Neither taxon is currently monitored by heritage
programs in Arkansas, Missouri or Oklahoma. Based on morphological analy-
sis of Viburnum ozarkense and V. molle, lrecognize these taxa as distinct, pro-
vide a key to differentiate them, and map the current distribution of both taxa
in Arkansas, Missouri and Oklahoma.
MATERIALS AND METHODS
Loans were requested from herbaria with Ashe collections: A, GH, ILL, L, LCU,
MO, NCU, POM, and US, and certain herbaria having Arkansas, Missouri and
Oklahoma Viburnum material: APCR, BRIT, NA, NYBG, OKL, SMS, UAM,
UARK, and UMO. Additional specimens were collected during field work in
Arkansas, Missouri and Oklahoma in 1999-2001.
Measurements were recorded for 14 quantitative characteristics: leaf size,
toothing, venation, length of leaf tip, petiole length on fertile and vegetative
shoots, peduncle length, number of rays, inflorescence width and height, co-
rolla width, stamen length and fruit length and width. Qualitative characteris-
tics including leaf shape, leaf and petiole pubescence, the presence, placement
and pubescence on stipules and inflorescence bracts, flowering times and shoot
pubescence and color were noted. Mensural and qualitative data were used to
prepare a diagnostic key for the species.
Habitat and associate information was gleaned from vouchers and from
field work. A distribution map of all known populations of V.ozarkense and V.
molle occurring in Arkansas, Missouri and Oklahoma was prepared from an-
notated loans and new collections.
—_—
RESULTS AND DISCUSSION
Loans resulted in 236 vouchers for study. An additional 119 collections were
produced from field work during 1999-2001 in Arkansas, Missouri and Okla-
homa. Of these, 193 were referable to Viburnum ozarkense and 162 to V. molle.
They are all listed in the specimens examined section for each taxon.
Mensural results
Mensural analysis indicates that the most useful features for differentiating
Viburnumozarkense from V. molle are leaf width, number of teeth per half leaf,
number of veins per half leaf, leaf tip length, petiole length below an inflores-
cence or at the apex of a vegetative shoot, corolla width and fruit length. The
means of these characters were all statistically significantly different (Table 1).
Measured features not significantly different between the species were leaf
length, peduncle length, inflorescence width and height, stamen length and
fruit width.
WECKMAN, REINSTATEMENT OF VIBURNUM OZARKENSE 851
Based on my measurements leaves in Viburnum ozarkense averaged 7.2
cm wide, with 5.3 veins and 13.9 teeth per half leaf, and had an acuminate apex
averaging 7.7 mm long. Leaves of V. molle were wider, averaging 8.2 cm broad,
with 6.1 veins; more toothed, averaging 25.6 teeth per half leaf;and with an acute
apex averaging 2.9 mm long. Viburnum ozarkense petioles on both flowering
and sterile shoots are shorter than those of V. molle. The first petioles below an
inflorescence averaged 1.lcm in V.ozarkense and 2.4 cm for V. molle. Petioles on
the distal ends of vegetative shoots are similarly shorter on V.ozarkense shoots
(x= 2.1 cm) than V. molle shoots (x= 3.4 cm). Corollas were significantly wider
in V.ozarkense flowers (x= 6.6 mm) than in V. molle flowers (x= 4.4 mm), but
these data were derived from a small sample (n= 40). Mature fruits were signifi-
cantly longer in V ozarkense (x= 1.1cm) than in V. molle (x= 0.9 cm), but again,
sample size was small (n= 41). Vegetative specimens of these taxa are differenti-
ated based on the number of teeth per half leaf. This characteristic overlaps in
only a few individuals with distinct peaks well separated (Fig. 1).
Qualitative characteristics
Qualitative features such as leaf shape, leaf and petiole pubescence, color of
current year shoots and bark features also provided useful characteristics to
separate the species (Table Ll). In outline, Viburnum molle leaves are broadly
ovate or orbicular; those of V. ozarkense are narrower and appear ovate in out-
line. Adaxial leaf surfaces of V.ozarkense are uniformly hirsute, with short hairs
(ca. 0.25 mm long) across the surface. In contrast, adaxial surfaces of V. molle
leaves are glabrous or may have a few scattered red, glandular hairs. Similarly,
petioles of V.ozarkense are pubescent, covered with moderate to dense strigose
and glandular hairs and an open furrow. Petioles of V. molle are essentially gla-
brous and any observed pubescence is typically manifested as a few glandular
hairs at the distal end of the petiole or asa few villous hairs along the margin of
the closed petiole furrow. Stipules are present in both species; in V. molle these
occur in single pairs, while in V. ozarkanse two pairs of stipules per leaf stalk
are often observed. In cross section, V. ozarkense petioles appear oval while V.
molle petioles appear terete. The color of current year shoots at mid-season also
provides a useful way to differentiate these taxa. For V.ozarkense they are red-
dish-brown, in contrast to the yellow-tan shoots of V. molle. The most obvious
field character separating these viburnums is the presence of exfoliating bark
persistent on stems after the second year of growth on V. molle shoots verses
the tight bark throughout the branching of V. ozarkense. Although this is a
prominent field character, it is not consistently present on herbarium sheets, as
many collections do not show enough branch to capture it.
DIAGNOSTIC KEY
Leaves ovate, adaxially hirsute, 8 to 21 (mode 14) marginal teeth per half leaf; petioles
of first pair of leaves below inflorescence X= 1.0 cm, X= 2.0 cm at tips of vegeta-
852 BRIT.ORG/SIDA 20(2)
Taste 1.Comparison of characteristics useful in differentiating V.ozarkense and V.molle.N = number
of newanale measured or sampled. Data = range with means cae standard deviations in paren-
these = values significantly different at p <
Character N Viburnum ozarkense N Viburnum molle
leaf shape 190 ovate 158 broadly ovate, orbicular
*leaf width (cm) 190 4-14 (7.2+1.4) 158 5-12 (8.2+1.3)
*number of veins/ 192 4-8 (5.4+0.65) 162 5-8 (6.1£0.69)
half leaf
adaxial leaf surface 190 hirsute 158 glabrous
pubescence
*number of teeth/ 192 8-21 (13.842.3) 162 17-44 (25.6+4.2)
half leaf
*length of 1 petiole below 82 0.5-2.2 (1.140.37) 68 1-4.2 (2.40.52)
inflorescence (cm)
*length of 1° petiole 109 0.9-3.4 (2.140.56) 90 1.8-4.9 (3.4+0.72)
petiole pubescence 190 strigose and glandular 158 — glabrous to sparse glandular
*leaf tip length (mm) 184 3-15 (7.7+2.6) 160 1-7 (2.91.1)
*corolla width (mm) 19 4-9 (6.6+1.5) 21 3-6 (4.4+0.78)
*fruit length (cm) 16 0.8-1.3 (1.140.15) 25 0.7-1.1 (0.90.11)
bark tight throughout ie on >2 year old
0d
current season shoot color reddish-brown ion tan
tive shoots, oval in cross-section, moderately to det isely strigose and glandular, fur-
rows open; stipules often 2 pair per leaf stalk; shoots of current season reddish-
brown; twigs and stems with tight bark throughout Viburnum ozarkense
Leaves broadly ovate to orbicular, adaxially glabrous, 17 to 44 (mode 24) m Meas
teeth per half leaf; petioles of first pair leaves below inflorescence X= 2. x
3cmat tips of vegetative shoots, terete in cross section, glabrous or rie ne
dular, furrow closed; stipules in single pairs only; shoots of Current season yellow-
tan; older (>2 years) twigs and stems with exfoliating bark Viburnum molle
Phenology
Flowering material (about 5% each of Viburnum molle and V.ozarkense vouch-
ers) for the plants that occurred in Arkansas, Missouri and Oklahoma indicated
that V. ozarkense flowers earlier than V. molle. Based on the material | exam-
ined, V. ozarkense flowered between May 6 and June 7, with a peak flowering
time of May 19. Viburnum molle flowered between May 15 and May 29, witha
peak on May 27-29 (Fig. 2).
Habitat and Associates
Information gleaned from herbarium labels and field experience indicated that
Viburnum ozarkense and V. molle occupied similar habitats and had a similar
set of associates. Most associate data were compiled from field notes of the author,
WECKMAN, REINSTATEMENT OF VIBURNUM OZARKENSE 853
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Viburnum ozarkense
Populations of Viburnum ozarkense in Arkansas are most often encountered
on north-facing ridges, rocky limestone hillsides and wooded slopes in rich
woods. It also occurs on ledges and in rocky woods above and along creeks (ex.
Alum Fork, Bard Springs, Buffalo River, Cap Fork, Little Red River and Stepp
Creek). Associates of V. ozarkense in Arkansas include: Acer saccharum,
Arundinaria gigantea, Berchemia scandens, Carpinus caroliniana, Carya
cordiformis, Chionanthus virginicus, Cornus florida, Diospyros virginiana, Dirca
palustris, Hamamelis vernalis, ae arborescens, Magnolia ceumninare,
Philadelphus pubescens, Quercus i,Q. stellata, RI
Rhus aromatica, Sideroxylon lanuginosum, Staphyl ea trifolia, Tilia americana,
Vaccinium arboreum and Viburnum rufidulum.
In Missouri, Viburnumozarkense has been collected from north and north-
east facing limestone ledges and bluffs above the Eleven Point River, from lime-
stone slopes along Myatt Creek, and north facing limestone slopes along the
South Fork of the Spring River. These are often mid-slope locations with fil-
tered light. Associates of V. ozarkense in Missouri included: Acer saccharum,
Berchemia scandens, Carpinus caroliniana, Cornus florida, Fraxinus
quadrangulata, Lindera benzoin, Ostrya virginiana, Quercus alba, Q.
muhlenbergii, Q. rubra, Sideroxylon lanuginosum, and Viburnum rufidulum.
In Oklahoma, Viburnum ozarkense has been collected froma rocky north
facing chert slope above Caney Creek in Adair County, and wooded oak-pine
hillsides in McCurtain County. Associates of V. ozarkense in Oklahoma at the
—
854 BRIT.ORG/SIDA 20(2)
Pare he: f 12 WERT iagim a> 2@ and VYhbuwraum mele alante
4
3
22
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8 12 16 | 20 24 28 June 1 5
May 6 10 14 18 22 26 30 3 7
Cl - L | £UL L J UssLb i . Aryl RA’ - IALI-L
Fic. 2 of
Adair County site included: Acer saccharum, A. rubrum, Asimina triloba, Cercis
canadensis, Corylus americana, Hydrangea arborescens, Lindera benzoin, Nyssa
sylvatica, Quercus alba, Sassafras albidum, Staphylea trifolia, Tilia americana
and Viburnum rufidulum.
Viburnum molle
Viburnum molle occurs in Arkansas on chert slopes at the base of north facing
bluffs along War Eagle Creek in Madison County. A single collection by EJ.
Palmer documented V. molle in Boone County from “rocky ledges, Harrison.”
Associates of V. molle in Arkansas at the War Eagle Creek site included: Acer
saccharum, Asimina triloba, Hydra ngea arborescens, Juglans nigra, Lindera ben-
zoin, Magnolia acuminata, Quercus muhlenbergii, Q. rubra, Staphylea trifolia,
and Tilia americana.
Habitats in Missouri which support Viburnum molle include north facing
slopes, bluffs and limestone ledges along creeks and rivers (ex. Castor River,
Cole Camp Creek, Gasconade River, Jacks Fork of Current River, James River,
Jonca Creek, Lost Creek, Meramec River, Piney Creek, River Aux Vases, Stinson
Creek, Swan Creek, West Fork Cuivre River). Along these creeks and rivers, V.
molle plants typically occur as understory elements receiving filtered or part
day sunlight, at the mid-slope point, neither in the floodplain nor at the more
xeric ridge crest. In Missouri associates included: Acer saccharum, Asimina
triloba, Carpinus caroliniana, Cornus florida, Dirca palustris, Juniperus
WECKMAN, REINSTATEMENT OF VIBURNUM OZARKENSE 855
virginiana, Quercus muhlenbergii, Sideroxylon lanuginosum, Tilia americana,
and Viburnum rufidulum.
Viburnum molle is not known to occur in Oklahoma.
Distribution and rarity
Distribution of Viburnum ozarkense was compiled from annotated herbarium
vouchers and new collections produced for this study (Fig. 3). Viburnum
ozarkense is confined to the Ouachita and Boston Mountains and Ozark pla-
teau, and is documented from 14 counties in Arkansas, Howell and Oregon coun-
ties in Missouri, and Adair and McCurtain counties in Oklahoma. While some
populations have been sampled repeatedly over the past 75 years, (ex. type popu-
lation in Stone Co., Arkansas), others are documented by single collections. Based
on herbarium material, 50% of the known populations of V.ozarkense have not
been sampled for more than 30 years, and some populations not sampled for as
long as 80 years. Some Arkansas populations have conservation protection in
the Ozark National Forest, and along the Buffalo National River.
Viburnumozarkense populations in Missouri are afforded some protection
at sites along the Eleven Point River in the Mark Twain National Forest. A popu-
lation examined in 2000, at the western edge of the range in Adair County,
Oklahoma, was composed of about 15 individuals and is not known to have
conservation protection. The size and conservation status of the McCurtain
County, Oklahoma population is unknown; this population has not been
sampled since 1966. With a known distribution of 18 counties in three states, V.
ozarkense clearly falls under the 26 county upper limit for rare southeastern
endemics established by Estill and Cruzan 2O00D. Viburnum ozarkense should
be considered for state listing in states where it occurs and attempts should be
made to locate additional populations.
The commonness or rarity of V. molle throughout its range has not been
critically reviewed. In Arkansas, V. molle is restricted to Madison and Boone
counties (Fig. 3). The Madison County population of V. molle, was discovered
by Shepherd in 1987 while working for the Arkansas Natural Heritage Com-
mission. This population was re-visited in 2001 by Weckman and Shepherd
and is estimated to consist of 35-50 individuals. It is on private land with no
current conservation protection. An attempt by the author to relocate the Boone
County population collected by Palmer in 1914 was not successful in 2000. The
terse label information, “rocky ledges, Harrison” gave few clues as to its loca-
tion. This population may no longer be extant. This research indicates V. molle
is the rarest Viburnum species in Arkansas.
Viburnum molle occurs in 16 counties in Missouri. A population observed
in Oregon County in 2001 was estimated to consist of about 12 individuals. La-
bels typically lack information about numbers of plants in a population; thus
sizes of other populations are unknown. In the three state study area, herbarium
856 BRIT.ORG/SIDA 20(2)
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Fic. 3. Distribution of Vit kense Ashe (@) and Vit He Michx. (a) in Arkansas, Missouri and Oklat
based on annotated herbarium vouchers.
vouchers examined of V. molle reveal that 50% of the counties were last col-
lected in or before the 1950s. Conservation status of the Missouri populations
of V. molle are unknown but some may be afforded protection in the various
units of the Mark Twain National Forest and along the Eleven Point River.
Overlap of morphology and geographic range
For the mensural characters used in this study, ranges of the measurements
overlap, however 8 of 14 characters were statistically significantly different at
p= <0.05. If single character taxonomy is desired in the field, the sine qua non
for Viburnum molle is the presence of exfoliating bark and conversely the ab-
sence of exfoliating bark defines V. ozarkense. This is an inviolate Viburnum
molle character, used by Michaux in his type description, and observed by the
author in the field for plants in Kentucky, Ohio, Missouri, and Arkansas, and on
herbarium specimens of V. molle through its range in Tennessee, Indiana, IIli-
nois, and lowa. For determination of herbarium material collected without older
stems, the suite of abundantly toothed leaf margins, short apex, and glabrous,
terete petioles on yellow-tan twigs would lead one to Viburnum molle. Conversely,
a specimen with moderately toothed leaves, longer, acute tips, and glandular-
—
WECKMAN, REINSTATEMENT OF VIBURNUM OZARKENSE 857
pubescent, oval petioles on reddish-brown twigs would lead one to Viburnum
ozarkense. These suites of characters allow for the differentiation of these two
related species.
Viburnum ozarkense and V. molle co-occur in Madison County, Arkansas
and Oregon County, Missouri, at the southwest edge of the range of V. molle
(Fig. 3). In Arkansas the populations are about 30 air miles apart, while in Mis-
souri the populations are about 12 river miles apart. No sympatric populations
of Viburnum molle and V. ozarkense were located by the author, nor were any
individuals appearing intermediate between these species observed in the field
or voucher material examined.
Specimens examined: V As RKANSAS: B Co.: Baxter Co..6 Jun 1931, Ashe
s.n. (NCU); 7 Jun 1931, ren s.n. (NCU); cae 1931, Ashe s.n. (NCU, NY). Benton Co.: Devils Eyebrow,
22 Apr 1928, Demaree 4842 (UARK). Franklin Co.: Devils H Hollow, 3 Jun 1978, Barber 849 (UAM, UARK).
Garland Co.: creek banks, Ouachita River, 28 Aug 1939, Demaree 20517 (MO), Walnut Creek Recre-
ation Area, 10 Aug 1951, Moore 510786 (UARK); rich woods, Charlton Recreation Area, 23 Apr 1994,
Sundell et al 10759 (NA, UAM). Johnson Co.: creek bank, Ozark National Forest, 14 Sep eee
Kirshberger & Davis 101(UAM, DARK) eevee Co.: oe co S of Red oi 18 Apr 19
Kirshberger 12 (UAM); mesic woods, \ n, 17 Nov 1973, Tucker 12461 (SMU) Little vee
berry Creek, S of Red Star, 3 Jun 2 vai We eman 601 -6419 (EKY). Newton Co.: Stepp Creek n
Swain, 20 Jul 1979, Bonar s.n.(UARK); Boxley, 22 May 1948, Moore 480226 (UARK); limestone ae
above Buffalo River, 20 May 2000, Weckman 5597A,B- -5599 (EKY); upland woods Buffalo River, near
Boxley, 20 May 2000, Weckman 5603(EKY). Polk Co.: creek banks, Bard Springs, 4 May 1955, Demaree
36632 (SMU). Pope Co.: Snow Creek, N of Hector, 31 Oct 1978, Boatright & Utley 102 (APCR); along
creek bank, Hector, 17 Oct 1932, Merrill 89 (A); creek bank, Nogo, 11 May 1933, Merrill 271 ILL, MO,
UARK); rugged hills NE of Russellville, Aug 1955, Moore et al 55-432 (UARK); rich woods along In-
dian Creek, SW of Pelsor, 10 May 1969, Tucker 7926 (AI am Saline Co.: bluffs, Alum Fork, Saline
River, 28 May 1954, Demaree 35333 (NCU, US); the Narr of Crows, 24 Sep 1995, Sundell G Pagan
11726 (UAM); narrow ridge, Saline River, 11 Jul 1997, eee oe (UARK),; N face slope above
Alum Fork, 6 Jul 1999, Weckman & Walker 5110, 5111,5113,5131 (EK Y). Searcy Co.: rocky creek bluff,
Harriet, 9 May 1970, Demaree 61712 (MO); hillside, S of Marshall, 29 Apr 1967, Graham 343 (NCU).
Stone Co.: near Sylamore, n.d., Ashe s.n. (NCU); near Sylamore, 18 May 1924, Ashe s.n. (NCU); bottom
aE NiSilamore Cee 7 Jun 1931, Ashe s.n.(NCU); along Cap Fork Creek, 6 May 1981, Davis 2975 (APCR);
creek bottom in woods, Mountain View, 10 May 1952, Demaree 31751 (US, VDB); rocky hillsides
Sasa Springs, 19 May 1952, Demaree 31850 (NCU, US, VDB); rocky open woods, bluffs, Sylamore
eek, 19 May 1965, Demaree 52296 (NCU); Barkshed Recreation Area, along N Sylamore Creek, 29
he 1967,Graham ae emacs Blanchard Springs, 18 Jun 1945, Moore 450600 (UARK); Sylamore
24 Jun 1945, Moore 450629 (UA Sylamore Forest, 24 Jun 1945, Moore 450646 (UARK), Blanchard
Springs, 23 Jun 1945, Moore 450721 (WARK): Blanchard Springs, 18 Aug 1948, Moore 480735 (UARK),
Caney Creek, N of Mt. View, 21 Jun 1951, Moore 510347 (UARK); wooded N facing slope, Gunner Pool
Recreation Area, 19-21 May 1978, Redfearn 31673 (SMS); Big Creek, 9 May 1970, Smith 1462 (UARK);
above creek bed, vicinity of Blanchard Springs, 22 May 1984, Sundell 6308 (UAM), along Sylamore
Creek, 8 Oct 1966, Tucker & Demaree 3578 (APCR),; alluvial woods, Barkshed Recreation Area, 14 Oct
1967, Tucker 6862 (APCR); alluvial woods along N aes Creek, 14 Oct 1967, Tucker 6867 (APCR,
NCU), ees Blanchard Springs Recreation Area, 2 Oct 1969, Tucker 8225 (APCR), rocky
streamban anchard Springs, 5 Aug 1972, Tucker 10418 (UARK); rocky slope, Gunner Pool, 5 Aug
1972, Serie PCR); along N Sylamore Creek, 8 Oct 1972, Tucker 10605 (APCR - 2x, SMU, VDB);
Cap Fork, Barkshed Recreation Area, 7 Jul 1999, Weckman 5151, 5160-5168 (EKY ); Blanchard Springs,
—
ory
858 BRIT.ORG/SIDA 20(2)
7 Jul 1999, Weckman 5172-5176, 5178-5184 (EKY). Van Buren Co.: sandstone bluffs, Little Red River,
near Shirley, 28 May 1924, Palmer 25197 (A, MO, UARK). ag Co.: Cove Creek Valley, 17 Aug
1954, Hite 102 CUARK). Unknown Co. Arkansas, n.d., anon. sn, (NC
MISSOURI: Howell Co.: limeston slopes along Mya att Creek, es Apr 1938, Ste ye rmark 5186 (F
MO); lime bluffs, Eleven Point River, 12 Aug 1934, Steyermark 14503 (MO, NA,US):; shaded lime bluff,
Eleven Point River, NE of Peace Valley, 12 Aug 1934, Steyermark 14503A, 14503B (UMO); Eleven Point
River near Blowing Springs, l4 Apr 1935, Steyermark 18614 (UOMO); S Fork Spring River, 26 Jun 1955,
Ste yermark 78762 (UMO); N face limestone slope, S F i Spring River, 26 Jun 1955, Steyermark 78763
(UMO); limestone bluffs, S Fork Spring River, 26 Jun 1955, Steyermark 78765 (GH, MO, UMO); along $
Fork Creek, 19 Jul 1979, Summers 527 (MO); ledges, bl if dolomite glade, SE of Mt. View, 27 Jun 1990
Summers 3394 (MO), N face dolomite bluff, S Fork Spring River, 4 Jul 1990, Summers 3431 (MO). N
face dolomite bluff, S Fork Spring River, 4 Jul 1990, Summers 3432 (MO); Eleven Point River at Blow-
ing Springs, 14 Sep 1994, Summers 7102 (MO); Eleven Point River at Blowing Springs, 2 Jun 2001,
Weckman 6360-6365, 6371-6385 (EKY), S Fork Spring River, N of Lanton, 2 Jun 2001, es 6390-
6393 (EKY). Oregon Co.: SW facing dolomite glade and bluff, 4 Aug 1993, $ MO).
OKLAHOMA: Adair Co.: rocky hillside, 14 May 1980, Huft & Goodman 1160 (OKL); shen SecE above
Caney Creek, 27 Apr 1968, Perino & Pierson 168 (OKL): NW of Stilwell, 13 Jun 1957, Rice s.n. (GH,
OKL, OKLA); NE of Stilwell, 2 May 1959, Rice s.n. (OKLA); chert slope above Caney Creek, 19 May
2000, Weckman 5577- -5587 (EKY). MeCurtain Co.: oak-pine woods in mountains, $ of Bethel, 9 Aug
1948, Waterfall 8515 (OKL, OKLA): wooded hill, S of Bethel, 4 Jun 1949, Waterfall 8826 (OKLA); open
woods, base of mountains, N of Broken Bow, 24 Jun 1966, Waterfall 17228 (NCU, OKLA).
Viburnum molle Michx. ARKANSAS: Boone Co.: rocky ledges, Harrison, 25 Oct 1914, Palmer 6914 (A,
MO, US). Madison Co.: ledge, limestone bluff, War Eagle Creek, 15 May 1987, Shepherd & Smith 286
(UARK); base of blulf, War Eagle Creek, N of Huntsville, 4 Jun 2001, Weckman & Shepherd ene
6441 EKY). MISSOURI: Barry Co.: wooded slopes of Piney Creek, 22 Jun 1937, Steyermark 22597 (
F MO). Benton Co.: bluffs of Cole Camp Creek, 28 May 1896, Demetrio s.n.(MU, NCU, US): bluffs i
Cole Camp Creek, vicinity of the Big Cave, 27 May 1897, Demetrio s.n. (GH); Cole Camp Creek, May
1905, Fulleraf sn (MO); Cole Camp Creek, 30 Aug 1896, Heck s.n. (GH); Bluffs of Cole ae Creek,
vicinity of Big Cave, 9 May 1897, Heck s.n. (GH); N facing slopes, ledges, Big Cave bluff, 2 Oct 1924,
Palmer 26370 (A, GH, MO, MOR, UMO); N facing ledges, Big Cave bluff, 12 May 1926, Palmer 30061
(A, MOR, UMO); bluffs, Cole Camp Creek, 24 Sep 1938, Steyermark 7289 (F), Cole Camp Creek, 19
May 1904, Trelease 4 (A); Big Cave, Cole Camp Creek, 12 Jul 1897, Trelease 383 (NCU); Big Cave, Cole
Camp Creek, 12 Jul 1897, Trelease 384 (A, MO); Cole Camp Creek, n.d., anon. s.n. (A). Callaway Co.: N.
limestone slopes along Middle River, 12 Sep 1937, Steyermark 26203 (F MO) steep slopes, Stinson
Creek, 12 Sep 1937, Steyermark 26235 (F, MO); N slopes, Stinson Creek, 25 May 1948, Ste yermark 65464
(F). Christian Co.: along Billin Creek, 28 Sep 1905, Bush 3480 (A, MO); rich woods, 21 May 1907, Bush
4625 (A, MO); bluffs, Swan Creek near Garrison, 28 Oct 1955, Palmer 61802 (UMO); N facing wooded
slope along Woods Fork, 9 May 1981, Redfearn 32647 (SMS, UMO); E and NW facing bluffs, Swan
Creek, 6 Jul 1937, Ste yermark 23039(E NY). Crawford Co.: rocky woods along stream, Z: ahorsky Woods,
25 Aug 1983, Christ sn. (MO); cliff base, Meramec River, 12 Sep 1986, oe . (MO), hillside near
Steelville, May 1929, Shoop s.n. (UMO): lime bluffs, Meramee River, 21 ¢ 934, Steyermark 16317
(MO, US): lime bluffs, SE of Bourbon, 21 Oct 193 = ae 16317A, a (UMO), steep slopes,
Meramec River, 15 Jun 1941, Steyermark 41332 (EF S side Meramec River, 15 Jun 1941,
Steyermark 41339 (F GH, MO, US): base of slaps, “Me ramec a 15 Jun 1941, mal 413415
GH); NE face, Meramec River, 1] Jun 1955, Steyermark 78602 (UMO). Lincoln Co.: N face bluffs W
Fork Cuivre River, 28 Apr 1941, Steyermark 28516 (F GH, MO); McDonald Co.: ich hillsides, Noel, 2
Apr 1909, Bush 5531 (A); rich woods, Noel, 27 May 1909, Bush 5763 (A). Madison Co.: limestone bluff,
outcrops, Castor River, 27 Aug 1948, Ste yermark 66097 (F). Oregon Co.: limestone bluffs, Eleven Point
River, | May 1986, Summers 1589 (MO); Cane Bluff, Eleven Point River, | Jun 2001, Weckman 6350-
Oo
WECKMAN, REINSTATEMENT OF VIBURNUM OZARKENSE 859
6359 (EKY). Pulaski Co.: N facing li | le River, 24 Aug Se pea reie
(F, MO); N limestone bluffs, Gasconade River, 10 Se ep 1956, S k 82532A,82532B(UMO). Phelps
Co.: N facing bluff, Gasconade River, 2 May 1991, Summers 4248 (MO). St. Genevieve Co.: talus slopes,
ledges, Jonca Creek, 17 Jun 1993, Brant 2398 (MO), NW facing lime bluff, River Aux Vases, 6 Jul 1946,
Ste yermark 63872 (F), NW facing lime bluff, River Aux Vases, 6 Jul 1946, Steyermark 63873 (F). Stone
Co.: cliffs of James River, 15 Oct 1913, Palmer 4671 (A, US); high limestone bluffs, James River, 20 May
1914, Palmer 5671 (NY, US); high limestone bluff, James River, 11 Sep 1918, Palmer 14361 (A); high
limestone bluffs, James River, 17 Apr 1920, Palmer 17226 (A), N high lime bluffs, James River, 23 May
1923, Palmer 22816 (A, MO, MOR, UMO, US); high limestone blutfs, James River, 24 Sep 1923, Palmer
23872 (A, SIU); high limestone bluffs, James River, 24 Sep 1923, Palmer 23872a (A, MOR); N face,
imestone ledges, James River, 16 Sep 1924, Palmer 26158 (A, MOR); N face limestone bluffs, James
River, 16 Sep 1924, Palmer 26158a (A, MO, MOR); mesic forested dolomite bluff, James River, 23 Jun
1988, Smith 2733 (MO, SMS). wooded blulf eae River, 22 Jun 1941, Ste yermark 40133 (F). Taney
. bluffs along Swan, 9 Oct 1899, Bush 798 (GH, MO, NCU, ND-G, NY, US); rich banks, Swan, 26 Sep
ath Bush 3449 (A, GH, NY, US); cliffs of Swan ‘River 8 Oct 1894, Sargent s.n. (A). Texas Co.: Jacks
Fork, Current River, 26 Aug 1939, Bauer 867 (F); rocky slopes, dolomitic bluff, Jacks Fork, | Aug 1969,
Red ae et al 899 (MO, NCU, SMS); lime bluffs, slopes, Jacks Fork Current River, 15 Aug 1934,
ae) yer ea 14574 (GH, MO); lime bluffs, euiee Jacks Fork Current River, 15 Aug 1934, Steyermark
14574B (UMO); N face limestone bluff, N Prong of Jacks Fork, 27 Apr 1952, Steyermark 73LI7
(F, eer N face dolomite bluff, Jacks Fork, - Aug 1990, Summers 3625 (MO). Warren Co.: Lost
Creek nae ones 4 pun 1987, Christ s.n. (MO); limestone bluffs along Lost Creek, 9 Jun 1946,
Ste y
—
i
ACKNOWLEDGMENTS
I would like to thank Stephen Walker who introduced me to the unusual “V.
molle” of Arkansas and Bill Shepherd for facilitating re-location the V. molle
population he discovered in Arkansas in 1987. Additionally, Johnnie Gentry,
Burnetta Hinterthuer, Philip Hyatt, Lang Kirshberger, Gene Leeds, Janet Self,
Karen Tinkle and Gary Tucker shared valuable locality information and pro-
vided logistical support in Arkansas. In Missouri, essential locality informa-
tion was provided by Bill Summers and George Yatskievych. Analysis of statis-
tical data was greatly facilitated by Judith Weckman. This work was supported
in part by funding from the Office of the Dean, College of Arts and Sciences,
Eastern Kentucky University. Finally, | am indebted to the curators of the her-
baria listed above who generously lent from the specimens in their care.
REFERENCES
AsHe, W.W. 1928. Notes on southeastern woody plants. Bull. Torrey Bot. Club 55:463-464.
Coker, W.C., J.S. Houmes, and C.F. Korstian. 1932.William Willard Ashe. Elisha Mitchell Sci. Soc.
48:40-47.
Esti, J.C. and M.B. Cruzan. 2001. Phytogeography of rare plant species endemic to the
Southeastern United States. Castanea. 66:3-23
Hotmoren, PK., N.H. Hotmaren, and L.C. Barnett (eds.). 1990. Index Herbariorum. Part 1: The
Herbaria of the World, 8'" edition. New York Botanical Garden, Bronx, NY.
860 BRIT.ORG/SIDA 20(2)
Massey, J. 2001. Personal communication
McAtee, W.L. 1956. A review of the nearctic Viburnum. Privately published by the author.
Chapel Hill, North Carolina.
Michaux, A. 1803. Flora Boreali-Americana. 2 volumes. Paris and Strasbourg. Facsimile re-
print.Introduction by J.Ewan 1974. Hafner Reprint, New York.
Moore, D.M. 1941.A checklist of the ligneous flora of Arkansas. Proc. Arkansas Acad. Sci.
1:41-55.
Smith, E.B. 1988. An atlas and annotated list of the vascular plants of Arkansas. Privately
published by the author. Fayetteville, Arkansas.
STEYERMARK, J.A. 1963. Flora of Missouri. lowa State University Press, Ames.
Tucker, G.E. 1976. A guide to the woody flora of Arkansas. PhD dissertation, University of
Arkansas, Fayetteville.
USDA, NRCS. 2002. The Plants Database, Version 3.5 (Attp://plants.usda.gov). National Plant
Data Center, Baton Rouge, LA 70874-4490 USA.
CELEBRATION HONORS PIONEER FIELD
BOTANIST ANDRE MICHAUX
William R. Burk
John N. Couch Biology Library
Ur liversity of North Carolina
CB#3280 Coker Hall
Chapel Hill, NC 27599-3280, U.S.A.
André Michaux, noted 18th century North American plant explorer extra-
ordinaire, died on the island of Madagascar two hundred years ago. On the oc-
casion of the bicentennial of his death, Michaux’s life and contributions to North
American botany were highlighted and elucidated at “A Michaux Celebration,”
attracting nearly 4,000 people. The events began with the André Michaux In-
ternational Symposium (AMIS), held May 15-17, 2002 in Gaston County, North
Carolina. The subsequent two days—called “Celebrate France! Festival Foire!”—
occurred at the Daniel Stowe Botanical Garden, a relatively young garden with
strong public programs. AMIS constituted the official symposium for the South-
ern Appalachian Botanical Society (SABS), held for the first time separately from
the Association of Southeastern Biologists’ annual meeting. Belmont Abbey
College (BAC), the Daniel Stowe Botanical Garden (DSBG), and Gaston Day
School (GDS) were the hosts of the celebration, three years in the planning. Li-
brarian Charlie Williams was the key architect and organizer of the Celebra-
tion, whereas Jeanne Miller was the coordinator. Michael J. Baranski (SABS)
served as Program Chair for AMIS. Others were Carol Brooks, Dean de la Motte
& Robert Tompkins (BAC); Mike Bush (DSBG), Martha Mayberry (Mint Mu-
seum of Charlotte); Richard Rankin (GDS); and Jean-Pierre Riviere (French-
American Chamber of Commerce, NC).
Setting the tone for the first day were two field trips, enticing and piquing
attendees curiosities and expectations of Michauxiana tocome. That evening ethno-
botanist Mark Plotkin delivered the opening address at the Gaston Day School.
We learned about the importance of natural products from tropical Brazil and
Surinam, such asa potential painkiller from the poison dart frog. He emphasized
the need for a holistic approach to conserving tropical flora and fauna. Just as pio-
neer Michaux persevered under the rigors of exploration, conservation efforts
require persistent leadership under equally challenging circumstances
The fast-paced sequence of 28 fact-packed presentations of May 16 and 17,
enlightened our knowledge of Michaux. James L. Reveal, historian of botany,
delivered the plenary talk, “No Man isan Island: The Lives and Times of André
Michaux.” After placing into perspective the accomplishments of naturalists
prior to Michaux’s arrival in the United States, he discussed Michaux’s contri-
SIDA 20(2): 861-864. 2002
862 BRIT.ORG/SIDA 20(2)
Walter K. Taylor as André Michaux at AMIS, 16 May 2002, Bel Abbey College, Belmont, North Carolina. (Photograph
by Ronald L. Stuckey, Columbus, Ohio)
BURK 3 863
butions to field botany. Subsequent paper presentations were of two types: in-
vited (9) and contributed (19). Two other scheduled papers on “The Lowcountry
Connection of André Michaux” and “Nature and History of the Botanical Ma-
terials of André Michaux in the Herbarium of the Academy of Natural Sciences
of Philadelphia” were not presented.
Each presentation was virtually unique in topic, overlapping little with
others. Among the varied themes on Michaux and his activities, were: his con-
nection with the Bartrams of Philadelphia; his Canadian journey; several ac-
counts on plants—named as new to science by him—including Magnolia
macrophylla; the rediscovery of the site of his botanical garden in Charleston,
South Carolina by archaeologists from the College of Charleston; his involve-
ment with the intriguing Edmund Charles Genet Affair in American history;
and his excursion to Madagascar where he died on October 11, 1802 (confirmed
by anewly discovered death certificate from Mauritius). When speaker/author
Walter K. Taylor gave his talk on Michaux in Spanish East Florida during 1788,
he appeared in a period costume, eliciting surprise and delight from the audi-
ence. In closing the symposium, Francoise Winieska, noted photographer of
gardens and plants representing France at the program, gave a slide presenta-
tion, “18" century French Gardens in the Domains of Versailles and Rambouil-
let,” sharing her striking photographs with the captivated viewers.
Additional field trips were held during the symposium and at its conclu-
sion. We enjoyed two dinners at the Stowe Botanical Garden, the evening of May
16 being a “French Gala’ of cuisine, eighteenth century period music and dance.
On the last evening of AMISa special treat was an exhibit of engravings and water-
colors by eighteenth century botanical artist Pierre-Joseph Redouté at the Mint
Museum in Charlotte. Opening in March 2002, the exhibit was specially devel-
oped as an adjunct to the Michaux symposium and included five associated talks,
among them, “Botanical Illustration: Redouté and the Michaux,” by librarian/
historian lan McPhail. The exhibit will close this year on Bastille Day, July 14.
During breaks and other free periods of time, attendees were lured by ex-
hibits, such as the book displays by the Missouri Botanical Garden and by sym-
posium speakers and other participants. Among the latter were books and au-
dio-cassettes by C. Ritchie Bell; Gail Fishman; Walter K. Taylor and Eliane M.
Norman; and Charlie Williams. Locally produced and limited in distribution,
the booklet “Memoirs of the Life and Botanical Travels of André Michaux,” by
J.PE Deleuze is of special note to librarians and historians. Conceived and ed-
ited by Charlie and commemoratively published for the symposium, it contains
four appendices, among them a supplementary bibliography by Ronald
Gilmour. Charlie’s role as editor isa misnomer of the extent of his contribution,
as he is also the writer of two of the appendices. The Catawba Lands Conser-
vancy prepared the useful color map folded in the back of the text. The map
traces nine journeys that Michaux made in North America from 1787 to 1796.
864 BRIT.ORG/SIDA 20(2)
The publication is available only from Charlie Williams, 6720 Wheeler Drive,
Charlotte, NC 28211. Checks are to be written for $16.50 (which includes post-
age/handling) to “Belmont Abbey College.” Artist and school administrator Erin
Whitener had a popular display of her line of note cards, featuring color illus-
trations of plants discovered by Michaux, which sold out early in the program.
Her design of the Magnolia macrophylla, leaf or fruit, appeared on symposium
stationery, handouts, and tote bags given to attendees
Two AMIS booklets in our orientation bags may be of interest to others,
particularly librarians. These were an “Attendee Roster” containing names, titles,
addresses, phone, fax, and email addresses of all attendees; and “Abstracts,” pro-
viding succinct summaries of each presentation. Fortunately, the proceedings
of AMIS are planned to be published in Castanea and its Occasional papers.
The French Festival featured special talks, French cuisine, and vendors of
merchandise such as artwork, jewelry, and Limoge porcelain. This Festival was
open to the public and included activities for children such as face painting,
games, and guignol (a Punch and Judy show). Sponsored by the French-American
Chamber of Commerce (NC), this event took place at the Stowe Garden on May
18 and 19.
The Michaux Symposium is a model for future theme-type conferences.
Both the Symposium and the highly popular “Celebrate France!” were resound-
ing successes. The fervor of symposium goers was evident as noted by conver-
sations sparked by the presentations and scholarly interests of those in atten-
dance. Special thanks are extended to the organizers, hosts, and myriad
behind-the-scenes people for bringing together sucha distinguished assemblage
of Michaux scholars and providing a wide array of events.
Historians of botany will have access to a collection of information sheets
and brochures available at the conference that have been assembled by The Ohio
State University botanist Ronald L. Stuckey. He has deposited the compilation
in the Hunt Institute for Botanical Documentation, Pittsburgh. This contribu-
tion also includes all of the photographs he took of the invited speakers par-
ticipating in AMIS.
The symposium will be a lasting contribution to the history of botany,
bringing visibility and recognition to the great botanical pioneer André Michaux.
“Celebrate France!” accentuated the important French contribution to Ameri-
can culture. “Celebrate Michaux” merged science and culture to make for a fas-
cinating five days of learning and entertainment. Vive Michaux! The influence
of Michaux also was recently recognized by the NC Department of Highways,
which landscaped an intersection of Interstate 85 in Gastonia with plants first
discovered by Michaux. The plot is identified with a permanent marker.
——
ACKNOWLEDGMENTS
Ronald L. Stuckey, The Ohio State University, Columbus, assisted with writing
the paper (May 24, 2002).
DEL WENIGER (1923-1999),
REMARKABLE TEACHER AND NATURALIST
Marjorie Fohn Epler
924 East 50th Street
Austin, TX 78751, U.S.A.
In 1960, Del Weniger joined the faculty of Our Lady of the Lake College in San
Antonio, Texas.
He was later to become the chairman of the Biology Department. After
teaching general biology and ecology for nearly forty years, Del retired from
O.L.L. in 1988 as Professor Emeritus.
Although born in Kingman, Kansas, Del thought of himself as an adopted
Texan. Appropriately so, since he researched the plants and animals of the state
for half a century, this resulting in numerous published articles, and his first
two books, Cacti of the Southwest and Cacti of Texas and Adjacent States. The
first book received, in 1970, Theta Sigma Phi’s “Author of the Year's Best Book By
A Texas Publisher” award, as well as accolades from numerous organizations
and reviewers. Echinocereus wenigeri, a Texas cactus was named for him.
His first two books were followed by Explorers Texas, Vol. 1, which the Texas
Historical Commission judged to be the “Best Historical Publication of 1984.”
Vol. II of the series soon followed. Volumes III and IV were in progress at the
time of Del’s death. He also co-authored the book, San Antonio in the Eighteenth
Century, which received the Texas Historical Commission's 1978 “Official Texas
Award For The Best Publication of the Year in Local or Regional History.”
Del spent many | hing whatever was written down by the early
explorers and pioneers of Texas. His fami ly and friends remember the great plea-
sure he took in finding new journals or letters of early explorers or settlers who
recorded what they saw as they traveled Texas. His deep sense of stewardship
of the earth and its gifts was evident in his teaching, his lectures, his volunteer
work, and his ecological consultation for federal and local agencies. He stressed
responsible, respectful use of our natural heritage as he served as a founding
member of the San Antonio Botanical Center, and on local, state and regional
boards of the Nature Conservancy and Sierra Club.
A thorough-going naturalist with a respect for all forms of life, Del exhib-
ited an educator's affection for those of his students who d trated enthu-
siasm for the study of biology. As a former student I can attest to his effective-
ness in teaching both the fundamentals of ecology and the delight of field work
associated with such studies. Professor Weniger loved teaching as much as he
loved learning and always took extra time for a student's interest or inquiry.
SIDA 20(2): 865- 866. 2002
866 BRIT.ORG/SIDA 20(2)
Those of us who were familiar with Del Weniger’s dedicated working hab-
its and regular good humor were amazed to learn, upon his death, that he had
been living, since birth, with an illness called Eisenmenger Complex, a condi-
tion whereby a person is born with only a 3-chambered heart. Medical experts
think that, because of the heart having to work so much harder, one’s life ex-
pectancy is lowered to less than 40 years. Delbert Kenneth Weniger, born May
LO, 1923, died at the age of 76 on July 10, 1999.
Del Weniger (1923-1999)
BOOK REVIEWS 867
Book REVIEWS
THE NATIONAL BIODIVERSITY TEAM OF THE REPUBLIC OF THE MARSHALL ISLANDS. 2000. The
Marshall Islands, Living Atolls Amidst the Living Seas: The National Biodiversity
Report of the Republic of the Marshall Islands. (ISBN 982-9045-02-1, pbk.)
St. Hildegard Publishing Company. Santa Clarita, CA 85719, U.S.A. Price
not given, 345 pp, numerous illustrations, 8 1/2" x 11"
During the Earth Summit in Rio de Janeiro, Brazil in 1992, the Republic of the Marshall Islands was
among the first to sign the Convention on Biodiversity. Consequently, the Republic sanctioned a re-
port on the biota of its atolls with the hope of raising public awareness to the various aspects of
biodiversity. The goal of this report is to “provide background about the Marshall Islands, its history
especially in relation to plants, animals and ecosystems, to give an overview of its biodiversity and
biological resources” (p.3). lt is mainly a compilation of ‘ological noon previously published
by other scientists, place nasinge format thereby making it he residents of the
Marshall Islands. Even the writing style is informal, “with the idea of making the information avail-
able to all sorts of people, even high school student
While on the surface this book might seem aoe for only asmall target audience, only a
brief look into this book reveals many informative features. Many of the organisms are illustrated,
such as the reef crab and giant ereupe with the Marshallese name and derivation prouiedl Text
ent important biological concepts. A section on “What is a Coral”
a
boxes Ramune a special topic pre
informs the reader about ae groups of corals and coral li | isd s. Likewise, a section
on endemism defines the topic and the term “invasive species” is discussed in some c
The first part of the book is comprised of three chapters, apes deal with the history of the
Qo.
etail.
Marshall Islands ane its people, biological resources, and threats to biodiversity. Chapter one nace ns
witha short historical account of the Marshall Islands and its people, history about the f
the Marshall island atolls and their relatively recent colonization by terrestrial life only 3, 000- 4, a
he detrimental effects people have had on the atolls, with the introduced
‘several species, including the en-
—
years ago. Furthermore, t
plants and animals they brought, has resulted in the extinction o
demic flightless Wake rail, the purple-capped fruit dove and the Micronesian pigeon. The history of
government reveals that the Marshalls have been under the control of several countries, including
Germany, Japan and the United States, before becoming their own nila the in 1990 and j ae
ing the United Nations. This chapter also provides information on the individual atolls
the Marshalls, with the Marshallese names of each, the number of square land ae they neeiey
and brief historical information.
Chapter two, entitled “Biodiversity and Biological Resources,” gives a brief overview of the eco-
systems on the Marshall Islands, listing each by plant community and including information on
location, importance, status and threats. Following is a concise overview of the biota of the Islands,
o
divided into plants and animals, with tables to give quick information about total number of species,
percent native species, percent endemism, muni: of threatened and endangered species, etc. The
fragility of these islands and invasive nature of newly arrived exotics is evident in these tables, since
only 18% of vascular plants and 23% of animals are native.
versity, which include nuclear
threat
C
Chapter three deals with current and historical thre:
testing, the invasion of exotics, changes in population and lifestyle of the native people and global
to hiod
climate chan
The second half of the book is poceieated to tables and lists . ae epee anal sc
| iders, insects bi
for plants, pulmonate and non
and mammals. For each of the species in these ae a name, Enelish name, Marsh ae
SIDA 20(2): 867. 2002
868 BRIT.ORG/SIDA 20(2)
name, status and traditional uses are reported. Next, ee on ee conse worthy of conser-
vation concerns with definitions of vs. endangered)
In addition, a table of endemic species, one on seslilerous: and faves species and those species
protected by Marshall Island law ease the reader in identifying biota of critical importance. Illustra-
tions are found throughout the book, often with scientific and Marshallese name, and five color fig-
ures with keys to identify the organisms in an important community, such as a coral reef. These
illustrations enhance the book’s properties, helping reader appreciate the biodiversity on the Mar-
shall Islands.~Amy lrauth Nare, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth,
TX, 76102 U.S.A, amy_trauth@yahoo.com.
CHARLES H. Smit. 2000. Biodiversity Studies: A Bibliographic Review. (ISBN 0-
8108-3754-4, hbk.) Scarecrow Press, 4720 Boston Way, Lanham, MD 20706.
US.A. http:// wwwscarecrowpress.com (Orders: The Rowman & Littlefield
Publishing Group, 15200 NBN Way, Blue Ridge Summit, PA 17214-0191, U.S.A,
800-462-6420). $69.00, 449 pp, 51/2" x 81/2".
Charles H. Smith, Science Librarian and Associate Professor of Library Public Services at Western
Kentucky University, cites literature both on the biological study of diversity itself, and on the socio-
natural science of diversity conservation. His effort has received two very prestigious recognitions.
His work was listed in Library Journal's Best Reference Sources for eae si it won the biannual
2001 Oberly Award for Bibliography in the Agricultural or aa Scien
Two bibliographies are included in this work. Bibliography I, “Books and Articles,” lists approxi-
mately 1,200 monographs and 4,500 articles, while Biblicee oy II, sce ssues,” provides a selec-
tive accounting of special issues of journals/serials dealing exclusively with biodiversity in all aspects.
His emphasis is on the literature of the investigation of Plociversiyy the written record of efforts to
measure, understand, monitor, and preserve it. Some self-imposed restric Hens - a to Be eens
during his exhaustive literature search, We included over one hundr
an equal number of print bi pilee apa and references cited lists. Because of space limitations indi-
vidual papers appearing in edit s were excluded, as were electronic sources, theses, dis-
sertations, reference works, systematic or floral/faunal revisions, and juvenile literature. Almost every
entry in the list is in the English language. What is featured is the primary literature of biodiversity-
directed sae science, social science, and humanities subjects, with a select amount of science
el ages erage centers on the period 1986 to 1998, extends to many works published prior
to that date o remain relevant to trends occurring during it.
s basic goal was to provide as much hacia: as possible in as little space as aaa for
each entry. Each citation is enhanced with bibliometric ratings, brief annotations, and/or a list of
descriptive “added terms.” The bibliometric malne se spores s appears asa l to 3 at according to
~ average number of times they have bee ra several- to ten- or twelve-year period
n the combined Citation Index portions of Science Citation index Social ee Citation Index,
a Humanities Citation Index. Monographs are rated on the same 1-3 scale but on three different
factors: (1) the number of OCLC libraries who hold the item; (2) number of times the work was re-
viewed after ia reeiiaa and (3) number of citations it has generated. Both ratings appear in brack-
ea after the pagination in mene onion: ra Pom a appears in his es for the ten most
uentl ae que tly cite ed. The addi-
tion of ie bibliometric sae makes this bibliography not only mate ae also sojecpe cally useful
the
v
due t relative i nportance or value of each entr
The ee indexes, covering general, geographical, and peanteral subjects, respectively, gen-
erate over 20, 000 retertals to the items in ie e peat Index | is alphabetically tas by
g ndex Ill contains nine taxonomic lists. The index-
cross-referrals | y of the per concepts in the devel oping field
su abject Index
fo)
SIDA? 20(2 ): 868. 2002
BOOK REVIEWS 869
of biodiversity are associational. The other major feature of the indexes derives from the bibliometric
approach taken in Bibliography I. Item numbers in the Indexes that are underlined represent works
which were rated with the higher scores. This feature will male it easier to pHonnize readings. The
work also surveys closely related topics from such allied f , nature phi-
losophy, environmental law, a erase ecology, evolutionary biology, biometrics, sustainable
development, environmental policy, forestry, pone education, climatology, and paleobiology. It
provides a good or oy erview of important research and will make the literature review process easier.
While‘ Studies: A Bibliographic Review,” is highly recommended for scientists, eae
cators and professionals-to-be, it cites a fair amount of material that can be understood by under-
graduates or even high school students. This book would be a valuable addition to library ae
collections and should be in all graduate and professional ea, ais —Gary L. Jennings,
Botanical Research Institute of Texas, Fort Worth, TX 76102-4060, U.S
DIOQILVEToIL
Mary Toomey and Everett Leeps. 2002. An Illustrated Encyclopedia of Clematis.
(ISBN 0-88192-508-X, hbk.). Timber Press, Inc., 133 S.W. Second Avenue, Suite
450, Portland, OR 97204-3527, U.S.A. (Orders: www.timberpress.com, 800-
327-5680, 503-227-2878, 503-227-3070 fax). $59.95, 428 pp, 652 color photos,
5 b/w illustrations, 15] line illustrations, 2 maps, 81/2" x 11"
An Illustrated Encyclopedia of Clematis was written by distinguished Clematis experts Mary Toomey
and Everett Leeds and supported in spirit and publication by the British Clematis Society. Their goal
was to produce a reliable, substantial and definitive work on the genus. This comprehensively illus-
trated work will meet those needs, particularly of passionate ericenets, Ege and nurseries.
The popularity of the genus Clematis as a garden plant r in the s in part
because of the extraordinary diversity of its flowers and in part because sadeners were exposed toa
better and wider — one age powered cultivars from all corners of the world. Clematis, be they
cultivars or sp n bloom almost every month of the year. Add the astonishing
range of fe ae and the size nd shape of the flowers and it becomes apparent why clematis
has been justifiably crowned queen of the climbers. They are very accommodating garden plants.
Drama, rhythm, and movement can be created in a garden of any size or ae Widely known as
climbers, the genus also includes shrubby, and herbaceous species and cultiva
It is the first-ever encyclopedia on the genus and includes more than Se rca: worthy spe-
cies and hybrids of the more that 800 listed in the Hoye Hortic ache Society’ Plant Finder (1999-
2000 ed.). Prof a aie ae isan full ~color il stration for each of the species and cultivars
described located n th
This long- ae book is sateen into two Parts. The first Part is devoted to the care and
cultivation : clematis, history and botany, pruning, propagation and hybridization, pests and dis-
eases, anda chapter devoted to growing clematis in North America. The second Part provides an A-
Z directory of more than 550 plants, from the acclaimed _ flowered forms to the less familiar
shrubby, sub shrubby, and herbaceous pened t are beginning to be available. Using everyday lan-
guage, the authors provide plant ee ihe include information on synonyms, origin, plant
habit and oe flowers, nin Broup, flowering period, cultural requirements, recommended
uses in the gar and h s. Several usemua ppendices assist gardeners in choosing the
ae one. for “a ane site and in obtaining reliable p d seeds. A list of nurseries is included
which specialize in ae order, both foreign and domestic. There is also information included to en-
clematis by name or by color. The text is accompanied by more than 650
SIDA 20(2): 869. 2002
870 BRIT.ORG/SIDA 20(2)
Lad ] 1;
magnificent color photos indispensable reference and guide. This book
is written by gardeners with a passion for ices wonderful, colorful plants for gardeners with a pas-
sion to ey nem It is een al ae for all gardeners and amateur growers, as well as the nursery
libraries serving them.—Gary L. Jennings, Botanical Research Institute of Texas,
For Worth, TX 76102-4060, U.
MAGNUS JOHNSON. 2001. The Genus Clematis. Edited and Translated by Bengt
Sundstrom. (ISBN 91-631-1030-X, hbk.). Magnus Johnson Plantskola ABand
Bengt Sundstrom, Branningestrandsvagen 63, 151 39 SODERTALJE, Swe-
den. (Orders: 08-550 979 77; +468 550 97077). $110.00, 896 pp, 24 gouaches,
about 100 illustrations from classical literature and ink drawings by the
author, about 170 color photos, 6 3/4" x 91/2"
Che Genus Clematis, by Magnus Johnson, isasummary of the knowledge the author acquired during
his career as a botanist, picau ok een and nurseryman about clematis. He purpose was to pro-
v = a histor ical review of i S tivation in gardens and to present an ordered arrangement
sof tl een eu ieane
ol d Cie >| Ut C
There are pli Ants esteemec “dies their Gees BHEsO! ane vanes, others because they are
ted itl af ] -__ ] } teof cnr o ] )]
3 - ° oO
prairies reaching to the sun, and still others charged with sy aliclicm—s hite ies red poinsettias
and blue forget-me-nots. A clematis has its effect by its immediate beauty. The shape of the flowers,
the infinite variation of color, their graceful and elegant foliage. One may experience this through
the beauty and structure exhibited in the gouaches and ink drawings of vera ane s repro-
duced in this work. He was awarded the Grenfell Medal by the Royal Horticultural Society for his
gouaches.
The book is presented in three parts. The first deals with the history, horticultural aspects, and
the cultivation and propagation of clematis. The second part is a scientific description of clematis
and gives an overview of different characteristics in text and drawings. It also includes a historical
review of the genus classification by different scientists cumulating with the author's view. In the
third part, each section is introduced by a description of general characteristics, maps showing the
distribution of each section, and comments on the use of Clematis in gardening. Each description
begins with references to the literature cited, with a complete bibliography oe the descrip-
tion section. A total of 325 species are described with around 180 varieties and 1,400 cultivars treated.
Ab
glish-« German-French-Latin and -Swedish listed.
dasic terminology of particular words used in the text is included, with ane pee words, in En-
Originally published in 1996 in Sweden, this transl ation undertaken by Johnson's son-in-law.
He has delivered with Hair both the charm and int c beauty of the work. Dr. Mary Toomey as-
sisted in turning the translation into readable ee ae in developing the chapter on history, tax-
onomy and characteristics.
This book is recommended for serious gardeners, nursery professionals, and botanists and re-
searchers who wish to understand clematis heed Highly recommended for libraries serving this
population. It is too early to declare that this is a classic but it bears all the al of one.—Gary |
Jennings, Botanical Research Institute of Texas, Fort Worth, TX 76102-4060, U.S
SIDA 20(2): 870. 2002
BOOK REVIEWS 871
JAMES F. FRALISH and Scott B. FRANKLIN. 2002. Taxonomy and Ecology of Woody
Plants of North American Forests (excluding Mexico and Subtropical Florida).
Illustrations by Hyosun Park. (ISBN 0-471-16158-6, hbk.) John Wiley and
Sons Inc. One Wiley Drive, Somerset, NJ 08875, U.S.A. (Orders: 732-469-
4400, 732-302-2300 fax). $100.00, 624 pp, 550 line drawings.
Advances and developments in forestry and wildlife management in recent years have been borne
out of vast amounts ef seenhlic. information and heightened public awareness of the value of our
a} |
forests
he taxonomy and community ecology of North
American forests. It begins witht I t that appropriately introduce reader to taxonomy and
forest ecology. Key Sole ea terms, sack a8 population vs. community, are Canty defined, a section
on forest comm unity ecology
introduced. Taxono be one of the major themes: in this book, with more scuoliagien on the
ee ui ee, et ee on iaicle sight identification of only important commercial trees.
he book is generally divided into four major parts: 1. Taxonomic Concepts and Methodology,
Il. a nosperms (Division Pinophyta), If. Woody Angiosperms (Division Magnoliophyta), and IV.
Forest Community Ecology: Combining Species into Communities. Part I begins with the funda-
mentals of plant classification. Definitions of family, genera, species are provided, as well as explana-
tions of latin binomials and author names. Proper techniques for collecting, labeling and preserving
plant specimens are explained. Finally, the authors instruct on how to construct and use taxonomic
keys.
th
In Part Il, gymnosperms are considered. The section begins with an overview of gymnosperms,
’ Ds
which includes, for example, sections on evolu
ionary history, wood composition and g gecerapute
range. The following ee is a discussion of vegetative and reproductive structure
—
four chapters are comprised of taxonomic descriptions of gymnosperm species in the baie Taxales,
Ginkgoales and Cycadales.
Part I deals with woody angiosperms, beginning like part I] with an overview of the group.
Information is included on life form, morphology and the taxonomic classification that follows
Cronquist. There are two chapters on vegetative morphology and one chapter on reproductive mor-
phology. The following eight chapters are dedicated to taxonomic descriptions. As in part II, taxo-
nomic descriptions begin with a short discussion of famihal characteristics, the number of taxa and
sd vith common Names, a
fab)
taxonomic affinities. After each family are individ
comprehensive, yet not overly complex ca. a eee, oe economic cae
tance, ele distribution and cover types. Cover types can be cross-referenced to pa
t, Part Bvt eepieto lores comtaimity eCO08), integrating the gymnosperm and angiosperm
tl} AGE,
species eons discussed int ties. The first eee in this section explains the differ
ent factors that contribute to forest com scsition: Factors such as climate, soil, topography and geol-
ogy are considered. Moreover, a discussion of forest ee systems is put forth. To identify
communities within natural vegetation regions, the authors employed the National Vegetation Clas-
sification Standard developed by the Nature Conservancy and developed their own naming strategy
=
or the various communities described, which coe! mos on Society of American Foresters’ cover
type system. The last five chapters of this book des rent forests in North America. Gener-
ally, these include Northern Conifer-Hard wood, Cael Hardwood Southern Pine-Hardwood, Rocky
Mountain eae and pes and Southern Coniter Forests. The pools ends with g oe of tech-
nic cal ter names. anc ] Su bje Cc t in ndice
ae review would be oe se without some r ecognition of As note provided in
this book. With 550 line drawings, the majority of the species des erein are illustrated. These
SIDA 20(2): 871. 2002
872 BRIT.ORG/SIDA 20(2)
illustrations are informative, expository and beautiful. While they enhance the aesthetics of the book,
they also tremen lously increase the scientific value by providing a clear identification tool to sup-
port each species description. With more than 800 species described, this book is absolutely packed
with information on woody ae ae sul caren oa nature of this book makes it invaluable and
a necessary text for any classes | gy, taxonomy, forestry, botany
or environmental studies are taught. Fralish and Franklin have clearly set a high senda | or the
future of dendrology texts.—Amy Trauth Nare, Botanical Research Institute of Texas, 509 Pecan Street,
Fort Worth, TX 76102, U.S.A, amy_trauth@yahoo.com.
Donatp Watts. 2000. Elsevier’s Dictionary of Plant Names and their Origin. (ISBN:
0444503560, hbk.). Elsevier Science B.V., Sara Burgerhartstraat 25, P.O.
Box 211, 1000 AE Amsterdam, The Netherlands. $209.50, 1001 pp, 6 1/2" x
91/2",
dictionary contains about 30,000 elas literary names of pene\ pean of ene ish
origin, a although American, Australian, New Zealand, and South African names are included), both
ild ted, with their botanical name and a brief account ot sa names desis e known,
Knowledge of cee insight into largely forg liefs were prol
ably the result of some taboo aint ie: the pant or eating the fruit. Names may also be purely
descriptive, and ve the botanical name. Literary, or “book” names,
have also been included in this licens as being a very important part of the whole. That is, names
f plants that were written but never used in the vernacular. Names in many instances provide links
to oe transmission of words through the ages.
The v lation of names from a wide diversity of sources. An
entry is nee by the vernacular name followed by its botanical (Latin) synonym. This may be
expanded by the addition of a vernacular synonym and the source of the primary epithet. Users
must reference the source by referring to the bibliography at the front of the dictionary. There is
hen a vernacular name is known but not the converse. There is no index
access toa botanical name w
of botanical names. There is me emain y lor vernacular ‘binomials’. Each part has its own
try. There are no cross-references, thus Aaron’s Beard and Beard,
complete, and | largely duplicated, e
Aaron's share the information. A All of the vernacular synonyms have their own individual entries
rather than being grouped together under a common botanical name
work was conceived as part of the author's wider interest in plant and tree lore, and ethno-
botanical studies. A labor of love, a monument to the author, but its bulk, poor editing, out-of-date
botanical nomenclature, and oe oes serve to make this less than an adequate choice for most
I lso weighs against a potential purchase. It is
7
libraries or researchers. I
not only British in scope, but didi iaecd towards southern Britain. Although many of the names
listed are current, most belong in the past, and the author makes no attempt to indicate which are
current. This work is recommended only for those very large libraries that must have everything
available on the subject.—Gary L_ Jennings, Botanical Research Institute of Texas, Fort Worth, TX 76102-
4060, U.S.A
SIDA 20(2): 872. 2002
BOOK REVIEWS 873
Davin J. SCHMIDLY. 2002. Texas Natural History: A Century of Change. (ISBN O-
89672-469-7, hbk.). Texas Tech University Press, PO. Box 41037, Lubbock,
TX 79409-1037, US.A. (Orders: 800-832-4042, wwwittup.ttuedu/). $39.95,
576 pp, 144 b/w figures, 43 maps, notes, 7" x 11".
With the publication of “Texas Natural history: A Century of Change,” Dr. Schmidly has done a great
service for both professional and amateur naturalists in Texas. The bulk of this volume is a re-publi-
cation of Vernon Bailey's “Biological Survey of Texas.” The survey was conducted between 1889 and
1905 by Bailey and twelve “federal agents” and was published in 1905. The Survey has been out of
print since the 1920s, so it has been inaccessible to older naturalists and essentially unknown to
younger naturalists. The Survey contains a wealth of information that is valuable for the historical
perspective it as well as forming a basis of comparison with the current status of biological
diversity in T
The first chapter of this seven-chapter b
o the initiation ae the biological suey of te Dr. eebnnehy ae a detailed account of ne
| he historical setti J} event leading up
ves. I
a involved, their personalities, and their }
information that it may be bewildering for readers who have little “ ledge of the important fig-
ures in the history of natural history. In spite of the voluminous information in this chapter, some
details that would be of considerable interest are lacking. In addition to the people, Dr. Schmidly
covers the political situation and the government agencies involved in the effort to document the
nation’s biological resources. One of the motivations of this effort was agricultural concerns, espe-
cay in the western states, so much of the survey was focused on plant communities and mammals,
| impac ts are potentially great. Thus Dr. Schmidl ys approach i is de-
cidedly, admittedly, and appropriately centered on mammals
he second chapter is the re-publication of the Biological Survey of Texas, which covers pages
51-266 of the book. The first 37 pages of the Survey describe “faunal areas” or “life zones” of Texas,
and outlines their characteristic flora and fauna. This section of the Survey contains much valuable
information for those interested in the capa zones =e) me state and their asroniated vertebrate
fauna. The next several pages lizard
ae omitted. The — of the Survey is a detailed Selanne of the biology distribution, a
eonsenya on tine of te ii a of mamima)s By a any ptandard, ea isan imp to
1undet diti vere primi-
tive at best. pager nine new taxa of mammals were sae ribed ei Gaoucs the efforts of the survey team,
nine of which were first described in the Survey itself. As Dr. Schmidly points out, all but nine of
these species are currently recognized as valid, which is testament to the scientific ability of the
federal agents. Obviously the mammal section of the Survey contains much information of histori-
a ae sue as ae of large carnivores and ungulates, but it also contains information that
tribution and abundance of rodents, bats, “varmints,” and other species of
current ee interest.
n chapter 3, “Annotations of the Biological Survey of Texas” Dr. Schmidly updates the informa-
tion presented in the Survey, and includes changes in taxonomy, nomenclature, distribution, and
undance, as well as comments on conservation status. The information comes from the scientific
oe and the author's decades of experience with Texas mammals. This chapter will be of con-
siderable use to professional mammalogists in comparing the information in the survey with cur-
rent Ree des ane in Lay eta in 1 ae of craeaeuae and Seca For ae in-
d biolog
sae 3 givesa io at the give a =e ae which taxonomy and systematics proceeds, and
SIDA 20(2): 873. 2002
874 BRIT.ORG/SIDA 20(2)
at changes in the fauna as well as changes in the scientific study of the fauna. We al sional]
get a glimpse into the past that highlights the professional activities of Siew or old friends.
hapter 4, “Texas Landscapes, 1889-1905” gives Bailey and the federal agents the opportunity
to tell us what they saw in their own words, and through their photographs show us the Texas land-
scapes they saw. This chapter will fascinate anyone interested in biogeography and the biotic dis-
tricts of Texas. The many ees and first-hand descriptions reveal landscapes that were in
many instances already heavily transformed by human activity. According to Dr. Schmidly, “Of all
t
blackland prairies of north central Texas are barely mentioned in the survey. This should not be
ne community types..the greatest ae impacts appear to have been on grasslands.” In fact, the
surprising because the blacklands are among the most fertile areas of the country, and had been
converted to croplands long before the es of Hie pieloeiea, survey. In reading the a accounts and
seeing the photographs, it appears that some Tk vere more heay ily transformed in the
late
800s than they are today. This realization gives hope a at remnants ol each of the “life zones” of
Texas may be er Unfortunately, other information in this chapter makes it clear that there is
little hope that Texas biological communities can be restored toa state even vaguely similar to their
condition before settlement by Europeans
Chapter 3, “Twentieth Century Changes in Texas Landscapes and Land Uses” continues the
development of the idea that although the 20" centur ? was one ol dramatic change in Texas biota,
|
the reader should not assume that there had been no changes | tothe years of the survey. In fact,
one of the most important points of this work is that m ae of the Texas landscape had been altered
substantially belore the survey was conducted. Another important point is that not all of the changes
t
jat have occurred since the survey were lor the worse. Many changes in the 20" ay Have come
about through conservation and restoration efforts by government agencies, private organizations
and concerned landowners
r 6 on “Twentieth Century Changes in the Texas Mammal Fauna’ will probably prove to
bet
that
1e Most useful to prolessional mammalogists. T his chap pter discusses several categories of changes
nave occurred, including local (e.g. within Texas) t dramatic declines in abundance,
changes in distributions, and discovery of new ta
ee on the introduction of exotic em for show and hunting (the “texotics”) and on
the | | tof mammalogy in Texas will interest the amateur naturalist. The section
on iptioructog of sonaiee nous species” appears to take Dr. Schmidly pretty far afield; why not
just say “exotic” rather than “nonindigenous”? The discussions of the red imported fire ant and the
brown tree snake barely scratch the surface of the literature on these exotic species and their effects
on endemic biota
Finally, in chapter 7, Dr. Schmidly discusses “Challenges for Wildlife Conservation in Texas.”
This chapter is insightful in some places but frustratingly vague and incomplete in other places. In
the first pages the discussion leaves unanswered or glosses over some potentially substantial areas of
controversy, for instance fhe serie (and conflicting) uses of the term “sustainable” and how
sustainability figures into conservation strategies. The chapter discusses “new perspectives in con-
servation,” many of which are strategies that have been discussed in conservation circles for years.
The focus in the latter part of the chapter seems to be on “managing wildlife diversity,” alt
though
what oe as wildlife needs to be managed is not made clear. The emphasis seems to be on large
1]
r} “]
rgame animals, but this is not specified. The discussion of
ing wildlife diversity” armen series of strategies for pr servation of biological cae
sity which have appeared in the conservation literature in the recent
pa
It is impossible to oo a book of this scope without errors creeping in, and this work is no
exception. Some errors are a result of inattentive editing, aa) asty Se eens of which
are quite noticeable), but others are a result o les on page 47
oO Jt oO
SIDA 20(2): 874. 2002
BOOK REVIEWS 875
and captions for figures that a match the figures. Many of the black and white photos repro-
duced in the book provide a valuable perspective, but there are several that are of such poor se
that they detract from the presentation more than they add. In some sections is seems
Schmidly is unsure whether his audience is prolesslonal Pighogists or informed amateurs. oe
sionals would like to see more precise use of scientific vocabulary (e.g. variation, communities, adap-
tation) and more documentation from the primary literature, where amateurs would prefer clear
and concise eee of events and processes without tecnica yar eon
] ] | Ba |
In spite of these minor criticisms, this book i
understanding of Texas natural history, and provides a perspective on changes in the biota ae Texas
with applicability far beyond its borders. Because “east meets west and north meets south” in Texas,
| have occurred in other
many of the changes that occurred and the processes that caused them wi
places, and the understanding of the changes and processes that this book provides will inform sci-
A great serv ice for professional natural-
3
entists gee in those areas. Dr. Schmi dly has ind
ists in Texas and elsewhere.—Steven Goldsmith, De partment of Biol ogy, Austin College, Sherman, TX
75090, U.S.A. sgoldsmith@austincollege.edu
Q.
SIDA 20(2): 875. 2002
876 BRIT.ORG/SIDA 20(2)
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TABLE OF CONTENTS
Systematics
The resurrection and lectotypification of Tetrazygia fadyenii (Melastomataceae:
Miconieae): a hummingbird-pollinated treelet endemic to Jamaica
Darin S. PENNEYS and WALTER S. JUDD—877
Nomenclatural adjustments and comments in Abronia and Acleisanthes
(Nyctaginaceae)
RICHARD SPELLENBERG and JAckiE M. Poo_E—885
Yucca cernua (Agavaceae: series Rupicolae), a new species from Newton and
Jasper counties in eastern Texas
Eric L. KeitH—891
Two new species of Parathesis (Myrsinaceae) from Mesoamerica and neotypi-
fication of Parathesis rothschuhiana
Jon M. RickETSON and JOHN J. Pipoty I1I—899
Lectotypification of Schlechtendal Myrsinaceae from Schiede and Deppe
collections
Jon M. RickETSON and JOHN J. Piroty I1I—909
A synopsis and nomenclator of the genus Parathesis (Myrsinaceae)
Jon M. RickETSON and Joun J. Pipoty [HI—913
A new species of Stylogyne (Myrsinaceae) from Darién, Panama
Jon M. RickeETson and Joun J. Pipoty II]—919
Pteroceras monsooniae (Orchidaceae), a new species from India
N. SASIDHARAN and P. SuJANAPAL—923
Symplocos kothayarensis (Symplocaceae), a new species from peninsular India
V. SUNDARESAN, S. RAJKUMAR, G.J. JorHi, and V.S. MANICKAM—927
Sinopsis del género Meliosma (Sabiaceae) en Costa Rica y Panama, con tres
nuevas especies
J. FRANCISCO MORALES—931
A new species of Packera (Asteraceae: Senecioneae) from the Edwards Plateau
of Texas
Depra K. Trock and Ropert J. OKENNON—945
Studies in Glandularia (Verbenaceae): the Glandularia quadrangulata complex
JAMES HENRICKSON—953
The bracteoles in Amaranthus (Amaranthaceae): their morphology, structure,
function, and taxonomic significance
Minal Costed and Francois J. TARDIF—969
Nomenclatural changes in the genus Polygonum section Polygonum
(Polygonaceae)
Minat Costea and FrANcols J. TARDIF—987
Taxodiomeria (Taxodiaceae), an intergeneric hybrid between Taxodium and
Cryptomeria from Shanghai, People’s Republic of China
ZHANG JIAN-JUN, PAN SHI-Hua ZHu, WEI-JiE, Niu Hur-Juan, YE ZHENG-J1, ZHU JIAN-Hua,
and Hsu PING-SHENG—999
A new species of Dioscorea (Dioscoreaceac) from the Andes of Venezuela
LJ. Dorr and Basi STERGIOS—1007
Festuca dentiflora (Poaceae: Loliinae: sect. Glabricarpae), a new species from
Peru and taxonomic status of FE. presliana
DaNieL STANCiK and PAuL M. PETERSON—1015
The genus Packera (Asteraceae: Senecioneae) in Colorado, U.S.A.
Desra K. TROocK—1023
Nomenclatural combinations in North American Panicum subgenus
Dichanthelium and subgenus Agrostoides (Poaceae: Paniceae)
JosePH K. WipFF—1043
Development and Structure
Perforation plate diversity in Mlicitum floridanum (I]liciaceae) with respect to
organs, provenance, and microtechnical methods
Epwarb L. SCHNEIDER and SHERWIN CARLQUIST—1047
—
Comparison of the morphology, flowering phenology, and life cycle type in plants
of Grindelia lanceolata (Asteraceae) from cedar glades in middle Tennessee
and northern Alabama: a common garden study
CuristopHER A. ADAMS, JERRY M. Baskin, and Carot C. BASKIN—1059
Botanical History
The name of the amaranth: histories of meaning
MinAl Costea and FRANcoIs J. TARDIF—1073
Documented Chromosome Numbers
Documented chromosome numbers 2003: 1. Chromosome number of
Toxicoscordion nuttallii (Liliales: Melanthiaceae) and clarification of the genus
Wenpy B. ZOMLEFER—1085
Floristics and Ecology
A community-level floristic reconnaissance of the Raja Ampat Islands in
New Guinea
W. TAKEUCHI~—1093
Soil-dependent fire frequency: a new hypothesis for the distribution of prairies
and oak woodlands/savannas in North Central and East Texas
GeorGE M. Diccs, Jr. and PETER C. SCHULZE—1139
Flora and vegetation of the Mohawk Dunes, Arizona
RICHARD STEPHEN FELGER, DALE SCOTT TURNER, and MICHAEL EF WitsoN—1155
A floristic survey of Big Branch Marsh National Wildlife Refuge, St. Tammany
Parish, Louisiana
Davip J. Rosen and STANLEY D. JoNEsS—1189
The vascular flora of the Ogallala ecotone on the Dempsey Divide, Roger
Mills County, Oklahoma
CRralG C. FREEMAN and Cates A. MorsE—1217
West Gulf Coastal Plain ecoregions
MicuaeL H. MacRoserts and BarBara R. MACRoBERTS—1247
The John D. Freeman Herbarium (AUA): a history and catalogue of vascular
plant type specimens
Curtis J. HANSEN—1277
New vascular plant records for the grassland biome of central North America
CRAIG C. FREEMAN, CALEB A. Morse, and RONALD L. MCGREGOR—1289
New records of Corchorus (Tiliaceae) for Mexico
Jost AURELIO COLMENERO Roses and RAFAEL FERNANDEZ Nava—1299
Addition to Flora Mesoamericana: a new record of Thelypteris
(Thelypteridaceae) for Chiapas, Mexico
MicueL A. PEREZ FARRERA, RAMON RipA, and Maria E. Lopez-MoLiNna—1311
Polygonum douglasii subsp. johnstonii (Polygonaceae) new for Canada
Mrinal Costes and Francois J. TARDIF—1317
Koenigia islandica (Polygonaceae) new for Utah
Minal Coste and FRANCOIS J. TARDIF—1319
} ]
Notes on two species of Agalinis (Sc ropnulat iaceae) including Agalinis
harperi, new to Texas
Eric L. KeitH and JouN Hays—1321
Oxypolis ternata (Apiaceae) deleted from the Texas flora
Bruce A. SorriE, MICHAEL H. MAcRoserts, BARBARA R. MACRoserts, and SUZANNE
BIRMINGHAM WALKER—1323
Another juniper (Cupressaceae) in Jeff Davis County Texas? We're not convinced
JOHN P. Karces and JAMes C. ZECH—1325
A rockhouse microhabitat in the West Cross Timbers of North Central Texas
GeorGE M. Dicas, Jk. and Ropert J. OKENNON—1327
Commemorative
William Dean Reese (1928-2002)
RONALD A. PurseL_t and PAuL L. REDFEARN—1335
Book reviews and notices 890, 908, 912,944, 952, 968, 986, 998, 1014, 1042,
1058, 1072, 1084, 1154, 1188, 1246, 1288, 1298, 1310, 1316, 1320, 1326, 1334, 1339
Announcements 1341
Index to new names and new combinations in Sida 20(3), 2003
Abronia umbellata var. breviflora (Standley) L.A. Galloway, comb. et stat. nov.—888
Acleisanthes lanceolata var. megaphylla (Fowler & B.L. Turner) Spellenb. &
J. Poole, comb. nov.—886
Dioscorea lisae Dorr & Stergios, sp. nov.—1007
Festuca dentiflora E.B. Alexeev ex Stancik & P.M. Peterson, sp. nov.—1016
Meliosma chiriquensis J.F. Morales, sp. nov.—934
Meliosma clandestina J.- Morales, sp. nov.—934
Meliosma depressiva J. Morales, sp. nov.—937
Packera texensis R.J. O’Kennon @ D.K. Trock, sp. nov.—945
Panicum aciculare var. neuranthum (Griseb.) Wipff, comb et stat. nov.—1044
Panicum aciculare var. ramosum (Griseb.) Wiplf, comb nov.—1044
Panicum acuminatum var. ferventicola (Schmoll) Wipff, comb. et stat. nov.—1044
Panicum acuminatum var. thermale (Bol.) Wipf, comb. nov.—1044
Panicum commutatum var. equilaterale (Scribner) Wipff, comb. et stat. nov.—1044
Panicum ovale var. praecocius (Hitchcock @ Chase) Wipff, comb. nov.—1044
Panicum rigidulum var. abscissum (Swallen) Wipff, stat. nov—1045
Parathesis pseudocalophylla Ricketson & Pipoly, sp. nov.—899
Parathesis pseudocrassiramea Ricketson & Pipoly, sp. nov.—902
Parathesis subg. Laterales Ricketson & Pipoly, subg. nov.—915
Parathesis subg. Laterales sect. Calophyllae Ricketson & Pipoly, sect. nov.—915
Parathesis subg. Laterales sect. Laterales Ricketson & Pipoly, sect. nov.-915
Parathesis subg. Parathesis sect. Cubanae Ricketson & Pipoly, sect. nov.—916
Parathesis subg. Parathesis sect. Versatiles Ricketson & Pipoly, sect. nov.—917
Polygonum aviculare subsp. buxiforme (Small) Costea & Tardif, comb. et stat.
nov.—988
Polygonum fowleri subsp. hudsonianum (Wolf & McNeill) Costea & Tardif,
comb. nov.—994
Polygonum humifusum subsp. caurianum (Robinson) Costea & Tardif, comb.
et stat. nov.—995
Polygonum ramossisimum subsp. prolificam (Small) Costea & Tardif, comb.
et stat. nov.—995
Polygonum striatulum vat. texense (M.C. Johnston) Costea & Tardif, comb. et
Stat. nov.—996
Pteroceras monsooniae Sasidharan & Sujanapal, sp. nov.—923
Stylogyne pucuroensis Ricketson & Pipoly, sp. nov.—919
Symplocos kothayarensis Sundaresan, Jothi, Rajkumar & Manickam, sp. nov.—928
x Taxodiomeria peizhongii Z.). Ye, J.J. Zhang et S.H. Pan, gen. et sp. nov.—1001
Yucca cernua Keith, sp. nov.—892
THE RESURRECTION AND LECTOT YPIFIGATION OF
TETRAZYGIA FADYENII (MELASTOMATACEAE:
MICONIEAE): A HUMMINGBIRD-POLLINATED TREELET
ENDEMIC TO JAMAICA
Darin S. Penneys Walter S. Judd
Ur river sity of Flor ida Ur dive! sity Ul Flor ida
wi tment of Botany Department of Botany
220 Bartram Hall, PO. Box 118526 220 Bartram Hall, PO. Box 118526
Gainesville Fl 32611-8526, U.S.A. Gainesville, FL 32611-8526, U.S.A.
dpenneys@ufl.edu wjudd@botany.ufl.edu
ABSTRACT
In the course of a cladistic analysis and systematic revision of Charianthus (Melastomataceae:
Miconieae), it was discovered that one species, C. fadyenii (Hook.) Griseb., is unrelated to the other
members of that genus, and this species is thus reassigned to Tetrazygia
RESUMEN
En el curso de un analisis cladistico y revision sistematica de Charianthus (Melastomataceae:
Miconieae), se descubrié que una aces © ie venti (Hook.) Griseb., no esta relacionada a los otros
miembros de este género y por signadoa letrazygia.
INTRODUCTION
Tetrazygia fadyenii is a very attractive melastome endemic to the Greater
Antillean island of Jamaica. This species was first described by W. J. Hooker
(1849) as Tetrazyeia fadyenii. A. H. R. Grisebach (1864), misled by superficial
morphological adaptations common to hummingbird-pollinated plants, trans-
ferred the species to Charianthus Don. The incorrect placement of this species
(as C. fadyenii (Hook.) Griseb.) persisted through subsequent classifications and
floristic treatments by Triana (1871), Cogniaux (1891), Hodge (1941), Adams
(1972), and Howard (1989). Judd and Skean (1991) recognized the non-mono-
phyletic circumscription of Charianthus, and transferred this species to Miconia
sect. Miconia. Convergent evolution, likely driven by parallel shifts to bird pol-
lination in Charianthus and Tetrazygia fadyenti, has resulted in the presence
of relatively large, red, pseudocampanulate corollas and nectar production in
both taxa. We note that J. Dan Skean, Jr has observed hummingbirds visiting
flowers of T. fadyenii (pers. comm.; Skean & Samuels 1840).
In the course of cladistic analyses of Charianthus based on both morphol-
ogy and nuclear ribosomal ITS sequences (Penneys 2001, in prep.), it was dis-
covered that numerous morphological and molecular synapomorphies unite
SIDA 20(3): 877-884. 2003
878 BRIT.ORG/SIDA 20(3)
the six species of that clade. As newly circumscribed (Penneys 2001),
Charianthus is the only plant genus endemic to the Lesser Antilles.
Additionally, Tetrazygia fadyenii was found to form a clade with T. bicolor
and T. crotonifolia (and presumably, other species of Tetrazygia). Although T.
fadyenii is currently best placed within Tetrazygia, ongoing investigations by
the authors (and collaborators) on the phylogenetics of the Miconieae are needed
to confirm this placement. The monophyly of Tetrazygia is supported in our
preliminary analyses, but additional investigations, using more species, are
needed. The monophyly of Miconia is doubtful, and recognition of other gen-
—
era within the tribe may make Miconia paraphyletic, but this matter is beyond
the scope of this paper.
Some morphological characters distinguishing Tetrazygia fadyenii from
Charianthus and uniting it with T. bicolor include: the presence of druse crys-
tals (vs. these lacking), anthers opening by terminal pores (vs. slits), a 2-locu-
late (vs. 4-loculate) ovary, presence of mite domatia (vs. such structures lack-
ing), spatulate (vs. broadly ovate) petals, the calyx constricted above the ovary
(vs. not constricted), an adaxial thickening of the anther base (vs. anther base
non-thickened), and rounded branches (vs. flat branches) of the stellate hairs.
The presence of a hypanthium strongly constricted in fruit above the ovary
may be a synapomorphy for Tetrazygia, and is clearly evident in T. fadyenii.
Thus, this distinctive species should again be known by its basionym, I. fadyenii.
Its resemblance to Charianthus is superficial. As outlined above, these simi-
larities presumably reflect convergent adaptations to bird pollination.
It is also of interest that the species of Tetrazygia occur chiefly over lime-
stone, as does T. fadyenii, while the species of Charianthus sensu stricto grow
on volcanic substrates.
Jamaica is home to four species of Tetrazygia, of which three are consid-
ered to be endemic (Adams 1972). This pattern of endemism is the norm for
Tetrazygia throughout its range; the majority of species in the genus are single-
island endemics. Ietrazygia fadyenii can be distinguished from other Jamai-
can members of the genus by the combination of elliptic (rarely widely elliptic
or narrow obovate), essentially glabrous leaves, and bright red petals that are
9.5 to 13.5 mm long.
TAXONOMIC TREATMENT OF TETRAZYGIA FADYENII
Tetrazygia fadyenii Hook., Hooker's J. Bot. Kew Gard. Misc. 1:379, t. 12. 1849.
Charianthus fadyenii Hook.) Grisebach, Fl. Br. W. Ind. Isl. 264. 1864. Miconia fadyenii (Hook.)
W. Judd & Skean, Bull. Florida Mus. Nat. Hist. 36(2):25-84. 1991. LECTOTYPE, designated here:
“Melastoma. Hab - [illegible] District St. Anns [sic]. This is the most beautiful plant | have seen
in Jamaica. Seeds 227.” Macladyen (Kk, H2001/02842-1, fragment Al). ISOLECTOYPES, designated
dadyenii.” (kK, H2001/02842-2, fragment Db);
“Tetrazygia? fadyenit Hook. Kew Journ. 1849. Jamaica. Macfadyen. Hooker 1849." (kx, H2001/
02842-3, fragment Gl).
nere: “Jamaica. Dr. McFadyen Tetrazygia.
an
PENNEYS AND JUDD 879
Nomenclatural note.—Three sheets located at K must be considered in the
lectotypification of Tetrazygia fadyenii Hook. Each of the three sheets contain
three twigs, which have been designated with consecutive letters for clarity,
thus, sheet H2001/02842-1 has fragments labeled A through C, H2001/02842-
2 with fragments D through F and H2001/02842-3 with fragments G through
I. All specimens designated here as lectotype material were seen by Hooker in
1849, or earlier, and were collected by MacFadyen. Proof of this is demonstrated
either by their label data or by representation in the illustration of the species
published by Hooker in 1849.
The BM collection H2001/02842-1 has attached to it the illustration that
accompanied Hooker’s description of the new species (Hooker's J. Bot. Kew Gard.
Misc. 1:379, t. 12.1849.). In that description, Hooker noted that MacFadyen pro-
vided him with specimens of the species, which he then named in his honor.
This sheet also has a label, presumably written in the field by MacFadyen, that
reads, “Melastoma sp. Hab - lillegible] District St. Anns|sic]. This is the most beau-
tiful plant | have seen in Jamaica. Seeds 227.” Written in a different hand at the
top of that label is “Charianthus sp. nov.” Fragment A, a leafy twig in fruiting
condition, is here selected as the lectotype of Tetrazygia fadyenii Hook. This
collection is from the Hooker herbarium, and was annotated by Hooker as T.
fadyenii Hook. A fruit and seed of fragment A are represented as figures six and
seven in the al ioned illustration. Fragment B is in bud and flower, but
cannot be definitively considered to have been seen by Hooker. Fragment C isa
poor, sterile specimen.
Label data are completely lacking on H2001/02842-2. However, fragment
D, at least, was observed by Hooker, who wrote next to that flowering twig, “Ja-
maica. Dr. McFadyen Tetrazygia fadyenii.” This fragment was likely used for
the habit (figure unnumbered) and flower figures two and three of the illustra-
tion cited above, which it very closely resembles, and we consider it to be an
isolectotype. Fragment E is a smaller flowering twig, and F is nothing more
than a scrap with five mature fruits and the date 1850. The sheet is annotated
in an unidentified hand as Charianthus fadyenii Griseb.
From the sheet H2001/02842-3, it may be presumed that Hooker sent material
of Tetrazygia fadyenii to Bentham as this sheet has “Herbarium Benthamianum
1854” stamps next to fragments G and |. Fragment G isa small flowering speci-
men overlaid by a label stating: “Tetrazygia? Fadyenii Hook. Kew Journ. 1849.
Jamaica Macfadyen. Hooker 1849.” It, therefore, appears that this fragment was
included with the material originally under consideration by Hooker when he
described the species. We consider it to be a possible additional isolectotype.
The fruiting (and sparsely budding) twig H cannot definitively be assigned to
either of the other two samples of this sheet. Fragment lis a budding twig with
a label glued over it on which it is written, “Jamaica. Hooker 1843.” The collec-
880 BRIT.ORG/SIDA 20(3)
tor of fragment | is unknown. This sheet is annotated below as “Charianthus
fadyenii Griseb. Triana in Linn. Trans. Vol. 28, p. 99.”
Description.—Evergreen shrubs or small trees to 10 m high. Druse crystals
present. Bark smooth to shallowly, longitudinally fissured; young twigs terete
(slightly quadrangular), gray-brown, essentially glabrous but witha very sparse
indumentum of inconspicuous, stellate and unicellular, uniseriate hairs, the
stellate hairs ca. 0.09-0.13 mm in diameter, nodes (1.93-)2.13-3(-5) mm in di-
ameter, a slightly expanded, inconspicuous interpetiolar ridge present, with
inconspicuous stellate pubescence, the hairs (0.05-)0.08-0.19(-0.21) mm in
diameter. Leaf blade coriaceous, elliptic (widely elliptic or narrow obovate), (3.2)
4.4-6.5(-8) cm long, (1.8-)2.2-3.1(-3.5) cm wide, blades somewhat V-shaped,
margins flat (very slightly revolute), light green, yellowish-green below, with
midrib, secondary veins, and margins sometimes reddish; apex acute (barely
obtuse), base acute, acute cuneate or cuneate (obtuse); adaxial surface of young
blades with sparse stellate hairs (some sunken in pits) and unicellular hairs,
the stellate hairs (0.05-)0.08-0.19 mm in diameter, mature leaves glabrous with
extremely sparse stellate and unicellular hairs, abaxial surface of young blades
with few unicellular or multicellular uniseriate hairs, sometimes with very
dense stellate hairs (0.05-)0.08-0.19(-0.24) mm in diameter, mature leaves ap-
pearing glabrous but with few stellate hairs, often sunken in pits and appear-
ing as punctations (0.05-)0.08-0.24(-0.25) mm in diameter; domatia (occasion-
ally absent) of primary-axillary hair tufts, the domatial hairs multicellular
multiseriate dendritic setae, domatia (0.48-)0.56-2.34(-2.80) mm long, (0.32-)
0.33-1.47(-2.54) mm wide; margin entire; venation opie basal (basal) with (-)
2 pairs of secondary veins, and | pair of i ginal secondary
veins, the tertiary veins very indistinct (10- es 19-21), intergrading with qua-
ternary veins at base and apex; midrib and secondary veins flat to slightly im-
pressed adaxially, tertiary and quaternary veins very indistinct adaxially, mid-
rib raised abaxially, secondary veins slightly raised to flat abaxially, tertiary
and quaternary veins flat abaxially. Petiole (0.51-)0.60-1.35(-1.80) cm long, shal-
lowly canaliculate, often reddish, moderately stellate pubescent, with the hairs
+ evenly distributed on all parts of petiole or somewhat denser abaxially, these
(0.06-)0.08-0.19(-0.25) mm in diameter. Inflorescence terminal, cymose, pan-
iculate (loosely corymbose), (3.6-)5.7-8.8(9.5) cm long, (3.4-)3.5-5.4(-5.5) em
wide, with 2-3 orders of branching, the caducous bracts to ca 3 cm long; pe-
duncle (1.9-)2.2-2.9(-3.6) cm long, first internode above peduncle (1.03-)1.30-
1.80(-2.20) cm long, first lateral internode (1-)1.1-L7(-2.2) cm long, the flowers
(21-)27-43(-76) per inflorescence. Pedicels (4.75-)5.25-7.75(-10.50) mm long,
with stellate hairs (0.09-)0.11-0.16(-0.18) mm in diameter. Hypanthium cylin-
drical to suborbicular around ovary, tubular above ovary, (2.09-)2.75-3.60(-3.63)
mm long as measured from base of hypanthium to torus, (2.07-)2.16-3.14(-3.47)
mm wide, indumentum of stellate hairs, (0.05-)0.06-0.11(-0.18) mm in diam-
—_—
jay
—
eter, the portion of hypanthium free from ovary (0.97-)1.11-L.38(-L.57) mm long;
inner hypanthium smooth (inco! ISspic uously ridged), torus to calyx apex length
(1.72-)2.50-3.54(-3.73) mm, torus to calyx notch length (1.63-)2.19-3.27(-3.32)
mm. Calyx lobes 4, (0.09-)0.16-0.53(-0.69) mm long, (2.13-)2.32-2.97(-3.13) mm
wide, lobes very broadly triangular (often to such a degree that the lobes are
difficult to discern), the indumentum similar to that of hypanthium, the exter-
nal calyx teeth (0.25-)0.33-0.63(-0.67) mm long, usually only a blunt knob on
rim of hypanthium. Petals 4, bright red, oblong spatulate, the apex truncate,
(9.50-)9.88-13.13(-13.63) mm long, (3-)3.13-4(-4.31)) mm wide, glabrous, many-
veined, erect or slightly spreading. Stamens 8, folded adaxially in bud, in two
series, isomorphic, (19-)20-21.5(-22) mm long; filaments pink, slightly tapered
towards apex, 14.5-16 mm long, with proximal filament segment (0.38-)0.41-
0.60(-0.67) mm wide, distal segment (0.22-)0.28-0.50(-0.53) mm wide; anthers
yellow, (4.46-)4.94-5.94(-6.07) mm long, (0.53-)0.60-0.81(-0.82) mm wide, el-
liptic, slightly curving, opening by a single apical pore (0.19-)0.27-0.37(-0.45)
mm wide. Ovary inferior, 2-loculate, (1.94-)1.97-2.94(-3) mm long, (1-)1.13-
141-187) mm wide, with an apical appendage (0.94-)0.97-1.22(-1.25) mm long,
with deeply intruded axile placentation; style pink or red, straight to shallowly
S-shaped, glabrous, (20.5-)21-24(-24.2) mm long, (0.38-)0.41-0.57(-0.60) mm
wide; stigma not expanded, papillose. Berries globose-urceolate, 4-7 mm in di-
ameter, urceolate, deep purple. Seeds pyramidal, 1.35-1.50 mm long, 0.75-1.03
mm wide, the testa smooth. See Figure | (see also Hodge 1941, plate 6, figures 5
and 6).
Distribution and ecology.—Tetrazygia fadyenii is endemic to Jamaica. It
occurs on limestone hills in xeric to mesic woodlands and forest edges from
450 to 950 meters elev. where it may be locally common (Fig. 2).
Phenology.—Tetrazygia fadyenii has been collected in reproductive condi-
tion throughout the year, with the exception of the month of November.
Specimens examined: JAMAICA. Clarendon: 1.5 mi SSW of junction at Cave Valley, 17 Jul 1977,
is (FLAS); Peckham Woods, Upper Clarendon, 7 Jul 1911], ee 10997 (NY, US); Peckham
Upper Clarendon, 27 Sep 1912, Harris 11181 (NY, US); Bird Rock, Glenwood Springs,
ee Sunbury Road, 4 ee om Morley & Whiteford 967 (A); aes ee district, be-
n Balcarres and Sunbury, 18 Aug 1973, Proctor 33485 (IJ). Hanover: Dolphin Head, Aay 1906,
ee 9234 (A, US); Dolphin Head, SW of Askenish, ca. 7 mi S of Lucea, 12 May ine es & Samuels
5300 (FLAS, NY); 12 May 1987, Judd & Samuels 5311 (FLAS, NY); near Askenish, Dolphin H
1975, Judd, Kress & Clarkson 75-141 (A); summit of Dolphin Head, 29 Oct 1952, Proctor sae ai a
Dolphin Head, on the top, 9 Jul 1986, Skean & Samuels 1840 (NY, US); Dolphin Head Mountain, 8 mi
SSE by road from Lucea, 1 Aug 1967, Weaver 1278 (GH, MO); Dolphin Head, 28 Jul 1954, Webster &
Wilson 5071 (A, US); slopes of Dolphin Head, 24 Jun 1959, Webster et al. 8563 (S). Manchester: at Ba-
nana Ground, 14 Jul 1963, Crosby, Hespenheide & Anderson 673 (GH, NY); 0.5 mis NW of Christiana,
3 Jul 1955, Howard & Proctor 14331 (A); between Fairfield and H anlene Jan 1957, Proctor 16106 (GH);
Gourie Forest, ca. 1 mi SW of Coleyville, 18 Apr 1971, Proctor 31748 (J); Huntley Hill, 20 Jul pale
Proctor 37906 (MO). Middlesex: St. Anne. Mt. Diablo, 22 Jan 1938, Hunnewell 15322 (GH). St.
Albion, 14 Mar 1850, Grisebach s.n. (GOET). Holly Mount, near Ewarton, 1] Aug 1896, Harris ie
882 BRIT.ORG/SIDA 20(3)
Fic. 1. Te ia fadyenii. A. Habit. B. Inflorescences. C. Flower. D. Fruits
© ee FADYENII
Fic. 2. Distribution of Tetrazygia fadyenii.
(NY); Holly Mount, Mt. Diablo, 29 Aug 1905, Harris 8994 (NY); Mt. Diablo area; ca. 1 mi along road
towards Resource, 11 Aug 1965, Hespenheide 973 (FLAS, GH, MO, NY, US); Mt. Diablo, 27 Jan 1952,
Snes aS Diablo region, lower slopes of Hollymount, 28 May 1987, Judd 5509 (FLAS,
NY); Mt. Diablo, Jul 1849, McNab s.n. (US); vicinity of Mosely Hall Cave. 1 mi W of Blackstonedge, 14
Dec 1952, Proctor 7480) ie Diablo, Grier Mount, 29 Aug 1954, Webster & Proctor 5640 (A, US); Mt.
Diablo, E gh, 25 Jun 1959, Webster etal. 8530 (S). St. Catherine: Hollymount,
22 Sep 1902, Adams] 1725 (MO), near summit of Hollymount, 5 Jul ney inderson & Sternberg 3048
(US oR Grier Mount, Mt. Diab lo, 20-31 Dec 1953, I foward & Proctor 1360 ; A, I); Grier Mount, sie Diablo,
20-31 Dec 1953, Howard & Proctor 13617 (A). St. Elizabeth: M hae ne 1916, eae 12373 (MO,
NY, P). St. Mary: forests at Union Hill near ens 13 Jul 1950, Howard 12019 (A
a evel District, SE end of John Crow Mts., 16 Mar 1956, Stearn 508 (A); between Row
Big Level; John Crow Mts., 15 Jun 1976, Thorne & Proctor 48305 (NY); John ane Mts., Big Level, above
ani field, 16 Jun 1959, Webster et al. 8281 (GH, VJ, S. US). Trelawny: between Burnt Hill and
Ramgoat Cave, 20 Jul 1997, Acevedo-Rdgz. 9558 (US): the Cockpit Country, near Barbecue Bottom, 14
Aug 1963, Crosby & Anderson 1212 (GH); 2-3 mi N by road of Burnt Hill crossroads, 16 Aug 1965,
Hespenheide 1267 (FLAS, MO, NY); at Ramgoat Cave on road from St. Vincents to Kimloss, 26 Sep
1954, Howard 14130 (A); along road to Mango Tree Hill from Burnt Hill, 27 May 1967, Read 1937 (GH,
US). Location unknown: Apr 1903, Alexander s.n.(GH, NY); Grisebach s.n. (GOET); 15 Feb 1905, Har-
ris 8883 (NY); 27 Sep 1912, Harris 11190 (NY, US).
oy
a
wn
cae field and
Illustrations of this species can be found in Hooker (1849) and Fawcett and
Rendle (1910-1936). Hodge (1941) provided photos of flowers and fruits in plate
six, figures five and six.
ACKNOWLEDGMENTS
We gratefully acknowledge the following individuals: Norris H. Williams and
W. Mark Whitten provided vouchers and material freshly collected in silica gel
that was used in the molecular analyses. Kent Perkins and Trudy Linder at FLAS
kindly provided curatorial services. Thanks also to the staff at A,GH, GOET, JJ,
884 BRIT.ORG/SIDA 20(3)
MO, NY, PS, US. The authors are especially grateful to the staff of Kew for fur-
nishing taxonomically critical specimens from their type collection.
REFERENCES
Apams, C.D. 1972. Flowering plants of Jamaica. University of the West Indies, Mona,
Jamaica.
Coanidux, C.A. 1891. Melastomaceae. In: A. de Candolle and C. de Candolle, eds.
Monographiae phanerogamarum vol. 7.G. Masson. Paris.
Fawcett, W. and A.B. Renote. 1910-1936. Flora of Jamaica. Vol. 5 British Museum (Natural
History), London.
Grisegach, A.H.R. 1864. In: Flora of the British West Indian Islands. J. Cramer, Weinheim.
Hooce, W.H. 1941. The genus Charianthus. Contr. Gray Herb. 135:115-133.
Hooker, W.J. 1849. Hooker's J. Bot. Kew Gard. Misc. 1:378-379.
Howarb, R.A. 1989. Flora of the Lesser Antilles. Vol. 5, Part 2 [Melastomataceae: 532-579].
Armold Arboretum, Harvard Univ., Jamaica Plain, Massachusetts.
Jupp, W.S. and J.D.SkeAn, Jr. 1991. Taxonomic studies in the Miconieae (Melastomataceae).
IV.Generic realignments among terminal-flowered taxa. Bull. Florida Mus. Nat. Hist. 36(2):
—84,
Penneys, D.S. 2001.A systematic revision and cladistic analysis of Charianthus (Miconieae:
Melastomataceae) using morphological and molecular characters. Unpublished M.S.
Thesis, University of Florida, Gainesville, FL.
TRIANA, J.J. 1871. Les Mélastomaceées. Trans. Linn. Soc. London, Bot. 28:1-188.
NOMENCLATURAL ADJUSTMENTS AND COMMENTS IN
ABRONIA AND ACLEISANTHES (NYCTAGINACEAE)
Richard Spellenberg Jackie M. Poole
Department of Biology Texas oi ae Wildlife Department
New Mexico State University 3000 I-35 South, Su i. ie
Las Cruces, NM 88003, U.S.A. ee TX 78704,
rspellen@nmsu.edu Jackie.Poole@tpwd.state.tx.us
ABSTRACT
A new combination is made for A Flora of the Chihuahuan Desert Region and for Flora of North America
North of Mexico in the now more broadly eae Acleisanthes (Nyctaginaceae): Acleisanthes
lanceolata (Wooton) R.A. Levin var. megaphylla (Fowler & B.L. Turner) Spellenb. & J. Poole,
This is based on evidence published Cae . Levin (2000) showing that neither eee nor
os as traditionally treated is eine but together they form a well-defined clade in
Nyctaginaceae. We also provide a short review of the literature showing t
dat Commicarpus
ee (Chiov.) Lebrun a cn is eet an Acleisanthes (sensu lato), based on Selinocarpus
somalensis Chiov. Due to need for consistency in level of infraspecific rank in treatments in Flora o
North America North of Mexico Abronia umbellata Lam. subsp. breviflora (Standl.) Munz is recom-
bined as Abronia umbellata Lam. var. breviflora (Standl.) L.A. Galloway, comb. et stat. nov
RESUMEN
Se hace una combinacion nueva para “A Flora of the Chihuahuan Desert Region” y para “Flora of
North America North of Mexico” en el género Acleisanthes, ahora construido mas ampliamente:
Acleisanthes lanceolata (Wooton) R.A. Levin var. es phylla (Fowler & B.L. Turner) Spellenb. & J. Poole,
mb mbinacion resulta de la evidencia recientemente publicada por Levin (2000),
mostrando que ni Acleisanthes ni Selina como se han tratado ac) sonamnents, son
monofiléticos, pero juntos forman lo en las Nyctaginaceae. También, ofrecemos
citaciones bibliograficas declar sie que Commicarpus somalensis (Chiov.) L oa & Stork es
acral ncnte una Acleisanthe s peas lato), bi aeaeo en Senecanpus somalensis Chiov. A causa de la
los tratamientos de “Flora of North
America North of Mexico” Abronia pinball L: am. subsp. breviflora (Standl.) Munz se ha
recombinado como Abronia umbellata |.am. var. breviflora (Standl.) L.A. Ga
loway, comb. et stat. nov.
INTRODUCTION
The most recent taxonomic revisions of Acleisanthes, Ammocodon, and
Selinocarpus have maintained the genera as distinct (Fowler &@ Turner 1977;
Smith 1976). Acleisanthes (7 spp.) and Selinocarpus (9 spp.) have been consid-
ered distinct genera since their inception; Ammocodon (1 sp.) was originally
described asa Selinocarpus (Torrey 1859). All are from arid or semi-arid regions
in southwestern United States and northern Mexico, mostly on the Chihuahuan
Desert or adjacent semi-arid closely allied floristic areas, except for S.somalensis
Chiov. which occurs in northeastern Africa.
SIDA 20(3): 885-889, 2003
886 BRIT.ORG/SIDA 20(3)
Recently Levin (2000), through the use of nuclear and chloroplast genes,
demonstrated that Acleisanthes and Selinocarpus formed a well defined clade
within the Nyctaginaceae, but individually the genera were not monophyletic.
Levin (2002) then transferred those species traditionally recognized as
Selinocarpus and Ammocodon, and which she recognized as valid taxa, to
Acleisanthes. Levin did not recognize varieties in her study, considering them
insignificant for her work (pers. comm. to J. Henrickson, 2002). Although the
winged fruits of Selinocarpus and Ammocodon are conspicuous in their differ-
ences from the round-ribbed fruits of Acleisanthes, it might be inferred from
Willson and Spellenberg (1977) that a change from ribbed to winged fruit is
not difficult. As discussed by Levin and others, most Acleisanthes are herba-
ceous, whereas many Selinocarpus are suffrutescent or woody. Ammocodon is
herbaceous. All genera produce cleistogamous flowers. In addition, Selinocarpus
diffusus is very difficult to distinguish from A. wrightii vegetatively.
pada
NOMENCLATURE
For an upcoming volume of Flora of North America North of Mexico and for A
Flora of the Chihuahuan Desert Region, we are adopting Levin’s interpretation
of this portion of the Nyctaginaceae, recognizing those taxa traditionally called
Ammocodon or Selinocarpus as Acleisanthes. Turner (in Turner et al. 2003) has
also adopted an expanded concept of Acleisanthes and has made some nomen-
clatural transfers there. Levin (2002) made only species level nomenclatural
transfers. An easily missed error occurred in her work when specific epithets
were transferred in the masculine from Selinocarpus|masculine]to Acleisanthes
feminine]. Corrections necessary are considered orthographic and will be made
in upcoming treatments.
Acleisanthes lanceolata (Wooton) R.A. Levin var. megaphylla (Fowler & BL.
urner) Spellenb. & J. Poole, comb. nov. Basiony: Selinocarpus lanceolatus Wooton
var. megaphyllus Fowlers ee Turner, Phytologia 37:181. 1977. Type: MEXICO. CHIHUAHUA:
Jurassic gypsum ca. 15 mi SW of Estacion Moreon on Rio Conchos Lake Road, Sierra de la
Monillas, 25 May 1971, a 2105 (HOLOTYPE: LL; ISOTYPE: TEX!)
Acleisanthes beso a a (Fowler & B.L. Turner) B.L. Turner in B.L. Turner et al., Atlas Vasc. Pl.
s 7.2003. Selinocarpus megaphyllus (Fowler & B.L. Turner) B.L. Turner. Phytologia 75:242.
1993
The Acleisanthes lanceolata complex consists of obligate gypsophiles distrib-
uted in islandlike populations from northern New Mexico through western
Texas to east-central Chihuahua. There is an irregular cline of increasing leaf
width and less noticeably, perianth size, from north to south, with a more or
less abrupt difference evident in populations south of the Mexico-U.S. border.
Acleisanthes lanceolata var. lanceolata has lanceolate to broadly lanceolate
leaves (larger leaves 5-18 mm wide), whereas A. lanceolata var. megaphylla has
NOMENCILATURAT ANT ACI EICANTUEC 887
ovate to round leaves (larger leaves 12-41 mm wide). In the var. megaphylla pro-
portionately broadest leaves occur especially on the basal half of the stem. It is
this variation that Fowler and Turner were attempting to circumscribe
nomenclaturally when they described the new variety, Selinocarpus lanceolatus
var. megaphyllus. Turner went even further in 1993, elevating the var.
megaphyllus to species level as S. megaphyllus Fowler @ B.L. Turner) B.L. Turner
and later (2003) as Acleisanthes megaphylla (Fowler & B.L. Turner) B.L. Turner
in B.L. Turner et al. He formally recognized intermediate populations in west-
ern Texas and eastern Chihuahua as S. maloneanus B.L. Turner, reducing it to
varietal status again in 2003 as Acleisanthes lanceolata var. maloneana (BLL.
Turner) B.L. Turner. Levin (2002) lists all names in the complex that were avail-
able at the time as synonyms of A. lanceolata.
For A Flora of the Chihuahua Desert Region we will recognize the extremes
of variation in the Acleisanthes lanceolata complex, which more or less corre-
late with geography, as two varieties in A. lanceolata, the var. lanceolata (which
includes A. lanceolata var. maloneana) and var. megaphylla. One collection from
the southern part of the range of A. lanceolata supports the view that these
taxa better represent varieties than species. Within a single population are
round-leaved plants, intermediates, and lanceolate-leaved plants, the last un-
common in the population (Mexico, Chihuahua, Mcpio. Camargo, 13.6 km W
of Camargo on road to Lago Boquilla, on gypsum river terraces, 14 Aug 1974,
Spellenberg and Syvertsen 3758, CHDIR, IEB, MEXU, NMC [sheets 46553, 628141,
TEX. So that future confusion might be avoided, two replicates of the same col-
lection (3758) were independently prepared and accessioned at NMC several
years apart, the first with the label stating distance of the site from Camargo as
8.5 miles rather than 13.6 km, the second stating the distance in kilometers).
Jim Henrickson relates to us in his review of this manuscript that the stated
type locality of Selinocarpus lanceolatus var. megaphyllus is problematic. He
states that Estacion Moreon is correctly Estacion Morrion. There is no Sierra de
Monillas, but there is the Sierra Morrion (misspelled on Henrickson’s gazetteer
map [Henrickson & Straw 1976]as Sierra Gorrion). He writes in his review, “But
there is a lake behind Presa Luis. L. Leon along the Rio Conchos and there is a
road to the south end of the lake about 15 mi SW of Est. Morrién and that is
probably the type locality.
Acleisanthes somalensis (Chiov.) R.A. Levin: To trace the interpretation of
Commicarpus somalensis (Chiov.) Lebrun & Stork (1989) requires access to lit-
erature difficult to obtain by those working in the Chihuahuan Desert region.
We review the situation here. Lebrun and Stork, in making their new combina-
tion, examined a photograph in Chiovenda (1929, Tab. XX XIII, fig. 1) that was
labeled as the type of Selinocarpus somalensis. The type of that taxon, however,
is actually pictured in the adjacent photo (Fig. 2), labeled as Boerhaavia
888 BRIT.ORG/SIDA 20(3)
reniformis [now considered Commicarpus reniformis (Chiov.) Cufod.| The cap-
tions were switched in Chiovenda’s work, as pointed out by Thulin (1994, p.
LIL), misleading Lebrun and Stork. The descriptions are valid and the type is
indeed a Selinocarpus [now as Acleisanthes somalensisl, as indicated by the
ecular data of Levin (2000).
=
mo
Abronia umbellata var. breviflora (Standley) L.A. Galloway, comb. et stat. nov.
BASIONYM: A bronia breviflora Standley, Contr. U.S. Natl. Herb. 12:312, plate 30.1909. Typ: U.S.A.
CALIFORNIA. MENDOCINO Co.: Mendocino, Jun 1989, H.E. Brown 833 (HOLOTYPE: US, not seen;
ISOTYPE: MO}).
With little exception, the rank of variety has been used for infraspecific taxa in
the Nyctaginaceae. The Flora of North America North of Mexico project re-
quires that all infraspecific taxa within a generic treatment be at the same rank.
In Abronia,and the Nyctaginaceae in general, the rank of variety has been most
used and to continue to use rank results in the least nomenclatural disturbance.
One transfer from subspecies to variety is necessary. This new combination
makes the FNA Abronia treatment consistent, replacing Abronia umbellata
subsp. breviflora (Standl.) Munz (Aliso 4:90. 1958).
ACKNOWLEDGMENTS
We thank the following: Jim Zarruchi for providing literature from the Mis-
souri Botanical Garden regarding Commicarpus somaliensis and copies of rel-
evant pages of Atlas of Vascular Plants of Texas, Neil Harriman for his opinion
regarding the gender of specific epithets when transferred from Selinocarpus
to Acleisanthes, Jim Henrickson and Guy Nesom for helpful comments on a
draft manuscript, Roy Gereau for locating an isotype of Abronia umbellata,and
to Leo Galloway for his patience and attention during the long process of pre-
paring FNA Nyctaginaceae treatment. Leo Galloway has asked us to include
this nomenclatural transfer in this note, thus this combination should be re-
garded as “L.A. Galloway in Spellenb. & Poole.”
REFERENCES
CHiovenba, E. 1929. Flora Somala. Sindacato Italiano Arti Garfiche, Rome.
Fow er, B.A. and B.L. Turner. 1977. Taxonomy of Selinocarpus and Ammocodon
(Nyctaginaceae). Phytologia 37:177-208.
Henrickson, J.,and R.M. Straw. 1976.A gazetteer of the Chihuahuan Desert region;a supple-
ment to the Chihuahuan Desert flora. Privately published.
Lesrun, J.-P. and A.L. Stork. 1989. La véritable identité du Selinocarpus somalensis Chiov.
(Nyctaginaceae). Bull. Mus. Natl. Hist. Nat., B, Adansonia 2:213-215
Levin, R.A. 2000. Phylogenetic relationships within Nyctaginaceae vibe Nyctagineae: evi-
dence from nuclear and chloroplast genomes. Syst. Bot. 25:738-/50.
PELLENBERG AND POOLE, NOMENCLATURAL AND ACLEISANTHES 889
Levin, R.A. 2002. Taxonomic status of Acleisanthes, Selinocarpus, and Ammocodon
(Nyctaginaceae). Novon 12:58-63.
Smith, JM. 1976. A taxonomic study of Acleisanthes (Nyctaginaceae). Wrightia 5:261-276.
Tuuun, M. 1994. Aspects of disjunct distribution and endemism in the arid parts of the
Horn of Africa, particularly Somalia. In: JH. Sayani and A.C. Chikuni, eds. Proc. XIlI" Ple-
nary Meeting AETFAT, Malawi.
Torrey, J. 1859. Botany of the boundary. |n:W.H. Emory. United States and Mexican Bound-
ary Survey. P. 170.
Turner, B.L. 1993. New species and combinations in Selinocarpus (Nyctaginaceae
Phytologia 75:239-242.
Turner, B.L., H. NicHots, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas, vol.
1. Sida, Bot. Misc. 24.
Wittson, J. and R. SpettenserG. 1977. Observations on anthocarp anatomy in the subtribe
Mirabilinae (Nyctaginaceae). Madrono 24:104-111.
eer
BRIT.ORG/SIDA 20(3)
Book REVIEW
LINCOLN Taiz and Epuarpo ZriceR. 2002. Plant Physiology, Third Edition. (ISBN
0-87893-823-0, hdbk.). Sinauer Associates, Inc., 23 Plumtree Road, PO. Box
407, Sunderland, MA 01375.(Orders: wwwsinauer.com, 413-549-1118 fax).
$104.95, 690 pp., 542 illustrations, 81/2" x 11".
The third edition of this highly regarded undergraduate textbook has been revised, updated, and
given a facelift. Though preceded by two hefty editions, this third edition is more than 100 pages
shorter than the second. The authors, Tiaz and Zeiger, elected to place two chapters on a dedicated
wera By using ae bs net as a communication tool for some Epon tant plant phy siology topics,
they |
tronic form. Clearly, the Keane Sree the authors’ wilteanes to keep the rard copy version of the
text a wor Bae size ewitnoe r sing its content. Additionally, a
] } |} .
KOC lat
version
wealth of information pertinent to the chapter.
The book is generally divided into three units: ee Traneport and od elouee of Water and Sol-
utes, Biochemistry and Metabolism, and Growth and Developmen e first unit begins appropri-
ately with chapters on plant cells, water transport ie balance. aioe chapters included in this unit
are mineral nutrition and solute transport. Unit two
s filled with information on metabolic reac-
tions. Chapters in this unit are respiration, aon of mineral nutrients, secondary metabolites,
and three
chapters on photosynthesis. Unit three has chapters on cell walls, hormones, control of
flowering, stress physiology, oe rome and light control of development.
Each chapter is subsequently broken into sections and subsections with headings. These head-
ings would undoubtedly prove ee or the student, since the headings are written in the form of
a single sentence summary, giving the student a ‘heads up’ on what they are about to re
anion wee ae net, not only contains two ned not oe in
the textbook, but
L-Thhatter covered in
the text. A list of web topics and essays at ihe end of each chapter in text om to these readings
on the websi
Certainly this varvapiew text has b ignifi l pnporess for i ird edition. The compan-
]
P nee
1QO11
]
i g i rue ae of fess textbooks. ai Physi-
} ] ] -] A; 1] 4 -]
Sy y § their
revisions and editorial work. Without doubt, this book should be a standard text in botany and biol-
y departments —Amy Trauth Nare, Botanical Research Institute of Texas, 509 Pecan Street, Fort
Worth, TX, 76102 U.S.A, amy_trauth@yahoo.com.
SIDA 20(3): 890, 2003
YUCCA CERNUA (AGAVACEAE: SERIES RUPICOLAE),
A NEW SPECIES FROM NEWTON AND JASPER COUNTIES
IN EASTERN TEXAS
Eric L. Keith
Raven Environmental Services Inc.
PO. Box 6482
Huntsville TX 77342, U.S.A.
eric. keith@excite.com
ABSTRACT
A new species of Yucca from the series Rupicolae i is cesctiped nom Jesper ane Newton counties in
b 1 beaked
eastern Texas. Series Rupicolae is distinguished
fruits. This new species, Yucca cernua, sp
acaulescent species in ser. Rupicolae. The combined seen Me a tall flowering scape; the ten-
| ibed, illustrated d disti ished from other
dency of the branches of the i p engine ; the moderately to
densely f] fl large | picall i lati ort capsules, pistils
and stamens; dee habitat; and disjunct distributi mn distinguish this Spec from other acaules-
idate for
ent es in ser. Rupicolae. The rarity of this speci
state aud federal ape protection. Also ieliided is a key of the entire or denticulate, non-
filiferous leaved North American species of Yucca.
RESUMEN
Se describe una nueva especie de Yucca de la serie Rupicolae de los condados Jasper y Newton en el
Este de Texas. La serie Rupicolae se distingue por las hojas denticuladas y frutos picudos. Esta nueva
especie, Yucca cernua sp. nov. se describe, se ilustra y diferencia de las otras especies de la serie
Hapenae ca coment ae varias caracteristicas edistneue esta especie de las otras especies
a presenela ane un ecape ate tocon flores,
] j H dp] ae | ee
de means a pCO AEN Heroes, ayes pistilos Y ee eeuwamente cortos, las hojas
n habitat tinico, y su distribuci6n disyunta.
L r
eee za de est ie justifi ideracion c lidata para la regulacion de proteccién
estatal y federal. iambic se incluye una clave de todas las especies de Yucca norteamericanas con
hojas denticuladas y filiferas.
In May of 2001, an unusual population of Yucca was discovered along U.S. High-
way 190 in Newton County, Texas. It appeared distinctive because of its tall in-
florescence (up to four meters) and the characteristic tendency of the inflores-
cence branches to recurve and droopas they lengthened. On closer investigation,
these plants apy 1 most similar to Y. pallida McKelvey with mostly flat, glau-
cous, denticulate leaves. No plants were collected because the significance of
the population was not recognized at the time. Plants were later collected in
flower in June of 2002 and in fruit in November of 2002. It was unknown
whether this population was escaped from cultivation, disjunct, or represented
SIDA 20(3): 891-898. 2003
892 BRIT.ORG/SIDA 20(3)
an undescribed species. After examination of live plants and numerous her-
barium specimens of its apparent closest relatives, it was determined to repre-
sent a new species. Acronyms of the herbaria cited follow Index Herbariorum
(Holmgren et al. 1998).
Yucca cernua Keith, sp. nov. (Figs. 1, 2). Type: US.A. TEXAS. NEWTON Co: brown, clayey
roadside on US Hwy 190, 3/8 mi E of Newton-Jasper County line, plants ranging from 1.5-4
m tall, branches of inflorescence drooping with age, 11 Jun 2002, Eric L. Keith 183 (HOLOTYPE:
TEX: ISOTYPES: BRIT, SHSU)
A Yuccae pallidae ac Yuccae rupicolae similis fructibus rostratis et marginibus serrulatis foliorum;
) utroque differt inflorescentia elatiore (usque ad 4 m) ramis moderate vel dense floccosis
| | ; lialaweentat falii st
pet)
c
Jan
us ad 80 cm longis ac 6.5cm
TCCUL
oO a 1
latis), pistillis brevioribus (2-2.7 cm), et staminibus brevioribus (1.2-1.7 cm).
Acaulescent, forming large solitary clumps and spreading from thick rhizomes,
1-2 cm diameter. Leaves (30-)40-70(-80) cm long, (3-)3.5-6.5 cm wide at wid-
est point, greatly narrowed (0.7-1.2 cm) to just above union with base, young
leaves glaucous, the lowermost leaves usually undulate and twisting, older leaves
becoming yellowish-green to olive green and flat or slightly concave, turning
purplish or brown on drying, acute or slightly acuminate, becoming concave
and inrolled 0.7-2.5 cm below tip; leaf margins corneous, yellowish and mi-
nutely denticulate; spines, 0.3-1 cm long, pungent, yellowish at base and red-
dish-brown at tip. Inflorescence long-exserted, (1.5-) 2-4 m tall, scape (1.2-)1.5-
3.2 m in length; inflorescence panicled, moderately to densely floccose,
branched from base to racemose tip; inflorescence branches characteristically
recurved and drooping as they lengthen, longest branches to 35cm long. Flowers
mostly white or slightly greenish white, campanulate, pendant; tepals free, gla-
brous, outer whorl narrowly elliptic or lanceolate, 3.4-5(-5.8) cm long, 0.75-L7
cm wide, inner whorl elliptic 3.5-5(-5.8) cm long and 1-1.8 cm wide; stamens
|.2-1.9(-2.2) cm long; filaments reflexed or even slightly twisted or contorted
especially on drying, echinate; anthers to 0.7 mm long, sagittate; pistil 2-3
(-3.3) cm long as measured from base of ovary to tip of stigmas; ovary slender,
oblong-cylindric, 0.4-0.8 cm diameter; styles 3, 0.8-1.5(-L7) cm long, erect to
slightly spreading, branches usually papillose. Fruit dry, dehiscent, pendant,
3.2-4.5cm long, 2-2.5cm in diameter, symmetrical, oblong-cylindric, at dehis-
cence splitting for entire length on primary fissures and usually !/4 to !/2 of
the length of capsule on secondary fissures; beaks 0.5-1 cm in length, usually
turned inward at 45° angles where split on secondary fissures (Fig. 1). Seeds
polymorphic, obovate to D-shaped 5-7 mm long and 4-6 mm wide, thicker to-
ward edges, 0.5-1 mm thick.
Etymology.—The epithet refers to the characteristic tendency of the inflo-
rescence branches to recurve and droop as they lengthen, an apparently dis-
tinctive feature separating it from all other Yucca species.
Habitat and Distribution.— Yucca cernua occurs in open or slightly shaded
KEITH, A NEW SPECIES OF YUCCA FROM EAST TEXAS
893
pape dace anal
ae cathy fo
agente 7
et Pe,
Sy yo
F<
otros aah
What AA VE
nde te
gas tite
Fic. 1. Yucca cernua. A. capsule B. Habit. C. ae leaf quan elie ait fu margin magnified 11x. D.
Stamen, fi slig htly enlar :
areas apparently restricted to brownish, acidic clayey soils of the Redco Soil
Series in Jasper and Newton counties in eastern Texas (Neitsch 1982). The Redco
Soil Series occurs in southeastern Texas from Newton County westward to
Walker County (McClintock 1979; Neitsch 1982; USDA 2002). This species is
currently known only from an area approximately 6-km* centered in west-
central Newton County and just into the far eastern portion of Jasper County.
One large population of approximately 1000 plants and six small populations
894 BRIT.ORG/SIDA 20(3)
apenas
Fig. 2. Yucca cernua Keith. A. eee eee glaucous. Note lewer seas ie manor and twisting. B. Older leaves
yellowish-green or
KEITH YUCCA FROM EAST TEXAS 895
of less than a 100 plants each are known. In 1987, Thomas and Allen collected
this species (labeled as Yucca sp.) at the type locality. A 1951 collection of Y.
cernua (labeled as Yucca) at the Botanical Research Institute of Texas (BRIT)
from Jasper County indicates that it was collected in the town of Jasper. This
population, if still extant, is at least five miles west of any known location. No
other collections of this species are known. This species is apparently tolerant
of soil disturbance. All of the plants are found along roadsides and other dis-
turbed areas, and it rapidly colonizes areas with bare soil. It is hoped that more
populations will be found throughout southeastern Texas in similar habitats.
Associated species—Eurybia hemisphaerica (Alexander) Nesom, Helianthus
angustifolius L., Panicum anceps Michx., Solidago canadensis L., Tridens strictus
(Nutt.) Nash, Smilax bona-nox L., Pinus taeda L., Symphyotrichum lateriflorum
(L.) A.SD. Love, Symphyotrichum dumosum (L.) Nesom, Carex flaccosperma
Dewey, Liquidambar styraciflua L., Ilex vomitoria Ait., and Coreopsis sp.
Additional collections examined: U.S.A. TEXAS. Jasper Co.: Jasper, | 69 cm long and 4. cm
ee terminal gee DF a mm none 20 Jul 1951, H.R. Reed 1238 (BRIT). Nee toii Co.: on S side of US.
mi E of 1 near Jasper and 4.3 mi W of Newton, Texas, top of dry, sandy
ee S side of U.S. 190, 07 Sep 1987, R. Dale Thomas & Charles M. Allen 101823 (NLU).
Yucca cernua is a very distinctive species adapted to a unique soil type. It ap-
pears to be closely related to Y. pallida and Y. rupicola Scheele in the series
Rupicolae and which it most closely resembles by its acaulescent habit, den-
ticulate leaves, and beaked capsules (Hess & Robbins 2002; McKelvey 1947;
Trelease 1902). The closest known populations of either species are over 300
km to the west and northwest of Y. cernua (Correll & Johnston 1970, Diggs et al.
1999; Hochstaetter 2000; McKelvey 1947).
Table 1 provides a list of the major differences between these three species.
To date, our understanding of the species indicates that it is endemic to this
region and is extremely rare. Adverse impacts to this population described above
could pose serious threats to the continued existence of the species. All known
populations of Yucca cernua are found on private land and public highway
rights-of-way and are not currently protected. Fortunately, a preliminary agree-
ment with a private landowner will set aside a preserve along both sides of
Highway 190 where the largest population occurs. Since this species appears to
have a very restricted distribution based on unique soil requirements, it should
be considered as a candidate for federal and state protected species status.
Included herein is a key of the entire or denticulate, non-filiferous leaved
North American species of Yucca. Key is derived from Hess and Robbins (2002)
and personal observations.
. Mature plants caulescent; rosettes each with more than 100 leaves; distribution in
western Texas.
2. Mature plants 2.5-3.6 m, not including inflorescence; leaf blade 25-60 x 1.2-1.7
cm, widest cor siderably above middle, smooth Yucca rostrata
Tas.e 1.4 comparison of Yucca cernua, Yucca pallida,and Yucca rupicola. Characters are taken from p lob ti
(2002), and McKelvey (1947).
,Diggs et.al (1999), Hess and Robbins
Feature
Yucca cernua
Yucca pallida
Yucca rupicola
Distribution
Habitat
Leaf dimensions
Leaf characteristics
Spine characteristics
Inflorescence height
Inflorescence
characteristics
Tepal dimensions
Pistils
Stamens
Capsules
Capsule beaks
Jasper and Newton counties in eastern
Texas
Acidic clays of the Redco Soil Series
(30-)40-70(-80) cm long, (3-)3.5-6.5 cm
wide at widest point
Most leaves flat except green lowermost
leaves which are usually undulate
twisted, young leaves glaucou
becoming olive or yellowish ae age
Yellow at base, reddish brown at tip
(1.5-)2-4 m tall
Moderately to densely and floccose,
branches recurved and drooping
3.4-5(-5.8) cm long, 0.75-1.8 cm wide
2-3(-3.3) cm long
1.2-1.9(-2.2) cm long
3.2-4.5 cm long
0.5-1 cm long, usually turned inward
at 45° angles where split on secondary
fissures
Central Texas northward to North
Central Texas
Open areas on limestone outcrops
18-35(-56) cm long, (1.0-
cm wide at widest point
) 2.5-3.2(-4.5)
Most leaves flat and persistently
blue-green glaucous
Mostly all yellow, occasionally
slightly reddish near base
1.3-2.5 m tall
Glabrous and glat ICOUS,
branches wide-spreading
(3.6-)5-6.5 cm long, (1.5-)
2-3.2 cm wide
2.4-4 cm long
2.1-3.2 cm long
3.8-5 cm long
0.8-1.2 cm long, usually
erect
Edwards Plateau
On limestone ledges and plains
25-60(-82) cm long, (1.7-) 2.3-5(-6
cm wide at widest point
Most leaves olive-green and
undulate and twisted
Orange, yellowish-brown, or
reddish-brown
1.5-3.6 m tall
Glabrous or slightly pubescent,
branches erect-ascending
(4-)5.5-8.2 cm long, (1.4-)
2-3.2 cm wide
2.7-4 cm long
1.8-2.8 cm long
4.8-7.5 cm long
0.9-1.2 cm long, erect or slightly
reflexed
(€)0z vals/OuO'LINd
KEITH YUCCA FROM EAST TEXAS 897
2. Mature plants 0.7-2.5 m, not including inflorescence, leaf blade 20-30(-45) x
Yuc
.2cm,widest at or above middle, + scabrous cca thompsoniana
1. Mature Lene acaulescent, usually forming colonies; rosettes each with fewer than
100 leaves.
3. Mature leaf blade undulate and twisted, strongly concave, margins orange
or yellowish-brown to reddish-brown; endemic to eastern Edwards Plateau
Yucca rupicola
3. Mature leaf blade ee Vaan leaves a twisted in. ¥. cernua), flat or
slightly concave, margins yellow brow
4. Mature eee me 2 sem ae a nleeees densely floc-
cose with branchlets spreading or ascending; endemic to western Edwards
Plateau Yucca reverchonii
. Mature leaf blades (1-)2.5-6.5 cm at greatest width, inflorescence either mod-
erately to densely floccose with recurved and drooping branchlets or gla-
brous with wide-spreading branchlets.
. Plants in colonies of 10-30 rosettes each, leaves (1—-)2.5-3.2(-4.5) cm at
greatest width, persistently blue-green glaucous, inflorescence glabrous
with wide-spreading branchlets;endemic to central and north-central Texas
aes
[eal
ucca pallida
5. Plants usually solitary, leaves (3—)3.5-6.5 cm at greatest width, young leaves
glaucous becoming olive or yellowish-green with age, inflorescence
floccose with recurved and drooping branchlets; apparently endemic to
eastern Texas Yucca cernua
ACKNOWLEDGMENTS
I would like to thank Karen Clary of Austin, Texas for all her advice and en-
couragement, for sharing her expertise on Yuccas, and for critical review of the
paper. William J. Hess also gave critical review of the paper and offered helpful
suggestions. I would also like to thank Guy Nesom of the Botanical Research
Institute of Texas (BRIT) for all his help and for providing the Latin diagnosis.
University of Texas Herbarium curator Tom Wendt, interim collections manager
Amy Trauth Nare at BRIT, and Justin Williams at Sam Houston State Univer-
sity kindly allowed me to use the collections and were very helpful. In addition,
lexamined Yucca loan material at TX/LL from A, ARIZ, BAY, DS, GA, GH, MO,
NLUUNY TABS Lx. UC and Us:
I give special thanks to my botany professors at Stephen F. Austin State
University, Elray Nixon and James Van Kley, for instilling my interest in plant
taxonomy and ecology and to Larry Brown of Houston Community College and
Spring Branch Science Center for his enthusiastic tutelage on the flora of Texas.
Lalso thank my co-workers, Ross Carrie and Kevin Mundorff, for their as-
sistance including critical review of the paper. Richard Lane of the Huntsville
Public Library and Vinessa Mundorff of Sam Houston State University were
also very helpful in acquiring most of my reference material.
Most of all, 1 would like to thank my family for all their love, support, and
patience during my botanical pursuits including my wife, Elizabeth (who also
edited the paper), and two sons, Lance and Will. My father, William Keith of
BRIT.ORG/SIDA 20(3)
Lufkin, Texas, provided the Spanish translation of the abstract, and my uncle,
Larry Keith of Houston, Texas, provided the illustrations.
REFERENCES
Corrett, D.S. and M.C. JoHNston. 1970. Manual of the vascular plants of Texas. Texas Re-
search Foundation, Renner, TX.
Diccs, G.M., B.L. Liescome, and RJ. O’Kennon. 1999. Shinners & Mahler's illustrated flora of
north central Texas. Sida, Bot. Misc. 16:1-1626.
Hess, WJ. and R.L. Rossins. 2002. Yucca. Flora of North America Editorial Committee, eds.
Flora of North America North of Mexico, Vol. 26, Magnoliophyta: Liliidae: Liliales and
Orchidales. Oxford University Press.
HOcHsTAETTER, F. 2000. Yucca (Vol.|), in the Southwest and Midwest of the USA and Canada.
Published by the author.
Houmeren, P-K., N.H. Houmoren, and L.C. Barnett. 1998. On line updated version of Index
Herbariorum, Part |: The herbaria of the world, 8th ed. [http://www.nybg.org/bsci/ih/
ih.Atm|]
McCuntock,W.R. 1979. Soil survey of Walker County, Texas. Published by the U.S. Depart-
ment of Agriculture, Soil Conservation Service, in cooperation with the Texas Agricul-
tural Experiment Station.
McKetvey, S.D. 1947. Yuccas of the southwestern United States, Part Il. The Arnold Arbore-
tum of Harvard University, Jamaica Plain, MA.
NeitscH, C. 1982. Soil survey of Jasper and Newton counties, Texas. Published by the U.S.
Department of Agriculture, Soil Conservation Service, in cooperation with the Texas
Agricultural Experiment Station.
TrRELEASE, W. 1902. The Yucceae. Rep. (Annual) Missouri Bot. Gard. 13:65-71.
United States Department Agriculture (USDA). 2002. Natural Resources Conservation
Service Website. http://ortho.ftw.nrcs.usda.gov/osd/dat/R/REDCO.html
TWO NEW SPECIES OF PARATHESIS (MYRSINACEAE)
FROM MESOAMERICA AND
NEOTYPIFICATION OF PARATHESIS ROTHSCHUHIANA
Jon M.Ricketson John J. Pipoly Ill
Missouri Botanical Garden Fairchild Tropical Garden
PO. Box 299 aie Old Cutler Road
St. Louis MO 63166-0299, U.S.A. Coral Gabl i) Fl Se ices U. S. A
jon.ricketson@mobot.org jpipoly@fairchildgard
ABSTRACT
Two new species of Parathesis, P. pseudocalophylla Ricketson & Pipoly from Chiapas, Mexico, and P.
pseudocrassiramea Ricketson & Pipoly from San José, Costa Rica, are described and illustrated. Fur-
ther revision of the genus has revealed the necessity of neotypifying P. rothschuhiana Mez.
Qa
RESUMEN
Se describen d i l gé P vesis, P pseudocalophylla
Ricketson et Lone Vale peciinles rassiramea pieleman et Pipoly, provenientes de Chiapas, México y
San José, Costa Rica. Ademas, se neotipifica P rothschuiana Mez.
mags
arat
The genus Parathesis (A. DC.) Hook. f. contains 95 species distributed from
northern Mexico to Panama, the Caribbean, and throughout the Andes from
Venezuela to Bolivia and adjacent Brazil. The genus is defined by the unique
glandular papillae of the corolla lobes, and bright yellow anthers. In preparing
the treatment of the genus Parathesis for Flora Mesoamericana, two new spe-
cies were discovered, and are described herewith.
Parathesis pseudocalophylla Ricketson S Pipoly, Sp. Nov. oe 1). TYPE. MEXICO,
I -aM po
Alegre, 2,300 m, 3 Jul 1981 (EL), D. Breedlove S131] 4 ¢ lOLOTY PE: MO; ISOTYPES: CAS, LL).
uose pana pula in
ah am lateralem P.calophyllae valde affinis, sed ab ea ramulis
(non adpresse ferrugineo-) arachnoideo- dendroideo-tomentosis,
aol 0.7-1.5 (non 26 3.4) cm longis, c: ree L.0-L8 ie 2.2-2.4) mm longo, corolla 6.1-6.3 (non
7.2-75)mm longa, atque staminium 4.5-4.7 (non 4.3-4.5) mm longarum, filamentis 2.9-3.1 (non 2.5-
2.7)mm longis, ac antheris lanceolatis (non ovatis) 2-2.2 (nec 1.8-2.1) mm longis statim separabilis.
Trees to 6.1 m tall. Branchlets slender, angulate, 3-6 mm in diam., densely and
stiffly rufous arachnoid and dendroid tomentose. Leaves: with blades
chartaceous to subcoriaceous, elliptic, 8.8-14.4 cm long, 2.2-4.7 cm wide,
apically acuminate, the acumen 0.6-16 cm long, basally acute, decurrent on
the petiole, conspicuously and prominently punctate and punctate-lineate, gla-
brous above except scattered to densely rufous dendroid tomentose at the base
and along the midrib, with scattered arachnoid and dendroid trichomes below
SIDA 20(3): 899-907. 2003
900 BRIT.ORG/SIDA 20(3)
Fic. 1. Parathesis pseudocalophylia Rick & Pipoly. A. Flowering branch. B, Detail of abaxial leaf surface. C. Detail of
inflorescence. D. Pistil. A-D drawn from holotype, D. Breedlove 51374 (MO).
901
except densely so along the midrib, the midrib impressed above, prominently
raised below, the secondary veins 20-27 pairs, slightly bullate above, promi-
nently raised below, the margins entire; petioles slender, marginate, 0.7-1.5 cm
long, vestiture as in the branchlets. Inflorescences lateral, pinnate to bipinnately
paniculate, 3.5-9.5 cm long, 2-5 cm wide, pyramidal, shorter than the leaves,
the rachis with a vestiture as in the branchlets, the branches 3-7-flowered
corymbs; peduncles 2.2-5.1 cm long; inflorescence bracts unknown: inflores-
cence branch bracts caducous, membranous, ovate to lanceolate, 1.8-2.1 cm long,
0.7-0.9mm wide, apically acute, conspicuously and prominently punctate and
punctate-lineate, glabrous adaxially, densely arachnoid and dendroid tomen-
tose abaxially, the veins inconspicuous, the margins entire; floral bracts simi-
lar to the inflorescence branch bracts but, 0.9-1.6 mm long, 0.3-0.5 mm wide;
pedicels slender, angulate, 4.3-5.5 mm long, conspicuously and prominently
punctate and punctate-lineate, vestiture as in the branchlets. Flowers 5-merous;
calyx chartaceous, 1.6-1.8 mm long, the tube 0.3-0.5 mm long, the lobes ovate,
1.1-1.3mm long, 0.8-1 mm wide, apically acute, conspicuously and prominently
punctate and punctate-lineate, glabrous within, scattered to densely pubescent
with simple trichomes, the margins entire; corolla pink, chartaceous, 6.1-6.3 mm
long, the tube 0.6-0.8 mm long, the lobes lanceolate, 5.4-5.7 mm long, 12-15
mm wide, apically attenuate, conspicuously and prominently punctate and
punctate-lineate, glandular-papillose within except glabrous at the very base,
scattered to densely pubescent with simple trichomes, the margins entire; sta-
mens 4.5-4.7 mm long, the filaments 2.9-3.1 mm long, the staminal tube 0.7-
0.8 mm long, the apically free portion 2.1-2.3 mm long, conspicuously and
prominently punctate and punctate-lineate, glabrous, the anthers yellow, erect,
basifixed, lanceolate, 2-2.2 mm long, 0.7-0.9 mm wide, apically acute, apicu-
late, basally cordate, the connective conspicuously and prominently punctate
and punctate-lineate; pistil 5.9-6.2 mm long, the ovary ovate, 1.1-1.3 mm long,
inconspicuously punctate and punctate-lineate, glabrous except apically of
scattered to densely villous or pilose of simple multicellular trichomes, the style
4.7-4.9 mm long, conspicuously and prominently punctate and punctate-
lineate, glabrous except basally with scattered to densely villous or pilose of
simple multicellular trichomes, the ovules 6-8, uniseriate. Fruits unknown.
Distribution.—Parathesis pseudocalophylla is only known from the type
collection, between Las Margaritas to Campo Alegre, in the Mpio. of La
Independencia, Chiapas, Mexico, growing at 2300 m elevation.
Ecology and conservation status.—Parathesis pseudocalophylla occurs in
evergreen cloud forests, with Pinus, Quercus, Magnolia, Podocarpus, Photinia and
Olmediella. Because it was collected along a logging road, it should be consid-
ered threatened.
Etymology.—The specific epithet comes from it close affinity to Parathesis
calophylla.
—
902 BRIT.ORG/SIDA 20(3)
Parathesis pseudocalophylla wasat first confused with P.calophylla because
of the chartaceous to subcoriaceous, slightly bullate leaves, and lateral inflo-
rescences. However, P.calophylla has densely appressed ferrugineous arachnoid
or dendroid trichomes forming a tomentum, while in P. pseudocalophylla, the
hairs are erect, stiff and rufous. In addition, P pseudocalophylla has shorter peti-
oles and perianth parts, but significantly longer stamens to 4.7 mm long, and
larger lanceolate instead of ovate anthers to 2.2 x 0.9mm.
Parathesis pseudocrassiramea Ricketson & Pipoly, sp. nov. (Fig. 2). Type. COSTA
RICA. SAN JOSE: Canton de Pérez Zeledon, Parque Nacional Chirripo, Cuenca Térraba-Sierpe,
Estacion Santa Elena, 09 23' 36" N, 083° 35' 21" W, 1300-1400 m, 17 Sep 1997 Cr), E. Alfaro 1431
CHOLOTYPE: INB; ISOTYPES: CR, MQ).
Ob laminam bullatam int] i inal i | |
L Lf
P.crassirameae valde affinis, sed ab ea ramulis 8-10 (non 2-5) mm diametris minute papilloso- vel
glandulari granuloso ( Iticellul vel stellato ) tomentosis, amina foliari 18-25 (non 5.8-18.2)
cm longa, 7.9-10.5 (nec 1.8-6.3) cm lata, ad apicem acumine 1-18 (nec 0.3-1) cm longo, calyce 12-17
(non 1.7-L9) mm longe, ant
oy
oO
wn
U1
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oe)
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o
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~
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5
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2.2-2. le recognoscitur.
Trees 6 m tall. Branchlets stout, angulate, 8-10 mm in diam, densely appressed
papillose- or glandular-granulose-tomentose, glabrescent with age. Leaves: with
blades coriaceous, elliptic to oblong, 18-25 cm long, 7.9-10.5 cm wide, apically
obtuse, abruptly acuminate, the acumen 1-1.8 cm long, basally obtuse to
rounded, decurrent on the petiole, bullate, conspicuously and prominently
punctate and punctate-lineate, glabrous above, essentially glabrous below but
with sparse papillae or glandular granules at the very base, the midrib im-
pressed above, prominently raised below, the secondary veins 80-110 pairs, con-
spicuously impressed above, prominently raised below, the margins entire, en-
rolled; petioles slender, marginate, 2.8-3.6 cm long, glabrous above, appressed
papillose- or glandular-granulose-tomentose below, glabrescent with age. In-
florescences terminal and in the axils of the uppermost leaves, pinnate to bipin-
nately paniculate, 27-28 cm long, 22-25 cm wide, pyramidal, as long or longer
than the leaves, the rachis tomentose as in the branchlets, the branches 3-7-
{lowered corymbs; peduncles nearly absent to 1 cm long, the lower branches
subtended by leaves; inflorescence bracts unknown; inflorescence branch bracts
unknown; floral bracts caducous, membranous to chartaceous, ovate, 0.9-1.5
mim long, 0.5-1 mm wide, apically acuminate, basally sessile, conspicuously
and prominently punctate and punctate-lineate, glabrous adaxially, vestiture
as in the branchlets abaxially, the veins inconspicuous, the margins entire;
pedicels stout, angulate, 1-2.5 mm long, conspicuously and prominently punc-
tate and punctate-lineate, vestiture as in the branchlets. Flowers 5- to 6-merous;
calyx chartaceous, 1.2-1.7 mm long, the tube 0.2-0.4 mm long, the lobes ovate,
0.9-1.5 mm long, 1-14 mm wide, apically long acuminate, conspicuously and
prominently punctate and punctate-lineate, glabrous within, vestiture as in the
branchlets, the margins entire; corolla color unknown, chartaceous to coria-
: ae
= AN or
oe ¥ eg Lap of \
Re a Yo IS \
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— C% PO \\\\\\
— Te pe a E\M\\\\\\
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5 }
y t2 [Ss Ki
RIG
SERS
Cae
C. Detail of
flower, showing corolla lobe with glandular papillae and stamen. D. Flower with immature fruit. E. Fruit. A-E drawn
from isotype, £. Alfaro 1437 (MO).
Fic 2. Parathes { ; Rick ® Pingly A. Flower ina branch B.D
903
904 BRIT.ORG/SIDA 20(3)
ceous, 5-5.2 mm long, the tube, 0.8-1 mm long, the lobes linear-lanceolate, 4.1-
4.3 mm long, l-1.2 mm wide, apically long acuminate, conspicuously and
prominently punctate and punctate-lineate, densely tomentose near the base
within, the margins entire; stamens 2.8-3 mm long, the filaments 1.8-2 mm long,
the staminal tube 0.5-0.7 mm long, the apically free portion 1.2-1.4 mm long,
conspicuously and prominently punctate and punctate-lineate, glabrous, the
anthers yellow, 1.5-1.8 mm long, 0.6-0.8 mm wide, apically acuminate, apicu-
late, basally sagittate, the connective, conspicuously and prominently punc-
tate and punctate-lineate; pistil 4.3-4.6 mm long, the ovary ovate, 0.9-1 mm
long, conspicuously and prominently punctate and punctate-lineate, densely
tomentose with appressed simple or stellate trichomes, the style 3.4-3.6 mm
long, conspicuously and prominently punctate and punctate-lineate, glabrous,
the ovules unknown. Fruits depressed globose, 4-5 mm long, 6-9 mm wide, con-
spicuously and prominently punctate and punctate-lineate, densely puberu-
lent of finely appressed simple or stellate trichomes, inconspicuously costate.
Distribution.—Parathesis pseudocrassiramea is only known from the type
collection in the Parque Nacional Chirripo, San José, Costa Rica, growing from
1300-1400 m elevation.
Ecology and conservation status.—Parathesis pseudocrassiramea occurs in
premontane forests. Although it seems protected in the Chirripo National Park,
itisonly known from the type collection and should be considered threatened
at this time.
Etymology.—The specific epithet comes from it ¢
crassiramed.
The type is in fruit with few partial remnant flowers and buds. The re-
maining buds appear to house insects, thus bud measurements are inaccurate.
The fruits are unusual in the genus and appear as a “cauliflower” shape, how-
ever this may be strictly because they were “juicy” at the time of collecting and
the “wrinkled” nature may be caused by the drying process.
Parathesis pseudocrassiramea is closely related to P. crassiramea because
of the bullate leaves, terminal inflorescences, and the ovary densely puberu-
lent with finely appressed simple or stellate trichomes. The leaf blades of P
pseudocrassiramead are larger, to 25 x 10.5 cm, and essentially glabrous below
but with a few papillae or glandular granules at the very base, while in P
crassiramea they are inconspicuously to conspicuously bizonal below with the
marginal zone glabrous and the costal zone sparsely to scattered fine appressed
with whitish stellate trichomes. In fruit, P pseudocrassiramea may also be con-
fused with P cartagoana; however, the vestiture of the lower leaf surface is in-
conspicuously or conspicuously bizonal and the marginal zone is strongly ap-
pressed with dense, canescent stellate or dendroid trichomes and the costal zone
has a mixture of appressed, dense, canescent or ferrugineous to tomentose stel-
—
ose affinity to Parathesis
905
C. Flower. D. Fruit. A-D drawn from
Fic. 3. Pi
J. Atwood & D. Neill 14 (MQ).
906 BRIT.ORG/SIDA 20(3)
late and/or dendroid trichomes with scattered erect reddish dendroid tri-
chomes, which aialaass iately ce the two taxa.
In addition to the af tioned novelties, revision of the genus disclosed
the necessity of ene fying Parathesis rothschuhiana Mez.
—— nore gua Mez, Pflanzenr. lV. 236 (Heft 9):176. 1902. (Fig. 3). Type.
NICARAGUA. MANAGUA: Grenze der Berg und savannenregion, 550 m, 9 Mar 1894 ({1., fr), E.
tar rype: B destr. 1943, F neg.4875); NEOTYPE. NICARAGUA. NUEVA SEGOVIA:
a la orilla del Rio Achuapa al S del Cerro Mogoton, 1,500 m, 12 Jun 1975 (fl), J. Atwood & D.
Neill 14 (NEOTYPE, here selected: MO; ISONEOTYPE: FTG).
Ardisia fusca Oerst. var. glabrata Oerst., Vidensk. Meee’ Dansk Naturhist. Foren Spben have
1861:128. 1862. Type. NICARAGUA. JINOTECA: sylvis humidis monte Pantasmo, [13° 28'N, 85°
54’ W], 3,000-4,000 leg [914-1219 ml], Jan 1848 ( (f |.) A. Oersted 37A (HOLOTYPE: CG ISOTYPE: LL).
The holotype of Parathesis rothschuhiana was destroyed at Berlin (B) in March
1943, thus necessitating the need to located an isotype. Unfortunately, the E.
Rothschuh specimens were not widely distributed, the only other institutions
known to house his material are Leiden (L) and the Smithsonian (US). Searches
of these herbaria, as well as the National Herbarium of Nicaragua (HNMN),
have failed to locate a duplicate specimen. In the absence of any original type
material a neotype must be selected. The J. Atwood & D. Neill 14 material is in
good condition with mature flowers and young fruits. Unfortunately, a search
by Alfredo Guijalva CHNMN) of Nicaraguan herbaria has failed to locate a du-
plicate of this specimen. Thus we hereby designate the J. Atwood & D. Neill 14
sheet at MO as the neotype.
Oersted (1862) was the first to recognize the unique characters of this taxon,
when he described Ardisid fusca Oerst. var. glabrata Oerst. However, Mez (1902)
misunderstood the material and synonymized the variety under Parathesis
fusca (Oerst.) Mez and failed to recognize the similarities between the type of
Ardisia fusca Oerst. var. glabrata Oerst. with his new species P. rothschu hidna
Mez. Lundell (1966) in his revision of the genus was the first to synonymize the
taxon under Parathesis rothschuhiana but was unable to locate any type material.
ACKNOWLEDGMENTS
We wish to thank the Missouri Botanical Garden (MO), Gerrit Davidse (MO)
and the Flora Mesoamericana Project for its support, to bring Ricketson and
Pipoly together to work on the project. We gratefully acknowledge the loans
from the herbaria that made this study possible, along with our collaborators
who supplied us with much critical material. We also gratefully wish to thank
Alfredo Guijalva (HNMN) for his diligent search for specimens in Nicaragua.
Thanks are due to Roy Gereau (MO) for comments on the Latin diagnosis. Gerrit
Davidse and Zachary Rogers (MO) generously gave of their time to provide thor-
ough reviews of the manuscript.
—_
REFERENCES
LUNDELL, C.L. 1966. The genus Parathesis of the Myrsinaceae. Contr. Texas Res. Found., Bot.
Stud. 5:1-206.
Mez, C. 1902. Myrsinaceae. In: A. Engler, ed. Das Pflanzenreich IV. 236(Heft 9):1-437.
Oerstep, A.S. 1862. Myrsineae Centroamericanae et Mexicanae. Vidensk. Meddel. Dansk
Naturhist. Faren Kjabenhavn 1861:117-142, tabs. 1 & 2.
908 BRIT.ORG/SIDA 20(3)
Book REVIEW
WILLIAM . ha TARN, pie ane Genus Epimedium and Other Herbaceous Berberidaceae
lophyllum. (ISBN 0-88192-543-8, hdbk.) Timber Press,
Ine, 133 SW Second Ave, Suite 450, Portland, OR 97204-3527. (Orders:
eaten enn: 503-227-2878, 1-800-327-5680, 503-227-3070 fax).
$49.95, 354 pp, 27 color paintings, 75 color photos, 31 line drawings, 7 b/w
photos, 7 3/8" x 95/8".
Though Epimedium species have been cultivated for centuries in Asia, it is only relatively recently
that the taxonomy has been understood, indeed in great part to the author of this book. William
Stearn, who is probably best known for his other books, Bot amcalt Latinand Dictionary of Plant Names
for Gardeners, provides in his ive monograph of Epimedium
species and cultivated hybr ia Other her eee Berberid. aceae genera are treated in this book as
well, including Achlys, Bongardia, Caulopyhllum, Diphylleia Jeffersonia, Leontice,
Ranzania,and Vancouveria. A treatment of 14 species of Paeyiotlinnd is provided by Julian Shaw.
\ historical introduction begins the book, in addition to a general description of the herba-
ceous genera in Berberidaceae and a key to these genera. The second chapter is filled with facts on
touted medicinal properties, discovery and treatment of new species by 18" century botanists, and
modern hybridization of species for cultivation. Interestingly, the genus was first described by
Linneaus in 1753 when he described Epimedium alpinum, but it remained monotypic until 1821.
Chapter three is | { morphologic characters, including rhizome, stem, foliage
P g ng
flowers and fruits. Stearn debe on the importance of floral morphology with three extensive
figures that include line drawings of the [lowers and spurred petals of each species. Next, Stearn
explains his classification system for Epimedium, which comprises two subgenera and four sections
and his biogeographical hypotheses for present distributions of Epimedium. The taxonomic treat-
ment of Epimec ce in chapter seven is comprehensive, beginning ap pues with a key to spe-
cies and following tions of 54 se and 20 hybrids. Each species description includes
eponymy, synonymy, disenoktie characteristics, illustration citations ne geographic distribution.
In addition, each species has an illustration rte a color photogra dh to paceomne any the text. The
]
_
rest of the book is dedicated to description of the other Descriptions
of these taxa follow the same format as Epimedium.
Commonly called Barren-worts, Epimedium species are increasingly popular in North Ameri-
can landscapes and woodland gardens. Clearly, Stearn himself was interested in the horticulture of
herbaceous Berberidaceae, since this book is littered with comment the history of their cultiva-
tion. The comprehensive information provided here by Stearn and collaborators makes it a worth-
while read for a variety of audiences. The importance of this book to taxonomy notwithstanding,
people interested in species of herbaceous Berberidaceae, the origins of Epimedium and other taxa,
and the cultivation of hybrids should consider adding this book to their library.—Amy Trauth Nare,
Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX, 76102-4060, U.S.A
amy trauth@yahoo.com.
SIDA 20(3): 908. 2003
LECTOTYPIFICATION OF SCHLECHTENDAL MYRSINACEAE
PROM SGEIEDE AND DEPPE COLLECTIONS
Jon M.Ricketson John J. Pipoly Ill
Missouri Botanical Garden Fairchild Tropical Garden
O. Box 299 11935 Old Cutler Road
St. Louis MO 63166-0299, U.S.A. Coral Gables (Miami) FL 33156-4299, U.S.A.
jon.ricketson@mobot.org |pipoly@fairchildgard fusia@/ati |
ABSTRACT
Ardisia escallonoides, Myrsine myricoides and Parathesis melanostricta (Myrsinaceae), are
lectotypified, based on Schiede and Deppe collections housed at HAL
RESUMEN
Cal ifs A AS aay orally ae | AA D +] 5 il
en base a ee de Schiede y Deppe eaardados enel hes bario HAL.
INTRODUCTION
C.J.W. Schiede and F Deppe collected in central Mexico between 1828 and 1836,
and sent those collections to D.EL. von Schlechtendal in Berlin (now B).
Schlechtendal with A. von Chamisso determined the material and published
the results in a series of papers in Linnaea. However, in 1833 Schlechtendal
moved from Berlin to become the Director of the Botanical Garden and profes-
sor of botany at Friedrichs-Universitat (since 1933 Martin-Luther-Universitat)
in Halle, Germany. He transferred his private herbarium to Halle, which con-
tained numerous duplicates of collections from B. After his death in 1866, his
private herbarium was acquired by the university and merged with the univer-
sity herbarium (now known as HAL). Given that the herbarium in Berlin was
destroyed in 1943 during World War II, we cannot be sure whether
Schlechtendal transferred the types for the new species of Myrsinaceae to HAL
when he moved, or brought only duplicates. Therefore, the only choice we have
is to lecotypify his names with the duplicates at HAL that bear notes hand-
written by Schlechtendal. Lecotypifications are as follows:
Ardisia oe Schltdl. @ Cham., Linnaea 6:393. 1831. Tinus escallonioides
(Sch ) Kuntze, Revis. Gen. PI. 2074. 1891. TYPE: MEXICO. VERACRUZ: in sylvis
enn apm without elev. Dec 1828 (f1, fr), C. Schiede & F Deppe 1226 (LECTOTYPE,
accession number 89724-A + B; ISOLECTOTYPES: K, MO (LL neg. 1971-
39), W).
We here by designate the collection at HAL, accession number 89724 A+B as
the lectotype.
SIDA 20(3): 909-911. 2003
910 BRIT.ORG/SIDA 20(3)
Myrsine coriacea (Sw.) R. Br ex Roem. & Schult. Myrsine myricoides Schltdl, Linnaea
5.1834. Rapanea myricoides (Schltdl) Lundell, Wrightia 3:109. 1964. Type, MEXICO. VE
RACRUZ: Jalapae [Jalapal, without elev, May 1829 (fr), C. Schiede 175 (LECTOTYPE, here eae
nated: HAL accession number 89723-Bogen A/AB; ISOLECTOTYPES: HAL 89723-Bogen B/AB,
MO).
Myrsine myricoides Schltdl. has been considered a synonym of M.coriacea (Sw.)
R. Br. ex Roem. & Schult. for some time. In our previous publications Pipoly
(1992a), Ricketson & Pipoly (1997,1999) we erroneously attributed the type to
H. Galeotti 521,522 and 526 based on Mez’s (1902) list of species. Lundell (1964)
also failed to designate a lectotype when he placed this species in the now syn-
onymized genus Rapanea, see: C. Chen and Pipoly (1996), Pipoly (1991, 1992a,
1992b, 1996) and Pipoly and C. Chen (1995). It is now clear that Schlechtendal
based his new species solely on the C. Schiede 175 collection. As noted above,
HAL has two specimens of this collection, we here by designate the specimen
with the hand written label (in Schiede’s hand) that states, “175, Myrica
jalapensis form., Jalapae, Maj. 29.” This specimen also includes a label from
Schlechtendal “Myrsine myricoides n. sp.” and a determination slip from Carl
Mez, 1901 determined as Rapanea ferruginea |(Ruiz & Pav.)| Mez. The HAL
isolectotype does not have a locality label with only the Schlechtendal’s hand
written label stating “175, Myrsine myricoides” without the “n. sp.” addition. This
lends further evidence for selecting the specimen with the hand written local-
ity label.
Parathesis moe ee ) Hemsl., Biol. Centr-Amer, Bot. 2:291. 1881.
Ardisia mel Itdl., Linnaea 8:526. 1834. Type. MEXICO. VERACRUZ: In sylvis inter
San Salvador et Tale apam, Jun 1829 (1), C. Se be ae rOTYPE: HAL; ISOLECTOTYPES: G (F
neg. no. 20085), GH, MO, NY (LL neg. no. 1217), \
We here by designate the HAL al accession number 95759, as the
lectotype.
ACKNOWLEDGMENTS
We are most grateful to Uwe Braun of Martin-Luther-Universitat, Curator of
the HAL herbarium, for his advice and photocopying of the Schlechtendal speci-
mens. We are also grateful to Gerrit Davidse and the Flora Mesoamericana
project (MO) and Fairchild Tropical Garden for support of our joint work. Both
Braun and Davidse generously gave of their time to provide thorough reviews
of the manuscript.
REFERENCES
C.CHen, C.and J. Piroty. 1996. Myrsinaceae. In: Wu Zheng-yi and P. Raven, eds. Flora of China.
Science Press, Beijing and Missouri Botanical Garden, St. Louis, MO. 15:34-38.
Lunpett, C. 1964. Studies of the American Myrsinaceae-ll. Wrightia 3:97-114.
Mez, C. 1902. Myrsinaceae. In: A. Engler, ed. Das Pflanzenreich IV. 236(Heft 9):1-437.
Pipoty, J. 1991. Systematic studies in the genus Myrsine L.(Myrsinaceae) in Guayana.Novon
1:204-210.
Pipovy, J.1992a. Estudios en el género/ (Myrsinaceae) in Colombia. Caldasia 17:3-9.
Pipovy, J.1992b. Notes on the genus ie ine (Myrsinaceae) in Peru. Novon 2:392-407.
Pipoty, J. 1996. Contributions toward a new flora of the Philippines: |. A synopsis of the
genus Myrsine (Myrsinaceae). Sida 17:115-162.
Pipowy, J. and C.C. CHen. 1995. Nomenclatural notes on the Myrsinaceae of China. Novon
599 /=361,
Ricketson, J. and J. Pipoty. 1997. Nomenclatural notes and a synopsis of the genus Myrsine
(Myrsinaceae) in Mesoamericana. Sida 17:579-589.
Ricketson, J. and J. Pipoty. 1999. The genus Myrsine (Myrsinaceae) in Venezuela. Sida 18:
1095-1144.
AA
912 BRIT.ORG/SIDA 20(3)
BOOK NOTICES
David YETMAN. 2002. The Guarijios of the Sierra Madre: hidden People of North-
western Mexico. (ISBN 0-8263-2234-4). University of New Mexico Press,
1720 Lomas Boulevard NE, Albuquerque, NM 87131-1591, U.S.A. (Orders:
1-800-249-7737; 1-505-277-9270 fax.) $49.95, 270 pp., numerous b/w photos,
Tmnap, 6 L722" 1s,
The Guarijios are a native tribe of Mexico's remote Sonora desert, a rugged land of mountains, can-
yons, plains, and the all-important river. It isa land perhaps best known to botanists through Howard
S. Gentry’s Rio Mayo Plants 1942) and Gentry’s Rio Mayo Plants, by PS. Martin, Yetman, et al. (1998.)
Yetman journeyed there sae to study the plants and the ways that the native use the plants. Actu-
ally, by showing an interest in the plants, he became acquainted with the people and their lives so
hat his book is more of an aie ea study than a botanical one. Nevertheless plants figure
predominantly, though it is in no way an attempt to update the earlier works. Rather, Yetman con-
centrates on the usage (food, medicine, lumber, etc.) with an introductory "plant primer” of nine un-
usual plants. He culminates his findings with a list of plants, the “ethnoflora” of the Guarijios.—Joann
Karges, Botanical Research Institute of Texas, Fort Worth, TX, U.S.A.
Davip YeTMANand THomas R. VAN DeveNDER. 2002. Mayo Ethnobotany: Land, History,
and Traditional Knowledge in Northwest Mexico. (ISBN 0-520-22721-2, hbk.).
University of California Press, Berkeley,CA. U.S.A. (Orders: California Princeton
Fulfillment Services, 609-883-1759; Fax 609-883-7413) $48.00 hbk. 359 pp.,
numerous black and white photographs.
In more than fifty trips of four to ten days between the early 1990s and 2000 the authors visited the
Mayos of Sonora and Sinaloa, studying their land, hi heir customs, and scpactally cheit use of plants.
a
most 4 the pan in one
f
way or another in their daily living. After comprehensive execs of the of the geography and
These are people who for the most part live off th
foe of the land and he eth hy of the Mayos, the authors describe a plants that are
lost important to the Mayos, plants ie mi ae the Mayos Mayos.” Following isa list of the plants of
ae areas, ar raneee By amily, and PEOy iding the M ayo and Spanish names as well as the binomial.
TI ch plant and cite its —some vital] y impor-
tant (construction, food, ee and some a (such as kindling or a a. seed-spitting
game).
With this work, Yetman and Van Devender have contributed another in their growing body of
publications that lead to an understanding of the Mayos and ane ie as it relates to plants.—
Joann Karges, Botanical Research Institute of Texas, Fort Worth, T
SIDA 20(3): 912. 2003
A SYNOPSIS AND NOMENCLATOR OF THE
GENUS PARATHESIS (MYRSINACEAE)
Jon M.Ricketson John J. Pipoly Ill
Missouri Botanical Garden Fairchild Tropical Garden
PO. Box 299 11935 Old Cutler Road
St. Louis MO 63166-0299, U.S.A, Coral Gables (Miami) FL 33156-4299, U.S.A.
jon.ricketson@mobot.org jpipoly@fairchildgard lusia@/ati il
ABSTRACT
A new infrageneric classification for the genus Parathesis is proposed. The genus is divided into two
subgenera, Parathesis subg. Laterales Ricketson & Pipoly, and subg. Parathesis. Parathesis subgenus
Laterales is Hanes ay Lage into sections Se dlaphy ide RIGKESOD & r Pipoly, a and Laterales Ricketson
& Pipoly. I ided i ubanae Ricketson & Pipoly,
Parathesis, and Versatiles Ricketson & Pipoly. All 95 species are listed in their respective sections.
RESUMEN
Se propone una nueva clasificacion infragenérica para el género Parathesis. Se divide el género en dos
subgéneros, Parathesis subg. Laterales Ricketson & Pipoly, y subg. Parathesis. Se divide el Parathesis
subgénero Laterales en secciones: Laterales Ricketson & Pipoly, y Calophyllae Rickeston & Pipoly. Se
divide Parathesis subg. Parathesis en secciones, aanas Ricks ison & Pipoly, Parathesis y Versatiles
Ricketson & Pipoly. Se enumeran toda
INTRODUCTION
The genus Parathesis (A.DC.) Hook. f. was first described as a section of Ardisia
Sw. by Alphonse de Candolle (1844) and included five species: Ardisia cubana
A. DC.(P cubana (A. DC.) Molinet & M. Gomez), A.crenulata Vent. (P crenulata
(Vent.) Hook. f.ex Hemsl.), A. serrulata Sw. (P serrulata (Sw.) Mez), A. lateriflora
Sw. (= Stylogyne lateriflora (Sw.) Mez) and A. lepidota Kunth. (= Geissanthus
lepidotus (Kunth) Mez). Parathesis was elevated to the generic level by J.D.
Hooker (1876).
In the first in-depth study of the genus, Carl Mez (1902) listed 20 species of
Parathesis, including seven new species. However, it wasn’t until CL. Lundell
(1966) published his monograph on the genus Parathesis that a usable classi-
fication was developed. Lundell listed 71 species divided into two “Sections” and
23 “Series”. Admittedly Lundell stated (1966: p. 8) that his infrageneric classifi-
cation was “designed to expedite the use of the general key,” and therefore com-
prises a numerical listing rather than a formal classification
Since Lundell’s monograph was published, numerous species have been
described, usually with no refernece to Lundell’s classification system. Cur-
rently, we recognize approximately 150 taxa of Parathesis, 95 at the rank of spe-
cies. Although Lundell’s system is helpful, it is far too large and cumbersome to
—
SIDA 20(3): 913-918. 2003
914 BRIT.ORG/SIDA 20(3)
be useful. In preparation for our treatment of the genus Parathesis for Flora
Mesoamericana, a new infrageneric classification system was needed, which is
formalized herewith. The species listed reflect our current understanding of
the genus. Complete synonymies will appear in the Flora Mesoamericanad and
Flora Neotropica.
TAXONOMIC TREATMENT
Parathesis (A. DC.) Hook. f. in Benth. &@ Hook. f., Gen. pl. 2:045. 1876. Ardisia sect.
Parathesis A. DC., Prodr. 8:120. 1844. Tyre: Parathesis serrulata (Sw.) Mez.
Terrestrial polyaxial shrubs and trees. Young branchlets frequently ferrugine-
ous-tomentose with stellate or dendroid trichomes, often glabrescent. Leaves
monomorphic, alternate, the blades usually pubescent on the lower surface, the
trichomes stellate or dendroid, often appressed and bizonal, often early gla-
brescent, or glabrous, usually paler beneath; costa and primary lateral veins
usually elevated beneath, smooth or impressed above; margins entire, crenu-
late, dentate or serrate; petioles marginate or canaliculate. Inflorescences ter-
minal or lateral, paniculate and pyramidal, the branches umbellate, corym-
bose or subcorymbose-racemose; pedicels accrescent. Flowers bisexual, (4-or)
-5 (or-6)-merous; calyx connate at the base, open in bud, usually glandular
tomentulose and papillose; corolla rotate, valvate, usually narrowly lanceolate,
white to pink, usually tomentulose outside, glandular papillose-tomentose in-
side at least along margins and apically, connate basally to form a tube; sta-
mens included, inserted near the base of corolla tube, the filaments well devel-
oped, slender to stout, usually connate basally to forma tube, the bright yellow
anthers erect or versatile, usually dorsifixed above base, dehiscent by introrse
slits or apical pores, sagittate, lanceolate to ovate, acute, mucronate, apiculate
or obtuse at the apex, dorsally punctate or epunctate; pistil obturbinate, the
ovary ovoid or subglobose, the style long and slender, the stigma punctiform,
ovules few to numerous, usually uniseriate, sometimes biseriate or rarely
pluriseriate on the placenta. Fruits drupaceous, |-seeded, usually depressed-glo-
bose, the endocarp crustaceous, the embryo cylindric, transverse.
A genus of ca. 95 species ranging from northern Mexico to Panama, the
Carribean and throughout the Andes from Venezuela to Bolivia. Members of
the genus occur from coastal mangroves to elfin forest formations at high el-
evations. The genus is diagnosed by the unique glandular papillae of the calyx
and corolla lobes, and the bright yellow anthers.
KEY TO THE SUBGENERA AND SECTIONS OF PARATHESIS
1. Inflorescences lateral, or terminal and lateral Subg. Laterales
2. Tomentum of branchlets and the undersurface of the leaves fine, mostly stellate
and usually closely appressed, sometimes sub-appressed and rather loose,
often glabrescent Sect. Laterales
TARALTIDOI 915
2. Tomentum of branchlets and undersurface of the leaves usually coarse and con-
spicuous and of distinct reddish trichomes, the trichomes dendroid and sessile
or stipitate, or the trichomes stellate with erect and spreading rays, the stellate
trichomes either. sessile, subsessile, or ee
. Inflorescences terminal, p l-pan
Sect. Calophyllae
Subg. Parathesis
Anthers versatile, eae pal hee nati slits, dorsifixed at or about
the middle; filaments usually longer than the Sect. Versatiles
3. Anthers erect, with longitudinal slits and/or seiner pores, usually dorsifixed
near the base, never versatile; filaments shorter than the anthers, often one-half
as lon
4. Tomentum of branchlets and abaxial leaf surface fine, mostly sessile or
subsessile stellate and usually closely appressed, sometimes sub-appressed
and rather loose, often glabrescent Sect. Cubanae
4. Tome ntum of branchlets and undersurface of the leaves usually coarse and
cons id of distinct reddish trichomes, the trichomes dendroid, sessile
or ee or “ihe trichomes sessile, subsessile or stipitate stellate with erect
and spreading rays, usually persistent.
Sect. Parathesis
Parathesis (A. DC.) Hook. io in Benth. & Hook. ft subg, Laterales ee &
Pipoly, subg. nov. Typ Hook. f. ex Mez.
oc subgenus a subgenere Parathese inflorescentiis lateralibus (axillaribus) vel terminalibus et
lateralibus perfacile separabilis.
Inflorescences lateral (axillary), or terminal and lateral (axillary).
Parathesis subg. Laterales as circumscribed here consists of two sections
(sect. Calophyllae and sect. Laterales) and includes 42 species.
Parathesis subg. Laterales Ricketson & Pipoly sect. Calophyllae Ricketson &
Pipoly, sect. nov. Tyre: Parathesis calophylla Lundell.
Sectio haec a sectione | lil
rq persistente statim distat.
nflorescences axillary, or terminal and seals tomentum of branchlets and
eae of the leaves usually coarse and conspicuous and of distinct red-
dish trichomes, the trichomes dendroid and sessile or stipitate, or the trichomes
stellate with erect and spreading rays, the stellate trichomes either sessile
subsessile, or stipitate.
Parathesis subg. Lateralis sect. Calophyllae contains 9 species:
Parathesis calophylla Donn. Sm. P. reflexa Brandegee
Poerstediana Mez P. schultesii Lundell
P. pipolyana Ricketson P.travisae Lundell
P. pleurobotryosa Donn. Sm. P. vestita Lundell
P. pseudocalophylla Ricketson & Pipoly
Parathesis subg. Laterales Ricketson & Pipoly sect. Laterales Ricketson & Pipoly,
ct. nov. TyPE: Parathesis adenanthera (Miq.) Hook. f. ex Me
Sectio haec sectionem Laterali inflorescentiis lateralibus vel lateralibus et terminalibus, tomentio
subtiliter adpresso vel subadpresso atque plerumque glabrescentes statim distat.
916 BRIT.ORG/SIDA 20(3)
Inflorescences lateral, or terminal and lateral; tomentum of branchlets and
undersurface of leaves fine, mostly stellate and usually closely appressed, some-
times sub-appressed and rather loose, often glabrescent.
Parathesis subg. Laterales sect. Laterales contains 33 species
Parathesis adenanthera (Mig.) Hook.f. P. montana Lundell
ex Mez P moritziana Mez
P.angustifolia Lundell P multiflora Lundell
P.aurantidaca Lundell P. psychotrioides Lundell
P.calzadae Lundell P. rekoi Standl.
P.candolleana Mez P. seibertii Lundell
P.cintalapana Lundell P. skutchii Lundell
P.conzattii (S.F Blake) Lundell P.subulata Lundell
P.cuspidata Lundell P.tartarea Lundell
P. glaberrima Lundell P. tenuis Stand.
P. glabra Donn. Sm. P. tomentosa Lundell
P. gracilis Lundell P. venezuelana Mez
P. leptopa Lundell P. villosa Lundell
P. macrantha Lundell P. vulgata Lundell
P macronema Bullock P. wendtii Lundell
P. melanosticta (Schltdl.) Hemsl. P. rekoi StandL.
P minutiflora Lundell P seibertii Lundell
Parathesis (A. DC.) Hook. f. subg. Parathesis.
Inflorescences terminal, oe
Parathesis subg. Parathesis as circumscribed here consists of three sections
(sect. Cubanae, sect. Parathesis and sect. Versatiles) and includes 53 species.
Parathesis subg. Parathesis sect. Cubanae Ricketson & Pipol y, sect. Nov. TyPE:
Parathesis cubana (A. DC.) Molinet & M. Gomez.
=
] ] ~af;
Sectio haec quoad vel subapicaliter dehiscentes necnon
filamenta antheris breviora, sect. Parathesi v aide affnis, sed ab ea foliis ramulisque subtiliter et
adpresse- (nec grosse et erecte) tomentosis perclare distinguitur,
Inflorescence terminal, pyramidal-paniculate; anthers erect, with longitudinal
slits and/or subterminal pores, usually dorsifixed near base, never versatile; fila-
ments shorter than the anthers, usually one-half as long; tomentum of
branchlets and undersurface of leaves fine, mostly stellate and usually closely
appressed, sometimes subappressed and rather loose, often glabrescent.
Parathesis subg. Parathesis sect. Cubanae contains 26 species:
Parathesis agostiniana Lundell P.crenulata (Vent.) Hook. f.ex Hemsl.
P. bracteolata Lundell P.cubana(A. DC.) Molinet @ M. Gomez
P. breedlovei Lundell P. donnellsmithii Mez
P.crassiramea Lundell P.emarginata Mez
TARALACIS 917
P. ferruginea Lundell Ppseudocrassiramea Ricketson & Pipoly
P lanceolata Brandegee P. reticulata Lundell
P. laxa Lundell P. rosea Lunde
P lenticellata Lundell P rothschuhiana Mez
P. mexicana Lundell P.sinuata (Lundell) Ricketson & Pipoly
P microcalyx Donn. Sm. P subcoriacea Lundell
P. obtusa Lundell P.tetramera Bullock
P. parvissima Lundell P. williamsii Lundell
P. perpunctata Lundell P. zuliana Lundell
Parathesis subg. Parathesis sect. Parathesis
Inflorescences terminal, pyramidal-paniculate; anthers erect, with longitudi-
nal slits and/or subterminal pores, usually dorsifixed near base, never versa-
tile; filaments shorter than anthers, usually one-half as long; tomentum of
branchlets and undersurface of leaves usually coarse and conspicuous and of
distinct reddish trichomes, the trichomes dendroid and sessile or stipitate, or
the trichomes stellate with erect and spreading rays, the stellate trichomes ei-
ther sessile, subsessile, or stipitate.
Parathesis subg. Parathesis sect. Parathesis contains 15 species:
Parathesis acostensis J.F Morales P. neei Lundell
P. aeruginosa Stand. P. prionophylla Stand.
P. amazonica Mez P rufa Lundell
P. chiapensis Fernald P. serrulata (Sw.) Mez
P.columnaris Lundell P sessilifolia Donn. Sm.
P eggersiana Mez P. tenorioi Lundell
P fusca (Oerst.) Mez P. villalobosii Lundell
P. kochii Lundell
Parathesis subg. Parathesis sect. Versatiles Ricketson @ Pipoly, sect. nov. Tyre:
Parathesis trichogyne Hemsl.
Sectio haec inter alias antheris versatilis longitudinaliter deshiscentibus et dorsifixis, filamentis
antheris longioribus statim diagnoscenda.
Inflorescences terminal, pyramidal-paniculate; anthers versatile, or tardily ver-
satile, with longitudinal slits, dorsifixed at or about the middle; filaments usu-
ally longer than the anther.
Parathesis subg. Parathesis sect. Versatiles contains 12 species:
Parathesis acuminata Lundell P longipedicellata Ricketson
P.amplifolia Lundell P. palaciosii Pipoly
P.cartagoana Lundell P. panamensis Lundell
P. costaricensis Ricketson P papillosa Lundell
P.croatii Lundell P. pyramidalis Lundell
P glendae Ricketson P.trichogyne Hemsl.
918 BRIT.ORG/SIDA 20(3)
EXCLUDED TAXA
Parathesis acutissima Cuatrec. = Hymenandra acutissima (Cuatrec.) Pipoly &
Ricketson
Parathesis micranthera Donn. Sm. = Gentlea micranthera (Donn. Sm.) Lundell
Parathesis viridis Lundell = Stylogyne viridis (Lundell) Ricketson & Pipoly
—
ACKNOWLEDGMENTS
We thank the Missouri Botanical Garden and the Flora Mesoamericana Project
for its support to bring Ricketson and Pipoly together to work on the project.
We gratefully acknowledge the loans from many herbaria that made the study
possible, along with our collaborators who supplied us with much critical ma-
terial. Gerrit Davidse and Roy Gereau generously gave of their time to provide
thorough reviews of the manuscript.
REFERENCES
CANDOLLE, A.L.P. de. 1844. Myrsinaceae. In: A.P. de Candolle, Prodr. 8:76-143.
Hooker, J.D. 1876. Myrsineae. In:G. Bentham and J.D. Hooker, Gen. pl. 2:639-648.
Lunbett, C.L. 1966. The genus Parathesis of the Myrsinaceae. Contr. Texas. Res. Found., Bot.
Stud. 5:1-206.
Mez, C. 1902. Myrsinaceae. In: A. Engler, ed. Das Pflanzenreich lV. 236(Heft 9):1-437.
A NEW SPECIES OF STYLOGYNE (MYRSINACEAE) FROM
DARIEN, PANAMA
Jon M. Ricketson John J. Pipoly Ill
Missouri Botanical Garden Fairchild Tropical Garden
PO, Box 299 11935 Old Cutler Road
St. Louis MO 63166-0299, U.S.A. Coral Gables (Miami) FL 33156-4299, U.S.A.
jon.ricketson@mobot.org jpipoly@fairchildgard lusia@lati i
ABSTRACT
Preparation of a taxonomic treatment of the Myrsinaceae - oe ele pespamencana proree re-
sulted in the discovery of a new species, S
and illustrated. Its closest relationship is wih S. aiaeciaes A. DC. Siloeyne glomeruliflora is re-
moved from the project area.
RESUMEN
Al preparar un tratamiento taxonémico de la familia Myrsinaceae para Flora Mesoamericana, se
descubrié una especie nueva para ee ciencia, Stylogyne pucuroensis Reson = Pipl: Se GeSerDe,
se ilustra y se discute el Se elimina | ifl
como registro para Mesoamerica como feaultads del estudio.
£
Stylogyne pucuroensis Ricketson & Pipoly, sp. nov. (Fig. 1). Type. PANAMA. Darien:
Parque Nacional del Darién, ridge between N & S branches of Rio Pucuro, in forest N of old
village of Tacarcuna, ca. 18 km E of Pucuro, 077° 16' W, 08" 05' N, 600-800 m, 24 Oct 1987 (fr),
B. Hammel, G. de Nevers, H. Cuadros & H. Herrera 16469 (HOLOTYPE: MO; ISOTYPES: FTG, LL,
PMA).
Quoad inf] i 1 |] is iol | f] 5- partitos S. ardisioideo valde arcte
affinis, sed abea lliptici bl ioli
vel eeenEe (non subdeltatis vel del bate) ir 15 (nec L6- D 0) mm longis atro- (nec pallugida: =)
punctatis I caduco) perfacile recognoscitur.
Shrubs to 3.5 m tall. Branchlets slender, terete, 1-3 mm in diam., glabrous. Leaves:
with blades membranous, elliptic, 8.8-14.6 cm long, 2.8-5.6 cm wide, apically
acuminate, with an acumen 1-1.6cm long, basally acute, decurrent on the peti-
ole, conspicuously and prominently punctate and punctate-lineate, glabrous,
the midrib impressed above, prominently raised below, the secondary veins 37-
47 pairs, inconspicuous above and below, the margins entire, flat; petioles slen-
der, marginate, 0.5-1 cm long, glabrous. Inflorescences lateral, appearing pan-
iculate, 3-5-racemose branches from a common fascicle, 0.5-1 cm long, 1-15
cm wide, shorter than the leaves, the rachis glabrous, the branches 3-7-flow-
ered racemes; peduncles ne obsolete, glabrous; inflorescence bracts un-
known: floral bracts persistent, chartaceous, ovate, 0.5-0.6 mm long, 0.5-0.8 mm
wide, apically acute, Tomine) punctate and punctate-lineate, glabrous, the
veins inconspicuous, the margins entire; pedicels slender, terete, 2.3-3.2 mm long,
SIDA 20(3): 919-922, 2003
920 BRIT.ORG/SIDA 20(3)
Fic. 1. Styl is Rick & Pipoly. A. Flowering branch. B. Fruiting infl dD f hol RB
Pury J ’
Hammel et al. 16469 (MO).
prominently punctate and punctate-lineate, glabrous. Flowers 5-merous, calyx
membranous, the sepals ovate to oblong, 1.3-1.5 mm long, 0.7-0.9 mm wide,
apically acute, prominently punctate and punctate-lineate, glabrous, the mar-
gins entire; corolla, stamens and pistil unknown. Fruits red, globose, 4.5-5.3 mm
in diam., conspicuously and prominently punctate and punctate-lineate, gla-
brous, conspicuously costate, the style base persistent.
Distribution.—Known only from the type collection in the Parque Nacio-
nal del Darién, adjacent to the lower slopes of Cerro Tacarcuna, in Darién,
Panama, growing between 600-800 m elevation.
Ecology and conservation status.—Stylogyne pucuroensis occurs in
premontane pluvial forest. Because it occurs in a remote area of the Darién near
the Panama-Colombia border and within the National park, this species should
not be considered threatened at this time.
Etymology.—The specific epithet comes from its locality, near the Rio
Pucuro.
Because of its lateral inf] equal to the petioles and 5-merous flow-
ers, Stylogyne pucuroensis is most closely related to S. ardisioides. However, S.
pucuroensis has elliptic (not oblong) leaf blades, marginate (not canaliculate)
petioles, ovate to oblong (not subdeltate to deltate) sepals 1.3-1.5 (not 16-2) mm
long with black (not pellucid) punctations. Finally in S. pucuroensis the persis-
tent (not caducous) style on the fruit permit easy recognition.
In our previous paper (Ricketson & Pipoly 1997) this collection was con-
fused with Stylogyne glomeruliflora Cuatrec., a species from the Choco Floris-
tic Province. However, further studies of the genus Stylogyne have shown that S.
glomeruliflora is a member of a small group of taxa with 4-merous flowers,
while S. pucuroensis has a 5-merous flower. Although the small lateral inflo-
rescences are similar, a number of taxa possess this character, including S.
darienensis, which has much longer, racemose branches. Stylogyne
glomeruliflora should not be considered a member of either the Panamanian
or Mesoamerican floras. We therefore provide a revised key to the species of
Stylogyne in Mesoamerica.
KEY TO MESOAMERICAN SPECIES OF STYLOGYNE
1. Leaf blades bullate, pubescent, tne margins crenate Stylogyne hayesii Mez
1. Leaf blades smooth, glabrous, the at entire.
2. Inflorescences lateral, sessile or sub y racemose, 3-/-racemes
Ho a common fascicle.
3. Inflore elongated racemes or corymps, 2-4.5 cm long; pedicels 3-3.5
mm; anthers a longitudinally dehiscent; fruits 5.5-6.5 mm in diam.
Stylogyne darienensis Lundell
3. Inflorescences short racemes, 0.5-1 cm long; pedicels 0.5-2 mm; anthers
dehiscent by pores opening into longitudinal slits; fruits 4-5 mm in diam
Stylogyne pucuroensis Ricketson & Pipoly
BRIT.ORG/SIDA 20(3)
2. Inflorescences terminal and/or lateral, paniculate Stylogyne turbacensis
(Kunth) Mez
4. Leaf blades obovate to oblanceolate, subcoriaceous, the punctations con-
spicuous above; inflorescences pyramidal-paniculate, mostly terminal, or oc-
casionally terminal with few reduced Hee inflorescences eit turbacensis
urbacensis
4. Leaf blades elliptic to oblong, coriaceous, the punctati i icu “a
above; inflorescences columnar-paniculate, mostly lateral or rarely lateral and
terminal or pseudoterminal tylogyne turbacensis subsp.
laevis (Oerst.) Ricketson & Pipoly
ACKNOWLEDGMENTS
We are grateful to Tom Wendt (TEX-LL) for loaning specimens, to the Missouri
Botanical Garden (MO), to Gerrit Davidse (MO) as well as the Flora
Mesoamericana Project, and finally Fairchild Tropical Garden (FTG) for their
support of this inter-institutional research effort. Many thanks are due to Roy
Gereau (MO) for reviewing the Latin diagnosis. Gerrit Davidse and James So-
lomon (MO) generously gave of their time to provide thorough reviews of the
manuscript. The drawing was prepared by Jon Ricketson.
REFERENCES
RickeTson, JM. and J.J. Pirovy. 1997. Nomenclatural notes and a synopsis of Mesoamerican
Stylogyne (Myrsinaceae). Sida 17:591-597
PTEROCERAS MONSOONIAE (ORCHIDACEAE),
A NEW SPECIES FROM INDIA
N. Sasidharan and P. Sujanapal
Kerala Forest Research Institute
Peechi 6806
Kerala, INDIA
ABSTRACT
A new species of the South East Asian Genus Pteroceras Hassk. is described, illustrated and com-
pared to its closest relative Pteroceras johorense.
RESUMEN
bn “4 vale we 2) H kk. del sureste de Asia. se ilustt Ay se compara
con su pariente mas proximo Pteroceras johorense.
INTRODUCTION
Pteroceras Hassk. is reported to have 19 species, mostly inhabited in the warm
humid regions of Southeast Asia, and is best represented in Borneo with 10 spe-
cies (Pedersen 1993). From Peninsular India, only one species, Pteroceras
leopardinum (Parish & Reichb. f.) Seidenf. & Smitin., is reported. During our
recent explorations in the Parambikulam Wildlife Sanctuary of Kerala inter-
esting specimens of Pteroceras collected. The specimens were found to dif-
fer from the South Indian species. pol owas the monograph of the genus
(Pedersen 1993) tl I similarities with the little known Pteroceras
johorense (Holttum) Holttum that seems to be endemic to Johor in Peninsular
Malesia. However, they differ in the leaf form and size; texture, form and size of
he peduncle, size of the bracts and floral parts; structure of labellum and pol-
linia (Table 1). Pedersen (1993) recognized four main types of inflorescences in
the genus. Our species agrees with Pteroceras asperatum (Schltr) PE Hunt, P.
hirsutum (Hook. f.) Holttum, P johorense (Holttum) Holttum, P muriculatum
(Reichb. f.) PE Hunt, P. philippinense (Ames) Garay and P. spathibrachiatum
(J. Smith) Garay by having a muricate or prickly (sub)terete rachis without
any keel or wings. Drawings of our material were sent to Pedersen who sup-
ported our view that it should be described and illustrated as a new species.
Pteroceras monsooniae Sasidharan @ Sujanapal, sp. nov. (Fig. 1). Type: INDIA.
LA. PALAKKAD DISTRICT: Parambikulam Wildlife Sanctuary, 10°32'226" N, 76°86'552" E,
1180m, 28 May 2000, P Sujanapal KFRI 30407 (HOLOTYPE: KFRE; IsOTYPES: BRIT, CALI, MH,
G
Pt ti jol i similis in rhachid l hio oblong-ovato acuto vel rotundato,
sed folii apice latus facto bilobato, pedunculo crasso brevi prominente aculeato, floribus
SIDA 20(3): 923-926. 2003
924 BRIT.ORG/SIDA 20(3)
Taste 1. Comparison of Pteroceras monsooniae with P johorense (Holttum) Holttum.
Character Pteroceras johorense! Pteroceras monsooniae
Leaves ca.5 on a flowering shoot ca.4ona sabe shoot ie or
linear-lanceolate, 5-8 x 1-1.5 cm, elliptic-lanceolate, to 4-8 x 1-2
apex narrowed, obtuse or unequally cm, apex broad, unequally cae
bilobed, lobes very small lobes broad and rounded
Roots 0.6—1 mm diameter 4-6 mm diameter
Inflorescence to 3 per plant 2-7 per plant
Erect, 3.7-4.1 cm long at right angles to the stem, 1-3 cm long
Flowers 2-8, arranged towards the apex 2-4, equidistantly ene in the raceme
F the raceme peduncle 2-4 mm diamete
peduncle 0.4-0.6 mm diameter
Bracts ca.1 x 1 mm, subtriangular, not ca.6 mm long, broadly ovate,
embracing the peduncle base 6 mm embracing the peduncle,
broad hairy on abaxial side prickly on abaxial side
apex not spinulate apex spinulate
Floral parts median sepal, 8.5-11 x 4.5-5 mm median sepal to 18 x 10 mm
petals 8.3-10 x 4.6-5.2 mm petals to 14 x 8mm
spur without ridges spur with two ridges
anther ovate with an acute Anther triangular with a caudate apex
triangular apex, pollinia subspherical, —_ pollinia obovoid, 1.5-1.8 mm long,
1—1.2 mm diameter 1 mm broad
‘Characters based on Pedersen’s (1993) description. The description of Pteroceras johorense is rather
poor due to the rarity of this little known species which is known from two collections only.
aequidistantibus, bractea magna aculeata apice spinuloso, calcare bi-porcato, polliniis obovoideis
rima L-lorma differt.
Epiphytic herb, stem to 2 cm tall, erect, unbranched. Leaves 4, distichously al-
ternate, +-8 x 1-2 cm, sessile, articulate, linear to elliptic-lanceolate, slightly
wider towards the apex, unequally bilobed, lobes rounded; leaf sheaths imbri-
cate, glabrous, covering the internodes. Inflorescences racemous, from the ax-
ils of older leaf sheaths, 2-7, unbranched, at right angles to the stem, 1-3 cm
long; peduncle terete, ca. 4 mm diameter, gray, densely covered with soft brown
prickles; bracts ca. 6 mm long, persistent, broadly ovate, spinulose at apex,
prickly on abaxial side; base 6 mm wide, more or less sheathing the peduncle;
flowers 2-4, sessile, spirally alternating (one open at a time). Sepals and petals
free, spreading, sparsely prickly on abaxial side, grayish-white; median sepal
ca. 1.8 x lcm, laterals adnate to the column foot, elliptic-obovate, apex acute;
petals ca. 1.4 x 0.8 cm, oblong-obovate, base rounded, glabrous or slightly
muricate on abaxial side. Labellum to 6 mm long, side lobes 4 x 3mm, embrac-
ing the column before anthesis and spreading (open) after anthesis, apical part
widened, apex obliquely rounded, midlobe small, with a porrect spur. Spur sac-
cate, ca. 3mm across and 2-toothed at mouth, apex rounded with 2 dorsal ridges,
SASIDHARAN AND SUJANAPAL, A NEW SPECIES OF PTEROCERAS FROM INDIA 925
ig.1.P iae Sasidt & Sujanapal. A. Habit: B. Flower: C. & D. Col ith lip (d
view); E. Flower L.S. (sepals and petal 1); F. Column; G. Bract; H. Sepal; I. Petal; J. & K. Stipe with viscidium
(dorsal and ventral view); L.& M. Pollinia.
926 BRIT.ORG/SIDA 20(3)
lower ridge yellowish, ca. 2 mm; rostellum ca. 3 mm, deeply bifid 3 mm,
clinandrium 3 mm diameter, apiculum 2 mm long. Anther terminal, triangu-
lar with a caudate apex, strongly incumbent, dehiscing ventrally, to form 2 flaps;
pollinia 2, obovoid, 1.5-1.8 x 1mm, deeply cleft, stipe ca. 2 mm long, narrowed
towards the base, bilobed at apex, lobes rounded, viscidium obovate, to | mm
long, broader than the stipe. Ovary ca. 1 cm, terete, twisted to the left, slightly
6-ribbed, sparsely echinate. Capsule ellipsoid, to 2.8 x 1.6 cm, 6-ribbed depressed
at apex (Fig. 1).
Note.—The flower has a two-ridged spur with an yellowish neck (like the
head of a honey bee) which attracts insects during anthesis. The flowers are
ephemeral and withering after 8-10 hours. However, the floral parts continue
to be attached to the ovary during the fruit development.
Flowering and Fruiting —May-June, usually flowering with the onset of
the southwest monsoon.
Etymology.—Specific epithet of the new taxon is based on the flowering
season.
Distribution —Known only from the type locality
Habitat.—This small monopodial epiphyte is growing + L100 m above sea
level on trees like Phyllanthus emblica L.and Syzygium cumini(L. Skeels along
the ecotone area of grassland. Other common trees in the locality are
Cinnamomum sulphuratum Nees, Ficus tinctoria Forst. f. ssp. parasitica (Koen.
ex Willd.) Corner, Xantolis tomentosa (Roxb.) Rafin. var. elengioides (DC.) Vajra.,
etc. The locality constitutes a continuation of the Coimbatore plateau of Deccan
which is subjected to high velocity of wind and humidity. Therefore the trees
are mostly stunted and clothed with dense moss growth.
Paratype: INDIA. Kerala. Palakkad di P bikulam Wild
E, 1180 m, 19 May 2001 Sujanapal KFRI 30596 (KERD.
life Sanc tuary, 10 32'226' N, 76°86'552"
ACKNOWLEDGMENTS
Our sincere thanks to the Kerala Forest Department (Wildlife) for financial as-
sistance, Wildlife Warden and Staff of Parambikulam Wildlife Sanctuary for
the help rendered during the study. We are also thankful to C. Sathish Kumar,
TBGRI, Thiruvananthapuram for providing literature and critical comments,
Henrik Atrenlund Pedersen, University of Copenhagen, Denmark for comment-
ing on our material, J. Veldkamp, Rijksherbarium for the Latin diagnosis, and
V.B. Sajeev for the illustration.
REFERENCE
Pepersen, HA. 1993. The genus Pteroceras (Orchidaceae) — a taxonomic revision. Opera Bot.
117:1-64. Copenhagen.
SYMPLOCOS KOTHAYARENSIS (SYMPLOCACEAE),
A NEW SPECIES FROM PENINSULAR INDIA
V. Sundaresan S.Rajkumar
Genetic Resources ane elorecnnelegy Division Genetic resources and Biotechnology Division
al! itute of Medicinal Central Institute of Medicinal
Sirona Plants (CIMAP) & Aromatic Plants (CIMAP)
Lucknow - 226 015 Lucknow -226 015
Uttar Pradesh, INDIA Uttar Pradesh, INDIA
resanvs@rediffmail.com rajgoa@rediffmail.com
GJ. Jothi V.S.Manickam
Centre for eae and aes Centre for Biodiversity and Biotechnology
of Bot Depar otany
- ae ae St. Xavier's College
Palayam kottai - 627 002 Palayam kottai - 627 002
Tamil Nadu, INDIA Tamil Nadu, INDIA
jothigj@yahoo.com
ABSTRACT
A new species of Symplocaceae, S | isSund Jothi, Rajkumar &@ Manickam
from the Southern Western Chars of I ‘ramil Ni adu, India is described and illustrated.
Key W Sympl Symplocos kothayarensis, Tamil Nadu, India.
RESUMEN
Se describe y se ilustra una especie nueva de Symplocaceae, Symplocos kothayarensis Sundaresan,
Jothi, Rajkumar & Manickam del suroeste de Ghats de Tamil Nadu, India.
INTRODUCTION
The family Symplocaceae comprises four genera (fide Index Nominum
Genericorum; ex: Bobu, Dicalix, Cordyloblaste) and 500 species distributed in
the tropical and subtropical regions, except Africa (Ahmedullah & Nayar 1986).
They occur in the eastern parts of the Old World, reaching from Bombay in the
West to Fiji in the east and from Manchuria in the north to New South Wales
and Lord Howe Island in the south. In the New World the species occur from
the state of Washington to southern Brazil (Nooteboom 1981).
The genus Symplocos Jacq. is represented in India by ca. 45 species. Among
these, 13 species and one variety are endemic to peninsular India (Ahmedullah
& Nayar 1986). In Tamil Nadu alone the genus Symplocos is represented by 10
species and six subspecies (Gopalan 1987).
The southern section of the Western Ghats in India is by far the richest
floristic area with a concentration of endemic taxa (Ahmedullah & Nayar 1986).
The Tirunelveli hills which include the “Agasthiyamalai hills, is a remarkable
SIDA 20(3): 927-930. 2003
928 BRIT.ORG/SIDA 20(3)
micro endemic center and one of the hot spots of the Indian flora (Nayar 1996)
and attested by IUCN as CPD CITE IS7 (Davis et al. 1994).
During the preparation of the family Symplocaceae for the flora of
Tirunelveli hills, one specimen was observed with striking differences belong-
ing to the genus Symplocos Jacq. On critical examination, it was found to be a
new species, and is described and illustrated here.
Symplocos spell ais Sundaresan, Jothi, Rajkumar @ Manickam, sp. nov.
(Fi E: INDIA. TAMIL NADU. TIRUNELVELI HILLS. KANYAKUMARI DT. Kothayar near
eee ayal, 1450 m, 12 Mar 1998, Sundaresan 14968 (HOLOTYPE: NCH; ISOTYPE: MH).
Symplocos pulchrae arte affinis, sed foliis obovato-oblongis, obtusis; racemes curtioribus; disco
glabrato et stylo ad basim glabrato diffe
Small trees to 2(-3)m tall; branchlets fulvous tawny hairy. Leaves simple, alter-
nate, coriaceous, oblong-obovate, 6-9 x 2-4 cm, entire or serrate or half serrate
at margin, the nerves ca. 6 pairs, faintly prominent, glabrous above, sparsely
hairy beneath, the apex obtuse, emarginate, base acute-subobtuse; petioles ca.
2mm long, thick, bristly tawny hairy. Fowers ca. 15 in condensed axillary
racemes, ca. lem long; bracts ovate, ca. 2.3 x L.2mm long, obtuse at apex, densely
villous without, glabrous within, persistent; flowers sessile or shortly pedicel-
late. Calyx tube ca. | mm long, with long appressed hairs; lobes 5, ovate, obtuse
or subacute at apex, appressed hairy without, glabrous within. Corolla white,
lobes 5, ovate-elliptic, apex subacute-obtuse, ca. 3.2 mm long. Stamens ca. 30,
connate at the base, filaments free, ca. 2-2.5mm long; anthers 2-celled, ca. 0.5
mm long, Ovary ca. 1.3 mm long, hairy, 3-locular; ovules 2-3; styles stout, ca.
3mm long; stigma capitate; disc 5-lobed, glandular, glabrous. Fruits not seen.
Distribution.—Symplocos kothayarensis is endemic to the southern part of
Western Ghats, Tamil Nadu, India and is known only from the type locality,
Kothayar hills of Tirunelveli hills near Muthukuzhivayal in Kanyakumari Dis-
trict of Tamil Nadu.
Ecology and phenology.—This species grows at 1450 m elevation in the dense
evergreen forests along with the species like Cullenia exarillata Robyns,
Elaeocarpus serratus L., Mallotus philippensis (Lam.) Muell.-Arg. var.
philippensis, Artocarpus heterophyllus Lam. Symplocos cochinchinensis (Lout.)
Moore subsp. laurina (Retz.) Nooteb. etc... flowering late February to April.
Etymology.—The species is named after its type locality.
Symplocos kothayare nsis is closely allied to S. pulchra Wight but it differs in
the obovate-oblong, obtuse leaves, short racemes, and glabrous disc and style base.
a)
IDENTIFICATION KEY TO SYMPLOCOS KOTHAYARENSIS
|. Diffuse shrubs. Leaves membranous, oblong lanceolate, apex long acuminate, base
rounded or subcordate. Racemes > 2 cm. Disc and style base hairy S. pulchra
. Small trees, Leaves coriaceous, obovate-oblong, apex obtuse, emarginate, base acute
subobtuse. Racemes < 1 cm.Disc and style base glabrous S. kothayarensis
SUNDARESAN ET AL., A NEW SPECIES OF SYMPLOCOS FROM INDIA 929
“!
‘ oh fh i
pe | /
rns
ia = en,
LASS o ae
a ae,
fd tee
fee ee ae
Fic. 1. Sympl k y A Flowering shoot. B. Flowers. C. Bracts. D Calyx E. Coroll plit i F. Stamens. G
pistil with disc. H. Ovary (long. section). 1. Ovary ( ion) d f he type by H. Sankar.
ACKNOWLEDGMENTS
We would like to thank VJ. Nair, Ex Joint Director, Botanical Survey of India for
his valuable comments on our specimen and for the Latin diagnosis. We are
thankful to the Tamil Nadu Forest Department for allowing us to undertake
fieldwork. This work was supported by University Grants Commission (UGC).
The illustration was prepared by H. Sankar, CBB, St. Xavier’s College,
Palayamkottai. We also extend our thanks to the two anonymous referees for
their suggestions in the earlier manuscript.
930 BRIT.ORG/SIDA 20(3)
REFERENCES
AHMEDULLAH, M.and M.P. Nayar, 1986.Endemic plants of the Indian region. Botanical Survey
of India, Calcutta.
Dawis, S.D., V.H. Heywoop, and A.C. Hamitton 1994, Centres of plant diversity. IUCN. The World
Conservation Union. Oxford, U.K-Vol. IL.
Goratan, R. 1987. Symplocaceae. In:A.N.Henry,G.R. Kumari and V.Chithra eds. Flora of Tamil
Nadu, India. Bot. Survey India. Coimbatore. Vol. II. Pp. 68-69.
Nayar, M.P. 1996. Hot spots of endemic plants of India, Nepal and Bhutan. Tropical Botanic
Garden & Research Institute, Thiruvananthapuram.
Nootesoom, H.P. 1981. Symplocaceae. In: M.D.Dassanayake and F.R. Fosberg eds. Flora of
Ceylon. Amerind Publishing Co., New Delhi.
SINOPSIS DEL GENERO MELIOSMA (SABIACEAE) EN
COSTA RICA Y PANAMA, CON TRES NUEVAS ESPECIES
J.Francisco Morales
Instituto Nacional de Biodiversidad (INBio)
Apto 22-3100
Santo Nom! g deH Ji 1, COSTA RICA
RESUMEN
do] 4 NAb]; Cale )
I I g _osta Rica y Panama. Tres nuevas especies,
Meliosma chiriquensis, M. clandestina y M. dept lescritas e i S
£
ABSTRACT
A synopsis of the Costa Rican and Panamanian species of Meliosma (Sabiaceae) is presented here
Meliosma chiriquensis, M. clandestina, and M. depressiva are described and illustrated and their re-
lationships are discussed.
El género Meliosma (Sabiaceae) es un género respresentado por cerca de 15
especies en los trdpicos asiaticos y 40 en el neotrdopico (Gentry 1980). Todas las
especies estan restringidas a zonas de bosque humedo, donde se pueden
reconocer facilmente por sus hojas simples alternas a subopuestas 0 raramente
subverticiladas, con peciolos pulvinulados y por sus frutos, con secrecién
gomosa al cortarse cuando maduros, persistentes por largo tiempo en el arbol,
inclusive hasta 1 ano desputes de madurar.
En vista de la ausencia de trabajos t smi aficos recientes para
la region de Mesoamérica y la presencia de varias especies atin no descritas
encontradas en el proceso de la elaboracion del tratamiento de Sabiaceae para
el Manual de las Plantas de Costa Rica, una sinopsis del género Meliosma en
Costa Rica y Panama se presenta a continuacion, incluyendo distribucion y datos
fenoldgicos; en aquellos casos de especies recientemente registradas, nuevos
reportes o de complejos taxonomicos de especies afines, se incluyen los
especimenes examinados mas representativos. En virtud que los frutos en este
género persisten durante largo tiempo en el arbol despues de producidos y las
colecciones no clarifican la presencia de frutos maduros o ya secos desde hace
largo tiempo, he decidido no incluir datos sobre la fructificacion en las
] : :
GeEsCripcirories.
CLAVE PARA LAS ESPECIES DE MELIOSMA EN COSTA RICA Y PANAMA
1. Hojas largamente pecioladas, la mayoria de los peciolos mas de 4.5 cm largo (rara
vez 4cm en hojas muy jovenes).
2. Hojas cordadas basalmente; sépalos 1.5 mm largo M. cordata
SIDA 20(3): 931-943. 2003
BRIT.ORG/SIDA 20(3)
2. Hojas cuneadas, obtusas a redondeadas basalmente; sépalos 0.5—1 mm largo.
3. Hojas densamente puberulentas en el envés, mas evidente a lo largo de los
nervios; flores rojo-vino a rojo-marron; bosques en Cordilleras 1600-2000 m
M. depressiva
3. Hojas glabras o glabrecentes en el envés; flores anaranjado-crema a
anaranjadas; bosques en llanuras humedas y filas costeras bajo los 600
M. allenii
1. Hojas subsésiles a muy cortamente pecioladas, peciolos usualmente menos de 4
cm largo.
4. Hojas densa a moderadamente puberulentas en el envés, la pubescencia
diminuta, pero regularmente esparcida por toda la lamina.
5. Laminas levemente cordadas basalmente M. subcordata
5. Laminas cuneadas a obtusas basalmente.
6. Peciolos menos de 9mm M. brenesii
6. Peciolos 12-35 mm largo M. occidentalis
4. Hojas glabras a glabrad | : la pubescencia restringida solo a las
nervaduras (M. occidentalis)
7. Hojas largamente lineares M. linearifolia
7. Hojas no lineares, de variadas formas.
8 Flores conspicuamente pediceladas, flores no dispuestas en glomérulos;
frutos menos 1 cm diametro, globosos
M.idiopoda
8. Flores sésiles, subsésiles o inconspicuamente pediceladas, generalmente
dispuestas en glomérulos; frutos mas 1.5 cm diametro, obovoides a
piriformes.
9. Ramos laterales de la inflorescencias inconspicuos, inflorescencias
pareciendo racemosas 0 espiciformes
M. grandiflora
. Ramos laterales de la inflorescencias conspicuamente desarrollados,
inflorescencias claramente paniculadas,
16 La rf fA ]j H | ||
en toda su longitud
M. frondosa
10. Laminas foliares no bulladas o solo muy levemente, los margenes
no revolutos o solo justo antes de la union de la lamina con el
peciolo.
11. Laminas foliares usualmente revolutas hacia la base, justo antes
de la union del peciolo y la lamina
Laminas foliares no revolutas basalmente.
12. Inflorescencias mayormente caulifloras, mas raramente
axilares, glabras, glabradas
puberulentas, no ferrugineas
M. hartshornii
11.
a muy esparcidamente
N
M. glabrata
Inflorescencias mayormente terminales o subterminales,
puberulentas o diminutamente ferrugineo-puberulentas
en la antésis.
13. Hojas ubsésiles peci slo mer
) 9mm largo (hasta
12 mm en hojas muy desarrolladas); lamina obovada
a angostamente obovada, gradualmente atenuadas
hacia el peciolo, con la base obtusa o redondeada;
tallos jovenes algo angulados.
14. Lamina (28-)34-68 3 7.5-18cm;frutos 2.2-2.4cm
largo; pétalos |
argo
—1.5 mmi;inflorescencias 17-65 cm
M. donnellsmithii
MORALES, MELIOSMA EN COSTA RICA Y PANAMA 933
14. Lamina (7—)10-33(-35) 3 (2-)4-8.5 cm;frutos 1.7-
1.8(-2) cm largo; pétalos 2-2.5 mm;inflorescencias
17-26 cm largo M. clandestina
Hojas pecioladas, pecfolo 1-3.5 cm largo; lamina
San con la base cuneada; tallos jovenes teretes a
subte
15. ee once. con las flores laxamente distri-
buidas; 100-800(—1200) m M. occidentalis
15. Inflorescencias con las flores densamente
aglomeradas; 1000-2300 m.
16. Inflorescencias con el eje principal glabro a
glabrado y los ejes axilares esparcidamente
puberulentos, la pubescencia no ferruginea;
1000-1100 m M. chiriquensis
16. Inflorescencias con el eje principal y los ejes
axilares ferrugineo-puberulento a densa-
mente ferrugineo-puberulentas; 1100-2300 m
M. vernicosa
Ww
1. Meliosma allenii Standley & L.O. Williams, Ceiba 3:213. 1953. ele maxima
Standley & L.O. Williams, Ceiba 1242. 1951, non Standley & Steyermark, 1944. Meliosma
oe Standley & L.O. Williams ex Brizicky, Trop. Woods 110:39. me co Costa
ca. Puntarenas: Palmar Nor te, Allen 5648 (HOLOTIPO: F; ISOTIPOS: F, MO, US).
Distribucién.—Costa Rica y Panama, 0-600(-1000) m. Este nombre ha sido
utilizado para algunos especimenes provenientes de Pert y depositados en el
Jardin Botanico de Missouri (MO). Sin embargo, probablemente representen otra
entidad, dado que la morfologia de las hojas y los frutos es diferente. En adicion,
la mayoria de los especimenes son estériles.
Fenologia.—Especimenes con flores han sido recolectados entre Marzo y
Junio. Especimenes con frutos se reportan durante Abril a Diciembre.
Una eae a especies mas comunes en zonas ae donde puede llegar a ser
bastante comun na lo largo de cauces de agua. Solamente existe otra especie con
hojas ees pecioladas en Costa Rica y Panama, Meliosma depressiva
(descrita aqui), pero M. allenii se reconoce por sus hojas con laminas glabras a
glabradas en el envés, flores usualmente anaranjadas a crema-anaranjadas y
distribucion altitudinal diferente.
ray
2. i brenesii Standley, Publ. Field Mus. Nat. Hist., Bot Ser. 18:648. 1937.
TA RICA. ALAJUELA: La Palma de San Ramon, 1050 m, Brenes 5556 (HOLOTIPO: F;
ISOTIPOS: CR, NY).
Distribucién.—Costa Rica y el Norte de Panama, 200 1200(-1700) m.
Fenologia.—especimenes flores se hanr tado entre Junio y Agosto.
Dentro del grupo de especies con hojas oo y peciolos cortos
presentes en Costa Rica y Panama, Meliosma brenesii puede ser confundida con
M. subcordata, con la cual comparte laminas foliares, areas de distribucion
geograficas y pref logicas similares. Sin embargo, M. brenesii se puede
934 BRIT.ORG/SIDA 20(3)
reconocer facilmente por sus hojas con la lamina cuneadasa obtusas en la base,
no cordadas en la base.
3. Meliosma chiriquensis ].. Morales, sp. Nov. (Fig. 1). Tipo. PANAMA. CHIRIQUI: vicin-
ity of Fortuna Dam, 1] Feb 1987 (fl), McPherson 10480 (HOLOTIPO: INB; IsoTIPO: MO).
A Meliosma vernicosa cui affinis, foliis subopuesta vel verticillatis, inflorescenti | lis differt
Arbor. Folia simplicia, 7.7-11.6 ¢ eae 6.2 cm, enpUCS eae subopuesta vel verticillata.
Inflorescentia ramiflora, nee paniculata, terminales ilis, sepala
ciliata, glabrata. Fructus ignotus.
Arbol 18 m; ramas glabras o glabradas. Hojas subopuestas a subverticiladas;
peciolo 2-2.7 cm largo, glabro a glabrado, lamina 7.7-11.6 x 4-6.2 cm, elipticas,
redondeadas a raramente emarginadas apicalmente, cuneadas en la base,
lak
subcoriaceas, enteras, g oglabradas. Inflorescenicas ramifloras, terminales
asubterminales, mas raramente axilares, piramidal paniculada, esparcidamente
puberulentas, especialmente los ejes oo 7-l1cm largo; flores sésiles a
| obtusos apic almente, ciliados
marginalmente, pero el resto glabros a glabrados: | pétalos blancos, 1.8-2 mm
largo, anchamente ovados, glabros; ovario menos de 1 mm largo. Frutos
desconocidos.
Distribuci6n.—endémica por el momento al norte de Panama, pero podria
estar presente en el sur de Costa Rica, LOOO-1100 m. Conocida Unicamente por
la coleccion tipo.
Fenologia.—Floracion en Enero y Febrero.
Esta especie se encuentra algo relacionada con Meliosma vernicosa. Ambas
especies comparten inflorescencias con estructuras morfol6gicas y preferencias
ecoldgicas simi ilares, pero M. chiriquensis se reconoce por sus hojas mayormente
subopuestas o subverticiladas e inflorescencias mucho mas cortas,
esparcidamente puberulentas y con la pubescencia no ferruginea.
subsésiles, sépalos 5, ovados, anc
4. Meliosma clandestina J.) Morales, sp. nov. (Fig. 2). Tipo. COSTA RICA. SAN Jose:
Canton de Dota, Fila Dota, camino entre San Joaquin y Naranjito, 600 m, Mar 2001 (fr),
Morales 762 3; ISOTIPOS: COL, CR, FG, K, MEXU, MO, NY).
Za
A Meliosma donnellsmithii cui affinis, folia minor, fructus major differt. Arbor. Folia simplicia, (7-)10-
33(-35) em X So a 8.5.cm pebovals glabrata. Inflorescentia ramiflora, paniculata, terminales vel
subterminales. { jala ciliata gl abrata. Fructus 1.7-1.8(-2) cm X 1.4-1.7 em, obovoideus.
Arbol 5-11 m; ramas Ages o muy inconspicua y esparcidamente
puberulentas. Hojas alternas a subopuestas en el final de las ramitas; peciolo
0.2-0.5 cm largo, glabro a glabrado; lamina (7-)10-33(-35) x -)4-8.5 cm,
obovadas a angostamente obovadas, agudas a acuminadas (raramente obtusas)
apicalmente, largamente atenuadas hacia la base, pero redondeadas u obtusas
en la base, cartaceas a subcoriaceas, enteras o subenteras y con muy pocos y
casi inconspicuos dientes, glabras o glabradas y con solo el nervio central
puberulento. Inflorescencias ramifloras, terminalesa subterminales, paniculadas,
MORALES, MELIOSMA EN COSTA RICA Y PANAMA 935
Fic. 1. Meliosma chiriquensis J.F. Morales (McPherson 10480, INB). A. Habito.
puberulentas, no ferrugineas, (8-)17-26 cm largo; flores sésiles a subsésiles y
con pedicelos de hasta 1 mm; sépalos 5, ovados, anchamente agudos a obtusos
apicalmente, ciliados marginalmente, pero el resto glabro; pétalos amarillos a
crema, 2-2.5 mm largo, anchamente ovados, glabros; ovario ca. 1 mm largo.
Frutos 1.7-1.8(-2) x 1.4-1.7 cm, obovoide.
3cm
A.
*A
Fic. 2. Melfi landestina J.F. Morales (Morales 7626, INB). A. Habito, infl ia y fruto.
BRIT.ORG/SIDA 20(3)
Distribuci6n.—Endémica a Costa Rica, en las faldas costenas del Pacifico
Central y Sur y la Peninsula de Osa, 100-900 m.
Fenologia.—Especimenes con flores y frutos han sido recolectados entre
Julio, Setiembre y Febrero.
Algunos especimenes de esta especie fueron tratados bajo el concepto de
MORALES, MELIOSMA EN COSTA RICA Y PANAMA 937
Meliosma donnellsmithii en el tratamiento de Sabiaceae para la Flora de Panama,
pero tal y como Gentry lo mencion6 (1980), existian caracteres que sugerian
que el material de Costa Rica de la Peninsula de Osa en realidad correspondiaa
otra especie diferente. Colecciones adicionales acumuladas desde esa época
muestran que en efecto se trataba de una ees sin a Los caracteres
mas importantes para separar M. clandestina de ithii, incluyen
frutos y hojas conspicuamente mas pequenos y f = mas grandes.
Paratiros. COSTA RICA. Puntarenas: forested slopes above Golfito, 27-28 Ene 1967, Burger & Mata 4696
7MO):P. 1 Corcovado, Llorona, 13 Jul 1977, Liesner 3227 (CR, MO). San José: Canton de
Puriscal, Cuenc adel Tulin, Fila Cangreja, 15 Set 1998, Acosta 22(INB),R Forestal La Cangreja, Mastatal
de Puriscal, 22 Jul 1988, Hammel et al. 17116 (NB, MO): Pérez Zeled6n, cuenca ae Bart, Fila Tinamastes, 9
Set 1996, Hammel et al. 20431 (INB, MO); Santa Rosa de Puriscal, 4 Feb 1973, Poveda 480 (CR).
5. ne cordata A. Gentry, Ann. Missouri Bot. Gard. 67:954. 1980 [1981]. Tipo.
NAMA. VERAGUAS: lower slopes of Cerro Tute, ca. 10. km NW of Santa Fe, 750-1000 m, 19
a 1975 (fr), Mori 6297 (HOLOTIPO: MO [2 partes).
Distribucién.—Sur de Costa Rica y norte de Panama, 750-1850 m.
Fenologia.—Especimenes con flores han sido recolectados entre Marzo y
Abril. Frutos se reportan de Marzo a ae
Esta especie era anterior te conocida tnicamente por la colecci6n tipo,
pero ahora es reportada de Costa Rica. Se encuentra bastante relacionada con
Meliosma depressiva (descrita aqui), pero se reconoce por sus hojas con la lamina
basalmente cordada y frutos mas pequenos; en adicion, sus sépalos y pétalos
son mas grandes.
Especimenes representativos examinados. COSTA RICA. Puntarenas: foothills of the Cordillera de
Talamanca, around Tres Colinas, 1800-1850 m, 20 Mar 1984, Davidse et al. 25672 (NB, MO): Canton
de Buenos Aires, Puesto Tres Colinas, Rio Guineo, 17 Set 1996, Gamboa & Picado 649 (INB, MO).
6. Meliosma depressiva J. Morales, sp. nov. (Fig. 3). Tipo. COSTA RICA. SAN Jose: Canton
de Aserri, Fila Bustamante, Hacienda Tiquires, Fila Aguabuena, bosque a camino al Alto La
Arana, 1900-2000 m, 11 May 1996 (fD, Morales & Ulate 5379 (HOLOTIPO: INB; ISOTIPOs: CR, F MO).
At
Inflorescentia ramiflora vel cauliflora, paniculata, floribus pedicellatis, sepala tomentosa. Fructus
2.2-2.7 cm X 16-18 cm, obovoider
pas
rbor parva. Folia simplicia, 9-30 cm x 5-12 cm, eliptica, obovata-eliptica vel lanceolata, puberula.
Arboles 0 arbustos 3-7 m; ramas densamente ferrugineo-tomentulosos 0
densamente puberulentos cuando jovenes, glabrados y muy esparcida e
inconspicuamente puberulentos con la edad. Hojas alternas a subopuestas en
el final de las ramitas; peciolo 4.5-12 cm largo, raramente menos de 4 cm en
hojas inmaduras y jovenes, moderada a esparcidamente puberulento, muy rara
vez glabrado; lamina 9-30 x a 7 cm en ilcas. obovado-elipticas a anchamente
lanceoladas, agudas a obtusas en la base, subcoriaceas,
enteras 0 con muy pocose inconspicuos dientes, esparcidamente puberulentas
en el envés, la lamina glabrada cuando vieja, y con la pubescencia restringida
al nervio central puberulento. Inflorescencias ramifloras a caulifloras, axilares,
938 BRIT.ORG/SIDA 20(3)
paniculadas, densamente puberulentas a tomentuloso-puberulentas, 20-75 cm
largo; flores subsésiles y con pedicelos 0.5-1.5 mm; sépalos 5, ovados, anchamente
agudos apicalmente, densamente puberulentos a tomentulos-puberulentos;
pétalos rojo.vino a rojo-marron, ca. 2mm largo, anchamente ovados, glabros;
ovario ca. | mm largo. Frutos 2.2-2.7 « 1.6-L8 cm, subesférico.
MORALES, MELIOSMA EN COSTA RICA Y PANAMA 939
Distribuci6n.—Endémica a Costa Rica, donde se encuentra en la Cordillera
Volcanica Central, Cordillera de Talamanca, Zona Protectora Cerros de Turrubares
y en la Fila Aguabuena, en las vertiente pacifica, 1300-1900 m. Probablemente
también en Panama, en las provincias de Chiriqui y Bocas del Toro.
Fenologia.—flores y frutos observadas de Enero a Setiembre, pero
mayormente entre Abril y Agosto.
Meliosma depressiva puede distinguirse por sus hojas largamente
pecioladas, inf muy largas y generalmente pendulosas,
las cuales pueden alcanzar mas de 75cm de largo, con las flores rojo vino a rojo
oscuro y distribucion altitudinal sobre los 1300 m. Varias de las colecciones de
M. depressiva se encuentran erroneamente identificadas como M. vernicosa, ya
que ambas especies comparten caracteristicas morfoldgicas similares, pero esta
altima se distingue por sus hojas con peciolos mas cortos, y con la lamina foliar
glabra a glabrada. En Costa Rica y Panama, M. depressiva es también similar a
M. allenii (vease discusi6n previa).
En Mesoamérica, M. depressiva puede ser confundida con M. grandifolia,
distribuida de México a Nicaragua (Matagalpa), pero esta Ultima especie tiene
la pubescencia de sus hojas conspicuamente hirsuta y erecta, no adpresa como
en M. depressiva. En adicion esta nueva especie se encuentra algo relacionada
con M. pittieriana, distribuida en Colombia, Venezuela, Ecuador y Peru, pero
sus laminas foliares son mas angostas y sus flores son rosadas.
Paratipos. COSTA RICA. Cartago: near Rio Naranjo, E of Cachi, 11 Jul 1971, Lent 2002 (F, MO). Heredia:
between Trinidad and Coliblanco, 25 Jull975, Utley & Utley 2718 (CR, EK MO [2 laminas]). Puntarenas:
Buenos Aires, Parque Internacional La Amistad, Altamira, 6 Mar 1999, Alfaro & Alfaro 2174 (CR, INB,
O); Buenos Aires, estacion Altamira, 6 Dic 1999, Alfaro 2604 (CR, INB, MO); Parque Internacional a
Amistad, Cordillera de Talamanca, Pittier, 8 Jun 1995, Angulo 329 (INB); foothills of the Cordillera de
Talamanca, around Tres Colinas, 20 Mar 1984, Davidse et al. 25672 (INB, MO), foothills of the Cordillera
de Talamanca. between Agua Caliente and Rio Canasta. 8 Set 1984. Davidse et al. 28306 (CR. MO): around
Rio Canasta, between Cerro Frantzius and Cerro Pittier, 6 Set 1984, Davidse et al. 28409 (MO): Parque La
Amistad, Pittier, sendero a Rio Gemelo, 30 Ene 1995, Fletes 29 INB, MO). La Amistad, sendero a Cerro
Pittier, 15 Jun 1995, Gamboa 252 (INB, MO): Zona Protectora Las Tablas, near Las Alturaas biological
station, Ene 1996, Holl 1131 (CR); Parque Internacional La Amistad, estacion Pittier, 1 Ago 1995, Moraga
266 (INB); Parque Internacional La Amistad, estacion Altamira, Rio Colorado, 9 Abr 1996, Villalobos
301 os INB, MO). San José: Pérez Zeled6n, Santa Elena, 13 Set 1997, Alfaro & Segura 1382 (INB, MO),
Pér ledon, Chumasquera, 16 Feb 2000, Alfaro & Castro 2836 (INB, MO), Zona Protectora Cerros de
Seas faldas Cerro Turrtibares, 31 Mar 1990, Zuniga et al. 166 (CR, MO).
=
7. Meliosma donnellsmithii Urban, Bot. Gaz. 37:214. 1904. Tipo. COSTA RICA. CARTAGO:
Rio Turrialba, 500 m, Mar 1896 (f1), Donnell Smith 6852 (HOLOTIPO: US; ISOTIPO: NY).
Distribucién.—Nicaragua y Costa Rica, 0-750 m.
Fenologia.—Floracion de Abril a Julio.
Esta especie se reconoce facilmente por el tamano y forma de sus hojas,
usualmente subsésiles o muy cortamente pecioladas (menos 12 mm).
Anteriormente este nombre fue utilizado para dos especies diferentes, una de
ellas previamente no descrita (vease discusion en M. clandestina).
940 BRIT.ORG/SIDA 20(3)
8. Meliosma frondosa Cuatrecasas & Idrobo, Caldasia 7:193. 1955. Tivo. COLOM-
BIA. Cundinamarca: Zipacon, cabeceras del Rio Apulo, entre la Hacienda Sebastopol y Dos
Caminos, 1900-2100 m, 24 Dic 1953 C1), Idrobo & Hernandez 1532 (HOLOTIPO: COL; ISOTIPOS:
FE NY, US).
—
Distribucion.—Panama, Colombia, Ecuador, Venezuela, Pert y Bolivia, 1600-3100 m.
Fenologia.—Floracion entre Diciembre y Enero.
Esta especie se reconoce facilmente por sus hojas conspicuamente bulladas
y algunas veces revolutas marginalmente. La wnica especie con hojas
conspicuamente bulladas y revolutas marginalmente en Mesoamérica es M.
linearifolia, pero la forma de las laminas es totalmente diferente.
9. suse L glabrata (Liebmann) Urban, Ber. Deutsch. Bot. Ges. 13:212. 1895.
Liebm., Vidensk. Meddel. Dansk Naturhist. Foren. Kjobenhavn 1850: 71
me Tipo. C osta Rica. Cartago: Turrialba, 1845 ({1), Oersted s.n. (HOLOTIPO: C; photograph Field
negative 13373, INB).
Lorenzanea ira Liebm., Vidensk. Meddel. Dansk Naturhist. Foren. Kjobenhavn 1850:71. 1850.
Meliosma ira (Liebm,) L.O. Williams, Fieldiana, Bot. 31:261-262. 1967. Tipo. COSTA RICA.
ALAJUELA: Naranjo, May 1845 ({), Oersted s.n. (HOLOTIPO: C, no examinado).
Meliosma tonduzii Donnell Smith, Bot. Gaz. 55:432. 1913. Tipo. COSTA RICA. CARTAGO: Tucurrique,
635-700 m, Tonduz 13368 (HOLOTIPO: US; IsoTIPO: US).
Meliosma panamensis Standley, Trop. Woods. 10:49. 1927. Tipo. PANAMA. Bocas DEL TORO:
Almirante region, Changuinola valley, 1927 (fr), Copper & Slater 29 (HOLOTIPO: US; ISOTIPOS: F,
MO, NY).
Distribucién.—México, Nicaragua, Costa Rica a Colombia, Ecuador y Pert, 500-
1300 m.
Fenologia.—floracion Febrero a Junio.
Esta especie se reconoce facilmente por tener inflorescencias mayormente
caulifloras. Aunque existen especimenes que pueden tener inflorescencias
axilares, sus inflorescencias glabras o casi glabras las separan al instante de
especies similares. Adicionalmente, puede confundirse con M. occidentalis, pero
esta especie se reconoce por las laminas foliares usualmente pubescentes por el
envés y las inflorescencias con flores muy laxamente distribuidas (vrs
densamente aglomeradas en M. glabrata).
10. Meliosma grandiflora Morton ex A.H. Gentry, Ann. Missouri Bot. Gard. 67:957,
fig. 1 C-D. 1980 [1981]. Tipo. COSTA RICA: vicinity of El General, 730 m, Skutch 4049
(HOLOTIPO: MO; IsOTIPOS: CR, NY)
Distribucidn.—Costa Rica y Panama, 0-1800 m.
Fenologia.—flores observadas de Enero a Junio.
En Mesoamérica es la especie que tiene los pétalos mas grandes. Se puede
confundir con Meliosma glabrata, pero reconoce facilmente por sus
inflorescencias racemosas 0 espiciformes,con los ramos laterales conspicuos y
por sus pétalos mucho mas grandes.
MORALES, MELIOSMA EN COSTA RICA Y PANAMA 941
11. Meliosma hartshornii A.H. Gentry, Ann. Missouri Bot. Gard. 73:820. 1986
[1987]. Tipo. COSTA RICA. HEREDIA: road to Volcan Barba, 2 km N of Sacramento, 2750 m, 29
Dic 1974 (f1), Hartshorn 1608 (HOLOTIPO: MO; ISOTIPO: CR).
Distribuci6n.—Costa Rica y Panama, 2300-3000 m.
Fenologia.—floracién de Octubre a Marzo y Junio.
Restringida a las regiones mas altas de la Cordilleras, esta especie se
reconoce por sus hojas glabras a glabradas, con la lamina conspicuamente
revoluta basalmente e inflorescencias claramente paniculadas, con las flores
sésiles a subsésiles.
12. Meliosma idiopoda Blake, J. Wash. Acad. Sci. 14:289. 1924. Tipo. COSTA RICA.
CARTAGO: Tucurrique, 635-700 m, Tonduz 13372 (HOLOTIPO: US).
Meliosma a Standley & Steyermark, Publ. Field Mus. Nat. Hist., Bot. Ser. 23:60-61. 1944. TIPO.
GUATEMALA. acumen eastern slopes of Volcan de Santa Clara, above Chiracao, May
1942 i , Steyermark 467
Distribucién.—México a Panama, (500-) 100-2000 m.
Fenologia.—Especimenes con flores han sido recolectados de Noviembre a
Abril.
Meliosma idiopoda es facilmente reconocible por sus hojas cortamente
pecioladas, esencialmente glabras, inflorescencias con flores conspicuamente
pediceladas y frutos muy pequefios, menores a 1 cm de diametro. Llega a ser
localmente abundante en las regiones donde crece y es sumamente atractiva
durante la floracién, por las numerosas inflorescencias que produce y la
coloracion blanca de sus flores.
13. Meliosma linearifolia A. Gentry, Ann. Missouri Bot. Gard. 67:960, fig. 1 A-B.
1980 [1981]. Tireo. PANAMA. PANAMA: Localidad exacta preservada, 12 Mar 1969 (f1), Correa
& Dressler 1148 (HOLOTIPO: MO; ISOTIPO: NY).
Distribuci6n.—Endémica a Panama.
Fenologia.—El unico especimen conocido hasta el momento, fue recolectado
con flores en Marzo.
Una de las especies mas distintivas especies en el género que se reconoce al
instante por sus hojas lineares y muy largas. Desptes de la coleccién tipo, esta
especie ha sido raramente colectada y ha sido incluida en la lista roja de especies
amenazadas de la Union Internacional para la Conservacion de la Naturaleza
(Hilton-Taylor, 2000). Por lo tanto, su localidad especifica se omite.
14. agen occidentalis Cuatrecasas, Lloydia LI: 217. 1948. Tipo. COLOMBIA. VALLE:
nchicaya, 230-260 m, sin fecha (f{1), Cuatrecasas 1. F; os: COL, US).
Distribucién.—México (2), Costa Rica, Panama, Colombia y Ea 100-800
(-1200) m.
Fenologia.—floracion en Setiembre y Octubre.
Este nombre ha sido utilizado en forma comun y erronea, para identificar
BRIT.ORG/SIDA 20(3)
especime nes pervtenec ientesa Meliosma vernicosa yen algunos CasoOs M. glabrata.
Prefiero no utilizar los caracteres de venacion pubescente empleados por Gen-
try (1980) y Duran (1997), ya que aunque es notable que esta especie tiene
mayormente nervios y peciolos diminutamente puberulentos, también he
encontrado al menos 2 especimenes con hojas totalmente glabras. El caracter
mas distintivo para separarla de las especies relacionadas son las flores
laxamente dispuestas de sus inflorescencias.
También puede ser confundida con M. depressiva, con quién comparte
formas de hoja y pubescencia similares, sobretodo en especimenes con brotes
muy jovenes de ramas, pero se reconoce por sus peciolos mas cortos, flores mas
grandes y distribucion geografica usualmente diferente.
Aunque he revisado bastante material de Costa Rica y Panama, solamente
he podido ubicar tres especimenes con flores que encajan con la descripcion
original; sin embargo, uno de ellos que difiere por sus hojas mas pequenas y
totalmente glabras, este especimen de la Peninsula de Osa es ubicado
tentativamente bajo este nombre e incluso podria representar otra especie
diferente de Suramérica. Mucho otro material presente en herbarios e
identificado con este nombre esta estéril y no se puede tener certeza sobre su
identificacion.
Especimenes representativos examinados. COSTA RICA. Heredia: Cerros Sardinal, N of Chilamate,
10 Jul 1982 (f(D), Hammel & Trainer 13119 (DUKE, INB). Limon: Braulio Carrillo National Park, path
beyond Rio Sucio, May 1984, Gomez et al. 22799 (CR, INB, MO). Puntarenas: Reserva Forestal Golfo
Dulce, Playa Campanario, Sierpe, 12 Oct 1991, Harmon 273 (NB).
15. Meliosma schlimii (Turcz.) Triana, Ann. Sci. Nat., Bot., ser. 5, 16:363. 1872.
Oligostemom schlimii Turcz., Bull. Soc. Imp. Naturalistes Moscou 3k: 447, 1858. TIPO. Colom-
bia. Santander del Norte: localidad perdida, Schlim 702 (HOLOTIPO: G-DC, fotografia negativo
Field 23600, INB).
Distribucié6n.—Costa Rica a Colombia y Ecuador, 0-750 m.
Fenologia.—Especimenes con flores han sido recolectados en Junio y Julio.
Se reconoce por la dimension de sus hojas, densamente ferrugineas a marron
puberulentas en el envés, subsésiles y atenuadas basalmente. En Mesoamerica,
no esta relacionada cercanamente con ninguna otra especie.
16. Meliosma subcordata Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 18:649.
1937. Tipo. COSTA RICA. ALAJUELA: La Palma de San Ramon, 26 Oct 1922 (fl), Brenes 378
CHOLOTIPO: F; ISOTIPO: NY).
Distribucién.—Endémica a Costa Rica, donde se encuentra restringida a la Cor-
dillera de Tilaran, 1400-1600 m.
Fenologia.—Floracion Junio a Agosto.
No es una especie muy comun en el campo, pero se reconoce al instante
por sus hojas cortamente pecioladas, con la lamina densamente puberulenta
enelenvés y levemente cordadas basalmente. Aunque se podria considerar como
MORALES, MELIOSMA EN COSTA RICA Y PANAMA 943
una variedad morfologica de M. brenesii, no se han encontrado variaciones
7
mortfoldgicas y pes entre los caracteres claves en las diferentes poblaciones
examinadas en el campo que asi lo sugieran.
18. Meliosma vernicosa (Liebm.) Griseb., Cat. Pl. Cub. 47.1866. Lorenzanea vernicosa
Liebm., Vidensk. Meddel. pane Naturhist, — Kjobenhavn 1850: 72. he TES: sone
IC LAJUELA: Naranjo Jersted s.n. (HOLOTIPO: C f
13379, INB).
Oo
Distribuci6n.—Nicaragua, Costa Rica & Panama, 1100-2300 m.
Fenologia.—Flores observadas entre Junio y Julio.
Meliosma vernicosa ha sido comanmente confundida con M. depressiva,
pero se distingue por varios caracteres. En primer lugar, esta especie tiene hojas
con peciolos menos de 3.5 cm largo, en contraposicion de M. depressiva, que
tiene peciolos usualmente mas de 4.5 cm largo. La pubescencia de las hojas es
también diferente siendo las hojas de M. vernicosa glabras a glabradas.
Usualmente tiene inflorescencias mas compactas, con flores cremas a blancas.
Es importante aclarar para futuros monografiadores, que aunque la localidad
del especimen tipo exacta es desconocida, este fue probablemente recolectada
por Oersted en las faldas del Volcan Barva, en la Cordillera Volcanica Central,
donde realiz6 varias colecciones. En esta zona, M. vernicosa es localmente
abundante.
AGRADECIMIENTOS
Deseo agradecer a los curadores y directores de COL, CR, F HUA, JAUM, K,
MEDEL, MO, NY, US] por el préstamo y uso de colecciones. Asimismo, deseo dar
las gracias al personal del Herbario de la Universidad de Antioquia (HUA) y del
Herbario de la Fundacion Jardin Botanico Javier Antonio Uribe J AUM), Co-
lombia, por las facilidades logisticas brindadas durante mis visitas a Medellin.
Esta publicacién fue financiada por la Asistencia Holandesa para el Desarrollo
(NEDA)a través del proyecto “Desarrollo del Conocimiento de la Biodiversidad
Uso Sostenible en Costa Rica” conducido por el Instituto Nacional de
Biodiversidad (INBio). Ademas, la investigacion fue posible ademas gracias al
convenio de cooperacion entre el Ministerio de Ambiente y Energia (MINAE) e
INBio con el fin de completar en Inventario Nacional de Biodiversidad.
REFERENCIAS
Duran, C. 1997. Sabiaceae. In:V. Sosa et al.,eds. Flora de Veracruz. Fasciculo 96. Instituto de
Ecologia, Xalapa, Veracruz, México. Pp. 1-15.
Gentry, A.H. 1980. Sabiaceae.In:R. Woodson et al.,eds. Flora of Panama. Ann. Missouri Bot.
Gard. 67:949-963.
HitTon-TayLor, C (compilador). 2000. 2000 UICN Red list or threatened species. |UCN, Gland,
Switzerland and Cambridge, UK.
944 BRIT.ORG/SIDA 20(3)
BOOK NOTICES
Mark BLUMENTHAL. 2003. The ABC Clinical Guide to Herbs. (ISBN 1-58890-157-2
dk.). Thieme New York, 333 Seventh Avenue, New York, NY 10001, U.S.A.
(Or ( le PS: CUSLOTTIECTSer V ice@theime.con Jee 800-782-3488, 212-760-0888). $4995,
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This book must be the ultimate resource for information about each of the twenty-nine most fre-
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ucts which have been the subject of clinical studie
There are two pages of clinical overviews, a eee information sheet, and a comprehensive
I ach herb as well as two photographs and a line drawing. The primary and potential
uses, has macological actions, dosage and administration, adverse effects and the regulatory status
in various countries around the world are spelled out plainly for each herb. In addition, abundant
references are included.
This should be a valuable book for the lay person as well as the clinician at a time when alter-
native medicines are being widely discussed.—Ruth Ginsburg, Botanical Research Institute of Texas,
Fort Worth, TX, U.S.A
Gordon R. Hanks (Editor). 2002. Narcissus and Daffodil: The Genus Narcissus.
(ISBN 0-415-27344-7, hdk.). Taylor & Francis, |] New Fetter Lane, London
EC4P 4EFE, UK and 29 West 35" Street, New York, NY 10001, U.S.A. (Or-
ders: info@tandf.co.uk, wwwtandf.co.uk, 44 (0) 207 583 9855, fax 44 (0)
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pp, 16 color photos, 13 b/w photos, 98 b/w drawings, 31 tables, 7" x 9 3/4".
This is volume 21 of the Medicinal and Aromatic Plants—Industrial Profiles, a series aimed at those
engaged in research in botany, agriculture, pharmacognosy, phytochemistry, the food flavor, cos-
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ences. “Narcissus and Daffodil is the first book to provide a complete overview of the genus Narcis-
There are 19 chapters presenting original research material and information. Each chapter is
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couislels with references; some 1,700 references are scattered t
SIDA 20(3): 944. 2003
A NEW SPECIES OF PACKERA (ASTERACEAE:
SENECIONEAE) FROM THE EDWARDS PLATEAU OF TEXAS
Debra K.Trock' Robert J.O’Kennon
Botanical Research Institute of Texas Botanical Research Institute of Texas
509 Pecan Street 09 Pecan Street
Fort Worth, TX 76102-4060 U.S.A. Fort Worth, TX 76102-4060 U.S.A.
ABSTRACT
Packera texensis, sp. nov. is described and illustrated. It is endemic to the granitic sands of the Central
Mineral Region within the Edwards Plateau of central Texas where it is known from four counties.
RESUMEN
Se describe y se ilustra Packera texensis, sp. nov. Es ee os = arenas ee de la‘ a
Mineral Region” en la Meseta Edwards del centro de Texas
INTRODUCTION
Packera isa genus of 63 species that occur almost exclusively in North America
and that have been treated historically as the “Aureoid” complex of the genus
Senecio. Love and Léve (1976) pointed out that this anomalous group of species
has a chromosome base number of x = 22 or x = 23, unlike other members of
Senecio which are characterized by having a number of x = 10. They proposed
that Barkley’s “Aureoid Senecios” be placed in a new genus, Packera, based pri-
marily on gross morphology and chromosome number. The group has notably
imprecise species boundaries and a complex taxonomic history (Bain 1988; Bar-
kley 1962, 1963, 1968a, 1968b, 1978, 1980, 1988; Freeman & Barkley 1995; Kowal
1975; Packer 1972). During field work in central Texas, a species of Packera has
been discovered that is distinct from others currently known.
DESCRIPTION
Packera texensis R.J.O’Kennon & D.K. Trock, sp. nov. (Fig. 1). Type: U.S.A. TEXAS.
GILLESPIE Co.: Keese Road, 0.7 mi (11 km) N of Keese-Sagebeil Road, ca. 18 mi (29 km) N of
at bare in soils derived from weathered granite and among granite boulders, 1720 ft
(524 m), 5 Mar 2000, RJ. O’Kennon & D.K. Trock 11059 (HOLOTYPE: BRIT, ISOTYPES: MEX, MSC,
NY, TAES).
opter laminas 1.5-4 cm latas lobis terminalibus grandibus, denique petiolos distinguibiles a P.
plattensis arcte similans, sed am ca laminarum lateralibus irregularibus (non dentatis vel laceratis),
caudice usque ad 5mm (non 1-1.2) cm diametre, flosculis radii 13 (non 8), nec non habitu pereni (non
bieni) prae clare distat
Robust herbaceous perennial, arising from an erect woody taproot-like caudex
6-12 mm thick, gl to a {loccose about nodes and among heads of
1q 1d Mi hig 5 Uni i Biol East Lansing MI 48824-1312, U.S.A.
J
SIDA 20(3): 945-951. 2003
BRIT.ORG/SIDA 20(3)
946
\e Ae
BRIT)
Fic. 1. Hat
TROCK AND O’KENNON, A NEW SPECIES OF PACKERA FROM TEXAS 947
capitulescence. Stems 2-4(-5) dim, single or 2-5, rarely in clusters of up to 20.
Basal leaves petiolate, blades elliptic-ovate, broadly oblanceolate or lyrate, bases
tapering, margins irregularly and deeply parted to irregularly lobed with 3-9
lobes on each side and again incised at apex, 4-7 cm long and 1.5-2.5 cm wide,
usually about 2 times longer than wide, the petiole subequal to the blade; cauline
leaves gradually or sometimes abruptly reduced distally, similar in aspect to
the basal leaves. Capitulescence an open or congested corymbiform cyme of 4-
20(-30) heads. Capitula campanulate; phyllaries 13 or 21, 5-7 mm long. Ray
florets conspicuous, golden-yellow, ca. 13 in number, ligule 7-9(-10) mm long.
Cypselae, 2.5-3.0 mm long, 0.5mm wide, with thick appressed simple trichomes
along the angles, pappi 5-7 mm long.
Habitat and Phenology.—Gillespie County lies on the northern boundary
of the Balcones Canyonlands Association, an area of limestone plateaus deeply
dissected by creeks and rivers on the Edwards Plateau. The county is on the
southern boundary of the Llano Uplift or Central Mineral Region, where the
soil is primarily made up of granite, sandstone, gneiss, schist, and granitic-de-
rived sands. The Edwards Plateau is botanically well known and has long been
recognized as a region of endemism (Correll & Johnston 1979).
Recent collections of Packera texensis were made in an area of dry granitic
gruss near the boundary of Gillespie and Llano counties. The plants are site
specific and relatively uncommon, and the species is currently known from
only a few localities. Packera texensis arises quickly from evergreen winter ro-
settes during the January rains and blooms as early as the first week in Febru-
ary. Plants begin to senesce by the end of March during a rapid drying out of
the substrate prior to the spring mia season that beings in late April.
Additional collections jaca U.S.A. TEXAS. Callahan Co.: I-20 (old Rt. 80) frontage road N side,
1 km E of County Road CR 115, 15 Apr 2002, RY. ye Kennon 16020 (BRIT); 3 mi S of Eula along Hwy
603, 20 Mar 1963, N.C. ae rson 63-45 (BRIT); 2 mi E of Clyde along Hwy 80, sandy roadside, 8 Apr
1962, N.C. Henderson 62-67 (BRIT). Gillespie Co.: 3.5 mi N of Eckert on Hwy 16, 17 Mar 1964, DS.
Correll 29037 (BRIT), Keese Road, 0.7 mi (11 km) N of Keese-Sagebeil Road, ca. 18 mi (29 km) N of
Fredericksburg, decaying granitic gruss soils among granite boulders, 1720 ft (524 m), 20 Mar 1990,
RJ.O’Kennon 6577 (BRIT); 26 Feb 1991, RJ. O’Kennon 8373 (BRIT); old Llano Road N of Fredericksburg,
21 Feb 1931, E. Whitehouse 12050 (BRIT). Llano Co.: Enchanted Rock, granitic soil, partial shade, 18
Apr 1931, E. alae 12054 (BRIT). Mason Co.: | mi S of Katemcy, granite outcrop, 5 Apr 1984, S.
Pence 82 (BRIT
DISCUSSION
Packera texensis most closely resembles P. tridenticulata (Rydb.) W.A. Weber
S& A. Love but it shares characteristics of P tampicana (DC) C.Jeffrey as well as
P plattensis (Nutt.) WA. Weber & Léve. Most specimens would key to Senecio
tridenticulata (Rydb.) in Correll and Johnston’s Manual of the Vascular Plants
of Texas (1979). Some of the more pubescent specimens would key to Senecio
plattensis (Nutt.).
948 BRIT.ORG/SIDA 20(3)
Morphologically the new species differs from P. tridenticulata mainly in
height, shape and size of leaves, leaf margins, ratio of petiole length to blade
length, length of phyllaries, length and pubescence of achenes. It differs from P
plattensis in number of stems, type of perennating structures, amount of loca-
tion of pubescence, number of ray florets, length of ligules, length and pubes-
cence of achenes. It differs from P tampicana in duration, number of stems,
pubescence, overall leaf shape and degree of lobing of the margins, and num-
ber of ray florets. Characters useful in recognizing these species are listed in
Table 1. Data in this table were obtained by the senior author from measure-
ments made on dozens of herbarium specimens.
There are three other species of Packera in Texas, P.glabella, P obovata and
P. tomentosa. There is little likelihood that any of these species could be mis-
taken for P texensis. Packera glabella has distinctly striated hollow stems, a short
fibrous-rooted caudex, and leaves with margins pinnately lobed. Packera
ovobata has abundant stolens, a rhizomatous caudex, and distinctly obovate
leaves and P tomentosa has persistent floccose pubescence covering all of the
vegetative surfaces. In contrast to these three species P. texensis lacks the hol-
low striated stems, has a distinctly tap-rooted caudex, no stolons, leaves that
are elliptic-ovate to oblanceolate with margins that are at most irregularly lobed
and pubescence restricted to the leaf axils, lower petioles and inflorescence.
There are habitat and substrate preferences that may also be used to dis-
tinguish these four species. Packera texensis has been collected only on or near
the Edwards Plateau in soils derived from igneous or metamorphosed igneous
rocks such as granite, gneiss and schists. Packera plattensis has been collected
from a wide variety of soil types, but is most abundant in the limestone and
sandstone-derived soils of the Great Plains. Packera tridenticulata isa plant of
the High Plains and foothills of the Rocky Mountains, while P tampicana is
—
en
always found in disturbed wet sandy or muddy sites. Distribution maps for the
four species are provided in Figure 2.
Hybridization is apparently common in this genus, but there is no evidence
that P texensis hybridizes with any of its congeners. It is consistently distinct
in morphology, habitat and substrate preference, and flowering phenologies.
There is also good distributional data to support the recognition of the new
species. Additional field and laboratory studies will undoubtedly contribute
to our understanding of the relationship of P.texensis to other morphologically
similar species.
KEY TO PACKERA TEXENSIS AND ITS IMMEDIATE ALLIES
Plants permanently floccose a at the base of the stem and in the axils of
the leaves, lightly tomentose el P. plattensis
. Plants with light tomentum at rie ie of the stem and in leaf axils, glabrous
elsewhere.
Tasle 1. Comparison of Packera texensis with selected congeners.
P. texensis
P. tridenticulata
P. plattensis
P.tampicana
Duration and habit
Height
Number of stems
Pubescence
Leaf dimensions
Petiole to
blade ratio
Leaf blade shape
Leaf margins
Capitulescence
Phyllaries
Ray florets
Ligule length
Cypselae
Perennial, woody caudex,
4-12 mm thick, no stolons
3-4(-5) dm
Single to several
Glabrous to lightly
loccose near nodes
and among capitula
Blades 4—8(-10) cm
long, 1.5-2.5 cm wide
Petioles subequal to blade
Elliptic-ovate to
broadly oblanceolate
Irregularly incised or
dissected to deeply
arted or pinnatisect,
occasionally lateral
margins irregularly lobed
4—20(-30) capitula
13 or 21,5-7 mm long,
green, glabrous
About 13
5-6(-7) mm
1-1.5 mm long, thick
appressed trichomes on
the angles
Perennial, caudex,
2-4 mm thick, terminally
branching taproot
—3(-4) dm
Single to several
Glabrous to lightly
floccose near nodes
Blades 2-4(—5) cm long,
0.5-1.5 cm wide
1-2 times longer
than blade
Lanceolate to narrowly
oblanceoate
Entire, subentire or
dentate near the apex
rarely sub- pinnatisect.
4—12(-20) capitula
13 or 21,6-10 mm,
green, sparsely tomentose
proximally
8-10, rarely 13
5-8(-9) mm
1.5-2.5 mm long,
glabrous or lightly
hirtellous on the angles
Biennial or perennial from
erect or suberect caudex,
may be stoloniferous
2-6(-7) dm
Usually single, rarely 2-3
Floccose-tomentose at
base of stem, in nodes,
abaxial leaf surface and
among capitula
Blades 2-7(-8) cm long,
1-3(-4) cm wide
1-1.5 times longer than
blade
Elliptic-ovate to oblanceo-
late or rarely suborbicular
Subentire to crenate,
serrate-dentate or
sublyrate
13 or 21,5-6 mm,
green, - nsely
tomentose Samal)
About 8
9-10(-11) mm
1.5-2.5 mm, hirtellous
on the angles or
rarely glabrous
Annual from a
slender taproot
—5(-6) dm
Usually single, or rarely 2-6 clustered
Glabrous throughout
Blades 4-12 cm long,
1-3(-4) cm wide
Petioles subequal to
blade
Oblanceolate to spatulate
Deeply pinnate, 1-6
pairs of lateral lobes,
large terminal lobes
—15(-20) capitula
13 or 21,3-7 mm
green with reddish
apices, glabrous
8 or 13
3-7 mm
1-1.5 mm long,
hirtellous on the
angles
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Fic. 2. Geographic distribution of f phologically similar species of Packera in Texas. Circles = P texensis; triangles
D ¢etd nl mT D nlate. De Nicerthut h d ' ott £ 2nn
herbarium specimens.
2. Basal leaves lanceolate or narrowly oblanceolate, margins entire, subentire to
oe near the apex or occasionally sub-pinnatise P. tridenticulata
N
. Basal leaves elliptic-ovate, obovate or lyrate, margins lobed, parted or deeply
incised.
3. Annual, from a slender taproot
3. Perennial, from a stout, woody tap rootec
P. tampicana
caudex P. texensis
2).
ACKNOWLEDGMENTS
The authors would like to acknowledge Ted Barkley and Alan Prather for their
assistance with various drafts of this manuscript. Also thanks to Guy Nesom
and John Bain for providing their insightful comments. Special thanks go to
Linny Heagy for her wonderful illustrations and to John Pipoly for providing
the Latin diagnosis. We would also like to thank the following herbaria and
their curators for the loan of specimens: BRIT, BRY, KANU, KSC, LSU, MICH,
MO, MONT, MSC, MU, NDA, NLU, OSC, RSA, TEX, UC, UNM, VDB
TROCK AND O’KENNON, A NEW SPECIES OF PACKERA FROM TEXAS 951
REFERENCES
Bain, J.F. 1988. Taxonomy of Senecio streptanthifolius Greene. Rnodora 90:277-312.
Barkiey, 1.M.1962.A revision of Senecio aureus L.and allied species. Trans. Kansas Acad. Sci.
65:318-408.
Bark ey, 1.M. 1963.The intergradation of Senecio plattensis and Senecio pauperculus in Wis-
consin. Rhodora 65:65-67.
Barkvey, .M.1968a.Intergradation of Senecio sections Aurel, fomentosi,and Lobati through
Senecio mutabilis. Southw. Naturalist 13:109-115.
Barktey, T.M. 1968b. Taxonomy of Senecio multilobatus and its allies. Brittonia 20:267-284.
Barktey, T.M. 1978. Senecio N. Amer. flora II. 10:50-139.
Bark.ey, T.M. 1980. Taxonomic notes on Senecio tomentosus and its allies (Asteraceae).
Brittonia 32:291-308.
Barktey, .M. 1988. Variation among the aureoid Senecios of North America:a geohistorical
interpretation. Bot. Rev. 54:82-106.
Corrett, D.S.and M.C. JOHNSTON. 1979. Manual of the vascular plants of Texas. Univ. of Texas,
Dallas.
Freeman, C.C. and T.M. Barkley. 1995. A synopsis of the genus Packera (Asteraceae:
Senecioneae) in Mexico. Sida 16:699-709.
Kowat,R.R. 1975. Systematics of Senecio aureus and allied species on the Gaspe’ Penninsula,
Quebec. Mem. Torrey Bot. Club 23:1-113.
Love, A.and D. Love. 1976. Nomenclatural notes on Arctic plants. Bot. Not. 128:497-523.
Packer, J.G. 1972. A taxonomic and phytogeographical review of some arctic and alpine
Senecio species. Canad. J. Bot. 50:507-518.
952 BRIT.ORG/SIDA 20(3)
BOOK NOTICES
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6 b/w photos, 60 b/w drawings, 45 tables, 7" x 93/4",
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SARA OLDFIELD. 2003. Rainforests. (ISBN 0-262-15106-5 hbk.). The MIT Press, 5
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ened trees.—R
SIDA 20(3): 952. 2003
STUDIES IN GLANDULARIA (VERBENACEAE):
THE GLANDULARIA QUADRANGULATA COMPLEX
James Henrickson
Department of Biological Sciences
California State University
Los Angeles, CA 90032, U.S.A.
ABSTRACT
This paper responds to Turner's inclusion of Shandon vere cunda and G. racemosa into G,
]
quadrangulata. The latter |
outward at the base; G. verecunda, is very similat veperat ce has similar nutlets, but they ck the
distinctive terminal appendage. Pollen/ovule ratios indicate that both taxa are facultative autoga-
us and both taxa have distinct ranges with limited areas of sympally, ne inter paecliates are known,
oa intermediate characteristics may not exist. They also differ
porting their recognition as distinct species. Glandularia racemosa, with larger noel ¢ a averent
nutlet type, and leaf structure, is a hexaploid (not tetraploid as in the other taxa) and represents a
separate group within the genus and should not have been submerged into G. quadrangulata.
RESUMEN
Este articulo responde a la inclusion de Turner de Glandularia pane G. racemosa en G.
Pee Esta ultima tiene apéndices terminales distin n las nuculas y el pericarpo se
acampana hacia el exterior en la base; G. verecunda, es muy similar vegetativamente, tiene nuculas
similares, pero le falta el apéndice terminal distintivo. La relacion polen/ovulos indica que ambos
taxa son autogamos facultativos y ae taxa tienen irene eiuanucien con areas de simpatria
limitada. No se conocen intermedios, y los no existan. También se
diferencian en su dotacion de [lavonoides que apoyan su ~anotee mien como especies distintas.
Glandularia racemosa, con fede mas grandes, un tipo diferente de nucula, y estructura de la hoja,
es un hexaploide (no tetraploide como en los otros taxa) y grupo separado en el género
y no deberia haber sido incluida en G. quadrangu
[
atd.
INTRODUCTION
Turner (1998) presented a new classification Glandularia quadrangulata
(Heller) Umber submerging both G. verecunda Umber and G. racemosa (Eggert)
Umber into G. quadrangulata. All three taxa are similar vegetatively. They are
low-growing, branched, hairy, usually perennial herbs with broad, distally dis-
sected leaves and have small, inconspicuous flowers with corolla limbs only
2.5-9 mm wide. Turner (1998) however, failed to take into consideration basic
differences in nutlet structure, ploidy levels, and breeding systems of these taxa
that support a different classification.
MATERIALS AND METHODS
The data presented here are based on empirical observations on specimens from
the TEX-LL, SRSC herbaria. Direct counts of pollen grains were made from
SIDA 20(3): 953-967. 2003
954 BRIT.ORG/SIDA 20(3)
unopened buds that were softened by boiling; the four anthers were opened
directly on microscope slide.
RESULTS AND DISCUSSION
Nutlet structure.—Nutlets of G. quadrangulata are the most distinctive (Fig. |
D). The nutlets are 3.3-4.2 mm long, the central bodies are slender, 0.55-0.65
mm wide, at the base and the outer pericarp wall greatly expands and flares
outward to 1.2-L6 mm in width. At the inner tip they have a large, thin, some-
what concave, ovate appendage 1.0-1.3 mm long. The inner or commissural sur-
ace of the nutlet is cream-white and covered with distinct papillate-aculeate
excrescences. The commissural surface is nearly as wide as the nutlet in the
central portion and extends to the top of the ovate appendage. The outer peri-
carp walls, as in most glandularias, are black, with a broad reticulate, raised
pattern that extends from the tip about half way down the nutlets where it
disorganizes into a series of vertical striae that extend to the nutlet base. The
quartet of nutlets are so arranged that the flared bases expand over the recep-
tacle and the distal appendages form a valvate dome over the developing nut-
lets through which the style extends from the tip of the united appendages.
When the nutlets separate, they leave no distinctive receptacular rim.
The fruits of Glandularia verecunda are similar to those of G.
quadrangulata except they lack the conspicuous terminal appendage (Fig. |
H). They are (2.6-)2.8-3.2(-3.3) mm long, and the central body is also slender,
0.5-0.65 mm wide. The outer pericarp also abruptly flares out at the base toa
width of 10-13 mm although this characteristic is not present in immature
fruit. Again the inner whitish commissural face is nearly as wide as the central
nutlet itself. The aculeate surface continues to the distal portion of the nutlet
where its distal margin is rather truncated, sometimes retuse, marked by small
horizontal beak that extends inward. The horizontal beak may be homologous
tothe much-expanded vertical beak in G. quadrangulata. In their broadly flared
base and slender body, the broad commissural face of the nutlets are identical
to those of G. quadrangulata.
Nutlets with similar slender bodies and greatly expanded bases occur ina
small group of North American Glandularia, namely G. polyantha Umber, G.
delticola (Small) Umber, and in two other taxa that also have greatly expanded
air spaces in the adaxial portion of the nutlet: G. tumidula (Perry) Umber, and
G. brachyrhynchos Nesom & Vorobik. Of note, the distinctive nutlet appendage
is not unique to G. quadrangulata as it also occurs in species of southern South
America (Uruguay, Brazil and Argentina) where Glandularia tenera (Spreng)
Cabrera has an appendage similar to that of G. quadrangulata and G. pulchella
(Sweet) Tronc. has a shorter, less-developed appendage. Glandularia pulchella
is adventive in the United States from Florida into Texas and in northeastern
—
—
HENRICKSON, STUDIES IN GLANDULARIA 955
dark J J
verecunda. AD. G. quadrangulata A. Stem, showi 1 infl
heen LL); B. Leaf, adaxial view, (Lundell 9790, LL); C.-Leaf, adanialviewltuner aT: 112 , TEX); D. Mature nutlets
showing the distinctive ovate terminal appendage and broadly flaring basal pericarp as seen in abaxial view (left)
showing outer reticulate tee (note broad expansion of nutlet base), in adaxial view (central) showing aculeate
inner commissural surface nearly as wide as the nutlet, _ in ai view Adah) Sue adil ioeation of the
Sea E an 15923, TEX).E-H.G. verecunda. E
TE) . poo acanelvey iS Onmeyer he TEX); G aes Jaxial vi a kson 22775, TEX); H.M lets st
wide as the ull nutlet and continuing 6 near ou OuueR tip, abaxial view ( 1) showing abaxial nutlet surface
I p p llel lines below), and | | view (right) show-
th I haal iil oe tha tin fall 2 © Donn 22 Try) MA a . a ous
P\ ’ } bo
956 BRIT.ORG/SIDA 20(3)
Mexico; most plants show moderate appendages, in some plants they are poorly
developed, shorter and more perpendicular to the nutlet axis. Nutlets of both
species are illustrated in Burkart and Troncoso (1979).
The nutlets of Glandularia racemosa, however, are strongly different and
are of atype widely occurring through most North American Glandularia (Fig.
pa
the basal pericarps are only slightly expanded and the nutlets are rounded at
the tip. Furthermore the whitish aculeate or otherwise marked commissural
surfaces are much narrower than the width of the nutlets (often 0.25-0.3(-0.35)
mm wide, with the nutlet being (0.55-)0.6-0.77 mm wide), and this whitish
commissural surfaces extends to within 0.2-0.4 mm of the dome-like nutlet
tips where they are distinctly rounded, not truncated or retuse. The blackened
pericarp, however, has a similar raised reticulate pattern in the distal half that
disorganizes into a series of vertical striae as in the other two species.
Cytology.—Umber (1977, 1979) presents chromosome counts for the three
taxa. Both Glandularia quadrangulataand G. verecundaare tetraploid with n =
LO. Glandularia racemosa in contrast, is hexaploid with n = 15. An older count
by Dermen (1936) also shows hexaploidy for G. racemosa (2n = 30). This would
not support submersion of G. racemosa into G. quadrangulata. Lewis and Ol-
iver (1961) report n = 10 (tetraploid) for G. racemosa based on a Lewis & Oliver
5416 collection from Coahuila, Mexico, however, Umber (1977) correctly notes
this collection is G. verecunda, not G. racemosa.
Phytochemistry.—Umber (1977) notes that the west Texas populations of
Glandularia verecunda and G. racemosa are similar in their flavanoids both
producing 7-0-gentiobiosides and 7-O-sophorosides of apigenin, luteolin and
chrysoeriol, while six south-Texas samples G. quadrangulata produced only 7-
0-gentiobiosides (but not 7-0-sophorosides) of apigenin (its dominant fla-
vonoid), with only trace amounts of 7-O-gentiobiosides of luteolin and
chrysoeriol. The south Texas sample of G. verecunda, however, also produced
only 7-O-gentiobiosides of apigenin, luteolin and chrysoeriol as in G.
quadrangulata.
Biogeography.—Each of these three taxa have distinct ranges: Glandularia
quadrangulata occurs from south-central Texas west to the Pecos River and to
the border region of Mexico (Fig. 3). Glandularia verecunda occurs in trans-
Pecos Texas, scattered in southern Texas, and continues south into Mexico in
eastern Chihuahua, eastern Coahuila, south to near Monterrey in Nuevo Leon.
Glandularia racemosa is restricted to trans-Pecos Texas (Fig. 3).
Breeding systems.—Flowers in both Glandularia quadrangulata and G.
verecunda are very small having white to lavender corollas with very slender
tubes 5.2-6.5 mm in length and limbs 2.5-5.2 mm in diameter. Within the tube,
as in all glandularias, the style is initially situated between the two pair of in-
trorse anthers among a series of downward-pointing stiff hairs (see Fig. 2 E-H).
HENRICKSON, STUDIES IN GLANDULARIA 957
A
. »
Ax i Se
EN
21- 212, TEX); C. leak adaxial view (Turner 97-28, TEX); D.-H. Floral
structure Sace re-expanded flowers of Turner 21- 74 TEX and others) D. Corolla, face view, showing larger, more emar-
alnate gel corals limb, and ey ake ae is E. mene ae uion of le ae Svar of
Fic. . alee racemosa. A. cre habit of y
21-2
e of the tube
are stiffer, more busta: F. Medial section of calyx, showi tyle and stigma (offset). Tt g
p * G. Distal stamens, showi g| teral qlandul . | U Nictal fee
1 associated hairs ( tive appendage on the distal st offset) il al nutlts showing
lett | relativel i If lateral view (left) showi f adaxial
/ wl
rt re Pace | Yer | H +} . f | ee ec ype a ee | * L Les | £ +
VIEW | VICW |
that is less organized below (Warnock 46135, TEX). Magnificati {i 1. Scale in D, holds for D, E, and F.
958 BRIT.ORG/SIDA 20(3)
Glandularia quandrangulata
Glandularia verecunda
Glandularia racemosa
O.
nt hy 4 , er a 4 { ee ie 1 \ 4 (hk i _ 1 \ Pe al
Fic. 3
Texas and Mexico, from collections at TEX-LL and SRSC.
The basal two anthers are 0.3-0.6 mm long, with filaments to 0.3 mm long and
contain fewer pollen grains than present in the slightly larger distal pair of anthers
that are 0.4-0.8 mm long with filaments 0.3 mm long. As the flower matures,
the style elongates pushing the stigma across the distal pair of anthers where
pollen easily contacts the stigma. Unless some self-incompatability factors are
at work, autogamy (inbreeding) is certainly possible. Regarding self incompat-
ibility, | have observed that even isolated plants of G. quadrangulata set a full
compliment of nutlets, and in most herbarium specimens, the plants have full
HENRICKSON, STUDIES IN GLANDULARIA 959
fruit set, leading me to believe that there is no self incompatibility involved in
the reproduction of the taxon.
Cruden (1977) noted that pollen-ovule ratios can be used as an indicator of
the breeding system in flowering plants. In his study, he indicated average pol-
len/ovule ratios of 4.7 (range 3-7) for cleistogamous taxa; 27.7 (range18-30) for
obligate autogamous taxa; 168.5 (range 76-396) for facultative autogamous taxa;
796.6 (range 160-2,558) for facultative xenogamous (outbreeding) taxa; and 5859
(range 1,648-16,807) for obligate xenogamous taxa using his outcrossing index
as a relative measure of inbreeding vs. outcrossing.
Glandularia species in North America differ strongly in corolla size, with
larger flowers having larger anthers with more pollen. 1 measured pollen grain
numbers per flower and correlated that with overall flower size (using maxi-
mum expanded diameter of the corolla limb asa gross measurement of corolla
size) in several species of Glandularia. | found that plants with corolla limbs 9-
13 mm in diameter had 2168-2077 pollen grains (5 counts) (average 2344 per
flower, 586 pollen grains per ovule-four ovules per ovary). Flowers with corolla
limbs 4-7 mm in diameter had 956-1410 pollen grains per flower (8
counts)(average 1275 per flower, 318 pollens per ovule) and those with corolla
limbs 2-4 mm in diameter, had 580-1149 pollen grains (7 counts) (average 748
per flower, 187 pollen grains per ovule) indicating that most Glandularia were
in the range of facultative autogamous. As noted above, the stigma is also pro-
duced in close association with the anthers in the distal corolla beg Pa the
species here in question, I found in four collections of G. quadrangulata, 580,
626,615, 706 pollen per flower (mean of 157 per ovule-4 ov les per Slower) (wet-
ted corolla limb diameters 2.5, 2.5, 2.7, 2.5 mm respectively-see Table 1); for G.
verecunda 1149, 870, 740, 692, pollen per flowers (mean of 218 per ovule), (co-
rolla limb diameters of 2.5, 3.5, 5.2, 3.5 mm) and G. racemosa 2184, 2125, 1728,
1803 per flower (mean of 490 per ovule), (corolla limb diameters of 9.0, 8.5, 8.0,
7.8mm). This would imply that the taxa, at least for G. quadrangulata and G.
verecunda, are in the range of facultative autogamous taxa and G. racemosa is
in the facultative xenogamous range.
Also, most plants, even isolated plants, exhibit full or nearly full fruit set.
Umber, (pers. comm.) notes that in many Glandularia species, greenhouse
grown plants produced full fruit set, in some species you had to push the flow-
ering branches back and forth, similar toa wind action, to insure good fruit set.
Even the few, widely isolated specimens of G. quadrangulata | observed in
Kleberg Co., Texas, all had near complete fruit maturation. Though Umber 1977
notes that some populations of his G. bipinnatifida have low fruit set, indicat-
ing the possibility of local self incompatability. My conclusion is that G.
quandrangulata and G. verecunda are capable of reproducing by autogamy, and
that self incompatibility does not restrict fruit set, so that even if they occur in
sympatric populations, gene exchange between them may be limited and inci-
960 BRIT.ORG/SIDA 20(3)
Taste 1.Pollen/ovule ratios of Glandularia species.
Total pollen p/o Corolla
per fl. ratio diameter
Glandularia quadrangulata
Turner 97-01 580 145 2.5
Crockett Co., Tex.
WR. Carr 11723 615 153 25
McMullen Co., Tex.
Williges 359 626 156 2.7
San Patricio Co,, Tex.
V. Villarreal 706 176 2.5
Kleberg Co., Tex.
Glandularia verecunda
Correll 20699 1149 287 2.5
rewster Co,, Tex.
Powell, Turner & Sikes 870 217 35
Ojinaga, Chih., Mex.
Turner 97-86 780 195 52
Terrell Co., Tex.
Lundell & Lundell 10071 692 173 35
Brooks Co., Tex.
Glandularia racemosa
Turner 97-28 2184 546 9.0
Pecos Co,, Tex.
Turner 97-2712 2125 531 8.5
Culberson Co,, Tex.
Turner 97-76 1728 432 8.0
Brewster Co., Tex.
Tharp& Havard 49359 1803 451 78
Pecos Co., Tex.
Average 490
dental. Certainly both taxa produce flowers with small corolla tubes that may
be attractive to long-tongued flies J. Neff, pers. comm.) but are certainly ca-
pable of self pollination.
Classification.—The relationship of Glandularia racemosa to the other two
taxa, is very clear. While G. racemosa is somewhat vegetatively similar to both
G. quadrangulata and G. verecunda, the leaves of G. racemosa are more deeply
divided into linear segments (except in some large, perhaps shaded leaves of
rapidly-growing specimens), its growth habit is different, initially briefly de-
cumbentat the base with erect-ascending stems, but with age it becomes more
spreading. Furthermore the fruits of G. racemosa are completely different as
—_—
HENRICKSON, STUDIES IN GLANDULARIA 961
cliscussed above and are of a type found in most North American Glandularia.
Also both G. quadrangulata and G. verecunda are tetraploid (n = 10), while G.
racemosa is hexaploid (n = 15). In addition the corollas of G. racemosa are larger
[(5.5-)6-8(-9.0) mm in limb diameter] than those of G. quadrangulata|(2.0-)2.5-
3.5(-4.5) mm diameter] and G. verecunda [3.2-4.5(-5.5) mm diameter]. Corolla
limbs of G. racemosa are also usually white in color [rarely tinged with blue—
which Umber (1977, 1979) considers indication of introgression with G.
bipinnatifidal, while those of G. quadrangulataand G. verecunda are more com-
monly lavender, only occasionally white in color. It isapparent that G. racemosa
is not a taxonomic synonym of the G. qudadrangulata-verecunda group and isa
distinct, endemic species in trans-Pecos Texas. I find no support for Turner
inclusion of G. racemosa into his concept of G. quadrangulata other than gross
vegetative similarity.
While Glandularia quadrangulata and G. verecunda have largely allopat-
ric ranges, mixed populations occur near the Pecos River and in southern Texas.
In fact there are three herbarium collections at TEX-LL that contain material
of both taxa: two from Upton County, Texas, 3.1 miles east of McCamey, 21 Apr
1999, Turner 99-179 and 99-200 (TEX) and one from southern Texas, Kleberg
Co., Texas; Hwy. 77, 2 mi W of Riviera, 15 Mar 1965, Ramerez & Cardenas 78
(TEX). An additional pair of specimens from SRSC (Brewster Co., Texas, sandy
soil along the Rio Grande at Boquillas, 10 May 1933, H.T. Fletcher 1224 (G.
quandrangulata) 1226(G. verecunda) were apparently collected in the same area.
In vestiture, leaf size and shape the two taxa are very similar, if not identical.
The overall similarity and occasional sympatry of Glandularia
quadrangulata and G. verecunda caused Turner (1998) to conclude that G.
verecunda was only a ex-appendiculate form of V. quadrangulata unworthy of
formal nomenclatural recognition although he presented no evidence of intro-
gression. But we must ask, what characteristics would be expected in hybrids?
Would they have intermediate appendages or no appendage at all and be iden-
tical with G. verecunda?
No plants have every been discovered in nature with nutlets intermediate
in structure between those of G. quadrangulata and G. verecunda. All plants
either have nutlets with or without the distinctive distal appendage. Further-
more, no artificial hybridization has been done to determine the condition that
would be expressed ina hybrid. The pattern found with G. quadrangulata and
G. verecunda presents an interesting conundrum. Are the two taxa merely forms
of one other, as Turner espouses, separated by only a single character that is
either expressed or not, so that hybrids would be identical to one taxon or an-
other? Or are they two separate species that do not or seldom hybridize? Or
should they be recognized at least at the infraspecific level?
It remains that the two taxa are nearly identical vegetatively and florally
except for the distal nutlet appendage on G. quadrangulata. | concede that the
962 BRIT.ORG/SIDA 20(3)
two are most-closely related or sister taxa; to think otherwise would be diffi-
cult to defend. But there seems to be a stability in their lineages as indicated by
their respective large and mostly independent geographical ranges.
The genetic basis of the nutlet-appendage character is not known. If itisa
simple Mendelian trait, it would appear not to have a high mutation rate. If the
appendaged nutlet type in Glandularia quadrangulata were pleisomorphic and
dominant state (.e., AA, or Aa) and the non-appendaged nutlets represented a
loss of this character (ie., aa), then once a mutation (a) forms, or once hybrid-
ization occurs, one would expect the heteozygotic plants to form seeds at a 1:2:]
genotypic ratio (AA:2Aa:laa), a phenotypic ratio of 3 appendaged to | non-
appendaged character state while the homozygous plants would breed true over
successive generations. The same pattern would be expected if the appendage
were a recessive character state. Whether this is occurring in certain localities
is unknown. But it remains that large geographical regions are occupied by
plants with either appendaged or non-appendaged nutlets indicating a low rate
of mutation or hybridization. If mutation rates were higher, or hybridization
more frequent, homozygous recessive traits would be expected throughout the
dominant-trait populations.
Turner (1989) has treated the two taxa as unnamed forms, relegating
Umber’s G. verecunda to obscurity. But if the two taxa are facultative autoga-
mous, they could very well become sympatric without mixing and thus be-
have much like species. Likewise if they are allopatric, and mutation rate is
very low, again they would breed true to form and behave as species. To relegate
the two taxa to unnamed forma, is not reflective of the conditions observed in
nature. In typical formas, the homozygous recessive character states usually
occur scattered in some or many populations, their frequency reflecting the
frequency of the recessive trait. These two taxa behave more like species. They
each have broad geographical ranges. The distinguishing character state is con-
spicuous. There is a possibility that in some populations, where both taxa are
present, they may have introgressed and mixed populations are being produced
through recombination. But as there is no intermediate character state, the popu-
lations would appear to be consisting of two sympatric separate taxa. If they
actually are interbreeding, there would be no way to tell except by growing the
nutlets out from these mixed populations to see if they breed true to form. |
have visited two populations where the two taxa are sympatric, west of Riviera,
in Kleberg Co., Texas and west of McCamby, in Upton Co., Texas in the late spring
of 2001. In the former the roadside populations are largely being replaced by
Bermuda Grass, but some nutlets were collected from scattered specimens of
G. quadrangulata. In the Upton County populations, no plants were found pre-
sumably due to local rainfall conditions.
Of the three nomenclatural possibilities: 1) being inconsequential forms
of each other; (2) being geographical subunits of a species, i.e., subspecies or
HENRICKSON, STUDIES IN GLANDULARIA 963
varieties of one species; or (3) being separate species that merely co-occur in
some areas. I consider that the first choice is least supportable. Mixed popula-
tions are few, no plants with intermediate fruit conditions are known to exist,
plants with the two fruit types occur in broad, somewhat discrete geographical
ranges. This treatment would ignore the pattern of variation as it would not be
reflected in the nomenclature. One could argue, just as strongly, that the two
taxa are distinct species that are very similar vegetatively that can co-occur
without interbreeding as no intermediates have every been found. But the lack
of interbreeding may be based in autogamy more than some other type of iso-
lating mechanism.
If there was evidence of interbreeding, they the two taxa could be recog-
nized at the infraspecific level. But this does not exist. Therefore the two taxa
will here be recognized as distinct species.
Glandularia quadrangulata (Heller) Umber (Fig. 1 A-D). Basionym: Verbena
quadrangulata Heller, Contr. Herb. Franklin Marshall Coll. 1:84. pl. 6.1895. Type: U.S.A. TEXAS.
NUECES Co. Corpus Christi, 5 Mar 1894, Heller 1388 (HOLOTYPE: (in part) NY; ISOTYPES: GH, MQ).
Verbena pumila ft. albiflora StandL, Field. Mus. Publ. Bot. 4: 256.1929. Type: U.S.A. TEXAS. BEXAR
Co. San Antonio, L.E. Pagel 2208 (HOLOTYPE: F).
Weak-stemmed annuals 0.5-2 dm tall; stems several from the base, 1-3(-7) dm
long, initially briefly decumbent, erect above, with age spreading, decumbent,
ascending distally, the internodes (1.2-)2-5(-11) cm long, green to tan, some-
times purple with age, grooved on two sides when young , this obscured when
larger, moderately (to strongly below inflorescence) hirtellous to pilose with
slender, tapering erect, mostly straight hairs 0.2-1.2(-L.8) mm long, sometimes
with stipitate glands to 0.2 mm long. Leaves ovate to ovate-rhombic in outline
(12-)15-25(-38) mm long, (8.5-)10-20(-30) mm wide, usually much shorter than
the internodes, with petioles 2-1] mm long, the blades 3-5 cleft about half, two-
thirds way to the midrib, the clefts entire to few to strongly toothed and lobed,
usually moreso on the outer margins, ultimately the leaf with 7-22 oblong to
oblong-obovate, obtuse-rounded, sometimes acute lobes 1-2.5(-3.5) mm wide,
the blades flat to usually somewhat concave and with impressed primary-sec-
ondary veins above, with rather stiff, tapering unicellular hairs 0.3-0.8-L.1) mm
long on both surfaces, these generally dispersed and ascending above, mostly
along the yellowish raised veins and the slightly revolute margins (or + uni-
formly scattered) beneath, those of the upper sur face sometimes pustulate based.
Spikes terminal on all branches, initially crowded with a short peduncle, 9-11
mm wide, with age the inflorescence axis elongating below, and the lowermost
developing fruit becoming increasingly more remote, until separated by 5-7
mm along the axis, the inflorescences then extending 4-13 cm long with a pe-
duncle to5cm long, the spike rachis + densely pilose with hairs to 1.8 mm long;
bracts ascending, linear-lanceolate, 3.2-5.5 mm long, hirsute along margins and
964 BRIT.ORG/SIDA 20(3)
midvein with hairs 0.2-1.6 mm long, 59-79 percent as long as the calyx; calyces
initially 5.0-6.5 mm long, membranous between the 5 green veins, hirtellous
and hirsute with hairs 0.2-1.2 mm long, without or without stipitate glands,
the lobes unequal, 0.4-0.8 mm long, green or purple tipped, in fruit the calyces
expanding around the nutlet becoming strongly trunctated and to 2.5mm broad
at the base; corollas slaverform, the slender tubes 5.2-6.6 mm long, 0.4-0.6 mm
wide at the throat, the limbs white to lavender, (2.0-)2.5-3.5¢4.5) mm wide, the
5 lobes all retuse, or the upper two lobes not retuse, |.0-1.2 mm long, 0.5-1.3 mm
wide; the tube orifice with gold-white moniliform hairs to 0.5 mm long; distal
anthers 0.4-0.5 mm long, the anther gland obscure, with filaments 0.3 mm long,
the proximal anthers 0.3 mm long, with filaments 0.2 mm long; style 4-5.2 mm
long. Mature fruit (3.3-)3.5-4.2 mm long, the distal, adaxial, obtuse, ovate ap-
pendage 1.0-1.4 mm long, 0.8-1.0 mm wide, the central body cylindrical, 0.55-
0.65 mm wide, the cream-colored commissural face 0.38-0.55 mm wide, this
—
—
extending throughout the distal appendage, strongly aculeate with papillae to
0.25 mm long, the basal pericarp expanded, forming a hollow chamber to L.0-
1.6mm wide, the outer pericarp black, shallowly reticulate in the distal half,
the reticulate pattern disorganizing into vertical striae in the lower half, the
post-dehiscent receptacle small, not whitish or whitish-cream.
The species is characterized by its moderate to strongly incised leaves that
are relatively broad along the ae (often cleft to only 1/3 to total width),
by the small corollas, and by distinctive nutlets that have narrow central bod-
ies, with the pericarp greatly expanding at the base, with a conspicuous, spoon-
like appendage nearly | mm high at the tip, with the commissural face nearly
as wide as the mid nutlet, continuing into the distal appendage. Relationships
of the species may also lie with two other species in North america have similar
nutlets with broad commissural faces: Glandularia shrevei (Moldenke) Umber
and possibly Glandularia bajacalifornica (Moldenke) Umber (Umber 1979).
Widespread in southern Texas and closely adjacent Mexico also in west
Texas near the Pecos River (Fig. 3), in limestone, caliche, gravelly to sandy, clay-
ish soils in coastal prairie, thorn scrub, in cis-Pecos Texas in desertic scrub, of -
ten roadsides, open areas, 10 to 1000 m, flowering late February to April.
aaa verecunda Umber, Syst. Bot. 4:99. 1979. (Fig. 1 E-H). Tyr US.A. TEXAS.
REEVES Co. 2.5 mi F of Toyah, R.& J. Umber 379 (HOLOTYPE: GH; ISOTYPES: TEX!, NY)
Verbena pumila t. albida Moldenke, Phytologia 9:188. 1963. Type: U.S.A. TEXAS. KENEDY Co. a
the edge of Kenedy Co., — ay 96 at the Willacy Co. line, 18 Apr 1941, Runyon 2499 (HOLO-
rYPE: NY; ISOTYPE: TEX
Vegetatively similar to G. caine rangulata but the leaves more ovate to obovate
in outline (12-)15-25(-42) mm long, (9.5-)10-20(-25) mm wide, with petioles
4-8 mm long, the lateral clefts more often entire or with fewer lateral teeth-
lobes, and the hairs more often with pustulate bases. Inflorescence bracts lin-
HENRICKSON, STUDIES IN GLANDULARIA 965
ear-lanceolate to lanceolate, 3.2-6.5 mm long, 49-87 percent as long as the ca-
ee 5.5-7.0mm long, the lobes, 0.4-L.1 mm long, in fruit the calyces to
L.7-2.5 mm broad at the base; corolla tubes 5.5-7.0 mm long, 0.5-0.9 mm wide,
the limbs white to lavender, 3.2-4.5(-5.5) mm wide, the 5 lobes all retuse. or the
upper two lobes not retuse, 1.4-18 mm long, 0.8-2.0 mm wide; the distal an-
thers 0.6-0.8 mm long, with filaments 0.5-0.7 mm long, the proximal anthers
0.4-0.6 mm long, with filaments 0.2 mm long; styles 5.0-5.2 mm long. Mature
nutlets (2.6-)2.8-3.3 mm long, lacking a terminal appendage, the central body
0.5-0.6(-0.65) mm wide, the cream-colored commissural face 0.35-0.5 mm wide,
this extending to within 0.05-0.2 of the tip of the fruit, the distal commissural
surface truncate or slightly retuse, the inner tip of the fruit with a short inward
pointing, tapering, horizontal appendage to 0.2 mm long, the commissural face
aculeate with papillae to 0.15 mm long, the basal pericarp expanded, forminga
hollow chamber to 10-13 mm wide, the post-dehiscent receptacle 1.0-1.2 mm
wide, whitish-cream in color.
South Texas and trans-Pecos Texas south into eastern Chihuahua, also east-
ern Coahuila and adjacent Nuevo Leon (Fig. 3) on calcareous and limestone
derived clays, sandy soils, grasslands, thorn scrub, to Larrea, Acacia, Hectia
scrub, also disturbed flats, roadsides, milpas; 5-800 m elevation; flowering,
March through May, also in Fall with rains.
The taxon is vegetatively similar to that of G. quadrangulata has less strong
cleft leaves, slightly larger corollas, anthers, and similar nutlets that, however,
lack the distinctive distal appendage.
Glandularia racemosa (Eggert) Umber, Syst. Bot. 4:99. 1979. (Fig. 2). Verbena
racemosa Eggert, Torreya 2:123. 1902. Typr: U.S.A. TEXAS. EL PASO (now Hudspeth) Co.: low
andy valley, Sierra Blanca, 15 May 1901, H.K.D. Eggert s.n. (HOLOTYPE: NY, ISOTYPES: GH, MO).
Verbena pulchella Greene, Pittonia 5.136. 1903. Typr: U.S.A. TEXAS. JEFF DAVIs Co: Foothills of
the Davis Mountains, Tracy & Earl 106d (HOLOTYPE: ND-G?; ISOTYPES: NY, GH).
Usually thick-stemmed annuals 0.5-2.5 dim tall; stems initially several from
the base, briefly decumbent and erect above, with age more spreading, decum-
bent, ascending distally, 1-3(-7) dm long, and the plants becoming bushy, the
internodes (1.0-)2-4(-6.5) cm long, green, sometimes purple with age, 4-angled,
obscurely grooved on two sides when young, moderately to strongly (below
inflorescence) hirtellous to pilose with slender, tapering erect, mostly straight
hairs 0.2-1.2(-1.4) mm long, sometimes with stipitate glands to 0.2 mm long
below the inflorescence. Leaves site and in axillary clusters, broadly ovate
to ovate-rhombic in outline, - 25(- 45) mm long, (6-)12-22(-26) mm wide,
shorter than the internodes, with petioles 2-6 mm long, the blades 2-3 pinnati-
fid, deeply divided nearly to the midrib, the divisions variously revolute, ob-
long to lanceolate, 1-7 mm long, 0.8-1.5(-3.5) mm wide, with 7-28 ultimate, acute
to obtuse lobes, the blades flat to usually conduplicate with the lobes ascend-
—
966 BRIT.ORG/SIDA 20(3)
ing, with rather stiff, tapering unicellular hairs 0.2-0.5(-0.8) mm long on both
surfaces, these generally dispersed and ascending on both surfaces, the hairs
sometimes with pustulate bases, the veins impressed above, broadened beneath,
the blades green to grayish due to the vestiture. Spikes terminal on all branches,
10-25(-35) mm long, 11-12 mm wide, densely crowded, the inflorscence axis
seldom elongating at the base; peduncles 14-24 mm long, or the spikes closely
subtended by leaves, the spike rachis + pilose with hairs to 1.4 mm long and
often with stipitate glands; bracts ascending, linear-lanceolate, 3.3-6.1 mm long,
to 0.6-1 mm wide, hirsute along margins and midvein with hairs 0.2-1.1 mm
long and with stipitate glands, 51-89 percent as long as the calyx; calyces ini-
tially 5-7 mm long, membranous between the 5 green veins, hirtellous and hir-
sute with hairs 0.2-0.8 mm long, usually with stipitate glands 0.1-0.2 mm long,
the lobes unequal, 0.4-1.4 mm long, green or purple tipped, in fruit the calyces
becoming trunctated and to 1.5 mm broad at the base; corollas slaverform, the
tubes 6-7.5 mm long, expanding to 1.0-L3 mm wide at the throat, the limbs
white or tinged with lavender, (5.5-)6.0-9.0 mm wide, the 5 lobes all retuse, 1.6-
3.5 mm long, 1.7-3.8 mm wide; the tube orifice with white or yellow monili-
form hairs to 0.7 mm long; distal anterior anthers 0.6-0.8 mm long, the anther
gland obscure or extending above the anthers, the filaments 0.5-0.7 mm long,
the proximal posterior anthers 0.5-0.6 mm long, the filaments 0.2-0.3 mm long;
style 5.5-6.2 mm long. Mature nutlets (2.0-)2.2-2.7 mm long, rounded at the tip,
the central body 0.55-0.75 mm wide, the whitish commissural face 0.25-0.3(-
0.35) mm wide, this rounded at the tip and extending to within 0.15-0.4 mm of
the rounded nutlet tip, weakly aculeate with papillae to 0.1 mm long, the basal
pericarp slightly expanded, hollow, 0.6-0.85 mm wide, the outer pericarp black,
shallowly reticulate in the distal half, the reticulate pattern disorganizing into
vertical striae in the lower half, the post-dehiscent receptacle yellowish, 4-
angled, distinct.
Central trans-Pecos Texas (Fig. 3), mostly on limestone derived, calcareous
soils, roadsides, mesquite flats, grasslands, with specimens verified from
Brewster, Pecos, Hudspeth, Terrell and Reeves counties, Texas, flowering March
and April, 2500-4000 ft. elevation. Moldenke 1964b sites additional specimens
from Houston, Montgomery, Glasscock, Presidio, ValVerde, LaSalle counties,
Texas, as well as San Miguel Co., New Mexicoand the Mexican states of Coahuila
and Nuevo Leon. His Montgomery Co., was from Stockton (= Ft. Stockton, Pecos
Co., Tex.) His San Miguel Co., New Mexico was from Pecos Valley (= Pecos Co.,
Tex.) Others were G. verecunda or G. pumila.
The species has more strongly divided leaf blades, with the blade narrowed
to 1/10 of the leaf width at the midvein, and sometimes have a denser vestiture.
They have large corollas with tubes 6-7.5 mm long, and limbs 6.2-9.0 mm wide,
and nutlets are shorter, rounded at the tip, not strongly expanded at the base,
the commissural face is narrower than the nutlet width, and its distal surface is
——
HENRICKSON, STUDIES IN GLANDULARIA 967
rounded, not retuse, and the aculeae on the commissural face are reduced to
only 0.1 mm in total length.
Glandularia racemosa is easily separated from G. quadrangulata and G.
verecunda, as leaves are more deeply divided, often with a denser vestiture, the
corollas limbs are larger, and the nutlets are much shorter, without a broadly
outwardly flaring base, the commissural face is notably narrower than the cen-
tral portion of the nutlet, and the distal portion of the commissural face is
rounded, not retuse and occurs 0.15-0.3 below the rounded fruit tip. Umber (1977,
1979) notes relates this species to the Glandularia bipinnatifida complex, and
reports hybridization between G. racemosa and G. bipinnatifida documented
by mixed flavonoid profiles in the hybrids (Umber 1977).
A separate paper will deal with Turner’s (1999) reclassification of the
Glandularia bipinnatifida complex.
ACKNOWLEDGMENTS
[thank Ray Umber for valuable discussions and a personal loan of a copy of his
dissertation, Bobbi Angell for the illustrations, Jack Neff for discussions on po-
tential pollinators, Fernando Chiang for the Spanish abstract translation, SRSC
for a loan of specimens, and the University of Texas, Plant Resources Center, for
use of facilities.
REFERENCES
Burkart, A. and N.S. Troncoso, 1979. Verbenaceae. In: Flora Ilustrada de Entre Rios (Argen-
tina) A. Burkart, ed. Parte V: pp. 229-294. Buenos Aires.
Crupen, R.W. 1977. Pollen-ovule ratios: a conservative indicator of breeding systems in
flowering plants. Evolution 32:32—-46.
Dermen, H. 1936.Cytological study and hybridization in two sections of Verbena. Cytologia
7:160-175.
Lewis, W.H. and R.L. Ouiver. 1961. Cytogeography and phylogeny of the North American
species of Verbena. Amer. J. Bot. 48:638-642.
Motoenke, H.N. 1964a. Verbena quadrangulata Heller, pp.40—46.In: Materials toward a mono-
graph of the genus Verbena. XXIll. Phytologia 11:1-64.
Mo pene, H.N. 1964b. Verbena racemosa Eggert, pp. 46-50. In: Materials toward a mono-
graph of the genus Verbena. XXIll. Phytologia 11:1-64.
Perry, L.M. 1933. A revision of the North American species of Verbena. Ann. Missouri Bot.
Gard. 20:239-356.
Turner, B.L. 1998. Texas species of Glandularia (Verbenaceae). Lundellia 1:3-16.
Turner, B.L. 1999. A reevaluation of the Mexican species of Glandularia (Verbenaceae)
Lundellia 2:55-71.
Umer, R.E. 1977. Biosystematic studies in the genus Glandularia (Verbenaceae) of North
America. Ph.D. dissertation. Harvard University, Cambridge.
Umer, R.E. 1979. The genus Glandularia (Verbenaceae) in North America. Syst. Bot.4:72-102.
968 BRIT.ORG/SIDA 20(3)
TIMBER PRESS
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Key Words: Dahlia, history, cultivation, propagation, gardening.
PLANTFINDER’S GUIDE
ROGER GROUNDS. 2003. The Plantfinder’s Guide to Ornamental Grasses. (ISBN O-
88192-566-7, pbk.). Timber Press, Inc., 133 5.W. Second Avenue, Suite 450,
Portland, OR 97204-3527, U.S.A. (Orders: www.timbet press.com, 800-327-
5680, 503-227-2878, 503-227-3070 fax). $19.95 (pbk, 2003), 192 pp., 106 color
photos, 5 b/w line drawings, 7 1/2" x 10 1/4".
lable of Contents: Introduction to Grasses, Colour a leal Grasses, Flowering grasses, Orchestrating
the Grasses, Plant Care, Appendices, Inc
ex,and A
MArtTIN RickARD. 2003. The Plantfinder’s Guide to Garden Ferns. (ISBN 0-88192-
507-5, pbk.). Timber Press, Inc., 133 S.W. Second Avenue, Suite 450, Portland,
OR 97204-3527, US.A. (Orders: www.timberpress.com, 800-327-5680, 503-
227-2878, 503-227-3070 fax). $19.95 (pbk, 2003), 192 pp., 121 color photos,
20 b/w line drawings, 2 color maps, 7 1/2" x 10 1/4".
—
Table of Contents: Introduction to Ferns, The Plants, Plant Care, Appendices, Index, and
A | le =
SIDA 20(3): 968. 2003
THE BRACTEOLES IN AMARANTHUS (AMARANTHACEAE):
THEIR MORPHOLOGY, STRUCTURE, FUNCTION, AND
TAXONOMIC SIGNIFICANCE
Mihai Costea'! Francois J. Tardif
Department of Plant Agriculture papaKiaeh It eee Ruegieth ture
University of Guelph University of Guelph
Guelph, Ontario, NIG 2W1, CANADA Guelph, Onans NIG 2W1, CANADA
coste_amihai@hotmail.com ftardif@uoguelph.ca
ABSTRACT
The morphology and anatomy of bracteoles were surveyed in 20 Amaranthus taxa in order to deter-
mine their taxonomic significance and role. Three types of bracteoles were distinguished: spinose, fo-
liaceous and membranous. pe bri acicele® retain ee _ structure of the leaves, but they differ from the
latter by the presence of well | {scl I shaped 1 Kranz bundle- sheath. Pro-
tection and photosynthesis are the two functions of bracteoles inferred from their structure. Based on
their structure, foliaceous and membranous bracteoles are considered symplesiomorphic, while spi-
nose bracteoles are interpreted as synapomorphic. The spines present at the base of each leaf in A.
spinosus L. are metamorphosed bracteoles. The shortness of bracteoles in the grain amaranths (A
caudatus L., A.cruentus L.and A. hypochondriacus L.) was found to bea reliable character, which sepa-
rates them from their wild relatives (A. hvbridusL. and A.powellii S. Watson). The structure of bracteoles
may be significant for recognition of taxa in only a few cases: A. hybridus L.and A. powellii S. Wats: A.
powellii subsp. powellii and subsp. bouchonii (Thell.) Costea & Carretero and A. blitum L. agg
RESUMEN
La mioriologis anatomia de las bracteolas or 20 taxa de Armarantin us mueron examin: ada is para
1 2% 4
{
SU TUNCION
foliaceas y sem branes Las acids retienen la estructura C, de las hoes pero difieren de estas
ultimas por la presencia de un escleré ae Sa desarrollado y en la forma arqueada de la vaina de
los sag ue pee Kranz. Las dos funcione ; olas son la proteccion y la fotosintesis segun
Basandonos en suestructura. ates bri acieolas! ounce, pu cinenanOsas son
aes ] . ] ‘ Es |
mientra que
[
sinapomorficas. Las espinas presentes en la base de cada hoja de A. spinosus L. son bracteolas
metamortoseadas. La cortedad de las bracteolas en los amarantos cultivados como pseudocereales
(A. caudatus L., A.cruentus L. y A. hypochondriacus L.) fue encontrada como un caracter que los separa
de sus parientes silvestres (A. hybr a is ye . iets Wi aon La estructura de las bracteolas
le ser significativa parae
A. hybridus L. y A. powellii
DILUM age.
5,
S. Wats.; A. powellii subsp. powellii y subsp. bouchonii (Tell) C ‘ostea & C arretero y A,
INTRODUCTION
The genus Amaranthus is a group of extremely high economical value. Some
species are grown as vegetables and pseudocereals (reviewed by Brenner et al.
2000), while others are among the most important weeds of the world (Holm et
el
is orresponding author.
SIDA20(3 \:9 969- 985. 2003
970 BRIT.ORG/SIDA 20(3
al. 1997). Additionally, some species are widespread ornamentals and they have
a potential as forage crops and as a source of red food colorants (Cai et al. 1988).
The taxonomical treatment of some species has proven to be problematic (e.g.
Costea et al. 2001; Xu & Sun 2001), and it is proposed that characters of
bracteoles may bring forward additional data useful for the classification of the
species within this genus. Bracts are modified leaves without an axillary bud,
associated with the inflorescence or with individual flowers (McCusker 1999).
Bracteoles are secondary bracts borne singly (usually in monocotyledons) or
in pairs on the pedicel, the calyx or the perianth of a flower (McCusker 1999). In
Amaranthus, bracteoles should not be mistaken with the tepals. The later are
the units of perianth, which is not differentiated in a calyx and a corolla
(McCusker 1999). As Brenan (1961) wrote: “the size and shape of longer bracteoles
in the inflorescence, and particularly their length relative to the flowers they
subtend, are of great importance in distinguishing several species of the genus
Amaranthus.” In the difficult A. hybridus aggregate, the morphology of
bracteoles has been used to differentiate grain amaranths (Amaranthus
caudatus, A. cruentus and A. hypochondriacus) from their wild relatives (A.
hybridus subsp. quitensis, A. hybridus subsp. hybridus and A. powellii subsp.
powellii) (reviewed by Costea et al. 2001). The length of bracteoles has also tra-
ditionally been used to classify the infraspecific variability of some species such
as A. retroflexus (e.g., Thellung 1914). We have proposed to reevaluate the
bracteoles in Amaranthus in order to determine their taxonomic significance.
Amaranthus spp. are C4 plants and, therefore, exhibit the characteristic Kranz
anatomy in the leaves (best described in A. retroflexus by Fischer & Evert 1982)
and cotyledons (Wang et al. 1993). No anatomical study of bracteoles has been
published in Amaranthus, and it would be interesting to compare their anatomy
with that of the leaves. Furthermore, the structure may indicate their function
and could provide new relevant data regarding the evolution of the genus.
Amaranthus spinosus is the only species of the genus that exhibits two
spinesat the base of each leaf,and Thellung (1914) suggested that the two spines
are actually metamorphosed bracteoles. Testing this hypothesis was another
objective of this study.
MATERIALS AND METHODS
Morphology of female bracteoles was examined both on herbarium and fresh
material. Only the bracteoles subtending the flowers during fructification were
taken into consideration. The following herbaria collections were surveyed for
the nine qualitative characteristics (see below): BH, BP BPI, BRIT, CAS, CLA, CM,
DAO, DAV, DS, DUKE, FLAS, HAM, I, IBE, ISC, LA, LIL, MICH, MIL, MIN, MO,
MSC, MT, NA, OAC, OKL, PRH, QFA, RB, RSA, SMU, SOM, TEX, UCR, UMO, US,
UTEP, VAB, VDB, VF and WIS. Additionally, bracteoles were collected from the
wild flora (mainly from Romania and Spain) and from cultivated accessions
971
COSTEA , TAXONOMIC SIGNIFICANCE OF
provided by the USDA and Gatersleben germplasm collections (Table 1). Voucher
specimens are preserved in the BUAG herbarium collection, except for the USDA
accessions, which are preserved in the US collection. For the anatomical study,
twenty bracteoles were collected from each plant, and 15-20 different plants
for each species were examined. Bracteoles were fixed in formalin-acetic acid
alcohol (5:5:90) (FAA) and embedded in paraffin. Transverse sections were made
at 5-7 mm in thickness and stained with Toluidine Blue. The structure of the
bracteoles varies along with their length. The most complete region is usually
at the base (0.2-0.4 mm) of bracteoles, and characters refer to this region. Draw-
ings were prepared with a Reichart camera lucida.
List of bracteole characters examined (See the results section for additional ex-
planations.):
a) qualitative characters of bracteoles (Table 2)
1. Bracteole type:
2. Branching of mid-vein:
+, branched;
- unbranched;
3. Pattern of mesophyll development a the bracteole’s mid-vein:
uniform, mesophyll more or less uniformly distributed along the mid-vein;
-, not union m, mesophyll is more developed in certain regions, usually at the middle or above
4. Color of the mid-vein ( ined on dry material):
g, green (usually dark-green);
y, yellow to yellowish-brown or reddish;
5. Shape of the median part of bracteoles (vascular tissue + sclerenchyma + mesophyll) as seen in
cross-sections (examined only for spinose bracteoles):
cir, semicircular;
el, semi- elli ptic;
lee peel ] =e es ae ee
velns):
6. Spatial pattern of | bundle sheat!
sep, Kranz sheath separately surrounds a branch of the mid-vein;
tog, Kranz sheath encloses all the branches together;
7. Shape of “Kranz cells” as seen in cross-section:
ro, rounded or square;
pa, palisade like;
8. Bundle sheath:
bundle sheath in contact a the cas epigennisa at aah in some pos
a
cot,
co-, additional layers of
9. Mesophyll cells (when present)
pa, palisade-like
1
spo, spongy parenchyma-
h
pla
ei
ike;
: il ] aed
tan, Cangenttally e1ongated,
BRIT.ORG/SIDA 20(3)
Taste 1.Provenance of Amaranthus taxa (Amaranthaceae) examined for anatomy of bracteoles (Ger
many* = accessions from Gatersleben).
Taxa Voucher no. or accession Provenance
Subgenus Acnida (L.) Aellen ex K.R. Robertson
A.palmeri S.Wats.
Subgenus Amaranthus (= section Amaranthus)
A.caudatus L.
A.cruentus L.
A. hypochondriacus L.
A. powellii S. Wats. subsp. powell
A. powellii subsp. bouchonii (Thell.) Costea & Carretero
A. hybridus L.subsp. hybridus
Costea & Carretero
A. hybridus subsp. quitensis (Thell.
A. spinosus L.
A. retroflexus L.
Subgenus Albersia (Kunth) Gren. & Godr. (= Section Blitop
-albus L.
A. blitum L. subsp. blitum
A. blitum subsp. oleraceus (L.) Costea
A.emarginatus Moq. ex Uline & Bray
= A. blitum subsp. emarginatus (Moq. ex Uline & Bray)
Carretero, Munoz Garmendia & Pedrol
A. viridis L.
23522 (a-j)
23050 (a-j)
k—s
(
Ames 2026
P| 16604
P| 490440
23037 (a-})
P| 566896
P| 566897
PI 511919
22769 (a-})
k
sis Dumort.)
21800 (a-j)
(k-s)
Ames 13788
22996 (a-})
(k-s)
Ames 23387
Ames 14964
Romania
Romania
Germany*
*
Romania
Arizona, USA
India
Guatemala
Romania
Spain
Romania
Spain
Romania
Spain
Romania
x
Germany*
Germany”
Romania
pain
Romania
Spain
Canada
Romania
Germany”
Germany”
Bangladesh
Romania
Spain
Brazil
India
Romania
Spain
COSTEA AND TARDIF, TAXONOMIC SIGNIFICANCE OF 973
Tasce 1. continued
Taxa Voucher no.or accession Provenance
A. blitoides S.Wats. 23045 (a-j) Romania
(k-s) Spain
P1553 059 Canada
P| 608 663 USA
A, oe i. 9. & Thev.) N. Terraciano 22225 (a-j) Romania
A. deflex 22228 (a-j) Romania
(k—s) Spain
10. Sclerenchyma (only for bracteoles with branched mid-veins):
com, compact, enc losing all the vascular tissue:
frag, fragmented, separately accompanying each vein:
ll. Base of the
ms, with mesophyl + -sclerenchy ma;
=
m, only with mesoph
s, only with Nene t
: neither mesophyll nor sclerenchyma is on
b) quantitative ¢ hat acters of bracteoles (Table 3
1
1. Length (mm);
2. Maximum width at the base (mm);
3. Length of the spinose apex—only for spinose bracteoles (um):
+. Angle between w me and the median part
Thickness at the base (measured on cross i (
6. re kness of the ee bundle or of the main Pata al bundle in the case of branched mid-
veins (um);
7. Thickness of mesophyll (um);
8. Thickness of bundle sheath cells (um);
9. Ratio: bracteoles length/ perianth length.
RESULTS
In all the species examined, each flower is subtended by two bracteoles (some-
times only one, or up to four) which are more or less modified compared to the
leaves. The bracteoles in Amaranthus can be classified as follows:
a) Spinose bracteoles (Fig. 1, A-F; Fig. 3A).—Are rigid and have a spiny tip.
They have a median part that includes the mid-vein surrounded by scleren-
chyma and mesophyll, and two membranous lateral parts that we shall name
“wings,” due to of their membranous appearance. The sclerenchyma is more
developed than the mesophyll, conferring the hard and rigid appearance of
bracteoles. As seen in cross-sections, the median part can be semicircular (Fig,
IC, E) semi-elliptic (Fig. 1D), deltoid (Fig. 1B, F) or even trilobed (Fig. 2A) (Table
2). The wings are not arranged in the same plane with the median part. They
form a characteristic angle with the median part, which is maintained even in
cross-sections (Table 3). Representative species: Amaranthus powellii (Fig. LE).
974 BRIT.ORG/SIDA 20(3)
Taste 2. Data matrix for qualitative characters in some species of the genus Amaranthus
(Amaranthaceae). See “material and methods” for the numbered characters; na = not aplicable.
Taxa/Character 1 2 3 4 5 6 7 8 9 10 11
A. palmer! 5S + + @Q de tog na na spo com s,m+s
A. caudatus S + YY cir na ro (ae) Spo na m
A.cruentus s 4 y el na ro co spo na om+t+s
A. hypochondriacus S + oy cir na pa co pa na m
A.powellii subst S + oy cir na pa COo- pa na m
powellii
A. powellii subsp. S $+ oy de na ro/pa cot spo na = mts
bouchonil
A. hybridus subsp. 5 + 9g de na ro co- tan no m-+s
hybridus
A. hybridus subsp. S + g de na ro co tan no m-+s
quitensis
A, Spinosus 5 t + dir tog na na spo com s,m+s
A. retroflexus S + y cit/de na ro/pa cot spo na 5,M+S
A. albus s + y de/tri na ro co- pa na m
A. blitum subsp. f/m - g na na ro cO- spo na na
blitum
A. blitum subsp. f + - g na sep ro cO- spo. sep. sna
sleraceus
A. blitum subsp. m = G na na ro cot Na-spo na na
emarginatus
A. viridis f/m - og na na ro co spo na
A. blitoides blitoides f+ - og na sep/tog 1o/pa co- spo com na
A. crispus mM = 9g na na rO cot na-spo na na
A. deflexus m - g na na ro cot Na-spo na na
b) Foliaceous bracteoles (Fig. 2C-F).—Resemble more a small leaf, and they
do not exhibit an obvious differentiation between the median and lateral parts.
The mesophyll is more developed than the sclerenchyma, and as a result, the
bracteoles are mostly green. The wings are narrow. Representative species. A.
blitoides (Fig. 2E-F).
c) Membranous bracteoles (Fig. 2B).—Resemble the foliaceous bracteoles,
but the mesophyll is poorly developed, restricted along the mid-veins, and is
often reduced to the one-layered bundle-sheath. Sclerenchyma is also scarcely
developed. The rest of the bracteole is thin, and membranous (as in the wings
of the spinose bracteoles). Representative species: A. crispus. Transitional forms
between the foliaceous and the membranous types occur in several species (A.
plitum subsp. blitum and A. viridis).
The structure of bracteoles is simple: they have a dorsiventral structure, and
they are composed of epidermis, mesophyll, vascular tissue and sclerenchyma.
The marginal-meristem activity is intense; its cells proliferate as monolayers
~~
975
COSTEA AND TARDIF, TAXONOMIC
=e CO
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ee
Za
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oD as
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B.A. hybridus (incl ssp quitensis) CA hypochondriacus,
F. A. powellii ssp. bouchonii. bs-kranz bundle-sheath, ep-epidermis, m-meso-
D. A. cruentus, E. A. powellii ssp. powellii
i
w-wing. Scale bar = 20 um.
),
phyll, s-sclerenchyma, v-vascular tissue (collateral bundle
976 BRIT.ORG/SIDA 20(3)
Ks
reg D4
a aay gre=.
AMS
tS)
¢
SU Qe
sete y
yy
Fic. 2. Struct f bracteoles. A spinose in {| i lbus, B. membranous in A blitum ssp emarginatus C-F folia-
ceous in: C. A. blitum ssp. blitum, D. A. blitum ssp. oleraceus, E-F. A. blitoides, E i 1 at tl
the base. bs-kranz bundle-sheath, ep-epidermis, m-mesophyll, st-stomata, s-sclerenchyma, v-vascular tissue (collat-
eral bundle), w-wing. Scale bar = 20 tm.
ddle, F. sectioned at
COSTEA TAXONOMIC
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Fic. 3. Struct f spi ind
| | early stage, bracteole-like, B. mature spine. bs-kranz bundle-sheath,
ep-epidermis, m-mesophyll, par-parenchyma, s-sclerenchyma, v-vascular tissue (collateral bundle), w-wing. Scale bar
=20um.
Taste 3. Data matrix for the quantitative characteristics of bracteoles in some Amaranthus taxa (Amaranthaceae). See material and methods for numbered
characters.
Taxa/Character 1 2 3 4 5 6 7 8 9
A. palmeri 554] 14+0.1 25220:5 135-5 320 + 20 300 + 25 22.5425 22.5425 1.3-2.]
A. caudatus 3.254175 047+0.1 0.67 + 0.1 137.5425 215 +30 155+15 55+5 22.5425 12-15
A. cruentus 2.254025 095+0.1 0.65+0.5 135+5 175425 12545 50+ 10 225425 1-1.25
A. hypochondriacus 3.5+0.5 1.2+0.25 140.15 135.9 210+ 10 160 + 10 100 + 20 20415 1.25-1.6
A. powellii subsp. powellii 65215 2.4+0.25 2.5 +05 132.5475 320 + 20 210+ 10 100 + 10 55:5 2-3
A. powellii subsp.bouchonii 4.25+0.75 1.15+0.25 1.2+0.3 12/54 75 175425 12252125 40 + 20 22+25 2-2.2
A. hybridus subsp. hybridus 3.754115 1.15+0.25 1+0.2 95.545 165+15 110+ 10 45+5 2242.5 12-2
A. hybridus subsp. quitensis 3.75+075 1.10+0.23 0.67 +0.5 954+5 165415 110+ 10 45+5 224 25 1.2-2
A. spinosus 3.6+0.9 1.2540.25 1.024027 105+ 15 270+ 220 + 20 71+#11 2142.2 1.5-2
A. retroflexus 5.25+1.25 1.3+40.5 2.174017 135+5 230.5 + 30.5 203 +6 25425 20+ 15 1.1-2.8
A. albus 42+18 1.12 +0. 22 140.2 165+5 240420 16254125 76.544 20+ 15 2-2.5
A. blitum. subsp. blitum 1.03+0.25 0.75+0.15 56+6 3242 325475 12.5+2.5 0.6-0.9
A. blitum subsp. oleraceus 1.25 +0.35 0.8+0.1 - - 80+ 10 375+42.5 4545 16.5 + 1 0.6-0.9
A.blitum subsp.emarginatus 05+0.1 0.45+0.5 Ss 425+75 6+4 12:5 4125 10414 0.4-0.7
A. viridis 0.62 + 0.7 05+0.1 - 45+5 D254 25 25425 lao 4t25 0.6-0.9
A. blitoides 2.15+0.15 1.65+0.34 - 17754125 744128 7645 32.7424 0.8-0.95
A. crispus 1+0.1 0.45+5 - - 4254125 30.5 +11 10+ 10 9+1.1 0.6-0.9
A. deflexus 1.15405 0.8+0.1 - - 52.5+7.5 32.5425 1754175 125425 0.73-0.95
816
(€)02 VdIS/9¥O'LINE
COSTEA AND TARDIF, TAXONOMIC AMARANTH) on
and give rise to the wings. Venation is poor, but the vascular tissue is accompa-
nied by well-developed sclerenchymaa. Particularly the lower epidermis may
have anomocytic stomata, with three to six neighboring cells surrounding the
guard cells. Sparse, multicellular, uniseriate trichomes may occur in all species,
especially toward the base of the bracteoles. The vascular tissue consists of a
single colateral bundle surrounded by sclerenchyma and a bundle sheath. In
some of the species examined—A. spinosus, A. palmeri, A. blitoides and A. blitum
subsp. oleraceus—the mid-vein branches (Fig. 2D-F; Table 2). In all species, the
bundle sheath exhibits the “Kranz syndrome” characteristic of the leaves. The
fundamental difference is that the bundle-sheath is incomplete, with the cells
arranged in an arc around the sclerenchyma. The bundle sheath cells are tightly
packed together, they have thicker walls, and their chloroplasts are larger than
those of the mesophyll cells. In the species with bracteoles possessing a branched
mid-vein, the “Kranz” bundle sheath can separately follow each branch (A.
blitum subsp. oleraceus, Fig. 2D), or it can enclose all the branches together (A.
palmeriand A. spinosus, Fig. 3A). In A. blitoides both situations may be encoun-
tered in the same bracteole: the Kranz eae sheath may be continuous at the
base of the bracteoles (Fig. 2E) and fragmented above their middle (Fig. 2F).
The mesophyll is homogenous and distributed only along the abaxial face. Its
cells, arranged in up to four layers, can be palisade-like (Fig. 1C, FE; Fig. 2A),
spongy parenchymaclike (e.g. Fig. 1D) or, tangentially elongated (e.g. Fig. 1B)
(Table 2). The mesophyll can be uniformly distributed along the mid-vein, or
the mesophyll may develop preponderantly along certain regions of the
bracteoles (usually in the middle and above). In the first case, the mid-vein
morphologically appears “constant,” gradually decreasing along its course to-
wards the apex. In the second case, the mid-vein appears enlarged above the
middle of the bracteoles. The sclerenchyma in spinose bracts is strongly devel-
oped, compact, engulfing the mid-vein and, when present, its secondary
branches (e.g. in A. spinosus and A. palmeri). In foliaceous and membranous
bracts with branched veins, sclerenchyma can be compact or fragmented, sepa-
rately accompanying each branch (Fig 2D) (Table 2). The wings have several
layers of cells at their base and they are one-layered in the rest. The base of the
wings may exhibit only mesophyll cells, only sclerenchyma, mesophyll and
sclerenchyma cells or none one of these (Fig.|; Table 2). The monolayer of wings
has thickened cell walls and may contain calcium oxalate crystals.
Origin of spines in Amaranthus spinosus.—This study supports Thellung’s
(1914) hypothesis. The two spines present at the base of each leaf are the meta-
morphosed bracteoles of the first flower within the first dichazial cyme, whose
development is suppressed. No floral rudiments were observed in the vicinity
of the developing spines which indicates that the suppression is achieved very
early in the ontogeny. In an incipient stage of development, spines are similar
to bracteoles (Fig. 1A). However, in bracteoles, the development of sclerenchyma
ae)
980 BRIT.ORG/SIDA 20(3)
is rapidly completed. The growth of spines involves primarily cell expansion.
clerosis is delayed until the parenchyma cells enlarge up to five times com-
pared to the parenchyma cells of the bracteoles. Only then is the development
of sclerenchyma completed (Fig. 3B). The marginal meristems make at most a
limited contribution to the development of the spines, and consequently the
wings are reduced or entirely absent (Fig. 3A, B). Mesophyll is present at least
towards the base of spines and the vascular supply is similar to that of the
bracteoles (Fig. 3B). Amaranthus spinosus has a form—f. inermis Lautersbach
and Schumann-—that has no spines, or with spines that are bracteole-like, sub-
tending the first dichazial cyme of each leaf. We believe this form can be re-
garded as the ancestral type, from which the spiny plants derived. We observed
a similar tendency of bracteoles’ transformation in some individuals of A.
powellii. However, the degree of sclerenchyma development never attains the
levels observed in A. spinosus, and the process is not accompanied by the devel-
opmental suppression of the first cyme.
WN
DISCUSSION
Taxonomic significance of bracteoles.—Taking into account the foliar origin of
bracteoles, their phylogeny can be speculated upon. The primitive condition in
Amaranthusis represented by the foliaceous and membranous bracteoles, while
the spinose bracteoles can be considered a synapomorphy. Consequently, the
most primitive Amaranthus species are those classified in the subgenus
Albersia, while those from the subgenus Amaranthus are more evolved. This
hypothesis is also supported by the evolution of other characteristics such as
the dehiscence-indehiscence of fruits (Costea et al. in press) and the vascular
system (Costea & DeMason 2001).
Based on the populations and accessions used, only a few of the taxonomi-
cally difficult species with spinose and foliaceous bracteoles could be separated
using anatomical characteristics. Furthermore, such characteristics are even
less significant in the species with membranous bracteoles, due to their extreme
reduction. The anatomic differences between grain amaranths and their wild
relatives are minute (Table 2, 3; Fig. 1), and it is possible that these differences
will be found to be even less significant when more accessions are examined.
There are no differences between A. hybridus subsp hybridus and A. hybridus
subsp. quitensis which are considered different species by some authors (e.g.
Xu& Sun 2001). Inthe A. hybridusagg., Townsend (1988) and Stace (1997) con-
sider the names A. powellii and A. hybridus as taxonomic synonyms. Even if
other morphological characters can easily differentiate these taxa, the distinc-
tiveness of their bracteole anatomy supports their separate recognition (Fig.|
D-E; Table 2, 3). Another taxonomic controversy exists between A. powellii and
A. bouchonii (reviewed by Costea et al. 2001). The morphologic and anatomic
differences between their bracteoles support the recognition of A. bouchoniias
COSTEA TAXONOMIC 981
an infraspecific taxon of A. powellii (Costea et al. 2001) (Fig. 1 E-F; Table 2, 3). In
Amaranthus blitum agg,, the three subspecies—subsp. blitum, subsp. oleraceus
and subsp. emarginatusare dissimilar with respect toanatomy of the bracteoles:
Fig. 2D)
—
1. Mid-vein of (foliaceous) bracteoles branched A. blitum subsp. oleraceus
1. Mid-vein of bracteoles unbranched.
2. Bracteoles foliaceous (rarely membranous). Cells of the Kranz bundle sheath are
separated by the cells of the lower epidermis by a 1-2 layered mesophyll A. blitum
subsp. blitum (Fig. 2C
2. Bracteoles membranous. Cells of the Kranz bundle sheath are in contact with
the cells of the lower epidermis (mesophyll is absent) A. blitum
subsp. emarginatus (Fig. 2B)
The length of bracteoles and the ratio between bracteole and tepal lengths have
been used in the past to classify the infraspecific variability of some Amaranthus
species (e.g. Thellung 1914; Brenan 1961). For example, the individuals with short
bracteoles (equaling the flower perianth) were designated as var. delilei (Rich.
& Lor.) Thell. in A. retroflexus. Such forms are not encountered at the popula-
tion level, presumably because the short bracteole type is selected against. Based
on our current knowledge, such plants cannot even be considered ecophenes,
because the shortness of bracteoles can hardly be associated with any ecologi-
cal advantage. In contrast, the morphology of the bracteoles is one of the few
characters that differentiate the grain amaranths (especially Amaranthus
caudatus and A. cruentus) from their wild relatives A. hybridus (incl. A.
quitensis) and A. powellii. Thus, the spinose bracteoles of grain amaranths are
shorter and thinner compared to those in their wild relatives. This may seem a
contradiction with the previous conclusion, but in this particular case the mor-
phology of the bracteoles is important. Domestication of grain amaranths goes
back in time more than 7000 years (Sauer 1993), and this character—even if
quantitative—is probably genetically fixed. The shorter and softer-spined
bracteoles were selected for under domestication because they facilitate har-
vesting. The survey of many herbarium specimens revealed that cultivated grain
amaranths consistently have short bracteoles. Only some specimens of A.
hypochondriacus may exhibit long bracteoles (comparable with those of A.
powellii), similarly as in A. retroflexus some individuals may have short
bracteoles. The distinctiveness of grain amaranths is also supported by mor-
phological (Costea et al. 2001), anatomical (Costea & DeMason 2001), cytologi-
cal (reviewed by Greizerstein et al. 1997) and molecular data (Xu & Sun 2001).
Bracteoles versus foliage leaves in Amaranthus.—In many plants (e.g.,
Chrysosplenium spp., Euphorbia spp., Melampyrum spp., Costea unpublished)
a continuous morphological and structural transition occurs between foliage
leaves and bracts on the same plant. In amaranths, such a transition does not
exist. Bracteoles in amaranths are sharply differentiated from the leaves. Several
tissues of bracteoles show a reductive tendency. Bracteoles have fewer stomata,
982 BRIT.ORG/SIDA 20(3)
their mesophyll and vascular tissue are less developed, and the Kranz bundle
sheath is incomplete. In addition to this developmental abbreviation, lorma-
tion of new structures without a counterpart in leaves occurs. An example is
the sclerenchyma that is absent in Amaranthus leaves (e.g. Fisher & Evert 1982;
Viana 1993; Esparza-Sandoval et al. 1996), but is an important tissue in the
-meristem activity is intense, giving rise to the one-layered
—
bracteoles. Margina
membranous wings, which are also absent from the leaves.
Based on their structure, Amaranthus spp. leaves belong to a variant of the
Atriplicoid type, having a continuous Kranz sheath around the bundles (reviewed
by Dengler & Nelson 1999). However, in the bracteoles, the organization of the
Kranz cells in arcs is entirely different from this type, and it calls to mind other
C4 leaf structures described in Chenopodiaceae suchas the kochioid and solsoid
types (reviewed by ie & Nelson 1999; Jacobs et al. 2001). Since bracteoles
are ontogenetically abbreviated leaves, one cannot refrain from speculating on
the old idea that‘ amen aerate phylogeny” (See Gould 1977). It is a fact
that Amaranthus shares close morphological, structural (Costea @ DeMason
2001) and molecular affinities with Chenopodiaceae (reviewed by Cuénoud et al.
2002) and these new data provide additional evidence in this respect.
Functions of bracteoles in amaranths.—Two functions can be inferred from
the structure of bracteoles in Amaranthus: photosynthesis and protection. Bracts
(in general) are metamorphosed leaves. Although the degree of modification
varies in different genera and families, the other functions of the bracts are sub-
sequently added tothe primordial role of photosynthesis. When present, bracts
may participate in the development of flowers, fruits and seeds. For example,
the contribution of bracts to the achene maturation in Carpinus spp. (Hori &
Tsuge 1993) or to the development of buds in Gossypium spp. (Zhao & Oosterhuis
1999) was found to be significant. Studies made on Gossypium spp. showed that
the removal of bracts reduced the cotton-boll size and the number of seeds per
boll (e.g. Bangal et al. 1985; Ahmed 1994). Similar results—a lower seed yield—
were reported after removing the phyllaries from the antodia of Carthamnus
tinctorius and Helianthus annuus (e.g. Dhopte & Lall 1981; Hayashi @ Hanada
1986). In Amaranthus, due to the nee number of flowers that develop in the
inflorescences, the photosynthetic role of bracteoles should not be underesti-
mated. A single plant of Amaranthus retroflexus can produce approximately |
000 000 seeds (Hanf 1983). Assuming that the number of female flowers in the
inflorescence is at least equal to the number of produced seeds, this would mean
there are at least 2 million bracteoles in one inflorescence. Therefore, photo-
synthesis in bracteoles may significantly contribute to the development of the
enormous number of small seeds and consequently to the ecological success of
amaranths, especially as weeds. However, this hypothesis needs to be verified
and the contribution of bracteoles to total photosynthesis to be established.
In many plants, protection is the most common function acquired by bracts,
—
ph
—
COSTEA AND TARDIF, TAXONOMIC SIGNIFICANCE OF 983
which is performed in a multitude of ways, against a multitude of factors. Pro-
tection can be limited to the inflorescence and flowers, or it can also be ex-
tended to the developing fruits as well. In Heliconia the liquid produced in the
bracts contributes to the protection against herbivores (Wooton & Sun 1990).
Association with ants as a result of extrafloral nectaries developed on bracts
provides a defense mechanism against a wide variety of predators (e.g. Scott
1981; McLain 1983). In Rheum nobile, an alpine plant, bracts absorb UV radia-
tion and keep the reproductive organs warm (Omori et al. 2000). Mechanical
protection of both flowers and fruits against herbivores is obvious for the spi-
nose bracteoles in Amaranthus. There seems to be a correlation between the
presence of spinose bracteoles and the dehiscence of fruits: species with dehis-
cent fruits have spinose bracteoles, while species with indehiscent fruits have
membranous or foliaceous bracteoles (although a few exceptions exist). One of
these exceptions is A. powellii subsp. bouchonii that has indehiscent fruits and
spinose bracteoles. Yet, this taxon has shorter and thinner bracts compared to
A. powellii subsp. powellii, its closest relative with circumscissile fruits (Costea
et al. 2001). Furthermore, in monoecious and some dioecious species (e.g. A.
palmeri) the bracteoles of the female flowers are more developed than the
bracteoles of the male flowers. The abortive female flowers of hybrids are sub-
tended by longer (than normally) bracteoles (Brenner, unpublished).
In conclusion, based on the populations and accessions examined,
bracteoles are important for the species delimitation in A. hybridus agg. and for
the separation of subspecies in A. blitum. The anatomical study of bracteoles,
along with other characters (the vascular system and the structure of fruits)
indicates that subgenus Albersia has a basal phylogeny within Amaranthus,
while the species of the subgenus Amaranthus are derived. Protection and pho-
tosynthesis are the main functions deduced from the structure of the bracteoles.
ACKNOWLEDGMENTS
We owe thanks to David Brenner, to an anonymous reviewer and to Melissa
Wheeler who reviewed earlier versions of the manuscript providing valuable
comments and suggestions.
REFERENCES
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Ca¥, M.Sun,and H.Corke. 1998.Colorant properties and stability of Amaranthus betacyan
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phylogenetics of Caryophyllales based on nuclear 18S rDNA and plastid rbcL, atpB, and
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relation to its chlorophyll content and dry matter in nine varieties of safflower. Punjabrao
Krishi Vidyapeeth Res. J. 5:6-10.
Esparza-Sandoval, S., G. ALEJANDRE-ITuRBIDE, and Y. HerreRA-Arric ta. 1996. Foliar anatomy and
morphology of seeds in some Mexican species of Amaranthus. Phytologia 81:273-
FiscHer, D.G. and R.F. Evert. 1982. Studies on the leaf of Amaranthus retroflexus L.: morphol-
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Goutp, $.J.1977.Ontogeny and phylogeny. Cambridge University Press.
GreizersTeIN, E.,C.A. Naranio, and L. Poaaio. 1997. Karyological studies in five wild species of
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HayasHi, H. and K. HANaba. 1986. Contribution of leaves and bracts to the seed yield and
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Hom, G.L., J. Dott, E. Hou, J. Hersercer, and J. PancHo. 1997. World weeds. Natural histories
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Jacoes, S.W.L. 2001. Review of leaf anatomy and ultrastructure in the Chenopodiaceae. J.
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McLain, D.K. 1983. Ants, extrafloral nectaries and herbivory on the passion vine, Passiflora
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Omori, Y.H. Takayama and H. Ouea. 2000. Selective light transmittance of translucent bracts
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in the Himalayan giant glasshouse plant Rheum nobile Hook.f. and Thomson
(Polygonaceae). Bot. J. Linn. Soc. 132:19-27.
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BRIT.ORG/SIDA 20(3)
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the various stages ol its life cycle and the damage it does to specific plants. Color photographs lead to
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SIDA 20(3): 986. 2003
NOMENCLATURAL CHANGES IN THE GENUS
POLYGONUM SECTION POLYGONUM (POLYGONACEAE)
Mihai Costea! Francois J. Tardif
Department er eagne ag! culture Department of Plant Agriculture
University of Guelph Un iversity of Guelp
Guelph, Ontario, NIG 2W1, CANADA Guelph, Ontario, NIG 2W1, CANADA
coste_amihai@hotmail.com ftardif@uoguelph.ca
ABSTRACT
Five new combinations are proposed in the genus Polygonum s. str. for the forthcoming treatment of
Flora of North America: P. aviculare subsp. buxiforme, P fowleri subsp. hudsonianum, P humifusum
subsp. caurianum, P. ramosissimum subsp. prolificum and P striatulum var. texense. The taxonomy
of each taxon is briefly discussed and a diagnostic table is provided for the subspecies of Polygonum
aviculare.
RESUMEN
L:
| género Polygonum para el futuro tratamiento de este
seven en la Flora de Norteamérica: P. aviculare subsp. buxiforme, P.fowleri subsp. hudsonianum, P
humifusum subsp. caurianum, P. ramosissimum subsp. prolificum y P. striatulum var. texense. La
t 4 ] qd ]; } 3 ] hI 1 f . Lae } 7
\/
/ } (=) £ if
de Polygonum aviculare
INTRODUCTION
During the preparation of Polygonum s. str. for Flora of North America it was
concluded that several new combinations are required to reflect the new taxo-
nomic knowledge in this genus. The changes involve P. buxiforme Small, P.
hudsonianum (Wolf & McNeill) Hinds, P caurianum Robinson, P. prolificum
(Small) Robinson and P. texense M.C. Johnston.
1. Polygonum aviculare complex
Polygonum aviculare isa taxonomically controversial polyploid complex of pre-
dominantly selfing annuals. Lindman (1912) recognized two species: P.
heterophyllum (= P. aviculare subsp. aviculare) and P. aequale (= P. aviculare
subsp. depressum), while Chrtek (1956) accepted only one species, P aviculare,
with several subspecies. However, European and North American taxonomists
have frequently recognized several distinct species within this complex: P.
aviculare s.str, P. arenastrum Boreau, P. boreale Lange, P. rurivagum Jord. ex
Boreau, P. neglectum Besser and P. buxiforme Small (e.g. Styles 1962; Mertens &
Raven 1965; Scholz 1977; Wolf @ McNeill 1986). The recognition of the former
four species has been mainly based on Styles (1962) who studied British popu-
\Corresnondina author
I
SIDA 20(3): 987-997. 2003
988 BRIT.ORG/SIDA 20(3)
lations. Styles (1962) found these four species to be morphologically discontinu-
ous and with different chromosome numbers: P. aviculare and P. rurivagum were
hexaploids(2n = 60), while P. boreale and P.arenastrum were tetraploids 2n=40).
Studies done in North America initially confirmed these observations and the
recognition of these species relied heavily on these different chromosome num-
bers (e.g. Mertens & Raven 1965; Moore et al. 1970; McNeill 1981). More recent
studies have challenged both the morphological and the cytological criteria.
Multivariate analysis and isoenzyme studies using many populations have
proved that these taxa are not well-differentiated and populations with inter-
mediate characteristics may occur (Meerts et al. 1990). Other chromosome
counts, obtained both in Europe and North America, reported hexaploid popu-
lations among P.arenastrum and tetraploids among P.aviculare (e.g. Gasquez et
al. 1978; Wolf & McNeill 1987; Meerts et al. 1998). Isoenzyme studies showed
that the P aviculare complex has probably an allopolyploid origin (Meerts et al.
1998) and it has evolved as a swarm of inbreeding lines (‘Jordanons”) (Gasquez
et al. 1978). Yurtseva (2001) suggested that the patterns of variation in the peri-
carp surface support the hybrid origin of some species. For example, P.
arenastrum presumably originated as a hybrid between P. aviculare and P.
calcatum (= P.aviculare subsp. depressum in this study). Polygonum neglectum
resulted from the hybridization of P aviculare and P.arenastrum. Although the
members of P. aviculare are predominantly inbreeders, some structures that
make possible cross-pollination in other Polygonaceae are present here as well.
Cleistogamous and chasmogamous flowers, the heterostyly, protandry, and the
capacity to secrete nectar may be regarded as remnants of an ancestral mixed
mating system (Costea and Tardif, unpublished). Bugg et al. (1987), Yurtseva
(1998) reported that flowers of P.aviculare agg. are frequently visited by insects
and thus introgression may occur to some extent. This would explain the com-
plex intermediate variation patterns that occur between its members. In such
conditions, recognition of these taxa at species level is unrealistic and unprac-
tical. The subspecific rank is proposed for P. buxiforme because of the interme-
diates observed mainly with P aviculare subsp. aviculare.
Polygonum aviculare |. Sp. P1359. 1753 subsp. buxiforme (Small) Costea & Tardif,
comb. et stat. nov. (Fig. 1.). BASIONYM: Poly ee buxiforme Small, Bull. Torrey Bot.
Club 33:56. 1906. Polygonum littorale auct., non Link: lare var. littorale auct.,
non Mertens & Koch: TYPE: pene tees LEE ss Os Ee Cumberland Gap, 27 Jul 1892,
forma | here, Katz @ McNeill, in herb 1988, NY 3237541).
any
—
JAK. Small s.n. LECTOTYPE 1
Polygonum buxiforme {. montanum Brenckle, Phytologia 3:363. 1950.
_~
Polygonum aviculare includes at least six subspecies in North America, which
are distinguished only by acombination of characters (Table |). Some extensively
employed characters may be misleading if not examined properly. Heterophylly
has been considered by all authors an important feature that distinguishes for
COSTEA 989
x pe Rippon Sear forme oy suid
eae * of Berne “he of = ie
i one er Sef is t]
S: fates To Me Mer/l ie
¥| |
“a Polygonus bus iferor Sertt |
68
. ¥ YQ Der. Deboroh S$. Kats Boe |
é CUMBERLAND MOUNTAIN FLORA, oe Ww. ee
ps COLLECTED amour oueneay LAND
k we Bea
) a A
aly g tise Ctdliy i le oe reed,
//
AL nTWOr. HOO Frey
Fic. 1. Pol cat h hiyify 5 he| I U.K. Small s.n. NY 323754.)
Taste 1.Comparison between the subspecies of Polygonum aviculare. Mature early season plants with leaves, flowers and achenes are necessary for accu-
rate determinations. Measurements of leaves and ochreas
were made at the m
iddle of mai
066
n stem. Description of perianth refers to fruiting perianth, which
was measured from the joint with the pedicel. Description of achene uses the terminology of Wolf and McNeill (1986).
Polygonum aviculare subspecies
Characters
STEMS; Number
Position of main stems
OCREAE; Size
Appearance
ssp. aviculare
= Pmonspeliense
Pers.
=P heterophyllum
Lindm
1-3
ascending-erect;
rarely procumbent
(5-)6-10(-12) mm
free part soon
dissolved into
fibers
ssp. boreale
(Lange)
Karlsson
= P boreale Lange
i
procumbent-
ascending
3.5-6.5(-8) mm
free part disinte-
grating with
almost no fibrous
remnants
spp. buxiforme
Small)
Costea and Tardif
—3 or numerous
procumbent-
ascending
3.5-6.5(—8) mm free
part hee relatively
persist ith
inconspic
after disintegrating
with or without
fibrous remnants
UOUS veins,
ssp. rurivagum ssp. neglectum
(Jord. ex Boreau) (Besser) Arcang.
Berher
=Prurivagum Jord. =Pneglectum
ex Boreau Besser
= Paviculare ssp.
= Paviculare var. rectum Chrtek
angustissimum
ssp. depressum
(Meissn.) Arcang.
= Paviculare ssp.
microspermum (Jord.
ex Boreau) Berher
= Parenastrum Boreau
= Paequale Lindm.
Meisn = Pcalcatum Lindm.
=P microspermum
Jord. ex Boreau.
= P montereyense
Brenckle
numerous numerous numerous
procumbent- procumbent- procumbent, rarel
ascending ascending; ascending
rarely erect
(6.5-)8-12.5 mm (3.6-)4-8.1mm a) 3-5.5 orb) 8-12 mm
free part with free part with in- a) free part with
strong and brown — conspicuous veins, inconspicuous veins
after disintegrating
with almost no
fibrous remnants
veins, persistent
after disintegrating
after denteratre
leaving almost n
fibrous remnants
(€)0z vals/O¥O'LINa
LEAVES; Shape of
CYMES; Nr. of flowers
elliptic to
lamina oblanceolate
Shape of lamina — acute or obtuse
apex
Length (10-)20-55 (-65)
mm
Width 6-20(-26) mm
Ratio length : width 2-4.5(-5.5)
Ratio length (1.1-)1.5-4
of middle st
leaves and adjacent
branch leaves
Color pure green
1-6.5 mm
(2-)4-8
Distribution of congested at the
cymes on the top stems and
stems branches
Petiole
PERIANTH; Size (2.3-)2.6-4.8(-5)
mm
Ratio length :width 1.8-2.8
spathulate to
obovate
obtuse to rounded
(10-)15-45(-50)
mm
5-20(-26) mm
2-4(-5
pure green
1-7.5mm
(2-)4-8
+ uniformly
(3-)3.3-5.5 mm
1.6-2.8
elliptic to obovate;
narrow elliptic to
lanceolate or
oblanceolate
obtuse to acute
6-30(-45) mm
3-6(-13) mm
2.5-5.6(-10)
2.5(—3.3)
gray-green
0.3-2(-3.5) mm
2-6(-8)
+ uniformly, rarely
congested at the
top stems and
branches
(2-)2.3-3.4(-3.6)
0.9-1.3 (-1.5)
very narrow-elliptic narrow-elliptic to
to linear-lanceolate
acute
(10-
2.5-5(-
a
)15-30 mm
9) mm
)5-13.5(-15)
1.2-2.4(-3.2)
pure green
0.3-2 mm
1-3
+ uniformly; rarely
congested at the
top stems and
branches
(2
—)2.3-3.2 mm
1.6-2.6
oblanceolate
acute
(7.5-)12-35(-45
mm
—6(-10) mm
2-6
(3.5-)4.3-8.5(-10)
1-2
—2.5(-3.3)
pure green
0.5-3(-5) mm
=A(=6)
+ uniformly
(2-)2.2-3.4(-
mm
1.6-2.6
)
3.6)
b) free part persistent,
silvery, overlapping
oward the stem and
branch apices
elliptic to narrow-
elliptic or oblanceolate
obtuse or acute
7.5-30 mm
i a) Cars)
+ uniformly; rarely
congested at the top
stems and branches
—
2-)2.2-3.5(-4) mm
1.5-2.9
VILIUS
166
Taste 1. continued
Polygonum aviculare subspecies
The perianth tube
represents (%) of
perianth length
Outer 3 tepals
ACHENE; Size
Morphology
0-)20-36%
laterally overlap-
-ping;oblong,
cucullate, green
with broad pink,
purple, red or
white borders;
veins dendriticly
branched, strongly
thickened
(2,2-)2.4-3.7 mm
enclosed in the
dark-brown, with 3
subequal, concave,
coarsely striate-
papillose faces;
apex straight
(15-)25-36(-40)%
laterally overlap-
white (sometimes
pink) margins;
veins ee
branched, moder
ately to strongly
tickened
(2.4-)2.7-4 mm
enclosed in the
aa dark-
brown, w
ea concave,
apex straight
20-36(—40)%
laterally overlap-
ping; cucullate,
oblong; the base
with a poach like
protrusion; green
with narrow, usually
20-36(-40)%
not overlapping,
oblong; + cucullate;
green with usually
ink borders; veins
dendriticly
branched, strongly
white borders; veins
dendriticly branched,
moderately to
strongly thickened
(1.8-)2-2.8 (-3) mm
usually enclosed;
dark-brown to light-
brown, with 3
ubequal, concave
to flat faces; coarsely
striate- papillose to
straig ht, sometimes
slightly beaked
tickene
(1.6-)2-2.6 (3) mm
usually exerted
(achene edges are
visible); dark-brown;
with 3 subequal,
concave to flat and
coarsely striate-
papillose faces;
sometimes one
face is narrower;
apex straight
25-45(-50)%
not Overlapping to
sligt Itly 2U,OVIVGIIG,
+ cucullate to flat;
green with usually
pink borders
veins dendriticly
ranched; moder
th to strongly
thickened
(1.6-)1.8-2.8(-3) mm
usually exerted
(achene edges are
isible): dark-brown;
often with one
face narrower; faces
oncave to flat
coarsely straite-
papillose faces,
(rarely only
roughened); apex
straight or a little
bent toward the
narrow face
40-58%
not overlapping;
oblong; obscurely
cucullate to flat;
green with white
margins which may
be reddish-brown in
the innermost part;
veins unbranched,
not thickened
(1.6-)1.8-2.8 mm
I] ton-d |
UsUallY CACTLEG, Gal kK
or light-brown; with
one face narrower or
sometimes biconvex;
faces flat, almost
smooth to coursely
striate-papillose; apex
straight or a little
bent toward the
narrow face
766
(€)0z vals/o¥O'LINa
Chromosome
numbers
Intergrades with:
Origin
Distribution
Ecology
2n = 40, 60
sp. neglectum, ssp
ae ssp.
oreale
Eurasia
frequent across all
of North America
weed in ruderal
places and culti-
vated fields; more
trampling than
ssp. depressum
2n= 40
ssp. aviculare ssp.
buxiforme?
Northwestern
urope, Greenland
Greenland, rare in
Labrador
sea-shores in open
sand, drift banks
and rocky shores
hores
2n=60
ssp. aviculare, ssp
2n = 60
ssp. neglectum, ssp
2n = 40, 60
ssp. rurivagum, ssp
2n = 40, 60
boreale?
North America
frequent across all of
North America
weed in ruderal
habitats or some-
times agrestal;
nondrifting sands,
borders of marshes
and dunes
depressum
Europe
rare in North
America (Costea &
Tardif, in ed.)
dry and sunny
habitats; mostly
ruderal growing
in pavement cracks
and grave
depressum
Eurasia
across most of
North America, but
less frequent than
ssp. depressum
growing in all types
of ruderal and
agrestal habitats
ssp. rurivagum, ssp
I
ctum,
ssp. buxiforme?
common across all of
rica
North A
agrestal and ruderal
in heavily trampled
habitats; sometimes
in turf or grazed
grasslands
VIALS
£66
994 BRIT.ORG/SIDA 20(3)
example subsp. aviculare (= P. aviculare s. str.) from the homophyllous taxa:
subsp. depressum (= P. arenastrum) and subsp. neglectum (= P. neglectum). It was
stated that heterophyllous plants have cauline leaves which are more than three
times as long as the branch leaves, while homophyllous plants have stem and
branch leaves of about the same size (e.g. Wolf & McNeill 1986). The ratio be-
tween the lengths of middle stem and branch leaves (Table 1) shows that using
the above definitions, some plants of subsp. aviculare may be considered
homophyllous, while some of the subsp. depressum and subsp. neglectum may
be characterized as heterophyllous. Heterophylly is not alwaysa clear cut char-
acter and it can be efficiently used by taking into account the mentioned ratios
(Table 1). In other words, keeping in mind that some taxa are “more heterophyl-
lous” than others. Another potential source of error is the morphology of the
tepals. Cucullate tepals are said to be distinctive for North American taxa such
as subsp. buxiforme. In contrast, the non-native taxa (e.g. subsp. aviculare) are
said to possess flat tepals (Mitchell & Dean 1978). This is not true as subsp.
aviculare, subsp. neglectum and often subsp. depressum may also exhibit cu-
cullate tepals in fruit (Table D.
2. Polygonum hudsonianum (Wolf &@ McNeill) Hinds
Wolf and McNeill (1986) described this taxon as a subspecies of P caurianum.
Later, Hinds (1995) observed that subsp. hudsonianum was clearly differenti-
ated from P caurianum and he raised the former to specific rank. However, the
author noted that the “separation of P fowleri and P. hudsonianum is [..] more
difficult” (Hinds 1995). Indeed, intermediate plants may occur between the two
taxa, making their identification problematic at times. Therefore, taking into
account that P caurianum is distinct from both (see below), we propose the
classification of P hudsonianumas a subspecies of P. fowleri:
Polygonum fowleri Robinson subsp. } isoni Wolf & Sen Costea &
Tar if, omb. nov. BA P i I sonianum Wolf
& McNeill, Rhodora 88: 469. 1986. Be byeonmun lsoni (Wolf & an Hinds, Novon
5165. 1995. Type: CANADA. QUEBEC: Fort George, 30 Aug 1950, E. Lepage 12863 (HOLOTYPE:
DAO 539717!; ISOTYPE: QFAD).
3. Polygonum humifusum Merk ex Koch
Hultén (1968) observed that P caurianum Robinson might be “closely related
or identical” to P humifusum Merck ex Koch from Siberia and Kamtschatka.
The BONAP list (1998) explicitly considered the latter name as a nomencla-
tural synonym of the former. However, it must be noted that P. humifusum is
the correct binomial because it has priority. This species has a unique feature:
the lower leaves are often opposite, and Tzvelev (1979) even separated it in a
distinct subsection - Humifusa. The closest North American species is P. fowleri,
which may rarely (especially subsp. hudsonianum) have a few leaves that are
opposite. For differences, see Hinds (1995). The comparison of Russian and North
(NCTEA 995
American specimens of P. humifusum revealed morphological differences,
which together with the disjunct geographical distribution allow the recogni-
tion of two subspecies:
Polygonum humifusum Merk ex Koch, Linnaea 22:205. 1849.
a. subsp. humifusum
Stems and leaves green. Perianth 1.5-2.0 mm, with + erect tepals, green with
white margins, only in some flowers with pink margins. Achene 2.1-2.7 mm, +
rostrate, the apex exserted 0.9-1.3 mm. 2n = 20. (Tzvelev 1979). Siberia and
Kamtschatka.
b. subsp. caurianum (Robinson) Costea & Tardif, comb. et stat. nov. BasionyM:
Polygonum caurianum Robinson Proc. Bost. Soc. Nat. Hist. 31:264.1904. TYPE: U.S.A. ALA
Nushagak, 59° N 158° W, Aug 1882, CL. McKay s.n. (HOLOTYPE: GH 326031).
Stems and leaves reddish or purplish tinged (if leaves are green, at least the
margins are reddish or purplish). Perianth 1.5-2.3(-3) mm, with + outcurved
tepals, green or purple with the margins pink to purple. Achene 1.4-1.6(-2.2)
mm, not or obscurely beaked, with the apex exerted 0.1-0.5 mm. Chromosome
number unknown. NW Alaska, Yukon and NW Territories.
4. Polygonum prolificum (Small) B.L. Robinson
Polygonum prolificum was originally described by Small (1894) as a variety of
P. ramosissimum, and later it was raised to species rank by Robinson (1902).
Mitchell and Dean (1978) mentioned that the specimens of the two taxa “show
too much intergradation to be separated with satisfaction” and we confirm their
findings. The subspecific rank is proposed for P. prolificum.
Polygonum ramossisimum Michx. subsp. prolificum ee Costea & Tardif,
comb. et stat. nov. B rolificum Small, ae
ae Bot. Club 21:171. 1894. Belyoonun prolificum (Small) neeeoe ae dora 4:68. 1
TYPE: U.S.A. NEBRASKA: Exeter, J.H. Wibbe s.n. (HOLOTYPE: NY 323808).
Polygonum prolificum var. autumnale Brenckle, Phytologia 2:404. 1948
Polygonum prolificum var. profusum Brenckle, Phytologia 2:404. 1948.
For differences between the two taxa see Mitchell and Dean (1978) and Wolf
and McNeill (1986). Subsp. prolificum is less variable and less common than
subsp. ramossisimum, growing mostly in wet, saline places, which are scattered
across all of North America.
5. Polygonum texense M.C. Johnston
Recently, P texense has been considered a synonym of P. striatulum (BONAP
1998; Hatch et al. 2002). Although the morphology of the flowers and fruits is
very similar, the two perennial taxa are distinct on the basis of their leaf mor-
phology and their different phenology and ecology. The varietal rank is pro-
posed here for P texense:
996 BRIT.ORG/SIDA 20(3)
Polygonum striatulum Robinson var. texense (M.C. Johnston) Costea & Tardif,
comb. et stat. NOV. BASIONYM: Polygonum texense M.C. Johnston, Southw. Naturalist 14:257,
1969. Type: U.S.A. TEXAS, J. Scudday s.n, (HOLOTYPE: LL. ISOTYPE: LL)
For morphological differences between var. striatulum and var. texense see
Correll and Johnston (1970). Plants of var. striatulum grow in eastern and south-
eastern Texas, while plants of var. texense occur in central-western Texas and
southeastern New Mexico.
ACKNOWLEDGMENTS
The authors thank John McNeill and Guy Nesom for their prompt and most
helpful suggestions on the manuscript. We also thank curators from ACAD,
ALTA, CAS, DAO, DC, DS, GH, F JEPS, LL, MT] MTMG, NFLD, NSPM, NY, PO,
QFA, QUE, RSA, SASK, TEX, UBC, UC and US for preparing the (usually) volu-
minous loans.
REFERENCES
BONAP. 1998. A synonymized checklist of the vascular flora of the United States, Puerto
Rico, and the Virgin Islands. http://www.csdl.tamu.edu/FLORA/b98/check98.htm.
Buaa,R.L., LE. EHter, and L.T.Witson. 1987. Effect of common knotweed (Polygonum aviculare)
on abundance and efficiency of insect predators of crop pests. Hilgardia 55:1-53.
Curtek, J. 1956. Promenlivost druhu Polygonum aviculare L.v CSR. Preslia 28:362—368.
Corrett, D.S. and M.C. JoHnston. 1970. Manual of the vascular plants of Texas. Texas Re-
search Foundation, Renner, Texas.
Gasquez, J., J.P. COMPOINT, G. BARRALIAS, R. CHADOEUF, and J.P LoncHamp. 1978. Essai de taxonomie
d'une espéce adventice annuelle: Polygonum aviculare L. Ann. Amélior. Pl. 28:565—577.
Hatcu, S.L., KN. GANDHI, and L.E. Brown. 2002. A checklist of the vascular plants of Texas.
http://www.csdl.tamu.edu/FLORA/taes/tracy/regeco.html
Hinos, H. 1995. Nomenclatural changes in Polygonum, Persicaria, and Aconogonon
(Polygonaceae). Novon 5:165-166.
Hutten, E. 1968. Flora of Alaska and neighboring territories. Stanford University Press.
Stanford, CA.
Karisson, T. 2000. The genus Polygonum L.|n:Jonsell, B.ed., Flora Nordica vol. 1, Stockholm.
Pp: 255-273.
LinbMAN, C.A.M. 1912.Wie ist die kollektivart Polygonum aviculare zu spalten. Svensk Bot.
Tid. 6:673-696.
McNett, J. 1981. The taxonomy and distribution in eastern Canada of Polygonum
arenastrum (4x = 40) and P monspeliense (6x = 60), introduced members of the P
aviculare complex. Canad. J. Bot. 59:2744—-2751.
Meets, P., J.-P. BRiANe, and C. Lerépre. 1990. A numerical taxonomic study of Polygonum
aviculare complex (Polygonaceae) in Belgium. PI. Syst. Evol. 173:71-89.
Meerts, P,, T. Bava, and C. Leresre. 1998. Allozyme variation in the annual weed species com-
plex Polygonum aviculare (Polygonaceae) in relation to ploidy level and colonizing
ability. Pl. Syst. Evol. 211:239-256.
Mertens, T.R. and PH. Raven. 1965. Taxonomy of Polygonum section Polygonum (Avicularia)
in North America. Madrono. 18:85-92.
MitcHett, R.S. and J.K. Dean. 1978. Polygonaceae (buckwheat family) of New York State. In:
R.S. Mitchell ed.,Contributions to a flora of New York State I.N.Y. State Mus. Bull. No. 431.
Moore, D.M., T.R. Mertens, and J.E. HicHwoop. 1970. Cytotaxonomic notes of the genus
Polygonum, section Polygonum. Proc. Indiana Acad. Sci. 79:396-400.
Rosinson, B.L. 1902. The New England Polygonums of the section Avicularia. Rhodora 40:
65-73.
ScHoiz, H. 1977. Bemerkungen zur Markmalsgeographie des Polygonum aviculare,
insbesondere des Polygonum arenastrum. Verh. Bot. Ver. Prov. Brandemburg 98/100:
180-182
Small, J.K. 1894. New and interesting species of Polygonum. Bull. Torrey Bot. Club 21:
168-173
Sty.es, B.T. 1962. The taxonomy of Polygonum aviculare and its allies in Britain. Watsonia.
5:177-214.
Tzvetev, N.N.1979.On species of Polygonum L., section Polygonum of the European part of
the U.S.S.R.Novitates Systematicae Plantarum Vascularium 1978, 15;Nauka, Leningrad,
Pp.:128-142.
Wotr, S.J.and J. McNeit. 1986. Synopsis and achene morphology of Polygonum section
Polygonum (Polygonaceae) in Canada. Rhodora 88:457-479.
Wotr,S.J.and J. McNett 1987.Cytotaxonomic studies on Polygonum section Polygonum in
eastern Canada and the adjacent United States. Canad. J. Bot.65:647-652.
Yurtseva, O.V. 1998. Self-pollination in species of Polygonum subsection Polygonum. Bull.
Moscow Soc. Nat. 5:61-67.
Yurtseva, O.V. 2001. Ultrastructure of achene surface in Polygonum section Polygonum
(Polygonaceae) in Russia. Nordic J. Bot. 21:513-528.
>
998 BRIT.ORG/SIDA 20(3
BOOK NOTICE
CHARLES B. HEIseR. 2003. Weeds in My Garden: Observations on Some Misunder-
stood Plants. (ISBN 0-88192-562-4, hbk.). Timber Press, Inc., 133 S.W. Sec-
ond Avenue, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders:
www.imberpress.com, 800-327-5680, 503-227-2878, 503-227-3070 fax).
$22.95, 260 pp., 29 color photos, 30 line drawings, 51/2" x 8"
From the dust jacket: “Weeds, by their nature, get around. This book treats about 140 weeds, and more
than half of these may be found throughout most of North America. Many were originally native to
eastern North America, and others first came from Eurasia. The author chooses weeds that have found
their way into “his garden,” the Botany Experimental Field at Indiana University where he has grown
plants for more than half a century.”
Reading paneueys Heiser’s wonderful book on weeds brought back pleasant childhood memo-
ries. For | know adilferent kind of “garden,” my dad’s cotton field in southwestern Okla-
homa. Dad was a cotton farmer and a good one at that. Being born ae cotton harvest time (in the
)in en middle of the 20" century, mother tells me | made my first trip to a cotton field at the
) |
nad my first
fal
a
>
early age of a few days to one week. It was there in those cotton fields back home where
and up close encounters with plants, albeit cotton plants. I didn’t do any collecting (not that | re-
member) except for lots of sleep in the shade of the cotton wagon. | occasionally took a ride down the
cotton rows on the top of mother’s cotton sack. By the time cotton is ready to harvest most of the
weeds have been pretty much eliminated from the “garden” by either a cultivator or by a hoe. As I got
older | discovered weeds through “c hopping cotton” with a hoe, and it didn’t take long to learn that
“chopping cotton,” meant chopping weeds. Dad taught me—early in life—my first lessons in identify-
ing plants (weeds). Positive oe - cotton field ‘ weeds” was of the utmost importance;
dac manes sure of that, oe ‘chopy ping c cotton” was all about vee Eauueapene that is, all of the
“unwante y\ ith those s. What isa weed?
I
For me it was re slat Bue cotton?—Barney Lipscomb, Botanical Researc h Institute of Texas, 509
Pecan Street, Fort Worth, TX 76102-4060, U.S.A. barney@britorg.
SIDA 20(3): 998. 2003
TAXODIOMERIA (TAXODIACEAE), AN INTERGENERIC
HYBRID BETWEEN TAXODIUM AND CRYPTOMERIA
FROM SHANGHAI, PEOPLE’S REPUBLIC OF CHINA
Zhang Jian-Jun!
Shanghai Academy of Agricultural
Sciences,
Shang ai 210006
=
Peop
Zhu Jian-Hua
Shanghai Seed Industry Company
Shanghai 200233
e’s Republic of China
Pan Shi-Hua, Zhu Wei-Jie
Niu Hui-Juan, and Ye Zheng-Ji
Shanghai Forestry Station
anghai 200
People’s Republic of China
Hsu Ping-Sheng'
Institute of Biodiversity
Science, Fudan eon ity
People’s Republic of China Shanghai
People’s Phe ic sie
ABSTRACT
We describe in this paper the hybrid genus x Taxodiomeria ZJ. Ye, J.J. Zhang et S.H. Pan, gen. nov.
(Taxodium Rich. x Cryptomeria D. Don) and the hybrid species xT. peizhongii ZJ. Ye, JJ. Zhang et
H. Pan, sp. nov. (Taxodium mucronatum Ten. x Cryptomeria fortunei Hooibrenk ex Otto et Dietr.)
(Taxodiaceae). The hybrid originated in Nanjing in 1963 and was introduced into Shanghai in 1975; it
has been propagated vegetatively since then. These hybrids, about 2000 in all, were hardly known
but are now big trees and highly valued for their fine shape, faster growth rate, and adaptation to
urban environments compared to Taxodium and Metasequoia. Morphologically, the hybrid species
strongly resembles Taxodium mucronatum, differing slightly in a few vegetative characteristics.
ABSTRACT (CHINESE)
AS SORT ARE IR Taxodiomeria Z.J. Ye, JJ. Zhang et S.H. Pan, gen. nov. (Taxodium Rich. x
Cryptomeria D. Don) FUARACAH T. pedo Z.J. Ye, J.J. ee et S.H. Pan, sp. nov. (Tascoaium
mucronatum Yen. X Cryptomeria fortune’ Hooibrenk ex Otto et Dietr.) (TL; axodiacea e) VES ik.
BARA 1963 AFTER RARE, 1975 #5] A, Seubtr chi semn mt, La Ke 2000
PK. LAL EE AAR, (n> SCRE LEM, REP PE ( Taxodium)
PUIKAZIR (Metasequoia) RP AE REE VT ARE, SEBEL. BARA EIB A E
5 SAPS BEANE (Taxodinm mucronatum) (RAM, (ZETA RHIE ES a Ac »
A new intergeneric hybrid “Taxodiomeria peizhongii ZJ. Ye, JJ. Zhang et S.H.
Pan of Taxodiaceae, described below, arose from a cross between Taxodium
mucronatum Ten.(?) and Cryptomeria fortunei Hooibrenk ex Otto et Dietr.(d).
It is said that a number of individuals of Taxodium mucronatum were intro-
duced into China around 1925, but now only two were found living. One was
on the campus of Dongnan University of Nanjing, the maternal plant of the
hybrid species. Cryptomeria fortunei, a native of southeastern China, is widely
1 i £ q 7
Jian-Jun,zjjun07141 ing
SIDA 20(3): 999-1006. 2003
1000 BRIT.ORG/SIDA 20(3)
cultivated in the area south of the Yangtze River, but the paternal tree of the
hybrids has not been found. The hybrid cross was made in Nanjing in 1963 by
the late Prof. Ye Pei-Zhong, a prominent botanist specializing in the cross-breed-
ing of trees. Finally he was able to produce three cones and a number of hybrid
seedlings, but only five of the 12 F) seedlings survived (Anonymous 1979; Miao
et al. 1982). About 6000 cuttings were prepared in successive years up until 1972.
Some of these were introduced to Shanghai, Hangzhou, Wuhan, and other cit-
ies in China. It seems that the hybrid saplings had not shown any valuable sil-
vicultural characteristics superior to the related species of Taxodium, Metase-
quoid, and Cryptomeria. Therefore, they did not excite much interest among
dendrologists in Nanjing. This is probably why there are only a limited num-
ber of hybrid trees growing there today.
Some 260 hybrid saplings were introduced into Shanghai for the first time
in 1975 under the supervision of professors Cheng Xu-Ke and Jin Guo-Pei. From
1975 to 1983, thousands of young plants were propagated vegetatively from these
saplings in the tree nurseries of Songjiang and Yangjing while Ye Zheng-Ji, the
fifth author of this paper, was in charge of the work of cultivation and propa-
gation of the hybrids in Songjiang. However, for lack of sufficient knowledge as
to the merits and potentialities of the hybrid trees and also for some other rea-
sons, the number of trees gradually decreased; these were hardly known to the
public until the end of the 1990s.
The discovery of the hybrids was the result of a survey for tree resources of
Shanghai carried out by Ye Zheng-Ji and his colleagues from 1998 to 2001. They
have found that the 20-30 year-old F) hy brids—about 2000 in all—have proven
to be very successful in Shanghai compared to Taxodium distichum (L.) Rich.,
T.ascendens Brongn., Metasequoia glyptostroboides Hu et Cheng, and Cryptom-
eria fortunei Hooibrenk ex Otto et Dietr. These hybrids are semi-evergreen, grow
fast, hold up to strong winds, and thrive not only in ordinary garden soil but
also in wetlands and saline sea-shores with a soil pH ranging from 65 to 86.
The trees can grow in saline soil with 0.4% salt. A small plantation of the hy-
brids was seen growing well in a lake in Songjiang. These data are congruent
with the statements that hybrids sometimes respond better than their parents
(Stebbins 1969) and that they may show new characters not present in either
parent (Knobloch 1972). The stand of hybrids in Chuansha Tree Nursery, the
largest one in Shanghai—with about 1500 individuals covering an area of about
L8 ha.—was designated “The Protected Forest of Germplasm Resources of Shang-
hai” by the municipal authorities in 2001. These trees are 30-40 cm. in diam-
eter at breast-height, and the largest is about 62.5 cm. in diameter. The hybrid
species is useful for landscape planting as well as for large-scale windbreaks in
the riverine and coastal regions of Shanghai and neighboring provinces. Since
the climatic and edaphic conditions (the more or less alkaline soil and the high
groundwater level) in Shanghai are not very favorable for growing many kinds
ZHANG ET AL., ANEW HYBRID GENUS x TAXODIOMERIA 1001
of trees, these new hybrids are bound to play an important role in the rapid
development of gardening and urban forestry in the city.
<Taxodiomeria peizhongii Z.). Ye, J.J. Zhang et S.H. Pan, gen. et sp. nov. (Figs. 12
3). ium mucronatum Ten.(2)and Cryptomeria fortunei Hooibrenk
ex Otto et Dietr. (d). Type: CHINA. SHANGHAL Jinshan, Jinshan General Petrochemical
Works, cultivated tree more than 20 m high, diameter breast-height ca. 50 cm, bearing juv
nile male strobili, strips of bark transversely fissured, 25 Sep 2001, SH. Pan 01563A a
TYPE: PE; ISOTYPES: NAS, SHM).
Haec species biter speciel materno ote xodio aaa Ten. valde similis, a i a ae differt
planta ut i g mida, trunco integro
c
5-8 metrorum ae aut aes ramulis
email aetancieacin et pl i Ititudi
primariis diviso raro mee et sralice corticis schedis longis angustisque saepe transversaliter
fissuratis tecto, et surculis mass in n ramulis primariis vel etiam in trunco instructa. Strobili a nobis
non visi praeter Jinsl immaturis 6-12 cm longis
secus ramulos cone. dispositis paniculam pendulam formantibus instructis. Strobili masc.
immaturi unusquisque 6 staminibus praediti.
I
n cultura in solo usitato horti et in terris humidis et etiam in littoribus salsis.
- atype: CHINA. Shanghai: Songjiang ee hi Park ey ated, 21 Sep 2001, S.H. Pan 01540(SHM);
udong, Yan N ultivated d 2 Aug 2001, S.H se cctlgsti dee el
91400 (SH
. Pudong, Cn Tree Nursery, in a stand alone = coast, | Sep 2001, S.H
The specific epithet peizhongii honors Prof. Ye Pei-Zhong of the former Nan-
jing Forestry College, who made this hybrid by crossing Taxodium mucronatum
(2) and Cryptomeria fortunei(d).
This hybrid species resembles the maternal species Taxodium mucronatum
Ten. in the spirally arranged, drooping, annual leafy shoots that drop shortly
before the new shoots leaf out in spring and in having closely arranged, disti-
chous, short (<lcem long) leaves (Fig. 11). The phenomenon of matro-clinal in-
heritance of the hybrids has recently been confirmed by RAPD analysis (Chen
et al. 2002). This report has shown that the genetic distance of the hybrids from
I. mucronatum (0.210-0.246) is much shorter than that of the hybrids from
Cryptomeria fortunei, their paternal species (0.757-0.764). Morphologically, the
hybrid differs from the maternal species chiefly in a few significant characters.
It does not have butswells (knee-like growths) and buttresses (as in the pater-
nal species), the trunk is usually divided at a height of 5-8 m into two or more
primary branches and, consequently, the crown has forms such as conical, cy-
lindric, elipsoid, and pyriform, etc., all of them being more or less round-topped.
Rarely is the trunk straight and undivided. The narrow, long strips of bark are
usually transversely fissured. There are many shoots on the primary branches
or even on the trunk.
Buttresses and butswells are character istic of the genus Taxodium (Harri-
son 1966; Hart & Price 1990), though butswells are only occasionally seen in T.
mucronatum (Hart & Price 1990; Martinez 1950), and their presence depends to
a great extent on the environmental conditions in which the plants grow (Kurz
1002 BRIT.ORG/SIDA 20(3)
Fic. 1. Base of the trunk of x Taxodiomeria peizhongii growing in Chuansha Tree Nursery, Shanghai, showing lack of
buttresses
Fic. 2. Base of the trunk of Jaxodium mucronatum Ten. ( | plant of the hybrid) cultivated in tl pus of Dong
University, Nanjing, showing the buttresses.
Fic 3 op feb L { T, 4; Ly g = D L Lil L g DC TL L L
t
+h kj *
£4
VIATS Papel
ZHANG ET AL., ANEW HYBRID GENUS x TAXODIOMERIA 1003
Fic. 4. Close shot of the bark of x Taxodiomeria peizhongii growing in Chuansha Tree Nursery, Shanghai, showing the
transversely fissured strips.
Fic. 5. Close shot of the bark of = Taxodiomeria peizhongii growing in Chuansha Tree Nursery, Shanghai, showing the
transversely fissured strips.
Fic. 6. Close shot of the lower part of
the divided trunk.
Fic. 7. Individuals of
ye . 4 ee : oc -
Tree Nursery, Shanghai, showing
T, ye * oy oe ’ cov. oe T, AI Ch | Pee ae | . L oe yy 1
1004 BRIT.ORG/SIDA 20(3)
! W th
=n)
Fic. 8. Two individuals of = Taxodiomeria peizhongii growing in Yangjing Tree Nursery, Shanghai,
h 1 th
ul
trunks.
Fig. 9. x ane ipeenongit in nual Mee Nursery, Shanghai.
near Yangjing Tree Nursery, Shangh
UIVIGeU
Fic. 10
Fic. 11. Close shot of a ‘leafy branch and branchlets of = Taxodiomeria peizhongii growing in Sanaa Tree Nursery,
Shanghai
ZHANG ET AL., A NEW HYBRID GENUS x TAXODIOMERIA 1005
i re
};
c. 12. Close shot of branchlets with male strobili of < Taxodiomeria peizhongii taken from the type specimen tree
be in Jinshan, hai
Fic. 13. Holotype of x aa peizhongii.
& Demaree 1934). In the present hybrid species, however, the trunk is integral
and buttresses and butswells have never been seen even in plants growing ei-
ther in wetlands or in seasonally flooded lowlands. Thus, it can be seen that
there is a difference between the hybrid species and Taxodium mucronatum
concerning the presence or absence of butswells or buttresses.
It seems that different tree forms have something todo withage in Taxodium,
from columnar, to more conical, and to flat topped a Li, pers.comm. 2001). The
hybrid trees are of conical habit when young but develop a wide and often flat-
tened crown when old (Harrison 1966; Hart & Price 1990). But no matter how
great the variation of the tree forms may be, the trees maintain their straight
trunks and do not form double multiple leaders as do most of our hybrid trees.
Although the hybrids generally have no strobili, at least now, some indi-
viduals cultivated in Jinshan have been found to bear male catkins 6-12 cm
long assembled into pendulous panicles on the present year’s shoots in autumn
[2002]. These immature male strobili possess six microsporophylls each and
are expected to open in spring 2003.
1006 BRIT.ORG/SIDA 20(3)
KEY TO VEGETATIVE DIFFERENCES BETWEEN
x TAXODIOMERIA PEIZHONGIL AND TAXODIUM MUCRONATUM
Base (trunks) of adult trees in either wetlands or in saline soil without buttresses
(Fig. 1), butswells (knee-like growths) absent; long narrow strips of bark usually
transversely fissured (Figs. 4, 5); tree usually divided at a height of 5-8 m into 2 or
more primary branches (Figs. 6, 7, 8); crown with many different forms, conical, pe
lindric, ellipsoid, pyriform, etc.—but with more or less a rounded top (Figs. 9
inne peizhongii
. Base (trunks) of adult trees always with a swollen and often buttressed bases (Figs.
2, 3), butswells (knee-like growths) frequently present in flooded areas; long nar-
row strips of bark never transversely fissured; tree conical in habit when young, but
developing a wide and often flattened crown when old Taxodium mucronatum
ACKNOWLEDGMENTS
We thank Li Jian-Hua and Peter Del Tredici of the Arnold Arboretum of Har-
vard University for their helpful suggestions and for sending us needed litera-
ture; Verne Grant, Barney L. Lipscomb, and John W. Thieret for offering sugges-
tions for improvement; and Prof. Ni De-Xiang of Fudan University for his
participation in our investigation and helpful comments.
REFERENCES
Anonymous. 1979. Silvicultural techniques for the principal trees in China. Agricultural Press,
Beijing (in Chinese).
Cuen, YP, S.H. Pan, J.J. ZHANG, W.J. ZHU, HJ. Niu, Z.C. Qu, J.W. Wana, D.L. SHEN, and Z.J. Ye. 2002.
RAPD analysis of genetic relationships among the natural populations of hybrid
Taxodium mucronatum Tenore. J. Fudan Univ. (Nat. Sci.) 41(6):641-645
Harrison, 5.G.1966.A handbook of Coniferae and Ginkgoaceae. Edward Arnold Publishers
Ltd., London.
Hart, J.A.and R.A. Price. 1990. The genera of Cupressaceae (including Taxodiaceae) in the
southeastern United States. J. Arnold Arbor. 71:275-322.
KNos.ocn, LM. 1972. Intergeneric hybridization in flowering plants. Taxon 21:97-103.
Kurz, Hand D. Demarer. 1934. Cypress buttresses and knees in relation to water and air.
Ecology 15:36-41.
Martinez, M. 1950. El ahuchuete (Taxodium mucronatum Ten.). Anales Inst. Biol. Mexico
21:25-82.
Miao, M.Z.,Z.Q. ZHANG, Y.C. FANG, and R.R. Ren. 1982. The principal economical trees of Hunan.
Hunan Science and Technology Publishers (in Chinese).
Stessins, Jr., G.L. 1969. The significance of hybridization for plant taxonomy. Taxon 18:
26-35.
A NEW SPECIES OF DIOSCOREA (DIOSCOREACEAE)
FROM THE ANDES OF VENEZUELA
LJ.Dorr Basil Stergios
National Museum of Natural History Universidad Nacional Experimental
Dept. Biology (Botany), MRC—166 de los Llanos Occidentale
Smithsonian Institution, PO, Box 37012 “Ezequiel Zamora” (UNELLEZ)
Washington, DC 20013-7012, U.S.A. Mesa de Cavacas, Guanare
dorr.laurence@nmnh.si.edu Estado Be aie 3323, VENEZUELA
basilven@cantv.net
ABSTRACT
A new saan of Dioscorea, D. es os r& as is described and illustrated. Relationships, dis-
ters, and s are discussed. Information also is provided on the ecol-
ogy meee ge a al distribution of this new species.
Key Worps: Dioscoreaceae, Dioscorea, Venezuela, Andes
RESUMEN
Se describe y se ilustra una nueva especie de Dioscorea, D. lisae Dorr & stergios. Se ase las
relaciones, caracteres particulares y la distribucion del sexo. Ademas, se presenta inform la
ecologia y distribucion geografica de esta especie nueva.
PALABRAS CLAVES: Dioscoreaceae, Dioscorea, Venezuela, Andes
Four species of Dioscorea L. (Dioscoreaceae) have been collected in Guaramacal
National Park, which protects an area of ca. 200 km2 of montane forest, cloud
forest, and paramo near the northern end of the Cordillera de Mérida in
Portuguesa and Trujillo states, Venezuela (Dorr et al. 2000). Two of the species,
D. lehmannii Uline and D. meridensis Kunth, are well-circumscribed although
infrequently collected and with wide ranges in the Andes, extending south to
Ecuador and Peru. When we published a catalogue of the vascular plants of
Guaramacal, we had not yet identified two of the four species, and we listed
them as Dioscorea sp. A and Dioscorea sp. B (Dorr et al. 2000). The latter ap-
pears to be D. coriacea Humb. & Bonpl. ex Willd.,a variable species with a num-
ber of synonyms and an extensive range in the Andes (Colombia and Venezu-
ela to Bolivia). The former, however, cannot be associated with any known species
and is described here as new.
Dioscorea lisae Dorr & Stergios Sp. Nov. ee 1- 2). Type: VENEZUELA. TRUJILLO: Mpio,
Boconé, P oad fro -amacal, SE of Bocon6, N slope
of mountain, 1, 09 13'N, 070°12'W, 2000 m, 13 oe 1995 (3,2, fls), Dorr & Barnett 8057 (HOLO-
).
TYPE: PORT; IsOTYPES: MEXU, US,
atfi i imis differt tereticaulibus licellis | ioril folii
tL Ee I oO
an Piesconca cori
5-7-ne
SIDA 20(3): 1007-1013. 2003
1008 BRIT.ORG/SIDA 20(3)
rat
Fic. 1 ni It Nn 0 Ct gi A Habit (of pl +L 7 ers] tel n
. 1
t i fi ).B.S i fl C. Pistillate fl (A, Dorr et al. 8921, US;
B, Dorr & Barnett 8057, PORT; C, Dorr & Barnett 8057, MEXU).
DORR AND STERGIOS, A NEW SPECIES OF DIOSCOREA FROM VENEZUELA 1009
LH 4)
NY)
NG
4
y
YY }
iom
1cm
tom
ART aR 2003 D
Fic. 2. Dit lisae Dorr & Sterqios. A. F le infl B Capsule C Caps f frui ll | h
tation of seeds and seed wings. D, E. Seeds. (A, Dorr et al. 8057, US; B— a et al. 17034, US).
Herbaceous twining vines; dioecious (or monoecious, with staminate inflores-
cences found on pistillate plants in fruit). Stems delicate, terete, twisting to the
right (dextrorse), glabrous. Leaves alternate, entire. Leaf blades ovate-lanceolate,
7-11 x 3.2-5 cm, 5-7-nerved, the base cordate to truncate, the apex acuminate
to long-acuminate, glabrous, membranous; petioles 1.5-3.5 cm long, glabrous,
faintly glandular-dotted below. Staminate inflorescences paniculate, pendent,
8-13cm long, with 1-4 racemose branches, flowers solitary, the rachis minutely-
papillate, delicate; each flower subtended by a pair of unequal, acicular
bracteoles, ca. 1-1.25 mm long; pedicels 5-8 mm long, minutely-papillate to
1010 BRIT.ORG/SIDA 20(3)
smooth. Staminate flowers greenish-red (Licata & Cuello 217) to purple (van
der Werff & Ortega 6066), rotate at anthesis, the tepals equal, distinct, 1.25-L5 x
0.75 mim, lanceolate, glabrous; stamens 3, borne on a flattened, annular disc,
filaments very short, ca. 0.25 mm tall, thickened, anther cells ovoid, free (or con-
nivent), introrsely dehiscent; staminodia absent. Pistillate inflorescences spi-
cate, pendent, to 14 cm long, solitary; each flower subtended by a pair of un-
equal, acicular bracteoles ca. 1.5-1.75 mm long. Pistillate flowers with tepals
equal, distinct, ca. 1-1.25 « 0.5 mm, lanceolate, glabrous; styles 3, fused at the
base and free towards the apical stigmatic surfaces; staminodia absent. Cap-
sules 2-3 « 1.2-L.4cm, elliptic, the apex and base rounded, flattened, green (dry-
ing light brown) with dark, elliptic maculations, glabrous; seeds 2 per locule,
ca. 0.5 x 0.5cm, irregularly lenticular to ovate-lenticular, compressed, winged
dorsally, the wing to lcm long, membranous.
PARATYPES. VENEZUELA. Trujillo: Mpio. Bocono, Parque Nacional Guaramacal, vertiente norte,
subiendo por la Laguna Negra, 2620 m, Jun 2002 (4 [ls, fr), Cuello 2433 (PORT); Parque Nacional
Guaramacal, road from Boconé to Guaramacal, SE of Boconoé and just above sae sce alee N
slope of mountain, 09° 13'N, 070° 12'W, 5 Mw 1995 (¢ fs), Dorr et al. 7I8ICNY, PC US); Parque Na-
cional Guaramacal, road from Boconé amacal, SE of Bocond, N slope of mountain (09 13'N,
070° 12'W), 29 Oct 1998 (3,2, fls), Dorr et al.8 we PORT, US); Parque Nacional Guaramacal, trail
from El Cafenol (E of Mosquey) to Fila Los Recost: 5, 1790-2200 m, oe Jun 2001 (6,2, fls), Dorr et
al. 8896 (PORT, US), Ibid (dfs, fr), Dorr et al. 8921 (PORT, US); Parque Nacional Guaramacal, en la
vertiente norte, 09° 25'7"N, 070° 1334'"W, 1950 m, 4 Jun 1995 fs), Licata & Cuello 217 (PORT unicate );
Parque Nacional Guaramacal, sector vertiente sur, carretera al caserio Guaramacal, 2000-2750 m,
1997 ({r), Stergios et al. 17034 (PORT, US); entre Bocono y Guaramacal, 2100-2300 m, 4 Sep 1966
é [ls), Steyermark & Rabe 97309 (US, VEN); near Bocono, Paramo de Guaramacal, 09° 14'N, 070°
if W, 2600-3100 m, 22 Nov 1984 (¢ Els), van der Werff & Ortega 6066 (MO, PORT
Distribution and ecology.—Dioscorea lisae is presently known oe from the
Ramal de Guaramacal in Trujillo state, Venezuela. This ridge is a 40 km long
offshoot of the more extensive Cordillera de Mérida, which extends 325 km
lare from the north
el
from Tachira to Lara. All but one of the collections
slope (mostly Qda. Segovia) of the Cerro de Guaramacal:; the sole exception isa
collection from the south slope (Qda. Jirajara) of the same mountain. Dioscorea
lisae has been collected in montane and upper-montane forest; 1790-2750
(-3100) m. Flowering June, July, September, and November; fruiting February.
The US sheet of Steyermark & Rabe 97309 is mislabeled with a locality in
Tachira and the date 27 August 1966. Bono (1996; 526) also reported this collec-
tion (as Dioscorea coriacea) from Tachira, which suggests other duplicates may
be mislabeled. Steyermark’s field notes are unambiguous about the fact that he
and Marvin Rabe collected a Dioscorea in Guaramacal on 4 September 1966
and that the collection was numbered 97309. Since all of the data on the labels
used by Steyermark are printed save for the collection number and determina-
tion, which are typed, it is easy to imagine a typist grabbing one or more labels
from the wrong stack when preparing the distribution of duplicate sheets.
DORR AND STERGIOS, A NEW SPECIES OF DIOSCOREA FROM VENEZUELA 1011
Etymology.—The epithet honors Lisa Ceryle Barnett, botanist, science ad-
ministrator, and wife of LJD. She has participated in many collecting trips to
Guaramacal as well as numerous other expeditions by foot, mule, horse, or jeep
to remote forests and paramos in the Venezuelan Andes.
DISCUSSION
Following the classification of Knuth (1924), Dioscorea lisae belongs in subg.
Helmia (Knuth) Benth, the ovules are attached near the distal (stylar) end of
the ovary and the wing is developed unilaterally toward the proximal end (ie.,
toward the base of the capsule). Several characters, including the presence in
male flowers of a large glandular disc (not always readily apparent) and race-
mose male inflorescence branches with solitary flowers that have 3 anthers and
lack staminodia, suggest a placement in sect. Cycladenium Uline of Knuth’s
(1924) scheme. In Huber’s (1998) classification, D. lisae keys to sect. Trigonobasis
Uline of Mexico and Central America, which he also associates with sect.
Cycladenium.
Dioscorea lisae differs from other species in sect. Cycladenium in that its
male flowers have very long pedicels, which greatly exceed the bracteoles in
length, and its female flowers produce relatively large capsules, which have
rounded (versus acute) apices and bases. As noted above, at least one of the speci-
mens (Steyermark & Rabe 97309) we refer to D. lisae was misidentified earlier
as D.coriacea, but D. lisae differs from that species in a number of morphologi-
cal characters summarized in Table L.
Dioscorea lisae apparently is not strictly dioecious. One of our collections
(Dorr et al. 8921, US) has fruit and male inflorescences and it would appear that
the sexes are separated temporally. In this case the male inflorescences evidently
arise well after the female inflorescences have produced flowers and set fruit.
Condon and Gilbert (1988, 1990) discuss what might be an analogous situation
in the Cucurbitaceae. The genera Gurania (Schltdl.) Cogn. and Psiguria Arn.,
both neotropical vines that were long thought to be dioecious, are actually
monoecious and only temporally dioecious. They documented that these gen-
era have size-related and intra-seasonal sex changes. They also discovered
strongly male-biased sex ratios. Our limited sample size (ten collections) hints
at a male-biased sex ratio in D. lisae: nine collections have male flowers and six
have female flowers and/or fruit. Nonetheless, the exact nature of sex expres-
sion in this species and factors controlling the same are not at all clear and will
not be resolved with herbarium specimens alone.
While Dioscorea is almost universally considered to be dioecious, there are
a number of other reports, usually dismissed, that this is not true of all species.
Burkill (1960: 327) summarized the prevailing view by stating categorically “to
date we do not havea single proved case of the two sexes being fertile on the one
individual: what we have is records of imperfect male organs and imperfect
—
—
1012 BRIT.ORG/SIDA 20(3)
Taste 1. Selected morphological characters distinguishing Dioscorea lisae and D. coriacea.
D.lisae D. coriacea
stems terete angular
leaves 5-7-nerved 7-9-nerved
pedicel length 5-8 mm 2-4mm
capsule size 2-3 X 1.2-14cm 1-2 x 0.5-0.8(-1.2) cm
capsule apex rounded acute
female organs intruded into what may be fertile individuals of the other sex.”
Al-Shehbaz and Schubert (1989: 67) referred to the reports of such deviation as
“abnormally-monoecious”, Huber (1998: 220) cited a “a few reports of monoecy,
most of them referring to American species ...”, and Caddick et al. (2002: 109)
wrote that plants of Dioscorea are “dioecious but vestigial male/female parts
[are] usually present.” Xifreda (1982a, b; 1983; 1984), studying Argentinian spe-
cies of Dioscorea, described D. furcata Griseb., D. hieronymi Uline ex R. Knuth,
D. megalantha Griseb., D. stenopetala Hauman (all sect. Cycladenium), D. de-
mourae Uline ex R. Knuth, D. haumanii Xifreda (both sect. Monadelpha Uline),
and D. pilcomayensis Haumen (sect. Centrostemon Griseb.) as monoecious. She
also observed infraspecific variation in male flowers of several species, notably
D. megalantha; in some collections the glandular disc is not readily apparent or
is incompletely developed, while in others it is fully accrescent. She attributed
this anomaly to collections being made at different stages in floral develop-
ment. Hammel (2000), relying upon his field knowledge of populations of D.
lepida C.V. Morton (sect. Irigonobasis), recently reported that this Central
American species is monoecious and that it flowers precociously with stami-
nate flowers, then produces pistillate flowers toward the apex of an inflores-
cence and even some hermaphroditic flowers. Plants of D. lepida observed ina
later {lowering or fruiting stage consequently show little or no evidence of the
staminate inflorescences. Given the evidence presented here and by Hammel
(2000), it seems clear that careful field observations need to be coupled with
be a wide
—
herbarium, laboratory, and greenhouse studies to clarify what could
range of different floral sex strategies in Dioscorea.
ACKNOWLEDGMENTS
Field work was supported by funds from the Biological Survey and Inventories
Program of the National Museum of Natural History, and herbarium work by
the Andrew W. Mellon Foundation. We are indebted to the staff of Guaramacal
National Park for generous assistance in the field; the Ministerio del Ambiente
and INPARQUES for collecting permits; the curators of MO, NY, and VEN for
the use of their collections; the staff of the Missouri Botanical Garden Library
for photocopies of the field books of J.A. Steyermark; and Barry Hammel (MO),
DORR AND STERGIOS, A NEW SPECIES OF DIOSCOREA FROM VENEZUELA 1013
Lauren Raz(NY),andan mous reviewer for critical comments. The illus-
trations were prepared by Alice Tangerini.
REFERENCES
AL-SHEHBAZ, LA. and B.G. ScHuserT. 1989. The Dioscoreaceae in the southeastern United
States. J. Arnold Arbor. 70:57-95.
Bono, G. 1996. Flora y vegetacion del Estado Tachira, Venezuela. Museo Regionale di Scienze
Naturali (Torino), Monografie 20:1-951.
Burkitt, LH. 1960. The organography and the evolution of Dioscoreaceae, the family of the
yams. J. Linn. Soc. Bot. 56:319-412
Capoick, L.R., P. Witkin, PJ. Rubatt, LAJ. Heoperson, and M.W. Crase. 2002. Yams reclassified: A
recircumscription of Dioscoreaceae and Dioscoreales. Taxon 51:103-114.
Conpon, M.A.and L.E.Giiperr. 1988. Sex expression of Gurania and Psiguria (Cucurbitaceae):
Neotropical vines that change sex. Amer. J. Bot. 75:875-884.
Condon, M.A.and L.E. Gitgert. 1990. Reproductive biology and natural history of neotropical
vines Gurania and Psiguria.In:D.M. Bates, R.W. Robinson, and C. Jeffrey, eds. Biology and
chemistry of the Cucurbitaceae. Cornell University Press, Ithaca, New York. Pp.
150-166.
Dorr, LJ., B. Stercios, A.R. Smith, and N.L. Curito A. 2000 [2001]. Catalogue of the vascular
plants of Guaramacal National Park, Portuguesa and Trujillo states, Venezuela. Contrib.
U.S. Natl. Herb. 40:1-155.
HAMMEL, B.E. 2000. A new species of Dioscorea from Costa Rica. Novon 10:378-381.
Huser, H. 1998. Dioscoreaceae. In: K. Kubitzki, ed. The families and genera of vascular plants
3:216-235,
KNUTH, R. 1924. Dioscoreaceae. In: A. Engler, ed. Das Pflanzenr. IV, 43 (Heft 87):1-387.
XIFREDA, C.C. 1982a. Estudios en Dioscoreaceae I. Bol. Soc. Argent. Bot. 20:31 7-327.
XIFREDA, C.C. 1982b. Estudios en Dioscoreaceae II. Bol. Soc. Argent. Bot. 21:273-298.
XIFREDA, C.C. 1983. Estudios en Dioscoreaceae. Ill. Bol. Soc. Argent. Bot. 22:337-344.
XiFREDA, C.C. 1984. Estudios en Dioscoreaceae. lV. Kurtziana 17:119-123.
1014 BRIT.ORG/SIDA 20(3)
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SIDA 20(3): 1014. 2003
FESTUCA DENTIFLORA (POACEAE: LOLIINAE: SECT.
GLABRICARPAE), A NEW SPECIES FROM PERU AND
TAXONOMIC STATUS OF F. PRESLIANA
Daniel Stancik Paul M. Peterson
Department of Botany, Faculty of Science Department of Systematic Biology - Botany
Charles University of Prague National Museum of Natural History, MRC-166
Benatska 2, 128 01 Prague Smithsonian Institution
ZECH REPUBLIC Washington, DC 20013-7012, U.S.A.
dan_stancik@yahoo.com peterson.paul@nmnh.si.edu
ABSTRACT
A new species, Festuca dentiflora E.B. Alexeev ex Stancik & P.M. Peterson from the Andes of Peru, is
described and illustrated. The new species is a member of Festuca subg. Subulatae SlevEre v) EB.
Alexeev sect. Glabricarpae E.B. Alexeev and is apparently closely related to Festuca calc 1th)
Kunth from Ecuador and Colombia. A key to the species of Festuca sect. Glabricarpae is saucluded In
addition, the taxonomic status and geographical origin of F presliana is discussed.
RESUMEN
Se describe e ilustra una especie nueva de Festuca de los Andes de Pert: F. dentiflora E.B. Alexeev ex
Stancik & PM. Peterson. La nueva especie pertenece a Festuca Ube subuldiae (Tzvelev) E.B. Alexeev
slabric EB la especie
caldasi1 (Kunth) Kunth de ecuaday y Co
Glabricarpae. Adem
ombia. Se Hee tambien una clave Dae la
y origen geografico de presliana.
The Russian botanist, E.B. Alexeev, recognized several specimens with unique
features while studying South American material of Festuca in the United States
National Herbarium (US). Alexeev marked many sp species with
herbarium names, however, some of these were never published. The status of
some of his proposed new species is problematic since, in many cases, only type
material exists, and species concepts in Festuca are highly variable. In this pa-
per, we describe a new grass species from Peru based upon a specimen that was
selected and given an herbarium name by E.B. Alexeev.
The new species, Festuca dentiflora E.B. Alexeev ex Stancik & P.M. Peter-
son, appears to belong in Festuca subg. Subulatae (Tzvelev) E.B. Alexeev, sect.
Glabricarpae E.B. Alexeey, in the subtribe Loliinae, tribe Poeae, and subfamily
Pooideae (Soreng et al. 2000). Section Glabricarpae is one of four sections (sect.
Elmera EB. Alexeey, sect. Glabricarpae, sect. Longiglumes S.L. Lu, and sect.
Subulatae E.B. Alexeev) in subg. Subulatae (Tzvelev) E.B. Alexeev. Section
Glabricarpae differs from sect. Elmera by having flat leaf blades without promi-
nent ribs, and differs from most species of sect. Subulatae by having ovate spike-
—_—
ee
SIDA 20(3): 1015-1022. 2003
1016 BRIT.ORG/SIDA 20(3)
lets that are significantly longer (> 11 mm long), scabrous lemmas (usually gla-
brous in sect. Subulatae), significantly longer anthers (3.5-5 mm long), and gla-
brous ovary apices. While the distribution of species in sect. Elmera is restricted
to the southwestern North America, sect. Longiglumes is known only from
China and sect. Subulateae is sub-cosmopolitan in its distribution (Lu 1992;
Stancik & Peterson 2002). By inclusion of the new species, sect. Glabriformae
consists of five species, two (F. brevighumis Swallen, F chiriquensis Swallen) re-
stricted to Mesoamerica and three CF caldasii, F dentiflora, F woodii Stancik)
found in northern South America (Alexeev 1980, 1982, 1986; Stancik 2001, 2003).
Based on morphological comparisons, F. dentiflora appears closely related to F
caldasii (Kunth) Kunth.
a aan E.B. Alexeev ex Stancik & P.M. Peterson, sp. nov. (Fig. 1). Tyr:
). DEPARTAMENTO HUANUCO. Provincia Banos: U.S. Exploring Expedition of Capt. Wilkes
5 (HOLOTYPE: US-1006459)),
Haec species a Festuca caldasii Kunth) Kunth spiculis l-12 (non 15-17) mm long., glumis majoribus
(inferioribus 4.5-6 non 3.5-4 nlo ong., peepee 8 non 45-6.5mm long.), | lemmate minoribus
(8.5-10 non 10-14 mm on aristis maj s (6-8 non 1-3 mm long.) differt.
Perennial, loosely tufted. Culms about 100 cm tall, erect, glabrous, with 2-
nodes in basal half. Sheaths membranous, glabrous, margins free, innovations
extravaginal; auricles absent. Ligules 3.5-4 mm long, membranous, acute, lac-
erate. Blades (only a fragment seen) 3-4 mm wide, flat, green, glabrous. Panicles
about 20 cm long, 10-15 cm wide, ovate; branches erect or sometimes spread-
ing, in pairs at each node, scabrous. Spikelets 11-12 mm long, ovate; florets 4-5
(perfect); rachillas 1.2-1.8 mm long, densely pilose. Glumes lanceolate, brown-
ish-green with membranous margins; lower glume 4.5-6 mm long, |-veined;
upper glume 7-8 mm long, 3-veined. Lowermost lemmas 8.5-10 mm long, lan-
ceolate, coriaceous-membranous, green, hairy, 5-veined; apex bi-dentate with
an awn 6-8 mm long; callus glabrous. Paleas almost as long as lemma, mem-
branous, two-keeled, scabrous; apex two-dentate, hairy. Lodicules oblong-lan-
ceolate. Stamens 3; anthers 4.5-5 mm long. Ovary apex glabrous. Caryopses not
observed.
Transversal section of leaf blade.—Leaf section with many vascular bundles,
without ribs; bulliform cells not observed; sclerenchyma under both abaxial
and adaxial epidermis discontinuous, without hairs.
Distribution.—Known only from the type locality in Peru.
Etymology.—The specific epithet, as marked on the specimen by E.B. Alexeey,
probably refers to the bi-dentate lemma and palea.
—_—
—
peel
DISCUSSION
Section Glabricarpae is quite homogenous and all five species (FE breviglumis,
F-caldasii, Echiriquensis, F dentiflora, F woodii) are morphologically very simi-
STANCIK AND PETERSON, A NEW SPECIES OF FESTUCA FROM PERU 1017
Fic. 1. Festuca dentiflora (Wilkes 5). A. Culm (portion of). B. Sheath, ligule, and portion of the blade. C. Inflorescence. D.
Spikelet. E. Floret. F. Lower glume, dorsal view. G. Upper glume, dorsal view. H. Lemma, dorsal view. |. Palea, dorsal view.
J. Palea, ventral view. K. Palea, lodicules, stamens, and pistil. L. Lodicules (2), M. Stamens and pistil.
1018 BRIT.ORG/SIDA 20(3)
lar (Table 1). In comparison with other members of this section, F. dentiflora
and F woodii have shorter spikelets (11-14 mm long verses 15-20 mm long), F.
dentiflora and F.caldasii have longer ligules (2.5-4 mm long verses 0.5-1.1.mm
long), and F. dentiflora and FE. breviglumis have longer awns (6.5-15 mm long
verses 0.5-4 mm long).
KEY TO THE SPECIES OF FESTUCA SECT. GLABRICARPAE IN SOUTH AMERICA
|. Spikelets 10-14 mm long, lemmas 10-16 mm long.
Lower glumes 7-8.5 mm long, upper glumes 9-12 mm long Festuca chiriquensis
2. Lower glumes 3.5—5(-6.5) mm long, upper glumes 4.5-8.5 mm long.
3. Ligules 0.5-0.8 mm long, lemma awn 7-15 mm long ___ Festuca breviglumis
3. Ligules 0.5-3 mm long, lemma awn 1-3 mm long Festuca caldasii
1. Spikelets 15-20 mm long, lemmas 8.5-9.5 mm long.
4. Ligules 0.5 mm long, lemma awn 0.5-1 mm long, anthers 3.5-3.7 mm long
Festuca woodii
4. Ligules 3.5-4.5 mm long, lemma awn 0.5-1 mm long, anthers 3.5-3.7 mm long
Festuca dentiflora
Alexeev (1986) also included F. presliana Hitche. in sect. Glabricarpae, but
the status of this species is not clear (see Table Ll). The type specimen of F:
presliana differs from the other members of sect. Glabricarpae by having 3-
veined, membranous, and papillate or short scabrous (not long scabrous) lem-
mas. Since F presliana has lanceolate lemmas with long awns and short an-
thers it may be more correctly placed in Festuca subg. Subulatae sect. Subulatae.
The type specimen of F. presliana was originally described as Bromus
depauperatus J. Presl [Reliq. Haenk. 1(4-5): 263.1830] from a specimen (Haenke
s.n.) collected by the Czech botanist, Thaddaeus Peregrinus Xaverius Haenke,
as a member of Malaspina Expedition (1789-1793). Haenke collected about
15,000 plant specimens on this expedition. However, a large part of this mate-
rial was destroyed during transportation (Stern 1973) and only a small part of
the collection reached the European continent and was finally acquired by
Kaspar von Sternberg in 1821 for the National Museum in Prague. Under the
leadership of K.B. Presl, processing and determination of this material and re-
sults were published in Reliquiae Haenkeanae (Presl 1825-30, 1831-35). New
grass species were described by J.S. Presl. A major part of this collection has
recently been deposited in the National Museum Herbarium in Prague (PR),
and duplicate sets can be found in the Herbarium of Charles University in Pra-
gue (PRC) and in other herbaria (k, MO, US, W).
A taxonomic revision of Festuca in Colombia has recently been completed
by the senior author (Stancik 2003), and the status of F presliana is still un-
clear. While Presl described the type specimen as Bromus depauperatus and
mentioned “sinu Nootka” (=Vancouver Island) as the type locality, Hitchcock
(1927) in his treatment of South American grasses, transferred this taxon to the
genus Festuca and created a new name F. presliana Hitchcock to replacing the
—
Taste 1. Morphological comparison of the South American species of Festuca sect. Glabricarpae and F. presliana.
Taxon F. caldasii F. woodi F. breviglumis F. chiriquensis F. dentiflom F. venezuelana F. presliana
(in press) (in press)
Herbarium PRC COL US Taken from US US PR
description in
Davidse (1994)
Collector S. Laegaard JR1 Wood R. McVaugh C. Wilkes E.P Killip T.PX. Haenke
20405 5254 24827 D: 19719 a
Geographical distribution = Colombia, Colombia Mesoameérica Mesoamérica Peru Colombia, not clear
Ecuador (Mexico- (Mexico Venezuela (see discussion)
Panama) Panama)
Culm height (cm) 70-150 80 55-130 80-130 100 120 70
Sheath base slightly fibrous — fibrous slightly fibrous — slightly fibrous? fibrous fibrous
Ligule size (mm) 2.5-3 0.5 0.5-0.8 5-11 3.5-4 - 5-0.€
Ligule apex acute truncate ? ? acute truncate truncate
Blade length (cm) 20-35 50-60 19-35 15-50 ?
Blade width (mm) 3-9 2.5-5 5-8 4-7 35 : 5.5-6.5
Panicle size, 13-17x5-8 15x 20 15 x 17-22 10-15 x 15-30 20 x 10-15 20 x 17 17 x 10
length x width, (cm)
Spikelet length (mm) 15-17 12-14 15-20 15-20 11-12 12 10-12
Number of florets 5-7 4-5 4-5 ? 4-5 5 2 (not complete)
Lower glume length (mm) 3.5-4(5) 45-55 3-5(6.5) 7-8.5 45 5-6 -
Upper glumelength(mm) = 4.5-6.5 6-7.5 5-8.5 9-12 7-8 6-7 5.5-6.5
Lemma length (mm) 10-14 8.5-9 oat) 4-16 5-9. 8-9 9.5-10
emma apex two-dentate two-dentate two-dentate two-dentate two-dentate two dentate two-dentate
Number of lemma veins S 5 5 3
Awn length (mm) 1-3 0.5-1 7-15 1-4 6.5-8 6-9 8
Anther length (mm) 5-3. 2.5-5.2 3-45 45 1.7-2 25
Ovary apex glabrous glabrous glabrous glabrous glabrous not seen glabrous
Nad WOUd VINLSI4 40 S31D9dS MAN V ‘NOSHILId NV HIDNVLS
6LOL
1020 BRIT.ORG/SIDA 20(3)
blocking name, F depauperata Bertol (FI. Ital.l: 620.1834). Hitchcock also dis-
cussed the geographical origin of this specimen comparing Haenke’s collec-
tion with the specimen of Wilkes Expedition (type specimen described here as
F dentiflora). Hitchcock concluded that the type locality published by Pres!
was in error because neither the specimens nor the description corresponds
with any North-American species of Festuca. Hitchcock, suspected the “true
identity” of the F presliana type to be from Banos, Peru. This opinion, at first,
appeared highly probable because a large portion Haenke’s specimens come
from Huanuco region of central Peru (Fig. lb in Kthnel 1960). In the Flora of
Peru, Macbride (1936) also accepted Hitchcock’s opinion.
Alexeev (1986) opened a new discussion on the geographical origin
Haenke’s specimen and compared the northern South American collections of
Festuca with a specimen of Killip [Killip & Smith 19719 (US)| from the Cordil-
lera Oriental of Colombia. Alexeev concluded that Hitchcock's opinion of the
Peruvian origin was erroneous and that Festuca presliana is a species endemic
to Colombia.
There are no doubts about the placement of Haenke’s specimen in the ge-
nus Festuca since it has an open sheath, glabrous ovary, and a caryopsis that
does not adhere to the lemma. A comparison of the type specimens of Haenke
sn.CF presliana) and Wilkes 5(F dentiflora) with the Killip & Smith specimen,
suggests that Peruvian and Colombian origin of F presliana are in error. The
specimens do not represent the same species (see Table | for comparison). The
Peruvian specimen (Wilkes 5) is described here as a new species (F dentiflora),
and the Killip & Smith specimen represents a new species (FE venezuelana) from
Venezuela (Staneik 2003). In addition to the specimen comparisons above, a re-
construction of Haenke’s itinerary in South America indicates (Fig. la & b in
Kitihnel 1960) that he never reached the Cordillera Oriental of Colombia. A re-
examination of Haenke’s specimen using Hitchcock and Chase (1951),
Soderstrom and Beaman (1968), Stewart and Hebda (2000), Serna and Lopez-
Ferrari (2000), Beetle 1991), Macbride (1936), Pinto-Escobar (1981), and Davidse
(1994) also excluded the possibility that it is of American origin.
The most important morphological feature distinguishing F presliana is
he 3-veined lemma, that is commonly 5-15-veined in Bromus and usually 5-
veined in Festuca. Interestingly, there are only five known species of Festuca
with 3-veined lemmas. In addition to FE presliana, Clayton and Renvoize (1986)
mentioned the following four species with 3-veined lemmas: F africana (Hack.)
Clayton and EF engleri Pilg. From Africa, F altissima Boiss. from Eurasia, and F-
asthenica Hook.f. from Asia. All of these species come from the Old World. The
two African species are placed in the wide-spread, Festuca ape Subulatae
(Stancik in prep.), whereas the Eurasian/ Asian species are alligned with Festuca
Subg. Drymanthele (Aleexev 1977). The occurrence of taxa with 3-veined lem-
——t
co
—
—
—
STANCIK AND PETERSON, A NEW SPECIES OF FESTUCA FROM PERU 1021
mas in the Old World is remarkable. However, use of this observation as sup-
port for the extra-American origin of this Haenke type is probably insufficient.
Other cases of confusion in the location of Haenke’s collections are known, for
example, Alston (1934), Lamson-Scribner (1899), Merrill (1906), and Veldkamp
et al. (1991). Frequently there is confusion with reports of South American spe-
cies as occurring in Luzon (Philippines) and vice versa. Revision of Bromus and
Festuca in the Philippines is needed and could clear up some of this confusion.
ACKNOWLEDGMENTS
We wish to thank the Smithsonian Institutions, Fellowships and Grants for
supporting a research visit by the senior author to the United States National
Herbarium and curators from the following herbaria: AAU, B, COL, LPB, MA,
PR, PRC, QCA, QCNA, VEN, and W Financial aide to visit herbarium collec-
tions was made possible by support from GACR (No.206/01/0687) and a grant
from the Ministry of Education of Czech Republic (No.313004). Appreciation
is extended to Alice R. Tangerini for providing the illustrations; Dan Nicolson
for correcting the Latin diagnoses; and to Kelly W. Allred, Barney Lipscomb,
and Stephan L. Hatch for reviewing the manuscript.
REPERENCES
Acextev, E.B.1977.To the systematics of Aisian Fescues ( Festuca).|. subgenera Drymanthele,
Subulatae, Schedonorus, Leucopoa. Bjull. Mock Izpit. Prirod. 82:95-102.
Avexeev, E.B. 1980. Festuca L.subgenera et sectiones novae ex America et Mexica. Novosti
Sist. Vyss. Rast.17:42—53.
Atexeev, E.B.1982.A new section and three new species of the genus Festuca from Mexico
and Central America. Bot. Zhurn. (Moscow & Leningrad) 67:1289-1292.
Atexeev, E.B. 1986. Festuca L. (Poaceae) in Venezuela, Colombia et Ecuador. Novosti Sist.
Vysé. Rast. 23:5-23.
AtsTON, A.H.G.1934.Notes on Selaginella.VI.The Selaginellae collected by Thaddeus Haenke
and described by K.B. Presi. J. Bot. (London) 72:223-230.
BeetLe, A.A. 1991. Festuca L.In: Beetle, A.A.,ed. Las Gramineas de Mexico, Vol.Ill. Secretaria de
Agricultura y Recursos Hidraulicos: COTECOCA, México. Pp. 124-144.
CtayTon, W.D.and Renvoize, S.A. 1986.Genera Graminum. Grasses of the world. Her Majesty's
Stationery Office, London P. 93-94.
Daviose, G. 1994. Festuca L. In: G. Daviose, M. Sousa S., A.O. Cuarer, eds. Flora Mesoamericana
Vol.6.Alismataceae a Cyperaceae. Universidad Nacional Autonoma de México, México,
DiEPO.wes=227,
HitcHcock, A.S. 1927. The grasses of Ecuador, Peru, and Bolivia. Contr. U.S. Natl. Herb. 24:
291-556.
HitcHcock, A.S. and A. CHase. 1951. Manual of grasses of the United States, Second Edition.
USDA Misc. Publ. 200:1-1051.
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=
KUHNeL, J. 1960. Thaddaeus Haenke, Leben und Wirken eines Forschers. Munchen
LamsON-ScRrIBNER, F. 1899. Notes on the grasses in the Bernhardi Herbarium, collected by
Thaddeus Haenke, and described by J.S. Presl. Report (Annual) Missouri Bot. Garden
10:35-59.
Lu, L.S. 1992. New Taxa of Festuca L. from China. Acta Phytotax. Sinica 30:529-540.
Maceribe, J.F. 1936. Gramineae, Flora of Peru. Field Mus. Nat. Hist. Bot. Ser. 13:96-261.
Merritt, E.D. 1906. An enumeration of Philippine Gramineae with keys to genera and spe-
cies. Philipp. J. Sci. 1, Suppl. 5:307-392.
Pinto-Escorar, P. 1981. The genus Bromus in northern South America. Bot. Jahrb. Syst.
102:445-45 7.
Presi, K.B. 1825-30. Reliquiae Haenkeanae. Prague. 1:1-356.
Presi, K.B. 1831-35. Reliquiae Haenkeanae. Prague. 2:1-152.
Serna, A.E. and A.R. Lorez-Ferrari. 2000. Las Monocotiledoneas Mexicanas, una Sinopsis
Flor{stica. 1. Lista de referencia, partes IX a XI, Pandanaceae a Zosteraceae. Consejo
Nacional de la Flora de México, A.C., Mexico D.F.
Soperstrom, TR. and J.H. Beaman. 1968. The genus Bromus (Gramineae) in Mexico and Cen-
tral America. Publ. Mus. Michigan State Univ., Biol. Ser. 3:465-519.
SorenG, R.J., G. Daviose, PM. Peterson, F.O. Zuoaea, E.J. Juoziewicz, and T.S. Fitaueiras. 2000. Cata-
logue of New World grasses (Poaceae), suprageneric classification (updated Oct 2002).
http://mobot.mobot.org/W3T/Search/nwgc.htm|
STANeik, D. 2001.New records of the genus Festuca L. (Poaceae) for Colombia and Venezu-
ela. Caldasia 23:337-339.
Stancik, D, 2003. Las especies del género Festuca L. (Poaceae) en Colombia. Darwiniana, in
press.
Stanéik, D.and PM. Peterson. 2002. Two new species of Festuca from South America (Poaceae:
Loliinae: sect, Subulatae). Sida: 20:21-29.,
Stearn, W.T. 1973. Introduction in Presl, K.B., Reliquiae Haenkeanae (reprint of original edi-
tion). Prague 1830-1831, vol.1, 356 pp. Asher and Company, Amsterdam.
Stewart, H.and RJ. Hespa. 2000. Grasses of the Colombia Basin of British Columbia. British
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VetoKamp, J.F., M. Eriks, and S.S. Smith. 1991. Bromus (Gramineae) in Malesia. Blumea 35:
THE GENUS PACKERA (ASTERACEAE: SENECIONEAE)
IN COLORADO, USS.A.
Debra K. Trock'!
Botanical Research Institute of Texas
509 n Street
Fort Worth, ne an 4060, U.S.A
ABSTRACT
Fourteen species of Packera are known to occur in Colorado. Recently published floras for the state
have treated these species variously as Packera, Senecio and Ligularia. While preparing a treatment
of this genus for the Flora of North Americ a ia the auenor discovered some confusion among
} fe) P|
a a al b botanists Who rely on those {floras for this dif ficult group. This paper pre-
“Tr ipti
ns for each of the species that occur in the state, comments on how to differentiate
them in the field, and identification keys.
RESUMEN
Se conocen catorce epee de Pa va ra en Colorado. Las floras del estado publicadas recientemente
han trat | Packera, Senecio y Ligularia. Mientras preparaba
]
te género para la Flora de Norte América, el autor encontr6 confusion entre los
botanicos locales que usan estas floras para las identificaciones de este eyine dificil. ae artic ule
presenta descripciones de las especies que viven en el estado
enel campo, y claves de identificacion
INTRODUCTION
What Distinguishes a Packera from a Senecio?
Traditionally, species now included in Packera were relegated to the genus Senecio,
a large genus of over 3,000 species. In the early 1800s DeCandolle (1838) pro-
vided a worldwide treatment of Senecio. He recognized 16 informal groups, based
largely on geographic distribution. Greenman (1915, 1916, 1917, 1918) and Rydberg
(1900) had varying definitions and numbers of informal “species groups.”
Through the 1960s into the 1980s Barkley (1962, 1968a, 1968b, 1978, 1980) pub-
lished several papers on three of these informal groups, the Aurei, the Lobati
and the Tomentosi, which he collectively referred to as the “Aureoid Senecios.”
Within Senecio sl, most of the taxa have a base chromosome number of
x=10 or derivatives thereof. However, Love and Léve (1976) pointed out that there
was a group of anomalous species whose chromosomal base number was ei-
ther x=22 or x=23 and hada suite of unique morphological characters. Further-
more those taxa corresponded to three New World groups informally recog-
nized by Barkley: the Aurei, the Lobati, and the Tomentosi. They proposed that
these species be placed in a new genus, Packera, based primarily on gross mor-
phology and chromosome number.
'Current address: Michigan State University, 166 Plant Biology, East Lansing, Ml 48824-1312, U.S.A.
SIDA 20(3): 1023-1041. 2003
1024 BRIT.ORG/SIDA 20(3)
The recognition of Packera was not immediately accepted and still is not
recognized by some workers. However, support for recognizing Packera has
grown over the past 25 years. Molecular, palynological, and cytological data,
while doing little to help clarify relationships between species in this group,
have all lent support for recognizing the genus Packera. These types of data are
acquired by laboratory work, but how does a botanist in the field decide if he/
she is looking at a Packera or a Senecio? Table | provides a comparison of char-
acters that are useful for field or herbarium identifications. Often a combina-
tion of these features will be needed to make a determination.
MATERIALS AND METHODS
While preparing a treatment of the genus Packera for the Flora of North
America project, | borrowed specimens from many regional herbaria, and vis-
ited several others to obtain both accurate distribution records, and a good un-
derstanding of the morphological variation that is characteristic of these spe-
cies. | have also made four collecting trips through the central part of Colorado,
where the diversity of Packera is the highest. During the course of that work it
became apparent that there was confusion among local botanists concerning
which characters provide for accurate identification, and about which species
belong in the genus Packera, and which are still treated as Senecio s.l.
have examined over 1100 Colorado specimens from the following herbaria:
Adams State College (ALAM); Botanical Research Institute of Texas (BRIT);
Brigham Young University (BRY); Canadian Museum of Nature (CAN); Colo-
rado College (COCO); Colorado State University (CS); Kansas State University
(KSC): Miami University (MU); Missouri Botanical Garden (MO); Montana State
University (MONT); New York Botanical Garden (NY); North Dakota State Uni-
versity (NDA); Northeast Louisiana University (NLU); Oregon State University
(OSC); Rancho Santa Ana Botanic Garden (RSA); Rocky Mountain Herbarium
(RM); University of Alaska Museum (ALA); University of British Columbia
(UBC); University of California (UC); University of Colorado (COLO), Univer-
sity of Kansas (KANU); University of New Mexico (UNM); University of South-
ern Colorado (PUSC); University of Washington (WTU); and Vanderbilt Uni-
versity (VDB).
The goal of this paper is to provide descriptions, keys, common synonyms,
and a discussion of the most useful morphological characters to assist bota-
nists in their efforts to accurately identify species in this genus. Dot maps pro-
viding distribution information for each species are presented in Figures |, 2
and 3.
Generic characters of Colorado Packera species
All taxa of Packera are exclusively herbaceous, and most are perennials. Three
of the Colorado species, P. multilobata, P. plattensis,and P neomexicana can also
TROCK, THE GENUS PACKERA IN COLORADO 1025
Taste 1. Comparison of characters useful for field or herbarium identifications of Packera and
Senecio.
Packera Senecio Informal Groups of
Barkley within Senecio
Leaves progressively Leaves concentrated distally, or Suffruticosi
reduced distally equally distributed along the stem
Heads erect on peduncles Heads nodding Amplectentes
Phyllaries occasionally with Phyllaries with prominent black tips, Amplectentes
reddish tips. Rarely black- AND middle cauline leaves clasping
tipped, and if so, middle cauline
leaves NOT clasping
Caudices taprooted or Caudices button-like or lateral Lugentes, Boreali,
rhizomatous, with branching rhizomes, with fibrous roots Integerrimi
fibrous lateral roots unbranched and fleshy
be biennials, and P neomexicana may occasionally be a winter annual. Stems of
all Packeras consistently grow from either a single or a cluster of basal rosettes.
The perennating structure is a woody caudex or a shortened rhizome. A
caudex is the persistent base of an herbaceous stem that normally arises froma
taproot. In some species the caudex is short, thin, and weak, while in others it
can become elongated, branching, stout, and woody. Species that inhabit steep
rocky slopes have creeping rhizomatous caudices. Taxa growing in wet meadows
or other mesic environments frequently have abundant fibrous roots, which
may almost completely disguise the presence of the caudex.
Stems are normally single from the caudex or taproot. However, because in
many species the caudex is more or less branching, it may appear that there are
several stems clustered together, but usually no more than four or five. Some
species may have stems that display anthocyanic (reddish) pigments. P.
multilobata, growing in open sagebrush habitats, and P. porteri, growing on ta-
lus slopes both exhibit this characteristic.
There is considerable variation in the amount of pubescence along the
stems. In Colorado, P. cana, P fendleri, and P. werneriifolia are nearly always
densely tomentose. Other species vary widely in the amount and persistence
of tomentum. Almost all Packeras have at least some pubescence restricted to
the base of the stem and to the axils of the leaves when young. Although the
presence or absence and location of pubescence is commonly used in keys and
descriptions, it should only be used in conjunction with other characters when
identifying specimens.
The shape and texture of basal leaves have long been of key taxonomic
importance in the genus. The shape of the blade, base of the blade, and blade
margins are the three most important characters. In general the basal leaves are
1026 BRIT.ORG/SIDA 20(3)
Packera cana Packera crocata
Packera dimorphophylla var. dimorphophylla
petiolate. The only exception among Colorado Packeras is P. werneriifolia. Some
individuals of this species have basal leaves that are sessile.
The shape and size of cauline leaves can also be useful, but there may be
extreme variability within species. In all species of Packera, the basal leaves
are arranged in rosettes, while the cauline leaves are either gradually, progres-
1027
TROCK, THE GENUS PACKERA IN COLORADO
Packera paupercula
Packera neomexicanavar. mutabilis
Packera plattensis Packera porteri
Packera streptanthifoliavar. streptanthifolia
oe ae | ay < d\
}
Fic. 2. Distributi
r r
sively, or abruptly reduced upwards on the stem. In all species, the uppermost
cauline leaves are sessile and most are bract-like.
The heads in the inflorescence are arranged in either subumbellate or
corymbiform cymes. Two species (P. porteri and P. werneriifolia) nearly always
have only one large terminal head or at most two or three smaller heads.
Most species of Packera have 8 or 10 or 13 ray florets per head. However, P.
1028
BRIT.ORG/SIDA 20(3)
°
e
s
e e ° e .
°
.
2 °
it _ |.
J
. - °
e °
°
se
7 e
Packera tridenticulata
Fi. 2 Di 1 . £.. Deel er ae | 1.7 . 4)
r r A /
debilis, is typically rayless. A few others, including P. paupercula, P. porteri, P.
pseudaurea, and P. werneriifolia are occasionally rayless. In most species the
florets are a bright yellow. P dimorphophylla var. dimorphophylla and P. crocata
frequently have florets that are deep orange-yellow to almost brick-red. P.
streptanthifolia may have rays that are a deep orange-yellow as well.
Another character that is frequently used in descriptions is size and pu-
bescence of the cypselae. While there is variation in size, P. cana has noticeably
large fruits and P. dimorphophylla has noticeably small ones. Many species have
glabrous fruits, but a few have tiny hairs along the ribs of the cypselae.
—
KEY TO THE PACKERAS OF COLORADO
1, Plants distinctly tomentose throughout, or occasionally glabrate.
2. Margins of leaf blades shallowly but evenly pinnatifid to pinnatisect or merely
Wavy
P. fendleri
2 Hes ins of leaf blades variously entire or dentate, but not pinnatifid
. Plants with 1-3(—5) large heads, alpine to sub-alpine, scapose or with cauline
leaves reduced to mere bracts P. werneriifolia
3. Plants without the above combination of characters.
4, Plants persistently lanate or canescent, leaf blades ovate or elliptic, or rarely
broadly lanceolate, leaf margins entire or merely w P.cana
4. Plants with persistent dense eee at least at Ene base of the stem
and in leaf axils. Lightly tomentose elsev glabrescent with a
Basal leaves lanceolate, oblanc re or an elliptic, nee cauline
leaves similar to basal leaves, calyculate bracts long and conspicuous.
Plants of Ra meadows and coniferous woodlands at mid to high
elevat P.ne xicana var. mutabilis
Basal ae elliptic to ovate or oblanceolate, lower cauline ieee lyr
to pinnatisect, a bracts few and inconspicuous. Plants ae
prairies and gras S
1. Plants glabrous, may
nN
P. plattensis
ye tomentose when young, or persistently so in the axils of
and at the base of the stem; or rarely lightly pubescent throughout.
6. Heads rayless
ray florets much reduced, involucres greenish with anthocyanic
TROCK, THE GENUS PACKERA IN COLORADO
1029
tips; lower and mid-cauline leaves pinnately lobed with deep rounded sinuses
P. debilis
6. Heads radiate, or if rays lacking only 1 head per stem and involucres deeply
anthocyan
7. Plants es a single large head; involucres deeply anthocyanic, basal leaves
reniform to subreniform, cauline leaves reduced to bracts, plants of alpine
scree slopes P. porteri
7. Plants with few to several heads, involucres and leaves without the above
combination of characters.
8. Basal leaf blades 1 road | bl lat
narrowly elliptic.
9. iewereailing ies ee hoe leaves, upper ones becoming sessile;
calyculate bracts conspicuous and numerous but small; cypselae hirtel-
lous along the angles; plants of prairies and sagebrush habitats
P. tridenticulata
9, Lower cauline cee various; calyculate bracts either few and incon-
spicuous or large a nspicuous; cypselae glabrous; plants of moun-
tain or wet me sdew| habitats.
10. Involucres dark green, often with anthocyanic tips; heads with n
merous large conspicuous calyculate bracts P. nie nhiole var.
streptanthifolia
10. Involucres light green, tips not distinctly anthocyanic; calyculate
bracts ew ene InEOnS PICHOUS:
{| f |
eaf blades broadly lanceolate b leaf blades trun-
cate to subcordate or occasionally abs lower cauline leaves
sublyrate to subentire; middle and upper cauline leaves sessile
and frequently clasping P. pseudaurea var flavula
11. Leaf blades lanceolate to narrowly elliptic; base of leaf blades
tapering to cuneate or obtuse, but never subcordate; lower
cauline leaves often sublyrate or with margins variously dis-
sected, but rarely subentire; middle and upper cauline leaves
mere sessile bracts P. paupercula
8 Basal leaf blades broadly elliptic, obovate or oblanceolate to ovate or
orbicular
12. Leaf margins lobed or pinnatifid; plants of sandy sagebrush habitats
P. multilobata
12
. Leaf margins var o crenate or shallowly lobed but never
pinnatifid; slats of alone to ene habitat
13. Lower and middle cauline leaves sessile, acids clasping
and auriculat P. dimorphophylla var. cinoncharholls
13. Lower and ace cauline leaves sessile or petiolate, may be weakly
clasping but never auriculate
14. Leaf blades thin, margins entire to weakly crenate or merely
wavy; upper cauline leaves sessile and weakly clasping; rays
often deep orange yellow or brick red;calyculate bracts few or
absent; plants of sub-alpine meadows P. crocata
14. Leaf blades thick and turgid, margins variously entire to con-
spicuously dentate; upper cauline leaves sessile and reduced
to mere bracts; rays deep yellow, but never reddish; calyculate
bracts conspicuous; habitats from damp to drying ___ P. streptanthifolia
var. streptanthifolia
1030 BRIT.ORG/SIDA 20(3)
SPECIES DESCRIPTIONS AND COMMENTS ON IDENTIFICATION
Packera cana (Hook.) W.A. Weber & A. Love, Phytologia. +9:46. 1981. Senecio canus Hook.
: shi s Nutt; Senecio canus var. purshianus A, Nelson; Senecio canus var. eradiatus
De ae iene onus Senecio hallii Britton; Senecio harbourii Rydb.: Senecio
nvallium ! Senecio hallii var. discoidea WA. Weber
]
Perennials. Caudices stout, branching r] tous, suberect. St from
a rosette, or clustered rosettes, densely lanate-pubescent or canescent. Basal
leaves petiolate, blades ovate or elliptic to lanceolate, 2.5-5+ x 1-3 cm, bases
tapering, margins entire or irregularly undulate to weakly dentate, persistently
canescent on lower surface, canescent to subglabrescent on upper surface.
Cauline leaves petiolate to sessile and weakly clasping, progressively reduced
distally, elliptic to lanceolate, margins entire to weakly dentate. Heads radiate,
8-15+ in corymbiform arrays. Peduncles bracteolate, densely lanate to floccose
tomentose, subglabrescent in some populations. Calyculi present, inconspicu-
ous, tomentose. Phyllaries 13 or 21, green, 5-8+ mm, densely tomentose. Ray flo-
rets 8-10 or 13, laminae 8-10+ mm. Disc florets 35-50+4; corolla tubes 2.5-3.5
mm, limbs 3.5-4.5 mm. Cypselae 2.5-3.5+ mm long, glabrous, pappi 4.5-7 mm
long. 2n=46.
Packera cana is found on open high plains and in sagebrush associations.
It is also collected from dry rocky slopes and from crevices on both granitic
and limestone outcrops. It [lowers from late June into August, depending on
latitude and elevation and is very abundant in Colorado. Although this species
is found on grasslands in Wyoming, Montana, North Dakota, South Dakota, and
Nebraska, it is rarely found on the high plains of the eastern part of Colorado.
Probably the only other species in Colorado with which it can be confused
is P werneriifolia. Two features contribute to this potential confusion: 1) At very
high elevations P cana may be very short (< 10 cm); 2) P. werneriifolia can be
nearly as pubescent as P. cana.
There are two features distinguishing these two species: 1) P cana nearly
always has more than 3 heads in the inflorescence. At high elevations the inflo-
rescence is often very compact, and the heads are small. Packera werneriifolia,
on the other hand, normally has only | or two large heads. It can occasionally
have 3-5 heads, in which case they are smaller than normal. 2) P. cana always
has some cauline leaves even if they are very reduced and mostly on the lower
portion of the stem, while P. werneriifolia is nearly always scapose, or in some
very robust specimens, may have a few bract-like leaves on the stem.
Packera crocata (Ryd b.) Weber & Léve, Phytologia. 49:46. 1981. Senecio crocatus Rydb.
Senecio dureus var. croceus A. Gray; Senecio pyrrochrous Greene; Senecio tracyi Rydb.
Perennials. Caudices long, stout, horizontal to ascending. Stems 2-6+ dm, single
or rarely 2-3 clustered, glabrous. Basal leaves thin petiolate, blades narrowly
lanceolate or ovate to oblong ovate, 2-6+ x 1-4+cm, bases abruptly contracted
TROCK, THE GENUS PACKERA IN COLORADO 1031
to tapering, margins subentire to crenate-dentate. Cauline leaves petiolate to
sessile and weakly clasping, progressively reduced distally, lanceolate to ob-
long, margins sublyrate or lobed. Heads radiate, 7-15+ in open corymbiform
arrays. Peduncles absent or small and inconspicuous, glabrous. Calyculi absent
or few and small. Phyllaries 13 or 21, light green or with reddish tips, 4-8 mm,
glabrous. Ray florets 8 or 13, corolla laminae 6-8+ mm, deep yellow to orange-
red. Dise florets 60-80+; corolla tubes 4.5-5.5 mm, limbs 2.5-3.5 mm. Cypselae
1-15 mm long, glabrous, pappi 3-5 mm long. 2n=46.
This species grows in wet meadows often associated with grasses, sedges
and willows, along trails and in rocky outcrops at middle to subalpine eleva-
tions and flowers from early July to mid-August. The only species that it may
be confused with is P dimorphophylla var. dimorphophylla. Three features dis-
tinguish these two species: 1) P.crocata has an open inflorescence, while that of
P. dimorphophylla var. dimorphophylla is compact and congested. 2) P.
dimorphophylla has dramatically large, auriculate, and conspicuously clasp-
ing middle and upper cauline leaves. While P crocata may have larger than nor-
mal cauline leaves, they are at most weakly clasping. 3) While both species gen-
erally grow in wet places, P. crocata grows at elevations of ca. 8,000 - 10,000 feet
and is fairly tall and thin, while P dimorphophylla var. dimorphophylla is al-
most never found below 10,000 feet and is normally a more compact plant.
It is frequently noted in the literature that P. crocata has orange to reddish
ray florets. However, P crocata sometimes produces deep yellow ray florets, and
P. dimorphophylla var. dimorphophylla may produce orangish ray florets.
Packera debilis (Nutt.) WA. Weber & A. Love, Phytologia. 49:46. 1981. Senecio
debilis Nutt.
Senecio fedifolius Rydb.
Perennials. Caudices short, weakly branching. Stems 20-50+, single or 2-4 clus-
tered, bases lightly floccose-tomentose or glabrous, leaf axils tomentose. Basal
leaves petiolate and turgid, blades elliptic to ovate or subreniform, 2-4+ cm x 1.5-
3+ cm, bases cuneate to subcordate, margins subentire or crenate to crenate-den-
tate. Cauline leaves petiolate to sessile, progressively reduced distally, blades pin-
nate-lobed, sinuses deep and rounded, lobes with entire to subentire margins.
Heads eradiate, 6-20+ in open or compact corymbiform arrays. Peduncles present,
short and conspicuous or absent, glabrous or lightly pubescent. Calyculi present,
few and inconspicuous. Phyllaries 13 or 21, green with reddish tips, 6-8+ mm,
glabrous. Ray florets absent. Disc florets +5-65+, corolla tubes 3-4 mm, limbs 2-
3 mm. Cypselae 1-2 mm long, glabrous, pappi 4.5-5.5 mm long. 2n=46.
This is a distinctive plant of central and northern Colorado. It is collected
frequently in Park County and less frequently in northern counties at mid to
subalpine elevations in open meadows with alkaline soils. Packera debilis is
sometimes identified as P pauciflora, which is alsoa rayless plant with reddish
1032 BRIT.ORG/SIDA 20(3)
involucres. However, P. pauciflora isa plant witha more northerly distribution
(it does not occur in Colorado) and a suite of characteristics that distinguish it
from P. debilis. 1) Packera debilis generally has more than 6 heads in the inflo-
rescence while P. pauciflora normally has 6 or fewer large heads. 2) Packera
debilis has very short bracteoles on the peduncles, while P. pauciflora has very
large conspicuous ones. 3) Packera debilis, has very few greenish, inconspicu-
ous calyculate bracts. P pauciflora,on the other hand, has long calyculate bracts
that are normally quite reddish in color or at least have red tips. 4+) The involu-
cral bracts of P debilis may display reddish coloration on the upper one-half to
one-third of the tips, but the involucres of P. pauciflora are normally deeply
anthocyanic throughout. 5) Probably the most easily distinguished character-
istic concerns the lobing of the mid-cauline leaves. The terminal lobe of the
leaves on P debilisis narrow and pointed, and the sinuses of the individual lobes
are deep and rounded.
Although P. debilis has been known at least from Park, Larimer, and Jack-
son counties for many years, it is very infrequently collected and is confined to
alkaline meadow habitats. It may indeed be worthy of special note, and an ef-
fort should be made to document existing populations.
Packera dimorphophylla (Greene) var. dimorphophylla W.A. Weber & A. Love,
Phytologia. 49:46. 1981. Senecio dimorphophyllus Greene var. dimorphophyllus
Perennials. Caudices short, stout, fibrous-rooted, horizontal to erect. Stems |-
3+ dm, single or rarely 2-3 clustered, simple or branched, glabrous. Leaves: basal
leaves petiolate, thick and turgid, blades varying from ovate to subreniform or
occasionally oblanceolate, bases tapering to a broad-winged petiole, margins
entire to crenate; lower and mid-cauline leaves sessile, conspicuously clasping
and auriculate, blades as large or larger than basal leaves, oblanceolate, obovate
or lyrate, margins subentire to bluntly and irregularly dissected; upper cauline
leaves usually reduced to sessile bracts, occasionally large and conspicuous.
Heads radiate, 1-6+ in congested corymbiform or subumbellate arrays. Pe-
duncles inconspicuously bracteolate or bracteoles absent, glabrous. Calyculi
conspicuous, bases swollen. Phyllaries 13 or 21, occasionally fewer, green with
anthocyanic apices, 5-7+ mm, glabrous. Ray florets 8 or 13, deep yellow to or-
ange, corolla laminae 5-8+ mm. Disc florets 45-60+, corolla tubes 2-3 mm, limbs
2.5-3.5 mm. Cypselae 0.75-1.5 mim long, glabrous, pappi 3-4 mm long. 2n = 46.
This species is quite distinctive and is only rarely confused with one other
entity in Colorado, P. crocata. A discussion of the characters used to separate
the two species is found with the description of P. crocata. It is found at eleva-
tions generally over 10,000 feet in damp meadows or in seeps and crevices on
hillsides. It flowers from early July to mid-August.
Packera fendleri (A. Gray) W.A. Weber & A. Love, Phytologia. 49:46. 1981. Senecio
fendleri A. Gray
TROCK, THE GENUS PACKERA IN COLORADO 1033
Senecio nelsonii Rydb.; Senecio salicinus Rydb,; Seneci Rydb.; Senecio rosulatus Rydb:
Senecio fendleri var. molestus Greenm.
Perennials. Caudices rhizomatous, prominent lateral branches, horizontal to
suberect. Stems 1-4+ dm, single or crowded to subcaespitose in some popula-
tions, floccose-tomentose or glabrescent. Basal leaves petiolate, blades lanceolate
to oblanceolate, 3-6+ x 1-3+ cm, bases tapering, margins shallowly but evenly
pinnatifid to pinnatisect or merely wavy, permanently floccose-tomentose or
subglabrescent on upper surface. Cauline leaves sessile, gradually reduced dis-
tally, lanceolate to oblanceolate, margins pinnatisect to wavy. Heads radiate, 6-
25+ in open or compact corymbiform arrays. Peduncles with numerous small
bracteoles, heavily to irregularly floccose. Calyeuli present but few and short,
red tinged. Phyllaries 13, green, 5-7 mm, floccose proximally to glabrescent dis-
tally. Ray florets 6-8+, corolla laminae 5-7 mm. Disc florets 30-40+; corolla tubes
2.5-3 mm, limbs, 2.5-3.5 mm. Cypselae 2.5-3 mm long, glabrous, pappi 4-5 mm
long. 2n=46.
This species is rarely confused with other Colorado Packeras. Some indi-
viduals, with merely wavy leaves could be mistaken for P. cana, but close in-
spection should easily separate the two. Packera fendleri is very abundant and
almost weedy where it occurs. It prefers steep slopes in loose, dry rocky or grav-
elly soil, streamsides, open forested areas, and disturbed sites such as roadsides
and picnic areas. It flowers any time between the end of May and early October
throughout its range, depending on elevation.
Packera multilobata (Torr. SA — ex A. oe W.A. Weber & A. Léve,
Phytologia. 49:47. 1981. s ts Torr. & A. Gray ex. A. as
Senecio lynceus Greene; Senecio nels rar. uintahensis A. Nelson; S 5
23 necio stygius Greene; Senecio ol xus Greenm.; Senecio leucoreus Greenm.; Senecio
PHPErL OLeCHilh.
Perennials or biennials. Caudices weakly branching, taprooted, ascending to
erect. Stems 2-4+ dm, single or 2-5 loosely clustered, glabrous or glabrescent,
occasionally lightly tomentose throughout, axils of basal leaves with tufts o
tomentum. Leaves: basal and lower cauline leaves petiolate, blades obovate,
oblanceolate or spatulate, 4-8+ « 1-3+ cm, bases tapering, margins deeply pin-
natifid or lyrate to sublyrate, leaf segments variously toothed and rounded;
middle and upper cauline leaves progressively reduced distally, sessile. Heads
radiate, 10-30+ in open corymbiform or subumbellate arrays. Peduncles con-
spicuously bracteolate, glabrous or with tufts of tomentum. Calyculi present
and obvious. Phyllaries 13-21, green often with yellow tips, 4-9+ mm, glabrous
to lightly tomentose proximally. Ray florets 8-13, corolla laminae 7-10 mm. Dise
florets 40-50+, corolla tubes 4-5 mm, limbs 3-4 mm. Cypselae 2-3 mm long,
glabrous or hirtellous on the angles, pappi 5-6 mm long. 2n = 46 or 92
This species is also rarely confused with any others in Colorado. P.
1034 BRIT.ORG/SIDA 20(3)
multilobata and P.fendleri are the only distinctly lobed or pinnatifid Packeras
in the state. Packera fendleri is consistently very pubescent and P. multilobata
is nearly always glabrous. Packera multilobata occurs on the western slope but
thus far has not been recorded from counties along the Rocky Mountain Front.
It prefers dry rocky or sandy substrates in habitats as varied as desert sagebrush,
juniper woodlands, and subalpine forests. It flowers in early May at lower eleva-
tions and as late as mid-July at higher elevations.
Packera neomexicana iGreene) var, mutabilis WA. Weber SAL Ok lec
eg eal lols. ;
neomexicanus va ) TM.
lisG re
Barkley
Perennials. Caudices branched, weakly spreading, horizontal or ascending to
erect. Stems 2-5+ dm, single or 2-3 clustered, densely lanate or arachnoid to-
mentose or glabrescent. Basal leaves petiolate, blades lanceolate or narrowly
lanceolate, 2-6+ x 1-3+ cm, bases tapering, margins subentire to dentate, per-
manently but loosely tomentose throughout. Cauline leaves progressively re-
duced distally; lower leaves petiolate and similar to basal leaves; mid to upper
cauline leaves becoming sessile, blades lanceolate, margins entire. Heads radi-
ate, 3-20+ in open or compact corymbiform to subumbelliform arrays, sub-
tended by smaller cymes arising from leaf axils. Peduncles conspicuously
bracteolate, normally pubescent. Cayleuli present and conspicuous. Phyllaries
13 or 21, green or yellowish, 4-7 mm, tomentose to glabrescent. Ray florets 8 or
13, occasionally 5, corolla laminae 4-10 mm. Disc florets 40-60+, corolla tubes
1.5-2.5mm, limbs 3.5-4.5 mm. Cypselae 1.5-2.5 mm long, hirtellous on the angles
or infrequently glabrous, pappi 5-6+ mm long. 2n = 46 or 92
This species flowers from late April to mid-July. It is frequently confused
with Packera tridenticulata. Four characteristics, when taken in combination,
can help to distinguish these two entities.
1) P.tridenticulata is a plant of the high plains and foothills in Colorado. In a
few cases, it has apparently “jumped” the Rocky Mountain Front and has been
collected growing in some of the interior valleys, usually in grassland/sage-
brush associations. On the other hand, P neomexicana var. mutabilis is gener-
ally found growing in rocky soils in open meadows and coniferous woodlands
at high elevations.
2) P. tridenticulata usually grows in obvious clumps from a stout taprooted
caudex, while P. neomexicana var. mutabilis normally has no more than 2-3
stems arising from a branching caudex
3) P. tridenticulata tends to have slightly smaller achenes that are normally
glabrous or at most lightly hirtellous along the angles, while P neomexicana
var. mutabilis has achenes that are a bit larger and normally distinctly hirtel-
lous along the angles.
TROCK, THE GENUS PACKERA IN COLORADO 1035
4) P.tridenticulata collected from the eastern part of its range is always glabrous.
In the western part of its range, and in the interior valleys west of the Rocky
Mountain Front, entities that would otherwise be easily identified as P
tridenticulata often show some very light pubescence at stem bases and in leaf
axils. Packera neomexicana var. mutabilisisalways loosely but persistently tomen-
tose at least at the base of the stem, in the axils of the leaves, and at the base of the
inflorescence. The lower leaf blades are also almost always lightly pubescent.
The center of distribution for P neomexicana var. mutabilis is in the four-cor-
ners region, while P. tridenticulata is primarily a plant of the high plains and
short-grass prairies. There is good evidence for hybridization between these two
entities, primarily in the high valleys around Alamosa in south-central Colo-
rado. There are several specimens in the herbaria at Adams State College in
Alamosa and Colorado College in Colorado Springs with characters that are
intermediate between P neomexicana and P tridenticulata. Several specimens
display all the cl teristics of P tridenticulata, but are unusually and persis-
tently tomentose, while others that are obviously glabrous have hirtellous
achenes. Biosystematic work on these two species throughout the central val-
leys of Colorado would undoubtedly shed light on the complicated relation-
ships in this complex.
Packera neomexicana var. mutabilis is infrequently confused, with P
streptanthifolia and P.fendleri. Some specimens of P.fendleri with merely wavy
leaves and less than the normal amount of tomentum have been misidentified as
P. neomexicana var. mutabilis. Packera streptanthifolia is a highly variable spe-
cies with a number of identifiable “phases.” The narrow-leaved phase of P.
streptanthifolia is sometimes difficult to distinguish, but once again, careful ob-
servation of acombination of characters should provide accurate identifications.
Packera paupercula (Michx.) Love & Léve, Bot. Not. 128:520. 1976. Senecio
pauperculus Michx.
fc i Britton; Sene-
Senecio balsamitae vee ex Wi nes Senecio plavew irens Rydb.;
cio tweedyi Rydb. sis Greenm.; Senecio gaspensis Greenm.
Perennials. Caudices sub-rhizomatous, weakly branching, ascending to erect.
Stems 2-4.5+ dm, single or 2-4 loosely clustered, glabrous or lightly tomentose
proximally. Leaves: basal leaves petiolate, blades lanceolate to narrowly elliptic
or oblanceolate, 3-6+ x 1-2+ cm, bases tapering to cuneate or obtuse, margins
subentire to variously dentate or serrate. Cauline leaves progressively reduced
distally; lower ones petiolate, blades sublyrate; middle ones sessile, blades lan-
ceolate, margins variously dissected, incised or lacerate; upper ones sessile
bracts. Heads radiate, 2-10+ in loose or compact corymbiform arrays. Peduncles
bracteolate or bracteoles occasionally absent, glabrous. Calyculi present and
inconspicuous. Phyllaries 13 or 21, green, 5-8+ mm, glabrous. Ray florets 8 or 13,
or occasionally absent, corolla laminae 5-10+ mm. Disc florets 50-65+, corolla
1036 BRIT.ORG/SIDA 20(3)
tubes 2-3 mm, limbs 2-3 mm. Cypselae |-2 mm long, glabrous or occasionally
hispidulous on the angles, pappi 3.5-4.5 mm long. 2n = 44 or 46.
Packera paupercula is the most ecologically and morphologically variable
Packera in North America. Although widespread, it has been infrequently col-
lected in Colorado. It flowers from late June to late July in Colorado and can be
found in wet meadows, open woodlands, and along streambanks. In areas of
sympatry, P paupercula can be confused with P plattensis, but that is not a prob-
lem in Colorado because P. plattensis is confined to the high plains and P
paupercula has only been collected from mountainous regions.
The other species with which P. paupercula could easily be confused is P.
pseudaurea var. flavula. There are six characteristics that can be used in combi-
nation to separate these two entities: 1) Basal and lower leaf blades of P.
paupercula are lanceolate to narrowly elliptic, and normally |.5-3 times longer
than wide. Those of P. pseudaurea var. flavulda are broadly lanceolate to ovate,
and |-2 times longer than wide; 2) Bases of the leaf blades of P paupercula are
tapering, those of the P pseudaurea var. flavula are truncate to subcordate; 3)
Upper cauline leaves of the former are sessile, those of the latter are weakly
clasping; +) Phyllaries of P. paupercula specimens from Colorado normally
number 13 and are 5-8 mm long, those of P. pseudaurea var.flavula usually num-
ber 2L and are 3-5 mm long; 5) P. paupercula normally has 40-50 disc florets in
a one head, P. pseudaurea var. flavula has 60-80; 6) Corolla tubes of the disc
florets in P. paupercula are about the same length as the limbs, 2-3 mm, how-
ever in P. pseudaurea var. flavula, the corolla tubes are much longer than the
limbs, 3.5-4.5 mm and 2-3 mm, respectively.
—
Packera plattensis (Nutt.) W.A. Weber & Love, Phytologia. 49:48. 1981. Senecio
plattensis Nutt.
Senecio pseudotomentosus Mackenzie & Bush,
Perennials or biennials. Caudices stoloniferous in eastern populations, fibrous
rooted elsewhere, erect or suberect. Stems 2-6+ dim, single or 2-3 clustered, per-
sistently floccose-tomentose proximally and in leaf axils, lightly tomentose or
glabrescent elsewhere. Leaves: basal and lower cauline leaves petiolate, blades
narrowly elliptic to elliptic-ovate or oblanceolate to suborbicular, abaxial sur-
face subpersistently Floccose-tomentose, especially along the mid-vein, 2-7+ x
1-3+ cm, bases tapering to rounded or abruptly contracted, margins subentire
to crenate, serrate-dentate, or sublyrate; middle cauline leaves progressively
reduced distally, short petiolate, blades sub-lyrate to pinnatisect, lightly pu-
bescent on the abaxial surface; upper cauline leaves, sessile, subentire to irregu-
larly dissected. Heads radiate, 6-20+ in open or congested corymbiform arrays.
Peduncles abundantly bracteolate, lightly to heavily tomentose. Calyculi nu-
merous but inconspicuous. Phyllaries 13 or 21, green or occasionally with red-
dish tips, 5-6+ mm, densely tomentose distally, glabrescent above. Ray florets
TROCK, THE GENUS PACKERA IN COLORADO 1037
8-10, corolla laminae 9-10 mm. Disc florets 60-70+, corolla tubes 2.5-3.5 mm,
limbs 3.5-4.5 mm. Cypselae 1.5-2.5 mm long, hirtellous or occasionally glabrous,
pappi, 6.5-7.5 mm long. 2n = 46 or 92.
Packera plattensis inhabits rocky, gravelly, or sandy/loam soils in open
prairies and grasslands. It is found throughout eastern Colorado but in dry years
may not be abundant. It occurs in the foothills of the Front Range, but to my
knowledge has never been collected further west than that. Packera plattensis
flowers from mid-May to mid-June in Colorado. The only other high plains
Packera species in Colorado is P tridenticulata, and these two species are rarely
confused. Packera plattensis is always densely floccose-tomentose at the base
of the stem and in the leaf axils and often persistently lightly tomentose else-
where. Packeratridenticulata collected on the high plains is nearly always gla-
brous or at most very lightly tomentose at the base of the stem.
Packera porteri (Greene) C. eae a Bull. 47:101. 1992. Senecio porteri Greene:
Ligularia porteri (Greene) WA. \
py ie 1 D. C
Senecio ‘i ‘eus var. alpinus A. Gray.
Perennials. Caudices with long slender branching rhizomes. Stems 3-10+ cm,
single, often anthocyanic, glabrous. Leaves: basal leaves petiolate and turgid,
blades reniform to sub-reniform, 0.5-1.5+ x 0.5-2.5+ cm, abaxial surface dis-
tinctly anthocyanic, bases abruptly contracted to cordate, margins distinctly
crenate or occasionally wavy; cauline leaves abruptly reduced distally to 1-4
sessile bracts with entire margins. Heads radiate, | large terminal head, 10-15+
mm high. Calyculi present and conspicuous, often anthocyanic. Phyllaries 13
or 21, deep red, 8-10+ mm, glabrous. Ray florets 8 or 13, corolla laminae 8-10+
mm. Disc florets 40-50+, corolla tubes 2.5-3.5 mm, limbs 2.5-3.5 mm. Cypselae
1.5-2.5 mm long, glabrous, pappi +-5.5 mm long.
Packera porteri is undoubtedly the most morphologically distinctive
Packera in Colorado and the most restricted in range and habitat. It is confined
to steep talus slopes in alpine habitats, rarely below 11,000 feet. It is most fre-
quently collected west of the Continental Divide in two general locations:
Gunnison and nearby counties, in the Elk Mountains around Gothic and in
Maroon Bells/Snowmass Wilderness areas; and in San Juan and Hinsdale coun-
ties. However, it has also been collected east of the divide in Chaffee County.
Due to its diminutive form and remote location, it may have been overlooked
elsewhere. Packera porteri flowers from mid-July to mid-August.
Packera pseudaurea (Greene) var. flavula D.K. Trock & T.M. Barkley, Phytologia.
49:48. 1981. Senecio pseudaureus var. flavulus (Greene) Greenm.; Senecio flavulus Greene.
Perennials. Caudices simple or branched, fibrous-rooted, horizontal to erect.
Stems 2-5+ dm, thin and slender, single or occasionally 2-4 clustered, glabrous
or lightly tomentose proximally. Leaves: basal leaves petiolate, blades broadly
1038 BRIT.ORG/SIDA 20(3)
lanceolate to ovate or occasionally sub-hastate, 2-4+ x 2-3+ cm, bases trun-
cate to subcordate or obtuse, margins bluntly dentate or denticulate, rarely
sharply toothed; lower cauline leaves petiolate, blades sublyrate to lacinate or
occasionally subentire; middle and upper cauline leaves progressively reduced
distally, becoming sessile, occasionally clasping. Heads radiate, 5-12+ heads in
a congested cyme. Peduncles inconspicuously bracteolate, glabrous. Calyculi
present. Phyllaries 21 or occasionally 13, rarely 30 or more, light green, 3-6 mm,
glabrous. Ray florets 8 or 13, corolla laminae 6-10+ mm, or occasionally absent.
Disc florets 60-804, corolla tubes 3.5-4.5 mm, limbs 2-3 mm. Cypselae 1-1.5
mm long, glabrous, pappi 4.5-5.5 mm long. 2n = 46
This variety of P. pseudaurea is found in the southern Rocky Mountains
and is most abundant in Colorado. It inhabits the same habitat type as P.
paupercula, and its gross morphology is similar. The comments following the
description of P. paupercula provide a suite of six characteristics that can be
used to distinguish the two entities. Packera pseudaurea var.flavula flowers from
ate May to early July. Where the two occur at similar elevations and in similar
habitats, P pseudaurea var. flavula usually flowers about one month earlier than
P. paupercula.
—
—
Packera streptanthifolia (Greene) W.A. Weber & Love var. streptanthifolia
Phytologia. 49:48. 1981. Senecio streptanthifolius Greene; Senecio cymbalarioides var.
streptanthifolius (Greene) Greenm
Senecio ¢ ean Nutt.; Se necto ian id var, ae pence nee) J.E Bain; Senecio
t] if TRS }
—
* Bain; ; E tus Greene; Senecio wardti
Ur.
Greene; Senecio oa Howell; Senecio su were Greenm.; conse longipetiolatus Rydb.
Senecio fulgens Rydb. Senecio rydbergii A. Nelson; Senecio jonesii Rydb.; Senecio subcuneatus
Rydb., Senecio platylobus Rydb. Senecio streptanthifolius var. oodes (Rydb.) J.F. Bain; Senecio
oodes Rydb., Senecio leonardii Rydb.; Senecio streptanthifolius var. kluanei J.F Bain; Senecio
streptanthifolius var. wallowensis J.F Bain.
rete ane weak to stout, fibrous-rooted, horizontal to suberect. Stems
2-4+ dm, single or 2-5 clustered, glabrous or occasionally lightly floccose to-
mentose proximally and in leaf axils. Leaves: basal and lower cauline leaves,
petiolate, thick and turgid, blades varying from oblanceolate or ovate to orbicu-
lar, 2-4+ x 1-3+ cm, bases tapering or abruptly contracted, margins entire,
crenate, dentate or weakly lobulate, normally glabrous or sometimes persis-
tently pubescent; middle and upper cauline leaves gradually reduced distally,
becoming sessile, margins entire to sub-entire. Heads radiate, 6-15+ in loose
corymbiform or subumbellate arrays. Peduncles variably bracteolate, glabrous
or lightly tomentose. Calyculi present and conspicuous. Phyllaries 13 or 21, oc-
casionally 8, green or with anthocyanic tips, 4+-7+ mm, glabrous. Ray florets 8
or 13, corolla laminae 5-10 mm. Disc florets 35-604, corolla tubes 2-4 mm, limbs
2.5-4 mm. Cypselae |-2.5 mm long, glabrous, pappi, 3-6 mm long. 2n = 46.
Packera streptanthifolia grows in a wide variety of habitats, from forested
TROCK, THE GENUS PACKERA IN COLORADO 1039
areas to open meadows and valleys and in soils ranging from dry and rocky to
damp and loamy. It also occurs at a wide range of elevations, from 6500 feet to
above timberline. The species is complex and has been divided by some research-
ers (Bain 1983, 1988) into as many as seven distinct taxa—five species and two
varieties or two species and five varieties. This author recognizes only one spe-
cies and two varieties, while acknowledging that there is a great amount of
morphological variation. There is less morphological variation within this spe-
cies in Colorado than there is in other parts of its range.
The narrow-leaved phase of P. streptanthifolia could superficially be con-
fused with P. neomexicana var. mutabilis. However, the former has glabrous
achenes, while achenes of the latter are normally hirtellous. Packera streptan-
thifolia in Colorado normally has leaves that are ovate to orbicular. Probably
the most reliable characteristics are the thick and turgid leaf blades, and a very
stout fibrous-rooted caudex. In Colorado, this species flowers from early June
to early July at lower elevations and from mid-July to mid-August at high
elevations.
Packera tridenticulata (Rydb.) W.A. Weber & Léve, Phytologia. 49:48. 1981. sene-
cio tridenticulatus Rydb.
S s Rydb.; Senecio densus Greene; Senecio oblanceolatus Rydb; $
aydb. Sonieet manitobensis Greenm.; Senecio remifolius Wooton & Standle
tan
Perennials. Caudices t ascending to erect. Stems |-3+ dm, a or fre-
quently several clustered, glabrous or rarely lightly floccose-tomentose, often
with tufts of tomentum in leaf axils. Leaves: basal and lower cauline leaves peti-
olate, blades thick and turgid, lanceolate or narrowly oblanceolate, 2-4+ x 0.5-
15 cm, bases tapering, margins entire, subentire or dentate at the apex, occea-
sionally sub-pinnatisect; middle and upper cauline leaves gradually or
progressively reduced distally, becoming sessile. Heads radiate, 4-15+ heads in
corymbiform arrays. Penduncles bracteolate, glabrous to lightly tomentose.
Calyculi numerous but small. Phyllaries 13-21, green, 6-10 mm, sparsely tomen-
tose proximally, glabrous distally. Ray florets 8-10 or occasionally 13, corolla
laminae 5-8+ mm. Disc florets, 45-604, corolla tubes 3-3.5 mm, limbs 4-5 mm.
Cypselae 1.5-2.5mm long, glabrous or lightly hirtellous on the angles, pappi 5-
6mm long. 2n = 46.
This species can be found in shortgrass prairies or sagebrush scrub habi-
tats, growing in dry gravelly or sandy soils. It is primarily a plant of the central
High Plains, but its range extends into the foothills of the Rockies and into some
of the high valleys west of the Rocky Mountain Front in Colorado. It tends to
grow in clumps with several stems arising from a single taprooted, woody cau-
dex. The only other species in Colorado with which it may be confused is P
neomexicana var. mutabilis. That relationship is discussed in detail under P
neomexicana. Packera tridenticulata flowers from late May to early July.
1040 BRIT.ORG/SIDA 20(3)
Packera werneriifolia (A. Gray) W.A. Weber & Love, Ph ytologia 49:48. L981. Senecio
werneridefolius A. Gray.
Senecio saxosus Klatt; Se fae Sues Greene; . pass eee Nelson; S
A. Nelson; Senecio muirii Greenm.; sGreenm.
Perennials. Caudices branching, sub-rhizomatous, sometimes several densely
crowded and caespitose. Stems 7-15+ cm, single or 3-5 clustered, floccose or
lanate-tomentose, canescent or occasionally glabrate. Basal Leaves sessile or
with very short petioles, blades narrowly lanceolate to elliptic, 1.5-4+ x 0.5-2.5
cm, bases tapering, margins entire or dentate toward the apex, frequently revo-
lute; or in some populations petiolate with blades ovate to orbicular, 1-2 x 0.5-
15cm, bases tapering to abruptly contracted, margins entire or wavy, occasion-
ally dentate toward the apex. Cauline leaves reduced to mere bracts, plant with
a scapose aspect. Heads radiate, 1-5 or in some populations 3-8. Peduncles in-
conspicuously bracteolate, glabrous to densely pubescent. Calyculi conspicu-
ous, frequently anthocyanic. Phyllaries 13 or 21, green or with anthocyanic api-
ces, 4-10 mm, glabrous to pubescent. Ray florets 8 or 13, corolla laminae 5-10
mm; or occasionally absent. Dise florets 30-50+, corolla tubes 2.5-3.5 mm, limbs
3-4 mm. Cypselae 1.5-2 mm long, glabrous, pappi 5-6 mm long. 2n = 44 or 46.
This species grows on rocky talus slopes or in sandy soil in forest openings
near or above timberline. Flowering time varies greatly with elevation but is
generally from mid-June to mid-August. Some specimens have basal leaf blades
that are sessile and very narrow, while the basal leaves of others are petiolate
and more ovate in outline. Some specimens are densely tomentose, while oth-
ersare only lightly pubescent, even glabrate. But the feature that all of the speci-
mens have in common is their scapose aspect.
The only species in Colorado with which it could be confused is P. cana,
and that relationship is discussed in detail under the comments for P. cana.
a.
>
a
ACKNOWLEDGMENTS
lam grateful to Ted Barkley, Guy Nesom, and Ron Hartman for their comments
and helpful suggestions. I would also like to thank Catherine Kleier and Tass
Kelso for providing their insights on the Colorado flora and for their assistance
with herbarium specimens. Thanks also go to all of the institutions mentioned
in this paper for their loan of specimens.
REFERENCES
Bain, J.F. 1983. A biosystematic study of the Senecio streptanthifolius complex. Doctoral
dissertation. Edmonton: The University of Alberta.
Bain, J.F. 1988. Taxonomy of Senecio streptanthifolius Greene. Rhodora 90:277-312.
Bark ey, I.M.1962.A revision of Senecio aureus L.and allied species. Trans. Kansas Acad. Sci.
65:318-408.
TROCK, THE GENUS PACKERA IN COLORADO 1041
Barktey, T.M. 1968a.Intergradation of Senecio sections Aurei, Tomentosi, and Lobati through
Senecio mutabilis. Southw. Naturalist 13:109-115.
Barkley, I.M. 1968b. Taxonomy of Senecio multilobatus and its allies. Brittonia 20:267-284.
Barkley, 1.M. 1978. Senecio. N. Amer. Flora Il. 10:50-139.
Barkley, T.M. 1980. Taxonomic notes on Senecio tomentosus and its allies (Asteraceae).
Brittonia 32:291-308.
DeCanvoute, A.P.1838.Prodromus systematis naturalis reqni vegetabilis vol. 7. Paris: Treuttel
& Wurtz.
GREENMAN, J.M. 1915. Monograph of the North and Central American species of the genus
Senecio, Part ll. Ann. Missouri Bot. Gard. 2:543-626.
GREENMAN, J.M. 1916. Monograph of the North and Central American species of the genus
Senecio, Part Il, Sect.6 Ann. Missouri Bot. Gard. 3:85-194.
Greenman, J.M.1917.Monograph of the North and Central American species of the genus
Senecio, Part Il. Ann. Missouri Bot. Gard. 4:15-36.
Greenman, J.M.1918.Monograph of the North and Central American species of the genus
Senecio, Part Il. Ann. Missouri Bot. Gard. 5:37-103.
Love, A.and D. Love. 1976.Nomenclatural notes on arctic plants. Bot. Notiser 128:497-523.
RybserG, P.A. 1900. Studies on the Rocky Mountain flora |. Bull. Torrey Bot. Club 27:169-189.
BRIT.ORG/SIDA 20
=
1042 3)
BOOK NOTICES
ELIZABETH SILVERTHORNE. 2002. Legends and Lore of Texas Wildflowers. (ISBN 1-
58544-230, pbk.). Texas A&M University Press, John H. Lindsey Bldg,,
Lewis St. 4354 TAMU, College Station, TX 77843-4354, U.S.A. (Orders:
wlawrence@tamuedu, http://wwwtamuedu/upress, 800-826-8911). $24.95
hbk., $16.95 pbk., 264 pp., 18 line drawings, Bib. Index, 7" x 9".
Forty-four Texas wildflowers are discussed in this book. Their botanical names as well as their com-
mon names are given, and stories about the origins and lore surrounding them are the focus. This is
not a field guide or identification book, but rather a collection 0
interesting tales about these par-
ticular flowers.
The author does deal with the habitats, descriptions, uses—medicinal ane scabies —of the
ith i told in non-
plants, and tells something of the early botanists. In fact it is crammed w
the flowers are illustrated, although the artist is not credited. Itisa siege
technical terms. Some of
addition to literature of wild
Worth, TX, l
flowers.—Ruth Ginsburg, Botanical Research Institute of Texas, Fort
Roy L. LEHMAN, RutTH O’Brien, and TAMMY Wuite. 2002. Plants of Webb County,
Texas. (ISBN not given, pbk.). Texas AGM University-Corpus Christi, Dept.
of Physical and Life Sciences, 6300 Ocean Drive, Corpus Christi, TX 78412,
S.A. (Orders: 512-994-2754, fax 512-994-2742). Price not given, 47 pp.,
numerous color photos, | map, index, 51/2" x 81/2"
According to the foreword, “This booklet was first designe ‘ to bs a auc for secondary school, col-
lege students and for the laymen interested in flowers and | the Galvan Ranch,” It was
decided that, as the ranch was no longer accessible to the seul public, that the guide be reformat-
ted and titled to include common sae interesting plants from the county [ Webb.”
‘the species covered in the book includes a color photograph, common name(s), Latin
name and some synonyms, followed by a brief description including habitat and [lowering time.
The authors are to be congratulated for all the hard work on this book. It is especially nice because of
the lack of floras for that part of Texas. A big thank you goes to the Ed Rachel Foundation in Corpus
Christi for their financial support of the project. The Foundation’s support of research on Texas na-
{ Webb County and Texas.—
peas
fay)
oad
S)
tive plants is truly appreciated; what an incredible gift to the peop
Barney Lipscomb, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060,
U.S.A, barney@britorg.
SIDA 20(3): 1042. 2003
NOMENCLATURAL COMBINATIONS IN NORTH AMERICAN
PANICUM SUBGENUS DICHANTHELIUM AND SUBGENUS
AGROSTOIDES (POACEAE: PANICEAE)
Joseph K.Wipff
Herbarium
Pure Seed Testing, Inc.
Canby, OR 97013, U.S.A.
ABSTRACT
new combinations made here are: Panicum aciculare var. ramosum, Panicum aciculare var.
neuranthum; Panicum dcuminatum var. ferventicola; Panicum acuminatum var. thermale; Pani-
cum commutatum var. equilaterale; Panicum ovale var. praecocius, and Panicum rigidulum var.
abscissum.
RESUMEN
Se hacen las siguientes nuevas combinaciones: Panicum aciculare var. ramosum; Panicum aciculare
var. neuranthum: Panicum acuminatum var. ferventicola; Panicum acuminatum var. thermale; Pani-
cum commutatum var. equilaterale; Panicum ovale var. praecocius; y Panicum rigidulum var.
abscissum.
The debate over whether Dichanthelium should be recognized as a segregate
genus from Panicum or as a subgenus of Panicum will probably never be re-
solved, with both ‘camps’ having data to support their views. It is a debate that
I will not get into here, but will only provide references for both treatments, so
the reader can form their own opinion. Gould (1974), Clark and Gould (1975),
Gould and Clark (1978), Gould (1980) and Giussani et. al (2001) support the
recognition of Dichantheliumasa segregate genus. Whereas, Pohl (1980), Lelong
(1984), Zuloaga (1987), Morrone and Giese (1901), Zuloaga et al. (1991) and
Zuloaga et al. (1993) support the recognition of Dichanthelium as a subgenus
of Panicum; a classification with which | also agree.
Freckmann and Lelong (2002) published fifty-two new combinations in
Dichantheliumand Panicum while eee their treatment for Volume 25 of
The Flora of North America (Barkworth et al. [eds.], in press). Some of the taxa
treated in Dichanthelium by Freckmann and Lelong (2002) do not have an avail-
able name in Panicum, which are required for use in the Vascular Plants of Texas:
A Comprehensive Checklist including Synonymy, Bibliography, and Index Jones
et al. 1997), revision in progress, as well as, for on going work on Panicum subg.
Dichanthelium in North America. In addition, Panicum abscissum Swallen is
recognized at the rank of variety and not subspecies, following the treatment
of Panicum rigidulum by Lelong (1984), since no data were presented in Freckmann
SIDA 20(3): 1043-1046. 2003
1044 BRIT.ORG/SIDA 20(3)
and Lelong (2002) warranting the treatment of these taxa at the subspecific
rank.
Panicum aciculare Desvaux ex Poiret Var. ramosum a ) Wi fee comb nov.
BASIONYM: Panicum neuranthum Griseb. var. ramosum Griseb., Cat. pl. C es 2. 1866. Pani-
cum fusiforme Hitche., Contr. - S.Natl. Herb. 12222. 1909. Dichant ae forme (Hitche.)
Harvill, Castanea 42:177. 1977. Dichanthelium aciculare var. ramosum (Griseb.) Davidse,
Novon 2:104. 1992. ener a aciculare Desvaux ex Poiret subsp. fusiforme (Hitchc.)
Freckmann & Lelong, Sida 20:167. 2002. Typr: CUBA: western Cuba, 1863, C. Wright 3454
(HOLOTYPE : GOET, fragment US).
Panicum aciculare Desvaux ex Foret: vay neuranthom (Griseb.) Wipf ‘f,combet
Stat. NOV. BASIONYM: Panicum neuranthu seb., Cat. pl. Cub, 232. 1866. Dichanthelium
a 20:167
aciculare Desvaux ex Poiret subsp. ne ae Gri ise Bo Fredman & Lelong, Sic
2002. TypF: CUBA: eastern Cuba, 18600, C. Wright 3453 (HOLOTYPE : US).
Panicum acuminatum Sw. vat. ferventicola (Schmoll) Wipff, comb. et stat. nov.
BASIONYM: Panicum ferventicola Schmoll, Madrono 5:92. 1939; <- the autonym created by
Panicum ferventicola var. papillosum Schmoll, Madrono 5:94 and Panicum ferventicola
Schmoll var. sericeum Schmoll, Madrono 5:92. 1939. TyPr: U.S.A. WYOMING: Yellowstone Na-
tional Park, in fine gravel in warm soil close toa boiling spring, bathed in hot vapor, 5 mi N of
Norris Geyser Basin, LO Aug 1908, A. Chase 5252 (HOLOTYPE: G
enue ferve nticola oe var. sericeum Schmoll, Madrono 5:92. 1939. Dichanthelium
(Elliott) EW. Gould var. sericeum (Schmoll) Spellenberg, Madrono 23:150. 1975.
Dicheoiheltum acuminatum (Swartz) FW. Gould & CA. Clark var. sericeum (Schmoll)
Freckmann, Bae ae is Dichanthelium dacuminatum (Swartz) FW. Gould & CA.
Clark subsp. sericeum (Schmoll Freckmann & Lelong, Sida 20:168. 2002. Type: U.S.A. WyYo-
MING: in wet ane sai hot springs, Mammoth Hot Springs, Yellowstone National Park, 21
Jul 1889, A. Nelson & E. Nelson 6037 (HOLOTYPE: RM 20108).
Panicum - rventicola Schmoll var. papillosum Schmoll, Madrono 5:94. 1939. TypE: CANADA.
ALBERTA: Banff, on declivity wet with water from hot spring, 31 Jul 1914, A.S. Hitchcock HS5I1
No
(HOLOTYPE: G
Panicum acuminatum ae var. thermale (Bol.) Wipff, comb. nov. BasionyM: Panicum
thermale Bol. Proc. . Acad. Sci. 2:181. 1862. Dichanthelium lanuginosum (Elliott) FW.
Gould var. thermale Gal ) eanens Madrono 23:15]. 1975. Dichanthelium acuminatum
(Swartz) EW. Gould & CA. Clark var. thermale (Bol) See EA OLE)4 48:107. 1981.
nel acuminatum (Swartz) EW. Gould & Clark subsp. thermale (Bol.)
Freckmann & Lelong, Sida 20:168. 2002. TYPE: Us 5A. ne IFORNIA: ae part of Sonoma
County, on hot rocks and in hot water flowing from Geyser Springs and Geyser Mountains,
H.N. Bolander 3941 (HOLOTYPE: G
—
Panicum commutatum Schultes var. equilaterale (Scribner) Wipff, comb. et stat.
ao ees Panicum megane ae seribner, Bull. Div. Scene US.D.A. 11:42. 1898.
a tees y ril
SU ubsp : equilate rale (Scribner) | “reckmann &
alone Sida 20:169. 2002. TYPE: U. S.A. FLORIDA: Lake Co. scrubby feck lands, Eustis, 16-
25 Aug 1894, G.V. Nash poe LECTOTYPE: US 743929 (designated by Hitche. & Chase, Contr.
U.S. Natl. Herb. 15:310. 1910)].
Panicum ovale F|liott var. praecocius (Hitchcock & Chase) Wiplf, comb. nov.
BASIONYM: Panicum praecocius Hitche. & Chase, Rhodora 8:206. 1906. Panicum lanuginosum
Elliott var. praecocius (Hitche. & Chase) Dore in McNeill & Dore, Naturaliste Canad. 103:502.
1977. Dichanthelium villosissimum (Nash) R.W. Freckman var. praecocius (Hitchc. & Chase)
1045
Freckmann, Phytologia 39:270. 1978. Dichanthelium ovale (Elliott) FW. Gould & C.A. Clark
subsp. praecocius (Hitchc. @ Chase) Freckmann & Lelong, Sida 20:170. 2002. TyPrF: U.S.A.
ILLINOIS: Stark Co. dry bank, near Wady Petra, 30 Jun 1900, V.H. Chase 649 (HOLOTYPE: US
558686).
Panicum rigidulum Nees var. abscissum (Swallen) Wipff, stat. nov. Basionym: Pani-
cum abscissum Swallen, J. Wash. Acad. Sci. 30:215, f. 4.1940. Panicum rigidulum Bosc ex Nees
subsp. abscissum (Swallen) Freckmann @ Lelong, Sida 20:172. 2002. Type: U.S.A. FLORIDA:
Sebring, in dry sand at tourist camp near lake, 3 Oct 1925, P Weatherwax s.n. (HOLOTYPE: US
1259952).
ACKNOWLEDGMENTS
Iam grateful to K.N. Gandhi (GH) for his insightful review and comments re-
garding the nomenclature. | also would like to thank S.D. Jones (BRCH) and
two anonymous reviewers for their helpful suggestions and comments on the
manuscript.
REFERENCES
Barkworth, M.E., K.M. Carets, S. Lona, and M.B. Plep (eds.). In Press. Magnoliophyta:
Commelinidae (in part): Poaceae, part 2. Flora of North America North of Mexico, vol-
ume 25.Oxford University Press, New York.
Ciark,C.A.and F.W.GouLp. 1975.Some epidermal characteristics of Panicum, Dichanthelium
and Echinochloa. Amer. J. Bot. 62:743-748.
FRecKMANN, R.W. and M.G. Letona. 2002. Nomenclatural changes and innovations in Pani-
cum and Dichanthelium (Poaceae: Paniceae). Sida 20:161-174.
Giussanl, L.M., J.H. Cota-SANncHez, F.O. ZuLoaca, and E.A. Keto. 2001.A molecular phylogeny
of the grass subfamily Panicoideae (Poaceae) shows multiple origins of C4 photosyn-
thesis. Amer. J. Bot. 88:1993-2001.
Gout, FW. 1974.Nomenclatural changes in Dichanthelium (Poaceae). Phytologia 39:268-
GouLp, FW. 1980.The Mexican species of Dichanthelium (Poaceae). Brittonia 32:353-364.
Goutb, FW.and C.A.Ciark. 1978. Dichanthelium (Poaceae) in the United States and Canada.
Ann. Missouri Bot. Gard. 65:1088-1132.
Jones, S.D., J.K. Wiper, and PM. Montaomery. 1997. Vascular plants of Texas: A comprehensive
checklist including synonymy, bibliography, and index. Austin, University of Texas Press.
Letonc, M.G. 1984. New combinations for Panicum subgenus Panicum and subgenus
Dichanthelium (Poaceae) of the southeastern United States. Brittonia 36:262-273.
Morrone, O.and F.0. Zutoaca. 1991.Estudios morpholdgicos en el subgenera Dichanthelium
de Panicum (Poaceae: Panicoideae: Paniceae), con especial referencia a Panicum
sabulorum. Ann. Missouri Bot. Gard. 78:91 5-927.
POHL, R.W. 1980. Gramineae. In: Flora Costaricensis.W. Burger, ed. Fieldiana: Botany 4:1-608.
ZULOAGA, F.O. 1987. Systematics of the New World species of Panicum (Poaceae: Paniceae).
In: Grass systematics and Evolution, T.R. Soderstrom, K.W. Hilu, C.S. Campbell, and M.E.
Barkworth, eds. Smithsonian Institution Press, Washington, D.C. Pp. 287-306.
1046 BRIT.ORG/SIDA 20(3)
ZULOAGA, F.O., J. Dugcovsky, and O. Morrone. 1991. Infrageneric phenetic relations in New
World Panicum (Poaceae: Panicoideae: Paniceae): a numerical analysis. Canad. J. Bot.
71:1312-1327.
ZULOAGA, F.O., R.P. Etuis, and O. Morrone. 1993. A revision of Panicum subg. Dichanthelium
sect. Dichanthelium (Poaceae: Panicoideae: Paniceae) in Mesoamerica, the West Indies,
and South America. Ann. Missouri Bot. Gard. 80:1 19-190.
PERFORATION PLATE DIVERSITY IN ILLICIUM
FLORIDANUM (ULLICIACEAE) WITH RESPECT TO ORGANS,
PROVENANCE, AND MICROTECHNICAL METHODS
Edward L. Schneider and Sherwin Carlquist
Santa Barbara Botanic Garden
1212 Mission Canyon Road
Santa Barbara, CA e 05, U.S.A.
ABSTRACT
Pit membrane remnants in scalariform perforations of mature Illicium floridanum vessels of stems
and roots from three sources were studied with scanning electron microscope (SEM). Several meth-
ods of preservation and microtechnical treatment were used; results for all of these methods, for the
three sources, and for the two organs revealed similar patterns. The perforation pit membranes illus-
trated are believed to be relatively free of artifacts. Pit membrane presence in eee of all col-
lections of L fl Pa studied ranges from intact non-porose pit membranes to near-absence of
any pit membrane fragments. However, the extreme conditions are relatively infrequent, and the
most common condition is presence of less than 50% of the pit membrane; presence of more than
50% of the pit membrane is somewhat less common. The phylogenetic significance of pit membrane
presence in perforations is considered a symplesiomorphic feature for angiosperms. Presence of pit
membrane remnants in perforations is valuable in phylogenetic analyses of angiosperms.
Key Worps: basal angiosperms, Illiciales, perforation plates, pit membrane remnants, tracheid-ves-
sel transition, vessel evolution, wood evolution
RESUMEN
Se estudiaron al microscopio electronico de barrido (SEM) restos de las membranas de punteaduras
e las perforaciones escalariformes de vasos = ane y raices de Illicium fi oridanum eee
procedentes de tres fuentes. Se usaro ic vacion y
obteniendo patrones similares con ae los metodos, para las tres fuentes y los dos organos. Las
ee de las membranas de las | jue se ilustran se cree que no tienen artefactos.
a de punteaduras de Peete en las perforaciones de todas las colecciones de 1.
floridanum estudiadas varian desde punteaduras intactas no- eda BES la casi ausencia de
fragmentos de membrana. Sin embargo, las condiciones extremas te infrecuentes, y
las condiciones mas comunes son la presencia de menos del 50% de membrana de la punteadura; la
presencia de mas del 50% de la eee de la punteadura es algo menos frecuente. El significado
filogenético de la presencia de la membrana de la punteadura en las perforaciones se considera una
caracteristica simplesiomorfica en angiospermas. La presencia de restos de la membrana de la
punteadura en las perforaciones es valiosa en los analisis filogenéticos en angiospermas.
INTRODUCTION
Presence of pit membrane remnants in perforations of primitive woody dicoty-
ledons with scalariform perforation plates was figured by Meylan & Butterfield
(1978) for several species of New Zealand woods, suchas Ascarina lucida Hook f.,
Carpodetus serratusJ.R.&G. Forst., Quintinia acutifolia Kirk,and Weinmannia
SIDA 20(3): 1047-1057. 2003
1048 BRIT.ORG/SIDA 20(3)
racemosa L.{. Similar figures were presented by Takahashi (e.g., 1985). Meylan
and Butterfield (1978) refer to threadlike pit membrane remnants as “mi-
crofibrillar webs,” and in some cases this describes the appearances well. In oth-
ers (Ascarina, Weinmannia), the pit membranes look essentially intact; they
contain circular pores of various sizes. Pit membrane remnants ina simple per-
foration plate of Knightia (Proteaceae) were figured by Butterfield and Meylan
(1982).
Ina review of this phenomenon (Carlquist 1992a), the term “pit membrane
remnants” was proposed to describe the diversity of pit membranes or mem-
brane portions that occur as threads as well as those that occur as persistent
sheets in which component microfibrils are not evident. Pit membrane rem-
nants occur in a number of dicotyledon families (Carlquist 1992a) as well as in
some monocotyledons (see illustrations and literature cited in Carlquist &
Schneider 1997, 1998; Schneider & Carlquist 1998) and ferns (Carlquist &
Schneider 2001 and literature cited therein).
Pit membrane remnants appear characteristic of particular families, such
as Chloranthaceae (Carlquist 1990, 1992b, 1992c). Among the families that cur-
rently constitute Illiciales, pit membrane remnants are most conspicuous in
Illicium (Carlquist 1992b; Carlquist &@ Schneider 2002b), sole genus of
Illiciaceae, as well as in the monospecific family Austrobaileyaceae (Carlquist
2001) and both genera of Schisandraceae (Carlquist 1999). The survey of the
genus Illicium with respect to pit membrane remnants (Carlquist 1992a) showed
diversity but not the causes for this diversity. Conceivably, the sources lor dif-
ferent degrees and types of pit membrane remnant presence in Illicium can be
related to differences among species, differences in provenance, differences
among parts of a single plant, or differences in methods of preparation. The
present paper has selected one readily available species, Illicium floridanum
Ellis, because pit membrane remnants in perforations appeared abundant and
varied on the basis of preliminary observations (Carlquist 1992a), and because
material of living plants is readily available.
An understanding of the diversity of pit membrane remnants in Illicium
floridanum is needed as a baseline against which to compare diversity in such
remnants in other dicotyledons. Do pit membrane remnants indicate a primi-
tive character state in phylads of angiosperms as suggested by Carlquist (1992a)?
Can degrees of pit membrane presence be indicative of departure from trac-
heid-like tracheary elements, and if so, can these degrees be used cladistically
and can they help us understand the nature of the tracheid-vessel element tran-
sition (Carlquist & Schneider, in press)?
a
MATERIAL AND METHODS
The dried Illicitum floridanum stem (Palmer 19768; “Florida” without further
locality) was derived from a specimen 12 mm in transectional diameter in the
1049
RSAw collection. This specimen had been donated to Rancho Santa Ana Botanic
Garden by the USS. National wood collection (USw). A portion of this specimen
was boiled in water, stored in 50% aqueous ethanol, and sectioned on a sliding
microtome. Some of the sections were dried between clean glass slides, sputter
coated with gold and examined with an ISI WB-6 scanning electron microscope
(SEM) at Rancho Santa Ana Botanic Garden. Living stems (8 mm in diameter)
of the species were obtained froma cultivated plant in the lathhouse at Univer-
sity of California, Santa Barbara, and were preserved in 50% aqueous ethanol. A
stem 5 mm in diameter was obtained from the University of Florida, Gaines-
ville. Living roots of the species (10 mm in diameter) were provided by the US.
National Arboretum, Washington, D.C., and were preserved in 50% aqueous
ethanol. Root and stem portions from living specimens were preserved in alco-
hol without the use of any other reagents in order to minimize artifact forma-
tion in the pit membranes of perforation plates. Critical-point drying was not
employed because in mature woods, many vessels are already filled with air
and critical-point drying would be redundant (Butterfield @ Meylan 1982). The
stems and roots derived from living specimens were sectioned ona sliding mi-
crotome (only radial sections were prepared). Sections were dried between glass
slides, sputter-coated with gold, and observed with a Bausch @ Lomb Nanolab
200 SEM at Santa Barbara Botanic Garden. Thick sections (ca. 50 um) were cut
in order to obtain complete sheets extending from cambium to pith rather than
fragmentary sections. Thick sections offer no disadvantage where SEM is in-
volved. Thick sections, on the contrary, are more likely than thin sections to
have long portions of perforation plates or even entire perforation plates.
Low accelerating voltage was applied to the SEM-observed sections when-
ever possible in order to minimize the possibility of beam damage to the pit
membranes. We believe that tearing of pit membranes from sources other than
sectioning (the effects of which can easily be detected) was negligible. Our re-
sults seem comparable to the photograph of pit membranes for Weinmannia
racemosa (Meylan & Butterfield 1978). Current literature on wood anatomy does
not consider these appearances to be instances of artifact formation. In order to
stress the nature of pit membrane remnants, our illustrations are at high mag-
nifications. Our equipment does not offer maximal resolution compared to
newer machinery, unfortunately. Also, the nature of SEM involves lowered re-
trieval of electrons from depressed areas such as perforations, and thus resolu-
tion is less than for convex structures.
—
RESULTS
Perforation plates in Illicium floridanum (e.g., Fig. are scalariform and are
easily distinguished from lateral wall pitting, which consists of pits with rela-
tively short length, separated from each other by bands of wall material appre-
ciably wider than the bars of a perforation plate. Pit membranes of pits on lat-
1050 BRIT.ORG/SIDA 20(3)
Tk a
re ah ge
Fics. 1-4. SEM photographs of radial sections of Ilicium floridanum vessels. Figs. 1-3. HOrhlons of peaiayer plates
from dried stem wood sample. Fig. 1. About a quarter of a pertoranion pate) it remnants ts visible j in the
perforations Fig. 2. perforations sini to those of F Hg: , Showing i b g. 3. Perfora-
| febh Acin th £ H .
be J
an artitark tad I
y enn of wood empict or sectioning: Fig: bi Pits on lateral wall a vessels, showing abeene: of
7 ft t | y 4 5 ym.
1051
eral walls of vessel elements were intact and devoid of any pores (Fig. 4); nearly
all of the pit membranes in perforations have pores in a sheetlike membrane,
or threadlike or strandlike remnants of a pit membrane (Figs. 1-3, 5-11). We
have presented a photograph of an exceptional pit membrane from a perfora-
tion plate in which pit membranes that lack visible pores or holes were retained
(Fig. 12); such non-porose perforation pit membranes were infrequent, present
in fewer than 5% of the perforation plates we examined, with the exception of
the uppermost (or lowermost) perforation in some perforation plates. One or
two such terminal perforations, which may be considered transitional to lat-
eral wall pitting, may be seen in numerous perforation plates, but no such non-
porose perforations occur in the remainder of most perforation plates. Our com-
ments, unless otherwise stated, refer to perforation plates only, and describe
conditions other than the exceptional type of perforation pit membranes illus-
trated in Figure 12.
Pit membrane remnants are retained in perforations of the dried wood
sample that was boiled in water prior to sectioning (Fig. 1-3). The durability of
these membranes is attested by retention of seemingly delicate weblike mem-
brane portions (Figs. 1, 2). Also preserved by this method are strandlike pit
membrane remnants that run parallel to the long axis of the vessel element
(Fig. 3).
Study of wood from living specimens of stems fixed in 50% ethanol reveals
a wide range of pit membrane remnants in perforations (Figs. 5-8). Circular to
oval holes in pit membranes characterize numerous perforation plates (Fig. 5).
This type of pit membrane may coexist in a perforation plate with pit mem-
branes in the form of strands (Fig. 6). Some pit membranes in the form of web-
like remnants can be found (Fig. 7). Above in Figure 7 is a pit membrane frag-
ment containing pores; this illustrates the transition between porose sheets of
pit membranes and weblike pit membranes. In Figure 8, pit membrane rem-
nants are restricted to the ends of perforations. The remnants shown contain
small pores of various sizes and shapes. The perforations shown in Figure 8 are,
because of restriction of pit membranes to ends of perforations, mostly clear,
Perforations with small vestiges of pit membranes of this sort, but with most
perforation area clear of membrane remnants, characterized about 10% of the
perforation plates in this material. Also present in the liquid-preserved stems
from living plants were a few perforation plates in which pit membranes were
intact, like those shown in Figure 12. Such intact pit membranes are not from
immature vessel elements, since they can occur at various points between pith
and cambium.
Liquid-preserved roots from living cultivated plants also revealed a wide
range of pit membrane conditions (Figs. 9-12). In Figure 9, portions of two per-
foration pit membranes are illustrated. As suggested by the irregular distribu-
tion of pores and holes, pit membranes in this perforation plate possessed some
1052 BRIT.ORG/SIDA 20(3)
Fics 5-8 COM L rs L £ <. = 1 - J 1 £ j 1 fie £i. Bi
(plant culti Jat Uni ity of California, Santa Barbara) Fig. 5. Perforati itl | holes in pi I (up
per portion of holes obscured by oblique view of perforations). Fig. 6. Perforations with pit membranes containing
kK | 4 ee J ( pp p 4 £ } aan | kK J 4 y L li i W ff £ : 7 Perfora-
ra 4 Lye kKItl x L ie Z \ Wl L A Fidq Q dD £ =e 4
r r - r
porose remnants at the ends of the perforations. Magnificati i g 5 um.
1053
Fics. 9-12, SEM ee : ; ae : : eae i
IGS. if y t t J t ania ‘ ce Es
ti n f Lu tI J I n | WC RN TArl Fin.9 P } }
Lh e Tl i$ ! 1 Es by > e D*. L ral +b 4
have non Fig. 10
p
lar t | hol portions of pi I in the form of strands traversing the perforation. Fig. 11. Pit
+l I d nsetunlitn hat, £ ct ds Eia. 1? +i £ £ i | ill indi i
y g I I g I
membranes, | ibl Magnifi i 5pm.
1054 BRIT.ORG/SIDA 20(3)
areas apparently devoid of pores, whereas others had small pores and others
larger holes. A further stage in development of open areas within the pit mem-
brane remnants is represented in Figure 10. The areas apparently devoid of holes
in Figure 10 may result from inhibition of lysis, perhaps by adherence of some
overlying substance, a possibility observed in Ascarina (Carlquist 1990). In Fig-
ure 11, however, minimal inhibition of lysis is evident, and almost all of the pit
membrane remnants are in the form of delicate strands. Portions of an uncom-
mon type of perforation in which hydrolysis producing pores did not occur are
illustrated in Figure 12. As mentioned in the preceding paragraph, this type of
perforation plate is probably not immature, because perforation plates with
intact pit membranes can occur at any point between the cambium and the
pith. A few perforation plates with minimal pit membrane remnants were also
observed in the root material.
DISCUSSION AND CONCLUSIONS
Before examining the significance of the various manifestations of pit mem-
brane presence in perforations, the issue of possible artifact formation must be
addressed. Certainly tears due to sectioning or handling do occur, but rips in
pit membranes attributable to damage can easily be identified by their con-
tours, and are not mentioned in the above descriptions. The appearances we
describe are believed to represent naturally-occurring conditions. The living
stem and root portions were subjected only to dilute alcohol. The entirely in-
tact nature of lateral wall pits of vessels in these materials serves as a control,
certifying that application of this single reagent does not create artifacts. Di-
lute ethanol has never, to our knowledge, been claimed to create pores or holes
in pit membranes.
We did observe that prolonged exposure of perforation pit membranes at
high magnification and moderate to high accelerating voltage resulted in tear-
ing. This tearing, like that due to processing, is easily identifiable, in our opin-
ion. The pit membranes in perforations of Ilicium floridanum seemed to resist
deformation by the electron beam better than perforation pit membranes of
other dicotyledons we have studied, notably Amborella (Carlquist & Schneider
2001b).
A source of pit membrane destruction in perforations that is difficult to
estimate is the potential action of the conduction stream. One might attribute
distortions like tears to pit membranes altered by the conductive stream or by
formation of air embolisms.
Assuming that the appearances illustrated in the present paper are rela-
tively free from artifacts, how can one interpret the phenomenon of pit mem-
brane remnants in perforation plates? The distribution of dicotyledons with
pit membrane remnants in perforations (Carlquist 1992a) suggests that this isa
relictual feature, as though vessel elements were not definitively differentiated.
1055
The listing given in Carlquist (1992a), if compared to recent phylogenies, such
as that of Soltis et al. (2000), shows that pit membrane remnants are more com-
mon in the families of basal angiosperms, as well as in lower branches of eudicot
clades and monocotyledons (Carlquist & Schneider 1998; Schneider & Carlquist
1998) on the basis of admittedly incomplete data. Many additional instances of
pit membrane presence in perforations in dicotyledons have yet to be reported.
However, the rarity of pit membrane remnants in the woods sampled by Meylan
and Butterfield (1978) is likely indicative of the frequency of this phenomenon
if a large sampling of dicotyledons were to be surveyed.
Pit I dissolution in perforations is presumably an easily achieved
process, involving the action of enzymes. However, we do not know whether
this process can be phylogenetically reversed or not. Quite conceivably, a few
instances of pit membrane retention in perforations might be achieved second-
arily. The occurrence of pit membranes in perforations of some Myrothamnus
vessels might be such an instance (Carlquist 1988). However, restoration of pit
membrane remnants phylogenetically seems likely to be infrequent, an occur-
rence only in woods with scalariform perforation plates. Once a simple perfo-
ration plate has been achieved, retention of portions of a pit membrane in the
perforation plate seems unlikely, and indeed, apparently no cases of pit mem-
brane remnants in simple perforation plates have been reported.
Degrees of pit membrane presence in perforations have been cited as indi-
cating character state changes (Carlquist & Schneider 2002a. Five character
states are designated in that paper (O = pit membrane with no visible porosi-
ties; | = pit membrane with small porosities; 2 = pit membrane with less than
50% perforated; 3 = pit membrane present, but with more than 50% absent; 4 =
pit membrane absent or essentially so. All stem and root wood sections of II]i-
cium floridanum represented here cover the complete gamut from 0 to 4, in-
clusive. However, character states 0 and 4 are relatively infrequent, and could
be neglected if one were constructing a data matrix for this species. The major-
ity of perforation plates in I. floridanum would fall into character state 3, with
character state 2 less common. We see no Paes in designating the most com-
Lay
mon characte SlaALe for a specimen of a Pal ticular species, even if other character
states coexist with the most common manifestation. End wall angles of vessel
elements and number of bars per perforation plate are features with wide ranges
of expression within a single wood sample, yet they are frequently cited (often
in the form of means) as indicators of degree of phylogenetic specialization
ACKNOWLEDGMENTS
We thank Barbara Carlsward and William L. Stern, Department of Botany, Uni-
versity of Florida, Gainesville, for collecting stem and root material. Barron
Rugge, greenhouse manager at the University of California, Santa Barbara,
kindly permitted us to take root and stem portions from three cultivated speci-
1056 BRIT.ORG/SIDA 20(3)
mens. Thomas S. Elias, Executive Director, U.S. National Arboretum, Washing-
ton, D.C., graciously arranged for the excavation of a substantial portion of a
root system, which provided the large-diameter root used in this study.
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Araceae subfamily Philodendroideae. J. Torrey Bot. Soc. 125:253-260.
Sottis, D.E., PS. Soutis, M.W. CHase, M.E. Mort, D.C. Atgach, M. Zanis, V. SAVOLAINEN, WH. HAHN, S.B.
Hoot, M.F. Fay, M. Axtett, SM. Swensen, L.M. Prince, WJ. Kress, K.C. Nixon, and J.S. Ferris. 2000.
1057
Angiosperm phylogeny inferred from 185 rDNA, rbcL, and atpB sequences. Bot. J. Lin-
nean Soc. 133:381-461.
TAKAHASHI, A. 1985.Wood anatomical studies of Polycarpicae. |. Magnoliales. Sci.Rep. Osaka
Univ. 34:29-83.
1058 BRIT.ORG/SIDA 20(3)
BOOK NOTICES
RICHARD SPELLENBERG. 2003. Sonoran Desert Wildflowers. (ISBN 0-7627-1136-1, pbk.).
The Globe Pequot Press, 246 Goose Lane, PO Box 480, Guilford, CT 06437,
USS.A. (Orders: 800 826-8911, fax 800-820-2329). $24.95 pbk., 246 pp., nu-
merous color photos, 7 b/w figures, | map, | graph, 7" x 9".
From the back cover: Sonoran Wildflowers is the ultimate field guide to wildflowers of North
America’s most biologically diverse desert, which straddles the Gulf of California between the Baja
Peninsula and northwestern Sonora and stretches north into California and Arizona. Packed with
vivid color photos and informative text, this valuable reference will help you identify and appreciate
the varied flora of this vast region's six different climates.” The author's photographs are exception-
ally crisp and vibrant. Botanists and wildflower enthusiasts alike cannot go wrong with having this
book in their library.—Barney Lipscomb, Botanical Research Institute of Texas, 509 Pecan Street, Fort
Worth, 1X 76102-4060, U.S.A, barney@britorg.
EvizABeTH A. (BERNDT) GEorGE, watercolour paintings and illustrations by Mar-
GARET PIFRONI. 2002. Verticordia: The Turner of Hearts. (ISBN 1-876268-46-
hbk.). University of Western Australia Press (ww w.uwapress.uwa.edu.au),
Crawley, Western Australia 6009 in association with Australian Biologi-
cal Resources Study, GPO Box 787, Canberra, Australian Capital Territory
2601. (Orders: International Specialized Book Services, 5824 NW Hassalo
St., Portland, OR 97213-3644, U.S.A.; 503-287-3093, fax 503-280-8832,
wwwisbs.com, email: info@isbs.com). $84.95 hbk., 422 pp., 102 plates of
or paintings with line drawings, maps, 8 1/4" « 1114".
—
—
waterco
From the back cover: “Verticordia: The Turner of Hearts brings together for the first time available
information on all the described taxa of Verticordia 0] peal 13 subspecies and 30 varieties) pro-
viding a comprehensive, user- Buea) ae to their id é and ae Verticordia isa
member of the Myrtaceae family ith a description followed by information on
the distribution, habitat, flowering period distinguishing characteristics, other names used (syn-
onyms), cultivation, and conservation status. A key to infrageneric taxa and species is provided.—
Barney Lipscomb, Botanical Research Institute of Texas, 509 Pecan Street, ~ Worth, TX 76102-4060,
U.S.A, barney@britorg.
SIDA 20(3): 1058. 2003
COMPARISON OF THE MORPHOLOGY, FLOWERING
PHENOLOGY, AND LIFE CYCLE TYPE IN PLANTS OF
GRINDELIA LANCEOLATA (ASTERACEAE) FROM CEDAR
GLADES IN MIDDLE TENNESSEE AND
NORTHERN ALABAMA: A COMMON GARDEN STUDY
Christopher A.Adams! — Jerry M. Baskin Carol C. Baskin
Department of Biology Department of Bi ahs;
y Department of Biology
University of Kentucky University of Kentucky University of Kentucky
Lexington, KY 40506, U.S.A. Lexington, KY ie USA. Lexington, KY 40506, U.S.A.
and
Department of Agronomy
University of Kentucky
Lexington, KY 40546, U.S.A.
ABSTRACT
The life cycle type of Grindelia lanceolata Nutt. has been described as biennial, short-lived mono-
carpic perennial, and (polycarpic) perennial in the taxonomic literature. Plants of this species in
iddle Tennessee cedar glades clearly are monocarpic. However, field observations suggested that
those from glades in northern Alabama are at least dicarpic, and further that they differ morpho-
logically and Hower later han Hose in Tennessee glades.
The purpose of this study was to deter-
mine if diffe
phenology, and/or life cycle type of Tennessee and Ala-
bama plants are retained ie ee from sane in a “common garcen: -ie, ina honleated
greenhouse in Lexington, Kentuc rpholog -(
G. lanceolata plants from a ad Alabama were measure in the llowne (1) size oe both
osette and stem leaves; (2) number of secondary basal stems; (3) height of primary stem; (4) number
al capitula per plant; (5) number of ray and of disk flowers per capitulum: (6) diameter of aaualan:
and (7) length of ray flower corolla . Tennessee plants began flowering about | month earlier than
Alabama plants, and none of them produced basal rosettes a
‘ter they flowered once (in their 2"
year), confirming that they are strictly so a aa Alabama plants also flowered first in their 2”
year, however, 66% of them have produced basal rosettes (which bolted and flowered) for five con-
secutive growing seasons, confirming that an) are polycarpic. Also, individual Tennessee plants
potentially can produce twice as many seeds as individual Alabam
ing/fruiting period. Thes
a plants during a single flower-
ese results sonal suggest genetic differences exist in vegetative and floral
morphology, pLOWeHIE phenology, anh fe cycle type between paisa
lanceolata. We Id be associated wit]
origins: Pec plants from a monocarpic race in ae Ozarks and Alabama plants from a pcre
nial race in the Southwest.
e and Alabama peas oe -
<et
RESUMEN
El tipo de ciclo de vida de Grindelia lanceolata Nutt. ha sido descrito, en la esate taxondmica,
como bianual, como perenne monocarpico de vida corta y com renne (f I as plantas
‘Christopher A. Adams, corresponding author: Telephone (859) 257-3996; FAX (859) 257-1717; email:
caadam0@uky.edu
SIDA 20(3): 1059-1071. 2003
1060 BRIT.ORG/SIDA 20(3)
de esta especie presentes en clatos Sakis del area a Cenk al de Tennessee son claramente
monoc arpic as. Sin embargo obset ll d la zona norte ¢ de Ala-
bama son, sige menos, cane y mas alla, que ateena mor fologicamente y a f picaues mas tarde
]
| reos de Tennesssee. El proy si las
diecencias en mor ae fenologia floral y/o tipo " ciclo de vida de plantas de Tennessee y Ala-
be bama se mantienen ncuando las plantas crecen a pa tir se ibaa enun asus comun” - ie, en un
Lexington, Kentucky. |
plantas de G. lanceolata de Tennessee y de Alabama incluyen: (1) el tamano de
sins dlases de hojas, de roseta y de tallos; (2) numero de tallos basales secundarios; (3) altura del
tallo He (4) namero de capitulos por planta; (5) numero de flores radiales y de disco por
(6) tro del capitulo; y (7) longitud de la corolla de las flores radiales. Las a. de
a . s)
capitulo,
Tennessee comenzaron a [lorecer apr Speeaaens 1 mes mas temprano que las plantas de A
I qaber florecido una vez (en su 2‘
f] 2D de 0
bama y ninguna de ellas prodiyjo rosetas
bien
confirmando que son monocarpicz Las a de Alabama tam
ano; sin embargo, 66% de ellas siguio produciendo rosetas basales (que desarrollaron tallo y
florecieron) por cinco periodos de crecimiento, confirmando que son policarpicas. Ademas, cada
individuo de Tennessee puede es elegans el doble se semillas que los individuos de
labama durante cada periodo de ; : Los resultados saa claramente que
hay diferencias genéticas en la morfologia vegetativa y floral, la oo floral, y los Epos de ciclo
dev i entre ae de G. oaks eolatade peewee de Alabam
las plantas ae a nes-
see, de una raza monoeameies en ales Ozarks: y las plantas de Ala barr ma, d |
INTRODUCTION
Grindelia lanceolata Nutt. (Asteraceae) is an herbaceous species that grows in
open habitats on shallow soils underlain by limestone (Steyermark 1934, 1937,
Baskin and Baskin 1979). Its geographical range extends from the Ozarks of
Missouri and southeastern Kansas, south through eastern Oklahoma and north-
ern and western Arkansas, and into northeastern and central Texas (Steyermark
1934, 1999: Correll and Johnston 1970; GPFA 1986; Smith 1988; Nesom 1990).
Nesom (1990) also reports the species from the Monterrey area in Nuevo Leon,
Mexico. Disjunct populations occur in the Central Basin of Tennessee (Chester
et al. 1997) and in northern Alabama (Small 1933; Harper 1944), where they are
associated closely with open cedar glades (Baskin @ Baskin 1979, 1996; Baskin
et al. 1995). Grindelia lanceolata has been reported from Louisiana (Rydberg
1932; Small 1933); however, Gandhi and Thomas (1989) do not list the species in
their recent treatment of the Asteraceae of Louisiana. The species also has been
reported from a single county in southeastern Ohio Jones 1943; Fisher 1988),
where, apparently, it has been introduced (Porter 1956).
In his taxonomic treatment of Texas species of Grindelia, Nesom (1990)
recognized three varieties of G. lanceolata: lanceolata, texana (Scheele) Shinners,
and greenei (Steyermark) Nesom, the latter known only from Mexico. However,
Correll and Johnston (1970) did not recognize any separate taxonomic entities
and thus included only G. lanceolata in their treatment. Julian A. Steyermark
listed two forms of G. lanceolata in his Flora of Missouri (1999): lanceolata and
ADAMS ET AL 1061
latifolia Steyerm., the latter known from only one county in Missouri. Small
(1933), Fernald (1950), Gleason (1963), GPFA (1986), Gleason and Cronquist
(1991), and Chester et al. (1997) do not recognize any intraspecific taxa in G.
lanceolata.
With regard to life cycle type, G. lanceolata has been reported to be bien-
nial (Rydberg 1932), short-lived monocarpic perennial (Baskin & Baskin 1979;
GPFA 1986; Gleason & Cronquist 1991), and (presumably polycarpic) perennial
(Small 1933; Correll and Johnston 1970; Enquist 1987; Nesom 1990). In their study
on the autecology and population biology of G. lanceolata in the limestone ce-
dar glades of the Central Basin of Tennessee, Baskin and Baskin (1979) found
that plants were short-lived monocarpic perennials (ie., plants lived for a few
years before they bolted and flowered once, and then died). The youngest plants
to flower in the Baskins’ study were in their third growing season (2+ years old).
Other plants in the study flowered in their fourth or fifth year; all plants died
after flowering once (i.e, monocarpic).
However, during field studies in the limestone cedar glades in northern
Alabama, Baskin and Baskin (pers. obs.) noticed that Alabama plants of G.
lanceolata differed in morphology, flowering time, and life cycle type. In con-
trast to plants in Tennessee, rosettes were present on those with dead flowering
stalks in Alabama, suggesting that this species is at least dicarpic in the cedar
glades of northern Alabama. The purpose of this study was to determine if there
are distinct measurable or observable differences in vegetative morphology, flo-
ral morphology, flowering phenology, and/or life cycle type between G.
lanceolata plants from Tennessee and Alabama cedar glades. To determine if
differences in morphology and life cycle type are genetically—or environmen-
tally—based, both Tennessee and Alabama plants were grown from seed ina
common environment, ie.,a common garden experiment.
MATERIALS AND METHODS
Growth Conditions
All plants used in this study were grown from seeds in a nonheated greenhouse
in Lexington, KY. Seeds were collected in autumn 1996 from G. lanceolata popu-
lations growing in cedar glades in Tennessee and Alabama and sown (sepa-
rately) on soil in metal flats. The greenhouse soil mix was a 3:1 (v/v) mixture of
limestone-derived topsoil and river sand. Following germination, juveniles were
transplanted to 15-cm-diameter plastic pots in spring 1997 and assigned a num-
ber. Morphological and life cycle features of 103 Alabama plants and of 88 Ten-
nessee plants were monitored in this study; plants of both groups were num-
bered consecutively.
Temperatures in the nonheated greenhouse were recorded continuously
with an electric thermograph for the duration of the five-year study period.
From these recordings, mean daily maximum and minimum temperatures for
1062 BRIT.ORG/SIDA 20(3)
TABLE bs cowl mean monthly maximum (+ SE) and minimum (+ SE) temperatures (°C) to which
G ta plants were exposed during the study.
Month Maximum Minimum
January 695 + 14 1.71 + 0.38
February 10.95 + 088 400 + 0.72
March 1448 + 140 565 + 1.10
April 21.92 + 1.30 11.81 + 0.64
May 286 + 077 18.22 + 0.79
June 3062 + 046 209 + 049
July 3213 + 1.20 23.15 + 0.85
August 31.68 + 092 2205. = 0/1
September 28.18 + 095 17.75 + 0.99
October 20.05 + 0.35 126 + 0.68
November 156 + 1.10 8.23 + 1.00
December 85 + 140 3.03 + 0.60
each month for each of the five years were determined. These temperatures then
were used to calculate mean maximum (+ SE) and minimum (+SE) monthly
temperatures (‘C) for the study (Table 1). Average daily photosynthetic photon
irradiance at plant level in the nonheated greenhouse, measured with a LI-COR
model LI-1000 data logger and three LI-190-SA quantum sensors, ranged from
6 mol md! on overcast days to 25 mol md! on clear days during the grow-
ing season (March to October) (Snyder et al. 1994).
Vegetative and Floral Morphology
To determine if there are differences in vegetative morphology, various leaf char-
acters were compared between the two groups of plants. Length, width, oven-
dry mass, and specific leaf area were determined for leaves collected from the
rosette and from the lower-, middle-, and upper Gust below the terminal ca-
pitulum) portions of the main stem. One leaf each from these four regions of
the shoot was removed and measured/ weighed for every plant in the study. Leal
length and width were measured to the nearest mm using a standard metric
ruler. The width measurement was taken at the widest part of the leaf. Leaves
were dried in an oven at 70°C for 24 hours, and their dry mass determined with
an analytical balance. Average values for each leaf character were calculated
for all plants in the study. Leaf prints were made using Diazo-type paper for
ammonia developing, and leaf area (one side of leaf) was determined by weight
of paper/area print relationships. Specific leaf area (SLA) was calculated using
the following equation: SLA = Aleat es where Aleaf is leaf area (one side only)
and Weaf is leaf dry weight. SLA was determined for one leaf from each of the
four shoot levels for each plant in the study, and then average values were cal-
culated for leaves of each position. In addition, number of secondary basal stems
ADAMS ET AL., A 1063
and height of each plant were determined and averages calculated. All means
were compared by t-tests (P=0.05).
Number of capitula for the main stem and for all secondary stems were
counted and averages calculated for each plant in the study. Floral measure-
ments were made on 15 Tennessee plants and on 15 Alabama plants selected
randomly using a random-numbers table. Number of ray and disk flowers,
length and width of ray flower corolla, and diameter were measured, to the near-
est mm, for the terminal capitulum of the main stem of each of the 30 plants.
The terminal capitulum is always the first to flower on a plant. Capitulum di-
ameter and corolla length and width were measured using a standard metric
ruler and means calculated. All means were compared by t-tests (P=0.05).
Flowering Phenology
The terminal capitula of the main stems were monitored for flowering. The flow-
ering period in this study extended from the beginning of flowering (indicated
by the first anther of the first disk flower to shed pollen; ray flowers are pistil-
late) in the first plant and ended with the beginning of flowering in the last
plant. Beginning of pollen shed was determined by brushing a finger across the
anthers and observing if clumps of the bright yellow pollen adhered to it.
Seed Production Potential
To assess the reproductive effort of G. lanceolata, potential number of seeds per
individual plant was calculated. Potential number of seeds per individual for
Alabama and for Tennessee plants was calculated as follows: Potential number
of seeds produced per plant = (Avg. no. capitula per main stem x Avg. no. ray
flowers per capitulum) + (Avg. no. capitula per main stem x Avg. no. disk flow-
ers per capitulum) + (Avg. no. capitula per secondary stem x Avg. no. secondary
stems x Avg. no. ray flowers per capitulum) + (Avg. no. capitula per secondary
stem x Avg. no. secondary stems x Avg. no. disk flowers per capitulum).
RESULTS
Vegetative and Floral Morphology
Length, width, dry weight, and SLA of leaves from the rosette, and from the
lower-, middle-, and upper portions of the main stem, differed significantly
between Tennessee and Alabama plants of G. lanceolata (Table 2). Only rosette
dry weight and length of leaves on the lower portion of the stem were nonsig-
nificant. In general, plants from Alabama had longer, wider, and heavier leaves,
a higher SLA, and were taller than Tennessee plants. However, Tennessee plants
produced significantly more secondary stems than Alabama plants (Table 2).
Tennessee plants produced significantly more capitula per plant on both
main stems and secondary, basal stems than did Alabama plants (Table 3). Av-
erage number of capitula per main stem and per secondary stem was signifi-
cantly higher for Tennessee plants. Alabama plants produced larger capitula
1064 BRIT.ORG/SIDA 20(3)
Taste 2. Comparison (mean + SE) of several vegetative and floral morphological characteristics of
Grindelia lanceolata plants from Tennessee and Alabama. An asterisk indicates means for a charac
ter are significantly different (t test, P = 0.05), whereas NS indicates that they are not significantly
different at this level.
Character Tennessee Signif. Level Alabama
Rosette Leaf
Length (cm) 8.6 + 0.2 x 73 & 02
Width (cm) 0.7 + 0.01 09 + 0.02
Weight (g) 0.038 + 0.006 NS 0.045 + 0.003
SLA (cm?/q) 159.81 +1] 211.37 +14
Lower Leaf
Length (cm) 9.3 + 01 NS 96 + 02
Width (cm) 0.7 + 02 * L2 2 03
Weight (g) 0.039 + 0.007 : 0.063 + 0.006
SLA (cm?°/q) 173.53. +15 * 200.97 +
Middle Leaf
Length (cm) 78 + 0.2 ; 89 + 02
Width (cm) 0.6 + 0.01 - 1.3 + 0.02
Weight (q) 0.031 + 0.005 : 0.070 + 0.002
SLA (cm°/q) 115.84 ane, ‘ 14746 +
Upper Leaf
Length (cm) 6.3 + 0.1 * 70 + 02
Width (cm) 0.5 + 0.02 : 10 + 003
Weight (q) 0.0396 + 0.002 0.057 + 0.004
SLA (cm?/q) 12563 + 4 135.86 +10
No. Secondary Stems 3.1 + 02 : 14 0.1
Height (cm) 55.39 F 0.2 86.09 + Q.2
Ray Flower Petals
Length (cm) 1.45 + 0.1 s 2.05 0.3
Width (cm) 0.33 + 0.06 NS 0.30 + 0.04
(i.e., greater diameter) and more ray flowers per capitulum than did Tennessee
plants. Number of disk flowers, however, did not differ significantly between
the two groups. Ray flowers from Alabama plants had longer petals than Ten-
nessee plants, but there was no difference in petal width between the two groups.
Flowering Phenology
All plants from both groups bolted and flowered in their second year. Flower-
ing in Tennessee plants began on | July 1998, and all terminal flowers of main
stems had flowered by 5 August 1998 (Fig. 1). In contrast, flowering in Alabama
plants began and ended on 26 July and 7 September, respectively. For both groups,
capitula elsewhere on the main stem and on secondary stems continued to
ADAMS ET AL.. A TANCENI ATA 1065
TABLE 3. emp alleen of number of capitula and of ray and disk flowers produced by Grindelia
lanceolata plar \ts from Tennessee and Alabama. All values are e pressed as means (+ SE) except for
total number of capitula produced by all ae An asterisk indicates means of each character are
significantly different (t test, P = 0.05), whereas NS indicates that they are not significantly different
at this level.
Character Tennessee Signif. Level Alabama
Total No. Capitula Produced
Main Stem 629 408
Secondary Stems 271 53
No. Capitula Produced
Main Stem 7a 3 . 40 + 1
Secondary Stems 22 + 05 7 1d O38
No. Ray Flowers 22 + 4 . 3] - ak 3
No. Disk Flowers 203 + 34 NS 252 4+ 71
Capitulum Diameter (cm) 44 + 0.2 7 Sf OS
flower 1-2 weeks following the beginning of anthesis of the last terminal main
stem capitulum.
After completion of flowering, all Tennessee plants (n=88) died; nota single
one produced a new basal rosette. However, many Alabama plants continued
to produce new rosettes and to bolt and flower over the course of the five-year
study period (Figs. 1, 2). One-hundred and three Alabama plants bolted and
flowered in 1998, 78 in 1999, 74 in 2000, 73 in 2001, and 68 in 2002 (Fig. 2).
Flowering period duration was 43 days in 1998, 48 in 1999, 43 in 2000, 39 in
2001, and 44 in 2002 (Fig. L). Length of flowering period in 1998 was 36 days for
Tennessee plants (Fig. 1). Flowering (both terminal and all other capitula) for
both groups had been completed by mid-September.
Seed Production Potential
An individual Tennessee plant had the potential to produce an average of 3,128
seeds, while a single Alabama plant had the potential to produce only 1,568 seeds.
DISCUSSION
Observations by Baskin and Baskin that Alabama G. lanceolata plants appeared
to be distinct in morphology, flowering time, and life cycle type from those in
Tennessee were supported by the results of this study. Alabama plants are taller
and have larger leaves than Tennessee plants. One of the more striking differ-
ences between the two groups is the number of secondary basal stems. Tennes-
see plants produce more than two times as many secondary basal stems as do
Alabama plants, giving them a suffrutescent-like appearance. Indeed, Tennessee
1066 BRIT.ORG/SIDA 20(3)
2b st
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Fic. 1. Flowering phenology for 88 lia! lata plants in 1998 ( @ ) and for 103 Grindelia lanceolata
Alabama plants in 1998 ( # ),1999( © ),2000(. ), 2001 (4), and 2002 (©).
and Alabama plants can be distinguished easily on the basis of height and
branching pattern.
Average diameter of the terminal capitulum of Alabama plants is greater,
by more than lcm, than that of Tennessee plants. Compared to Tennessee plants,
terminal capitula of Alabama plants produce significantly more ray flowers
and longer ray petals. On the other hand, Tennessee plants produce twice as
many capitula per secondary stem and nearly twice as many (1.85:1) per main
stem as do Alabama plants.
—
ADAMS ET AL., NCEOLATA 1067
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Fic. 2. Fate of 103 Alabama Grindelia lanceolata plants grown from seed in a non-heated greenhouse. All plants that
produced rosettes in 1998 were in their second year of growth.
1068 BRIT.ORG/SIDA 20(3)
Number of seeds per plant produced during a single reproductive event
was considerably higher in the monocarpic perennial (“biennial”) plants (Ten-
nessee) than in the polycarpic plants (Alabama). Salisbury (1942) reported that
biennials in Great Britain produced an average of more than four times as many
seeds as did polycarpic perennials. Duffy et al.1999) predicted that monocarpic
species should maximize the number of meristems (i.e., branches) devoted to
reproduction, which would allocate as many reserves as possible to reproduc-
tive effort since there is only one reproductive event. Conversely, they predicted
that polycarpic perennials should maximize the number of growth meristems
(but not reproductive ones) since there is a fitness premium on longevity and
competitive ability. Tennessee plants produced 2.2 times as many secondary
basal stems, all of which flowered, as did Alabama plants. Asa result, Tennes-
see plants produced, on average, significantly more capitula and had the po-
tential to produce almost twice as many seeds as did Alabama plants. Thus,
Tennessee plants obviously devoted more energy to reproductive meristems
than did Alabama plants, which devoted more energy to growth in height and
perennation. Whereas all Tennessee plants died after flowering once, the ma-
jority of Alabama plants continued to flower each year for several years. By pro-
ducing fewer flowers, Alabama plants have reserve energy for production of new
rosettes (i.e., perennation), which also should be considered growth meristems.
Thus, there is a tradeoff between number of seeds produced per reproductive
event and lifespan of an individual.
A very interesting difference between Tennessee and Alabama plants is life
——
cycle type. In the nonheated greenhouse, both Tennessee and Alabama plants
bolted and flowered in their second year. However, whereas all 88 Tennessee
plants died after they reproduced once, only 13 of 103 Alabama plants did so.
Thus, 100% of the Tennessee plants, but only 13% of the Alabama plants, be-
haved as biennials. Another 12% of the Alabama plants were dicarpic, 12% were
tricarpic, and 1% tetracarpic, dying after their third, fourth, and fifth years, re-
spectively. Sixty-six percent of the original 103 Alabama plants have flowered
in their second, third, fourth, fifth, and sixth years, and it is expected that most
=
of them could survive and flower for the next several growing seasons.
Apparently, differences in morphology, flowering time, and life cycle type
between Tennessee and Alabama plants are genetically-based, since they were
maintained in plants grown from seeds in a common environment. However,
the advantage, if any, lor G. lanceolata behaving as a short-lived monocarpic
perennial in Tennessee glades, and of it behaving as a polycarpic perennial in
the cedar glades in northern Alabama, is not known. Floristically and
vegetationally, cedar glades of northern Alabama are very similar to those in
central Tennessee (Baskin et al. 1995; Baskin & Baskin 1999, in press). Further,
soils in both areas are limestone-derived, and climatic differences are minimal;
ADAMS ET Al 1069
both areas are within Képpen’s Cfa climate type (ie., mild temperate rainy cli-
mate without a distinct dry season, and with a hot summer) (Ackerman 1941).
Soil moisture and microclimate in cedar glades cause stress in plants in Ten-
nessee glades (Baskin @ Baskin 1999), and one would expect the same condi-
tions in glades in Alabama. Thus, there are no apparent physical or biotic dif-
ferences between Tennessee and Alabama glades that would seem to favor a
monocarpic life cycle in one area and polycarpic life cycle in the other. A recip-
rocal transplant study would help to determine if, in fact, plants are subjected
to different sets of selective factors on life history in the two areas, one favoring
monocarpy and the other polycarpy.
In terms of rand K life history strategy (e.g., Pianka 1970), data collected in
this study indicate that Tennessee plants are more r-selected than are Alabama
plants, which are somewhat further along the r-K continuum to being K-se-
lected. Thus, compared to Alabama plants, Tennessee plants are monocarpic,
short-lived, smaller in stature, and much more productive in terms of numbers
of seeds per reproductive event.
Could differences in geographic origin of Tennessee and Alabama plants
account for the differences between them? Plants in Alabama glades could be
disjunct from ancestor populations in southwestern United States, specifically
in Texas. Texas floras (Correll & Johnston 1970; Enquist 1987; Nesom 1990) con-
sistently describe the species as a (presumably polycarpic) perennial. This ge-
netic race may have spread from Texas into Alabama, and the two groups be-
came separated through geologic time. On the other hand, it seems likely that
plants in the middle Tennessee cedar glades are disjunct from short-lived mono-
carpic or biennial races (Rydberg 1932; Wetter 1986; Gleason & Cronquist 1991)
in the Ozark Region. Several other species, e.g., Evolvulus nuttallianus R.andS.,
Oenothera macrocarpa Nutt. subsp. macrocarpa, Onosmodium molle Michx. var.
subsetosum (Mack. & Bush) Cronquist, and Solidago gattingeri Chapm. are dis-
junct between the limestone glades of Missouri and those in Tennessee, and none
of these occurs in Alabama glades (Baskin & Baskin 1986; Bridges & Orzell 1986;
Baskin et al. 1995; Baskin & Baskin in press).
—
ACKNOWLEDGMENT
The authors sincerely thank Mrs. Adrianna Sautu for translating the English
version of the abstract into Spanish.
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sperms: Dicots. The Center for Field Biology, Austin Peay State University. Clarksville, IN.
Corrett, D.S. and M.C. Johnston. 1970. Manual of the vascular plants of Texas. Texas Re-
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Durry, N.M., S.P. Bonser, and L.W. Aarssen. 1999. Patterns of variation in meristem allocation
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GANDHI, K.N. and R.D. THomas. 1989. Asteraceae of Louisiana. Sida, Bot. Misc. 4.
GLEASON, H.A. 1963. The new Britton and Brown illustrated flora of the northeastern United
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Gteason, H.A.and A.Cronauist. 1991.The vascular plants of northeastern United States and
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1072 BRIT.ORG/SIDA 20(3)
BOOK NOTICES
LEONID AVERYANOV, PHILLIP CRIBB, PHAN Ke Loc, and NGuYen Tien Hier, watercolours
by Caro Woopn. 2003. Slipper Orchids of Vietnam With an Introduction
to the Flora of Vietnam. (ISBN 0-88192-592-6, hbk.). Timber Press, Inc., 133
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Plo
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nam in all its remarkable anaes The main text provides comprehensive accounts of the history,
nomenclature and rel shit each of the 22 slipper orchid species and natural hybrids founc
—_
—
within the country. It also provides the first detailed accounts of their habitats, biology and ecology.
Ona more urgent note it eo highlights the imminent threat of extinction faced by many of them.
Further urgent measures are necessary pom within Vietnam and abroad to protect wh at remains.”
One new section is d phioy i Aver. et PJ. Cribb sect. nov. (p. 13).
Guy and LiLiANr GUsMAN. 2002. The Genus Arisaema: A Monograph for Botanists
d Nature Lovers. (ISBN 0-904144-91-X, hbk.). A.R.Gantner Verlag
Kommanditgesellschaft, FL 9491 Ruggel. (Orders: Koeltz Scientific Books,
PO Box 1360, D 61453 Konigstein, Germany, +49 6174 93720, fax +49 6174
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merous color photos, line drawings, 6 1/4" x 91/2".
From the dust jacket: Since Engler’s classification in 1910, this monograph is the first attempt to bring
order to the confused state of knowledge about the genus in the w hole of its i ee It
includes descr iptions, taxonomy, namesa and synonyms, critic the nomenclature,
geographic distributions and habitats together w ith ; pen the most up-to-
date account of the genus Arisdema available today.”
/names and new nomenclatural combinations Jaa Section Anomala G. & L. Gusm.
sect. nov. (© 61); Section Arisaema G. & L. Gusm. stat. nov. (p. 95); Section Lobata G. &@ L. Gusm. sect.
nov. (p. 215); Arisaema elephas var. handelii (Stapl ex nie) G. & L. Gusm. var. nov. (p. 110);
Section Pedatisecta subsection Pistillata (Engl.) G.& L. Gus. stat. nov. (p. 233); Section Pedatisecta
see ee pedatie ecta (Schott) G. & L. Gusm. stat. nov. a tile Secon Sinarisaema dae
306)
)G.& L.Gusm. stat. nov. (p
(Gara) G. & L. Gusm. stat. nov. (p. 296); Section Tortuosa subsection Attenuata ae i" pd G. & L.
Gusm. stat. nov. (p. 365); Arisaema yunnanense var. aridum (H. Li) G. @ L. Gusm. var. nov. (p. 110).
SIDA 20(3): 1072. 2003
THE NAME OF THE AMARANTH: HISTORIES OF MEANING!
Mihai Costea? Francois J. Tardif
Department of Plant Agricult Department of Plant Agriculture
University orcas) ph University of Guelph
Guelph, Ontario, NIG 2W1, CANADA Guelph, Ontario, NIG 2W1, CANADA
coste_amihai@hotmail.com ftardif@uoguelph.ca
—
ABSTRACT
The cultivated grain and the weedy Amaranthus L. species (section Amaranthus) origi din the
Americas. The Tal lage conquered the New World in the loth century. Conceuenty the North
American species of Amaranthus could not have been introduced in Europe before this date. H
ever, the name of the eens existed long before its association with plants in the genus eee
by Linnaeus. ae everlasting plants were named “amaranth” by the ancient Greeks and Romans,
and in the medieval times. Among these, Helychrisum - and Celosid argentea L. are the most com-
mon. The name “Amaranthus” has had multiple meanings and ramifications through time: botani-
cal, ethnobotanical, religious, literary and ey This paper explores this history of mean-
ings of the name “Amaranthus.”
RESUMEN
Las especies oy adas como pseudocereales y las malas hierbas de Amaranthus L. (seccion
ldranthus lorigenen las Américas. Los espanoles congisae el Nuevo Munda enel
silodied iséis. C haber
produces en Europa euiS de esta Fecha om embargo, el nombre del Se nthos existid mucho
; Linneo. Varias plantas fueron denominadas
Fs
“amaranth” por los ee y romanos 3S antiguos, asi como en les tiempos medievales. Entre éstos,
Helycl hrisum spp. y Celosia argented L.f El nombre “Amaranthus”
ha tenido multiples significados yr fanai eaciones a través del tiempo: botanicos, etnobotanicos,
religiosos, literarios y filoséficos. Este articulo explora esta historia de significados del nombre
“Amaranthus.”
INTRODUCTION
The genus Amaranthus of Linnaeus includes species that originated in the
Americas, Africa and Eurasia. The association between these plants and the
name Amaranthus is relatively recent. American cultivated grain amaranths
(Amaranthus caudatus L., A. hypochondriacus L. and A. cruentus L.) and their
wild relatives (A. Eines L. one A. powellii S. Wats.) were introduced in Eu-
rope inthe lothcentury, after Sp Is conquered the New World (Sauer 1950,
1967). Other Amaranthus species, such as A. tricolor L. from Asia, and A. blitum
L. from Eurasia were also associated with this name in or after the loth century
es 1633; Thellung 1914). Yet, the name of the amaranth had been used by
the ancient Greeks and Romans almost two millennia before. Therefore, the
"Stat (rosa) pristina nomine/Nomina nuda terminus" U. Eco, The name of the rose, the final Latin hexamer.
2Correspnondina author
= b J
SIDA 20(3): 1073-1083. 2003
1074 BRIT.ORG/SIDA 20(3)
question is: what was the ancient amaranth and how did the meaning of this
name changed in time? An investigation into the history of the name
“Amaranthus” revealed multiple meanings and ramifications through time:
botanical, ethnobotanical, religious, literary and philosophical. This paper is
an attempt to tell the story of the name of the amaranth.
Etymology and Nomenclature
Amaranthus L. Sp. pl.:989. 1753. = Amaranthus Adans. Fam. PL. 1, 269, 516. 1763.
The ancient Greek name used by Dioscorides (4:57, Gunther 1959), dué&pavtog
(amarantos) came from the adjective &uépavtog, unfading (alpha privative,
not + maraino, fading), alluding to the everlasting nature of the flowers and
inflorescences. In ancient Greece, Amaryntus was a village in Euboea, where
Amaryntium, the temple of Artemis Amaryntia was situated (Strabo, The Ge-
ography 10:10, 12; 1959-1961). However, these names probably derived from the
same etymology. The Latin form adopted by Pliny the Elder (Natural History
21:41, 1951) was “amarantus.” Instances when the word was not associated with
a plant can be found in the Greek New Testament, | Peter 5:4, “ton amarantinon
tes doxes stephanon” (‘the unfading crown of glory”) and 1 Peter 1:4, “kai
amaranton” will not fade away”) (see Rienecker 1980). Lucianus Samosatenis
described a fresco painting of a flowery meadow as “eternal spring and unfading
(amarantos) (The Hall 9, 1949).
Later, Tournelfort (1694) stated that the name came from the alpha priva-
tive (@)—“not,” and the words “marceo” (the Latin equivalent of “maraino”)—“to
fade,” meaning “never fading” and “anthos” Gv00G)—‘f lower.” As Ray (1686-1688)
observed, the second analogy was forced: “Amarantus male cum ‘th’ scribitur
Amaranthus.” Thellung (1914) also noted that if the word “anthos” had been
incorporated in the etymology, the name should have been “Amarananthus”
and not “Amaranthus.” Both Amarantus and Amaranthus were used in the loth
and 17th century (reviewed by Sprague 1928). However, Linnaeus knew both
forms and deliberately rejected the classical spelling Amarantus in favor of
Amaranthus (Sprague 1928). The retention of the original Amaranthus spell-
ing should be followed even if it is less desirable philologically (Saint Louis Code,
article 60.1).
The Huauhtli.—Amaranths were called huauhtli by the Aztecs (see Wimmer
2003). We wanted to see if “unfading” served as an etymological root for huauhtli
in the same way it did for Amaranthus. “Huauh-” in the Nahuatl of the loth
century was an independent nominal radical, connected with no other root.
“Hudqui, to dry, would give the name “hudctli,” a dried out thing, word which
unfortunately could not be retraced to any old uto-aztecan roots (Wimmer, pers.
com.). The history and ethnobotany of huauhtli are fascinating since its rela-
tionship with man goes back more than 6000 years (Sauer 1950; Cole 1979). The
COSTEA AND TARDIF, THE NAME OF THE AMARANTH 1075
uses of huauhtli, and its social, religious and economic significance during the
Aztec Empire, are well known today (e.g. Sauer 1950, 1967; Cole 1979; Early 1992).
Huauhtli played an important role in many Aztec religious ceremonies. Huge
idols were molded from the dough prepared from huauhtli flour and popped
grains, which were then mixed with honey and sometimes with blood. The idols
were dedicated to several gods, among which the war god Huitzilopochtli was
the most prominent. The rituals involved human sacrifices and the Spanish
conquistadors tried to suppress the “pagan” culture by prohibiting the cultiva-
tion of the huauhtli (Sauer 1950; Cole 1979). Crop fields were burned and Aztec
people killed if they were found to possess the plant or its grains. Despite their
efforts, the conquistadors could not eradicate huauhtli and the plants endured
the time, awaiting their new name.
The Amaranth: Several Everlasting Plants.—The name of the amaranth pre-
ceded the introduction of huauhtli in Europe in the loth century. It also pre-
ceded the introduction of A. tricolor L. from Asia in the loth century. Another
Euroasian species, Amaranthus blitum L., was known in ancient Greece as “blite”
(Theophrastus Book 1. 14:2, Book 7. 1:2-3, 3:2, 3:4, 14:2, 1916; Dioscorides 2:143,
see Gunther 1959), and it became associated with the name Amaranthus only
in the 17th century (Thellung 1914). The ancient amaranth was an everlasting
plant used by the ancient Greeks for garlands of deity statues and in death ritu-
als. Dioscorides (4:57, Gunther 1959) mentionséuapavtog (amarantos) asa syn-
onym for elichruson (Helychrisum spp., most probably H. arenarium (L.)
Moench), an everlasting plant with yellow flowers (phyllaries) used by the
Greeks “to crown idols.” “The hair (the inflorescences) being drank with wine
doth help Dysuries, & ye bitings of serpents, & Sciaticas, & ruptures, & it moves
ye menstrua, & consumes the clots of blood which are in ye bladder or ye belly,
being drank with mustum” (Dioscorides 4: 57, Gunther 1959). The yellow ama-
ranth mentioned several times in Vatsyayana’s Kama Sutra, may also be
Helychrisum (Burton 1963).
During the Roman Empire, Pliny the Elder described “Amarantus” as
“autem spica purpurea verius quam flos aliquis’ (Natural History 21:41, 1951).
The description indicates that the amaranth at that moment was probably
Celosia argentea L.,an Amaranthaceae from Asia and Africa that resembles
Amaranthus spp. in many respects. Indeed, the author distinguished the ama-
ranth from Helichrysum (21:168), and the latter retained all the characteristics
and uses previously mentioned by Theophrastus and Dioscorides. However, it
should be noted that the name of the amaranth was again associated with an
everlasting plant. The amaranth referred to by Ovid (4:435, 1931), Tibullus
Albius (3.4.33, 1962), Plutarch (Questiones convivales 3.13.648A, 1949) and Col-
umella (10:175, 1941) was also C.argentea. In the 15thand loth century, the name
Amaranthus (or Amarantus) was applied equally to Helychrisum spp., to the
1076 BRIT.ORG/SIDA 20(3)
amaranths introduced from the New World, to C. argentea, and even to
Gomphrena globosa L., another Asian Amaranthaceae. For example, in 1542
Fuchs described and illustrated two amaranths (“flos-amoris”): “Duplex est
Amarantus: unus luteus (Amarantus luteus = “Helychrison” = Helychrisum
arenarium)|..|, alter purpureus” (Amarantus purpureus = C. argentea) (De His-
toria Stirpium 98-101, 1542). Matthioli’s illustration of the amaranth from 1563
(in Krutch: 128, 1976; Fig. isalso probably C. argentea. Henry Fletcher, a painter
active in London around 1730, in a reproduction of a Pieter Casteels’ painting,
identified a plant of C. argentea as “yellow Amaranthus” (see Segal 1990).
“Floramore” or “Flour Amore” was equally a popular name for Amaranthus tri-
color, for Celosia argentea (= Amaranthus purpureus, Gerard 1633; Henry 1829,
Fisher 1998) and for the real amaranths “Amaranthus Pannicula Sparsa,” Ger-
ard 1633; see also Meyer et al. 1999). The confusion is understandable.
Amaranthus and Celosia are similar in many respects, and North America was
thought initially by Christopher Columbus to be a part of Asia. It is difficult to
ascertain who was the first botanist to distinguish the two genera and who was
the first to associate the name with today’s amaranths. The distinctive in
florescence of A. caudatus was described by Parkinson (1640. 753) and Ray
(1686-1688, |: 202). Although Ray did not mention Celosia, his “Amarantus” was
classified accurately in the group with “herbae florae imperfecto sexu carentes
sunt vel femine” (Methodus Plantarum, Tab. 5, 1684) (Celosia has bisexual flow-
ers). It was Linnaeus who legitimized the differences between the two genera
(Celosia Sp. Pl. 205. 1753; for Amaranthus see above).
The Amaranth: A Magical Plant.—Theophrastus, who preceded Dioscorides by
approximately 300 years, did not mention Amaranthus. He wrote about the
“magical properties” of the gold-flower (Helychrisum spp.), which was capable
of bringing fame and wealth to those who wreathed themselves with the flow-
ers sprinkled with an “unguent from a vessel of unfired gold” (Book 9, 19:2-4,
1916). However, Theophrastus was skeptical about “such tales ... (which) pro-
ceed from men who desire to glorify their own crafts.” Athenaeus of Naucratis
cited Theophrastus, but in a credulous tone, affirming that the plant really is
magical and capable of bringing fame and fortune (15:680, 1955-1961). Medi-
eval herbalists followed Dioscorides’ synonymy and transferred the virtues of
the “helichruson” to the amaranth. Joannes Ammonius Agricola called ama-
ranth “flos honoris’ (Medicinae Herbariae:35, 1539) because the flower was ca-
pable of bringing honor. Since “its flowers (are) dedicated to the gods and rarely
to living men,” dreaming garlands of amaranth (was) a bad sign for sick people
but a good one “for everyone, especially for people involved in lawsuits”
(Artemidorus Daldianus 1:77, 1975). English colonists sailing for America wore
magical amulets containing amaranth seeds to protect them (Cole 1979). Later,
in the L9th century Americans believed that the amaranth could attract light-
COSTEA AND TARDIF, THE NAME OF THE AMARANTH 1077
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Fic. 1.“Amaranthus” from Matthioli 1563, probably Ce/osia argentea L.
1078 BRIT.ORG/SIDA 20(3)
ning (Cole 1979). Today, hundreds of Web pages and magic books recommend
amaranth as the flower of “protection, healing and invisibility” (e.g. Cunning-
ham 1993a, b). Wearing a wreath of amaranth speeds healing (even of a broken
heart) and grants invisibility. Fragments of an amaranth plant, collected on
Friday night under a full moon, can even protect you against bullets... We won-
der what Theophrastus would say to this?
The Amaranth and the Types of Immortality.—“Eternity points, in its amaranth
bower” (Shelley, Bereavement, L901). The quest for eternity and immortality is
one of the major inquietudes of the human spirit. Physical or spiritual immor-
tality, everlasting love and friendship, eternal happiness and virtue are differ-
ent coordinates of the same obsessional geography of ideas. Amaranth was
chosen as a symbolic flower to capture all of these meanings. That is why an
exploration into the meanings of the amaranth name is at the same time a tax-
onomy of the types of immortality.
Immortal youth (life).—The first metamorphosis of the amaranth from an
everlasting plant into an immortality symbol can be found in one of Aesop’s
fables (the Rose and the Amaranth, 384, 1975). The Amaranth, envious of the
“beauty and perfume” of the Rose receives the following reply: “I indeed, dear
Amaranth, flourish but for a brief season! If no cruel hand pluck me from my
stem, yet | must perish by an early doom. But thou art immortal and dost never
fade, but bloomest for ever in renewed youth” (Aesop, Fables: The rose and the
amaranth, 384, 1975).
Immortal hero and everlasting virtue —At the funeral of Achilles, Thetis
and the Muses spread “immortal weeds’ on his grave (Homer, “Odyssey” 24:43-
29,1955). According to more recent sources that we could not trace in classical
authors, the Thessalians placed amaranth on the funerary monument of Achil-
les (Thylesius Antonius 1531) or wore amaranth crowns at his funeral (Genlis
1810-1811). For Cowper, “the only amaranthine flower on earth is virtue” (The
Task, Book 3: The garden, 1874) and for Sainte-Beuve the amaranth isa “symbol
of virtue that never fades” (Causeries du lundi, vol. 8:142, 1926 - 1947).
Immortal love and hope.—The following verses of Gombault were taken
from Henry’s Flora Historica (1829): “Je suis la fleur d’amour qu’amarante on
appelle/ Et qui vient de Julie adorner les beux yeux./ Roses, retirez-vous, j'ai le
nom d’immortelle,/ Il w appartient qu’d moi de couronner les dieux.” The old
Victoran name of A. caudatus, “love-lies-bleeding,” is in direct relation with the
shape and the blood red color of the inflorescence. “Hope without an object can-
not live” and the amaranth is for Coleridge a symbol of eternal hope (Work with-
out hope, 1972
Immortal soul.—Although Pliny’s description of the amaranth was utili-
tarian, his words were later interpreted ina symbolic or mystical way: “mireque,
postquam defecere cuncti flores, madefactus aqua revivescit et hibernas coronas
—
—~—
COSTEA AND TARDIF, THE NAME OF THE AMARANTH 1079
facit.summa natura eius in nomine est, appellati, quoniam non marcescat” (Natu-
ral History 21:41, 1951). In the Roman Empire of the second century AD, the
custom of wearing wreaths was regarded as a pagan practice by Tertullian (1992)
and Clement of Alexandria (1965-1983), a sacrilege to the crown of thorns worn
by Christ. For Clement, the amaranth became an imaginary immortal flower,
which grew only in Paradise and was offered as an eternal reward to good Chris-
tians. “The earth is not capable of producing this flower; heaven alone knows
how to grow it” (Pedagogos 2.8.73, 1965-1983). As Irwin (1990) pointed out:
“Clement... was certainly not simply transplanting an actual earthly flower to
heaven.” He was adding two new dimensions to Aesop’s symbol of immortality:
the spiritual and the sacred. More than 400 years after Clement, Milton envi-
sions the same heavenly amaranth, but his vision of the “immortal amaranth”
is purely metaphorical:
“With solemn adoration down they cast
Their Crowns inwove with Amarant and Gold,
Immortal Amarant, a Flour which once
In Paradise, fast by the Tree of Life
Began to bloom, but soon for mans offence
To Heav’n remov’d where first it cae there grows,
And flours aloft shading the Fount of Life,
And where the river of Bliss coud midst of Heaven
Rowls o’re Elisian Flours her Amber stream”
(Book H1, 351-360, and see also XI, 77-81, 1948).
Another, less solemn, type of heavenly amaranth can be found in Charlotte
Bronté: “Happiness is a glory shining far down upon us, out of heaven. It is a
divine dew which the soul, on certain of its summer evenings, feels dropping
upon it from the amaranth bloom and golden fruitage of Paradise” (Bronté,
Villete, Chapter 22, The letter, 1984).
acred amaranth.—In the 13th century, the name became an attribute of
the Virgin Mary, a symbol of immaculate immortality: “Amaranthus flos, sacro,
qui non marcet, honore vigens” Johanes Germanus 1460 in Marraccio 1693). In
a painting by Botticini, amaranth plants grow from the ruins of the palace of
David where Madona adores the sacred child (D’Ancona 1977). In Southern
Europe, amaranth was used to decorate churches on Ascension Day (Skinner
1925; Cole 1979), and some of the popular names included “the scourge of our
Blessed Lord” (Dowling 1900) and “Discipline de Religieuse” (Henry 1829). It
should noted that blood symbolistic, divine or not, is a convergence point be-
tween the Aztecs and the Old World.
Hermetic amaranth.—In 1653, Christina, Queen of Sweden, founded the
hermetic Order of the Amaranth (Akerman 1991). The emblem consisted of an
insignia with a double A, a crown of amaranth, and the inscription “dolce in
nella memoria.” Henry (1829) mentioned the above inscription as appearing
ona medal with an Amaranth in enamel. The exact type of immortality sought
1080 BRIT.ORG/SIDA 20(3)
by Christina can only be speculated: catholic ideal of virginity, everlasting and
imperishable glory or immortal mystical bond (see Akerman 1991). At least one
of the aims was achieved because the Queen is still “dolceinnella memoria.” The
emblem and the name have been embraced by the current Masonic Order of
the Amaranth (Order of the Amaranth 1994).
A Plant or a Symbol?—*..Sad Amaranthus, in whose purple gore/...To whom
sweet Poets hath given endlesse date” (Spenser, The Faerie Queene, book II, canto
VI, 4060-408, 1976). Hundreds of writers, poets and sometimes even musicians
have carved and deepened (or occasionally weakened) the name of the ama-
ranth. Some of them are probably not aware that the amaranth is an earthly
plant and not an abstract sign or symbol. This particular case illustrates the
disparity between the symbolism and the reality of many names. The dispute
over the presence of an existential relationship between a name and the named
thing recurs from Plato's Cratylus to modern semiotics. Whether the name isa
simple convention (as Hermogenes believed in Cratylus, Plato, 1926), a sign that
signifies (Wittgenstein, Tractatus logico-philosophicus:3, 1992), or more than
rom plant to plant
Pty
that is not for us to determine. This name traveled in time
until it found its perfect mach. Grain amaranths are “from the past for the fu-
ture” (Cole 1979) and as weeds they will be “a per petual source of trouble to
farmers” (Macoun 1883). At the same time, the meanings conveyed by the name
have probably changed our perception of the plants. Perhaps the amazing his-
tory of the huauhtli plants in the New World would have faded away without
the unfading symbolism of their new name. One thing is certain: the amaranth
will continue to grow, here or in Paradise as long as we do, or perhaps even longer.
ACKNOWLEDGMENTS
This paper would not had been written without the encouragement of Barney
Lipscomb, for which we are deeply grateful. David Brenner, Susan Weaver, Me-
lissa Wheeler and anonymous reviewer provided interesting observations,
which improved the quality of the manuscript. Alexis Wimmer helped us with
the Nahuatl language.
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1084 BRIT.ORG/SIDA 20(3)
BOOK NOTICES
JOHN G. SEMPLE, STEPHEN B.LHEARD, and Luc BrouiLLet. 2002. Cultivated and Native
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jesemple/uwbioserhtm). $20.00, 134 pp, 61 plates/line drawings, 8 3/8" x
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Fighteen new nomenclatural combinations are proposed.
Ropert R.HaAyNes and a 1TZ B. HOLM-NicLsen. 2003. Potamogetonaceae. (ISBN
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SIDA 20(3): 1084. 2003
DOCUMENTED CHROMOSOME NUMBERS 2003: 1.
CHROMOSOME NUMBER OF TOXICOSCORDION NUTTALLII
(LILIALES: MELANTHIACEAE)
AND CLARIFICATION OF THE GENUS
Wendy B. Zomlefer
Department of Plant Biology
University of Georgia
2502 Plant Sciences
Athens, GA 30602-7271, U.S.A.
ABSTRACT
The mitotic chromosome count for Toxicoscordion nuttallii (2n = 22), reported here, matches the
number formerly documented for other species in the genus. A previous unvouchered report of 2n =
32 for this species, therefore, likely represents a misidentification, and a base number of x = 11 is
synapomorphic for Toxicoscordion. An updated overview of this recently resurrected segregate ge-
nus is given.
RESUMEN
El recuento d m as mitoticos en Toxicoscordion nuttallii (2n = 22), citado aqui, concuerda
con el ntmero documentado anteriormente para otras especies del género. Un re e previo, sin
catalogar, de 2n = 32 para esta especie, probablemente representa una faces oe y el
numero base de x = 11 es sinapomorfico para Toxicoscordion. Aqui se presenta una revision de este
* 4] : . ]
[o)
Tribe Melanthieae (Liliales: Melanthi ) comprises seven genera (ca. 62-93 spp.)
of predominately woodland and/or alpine perennial herbs occurring mainly
in the temperate to Arctic zones of the Northern Hemisphere: Amianthium (1
sp.), Anticlea (9-11 spp.), Schoenocaulon (24 spp.), Stenanthium (2-3 spp.),
Toxicoscordion (8 SPP). Veratrum s.l. (7-45 spp.), and Zigadenuss.s.(1sp.). These
generic ptions (most novel) are supported by recent parsimony analy-
ses of trnL-F Pea DNA and ITS (nuclear ribosomal) sequence data (Zomlefer
et al. 2001, 2003). A significant consequence of the molecular studies was the
reassessment of the traditional Zigadenus s.L,a poorly defined assemblage with
a complex taxonomic history (summary in Zomlefer 1997). Although several
segregates of Zigadenus have been described, contemporary treatments (e.g.,
Schwartz 2002) have typically accepted only the monotypic segregate
Amianthium with the remaining ca. 19 species maintained in Zigadenus sl.
Based on these molecular data, however, Zigadenus s.l. is polyphyletic and forms
five strongly supported clades that correlate with certain geographical distri-
bution, morphological characters, and chromosome number.
SIDA 20(3): 1085-1092. 2003
1086 BRIT.ORG/SIDA 20(3)
One well-defined clade corresponds to the genus Toxicoscordion, originally
described by Rydberg (1903), and recognized as a distinctive taxon by both
Preece (1956) and Schwartz (1994) as Zigadenus sect. Chitonia (R.A. Salisbury)
J.G. Baker. This Zigadenus segregate comprises ca. eight species (Table 1) re-
stricted to the midwestern United States to western North America (Fig. LA)
and includes the well-known poisonous “death camas” plants of the rangelands
suchas T. nuttallii, T. paniculatum, and T. venenosum (see Marsh etal. 1915, 1926).
The resurrection of Toxicoscordion required one new combination (T. fontanum),
a species described within Zigadenus by Eastwood (1937), long after Rydberg
(1903) had circumscribed the genus and had transferred the appropriate con-
temporary Zigadenus species. Ioxicoscordion is defined by the morphological
synapomorphies of conspicuously clawed tepals (especially the inner three)
and one obovate gland per tepal (Fig.1B; Zomlefer et al. 2001; Zomlefer & Judd
2002). Although a discrete genus, specific and infraspecific limits are unclear,
especially within the variable T. micranthum-T. fremontii species complexes
and the intergrading I. paniculatum-T. venenosum. For example, in an isozyme
study of these species (Schwartz 1994), the monophyly of T. paniculatum is
suspect because the five sampled populations were separated from each other
in UPGMA cluster analyses and distance Wagner trees. These results warrant
further examination of the species complexes in the genus.
The chromosome number 2n = 22 (or n= 11) has been reported for all spe-
cies of Toxicoscordion (Table |; Fig. 2) except for one anomalous report of 2n =
32 for T. nuttallii. Since chromosome number is a significant and likely an in-
variable apomorphy for genera of tribe Melanthieae (e.g., Zomlefer and Smith
2002), 22 is the predicted mitotic count for all Zigadenus species now trans-
ferred to Toxicoscordion. This study challenges the earlier conflicting T nuttallii
report, thereby seeking to validate the diploid number of 22 as a consistent
synapomot! phy for the genus.
MATERIALS AND METHODS
Several bulbs of Toxicoscordion nuttallii were collected during April 2002 at
two localities in south-central Texas by J. Spangler and T. Wendt (see Table 2)
and transplanted to pots maintained at the Plant Biology Dept. Greenhouse
Facility at the University of Georgia. Dividing root tip cells were prepared lor
examination with some modification of the general protocols outlined by Flory
and Smith (1980) and Jones and Luchsinger (1986). Once the plants were well-
established, actively growing root tips were harvested at 8:00 AM and treated
with 0.2 % colchicine for + hours, rinsed in distilled water, and then fixed in
Carnoy’s solution G ethanol: | acetic acid) overnight or longer. Following this
fixation, the roots were rinsed in distilled water and stored under refrigeration
in 70% ethanol. The material was subsequently hydrolyzed in 1.0 N HCl at 53°
C for 5 minutes, rinsed several times again with water, and treated with 45%
ZOMLEFER, TOXICOSCORDION CHROMSOME NUMBER 1087
ENTRAL
NORTH AMERICA
Toxicoscordion
V
Fic. 1. Distinctive characteristics of Toxicoscordion.A.G | distributi p of I ( f 1940;
Preece 1956; Schwartz 1994, 2002; M. H. MacRoberts pers. comm.). The midwest lisj ge ( Oklahoma
Kansas—Missouri—Arkansas—Louisiana) comy he localities of Toxi li B. Adaxial surface of an out
(left) and an inner (right) tepal from 7. nuttallii (Rollins 5334, GH), showing the generi I phies: claw plus a
H +, | dA Nachad alli fil A : Th | £ +h * 1 eed | 1
than those of the outer whorl.
glacial acetic acid for 10 minutes. The growing tips were then removed, macer-
ated with a dissecting spatula ona glass mi pe slide, and stained with 1 %
aceto-orcein. After application of a cover slip, the slide was gently heated with
an alcohol lamp, placed between blotters, and subjected to additional pressure.
Slides were mounted in euparal for future reference. Well-spread metaphase (and
late prophase) chromosomes were traced under a Leica DMLB Research Micro-
scope with a camera lucida attachment. Herbarium specimen vouchers (Table
2) are deposited at GA and TEX.
RESULTS AND DISCUSSION
The mitotic chromosome number of 2n = 22 for Toxicoscordion nuttallii is con-
firmed with plants from populations in Hays and Williamson Counties, Texas
(Fig. 2E, F; Table 2). A citation of 2n = 32 for T. nuttallii, originally reported by
Zakharieva and Makushenko (1969) and subsequently cited in various indices
and floras (e.g., Fedorov 1969; Moore 1971, 1973; Churchill 1986; Yatskievych
1999), was based on an undescribed and unvouchered plant then growing at
the Munich Botanical Garden (original source not cited). This “Zigadenus” spe-
cies was likely an Anticlea, another Zigadenus segregate, but one character-
ized by x =8 (Zomlefer & Judd 2002).
Chromosome number is a useful taxonomic character for genera within
tribe Melanthieae, especially the synapomorphic 2n numbers of 20 for
1.The species of Toxicoscordion and their mitotic (2n) and/or meiotic (n) chromosome numbers. The single citation for 7. nuttallii, 2n = 32, is inconsistent
can reports for all other species in the genu
Taxon Chromosome Secondary Citation(s) Original Source(s)
Number in Indices and Floras
n
T. brevibracteatus (M.E. Jones) 11 — Cave (1960); Fedorov (1969); Moore Lewis (1959): Cave (1970)
R. Gates (1972, 1973); McNeal (1993)
T. exaltatum (Eastwood) A. Heller 11 — McNeal (1993) ?
T. fontanum (Eastwood) 1] — Moore (1972, 1973); McNeal (1993) Preece (1956, as Zigadenus venenosus
Zomlefer & Jt var. fontanus); Cave (1970)
T. fremontii (J. Torrey) Rydberg Wo— Darlington & Wylie (1956); Fedorov Miller (1930); Preece (1956, as Zigadenus
(1969); Moore (1972, 1973); McNeal fremontii var. fremontil); Cave (1970, as Z.
(1993) fremontil var. fremontii and var. inezianus)
T. micranthum (Eastwood) A. Heller 1 McNeal (1993) Preece (1956, as Zigadenus venenosus var.
T nuttallii (A. Gray) Rydberg _
=
T. paniculatum (Nuttall) Rydberg 1]
T. venenosum (S.Watson) Rydberg 1]
Fedorov (1969); Moore (1971, 1973);
190 \
ei /
Churchill (1986); Yatskievych (
McNeal (1993
Moore (1972, 1973); Goldblatt
(1981); Churchill (1986); McNeal
(1993)
peas
micranthus)
Zakharyeva & Makushenko (1969)
Preece (1956)
Preece aon as Zigadenus venenosus var.
gramineus and var. venenosus); Cave
(1 a ane & Taylor (1977, as Z.
venenosus var. gramineus); Hartman &
Crawford (1971, as Z venenosus var.
gramineus)
880L
(€)0Z VaIs/9¥O'LINd
ZOMLEFER, TOXICOSCORDION CHROMSOME NUMBER 1089
10 um t~
Fic 2. Summary illustration of previously documented meiotic (n = 11) and/or mitotic (2n = 22) chromosomes of
Toxicoscordion (A—D, G-l) and the new reports for 7. nuttalfii (E, F:2n = 22).A. 7. brevibracteatus, diakinesis (n = 11). B. 7.
fontanum, anaphase II (n = 11).€. . fremontii, metaphase | (n = 11).D. 7. micranthum, metaphase | (n = 11).E. 7. nuttallii
(Wendt 7246, TEX), metaphase (2n = 22). F. T. nuttallii (Spangler s.n., GA), late prophase (2n = 22). G. T. paniculatum,
metaphase (2n = 22).H. I. paniculatum, metaphase | (n = 11).1. I. venenosum, metaphase | (n = 11). Figures A,B, and |
ppl vig oe 970): Fiac. C.D.G and modified from P 1004)
(1
\
~—
Stenanthium (Zomlefer & Judd 2002) and 22 for Toxicoscordion. A probable
base chromosome number of x =8 is of ten cited for the tribe (Sen 1975; Tamura
1995; Lowry et al. 1987; Zomlefer 1997), and multiples of this number are preva-
lent in other genera of the tribe: Amianthium (2n = 32), Anticlea (including
Stenanthella; 2n = 32), Schoenocaulon (2n =16), and Veratrum (including
Melanthium; 2n = 16, 32, 64, 80, 96). Due to the small size of the chromosomes
1090 BRIT.ORG/SIDA 20(3)
Taste 2. Voucher specimens for the chromosome numbers of Joxicoscordion nuttallii reported in
this study.
Taxon Locality and collection number (voucher location) Mitotic chromosome
number (2n
Toxicoscordion nuttallii TEXAS. Hays Co.: Between Driftwood and
Dripping Springs, along FM 150, 3.7 mi 22
N of FM 1826, 15 Apr 2002, Wendt 7246 (TEX)
TEXAS. Wituiamson Co.: Austin, 0.25 mi from 8521 22
Foxhound Tr.,3 Apr 2002, Spangler s.n. (GA)
po
(ca. 2.0-4.0 ttm in length), however, the few detailed karyological studies (e.g.,
Lee 1985) lack the detail to infer mechanisms of chromosomal evolution, al-
though these chromosome numbers indicate the prevalence of polyploidy and/
or aneuploid variation of the prospective basic number. The validation of 2n =
22 for T. nuttallii strengthens support for the monophyly of Toxicoscordion, as
well as the phylogenetic significance of chromosome numbers as generic
synapomorphies for tribe Melanthieae.
ACKNOWLEDGMENTS
Jason Spangler and Tom Wendt (TEX) generously provided live bulbs of T.
nuttallii collected in the field. David E. Giannasi contributed his expertise with
the laboratory work, kindly shared reagents and equipment, and also reviewed a
draft of the manuscript. lam grateful also to Michael Boyd and Andrew W. Tull
(Plant Biology Dept. Greenhouse Facility, University of Georgia) for reviving and
maintaining plants shipped from the field; Walter S. Judd and Dale E. McNeal for
editorial comments; Emily Wood for assistance at GH; Robert J. Liebermann for
translating Russian text; James L. Zarucchi for providing a copy of the in-press
Zigadenus treatment from the Flora of North America; and William Carromero
for the Spanish translation of the abstract. Funds provided by the University of
Georgia Department of Plant Biology financed laboratory materials.
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A COMMUNITY-LEVEL FLORISTIC RECONNAISSANCE
OF THE RAJA AMPAT ISLANDS IN NEW GUINEA
W. Takeuch
Horhoarinq and Arnold Arhoret FI dl ini j
PIGUiVaGIid UFilver ily
Research Associate, PNG National Forest Authority
PNG Forest Research Institute
FO. Box 314
Lae, \ Morobe Province 41 i], PAPUA NEW GUINEA
ABSTRACT
The Raja Ampat district of Indonesian Papua is one of New Guinea's most unusual floristic environ-
ments. ie recent hap ic aesesstent Survey: examined s vegetation patterns across the district, charac-
g biodiversity tl tnreats . Mangroves, beach
jen
forest, iow aac hill forest, limestone karst, and ultr: ase! (serpentine) woodland were the principal
natural-growth habitat lored by the botanical team. Although several lowland areas are tei
degraded by concessional logging, most of the limestone communities remain in p1
There are substantial prospects for future taxonomic discovery.
KEY WORDS: botanical survey, limestone karst, Papua, serpentine, ultrabasics
ABSTRACT
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MO—-DCHS. BADFEYEFTLAAYhMBICHWTC. MROBENY —
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ARBRE OhERRABES CHS, BOMOGMAISHRRERICKLSBEOR
ALORSMNEM, BEORKST RRS RBEOKRSROTHU. DRFLO
MPS SHAMANS A ERM RC BbnS.
+—[7—K : BERR. GREANAL, NTT, te, BERLE
INTRODUCTION
Located at the far western end of New Guinea, the Raja Ampat islands (Figs. 1,
2) are one of Malesia's little-known environmental jewels. In addition to spec-
tacular coastal vistas deserving of World Heritage recognition, recent studies
have determined that the islands probably have the highest levels of coral and
reef fish diversity in the world (Reef base 2002). Floristically, the Raja Ampat
contain 3 of the 22 Papuan areas recognized as centers of very high endemism
(Supriatna 1999).
Most of western New Guinea (including Misool and northern Salawati) are
SIDA 20(3): 1093-1138. 2003
1094 BRIT.ORG/SIDA 20(3)
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Fic. 1.
part of a Papuan microcontinent which separated from the Gondwanic land-
mass during the early Mesozoic, subsequently drifting into its present position
independently of the Australian craton (Pigram & Panggabean 1984, Pigram
& Davies 1987; Hall 1998). Waigeo and adjacent islands comprise one of only
two oceanic terranes in this region of New Guinea. Especially in the
westernmost islands, ultrabasic and limestone substrates have developed dis-
tinctive plant communities containing many endemic species
lronically, considering the overall paucity of information for such envi-
ronments, the Raja Ampat was among the first areas to be visited by early ex-
plorer-naturalists to eastern Malesia. In 1792-93, the vessels Recherche and
Esperance obtained plant collections from Waigeo as part of a general program
of regional exploration. Botanical specimens were also taken during visits by
the Uranie and Physicienne in 1818-19, and during the cruise of the Astrolabe
(1826-29). Early collectors included Beccari (872-73) on Batanta and Kofiau,
and Micholitz in 1890, also on Batanta. Frodin and Gressitt 1982) provide a gen-
eral account of these early activities. Following the efforts of Cheesman on
Waigeo (1938-39) and by Royen on Batanta and Waigeo (Royen 1960), there was
little serious work in the district. Most of the interior sections remain unknown
because of the physical difficulties of access and the nearl
service infrastructures.
y total absence of
MATERIALS AND METHODS
The Nature Conservancy (TNC) survey of the Raja Ampat was designed pri-
marily to investigate the diverse reef ecosystems of the area, and secondarily to
identify linked land-marine conservation targets for possible community-based
initiatives. During a 3-week period (Oct. 30 to Nov. 22, 2002) the scientific team
TAKEUCHI, FLORISTICS OF THE RAJA AMPAT ISLANDS IN NEW GUINEA
1095
Kawe
Fofak Bay
Go Isthmus \, '
Kabare
Halmahera Sea
Waigeo
Batanta .. 9 ..w:.,,
Kasim-Waitama
Misool
o.. Jef Pelee
rn cy 5 en
Ay 5
~ Mesemta Bajampop
Wagmab
a
30 km
131
Fic. 2. Raia Amnat District. Princ f
t
4171 NWI nf YU 4
ATL IN VY
(respectively) 35 and 17 km
the TNC biological reconnaissance. Sayang and Wayag islands are
1096 BRIT.ORG/SIDA 20(3)
made numerous stopovers among all the main islands, using a chartered div-
ing vessel (the Pindito) as a live-aboard mobile base. As ecologist for the forest
assessment, the writer was responsible for describing the principal vegetation
types and ground-truthing perceived community distributions against a pre-
liminary classification developed from the RePPProT (Regional Physical Plan-
ning Program for Transmigration) plant typology maps (RePPProT 1990;
Hardiono 2002). The floristic evaluations and botanical collecting were con-
ducted with assistance by Duncan Neville (TNC Sulawesi Manager), Johanis
Mogea (Herbarium Bogoriense), and Fery Liuw (Departemen Kehutanan Papua).
Forest communities were examined in ad hoc fashion, via walk-through
assessment of areas accessed by dinghy or speedboat (Figs. 3-5). Community
variation was documented by numerous photographs referenced with readings
from a handheld GPS recorder. These onshore activities were necessarily con-
strained by the collateral marine studies and the corresponding movements of
the Pindito. Because of the constantly shifting itineraries of the marine team,
the forest participants could usually devote only one day to a given site, thus
restricting survey coverage to areas near the coast.
Botanical collections were typically made in sets of 5 duplicates whenever
fertile specimens were found. Gatherings were field-packed in newsprint and
plastic bags, then soaked with 70% ethanol for subsequent processing at Her-
barium Bogoriense (BO). Silica-dried samples for DNA sequencing were also
obtained if specialists had placed earlier requests for assistance. In order to sim-
plify the vouchering process, collections were numbered under the sequence of
botanist J. Mogea. Specimen distributions will occur from BO, with A, K, L,and
MANas the principal recipients (rawati pers. comm.). A taxonomic account of
the botanical findings is being prepared by Mogea for separate publication.
RESULTS
Botanical observation and collecting were complicated by drought conditions
caused by the recent El Nino disturbance. The collections tally (550 nos.: Mogea
et al. 7726-8276) includes about 100 sterile numbers because of the unfavor-
able phenology. Although the entire district showed signs of drought stress, the
severity of the disturbance was greatest in the outer islands (Misool, Kawe, and
Waigeo; Fig. 6). At Batanta and Salawati, forest canopies were mostly green (al-
though understories were very dry) and there were even brief periods of tor-
rential rain during the visit to those islands.
Among the habitats covered by the TNC-modified RePPProT forest classi-
fication, mangroves were apparently the only natural-growth community
which could be accurately mapped using Landsat reflectance patterns (see
Hardiono 2002). Topographic and geological maps proved of greater value than
any existing forest typing system. The lack of a GlS-based classification equiva-
lent to the Papua New Guinea Resource Information System (PNGRIS) and its
1097
TAKEUCHI, FLORISTICS OF THE RAJA AMPAT ISLANDS IN NEW GUINEA
Fic 2 1 ba L in Al Hl B® rz hie | iH iH £. + sel eH | laf Dalaa
group. The 75-ft Pindito li hored offsh Zodi 1 I It J f ided d ily transport for the land-
sea survey. Photo November 9, 2002.
Fic. 4 r ie I 1 g as tl . J I JL (I L WY = rT J ig oe | ly pl y i
ry & dD. I ies =: id at n . L . pee Oe L L W J f+] L LI £
4 L L J £ ll A y | gl L L VAL q Fy peed I pt AL L 4,
2002.
Fic. 5. Wagmab Island, SE Misool. Forester Fery Liuw searches for t ical speci in the dry scrub on limestone
I i ££ I | 4 : l = < J Aft let hans : £ +} £. kl
talus 7 the unfavorable
Photo N ber 4, 2002.
conditions
1098 BRIT.ORG/SIDA 20(3)
Fic. 6. Kawe. Drought-stri ti {land on steep sl Photo November 18, 2002.
generically related Forest Inventory Mapping System (FIMS), is a serious ob-
stacle to survey planning in western New Guinea (see Bellamy & McAlpine
1995; Hammermaster & Saunders 1995a, b). To facilitate comparisons with the
FIMS classification, the following community descriptions are prefaced by the
structural code (in parentheses) for the corresponding formations in East New
Guinea.
The expedition included 15 fulltime participants, of which 5 were journal-
ists or photographers. As a result of the intensive media coverage, survey im-
ages and preliminary findings are available on several websites (http://
www.teef base.org/RajaAmpat/,; http://www.AliensOf TheSea.com/HDTY;
http://www.projectbirdwatch.org/rajaampatrea/).
Principal Communities
Mangroves —(M) Of all the vegetation types encountered on the survey, the
mangrove communities are probably the best-studied (e.g. Backer & Steenis 1951;
Ding Hou 1957, 1958, 1960; Percival @ Womersley 1975; Floyd 1977; Duke &
Jackes 1987; Erftemeijer et al. 1989; Duke 1991; Mabberley et al. 1995). Unlike the
situation in nearby Telek Bintuni (cf. Erf{temeijer et al. 1989; Takeuchi et al. in
press), Raja Ampat mangroves are insignificant and markedly impoverished,
except ina few places where estuarine flats and tidal rivers have provided ample
habitat for the Bruguiera-Rhizophora associations (Fig. 7). Among investigated
TAKEUCHI, FLORISTICS OF THE RAJA AMPAT ISLANDS IN NEW GUINEA 1099
Fic. 7, Wayag Island. Interior view of a mangrove forest on limest
9) hi. 1 Ri: | h Photo November 16, 2002.
sites, the best examples of this community were seen on Pulau Misool, along
the lower Gam and Kasim rivers. At the second locality, there is a well-devel-
oped upstream sequence (in order) of Rhizophora mucronata ~ Ceriops tagal,
Bruguiera gymnorrhiza, Nypa fruticans, then a final freshwater assemblage
consisting of Xylocarpus granatum, Dolichandrone spathacea, and Heritiera
littoralis. However in most parts of the archipelago groves are represented
(if at all) only by isolated trees along a foreshore, with no horizontal develop-
ment of the community.
Swamp woodland.—(Wsw) Monodominant forests of Metroxylon sagu (sago)
are scattered through the Raja Ampat district, wherever soil flooding is severe.
The occurrences are insensitive to substrate. Sago swamps are found on lime-
stone at Kofiau (01°09'25S, 129°51'38E), and on mineral clays at Kapatlap,
Salawati. Although the floristic diversity is very low, sago communities are of
considerable subsistence value as a source of dietary starch obtained from the
Metroxylon trunks (cf. Powell 1976; Johns & Hay 1984).
Littoral or beach forest.—(B) A distinctive community along many coast-
lines, beach forests are mostly composed of widely distributed, even pantropical
taxa, but have been reduced over much of their former range because of an-
thropogenic pressures (Wikramanayake et al. 2001). The principal indicator
species for this community are Calophyllum inophyllum, Hibiscus tiliaceus,
BRIT.ORG/SIDA 20(3)
1100
Fic. 8.0
2002.
t
J
characteristic of other coastal areas. Photo November 4
TAKEUCHI, FLORISTICS OF THE RAJA AMPAT ISLANDS IN NEW GUINEA 1101
Pandanus tectorius sens. lat., lerminalia catappa,and Thespesia populnea (Fig.
8). In the Raja Ampat other associates often included Colubrina asiatica,
Parsonsia alboflavescens, Derris indica (Pongamia pinnata), Tacca
leontopetaloides, Ximenia americana var. americana, and Vigna marina. Beach
forest generally occurs on sand or coralline rubble behind the strand zone. A
particularly good example was seen at N Kofiau (O1'09'24S, 129°50'47E). On
uninhabited Sayang Island (0'16'245, 129°53'47E), a different type of beach as-
sociation (BCe) was recorded on sandy flats, consisting of Casuarina
equisetifolia, Scaevola sericea, Sophora tomentosa ssp. tomentosa, Spinifex
ittoreus, and Tournefortia argentea.
oy
Lowland forest on deep mineralized soil—(P1, Hm) The land team encoun-
tered this community type near Kapatlap on Salawati, and on the north and
south shores of Batanta, where the vegetation is predominantly a tall stature
rainforest similar to those from coastal lowlands of the Bomberai Peninsula.
Deep soil habitats (in the eastern Raja Ampat) were probably the most speciose
of all the surveyed environments, but were also characterized by plants com-
mon in adjacent mainland areas and thus of comparatively low conservation
interest (Figs. 9, 10). Intsia bijuga, Koordersiodendron pinnatum, Pometia
pinnata, and Terminalia cf. copelandii were dominant trees in such situations,
with Celtis, Ficus, Dysoxylum, Myristica, and Syzygium the best-represented
genera in the subcanopy. The regrowth phase included Alstonia scholaris,
Gastonia serratifolia, Morinda citrifolia, and Trema cannabina as the most
prominent species, at least within the logged-over tracts seen on Batanta and
Misool.
In the western Raja Ampat, deep soil habitats are generally absent except
in the flood plain of large rivers (e.g. Gam and Kasim rivers), or in the ravines
on limestone karst (e.g. the numerous islets near Mesemta Bajampop of SE
Misool). Such areas have tall forests comparable to the Batanta/Salawati forma-
tions, but are species-poor and usually less than | ha in size (Figs. 11-13). Fre-
quently, the stands are composed of Intsia bijuga, I. palembanica, Vatica rassak,
or Pometia pinnata; with Jagera javanica ssp. javanica, Trophis philippinensis,
Teijsmanniodendron bogoriense (uncommon), and Maniltoa spp. (M. plurijuga
and M. schefferi) underneath. Flindersia amboinensis and F laevigata var.
heterophylla are conspicuous emergents in these tall-forest pockets on the
smaller islands, particularly in the Misool group. Osmoxylon sessiliflorum was
especially prominent in subcanopies near rivers (Misool). At Jef Pelee (S. Misool)
a Homalium foetidum monodominant forest was encountered on ridge slopes
and crests (Figs. 14, 15; 02°01'09S, 130°01'28E).
The forest understories in the Raja Ampat are harder to characterize. Gen-
erally when the canopy has at least 25-50% closure the groundlayer is already
clear. Representative communities in the Misool group had either an open un-
derstory, or a palm-dominant layer of Licuala and rosette-stage Calamus (Figs.
i
1102 BRIT.ORG/SIDA 20(3)
Fic OF " the ta, generally species-rich i parison to tl ter Raja Ampat. Photo Novem-
ber 3, 2002.
Fic. 10. Batanta. Lowland If deep soils, in the | f i bsi gard Tall canopies
{ | Latakti tel A | tn th NI fant re | Av Ratanta 1 C>] a) b
\ y } AV LIC INV
tially absent on the outer islands. Photo November 3, 2002.
Fic. 11. Wagmab. F in broad ravine. C iti d taller, and richer than the surrounding vegetation on
higher slopes. Photo h ber 4, 2002.
TAKEUCHI, FLORISTICS OF THE RAJA AMPAT ISLANDS IN NEW GUINEA 1103
Fic. 12. Interior view of the forest shown in Fig. 11. Photo November 4, 2002.
Fic. 13. Alluvial f | he Kasim Ri f Misool. Photo November 10, 2002.
Fic. 14. Misool. Interi ive of a Homalium foetidum-dominant forest on ridgecrests. Photo November 9, 2002.
1104 BRIT.ORG/SIDA 20(3)
Ch : th | 4 d
>
Fic. 15. As for preceding figure
ity. Photo November 9, 2002.
TAKEUCHI, FLORISTICS OF THE RAJA AMPAT ISLANDS IN NEW GUINEA 1105
re
+
+
2
Fic. 16. Misool. Understory of a limestone forest. The near-g
Photo November 4, 2002.
16, 17). Ferns, herbs, and cryptogams were hardly ever seen, but it is not known
to what extent their absence was due to the severe drought conditions then pre-
vailing. Understories with diverse fern populations (Bolbitis, Davallia,
Drynaria, Nephrolepis, Sphaerostephanos, Tectaria) were found only in river-
ine forest near Fanfanlap (W Misool) and at Kofiau (e.g. 01°10'015, 129°50'30E).
Secondary forests.—(W ) A substantial but unknown percentage of the Raja
Ampat lowlands has been logged by industrial operators and is presently in
vari tages of recovery. The secondary examined by survey were pre-
viously cut at periods ranging from 15 to 30 years ago and are very distinct from
more mature communities in the same habitat.
At Kofiau (O1'11'03S, 129°43'21E) a 15-year regrowth (Figs. 18, 19) was a dep-
auperate woodland of Vitex cofassus, Morinda citrifolia, Conandrium
polyanthum, and Ficus spp. (mainly F microcarpa and F prasinocarpa). Lunasia
amara var. amara, Leea indica, and various Antidesma and Macaranga spp.
formed a sparse underlayer. In contrast, an adjacent section of unlogged forest
was a Closed canopy Pometia-dominant community with high frequencies of
Diospyros, Horsfieldia, and Knema.
On Sayang (a flat sandy island completely logged in 1984; 0°16'38N,
129°53'48E), the regrowth was of similarly depleted composition, consisting for
the most part of a remnant Artocarpus-Intsia~Pometia canopy with a second
CPS
t es a
ares | ‘ " i :
5 he
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a sa Nabe *
BRIT.ORG/SIDA 20(3
i
thal
»
1106
J
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Stems are gen-
Photo November 18, 2002.
bs |
Fic. 17. Kawe. Ct
TAKEUCHI, FLORISTICS OF THE RAJA AMPAT ISLANDS IN NEW GUINEA 1107
Fics. 18 and 19. Kofiau. Regrowth ity from a former forest clearfelled 15 years ago. Photo November 12, 2002.
ef
—
story of Calophyllum inophyllum and Aglaia argentea (the last species previ-
ously unrecorded for the Raja Ampat, cf. Pannell 1992). Conandrium
polyanthum, Codiaeum variegatum var. moluccanum, and Polyscias cumingiana
were undergrowth species. Dense seedling crops of Pometia pinnata often car-
peted the ground in spite of drought conditions.
On a number of small islands, repeated anthropogenic disturbances have
erased the former forest and a grassy fire disclimax (G) is now all that exists. At
Jef Pelee (coral platform: 02°07'03S, 130°19'00E) most of the land surface is cov-
ered by a seral community of Imperata conferta with Ischaemum muticum on
the seaward fringe (Fig. 20).
At Mesemta Bajampopa single example was seen of an early succession on
karst, consisting of Dracaena angustifolia and Commersonia bartramia, with
an extensive underlayer of Gahnia aspera (Fig. 21)!
Savanna.—(SaMl) Grassy savannas were recorded only on W Misool, at the
mouth of the Kasim River and further inland near the Waitama tributary
(01°47'235, 129°53'57E and 01°51'41S, 129°55'05E respectively). The Kasim com-
'These sort of communities are the result of localized disturbance and are too small to be mapped by forest
classification systems.
1108 BRIT.ORG/SIDA 20(3)
Fic. 20. Jef Pelee. Antt i land (/mp Ischaemum ia) i i d the Misool
mainland. Other seral taxa included De modium umbellatum, Pi tifolia, Morinda citrifolia, and Pandanus
tectorius. Photo November 8, 2002.
Fic. 21. Mesemta Bajampop. A seral community on poh consisting exclusively of Commersonia bartramia, Dracaena
angustifolia, and Gahnia aspera. Photo November 5, 200
TAKEUCHI, FLORISTICS OF THE RAJA AMPAT ISLANDS IN NEW GUINEA 1109
munities were Melaleuca dominant, but also included smaller individuals of
Pandanus tectorius and Timonius timon ina distinct second layer (Figs. 22, 23).
The dense 2 m+ groundcover was composed of Ischaemum barbatum and
Imperata conferta, together with scattered patches of taller Saccharum
spontaneum (Fig. 24). Indications of recent burn suggest that the Kasim savanna
is probably periodically fired.
The Waitama savannas are larger than the Kasim communities and include
Eucalyptus cf. papuana in addition to the more common Melaleuca
leucadendron sens. lat. (Fig. 25). Shrubs were commonly represented by Baeckea
frutescens, Decaspermum bracteatum, Melastoma malabathricum ssp.
malabathricum, and a Polyscias sp. The groundlayer of Ischaemum barbatum
and Rhynchospora rubra was habitat-partitioned, the latter species mainly oc-
cupying flat areas with the poorest drainage.
In appearance and composition the Waitama savannas are nearly identi-
cal to comparable formations on the Papuan mainland at Bomberai (cf. Takeuchi
et al. in press). The communities at both sites are apparently under substrate
control, with characteristic occurrences on flat or gently undulating terrain
underlain by hardpan and alumina deposits (pers. obs.). Although the savan-
nas in the Raja Ampat are affected to some degree by fire, the substrate patterns
suggest the communities are a longterm response to stable edaphic factors (cf.
Paijmans 1976).
Lowland forest on limestone karst.—(Hs, HsCp)* The limestone terrain is
one of the most visually stunning and unspoiled environments in the Raja
Ampat district. Much of this habitat is a deeply dissected karst which is often
extremely difficult and time-consuming to traverse. A small hill can take an
entire morning to ascend, so it is not hard to comprehend the relative lack of
botanical collections from such areas.
Excellent examples of karst vegetation are found in the Misool chain, par-
ticularly in the SW complex of small islets and at the western end of the Misool
mainland. Within the Waigeo group, extensive limestone habitats were explored
near Aljoei (0'11'435, 130°1539E) and at Wayag (0°10'21N, 130°01'17E).
Many of the smaller limestone islands are undercut at the highwater line
into toadstool platforms, or are sculpted into steep conical stacks (Fig. 26).
Oftentimes the sides of the taller islands have dizzying vertical faces that plunge
for hundreds of feet (Figs. 27, 28). On Wagmab, the stepped ledges have a stunted
woody vegetation, frequently wind-sheared, comprised of Stenocarpus moorei,
Exocarpos latifolius, Polyscias sp. nov., Wikstroemia androsaemifolia,
The FIMS code is partly based on the dominance of Gymnostoma papuanum (=Casuarina papuana) which is
replaced by Gymnostoma ee (=Casuarina Gaeeiane) in the Raja Ampat. The second specen! is a rela-
th EINAG
tively minor element i tk gent on ultrabasics.In conformity with FIMS
¢
convention, the Raja Am| | Idt as’HsCr',or'HsGr in any extension of the typing system to
Western New Guinea.
1110 BRIT.ORG/SIDA 20
3)
Fic. 22. Misool. Melal | lend |., the dominant stand-formi ies inthe Raja A t The whit-
ish-papery outer bark and drooping | haracteristic. Photo November 10, 2002.
1111
TAKEUCHI, FLORISTICS OF THE RAJA AMPAT ISLANDS IN NEW GUINEA
d, darker-leaved layer beneath taller Melaleuca.Photo
Fic. 23.Misool. K
November 10, 2002.
|. Photo November
Fic. 24. As for Fig. 23. Tt
7t
J
pe
10, 2002.
1112 BRIT.ORG/SIDA 20(3)
Fic. 25. Misool. Tt
Wai Photo November 11, 2002.
Fic. 26. Aljoei. Steep sided stacks and platforms, a typical geomorphic feature of the limestone terrain at Misool and
Waigeo. Photo November 14, 2002.
TAKEUCHI, FLORISTICS OF THE RAJA AMPAT ISLANDS IN NEW GUINEA 1113
f the underwater walls, Photo
Fic. 27 Aljoei Limestone cliffs
November 14, 2002.
J t J 7
BRIT.ORG/SIDA 20(3)
1114
+, 2002.
A
N h
hk Dhat
Fic. 28. Li
TAKEUCHI, FLORISTICS OF THE RAJA AMPAT ISLANDS IN NEW GUINEA 1115
Calophyllum spp. (including the novoguineense complex; cf. Stevens 1995), and
many vines of Alyxia purpureoclada (Fig. 29). The unusual Podocarpus
polystachyus is a dwarfed krummholz on solid rock, sprawling across ledges
and barely ascending to 1 m height. But at Wayag, crestline populations of the
same species grow as erect trees to 7 m height (Fig. 30; 0°10'25N, 130°01'22E and
0°10'42N, 130°01'16E).
On nearly all the limestone islands, Gulubia costata is a conspicuous emer-
gent, often forming dense stands along the higher slopes and crestlines.
Barringtonia, Dracaena, Ficus, Garcinia, Gynotroches, Myrsine (Rapanea),
Pouteria (Planchonella), Schefflera, Serianthes, and Sterculia are also common
subarborescent genera, but a disproportionate number of morphospecies were
documented only by sterile or scrappy vouchers and cannot be identified with
certainty. Of all the Raja Ampat habitats, the limestone vegetation was the most
severely affected by the El Nino drought, and effective documentation of such
communities was very difficult under the prevailing conditions. The canopies
on most islands were brown and withered, though probably not dead. Lime-
stone karst is undoubtedly a poor reservoir for soil moisture. Especially on small
islands, stands with green foliage were often restricted to the colluvial accu-
mulations in valleys and small draws (Figs. 1, 12).
Archidendron paluense, Lunasia amara var. amara, and the vinelike
Bauhinia binata (Lysiphyllum binatum) were common taxa in karst understo-
ries of the western Raja Ampat, especially on platforms and stacks around Misool
proper. At Mesemta Bajampop, substantial populations of Rauvolfia moluccana
and Monophyllaea were also noted. The Archidendron, Bauhinia, Monophyllaea,
and Rauvolfia were newly recorded for the district (cf. Burtt 1978; Verdcourt
1979; Nielsen et al. 1984; Hendrian & Middleton 1999). New registers for such
common or conspicuous plants are indications of the undercollected status of
the limestone, and show how poorly documented this flora still remains even
after more than a century of Papuan exploration.
Lowland ultrabasic scrub and forest.—(W, HsCp) Renowned in scientific lit-
erature, the Raja Ampat serpentine flora is also one of the region's most pictur-
esque environments. At Kawe (Figs. 31, 32) and Waigeo's north shore (Fig. 33)
the ultrabasic scrub affords breathtaking panoramas of turquoise reefs and red
laterite across extensive stretches of coastline.
The TNC survey explored the ultrabasic zone in a series of ascents along
steep-sided buttress ridges. Most of the vegetation consisted of xeromorphic
scrub or woodland with similar characteristics to communities described in
an earlier account of the eastern Waigeo peninsula (Royen 1960).
During the first landfalls on Kawe (ending at 0°03'07S, 130°08'05E), the for-
est team founda distinctive woodland of Ploiarium sessile, Exocarpos latifolius,
Gymnostoma rumphianum, Decaspermum bracteatum, Ixonanthes reticulata,
and Myrsine rawacensis (Fig. 34). The spreading sympodial crowns of Ploiarium
1116 BRIT.ORG/SIDA 20(3)
Fic. 29. Wagmab. Vegetation on limestone ledges. Photo November 4, 2002.
sessile were a particularly striking characteristic of this community (Fig. 35).
In the wide spaces between the larger trees were many shrubs of Myrtella
cae i, Styphelia abnormis and (in lesser numbers) Dodonaea viscosa. Vines
f Alyxia laurina were common climbers and scramblers. Patches of
Dic ranopteris linearis blanketed the ground nearly everywhere, with Dianella
ensifolia, Nepenthes danseri, Palhinhaea cernua, Machaerina disticha, and M.
glomerata scattered mainly over the bare spaces. Artocarpus, Livistona, and
Vitex were common on lower slopes and valley floors (Fig. 36; 0°03'05S,
130°08'24E).
On the Go Isthmus of Waigeo (Fofak Bay at 0°02'22S, 130°43'43E), the ultra-
basics were similar though somewhat richer, and also ee Arytera littoralis,
a small-leaved Gmelina, Melastoma malabathricum ssp. malabathricum,
Psychotriatripedunculata (Fig. 37),and Rhodamnia novoguineensisamong the
common woody plants. Parts of the scrubland above the bay had been burned,
exposing large patches of the distinctive red laterite (Fig. 38). Within such ar-
eas, the early fire succession consisted of a sparse association of Commersonia
bartramia, Myrtella beccarii, Scaevola oppositifolia, and Styphelia abnormis.
Although the ridgetop habitats are pees of open aspect, the vegetation in
the draws isa closed codominant forest of Dillenia alata and Calophyllum spp.
At scattered places in the ultrabasic zone (on both crestlines and depressional
areas), the scrub is replaced by a taller Gymnostoma rumphianum-Sapindaceous
—
TAKEUCHI, FLORISTICS OF THE RAJA AMPAT ISLANDS IN NEW GUINEA 1117
Fic. 30. Wayag. Podocarpus polystachyus (5-7 m tall) li ies. Photo November 16, 2002.
Fic. 31. East-central coastline of Kawe island, an ultraba
F be J r \
sic outcrop 15 km west of Waigeo. In contrast to the uneven and deeply dissected limestone, ultrabasics are characteristically
1977). Photo November 18, 2002.
SLLL
(€)0z vals/9¥0"LINa
TAKEUCHI, FLORISTICS OF THE RAJA AMPAT ISLANDS IN NEW GUINEA
1119
4
a!
~
al
Fic. 32. Kawe. Three
Tl Rahel L ot £ eh Wiel 1: L
mit
the hori f the last fi (to ESE). Photo November 17, 2002.
ee,
fee ies
Say i.
24
.
+
+
a
r
we
Fic. 33. Waigeo ultrabasic scrub at the entrance of Fofak Bay. Center of view to the southeast,
isla
2002.
- F J 31 Ka
Photo November 19,
OZLL
€)02 vaIs/O¥O'LINA
TAKEUCHI, FLORISTICS OF THE RAJA AMPAT ISLANDS IN NEW GUINEA 1121
Fic. 34. Myrsine rawacensis on Kawe ultrabasics. Photo November 17, 2002.
1122 BRIT.ORG/SIDA 20(3)
J 4 < L I+pa} . h Nhe | 1 tho £1 . 4°
Fic. 35. Ploiarium sessile, th
ately alsriiguish aE species from congenes (Kobuski ie oo recorded from Halmahera (ibid.), P sessile is
to November
canopy witha Myrtella beccarii understory (e.g. at Kabare, 0°04'18S, 130°56'36E:
Figs. 39, 40). Unlike the open growth, this taller community (at Kabare) is
densely stocked with 7-10 m pole-trees including the serpentine indicator
Ploiarium sessile. Gymnostoma rumphianum is a 25 m emergent in such situa-
tions. The ultrabasic vegetation is thus represented by several communities,
collectively forming an intergrading series of facies ranging from bare ‘blow-
outs’ (Fig. 41) through woodland of varying densities, then finally to a closed
multistoried forest.
DISCUSSION
—_—
Raja Ampat plant communities are primarily lowland environments. There are
no areas of significant size higher than the 900-1000 m level where montane
conditions generally begin (cf. Paijmans 1975, 1976; Johns 1977; Grubb & Stevens
1985; Hammermaster & Saunders 1995a). Although Royen (1960) describes a
mossy montane forest on Mt Bulfelhoorn, such habitats could not be visited
within the time allowed by the survey schedule. None of the daily excursions
from the coast were able to penetrate beyond 500 m elevation.
Within the limitations imposed by rapid assessment, it is apparent that
TAKEUCHI, FLORISTICS OF THE RAJA AMPAT ISLANDS IN NEW GUINEA
before
af
£
ing
a spr
te of the drought
In spi
entral Kawe.
Cc
£ L
VaHIty Ul SUULTING
Fic. 36.5
1123
\
}
J
fl * OL i |
hed. A locally-| r
J
J
other than small shrimp. Photo November 18, 2002.
rr
1124 BRIT.ORG/SIDA 20(3)
Photo Novem-
Fic. 37. The aptly-named Psychotria tripedunculata, one of tk
ber 19, 2002
ie ae Maigret fire suecesslonal community. Burning removes the horn ae of areca linearis and
to November 1
J U J 7
Fic. 39. i ised stand of pole stemmed Wo near me Mes succession shown | in Fig. 38. The ultrabasic forest facies is
Photo November 19, 2002
r J
1125
TAKEUCHI, FLORISTICS OF THE RAJA AMPAT ISLANDS IN NEW GUINEA
be J
Fic. 40. Ul
Myrtella codominant. Photo November 20, 2002.
1126 BRIT.ORG/SIDA 20(3)
tinctive orange-red late Th ional f babl sult of forest re-
moval: eltnough the oper communities convey a an impression of stability and continuity: the develonmnent of such
Photo November 19, 2002.
Fic. 41. Waigeo blowout. ie patches af sheet ded earth i tl h tk bland, exposing the dis-
e of th
.: t
the Raja Ampat flora is depauperate and disharmonic relative to adjacent main-
land environments and that these distinctions become more pronounced in the
outer island groups (e.g. Misool and Waigeo). Many of the most characteristic
Papuasian families, including Annonaceae, Elaeocarpaceae, Gesneriaceae,
Lauraceae, Melastomataceae, Meliaceae, Piperaceae, and Urticaceae, were poorly
represented in the lowland habitats where they are ordinarily prominent. Epi-
phytes and climbers were also generally scarce, as were several herbaceous fami-
lies usually found in New Guinea forest understories (e.g. Marantaceae,
Orchidaceae, Zingiberaceae).
This westward trend of diminishing diversity across the archipelago is cor-
related to a general reduction in rainfall and to the limiting nature of the west-
ern substrates. Rainfall is directly correlated with floristic diversity, more than
with any other abiotic factor (Gentry 1988). The perception of highest richness
in the near-mainland environments of Salawati and Batanta, and the impres-
sion of lower specie toward the outer groups, is consistent with annual
rainfalls (cf. Mangen 1993): Salawati and Batanta (3000-3500 mm); Misool
(2500-3000 mm); Waigeo and adjacent islands (1500-2000 mm). The floristic
trends expected from moisture availability are enhanced by the substrate dis-
TAKEUCHI, FLORISTICS OF THE RAJA AMPAT ISLANDS IN NEW GUINEA 1127
Fic. 42. Kamiali, Papua New Guinea. Successional scrub community on a 15-year old landslide. Plants on the exposed
Ara ced sel pee V ppreread | pa 2 sere i ni ic ii ic Mach j j M. rubiginosa
a 7 # f * r Lf ~s
The domi f Cyperaceae (p y ina) is a curious feature of the ground layer in situations where
= \WAlat 7.
ld 1°
J . ’ 1 n Th:
Ly J t
Feb. 5, 2003.
1128 BRIT.ORG/SIDA 20(3)
t t 7 PS.
Fic. 43. Kamiali, P New Gui S . Neo des afi
Kamiali
| Py J L )
L L = £.1] S\AI-A: Pe 4 j J iH . AA +
both districts. Photo October 6, 2002.
tinctions. Whereas most edaphic environments in the eastern islands are of
mineralized soil, the outer islands are predominantly limestone or ultrabasics,
infertile substrates that are known to be limiting for plant growth (Royen 1963;
Kruckeberg 1985; Brooks 1987). The synergistic combination of factors results
in diminished richness but increased endemism.
In terms of exploration priorities, the highest potential for taxonomic dis-
covery is probably with the Misool karst. Waigeo ultrabasics are comparatively
easy toaccess and the communities are usually of low stature and density, char-
acteristics which favor collection saturation. Most, if not all, of the ultrabasic
endemics have probably been discovered. In contrast, the Misool limestone is
wetter (notwiths conditions during the survey), very difficult to traverse,
and suitable collections are much harder to find. A significant number of un-
known taxa is likely to be present on the karst. The new distributional records
from the survey are suggestive of future opportunities.
Conservation Assets.—W hen judged on the basis of uniqueness, the ultra-
basic and limestone vegetation are the highest-value communities in the Raja
Ampat. As presently known, the ultrabasics have more species endemic to its
habitats than any of the other communities. Plants restricted to Waigeo ser-
ae
TAKEUCHI, FLORISTICS OF THE RAJA AMPAT ISLANDS IN NEW GUINEA 1129
Fic. 44. P H | ra th nf Eafal RB \Ala; In] eee n ay H rT ° re g. 33). Mapped
asa Gymnoeng (Casuarina) et in 11955, thls area is now an eroded and depauperate scrubland; A few ielictttal
Photo November 19, 2002.
pentine include Alstonia beatricis, Alyxia laurina, Archidendron royenii, Guioa
waigeoensis, and Maesa rheophytica. Psychotria tripedunculata is also known
primarily from the ultrabasics, with only one record originating elsewhere.
Waigeo's floristic patterns can be understood by comparison with similar
areas from other districts in New Guinea. In the Bowutu Mts of Morobe Prov-
ince, an elevational sequence of ultrabasic landscapes forms one of the largest
features of this type in Papuasia (ie. the Papuan Ultrabasic Belt, cf. Thompson
& Fisher 1965; Bain 1973; Loffler 1977). Especially within the Kamiali Wildlife
Management Area (KWMA), the serpentine flora extends across a series of
coastal communities beginning at sealevel, in a manner analogous to Waigeo.
K abitats are subject to frequent landslides because of a local com-
bination of humid climate and steep coastal ridges. In the recovery following
landslides, the community that develops is a scrub composed primarily of
Myrtella beccarii, Dicranopteris linearis, Machaerina glomerata, and M.
rubiginosa; with (at lesser frequencies) Stenocarpus moorei and Tristaniopsis
macrosperma (Fig. 42). In appearance and composition, the pioneer commu-
nity is very similar to the Waigeo scrubland and also occurs alongside a series
of taller forests including many of the same elements (Fig. 43). The KWMA
D
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(€)02 Vals/540'LINa
TAKEUCHI, FLORISTICS OF THE RAJA AMPAT ISLANDS IN NEW GUINEA 1131
community variation clearly represents a successional sequence, which can be
tracked because of the known history of specific sites.
Royen (1960: 41) described the Waigeo scrub as an edaphic climax, but its
spatial proximity and structural resemblance to other communities suggest
otherwise. By analogy to similar environments, most of the Waigeo ultrabasics
are probably an early stage in a successional sequence caused by fire. The prin-
cipal characteristics of the open scrub are hardly different from landslide seres
of the KWMA, except that the Waigeo communities are much larger, extend-
ing continuously over hundreds of hectares. The distinction in spatial scale can
be attributed to the presumed fire etiology, which would tend to act over much
larger areas than landslides. The etiologies can themselves be explained by the
climatic contrasts between the sites. Fires are a common ecological factor in
dry environments like Waigeo, while humid localities (KWMA rainfalls are
4000 mm per annum)are naturally susceptible to landslides during periods of
heavy rain. Royen's own observations provide evidence of instability in the Raja
Ampat ultrabasic vegetation (Royen 1960). The ridge in the foreground of Figs.
33 and 41 was a Gymnostoma (=Casuarina) forest in 1955 (ibid.: Fig. 5). As docu-
mented by the TNC survey, that forest now remains only as relictual trees sur-
rounded by scrubland (Fig. +4). These open areas are clearly a new develop-
ment and not a permanent edaphically-induced feature.
The frequent occurrence of fires is currently reflected in the patchy distri-
butions of ultrabasic forest on Waigeo, an expected pattern if seral sequences
are being continuously reset over large areas. Although the serpentine scru-
bland is one of Papuasia's most impressive environments, the rare and endemic
taxa are concentrated in the taller vegetation, and this latter habitat probably
represents a more valuable conservation asset. In an environment with fire-
induced succession, the closed forest (Figs. 39, 40) should be the richest com-
munity, and thus the more promising target for future exploration.
Apart from the biotic measures, aesthetic considerations are a legitimate
part of any environmental evaluation. Waigeo and Misool landscapes are very
photogenic, and can serve as focal assets for an ecotourism niche market in
combination with the marine attractions, as already demonstrated by the
Pindito (www.pindito.com).
Ecosystem Threats.—During the last 50 years, the areas of post-fire succes-
sion in northern Waigeo have expanded substantially (compare Royen 1960:
Figs. 3, 5). Several endemic plants are probably being threatened by the exist-
ence of this historical trend, particularly by the accompanying reductions of
taller communities. Among ultrabasic species, Archidendron royenii, Alstonia
beatricis, and Maesa rheophytica are still known only from their types, taken
in primary forest or older-growth woodland (see Nielsen et al. 1984; Sidiyasa
1998; Sleumer 1987). From the circumstances of their collection, these plants
are probably associated with the advanced stages of the ultrabasic succession,
1132 BRIT.ORG/SIDA 20(3)
—
and their apparent rarity is consistent with habitat reduction caused by in-
creased fire frequency. In contrast, the ultrabasic endemics Guiod waigeoensis
and Alyxia laurina are known by several collections from the open areas, and
are probably seral taxa (cf. Welzen 1989; Middleton 2000).
Naturalized alien plants were not found on serpentine, or on most of the
isolated karst islands at Aljoei and Mesemta Bajampop. Although limestone
areas near human settlements often have a few adventives (viz. Bidens pilosa,
Boerhavia erecta, Euphorbia heterophylla, E. hirta, Passiflora foetida,
Stachytarpheta jamaicensis, Tridax procumbens etc.) truly invasive taxa were
absent. The noxious Lantana camara and Piper aduncum were recorded only
on Batanta and Salawati. The same factors which make the western environ-
ments limiting for plant growth apparently act to discourage the establishment
of aliens.
A planted patch of Hibiscus rosa-sinensis was seen at uninhabited Wagmab,
and even if such occurrences are not threatening, deliberate introductions into
otherwise weed-free habitats need to be discouraged by proactive conservation
policy. Predictive guidelines for assessing possible invasive success of alien plants
have been developed for other island environments (cf. Pheloung et al. 1999;
Daehler & Denslow 2002). Their application should be considered by Papuan
managers admi nistering conservation tracts, as Most communities in the west-
ern Raja Ampat are of entirely indigenous or endemic composition. Nowadays,
ina world where the spread of aggressive species contributes to homogeniza-
tion of floras, areas such as the Misool karst are an increasingly rare ecological
resource.
Each of the principal islands (Batanta, Misool, Salawati, Waigeo) have siz-
able set-aside areas designated as nature preserves (Supriatna 1999). However
existing logging threats are substantial, and commercial operations were filmed
during the recent TNC assessment even within the so-called reserves. Although
the archipelago is known to have significant plant and faunal assets, many of
—
these are under threat and will require management action to ensure their con-
tinuity (ibid.). Establishing new incentives for conservation will not be easy,
since in most Raja Ampat forests the emergent canopy is composed of major
exportable timbers (Intsia bijuga, I. palembanica, Pometia pinnata, Flindersia
amboinensis, Vatica rassak; see Louman & Nicholls 1995) occurring at stocking
densities which favor profitable extraction (Departemen Pertanian Direktorat
1977). Although the current concessional areas are habitats with good site ca-
pacities for tree growth, the ultrabasics and limestone karst have stunted veg-
etation of little value as logging targets, and are thus at lesser economic risk.
Despite indications that climatic warming will cause substantial reduc-
tions in Papuasia's rainforest biome, there have been no attempts to define local
planning and management responses to this threat. Baselines are much needed
for determining the onset and direction of climate-induced change ina variety
TAKEUCHI, FLORISTICS OF THE RAJA AMPAT ISLANDS IN NEW GUINEA 1133
of communities. Because of their insular and pristine status, many Raja Ampat
environments would make appropriate stations for monitoring the floristic
effects of El Nino oscillations. The western islands are part of a forest continuum
spatially connected to perhumid mainland habitats, extending across a geo-
graphic sequence of sparsely inhabited landscape, tectonic, and biotic environ-
ments. The district is very suitable as a venue for ecological research.
Nearly all conservation programs in New Guinea are committed exclu-
sively to terrestrial or marine habitats, due to the disparate value of such envi-
ronments when they occur together at most sites. However at Misool and Waigeo,
highly endemic forest communities lie alongside some of the most diverse reef
ecosystems in the world (Fig. 45). This unusual combination of worldclass as-
sets will permit development of holistic strategies for managing the linked land-
sea resources. The information needs for integrated planning will be complex,
requiring data inputs on wildlife, floristic, recreational, cultural-political, land-
scape, and human subsistence values. But owing to the unique nature of the
environmental assets in this district, future initiatives have the opportunity of
being cost-effective, innovative, and compelling.
APPENDIX OF BINOMIALS CITED IN THE TEXT
Aglaia argentea Bl. (Meliaceae) Conandrium polyanthum (Laut. & K.Schum.)
Alstonia beatricis Sidiyasa (Apocynaceae) Mez iu lees
Alstonia scholaris (L.) R. Br. (Apocynaceae) Dec m (Roxb.) AJ. Scott
Alyxia laurina Gaudich. (Apocynaceae) ee
Alyxia purpureoclada Kaneh. & Hatus. Derris indica (Lam.) J.J. Bennett (Fabaceae)
(Apocynaceae) Des sel Genes (L.) DC. (Fabaceae)
Archidendron paluense (Kaneh.) Nielsen Dial / a) DC. (Liliaceae)
(Mimosaceae) Dicran biel s linearis (Burm. f.) Underw.
Archidendron royenii Kosterm. (Mimosaceae) (Gleicheniaceae)
Arytera littoralis Bl. (Sapindaceae Dillenia alata (R.Br.ex DC.) Martelli
aeckea frutescens L. (Myrtace = ae
Bauhinia backs Blanes 1) ane Dodonaea viscosa Jacq. (Sapindaceae)
Bidens pilosa L. (Asteraceae) Dolic! pie le a (Lf) K.Schum.
Boerhavia erecta L. eee PiguonieGeae)
Be as eas iza (L.) Lamk D lia Roxb. (Dracaenaceae)
(Rhizophoraceae) Eucalyptus anand White (Myrtaceae)
ie ee, um L. (Clusiaceae) Euphorbia heterophylla L. (Euphorbiaceae)
Casuarina equisetifolia Forst. & Forst. f. mesos ia Wine L. (Euphorbiaceae)
(Casuarinaceae) olius R. Br. (Santalaceae)
Casuarina papuana S. Moore (Casuarinaceae) Ficus microcarpa L. (Moraceae)
Casuarina rumphiana Miq. (Casuarinaceae) Ficus prasin 7 Elm. (Moraceae)
Ceriops tagal (Perr.) C.B. Rob. (Rhizophoraceae) Fl dersia amboi is Poir. (Rutaceae)
Codiaeum variegatum (L.) Bl. var. moluccanum Flindersia laeviga 1 Whit e& ae var. heter-
(Decne) Muell. Arg. (Euphorbiaceae) ophylla (Merr. & oe Hartley (Rutaceae)
ina asiatica Brogn. (Rhamnaceae) Gahnia aspera (R. Br.) Spreng. (Cyperaceae)
Commersonia bartramia (L.) Merr. Gastonia serratifolia ae Philipson
(Sterculiaceae) (Araliaceae)
1134
\A/
Guioa waigeoensis Welzen (Sapindaceae)
Gulubia costata Becc. (Arecaceae
sates papuanum (S.Moore) LASS.
n (Casuarinaceae)
aynnoston rumphianum (Mia.
Casuarinaceae)
Heri tiera ; care Ait. (Sterculiaceae)
cus rosa-sinensis L. ae
ne iscus tiliaceus L. (Malvaceae)
Homalium foetidum (Roxb.) an
(Flacourtiaceae)
Imperata conferta (Presl) Ohwi (Poaceae)
Intsia Sate (Colebr.) O. Kuntze
(Caesalpiniaceae)
Intsia ean ica Miq. (Caesalpiniaceae)
Iechaqemrrmy barhat
Ww
LAS.
ischaemum muticum L. (Poaceae)
Ixonanthes reticulata Jack (Ixonanthaceae)
Jagera javanica (Bl.) Bl.ex Kalkm. ssp. javanica
(Sapindaceae)
Koordersiodendron pinnatum (BI.) Merr.
(Anacardiaceae
ras
ee |
Leea indica (Burm. f.) Merr. (Leeaceae)
unasia amara Blanco var.amara (Rutaceae)
Lysiphyllum binatum (Blanco) de Wit
(Caesalpiniaceae)
Machaerina disticha (Clarke) Koyama
(Cyperaceae)
Machaerina glomerata (Gaud.) Koyama
Machaerina rubiginosa (Spreng.) Koyama
(Cyperaceae)
Maesa sls de ood ane a
aa niace
ie schefferi K. oon & Hollrung
(Cae eSulpinioes
Melaleuca | Hs 1 L.sens. lat. (Myrtaceae)
Melee one caaiobaiineans L.ssp.
malabathricum (Melastomataceae)
Metroxylon sagu Rottb. ena a
Morinda citrifolia L. (Rubiace
A.DC. sees
Myrtella beccarii (F.v.M.) Diels (Myrtaceae)
Nepenthes danserii Jepb & Cheek
Ne sak enthaceae
Nypa fruticans Wurmb (Arecaceae)
hanes sessiliflorum ion Philipson
(Araliaceae)
AA : ;
VIVE OTITIS (OVVOUCEH ots
wa
rea
BRIT.ORG/SIDA 20(3)
Dalh f
Palhi a cermud (L.) Vasc. & Franco
(L ee
Pandanus tectorius Parkinson sens. lat.
(Pandanaceae)
Parsonsia alboflavescens (Dennst.) Mabb.
(Apocynaceae)
Passiflora foetida L. (Passifloraceae)
Pie GENE .(Piperaceae)
Ploiarium sessile (Scheff.) Hall. £. (Theaceae)
Podocarpus polystachyus R. Br.ex Endl.
WOOP anaes
Pol a (Presl) F.-Vill. (Araliaceae)
Pometia pinnata JR. & G, Forst. (Sapindaceae)
Pongamia pinnata (L.) Pierre (Fabaceae
Premna serratifolia L. (Verbenaceae)
(Rubiaceae)
kgrf. (Apocynaceae)
Ri: izophora mucronata oe (Rhizophoraceae)
Rhodamnia novoguineensis AJ. Scott
(Myrtaceae)
Rhy nch pora rubra (Lour.) Makino
(Cyperaceae)
ee eae 1 L. (Poaceae)
cd R. Br. (Goodeniaceae)
Geneiola sericea Vah| (Goodeniaceae)
Sophora tomentosa L. ssp. tomentosa
(Fabaceae)
sas littoreus (Burm. f.) Merr. (Poaceae)
Stachytarpheta jamaicensis (L.) M.Vahl
(Verbenaceae)
Stenocarpus moorei F.v.M. (Proteaceae)
Styphelia abnormis (Sond.) JJ.Sm.
(Epacridacea
Tacca on inde (L.) O.K. (Taccaceae)
Teijsmanniodendron bogoriense Koorders
(Verbenaceae)
cir
Ke
—
ent Ie us dunculata Sohmer
Combretaceae)
Terminalia copelandii Elm. (Combretaceae)
Thespesia populnea (L.) Sol. ex Corr.
(Malvaceae)
Timonius timon (Spreng.) Merr. (Rubiaceae)
ae id pa ae = (Boraginaceae
Lour. (Ulmaceae)
Tridax procumbens L. een
Tristaniopsis macrosperma (F.v.M.) Wilson &
Waterh. (Myrtaceae)
Trophis philippinensis (Bur.) Corner (Moraceae)
Vatica rassak (Korth.) Bl. (Dipterocarpaceae)
Vigna marina (Burr.) Merr. (Fabaceae)
Vitex cofassus Reinw. ex Bl. (Verbenaceae)
SS
TAKEUCHI, FLORISTICS OF THE RAJA AMPAT ISLANDS IN NEW GUINEA 1135
Weinmannia fraxinea (D.Don) Mia. Ximenia americana L. var.americana
(Cunoniaceae) (Olacaceae)
Wikstroemia androsaemifolia Decne Xylocarpus granatum Koenig (Meliaceae)
——
(Thymelaeaceae
ACKNOWLEDGMENTS
I thank The Nature Conservancy of Indonesia, Herbarium Bogoriense, the Ar-
nold Arboretum, and Harvard Herbaria, for their institutional support of the
Raja Ampat survey.
also acknowledge Duncan Neville (TNC, land team coordinator), Johanis
Mogea (Herbarium Bogoriense), Fery Liuw (Departemen Kehutanan Papua),
Martin Hardiono (World Wildlife Fund Indonesia), and Victor Motombri (BP
Indonesia) for their contributions to the floristic work. Rod Salm (TNC) was
the overall expedition leader. Hitofumi Abe (Ecosystem Research Group, Uni-
versity of Western Australia) wrote the Japanese translation. Martin Hardiono
provided the map template for Fig. 2.
The other expedition participants included Pawel Achtel Jungle Run Pro-
ductions, photographer), Gerald Allen (Western Australian Museum), Ryan
Donnelly (environmental journalist), Fredy Frommenwiler (Pindito captain),
Morgan Gabereau Jungle Run Productions, producer and co-director), Creusa
Thetha Hitipeuw (World Wildlife Fund Sahul), Djuna Ivereigh (Project Bird
Watch), Agus Sumule (Universitas Negeri Papua), Yulianus Thebu (World Wild-
life Fund Sahul), Emre Turak (Australian Institute for Marine Science), and Joe
Yaggi Jungle Run Productions, team leader).
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SOIL-DEPENDENT FIRE FREQUENCY:
A NEW HYPOTHESIS FOR THE DISTRIBUTION
OF PRAIRIES AND OAK WOODLANDS/SAVANNAS
IN NORTH CENTRAL AND EAST TEXAS
George M. Diggs, Jr. Peter €,Scnulze
Department of Biology & Department of Biology &
Center for Environmental Studies Center ia Environmental Studies
ustin College Austin
Sherman, TX 75090, U.S.A. Sherman, TX 75090, U.S.A.
an Se ee ACen ege.edu
Botanical Research Institute of Texas
Fort Worth, TX 76102-4060
gdiggs@austincollege.edu
ABSTRACT
The distinctive historical vegetation pattern of alternating tallgrass prairies on clay soils and oak
woodlands/savannas on sandy soils in North Central Texas and East Texas has been described for
more thana century. Many authors have attributed this pattern to relatively high levels of soil mois-
ture available for tree growth on areas of sandy soil, and conversely, inadequate levels of soil mois-
ture fort clay soils. However, this explanation is not consistent with present day obser-
vations o
that the historical distribution of prairies and woodlands in North Central and East Texas can be
rept invasion of clay soils by woody vegetation. We propose an alternative hypothesis,
explained by soil-dependent variations in grass biomass and resulting differences in fire frequency
and intensity.
RESUMEN
El patron apa de ye enel 7 alternan praderas con hierbas altas en suelos calcareos y
robledales ro-norte e este de Texas ha sido descrito durante mas
de un ee Muchos autores sue atr pee este patron a los niveles rel altos de humedad
en el suelo disponible para los ae les en areas de suelo arenoso, y por el contrario,
niveles inadecuados die humedad en el suelo para el crecimiento de arboles en suelos calcareos. Sin
embargo, esta explicacion no es consistente con las observaciones actuales de invasion rapida de
suelos calcareos por vegetacion arborea. aiaeneee una hipotesis alternativa, que la distribucion
historica de las praderas y bosques en el centro-norte y este de Texas puede ser explicada por las
variaciones dependientes del ace en la biomasa de las gramineas y las diferencias resultantes en la
frecuencia e intensidad del fuego.
INTRODUCTION
Conditions at the time of European Settlement, Approximately 1800 A.D.
The vegetation of North Central and East Texas at the time of European settle-
ment (hereafter referred to as presettlement vegetation) was characterized by
well-defined zones of tallgrass prairie and oak woodlands/savannas (Fig. 1).
SIDA 20(3): 1139-1153. 2003
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DIGGS AND SCHULZE, SOIL-DEPENDENT FIRE FREQUENCY 1141
From west to east, the prairie bands are the Grand Prairie (composed of the
Fort Worth Prairie and the Lampasas Cut Plain), the main belt of the Blackland
Prairie, the San Antonio Prairie, and the Fayette Prairie, whereas the oak wood-
land/savanna belts are the West Cross Timbers, the East Cross Timbers, and
three bands of Post Oak Savanna. These were highly recognizable areas of veg-
etation, with the Cross Timbers appearing on the earliest maps and the
presettlement prairies described by numerous explorers, early settlers, and re-
searchers as almost devoid of trees (e.g. Gregg 1844; Brooke 1849; Parker 1856;
Dyksterhuis 1946; Thomas 1962; Correll & Johnston 1970; Hatch et al. 1990;
Diggs et al. 1999; Telfair 1999; Francaviglia 2000). Hereafter we refer to these
two vegetation types as Prairies and Woodlands. The current climate of the re-
gion, presumably relatively unchanged since presettlement times, is character-
ized by wet springs, dry summers with occasional thunderstorms, moderate to
strong winds, and periodic multi-year droughts. As in many areas of central
North America, prairie fires were common prior to settlement (Stewart 1951;
Komarek 1965, 1966; Wells 1970; Wright and Bailey 1982; Collins & Wallace
1990; Bragg 1995). The relative frequency of ignition by lightning versus Native
Americans is unclear.
The Woodlands represent the westernmost extension of the eastern decidu-
ous forest. In general, the vegetation was composed of an oak overstory with tall
grasses. The grasses were dominated by Schizachyrium scoparium (little
bluestem), with Andropogon gerardii (big bluestem) and Sorghastrum nutans (In-
dian grass) as lesser dominants. While varying locally, the woody vegetation was
in general dominated by two trees, Quercus stellata (post oak) and Quercus
marilandica (blackjack oak). The tree density of the oak woodlands was variable,
ranging from quite open to dense thickets. Some early accounts described wood-
lands through which wagons could easily pass (e.g. Marcy 1853, 1866), while oth-
ers described almost impenetrable thickets (e.g. Kendall 1845). Gregg (1844) ob-
served a variety of tree densities, noting that, “Most of the timber appears to be
kept small by the continual inroads of the ‘burning prairies; for, being killed al-
most annually, it is constantly replaced by scions of undergrowth; so that it be-
comes more and more dense every reproduction. In some places, however, the oaks
are of considerable size, and able to withstand the conflagrations.”
The presettlement Prairies were vast grasslands dominated by
Schizachyrium scoparium (little bluestem), with Andropogon gerardii (big
bluestem) and Sorghastrum nutans Undian grass) as lesser dominants, and
woody vegetation generally limited to areas along the larger watercourses, as
scattered mottes, or associated with locations that were protected from fire, such
as mesas and buttes (Smythe 1852; Parker 1856; Hill 1901; Diggs et al. 1999). For
example, Hill 887) described the Grand Prairie as “a prairie region, utterly
destitute of timber” and Kendall (1845) wrote, also of the Grand Prairie, “As far
as the eye could reach ..., nothing could be seen but a succession of smooth,
1142 BRIT.ORG/SIDA 20(3)
gently-undulating prairies.” Early accounts of the Blackland Prairie were simi-
lar. Smythe (1852) described the eastern edge of the Blackland Prairie as having
“..a view of almost boundless Prairie stretching to the north, as far as the eye
could reach...” and further, as “nearly destitute of trees.”
Soils of North Central and East Texas
Most soils of the Prairies of North Central and East Texas are derived from lime-
rich Upper Cretaceous rocks which weather to form soils with substantial lev-
els of clay. Outlying segments of the Blackland Prairie (Fayette and San Anto-
nio prairies) have soils developed from younger Tertiary age deposits. While
the majority of Tertiary deposits in East Texas are sandy in nature (ie., those
supporting the Pineywoods and Post Oak Savanna), those underlying Prairie
(e.g., Fleming, Oakville Sandstone, and Cook Mountain formations) in general
have a relatively hig
play the gilgai microtopography so typical of certain high clay soils
(Launchbaugh 1955; Smeins &@ Diamond 1983; Miller G Smeins 1988). Further,
where clay lenses are found in other geologic strata outcropping in isolated pock-
ets of the Cross Timbers, Post Oak Savanna, and Pineywoods, areas of prairie
vegetation can again be found (Dyksterhuis 1948; Hill 1991). It is thus clay that
appears crucial in the development and maintenance of the grassland vegeta-
=
—
h clay content and in some cases develop soils which dis-
tion characteristic of the Prairies. In some cases clay is abundant throughout
all soil horizons, while in others there is a clay-loam or loam surface layer—all
the Prairie soils, however, have significant amounts of clay (Godfrey et al. 1973;
Diamond & Smeins 1985). Conversely, the Woodlands are developed in general
from Cretaceous and Tertiary sandstone rocks of such geologic layers as the
Antlers/ Trinity, Woodbine, Carrizo,and Wilcox,and can be generally described
as sandy (Sellards et al. 1932; Hartmann @ Scranton 1992).
Prevailing Explanation for the Distribution of Prairies and Woodlands
As discussed above, the presettlement distribution of the Prairies corresponds
to the distribution of limestone parent material overlain by alkaline soils with
a high clay content, while the presettlement Woodlands occurred on either
sandy, slightly acidic soils or, in the westernmost part of the region, on gravelly
and rocky substrates (Sellards et al. 1932; Dyksterhuis 1948; Diggs et al. 1999).
The striking correspondence of vegetation and soil types led early writers to
propose that the distribution of Prairies and Woodlands was due to the differ-
ent water holding capacities of clays and sands. While the basic hypothesis has
been stated many ways, its essence is that greater infiltration and water stor age
at depth in sandy soils permits tree survival, while clay soils do not support
trees because of inadequate infiltration of water (and reduced root penetration)
due to the clay’s relative impermeability.
Hill (1887) appears to have been the first to propose this hypothesis. He
argued that,
DIGGS AND SCHULZE, SOIL-DEPENDENT FIRE FREQUENCY 1143
“The reason why the timber confines itself to [the sandy soils is that they]... afford a suitable
matrix for the penetration of the roots of trees, and a constant reservoir for moisture, thus fur-
nishing two of the greatest essentials to forest growth. ... The barrenness of the prairies, so far as
forest growth is concerned, is owing to the absence of the requisite structural conditions for
preservation of moisture, as well as the excess of carbonate of lime in their soils.”
Subsequent authors reiterated Hill’s contention. For example, Tharp (1926) sug-
gested that the sandy soils increased available soil moisture for tree growth.
Weaver and Clements (1938) argued that, “... the oaks ... have been able to main-
tain themselves against the competition af the grasses by virtue of the favor-
able chresard of the sandy soil.” Dyksterhuis (1948) summarized much of this
information, and apparently agreed with the general idea of sandy soil-mois-
ture availability favoring the growth of trees. Allred and Mitchell (1955) reiter-
ated these ideas:
a
“Sandy, gravelly and rocky areas ...also improve local soi
g
r
i
he grassland
co
formation. Rainfall intake is high on these soils, which also wield up moisture to plants more
readily and more completely than do heavier soils. ... In these areas, soil-moisture conditions
ve been improved sufficiently over the climatic soul so that trees or shrubs, p ‘aan
oaks, form savannah with the understory composed of grassland dominants of the True Prairi
This reasonably intuitive explanation for the historic distribution of the Prai-
ries and Woodlands has been nearly universally accepted, including by recent
authors (e.g., Diggs et al. 1999; Francaviglia 2000).
However, the hypothesis that the Woodlands are restricted to sandy soils
because of inadequate moisture availability on clay soils cannot be correct be-
cause, in fact, trees occur extensively on the clay soils. As noted above, early
accounts noted woody vegetation along the larger watercourses and as isolated
mottes or clumps of trees in scattered locations on the clay soils of the Prairies.
More impressive, however is the current invasion of trees onto extensive up-
land areas of clay soil that can be widely observed throughout the Prairies (e.g.,
Launchbaugh 1955; Smeins & Diamond 1986). Indeed, the few remaining prai-
rie remnants require active management to prevent loss to invading woody veg-
etation (Smeins & Diamond 1986). In the absence of fire, mowing, or some other
suitable disturbance, trees such as Juniperus virginiana (eastern red cedar),
Gleditsia triacanthos (honey-locust), Celtis laevigata (hackberry), Maclura
pomifera (bois-d’arc), Prosopis glandulosa (mesquite), and Ulmus crassifolia (ce-
dar elm) rapidly invade the native prairie and cause the vegetation to convert
toa thicket and then a woodland/forest. A glance from the road along almost
any major highway on the present day Blackland Prairie provides abundant
evidence of rapid invasion by woody species in the absence of disturbance (Fig.
2). In areas no longer cultivated or otherwise disturbed, the invasion by trees
can be observed within a relatively few years—this is particularly obvious in
the numerous areas that were cultivated until relatively recently. It therefore
seems clear that there is sufficient moisture for tree growth on the clay soils and
1144 BRIT.ORG/SIDA 20(3)
Fic. 2.0 £DI--Ll IDpatetn: la L ° . . L 1 * r +f
WUOUOU
conditions (photo by W. Cole Weatherby
—
some other mechanism or mechanisms must have been responsible for the his-
torical difference in the vegetation on the clay Prairies and sandy Woodlands.
It should be noted that the woody species present on the modern day Prai-
ries and Woodlands are largely different, probably due to the different soil re-
quirements of the various species, and we do not view woody plant encroach-
ment on the Prairies as simply an expansion of the Woodlands. Nonetheless,
woody vegetation is now extensive on many formerly Prairie areas. The woody
species currently encroaching on areas of Prairie vegetation were probably not
in general major components of the adjacent presettlement Woodland vegeta-
tion. Rather, they are probably species that in presettlement times were present
in low numbers on the Prairies themselves near streams or where topography
made fire unlikely. On the other hand, some species (e.g., Juniperus ashei, J.
virginiana) are at present highly invasive on both the Prairies and Woodlands.
HYPOTHESIS
Soil-dependent fire frequency
We propose that the presettlement distribution of Prairies and Woodlands in
North Central and East Texas was not due to insufficient moisture for tree
growth on clay soils, but rather to differences in fire frequency on different soil
DIGGS AND SCHULZE, SOIL-DEPENDENT FIRE FREQUENCY 1145
types. We hypothesize that higher fuel quantity on clay soils increased the fre-
quency and intensity of fire, and that fire in turn suppressed the growth of trees.
Prairie fires are fueled primarily by grasses, as opposed to forbs or woody veg-
etation, so an increase in grass biomass leads to an increase in the quantity of
fuel. We predict that grass biomass is typically higher on clay than on sandy
soils due to better moisture and nutrient availability at the shallower rooting
depth of grasses versus woody plants.
This situation would represent two alternative positive feedbacks. High fuel
quantity on clay would encourage fire, which would suppress woody vegeta-
tion and under certain conditions (e.g., depending on season of burn—Howe
1995) stimulate subsequent grass growth (by removing dead biomass which
hinders new growth), thereby maintaining high fuel quantity. Low fuel quan-
tity on sand would reduce the chance of fire, which would foster invasion by
trees that would then further suppress grass biomass (e.g., by shading or other
competition) and the subsequent frequency and intensity of fire. These alter-
native feedbacks would lead to alternative stable states, Prairies and Woodlands
(Fig. 3). This particular hypothesis for the distribution of Prairies and Wood-
lands is consistent with the more general conclusions of Scholes and Archer's
(1997) review of tree-grass interactions around the world. They write that “Moist
fertile environments le.g. our Prairies] support a vigorous grass growth that, if
not grazed, leads to frequent intense fires... Semi-arid environments on sandy,
low fertility soils [e.g. our Woodlands] are seldom treeless.”
Soil type (and its effects on grass biomass) is not the only variable that af-
fects the frequency or intensity of fire. Whether a fire ignites, how hot it burns,
and its ability to spread also depend on a number of other variables including
frequency of ignition events, season of year, rainfall, humidity, wind speed, to-
pography, and grazing (some of which also affect grass biomass). These vari-
ables would combine in a stochastic manner to increase or decrease the fre-
quency and/or intensity of fire in any given location at any particular time.
For example, fire frequency and intensity would be low during a wet summer,
but the resulting grass growth could combine witha subsequent windy drought
to increase fire likelihood and intensity the following year. Thus, the system-
atic effect of soil type on fire would be increased or decreased at any given loca-
tion at any particular time by the net effect of these other variables, with the
result being substantial variation in the time since the last fire in different ar-
eas of both the Prairies and the Woodlands. When this stochastic variation is
taken into account, the fire-frequency hypothesis can explain not only (1) the
historical distribution of Prairies and Woodlands, but also: (2) the historically
dominant tree species of the Woodlands; (3) the historical occurrence of iso-
lated groves of trees on clay soils; (4) the present invasion of Prairies by trees;
and (5) the difference between the historically dominant tree species of the
Woodlands and the species that are increasing in abundance on the Woodlands
and invading the Prairies today (e.g., Juniperus species).
BRIT.ORG/SIDA 20(3)
1146
Sandy or rocky soil Clay soil
Low fertility & low High fertility &
shallow moisture shallow moisture
Low grass High grass
biomass biomass
Low potential High potential
to carry fire to carry fire fepeansdieell
organic matter
Shade
Low probability of High probability of
frequent or intense fire frequent & intense fire
High tree High tree
seedling/sapling educa’ seedling/sapling Increased
eurvivel grass vigor mortality grass vigor
Fic. 3. Flow diag howing two f feedback t de atnativo-ckahlacksk
The Woodlands were historically dominated by oaks, species with thick
bark, stubby branches, and the ability to resprout from roots, features that im-
l extended periods without fire
part resilience to grass fires. Further, occasiona
re isolated groves of
on the Prairies would have allowed the establis
trees that were observed by Western settlers. Once established, such groves
would have been unlikely to burn due to the suppression of grass growth by
—
hment of t
the trees and the resistance of large trees to fire.
The present invasion of the Prairies by trees can be explained by a lack of
fire that has resulted from intentional fire suppression plus numerous and ex-
sy human activities (roads, cultivated
tensive fire breaks that have been created |
fields, overgrazed areas). Moreover, once this process begins, any Prairie area
that becomes substantially invaded by trees would lose grass biomass and come
to serve as an additional firebreak, thereby further reducing the likelihood of
present tree invasion of many
fire on adjacent remaining prairies. Finally, the |
lands understory)
(but certainly not all) areas of the Prairies (and of the Wood
is dominated by Juniperus virginiana (eastern red cedar) and Juniperus ashei
fs
iu
(Ashe’s juniper), species that are sensitive to fire (easily scorched/ignited an
unable to resprout from roots).
DIGGS AND SCHULZE, SOIL-DEPENDENT FIRE FREQUENCY 1147
The key assumption of our hypothesis is that the difference in grass biom-
ass was sufficiently higher on the Prairies than the Woodlands to raise the like-
lihood, frequency, and intensity of fire on the clay soils of the Prairies compared
to the sandy or rocky soils of the Woodlands. In addition, our hypothesis leads
to the prediction that patches of open grassland on sandy soil are very rare
except in instances of active management (e.g., suppression of woody species)
or immediate proximity to clay soils (which would result in more frequent fires
due to closeness to the fire-prone prairie vegetation).
We have not measured grass biomass on intact remnants of the Prairies
and Woodlands, and therefore do not have the data necessary to estimate dif-
ferences in a number of variables associated with fire (e.g., frequency, intensity,
extent, pattern, season, etc.). Further, while some biomass and fuel loading data
are available in the literature (e.g., Johnson @ Risser 1974; Engle & Stritzke 1995),
we have been unable to find directly comparable data for the area of study. How-
ever, substantial indirect evidence is consistent with the assumption of higher
grass biomass on clay soils. First, during dry periods clay soils generally hold
more water at grass rooting depths than do sandy soils. This is due to the rela-
tively large surface areas of the individual clay particles and the large number
of very small pores which act as billions of capillary tubes collectively holding
large amounts of water (Vankat 1979). The result of this increased water hold-
ing capacity is that plants rooted in such soils may continue active growth much
later in the dry season than plants rooted in coarser soils (Daubenmire 1974,
Burgess 1995; McAuliffe 1995; Tucker 1999; Greeves et al. 2000; Ball 2001). Fur-
thermore, undisturbed Blackland soils form gilgai, microtopographical surface
features that function like shallow basins, increasing water retention during
heavy rains (Hayward & Yelderman 1991; Diamond & Smeins 1993). Early set-
tler accounts and observations of existing prairie remnants (e.g., the Nature
Conservancy's Clymer Meadow preserve in Hunt County, the Matthews-
Cartwright-Roberts Prairie in Kaufman County, and Austin College’s Garnett
Prairie in Grayson County) suggest that these “hog wallows,” (as they were
known to early settlers) were abundant on Vertisols of the presettlement Black-
land Prairie. Temporary water storage in gilgai depressions of one-half acre foot
of water per acre of flat prairie have been estimated. As much as six inches of
rain could be temporarily trapped in these structures before runoff began (Hay-
ward & Yelderman 1991). Meanwhile, the high surface area and negative sur-
face charges of clay particles give clay soils high cation exchange capacity. This
allows these soils to hold more ionized minerals or nutrients, including those
essential for plant growth (Foth 1990; Whitehead 2000; Harpstead et al. 2001;
O'Connell 2001). It is therefore not surprising that indirect evidence, such as
agricultural productivity, suggests that the Blackland clay soils were among
the most fertile soils west of the Mississippi River (Haywood & Yelderman 1991).
In addition, the high below-ground biomass of Prairie vegetation serves to con-
1148 BRIT.ORG/SIDA 20(3)
tinually add organic matter to the soil, thereby functioning as a positive feed-
back mechanism to increase fertility and water holding capacity (in part due
to the surface area provided by the additional organic material). Conversely,
sandy soils have larger pores that allow water to drain more easily. They not
only dry earlier during dry periods but “the more water that percolates through
the soil, the more nutrients are washed out—particularly nitrogen, potassium
and sulfur” (Tucker 1999). Therefore, soils that are high in sand, like those of the
Woodlands, are often poor for plant growth since they are relatively infertile
and often too well-drained (Vankat 1979).
Confounding factors
Several other factors have the potential to operate synergistically or in opposi-
tion to the soil-mediated fire frequency hypothesis in influencing the balance
between grass-dominated and woody vegetation. Of particular importance is
the grazing regime. Van Auken (2000), lor example, has identified high levels
of herbivory by domestic animals as the primary cause of brush encroachment
—
in the semiarid grasslands of the southwestern U.S. This influence is probably
the result of acombination of disturbance and reduced fire frequency. The graz-
ing regime during presettlement times was certainly very different than at
present. Instead of domestic animals continuously confined to limited areas of
pasture, the primary herbivores were large dense herds of migratory bison free
to move over vast distances. The grazing regime would thus have been extremely
i
—
irregular both temporally and spatially. While a large herd could move through
an area of Blackland Prairie and crop the vegetation very short (thus prevent-
ing fires), in some seasons or years a given area would probably be missed en-
tirely leaving large amounts of standing biomass (heavy fuel load for fires).
Under such conditions fire would not be expected every year, nor would it be
essential every year for the maintenance of the grassland. Rather, fire would be
critical only with enough regularity to prevent trees from growing large enough
to become invulnerable to grass fires.
Another grazing related influence is the role of dense herds of grazers on
controlling woody vegetation and stimulating grasses. Savory (1998) has stressed
the importance of herding grazers on maintaining grasslands and has empha-
sized the extremely different consequences of typical human-controlled graz-
ing regimes (long periods of exposure to low densities of animals) and natural
systems (large migratory herds at high density due to threat of predation). He
has noted that extremely high densities of grazers (and thus damage to woody
plants) may be of critical importance in preventing encroachment of woody
vegetation and shifting the balance in favor of grazing adapted grasses. A dif-
ferent impact of grazers is that seed dispersal may be of major importance in
the invasion of grasslands by certain woody species. Data of Brown and Archer
(1999) suggest that rates and patterns of seed dispersal may be the primary de-
DIGGS AND SCHULZE, SOIL-DEPENDENT FIRE FREQUENCY 1149
terminants of encroachment by mesquite (Prosopis glandulosa) on present-day
landscapes in semi arid regions of the American southwest.
Still another confounding factor is the competition for resources between
grasses and woody species in an intact prairie ecosystem. Under certain condi-
tions (e.g., of soil, water, grazing, etc.) on an intact prairie, grasses might be able
to out compete woody species or at least resist their invasion, with or without
the influence of fire. However, the impact of fire and grazing are of critical im-
portance since under natural conditions it is unlikely that either of these fac-
tors would be absent for any significant length of time.
CONCLUSION
The soil-mediated fire frequency hypothesis is merely a particular case of a
group of mechanisms that, by reducing the frequency and intensity of fire, en-
able trees to grow where grass would otherwise dominate. The fire induced state
of grasslands on the Prairies is therefore apparently destabilized when fire is
suppressed for any of a number of reasons. In addition to the hypothesized ef-
fect of soil on grass biomass, other variables that can hinder fire and thus allow
trees to invade include precipitation, grazing, and topography (Collins &@ Wal-
lace 1990; McPherson 1995). Trees invade when rainfall prevents fire through-
out the year (e.g., in areas of eastern deciduous forest in the eastern U.S). Like-
wise, reduced grass biomass from grazing on the Prairies reduces fire likelihood,
which in turn enables invasion by trees (Smeins et al. 1982; Scholes & Archer
1997; Van Auken 2000). As noted by Van Auken (2000), “the driving force [for
brush encroachment] seems to be chronic, high levels of herbivory by domestic
animals. This herbivory has reduced the aboveground grass biomass, leading
to the reduction of fine fuel and a concomitant reduction or complete elimina-
tion of grassland fires. This combination of factors favors the encroachment,
establishment, survival and growth of woody plants.” Finally, where relief is
extreme, as on scarps or cliffs, woodlands are often present. This is due to both
the thin rocky soil (and hence low grass biomass) and the topography (e.g.,
abrupt scarps), which often creates natural firebreaks (Wells 1965, 1970; Axelrod
1985). Thus, any factor that reduces fire likelihood, frequency, or intensity can
be expected to allow trees to invade grasslands in areas where there is suffi-
cient moisture for tree growth. The soil-mediated fire frequency hypothesis for
the historical distribution of the Prairies and Woodlands of North Central Texas
and East Texas is consistent with both historical vegetation patterns and re-
cent changes in the vegetation, and is simply a special case of a generally ac-
cepted explanation for tree-grass interactions.
ACKNOWLEDGMENTS
Thanks to Steven Goldsmith for insightful comments on the manuscript, Rob-
ert J. George for valuable assistance with the figures, David M. Engle and Steve
1150 BRIT.ORG/SIDA 20(3
=
Windhager for reviewing the manuscript and making helpful suggestions, and
W. Cole Weatherby for the photograph.
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1154 BRIT.ORG/SIDA 20(3)
BOOK NOTICES
Howarb GARRETT. 2002. Howard Garrett’s Texas Trees. (ISBN 0-89123-076-9, hbk.)
Taylor Wilson, 1900 West Loop South, Suite 1900, Houston, TX 71027, USA,
(Orders:713-980-1950, fax 713-980-2429, 800-570-0082, wwwtaylorwilson.com).
$29.95, 253 pp., color throughout, 7" = 10".
Texas is a big state with a wide range of climate, soil and water quality, and knowing when to plant
the proper tree is crucial. Howard Garrett’s Texas Trees is a handy reference for everyone - rom your
botanist. According to Dr. Garrett “This book is about hort
as well as my opinions on weather and how to use the various trees available in Texas.”
basic gardener to degreec
Everything needed to identify over 100 varieties of Texas trees is here in this one source—along
with easy-to-follow directions for se
ecting, planting, and maintaining your trees. Howard takes a
very common sense approach (as with all his books) to planting, propagating and all things trees.
uded in the book is an in-depth Tree canis y with full details sons great color photos) as
as descriptions of natural habitats, ident t maintenance, propagation, prob-
—
=)
2)
wel
lems, — lowers, fruit, and foliage. Also included is a hi ae little a of pests, diseases, fun-
guses al > treatment methods, like the pon gis vee am, which Garrett developed.
Insummary, | would like to recommend < g this book to your librar : I'san excellent refer-
=
ence for native and introduced trees to the Been in —_ be a great reference guide for an in
the botanical world. —Asha McEl fish, Bot te of Texas, 509 es Street, Fort Worth,
[X 76102-4060, U.S.A
MicHAeL A. ARNOLD. 2002. Landscape Plants for Texas and Environs, Second Edition.
(ISBN 1-58874-153-2, hbk.). Stipes Publishing L.L.C., 202-204 West University
Ave. P.O Box 526 Champaign, II]. 61824-0526 (Orders: 217-356-8391,
stipes@soltec, wwwstipes.com). $68.00, 1088 pp., line drawings, maps, 8
Lf?” et".
Dry, but well-researched and to the point is how I would best describe Arnolds newest book, Land-
scape Plants for Texas and Environs. According to the book “The text was developed as a reference for
nursery and landscape professionals and as a study aide lor students of Texas landscaped plants.”
book probably won't win any design award, as the pages are laid out it in a plain black and
white blocked format with aon ine drawings and species descriptions; no color photographs are
found in the book except o
For each species eee it includes: Other Common Names, Family, Cold Hardiness, Foliage,
Flower, Fruit, Stem /Bark, Habit, Cultural Requirements, Pathological Problems, Ornamental Diseases,
Limitations & Liabilities, Landscape Utilization, Native habitat, and Related Taxa.
Personally I find this book to be lacking in the attention grabbing area of color, but it is easy to
read, well-researched, and you can find what you're looking for quickly. Definitely not a book you
puton your colfee table to show off, but one you will probably find yourself wanting when research-
ing the Texas flora for landscape plants.—Asha McElfish, Botanical Research Institute of Texas, 509
Pecan Street, Fort Worth, TX 76102-4060, U.S.A.
SIDA 20(3): 1154. 2003
FLORA AND VEGETATION OF THE
MOHAWK DUNES, ARIZONA
Richard Stephen Felger Dale Scott Turner = Michael F.Wilson
Drylands Institute The Nature Conservancy Drylands Institute
2509 North Campbell, #405 1510€. Fort Lowell 2509 North Campbell, #405
Tucson, AZ 85719, U.S.A. Tucson, AZ 85719, U.S.A. Tucson, AZ 85719, U.S.A.
rfelger@drylands.org
ABSTRACT
One-hundred twenty-two species of seed aa Fe presenting 95 genera and 35 families are docu-
mented for the 7,700 ha Mohawk Dune Field and di located in Yuma County,
Arizona, USA. Three major habitats were studied: dunes, adjacent sand foie and playa. The dunes
(including interdune swales) support 78 species, of which 13 do not occur on the adjacent non-dune
habitats. The adjacent non-dune habitats (sand flats and playa) support 109 species, of which 43
were not found on the dunes. The total flora has 81 annual species, or 66% of the flora. The dune flora
has 63 annual (ephemeral) species, or 81% of the flora—one of aa nee Dereon ee: fe)
my
- annuals
among any regional flora. Of these dune anes 33 species (84% g the cool season.
No plant taxon is endemic to the Mohawk region. There are 8 dune, or cad mae cied para
Cryptantha ganderi, Dimorphocarpa pinnatifida, Dicoria canescens, Ditaxis serrata, Pleuraphis
rigida, Psorothamnus emoryi, Stephanomeria schottii, Tiquilia plicata—all of which are found on
Two of them (C. ganderi and S. schottii) are of limited distribution, especially
in the USA, and have G2 Global Heritage Status Rank. There are four non-native species in the dune
flora (Brassica tournefortii, Mollugo cerviana, Sonchus asper, Schismus ae “ ane Brassica
and Schismus seem to pose serious threats to the dune ecosystem at this time. The Mohawk Dune
field is within the Lower Colorado Valley phytogeographic region of the Een Desert and located
within the Barry M. Goldwater Military Range. This unique dune system, largely unaffected by hu-
man activities, is rec minended| 2 as a site for baseline studies of plant migrations and population dy-
namics affected by al global cli ge.
RESUMEN
Ciento intid 1 | Sfi 5 ger 1eLros y 35 familias se documentan
para las 7.700 ha del Mohawk Dune Field ysusa irecedores en el condado de Yuma, Arizona, USA.S
estudiaron tres grandes habitats: dunas, arenales planos, y playas. Las dunas (incluyendo -
fern unitate tienen 78 ESPRCIES, ge las ane 13 no se dan en los habitats no dunares
adyace | laya) tienen 109 especies, de las
que pe no se encontraron en las ony: as. La flora total tiene 81 especies anuales, 0 el 66% de la flora. La
flora dunar oel 81% de la flora—uno de i Ss porcentajes mas altos
de anuales en las floes regionales. De estas anuales dunares, 53 (84%) se desarrollan durante la
estacion [ri
DUNE SYSTEMS AND ENVIRONMENT
Dunes inspire thoughts of barren desert, windblown and inhospitable. Yet dunes
can support substantial and unique plant life. Sand dunes occur around the globe
SIDA 20(3): 1155-1187. 2003
1156 BRIT.ORG/SIDA 20(3)
as both coastal and inland systems, and are not restricted to deserts but are often
prominent features of major deserts. Dunes are relatively rare in the Americas,
covering about 0.5% of the arid zones in North America (Bowers 1982, 1984). The
major Sonoran Desert dune fields are the Gran Desierto Dunes of northwestern
Sonora and the related Algodones Dunes of southeastern California. Hot-desert
dune plant communities must cope with an array of challenging growth condi-
tions, including burial, root exposure, wind shear, and low nutrient levels. As a
result, dune floras are frequently characterized by high levels of endemism and
specialized adaptations (Bowers 1982). Despite these challenges, Shreve (1951:127)
pointed out that “Sand..is a particularly favorable soil for the ephemerals of the
cool season, because of the deep infiltration of moisture and the rapid warming
of the surface early in the season and early in the day.”
This publication reports the composition and characteristics of the flora
and vegetation of the Mohawk Dunes, reflecting accumulated data from nu-
merous visits to the area by the authors spanning more than five years, as well
as herbarium records of other collectors. The first section describes dune sys-
tems and the physical environment, management, and global significance of
the Mohawk Dunes in particular. The second section is a description of the
Mohawk Dune vegetation and flora, and the third catalogues its flora, contain-
ing species accounts for each of the 123 species of the dune system and the im-
ou
mediately adjacent desert flats.
PHYSICAL CHARACTERISTICS OF THE MOHAWK DUNES
The Mohawk dune field is located about 65 km east of Yuma, Arizona, on the
eastern side of the Mohawk Valley in Yuma County (Broyles et al. 1997). The
region is within the Lower Colorado Valley phytogeographic subdivision of the
Sonoran Desert (Shreve 1951). The Mohawk Valley is roughly 28 km wide, form-
ing a nearly flat plain with moderately sandy soils. Steep, rocky mountain fronts
bound the valley to the east and west, similar plains stretch far to the south,
and the north end is bounded by agricultural fields along the now usually dry
Gila River. The valley lacks any well-defined central drainage channel, with
the mountain fronts draining into ephemeral stream channels, which rapidly
shrink to insignificance. The elongated dune system runs parallel to, and west
of the Mohawk Mountains, trending northwest to southeast (Fig. L). Small, scat-
tered outlier dunes occur in the San Cristobal Valley east of a pass in the Mohawk
Mountains. The Mohawk dune field is about 32 km long and varies in width up
to 3.2 km wide, covering approximately 7,770 ha. Maximum elevation is 207 m
near the center of the dunes, rising about 55 m above the surrounding desert.
The underlying desert floor rises gently from an elevation of 116 mat the north
end of the dunes to 192 m at the south end (Smith @ Bender 1973).
The dune contours consist of a fairly regular series of depressions surrounded
FELGER ET AL., FLORA , ARIZONA
1157
~~ Dateland
Tacna ae
ol -
ee e
‘ a)
: ¢ >
| % Ye
Goldwater Range 4 [fies By
Boundary %, 1 %
2\% “1%
@ \% f
S whe
44
7
N i ,
a
ie 3 i
Oi 2. 0.2 4 Miles a
Fic. 1. Location of Mohawk Dunes in sout! Ari
by ridges, resembling a honeycomb in aerial view. Swales—low areas between
dunes—and the surrounding crests form microhabitats with significantly differ-
ent plant and animal communities. The crests are characterized by shifting or
slightly stabilized sands and a dominance of sand-adapted species. In contrast,
the swales have soil textures and species assemblages that more closely resemble
the desert surrounding the dune field (Turner et al. 1997a). The swales are also
notable for having a much more stable surface. This is largely maintained by a
well-developed microbiotic crust dominated by the cyanobacterium Microcoleus
vaginatus and the lichen Collema sp. Jane Belnap, pers. comm.).
The Mohawk Dune system appears to have been in place for an extended
period. Extensive layers of calcium carbonate are evident on swale slopes, and
we have found numerous fossil remnants of tortoises and other vertebrates. The
dune sand is composed mainly of quartz and feldspar. According to Bowers
(1986:57), “.. the Mohawk Dunes have garnered all the sand available for their
formation: no more sand is being blown into the dune field.” It seems reason-
able to suspect that an earlier configuration of the Gila River was the source of
sand for these dunes, but that question remains unstudied.
1158 BRIT.ORG/SIDA 20(3)
A distinctive playa (the bed of an ephemeral lake), 1,400 m x 250 m, lies
along the eastern side of the dunes, about 8 km south from the dune field’s north
end (Fig. 1). It appears to fill toa depth of <1 m with a frequency of about once
per decade, most recently in winter 1992-93. Its more common aspect features
sun-baked clay with an extensive network of deep cracks.
Climate
Air temperature regimes for the area are typical of the lower Sonoran Desert,
with mean daily maxima of 40-43°C for June through September, and peaks in
excess of 49°C during midsummer (Sellers & Hill 1974). Surface temperatures
on the dunes reach greater extremes, with June and July maxima averaging
above 65°C in 1996. Relative humidity is generally low, dropping below 10% on
most days in the summer and many days in winter (Turner, unpubl. data).
Annual rainfall averaged 10.54 cm from 1900-195] at the nearest weather
station, Mohawk, ca. 5 km northeast from the north end of the dune field. About
half of the annual total, 4.85 cm, fell during the May-October warm season, pri-
marily during July-September, while 5.72 cm fell November- April (Sellers et al.
1985). Months with no rain are common, and summer storms can deliver a large
fraction of the total annual precipitation in a short period (Sellers & Hill 1974).
Although rainfall data from nearby stations may be indicative of regional
trends and normal year-to-year variation, they may not provide accurate data
on local precipitation. A six-year study in Yuma showed that summer rainfall
can vary by more than 2.5 cm for sites only 1.6 km apart (Turnage & Mallery
1941). Rain gauges at the Mohawk Dunes separated by 9.3 km recorded 2.54
and 0.38 cm for the same period of August through October 1994. The same
two gauges recorded 0.46 and 3.30 cm respectively for a single storm event in
July 1996 (Turner, unpubl. data). Such variation can cause dramatic differences
in vegetative composition and abundance between sites.
—
Mohawk Dunes as a Long-Term Research Site
The Mohawk Dunes and surrounding landscape are part of the Barry M.
Goldwater Military Range, jointly managed by 56th Range Management OI-
fice at Luke Air Force Base and the Range Management Department at Marine
Corps Air Station, Yuma (Felger et al. 1997; Tunnicliff et al. 19860). The western
part of the range including the Mohawk Dunes is managed by the Marine Corps
and lands to the east are managed by the Air Force. Civilian access requires
permission from the military. Military activities have impacted only approxi-
mately 5-12% of the entire 690,000 ha Goldwater Range and have had very little
apparent affect on the dune biota. To date the military has provided substantial
protection from the damaging recreational activities common to other dune
systems in the American Southwest, and the Mohawk Dunes remain largely
unaffected by human activity. However, future military and civilian effects on
the dunes will be determined by management plans now under discussion.
FELGER ET AL., FLORA , ARIZONA 1159
The dunes are a popular destination for the public during spring wildflower
displays, and there is occasional illegal off-road driving. Planning should as-
sure undisturbed areas for monitoring and research, while other areas have ac-
cess but with precautions to insure minimal impact.
Our understanding of global diversity is hindered by the lack of knowledge
of changes in species composition over time so that processes of colonization,
extinction, and evolution cannot be thoroughly described. This underlines the
importance of maintaining areas that remain undisturbed by human develop-
ment and that have protected status or the potential for protection due to land
ownership. The manageable size of the Mohawk Dunes makes them an excellent
candidate as a long-term research site. Careful stewardship of the dune field will
serve to protect their value to our understanding of ecosystem processes.
METHODS
This report reflects accumulated data from numerous trips to the area with
primary collections during 1995-1998. Specimens are at ARIZ, with duplicates
for most species variously going to other herbaria, e.g, ASU, RSA, SD, and the
Yuma District Office of the Bureau of Land Management. We also examined
additional specimens, primarily at ARIZ, ASU, and RSA.
As part of an ecological inventory and monitoring program (Turner et al.
1997 a, b), plant cover sampling was done at two sites in the dune system using
a stratified random sampling design to characterize the crest and swale plant
communities. This involved randomized placement of quadrats along curvi-
linear transects which stayed either on crests or in swales, along transects which
were previously established for a lizard monitoring program. Dune crests were
sampled with a l-km transect at a site on the east side of the dunes near the
playa and two 500-m transects on the west side. Swales were sampled with a
single l-km transect near the playa.
We used quadrats 0.5 x 2 m, to take 200 samples out of a 10-m-wide swath
along each |-km transect, or 100 samples for each 500-m transect, for a repre-
sentative sample of 1 ha for each sample site. Sampling was conducted during
March 1996, following an unusually dry year, and thus measured almost exclu-
sively perennial species.
VEGETATION AND FLORA
The total flora of the Mohawk Dune Region (dune and non-dune habitats) is
comprised of 122 vascular plant species in 95 genera and 35 families. The most
diverse families are the Asteraceae, Fabaceae, Poaceae, and Brassicaceae (Table
1). The most diverse genera are Camissonia with 4 species and two subspecies,
Cryptantha with 4 species, and Ambrosia and Boerhavia each with 3 species.
The growth-form spectrum is shown in Fig. 2. The different life forms are more
1160 BRIT.ORG/SIDA 20(3)
or less evenly distributed between the dunes and sand [lats except that all but
one (Prosopis) of the tree and large-shrub species (microphanerophytes) are
absent from the dunes (Table 2). The total flora has 81 annual or ephemeral
species, or a remarkable 66% of the flora (see discussion below for dunes). In
contrast the ephemeral/annual flora of northwestern Sonora comprises 46%
of the total flora, and across much of the Sonoran Desert the ephemerals make
up about 50% of the total flora (e.g., Felger 2000; Venable & Pake 1999).
During the warmer months, often April and early May, and again with sum-
mer-early fall rains, many of the common ephemerals can be ravaged by cater-
pillars of the white-lined sphinx moth (Hyles lineata) (Fig. 10). Plants favored
by these caterpillars include Allionia incarnata, Boerhavia spp., Euphorbia
abramsiana, Kallstroemia californica, Lupinus arizonicus, and Pectis papposa.
Vegetative cover by perennial plants
Vegetative cover data were gathered during an unusually dry spring, Eight plant
species were found living in 600 quadrats. We found almost no annual plants at
that time. Mean ground cover on the dune crests (“dunes”) ranged from 7-15%,
while that in the swales was about 9% (Table 3). Dominant perennial species on
the crests included Ambrosia dumosa, Aristida californica, Ephedra trifurca,
Pleuraphis rigida, and Psorothamnus emoryi. The swales had a similar suite of
species, with the addition of Larrea divaricata and a strong reduction in
Psorothamnus emoryi.
Habitats
Bowers (1982) found that plant communities on dunes are mixtures of dune
endemics, widely distributed plants of sandy soils, and plants found in adja-
cent, non-dune habitats. The desert flats, or sand flats, and the small playa are
the non-dune habitats adjacent to the Mohawk Dunes.
Dunes.—During years of favorable winter-spring rainfall, such as certain El
Nino years, the dunes support spectacular displays of wildflowers. Among the
most conspicuous are sand verbena (Abronia villosa), desert marigold (Baile ya
spp.), desert cryptantha (Cryptantha angusti tifolia), spectaclepod (Dimorphocarpa
pinnatifida), Arizona lupine (Lupinusar' iZonicus),evening- primrose (Oenothera
deltoides), and Spanish needles (Palafoxia arida). Notably prominent are plants
with dense whitish- or silvery-haired leaves and stems, e.g., Ambrosia dumosa,
Dicoria canescens, and Psorothamnus emoryi. Deep roots are likewise a common
feature, enabling plants to utilize moisture deep in the sand.
The floristic makeup of the dune communities, particularly among peren-
nial species, is highly predictable, with relatively little change in species com-
Psorothamnus emoryi, Tiquilia plicata, Ephemerals—A bronia villosa, Dicoria
canescens, Dimorphocarpa pinnatifida, Lupinus arizonicus, Oenothera plants
higher, shifting (moving or unstable) dunes are: Perennials—Ambrosia dumosa,
—
~
FELGER ET AL., FLORA ARIZONA 1161
Taste 1.Four largest families.
Family Genera Species
Asteraceae 17 2
Fabaceae 9 1]
Poaceae 7 9
Brassicaceae 7 7
60
50 4
&
jon)
|
No. of species
WwW
Oo
i
ho
So
}
we
Len]
4
jon)
pet
TW Ts TN ‘G H C N M E E
Fic. 2. Life-form spectrum of the Mohawk flora. TW = winter spring) or od herophytes ( Is or ephemer-
als): ie summer or nots -season me rophytes: | TN =non phytes; G = geophytes ( ials with m
ground) yt | (p ek 4 g a Rae 4) Near L a 4? . a =
L L { re a L Shit Ihe] Nc Thee L L
or shrub- sized iietennials usually 0.5-2 m tall) M= microphanerophytes (trees or r shrubs usually 2- Br m tall) L=
perennial liana or vine; E = epiphyte.
position across the dune fields. Among the most common and conspicuous on
Aristida californica, Ephedra trifurca, Pleuraphis rigida, deltoides,
Stephanomeria schottii,and many others. Characteristic common plants on low,
partially stabilized dunes as well as other sandy habitats include: Perennials—
Ambrosia dumosa, Aristida californica, Larrea divaricata, Pleuraphis rigida,
Tiquilia palmeri; Ephemerals—Abronia villosa, Brassica tournefortii,
Eriogonumtrichopes, Oenothera primiveris, Schi bicus, Stephanomeria
schottii, and many others.
The dunes support 78 species, of which 13 do not occur on the adjacent
non-dune habitats (Table 2). The dune flora includes 63 annual or ephemeral
1162
BRIT.ORG/SIDA 20(3)
Tasve 2. Distribution of the Mohawk Flora for three habitat types with growth-forms. See Fig. 2 for
explanation of life-form symbols (p = parasitic plants; s = succulent plants). Very rare occurrences
or extirpated species shown in parenthesis (-).
dunes Sandflats playa Growth form
sane ia villosa D S TW
D S TW
Allionia nae. D S TN
Amaranthus fimbriatus D S TS
Amaranthus - me S TS
ra S H
amie pie idea D S C
Ambrosia dumosa D S C
Amsinckia tessellata D S TW
Argemone gracilenta D H
Aristida adscensionis S TN
Arictid, lif ; D S H
Ascle as subulata S Ns
galus insularis Dp S TW
Astragalus nuttalliqnus D S TW
Baccharis salicifolia (P) N
Baccharis sarothroides (P) N
Baileya pauciradiata D S TW
Baileya pleniradiata D S TN
Boerhavia erecta S TS
Boerhavia spicatc D S TS
Boerhavia wrightil S TS
Bout eau aristi golaes D S TS
Bouteloua barbata D S TS
Brassica tournefortii D S TW
Camissonia boothii S (P) TW
Cami ia calif D S TW
Camissonia engnaenenones S TW
p. peeblesii D S TW
Cami ia claviformis ssp. Yumae S TW
Carnegiea gigantea (S) Ms
Castela emoryi S M
Caula ntrhise lasiophy Thrice D S TW
Chaenactis stevioides D S TW
Chorizanthe brevicornu D S TW
oo. ml jag S TW
D S Ns
hoe ie oa ifolia D 5 TW
Cryptantha gander D TW
€ nue maritim S TW
“| pt Intha micrant D TW
eee umbellata D 5 TSp
Dalea mollis D S TN
FELGER ET AL., FLORA ARIZONA 1163
TaBLe 2. continued
dunes Sandflats playa Growth form
Pana discolor t S TN
D TN
Dimerphecapa, pinnatifida D TW
Ditaxis se D S TN
Dithvren rcolifarnica D TW
Echinocactus polycephalus (S) Cs
Ephedra trifurca D S N
Pri t Ai ffi D S TW
Frigqetritm eremiciim S TW
Eriogonum trichopes D S TW
a iam lanosum S TW
dium texanum S TW
Ecce [zia glyptosperma S TW
Bee eee mi nut flora D S TW
D S TS
ephorb ee D S TN
us wislizeni (D) S Ns
FIEGOL arizonica S TW
Festuca octoflor S TW
poe slendens S M
S L
a canescen D 5 TW
Grusonia kunzei (S) Cs
Heliotropium convolvulaceum D TN
Hplint p i Ul) avicu 3) G
Hesperocallis undulata D S G
ne Unanp scale glauca P G
maa S P @
Kallstroemia californica D S Ts
Krameria erecta S C
Krameria grayi S N
Larrea divaricata D S N
Lepidium lasiocarpum D (S) TW
Linanthus bigelovii D S TW
Loeseliastrum schottii D S TW
Lupinus arizoni D S TW
Lycium parishii S N
Malacothrix glabrata D TW
Malvella sagittifolia P G
Mate rina lee D (S?) TN
n alhicarys li B) TW
Mentzelia multiflora D S TW
ollugo cerviana D TS
Monoptilon bellioid D S (P) TW
Muhlenbergia microsperma S TN
1164 BRIT.ORG/SIDA 20(3)
Taste 2. continued
dunes Sandflats playa Growth form
Nama hispidum D S (P)
Nemacaulis denudata D
Oenothera deltoides D
Oenothera primiveris D
Oligomeris linifolia D
Olneya tesota
Orobanche cooperi
Palafoxia arida D
Panicum alatum
Parkinsonia florida
Parkinsonia mic aad
tis pappo
Pe irya heter D
Pectocarya nes
Phacelia ambigua
PI Pal qf ~ 4] if
vylla
Oo
oO
ANNNNNHNNHNHUNHNHUNNYN
Physalis lobata P
Physaria tenella S
Plantago ovata D 5 Pp
Pleuraphis rigida
seer oe (P)
Telialital
Oo
wn
ase Sandals 5a
sorothamnus emoryl
Rafinesquia neomexicana
S his mirc arahircys
UODU0000
Sonchus asper
Sphaeralcea ambigua
ah
S
Steptanthela a longirostris
gS
}
3
=
58
ad
a
og 2
s
=
Stylocline micropoides
Tamarix ramosissima (P) N
Tidestromia hae
Tiquilia plicatc D (S) G
TOTALS: 122 els 78 101 I
5,
a
4
Ca)
species, representing 81% of the dune flora, one of the highest percentages of
annuals among any regional flora. Of these dune ephemerals, 53 species, or 84%
develop during the cool season (41 winter/spring and 12 non-seasonal species),
which supports Shreve’s (1951:127) statement that sand is a favorable soil for
cool-season ephemerals. Non-seasonal ephemerals are those that can germi-
nate and grow with sufficient soil moisture at any time of year (see Felger 2000).
Variation in plant cover and dune vegetation is shown in Figs. 3-5.
FELGER ET AL., FLORA ARIZONA 1165
Taste 3. Average percent ground cover at three localities in the Mohawk Dunes, March 1996.
Species E-side dune E-side swale W-side dune
Ambrosia dumosa 1.56 4.14 9.07
Aristid lif 0.00 0.89 1.28
Ephedra trifurca 0.97 1.12 1.30
Larrea divaricata 0.00 1.16 0.00
Pleuraphis rigida 3.35 1.41 3.50
Psorothamnus emoryi 0.89 0.24 0.08
nui au cata 0.01 0.00 0.01
0.01 0.01 0.02
Total live ie cover % 6.77 8.95 15.25
Playa.—The main expanse of the playa consists of silty-clay soil riddled
with deep fissures. Seventeen species were found on the playa and its margins.
Three herbaceous root-p bundant across the expanse of the playa:
Heliotropium curassavicum, Hoffmannseggia glauca, and Malvella sagittifolia.
Following the occasional times when the playa fills with water, Sphaeralcea
ambigua and other herbaceous plants may become locally abundant. Physalis
lobata, Oligomeris linifolia,and Prosopis glandulosa are locally common around
the margin of the playa. The remaining species are uncommon in this habitat
or on sandy soils at the playa margin. Four woody plants are known from one
or few individuals that probably germinated during the exceptionally wet year
of 1992-93: seep willow (Baccharis salicifolia), desert broom (B. sarothroides),
cottonwood (Populus fremontii), and tamarisk (Tamarix ramosissima). The cot-
tonwood and seep willow perished from drought by the end of 1994 and the
tamarisk by June 1996.
Sand flats. Areas surrounding the dune field vary from rolling hills and
low sand hummocks to occasionally extensive flats of sandy to silty-sand and
clay-sand soils. These habitats, collectively called “sand flats,” support 101 spe-
cies within ca. 1 km of the dune margins (Table 2). Nearly 30% of these species
were not found on the dunes.
Desert shrubs, namely creosotebush and the two bursages (Ambrosia deltoidea
and A.dumosa) are the most common, widespread elements in the landscape. Sev-
eral cacti and crucifixion thorn (Castela emoryi) are widely but thinly scattered
across the landscape, and patchy areas of ocotillo (Fouquieria splendens), ratany
(Krameria erecta), and several other perennials are locally prominent.
The sand flat habitats and vegetation grade into those of the dunes, and
less noticeably into the surrounding valley plains and vegetation. But common
shrubs such as creosotebush (Larrea divaricata) tend to be noticeably less dense
on these sandy areas than the surrounding plains.
1166 BRIT.ORG/SIDA 20(3)
Fic. 3. Mohawk Dunes, during a“ wet spring,” 28 February 1998. Oenothera deltoides (white fl ) Abronia vill
1 ‘AL €] n
P J J / J |
by Thomas R. Jones.
ps
>
Fic. 4. Same view as Fig. 3, but during a drought year, 12 March 2002. Th
deltoides shown in Fig. 3 left foreground. Photo by Thomas R. Jones.
FELGER ET AL., FLORA ARIZONA 1167
ne a: Nottie bart of Mohawk Dunes looking eastward towards Mohawk Mountains, April 1998. Abronia villosa
M forground). Photo by Charles Hedgcock.
J
Comparison to Other Dune Systems
In terms of plant diversity and species composition, the Mohawk Dunes, as ex-
pected, show the greatest affinity to other inland dune systems within the
Sonoran Desert. Of these, the greatest similarity exists with dunes of the Gran
Desierto, Sonora, the largest inland dune system in North America (5,000 km2),
centered approximately 70 km to the southwest of the Mohawk system (Felger
1980, 2000). The Pinta Sands of Cabeza Prieta National Wildlife Refuge, 38 km
to the south are the closest dune system, but these dunes are only a northern
extension of the Gran Desierto, being some 5 km from the northernmost dunes
of that system. The Algodones dunes (709 km) of southeastern California, 90
km to the west, share many plants with the Mohawk system (see WESTEC Ser-
vices 1977). Although no plant is endemic to the Mohawk Dune system alone,
Cryptantha ganderi, Dimorphocarpa pinnatifida, and Stephanomeria schottii
are endemic to dunes and adjacent sand flats of the Lower Colorado region of
the Sonoran Desert.
The Mohawk system has among the highest species richness for its size
1168 BRIT.ORG/SIDA 20(3)
among southwestern dunes. The enormous Gran Desierto dune system has a
‘lora of only 85 species (Felger 2000), amazingly few species for such an im-
mense area. The relatively low species richness probably can be attributed to
the low habitat diversity, extreme aridity, and harsh conditions brought about
by moving sand. The Mohawk Dunes, with less than 2% of the area, supports a
dune flora of 78 species (Table 2). The Algodones Dunes are reported to support
a flora of 97 species (Bowers 1984, interpreting WESTEC Services 1977; this
number, however, cannot be considered wholly accurate based on the available
data). The Kelso Dunes in the eastern Mohave Desert support 75 species (Thorne
et al. 1981), with some overlap with Mohawk Dune species. It is interesting to
note the similarity in species richness among these diverse dune systems.
Species composition of the dominant dune vegetation in the Mohawk
Dunes is similar to that of other Sonoran Desert dune systems (Felger 2000;
Warren & Laurenzi 1987). Five species typically comprise at least 90% of the
perennial vegetative cover on these systems, with Ambrosia dumosa and
Pleuraphis rigida the most abundant, and Psorothamnus emoryi, Ephedra
trifurca, and Tiquilia plicata as common associates.
Absent from the Mohawk Dunes are several dune-endemic species which
are relatively common in the Gran Desierto and/or Yuma Dunes and Algodones
Dunes, including Ascle pias erosa, Croton wigginsti, Drymaria viscosa, Eriogonum
deserticola, Euphorbia platysperma, Helianthus niveus, Heterotheca thiniicola,
Pholisma sonorae, Larrea divaricata var. arenaria, Palafoxia arida var. gigantea,
and Triteleiopsis palmeri (e.g., Felger 1980, 2000; Felger et al. 1997). This may be
due to the relative isolation of the Mohawk Dunes. Eriastrum eremicum, found
on the Mohawk Dunes is not present at the other dune systems; it is more char-
acteristic of Arizona Upland locales and has its southwestern limits near the
Mohawk Dunes.
Non-native Species
Five non-natives species, all native to the Old World, are recorded in the flora:
Brassica tournefortit, Mollugo cerviana, Schismus arabicus, Sonchus asper, and
Tamarix ramosissima. Tamarix is no longer present, and the remaining four
occur on the dunes and all but Mollugo are also found on the sand flats. The
Brassica, Mollugo, and Schismus are common and well established, while
Sonchus asper is widely scattered but not common. Additional invasive weedy
species undoubtedly will arrive on the scene. A number of non-native, weedy
plants are conspicuously absent from the Mohawk flora but are common in
nearby, mostly disturbed areas, e.g., buffelgrass (Pennisetum ciliare (L.) Link)
and tumbleweed (Salsola tragus L.).
A flora with only 4% non-native species is remarkable, since the average
for the Sonoran Desert is about 11.6% (Felger 2000; Wilson et al. 2002). It is not,
however, unusual for regional dune systems. The Mohawk Dunes and the enor-
FELGER ET AL., FLORA ARIZONA 1169
mous Gran Desierto dune system contain only the same four non-native spe-
cies and the abundance patterns are likewise similar (see Felger 2000:21). The
unusually low number of non-natives may be due to the uniqueness, isolation,
and relatively undisturbed condition of the habitat.
Although competition between plants on dunes is reported to be negligible
(Bowers 1982), Brassica tournefortii and Schismus arabicus certainly seem in-
vasive and detrimental to native species (Felger 2000; Wilson et al. 2002). Their
very rapid early growth seems to negatively affect native plants. On the Mohawk
dunes as well as elsewhere in the Sonoran Desert (e.g., Felger 2000), Brassica
rosette-leaves and Schismus plants spread over seedling and juvenile plants of
the apparently slower-growing native ephemerals, possibly smothering or shad-
ing them out, or preventing their germination (Fig. 6)
ANNOTATED SPECIES ACCOUNTS
The flora is listed alphabetically by family, genus, and species. Accepted scien-
tific names, including the variety (var) or subspecies (subsp.), are in bold. Se-
lected synonyms of scientific names are in italics within brackets [-]. Common
names follow the scientific names, and when known are given in English and
Spanish. Brief characterizations are provided for each species, including the
usual flowering time and in most cases the general or most conspicuous color
of the flower (flower color is often omitted for species with small, inconspicu-
ous flowers). Growth and flowering, of course, is almost entirely linked to rain-
fall and soil moisture. During extended drought, such as 2001 and 2002, many
plants fail to grow or flower. More detailed descriptions and distributional in-
formation can be found in many sources, e.g., Felger (1980, 2000), Hickman
(1993), Kearney and Peebles (1960), Wiggins (1964), and the various contribu-
tions to the Vascular Plants of Arizona (Vascular Plants of Arizona Editorial
Committee 2001).
The distributional information is based on field observations and for the
most part verified with herbarium specimens primarily at the herbaria of the
University of Arizona (ARIZ) and Arizona State University (ASU). Specimens
cited are at ARIZ, unless otherwise indicated, and in many cases duplicates,
especially those of Felger, are mostly at ASU, RSA, and SD. The five non-native
species are indicated with an asterisk (*). The area had abundant spring rains
during 1995 and 1998, and abundant summer rains during 1996 and 1997. In
our opinion we likely encountered plants of nearly all of the ey l species
present in the seed bank as well as all of the perennials.
AMARANTHACEAE - Amaranth Family oie Sand flats and dunes. Felger 97-63 et
Amaranthus fimbriatus (Torr.) Benth. ex S. ange 20 ery
Watson. Fringed pigweed; bledo, quelitillo.
Hot-weather annuals; flowers greenish and
n.Carelessweed,
pigweed; bledo, py te ive las aguas. Hot-
weather annuals. Sand flats, especially near
BRIT.ORG/SIDA 20(3
1170
Fic 6 Al +h rors | € AARL Ln 1 L? g | Cc if - late €, Ab (I ee |
rs) ni ee | : ay £, J 4 | H se | distance 4 J 4 L try ae AL
ril 1998. Photo by Charles Hedgcock.
a
Ap
FELGER ET AL., FLORA
ARIZONA 1171
the south end of the dune field. Felger 97-79
tal.
Tidestromia lanuginosa (Nutt.) Standl. subsp.
eliassoniana Sanchez-del Pino & Flores
Olvera. Honeysweet; hierba ceniza, hierba
lanuda. Hot-weather annuals; flowers yellow.
Widespread and common, sand flats and
dunes. Felger 97-71 et al.; Felger 96-149 &
Turner.
ASCLEPIADACEAE - Milkweed Family
ASCieDias supulat Bevis Desert EME SS,
aAlalifl
Perennials with multiple semi-succulent
stems; flowering during warmer months;
flowers waxy, cream- and yellow-white. Oc-
casional in washes in sand flats near north
side of dune field. Turner s.n., 12 Nov 2001.
Funastrum cynanchoides (Decne.) Schlt. var.
hartwegii (Vail) Krings [Sarcostemma
cynanchoides Decne. subsp. hartwegii (Vail)
R.W. Holm; Funastrum heterophyllum
(Engelm.) Standley]. Climbing milkweed;
guirote. Perennial vines; flowering at various
seasons; flowers maroon and white. Occa-
sional in washes in sand flats near the north-
ern part of the dunes. Felger & Turner, 24 Sep
1996, observation.
ASTERACEAE - Daisy or Composite Family
Ambrosia confertiflora DC. Slimleaf ragweed;
estafiate. Herbaceous perennials; flowering
late spring and in fall. Occasional in depres-
sions in sand flats near the dunes. Felger 97-
Ambrosia deltoidea (Torr.) W.W. Payne. Tri-
angle-leaf bursage; chamizo forrajero. Small
shrubs. Common on dunes, sand flats, and
playa margins. Summer dormant. Felger 96-
169 & Turner; Reina-G. 96-213 & Van Devender.
Ambrosia dumosa (A. Gray) W.W. Payne. White
bursage; chamizo. Small shrubs; flowering
September-April. Summer-dormant, new
leaves and flowers may appear as early as
September.Very common on sand flats and
dunes including higher dunes. Fe/ger 97-66
et al.; Felger 96-127 & Turner; Reina-G. 96-215
Van Devender.
Baccharis salicifolia (Ruiz & Pav.) Pers. [B.
glutinosa Pers.].Seep willow; batamote. Leafy
shrubs. About ten plants, mostly 2-2.5 m tall,
along the southern edge of the playa ap-
year of 1992-1993, and dle by the end
of 1994. Felger 96-154 & Tur
Baccharis sarothroides oe Desertbroom;
margo, romerillo. Broomlike shrub
One individual, ca. 2 m tall, at northeast mar-
gin of playa. Felger 96-150 & Turner.
Baileya pauciradiata Harv. & A. Gray ex A. Gray.
Few-flowered d
als; flowers yellow. Dunes and sand flats.
Reichenbacher 601; Salywon et al. 988 (ASU);
t marigold. Spring annu-
urner 95-26.
Baileya pleniradiata Harv. & A. Gray ex A. Gray.
Woolly renee marigold; tecomblate. Annu-
als, mostly sum-
er rains; ae: yellow. Dunes and sand
flats. Felger 98-56 & Evans; Salywon et al. 987
ASU); Turner 95-25, 95-30.
ed stevioides Hook. & Arn. Desert pin-
shion.Cool-season annuals; ce oa
or cream, pink in bud.Dune ially low
dunes, and sometimes on sand flats. ae
98-58 & Evans; Goldberg 77-87, Turner 95-5.
Dicoria canescens A. Gray subsp. canescens.
Bugseed. Annuals; germinating at least with
spring rains, their numbers variously re-
duced by late-spring and early summer
1 folly | Y eeee |
drought, recorded ft leafed-out in August,
spring sometimes with
—
and variously flowering from late October
through December; plants often robust to
1m tall and becoming tumbleweeds after
seeds ripen. Seedlings and juvenile plants
quickly produce a deep taproot and have
narrow, elongated leaves, the older plants
produce broad and shorter leaves. Restricted
to dunes, especially moving sands of higher
dunes. On 24 November 1994, Turner ob-
served plants in full bloom, abuzz with hon-
eybees but no native bees; the honeybees
probably were gathering pollen since the
plants are presumably wind-pollinated. On
23 June 1995, Turner found clipped stems
and leaves scattered around stripped
stems—evidence of herbivory by kangaroo
rats (Dipodomys sp.). Felger 98-59 & Evans;
Reichenbacher s.n., 7 Mar 1980; Turner 94-2.
Eriophyllum lanosum (A. Gray) A. Gray. Woolly
daisy. Small, cool-season annuals; disk flow-
1172
ers yellow, rays white. Sand flats. Felger 98-91
& Evans
Filago arizonica A. Gray. oe arizonica (A,
le Holub]. iets fluffwee nall, cool-
eason annuals areas in ee flats Felger
98-93 & Eva
Geraea canescens lorr. & A. Gray. Desert sun-
flower, desert gold. Coarse spring annuals;
flower heads sunflower-like, bright yellow
Dunes and sand flats. Goldberg 77-89; Turner
95
Isocoma acradenia (Greene) Greene var.
radenia_ [Hap/ a acradenius
oe F.Blake].Alka [ S
shrubs; flowering various seasons,at least in
spring and fall; flowers yellow. Occasional
and localized at the playa margin and sand
flats at roadsides, Felger 96-143 & Turner, 96
155; Reina-G. 96-214 & Van Devender.
Malacothrix glabrata (A. Gray ex D.C. Eaton) A
M. californica DC. var. glabrata A. Gray
ex D.C. Eaton] Smooth desert-dandelion.
Spring annuals; flower heads white with a
pale yellow center. Dune slopes and meee
dunes, scattered t metimes s nally
on. ie 98- Ae Evans; ae her
603; nee
Monoptilon aides (A. Grey) H.M. Hall
Desert star. Small, spring
turning pale lavender with age, the disk vel
low. Sand flats and sometimes on lower
some at playa margin. Felger 98-
1 & Evans, 98-96; Turner 95-21.
Roe arida BL. Turner & Ml. Morris var.
Hoe: spans neSdIe> Non sao nas annu
=}
~
ol
sand flats and dunes at all elevations Some
of the larger, robust plants on the dunes
approach var. gigantea (M.E. Jones) B.L.
Turner & Ml. Morris in having larger flower
heads and achenes. Felger 98-4 et al.; Felger
96-139& Turner; Turner 95-4
Pectis dias Harv. & A. Gray var. papposa
chinchweed; manzanilla de ayer
Ho t-w are annuals; herbage pungently
aromatic; flowers bright yellow. Sand flats
and dunes. Highly variable in size, drought
stressed plants can mature with a single
flower head. Felger 97-56 et al., 97-72; Felget
6-134 & Turner.
neh
BRIT.ORG/SIDA 20(3)
Rafinesquia neomexicana A. Gray. Desert
chicory. ll annuals; flowers white. Dunes,
| and swales al
especially dune olower
dunes and sand flats; mostly growing
through small shrubs such as Ambrosia
oo Felger 98-85 & Evans; Felger 98-12 et al.
onchus asper (L.) Hill subsp. asper. Spiny
een chinita. Winter-spring annuals;
flowers pale yellow. Occasional and widely
scattered in natural areas such as small ar-
royos and depression on sand flats, dune
swales, lower dunes, and rarely on higher
dunes. Felger 98-62 & Evans, 98-95.
Stephanomeria schottii (A. Gray) A. Gray.
Schott's wire-lettuce. Fig. 7. Late spring an-
nuals; flowers white tinged with evict Sea-
sonally abundant and wid
at all elevations and also iow stabilized
dunes of the nearby San Cristobal Valley.
Burgess 5067; Felger 98-16 et al.; Felger 98-63
& Evans; Reichenbacher 596; Turner 95-14. San
ristobal Valley dunes, Anderson 98-40, 98-54
(ASU).
Endemic to dunes and sand flats in the Gran
Desierto in northwestern Sonora and south-
western Arizona in Yuma County (Felger
206 20, pete 979). This species has a Global
e Status Rank of G2 (imperiled
in
ae iby.
Stylocline micropoides A. Gray. Desert
neststraw. Small, spring annuals. Locally and
seasonally common in swales and depres-
sions among dunes, and sometimes on
surrounding sand flats. Felger 98-83 & Evans.
BORAGINACEAE - Borage Family
Amsinckia tessellata A. Gray. Checker
fiddleneck.Cool-season annuals; flowers yel-
low. Common on dunes and sand flat
Turner 95-13.
Cryptantha angustifolia (Torr) Greene.
Narrowleaf cryptantha, desert cryptantha.
Cool-season annuals; corollas white with a
yellow throat.Widespread across the dunes
and sand flats;one of the most common and
ubiquitous spring annuals in the region.
Turner 95-12.
Cryptantha ganderi |.M. Joni neors season
annual sometimes
minute, white; fruits with 2 smooth nutlets,
FELGER ET AL., FLORA ARIZONA 1173
La
=
~
a)
ji, dunes, early morning, May 8, 1999. Photo by Charles Hedgcock.
1174
Lower dunes and steep dune slopes; often
beneath shrubs such as Ephedra. Felger 98-
64 & Eva
Dunes and oo flats of the Gran Desierto in
northwestern Sonora, and extending into
adjacent southwestern Arizona on the Pinta
Sands in Cabeza Prieta National Wildlife Ref-
uge, and disjunct on the Mohawk Dun
and similar habitats in southeastern ane
nia and Baja California. First confirmed for
sea in 1993 sce 2000; Felger et al.
| Heritage Sta-
tus Rank of G2 “imperiled globally).
Cryptantha maritima (Greene) Greene [in-
cludes var. maritima and var. pilosa |.M.
Johnst.].White-haired cryptantha. Cool-sea-
son annuals; corollas white. Widely scattered
on sand flats. Felger 98-99 & Evans
Cryptantha micrantha (Torr) |.M.Johnst.subsp.
micrantha. Dwarf cryptantha. Cool-season
annuals; flowers minute, white. Widespread
on dunes at all elevations. Fe/ger 96-164 &
2!
93). This species ha
—
Turner.
Heliotropium convolvulaceum (Nutt.) A. Gray
var.californicum (Greene) |.M.Johnst.Morn-
ing-glory heliotrope. Non-seasonal annuals
mostly germinating and growing during
warm weather with sufficient soil moisture;
flowers white and showy. Seasonally scat-
tered on dunes. Van ae ies 92-644,
Heliotropium curassavicum L_ [The plants are
var.oculatum (A. Heller) |.M.Johnst.ex Tidestr.,
t the variety is probably not worthy of
recognition, see Felger 2000]. Alkali helio-
trope; hierba del sapo.Semi-succulent peren-
nial herbs or facultative annuals; herbage
frost-sensitive; corollas white, the center yel-
low and turning purplish with age. Wide-
spread across the playa, sometimes locally
abundant. Felger 96-152 & Turner; Reina-G. 96-
211 & Van Devender, Turner 94-1.
Pectocarya heterocarpa {I.M. Johnst.) |.M.
Johnst. Mixednut comb-bur. Cool-season
Hao ei white with a yellow center.
ad and common, dunes and sand
i. Felger 98-65 & Evans
dee an anulie (Munz & ILM. Johnst.)
. Johnst. Broadwing comb-bur.
alae annuals; flowers white. Dunes
. sand flats. Felger 98-90A & Evans
BRIT.ORG/SIDA 20(3
“a plicata (Torr) A.T. Richardson. Fanleaf
inklemat.Perennial herbs from deep, thick
i black roots; flowering spring and sum-
mer; corollas lavender. Mostly on moving
dunes, occasional on low sand hills on sand
flats. Felger 97-70 et al.; Felger 96-136 & Turner,
Reichenbacher 599.
age ete (CRUCIFERAE) -
ard Family
“Brassica tournefortii Gouan. Sahara mustard,
wild turnip; mostaza. Cool-season annuals;
plants highly variable in size; flowers pale
yellow and apy ly selfing (Felger 2000).
Widespread and especially abundant on
dunes during years of high winter-spring
rains; sand flats and dunes including high-
est dunes. Larger plants with basal rosette
leaves each to 72 cm ene; Nios leaves
spreading close to ground
ing other spring aanualeahe leaves are bit-
ter-tasting except those of very small, young
lants. Felger 98-2 et al.; Turner 95-10.
Caulanthus lasiophyllus (Hook. & Arn.) Payson.
-ool-season annuals; flowers minute, white
to pinkish. Mostly in swales and on north-
facing dune slopes, locally common; also
sand flats in washes and swales, mostly at
the base of shrubs. Felger 98-67 & Evans
Dimorphocarpa pinnatifida Rollins [Dithyrea
wislizenii Engelm., in
spectaclepod. Fig. 8. Cool-season annuals,
sometimes more than 1 m across; flowers
white and fragrant. Dunes, especially the
higher dune crests. Endemic to dunes and
sand flats in Yuma County and the Gran
Desierto in northwestern Sonora. Felger 98-
6 et al.; Salywon et al. 985; Turner 95-7. San
Cristobal Valley dunes, Anderson 98-37 (ASU).
Dithyrea californica Harv. Spectaclepod. Cool-
season annuals; flowers cream-colored,
highly fragrant. Widespread across the
dunes, especially lower dunes and slopes.
Felger 98-5 et al.; Salywon et al. 984 (ASU).
Lepidium lasiocarpum Nutt. ex Torr. & A. Gray.
nd peppergrass. Cool-season annuals;
herbage peppery-tasting; flowers minute,
white. Widespread, mostly on lower dunes
and swales, and expected on sand flats.
Felger 98-3 et al.
FELGER ET AL., FLORA ARIZONA 1175
Fic. 8. Dimorphotheca pinnatifida, dunes, April 1998. Photo by Charles Hedgcock.
1176 BRIT.ORG/SIDA 20(3)
Physaria tenella (A. Nels.) O'Kane & Al-Shehbaz Large club cholla; flowers yellow, summer.
[Lesquerella tenella A. Nels.]. Desert One colony, mostly senescent, on low sand
bladderpod. Cool-season annuals; flowers hummock on sand flat near southeast mar-
bright yellow. Sand flats, often growing gin of dune field. Felger & Turner, 25 Sep 1996,
oes bursage sills spp.). Felger 98 observation
7 et al; Felger 98-98 & Eva
Scale longirostris if Watson) Rydb. CARYOPHYLLACEAE - Pink Family
Longbeak twistflower.Cool-season annuals; Achyronychia cooperi Jorr.& A.Gray.Sandmat,
herbage often glaucous and sometimes frostmat. Cool-season annuals; flowers
semisucculent; flowers minute, pale yellow papery white like nue uu ire Dunes, aus
ish-white. Widely scattered across dunes, cially lower d |
especially swales and north-facing slopes, and one. ats, Felger 98- 68 & Evans; a lywon
and less common on higher dunes; also etal. 991 (ASU),
sand di especially a the base of shrubs.
Felger Iger 98-7 etal; Salywon CONVOLVULACEAE — Morning-Glory Family
h. Desert BOGUS GIN:
etal. 993 (ASU), Cc Kallaéa:|
CACTACEAE - Cactus Family ee : ;
parasitic on Boerhavia 0 Seo aia
Carnegiea gigantea (Engelm.) Britton & Rose. dunes and sand flats. Felger 97-59 et al.
aguaro.Columnar cactus; one individual ca.
6 m tall with two large branches. Early sum EPHEDRACEAE - Ephedra Family
mer. Sand flat on east side of dun
Cylindropuntia echinocarpa (Engelm. & J.M.
Bigelow) F.M. Kn uth [Opuntia echinocarpa
Engelm.& J.M. Bigelow; O. wigginsii L.D. Ben-
sonl. sens Small chollas to 1+ m tall;
April; flowers silvery white with green fila
ments. Mostly lower dunes and swales, es
pecially with harder soils, and sand flats,
widely scattered, mostly in small colonies.
Felger & Turner, 24 Sep 1996, observation.
hi polycephalus Engelm.&J.M.Big- EUPHORBIACEAE - Spurge Family
elow var. polycephalus. Many-headed bar
rel cactus. Multiple-headed, mound-forming
barrel cactus; July; flowers yellow. One ma-
ture plant (14 living stems) on creosotebush
flat near west side of dunes. Turner, 27 Jul
1995, photo.
Ferocactus wislizeni (Engelm.) Britton & Rose.
Barrel cactus; biznaga. Unbranched barrel
cactus to ca.1.5 m tall; flowers orange to red-
orange, at least August and September; fruits
ae ts fleshy, pe in late summer a
Ephedra trifurca Torr. ex S. Watson. Longleaf
jointfir, Mormon tea; canutillo, tepopote. Ro
bust, spreading shrubs to ca. 1.5 m tall, the
trunks and major limbs thick and heavy.
Twigs with 3 elongated scale leaves at each
node. Dunes and sand flats. Pollen cones
mature in March; seeds later in spring. Felger
98-52 & Evans; Reina-G, 96-208 & Van Devender,
Salywon et al. 986 (ASU).
Ditaxis serrata (Torr) A. Heller var. serrata
Argythamnia serrata (Torr.) Mull. Arg.]. Non-
seasonal annuals; beans a lene
ite s.Felger
Pines
Pa
green and
97-51 et al., 98-216; sete 96-135 & Turner.
Euphorbia abramsiana L.C. Wheeler
[(Chamaesyce abramsiana (L.C. Wheeler)
Koutnik]. Golondrina. Hot-weather annuals;
petal-like appendages absent or minute.
Sand flats and dunes. Felger et al. 97-57, 97-
78 et al.; Felger 96-128 & Turner
Euphorbia polycarpa Benth. [Chamaesyce
polycarpa (Benth.) Millsp.; £. polycarpa var.
hirtella Boiss.]. Desert spurge; golondrina.
Non-seasonal annuals, but most common
during spring; petal-like appendages white,
sometimes reduced or absent. Sand flats and
early | widely scattered on
sand flats oP west side of dunes, and
rare on dunes. ey 97-52 et al.; Felger 96-
165 & Turner, 96-1
itil kunzei ee Pinkava [Opuntia kunzei
ose. O, stanleyi Engelm. ex B.D. Jackson var.
hn (Rose) L.D.Benson]. Desert club cholla.
FELGER ET AL., FLORA
ARIZONA 1177
=
dunes. Felger 97-53 et al., 97-69; Felger 96-130
& Turner, 96-144.
FABACEAE (LEGUMINOSAE) - Legume Family
Astragalus insularis Kellogg var. harwoodii
Munz & McBurney ex Munz. Sand locoweed.
Cool-season annuals; flowers rose-lavender
and white. Dunes, especially lower dunes, and
sand flats. Felger 98- 70 & Evans; Turner 95-24.
Astragalus nuttallianus DC. var. imperfectus
(Rydb.) Barneby. arene milkvetch.
Cool-season annuals; flowers blue and white.
Widely scattered; dunes, mostly swales and
wer dunes, and a flats. Felger 98-74 &
Evans; Felger 98-1
Dalea mollis Benth. ci dalea. Non-seasonal
annuals. Dunes, especially lower dunes and
dune slopes, and sand flats. Fe/ger 98-80 &
Evans; Reichenbacher 437; Turner 95-27.
lauca (Ortega) Eifert
ee
=]
flowering non-seasonally with warm
weather and sufficient soil moisture; flow-
ers yellow.Common across the playa. Felger
98-50 & Evans; Reina-G. 96-212 & Van
Devender, Turner 95-23.
Lupinus arizonicus (S. Watson) S. Watson [L.
arizonicus subsp. sonorensis J.A. Christian &
D.B. Dunn] Arizona lupine; /upino. Cool-sea-
son annuals, flowers lavender-pink. Wide-
spread and common, lower to higher dunes,
slopes, and dune swales, and sand flats.
Plants often ravaged by caterpillars of the
painted lady butterfly (Vanessa cardui) and
the white-lined sphinx moth (Hyles lineata;
Fig. 10). Felger 98-79 & ae Salywon et al.
993 (ASU); Turner 95-3
Marina parryi (Torr.& A Gtayy ex A.Gray) Barneby
[Dalea parryi Torr. & A. Gray ex A. Gray]. Non-
seasonal annuals (sometimes short-lived
herbaceous perennials elsewhere); flowers
dark blue. Dunes and expected on sand flats.
Reichenbacher, 7 Mar 1980, observation.
| tesota A | \:palo fiero. Tree
—
ca.6+ m tall;late April and May; flowers lav-
ender-pink. Washes in sand flats near the
northeast and southeast parts of dune field.
Felger & Turner, 25 Sep 1996, observation.
ene florida (Benth. ex A. Gray) S. Wat-
rcidium floridum Benth. ex A. Gray].
ae os verde; palo verde. Trees, ca. 5+ m
tall; ea flowers bright yellow. Large wash
in sand flats near north end of dune field.
ae s.n., 12 Nov 2001
Parkinsonia microphylla Torr. [Cercidium
microphyllum (Torr.) Rose & ILM. Johnst.].
Littleleaf palo verde; palo verde. Small trees;
April and May; flowers pale yellow with a
white banner. Scattered along washes in
sand flats ca. 1-2 km from northeast and
southwest margins of the dune field. Felger
& Turner, 24 Sep 1996, observat
Prosopis glandulosa Torr. var. ae (L.D.
Benson) M.C. Johnst. Western honey-mes-
quite; mezquite. Large shrubs to small trees;
flowering April and often sporadically
pl 7 f } Ps per 5 Pees : +}
if
a
surrounding desert, and rarely on dunes in-
avai dune swales ang crests, Mesquites
ith Pveliting
VVILCET ET. VOTCHCri
Wooton; most leaves have one pair of pin-
nae, as in Pglandulosa, although occasional
leaves have Ewe “Palle as in P VEMETa, and
+b f ba {
tne VIET
as inP veldiine (see Felger 2000), ee 96.
165 & Turner; Reina-G. 96-207 & Van Devender.
Psorothamnus emoryi (A. Gray) Rydb. var.
emoryi [Dalea emoryi A. Gray]. Emory
indigobush. Shrubs to ca.1 m tall; flowering
after rains, spring and summer; flowers
purple and white. Very common on dunes
at all elevations and to margins of sand flats.
Felger 96-141 & Turner, 96-163; Felger 97-65 et
al.; Reina-G. 96-209 & Van Devender.
FOUQUIERIACEAE - Ocotillo Family
Fouquieria splendens Engelm. subsp.
splendens. Ocotillo. Shrubs; spring; flowers
red-orange.Common on sand flats at north-
east margin of dune field. Felger & Turner, 24
Sep 1996, observation.
GERANIACEAE - Geranium Family
Erodium texanum A. Gray. False filaree, desert
stork-bill. Cool-season annuals; flowers pink-
purple. Sand flats. Felger 98-18 et al.
1178
BRIT.ORG/SIDA 20(3)
Fic. 9. Hoffmannseggia glauca, on the playa, June 1994. Photo by Charles Hedgcock.
HYDROPHYLLACEAE - Waterleaf Family
Nama hispidum A. Gray. Bristly nama. Fig. 11.
Cool-season annuals; flowers lavender.
Dunes and swales, playa margins, and sand
flats. Felger 98-71 & Evans; Turner 95-16
Phacelia ambigua ME. Jones [P crenulata Torr.
ex S.Watson; P crenulata var. ambigua (M.E.
Jones) J.F. Macbr.]. Desert heliotrope. Cool-
season annuals; flowers lavender-
Sand flats. Felger 98-89 & Evans; Turner 95-9.
KRAMERIACEAE - Ratany Family
Krameria erecta Willd. ex Schult. [K. parvifolia
Benth.]. Range ratany. Dwarf shrubs; flower-
ing at least spring and late summer; flowers
bright magenta-purple. lon sand
flats. Felger 96-148 & Turner, 96-172.
Krameria grayi Rose & Painter. White ratany;
cosahui. Shrubs; flowering various seasons;
flowers magenta-purple. Sand flats; appar
ently rare or at least uncommon in the
\ALA
Mohawk Dune region see Mos common in
nearby areas. Felger 96-147 & Turne
LILIACEAE (sensu lato) - Lily Family
Hesperocallis undulata A. Gray. Ajo lily, desert
lily; ajo silvestre. Fig. 12. Perennials from a
large, single bulb; flowering stalks some-
times reaching 1.5 m in height; February-
when fully open, opening late afternoon,
closing with daytime heat of the next morn-
ing. Widespread, dunes and sand flats; best
developed on dunes including highest
dunes. Felger 98-72 & Evans.
LOASACEAE - Stick-Leaf Family
Mentzelia albicaulis Douglas ex Hook. White-
stem blazingstar.Cool-season annuals; flow-
ers small with uniformly bright-yellow pet-
als. Dunes and especially in swales. Felger 98-
82 & Evans
FELGER ET AL., FLORA ARIZONA 1179
Fic. 10. Hyles lineata eating Lupinus arizonicus, April 1998. Photo by Charles Hedgcock.
1180
ek.
a4 ty ay
Fic. 11 Al e¢ A fel. Cok t } 41 O.
BRIT.ORG/SIDA 20(3
Mentzelia multiflora (Nutt.) A.Gray. Blazingstar.
Robust, cool-season annuals; flowers mod
erately large, yellow. Dunes and sometimes
on sand flats. Salywon & Shohet 558 (ASU):
Turner 92-2, 95-5,
MALVACEAE © Mallow Family
Malvella sagittifolia (A. Gray) Fryxell [Sida lepidota
A.Gray var. sagittifolia A.Gray].Narrowleaf mal
low.Perennial herbs from deeply seated roots,
spreading by rhizomes; growing and flower
ing with sufficient soil moisture any time of
year except the coldest months.Open area o
playa. Reina-G. 96-210 & Van Devender, Salywon
et al, 999 (ASU): Turner 95-18.
| April 1998. Photo by Charles Hedgcock.
Sphaeralcea ambigua A. Gray subsp.ambigua
Desert globemallow. Perennial subshrubs;
flowering various seas pecially spring;
flowers
orange. ci abundant at
playa margins, and occasional on sand flats
at northeast side of dunes. Fe/ger 98-21 etal.;
Turner s.n, 19 Apr 1998.
Sphaeralcea coulteri (S. Watson) A. Gray var.
coulteri. Annual globemallow; mal de ojo.
Non-seasonal annuals, generally growing
with winte! atl rains, non variable in
size, several cm t
ae
5m tall;
15 vere nie
yes and ond flats. Felger 98-75 & Evans;
Turner 95-17.
¢ TT | | |
Navy ADUNGANT, LO
FELGER ET AL., FLORA ARIZONA 1181
11998 +h | | 7 | 1 | th
’
Fic. 12. Hesperocallis undulata, flowers ca.6—7 cm long, Apri
sand. Photo by Charles Hedgcock.
1182
MARTYNIACEAE - Devil's Claw Family
Proboscidea altheaefolia (Benth.) Decne.
evil's claw; gato, una de gato, torito. Fig. 13.
Perennial herbs from a single tuberous root,
growing and flowering during warmer
months; flowers showy, bright yellow with
white and orange markings. Widely scat-
tered; dunes, sand flats, and playa. Felger 97-
63 et al.; Felger 96-146 & Turner.
MOLLUGINACEAE - Carpetweed Family
*Mollugo cerviana (L.) Sér. Thread-stem car-
petweed; Mela Seco Small, hot-
weathe green and
whitish, Seasonally common on dunes.
Felger 97-62 et al. 97-81.
NYCTAGINACEAE - Four-O'clock Family
Abronia villosa S. Watson var. villosa. Sand ver-
ena.Cool-season annuals; flowers lavender-
pink and fragrant. Widespread and often
seasonally abundant on dunes and sand
hummocks across the Mohawk Valley, often
producing spectacular floral displays in
spring. Salywon et al. 996 (ASU); Turner 95-15.
trailing
Allionia incarnata L. Ti Trailing windmill
four o'clock. Non-seasonal annuals, mostly
growing during warmer months, with trail-
i | | L 1* J ‘ |
violet-rose or magenta.Dunes and sand flats.
neiger 97- ae et a nee 96- 132 & Turner.
vari (M.E. Jones)
Kearney & Peebles [B. intermedia M.E. Jones;
B. triqueta S.Watson]. Spiderling. Hot-weather
annuals;flowers white to pale pink. Sand flats.
Felger 97-75 etal
Boerhavia spicata Choisy [B. spicata var.palmeri
S.Watson:B oe (Hook. ") S. viene Hot-
eather whit LO
pale pink.
Sand flats and eae poe ie uslye com-
The p ants
|; i} | pill (Hy loc hy \
Felger 97- 55 etal. Felger 96. (668 Turner
Boerhavia wrightii A. Gray. Largebract
spiderling. Hot-weather annuals; flowers
pinkish white to pink. Sand flats, often local-
ized in small depressions. Felger 97-76 et al.
ne
ONAGRACEAE - Evening Primrose Family
BRIT.ORG/SIDA 20(3)
Camissonia boothii (Douglas) PH.Raven subsp.
condensata (Munz) P.H. Raven. Woody
bottle-washer.Cool-season annuals; flowers
white. Skeletons may persistent for one year
or more. Mostly in localized colonies on sand
flats and accasionally at the playa edge.
Felger 98-92 & Evans; F 96-157 & Turner.
Camissonia californica (Torr. & A. Gray) PH
Raven. Cool-season annuals; flowers yellow,
changing to orange and drying pink. Local-
ized indiin pal =| cal qd
flats, ao ly along small
87 & Eva
Camissonia iamsenenuides (A. Gray) P.H.
Rav le evening-primrose. Cool-
season hae oe pink. Sand flats, gen-
erally along small washes or in shallow de-
pressions. This is ANE rmlesy flowered
Desert: the
Felger 98-
VaSIIes,.
=e} t
floral siraetute and modifications are char-
acteristic of self-pollinated flowers. Felger 98-
100 & Evans
Camissonia claviformis (Torr, & Frém.) PH.
Raven subsp. peeblesii (Munz) PH. Raven.
Cool-season annuals. Corollas white to pink
with maroon-brown inside the tube. Sand
flats, lower dunes, dune swales and slopes.
Felger 98-14 et al; Felger 98-94 & Evans; Turner
95-31.
Camissonia claviformis subsp. yumae (PH.
Raven) PH. Raven. Cool-season annuals. Co-
rollas yellow with maroon-brown inside the
tube. Sand flats near the dune field. Felger
98-94 & Evans. Although two subspecies oc-
cur in the area, they apparently do not oc-
cur intermixed and plants with intermedi-
ate features have not been found.
Oenothera deltoides Torr. & Frém. subsp.
deltoides. Dune evening-primrose, white
evening-primrose, devils’ lantern, Cool-sea-
son annuals; flowers large and white. The dry
skeletons persist for one or more years. Sand
flats and dunes. Often common and showy.
Turner 95-8.
Oenothera primiveris A. Gray subsp.
primiveris. Yellow desert evening-primrose.
Cool-season annuals; flowers large and yel-
low. The dry skeletons may persist for sev-
eral years. Mostly lower dunes, also swales
FELGER ET AL., FLORA
ARIZONA 1183
Fic. 13. Proboscidea altheaefolia, dunes, August 1996. Photo by Charles Hedgcock.
and sand flats. Felger 98-84 & Evans; Felger 96-
158 & Turner.
OROBANCHACEAE - Broomrape Family
Orobanche cooperi (A. Gray) A. Heller. Desert
broomrape; flor de tierra. Cool-season annu-
als parasitic on Ambrosia spp.,;stems thick and
succulent; flowers purple and white, the
throat often marked with yellow. piece
the plants may appear perennial because o
their robust habit, they are ee annu-
als (George Yatskievych, pers. comm.). Sand
flats. Goldberg 77-91; Turner 5.n., 2 ee 1998.
PAPAVERACEAE - Poppy Family
Argemone gracilenta Greene. Sonoran
pricklypoppy, fried-egg poppy; cardo. Short-
lived scene to 80+ cm tall, or probably
often facultative annuals in the flora region;
flowering late spring and in nearby areas
sometimes sporadically through the sum-
mer; flowers large, with white petals and nu-
merous yellow stamens. Lower dunes and
dune swales, scattered and seldom com-
mon. Felger 98-77 & Evans.
Eschscholzia glyptosperma Greene. Gold
poppy. Cool-season annuals. Locally com-
mon during years of exceptional winter-
spring rain; flowers yellow-orange. Sand flats
especially near the northern and eastern
part of the dune field. Felger 98-88 & Evans;
saan et al. ‘“ eu) Turner 95
S.Watson. | <n Id-
poppy. Cool-season annuals; flowers yellow-
orange. Dunes and locally on sand flats.
Turner 95-33.
1184
PLANTAGINACEAE - Plantain Family
Plantago ovata Forssk. [P insularis Eastw., not P.
insularis (Grenier & Godron) Nyman, P.
fastigiata E. Morris; P insularis var. fastigiata
(E. Morris) Jeps.J. Woolly plantain, Indian-
wheat; pastora. Winter-spring ephemerals,
highly variable in size depending on soil
moisture; flowers straw-colored.Widespread
across dunes and sand flats, but os
lower dunes and swales, and sometimes
the playa. Felger 98-1 et al; Goldberg 77- a
Salywon et al. 998 (ASU); Turner 95-19.
a,
POACEAE (GRAMINEAE) - Grass Family
Aristida adscensionis L. Sixweeks threeawn;
zacate tres barbas. Non-seasonal annuals.
and flats. Felger 96- veer
ancade californica [hurb. var. mis Cali-
fornia threeawn; tres barbas de california.
Tufted perennials, also flowering in first sea-
son during summer. Widespread, sand flats
and especially on dunes. Felger 98-
10etal.;Felger 96-142 & Turner, 96-167, 96-170.
Bouteloua aristidoides (Kunth) Griseb.
Sixweeks needle grama; aceitilla, navajita.
Summer annuals. Sand flats and dunes;
mmon.felger 97-80 et al.,
97-67; Felger 96-140 & Turner.
Bouteloua barbata Lag. Six-weeks grama;
navajita, zacate liebrero. Summer annuals.
ts and dunes. Felger 97-68 et al.; Felger
37 & Turner, 96-131.
Festuca octoflora Walter [Vulpia octoflora
(Walter) Rydb.; F octoflora var. hirtella Piper].
Sixweeks fescue, eight-flowered fescue.
Cool-season annuals; diminutive grass.Sand
flats, mostly beneath shrubs and in small
depressions at the northeast side of the
dune field. Felger 98-97 & Evc
Muhlenbergia cede (DC.) Trin
Littleseed muhly; liendrilla chica. Non-sea-
sonal annuals. Generally beneath shrubs in
small washes or depressions in sand flats.
Felger 98-18 et al.
Panicum alatum Zuloaga & Morrone var. mi-
nus ara Zuloaga & Morrone. Sum-
mer annuals. Shallow depression in sand
flats at edge of playa. Felger 98-8 et al. This
species is indistinguishable from the com
widespread and
I
—
n
BRIT.ORG/SIDA 20(3)
|
hirticaule pt for
n and widespread P
consistent spikelet characters. _ pres
Sonora and southwestern Arizona Palatum
has a limited and localized distribution, char-
acteristically in playas and similar habitats
with temporary standing water and clayish
soils, while P hirticaule is widespread and
common in many habitats throughout the
region (Felger ae do not seem to
occur togethe
Pleuraphis ae TUE (Hilaria rigida (Thurb.)
cribn.].Big galleta;galleta grande,
toboso. Large, tufted or bushy perennials.
Widespread and abundant, dunes
flats. Felger 96-137 & Turner.
*Schismus arabicus Nees. Arabian schismus.
ool-season annuals. Widespread and sea-
sonally abundant on sand flats and dunes,
and especially common on dune slopes and
wales. The plants at first grow close to the
sour some preventing the growth
of n ae (see Felger 2000).
ae cr
and sand
POLEMONIACEAE - Phlox Family
Eriastrum GITDSUmn (A. Giay) ch Mason. Cool-
Dunes and
sand flats. ae 98-55 & Evans.
Eriastrum eremicum (Jeps.) Hi Mason. Cool-
son Is-fl d lue. Sand flats
tern record for this spe-
cies,and the only plantin i veneer flora
not in the Gran Desierto or near the Pinta
Sands or Yuma Dunes. Burgess 5059; Turner
This is tl Ss outh
95-32.
Linanthus bigelovii (A. Gray) Greene [Gilia
bigelovii A. Gray, 1870; Gilia jonesii A. Gray,
1886; Linanthus bigelovii var. jonesii (A. Gray)
Brand; L. jonesii (A. Gray) Greene]. Cool-sea-
son annuals. Flowers white, nocturnal, open
ng at or near dusk and powerfully sweet-
ao sometime nauseously sweet, and
closing before sunrise. Dunes including
flats. Bowers 2993: Felger 98
54 & Evans; Reichenbacher 273.
Linanthus jonesii is distinguished by glandular-
stipitate een (sometimes quite densely
so) and seeds constricted near the middle
(a sae “pinched off"), whereas L. bigelovii
sensu stricto is distinguished by glabrous
as
wales and sand
FELGER ET AL., FLORA
ARIZONA
1185
calyces and a bladdery, ellipsoid seed. How-
ever, in southwestern Arizona the variation
seems continuous, the characters do not
seem correlated with each other, there ap-
pears to be no discernable correlation with
habitat or geography, and they do not seem
to be distinct taxa
Loeseliastrum schottii (Torr.) Timbrook
[Langloisia schottii (Torr.) Greene]. Winter-
spring annuals. Scattered, mostly in swales
and sand flats, sometimes extending to
higher dunes, seldom .Felger 98-53
& Evans; Galvan 77-83; Turner 95-34.
POLYGONACEAE - Buckwheat Family
Chorizanthe brevicornu’ Torr. subsp.
brevicornu. Brittle spineflower, shore horn
spineflower. Cool-season annuals; flowers
white and minute. Dunes, especially swales,
dune slopes, and lower dunes; also on sand
flats. Felger 98-81 & Evans.
Chorizanthe rigida (Torr.) Torr. & A. Gray. Rigid
spineflower. Cool-season annuals; the dry
skeletons persistent for several years; flow-
ers white, minute and cryptic.Common and
widespread on sand flats. Fe/ger $e et al,;
Felger 98-89 & Evans; Reichenbache
ate trichopes Torr. var. See Little
t. Cool-season annuals, occasionally
SUIV i the summer; flowers yellow. Dunes
and sand flats. Felger 98-57 & Evans; Salywon
et al. 989 (ASU).
Nemacaulis denudata Nutt. var. gracilis
Goodman &| p Benson.Woolly heads.Cool-
season ; Flowers minute, yellow, sur-
rounded by en green to white and pink
woolly bracts. Mostly in dune swales, also
lower to higher dunes. Felger 98-73 & Evans;
Salywon et al. 990 (ASU).
RESEDACEAE - Mignonette Family
Oligomeris linifolia (Vahl) J.-F. Macbr. Desert
cambess, slenderleaf cambess. Non-seasonal
annuals; flowers inconspicuous, white and
green. Playa, sand flats, and lower dunes.
Turner 95-20
SALICACEAE - Willow Family
ws \Wlatcon tAcn framoanti
Cottonwood; alamo. Several saplings along
the southern edge of the playa. Apparently
they grew during the El Nino year of 1992-
1993, and were still living by end of April
1994, having reached 2-2.5 m tall, but per-
ished by the end of 1994. Felger 96-153 &
Turner; Phil Rosen, 24 Apr 1994, photo.
SIMAROUBACEAE - Quassia Family
Castela emoryi (A. Gray) Moran & Felger
[Holacantha emoryi A. Gray]. Crucifixion
thorn;corona de cristo. Shrubs 2-3 m tall; May-
July; flowers cream-yellow to pink; fruits
densely clustered and persistent. Scattered
on sand flats,and also lowermost dunes sur-
rounding the playa. Felger 96-162 & Turner,
Reina-G. 96-216 & Van Devender.
SOLANACEAE - Nightshade Family
Datura discolor Bernh. Poisonous nightshade,
desert thorn-apple; toloache. Non-seasonal
annuals, the plants frost-sensitive; flowers 8-
cm long, corollas white with a purple
flush in the throat, nocturnal and fragrant.
Sand flats and probably on dunes. This is the
largest-flowered plant in the ae Felger
& Turner, 25 Sep 1996, observa
Lycium parishii A. Gray var. ser en wolf-
berry; salicieso. Shrubs; reproductive at vari-
ous seasons, especially late winter and early
near north end of dunes. Felger 98-20 et al.
Physalis lobata Torr. [Quincula lobata (Torr.)
Raf.]. Annual or perennial herbs with a deep
tap root; flowers lavender, after sufficient
ns. Locally at the playa
margins. Felger 98-51 & Evans; Turner 95-29.
rains at various seaso
TAMARICACEAE - Tamarisk Family
“Tamarix ramosissima Ledeb. [7. chinensis of
American authors, not Lour.?]. Saltce
tamarisk; salado, pino salado. One 6. at
the southwest edge of the playa, first seen
May 1995, and by June 1996 it had perished
from drought. Felger & Turner, 25 Sep 1996,
observation.
VISCACEAE - Mistletoe Family
Phoradendron californicum Nutt. Desert
1186
mistletoe; toji. Epiphytic perennial parasites;
flowers small, green or yellow, at various sea
BRIT.ORG/SIDA 20(3)
flats and dunes. Felger 97-58 et al.; Felger 96
129 & Turner, 96-133, 96-160
Larrea divaricata Cav.subsp. tridentata (Sesse
& Moc. ex DC.) Felger & C.H. Lowe [L.
tridentata (Sessé & Moc. ex DC.) Cov.].
Creosotebush; hediondilla, sane
Long-lived shrubs ca. 1-2 m tall w
hard wood; flowers eee various seasons,
Dominant shrub or “canopy” across most of
the sand flats and valley floor aoa
the dunes, and extending onto lower dunes
and dune swales.
Sons, venue sal and oe ey
ragranta attrac
insects; nee in same season. On widely
scattered mesquite trees on sand flats. Fe/ger
& Turner, 24 Sep 1996, observation.
ZYGOPHYLLACEAE - Caltrop Family
Kallstroemia californica (S. Watson) Vail. Cali-
fornia eae mal de ojo. Hot-weather an-
nuals; HOVVETS yellc VW LO Ye llov Orange. Sand
ACKNOWLEDGMENTS
For financial support we thank the Wallace Research Foundation, and the 56"
Range Management Office of Luke Air Force Base for funding this work under
Contract FO260497MS130 and under the Department of Defense Legacy Re-
source Management Program (Project 95-1009). We thank Bill Broyles, Luke
Evans, Julia Fonseca, Scotty Johnson, Cecil Schwalbe, Gerald Turner, Louise
Turner, and Peter Warren for generous assistance and Charles Hedgcock
(wwweharleshedgcock.com) and Thomas R. Jones for supplying photos. James
Henrickson, Wendy Hodgson, and John Anderson provided significant reviews.
Phil Jenkins and Michael Chamberland of the University of Arizona herbarium
greatly facilitated our work, and Andrew Salywon and Tom Van Devender
shared significant information.
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1188 BRIT.ORG/SIDA 20(3)
BOOK NOTICES
D.M,J.S. Bowman. 2000. Australian Rainforests: Islands of Green in a Land of Fire.
(ISBN 0-521-46568-0, hbk.). Cambridge University Press, The Edinburgh
Bldg., Cambridge CB2 2RU, U.K. (Orders: 40 West 20" St., New York, NY
10011-4211, U.S.A. wwwcup.org, www.cup.cam.ac.uk). $85.00, 345 pp., b/
w photos, line drawings, graphs, 6" x 9"
Contents: 1) Introduction; 2) What is Australian rainforest?, 3) The sclerophyll problem, 4) The
edaphic theory I. The control of rainforest by soil phosphorus; 5) The edaphic theory IL Soil types,
drainage, and fertility; 6) The climate theory I. Water stress; 7) The climate theory II. Light and tem-
perature; 8) The fire theory L. Field evidence; 9) The fire theory IL. Fire, nutrient cycling, and topogra-
phy; 10) The fire theory UL Fire frequency, succession, and ecological oe 11) The fire theory IV.
Aboriginal landscape burning; 12) The fire ny V. Aridity andth tion of flammable forests;
13) The fire aneOry VI. Fire management and st conservation; 14) Summary; followed by Rel-
erences and Index.
Ray L. Spectr and AuIson Sprcut. 2002. Australian Plant Communities: Dynam-
ics of Structure, Growth and Biodiversity. (ISBN 0-19-5516054-0, pbk.). Oxford
University Press, Department REF 198 Madison Avenue, New York, NY 10016,
U.S.A. (Orders: 2001 Evans Road, Cary, NC 27513, U.S.A. 800-451-7556, fax
919-677-1303). $40.00, 492 pp., b/w photos, line drawings, graphs, 6" x 9".
Contents: Part I Australian plant communities: Description: 1) Energy/biomass eae es mee
ics; 3) Structural cl: SICaliOn, 4) . loristic classification; 5) Australian plan
the Late Cretaceous; 6) luring the Cainozoic; 7) Australian oe com-
munities today; 8) Aboriginal ees Part 2 Australian a communities: Community physiol-
ogy; 9) Energetics; 10) Temperature; 11) Evaporative aerodynamics, 12) Available water, 13) Eco-
physiological leaf attributes; 14) Waterlogging; 15) Nutrient deficiencies; 16) Nutrient toxicities; 17)
Biodiversity and energetics, 18) Monitoring; 19) Scientific management, References; Systematic In-
dex; and General Index.
The authors believe this book will be an invaluable reference lor pisces environmental
scientists, planners, and managers. It will also be an excellent text for undergraduate and graduate
course in environmental science, resource Management, ae environmental planning and impact
assessment.
SIDA 20(3): 1188. 2003
A FLORISTIC SURVEY OF BIG BRANCH MARSH NATIONAL
WILDLIFE REFUGE, ST. TAMMANY PARISH, LOUISIANA
David J.Rosen Stanley D. Jones
U.S. Fish and Wildlife Service Herbarium
— a ee cal Services Field Office Botanical pee Center
9 Fl Camino Real, Suite 211 PO. Box 67
ae 7. 77058-3051, U.S.A. Bryan, TX 77805- a > U.S.A.
Virginia E. Rettig
U.S. Fish and Wildlife Service
Cat Island National Wildlife Refuge
St. Francisville LA 70775, U.S.A.
ABSTRACT
Big pes Mash a Wildlife See isa U.S. Department of me Interior Fish and Wildlife
Ser 370 dha (4 500 acres). A vey W as conducted
to provide ee staff ith a ede of v oe r plant species, along with species abundance and
habitat preference when one existed. Collecting trips made to the refuge from August 1999 through
April 2002 yielded 563 infraspecific taxa of vascular plants Rees 117 families and 309 gen-
era. The four largest families are Poaceae, Cyperaceae, Asteraceae, an e. Descriptions of sa-
lient plant communities are provided.
RESUMEN
El Refugio Nacional de Vida Salvaje de Big Branch Marsh es un complejo del Servicio del Departamento
de Pesca Interior y' becat aaa a os Estados Unidos que tiene aproximadamente 5,870 ha (14,500
acres). Se realizo\ t ee ara — al laa del Relu ao n listado de las especies
de plantas vasculares, junt b habitat cuando existia. Los
vale de recoleccion al refugio de agosto de 1999 a aul is 2002 cae 563 taxa infraespecificos de
plantas vasculares all7 eee y had ePnetOs: Las cuatro uamunaee mas pcs son
Poace e, Cyper aceae, Asteraceae y Fabac I vegetales
mas cen
INTRODUCTION
The National Wildlife Refuge System is administered by the U.S. Department
of the Interior, Fish and Wildlife Service. The mission of the Refuge System is
toadminister a national network of lands and waters for the conservation, man-
agement, and where appropriate, restoration of the fish, wildlife, and plant re-
sources and their habitats within the United States for the benefit of present
and future generations of Americans. Big Branch Marsh National Wildlife Ref-
uge (BBMNWR), which was established in 1994, is one of over 530 refuges
throughout the country and one of 22 refuges in Louisiana. At the onset of this
survey, BBMNWR was the last large undeveloped area on the rapidly develop-
SIDA 20(3): 1189-1216. 2003
1190 BRIT.ORG/SIDA 20(3)
—_
| Lake Pontchartrain. The management objectives of BBMNWR
with Big
ing north shore o
are to provide (1) habitat for a natural diversity of wildlife associatec
Branch Marsh, (2) wintering habitat for migratory waterfowl, (3) nesting habi-
tat for wood ducks, (4) habitat for non-game migratory birds, and (5) opportu-
nities for public outdoor recreation.
The forested areas of BBMNWR are primarily managed for the federally
listed endangered red-cockaded woodpecker (Picoides borealis). About 12 clus-
ters of these birds are currently active on the refuge. Red-cockaded wood peck-
ers build their nests in live pine trees and require an open understory necessi-
tating prescribed burns by refuge personnel every 3—4 years. Refuge marshes
are also burned every 2—3 years to improve habitat for wading birds and mi-
gratory waterfowl. The understanding and management of any animal com-
munity demands (1) defining the plant communities in which they live, 2)
defining the surrounding plant communities in which they move in and out of,
and (3) identifying the plant species utilized for food, cover, and nesting. As is
the case with most National Wildlife Refuges, no detailed floristic study had
been conducted on BBMNWR. Therelore, a floristic survey was conducted to
provide refuge staff with a checklist of vascular plant species and information
fou
on species abundance and habitat preference.
Study Area
BBMNWR is located between 30° 15' 35" and 30° 19'54"N latitude and 89° 54' 37"
and 90° 00'53"W longitude in St. Tammany Parish, Louisiana (Fig. 1). The ref-
uge lies within the Coastal Plain Province of the Atlantic Plain Division
(Fenneman 1928). The refuge headquarters and visitor center are located W of
Hwy. +34 at about 30° 19’ 17"N, 89° 560’ 14"W in the town of Lacombe. Several
minor tributaries border and traverse the refuge holdings. The refuge is bounded
to the W by Cane Bayou, bisected by Bayou Lacombe and its tributaries, and
traversed by Bayou Liberty, Bayou Bonfouca, Cypress Bayou, Bayou due Chien,
their tributaries, and other small unnamed tributaries. The most notable ref-
uge boundary is Lake Pontchartrain, a shallow, flat-bottomed fresh to brackish
water body with an average depth of approximately 3-m (Steinmayer 1939).
The refuge lies entirely within its floodplain between the 0 and 5 foot contour
lines. Lake salinity normally ranges from 1 to 10 °/oo. In recent drought years,
however, salinity has reached 20 °/0o (pers. obs.). The climate in St. Tammany
Parish is temperate with an average winter temperature of 12°C, and a daily
minimum average of 5°C. Average summer temperature is 27°C, and the aver-
age daily high is 33°C (Trahan et al. 1990). Total annual precipitation is 155 cm,
average relative humidity during the day is 60% and is 90% at dawn (Trahan et
al. 1990).
The approximately 5870 ha (14,500 acre) refuge is described by refuge staff
as predominately a mixture of intermediate and fresh marsh, pine flatwoods,
WILDLIFE REFUGE
ROSEN ET AL
BIG BRANCH MARSH
NATIONAL WILDLIFE REFUGE
PONTCHARTRAIN iS
P
f “
LAKE (40)
_ 3
J
ff PONTCHARTRAIN
/
ONAL ~~ —e | |
VICINITY MAP _ | — —— /
Fic. 1. Map showing i i ig I ing y
pine islands, pine ridges, and hardwood hammocks. The forests in the area were
almost completely clear cut early in the 20 century (Ellis 1981). Most of the
evergreen forests in the region were dominated by longleaf pine (Pinus palustris
Mill.) prior to modern silviculture operations and fire suppression. Currently,
either slash pine (P. elliottii Engelm. var. elliottii), loblolly pine (P. taeda L.), or
longleaf pine are dominant or co-dominant in refuge evergreen forests.
The most notable physiognomic feature of the refuge is a distinct line of
demarcation where marsh meets pine [latwoods. The prairie terrace is com-
prised of consolidated sediments formed during the Pleistocene resulting from
rising and falling sea levels and changes in the course of the Mississippi River
(Fisk & McFarlan 1955; Saucier 1963). The forested areas of the refuge overlay
the terrace. The entire refuge area is subsiding, but the effect is most noticeable
in the existing marshes. In some locations, the marsh-forest edge is lined with
dead stands of pine trees that have been inundated with brackish lake water.
METHODS
Topographical maps and color infrared aerial photos were examined and inter-
preted to locate different habitats and plan fieldwork. Collecting trips were made
to the refuge from August 1999 through April 2002. The objectives of each trip
1192 BRIT.ORG/SIDA 20(3)
were the collection of voucher specimens, careful notation of apparent habitat
preference, and estimates of abundance of each species. Voucher specimens are
housed at the Tulane University Herbarium (NO). Duplicate specimens, when
available, were provided to the staff of BBMNWR. Identifications were made
using regional manuals including Allen (1992), Correll and Johnston (1970),
Cronquist (1980), Gandhi and Thomas (1989), Godfrey (1988), Godfrey and
Wooten (1979, 1981), Gould (1975), Isely (1990), and Radford et al. (1968).
RESULTS
Floristics
Collecting trips yielded 563 infraspecific taxa of vascular plants representing
117 families and 309 genera (Table 1). The four families containing the most in-
fraspecific taxa (in parenthesis) are Poaceae (94 spp.), Cyperaceae (64 spp.),
Asteraceae (61 spp.), and Fabaceae (25 spp.). One species, Imperata brasiliensis
(L.) Beauv, is listed by the U.S. Department of Agriculture as a noxious weed.
Non native species accounted for 12.8 % of the total species. No Federally-listed
threatened or endangered plant species were found.
Plant Communities
The distribution and composition of plant communities on the refuge appear
most influenced by the salinity and tidal activity of Lake Pontchartrain and
larger tributaries. Other major influences are topography, geology, soil type,
weather, historic and ongoing disturbances, and management regimes. Plant
communities were described to indicate diversity and uniqueness and to aid in
future management practices. The following plant communities were defined
by comparing field observations of dominant woody and herbaceous species
and hydrological regimes to The Nature Conservancy's International Classifi-
cation of Ecological Communities: Terrestrial Vegetation of the Southeastern
United States ICEC) (Weakly et al. 1998). Plant community names of the ICEC
hierarchy are used. Following the ICEC alliance plant community name, the
name used by refuge staff is listed in parenthesis. Communities at the alliance
and association levels are reported. These are the two lowest communities in
the hierarchy of classification listed in the ICEC and are based on floristics.
Associations within alliances are discussed under the alliances. Conservation
ranks of G3 or lower are reported (Gl-critically imperiled globally; G2-imper-
iled globally; G3-rare or uncommon globally). A myriad of disturbed sites such
as borrow pits, ditches, roads, spoil areas, logging ruts, and mowed rights-of-
way may occur in each alliance or association, typically supporting a variety
of ruderal, wetland, and aquatic species.
Forest Alliances
Fagus grandifolia-Magnolia grandiflora Forest Alliance (Beech-Magnolia Woods).
A single representative association of this alliance was found on BBMNWR,
Tasle 1.Statistical taxonomic summary of the vascular flora of Big Branch Marsh NWR.
Infraspecific Taxa
Class Families Genera Native Introduced Total
Lycopodiopsida 1 ] ] 1
Isoétopsida 1 ] 1 = 1
Polypodiopsida 9 9 8 2 10
Pinopsida 2 2 6 - 6
Liliopsida 24 85 194 21 215
Magnoliopsida 80 211 281 49 330
Totals 117 309 49] 72 563
the Fagus grandifolia Ehrh. (American beech)-Magnolia grandiflora L. (south-
ern magnolia)-Quercus nigra L. (water oak)-Pinus glabra Walt. (spruce pine)/
Viburnum dentatum L. (southern arrow-wood) Association. This association
occurs on uplands fringed by the marshes adjacent to Cane Bayou. Common
trees in the subcanopy include Carpinus caroliniana Walt. (coastal ironwood),
Carya glabra (Mill.) Sweet. (pignut hickory), and Quercus falcata Michx. var.
falcata (southern red oak). Common shrubs are Aesculus pavia L. (red buck-
eye), Callicarpa americana L.(beauty berry), Erythrina herbacea L.(coralbean),
Halesia diptera Ellis. (two-wing silverbell), Symplocos tinctoria (L.) LHer.
(sweetleaf), and Vaccinium arboreum Marsh. (farkleberry). Southern arrow-
wood is apparently absent in this association at BBMNWR. Common herba-
ceous species include Chasmanthium laxum (L.) Yates var. laxum (slender
spikegrass), Carex spp. (caric-sedges), and Panicum spp. (panic-grasses). This
association is easily viewed at the Cane Bayou access area.
oo
Nyssa aquatica—(Taxodium distichum) Semipermanently Flooded Forest Alliance
(Floodplain Swamps). A single representative association of this alliance was
found on BBMNWR, the Nyssa aquatica L.(swamp tupelo)-Nyssa biflora Walt.
(swamp blackgum) Forest Association. This association occurs along a small,
meandering, unnamed tributary SE of Bremermann Road near the W bound-
ary of the refuge (Fig. 2A).Common trees in the subcanopy include Acer rubrum
L. var. drummondii (Hook. & Arn. ex Nutt.) Sarg. (swamp red maple) and
Fraxinus caroliniana Mill. (water ash). Common shrubs are Cephalanthus
occidentalis L. (buttonbush), Cornus foemina Mill. (gray dogwood), Itea
virginica L. (Virginia-willow), and Planera aquatica (Walt.) J.R Gmel.
(planertree). Common herbaceous species occurring in soils saturated for long
duration include Echinodorus cordifolius (L.) Griseb. (heart-leaf burhead), Iris
virginica L. (southern blue flag), Panicum gymnocarpon Ell. (swamp panic-
grass), Polygonum densiflorum Meisn. (denseflower knotweed), and Saururus
cernuus L. (izard’s tail). Woody and herbaceous species occuring on small hum-
1194 BRIT.ORG/SIDA 20(3
4 a Pay ae
al
Phy, Cahal
aquatica and N. biflora. B. N bi
7 t J 7 7
Crataegus opaca
g
and Sapium sebeferum. C. Example of hardwood | k along Boy $ Road Q ji D. Pine
J 7 ! XY
Wi oe We as Vlisch 1 hate * ID
J 7 J t I J
ROSEN ET AL., F LUKA UF WILDLIFE REFUGE 1195
mocks, tree bases, and other higher places within this forest association include
Carex debilis Michx. var. debilis. (weak caric-sedge), C. lonchocarpa Willd.
(spear-fruit caric-sedge), C. louisianica L.H. Bailey. (Louisiana caric-sedge), C.
reniformis (L.H. Bailey) Small (kidney-fruit caric-sedge), C. tribuloides Wahl.
var. sangamonensis Clokey. (sangamon blunt broom caric-sedge), Hypoxis
curtissii Rose (Curtiss yellow stargrass), Ilex verticillata (L.) Gray (winterberry),
and Rhododendron viscosum (L.) Torr. (swamp azalea).
Nyssa biflora Pond Seasonally Flooded Forest Alliance (Gum Swamps). A single
unnamed forest association was found within the swamp blackgum Pond Sea-
sonally Flooded Forest Alliance. This unnamed association would be called a
swamp blackgum/Crataegus opaca Hook. & Arn. (nayhaw) Pond Seasonally
Flooded Forest Association. The canopy of this association is dominated by
swamp blackgum with the mid-story dominated by mayhaw, and is rare at
BBMNWR. This association is a ponded wetland within an upland forest and
should have a conservation rank of G3. Another woody species present is Sapium
sebiferum (L.) Roxb. (Chinese tallow-tree). Lemna minor L. (lesser duckweed)
commonly formsa dense mat on the water’s surface. Characteristic herbaceous
species on pond margins can include Carex gigantea Rudge. (giant caric-sedge),
C. glaucescens Ell. (southern waxy sedge), C. joorii L. H. Bailey (cypress swamp
sedge), southern blue flag, Sabal minor Jacq.) Pers. (dwarf palmetto), Spiranthes
odorata (Nutt.) Lindl. (fragrant ladies’-tresses), and Woodwardia areolata (L.)
Moore. (netted chainfern). During times of drought or low water, exposed soils
are quickly colonized with species such as Eragrostis hypnoides (Lam.) B.S.P.
(creeping lovegrass), Heliotropium indicum L. (Indian heliotrope), and
Lindernia dubia (L.) Penn. (false pimpernel). An example of this alliance can
be viewed S of Boy Scout Road, near its terminus at Bayou Lacombe (Fig. 2B).
Pinus taeda Planted Forest Alliance (Pine Savanna). A Pinus taeda L. (loblolly
pine) Planted Forest Association borders the refuge boundaries N of Bayou
Paquet Road, just E of its intersection with Transmitter Road. Clearcut portions
of this association have developed into herbaceous bogs that overlap into clear-
ings and fire-brakes on refuge boundaries.
Pinus taeda Forest Alliance (Mixed Woods). This alliance serves asa placeholder
for refuge successional forests dominated by loblolly pine. We did not assign an
association name due to the great floristic variation in this alliance. However,
quanitative analysis would surely yield numerous associations. Other canopy
dominates include Liquidambar styraciflua L. (sweetgum), Nyssa sylvatica
Marsh. (blackgum), Quercus similis Ashe. (swamp post oak), Q. falcata Michx.
var. pagodifolia Ell. (cherrybark oak), and southern red oak. Shrub and herba-
ceous strata are well developed and species composition varies with
microtopography and hydrologic regime. See the annotated checklist for other
species found in this alliance.
1196 BRIT.ORG/SIDA 20(3)
Quercus laurifolia—Nyssa biflora Saturated Forest Alliance (Forest Swales). This
alliance best describes associations encountered in forest swales and drainages
within mixed woods. The swamp blackgum-Quercus laurifolia (diamondleaf
oak)-Quercus phellos (willow oak) / dwarf palmetto Forest Association (G3) is
common in the refuge. Other common woody species include swamp red maple,
mayhaw, and water oak. Herbaceous vegetation is often sparse or absent under
dense canopies. However, in clearings and forest openings herbaceous species
include Louisiana caric-sedge, southern blue flag, Leersia lenticularis Michx.
(catchfly grass), Rhynchospora corniculata (Lam.) Gray. (shortbristle horned
beaksedge), lizard’s tail, and netted chainfern.
Quercus (michauxii, falcata var. pagodifolia, shumardii)—Liquidambar styraciflua
Temporarily Flooded Forest Alliance (Oak-Hickory Ridge). This alliance best
describes unnamed associations that occupy the forested portions of BBMNWR
dominated by Quercus michauxii Nutt. (swamp chestnut oak) and sweetgum.
Quercus shumardii Buckl. (shumard oak) is absent on BBMNWR. A commu-
nity dominated by swamp chestnut oak, sweetgum, pignut hickory,and swamp
post oak exists on a broad ridge that spans NW of a small day-use parking area
at the intersection of Bertha Lane and Lucille Road. This association also oc-
curs as a transition or fringe allied with other forest associations.
Quercus virginiana Temporarily Flooded Forest Alliance (Hardwood Hammocks).
A single association was placed under this alliance, a Quercus virginiana Mill.
(live oak) / (dwarf palmetto, Serenoa repens (Bartram) J.K.Small [saw palmetto])
Forest Association. At the terminus of Boy Scout Road, a live oak forest occurs
on old archeological features, and grades into low lying areas that are tempo-
rarily flooded during storm events (Fig. 2C). The shrub stratum is poorly devel-
oped witha few scattered dwarf palmetto. Herbaceous cover is sparse with slen-
der spikegrass, and Elephantopus spp. (elephant’s-foot). Saw palmetto is absent
at BBMNWR.
Woodland Alliances
Pinus elliottii Saturated Temperate Woodland Alliance (Pine Flatwoods). This
alliance is represented by the Pinus elliottit Engel. var. elliottii (slash pine)/
Spartina patens (Ait.) Muhl. (saltmeadow cordgrass)-Juncus roemerianus
Scheele. (black needlerush)-(Panicum virgatum L. [switchgrass]) Woodland
Association (G3). These flat, low-lying wetlands adjacent to the refuge marshes
have an open to sometimes dense canopy of slash pine. Subcanopy and shrub
layers are usually absent, though species such as Morella cerifera(L.) Small (wax
myrtle), Iva frutescens L. (maritime marsh-elder), dwarf palmetto, Baccharis
halimifolia L. (groundsel-tree), B. angustifolia Michx. (saltwater false-willow),
and Chinese tallow tree may be present, particularly in areas not subjected to
fire. The herbaceous stratum is dense and is dominated by saltmeadow
ROSEN ET AL., FLORA OF WILDLIFE REFUGE 1197
cordgrass, switchgrass, Solidago sempervirens L. (seaside goldenrod), and
Euthamia leptocephala (Torr. & Gray) Greene. (bushy fragrant-goldenrod).
Other conspicuous though less common herbs include Symphyotrichum spp.,
Eryngium yuccifolium Michx. (rattlesnake master), Oxypolis filiformis (Walt.)
Britt. leafless cowbane), and Asclepias lanceolata Walt. lance-leaf milkweed).
This association is not inundated by normal daily tides, but receives inunda-
tion during storm events. Water tables are high due to landscape position and
soil texture. Within this association are a few scattered stands of Cladium
mariscus (L.) Pohl subsp. jamaicense (Crantz) Ktikenth. (sawgrass). The best
place to observe this association is along Boy Scout Road (Fig. 2D).
Pinus palustris—Pinus (echinata, taeda) Woodland Alliance (Longleaf Pine Sa-
vanna). A Pinus palustris Mill. longleaf pine)-Pinus (echinata |shortleaf pinel,
loblolly pine) / Schizachyrium scoparium (Michx.) Nash. (little bluestem) Wood-
land Association represents this alliance at BBMNWR. Subcanopy species in-
clude saplings of the nominal species and swamp post oak. Dominant shrubs
are Ilex vomitoria Ait. Gvaupon) and wax myrtle. Herbaceous cover is dense
and is dominated by little bluestem, Andropogon spp., and Aristida longespica
Poir. var. geniculata (Raf.) Fern. (plains threeawn). This forest occurs on mesic
flats near Lake Pontchartrain that are periodically inundated by storm events.
This community is somewhat inaccessible and located several miles from paved
roads in the Salmen Tract in the city of Slidell. Near the end of the study, a pre-
scribed burn management regime was initiated in this woodland alliance. This
will undoubtedly alter the community composition.
Herbaceous Alliances
Nymphaea odorata Permanently Flooded Temperate Herbaceous Alliance
(Sloughs). This alliance is represented by a single association, the Nymphea
odorata Ait. var. odorata (white water lily) Herbaceous Vegetation Association.
This association occurs as semipermanently flooded sloughs, ditches, borrow
ponds, and marsh ponds, usually within other communities. The best place to
observe this association is from an interpretive station along Boy Scout Road
near the parking area (Fig. 3A).
Sagittaria lancifolia Semipermanently Flooded Herbaceous Alliance (Fresh
Marsh). Two associations represent this alliance. The first is a Sagittaria
lancifolia L. (lanceleaf arrowhead) Herbaceous Vegetation Association (G3).
This shallow freshwater marsh is dominated by a monoculture of lanceleaf
arrowhead and is best observed along Boy Scout Road (Fig. 3B). The second is
an unnamed association that forms dense marsh fringes along Cane Bayou (Fig.
3C). This association, based on field observations, would be called a lanceleaf
arrowhead-Schoenoplectus tabernaemontani (K.C. Gmel.) Palla (softstem
bullrush)-Schoenoplectus americanus (Pers.) Volk. (olney threesquare) Herba-
1198 BRIT.ORG/SIDA 20(3)
Fic. 3 Dlant te ENAR HA | IR AS! hal R fal +p J oartth Al } 4. } Pee
rimmed with dead pines that have succumbed to subsidence. B. Fresh marsh along BO Scout Road dominated by
ae nine and aaa ane taonrs: C. Fresh marsh of 5 p j, S. americanus,
Ic] DS a £ . L .
ROSEN ET AL., FLORA OF WILDLIFE REFUGE 1199
ceous Vegetation Association with a proposed conservation rank of G3. Other
common herbaceous species include Eleocharis palustris(L.) R.&S. (great plains
spikerush), Ptilimnium capillaceum (Michx.) Raf. (thread-leaf mock bishop’s-
weed), and Alternanthera philoxeroides (Mart.) Griseb. (alligator weed). A few
widely scattered Taxodium ascendens Brongn. (pond cypress) are present in this
association.
Spartina alterniflora Tidal Herbaceous Alliance (Smooth Cordgrass Marsh). The
Spartina alterniflora Lois. (saltmarsh cordgrass) Louisianian Zone Herbaceous
Vegetation Association s monotypic bands of saltmarsh cordgrass along
tidal streams, canals, and littoral zones on the shore of Lake Pontchartrain.
Spartina patens—(Distichlis spicata) Tidal Herbaceous Alliance (Intermediate
Marsh). The saltmeadow cordgrass-Schoenoplectus (olney threesquare, pungens
[Vahl] Palla lcommon threesquare])-(Distichlis spicata (L.) Greene [saltgrass])
Herbaceous Vegetation Association (G3) inhabits the refuge marshes inundated
by spring tides. Bolboschoenus robustus (Pursh) Sojak (Alkali Bulrush) replaces
olney tl | and common threesquare as found elsewhere in marshy sites
at BBMNWR. Other common herbaceous species include switch grass,
Eleocharis cellulosa Torr. (Gulf Coast spikerush) and black needlerush. A band
of shrubs including groundsel-tree and maritime marsh-elder can occur as a
transition zone between this association and dryer communities or in small
scattered thickets. The best place to observe this community is from the obser-
vation platform on Boy Scout Road (Fig. 3D).
Vallisneria americana Permanently Flooded—Tidal Herbaceous Alliance (Grass
Beds). A Vallisneria americana Michx. (American eelgrass) Estuarine Bayou
Herbaceous Vegetation Association (G3) occurs along the shores of Bayou
Bonfuca, several tidal canals, sloughs, and marsh ponds. Other aquatic plants
common in this association include Myriophyllum sibiricum Kom. (siberian
water-milfoil), Ruppia maritima L. (wigeongrass), and Zannichellia palustris
L. (horned pondweed).
EXOTIC SPECIES OF CONCERN
The following non-native species are, or have the potential to become, serious
and difficult to control weeds and may threaten the diversity of plant commu-
nities of BBMNWR. Woody Species: Chinese tallow-tree, a native of China and
Japan, and a tenacious weed of the SE United States, was encountered in most
of the refuge forests and woodlands. This species is not well established in these
habitats, but should be monitored closely. Ligustrum sinense Lour. (common
privet), a native of Europe, is well established and frequent in mixed woods and
oak hickory forests. Poncirus trifoliata (L.) Raf. (trifoliate orange) is native to
Asia. This species has escaped from cultivation near an old homesite at the ter-
1200 BRIT.ORG/SIDA 20(3)
minus of Boyscout Road and has become well established in hardwood ham-
mocks. Armed with stout spines, this shrub forms inpenetrable thickets. An-
other native to Asia, Rosa bracteata J. Wendl. (Macartney rose), is occasional in
mixed woods. Woody and Herbaceous Vines: Lonicera japonica Thunb. Japa-
nese honeysuckle) isa climbing vine that was occasionally encountered in oak
hickory forests. A more troublesome herbaceous vine was Lygodium japonicum
(Thunb.) Sw. Japanese climbing fern), a native of Asia and the Austral Tropics.
This vine was well established in mixed and beech magnolia woods. Herbaceous
Species: Alternanthera philoxeroides (Mart.) Griseb. (alligator weed), native to
Australia, was frequent in fresh marsh and ditches. Colocasia esculenta (L.)
Schott (wild taro), a native to Asia, was restricted to roadside ditches SE of
Bremermann Road near the W boundary of the refuge. Imperata brasiliensis
(L.) Beauv. (Imperata) is Native to Tropical America and becoming well estab-
lished along Boyscout Road and other nearby locations. Other grasses occasional
to frequently abundant in disturbed areas with the potential to become a prob-
lem include Paspalum dilatatum Poir. (dallis-grass) and Paspalum urvillei
Steud. (Vasey-grass), both native to South America, Paspalum notatum Flugge.
(bahia-grass) native to Latin America, and Sorghum halepense (L.) Pers.
~~
VJohnson-grass), native to the Mediterranean.
ANNOTATED CHECKLIST OF SPECIES
Plant names are arranged by class, and then listed alphabetically within class
by family, genus, species, and infraspecific rank using the classification sys-
tems found in Cronquist (981), Kartesz (1994), Jones et al. (1997), or more re-
cent classifications found in monographs. Applicable synonymy for select spe-
cies is provided when considered useful. Each taxon is followed by a brief
description of its habitat type(s). Next is a subjective estimate of relative abun-
dance using the following scheme: very rare-known from only a single locale
with few individuals; rare-known from more than one locale, generally with
small populations or one locale with numerous individuals; infrequent-not
always in the stated community type(s), usually in small numbers; occasional-
often in the community type(s), but rarely in large numbers; frequent-usually
encountered in the community type(s), but not always in large numbers; and
abundant-expected in the type, usually in large numbers. Collectors and col-
lection numbers are David J. Rosen (DJR), James M. Joyner MJ), and Gary M.
Couret (GMC). Non-native taxa are based on Thomas and Allen (1993, 1996,
1998) and indicated by an asterisk (*).
ROSEN ET AL
WILDLIFE REFUGE 1201
LYCOPODIOPSIDA
LYCOPODIACEAE
ae fella alopecuroides (L.) R.Cranfill [Lycopo-
dium alopecuroides L.J, bogs, frequent, DJR
ISOETOPSIDA
SELAGINELLACEAE
Selaginella apoda (L : one oak hickory ridge,
occasional, DJR
POLYPODIOPSIDA
ASPLENIACEAE
Asplenium platyneuron (L.) Oakes. ex D.C. Eat., oak
hickory ridge, occasional, JMJ 1308
BLECHNACEAE
Woodwardia ee iy .) Moore, forest swales,
occasional, DJR 1
Woodwardia elie i Sm., mixed woods, oc
casional, DJR 1
DENNSTAEDTIACEAE
Pteridium aquilinum (L.) Kuhn, oak hickory ridge,
occasional, DJR 1397
LiMn 558
forest swales, infrequent, DJR
1388.
OSMUNDACEAE
Osmunda regalis |
1066
PARKERIACEAE
*Ceratopteris thalichtroides (L.) Brongn., gum
mops, very rare, DJR 1746
POLYPODIACEA
Pleopeltis po ue (L.) Andr
michauxiana (Weath.) Andrews &
Windham, epiphyte on live oaks, frequent,
'/ 1343
forest swales, occasional, DJR
1c RB \Alindham
LYGODIACEAE
*lygodium japonicum (Thunb.) Sw., mixed
woods, beech magnolia woods, occasional,
IMJ 1328.
THELYPTERIDACEAE
lai kunthii (Desv.) Morton., roadsides,
xed woods, frequent, DJR 1060.
PINOPSIDA
PINACEAE
Pinus elliottii Engelm. var. elliottii, pine flatwoods,
mixed woods, abundant, DJR 1953.
=“ me L.,, pine flatwoods, mixed woods,
eaf pine savanna, abundant, DJR 984.
ae Saas Walt, beech 1s, ridges,
occasional, DJR
Pinus palustris ae eee pine savanna, fre-
q
TAXODIACEAE
Taxodium eae, Brongn.,fresh marsh, infre-
be uen ae 303.
L., bayou edge, fresh marsh
occasional, JMJ 12917.
LILIOPSIDA
ALISMATACEAE
Echinodorus cordifolius (L.) Griseb., floodplain
requent, DJ
Sagittaria graminea Michx., borrow ditches, for-
est swales, frequent, JMJ 13175.
Sagittaria lancifolia L., fresh marsh, borrow
ditches, abundant, DJR 1356
ALLIACEAE
Allium canadense L., disturbed places, frequent,
DJR 1038.
AMARYLLIDACEAE
Crinum aan a ale W of Lake Road,
occasional, DJR
Hymenocallis lirosome ont, Shinners, forest
swales, occasional, D/JR 999
ARACEAE
Arisaema triphylla (L.) Schott. subsp. quinatum
Hutt., oak hickory ridge, infrequent,
IMJ 1273
naeeee
—
ok
oC
locasia esculenta (L.) Schott, borrow ditches,
rare, DJR 1225.
Peltandra virginica (L.) Schott, bayou edge, infre-
quent, DJR 10
Sabal minor (Jacq.) Pers., mixed woods, forest
wales, gum swamps, frequent, JMJ 1314
BROMELIACEAE
Tillandsia usneoides L., mixed woods, frequent,
1202
COMMELINACEAE
Commelina diffusa Burm.,forest swales, frequent
DJR 1020
Commelina virginica L., forest swales, occasional
Tradescantia ieuetore sel roadsides, mixed
woods asional. DJR 1392.
Tradescantia i Ly ot hickory ridge, oc-
casional, J
CYPERACEAE
Bolboschoenus robustus (Pursh) Sojak wills
robustus Pursh], intermediate marsh, fre
quent, DJR 1230
| ta Mack, oe vagnolia Vv 15
nal, DJR
Carex a a icans aie ex orene var. australis
reine J.Rettig, oak hickory ridge, frequent,
ne iene Schwein. beech magnolia
woods, infrequent 1374
Carex ae Torr. & Hoo k, oak
ri geen mixed woods, frequent, D/
Carex crebriflora Wieg., floodplain ae
quent, DJR 1990.
Carex debilis Michx. var. debilis, floodplain
swamps, infrequent, DJR 1437.
Carex digitalis Schkuhr ex Willd. var. floridana
(Bailey) R. Naczi & C. Bryson, beech magno-
lia woods, occasional, DJR 1370
Carex flaccosperma Dew., oak hickory ridge, fre-
,DIR 1025
th, forest swales, infrequent, DJR
Carex frankii|
IT10.
Carex gigantea Rudge, gum swamps, rare, DJR
1103
(a | 5 FIL pl f| 15 occasional
DIR 9
Paine: ic H. Bailey, gum swamps, occasional,
DIR 1744.
Carex se apes floodplain swamps, in
frequent, DJR
Carex longli ie. j hickory ridge, mixed
woods, frequent, DJR 1028.
Carex louisianica L.H. Bailey, forest swales, flood
pla ain swamps, infrequent DIR 1432
5
Carx lurida Wahl, borrow ditches, fahrecueni DJR
2003.
Carex nigromarginata >cnWeln. var. MeL ana
(Schwein,) Kukenth, b
acento 974,
BRIT.ORG/SIDA 20(3)
Carex reniformis (L. H. Bailey) oo floodplain
wamps, occasional, DJR 7
Carex striatula el ial cok: woods,
occasional, DJR |
Carex texensis (Torr.) L ] ae places,
infrequent, DJR 1041.
Carex triangularis ee — magnolia woods,
occasional, DJR |
Carex tribuloides ee var. sangamonensis
mai beech magnolia woods, floodplain
mops, occasional, DJR 1368.
rie mariscus (L.) Pohl subsp. jamaicense
(Crantz) Kukenth., fresh marsh, pine
flatwoods eniee DIR 1207.
Cyperus croceus Vahl, disturbed places, frequent,
Cyperus haspan L., forest swales, frequent, JMJ
1243.
Cyperus odoratus L. var. odoratus, Intermediate
sh, occasional, DJR 1672
Cyperus polystachyos Rottb., fresh marsh, occa-
sional, om 1259
Steud. var. pseudovegetus
“ghatanees places, frequent, DJR 1085.
Cyperus retrorsus Chapm., disturbed places, fre-
quent, DJR 952
Cyperus thyrsiflors Jung., disturbed places, fre-
quent, DJR 1084
Cyperus virens Michx.var.virens, disturbed places,
Forest swales, frequent, DJR 1083
Eleocharis cellulosa Torr., intermediate marsh,
pine flatwoods, frequent, DIR 896.
Fleocharis microcarpa Torr. var. Microcarpa, pine
flatwoods, frequent, DJR 901
cl baric tovi7l ra ears |
frequent, DJR 1000.
Fleocharis obtusa ae Schult. forest swales,
infrequent, DJR 1
Eleocharis olivacea io, tech marsh mud flats,
infrequer it, DIR 126
Eleocharis palustris (L.) a &S, fresh marsh, abun-
dant, D/JR 1984
Eleocharis parvula (R. & S.) Link var. anachaeta
rr.) Svens., fresh and intermediate marsh
ud flats, frequent, DJR 843.
Eleocharis deine es al one R.&S, borrow
ditches, rare, DJR |
Eleocharis tuberculosa “rae R. & S., pine
flatwoods, frequent, DJR 897
ROSEN ET AL., FLORA Of
WILDLIFE REFUGE 1203
Fimbristylis caroliniana (Lam.) Fern., pine
latwoods, frequent, DJR 1249
Fimbristylis castanea (Michx.) Vahl. intermediate
marsh h, frequent DIR 92?.
Fimbristylis dichotoma (L.) Vahl, disturbed places,
occasional, DJR 1279.
*Fimbristylis miliacea (L.) M. H. Vahl, disturbed
places, occasional, DJR 1706.
Furlena breviseta oe Cov., pine flatwoods, oc-
casional, DJR 8
Isolepis carinata ee & Arn. ex Torr. [Scirpus
kollolepis (Steud.) Gl.], wet, sandy forest roads,
TeqHent ee 1349.
Kylingia brevifolia Rottb., disturbed places, infre-
quent, DJR
cies ct Ell., mixed woods, fre-
10
’
Pres ie nate A.Gray, wet clearcuts,
frequ UR 1234
Rhynchospora colorata (L.) H. Pfeiffer
[Dichromena colorata (L.) Hitchc.], pine
flatwoods, occasional, DJR 9
ee pe corniculata (Lam.) A Gray, pine
woods, forest swales, frequent, JMJ 1327.
apa debilis Gale, bogs, frequent, DJR
i we elliottii D. Dietr., wet clearcuts, fre-
quent, DJR 1233
ss area fascicularis (Michx.) Vahl, pine
ds, freq 8.
Ss)
Go
flatwoo
Rhynchospora graci ae A. Gray, wet clearcuts,
frequent, DJR 1235
Rhynchospora inexpansa (Michx.) Vahl, pine
latwoods, frequent, DJR 890
Rhynchospora perplexa Britt., wet clearcuts, fre-
1176.
Rhynchospora rariflora (Michx.) Ell, wet clearcuts,
occasional, DJR 1120
Schoenoplectus americanus (Pers.) Volk. [Scirpus
americanus Pers.], fresh marsh, frequent, DJR
1082.
Schoenoplectus tabernaemontani (K.C. Gm
Palla [Scirpus validus Vahl], fresh marsh oe
quent, DJR 1087.
Scleria eee hx.,oak hickory ridge, infre-
=
Ce
AG
Scleria pauciflora rat ex ee var. pauciflora,
vanna, occasional, DJR 1955.
Scleria triglomerata Michx., wet oe fre-
qu 951.
,
Scleria verticillata Muhl. ex ee ee pine
avanna, infrequent, DJR
ERIOCAULACEAE
Eriocaulon decangulare L., bogs, occasional, JMJ
1320
HYDROCHARITACEAE
Vallisneria americana Michx., bayous, roadside
als, occasional, DJR 2005
HYPOXIDACEAE
Hypoxis curtissii Rose, floodplain swamps, infre-
,DIR 1997
Hypoxis hirsuta (L.) Cov., roadsides, infrequent,
DJR 990
IRIDACEAE
Iris hexagona Walt., borrow ditches, frequent, DJR
1075.
Ir aes gum swamps, forest swales, bayou
es, frequent, JMJ 1297.
; ei atlanticum Bickn., roadsides, fre-
quent, DJR 1009
*Sisyrinchium exile Bickn., disturbed places, fre-
quent, DJR 1012
JUNCACEAE
Juncus oe ey pine flatwoods,
occasional, DJR
Juncus bufonius L., a gees occasional, DJR
luncus ; Mack. forest swales, occasional
IR 1732.
Juncus He ae borrow ditches, oc-
asional, DJR 1
ae effusus L., oy ditches, frequent, DJR
1016.
Juncus elliottii Chapm., wet clearcuts, frequent,
IIR 101
0.
Juncus marginatus jee disturbed places, oc-
casional, 104
Juncus polycephalus ee borrow ditches, fre-
quent, DJR 10
Juncus repens ee forest swales, occasional,
DJR 1197
loncus roemenanus Sc heele j
abundant, DJR 986.
Juncus tenuis willd. var. eles = ) Wood,
est swales, occasional, DJR 1
incus tenuis Willd. var. tenuis, ane is fre-
quent, DJR 1994
1204
LEMNACEAE
Lemna minor L., gum swamps, frequent, DJR
NAJADACEAE
es guadalupensis (Spreng.) Magnus, Cane
Bayou canoe launch, frequent, DJR 1203.
ORCHIDACEAE
Spiranthes odorata (Nutt.) Lindl., gum swamps,
infrequent, 1745
Spiranthes praecox (Walt.) S. Wats., beech mag-
nolia woods, occasional, JMJ 1312
POACEAE
Agrostis hyemalis (Walt.) B.S.P., roadsides, occa-
sional, DJR 1068
Andropogon glomeratus (Walt.) B.S.P. var.
glaucopsis (Ell.) Mohr, longleaf pine savanna,
frequent, DJR 1664.
sta glomeratus (Walt.) B.S.P. var.pumilus
y, longleaf pine savanna, frequent, DJR
en
Andropogon gyrans Ashe var.stenophyllus (Hack.)
C. Campb., longleaf pine savanna, frequent
DJR 1676
Andropogon virginicus L. var. virginicus, mixed
ds, clearcuts, abundant, DJA 1684
Anthaenantia rufa (Ell.) Schult, bogs, infrequent,
=>
DIR 948.
Aristida longespica Poir.var.geniculata (Raf) Fern,,
longleaf pine savanna, occasional, DJR 893.
Arundinaria gigantea (Walt.) Muhl. subsp.
gigantea, mixed woods, occasional, DJR 1423.
a fissifolius ioe Kuhlm., pine
flatw s, occasional, DJR 899
*Briza minor L., roadsides, Hine DJR 1014
*Bromus catharticus Vahl, disturbed places, fre.
quent, DJR 1039
Chasmanthium laxum (L.) Yates var.laxum,mixed
ce aaa magnolia woods, abundant,
IR I
Chasmant ae laxum (L.) H. Vv
sessiliflorum (Poir.) Wipff & S.D. Jones, mixed
woods, infrequent, DJR 1731
Ctenium aromaticum (Walt.) Wood, bogs, infre
quent, DJR 947.
*Cynodon dacty a i Pers,, disturbed places,
Benen t,DUR |
Dig oe tz.
“DIR 1220.
edi roadsides, frequent
+a
BRIT.ORG/SIDA 20(3)
Distichlis spicata (L.) Greene, intermediate marsh,
frequent
“Echinochloa crus- eal (L.) Beauv. var. crus-galli,
fresh marsh, occasional DJR 1268.
Elymus virginicus A var. virginicus, roadsides, oc-
casional, DJR 110
Eragrostis ae S.Wats., pine flatwoods, occa-
sional, DJR 895
Eragrostis hypnoides (Lam.) B.S.P, gum swamps,
Fragrostis spectabilis (F. Pursh) E. von Steudel,
longleaf pine savanna, infrequent, DJR 1666.
*Eremochloa ophiuroides bees ica dis-
turbed places, frequent, JMJ 1
Horder;m pusilum Nutt roadsides pe DIR
O01.
*Imperata brasiliensis (L.) Beauv., roadsides, oc-
casional, DJR 1034. Federal noxious weed
Leersia hexandra Sw., borrow ditches, occasional
IMJ 1332
Leersia lenticularis Michx., forest swales, occa-
siona , 1718
Leersia ices ie Sw., borrow ditches, infre-
t, DIR 1
Leersia winginica Willd. »mixed woods, occasional
Leptochloa ices (L.) Kunth subsp. fascicularis
(Lam.) N. Snow., intermediate and fresh
marsh, frequent, DJR 1673.
*Lolium arundinaceum sete, S.J. ale
[Festuca eb], roadsides, oc
casional, DJR 1087.
*Lolium perenne ‘ disturbed places, frequent,
Melica mutica Wie disturbed roadside, occa-
ional, DIR |
h
1(Poir.) Trin., pine savanna,
infrequent DIR 946
Oplismenus hirtellus (L)B eauv.,, oak hickory ridge,
frequent, DJR 931
Panicum aciculare Desv.ex J. Poiret var.
angustifolium (Ell) Wipff & S.D.Jones., mixed
woods, frequent, DJR 1414.
Panicum acuminatum Sw. var.acuminatum, oak
hickor ridge infrequent, DJR
Panicum acuminatum SW. var. iaseiouiohin
a Lelong., mixed woods, frequent, DJR
Panicum anceps Michx., wet clearcuts, infre-
quent, DJR 954
ROSEN ET Al
WILDLIFE REFUGE 1205
Panicum boscii J.Poiret, beech magnolia woods,
frequent, DJR 1976
Panicum commutatum Schult var. jooril i
mixed woods, frequent, DJR |
eee
Panicum consanguineum Kunth, mixed aoe
frequent 343
Panicum dichotomi oo Michx. subsp.
lic} ifl ites, infrequent
DIR 1659.
Panicum dichotomum C. Linnaeus var. /ucidum
(Ashe) Lelong, beech magnolia woods, fre-
quent, DJR 1373.
Panicum dichotomum L. var. ramulosum (Torr.)
Lelong, oak hickory ridge, infrequent, DJR
1396.
Panicum dichotomum L. var. roanokense (Ashe)
Lelong, mixed woods, frequent, DJR 1346
Panicum A ed ails swamps,
occasional, DJR 17
Panicum hians EIl., sccm occasional, DJR
1069.
Panicum laxiflorum Lam., oak hickory ridge, oc-
casional, DJR 1375
Panicum portoricense Desv. ex Hamilt. var.
nashianum a .) Lelong, mixed woods,
frequent, DJR 1
Panicum repens L., vet leas ditch
abundant, DJR 12
Panicum a ee m - var. a pine
flatwoods occasional, DJR 1747.
SS
ee a rigidulum Nees var. ee (Vasey
Lelong, longleaf pine savanna, infrequent,
DIR 1691.
Panicum scoparium Lam., mixed Forests, fre-
quent, DJR 1193.
Panicum sphaerocarpon Ell, disturbed sites, fre-
quent, DJR 1417.
Panicum verrucosum Muhl., pine flatwoods, wet
clearcuts, frequent, DJR 905.
Panicum virgatum L., See este marsh, pine
flatwoods, frequent, DJ
*Parapholis incurva (L.) C. cae spoil levees,
frequent, DJR 1416.
Paspalum a pal Berg., disturbed places,
frequent, DJR 1269.
/ dil
tatum Poir,, disturbed places, fre-
aang DJR 1114
Paspalum deceit (L.) L., gum swamps, fre-
quent, DJR 1210
Paspalum floridanum Michx. var. floridanum, pine
flatwoods, occasional, DJR 891.
Paspalum floridanum Michx. var. glabratum
Engelm. ex Vasey, roadsides, frequent, DJR
1237.
Paspalum laeve Michx., pine flatwoods, occa-
sional, DJR 892.
Paspalum lividum Trin., borrow ditches, frequent,
1248.
*Paspalum notatum Fluggé, disturbed places,
occasional, JMJ 1326
ee plicatulum Mich var.plicatulum, pine
ods, occasional, DJR 1097.
Pas ae praecox Walt., wet clearcuts, occa-
sional, DJR 949.
ee setaceum ee a pine sa-
nna, frequent, D
Satie urvillei Steud., ee places, Oc-
casional, DJR 900.
Paspalum vaginatum Sw.,, spoil levees, frequent,
1716.
Phalaris a Nees ex Trin., disturbed road-
requent DIR 1402.
side
Phalaris sank Walt., disturbed areas, infre-
quent, DJR 1412.
Phragmites communis Trin., along canals, occa-
sional, DJR 1749.
*Poa annua nen frequent, DJR 981
eee monspeliensis (L.) Desf., disturbed
as, frequent, DJR 1090.
ea baldwinii Spreng. [Erianthus strictus
Baldw.], mixed woods, frequent, DJR 936
Saccharum giganteum (Walt.) Pers. [Erianthus
giganteus (Walt.) Muhl.], mixed woods, fre-
quent, DJR 1656.
Schizachyrium scoparium (Michx.) Nash. var.
scoparium, a flatwoods, mixed woods, fre-
quent
Setaria ree ae Kerguélen, roadsides, fre-
u 1]
*Sorghum halepens : Pers,, disturbed places,
occasional, DJR
Spartina Fe fete ot Intermediate marsh,
occasional, DJR 9
fs ee moses mr 3 oul intermediate
infrequent, DJR 1
Sparin ee (Ait.) Muhl., ae a in-
ediate marsh, abundant, DJR
ee sic obtusata a ales road-
sides, frequent, DJR 979.
1206
Sporobolus indicus (L.) R.Br, roadsides, frequent
DJR 1005
Sporobolus virginicus (L.) Kunth, spoil levees, fre
ent, DIR 1713.
*Stenotaphrum secundatum (Walt.) O. Ktze., dis
turbed places, frequent, JMJ 1341
Tridens ambiguus (Ell.) Schult, pine flatwoods,
roadsides, ‘aireeuenk DUA 50.
Iridens strictus (Nutt.) Nash, pine eee oc
casional, DJR 903.
‘
ae
Tripsacum si oides (L.) L. var. dactyloides, dis-
urbed sides infrequent DJR 1201.
ste her (Walt.) Rydb. var. octoflora, dis-
reas, occasional, DJR 1409
oe. miliacea (Michx.) Doell & Asch., bor-
ow ditches, fresh marsh, frequent, DJR 1052.
PONTEDERIACEAE
Pontederia cordata L., borrow ditches, frequent
R 1070
POTAMOGETONACEAE
Potamogeton diversifolius Raf, small lake at Boy
S infrequent, DJR 1428
RUPPIACEAE
— maritima L., ee Lacombe, marsh
ds, frequent, JMJ 1
SMILACACEAE
Smilax bona-nox L., mixed woods, infrequent,
DJR 1011,
Smilax glauca Walt., mixed woods, occasional,
DIR 1352.
Smilax laurifolia L., bogs, occasional, DJR 19517.
Smilax pumila Walt., mixed woods, occasional,
1385,
Smilax ons ial. sien n swamps, mixed
woods, frequent, DJA 9
Moronq las woods, frequent,
milax i
DIR 1351.
Smilax tamnoides L., mixed woods, occasional,
DIR 1954.
Smilax walteri Pursh, _ magnolia woods,
occasional, DJR 136:
TYPHACEAE
Typha latifolia L., oe marsh, borrow ditches,
frequent, DJR 1
BRIT.ORG/SIDA 20(3)
XYRIDACEAE
Xyris ah ee ex Kunth, pine savanna, oc-
casional, JMJ 1247.
Xyris laxifolia ie var. Iridifolia ia Kral,
DJR 1710.
pine savanna, occasional IR
Xyris stricta Chapm., pine savanna nen DJR
1198
ZANNICHELLIACEAE
Zannichellia palustris L., Lacombe, marsh
onds, frequent, JMJ 1
MAGNOLIOPSIDA
RENT HAGEDE
DIR 1053.
ACERACEAE
Acer rubrum L. var. drummondii (Hook. & Arn. ex
Nutt.) Sarg. floodplain swamps, forest swales,
frequent, JMJ
Acer rubrum L. vat. rubrum, es magnolia
O sional, DJR 13!
occasional
NR
Ga
\©o
Acer rubrum “ var. tri ne =a a A. Gray ex K.
orest swales, occasional, DJR sight
record.
ARANTHACEAE
*Alternanthera philoxeroides (Mart.) Griseb., bor-
row ditches, fresh marsh, frequent, DJR 975.
*Amaranthus cf. cannabinus (L.) Sauer, fresh
marsh, abundant, DJR 1264
ANACARDIACEAE
Rhus copallinum L., mixed woods, frequent, DJR
1255.
Toxicodendron radicans (L.) Kuntze, throughout
abundant, DJR 2006.
APIACEAE
*Centella erecta (L.f.) Fern., pine flatwoods, wet
clearcuts, frequent, JMJ 1220.
a tainturieri Hook., disturbed road-
occasional, DJR
“tse ieotephvllar (Pers) Sprag. ex
t.& Eis, roadsides neuen DJR he
FE ichx k
uent, DJR 1199.
Hydrocotyle verticillata Thunb., palustrine areas,
frequent, DJR 1042.
Oxypolis filiformis de Britt, pine flatwoods,
occasional, DJR
ROSEN ET AL., FLORA OF
WILDLIFE REFUGE
1207
Ptilimnium capillaceum (Michx.) Raf., floodplain
swamps, fresh marsh, frequent, DJR 2000.
Sanicula canadensis L., mixed woods, frequent,
DJR 1046
APOCYNACEAE
Trachelospermum difforme (Walt.) A.Gray., mixed
woods frequent DIR 1337
AQUIFOLIACEAE
Ilex cassine L., wet woods, very rare, DJR 909.
llex decidua T, Walter, wet woods, rare, DJR 1987.
Ilex glabra (L.) A. Gray, mixed woods, occasional
DIR 1348.
Ilex a ia Walt., mixed woods, frequent, DJR
llex opaca Ait., mixed woods, occasional, DJR 994
llex verticillata (L.) A. oS floodplain swamps,
infrequent, DJR 1
llex vomitoria Ait., eee woods, frequent, DJR
993.
ARALIACEAE
Aralia spinosa L., oak hickory ridge, frequent, JMJ
1344
ARISTOLOCHIACEAE
Aristolochia serpentaria L., beech magnolia
woods, infrequent, DJR 1978
ASCLEPIADACEAE
Asclepias lanceolata Walt., pine flatwoods, fre-
quent, JMJ 1304.
Asclepias longifolia Michx., bogs, infrequent, DJR
is (Michx.) Sherff, wet clearcuts, infre-
quent, oF 942.
Bigelowia nudata (Michx.) DC., pine flatwoods,
frequent, DJR 906.
Boltonia asteroides (L.) L'Her., pine flatwoods,
occasional, DJR 926.
Boltonia a EIl., pine flatwoods, occasional,
DJR 1
Bidens mit
ese odoratissimus (J. F. a Cass.,
we
t clearcuts, infrequent, JMJ
Carphephorus pseudoliatris Cass., wet eee
occasional, 945
Cirstumh
orridulum Michx., roadsides, abundant
IMJ 1262.
Conoclinum coelestinum (L.) DC. [Eupatorium
coelestinum L.], roadsides, frequent, DJR 927
*Conyza bonariensis (L.) Cronq., disturbed road-
sides one DJR
=
aa spoil levees, infre-
Suen DIR 1
Ec oot ae borrow ditches, frequent,
DIR 916
a8
woods, frequent, DJ
Elephantopus carolinianus Raeusch., mixed
687
eee a Bert., mixed woods, occa-
sional, DJR 1696.
Pe ene uc A.Gray, mixed
oy re
Friq
oods, OG-
phi oe roadsides, occasional
~JMJ a7
Erigeron vernus (L.) Torr. & A. Gray, pine savanna,
apna ee 956.
Prin
ad angustifolium Pers., pine flatwoods,
mediate marsh, frequent, DJR 1109.
Meee gonocarpos (Walt.) Shinners, beech
nolia woods, infrequent, JMU 1307
POLE RAS ENE
cmella oppositifolia (Lam.) R. Jansen var. repens
(Walt.) R. Jansen, disturbed roadsides, occa-
sional, DIR 1738
Ambrosia artemisi ela L., disturbed roadsides,
frequent, JMJ 1345
Baccharis angustifolia Michx, pine flatwoods,
occasional, DJR 928
pine
flatwoods, frequent DIR 925.
N bogs, infrequent, DJR 944.
Bidens aristosa (Michx.) N veinen forest swales
occasional, DJR 1708.
lifolium (Lam.) Small, disturbed
roadsides, eek places, frequent, JMJ 1227.
Eupatorium leucolepis (DC.) Torr. & A. Gray, pine
savanna, occasional, JMJ 1225
Eupatorium oe L., mixed woods, occa-
sional, DJR s.
Eupatorium rotundifolium L., pine flatwoods, oc-
casional, DJR 1273.
Eupatorium serotinum Michx., mixed woods, ac-
casional, DJR sight record.
Euthamia Silane sled i a mixed
woods, occasional, D
Euthamia leptocephala ee 4 - ni Greene,
pin
ne flatwoods, frequent, DJR 9
Gamochaeta purpurea (L.) Cabrera an, um
purpureum L.J, waste places, frequent, DJR
1208
Helenium amarum (Raf.) Rock, disturbed road
sid equent, JMJ 1226
Helenium flexuosum Raf., roadsides, pine
flatwoods, frequent, DJR 1067
Helianthus angustifolius L., pine savanna, fre-
JR
Helianthus heterophyllus Nutt
occasional, DJR 943.
reeds microcephala (Sch. Bip.) Cabrera
var. albiflora Ktze.) Cabrera, roadsides, oc-
casional, DJR 2002.
lva annua L., ae roadsides, frequent, DJR
pine savanna,
lva frutescens L., intermediate marsh, roadsides,
abundant, DIR 1242.
a (Raf.) Chambers, roadsides, fre-
“quent DIR 1003.
Mikania aries ia pine flatwoods, oc-
casional, JMJ 12
Krig
INI
Packera Hie la ns — C. Jeffrey [Senecio
glabellus J. oo disturbed areas, infre-
quent, DIR I
Pity ONSIS ifoly
(a Y
—
Michx.) Nutt., hummocks
on marsh edge, occasional, DJR 1698
Pluchea rosea Godfrey, roadsides frequent, DJR
1245,
Pyrrhopappus ae iflorus (D.Don) DC, roadsides,
1078.
occasiona ae
lidag 7 Ait., pine flatwoods, occasional,
IMJ 1223
Solidago canadensis L.var.scabra (Muhl.ex Willd.)
Torr. : - e disturbed places, occasional,
DIR 1
oko ag Mill., oak hickory ridge, frequent,
MJ 1
ees sempervirens L., intermediate marsh,
MJ 1305.
pine flatwoods, occasional, J
*Soliva sessilis R.& P, disturbed places, frequent,
DIR 1963
*Sonchus asper (L.) Hill, rodsides, frequent, DJR 976.
Stokesia laevis (Hill) Greene, pine savanna, infre-
q :
Symphyotrichum divaricatum (Nutt.) Nesom [As-
ter divaricatus (Nutt.) Torr. & A. Gray], mixed
s, frequent, DIR 1669.
pene dumosum (L.) Nesom [Aster
dumosus L.], pine flatwoods, occasional, DJR
933.
Symphyotrichum lateriflorum (L.) Love & Love
BRIT.ORG/SIDA 20(3)
oe lateriflorus (L.) Britt.], mixed woods,
casional, DJR s.
pc ae Aum patens (Ait.) Nesom [Aster pat-
Ait.], mixed woods frequent DIR 1699,
Symphyotrichum praealtum (Poir.) Nesom [Aster
praealtus Poir.J, mixed woods, frequent, DJR
1667.
Symphyotrichum subulatum (Michx.) Nesom [As-
ter subulatus Michx.], pine flatwoods, fre-
quent, DJA
Symphyotrichum tenuifolium (L.) Nesom [Aster
tenuifolius L.], intermediate Marsh, frequent
DIR 957
>
“Taraxacum officinale G.H. Weber ex Wiggers,
adsides, Common 1255
Verbesina virginica L., roadsides, frequent, DJR
sight record
Xanthium strumarium L., roadsides, infrequent,
711,
*Youngia japonica (L.) DC., roadsides, frequent,
BETULACEAE
Carpinus caroliniana Walt., mixed woods, beech
maanolia woods frequent DJR 1079.
BIGNONIACEAE
Bignonia capreolata L., mixed woods, frequent,
Campsis radicans (L.) Seem.ex Bur., mixed woods,
frequent, JMJ 1325.
BORAGINACEAE
“Heliotropium indicum |
gum swamps frequent
Myosotis macrosperma Engelm., mixed woods,
ent, DJR 1050
BRASSICACEAE
Lepidium ans um L., disturbed places, occa-
sional, 1048.
Rorippa palustris (L.) Bess
(Butt.& E. Abbe) B.E
sional, DJR 2001.
BUDDLEJACEAE
Polypremum procumbens L., pine flatwoods,
roadsides, occasional, DJR 1251
CALLITRICHACEAE
Callitriche heterophylla Pursh., borrow ditches,
occasional, DJR 1389.
. subsp. fernaldiana
Jonsell, roadsides, occa-
ROSEN ET AL., FLORA OF
WILDLIFE REFUGE 1209
CAMPANULACEA
Lobelia brevifloia re ex A. DC, ene pine
occasional, DJR 168
lobe Hct: Michx., ee ikon ridge, pine
ods, occasio
Poy spicata Lam., lene is woods,
occasional, DJR 1730
Triodanus perfoliata (L.) Nieuw., roadsides, fre-
quent, DJR 1043.
CAPRIFOLIACEAE
*Lonicera japonica Thunb., oak hickory ridge,
occasional, JMJ 1276.
Sambucus eee L., disturbed roadsides,
occasional, DJR 1105.
Viburnum cane lke i. —e ridge, mixed
woods, frequent, DJR
CARYOPHYLLACEAE
“Cerastium glomeratum Les disturbed road-
es, occasional, J
eae marina (L.) se spoil levees, fre-
quent, DJR 1410.
CERATOPHYLLACEAE
Ceratophyllum demersum L., Cane Bayou canoe
launch, frequent, DJR 1202
CISTACEAE
Lechea tenuifolia Michx., frquent, marsh edge,
R 1671
CLUSIACEAE
at cum Uta um (Spach) Steud, pine
s, frequent, DIR 1247.
bee cum en Lam., pine flatwoods, oc-
casional, JMJ 1221
Hypericum denticulatum Walt., pine flatwoods,
occasional, DJR 1250.
Hypericum gymnanthum Engelm. & A. Gray, for
est swale, occasional, D/R 1194.
Hypericum hypericoides (L.) Crantz, oak hickory
ridge, frequent, JMJ 1238
CONVOLVULACEAE
Calystegia sepium (L.) R.Br. roadsides, occasional
1313.
Dichondra ca i:
quent, DJR 1
Ipomoea es oe intermediate marsh, fre-
quent, DJA 1229.
is Michx., waste places, fre-
CORNACEAE
Cornus foemina Mill, floodplain swamps, fre-
R 1438
CUCURBITACEAE
Melothria dota a L., disturbed Forest, occa-
sional, DJR 1422
CUSCUTACEAE
Cuscuta indecora pe intermediate marsh,
abundant, JMJ
DROSERACEAE
Drosera capillaris Poir. var. capillaries, bogs, fre-
quent, 1959
EBENACEAE
Diospyros vriginiana L., mixed woods, frequent,
ERICACEAE
Rhododendron viscosum (L.) Torr., floodplain
swamps, infrequent, DJR 1988.
Vaccinium arboreum ale oak hickory ridge,
occasional, JMJ 1
Vaccinium conbosum a mixed woods, occa-
sional, DJR 134
Vaccinium stamineum L., mixed woods, occa-
sional, 1338
BUEHORBIACENE
Michx. var. lindheimeri (Engelm.
&A. Gravel Muell.-Arg., roadsides, frequent
DIR 1274.
Euphorbia humistrata Engelm., roadsides, dis-
turbed places, frequent, DJR 1243.
raphorb maculata L., roadsides, disturbed
s, frequent, DJR 917.
ape prostrata Ait., roadsides, disturbed
places, frequent, JMJ 1229
Phyllan thus nica: Walt, gum swamps, fre-
quent, DJR 1211.
*Phyllanthus urinaria st ee disturbed
places, occasional, D
*Sapium aut eite hee woods,gum
swamps, pine flatwoods, occasional, DJR
FABACEAE
x AILS) “eaen ee : ; | Nura
roadsides, infrequent
DIR 1213
Amorpha fruticosa L., floodplains, occasional, DJR
1210
Baptisia alba (L.) Vent. var. macrophylla (Larisey)
Isely (Baptisia man ye Thieret], mixed
woods, occasional, D
entrosema virginiatum . Benth., roadsides, fre-
quent, DJR 1253.
Chamaecrista fasciculata (Michx.) E. Greene,
mixed woods, frequent, DJR 1689.
Desmanthus illinoensis (Michx.) MacM., roadsides,
frequent, DJ 2
Desmodium pauciflorum (Nutt.) DC., mixed
wo 1058.
Desmodium sessilifolium Torr. & A. Gray, mixed
woods frequent DIR 1694
Erythrina herbacea L., tts woods, oak hickory
ridge, frequent, JMJ 1
sate triacanthos | fie frequent, DJR
1037.
hickory ridge, frequent, DJR
Lespedeza capitata Michx., mixed woods, fre-
quent, DJR 1695
*Medicago arabica (L.) Huds., roadsides, frequent
IMJ 1261.
*Medicago lupulina L., roadsides, occasional, DJR
1981,
*Melilotus indicus (L.) All, roadsides, infrequent,
IMJ 1263
ent,
Sesbania drummondii (Rydb.) Cory, Intermediate
Mars
Neptunia lutea (Leavenw.) Benth., roadsides, fre-
quent, DJR 122
equent DI j
*Sesbania punicea (Cav.) Benth. Floodplains, oc-
casional, JMJ 1283.
Tephrosia spicata (Walt.) Torr. & A. Gray, mixed
woods, occasional, DJR 1702
"Trifolium campestre Schreb.var. laa dis-
turbed place es, occasional, DJR
Trifolium dubium Sibth, disturbed ee fre-
nt, DIR 1045.
“Trifolium repens L., disturbed places, frequent,
IMJ 1280.
*Vicia sativa L., disturbed areas, frequent, DJR 1413.
Vicia tetrasperma (L.) rile disturbed road-
sides, occasional, JMJ 1282.
‘Vicia villosa Roth, ie roadsides, common,
SJiat
ede
DJR 1404.
Vigna luteola Jacq.) Benth, inter
occasional, JMJ 1299
Wisteria frutescens (L.) Poir, mixed woods, occa-
sional, DJR 1353
}
LC Tahoe,
BRIT.ORG/SIDA 20(3)
FAGACEAE
Fagus grandifolia Ehrh., beech magnolia woods,
f
requent, DJR 126
Quercus alba L., mixed woods, infrequent, DJR
1434,
Quercus falcata Michx. var. falcata, mixed woods,
eech magnolia woods, frequent, JMJ 1331.
pasiee falcata Michx. var. pagodifolia Ell., mixed
oods, frequent, DJR 1095.
avers laurifolia Michx., mixed woods, forest
5, frequent, DJR 995.
Quercus fue ndica Muenchh., hummocks on
arsh edge, rare, DJR 1700
see ichauxii Nutt. oak hickory ridge, mixed
woods, frequent, DJR 9
Dict nigra L., mixed Wooes forest swales,
occasional, DJR
Quercus phellos i tones swales, occasional, DJR
99].
Quercus similis Ashe, os Riek ridge, mixed
woods occasional, D
Quercus sinuata Walt. var. sinuata, forested
swales, rare, DJR 1113.
Quercus virginiana Mill. mixed woods, occa-
sional, DJR
GENTIANACEAE
C : Leb
If f + |
I
infrequent, JMJ 1307.
Eustoma exaltatum (L.) Salib. ex G. Don,, road-
sides, occasional, JMJ 1338
Sabatia calycina (Lam.) Heller, roadsides, occa-
sional, DJR 1071
Sabatia dodecandra (L.) B.S.P, roadsides, infre-
quent, JMJ 1339
GERANIACEAE
Geranium carolinianum L., roadsides, frequent,
DJR 1006
=
GROSSULARIACEAE
[tea virginica L., floodplain swamps, occasional,
DIR 1995
HALORAGACEAE
cee lum cf. aquaticum (Vell.) Verde., gum
infrequent, DJR 1430.
Myon lum pinnatum (Walt.) B.S.P., forest
wales, frequent, DJR 9835.
aay lum sibiricum i canals, marsh
ponds, abundant, DJA |
ROSEN ET AL., FLORA OF
WILDLIFE REFUGE
iat naca pectinata Lam., pine savanna, Oc-
sional, DJR 1117.
HAMAMELIDACEAE
Hamamelis virginiana L., oak hickory ridge, rare,
IMJ 12
Liquidambar styraciflua L., mixed woods, fre-
quent, DJR 987
HIPPOCASTANACEAE
Aesculus pavia L., beech magnolia woods, occa-
sional, DJR 1357.
HYDROPHYLLACEAE
ydrolea ovata Nutt. ex Choisy, palustirne, fre
quent, DJR 1254
JUGLAN
Carya si ae Sweet, mixed woods, oak
hickory ridge, frequent, DJR 1280.
Carya illinoiensis (F. von Wangenheim) K. Koch,
andoned homesite, very rare, JMJ 1281
LAMIACEAE
Hyptis alata (Raf.) Shinners, mixed woods, occa
sional, DJR sight record
Lycopus rubellus C. Moench, forest swales, occa
sional, DJR 1709
*Micromeria brownei (Sw,)
infrequent, JMJ 1286
*Peri la frutescens (L.) ie mixed woods mar
occasional, D
Salvia Her L., roadsides, frequent, JMJ 1258,
Scutellaria integrifolia L., mixed woods, frequent,
DJR 1056.
eee ovata Hill, mixed woods, frequent, DJR
096.
Bentham, roadsides,
QO =
oa s crenata Raf, roadsides, frequent, JMJ 1287.
Stachys floridana Shuttlw. ex Benth., disturbed
areas infrequent, DIR 1426.
Teucrium canadense L., mixed woods, infrequent
IMJ 1346.
Trichostema dichotomum |
sional, DJR 1690
LAURACEAE
Persea palustris (C. Rafinesque-Schmaltz Raf.)
Sarg., mixed woods, infrequent, DJR 1405.
LENTIBULARIACEAE
Utricularia foliosa L., borrow ponds and ditches,
occasional, DJR 1724
Utricularia gibba L., borrow ponds and ditches,
occasional, DJR 16
pine savanna, occa-
1211
Utricularia eee Walt., ale flatwood ponds,
borrow ditches, rare, DJR 8
Utri en radiata Small, pottow ponds and
ditches, occasional, DJR 9
Utricularia subulata |
LINACEAE
Linum medium (Planch.) Britt. pine flatwoods,
frequent, JMJ 1310
LOGANIACEAE
Gelsemium sempervirens St.-Hil., oak hickory
ridge, frequent, DJR 966.
Mitreola petiolata pee ) Torr.& A.Gray, roadsides,
frequent, DJR 12
LYTHRACEAE
Cuphea ae ate 5 Jacq.) Macbr.,roadsides,
frequent, DJR 9
bogs cue DJR 1960.
t h, frequent,
DJIR 1240.
MAGNOLIACEAE
Magnolia grandiflora L., beech magnolia woods,
frequent, DJR 1204.
pine savanna, occasional
NA fe . eae |
IVICACHTPUTICA VEFOITETCAT ECL L
MJ 1319
MALVACEAE
Hibiscus eee Walt., mixed woods, occa-
sional, DJR 1256.
Hibiscus ae L. subsp. lasiocarpos (A.
Cavanilles) O. Blanchard, fresh marsh, flood-
plains, frequent, DJR 1214
Kostelet kya virginica (L.) K. Presl ex sale inter-
mediate marsh, occasional JMJ |
Modiola sane (L.) G.Don sas ie OCCa-
sional, DJR
Sida rhombi | lia . oak hickory ridge, waste
places, abundant, JMJ 1236
MELASTOMATACEAE
exia mariana L., pine savanna margins, fre-
quent, JMU 1317
MORACEAE
Morus rubra |
MYRICACEAE
Morella caroliniensis (Mill.) Small (Myrica
heterophylla Raf.], mixed woods, occasional,
DJR 1350
floodplains, occasional DJR 1036.
—-
Morella cerifera (L.) Small (Myrica cerifera (L.)
mall], mixed woods, pine flatwoods, fre-
quent, DJR 9/4.
1212
—
YMPHACEAE
a advena (Ait.) Ait. (N. /utea (L.) J. E. Sm],
borrow ditches, frequent, DJR 1263.
ee odorata Ait.var.odorata, borrow ponds
and ditches, marsh ponds and sloughs,
abundant, DJR 907.
NYSSAC
Nyssa aquatica L., floodplain swamps, frequent
DIR 1979,
Nyssa biflora Hiss ee gum swamps, Fre
quent, JMJ 1
Nyssa sylvatica on mixed woods, frequent,
DIR 992.
OLEACEAE
Fraxinus caroliniana Mill. floodplain swamps,
occasional, DJR 1080.
ie citar sinense Lour., oak et ridge,
xed woods ane IMJ 12
ONAGRACEAE
Gaura parviflora Dougl. ex Lehm., roadsides, in
frequent, JMJ 1302
Ludwigia alterniflora L., pine flatwoods, occa-
sional, DJR 1272.
Ludwigia pwc Walt., forest swales, occa-
sional, DJR 1707.
Ludwigia ne Walt., pine flatwoods, infre-
quent, DJR 1257.
— octovalvis (Jacq.) Raven, borrow
s, frequent, DJR 12746.
iad a (L.) Ell, forest swale, infrequent,
DIR 1340
Ludwigia pilosa Walt., mixed woods, frequent,
DIR 1653
Oenothera laciniata ie disturbed roadsides,
occasional, DJR 140
Oenothera speciosa es disturbed places, oc-
C I 1296
asional, JM
OXALIDACEAE
*Oxalis oe Kunth, oak hickory ridge, frequent,
DJR
Oxalis ea eee disturbed roadsides, occa-
ional, DIR |
1
Oxalis stricta |
PASSIFLORACEAE
Passiflora lutea L., beech magnolia woods, infre-
quent, DJR 1361.
ore frequent, JMJ 1254.
BRIT.ORG/SIDA 20(3)
Silda aacioamis
PI ye 1 ericana L., roadsides, waste places,
occasional, JMJ 1330.
PLANTAGINACEAE
*Plantago lanceolata L., disturbed areas, infre-
DIR 1427.
*Plantago major L., roadsides, frequent, DJR 958.
Plantago virginica L., roadsides, frequent, DJR
1077
POLYGALACEAE
Polygala cruciata L., bogs, frequent, GMC s.n.
Polygala incarnata L., pine savanna, frequent, JMJ
1324
Dn | Io].
FOIYGUIG IC
( tocaulis Torr. & A. Gray, pine savanna,
infrequent, DJR 1115.
Polygala lutea L., pine savanna, occasional, JMJ
219.
Polygala ramosa FIl., pine savanna, infrequent,
R 1178.
POLYGONACEAE
brunnichia ovata (Walt.) Shinners, mixed woadls,
frequent, DJR 1362.
Polygonum caespitosum i aa oak hickory
t, DJR
ridge, frequen
es el pene nies Mein, floodplain
rps, frequent, DJR 1
KA
Pol | heen esic swales
in oak hickory ridge, occasional, DJR 1171.
a a punctatum Ell, fresh marsh, borrow
es frequent, DJR 908.
es one setaceum o w., disturbed places,
infrequent, DJR
*Rumex “noma Murr, waste places, in-
frequent, IMJ 1288.
*Rumex crispus L., disturbed roadside, occasional,
DIR 1394
PRIMULACEAE
Samolus valerandi L.subsp. parviflorus
roadsides, occasional, JMJ 1256
ss
—
Raf.) Hult,
RANUNCULACEAE
Clematis crispa L., floodplains, infrequent, JMJ
1333.
*Ranunculus marginatus d'Urv., roadsides, fre-
quent, DJK 1024.
*Ranunculus platensis Spreng., roadsides, fre-
uent, DJR 972
Ranunculus pusillus Poir., floodplains, frequent,
IIR 106
ROSEN ET AL., FLORA OF
WILDLIFE REFUGE 1213
*Ranunculus saa ie disturbed places,
pccasional DJR
var. sceleratus, floodplain
swamps, infrequent, DJ
Thalictrum dasycarpum Fisch. & Avé -Lall., dis-
turbed woods, frequent, DJR 1421]
RHAMNACEAE
erchemia scandens (Hill) K. Koch, floodplains,
frequent, DJR 1390.
SACEAE
Crataegus eta
casional, JMJ 1
Crataegus opaca can & Arn, forest swales,gum
amps, frequent, DJR 978.
Prunus serotina Ehrh., oak hickory ridge, occa-
ee oak hickory ridge, oc-
sional, JMJ 1264
*Rosa sen oe Wendl, mixed woods, occa-
sional, DJR 108
*Rosa laevi oe Michi floodplain swamps, oc-
casional, DJR 1224.
Rubus argutus Link, mixed woods, frequent, DJR
998,
Rubus trivialis Michx., roadsides, mixed woods,
frequent, DJR 975,
IACEAE
Cephalanthus occidentalis L.,floodplain swamps,
occasional, JMJ 1250.
Diodia teres V Walt., roadsides, frequent, JMJ 1298.
|
Diodia virgit HUEFICE Lt
pine savanna occasional, JM/
1244.
Galium aparine L., roadsides, frequent, DJR 982.
Galium obtusum J. nes places, fre-
Galium tinctorium ) ae os places,
swamps, frequent, DJR 1
Galium te Michx., oe magnolia
woods, infrequent, DJR 1979.
Mitchella repens L., mixed woods, infrequent, DJR
14
39)
Oldenlandia uniflora L., roadsides, infrequent, DJR
RUTACEAE
*Poncirus trifoliata (L.) Raf., disturbed places,
bund 1708
Zanthoxylum clava-herculis L., disturbed places,
infrequent, DJR 1055.
SALICACEAE
alix nigra Marsh., disturbed places, infrequent
SARRACENIACEAE
Sarracenia alata Wood, bogs, rare, DJR 1952.
SAURURACEAE
Saururus cernuus L., Ls sa swamps, forest
swales, frequent, DJ
SCROPHULARIACEAE
Agalinis fasciculata (Ell.) Raf., roadsides, frequent,
IMJ 1232.
Bacopa caroliniana (Walt.) Robins, borrow
ditches, abundant, JMJ 1309.
Bacopa monnieri (L.) Penn, mud flats, frequent,
J 1342.
Gratiola oo Michx., pine savanna, occasional,
DIR 1196.
Gratiola ramosa Walt., palustrine areas, frequent,
DIR 1008.
Lindernia dubia (L.) Penn,gum swamps, frequent
IR 1212
*Mazus pumilus (Burm.) C. van Steenis, oak
hickory ridge, infrequent, DJR 1021.
Mecardonia acuminata (Walt.) Small, disturbed
places, frequent, DJR 914.
Mecardonia ere (Mill.) Small, roadsides,
infrequent, DJR 1
Micranthemum iets (J. F. Gmel.) Blake,
mudflats, frequent, DJR 1061.
*Scoparia oo cis L., oak hickory ridge, frequent,
IMJ 12:
*Veronica arvensis L.,roadsides, frequent, JMJ 1289.
SOLANACEAE
Solanum carolinense L., roadsides, disturbed
places, frequent, DJR 1057
STYRACACEAE
Halesia diptera Ellis, beech magnolia woods, oc-
casional, DJR 1355
Styrax americanus Lam., oak hickory ridge, infre-
qu 1279
SYMPLOCACEAE
7 ocos tinctoria (L.) Ae iam magnolia
ods, occasional DJR 1
ULMACEAE
Celtis laevigata Willd.,, floodplains, frequent, JMJ
1284
1214 BRIT.ORG/SIDA 20(3)
Planera aquatica (Walt.) J. F. ee _ floodplain MVIOLACEAE
mps, occasional, DJR Viola lanceolata L., pine savanna, frequent, DJR
Ulmus alata Michx., beech eats woods, in- 971,
frequent, DJR 1736. Viola palmata L. var. trilobata (Schwein.) Gingins
URTICACEAE ex DC., beech magnolia woods, occasional,
: 2 1969
Boehmeria cylindrica (L.) Sw., roadsides, frequent non deze,
Dip 1044 Viola primuifolia L., oak hickory ridge, frequent,
DIR 964,
VALERIANACEAE Viola sororia Willd. var. sororia, oak hickory ridge,
Valerianella radiata (L.) Dufr., roadsides, frequent frequent, DJR 965.
DIR 1004
VITACEAE
VERBENACEAE eS arborea - .) KOehne, mixed woods,
Callicarpa americana L., oak hickory ridge, fre- frequent, DIR s.n.
quent, JMJ 1235 ee ee ica (Td Gagnepain, waste
“Lantana camara L., roadsides, infrequent, DJR places, rare, DJR 1226.
1092,
oe Stee quinquefolia (L.) Planch., mixed
woods, frequent, DJR 1395
Phyla n ipa tole (L.) Greene, roadsides, frequent
DJIR 9
Vitis astivalis Michx., mixed woods, frequent, DJR
*Verber ne siliensi waste places, roadsides
frequent, JMJ 130 Vitis cinerea (Engelm.) Engelm. ex Millardet,
te el ae places, occasional, mixed woods, occasional, DJR 1354.
J 1295. Vitis palmata Vahl, oak hickory ridge, occasional,
*Verbena rigida Spreng., disturbed places, occa- DIR 1400.
sional, JMJ 1316. Vitis rotundifolia Michx., oak hickory ridge, fre-
Verbena urticifolia L., roadsides, occasional, DJR quent, JMJ 1278
ACKNOWLEDGMENTS
We are grateful to James Harris, Charlotte Parker, Elizabeth Souheaver, and other
staff members with the Southeast Louisiana Refuge Complex for their assis-
tance, enthusiasm, and financial support. Special thanks to James Joyner who
made many collections and assisted with fieldwork during the early months of
this project, and Gary Couret and Todd Miller for their companionship afield. It
is a pleasure to thank Anne Bradburn and Steve Darwin at Tulane University
for their assistance. The following botanists provided determinations for vari-
ous specimens; Joseph Wipff (Poaceae), Billie Turner (Asteraceae and Fabaceae),
Guy Nesom (Asteraceae), Roland Roberts (Euphorbiaceae and Asteraceae),
Kevin Nixon (Quercus), James Phipps (Crataegus), Bill Reese (Lycopodium), and
Robert F.C. Naczi (Carex). The review and comments of Barney Lipscomb,
Charles Allen, and an anonymous reviewer were very helpful in preparing the
final manuscript presented here. Our thanks to Ms. Anna Edwards (USFWS)
lor preparing figure one.
REFERENCES
Aten, C.M. 1992. Grasses of Louisiana, 2nd ed. Cajun Prairie Habitat Preservation Society,
Eunice, Louisiana.
ROSEN ET AL WILDLIFE REFUGE 1215
Corrett, D.S. and M.C. JoHNsTON. 1970. Manual of the vascular plants of Texas. Texas Re-
search Foundation. Renner, Texas.
Cronauist, A. 1980. Vascular flora of the southeastern United States, Volume 1, Asteraceae.
Univ. of North Carolina Press, Chapel Hill.
Cronauist, A. 1981. An integrated system of classification of flowering plants. Columbia
Univ. Press, New York.
Evus, F.S.1981.St. Tammany Parish: L'autre cote du lac. Pelican Publishing Company, Gretna.
FeNNEMAN, N.M. 1928. Physiographic divisions of the United States. Annals Associ. Amer.
Geographers 18: 261-353.
Fisk, H.N. and E. McFartan, Jr. 1955. Late Quaternary Deltaic Deposits of the Mississippi
River. In: The crust of the earth. Geol. Soc. Amer., Special Pap. 62:279-302.
GANDHI, K.N. and R.D. THomas. 1989. Asteraceae of Louisiana. Sida, Bot. Misc. No. 4:1-202.
Goprrey, R.K. 1988. Trees, shrubs, and woody vines of Northern Florida and adjacent Geor-
gia and Alabama. Univ. of Georgia Press, Athens.
Goorrey, R.K. and J.W.Woorten. 1979. Aquatic and wetland plants of the southeastern United
States. Monocotyledons. Univ. of Georgia Press, Athens.
Goprrey, R.K.and J.W.Wooren. 1981. Aquatic and wetland plants of the southeastern United
States. Dicotyledons. Univ. of Georgia Press, Athens.
Goutb, FW. 1975. The grasses of Texas. Texas A&M Univ. Press, College Station.
Isety, D. 1990. Vascular flora of the southeastern United States, Volume 3, Part 2,Leguminosae
(Fabaceae). Univ. of North Carolina Press, Chapel Hill.
Jones, $.D., J.K. Wipre, and P.M. Montcomery. 1997. Vascular plants of Texas: A comprehensive
checklist including synonymy, bibliography, and index. University of Texas Press, Aus-
tin, Texas.
Kartesz, J.T. 1994.A synonomized checklist of the vascular flora of the United States, Canada,
and Greenland. Timber Press, Portland.
Levtincer, D.L.1985.A field manual of the ferns & fern-allies of the United States & Canada.
Smithsonian Institution Press, Washington, D.C.
LELONG, M.G. 1986.A taxonomic treatment of the genus Panicum (Poaceae) in Mississippi.
Phytologia 61:251-269.
Raprorb, A.E., H.E. AHLes, and C.R. Bet. 1968. Manual of the vascular flora of the Carolinas.
Univ. of North Carolina Press, Chapel Hill.
SAUCIER, R.T. 1963, Recent geomorphic history of the Pontchartrain Basin. Coastal Studies
Ser. No. 9. Louisiana State University Press, Baton Rouge.
STEINMAYER, R.A. 1939. Bottom sediments of Lake Pontchartrain. Bull. Amer. Assoc. Petrol.
Geols. 23:1-23.
THomas, R.D. and C.M. AtLen. 1993. Atlas of the vascular flora of Louisiana, Volume |: fern
allies, conifers, and monocotyledons. The Natural Heritage Program, Wildlife Div., Loui-
siana Dept. of Wildlife & Fisheries, Baton Rouge, LA.
THomaS, R.D.and C.M. Aten. 1996. Atlas of the vascular flora of Louisiana, Volume II: dicoty-
ledons, Acanthaceae-Euphorbiaceae. The Natural Heritage Program, Wildlife Div., Loui-
siana Dept. of Wildlife & Fisheries, Baton Rouge, LA.
1216 BRIT.ORG/SIDA 20(3)
THomas, R.D.and C.M. Atten. 1998. Atlas of the vascular flora of Louisiana, Volume Ill: dicoty-
ledons, Fabaceae-Zygophyllaceae. The Natural Heritage Program, Wildlife Div., Louisi-
ana Dept. of Wildlife & Fisheries, Baton Rouge, LA.
TRAHAN, L., J.J. Braotey, and L. Morais. 1990. Soil survey of St. Tammany Parish, Louisiana. Soil
Conservation Service, U.S. Dept. of Agriculture, Washington, DC.
Weaktey, A.S., K.D. Patterson, S. LANDAAL, M. Pyne, and orHers (compilers). 1998. International
classification of ecological communities: Terrestrial vegetation of the southeastern
United States. Working Draft of March 1998. The Nature Conservancy, Southeast Re-
gional Office, Southern Conservation Science Department, Community Ecology Group.
Chapel Hill, NC.
THE VASCULAR FLORA OF THE OGALLALA ECOTONE ON
THE DEMPSEY DIVIDE, ROGER MILLS COUNTY, OKLAHOMA
Craig C. Freeman Caleb A.Morse
R.L. McGregor Herbarium R.L. McGregor Herbarium
NaNUIE History Museum and Natural History Museum and
Biodiversity Research Center Biodiversity Research Center
University of Kansas sec ee
2045 Con Sane Menie 2045 Constant Avenue
Lawrence, KS 66047-3729, U.S.A. Lawrence, KS ae 3729, U.S.A.
ccfree@ku.edu cmorse@ku.edu
J. Peter Thurmond
Dempsey Divide Research Foundation, Inc.
Thurmond Ranch, In
Cheyenne, OK 73628-9729, U.S.A.
dempseydiv@aol.com
ABSTRACT
A floristic inventory of the 3,755-ha Thurmond Ranch and immediate vicinity, located in southern
Roger Mills County, Oklahoma, was conducted in 2001. Because of its unusual geologic and topo-
graphic setting, which has given rise to a heterogeneous environment and long history of human
habitation, the site has been the subject of intense archeological and paleoclimatological research
since the early 1980s. The pate documented 85 families, 286 genera, and 470 species of vascu-
lar plants in the study area. The five most species-rich families, Asteraceae, Poaceae, Fabaceae,
Euphorbiaceae, and Cyperaceae, collectively account for 51% of ene apse Nou: native species con-
stitute 10% of the total flora. Four No
1 by Euro-A meri-
cans, and Native Americans may have introduced nee species Populations of four species tracked
the Oklahoma Natural Heritage Inventory were discovered, and Epilobium leptophyllum is re-
ported for the first time in Oklahoma. Collection data show a marked floristic difference between
the Tertiary Ogallala eee and the Permian redbeds atone the ecotone. Nineteen naa vegeta-
tion alliances were identified based on observational data
seven
shrubland alliances, and eight herbaceous alliances. Mixed- grass i ie plant ommunities domi-
nate the study area landscape.
RESUMEN
Se realizé un io floristico de las 3,755-ha del oe R Raney adores localiz — en
el sur del condado de Roger Mills, Oklahoma, en 2001. Debido |
y topograficas, que dieron lugar a un ambiente es y eee historia de ocupacion bouiine el
lugar ha sido objeto de quiche uriveetigacren arqueologica y paleoclimatolégica desde principio de
los 1980. EL i 286 géneros, y +70 especies de eis vasculares en el
area de estudio. Las cinco familias mas ricas, Asteraceae, Poaceae, Fabaceae, Euphorbiaceae
Cyperaceae, tienen conjuntamente el 51% de las especies. Las especies no nativas comprenden el | 10% %
de la flora total. Cuatro especies norteamericanas fueron introducidas probablemente por Euro-
Americanos, y los nativos americanos puede que hayan introducido tres especies. Se descubrieron
SIDA 20(3): 1217-1245. 2003
1218 BRIT.ORG/SIDA 20(3)
poblaciones de cuatro especies seguidas por el Oklahoma Natural Heritage Inventory, y Epilobium
leptophyllum se cita por primera vez en Oklahoma. Los datos de colecci6n muestran una clara
diferencia floristica entre la formacion terciaria oe uae y las oe sedimentarias rojizas del
pérmico a lo largo del ecotono. Se identificaron alianzas de vegetacion basadas en datos
de observacion, que incluyen tres alianzas arboreas, siete ie y ocho herbaceas. Las
comunidades mixtas de pradera dominan el paisaje del area de estudio.
INTRODUCTION
The name “Dempsey Divide” was coined by Thurmond (1990) to refer to the
uplands between the Washita and North Fork of the Red Rivers in western Okla-
homa. Unusual geologic and topographic conditions in the area have given rise
to a heterogeneous environment, an ecology and biota that are distinctive in
the southern Great Plains, and a long history of human habitation. Conse-
quently, the Dempsey Divide has been the subject of intense archeological and
paleoclimatological research since the early 1980s (Bement 2000; Bement &
Buehler 1994, 1998; Buehler 1997; Thurmond 1990, 1991a, 1991b, 1991c, 1997;
Thurmond & Picarella 1999; Thurmond et al. 1998; Thurmond & Wyckoff 1998,
1999, 2001). To complement that work, a floristic investigation of the Dempsey
Divide was conducted in 2001. The primary objective was to document all na-
tive and naturalized vascular plants growing in the area. A secondary objective
was to gather baseline data about the plant communities present. This paper
summarizes the results of that work.
Study Area
Most of the research described here was conducted on the Thurmond Ranch, a
3,755-ha, contiguous block in southern Roger Mills County, Oklahoma (Fig. 1).
The ranch is southwest of Cheyenne, the county seat, and is situated at the south-
east edge of the High Plains physiographic province (Bruner 1931). It is on the
north side of the Dempsey Divide and extends from the crest of the divide north-
ward to the edge of the Washita River valley.
Brokenleg, Plum, and Sergeant Major Creeks, three short, north-flowing,
steep-gradient tributaries of the Washita River, drain the western, northern,
and central parts of the ranch, respectively. Killough Creek and Higgins Creek,
tributaries of Sergeant Major Creek, traverse the eastern part. These streams have
incised a dense, dendritic drainage network on the north side of the divide, cre-
ating a canyon system characterized by arroyo cut-and-fill cycles (Schumm &
Hadley 1957) operating on centennial and millennial time scales. The most re-
cent incision cycle began around the turn of the 20" Century, triggered by the
intense land use of the early post-1892 white settlement period. Maximum el-
evation on the ranch is 732 m above sea level in the southwest corner. Miminum
elevation is 601 m in the northeast corner along Sergeant Major Creek.
The climate of west-central Oklahoma is continental and typified by
marked seasonal variation (Curry 1970). Annual annual precipitation at Elk
City, roughly 25 km southeast of the ranch, is 57 cm (Oklahoma Climatological
FREEMAN ET AL., FLORA OF THE OGALLALA ECOTONE 1219
Roger bills
County
N
0 | 2 Miles nN
Th L Iker ALIa4l Nard DNaallals £ A nh Chsin£
J ,
Fic. 1
tion, light gray = Cloud Chief Formation.
Survey 2002). More than 75% of the annual precipitation occurs from April
through October, and precipitation is most abundant in May (12 cm) and least
abundant in January (2 cm). Average annual temperature at Elk City is 15.8’ C.
Average January temperature is 2.3’ C and average July temperature is 27.7’ C.
The average frost-free growing season is approximately 208 days (Curry 1970).
The ranch straddles a major geological and ecological boundary—the east-
ern outcrop edge of the Ogallala Formation, referred to here as the Ogallala
ecotone. Three major geological formations occur on the Dempsey Divide on
the ranch: the Tertiary period Ogallala Formation, and the Permian period Doxey
Shale and Cloud Chief formations (Fig. l). Thurmond et al. (2002) discuss the
1220 BRIT.ORG/SIDA 20(3)
geomorphology of each formation, but because differences among them have
profoundly affected the local topography, vegetation, and history of human use,
a brief summary follows.
The Ogallala Formation comprises late Miocene- and Pliocene-age, allu-
vial outwash from the Rocky Mountains. Its coarse sediments were deposited
across the western Great Plains by braided streams, many of which arose in the
Rocky Mountains where they were fed by episodically intense rainfall (Frye &
Leonard 1957; Gustavson & Finley 1985; Gutentag et al. 1984; Reeves & Reeves
1996; Ryder 1996; Seni 1980). Much of the lower Ogallala is extremely sandy. It
formsa gently rolling landscape atop the Dempsey Divide and was significantly
reworked by wind into dunes and deflation basins in the late Pleistocene
(Thurmond & Wyckoff 1998).
The Doxey Shale comprises discontinuous claystone and sandy siltstone
beds deposited during the Permian period on a tropical coastal floodplain. It
generally is dense, fine textured, and resists downward percolation of ground-
water. Consequently, many streams leading off the Ogallala Formation are
spring fed at the Ogallala-Doxey Shale contact. Seeps and small springs do oc-
cur in the Doxey Shale outcrop, but only in particularly wet years. The Doxey
Shale is fairly resistant to erosion, and its outcrop is typified by deep
canyonlands.
Beneath the Doxey Shale is the Permian Cloud Chief Formation (Carr &
Bergman 1976), which comprises soft, finely laminated sandstone with thin
layers of interbedded gypsum. The Cloud Chief erodes more evenly and readily
than does the Doxey Shale, yielding a more gently rolling topography. Upon
transition from the Doxey Shale to the Cloud Chief, slope gradients decrease
dramatically and stream valleys broaden.
Geologically, the Ogallala ecotone is an unstable and dynamic landscape.
Deposition of the Ogallala Formation ended in the Late Pliocene, and the east-
ern edge of the formation has been eroding episodically westward since. The
outcrop edge on the ranch is estimated to have retreated more than 1.5 km since
the end of the last glaciation, 15,000-10,000 years B.P. (Thurmond & Wyckoff
2001).
The complex geology and intensely dissected terrain on the north side of
the Dempsey Divide have yielded a patchwork of soils. Deep, calcareous, coarse-
textured mollisols dominate on the Ogallala and have a much higher effective
precipitation than do soils on the Permian outcrops. Extensive exposures of
weathering bedrock characterize the Doxey Shale, except where gully fills have
facilitated the development of isolated alfisols, inceptisols, and mollisols. These
same soil types achieve greater expression on the more gentle landscape of the
Cloud Chief outcrop, but evaporites discourage plant growth in many eroded
areas (Thurmond et al. 2002).
The geological and topographical setting of the Dempsey Divide has pro-
—
FREEMAN ET AL., FLORA OF THE OGALLALA ECOTONE 1221
vided unusual environmental conditions for the development of the region’s
biota. Diverse terrestrial, palustrine, and riverine natural communities inter-
digitate along the Ogallala ecotone, and humans have long taken advantage of
the natural resources available in this environment. A remarkable concentra-
tion of hunter-gatherer campsites is associated with the ecotone, and it has been
inferred that these sites were situated to make most efficient use of the diverse
floral and faunal communities along this boundary (Thurmond 1990, 1991b).
Human occupation of the Ogallala ecotone on the Dempsey Divide has
not been continuous over time. Prehistoric habitation was most intense during
the particularly wet periods of the 10" and 24 millennia before present
(Thurmond & Wyckoff 2001). Settlement by Euro-Americans began in earnest
in the late 1800s. Between the 1890s and the early 1900s, homesteaders exploited
the landscape for crop and livestock production. Approximately 17% (630 ha)
of the ranch was farmed, and cattle and horses intensively grazed the rest. Home-
steaders began to leave around 1915, and the Thurmond family purchased most
of the land composing the ranch between 1915 and 1950. The last of the culti-
vated land was reseeded to grass in the early 1960s, and land use has been lim-
ited to low-intensity grazing by cattle since that time. Deep drilling for natural
gas has caused localized disturbance since the late 1970s. Erosion on the ranch
has been controlled by revegetation efforts and the construction of erosion con-
trol structures (Thurmond et al. 2002).
METHODS
Four major collecting trips were made during 2001 by two of the authors (CCF
and CAM): 24-25 April, 30 May-1 June, 23-26 July, and 2-4 October. Sites with
vegetation representative of the major vegetation alliances on each of the three
major geologic formations were visited during each trip. For most species, at
least one voucher was collected on each formation on which it was encoun-
tered. Information was recorded about the abundance of species and the veg-
etation alliances in which they occurred. Opportunistic collecting by one of
the authors JPT) and by Hillary Loring of the Kansas Biological Survey netted
additional specimens. A complete set of vouchers is deposited in the R.L.
McGregor Herbarium (KANU), University of Kansas. Duplicate specimens for
many collections are deposited in the Robert Bebb Herbarium (OKL), Univer-
sity of Oklahoma.
Presence/absence data were recorded in a species x geological formation
matrix. These data were used to compare the floras of the formations using Dice’s
coefficient for 1) all species and 2) native species only. Coefficients were ob-
tained using SPSS 9.0 (SPSS Inc. 1998).
Major vegetation alliances tentatively were identified using observational
data. Dominant species, associated species, and habitat data were used to iden-
tify alliances following the hierarchical classification of Hoagland (2000).
1222 BRIT.ORG/SIDA 20(3)
RESULTS AND DISCUSSION
General Characteristics of the Flora
More than 1,400 numbers (3,100 speci including duplicates) were collected
in the study area. These represent 85 families, 286 ¢ genera, and 470 species of
vascular plants (Table 1). Four additional species were documented in the study
area (Thurmond et al. 2002), but vouchers for these were not verified at KANU
so they are excluded from this list. The five most species-rich families are
Asteraceae (81,5 non-native), Poaceae (77, 20 non-native), Fabaceae (44, 3 non-
native), Euphorbiaceae (19), and Cyperaceae (17). These five families collectively
account for 51% of the species currently known in the study area.
Non-natives compose 10% of all of the species documented. The vast ma-
jority of these are Eurasian species introduced accidentally or intentionally
since the time of Euro-American settlement. Campsis radicans, Carya
illinoinensis, Catalpa speciosa,and Maclura pomifera, although native to Okla-
homa, likely were introduced intentionally by homesteaders. Native Americans
may have introduced three other species (Acorus calamus, Echinochloa
muricata, and Juglans nigra) in the study area; Thurmond et al. (2002) discuss
these taxa in detail. Among the five largest families in the study area, intro-
duced species are proportionally highest in the Poaceae (26%). Three families
(Acoraceae, Bignoniaceae, and Tamaricaceae) and 32 genera are represented by
non-native species only. Non-native species dominate one plant community
on the ranch: the aquatic Nasturtium officinale herbaceous alliance.
Populations of four species tracked by the Oklahoma Natural Heritage
Inventory (Oklahoma Natural Heritage Inventory 2001) were discovered dur-
ing the inventory: Echinocereus reichenbachii, Eriogonum alatum, Monarda
pectinata, and Zinnia grandiflora. All grow on rocky, mixed-grass prairies on
the ranch, except Monarda pectinata, which is confined to sand prairies and
sandsage shrublands.
Epilobium leptophyllum is reported for the first time in Oklahoma. This
meio perennial grows in fens, marshes, and seeps throughout the north-
ern and central Great Plains (Great Plains Flora Association 1991) southward
to Meade County, Kansas (Great Plains Flora Association 1977). A small popu-
lation on a seepy, open slope near the Ogallala-Doxey Shale contact is 180 km
south-southeast of the nearest Kansas population (Freeman et al., in press).
The number of species documented on each of the three formations in the
study area is summarized in Figure 2. Nearly 900 occurrences (species x for-
mation combinations) were recorded. Despite the fact that all collections were
made during a single growing season, these records provide insights into the
general floristic patterns in the study area.
Thirty percent of all species (140) were documented on all three formations,
but there isa clear floristic difference on either side of the Ogallala ecotone (the
—
FREEMAN ET AL., FLORA OF THE OGALLALA ECOTONE 1223
Taste 1. Numerical summary of the vascular flora of the Ogallala ecotone on the Dempsey Divide.
Group Families Genera Species
Native Non-native Total
Pteridophytes 2 2 2 0 2
Gymnosperms ] 1 2 0 2
Angiosperms 82 283 418 48 466
Monocots 13 61 89 2| 110
Dicots 69 222 329 DF 356
Total 85 286 422 48 470
Tertiary /Permian outcrop boundary). Floristic similarity, as estimated by Dice’s
coefficient (D) using all species, was greater between the Cloud Chief and Doxey
Shale formations (D = 0.764) than between the Cloud Chief and Ogallala for-
mations (D = 0.625) or the Doxey Shale and Ogallala formations (D = 0.577). The
same calculation using native species only yielded slightly higher coefficients
but congruent results (D = 0.785 for Cloud Chief and Doxey Shale, D = 0.626 for
Cloud Chief and Ogallala, D = 0.594 for Doxey Shale and Ogallala). Twenty-five
percent of all species (119) were found only on the Ogallala. A large majority of
these are psammophytes. Mesophytes and hydrophytes account for roughly
25% of the Ogallala-only species. Non-native species are distributed roughly
proportionally to the species richness of each of the formations (Fig. 2).
Vegetation Alliances
Nineteen vegetation alliances are recognized, but accurate identification and
characterization of all alliances, as well as finer resolution of association-level
patterns, must await further studies. Given the dearth of vegetation studies for
western Oklahoma (Hoagland 2000), future work likely will be useful in refin-
ing the existing state vegetation classification.
Woodland Alliances.—A woodland is defined as an open stand of trees with
crowns not usually touching and canopy coverage of 25-60% (Hoagland 2000).
Structural diversity often is less than in a forest, and trees usually are taller than
5 m (Lauver et al. 1999). Three woodland alliances were identified on the ranch:
the Sapindus saponaria woodland alliance, the Populus deltoides woodland alli-
ance, and the Salix nigra woodland alliance. All three occur along riparian cor-
ridors, especially on the lower Doxey Shale and on the Cloud Chief formations.
Woodland alliances are best developed along the mainstems of Brokenleg, Plum,
and Sergeant Major Creeks, but they also extend up the larger tributaries and
occur near springs and seeps at the Ogallala-Doxey Shale contact.
Celtis reticulata, Juglans microcarpa, Juniperus virginiana, Robinia
pseudoacacia, Sideroxylon lanuginosum, and Ulmus americana are common
trees in these alliances. Cornus drummondii, Forestiera pubescens, Rhus
1224 BRIT.ORG/SIDA 20(3)
ZC Non-Native
@ Native
No. Species
lee)
oS
ODC O DC OC D C OD
_pormaneni=)
Fig. 2. Species occurrence by formation. 0 = Ogallala; D = Doxey Shale; € = Cloud Chief.
aromatica, Rhus glabra, and Salix exigua are typical shrub elements, with
Ampelopsis cordata, Parthenocissus quinquefolia, P. vitacea, Toxicodendron
radicans, and Vitis acerifolia the most common woody vines. Representative
herbaceous eae include Carex austrina, C. bulbostylis, C. gravida, Chenopo-
dium simplex, C. standleyanum, Elymus canadensis, E. virginicus, Galium
aparine, G circaezans, Geum canadense, Phytolacca americana, and Sanicula
canadensis. Species restricted to woodland habitats near the Ogallala-Doxey
Shale contact are Bidens bipinnatus, Botrychium virgi
Shrubland Alliances.—Shrubs or trees 0.5-5 m tall dominate a shrubland,
individuals or clumps do not touch or overlap, shrub canopy cover generally is
greater than 25%, and tree canopy cover is less than 25% (Hoagland 2000; Lauver
et al. 1999). Seven shrubland alliances were identified on the ranch. Some up-
land sites on the Doxey Shale and Cloud Chief formations support large popu-
lations of Mimosa borealis, and this likely represents an undescribed, xero-
phytic, deciduous, shrubland alliance.
The Quercus havardii shrubland alliance and the Artemisia filifolia
shrubland alliance are largely confined to deep dunes and alluvial sands above
the Ogallala Formation. Quercus havardii and Artemisia filifolia are the domi-
nant shrubs, respectively. Sporobolus cryptandrus and Schizachyrium scoparium
are codominant graminoids in both alliances. Characteristic species in both
types are Ambrosia psilostachya, Amphiachyris dracunculoides, Andropogon
hallii, Artemisia ludoviciana, Bouteloua curtipendula, B. hirsuta, Calylophus
and Rubus bushii
~—
FREEMAN ET AL., FLORA OF THE OGALLALA ECOTONE 1225
serrulatus, Cenchrus spinifex, Chamaesyce glyptosperma, Croptilon
hookerianum, Cyperus lupulinus, C. schweinitzii, Dalea candida, D.enneandra,
Eriogonum annuum, E. longifolium, Liatris punctata, Machaeranthera
pinnatifida, Mentzelia nuda, Mimosa borealis, Paronychia jamesii, Plantago
patagonica, Prionopsis ciliata, Rhus aromatica, Stenosiphon linifolius,
Thelesperma megapotamicum, Xanthisma texanum, and Yucca glauca.
Two other shrubland alliances, the Prunus angustifolia alliance and the
Rhus aromatica alliance, frequently are found on the Ogallala Formation, but
they also occur in sandy, gravelly, or silty sites on Permian substrates.
Schizachyrium scoparium occurs as a codominant in the former type and is a
regular associate in the latter. Other associates include Andropogon hallii,
Bouteloua curtipendula, Croptilon hookerianum, Eriogonum annuum, Plantago
patagonica, Prionopsis ciliata, Prunus gracilis, Rhus aromatica, Ioxicodendron
radicans, and Vitis acerifolia.
Three shrubland alliances occur along streams and shorelines of ponds and
reservoirs: the Salix exigua alliance, ee a occide nia alliance, and
Tamarix chinensis|= Tamarix 1 all three types
are Aster subulatus, Baccharis salicina, Distichlis spicata, Panicum virgatum,
Polygonum lapathifolium, Pascopyrum smithii, Pluchea odorata, and Populus
deltoides.
Herbaceous Alliances.—Herbaceous plants, usually with at least 25% cover,
dominate the herbaceous alliances. Trees and shrubs form less that 25% cover
(Hoagland 2000; Lauver et al. 1999). Herbaceous alliances, which include grass-
lands, are the predominant vegetation in the study area. Eight herbaceous alli-
ances were identified on the ranch, five of which are grassland types.
The Andropogon hallii herbaceous alliance, called sand prairie, isa tall sod
grassland alliance. The best examples occur on deep dune sands atop the
Ogallala Formation, with isolated occurrences on well-drained, sandy or grav-
elly soils on the Permian substrates. Artemisia filifolia, Prunus angustifolia,
Quercus havardii,and Rhus aromatica are typical shrubs. The sand prairie shares
many of the herbaceous associates found in the Quercus havardii shrubland
alliance and the Artemisia filifolia shrubland alliance. Additional associates
are Aristida purpurascens, Froelichia floridana, F. gracilis, Gaura villosa, Lechea
tenuifolia, Oenothera rhombipetala, Penstemon buckleyi, and Polygonum tenue.
Medium-tall sod grassland types, called mixed-grass prairies, dominate the
northern three-quarters of the ranch on Permian substrates. Three types were
identified: the Bothriochloa laguroides herbaceous alliance, Schizachyrium
scoparium-Bouteloua curtipendula herbaceous alliance, and Bouteloua
curtipendula herbaceous alliance. Additional work is needed to determine the
extent and composition of these alliances, which share many species.
Gutierrezia sarothrae and Mimosa borealis are characteristic shrubs. Herba-
ceous associates include Allium drummondii, Ambrosia psilostachya,
1226 BRIT.ORG/SIDA 20(3)
Ammoselinum popei, Amphiachyris dracunculoides, Aristida purpurea, Artemi-
sid ludoviciana, Aster ericoides, A. falcatus, A.fendleri, Bothriochloa ischaemum,
Bouteloua curtipendula, B. hirsuta, Buchloé dactyloides, Callirhoé involucrata,
Castilleja purpurea, Chaetopappa ericoides, Chamaesyce fendleri, Cirsium
ochrocentrum, C. undulatum, Croton texensis, Dalea candida, D. enneandra,
Eriogonum alatum, E. longifolium, Gaura coccinea, Grindelia squarrosa,
Lesquerella gordonii, L. ovalifolia, Liatris punctata, Linum rigidum, Lygodesmia
juncea, Machaeranthera pinnatifida, Penstemon albidus, Plantago patagonica,
Schedonnardus paniculatus, Schizachyrium scoparium, Sporobolus compositus,
S. cryptandrus, Stenosiphon linifolius, Tetraneuris scaposa, Thelesperma
megapotamicum, Vulpia octoflora, Xanthisma texanum, and Yucca glauca.
Two other grassland alliances occur along streams, shorelines of ponds and
reservoirs, and marshes. Intermittently flooded habitats support the
Pascopyrum smithii herbaceous alliance. Typical associates include Aster
subulatus, Cyperus odoratus, Echinochloa crusgalli, E. muricata, Hordeum
jubatum, Muhlenbergia a Phyla lanceolata, P. nodiflora, Pluchea
odorata ,Polypogon I nd Schoe nople ctus pungens Semipermanently
flooded areas support the i. pungens Gan sapalica as Schoenoplectus
americanus in Hoagland [2000] and many earlier floristic works) herbaceous
alliance. Associates are Ammannia coccinea, Aster subulatus, Distichlis spicata,
Cyperus odoratus, Echinochloa crusgalli, E. muricata, Eleocharis coloradoensis,
E. macrostachya, E. montevidensis, E. rostellata, Fuirena simplex, Leersia
oryzoides, Muhlenbergia asperifolia, Phalaris caroliniana, Poa arida, Juncus
torreyi, Phyla nodiflora, Pluchea odorata, Potentilla paradoxa, Ranunculus
cymbalaria, R. sceleratus, Rumex altissimus, Setaria parviflora, and Typha
domingensis.
The Lesquerella (gordonii, ovalifolia) herbaceous alliance occurs on rock
outcrops and rocky or gravelly slopes on the Doxey Shale and Cloud Chief for-
mations. Generally, perennial forbs contribute greater than 50% of the total
herbaceous cover in this alliance. Typical associates include Artemisia
campestris, Bouteloua curtipendula, B. hirsuta, Hedeoma drummondii, Hedyotis
nigricans, Heterotheca stenophylla, Krameria lanceolata, Oenothera
macrocarpa, Schizachyrium scoparium, Scutellaria resinosa, and Tetraneuris
SCaposa.
The Nasturtium officinale herbaceous alliance is found around springs and
along spring runs. It is dominated by hydromorphic, rooted vegetation that
grows in permanently flooded environments. Associates are Eleocharis
montevidensis, Leersia oryzoides, Muhlenbergia asperifolia,and Schoenoplectus
pungens. The abundance and persistence of many other species appear to de-
pend on local microhabit conditions.
Disturbed or degraded uplands on Permian formations often are similar to
Hoagland’s (2000) Amphiachyris dracunculoides herbaceous alliance. Char-
~
—
FREEMAN ET AL., FLORA OF THE OGALLALA ECOTONE 1227
acteristic species of this annual grassland alliance are Ambrosia psilostachya,
Aristida oligantha, A. purpurea, Bothriochloa ischaemum, B. laguroides,
Bouteloua curtipendula, B. hirsuta, Bromus japonicus, Chloris verticillata,
Eragrostis curvula, Schizachyrium scoparium, and Sporobolus cryptandrus.
Ruderal Sites.—Natural and semi-natural vegetation alliances typically
contain areas in which the native vegetation is disturbed asa result of localized
or widespread environmental or biological perturbations. Non-native species
often colonize these ruderal sites, and in some cases, aggressive non-natives may
spread and severely affect native plant communities. Parts of the ranch were
cultivated prior to 1915, but all such areas have since been reseeded to native
grasses and forbs. Many non-native species occupy these formerly cultivated
sites. Other non-native species are found on the ranch around old farmsteads
and newer building sites, along the shoulders of ranch roads, in corrals, and in
sites where cattle congregate.
The most frequent non-natives in ruderal sites are Aegilops cylindrica,
Bothriochloa ischaemum, B. laguroides, Bromus catharticus, B. japonicus,
Cynodon dactylon, Echinochloa crusgalli, Eragrostis curvula, Lactuca serriola,
Medicago minima, Melilotus alba, M. officinalis, Rumex crispus, Salsola tragus,
Setaria viridis, Sorghum halepense, Taraxacum laevigatum, and Tragopogon
dubius.
ANNOTATED CHECKLIST
Family names are arranged alphabetically within three major groups: pterido-
phytes, gymnosperms, and angiosperms. Genera, species, and infraspecific taxa
are arranged alphabetically within each family. Nomenclature for scientific
names and common names follows Freeman (2002). Names used in the Atlas
of the Flora of the Great Plains (Great Plains Flora Association 1977) and Flora
of the Great Plains (Great Plains Flora Association 1991) but synonymized in
Freeman (2002) are identified as “A” and “F”, respectively. Additional notes
clarify nomenclatural issues for some species. An asterisk (*) identifies non-
native species as identified by the Great Plains Flora Association (1991) and Free-
man (2002
Relative frequency of each species on the ranch is estimated from the num-
ber of populations seen by the authors using a slightly modified version of the
scale proposed by Palmer et al. (1995): abundant (5) = dominant or codominant
in one or more plant alliances; frequent (4) = easily seen or found in one or more
plant alliances (e.g., mixed-grass prairie or woodland) but not dominant in any
plant alliances; occasional (3) = widely scattered but not difficult to find in one
or more plant alliances; infrequent (2) = difficult to find with few individuals
or colonies but found in several locations in one or more plant alliances; and
rare (1) = very difficult to find and limited to one or few populations in one or
more plant alliances.
1228 BRIT.ORG/SIDA 20(3)
Alliance data for each species are summarized by major habitat types as
follows: w = woodlands (including riparian woodlands); s = sandy complex (in-
cluding sand prairies, sandsage shrublands, and shinnery oak shrublands); m =
mixed-grass prairies (including clayey, silty, sandy, gravelly, and rocky mixed-
grass prairies, rock outcrops, and rocky slopes); p = palustrine communities (in-
cluding ae and marshes); r = riverine sites (including spring runs and
streams); 1 = lacustrine communities (including ponds and reservoirs); and u =
ruderal sites (including abandoned homesites, roadsides, and well sites).
Geologic formations from which specimens were collected, or ina few cases
observed but not collected, are designated for each taxon by the following ab-
breviations: O = Tertiary Ogallala, D = Permian Doxey Shale, and C = Permian
Cloud Chief. Collection bers fc
voucher specimens are cited after any syn-
onyms. Abbreviations for collector names are F (Craig C. Freeman), L (Hillary
Loring), M (Caleb A. Morse), and T J. Peter Thurmond).
PTERIDOPHYTES
EQUISETACEAE
sala ee laevigatum A. Braun, Smooth scour-
ing-rush, 2; pt; 0)
OPHIOGLOSSACEAE
Botrychium virginianum (L. ) Sw. oe fern,
common grape fern, 1; w,p;O. MF59
GYMNOSPERMS
CUPRESSACEAE
Juniperus pin “hotii ste Pinchot juniper, lim;
DFI 6865, FMI
Juniperus virginiana i virginiana, Eastern red-
Cc 6935, FM1/689,
J ’ ' , 1 a i
FM17811, MF6386
ANGIOSPERMS
ACANTHACEAE
Ruellia humilis Nutt., Fringe-leaf wild-petunia, 2;
s;O. MF6061
AC
AGAVACEAE
Yucca glauca Nutt., Small soapweed, 4;5,m;0,D,C.
6186, MF6406, MF6421, MF6435, MF6501
ALISMATACEAE
Echinodorus berteroi (Spreng.) Fassett, Upright
burhead, 2;1;D,C. [E. rostratus (Nutt.) Engelm.
RACEAE
*Acorus calamus L., European sweetflag, 1; p,r;O.
2
ex A. Gray — A,F] FM17798,FM17881
pee Ni ineetae
maranthus albus L., bleweed amaranth, 2;
1u;O,D,C. FM 1 7 a 7930, FM18404
Amaranthus arenicola |.M. Johnst., Sandhill pig-
_1;5;0.FM17698
Amaranthus blitoides S. Watson, Prostrate pig-
,4,u; O,D,C. TA. graecizans L. — A, F]
ern FM17905, FM18339, MF6014,
70, MF6210
ial palmeri S.Watson,Palmer’s pigweed,
u;O,D. ae 51, MF6462
Amaranthr
exus L., Rough pigweed, Red-
root dai -u:O,D,C.FM17929, MF6392,
Froelichia floridana (Nutt.) Moq,, Field snake-cot-
on, 2;5;0.FM17714
Froelichia gracilis (Hook.) Moq., Slender snake-
cotton, 2;5,m;O0.FM17657, MF6487
ANACARDIACEAE
Rhus aromatica Aiton var. ees (Engelm.)
Shinners, Fragrant sumac, 2;w M17521
Rhus aromatica Aiton var. scnoung or
Rehder, Fragrant c, 3: w,m; O,D,C.
F16875, FM1 7662, ieee MF6277
Rhus aromatica Aiton var. trilobata (Nutt.) A.Gray,
Fragrant sumac, 1;m;D.MF6137
Rhus nate L., smooth sumac, 3; w,s,m; O,D,C.
FM17590, L2001-23, MF5945, MF6136,
F6
Toxicodendron radicans (L.) Kuntze subsp.
FREEMAN ET AL., FLORA OF THE OGALLALA ECOTONE
negundo (Greene) Gillis, Poison-ivy, 3; w,s,m;
FI 6899, FM17769, MF6397
Toicodendton rydbergii (Small ex Rydb.) Greene,
ora ivy, Western poison-ivy, 1;
Onn
APIACEAE
Ammoselinum popei Torr. & A. Gray, Plains sand-
parsley, 4; m; D,C. F16863, F16901, F16921,
MF6258
Cymopterus macrorhizus Buckley, Big-root
spring-parsley, 2; m; D,C. F16850, F16930,
F16945
Lomatium foeniculaceum (Nutt.) J.M. Coult. &
Rose var. daucifolium (Nutt.) Cronquist, Fen-
nel-leaf desert-parsley, 2; m; D.C. F16849,
F16951
Sanicula canadensis L. var. canadensis, Canadian
anicle, 3;w;D,C.FM17869, MF5934, MF6366
Spermolepis inermis (Nutt.ex DC.) Mathias & Con-
e, Spreading scaleseed, Western
scaleseed, 2:5;0,D. MF6004, MF6124, MF6473
*Torilis arvensis (Huds.) Link, Field hedge-parsley,
6 MF6364
, my
APOCY
arog Soa inumL., Hemp dogbane, Prai-
ed e,3;m,p;0,D,C.FM17735,MF5955,
MFé1 ee
ASCLEPIADACEAE
Asclepias asperula (Decne.) Woodson subsp.
ricornu (Woodson) Woodson, Antelope-
es milkweed, 3; s,m; O,D,C. [A. asperula
(Decne.) Woodson var. decumbens (Nutt.)
Shinners — A, F] MF6050, sacais: MF6222
Asclepias engelmanniana Woodson, Engel-
nn’s milkweed, 3; s,m; O,D,C. FM17541,
FM17666, FM17779, FM18396, FM18483,
FM18484, MF6140, MF6412
Asclepias latifolia (Torr.) Raf, Broad- a milkweed,
M17906, FM1832
Asclepes ees ‘ ee eae leaf milk-
reas Mars s eee interior Woodson
erfly milkweed, 2; 5; O. MF6008, MF6441
Asclepias viridiflora Raf., Green milkweed, 3; s,m;
O,D,C. FM17593, FM17702, FM17907,
MF6002, MF6144, MF6285, MF6449
Asclepias viridis Walter, Spider milkweed, 1
FM17672
1229
ASTERACEAE
Ambrosia psilostachya DC., Western ragweed, 4
.FM17604, FM17775, FM17873,
FM18264, FM18299, FM18393
Ambrosia trifida L., a ragweed,
M18325,FM18
Amphi fees 5 roanclit
eed, 5
t ' ped
1; wu; O,C.
—
a Nutt., Corm-
,D,C. [Gutierrezia
sracuncuta ides (D a ce Blake — A, F]
FM1 , FM17934, FM18263, FM18309,
aa
je oe skirrhobasis (DC.) Trel. var.
skirrhobasis, Parasol lazy-daisy, 2;s;O. MF6026,
9
MF643
Artemisia campestris L. subsp. caudata (Michx.)
H.M. Hall & Clem., Western sagewort, 2; s,m;
O.FM18307
Artemisia dracunculus L., Silky wormwort, Tarra-
gon, 2; m;D.FM18395, FM18420, L2001-118
Artemisia filifolia Torr., Sand sagewort, 5; s,m;
O,D,C. FM17626, FM17836, FM18340,
FM18475
ia ludovici Nutt. var./udoviciana, Loui-
siana sagewort, 4; s,m; O,D,C. FM 17522,
FM17899, FM17926, FM18265, FM18394,
FM18459
Aster ericoides L. subsp. ericoides var. ericoides,
Heath aster, White aster, 3; s,m; O,D,C.
FM18266, FM18268, FM18305, FM18408
Aster falcatus Lindl. subsp. commutatus (Torr. &
A. Gray) A.G. Jones, Western heath aster, 3;
m,p; O,D,C. FM18362, FM18416, FM18457
Aster fendleri A. Gray, Fendler’s aster, 3; m; D,C.
FM18267, FM18407, FM18417,L2001-16
Aster oblongifolius Nutt., Aromatic aster, 2; s,m;
O,D.FM18306b, FM18363, FM18418
Aster patens Aiton var. patentissimus (Lindl. ex
DC.) Torr. & A. Gray, Sky-drop aster, 2; 5; O.
FM17576, FM18306a
Aster subulatus Michx. var. ligulatus Shinners,
Saltmarsh aster, 4; p,rl; O,D,C. FM17767,
M18361, FM18428, FM18449
Baccharis salicina Torr.& A.Gray, Willow baccharis,
.D,C.FM17639, FM17721, aa.
1;
Berlandiera betonicifolia (Hook.) Small, Betony-
leaf berlandiera, 2; 5; O. [B. texana DC. - A, F]
MF6077, MF6431
Bidens bipinnatus L., Spanish needles, 1; w; O.
679
1230
Bidens comosus (A. Gray) ie Leafy-bract
eqgar-ticks, 1;1;C. FM
Bidens frondosus L., Devil's as 2; wl;C,
M18329, FM18435
Brickellia eupatorioides (L.) Shinners. var.
corymbulosa (Torr. & A. Gray) Shinners, East-
ern brickellbush, 2; s,m; O,C. [Kuhnia
eee L. var. corymbulosa Torr. & A.
— A,F] FM18303
ae ericoides (Torr.) G. Nesom, Heath
least daisy, 3; m; D,C. [Leucelene ericoides
(Torr.) Greene — A, F] F16826, F16866,
MF6148, MF6242
anne ochrocentrum A. Gray, Yellow-spine
e, 3;m;D,C.FM17851, MF6338, MF6360
see undulatum (Nutt.) Spreng., Wavy-leaf
thistle, 3; s,m; O,D,C. FM17562, FM1766/7,
Conyza canadensis (L.) Cronquist, Tall horseweed,
Canadian aia 3;5,U;0,D,C.FM17581,
FM17939, FM18424
Croptilon ane (Torr.& A.Gray) House var.
validum (Rydb.) E.B. Sm., Slende
goldenweed, 4; s,m; O. [Haplopappus validus
(Rydb.) Cory — A, F] FM17599, FM17713,
FM17920,FM18355
Cyclachaena xanthifolia (Nutt.) Fresen., Rag
sumpweed, eu! ee marshelder, 1; w,u; O.
=
ieee eee
CPE HPO INUILL
~A,F] F
bys a (Vent.) Hitchc., Prairie fetid-
ma ;m,u; D.C. FM17938, FM18385
Pe see ifolia DC. var. angustifolia, Nar-
row-leaf coneflower, 2; m,5; O,C. FM17528,
FM17611,MF6065
Eclipta prostrata (L.) L., Yerba de tajo, 2; 11; O,D,C.
FM17588, FM17754, FM18439
Engelmannia_ peristenia (Nutt.
Goodman & CA Lawson, Engelmann daisy,
-. [E. pinnatifida Nutt. — A, FJ
M1764, MF6149, MF6272
Frigeron modestus A. Gray, Plains fleabane, 2; m;
DC. F16875, F16932, FM18399, MF6131,
MF6316
Eger stigoss i ex Willd. eee hale
3;m,5;0,D, F6068, MF6130,M
lpaorom psa Clasping- he oe-pye-
eed et, 2; p,r;O.FM17719, FM18328
Bee See eel ides aoe e, Sticky
euthamia, 1; 1; O. [Solidago gramini ee. )
Salisb. var. pean (Greene) Croat
BRIT.ORG/SIDA 20(3)
- A; 5. graminifolia (L.) soe var. media
(Greene) S.K. Harris — A] FM1834
Evax prolifera Nutt. ex DC, Big- a rabbit-to-
sacco, 2; m,s; O,D,C. F16898, FM17755,
MF6238, MF6354, MF6470
Gaillardia pulchella Foug. var. pulchella, Indian
blanket-flower, Rose-ring blanket-flower, 3;
m; 0,D,C. MF6027, MF6094, MF6212
in suavis (A. Gray & Engelm.) Britton &
Rusby, Rayless blanket-flower, 2; m; D,C.
MF6127, MF6232
Grindelia squarrosa (Pursh) Dunal var.nuda (A.W.
Ww A. Gray, Curly-top gumweed, 3;5,m,u;
O,D,C.FM17921,FM18271, FM18312
Gutierrezia sarothrae (Pursh) Britton & Rusby,
room snakeweed, 4; s,m; 0,D,C. FM17894
18274, FM18301, FM18405, L2001-20
Helianthus annuus L., Common sunflower, 2; u;
M17693, FM17772, FM18380,
FM18482,L2001-101
Helianthus maximilianii Schrad., Maximilian’s
sunflower, 1;w;C.Ts.n.
Helianthus petiolaris Nutt.subsp. petiolaris, Plains
sunflower, 2; s,m; O0,C. FM17586, FM18308,
L2001-79, MF6028
Helianthus tuberosus L., Jerusalem-artichoke sun-
—
—
'
) 1
Heterotheca latifolia Buckley var. el ae SB.
Wagenkn., Broad-leaf golden-aster, 3; s
O,D,C.FM1 7606, FM17918,FM18304, bt.
76
Heterotheca stenophylla (A. Gray) Shinners var.
angustifolia (Rydb.) Semple, Narrow-leaf
golden-aster, 3; m; O,D,C. [Chrysopsis villosa
(Pursh) Nutt. ex DC. var. angustifolia (Rydb.)
Cronquist — A, F] FM17530, FM17807,
FM18310
Heterotheca stenophylla (A. Gray) Shinners var.
stenophylla, Narrow-leaf golden-aster, 2;5,m;
O,D. Chysopsisstenophyl ee sets Shinners
—A,F]F 838, FM
iymenopanps ae es a hill
olly-white, 3; s,m; 0,D,C. FM17629,
ve 7898, la MF6122, MF6206,
MF6419, MF64
Lactuca facies ne tt.) Soreesa lettuce,
Louisiana lettuce, 1; w;C. MF6378
*/actuca serriola L., Prickly ete 2, u; O,D.
FM17632,FM17791,FM17
Liatris punctata Hook.,
—
as dotted
FREEMAN ET AL., FLORA OF THE OGALLALA ECOTONE
gayfeather, 4;s,m;0,D,C.FM17893,FM18275,
FM18311,FM18411, L2001-14
a squarrosa (L.) Michx. var. glabrata (Rydb.)
Gates, ee eee 2;5;0.[L. glabrata
ne A] F
Lindheimera texana tee & Engelm., Texas-star,
1;m;C. MF6314
eevee eles (Pursh) D.Don ex Hook., Rush
eleto eed, 3;m; C.FM18270, FM18481,
eee
Machaeranthera pinnatifida (Hook.) Shinners var.
glaberrima (Rydb.) B.L. Turner & R.L. Hartm,,
Cut-leaf tansy-aster, 4;5,m;O,D,C. [Ha eae
spinulosus (Pursh) DC. subsp. glaberrimus
are H.M. Hall — F] FM17834, MF6071,
MF6090, MF
Machaeranthera pinnatifida (Hook, Shinners var.
pinnatifida, Cut-leaf tansy-aster,
[Haplopappus spinulosus (Pursh) DC. ee
australis (Greene) H.M. Hall — F; H. ‘ers
(Pursh) DC. subsp. cotu/a (Small) H.M. Hall —
F; H. spinulosus (Pursh) DC. subsp aimitestis
— A, F] MF6085, MF6207
Melampodium leu ee & A. Gray, Plains
blackfoot, 3; m; D,C. F16949, i 18270,
FM18392, ee MF6260, MF640
*Onopordum acanthium L., Scotch a SW,U;
F6330
Packera plattensis (Nutt.) W.A. Weber & A. Love,
Prairie pe 3;m;0,D,C. [Senecio plattensis
Nutt.— A, F] F16845
recs americanus (Nutt.) D.Don, Ameri-
napweed, Basket-flower, 3;m;D,C.[Cen-
taurea americana Nutt — A, F] FM17553,
FM17849, ee MF6146, MF6291
Pluchea odorata (L.) Cass., Purple marsh-fleabane,
rl; O,D,C. [2 purpurascens (Sw.) DC. — A]
FM17638, FM17876, FM18365, FM18448,
L2001-29
Prionopsis ciliata (Nutt.) Nutt., Wax-goldenweed,
[Haplopappus ciliatus (Nutt.) DC.- A,
F]FM17718, FM18272, FM18298
Pseudognaphalium obtusifolium (L.) Hilliard & B.L.
Burtt, Fragrant false-cudweed, 1; u; O
(Gnaphalium obtusifolium L.— A,F]FM18343
Psilostrophe tagetina (Nutt.) Greene var. cerifera
i Nelson) B.L. Turner, eee a flower,
im:C. [P villosa Rydb. - A, F] M
Pe eis grandiflorus (N ie a een
1231
false-dandelion, 2; s,m; O,D,C. F16848,
MF6037, MF6107, MF6195
Ratibida columnifera (Nutt.) Wooton & Standl.,
Upright prairie-coneflower, 2; s,m; O,D,C
MF6036, MF6125, MF6263
Senecio riddellii Torr.& A. Gray, Riddell’s grounsel,
3;5,m; O,D.FM18302,FM18415
Solidago altiplanities C.E.S. Taylor & R.J. Taylor,
High-plains goldenrod, 3;m;0,D,C.FM17557,
FM18273, FM18300, FM18403,FM18419
Solidago canadensis L.var.scabra (Muhl.ex Willd.)
orr. & A. Gray, Canadian reap 2; W,D,F;
O,C.FM17734,FM18327,FM18446
olidago gigantea Aiton, Late goldenrod, No-
vember goldenrod, 1; w,p,r; O.FM18346
Prairie goldenrod, 2; s,m; O,C. FM17709,
FM17877,FM18462, Ts.n
Solidago petiolaris Aiton var. pen iolaris, Downy
goldenrod, 1;5,m;0,D.FM18372, FM18402
*Sonchus asper (L.) Hill, Prickly sow-thistle, 2; u;
O,C. MF5 F6297
*Taraxacum laevigatum (Willd.) DC., Red-seed
dandeli O,C.F16913, MF6283
Tetraneuris scaposa (DC.) Greene var. scaposa,
Naked-stem tetraneuris, 4; m;
[Hymenoxys scaposa (DC.) K.L. Parker var.
glabra (Nutt.) KL. Parker — A, F; H. scaposa
(DC.) K.L. Parker var. scaposa — A, F] F16825,
F16876, F16952, MF6231, MF6418, MF6490
Thelesperma filifolium (Hook.) A. Gray var.
intermedium (Rydb.) Shinners, Thread-leaf
greenthread, 1;m;D.MF6120
Thelesperma megapotamicum (Sprenq.) Kuntze,
Rio Grande greenthread, 4; s,m; O,D,C.
ene 7835, MOOT MF6139, MF6233, MF6417
Western salsify, 2;m,u;
C. MF6080, MF6267
Verbesina encelioides (Cav.) Benth. & Hook. f.ex A.
sp.encelioides, Golden crownbeard,
elion, 2:mM,u:
eis
Verbesina encelioides (Cav) eae & Hook.f.ex A.
Gray subsp | Rob.& Greenm,)
J.R. Coleman, Golden Lean 2 ee:
M18386
Vernonia baldwinii Torr. subsp. interior (Small) W.Z.
Faust, Western ironweed, 3; s,m; O,D,C.
FM] 7547 FM17585, FM17762, FM17888,
FM1842
1232
Xanthisma texanum DC. subsp.drummondii (Torr.
& A.Gray) Semple, Sleepy daisy, 4;5,m;O,D,C.
[X. texanum DC. var. drummondii (Torr. & A.
Gray) A. Gray — A] FM17615, FM17917
FM18313, MF6044, MF6350, MF6451
Xanthium strumarium L., Common cocklebur, 3;
\C.FM17803, FM18276, FM18377
Zinnia grandiflora Nutt., Rocky Mountain zinnia,
2;m;D,C. MF6095, MF6336, MF6408
BIGNONICACEAE
*Campsis radicans (L.) Seem., Common trumpet-
creeper, Trumpetvine, 1; w,u;O.FM18335
*Catalpa speciosa Warder, Northern catalpa,
Catawba-tree, 2; w,u; 0. FM18336, MF5986
ee
L ii (Hornem.) Greene, Flat-spine
atl kseed, 3; s,m; O,D,C. F16853, F16922,
MF5940, MF6293
ne texana (Scheele) Britton, Cup-seed stick-
,2;5,m; D,C.F16927,F 1694
Pees caroliniense (Walter ex J.-F. Gmel.
MacMilL, rare gromwell, Carolina puc-
;5;O. MF6069
a
coon,
sae olbee incisum Lehrn., Plains gromwell,
; 0,D,C. F16878, F16944, MF6003,
vee 54, MF6254, MF6o469
BRASSICACEAE
*Camelina microcarpa Andrz. ex DC., Little-pod
false-flax, 1; u;C. MF6294, MF6494
*Capsella bursa-pastoris (L.) Medik., Common
shepherd's-purse, 2; u;O,D,C. F16838
Descurainia pinnata (Walter) Britton subsp.
halictorum (Cockerell) Detling, Pinnate
tansy-m i ee F16880
Dimorphocarpa candicans (Raf.) Rollins, Palmer's
spectacle-pod, 1; 5; O. [D. palmeri (Payson
Rollins — F; Dithyrea wislizenii Engelm. v
meri Payson — A] FM17654, MF6424
aa reptans (Lam.) Fernald, White whitlow-
D,C.F16832
Erysimum asperum (Nutt.) DC., Plains wallflower,
1;m;C. F16948
sae densiflorum ra Prairie pepper-
u; O,D. F
moe oblongun aT ee pepper-grass,
.F 16836, MF6257
iasturtioe ofcinaleR Br,Common watercress,
1; 1;O.F 16892, MF5970
Physaria gordonii (A. Gray) O'Kane & Al-Shehbaz,
S
ay See
=)
BRIT.ORG/SIDA 20(3)
Gordon's bladderpod, 4; m; D,C. [Lesquerella
ene (A. Gray) S. Watson — A, F] F16856
MF6
—s
Dh lifnl
Phy ifolia (Rydb.) O'Kane & Al-Shehbaz
Sib 5: sudifola Oval-leaf bladderpod, 4; m;
O,D,C. [Lesquerella ovalifolia Rydb. subsp.
ovalifolia — A, F] F16847, F16950, MF6404,
MF6488
CA EAE
C sist tha vivipara aa Britton & Rose var.
vipara, Pin- Ste oryphantha, 2; s,m;
Ce: eas . MF6331,MF6361
indropuntia imbricata (Haw.) F.M. Knuth var.
imbricata, Tree cholla, 1; m; C. [Opuntia
imbricata (Haw.) DC. var. imbricata — A, F]
ie
cy
FM18485
Echinocereus reichenbachii (Terscheck ex Walp.)
F. Haage var. reichenbachii, Lace hedgehog
cactus, 2;m;D,C.FM17559, MF6192, MF6413
Im.var.macrorhiza, Big-
root pricklypear, 3; s,m; O,D,C. FM17560,
MF6056, MF6332, MF6362, MF6436
Opuntia phaeacantha Engelm. var. major
Engelm., New Mexico pricklypear, 1; m; D.
FM17852
CAMPANULACEAE
Lobelia cardinalis L., Cardinal-flower, 1; p; O.
2001-61
Triodanis holzingeri McVaugh, Holzinger’s Venus’
looking-glass, 2; 5; O. MF6046, MF644
Triodanis mies arpa (Nutt.) Nieuwl. aes fruit
s'-looking-glass, 3; m; D,C. MF6119,
ce:
CAPPARACEAE
Polanisia dodecandra (L.) DC. subsp.
trachysperma (Torr. & A. Gray) H.H. IlItis,
Rough-seed clammyweed, 2; s,m; D,C.
MF6407
anisia jamesii (Torr. & A. Gray) H.H. IItis, James’
clammyweead, Cristatella, 2:5,m;O. [Cristatella
jamesii Torr.& A.Gray — A] FM17653,FM18282,
MF6073, MF6189
CARYOPHYLLACEAE
Loeflingia squarrosa Nutt. s
Barneb & Twis selm., Spreading loeflingia, 1;
Po
mets
>
ubsp. texana nee
Paronychi ia jamesii Torr.& A. Gray, James'nailwort,
4; 5,m; 0,D,C. FM17525, FM17597, MF6030,
MF6I 10, MF6268, MF 6478
FREEMAN ET AL., FLORA OF THE OGALLALA ECOTONE
Silene antirrhina L., Sleepy catchfly, 3; s,m; O,D,C.
F16829, MF6033, MF6161, MF6305, MF6447
*Stellaria pallida (Dumort.) Crép, Pale chickweed,
.F16911
CELASTRACEAE
Celastrus scandens L., American bittersweet, 1;w;
C.MF6379
CHENOPODIACEAE
Chenopodium berlandieri Mog. var. zschackii
(Murray) Murray ex Asch., Pit-seed goosefoot,
m,u; O,D. FM17904, FM17931, FM18341,
FM18382
eae a ial L., Oak-leaf goosefoot,
1;1;C.FM
Chenopodium missouriense Aellen, Missouri
goosefoot, 1;u;C.FM18466
Chenopodium setae aRydb.,Field goosefoot,
3; 5,m; O,D. F 578, FM17658, FM17776
MF6015, eos
Chenopodium si PIX (Torr) Raf., sed oa
goosefoot, 2;w
—F;C. hybridum L.- A] FM17728, FM] 19332
Chenopodium standleyanum Aellen, Standley’s
goosefoot, 3; w; O,D,C. FM17682, FM17727,
FM17817,FM18333, FM18412,FM18474
Cycloloma atriplicifolium (Spreng.) J.M. Coult.,
umble ringwing, 2;5;0.FM17616,FM18350,
MF6013
*Kochia scoparia
C.FM18465
Monolepis nuttalliana (Schult.
a (L.) Schrad., Broom kochia, 1; u;
LS
Greene, Nuttall’s
poverty-weed, 3; m,u; D,C. F16908, F16938,
MF6
*Salsola ae L., Common Russian-thistle, 3; u;
OD. rica Sennen & Pau — A, F]
FM17842, ee FM18373
ACEAE
Lechea tenuifolia Michx., Narrow- as 2;
s;O.FM17592,FM18315, MF6
COMMELINACEAE
ommelina erecta L. var. angustifolia Hest
Fernald, Erect dayflower, 3; 5; O. MF5990,
MF6024, MF6482
— occidentalis (Britton) Smyth & L.
alis, Western spiderwort,
nh cerns 3; s,m; O,D,C. MF6029
MF6176, MF6230, MF6444
LY
occident
1233
CONVOLVULACEAE
Convolvulus equitans Benth., Gray bindweed, 1;
m;C. MF6199
talli S hult., Nuttall’ |
; D.C. FM17529, MF6045, MF6108,
ee MF6228
Ipomoea leptophylla Torr., Bush morning-glory, 1;
s,m; D.MF6414
CORNACEAE
Cornus drummondii C.A. Mey., Rough-leaf dog-
wood, 4; w,m; O,D,C. FM17519, FM17661,
eet Meese
CUCURBITACEAE
ucurbita foetidissima Kunth, Buffalo gourd, 3:
s,m; O,D,C. MF6327, MF6420, MF6461
CUSCUTACEA
Cuscuta cuspidata Engelm., Cusp dodder, 2; s,m;
O,D,C.FM17656,FM18319, 2001-36, L2001-
ie4
CYPERACEAE
Carex austrina _ Southern sedge, 2;w,s;0,C.
MF6082, MF639
_— bul geen an Thick-style sedge, 2; w;
. [C. amphibola Steud. var. globosa (LH.
ie rea Taylor and Taylor (1994)]
MF5997, MF6370
Carex gravida LH. Bailey, Heavy sedge, 3; wl;
O,D,C. F16870, F16887, F16914, MF5943,
MF6224, MF6300, MF6358, MF6359, MF6375
ae lita Muhl.ex Willd., Woolly sedge, 1;p,1,l;
CIE lanuginosa Michx. — A, F] FM18463
Cyp Torr. & Hook., Tape-leaf flat-
sedge, 1;1;O.FM17646
Cyperus lupulinus (Spreng.) Marcks subsp.
lupulinus, Slender-stem flat-sedge, 2; s,m; O.
FM18364, MF5998
Cyperus pss Geise, Intermediate flat-
m;O.FM17647,MF5599
Cyperus as L., ae ae eee ms
O,D,C. F FM]
FM17933, F18379, FM18
Cyperus schweinitz ifTore a 's flat-sedge,
m; 0. 443
Eleocharis cloradoensi (Britton) Gilly, Colorado
spike-rush, 4; |: FM17739, FM17843,
rat Veg
Eleocharis erythropoda Steud., Bald spike-rush, 2;
|; O,C.FM17579, FM17859
,
1234
Eleocharis geniculata (L.) Roem. & Schult. Clus-
ter spike-rush, 1 a caribaea (Rottb,) S.F.
Blake — A,F] FM1/
Eleocharis eae Britton, Large-spike
spike-rush, 4; |; O,D,C. [E. xyridiformis Fernald
& Brackett — A,F] FM17650, MF6153, MF6357
Eleocharis montevidensis Kunth, Sand spike-rush,
4; p,r|; O,D,C. FM17717, FM17740, FM17858,
MF5971, MF6152, MF6373
Eleocharis palustris (L.) Roem. & Schult., Marsh
spike-rush, 1; 1; D. [E. smallii Britton — A, F]
Eleocharis rostellata (Torr.) Torr, Beaked spike-
rush, 2; p,l;0,C.FM17732,FM17861
Fuirena simplex Vahl var. aristulata (Torr.) Kral,
Western umbrella-grass, 2; p,rl; O,D,C.
FM17678,FM17759, FM17856
Schoenoplectus pungens (Vahl) Palla var.
longispicatus (Britton) S.G. Sm., Common
hreesquare twine-bulrush, 4; p,rl; O,D,C.
[Scirpus americanus Pers. — misapplied in A
and in Taylor and Taylor (1994); S. pungens
Vahl — F] FM17716,FM17863, MF5987
EBENACEAE
Diospyros virginiana L, Virginia persimmon, 2;W,p;
O,C.FM17725, FM17890
ELATINACEAE
Bergia texana (Hook.) Seub.ex Walp., Texas bergia
2:1. D,C. FM17800, FM1 7822
EUPHORBIACEA
Acalypha saa Riddell, Rough-pod
copperle a ea
Pees Torr.& Gray) Small, Fendler’s
mat-spurge, 4; m; O,D,C. Euphorey a ioe eri
an & A. Gray — A, F] F1693
ee -
Chamaesyce geyeri (Engelm.) Small, Geyer’s mat-
spurge, ene [Euphorbia geyeri Engelm.— A,
F]FM18
pa ee pean pce ) Small,
Ridge-seed mat-spur 75m; O,D,C. [Eu
phorbia al micsneniat: als elm. — A,
FM17613, FM17902, FM17941a, FM18280,
MF5979, MF6177, MF6299
Chamaesyce maculata (L.) Small, Spotted mat-
spurge, 3;1,u;0,D. ee oe L-A
FI FM17621,FM17747, MF599
Chamaesyce missurica (Raf.) Shinners, Missouri
Pala:
(one
BRIT.ORG/SIDA 20(3)
mat-spurge, 2;5,m;D,C. oa missurica
Raf.— A, F] FM17825, MF6o4
ier nos nutans eee ok Eyebane, Nod-
ding -spurge, 1; u; O. [Euphorbia nutans
Lag. - zx FM18357
hamaesyce prostrata (Aiton) Small, Prostrate
mat-spurge, 1;u;C. [Euphorbia prostrata Aiton
—~A,F] FM17941b, MF6491
Chamaesyce serpens (Kunth) Small, Round-leaf
mat-spurge, 1;1; C. [Euphorbia serpens Kunth
A, F] FM17823
Croton glandulosus L. var. lindheimeri MUll. Arg.,
i. oo 2;5,m; 0. FM17589, FM18283,
C
fe a neiean Michx., One-seed croton,
: (C.FM17774, FM17827, FM17896,
MF61
Croton texens a Mill. Arg., Texas croton,
M17538, FM17778, MF6022,
MF6168, fae
sauna ere Subils, Western toothed
mu; O,D,C.FM17780, FM17912
ete om icuris Scheele, Wedge-leaf spurge,
2,m;D,C.F 16859, MF6106B, MF6296
Euphorbia marginata Pursh, Snow-on-the-
mountain, 3; m; D,C. FM17546, FM17847,
279,FM18477
Euphorbia spathulata Lam.,Warty spurge, 2;5,™M;
D,C. MF6106A, MF6236
Stillingia sylvatica soe yes mre al
light stillingia, 2;s;O.FM17 6076
Tragia betonicifolia Nutt., se San 2:5;
M1 7642 ,FM17651, MF6047
ea ramosa Torr., Catnip noseburn, 3; m; D,C.
F16855,FM18410, vey MF6229, MF6405,
MF6489
FABACEAE
Acacia angustissima (Mill) Kuntze var. hirta (Nutt)
B.L.Rob., Prairie acacia, 3;s,m;O,D,C.FM17703,
FM17923, MF6290, MF6348, MF6471
Amorpha canescens Pursh, Leadplant, 1; s,m; O.
FM17610
os
nS
Amorpha fruticosa L., Bush wild-indigo, 2;m,1;D,C.
FM17840, FM17865, MF6317
=.
Astragalus crassicarpus Nutt. var. crassicarpus,
Ground-plum milk-vetch, 2; m;O,D.F16864
MF6081
So ilis Nutt, steteaedlags vetch, 2; mM;
DC. F 1, MF6092, MF62
FREEMAN ET AL., FLORA OF THE OGALLALA ECOTONE
Astragalus lotiflorus Hook., Lotus milk-vetch, 2;m;
D,C.F16827, MF6223, MF6343
Astragalus missouriensis Nutt. var. missouriensis,
Missouri milk-vetch, 2;m;D.F 16857, MF6250
Astragalus mollissimus Torr. var. mollissimus,
ie, ie: Raeccialo
MF6
Astragalus nuttallianus DC. var. austrinus (Small)
Barney, Small-flower milk-vetch, 1; m; D,C.
F16828, F16937
Astragalus plattensis Nutt.,Platte River milk-vetch,
2;5,m; O.F16886 6055
Baptisia australis (L.) R. Br. var. minor (Lehm.)
nald, Blue wild-indigo, 3; s,m; O,D,C.
F16868, F16954, FM1 7624, MF6191, MF6274
Baptisia bracteata Muhl.ex Elliott var. leucophaea
(Nutt.) Kartesz & Gandhi, Plains wild-indigo
m; O. [B. bracteata Muhl. ex Elliott var.
glabrescens (Larisey) Isley — F; B. leucophaea
Nutt. — A] F16719, FM17625, MF6075
Chamaecrista fasciculata (Michx. ) Spake oy
partridgepea
L.-F; Cossiafasccuat iictix = A) FMI 7587,
FM17910
Dalea aurea ee ex cine Golden prairie-clo-
ver, 3;m;0,D,C.FM17536,FM17752,FM17805
Dalea candida Michx. var. oligophylla (ett
Shinners,Western prairie-clover, 4;5,m
FM17567, FM17708, FM17832,M
Dalea enneandra Nutt., Nine-anther prairie-clo-
ver, 4; s,m; O,D,C. FM17535, FM17601,
FM17712, FM17751, FM17806, MF6287,
353
Dalea purpurea Vent. var. arenicola (Wemple)
arneby, Purple prairie-clover, 1;s;O. MF6007
MF6445
Dalea purpurea Vent. var. purpurea, Purple prai-
rie-clover, 3; s,m; O,D,C. FM17537, FM1 7668,
MF6116, MF6288, MF6398, MF6498
Dalea villosa (N
rie-clover, 1;5;O.FM17634
Desmanthuls illinoensis (Michx.) MacMill. ex B.L.
Rob. & Fernald, Illinois bundle-flower, ’ s,m;
O,D,C. FM17524, FM17665, FM17768
) Spreng. var. villosa, Silky prai-
Desmodium illinoense A. Gray, Illinois tick-clover,
2; w,S;O.FM17695, MF6051, MF6448
Desmodium sessilifolium (Torr.) Torr. & A. Gray,
Sessile-leaf tick-clover, 2; s; O. FM17600,
FM17925
1235
Gleditsia triacanthos L., Common honey-locust,
1;w 0)
Glycyrrhiza lepidota Pursh, American licorice, 3;
m,r,|; O.FM17608, MF5975
Gymnocladus dioica (L.) K. Koch, Kentucky
coffeetree, 2; w,m; C. F16916, FM17819,
MF6384
T;m;C.FM1754
Indigofera miniata Ort., Western indigo, 3;s;0. [L.
miniata Ort. var. /eptosepala (Nutt.) B.L. Turner
—A,F]FM17595, MF6010, MF6453
Lespedeza capitata Michx., Round-head bush-
clover, 3;5;0.FM17711
Lespedeza stuevei Nutt, Tall bush-clover, Stueve’s
bush-clover 4; 5;O.FM17584,FM18318
*Medicago minima (L.) L., Prickly sees : m,U;
D,C. F16841, F16910, MF6220, MF63
*Melilotus a Medik., White sweet- eee 2;U;
8348, MF6308
*Meli lotus pe alis (L : Pall, Yellow sweet-clo-
F6205, MF6390, MF6427
Mimosa borealis A. Gray, Fragrant mimosa, 4; s,m;
O 704, L200-52, MF6086, MF6198
Mimosa quadrivalvis L.var.nuttallii (DC.) L.S. Beard
ex Barneby, Cat-claw mimosa, 3; s,m; O,D,C.
[Schrankia nuttallii (DC.) Standl. — A, F
FM17607, MF6128, MF6197, MF6454
Oxytropis lambertii nee var. lambertii, —— S
C.F16862,FM1752
Pediomelum cusp! ae (Pursh) Rydb,, Tall- es
scurfpea, 3;m;D,C. [Psoralea cuspidata Pursh
— A, F] MF6104, MF6194
Pediomelum digitatum (Nutt.) Isely, Palm-leaf
scurfpea, 2; 5; O. oe oe itata Nutt. var.
digitata — A, F] FM1757 F6054
Pediomelum linearifolium a & A. Gray) J.W.
Gri Slim-leaf scurfpea, 3; s,m; wae
(iat Mel Died Redd
—
locoweed, 2;m;
nee ae Torr. & A. Gra A, F]
FM17558, 7833, MF6038, ie
MF6273
Prosopi “hile a Torr. var. glandulosa, Honey
mesquite, 1;m;O0,D.FM17561,L2001-112
seeds ea iflorum (Pursh) Rydb., Narrow-
leaf scurfpea, 3;5;O. [Psoralea tenuiflora Pursh
var. floribunda (Nutt.) Rydb.- A,F;P tenuiflora
ios var. tenuiflora — A,F] FM17924, MF6023,
A474
—
DAL
dia L., Black locust, 3;w;O,D,C
FI 16897, FM18423, MF6313
1236
a cae leiosperma (Torr. & A. Gray) Piper,
wildbean, 2; s; O,D. FM17598,
re oye
ee biflora (L.) Britton et al., wo-flower
pencil-flower, 1;5;O.FM1/572
Tephrosia virginiana (L.) Pers., Goat's-rue, 1; s; O.
MF6070
Vicia americana Muhl. ex Willd. var. minor Hook.,,
American vetch, 2;m;D.F16874
FAGACEAE
a oe Rydb., Shinnery oak, 5;5,m;O,D.
F16 F16955, F16956, FM17670, FM17916
oueras nee Rydb. x Quercus stellata
Wang , 3,.5,m; O. FM17671, L2000-104,
Quercus macrocarpa Michx., bur oak, 1;w;O.Ts.n.
FUMARIACEAE
Corydalis aurea Willd. subsp.occidentalis (Engelm.
G.B. Ownbey, Golden fumewort,
13m; 0.F 16879
GERANIACEAE
*Erodium cicutarium (L.) L’Her. ex Aiton, Califor-
nia filaree, 2; u;D,C.F16837, MF6304
Geranium carolinianum L., Carolina crane’s-bill, 2;
M,U; F5937,MF6262
GROSSULARIACEA
Sse asta H.WendI., Buffalo currant, 3;w,m;
O:F 1,MF5951
HYDROCHARITACEAE
Najas guadalupensis (Spreng.) Magnus subsp.
dalupensis, Common naiad, 4; |; D,C.
FM17738, FM17853
IRIDACEAE
Sisyrinchium ensigerum E.P. Bicknell, Sword-leaf
blue-e F1685
=
F16946, MF6261, MF6347, MF6372
JUGLANDACEAE
*Carya illinoinensis (Wangenh.) K. Koch, Pecan,
1; u;O.L2001-106
Juglans microcarpa Berland., Texas walnut, Little
walnut, 4; w,m; O,D,C. FM17692, FM17765,
FM18433, FM18461, MF6138, MF6312,
MF6385
* Juglans nigra L., Black walnut, 2;w;O.FM17691
BRIT.ORG/SIDA 20(3)
JUNCACEAE
Juncus dudleyi Wiegand, Dudley's rush, 2; p,r;O
6
Juncus torreyi Coville, Torrey’s rush, 3; p,rl;O,D,C.
M17645, FM17731,FM17743, FM1 7875
KRAMERIACEAE
Krameria lanceolata Torr, Trailing ratany, Lance-
leaf ratany, 3; s,m; O,D,C. FM18374, MF6083,
MF6280, MF6351
LAMIACEAE
Hedeoma drummondii Benth., Drummond's
false-penny-royal, 3; m; D,C. FM17527,
FM17773, FM17892, MF6188
Hedeoma hispida Pursh, Rough false-penny
royal, 3; m; O,D,C. F16918, MF6042, MF6111
56
*Lamium amplexicaule L., Henbit dead-nettle, 1;
u; C.F16909
Lycopus americanus Muhl. ex W.P.C. Barton,
American water- lel ale 3; pl; O,D
FM17636, F 726, FM17763, FM17889,
FM18360, ee
Monarda citriodora ie ex Lag., Lemon bee-
balm, eo m;D,C.F
da clinop
nan
dioides Gray, Basil bee-balm,
3,m;D,C. MF6091, MF6225
Monarda pectinata Nutt.,Plains bee-balm, 2;5;0.
Monarda punctata L.var.occidentalis (Epling) E.J.
oe Spotted bee-balm, 2;s;O.
MF6015 440
Salvia azurea ie Lam., Blue sage, Pitcher’s
C m;O,D,C.[S. pitcheri Torr.— A] FM17792,
FM18278, FM18314, FM18397, FM18422
Scutellaria drummondii Benth. var. edwardsiana
B.L. Turner, Drummond's skullcap, 1; m; D.
Torr., Resinous skullcap, 4m;
O,D,C. | 16947, MF6031, MF6098, MF6211,
MF641
CULEHUIG fESli!
0
L. var. canadense, American
germander, 2; p,rl; O,C. [T. canadense L. var.
virginicum (L.) Eaton — A] FM17685,FM1 7884,
FM18440
Teucrium laciniatum Torr, Cut-leaf germander, 1;
;D.MF6399
LILIACEAE
Allium canadense L. var. fraseri Ownbey, Canadian
onion, 2;s,w;O,C. MF6005, MF6374
FREEMAN ET AL., FLORA OF THE OGALLALA ECOTONE
Allium drummondii Regel, Drummond's onion, 5;
; 6824, F 16939
Nothoscordum bivalve (L.) Britton, Yellow false-
garlic, 1;m;D.F16840
LINACEAE
Linum berlandieri Hook. var berlandieri,
Berlandier’s flax, 2; m; O,C. [L. rigidum Pursh
var. berlandieri (Hook.) Torr. & A. Gray — A, F]
F16925,
HON toa Ea | {|
4;m;D,C. F16831, F16926, MF6410, MF6500
Linum rigidum Pursh var. rigidum, Stiff-stem flax,
4;m;D,C.F 16936, MF6143, MF6163
LOASACEAE
Mentzelia decapetala (Pursh) si i Gilg., Ten-
petal mentzelia, 1;m;D,C. F
Mentzelia nuda (Pursh) Torr. & pe one
mentzelia, Sand-lily, 4; s,m; O,D,C. FM17543,
FM17568, FM17705, FM17815,FM18480
Mentzelia oligosperma Nutt. ex Sims, Stick-leaf,
Chickenthief, 1; m;D. MF6400
LYTHRACEAE
Ammannia coccinea Rottb., Purple toothcup, 3:1;
D.FM17648, FM17801
Ammannia robusta Heer & Regel, Stout toothcup,
2;1;0,D,C. FM17746, FM17883
MALVACEAE
Callirhoé involucrata (Torr. & A. Gray) A. a var.
involucrata, Winecup, 1;m;D.MF61
Callirhoé involucrata (Torr. & A. Gray) A. an var.
lineariloba (Torr. & A. Gray) A. Gray, Winecup,
M17733, MF5994, MF6196
a ae coccinea (Nutt.) Rydb. var. coccinea,
Scarlet globe-mallow, 2; m; D,C. F16843,
F16943, MF6284
MENISPERMACEAE
Cocculus carolinus (L.) DC., Carolina snailseed, 1;
p; O.L2001-66
MOLLUGINACEAE
Mollugo ele illata L., Green carpetweed, 3;5,™,U;
M17652,FM17804,FM17826,MF6041
MORACEAE
*Maclura pomifera (Raf.) C.K. Schneid., Osage-or-
ange, Hedge-apple, 1;w,u;O [see Schambach
(2000) for a discussion of the pre-European
range of this species in Oklahoma].FM18337
*Morus alba L., White mulberry, 1; w;O.MF5953
feos
1237
Morus rubra L., Red mulberry, 1; w; C.FM17872,
MF6383
NYCTAGINACEAE
Mirabilis glat S.W. 11. Smooth four-
o'clock, 2;s;O.FM17614,FM17633
Mirabilis linearis (Pursh) Heimerl var. linearis, Nar-
(C.FM17839,
FM18464, L2001-09, MF6178, MF6301,
MF6422, MF6481
OLEACEAE
Forestiera pubescens Nutt., Downy forestiera, 4;
W,S; ee F16915, FM17628, FM17771,
MF63
ONAGRACEAE
Calylophus berlandieri Spach subsp. berlandieri
erlandier’s evening-primrose, 1; s; O. FM
17659, MF6437, MF6438
Calylophus hartwegii (Benth.) PH. Raven subsp.
pubescens (A. Gray) Towner & P.H. Raven,
Hartweg’s evening-primrose, 3; m; D,C.
F16904, F16941, MF6096, MF6202
a ee ifolius (Torr. & A. Gray) PH.
ender-leaf evening-primrose, 1;m;
ce ie
Calylophus serrulatus (Nutt.) PH. Raven, Plains
yellow evening-primrose, 4; s,m; O,D,C.
FM17548, FM17850, MF6009, MF6072,
MF6113, MF6201
Ae La Raf., American marsh
willo b, Slender-fruit willowherb, 1;9,1;O.
9
Gaura coccinea Pursh, Scarlet butterfly-weed, 3;
m; D,C. F16833, F16902, MF6239
Gaura parviflora Douglas, Velvet butterfly-weed,
3;5,m,U;O,D,C.FM17631,FM17753,FM17830,
FM18371,FM18454, MF6185, MF6289
Gaura villosa Torr. subsp. villosa, Hairy butterfly-
eed, 3;5,m;0.FM17594,FM18370, MF601/7,
MF6457
Oenothera laciniata Hill, Cut-leaf evening-prim-
rose, 2;5,m; O.F 16881, MF5948, MF6483
Oenothera macrocarpa Nutt. subsp. incana (A.
Gray) W.L.Wagner, Gray evening-primrose, 3;
m;D,C.F 16854, FM17542,FM17897,MF6129,
411
Oenothera rhombipetala Nutt.,
Bg) 0S PHosen isi oe ee
ea point
prim
rose, 1:m;C.F] 6928
1238
linifolius (Nutt.) =Heynh.,
s,m;D,C.FM17534,FM17750,
FM18391,
Stenosiphon
Stenosi
FM17812, FM17935, FM18281,
MF6102
OXALIDACEAE
Oxalis ei Jacq. subsp. dillenii, Gray-green
orrel, 2; m,u; O,D,C. F16929, MF6053,
eae FM18470
Oxalils violacea L.,Violet wood-sorrel, 1;5,m;0,D.
F16917
PAPAVERACEAE
Argemone polyanthemos (Fedde) G.B. Ownbey,
Plains prickly-poppy, 2; m; O,D. FM18432,
MF6058
PEDALIACEAE
Proboscidea louisianica (Mill.) Thell., Common
devil’s-claw, 1;1,u;D.FM17802
PHYTOL
Phytolacca americana L. var. americana, Ameti-
can pokeweed, 3; w,u; O,D. FM17736,
FM18414, MF5941
PLANTAGINACEAE
Plantago patagonica Jacq. var. patagonica,
olly plantain, 4; s,m; O,D,C. MF5959,
MF6016, MF61 12, MF6209, MF6340
Plantago patagonica Jacq. var. spinulosa (Decne.)
A. Gray, Bristle-bract plantain, 3; s,m; O,D.
MF6079, MF6121, MF6339
Plantago rhodosperma Decne., Red-seed plan-
u;0,D,C.F 16924, MF6052, MF6100,
MF6208, MF6389
tain, 3;5,.m
POACEAE
*Aegilops cylindrica Host, Jointed goat grass, 2;
/D,C.MF6183, MF6245
*Agrostis gigantea Roth, Spreading redtop, 1; 1;
M1768]
Avidropoaon hallii Hack., Sand bluestem, 5; s,m;
O,D,C. FM17550, FM17784, FM17908,
FM17928, FM18293, FM18406, FM18458
Aristida peat Engelm. ex Vasey, Fork-tip
thre ;5;0.FM18291, FM18369
Aristida on Michx., Old-field threeawn, 2;
s,m,u; O. FM17571, FM17644, FM18289,
FM18366
Aristida purpurascens Poir. var. purpurascens, Ar-
row-feather threeawn, 3; 5,m; O. FM17700,
FM18290
BRIT.ORG/SIDA 20(3)
Aristida purpurea Nutt. var. longiseta (Steud.)
Vasey, Purple threeawn, 1; m; D. [A. longiseta
Steud, — A; A. purpurea Nutt. var. robusta
(Merr.) A.H. Holmgren & N.H. Holmgren — F]
Aristida purpurea Nutt. var. purpurea, Purple
threeawn, 3; s,m; O,D,C. [A. reemeriana
Scheele — A, F] FM17701, FM17848, MF6049,
MF6213, MF6403, MF6477
Aristida purpurea Nutt. var. wrightii (Nash) Allred,
Purple threeawn, 5;m;0,D,C.[A. wrightii Nash
— A] FM17533,FM17556, FM17788, MF6181,
MF6478
*Avena fatua L. wild oats 1; u;D. MF6187
*Bothriochloa ischaemum (L.) Keng var.songarica
(Rupr. ex Fisch. & C.A. Mey.) Celarier & J.R.
Harlan, eee bluestem, 4; s,m,u; O,D,C.
[Andropogon ischaemum L. var. songaricus
Rupr. ex Fisch. ex C.A. Mey. — A, F] FM17660,
FM17901, FM18384, FM18476, MF5988,
MF6335, MF6368
Bothriochloa laguroides (DC.) Herter subsp.
torreyana (Steud.) Allred & Gould, Silver
bluestem, 3; m,u; O,D,C. [Andropogon
saccharoides Sw. - A; A. saccharoides Sw. vat.
torreyanus (Steud.) Hack. — F] FM17555,
ees O, FM17831, MF6021, MF6135,
MF624
Bou un curtipendula (Michx.) Nash var.
curtipendula, Side-oats grama, 5; 5,m; O,D,C.
FM17531, FM17566, FM17789, FM17813,
MF6171, MF6310
Bouteloua gracilis (Willd. ex Sela Lag. ex
ae Blue grama, 2; m; C. L2001-98,
ea i rsuta Lag. var. hirsuta, Hairy grama,
,C. FM17539, FM17609, FM17777,
MF6172
*Bromus pean ar.catharticus, Rescue
brome, 3; u;O,D,C. [B. uni aes Saale A, F]
16839, MES9S7, MF618 216
Se arora i snaiieee brome, 3;uU;
O,D,C. F16953, MF5958, MF6117, MF6215
MF64 a
Buchloé dactyloides (Nutt.) Engelm., Buffalo grass,
4; m; O,D,C. F16933, MF6043, MF6164,
MF6219
Cenchrus spinifex Cav., Coast sandbur, 4;5,u;0.[C.
incertus M.A.Curtis — A,F] FM17649, MF5993
FREEMAN ET AL., FLORA OF THE OGALLALA ECOTONE
Chloris verticillata Nutt., Whorled oe grass,
3; u;O,D,C. MF6166, MF6333,M
Chloris virgata Sw., Showy ae grass, POG.
FM 8913
*Cynodon dactylon (L.) Pers., Common Bermuda
g u;O,C. FM17882, MF5956
Dichanthelium oligosanthes (Schult.) Gould var.
scribnerianum (Nash) Gould, Scribner's pani-
O,C. MF5936, MF6035, MF636/7,
*Digitaria ciliaris (Retz.) Koeler, Southern crab
ass, 1;u;O.[D. adscendens (Kunth) Henrard
— A] FM17696
Digitaria feel (Schult.) Pilg. subsp. cognata,
Fall witch grass, 3; s,m; O. [Leptoloma
eee (Schult.) Chase — A, F] FM17619
FM18285, MF6466
ee ets ae .) Scop,, a crab grass,
>u;O.FM17909, FM18342, MF5978
*Echi deeeh crusgalli (L.) P. Beauv., Common
rnyard grass 3-1.U;0,C.FM17612,FM 17937,
FM18376
Echinochloa muricata (P. Beauv.) Fernald var.
microstachya Wiegand, Rough barnyard
grass, 2; p71; D,C.FM17794, FM18452, L2001-
31
Echinochloa muricata (P. Beauv.) Fernald var.
muricata, Rough barnyard grass, 3;p,r;O,D,C.
FM17686, MF6156, MF6249, MF6394
Elymus elas L. var. canadensis, Canadian
wild-rye, 3 . MF5989, MF6151,
MERE re 432
Elymus virginicus L. var. virginicus, Virginia wild-
Bis a ea rea ae -117,MF6377
“Ec oe Ie NAaAit |
grass, 1;u;0,C. ane 1a, FM18468
eae cilianensis (All.) Vignolo ex Janch.,
Stink grass, 2; u; O,D. FM17563, FM17793,
FM18334
Eragrostis curtipedicellata Buckley, Gummy love
2
*Eragrostis curvula (Schrad.) Nees var. conferta
Stapf, Weeping love grass, 3; s,m,u; O,D,C.
FM17569, FM17787, MF5963, MF6393,
MF6434, MF6455
Eragrostis pectinacea (Michx.) Nees var.
pectinacea, Carolina love grass, 3; u; O.
FM17602, FM18381b, MF5977
Eragrostis secundiflora C. Pres! subsp. oxylepis
1239
(Torr.) S.D. Koch, Red love grass, 1; 5; O. [E.
oxylepis (Torr.) Torr.— A] FM17564
Eragrostis sessilispica Buckley, Tumble love grass,
MF6048, MF6467
Eragrostis spectabilis (Pursh) Steud., Purple love
m;O,C. FM18286, FM18445
Eragrostis trichodes (Nutt.) A.W.Wood, Sand love
grass, 2;5;O.FM18288
Eriochloa contracta Hitchc., Prairie cup grass, 1;|;
D.FM17795
>=
Erioneuron pilosum (Buckley) Nash var. pilosum,
Hairy tridens, 4; s,m; O,D,C. [Tridens pilosus
(Buckley) Hitchc. — A, F] MF6087, MF6221,
F6464
ibatum L., Fox-tail barley, Squirrel-tail
grass, 3;5,m,l; D. MF5968, MF6184
Hordeum pusillum Nutt, Little barley, 4;5,m;0,D,C.
MF5992, MF6157, MF6218
ss oryzoides (L.) Sw., Rice cut grass, 3; p,1,1;0,C
M17688, FM18351, FM18447
ai jum perenne L. var. aristulatum Willd., Peren-
nial rye grass, 1;u;D. MF6180
Muhlenbergia asperifolia ae & Meyen ex Trin.)
P kali muhly, Scratch grass, 3; 1,1; O,D
FM17637,FM18347, FM18434
Muhlenbergia mexicana (L.) Trin., Mexican wire-
stem muhly, 1;w;C.FM17818
uhlenbergia racemosa (Michx.) Britton et al.,
Marsh muhly, 1; w,p,r; O. FM18344
hlenbergia sobolifera (Muhl. ex Willd) Trin.,
ck muhly, 1; w;O.FM18345
Panicum capillare L. var. barbipulvinatum (Nash)
gor, Common witch grass, 2; |; C.
FM18438
Panicum capillare L. var. brevifolium Vasey ex
Rydb. & Shear, Common witch grass, 2; w; O.
FM17675,FM18296
Panicum dichotomiflorum Michx., Fall panicum,
Hordets;m tt
J
aa
Panicum hillmanii Chase, Hillman’s panicum, 2;
Panicum virgatul n ke Switch grass, 2 mol; O,D,C.
FM17617,FM17782, FM17866, FM18287
Pascopyrum smithii (Rydb.) A.L6ve, Western pas-
MF6040, MF6159, MF6268, MF6396
Paspalum setaceum Michx. var. stramineum
(Nash) D.J. Banks, Thin paspalum, 3; s; O.
FM17643,MF5972, MF6479
1240
Phalaris caroliniana Walter, Reed canary grass, 2;
rLu;O,D. MF5961, MF6132
*Poa annua L., Annual blue grass, 1; m,u; C.
F16934
Poa arachnifera Torr, Texas blue grass, 1; s; O.
MF6034
Poa arida Vasey, Plains blue grass, 4; w,m; O,D.
F16870, F16885, F16903
*Polypogon monspeliensis (L.) Desf., Annual
rabbit’s-foot grass, 2; p,rl; O,C. MF5966,
MF6243
Schedonnardus paniculatus (Nutt.) Trel., Tumble
165,MF6217,MF6465
Schizachyrium scoparium (Michx.) Nash subsp.
scoparium, Little bluestem, 5; s,m; O,D,C.
[Andropogon scoparius Michx. — A, F] FM
17531, FM17596, FM17785, FM17895,
M18400
*Secale cereale L., Rye, 1;U;O. MF6012, MF6460
Setaria parviflora (Poir.) Kerguélen, Knot-root
bristle grass, 4; p,r,l;0,D,C. [S. geniculata (Lam.)
P Beauv.— A,F] FM17583,FM17723,FM17744,
M17886
*Setaria viridis (L.) P.Beauv. Green bristle grass, 3;
u;O0,C.FM17697,FM17911,MF5947,MF6376
Sorghastrum nutans (L.) Nash, Indian grass, 4;
/0,D,C.FM17786, FM18352,FM18460
*Sorghum halepense (L.) Pers., Johnson grass, 3;
u; O,D,C. FM17669, FM18326, MF6423,
MF6450, MF6492
Sphenopholis obtusata MITE DN) Scribn. var.
obtusata, Prairie wedgescale, 1;w;O.MF5962
Sporobolus clandestinus (Biehler) Hitchc., South-
eastern dropseed, 2; 5; O. [S. asper (P. Beauv.
Kunth var. donde coe Shinners —
A, FIFM18
Sporobolus rr (Poir.) Merr. var.
Senet Tall dropseed, 4; m; D. [S. asper (P.
uv.) Kunth — A, F] FM18401
rae us yeenr ae (Torr) A. Gray, Sand
dr ed, 4;5,m,u;0,D,C.FM17551,FM17915
2001-40, MF6179, MF6334, MF6485
Sporobolus vaginiflorus (Torr. & A. Gray) A.W.
Wood, ie ty dropseed, 3;u;O,C.FM18277,
FM1836
Tridens a — (Vasey) Wooton & Standl.,
ite tridens, 3;w,m,|D,C.FM17554,L2001-
34, MF61 60, MF6251
Tridens flavus (L.) Hitche. var. flavus, Purpletop, 3;
w,s,m,u;O,C.FM17677,FM18295,-FM1845 1
—_)
BRIT.ORG/SIDA 20(3)
Tridens muticus (Torr.) Nash var. elongatus
(Buckley) Shinners, Slim tridens, 1; m; C.
FM18450
Triplasis purpurea (Walter) Chapm. var. purpurea,
sand grass, 2;5;O.FM18292
Trisetum interruptum Buckley, Prairie trisetum, 3;
s,m; D,C. MF6088, MF6214
*Triticum aestivum L., Bread wheat, 1; u; D.
F6115
Vulpia octoflora (Walter) Rydb. var. glauca (Nutt)
F d weeks annual- ae 4;m;0,D,C.
Festuca aon Walter — A, F] FL16834,
FL16884, MF5960, MF6203, vie 75
POLEMONIACEAE
ee is longi it (Torr) V.E.Grant, Long-flower
mopsis, 2;5,m;0,C. FM18368, Ts.n
POLYGALACEAE
Polygala alba Nutt.,White milkwort, 3;5,m;0,D,C
F16923, MF6067, MF6105, MF6204
Polygala verticillata les Whorled milkwort, 2 S; O.
FM17674,FM17707, MF6063
POLYGONACEAE
Eriogonum alatum Torr. var. glabriusculum Torr.,
—
FM18390, FM18479, L2000-83, L2001-19,
L2001-75
Friogonum annuum Nutt., Annual wild-buck-
at, 4; s,m; O,D,C. FM17575, FM17891,
FM17922,FM18320, FM18388, L2001-88
Eriogonum longifolium Nutt. var. longifolium,
Lance-leaf wild-buckwheat, 4; s,m; O,D,C.
FM17545, FM17585, FM17756, FM17810,
oe 8389, Tee) Hels 15, Mroe°
Doorweed, 3; u; O,C. MF5996, MF6237,
MF6388
Polygonum lapathifolium L., Pale smartweed,
Dock-leaf smartweed, 2; 1,1;O,D,C. FM17749,
FM18353,L2001-84, MF6248
Polygonum ramosissimum Michx., Bushy knot-
weed, 2; w;0,D. FM17715,FM18354, L2001-
2
Polygonum tenue Michx., Pleat-leaf knotweed,
Slim knotweed, 2; 5; O. FM17635, FM17699,
18284
Rumex altissimus AW. Wood, Pale dock, 2; |; C.
6315
FREEMAN ET AL., FLORA OF THE OGALLALA ECOTONE
*Rumex crispus L.,,Curly dock, 2;1,u;0,D,C.MF5942,
MF6369
PORTIEREACERE
L., Common ei 2; 5,1,U;
O,D,C.FM17577, £17795 FM17
Talinum calycinum ee | eon
fameflower, 1;5;O0.FM17
POTAMOGETONACEAE
Meee tae Poir., ane ie pond-
32h M17854, FM18
ae Seles (L.) ee due
2;1,0,D,C. [Potam
FM17844,FM18375, MF6329
PRIMULACEAE
Androsace occidentalis Pursh, eae rock-ja
mine, 2;m; O,D.F16830, F16894
Samolus parviflorus ale cask water-pimpernel,
17724, FM17758, FM17857,
ogeton pectinatus L.— A, F]
FM18441, MF5967
RANUNCULACEAE
Anemone ata Pritz., Ten-petal anemone,
3;m;0,D,C. F16
Delphinium carolinianum Watler subsp. virescens
(Nutt.) R.E. Brooks, Plains larkspur, 2;5,m
(D. virescens Nutt.— A;D. virescens Nutt. ae
penardii (Huth) Ewan — F] MF6032, MF6145,
Ranunculus cymbalaria Pursh, Shore buttercup,
2:1; D.C.FM17880, FM18443
Ranunculus sceleratus L. var. sceleratus, Cursed
crowfoot, Blister buttercup, 2;1;0,C.FM18442,
MF5969, MF6282
RHAMNACEAE
Ceanothus herbaceus Raf., Inland ceanothus, 3;
im; O,D,C. [C. herbaceus Raf. var. pubescens
Cane A.Gray ex S.Watson) Shinners — A, F]
F16844,FM17520, FM17781, MF6001
ROSACEAE
Geum canadense Jacq., White avens, 2; w; O,C.
MF5935, MF6365
Potentilla paradoxa Nutt., Bushy cinquefoil, 2; |;
D,C.FM17742,FM18455
a ae ae Chickasaw plum,
F6060,MF6322,
ndhill plum
ee Dye al eee
Prunus gracilis Engelm.& A.Gray, Oklahoma plum,
2;5,0.F 16883, FM1 7627, MF6429, MF6458
1241
Rubus bushii LH. Bailey, Bush's blackberry, 1;w;O.
F5964
IACEAE
rele occidentalis L., Common button-
»m, rl; D,C.FM17764, FM1 7867
me ium aparine L., Catch- ae on 2; W,U;
Oc. F1691 2; Me ts
Gal ichx., Forest eae 3; Ww;
O,C.FM17870, en MF6363
Hedyotis nigricans (Lam.) Fosberg var. nigricans,
Narrow-leaf bluets, 3; s,m; O,D,C. FM17706,
MF6019, MF6097, MF6246, MF6416
SALICACEAE
ica deltoides W. Bartram ex Marshall subsp.
onilifera (Aiton) Eckenw., Plains cotton-
ood, 5; w,m; O,D,C. F16905, FM18383,
ees MF6456
Salix exigua Nutt. subsp. interior (Rowlee)
Cronquist, Sandbar willow, 2; w,rl; O,D,C.
F16906, FM17729, FM17766, FM18425
Salix nigra Marshall, Black willow, 2; w,rl; O,C.
F16889, MF6279
saaerseeh 3
C umbellata (L.) Nutt. subsp. pallida (A.
DC) PiehI, Umbellate bastard toad-flax, 2;m;
D,C.F16851, MF6295
SAPINDACEAE
Sapindus saponaria L. var. drummondii (Hook. &
rn.) L.D.Benson, Southern soapberry, 3;w,m;
D,C. [S. dtummondii Hook. & Arn. — A]
FM17820, MF6325
Sa ieants sa
aac on lanuginosum Michx. subsp.
blor rao Penn.,Woolly jungle-plum,
ee od, 3; w,m; O,D,C. [Bumelia
eee ae (Michx.) pers var. oblongifolia
(Nutt.) R.B. Clark — A, F] FM17816, FM18338,
FM18430, MF5995, MF6324
SCROPHULARIACEAE
Agalinis aspera (Douglas ex Benth.) Britton,
Rough agalinis, 2; m; D,C. FM18387, L2001-
10, been
Castilleja purpurea (Nutt.) G. Don var. citrina
(Pennell) Shinners, Citron paintbrush, 3; m;
D,C. F16860, F16942, MF6306, MF6497
Castilleja sessiliflora Pursh, Downy paintbrush, 3;
m; D.F16846
ceaed
1242
Nuttallanthus texanus (Scheele) D.A. Sutton,
as toad-flax, 3;5;O. [Linaria canadensis (L.)
svt ae texana (Scheele) Pennell — A, F]
F16888
Penstemon albidus Nutt., White oe 4;
m; D,C.F 16900, MF6253,
Penstemon buckleyi Pennell, Lone 5 beard-
tongue, 2;5;O.F16877, MF6452
Penstemon cobaea Nutt. var. cobaea, Cobaea
beardtongue, 3;m;D,C.FM17523, 2001-93,
MF6101, MF6190, MF6401
Penstemon fendleri Torr.& A.Gray, Fendler's beard-
tongue, 2;m;D.F 16823, MF6342
*Veronica catenata Pennell, Blue water speed-
well, 2; p,r;O. MF 5965
Veronica peregrina L.subsp.xalapensis (Kunth) H.
St.John & FA.Warren, Purslane speedwell, 2;
w,m,|; D.F 16842, MF6356
SMILACACEAE
cite Muhl.ex Torr, Bristly greenbrier, 1;
6382
=
1
SOLANACEAE
Chamaesaracha coniodes (Moric. ex Dunal)
ritton, Dusty false-nightshade, 2; m; D,C.
eee MF6169, MF6264
Datura inoxia Mill, Hairy thorn-apple, 1; w,u; O.
[D. meteloides DC. - A; D. wrightii Regel — A]
M1833]
Physalis cinerascens (Dunal) Hitchc. var.
cinerascens, Ashy ground-cher
ODC.IP vi aL.subsp ee Nee ae
var. cinerascens ne Waterf.— F;P viscosa
L _ cinerascens (Dunal) Waterf. — A]
FM1 7684, MF6109, MF6174, MF6240, MF6496
Physalis longifolia Nutt. var. longifolia, Long-leaf
ground-cherry, 3;5,m;O,C. MF6425, MF6426,
MF64
Quincula lobata (Torr) Raf, Purple ground-cherry,
m,u; D,C.F 16852, MF6175, MF6241
sas dimidiatum Raf., Western horse-nettle,
Bae oo FM18900, FM17927,
ee
Solanum an oe ee leaf night-
shade, 3; s,m; O,D, 829, MF5984,
re
MF6126, MF6227, M64 fi
Solanum interius Rydb., Plains black nightshade
1; u;C. FM18469
Solanum rostratum Dunal, Buffalo-bur night-
BRIT.ORG/SIDA 20(3)
aia 2; 5,m,l,u; O,D,C. FM17620, MF615
TAMARICACEAE
*Tamarix ramosissima Ledeb., Salt-cedar, 2; p,r]:
D,C.FM17760, MF5983, MF6321
TYPHACEAE
Typha domingensis Pers., Southern cat-tail, 4; pl;
O,C.FM17622,FM17687,FM17878,FM18378
ULMACEAE
Celtis laevigata Willd., a hackberry, Sugar-
berry, 1; w;C.
Celtis reti Sa ee ee leaf hackberry, 4; w,m;
O,D,C. F 2,FM18358, MF5954, MF6141,
Ulmus americana L., American elm, 4; w; O,D,C.
F16871, MF5952, MF6142, MF6320
URTICACEAE
Parietaria pensylvanica Muhl. ex Willd., Pennsyl-
vania pellitory, 2; wm; O,C.F16895, MF6281,
MF6371
VERBENACEAE
Glandularia bipinnatifida (Nutt.) Nutt. var.
bipinnatifida, Dakota vervain, 3;m; D,C. [Ver-
bena bipinnatifida Nutt. — A, F} F16920,
MF6147, MF6270
Phyla lanceolata (Michx.) Greene, Northern
fogfruit, Lance-leaf fogfruit, 2; p,l;O,C. (Lippia
lanceolata Michx. — F] FM17722,FM17887
Phyla nodiflora (L.) Greene, Turkey-tangle, Com-
mon fogfruit, 3; |; D.C. [Lippia nodiflora (L.)
abate var. nodiflora — F; P. incisa Small —
M17745, FM1 — FM17862, FM18453,
ee MF63
— bracteata “a & Rodr., Prostrate vervain,
2;U;0,D,C. L2001-24, MF5980, MF6271
Verbena hastata L., Blue vervain, 1;p,r;O.FM17720
Verbena stricta Vent., Hoary vervain, 2;5,m,U;O,C,
EMIZONE, ree
Verbena urticifolia L., Nettle-leaf vervain, 2; w; O.
FM17676
VIOLACEAE
Viola bicolor Pursh, Johnny-jump-up, 3; w,u; O,C.
LV. rafinesquii Greene — A,F] F16907, MF6078
Viola pratincola Greene, Meadow violet, 1;
F16893
VITACEAE
Ampelopsis cordata Michx., Heart-leaf raccoon-
sae
FREEMAN ET AL., FLORA OF THE OGALLALA ECOTONE
grape, 3; w; O,C. FM17690, ae
EMSS FM17871, MF5944, MF63
Pa is quinquefolia (L.) Planch., ai:
1243
Vitis acerifolia Raf, Bush grape, 3; w,s,m; O,D,C.
FM17864, FM18426, MF6059, MF6158,
MF6276
creeper, ne W; O,D,.C. FM17673, FM17770,
MF5946, MF6328, MF6387
Parthenocissus vitacea (Knerr) Hitchc., Thicket
creeper, Woodbind, 2; w; D.FM18427
ZYGOPHYLLACEAE
*Tribulus terrestris L., Spreading puncturevine,
Goat’s-head, 1; u; D.FM17943
ACKNOWLDEGMENTS
The Dempsey Divide Research Foundation, Inc., the R. L. McGregor Herbarium,
and the Kansas Biological Survey provided financial support for this study. We
thank Hillary Loring and Kelly Kindscher for their assistance in the field and
for their review of the draft manuscript. Ronald McGregor, Bruce Hoagland,
and Adam Ryburn also reviewed the manuscript and offered numerous helpful
suggestions. Jennifer Delisle kindly prepared the map.
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GREAT PLAINS FLoRA AssociaTION. 1991 (second printing). Flora of the Great Plains. University
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1246 BRIT.ORG/SIDA 20(3)
BOOK NOTICES
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Edition. (ISBN 06430065148, hbk.). CSIRO Publishing, PO Box 1139, Collingwood
Vic 3066, AUSTRALIA. (Orders: Antipodes Books & Beyond Ltd (USA &
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www.publish.csiro.aw). AU$195.00, 712 pp., color throughout, maps, 7 1/4"
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The first edition of Noxious Weeds of Australia was published in 1992. From the back cover: “Since
the first edition of Noxious Weeds of Australia, numerous changes have taken place throughout Aus-
tralia to the legal pain te i the control of proclaimed weeds and the manner in which they
In this secon n, the authors have completely updated this information for each
—_—
are classifie
State and _ ritory se have listed the newly proclaimed weeds in an addendum. The list of herbi-
cides and their properties) has also ai rev ages
led i ues agricultur al advisers, environmen-
The i
lies etek ane station agents, her-
Salsts, ecologists, be piescntalinesiohas agricultural chemica
ce of agricul | - but with
pas ie +}
ieioemation on over 200 species including nomenclature, habitat, description
and the occasional line enone, the book will also be useful to the botanist and taxonomists. Many
of the weeds covered in this book are widespread making the book useful beyond Australia.—Barney
Lipscomb, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A,
barney@britorg
LeNA StRUWeEand Victor A. Albert (Editors). 2002. Gentianaceae: Systematics and
Natural History. (ISBN 0-521-80999-1, hbk.). Cambridge University Press,
The Edinburgh Bldg., Cambridge CB2 2RU, U.K. (Orders: 40 West 20" ST,
New York, NY LOOL-4211, U.S.A. wwwcuporg, wwwcup.cam.ac.uk). $130.00,
652 pp., b/w photos, line drawings, graphs, 6" « 9".
This volume is a comprehensive review of the Gentianceae family including the latest seca ion
on phylogeny, classification, biogeography, palynology, phytochemistry, and morphology. ‘
tions of each of the 87 gentian genera are provided, as are discussions on morphological pert
and ee for each major ghscmeas! lineage.”
Co s: 1) Gentianaceae in context (V.A. Albert and L. Struwe); 2) Systematics, character evo-
lution, se hogs of Gentianaceae, including a new tribal and subtribal classification (L.
Struwe, J.W. Kadereit, J. Klackenberg, S. Nilsson, M. Thiv, K.V. von Hagen, and V.A. Albert): 3) Cladis-
tics of Gentianaceae: a morphological approach (S. Mészaros, J. De Laet, V. Goethals, E. Smets, and S.
Nilsson; 4) Gentianaceae:a review of palynology (S. Nilsson); 5) The seeds of Gentianaceae (F Bouman,
L. Cobb, N. Devente, V. Goethals, PJ.M. Maas, and E. Smets); 6) Che motaxonomy and phar eee of
Gentianaceae (S.R. Jensen and J. Schripsema); and Index.
SIDA 20(3): 1246. 2003
WEST GULF COASTAL PLAIN ECOREGIONS
Michael H.MacRoberts and Barbara R. MacRoberts
Bog Research
740 Columbia
Shreveport, LA 71104, U.S.A.
an
Herbarium, Museum of Life Sciences
Louisiana State ee Shreveport
Shreveport, LA 71115, U.S.A.
ABSTRACT
-] : ] 1 Sint
We review the literature on West Gulf Coastal Plain gy used
by ecoregion mappers. os tnapeme nae been done almost ny aout suena as-
sessment. We suggest that q I |
r
Key Worps: West Gulf Coastal Plain, Ecoregions
RESUMEN
Hemos revisado la bibliografia de las ecorregiones de la West Gulf Coastal Plain y examinado la
metodologia usada por los cartografos de ecorregiones. La cartografia ecorregional ha sido realizada
casi totalmente sin evaluacion cuantitativa. Sugerimos que las descripciones cuantitativas deben
reemplazar a las descripciones impresionistas.
INTRODUCTION
Half a century ago, Webb (1950) examined the methodology of biogeographers
who had mapped Texas and Oklahoma ecoregions. He found not only that
ecoregional mappers did not agree among themselves but that ecoregional
mapping was subjective. He suggested a remedy: “quantitative methods must
be developed before general agreement on the extent of the major [ecoregions]
can be reached. As long as personal opinion and individual juceme are the
only basis for judging the extent of the [ecoregions], f land insoluble
differences of opinion will continue’ (Webb 1950:246). Using vertebrates, he
quantitatively assessed the ecoregions of Texas and Oklahoma. His ecoregions
did not correspond with those developed by botanists and botanically oriented
ecologists. A decade and a half later, Gleason and Cronquist (1964:177), refer-
ring to the entire North American continent, reiterated Webb's point in almost
the same words but made no quantitative analysis. More recently, Hargrove and
Luxmoore (1998), again looking at the entirety of North America, recognized
the same thing: “Because the delineation lof ecoregions] is based on subjective
criteria, there are as many sets of ecoregions as there are experts.”
These assertions prompted us to examine the history of floristic ecoregion
delineation in the West Gulf Coastal Plain (WGCP). We will be concerned
SIDA 20(3): 1247-1276. 2003
1248 BRIT.ORG/SIDA 20(3)
mainly with three ecoregions: Longleaf Pine (LLP), Post Oak Savanna (POSa),
and Oak-Pine-Hickory (OPH). One of the traditional designations and locations
of these WGCP ecoregions is given in Figure I.
There are numerous descriptions of each of these ecoregions (e.g., Tharp
1939; Gould 1962; Correll & Johnston 1970; LBJ School of Public Affairs 1978;
Hatch et al. 1990; Telfair 1999). The lack of pines, the codominance of oaks and
hickories, and the ubiquitous presence of Quercus stellata Wang. are always
emphasized when the POSa region is described, the combination of pines, oaks,
and hickories is emphasized in the description of the OPH ecoregion, and the
LLP ecoregion is defined as being coterminous with the distribution of Pinus
palustris Mill.
METHODS
1. Ecoregional mappers.— We surveyed the ecoregional literature dealing with
the eastern part of the WGCP to see if Webb (1950), Gleason and Cronquist
(1964), and Hargrove and Luxmoore (1998) were correct in their assessment of
ecoregional mapping. The literature dealing directly or indirectly with the
ecoregions of this area is extensive (Cooper 1859; Transeau 1903; Harshberger
1911; Shreve 1917; Shantz & Zon 1924; Tharp 1926, 1939, 1952; Carter 1931;
Fenneman 1938; Dice 1943, 1968; Blair 1950; Braun 1950; Turner 1959; Gould 1962,
Gleason & Cronquist 1964; Kuchler 1964; Kuchler & McCormick 1965; LBJ
School of Public Relations 1978; Bailey 1980; McMahan et al. 1984; Omernik 1986;
Takhtajan 1986; Diamond et al. 1987; De Graaf et al 1988; Diamond & Smeins
1988; Greller 1988; Barbour & Christensen 1993; Thorne 1993; Bailey et al. 1994;
McNab & Avers 1994; Keys et al. 1995; Weakley et al. 1998; Diggs et al. 1999;
Ricketts et al. 1999a, 1999b; Delcourt &@ Delcourt 2000). Mappers are discussed
and their maps reproduced in the Appendix.
2. Floristics—We made a number of quantitative ecoregional compari-
sons to see how similar or different these regions are. Since there isno complete
floristic survey of any of the WGCP ecoregions, we had to rely on floristic lists
from each (e.g., Anon. n.d; McBryde 1933; Matos & Rudolph 1985; Nixon 1985;
Bridges & Orzell 1989; Hatch et al. 1990; Orzell 1990; Louisiana Natural Heri-
tage Program 1998; Turner et al. 1999; Van Kley 1999a, 1999b; MacRoberts &
MacRoberts 2001), which we have compared with distributional data given in
various floras, atlases, and plant lists (e.g., Smith 1988; MacRoberts 1989; Taylor
& Taylor 1989; Hatch et al. 1990; Thomas & Allen 1993-1998; Kartesz @ Meacham
1999; Turner 2003) and community classifications (Diamond et al. 1987; Loui-
siana Natural Heritage Program 1998; Bezanson 2000). To establish the exist-
ence of different floristic ecoregions one would expect there to be |) significant
regional floristic differences between ecoregions, 2) significant regional com-
munity differences between ecoregions, and 3) higher similarity within differ-
ent parts of an ecoregion than between ecoregions. We were unable to compare
species abundances between regions since the data are not available.
MACROBERTS AND MACROBERTS, WEST GULF COASTAL PLAIN ECOREGIONS 1249
Fic.1.$ loud |
3. Soils.—The interrelationship of soils and flora has long fascinated ecolo-
gists (Warner 1926). Eastern versus western United States is clearly marked by
a soil break (Carter 1931), and in the WGCP many plant communities are cor-
related directly with soils—e.g., blackland prairies, Weches Formation, and sand-
stone glades and outcrops. We surveyed the WGCP soil literature (Carter 1931;
Godfrey et al. 1973; Steila 1993) to see if soils correlate with floristic ecoregional
delineations.
1250 BRIT.ORG/SIDA 20(3)
4. Vertebrates.—A though our main focus is on the flora, we examined the
literature on ecoregional distribution of vertebrates (except fish) in the WGCP
in an attempt to see if animal distributions correlate with traditional plant
ecoregions (Blair 1950; Webb 1950; McCarley 1959; Neck 1986; Gehl bach 1991,
Wilkins 1992; Ward et al. 1994). Also, using lists of birds, reptiles, amphibians,
and mammals from the WGCP, we plotted vertebrate distribution to see if there
were any correlations with ecoregion.
RESULTS
1. Ecoregional mappers.—We confirmed the assessment of Webb (1950), Gleason
and Cronquist (1964), and Hargrove and Luxmoore (1998). There have been
dozens of WGCP ecoregional mappers, but, with one exception, there has been
no floristic mapping based on quantitative data. Instead, we found that
ecoregional mappers almost invariably base their maps on the presence or ab-
sence of one or a few “diagnostic” trees or a “suite” of trees, with little or no at-
tention paid to herbaceous and understory species. None of the modern arma-
ment of statistical ecology and sampling (e.g., Mueller-Dombois & Ellenberg
1974, Peet et al. 1998) has been applied although it is beginning to be used in
WGCP community ecology (Dale 1986; Turner et al. 1999). We also found little
agreement among mappers.
Only Turner (1959) used quantitative methods and, not surprisingly, his
results are unique (Fig. 2). Turner’s method was relatively simple: he calculated
“the percentage of species restricted to a given region Cendemic’ species) and
the percentage of species shared with other provinces (‘shared’ species). The
percentage of restriction, expressed as that fraction of the total species known
to occur in the province under consideration, was taken as the naturalness of
the floral region” (Turner 1959:5 [see also Webb 1950 for comparable method
using vertebrates]). He discovered that there were basically five regions in Texas
that fit his criterion of 40% or higher for “endemism.” Interestingly, he also found
that attempts to delineate clearly distinct, but smaller, subdivisions resulted in
such heterogeneous figures that these regions, ona floristic basis, would not be
strictly comparable. Turner found that floristically the POSa, LLP OPH, and
Cross Timbers are similar and, therefore, he lumped them (his Appalachian)
and separated from them the Prairie (his Campestrian) and South Texas Plains
(his Tamaulipan) because they are different. This is a singular result among
biogeographers, who invariably divide the WGCP into many ecoregions.
2. Floristics.—Using floristic and plant community lists from various ar-
eas within the POSa, OPH, and LLP regions, we compared floristic and commu-
nity similarities and differences throughout the LLP, POSa, and OPH region of
the WGCP. The result: all regions had greater than 90% of their flora in com-
mon. Virtually the same holds for communities. For example, pockets of prai-
rie occur throughout the OPH, POSa, and LLP regions, bogs occur in all three
MACROBERTS AND MACROBERTS, WEST GULF COASTAL PLAIN ECOREGIONS 1251
eee oo,
[tae etre nerner mend
Pann cree rnin teen mamaty arte oe
——
{ ON ep
7
j ae
j i
J |
‘
\
CAMPESTRIAN yee
“ax |
( TURNER 1959
Fig. 2.B.L. Turner’s (1959) Texas vegetation areas.
regions, notably in the LLP and POSa areas, hardwood-beech slopes are found
in both LLP and OPH regions, baygalls are common in all three regions,
grossarenic xeric sandylands and xeric oak woodlands are especially common
in the POSa region but present in the other two regions as well, sandstone out-
crops or barrens occur in the POSa and LLP regions, and so on. Since no one has
collected the data to perform an analysis of species abundance, very little can
be said about that parameter.
One of the predictions of biogeography (we assume) would be that differ-
ent sections of the same ecoregion should be floristically more similar to each
other than to different ecoregions. Biogeographers such as Kuchler (1964; see
also Gleason & Cronquist 1964; Cronquist 1982; Takhtajan 1986) have placed
states as distant as Maine, Michigan, and Texas in the same ecoregion or prov-
ince (Oak-Hickory Forest, Appalachian Province). We compared the floristics
between Kuchler’s Oak-Hickory Forest in Texas (POSa ecoregion) and Oak-
Hickory Forest in Missouri 300 miles away and the Oak-Hickory-Pine Forest
(OPH ecoregion) in Mississippi 300 miles away. We found that the different
ecoregions were more similar (Texas POSa and Mississippi OPH: 81% of species
1252 BRIT.ORG/SIDA 20(3)
in common) than areas within the “same” ecoregion (Texas POSa and Missouri
POSa: 67% of species in common). We also compared the differences between
Michigan and North Carolina and the POSa of east central Texas. Michigan and
North Carolina are both 1000 miles from east central Texas, but Michigan is
supposedly in the same province or ecoregion, while North Carolina is not. The
result: Michigan had 44% and North Carolina had 75% of the species in com-
mon with Texas.
3. Soils.—A survey of the soil literature uncovered no overall soil/floristic
correlations for the LLP, OPH, or the POSa ecoregions but instead a great inter-
digitation of soil types (largely Alfisols and Ultisols) throughout the region
(Tharp 1952; Godfrey et al. 1973; Smeins &@ Diamond 1986). However, the prairie
soils (both blackland and coastal) do fall out clearly along traditional floristic
boundaries. On a broader note, Hargrove and Luxmoore (1998), using multi-
variate analysis of nine edaphic variables (including: elevation, soil, water, tem-
perature, climate), produced a high resolution map of the United States. Over-
laying this on traditional WGCP vegetation maps shows no correlations. The
LLP, POSa, and OPH regions do not stand out individually but merge together
into one general area.
4. Vertebrates.—Zoologists have long known that no correlation exists between
vertebrates and the traditional floristic ecoregions. Instead, the prairie areas form
a break between eastern and western species. Central Texas is a transition zone
for vertebrates, but central Texas begins west of the POSa region (Webb 1950; Neck
1986; Gehlbach 1991; Wilkins 1992: Ward 1994). As far as vertebrates are con-
cerned, the eastern WGCP asa whole is significant, not its components. Our sur-
vey of vertebrate distribution confirms the zoologists’ contentions.
om
DISCUSSION
The most obvious finding of this study is that only one WGCP floristic
ecoregion map was developed using quantitative methods and, parenthetically,
it was subsequently ignored. All other schemesare subjective, the criticism made
by Webb (1950), Gleason and Cronquist (1964), and Hargrove and Luxmoore
(1998). There are no methodological descriptions, no quantitative data, no de-
marcation criteria, and no statistical analyses. Apparently, ecoregions are self -
evident and impressionistic description is considered sufficient. The question
Webb (1950:426) asked five decades ago is as pertinent today as it was then:
“How different must the fauna and flora of two areas be belore they are recog-
nized as examples of different ecoregions?”
Only Turner (1959) attempted an answer, and he found that the eastern
WGCP constituted a single floristic region. We have confirmed this finding:
between 90% and 99% of species occurring in the LLP, OPH, and POSa regions
also occur in the others, and the majority of plant communities are common to
all three regions. When traveling from one region to another, the botanist does
MACROBERTS AND MACROBERTS, WEST GULF COASTAL PLAIN ECOREGIONS 1253
not encounter a new flora, the zoologist a new fauna, and the ecologist a new
set of plant and animal communities. What differences exist are not of kind
but of degree. Sharp boundaries simply do not exist.
Many individuals have noted an east-west gradient between OPH and POSa
regions. Tharp (1926:48) pointed out that Quercus stellata and Q. marilandica
Muenchh. are often codominants with pine in the OPH ecoregion and that the
transition to POSa “.. merely means the gradual decrease of pine until it has
disappeared.” McCarley (1959:390) said: “The boundary between the pine-oak
region and the oak-hickory region is nowhere sharply marked, but is charac-
terized by a gradual decrease in pine and an increase in oaks and hickories.”
Wilson (1990:184) said: “At the western edge of the piney woods, the mixed pine-
oak forest gave way to...a hardwood forest similar in composition to the decidu-
ous component of the mixed forest,” and further, the pines disappear “from the
forest leaving a mixed hardwood forest.” Currie (1991), Nesom and Brown (1998)
and Nesom et al. (1997) documented the loss of tree species diversity, not just
pine, in the WGCP as one moves west. Although pockets of pines occur in the
POSa region almost to Austin and Paris, Texas (Warner 1926; Wilson 1989), pines
tend todrop out of the flora just before the true prairies are reached. Blair 1950)
clearly had trouble separating the OPH and LLP regions from the POSa and spoke
of an ecotone between them and the prairies farther west. Recent analysis of
the distribution of Pinus palustris in east Texas does not support the simple
story of savannas dominated by single species but rather tells a complex story
involving the occurrence of mixed longleaf-shortleaf pine savannas not only
on the periphery of the Pinus palustris range, but at its center (Evans 1997).
Monk et al. (1990:77), in their study of so-called oak-hickory communities across
the eastern United States, found that hickory was seldom, if ever,a dominant or
codominant: “No real evidence was found for a regional oak-hickory forest.” Dale
and Ware (1999), commenting on this matter, pointed out that in contrast to
these earlier findings, hickory may be a codominant in certain areas west of the
Mississippi River.
Few question how the lines get on ecoregional maps—why a given line
forms a polygon without internal polygons of other types. What is evident is
that ecoregion delineations leave out all the messy detail; they are idealizations
that actually describe very little of the area encircled. For example, at the macro-
level the Takhtajan (1986) and Cronquist and Gleason (1964) descriptions of
the Texas POSa region (their Appalachian and Eastern Deciduous Forest prov-
inces) are so far out of the range of the “type” that they are simply not the same
association of plants. At the micro-level, adjacent areas designated as different
are not floristically different at all. As far as we have been able to ascertain, the
LLP “ecoregion” is simply an autecological statement about the presettlement
distribution of Pinus palustris. There is no unique LLP “ecoregion” flora witha
substantial number of endemics and unique plant communities. The POSa
1254 BRIT.ORG/SIDA 20(3)
“ecoregion” is the western edge of the southeastern pine/oak forest and, as such,
many species drop out; notably pines. But again there is no floristic break or
large number of endemics or unique communities by which the POSa
“ecoregion” can be described. The OPH “ecoregion” is similarly undifferenti-
ated—aside from the absence of Pinus palustris, it is difficult to find anything
that warrants setting it off as a separate region.
While the construction of ecoregional maps might seem to have little or
no import, this is not entirely so. Ecoregions are the basis for deciding conser-
vation priorities. For example, Diamond et al. (1997), Ricketts et al. 1999a, 1999b),
Weakley et al. 1999), and Myers et al. (2000) all are concerned with priorities
for conservation of rare species, endemic species, and rare habitat, and deci-
sions about them are based on the concentration of rare species, endemics, and
so on, which in turn is based on which map is selected: Ricketts et al. (1999a,
1999b) chose Omernick (1986), Weakley et al. (1998) chose Bailey et al. 1994),
and Diamond et al. (1997) chose Gould (1962). Setting aside the almost insur-
mountable problems of how species numbers and their rareness are calculated
for these ecoregions, the fact remains that size of area and its location make a
lot of difference in deciding whether or not an ecoregion is an ecological “hot
spot” or “cold spot.” For example, Ricketts et al. 1999a) concluded that the OPH
plus LLP ecoregions (their “Piney Woods Forests”) had only 7 endemic plants
and the POSa region (their “East Central Texas Forests”) also had only 7 endemic
plants. However, by combining the POSa, OPH, and LLP regions, we calculate
that these three “regions” together have approximately 100 endemic or near-
endemic plants, not 14!-thereby instantly changing the WGCP from an eco-
logical cold spot to an ecological hot spot. A map that combines the POSa, LLP,
and OPH regions makes for very different biogeography and conservation than
one that divides them, and the decisions made by conservationists on the basis
of one map may be very different from decisions made using another map.
We are not suggesting that the WGCP or that part of it we have examined in
this paper is uniform. Quite the contrary. Previous biologists related something
they were familiar with and observed, and we, like our predecessors, can see what
was being referred to. If we are attentive, we can tell when we have entered the
LLP region by the presence of Pinus palustris, and traveling west, we also note
when pine vanishes and the landscape is dominated by oaks. We can see these
changes because we have been conditioned to look for them and because they are
there. We also have no doubt that in these different regions there are differences
in species abundance. Thus, the absence of pines from the POSa region and the
presence of pines in the OPH and LLP regions signals difference in canopy struc-
ture. But lacking in any of these ecoregional descriptions is hard data (floristic
lists and abundance data) and statistical procedures and operations for deter-
mining when two samples are ecoregionally the “same” or “different.” Instead,
cultural idealizations and stereotypy substitute for concrete description and
MACROBERTS AND MACROBERTS, WEST GULF COASTAL PLAIN ECOREGIONS 1255
analysis. Consequently, even if we wished to provide a brief summary descrip-
tion of any region inorder to recognize its most valuable or “true” features based
on previous delineations would be to rely on idealized and stereotyped descrip-
tions, and would be confounded by precisely what we are describing here.
It is premature to make any definitive statement on the WGCP ecoregions.
However, on the basis of our experience and limited floristic sampling, we would
say that there is certainly no reason to separate the LLP region from the OPH
region. There is nothing revolutionary in this since about half of the ecoregion
mappers have already lumped them. Our current thinking about the POSa re-
gion is that it is virtually identical floristically to the OPH and LLP regions
minus some tree species. But even this is difficult to maintain because, in north-
ern Texas, pine isa part of the POSa flora. Areas suchas the Gus Engeling Wild-
life Management Area (POSa) in Anderson County are identical edaphically
and floristically to Woods County (OPH): POSa habitat simply exists in Wood
County with pine on it (see Wilson 1990). This is also the case in such areas as
the “Lost Pines” near Bastrop. In conclusion, we find Turner’s (1959) analysis
approximately what we envision.
APPENDIX
This is a survey of the Spinel waist who have produced maps of the WGCP; they are repro-
duced so they can be understood and c red.
The literature describing and ane ecoregions was located by tracing bibliographical
entries and by consulting Kuchler and McCormick (1965). Vegetation mapping of the WGCP di
not begin in earnest until the twentieth century.
ral early botanists described WGCP vegetation but did not give maps.Without maps it is
very difficult, if not impossible, to interpret ecoregion locations precisely; therefore, these descrip-
tions are excluded from this review (e.g., Bray 1906; Warner 1926). Some authors present maps but
add little to previous descriptions; these are also excluded from the present discussion (e.g., Allred
& Mitchell 1955; McMahan et al. 1984). General accounts of early WGCP botanical research and
descriptions of vegetation are given by Harper (1920), Tharp (1926), and Geiser (1948).
P ecoregion terminology has never been standardized and any attempt to cross-index
all usages would be almost impossible. We have, therefore, taken the liberty of standardizing it for
this paper. Since we reproduce the maps, the reader need only see the relationship between vari-
ous maps to judge similarities and differences.
The following abbreviations are used.
CPM = Coastal Prairie and Marsh Ecoregion POSa = Post-Oak Savanna Ecoregion
EGCP = East Gulf Coastal Plai PRA = Prairie Ecoregion
LLP = Longleaf Pine Ecoregion a South Texas Plains Ecoregion
OPH = Oak-Pine-Hickory Ecoregion GCP = West Gulf Coastal Plain
We have redrawn all maps from the originals. This has been done to make them interpretable for
present purposes. We have taken some liberties to clarify complicated maps. Scholars should con-
sult original maps for detail and non terminology.
In this review—which is done chronologically d main|
areas at and below the ecoregional level. Mapping at higher levels (region, province subpro nce)
(Gleason & Cronquist 1964;Good 1974;Cronquist 1982;Takhtajan 1986) is not our concern, although,
1256 BRIT.ORG/SIDA 20(3)
since the“provinces’ of Dice (1 ei Takhtajan ae ite Gleason and Cronquist (1964) in the WGCP
are basically the same as ecoregions, we do disc
Cooper (1859; see Stuckey om ised one of the first province maps for North America
(Fig. 3). He first described the distribution of tree species in North America and, while performing
no statistical analysis, he ices that different areas had different combinations of species not
found in other regions. His map, not dissimilar from that of Takhtajan (1986) and Gleason and
Cronquist (1964), was still being seas almost a century later (Dayton 1949).He divided North
merica into provinces and regions. Two provinces interest us: the Appalachian (eastern wooded)
and the Campestrian (prairie). We ee separated these by a heavy line. He divided each into five
regions (we show only four for the Campestrian). The pattern later to be found in many ecoregion
and province maps ts clearly evident here
u (1903; see also Stuckey & Reese 1981; Delcourt & Delcourt 2000), following in the
footsteps of Sargent (1881), produced a simple map of North American vegetation (Fig. 4). His few
vegetation types are of interest only in that he included the Cross Timbers and POSa regions in the
eastern deciduous forest type and separated the coniferous forest type (what later would become
known as the LLP ecoregion).
Harshberger (1911) produced perhaps the first widely recognized phytogeographical map
of North America, according to which the Atlantic-Gulf Coastal region encompasses most of east
Texas and west Louisiana (Fig. 5). To the west of the Atlantic-Gulf Coastal region, Harshberger showed
what he called the Ozark area of the Alleghenian-Ozark district, which today would be called the
POSa and Cross Timbers ecoregions (POSa on map). Note that the area designated extends well
north of Texas. Also note that south and west of this “Ozark area” is a PRA region as wel as several
other vaguely described regions that extend to the Texas coast. The south¢ t, his “Gulf region
of the Mexican pce ete Oa province,” would be our STP. But the aie ae not follow current
usage. The western-most is his “Edwards Plateau forest, meeting ground for species of Atlantic,
Rocky er and Mexican forests” (EP on map). The center area is his “Transition Prairie-Forest
district, Comprising the oak openings" (TRAN on map). This region is a phytogeographic southern
disjunct from Wisconsin, lowa, and Illinois. The final region is his “Texas cross timber and coastal
plain belt of heavy live oak with prairies sandwiched between." This today constitutes part of the
CPM ecoregion but extends farther inland. Harshberger's use of “cross timber” here is confusing to
the modern biogeographer. Also note that the limits of longleaf pine (and many other trees) are
shown on his original map but that no longleaf pine ecoregion is recognized. But the salient fea-
tures of the Harshberger map are that east Texas fl lly is firmly joined to the Gulf and Atlantic
anne plains, and the POSs region is clearly recognized. There are several reprints of Harshberger
(1911) but with black and white maps. These are difficult to interpret, and only the colored map of
the first edition is satisfactory.
Shreve (1917) virtually established the modern WGCP pattern (Fig.6). He distinguished three
WGCP forest regions. The POSa region extends unbroken from south Texas to Michigan, the LLP
region extends from southeastern Texas to Florida and North Carolina, and the OPH region ex-
tends from Texas to Virginia. This pattern, as will be seen, was followed with very little modification
by such influential biogeographers as Braun (1950) and Kuchler (1964). Shreve's original map is in
color and cannot be interpreted from black and white reproductions of which there are man
ntz and Zon (1924), in their widely reproduced map, added essentially nothing to Shreve
(Fig. a Shantz and Zon map has been widely reprinted, sometimes with minor modifications
ot ayes 1962; Fowells 1965).
’s (1926) interest was regional (Fig. 8). He mapped Texas east of the 98th Meridian and
did extensive field work throughout east Texas. He provided extensive d tions of the vegeta-
tion regions. His captionless map (his plate Ill) is confusing, and the text jeaties careful study for
complete interpretation. But his map is the basic pattern that is familiar today: a LLP region in
southeast Texas, a coastal region, an OPH region to the north and west of the LLP region, and a
—
MACROBERTS AND MACROBERTS, WEST GULF COASTAL PLAIN ECOREGIONS 1257
‘ oh j \
—~L u | |
7 { 2
| 7 1
DACATOH {ILLINOIS
| L 4
i | Nat
rene |
eee aes = —
CAMPESTRIAN PROVINCE AP sree PROVINCE -
| a4 ‘ | \ os CAROLINIAN
AN Fd | TENNESSEEAN *
MS ME, | :
| \ {
COMANCHE \ MINSISSIPRIAN —
Sea
(
TS
COOPER 1859
Fic.3. Vegetation areas according to Cooper (1859).
ey re
fo a Se
if i 7
DECIDUOUS FOREST
( \
pcey |
eee \ |
|
i
i
TRANSEAU 1903
Fic. 4. Vegetation areas according to Transeau (1903).
1258 BRIT.ORG/SIDA 20(3)
i i \ v. i
/ ee
! po
| i f
! !
~ i j
ety Lo
POSa 5 i
PRA camaceacess
ATLANT 1G GULF COASTAL
ON
PRA — x
)
EP RAN CPM
SIP e
HARSHBERGER 1911
Fis. 5. Vegetati ling to Harshberger (1911).
POSa region to the west of the OPH region. Tharp divided the POSa region into two areas. To the
north, prairies are not Common but to the south is an “oak-prairie mictium," that is, a combination
of POSa and prairie southwest of the Brazos River. Later biogeographers showed this area as POSa
with islands of prairie in it (e.g.,Gould 1962; Diamond & Smeins 1988)
Carter (1931), the chief soil scientist for the Texas Agricultural Experiment Station, in his sur-
vey of the soils of Texas, produced a vegetation map for the whole state (Fig. 9). The accompanying
soil map, while not Weare: the vegetation patterns precisely, does delineate the distribution of
“main areas of deep sands’ that correspond closely with the POSa region. Descriptions of the re-
gions are given in text.
Fenneman (1938) Deas a slightly modified Shantz and Zon (1924) map (not reproduced
here). He filled in the non-for areas along the coast of Louisiana and Texas that they left blank.
A modified version of this map is in pennies (2000).
Tharp (1939) refined his regional vegetational map (Fig. 10). In this version, the lines that
were tentative in 1926 are bold. The WGCP is virtually identical to that given by Carter (1931), and
any comparison of Tharp's map to that of Gould twenty years later leaves no doubt about influ-
ences. Tharp’s brief descriptions of the vegetational regions are often folksy but never dull. Tharp’s
map was used by Wynd (1944)
Dice’s (1943, ses a Le aio one at a descriptive level above the ecoregion (Fig. 11).
‘ ee scheme, Dice divided east Texas into two regions: the Austroriparian and the Texan. The
mpasses ciel the entire Atlantic and Gulf Coastal Plain (OPH and LLP). The
MACROBERTS AND MACROBERTS, WEST GULF COASTAL PLAIN ECOREGIONS 1259
SN Ee Nese,
en ne nee warn gar ema,
ye een nt a serene pice
}
a
|
c
an
\
{oP ?
PRA poss A \ Ai ip
LLP i S
(“CPM tS
_— oe me
STP UF
SHREVE 1917
Fic. 6. Vegetation areas according to Shreve (1917).
Texan takes in the POSa as well as the prairies and cross timbers regions, and extends from central
coastal Texas to northern Oklahoma. Blair (1950) followed Dice in his description of the east Texas
biotic Seana but added nothinc
’s (1950) classic work on eastern forests adds little to our knowledge of the WGCP
region ae 12). Her map of"Forest Regions’ is essentially identical to that of Shantz and Zon (1924)
with only minor boundary shifts. She kept the three “regions” format: LLP, OPH, and POSa. While
Braun sampled vegetation widely across the east, and produced tables of percent composition
from many sites, but none from the WGCP she did not statistically compare regions, but left that to
her successors (e.g., Delcourt & Delcourt 2000).
rp’s (1952) final map (not reproduced here) of the vegetation regions represents revi-
sions and refinements of maps in his 1926 and 1939 publications. The greater detail is based on a
decade of additional field experience, but the refinements for the WGCP are en minor and
add little to his earlier work. Consequently, we do not pre ould be
noted inet his ae of each vegetation region are succinct and clea
Turner (1959) is unique among WGCP autor | in that he used Freee methods to de-
termine el he calls’ a provinces’ (Fig. 2).We ha 1 his work in the first part of this paper.
Gould’s (1962) map is clearly influenced by Caner (1931) and Tharp's (1926, 1939, 1952)
work (Fig. 13). The map and slight variants have been used extensively (e.g., Correll & Johnston
1970; McMahan et al. 1984; Hatch et al. 1990; Diggs et al. 1999) and one version of it presents subre-
gions (e.g. LBJ School of Public Relations 1978; Diamond et al. 1987; Telfair 1999)
1260 BRIT.ORG/SIDA 20(3)
ee mea,
SHANTZ & ZON 1924
vi . aS CL i 7 {107A)\
Fic. 7 tat
Ss] }
easonand hee eel detailed ten floristic provinces of North America, four of which
occur in the WGCP (Fig. 14). What is of interest to us in this map is the association of the POSa
region, notably regions as far south as Guadalupe County in Texas, with regions as far north as
Maine, southern Ontario, Michigan, and Minnesota: this entire area Gleason and Cronquist called
the Eastern Deciduous Forest Province. The floristic similarity among extreme parts of this large
region is very small indeed. At the same time, their Coastal Plain Province is nothing more than the
traditional Atlantic and Gulf Coastal Plain that extends from east Texas to Florida and north to
Virginia—a much more floristically homogeneous region than the aa ern Deciduous Forest Prov-
ince. Takhtajan (1986) produced a province map of North America ae chee to that of
¢ vaneatet (1964), which has been reproduced (Thorne 1993) (Fi
’s (1964) map,“Potential Natural Vegetation of the pi ; co States,"has
become the “central dogma” of North American biogeographers upon which all other maps are
judged (Fig. 16). It is widely cited and used (e.g., Martin et al. 1993). The Kuchler map presents no
novelties for the WGCP,; it is virtually identical with those of Shantz and Zon (1924) and Braun (1950).
Mahler (1981) presented an interesting variation on the Gould (1962) map (Fig. 17). Appar
ently Puente the lead of McLeod (1971), he designated not a LLP ecoregion in southeast Texas
but a Beech-Magnolia-Pine-Oak-Sweet Gum (Big Thic ket) region overlying most of the traditional
P ecoregion. hy Mahler (1981) chose a course is not clear, but it underlines the fact that other
ecoregion a can be consider
’s (1986) was the first ecoregion map where ecoregions no longer straddle the
Mississippi ie r (Fig. 18). Additionally, Omernik did not divide the pineywoods region into two
Gleason anc
=
oQ
=
MACROBERTS AND MACROBERTS, WEST GULF COASTAL PLAIN ECOREGIONS 1261
i !
i 3
|
i ‘
Eo
i
i
;
‘
4
THARP 1926
Fic. 8. Vegetation areas according to Tharp (1926).
ecoregions (OPH and LLP) but like Gould (1962) kept them together. Otherwise, the familiar pat-
tern is maintained. But what is sensi here is that the LLP region is not uniquely distinguished
in either the WGCP or EGCP. How Omernik distinguished EGCP ecoregions from WGCP ecoregions
is not evident since his brief ee are virtually of no help.
Greller (1988) produced a map that he claims is based on Braun (1950) but which shows no
closer relation to hers than it does to the maps of Shantz and Zon (1924) or Kuchler (1964) (Fig. 19).
Ms is as about Greller’s map is that it combines the POSa and the oe Kegions with no
ti n (but see Turner 1959). Greller did foi vever, retain the other »g
\
features, ach as the LLP ecoregion.
ur and Christensen (1993) presented a relatively simplified vegetational map of North
America (Fig. 20).We find in it the traditional LLP and OPH regions, but the POSa region has been
separated and joined into a vast region stretching from the Gulf of Mexico to Canada. This region is
described as “tallgrass prairie, including the prairie peninsula and oak savanna ecotone with the
eastern deciduous forest” (1993:98). Treating the POSa region as an“ecotone”and including it in the
prairie ecoregion has been commented on (Gehlbach 1991) and is reminiscent of Dice's (1943,
1968) and Blair's (1950) “Texan” biotic province.
Bailey et al. (1980, 1994, see also DeGraaf et al.
old Texan biotic province of Dice (1942, 1968) and Blair (1950), and then divided it into four subre
gions (Fig. 21). This ecoregion (“province” and “ecoregion” are apparently synonyms here) extends
1988; McNab & Avers 1994) resurrected the
1262 BRIT.ORG/SIDA 20(3)
| |
4 t
+ }
|
} ss
ae
ae a ae ee ee
| wae
=
}
. \
CARTER 1931
Fic. 9. Vegetation areas according to Carter (1931).
from the Texas coast to northern Oklahoma. The eastern most part is separated as the “Oak woods
and prairies subregion’ (255C), which is virtually identical to the POSa region of previous writers
except that it includes several major sections of the prairie region of east Texas, such as Fayette
Prairie. This makes the Bailey “Oak woods and prairies subregion” non-comparable with the maps of
most cial biogeographers. Like many before them Bailey et al. (1994) divided the remainder of
n two regions. The southern one corresponds with the LLP and the northern one corre
anne to ae At the subregional level, however, they separated both the OPH and the LLP re-
gions west of the Mississippi River from the OPH and LLP regions east of the ee River.What
distinguishes these areas, aside from geographical location, is not disclosed, and the McNab and
rs (1994) descriptions do not help.It should be further noted that the Bailey (1994) text nee the
McNab and Avers (1994) descriptive material are peppered with factual mistakes, something not
found in previous biogeographic descriptions
The map of Keys et al. (1995) and Bailey (1994) are virtually identical. Because the Key et al.
(1995) map is more detailed, the western boundary of the WGCP is more easily distinguished (Fig.
22). The northern POSa region may have been included in the OPH region while the southern
POSa may have been included with the Prairies sections. Bailey et al. (1994) may have done this
also, but that map is cruder and harder to decipher: they may have put the entire POSa region in
with the prairie group. The Keys et al. (1995) map divides ecoregions into dozens of "nested" sub-
sections that are not readily distinguishable on the basis of their descriptions (nor are they distin-
guished by regional ecologists and botanists). Also, the Keys et al. (1995) map is clearly based on a
poor knowledge of the WGCP. The text presents several factual mistakes.
==
MACROBERTS AND MACROBERTS, WEST GULF COASTAL PLAIN ECOREGIONS 1263
t
|
| |
| roe
a
PRA OPH
\
NY od
THARP 1939
Fic. 10. Vegetation areas according to Tharp (1939).
The Nature Conservancy (Weakley et al. 1998; Turner et al. 1999) relies heavily on Keys et
al.(1995) and Bailey et al. (1994), and it is through the Nature Conservancy maps that it is possible to
detail precisely where the Keys et al. (1995) ecoregions, provinces, sections, and subsections are
(Fig. 23). The POSa region has disappeared as a region. |In the north, it is divided between the prai-
ries region and the “Upper West Gulf Plain” region. In the south, the "West Gulf Coastal Plain’ region
is basically coterminous with Longleaf pine. This new terminology should lead to confusion
Ricketts et al. (1999a, 1999b) divided North America into two eee levels. The
first is ten Major Habitat Types in which most of the WGCP is lumped with the LLP region and not
with the OPH region of the east, and the POSa region is lumped with the majority of the east but is
geographically separated from it. At the finer level, in the WGCP the traditional LLP and OPH re-
gions are lumped into the “pineywoods region;” whereas in the EGCP. they are inexplicably kept
separate. The POSa region is separated from the “pineywoods.” In the EGCP, while the traditional
LLP and OPH regions are recognized, they are ignored in the WGCP.The descriptive sections of this
account are replete with factual mistakes, and it is clear that part of the problem is a heavy reliance
on Bailey et al. (1994). Ricketts et al. (1999a, 1999b) apparently believe that longleaf pine occurs
over the entire “pineywoods’ region. Since the Ricketts et al. maps of the WGCP are identical with
Omernik’s map, the reader is referred to that map for detail.
1264 BRIT.ORG/SIDA 20(3)
i 4
| / ro
| ——,
a |
i
|
|
t
}
e |
AUSTRORIPARIAN
\ / _ —
a ELA
{
DICE 1943
Fic. 11. Vegetation areas according to Dice 1943, 1968).
CONCLUSIONS
In summary, there has been relatively little evolution of the “ecoregion” maps
of the WGCP over the 20th century. The patterns, but not the precise bound-
aries, found in the earliest maps are found in the later ones. The main questions
appear to be whether or not to join the WGCP with the EGCP, whether or not to
join the LLP region with the OPH region, and what to do with the POSa region?
Should the latter be a part of a mega-region stretching from south Texas to
Michigan? Should it be combined with the prairie region to the west, should it
be kept separate from all other regions, or should it be cut in half and part of it
put with the prairies and part with the OPH region? About half of the mappers
recognize an LLP ecoregion, while the rest do not. lt would appear that bota-
nists who have constructed floras or distribution maps do not recognize a sepa-
rate LLP ecoregion, presumably because the vegetational distinctiveness is ei-
ther minor or virtually nonexistent (Correll @ Johnston 1970; Gould 1975; Vines
1977; Nixon 1985; Hatch et al. 1990; Nixon & Kell 1993). Turner 1959) is the prime
example of this. It is mainly ecologists who distinguish a separate LLP region.
MACROBERTS AND MACROBERTS, WEST GULF COASTAL PLAIN ECOREGIONS 1265
pees eerevey
tremens
frtbslnmtat ton eencaret utes
ig
188 0 heen nae ene g ae mel HI
r
\& BRAUN 1950
Fic. 12. Vegetation areas according to Braun (1950).
ACKNOWLEDGMENTS
Guy Nesom commented on several versions of the manuscript and brought B.L.
Turner’s ecoregion work to our attention. Billie Turner reviewed an earlier ver-
sion of the manuscript and made many useful comments. L.M. Hardy aided
with the maps.
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THE JOHN D. FREEMAN HERBARIUM (AUA): A HISTORY
AND
CATALOGUE OF VASCULAR PLANT TYPE SPECIMENS
Curtis J. Hansen
John D. Freeman Herbarium
ery Sciences Department
Life Sciences Bui Iding
uburn University
Auburn, AL 36849, U.S.A.
ABSTRACT
The John D. Freeman Herbarium at Auburn University can trace its roots almost back to the founda-
tion of the institution itself in 1856. By 1867 nee campus boastes a natural history museum that housed
botanical collections. This was most likely herbarium at Auburn University. Such
notable former faculty a as ES. Earle, L.M. LU nderwood ad ee PH. Mell, Jr, surmounted early
challenges, i fi 1887, to estapnst and maintain a a collection. By
1901, the herbarium noueed over 40,000 from A.H. Curtiss,
G.W. Carver, A.B. Langlois, and J. fou Tagialy the he baricra was again destroyed by fire in 1920.
Despite this, AUA again emerged fro e ashes to reestablish itself as an important state and re-
gional herbarium. Growth at AUA from 1995 was due t to the enone of ne palesone a Freeman,
who took the herbarium from obscurity to national g f hold-
ings during his tenure. The Freeman hore now contains over 60, 000 sheets and is one of the
largest and most modern collections in Alabama. The herbarium currently houses 26 documented
vascular plant type specimens that are catalogued.
RESUMEN
El Herbario John D. Freeman de la Universidad de Aul r | rT Ieeda lat ]
dela ee en 1856. En 1867 la ciudad universitaria disfrutal | le histori l
que albergaba colecciones botanicas. Es pr Span que éste no EHESE el primer ae organizado que
tuvo atin ersidad de Auburn 1 FS arle | LM Underwood,
PH. Mell, Jr, vencieron daddies temas incluso un Uae ce en se
ate ei el mantenimiento de la coleccion del herbario. En 1901, el h mas
40,000 especies, incluyendo colecciones importantes de A.H. Curtiss, G.W. Carver, A.B. Eaeiis -
Darby. meg camens 2 herbario destr mde una vez mas por un bance en ee - aig de la
]
segunda tragedia, el herbario volvida surgi
t
mas importantes del ee y ae la region. El crecimiento del herbari sde los anos 1968-199
debido a los eee del ex- euradon Jos nn D. nee man, quien tomo sia her pane desde sus pent um ae
ar
hasta set y quint UL Lic Cl
i
4 SV OOD pi qd
curador. Actualmente el herbario John D. Freeman cuenta con mas y es uno de
los herbarios mas grandes y modernos del estado de Alabama. Hasta la fecha A herbarie alberga 26
ejemplares tipo de plantas vasculares documentadas que estan propiamente enumeradas y
catalogadas
SIDA 20(3):: ): 1277- 1287. 2003
1278 BRIT.ORG/SIDA 20(3)
HISTORY
The early history of the John D. Freeman Herbarium can be traced back almost
to the foundation of Auburn University itself, and was directly affected by the
challenges of establishing an institution of higher education in the late 1850s.
Auburn University was originally chartered in 1856 as the East Alabama Male
College (EAMC),a Methodist college that officially opened its doors to students
in 1859 (Yeager & Stevenson 2000). Botany was the only life science course of-
fered at the college and was first taught by John Darby, A.M., the “Julia Ann
Hamiter” Professor of Natural Science (Davis 1992). Darby was a botanist, chem-
ist, and author of one of the earliest floristic works to focus on the southern
United States (Darby 1857). This work was intended as a botanical textbook for
college and high school students throughout the South and was surely used in
Darby’s own curriculum, along with other botanical texts by Gray and Chap-
man. While there is no evidence that an official herbarium collection existed
in 1859, Darby no doubt had a set of pressed and dried botanical specimens
—_
—
which he used for teaching botany and plant identification
Botany continued to be taught until 1861 when EAMC was closed due to
the Civil War, in which many of the faculty and students entered the Confed-
erate service (Draughon 1954). The Old Main building, predecessor to current-
day Samiord Hall, served as a Confederate hospital from 1864-1866 (Yeager &
Stevenson 2000). In 1866, the college reopened and Darby was again elected to
teach botany (Minutes July 9, 18606). Darby was given emeritus status in 1869
but continued to be listed as a faculty member until 1871. In the 1867 catalogue
of the EAMC it lists a “museum which is now being formed” as one of the assets
and student recruiting points of the college. This museum housed plants, min-
erals, fossils, insects, mammals, and Indian relics, among other items. Darby was
involved with the college museum and most likely organized the first herbarium
collection at Auburn in the late 1860s.
A depressed, post-Civil War economy led to serious financial difficulties
that plagued the college during the Reconstruction Period. In 1872, after much
debate and considerable objection, the Board of Directors (later Board of Trust-
ees) transferred the EAMC property and buildings to the state of Alabama to
establish a land grant college under the provisions of the Morrill Land Grant
Act passed by Congress in 1862 (Draughon 1954; Yeager & Stevenson 2000). This
act, extended to former Confederate States after the Civil War and ratified by
the Alabama Legislature in 1867, provided 30,000 acres of public land for each
senator and representative of a state as an endowment to establish a college
with the goal “to promote the liberal and practical education of the industrial
classes in the several pursuits and professions of life” (Draughon 1954). In 1872,
the college was officially named the Agricultural and Mechanical College of
iu
HANSEN, THE JOHN D. FREEMAN HERBARIUM 1279
Alabama (A & M College) and the herbarium became an integral part of Agri-
cultural Experiment Station work.
Along with the change from church-college to state-college came changes
in administration and some faculty members. In this turbulent time, J.B. Read
was selected as the first Professor of Natural Sciences, which included the re-
sponsibility of teaching botany. However, his tenure was cut short, when after
only eight months the Board of Directors reduced the number of faculty from
eight to six, presumably for financial reasons. For the next three years the pro-
fessorship of Natural Sciences was given to William C. Stubbs, a chemist. How-
ever Stubbs, who already taught chemistry, mineralogy and geology, probably
did not take on the added responsibility of teaching botany. There are indica-
tions that in 1876 botany classes were taught by Professor of Agriculture and
President of the young A & M College, Isaac T. Tichenor (botany classes were
required by the School of Agriculture course of study according to the early
and mid-1870s course catalogues [Yeager & Stevenson 2000)). In 1877, the Board
of Trustees hired E.Q. Thornton as Professor of Natural Sciences, but his tenure
lasted less than one year due to his unexpected death (Yeager & Stevenson 2000).
Finally, in 1878 the Trustees hired the talented Patrick H. Mell, Jr. to be Pro-
fessor of Natural Science (Yeager & Stevenson 2000). Under Mell’s direction and
leadership the college museum holdings were greatly expanded through ac-
quisition of specimens via purchases, exchanges with other institutions, and
donations from private collectors and citizens. During Mell’s tenure we find the
first documentation of the actual size of the herbarium. Reporting for the 1881-
1882 school year, Mell stated that the museum had increased in botanical col-
lections from 266 to 1577 (Mell 1882). The following year he reported an in-
crease of 442 botanical specimens for a grand total of 2019 (Mell 1883).
Tragically, in 1887, the herbarium was destroyed when Old Main caught
fire and burned. While the number of collections destroyed isn’t known, the
herbarium was indeed important and growing under Mell’s direction. He de-
scribed his work and the condition of the herbarium prior to the fire as follows:
“It was my pleasure to begin the preparation of this Botany of Alabama prior to 1887 but the
destruction of the Museum by fire that year lost the Institution a valuable herbarium rich in
plants collected from Alabama by Prof. Darby, Prof. Thornton, Prof. Stubbs and myself” (An-
nual Report 1896).
Despite the destruction of the herbarium, Mell published a list of cultivated
grasses in Alabama which included the first attempt to list all the grasses that
occurred in state (Mell 1889). Prior to the fire, Mell began compiling and writ-
ing the first flora of Alabama, to be issued in sections as they were completed
(Mell 1896). He was finally able to publish one section of his flora but, unfortu-
nately, no other sections were ever completed (Mell 1896). Incidentally, Charles
1280 BRIT.ORG/SIDA 20(3)
T. Mohr, who was concurrently writing his own Alabama flora, considered Mell
an upstart in the business of writing the flora of Alabama, and felt threatened
by his efforts (Davenport 1979).
Mell’s talents were not limited to teaching botany and directing museum
collections. Along with these responsibilities, he also served as the first Direc-
tor of the Alabama Weather Service from 1884-1893, an agency that he created.
He developed a system of flag signals used to relay weather conditions and fore-
casts around the state, a system that was soon adopted by the National Weather
Service (Yeager & Stevenson 2000). Later, Mell declined to have his name sub-
mitted to President Grover Cleveland to become Chief of the U.S. Weather Bu-
reau (see Mell Scrapbooks 1891-1906, Auburn Univ. Archives). In 1891, he was
also appointed Director of the Alabama Experiment Station where he served
until 1902 (Yeager & Stevenson 2000). In 1896, Mell was elected Chairman of
the Section of Botany and Horticulture of the American Association of Agri-
cultural Colleges and Experiment Stations (Anonymous 1899). After turning
down other prestigious offers, including the presidency of Mercer University,
Mell finally resigned from Auburn in 1902 to accept the presidency of Clem-
son College in South Carolina (Kerr 1985).
In late 1895, Lucien M. Underwood came to Auburn as a biologist and be-
gan working immediately on the “fungous flora of Alabama, since compara-
tively little is known of the extent and character of the rich cryptogamic flora
of Alabama” (Annual Report 1896). Underwood departed after only one year
but not before collecting a great deal and publishing a treatise on edible fungi
(Underwood 1896).
Frank S. Earle succeeded Underwood and served as biologist and horticul-
turist. Earle finished the final preparations on Underwood's “fungous flora of
Alabama” and in 1897 published their list of Alabama fungi with Underwood
as lead author (Underwood & Earle 1897). This list enumerated over 1100 spe-
cies of fungi known to occur in Alabama at the time. Also noted was the fact
that the college at Auburn was the only institution in the state that maintained
a course in botany. They also described some of the collectors and collections
in the herbarium at that time (Underwood & Earle 1897). Earle later provided
this list of fungi to Charles T. Mohr who included it in the fungi section of his
monumental work on the flora of Alabama (Mohr 1901). Earle also studied and
collected vascular plants from the unique habitats on and surrounding the meta-
morphic or granite outcrop regions of eastern Alabama (Earle 1902).
Earle provided excellent documentation of the growth and composition of
the herbarium collections from 1896 to 1900. Indications are that Mell’s admin-
istrative duties kept him away from the herbarium and that by the turn of the
20" century Earle was essentially managing the herbarium. On January 1, 1901,
Earle reported the following numbers and composition of the herbarium (An-
nual Report 1901):
—_—
—
HANSEN, THE JOHN D. FREEMAN HERBARIUM 1281
Flowering Plants, etc. 20,606
Fungi 16,950
Lichens 1,006
Mosses, etc 588
Algae 984
TOTAL 40,134
Considering that the herbarium burned in 1887 that was an impressive accu-
mulation of specimens in 13 years.
Earle left Auburn in 1901 and was replaced by Edwin M. Wilcox, Professor
of Botany, and self-described “Vegetable Physiologist and Pathologist” (Annual
Report 1901). Wilcox was the department head and taught botany classes from
1902-1908. After he arrived, Wilcox immediately began preparing what
amounted to the first plant atlas for the state of Alabama. He began a card in-
dex of all the plants, including a county map of the state pinpointing each col-
lection location (Annual Report 1902). It is doubtful the project was ever com-
pleted because during his six-year tenure Wilcox only referred to the herbarium
twice in his annual reports. Other than beginning the plant indexing project,
Wilcox’s only other reference to the herbarium was a request to get modern
cases to hold and preserve the collections (Annual Report 1904). While Wilcox
made some strides in maintaining the herbarium, no further reports document-
ing the size and growth of the herbarium exist after 1901. Most of Wilcox’s time
was spent working asa plant pathologist, not herbarium curator.
Francis E. Lloyd replaced Wilcox in 1908 as Professor of Botany. The her-
barium was apparently poorly maintained and had fallen into disuse during
Wilcox’s tenure. In Lloyd’s first report to the president he stated that, “Steps have
been taken toward...the rehabilitation of the herbarium” which, he continues,
“is of paramount importance for Experiment Station work as a source of infor-
mation in matters pertaining to the flora of this section of the country” (An-
nual Report 1908). Despite Lloyd’s emphasis on the importance and necessity
of the herbarium, not much was accomplished during his first year. The next
year, Lloyd again stressed the need to put the herbarium “into shape for use”
(Annual Report 1909). Furthermore, Wilcox’s 1904 request for modern her-
barium cases must have gone unfulfilled because in 1909 Lloyd stated that the
greatest need for the herbarium “is proper housing in the form of tightly con-
structed cases, since without these we cannot control the insect pests which
are peculiarly bothersome here” (Annual Report 1909).
After 1909, little documentation is found relating to the herbarium. It is
well known, however, that by 1910 the Botany Department (and herbarium),
along with many other life science departments, had moved toa new building
on campus. According to the 1910 bulletin, “A handsome and commodious
building has been erected for the departments of (1) agriculture, (2) horticul-
1282 BRIT.ORG/SIDA 20(3)
ture, (3) botany, (4) entomology, and (5) animal husbandry” (Davis 1992). Fur-
thermore, this building, named Comer Hall, was “thought to be superior or equal
to any building in the south” (Yeager & Stevenson 2000). By 1918, the Botany
Department occupied the western half of the third floor of Comer Hall. Incred-
ibly, the herbarium was again completely destroyed by flames when a fire en-
gulfed and gutted Comer Hall in October 1920! The total number of specimens
lost in this fire is unknown, but by extrapolating from Earle’s 1901 report of
40,134 total specimens, an estimate of greater than 50,000 specimens destroyed
would be conservative.
The composition of the herbarium before the Comer Hall fire is fairly well
documented, at least through the year 1900. The following is a list of some of
the valuable collections known to have existed in the herbarium prior to the
fire of 1920. Beyond simple documentation, this list serves only to sharpen the
tragedy of the loss. First and foremost, the losses included important historical
and scientific collections by G.F Atkinson, C.F Baker, G.W. Carver, J. Darby, J.F
Duggar, FS. Earle, A.B. Langlois, PH. Mell, Jr, W.C. Stubbs, E.Q. Thornton, L.M.
Underwood, and an entire set of collections documenting A.H. Curtiss’ work on
the flora of Florida. Additionally, a large collection of Alabama grasses, an ample
fades
seed collection, and 500 exchange specimens from Liverpool, England were also
consumed in the blaze. Finally, the loss of documentation for Mell’s work on the
Flora of Alabama, as well as many other research projects, was especially dis-
appointing (Davis 1992). Other departments that were partially or completely
destroyed included Agronomy, Plant Breeding, Soils, Plant Pathology, Horticul-
ture, Animal Husbandry, Entomology, and the administrative offices (Annual
Report 1921). The Comer Hall fire was a monumental loss in the history of the
herbarium and Auburn University. However, unlike Comer Hall, which was
rebuilt in two years, it took many decades to rebuild the herbarium.
While the herbarium was indeed reestablished, there is little information
regarding its growth during the years following the fire. In the 1950s, the heirs
of Elizabeth F Andrews, a botanist and author, donated her personal collection
of about 2500 plants to the herbarium which included collections by A.W. Chap-
man. This donation formed the nucleus for further rebuilding efforts. Donald E.
Davis, Professor of Botany, came to Auburn in the late 1940s and served as her-
barium curator for a time. Davis’ students in plant taxonomy and weed science
were required to submit specimens and thereby added many collections to the
herbarium from 1947-1968 (Davis 1992). Other taxonomists in the botany de-
partment during the 1950s and 1960s included Edward T. Browne, Jr. and Al-
fred C. Koelling (Don Davis, pers. comm.). Browne came to Auburn in 1956 and
was herbarium curator until about 1960. The herbarium grew very little dur-
ing his short tenure, but herbarium records do indicate there was an exchange
agreement with the University of Georgia. Browne himself only added about
two dozen of his personal collections to the herbarium. Later, Koelling served
HANSEN, THE JOHN D. FREEMAN HERBARIUM 1283
as herbarium curator from 1964-1967. In efforts to increase herbarium holdings,
Koelling initiated a number of exchanges with other institutions and also added
over 750 of his own collections. Herbarium growth from 1922 to 1968 was mod-
est, but by 1968 the herbarium contained a total of 11,300 sheets (Davis 1992).
Herbarium growth from 1968-1995 was due to the efforts of the late John
D. Freeman, who emphasized the importance of fieldwork and collecting, and
that the herbarium was an essential and valuable botanical research tool. He
initiated exchange programs with many institutions and was active in loaning
specimens out to various researchers. Freeman directed several graduate stu-
dent research studies that helped document the flora and assess plant distribu-
tions in critical areas throughout the state. These types of studies led to active
collecting efforts and, in turn, added greatly to the herbarium holdings. Under-
graduate students who took Freeman's systematic botany courses can testify
to the emphasis he placed on collecting. His students were required to turn in
large collections of 150-200 plants — pressed, identified and labeled — at the
conclusion of the course. That amount of collecting kept exchange programs
bolstered and greatly enlarged the herbarium holdings. In 1974, Freeman sum-
marized the growth of the herbarium during his first five years at Auburn as
follows (Freeman 1974):
Specimens mounted
and shelved
Year
1969 3,357
1970 Sor
1971 3,640
1972 2,100
1973 1,420
Total iO
Cumulative through 1968 11,300
Present holdings 29,594
Over his career, Freeman took the herbarium from relative obscurity to
national recognition. In 1974, the Advisory Committee for Systematic Resources
in Botany (1974) selected the Auburn University Herbarium as a National Re-
source Collection, a collection considered vital for continued science support
and research. This designation was given to only 105 (out of a total of 1127)
institutions nationwide. Freeman’s personal research interests likewise garnered
national attention as he focused on the genus Trillium. He collected Trillium
from all across the United States and Japan, naming seven new species includ-
ing the federally listed southeastern endemic Trillium reliquum.
Prior to his death, Freeman helped negotiate the acquisition of the her-
barium collection at St. Bernard College (SB) in Cullman County, Alabama. This
1284 BRIT.ORG/SIDA 20(3)
invaluable, historic collection of about 5000 sheets was the personal herbarium
of Wolfgang Wolf, a prolific botanist in the early 20" century and lay brother
of the Order of Saint Benedict. This collection contains not only Alabama plants
but also many collections from around the eastern United States and Europe.
This acquisition added greatly to the holdings of the Freeman Herbarium and
pushed the number of total collections to over 60,000. Ultimately, Freeman in-
creased the number of mounted specimens five-fold in 27 years. In January 1998,
the Board of Trustees, in recognition of his enormous botanical contributions,
unanimously passed a resolution officially naming the Auburn University
Herbarium in honor of Dr. John D. Freeman.
With no permanent curator in place after 1995, activity at the Freeman
Herbarium dropped dramatically. For the next three years Jerome Ward, a vis-
iting assistant professor, supervised herbarium activity ona part-time basis and,
with the help of students, began the long process of organizing and incorporat-
ing the St. Bernard collection.
From 1998 to 2002, Cynthia M. Morton held the post of plant taxonomist
and director of the Freeman Herbarium, and in 1999 the author was hired as
curator. Under Morton’s direction the Freeman Herbarium was brought back
to full activity with the resumption of many exchange agreements and in-
creased loan activity. The collections were refoldered using archival quality
genus covers, updated taxonomically, and organized geographically. Many mis-
filed collections were found and duplicate collections removed, which greatly
relieved limited cabinet space. Another major emphasis during this time was
—
——
to have the label information from each specimen entered into a computer da-
tabase. With the help of students and support from the State Lands Division of
the Alabama Natural Heritage Program, this project was completed in 2002.
The database is maintained as new specimens are accessioned and added to the
collection and is available via the World Wide Webat http://www.auburn.edu/
herbarium. The effort to database the herbarium collection has led to the pub-
lication of a checklist of Alabama plants compiled from the combined infor-
mation of the Freeman Herbarium and University of Alabama Herbarium
(UNA) (Morton et al. 2002).
The Freeman Herbarium is now one of the largest collections in Alabama
and isa vital resource for various natural science departments at Auburn Uni-
versity including the Agricultural Experiment Station, Biological Sciences, En-
tomology & Plant Pathology, Forestry, Horticulture, Pharmacy, Veterinary Medi-
cine, and Wildlife. The Freeman Herbarium is used throughout the region by
other colleges, universities, private companies, and government agencies, and
serves as the state herbarium (Alabama Code §1-2-12). The collection houses
valuable historic specimens that date to the late 19" and early 20" centuries,
including many by E.F Andrews, A.W. Chapman, and W. Wolf with the earliest
collections dating to the 1840s. More recent contributors who have added im-
HANSEN, THE JOHN D. FREEMAN HERBARIUM 1285
portant collections to the Freeman Herbarium include V.E. Crouch, A.R. Dia-
mond, J.D. Freeman, S.C. Gunn, R. Kral, H.D. Moore, D.W. Rutland, A.H. Sessler
and J.W. Short. While it is richest in specimens from Alabama and the southern
United States, the Freeman Herbarium also contains collections from around
the country and from many continents in the world. Growth and diversifica-
tion of the Freeman Herbarium are continuing through open gift and exchange
programs and fieldwork.
TYPE SPECIMENS
Currently, the Freeman Herbarium contains over 60,000 mounted vascular
plants, of which 26 are documented as types either through citation in original
literature or as annotated by a later researcher. This list includes 2 holotypes,
23 isotypes, 1 isoneotype and is arranged alphabetically by family, then by ge-
nus. The majority of type specimens are found in Trilliaceae (8) and Asteraceae
(7). The reference citation, location (restricted to country, state and county, or
equivalent), date, collector, collector number, AUA accession number, and the
holotype location, if known, is documented for each specimen.
CATALOGUE OF TYPE SPECIMENS
ARISTOLOCHIACEAE
Hexastylis shuttleworthii (Britten & Baker)
Small var. harperi Gaddy, isorvre. Sida 12:54.
1987.USA ae Madison Co.:9 May 1986,
Gaddy s.n. AUA 60732. (HoLoTyee: CLEMS).
ASTERACEAE
Coreopsis grandiflora Hogg ex Sweet var.
inclinata J. Allison, isotype. Castanea 66:162.
2001. USA. ALABAMA. Bigs Co.: 15 Jul 1999, Alli
son 12086. AUA 61466. (HoLoryee: US).
Erigeron strigosus Muhl. ex Willd. var.
Satan Allison, isorype. tie 66:169,
1, USA. Atasama. Bigs Co.: 10 Jun 2000, Alli-
son eee AUA 61464. NY).
Liatris xfreemaniana J. Allison, HOLOTYPE. ees
nea 66:179. 2001. USA. Atasama. Bigs 18
Jul 1993, Allison and Stevens 7804, AUA - 463.
Liatris x macdanieliana J. Allison, isorype. Casta-
nea 66:180. 2001. USA. Atasama. Bigs Co.: 20
Aug 1994, Allison 8559. AUA 61462. (HOLOTYPE:
US).
Liatris oligocephala J. Allison, isotyre. Castanea
66:175. 2001. USA. Atasama. Bigs Co.: 18 Jul
1993, Allison and Stevens 7802. AUA 61461.
(HOLOTYPE: GH).
scale oe Allison, isoryee. So
83. 2001. USA. ALasama. Biss Co.
ne aaa 12503. AUA 61459. (HOLOTYPE:
GH)
Silphium perplexum J. Allison, isotype. Castanea
2001.USA. ALABAMA. Daitas Co.: 18 Aug
1999, Allison 12153. AUA 61458. (HoLoTYPE: GH
BORAGINACEAE
Onosmodium decipiens J. Allison, isoryre. Cas-
tanea 66:180. 2001. USA. ALasama. Bigs Co.: 1
Nov 1993, Allison and Stevens 8139. AUA
61460. (HoLotyre: NY).
CYPERACEAE
Carex cumberlandensis Naczi, Kral, & Bryson,
isorype. Sida 19:994, 2001. USA. ALasama. Win-
sTON Co.:4 May 1991, Naczi 2868. AUA 62540.
(HoLoTYPE: DOV)
Fimbristylis argillicola Kral, isotyre. Sida 4:128.
1971. MEXICO. MicHoacan. 18 Aug 1965, Kral
25509. AUA 16998. (HoLoTyYPE: MICH).
FABACEA
Dalea aes Allison, isoryre. Castanea 66:166.
2001. USA. ALapama. Bigs Co.: 1 May 1994, Alli-
son and Stevens 8236. AUA 61465. (HOLOTYPE:
NY).
—
1286 BRIT.ORG/SIDA 20(3)
LAMIACEAE 13 Apr 1966, Kral 26254. AUA 21765. (NEOTYPE
Scutellaria montana Chapman, isorype (det. by GH).
Robert Kral 1979]. Bot. Gaz. 3:11. 1878. USA. ees ipiens J.D. Freeman, isorvee. Brittonia
GEORGIA. FLoYD Co.:no eatin 5.n. AUA 27:17. 1975. USA. Georaia. Earty Co.: 28 Mar
ie man 615 with Murrell lhara & lhara
PORTULACACEAE AU 56759, (ovate: GH)
: Trillium foetidissimum J.D. Freeman, \sorype.
alinum mengesii W. Wolf, Hovoryre [det. by Ken-
neth J.Wurdack 1988]. Amer. Mid. Naturalist
Brittonia 27:31.1975.USA. Mississipel. ADAMS Co.:
6:153. 1920. USA. ALABAMA. CULLMAN Co.:9 Aug ¢
1967, Any man 535 with Murrell &
1919, Wolfs.n. AUA 60731 ova, AUA 16713, GH)
ey src ] Freeman, isorype. Sida 3:289
RANUNCULACEAE _ USA. Texas. Sasine Co: 1 Apr 1966,
Clematis esse isoryee. Annals Mis aa ma 1039 with Freeman & Channel.
souri Gard. 74:665. 1987. USA. ALasama. AUA 568738. (HoLoTyPeE: GH)
ten oe 7 Jun “083, Kral 70176. AUA Trillium kurabayashii J.D. Freeman, isotype.
45730. (HoLoryPe: MQ). Brittonia 27:56. 1975. USA.Catirornia, Det Norte
SCROPHULARIACEAE Co.:8 May 1967, Freeman 594 with Freeman.
Castilleja caus J. Allison, isotype. Castanea AUA 56822. (HoLoTyPE: GH)
_ USA. ALABAMA. Bigs Co.: 15 Apr Trillium maculatum Raf. f. simulans J.D. Free-
anne ee 10466. AUA 61467. (Hol ennenys man, isotype, Brittonia 27:29. 1975.USA, SoutH
Carouna. DorcHesTer Co.: 10 Apr 1966, freeman
TRILEIRCERE 451a with Freeman. AUA 56864. (HOLOTYPE:
Trillium albidum J.D. freeman, isotype. Brittonia
27:48. 1975. USA. OREGON. JOSEPHINE CO.:9 May
1967, Freeman 596 with Freeman. AUA 56829.
VDB).
Trillium reliquum J.D. Freeman, |soryee. Brittonia
eF et GEORGIA. RICHMOND Co.: 20 M
(HOLOTYPE: G 1968, Free Murrell. lhara&| naa
bc channeliii | hee ne Sal & M. AUA Seay. ee GH).
ou, isotype. Novon 996, JAPAN.
ee bo. KusHiro Dist: on ay 1994, Fukuda XYRIDACEAE
and Itou s.n. AUA 57091. (HototyPe: MAK). Xyris tennesseensis Kral, \sorvr. ils
Trillium cuneatum Raf, Isoneotyee. Brittonia 80:444. 1978. USA. Tennessee. Lewis Co.:26 Aug
27:34.1975.USA.NortH CARoina. BuNcomBE Co.: 1969, Kral 36370. AUA 32229. (Hovoryre: US).
ACKNOWLEDGMENTS
lam grateful to Don Davis for sharing early memories of the Freeman Herbarium
and Botany Department at Auburn University, and to the staff of the Special
Collections &@ Archives Department at the Auburn University Library for their
helpful assistance in many hours spent researching records. | thank Terry Rod-
riguez of the Office of Communications at the Alabama Agricultural Experi-
ment Station for tracking down original copies of some very old extension bul-
letins and then sharing in the excitement of discovery. Roland Dute provided
helpful comments on an earlier draft of this manuscript. Critical reviews by
Larry Davenport and Robert Kral improved this paper. | thank Glenda Gil for
her Spanish translation of the abstract.
HANSEN, THE JOHN D. FREEMAN HERBARIUM 1287
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chanical College of Alabama. Auburn Univ. Archives, Auburn,
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Darey, J. 1856. Botany of the southern states. A.S. Barnes & Co., NY.
Davenport, LJ. 1979. Charles Mohr and Plant Life of Alabama. Sida 8:1-13.
Davis, D.E. 1992. History of the Department of Botany, Plant Pathology and Microbiology
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DraucHon, R.B. 1954. Alabama Polytechnic Institute. An address given at the 1953 Ala-
bama dinner of the Newcomen Society of England. The Newcomen Society in North
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Earte, FS. 1902. The flora of the metamorphic region of Alabama. Science Contributions
Alabama Agric. Exp. Sta. Bull. 119:43-120.
Freeman, J.D. 1974. The Auburn University Herbarium: a record of the plants of Alabama.
Auburn Univ. Agric. Exp. Sta. Highlights of Agric. Res. 21:12.
Kerr, N.A. 1985.A history of the Alabama Agricultural Experiment Station 1883-1983. Ala-
bama Agric. Exp. Sta., Auburn Univ. Brown Printing Co., Montgomery, AL.
Mett, PH., Jr. 1882. Professor Mell’s report to the Board of Trustees of the Alabama Agricul-
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Auburn, AL.
Met, PH., Jr. 1883. Extracts from the report of the Department of Natural History to the
Board of Trustees of the Alabama Agricultural and Mechanical College. Mell Scrap-
books 1891-1906. Auburn Univ. Archives, Auburn, AL.
Met, PH., Jk. 1889. Grasses and their cultivation. Alabama Agric. Exp. Sta. Bull. 6:149-186.
Mett, PH., Jk. 1896. The flora of Alabama. Part V: Leguminosae and Rosaceae. Alabama
Agric. Exp. Sta. Bull. 70:276-296.
Minutes. 1856-1957. The Board of Trustees at the Alabama Polytechnic Institute. Auburn
Univ. Archives, Auburn, AL.
Mone, C. 1901. Plant life of Alabama. U.S. Government Printing Office, Washington, D.C
Morton, C.M., C.J. Hansen, and G.Git.2002. Checklist of the vascular plants of Alabama from
the John D. Freeman Herbarium and the University of Alabama Herbarium. Carnegie
Mus. Nat. Hist. Spec. Pub. No. 21.
Unoerwooo, L.M. 1896. Edible fungi: a wasted food product. Alabama Agric. Exp. Sta. Bull.
73:337-346.
Unberwoon, L.M.and FS. Earte. 1897.A preliminary list of Alabama fungi. Science Contribu-
tions. Alabama Agric. Exp. Sta. Bull. 80:113-283
YeaceR, J.and G. Stevenson. 2000. Inside Ag Hill:the people and events that shaped Auburn's
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ENCYCLOPEDIAS
FROM MARCEL DEKKER
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SIDA 20(3): 1288. 2003
NEW VASCULAR: PLANT RECORDS FOR THE
GRASSLAND BIOME OF CENTRAL NORTH AMERICA
Craig C. Freeman Caleb A.Morse
RL. McGregor Herbarium Division by ee
1 a
RL. McGregor Herbarium Division of Botany
Natural History Museum and Natural ey Museum
see Research Center plover Research Center
ol of Kansas, 2045 i ila University of Kansas, 2045 Constant Avenue
ce, Ks 66047-3729, U.S.A. Lawrence, Ks 66047-3729, U.S.A
ae cmorse@ku.edu
Ronald L.McGregor
RL. MGGIEGOE! Herbarium, Division of Botany
Natural H um and Biodiversity Research Center
Uni ea ena 2045 Constant Avenue
Lawrence, KS 66047-3729, U.S.A.
ABSTRACT
Additions to the vascular floras of four states are reported herein: Colorado (Calibrachoa parviflora,
Chamaesyce prostrata, Gleditsia triacanthos, Oenothera engelmannii, Oenothera laciniata,
See eae, and Veronica persica), lowa (Deschampsia ae be Ohi Kansé
an albicans var. australis, Carex swanii, Car rvi, ae
oseus, ee diandrus, Dichentheliten dichotomum subsp. Poe: Dichanthelium
polyanthes, xElyhordeum montanense, Erigeron canus, Geranium dissectum, Geranium molle, Hi-
biscus syriacus, Phyllanthus urinaria subsp. urinaria, Pyrus calleryana, Sagina apetala, Sedum
sarmentosum, Silphium radula, and Tridens flavus var. chapmanii); and Oklahoma (Epilobium
leptophyllum).
RESUMEN
Se hacen adiciones a la flora vascular de cuatro estados: Colorado (Calibrachoa parviflora,
Chamaesyce prostrata, Gleditsia triacanthos, Oenothera engelmannii, Oenothera laciniata,
Trianthema porta lagasem) y Veronica persica), lowa (Deschampsia danthonioides); Kans
anthera caracasana, Carex albicans var. australis, Carex swanii, Carum carvi, ae
roseus, Cyperus diandrus, Aiea dichotomum subsp. dichotomum, Dichanthelium
polyanthes, xElyhordeum montanense, Erigeron canus, Geranium dissectum, Geranium molle, Hi-
biscus syriacus, Phyllanthus urinaria subsp. urinaria, Pyrus calleryana, Sagina apetala, Sedum
a, y Tridens flavus var. chapmanii); y Oklahoma (Epilobium
rmentosum, Silphium radu
eae m).
Continuing field and herbarium studies in the grassland biome of central North
America have produced additional records of vascular plants previously un-
documented in parts of the region. Herein we report 28 new state records for
vascular plants in Colorado (7), lowa (1), Kansas (19), and Oklahoma (1). Thir-
teen of these reports are considered to be native occurrences in the region; fif-
teen (marked with an asterisk) are considered the results of recent introduc-
SIDA 20(3): 1289-1297. 2003
1290 BRIT.ORG/SIDA 20(3
=
tions. These reports update the distribution data contained in Atlas of the Flora
of the Great Plains (Great Plains Flora Association 1977) and Flora of the Great
Plains (Great Plains Flora Association 1991). All voucher specimens are depos-
ited in the R. L. McGregor Herbarium (KANU), University of Kansas, unless
otherwise indicated.
*Alternanthera caracasana Kunth (Amaranthaceae). Native to South and Cen-
tral America, mat chaff-flower has been reported from throughout the south-
ern US. (Kartesz @ Meacham 1999). Previous reports record this species from
the panhandle of Texas and Oklahoma (Great Plains Flora Association 1991). A
single specimen confirms its presence in Kansas.
Voucher Bae ate KANSAS. Douglas Co.: Lawrence, University of Kansas C aaa T12S, — sec
36, SW !/4, SE!/4, near 38°57'29'N, 95°14'54"W, elev 305 m, weedy plantings and s Ik cr
campus, 30 Aug 2002, Morse 8534.
*Calibrachoa parviflora (Juss.) D'Arcy (Solanaceae). This prostrate annual oc-
curs throughout the southern U.S. and tropical America and is probably a re-
cent introduction into the Great Plains. Freeman et al. 1998) reported it from
central Kansas. Two populations were discovered in eastern Colorado in 2000,
both along the shorelines of reservoirs.
—
Voucher specimens: COLORADO, Bent Co.: Las Animas, 12 mi N, Adobe Creek Reservoir State Wild-
life Area, SE side of Adobe Creek Reservoir, T21S, R52W, sec 4, E!/2, SE!/4, elev 4.140 ft, level, a
shortgrass prairie and muddy to sandy shoreline, . 2000, Freeman 15345 & Morse. Kiow
Eads, 10 mi S, 4.5 mi E, Neenoshe ervoir, 5 side, T R47W, sec 9, SW !/4,SW!/4 & sec 16, ae
elev 3,930 ft, open, sandy to muddy shoreline and ad wooded perimeter of reservoir, 20 Aug
2000, Freeman 15982 & Morse.
lly
Carex albicans Willd. ex Spreng. var. australis (L.H. Bailey) Rettig (Cyperaceae).
Rettig (1990) recognized three varieties of C. albicans. Variety albicans occurs
in the Great Plains from eastern Nebraska south through eastern Kansas and
into central Oklahoma (Rolfsmeier &@ Wilson 1997; Taylor & Taylor 1994;
Yatskievych 1999). Recently, a population of approximately 100 individuals
clearly referable to var. australis was discovered in southeast Kansas in a sandy,
oak woodland. The nearest documented populations in Arkansas (Hyatt 1998)
and Missouri (Yatskievych 1999) are more than 250 km away.
Voucher specimens: KANSAS. Neosho Co.: Thayer, 0.5 mi W, Thayer City Lake, N side, 37°2901'N,
95°29'13"W, post ce blackjack oak woodland on sandstone, gentle S-facing ee witha mesic
ravines, 17 Apr 2C na 6776. ee 0.5 mi W,N side Thayer City La 29S, RI8E, sec 31,
NW !/4,NW !/4, re 7 305 m, mowed parkland, a area, and open blackjack sale post mt forest,
soil sandy, 23 Apr ae nine 5486 & W. oo
Carex swanii (Fernald) Mack. Cienieaes Carex swanii is broadly distrib-
uted throughout the eastern U.S. and southeastern Canada. A small population
recently discovered in eastern Kansas extends the range of this species by more
than 300 km from the nearest known stations in central Arkansas (Hyatt 1998)
and southeastern Missouri (Yatskievych 1999).
FREEMAN ET AL., NEW PLANT RECORDS OF CENTRAL NORTH AMERICA 1291
Voucher specimen: KANSAS. Douglas Co.: Junction of US Hwy 56 and Co Rd 1055 in Baldwin City, 3
mi N, 2.25 mi E, T145, R20E, sec 13, S!/2, SW !/4, near 38°49'30"N, 95°08'30"W, elev ca 290 m, open,
black-post-blackjack oak forest and upland tallgrass prairie on N- and W-facing slopes, surfacing
sandstone, sandy soil, 27 Jun 2002, Morse 8383. [Additional voucher at MICH
*Carum carvi L. (Apiaceae). Caraway is a Eurasian biennial that has become
established sporadically in waste places in the northern Great Plains (Great
Plains Flora Association 1991). The species now is added to the Kansas flora
based on a population recently discovered in the extreme eastern part of the
state.
Voucher specimen: KANSAS. Miami Co.: Somerset, | mi E, T175, R24E, sec 3, SW !/4, NW !/4, near
38°36'00"N, 94°45'00"W, elev 275 m, weedy floodplain forest and rocky gravel bars along S side South
Wea Creek, 5 Oct 2000, Morse 5186.
*Catharanthus roseus (L.) G. Don (Apocynaceae). A native of the Old World
tropics, rosy periwinkle has been planted extensively as an ornamental in the
US. Kartesz and Meacham (1999) mapped records from scattered states in the
southern U.S. Recent observations and a single collection confirm the presence
of C. roseus as an adventive species in Kansas.
Voucher specimen: KANSAS. Douglas Co.: Lawrence, University of Kansas Campus, T125, R20E, sec
36, SW !/4, SE!/4, near 38°57'29'N, 95°14'54"W, elev 305 m, weedy plantings and sidewalk cracks on
campus, 30 Aug 2002, Morse 8533.
| |
Chamaesyce prostrata (Aiton) Small (Euphorbiaceae). Prostrate mat-spurge is
a common annual in the southern half of the Great Plains. Kansas and Nebraska
records occur westward to the Colorado border (Great Plains Flora Association
1977), but there are no records for that state. It now may be included based ona
single collection from extreme east-central Colorado.
Voucher specimen: COLORADO. Kit Carson Co.: Burlington, 3 mi E along US 24, T8S, R43W, sec 35,
NW !/4,NW !/4, elev 4,010 ft, disturbed, open roadside just S of I-70 overpass, soil silty, 18 Aug 2000,
Freeman 15764 & Morse.
Cyperus diandrus Torr. (Cyperaceae). Cyperus diandrus is confined to the ex-
treme eastern Great Plains (Great Plains Flora Association 1977) except in Ne-
braska, where it is fairly widespread (Rolfsmeier 1995). In far western Missouri,
Yatskievych (1999) mapped records from Jackson and Dade counties. Previous
reports of the species in Kansas (Gates 1940) were based on misidentified speci-
mens of the superficially similar C. bipartitus Torr. The species now may be
included in the Kansas flora based on an overlooked specimen collected 40 years
ago. Several recent attempts to relocate this population have been unsuccess-
ful.
Voucher specimen: KANSAS. Atchison Co.: Mouth of Independence Creek, 28 Jul 1961, Blocker 759
[KSC],
*Deschampsia danthonioides (Trin.) Munro ex Benth. (Poaceae). This tiny an-
nual, native to western North America and South America (Cronquist et al. 1977),
1292 BRIT.ORG/SIDA 20(3)
has been reported as adventive in Ohio, New York, and Maine (Kartesz &
Meacham 1999). The first record from lowa, where it apparently wasa contami-
nant in grass seed, is reported here. The site has been revisited several times,
but there is no evidence that plants have persisted.
Voucher specimen: IOWA. Bremer Co.: Waverly, S edge of town, TIN, RI4W, sec 10, SE !/4, SE !/4,
42°42'03"N, 92°28'34'"W elev 990 ft, small area of reseeded lawn on W side of US 218 at 2416 4" Street
SW, exposed, loamy soil, 6 Jul 2001, Freeman 17518
Dichanthelium dichotomum (L.) Gould subsp. dichotomum (Poaceae).
Dichanthelium dicotomum is known from several counties in extreme western
Missouri (Yatskievych 1999), but has not been reported from Kansas. Recent
examination of specimens at KANU revealed a single specimen misidentified
as D. acuminatum (Sw.) Gould & Clark var. fasciculatum (Torr) Freckmann,
which represents the first verified record for the state.
Voucher specimen: KANSAS. Franklin Co.: | mi E, 3 mi S Homewood, ridge of N- a" cliffs above
small stream on W side of road, plants in sandy soil, dry, 20 Aug 1976, Brooks 1244
Dichanthelium polyanthes (Sc hult.) Mohlenbr. (Poaceae). Kartesz and Meacham
(1999) mapped records of this taxon, as D. sphaerocarpon (Ell) Gould var.
isophyllum (Scribn.) Angelo, from throughout the eastern U.S., westward to
Missouri, Oklahoma, and Texas. For its part, D. sphaerocarpon is common in
eastern Kansas, but no attempt has been made to distinguish the other mem-
bers of the complex to which that species belongs. Recent examination of Kan-
sas specimens identified as D. sphaerocarpon at KANU revealed a single speci-
en of D. polyanthes, which represents the first verified record for the state.
Voucher specimen: KANSAS. Cherokee Co.: 3.5 mi E Crestline, moist open woodland, 22 Jul 1982,
McGregor 33433.
xElyhordeum montanense (Scribn.) Bowden (Poaceae). This naturally occur-
ring hybrid between Elymus virginicus L. and Hordeum jubatum L. subsp.
jubatum has been reported from nearly a dozen states and provinces in the USS.
and Canada (Kartesz @ Meacham 1999). A recently redetermined specimen at
KANU is the first record for Kansas.
Voucher specimen: KANSAS. Republic Co.: T45, R2W, sec 28, elev 1,380 ft, silty clay loam, 25 Aug
1995, Elliott 162.
Epilobium leptophyllum Raf. (Onagraceae). Epilobium leptophyllum is an her-
baceous perennial found in fens, marshes, and seeps throughout the northern
and central Great Plains (Great Plains Flora Association 1991), occurring south-
ward to Meade County, Kansas (Great Plains Flora Association 1977). A small
population discovered in extreme western Oklahoma, on a seepy, open slope
near the Ogallala-Doxey contact is 180 km south-southeast of the southern-
most Kansas population.
—_—
Voucher specimen: OKLAHOMA. Roger Mills Co.: Cheyenne, 3.5 mi S, 4 mi W, Thurmond Ranch,
T12N, R24W, sec 4, NE !/4, 35°32'51'N, 99°44'27"W, elev 2,260-2,300 ft, gently rolling sandsage prai-
FREEMAN ET AL., NEW PLANT RECORDS OF CENTRAL NORTH AMERICA 1293
rie and mixed-grass prairie with abundant oak mottes, small pond at upper reaches of Sergeant Ma-
jor Creek, Tertiary Ogallala Formation, 3 Oct 2001, Freeman 18359 & Morse
Erigeron canus A. Gray (Asteraceae). Hoary fleabane grows on rocky, shortgrass
prairies in the High Plains from South Dakota and Wyoming, south to New
Mexico. Examination of specimens at KANU revealed plants misidentified as
E. pumilus Nutt. including six from extreme northwestern Kansas, which rep-
resent the first verified records of the species from the state.
Voucher specimens: KANSAS. Cheyenne Co.: St. Francis, 12 mi N,12 mi W, upland prairie pasture, top
of sandrock ledge, thin soil, 9 Jun 1967, Stephens 11449 & Brooks, St. Francis, 12 mi N, 12 mi W, high
prairie, hilltop, dry, rocky, gravel soil, 4 Jun 1969, Stephens 31383 & Brooks, St. Francis, 12 mi N, 12 mi
W, . of rocky, prairie hill, 5 Jun 1981, Stephens 90302. Rawlins Co.: Atwood, 8 mi SW, dry rocky
pra idge, 27 Jun 1954, McGregor 9449, Beardsley, 3.5 mi §, ae a hillside, 22 May 1975,
a pi McDonald, ca 1] mi S, 6 mi E, T5S, R35W, sec 7, E!/2 & 8, W!/2, elev 3,200-3,300 ft,
N-S trending canyon that joins Beaver Creek to the N, ae on shortgrass prairie, extensive
outcrops of Ogallala Formation in the area, 30 May 1991, Freeman & Brooks 3516.
*Geranium dissectum L. (Geraniaceae). Geranium dissectum is a native of Eu-
rope now established throughout the eastern and western U.S. and Canada
(Gleason & Cronquist 1991; Kartesz @ Meacham 1999). A single record confirms
its presence in Kansas.
Voucher specimen: KANSAS. Labette Co.: Parsons, E side, open grassy area by Labette Creek along US
160, 7 May 1998, Holland 9312.
*Geranium molle L. (Geraniaceae). Dove-foot crane’s-bill is a native of Europe
and western Asia (Gleason and Cronquist 1991). It is now a widespread weed in
the eastern and western USS. (Kartesz & Meacham 1999), but records from the
Great Plains are rare. A specimen recently was found at KANU among 80 speci-
mens of G. pusillum L. and is the only record for Kansas.
Voucher specimen: KANSAS. Wilson Co.: Fredonia, 6 mi N, 4 mi W, T285, RI4E, sec 7, SE!/4, SE!/4,
near old farm house it b oak woods, 12 Jun 1995, McGregor 41124.
*Gleditsia triacanthos L. (Fabaceae). Common honey-locust is native to the east-
ern US., apparently occurring naturally westward to central Nebraska and
Kansas (Great Plains Flora Association 1977, 1991). However, its native range in
the Great Plains has been obscured by its long history of cultivation as a shade
tree and in windbreaks and fencerows. The species was not included in the
Colorado flora by Weber and Wittmann (1992), but KANU now has two speci-
mens of this tree from the eastern part of the state. Both records are presumably
the result of introductions in recent decades.
Voucher specimens: COLORADO. Kit Carson Co.: Flagler, 3.5 mi E, Flagler State Wildlife Area, elev
4,710-4,770 ft, rocky, shortgrass prairie on the W side of Flagler Lake, 14 May 2000, Freeman 14319.
Yuma Co.: Beecher Island, 3.5 mi S, dry prairie hillside pasture, 18 Jun 1906, Stephens 4871
ry
*Hibiscus syriacus L. (Malvaceae). A native of east Asia, Rose-of-Sharon is widely
planted in the US, occasionally escaping from cultivation (Gleason & Cronquist
1991). Kartesz and Meacham (1999) mapped records from throughout the east-
1294 BRIT.ORG/SIDA 20(3)
ern US. and in Utah. Steyermark (1977) reported the species as rarely escaping
in Missouri. In the Great Plains, H. syriacus often persists after cultivation, but
escaped plants have not been recorded in our region. A plant discovered in east-
ern Kansas appears not to have been planted.
Voucher specimen: KANSAS. Douglas Co.: Lawrence, T13S, RIDE, sec 12, NW !/4, NW !/4, NE
38°56'30"N, 95 15'01"W, elev 255 m, weedy forest edge along Naismith Creek, 7 Sep 2001, Morse 73 oo
Oenothera engelmannii (Small) Munz (Onagraceae). Harrington (1954) included
Engelmann’s evening-primrose in his flora of Colorado; however, Weber and
Wittmann (1992) excluded it because the voucher for the report was
misidentified. A specimen at KANU confirms that the species occurs in Colo-
rado.
Voucher specimen: COLORADO. Baca Co.: Ca 8 mi W of Elkhart, KS, on the W side of Morton County
Rd B along the KS-CO border, $ of the Cimarron River, T35S, R41W, sec 10, NE 1/4 SE 1/4, elev 3,610-
3,620 ft, grazed, upland, rolling, sandsage prairie, soil sandy, 2 Jun 1989, Freeman 3023.
Oenothera laciniata Hill (Onagraceae). Weber and Wittmann (1992) stated that
cut-leaf evening-primrose must occur in eastern Colorado based on nearby col-
lections from Kansas, Nebraska, and Oklahoma. This species may be added to
the Colorado list based on a collection from the extreme southeast part of that
state.
Voucher specimen: COLORADO. Baca Co.: Ca 8 mi W of Elkhart, KS,on the W side of Morton County
Rd Balong the KS-CO border, S of the Cimarron River, T35S, R41 W, sec 10, NE 74 SE!/4, elev 3,610-
3,620 ft, grazec
upland, rolling, sandsage prairie, soil sandy, 2 Jun 1989, Freeman 3024.
*Phyllanthus urinaria L. subsp. urinaria (Euphorbiaceae). A native of tropical
east Asia, peewater leaf-flower was first documented i in North America in 1944
(Webster 1970). The species is now established in the southeastern U.S. and IIli-
nois (Kartesz @ Meacham 1999), where it is probably more common than her-
barium records suggest (Sundell 1986). Two recent collections confirm the pres-
ence of this species in southeast Kansas.
Voucher specimens: KANSAS. Neosho Co.: Chanute, S edge in Holiday Park, 29 Sep 1997, Holland
9286; St. Paul, near new Farmers National Bank, 2 Oct 1997, Holland 9289.
*Pyrus calleryana Decne. (Rosaceae). A native of China, Bradford pear is widely
cultivated as an ornamental tree in the US. Kartesz and Meacham (1999)
mapped records from more than a dozen states in the central, southeast, and
south-central U.S. Cultivated plants often persist after abandonment in the
Great Plains, and field observations suggest that the species has escaped in Kan-
sas more frequently than herbarium records indicate. A collection from the east-
ern part of the state comes froma tree that appears not to have been under prior
cultivation.
Voucher specimen: KANSAS. Douglas Co.: Lawrence, SE edge. Baker Wetlands, just W pull-in on N
side of site, T135, R20E, sec 18, NE!/4, NW !/4, 38°55'40"N, 95°14'00"W, elev 250 m, weedy woodland
along edge of wetland, 16 Nov 2000, Morse 5289, Freeman & Loring.
FREEMAN ET AL., NEW PLANT RECORDS OF CENTRAL NORTH AMERICA 1295
*Sagina apetala Ard. (Car yopl 1vilaceae) Crow (1978) mapped this diminutive,
introduced annual from along the west coast of the U.S. and at several widely
scattered sites in the southeast. Examination of KANU specimens recently re-
vealed two overlooked specimens of hairy-leaf pearl wort from Kansas.
Voucher sj KANSAS. Leavenworth Co.: US 24 highway at Basehor, sandy, gravelly soil around
roadside pull-off, exposed, + May 1983, Brooks 16545; US 24 at Basehor, roadside pull-olf area, grav-
elly disturbed site, 8 Jun 1983, Brooks 16638.
*Sedum sarmentosum Bunge (Crassulaceae). Stringy stonecrop is a succulent
ornamental native to China (Gleason & Cronquist 1991). Escaped plants have
been documented in most states east of the Great Plains (Kartesz & Meacham
1999). A small population recently discovered in eastern Kansas is the first
record for the state.
Voucher specimen: KANSAS. Miami Co.: Beagle, 2.5 mi N, 2 mi FE, T18, R22E, sec 25, 1/2, NE !/4,
38°27'22'N, 94'15'15"W, elev 275 m, grazed woodland along trail to S end of tallgrass prairie site, 28
May 2001, Morse 5856 & Michener.
Silphium radula Nutt. (Asteraceae). A member of the taxonomically difficult
Silphium integrifolium Michx. complex, rough-stem rosinweed is reported from
scattered populations throughout the southeastern US. (Kartesz @ Meacham
1999). Previously reported in the Great Plains from Oklahoma and Texas (Great
Plains Flora Association 1991), this species has been confirmed in southeast
Kansas based ona single collection.
Voucher specimen: KANSAS. Neosho Co.: Erie, 0.25 miS,5 mi E, along old MKT RR right of way, 17 Jul
2000, Holland 9959.
*Trianthema portulacastrum L. (Aizoaceac). Desert horse-purslane is an intro-
duced species known from widely scattered collections in the southern Great
Plains (Great Plains Flora Association 1977, 1991). Three collections were made
in 2000 in Kiowa County, Colorado, representing the first records for that state.
Voucher specimens: COLORADO. Kiowa Co.: Eads, 8.5 mi S, 4 mi E, Neenoshe Reservoir, NW side,
T20S, R47W, sec 6, NE!/4, elev 3,930 ft, shortgrass a airie immediately W of the reservoir and open
to wooded shoreline, 16 May 2000, Freeman 14545; Eads, 10 mi S, 4.5 mi E, Neenoshe Reservoir, S side,
T2058, R47W, sec 9, SW !/4, SW !/4 & sec 16,N!/2 ee ess open, sandy to muddy shoreline and
adjacent, wooded perimeter of reservoir, . a 2000, Freeman eae cee Eads, 8.5 mi S, 4 miE,
Neenoshe Reservoir, NW side, T20S, R47W, sec 6, NE !/4, elev 3,930 ft, shortgrass prairie immedi-
ately W of the reservoir and open to wooded s vee 20 Aug ee Fr reeman 16037 & Morse.
Tridens flavus (L.) Hitchc. var. chapmanii (Small) Shinners. (Poaceae). Tridens
flavus is broadly distributed throughout the eastern U.S., westward through
Wisconsin and southward to New Mexico. Variety flavus occurs in the Great
Plains from southeastern Nebraska, southward through eastern Kansas and into
Oklahoma (Great Plains Flora Association 1977). Variety chapmanii has been
reported from Oklahoma (Kartesz &@ Meachem 1999). Recent examination of
specimens from southern Kansas revealed that this variety also occurs in the
state.
1296 BRIT.ORG/SIDA 20(3)
Voucher specimens: KANSAS, Chautauqua Co.: Sedan, 1.5 mi NE, open blackjack oak woods, hillside,
dry rocky sandstone soil, 18 Oct 1969, Stephens 37321, Chautauqua, | mi S, 1.5 mi E, low woods along
small stream, soil sandy, moist, 14 Oct 1976, Brooks 12832. Cowley Co.: Winfield, 7 mi W, sandy prai-
rie, 3 Sep 1966, Koch 2100.
*Veronica persica Poir. (Scrophulariaceae). Though bird-eye speed well was over-
looked in the Flora of the Great Plains (Great Plains Flora Association 1991),
Freeman et al. (1998) reported this introduced annual from lowa, Kansas, Ne-
braska, South Dakota, and Wyoming in the Great Plains. According to Weber
and Wittmann (1992), previous reports of V. persica in Colorado were based on
misidentified specimens of V. polita Fr. However, a recent collection in eastern
Colorado confirms the presence of this species in the state.
Voucher sme COLORADO. Cheyenne Co.: Cheyenne Wells, S side of town, T145, R44W, sec 20,
ae elev 4, eee rest area along US 40 and adjacent Cheyenne County Fairgrounds, well-
li f ball diamond, 2 Oct 2000, Freeman 16705.
Wi ee
ACKNOWLEDGMENTS
We thank Carolyn Ferguson (KSC) for providing access to specimens at the
Kansas State University Herbarium and Anton A. Reznicek (MICH) for con-
firming our determination of Carex swanii. David Sutherland and an anony-
mous reviewer provided valuable comments about the manuscript. Support for
2000 fieldwork in Colorado was provided by a grant from the Colorado Natu-
ral Areas Program. Other studies were funded in part by the Kansas Biological
Survey and the University of Kansas Natural History Museum and Biodiversity
Research Center.
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Bulbostylis, Cyperus, Dulicium, Eleocharis, Eriophorum, Fimbristylis, Fuirena, Lipocarpha,and
Scirpus. Trans. Nebraska Acad. Sci. 22:27-42.
Rotrsmeier, 5.B.and B.Witson. 1997. Keys and distributional maps for Nebraska Cyperaceae,
Part 2: Carex and Scleria. Trans. Nebraska Acad. Sci. 24:5-26.
STEYERMARK, J.A. 1977. Flora of Missouri. 5th printing. lowa State University Press, Ames.
SUNDELL, E. 1986. Noteworthy vascular plants from Arkansas. Castanea 51:211-215.
Tayior, R.J.and C.E.S. Tayor. 1994. An annotated list of the ferns, fern allies, gymnosperms
and flowering plants of Oklahoma. Bio. Dept. Herbarium, Southeastern Oklahoma State
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Weeer, W.A. and R.C. Wittmann. 1992. Catalog of the Colorado flora: a biodiversity baseline.
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/www.Colorado.edu/CUMUSEUM/ research/botany/Catalog/Catalog.htm]
Weester, G.L. 1970. A revision of Phyllanthus (Euphorbiaceae) in the continental United
States. Brittonia 22:44-76.
YATsKIEVYCH, G. 1999. Steyermark’s flora of Missouri. Missouri Dept. of Conservation, Jeffer-
son City.
—=
1298 BRIT.ORG/SIDA 20(3)
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SIDA 20(3): 1298. 2003
NEW RECORDS OF CORCHORUS (TILIACEAE) FOR MEXICO
José Aurelio Colmenero Robles' Rafael Fernandez Nava!”
ordinacion General de Posgrado e Investigacion Laboratorio de Bot Fanerogamica
sae lité N / Escuela Naciona aeGeng 5 Bio oligicas
partado Postal 118-331 Instituto Polit
a : Villa, 07051, MEXICO, D.F. Prolongacién de Carpio y Plan de Ayala
Col. Santo Tomds, 11340, MEXICO, DF.
ABSTRACT
Two species of Corchorus (Tiliaceae) are reported for Mexico: Corchorus olitorius L. of Asian origin
and C. trilocularis L., native of the tropics of Africa and Asia. The first species was introduced into
Brazil and the Antilles toward the end of the 19th century and is now recorded for Mexico from the
state of Sinaloa. The second species is known from Veracruz and Chiapas, Mexico, and as well as
Panama and Ecuador. The exchange and trade of seeds from other continents could be an important
way for their introduction and that might explain the presence of these two species in Mexico. At
present SIX species s of Corchorus are known to occur in Mexico. A dichotomous key for the identifica-
tion of the cae papecies of the genus is provided, as well as descriptions of the two species, a map
of these species | in America, and their vet nacular names.
Oo oO [
Key worps: Corchorus olitorius, C. trilocularis, aloctonous species, Tiliaceae, Mexico.
RESUMEN
Sed | 31 de especies alc lel genero Corchorus (Tiliaceae): Corchorus
olitorius L. de origen asiatico y C. trilocularis L., nativa de los tropicos de Africa y Asia. La primera
especie fue introducido a sas yalas Ant a desde finales del siglo 19, en nuestro pais se reporta
por primera vez del loa; la segunda especie se registra de Veracruz 7 Chiapas, México,
ademas de Ecuador y Panama, se desconoce lak fecha exacta de su arribo al continente americano. Se
propone que el intercambio y comercio de E i de ot t { puede ser una
via importante para su introduccion, lo que podria ex pli n México. En la actualidad
son seis las especies reconocidas de este genero para México. Se ae aa as una clave dicotomica
pa epee Ce el género Corchorus de México, asi como la descripci species, un mapa
geografica para estas especies en América y sus nombres comunes.
Patasras CLAVE: Corchorus olitorius, C. trilocularis, especies aloctonas, Tiliaceae, México.
INTRODUCTION
The genus Corchorus (Tiliaceae) was originally described by Linnaeus in 1753,
who included four species in the genus: Corchorus capsularis L., C. hirsutus L.,
C.olitorius L., and C. siliquosus L. At the present time the genus is considered
pantropical with approximately +0 species, with about 10 species in the New
World (Howard 1989). Corchorus olitorius and C. capsularis, of Asian-African
Fellowship COFFA, IPN
Fellowship EDD, IPN
SIDA 20(3): 1299-1309. 2003
1300 BRIT.ORG/SIDA 20(3)
origin, have been cultivated since the XIX century in some tropical regions of
America for their well known jute fiber.
In Mexico, Standley (1923) registers the presence of three species: Corchorus
hirtus L., C. orinocensis Kunth, and C. siliquosus L.; later Standley and
Steyermark (1949) recognize Corchorus aestuans L. for Mexico and South
America; Fryxell (2001) recognize: C. destuans, C. siliquosos and C. hirtus for the
Flora Novo-Galiciana. In the present work C. olitorius and C. trilocularis L. are
added to the previous list, resulting in six species presently reported for Mexico.
The genus Corchorus is characterized as follows: annual or perennial herbs,
subshrubs, or shrubs, with simple, stellate, or stellate-dendritic hairs or gla-
brous; leaves alternate, simple, with stipules, persistent or deciduous; blades
dentate, serrate or crenate, occasionally with two teeth prolonged into long se-
tae at base; inflorescences lateral or leaf-opposed, bracteate, shortly peduncu-
late, I-to several-flowered umbellate or racemose cymes; flowers hermaphro-
ditic; calyx valvate, sepals 4-5, generally free, often apiculate or caudate at apex;
petals 4-5, yellow, free, usually shortly unguiculate, with an inconspicuous glan-
dular patch at the base of each petal; androgynophore usually present; stamens
4 to many, free, filaments free or connate; anthers with two theca, introrse,
dorsifixed; ovary 2-tol0-locular, ovules 2 to many per locule, placentation ax-
ile, parietal, or pendulous; style terete, papillate; capsules cylindrical to
subglobose, muricate, glabrous, glabrescent, or pubescent, 2- to 10-valved, valves
sometimes with transverse septae adaxially; seeds many and small.
METHODS
Specimens of Corchorus were studied inthe herbaria CHAPA, ENCB, MEXU, UAS,
and XAL. Specialized works on the genus were reviewed, with the morphological
characteristics of stem, leaves and fruits being compared with the respective de-
scriptions in the works of Martinez (1981), Edmons (1990), and Halford (1995).
For the particular case of C.trilocularis, the pubescence of the fruit was observed
in the compound microscope. The specimens were also compared with the
phototypes of the herbarium S-LINN, deposited in the ENCB herbarium. Syn-
onymies and vernacular names were taken from literature sources.
RESULTS
Two species of Corchorusare added to the family Tiliaceae in Mexico, C.olitorius
and C. trilocularis. The first one is native of the region of India and Malaysia
and is widely cultivated for its fiber, well known as jute. Corchorus trilocularis
is native to the region of Africa and Asia and is less important from a economic
point of view, not being acommercial crop. A dichotomous key for the identifi-
cation of all Mexican species of Corchorus is included below, followed by mor-
phological descriptions, uses, and specimens examined for each of the two newly
reported species, as well as a distribution map (Fig. 1).
ps
COLMENERO AND FERNANDEZ, NEW RECORDS OF CORCHORUS FOR MEXICO 1301
KEY TO THE SPECIES OF CORCHORUS IN MEXICO
1. Subligneous more than 1 m tall; ovary with 5-6 celled; capsules with 5-6 Corchorus
olitorius
Herbs smaller than 1 m tall; ovary with 2-4 celled; capsules with 2-4 valves.
Capsules 2 valved; ovary 2 celle
3. Shrubs; branchlets subglabrous; leaves glabrous below; ovary puberulent;
capsules aplanate with the apex obtuse and with 1-4 minute apical beaks ___- Corchorus
siliquosus
3. Herbs; branchlets densely tomentose and hirsute; leaves hirsute or pilose
below, ovary densely sericeous; capsules teretes without apical beaks.
4. Capsules hirsute with trichomes conspicuous ca. 1.5 mm long, lightly
curved; stipules and leaves hirsute; calyx hirsute; stamen 40; ovary hispid
Corchorus hirtus
4. Capsules glabrous or with BPIGHOINS INSOTSpletous: ca. 1 mm de largo,
right; stipules and leave lyg : g 0-
0; ovary appressed pubescent
2. Capsules 3-4 valved; ovary 3-4 celled.
5. Leaves mostly ovate, slightly cordate, base rounded and sometimes with a
pair of teeth prolonged; stamens 9-14, capsule oblongoid with 6-8 winged
and 3 or 4 apical bifid beak divergent and recurved Corchorus aestuans
ear, base obtuse or broadly cuneate never wit
Corchorus orinocensis
5. Leaves narro ‘ly ovate or line
pair of teeth prolonged, stamens 20-30, capsule torulose without win ue
and only one apex straight orchorus trilocularis
simak olitorius L. Sp. Pl. 529.1753. cache 1, 2). Type: ers eee Herb. Clif-
209, Corchortus no. 1, (LECTOTYPE: BM). Lectotype desig dsFl: Ftahes, 2:84.
ee
Corchorus quinquelocularis Moench, Méth. 248. 1794.
Corchorus decemangularis Roxb., Hort. Beng. 42. 1814.
Corchorus lanceolatus G. Don, Gen. Syst. 1:543. 1831.
Corchorus longicarpus G. Don, Gen. Syt. 1:543. 1831.
Corchorus cartharticus Blanco, Fl. Filip. Ed. 1:442. 183
Gorehors malchairi De Wild., Etudes Fl. Bangala & Chand 345. 1911.
ius var. australiensis Domin, Biblio. Bot. 89:380. 1927
Vernacular names inother countries, taken from the literature.—bunani waweza,
eteke nyamusiri, niimbaswa-lusali, otigu-kal, pombo mgunda (Africa); yute
(Argentina), gramigrain (Bahamas); yute, gringuele (Cuba); bogi, koshta, jute,
chehuncho, changhas, rajaan, tossa jutee (India); feuilles lalo, lalo, petit lalo, ti-
la-lo (Dominican Republic-Haiti).
Subligneous herb to (1-)2-4 m tall; branchlets glabrous, somewhat angu-
lar or sulcate when young, stipules 5-16 mm long, glabrous; leaves with peti-
oles terete, thin, (1-)3-4(-60) mm long, glabrous except for simple hairs on a
margin; stipules linear, (7-)10(-12) mm long, pilose or glabrous; blades narrowly
ovate to ovate, ovate-lanceolate, (1.5-)10(-12) cm long and (1-)2-5 cm wide, up-
per and lower surfaces usually glabrous, veins sometimes pilose, base rounded
and with two basal setaceous teeth to 10(-15) mm long, margin serrate to
crenate-serrate, apex acute to acuminate; inflorescences leaf-opposed, solitary
BRIT.ORG/SIDA 20(3)
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COLMENERO AND FERNANDEZ, NEW RECORDS OF CORCHORUS FOR MEXICO 1303
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1304 BRIT.ORG/SIDA 20(3)
at the nodes, geminate or ternate, pedicels very short, 1-2 mm long, glabrous,
bracts (1-)3-5 mm long, pilose o glabrous; sepals valvate, 7 mm long, linear-
lanceolate, concave and glabrous; petals obovate, imbricate, yellow, same length
as sepals, | mm long and 2-3 mm wide, glabrous but minutely ciliate on mar-
gin; androgynophore 2-5 mm long; annulus 5-lobed; stamens 30-50, borne in-
side of annulus; filaments filiform, 3-4 mm long; anthers dorsifixed, oblong;
ovary cylindrical, 5-6 celled, 10-sulcate, minutely setose and glandular, with
36-42 axial ovules per cell; style stout, glabrous, 1-2 mm long, stigma lobed,
fimbriate; capsule cylindrical, 3-)4-6(-8) cm long and (3-)4-5(-6) mm wide,
appressed to stem, sometimes torulose, dehiscent, 5-6 valvate, transversely sep-
tate, apex 3 to 4(-10) mm long; seeds more or less rhomboid, many, black, 1-3
mm (Fig. 2).
Specimens examined: MEXICO. Sinaloa: a 2 Km del ejido La Campana, km 25 carr. Culiacan
Guamuchil, municipio de Culiacan, Leobardo Chavez Montes y Fco. Javier Niebla Armenta 124 (UAS);
carr. Culiacan-Los Mochis, Km 19, al NE hacia el Cerro La Chiva, J. A. Prado V. y V. Llamas A. 201
UAS). pie del Cerro El Tule, lado W, orilla de la brechaa la torre de microondas, ca. 5 km de Culiacan,
0 Cebreros B.
Sy.
municipio de Culiacan, Rito Vega. 7295, Hipélito Aguiar H., Cenén Aguirre C. y Ricar
UAS), carretera internacional a Costa Rica Km 5, municipio de Culiacan, Salomé Ochoa Avalos y
- rman A. Bojorquez B. 162 (MEXU),; El Camichin, a 3 ae al aaa de ns ue de los Reyes,
nna de Cosala, Rito Vega 3487, German Bojorquez y Alejandro | z APA, UAS). 5 mi
of Mazatlan, D. Bates, O. tee PF ryxell 1543 (ENCB).
oO
This species is originally from India and Malaysia, with pantropical distribu-
tion where it is appreciated for its fiber Gute).
Corchorus olitorius can be distinguished from the rest of the species in
Mexico by being a semishrub 1-2 m, with the stem and leaves glabrous and the
inferior border of the leaves with two very long appendices, as well as by the
linear capsule, which is thick and glabrous with five valves. It is known from
the Pacific Coastal Plain, in the state of Sinaloa, where it grows on the edges of
cultivated fields, the banks of roads, in tropical deciduous forest and in thorn
forest, on alluvial soils and thin and stony soils, at altitudes that vary between
(30-)50 and 700 m. Its presence in America has been reported since the latter
part of the XIX century in most of the region of the Caribbean and Brazil
(Ricardo et al. 1995; Schumann 1886).
From very early times Corchorus olitorius was cultivated as edible in Africa
and the Middle East, while as a textile plant it was broadly domesticated in
India (Simmonds 1979). Well known as jute, it is presently cultivated in the west
of Bengal, Bangladesh, Burma, Nepal, China, Malaysia, and Sri Lanka (Maiti
1995). With the same objective it was introduced into Brazil and Cuba at the
end of the 19th century, and in the latter country it has become naturalized in
the western part (Schumann 1886; Ricardo et al. 1995; Rodriguez 2000). In Ar-
gentina its presence may be more recent (Dimitri & Rial 1952).
This species occurs in Africa in grassland and savannas, and diverse uses
are attributed to it there, such as paper production and the extraction of indus-
pai
COLMENERO AND FERNANDEZ, NEW RECORDS OF CORCHORUS FOR MEXICO 1305
trial alcohol (Irvine 1961; Edmons 1990). In India, the seeds are used as a purga-
tive and an infusion of the leaves is used as a tonic (Bhandari 1978).
Corchorus trilocularis L. Syst. Nat. ed. 12, 2:369.1767. (Figs. 1, 3). Typr: protologue
ARABIA: Forsskdl s.n. (LINN 691.2) type info. fide Flora of Tropical East Africa (2001).
Corchorus aestuans Forssk, Fl. Aegypt. ee 101.1775. non L.
Corchorus serraefolius DC, Prodr. 1:504
Coneniacu siecle: ee Flora 20: it ee
ort. Maurit. 43. 1837
Coichovusasplentenis® non Burch. ex Harv. & Sond., Fl. Cap. 1:229. 1800.
Corchorus somalicus Gandoger, Bull. Soc. Bot. France 69:348 1922
Corchorus rigidiusculus Domin, Biblioth. Bot. 89:381.1926
Vernacular names in other countries, taken from the literature.—apoik, lumere,
teke (Africa); hardikaket, kagle-ki-tambaku, karak (India).
Herbs, procumbent to ascending, up 50 cm tall; branchlets moderately cov-
ered with pilose pubescence; leaves simple, petioles 3-20 mm long with simple
hairs, stipules 2-3(-7) mm long; blades narrowly ovate or linear, (2.5-)4-9(-13)
cm long and 0.6-3.0 cm wide, base obtuse or broadly cuneate never with a pair
of teeth prolonged, margin dentate or crenate, sometimes with a pair of basal
teeth upwardly prolonged 5(-10) mm, apex obtuse or acute, upper surface with
scattered simple hairs, lower surface with simple hairs frequently on midvein
and secondary veins; inflorescence leaf-opposed, solitary at nodes, geminate or
ternate, peduncles and pedicels 1-2(-4) mm long, bracts up 33 mm long, cili-
ate; sepals 5, linear, (4-)5.5-(-6.3) mm long and 10 mm wide, pilose outside, con-
stricted toward base; petals 5, obovate, (4-)5-(6-)7 mm long and 20-25 mm wide,
minutely ciliate on margin; stamens 20-30; ovary cylindrical, 3-locular, 3 mm
long, shortly pubescent, stigma fimbriate; capsule torulose without winged and
only one apex straight, (2.5-)3.0-5.5(-7) cm long and (1-)2.5-3 mm wide, 3-val-
vate, pubescence of short simple and fasciculate or palmate hairs, the latter not
perceptible without magnification, at maturity more or less glabrous, apex
straight, acuminate, valves partially transversely septate adaxially; seeds more
or less rhomboid, dark brown to black, 0.8-1 mm diameter.
Specimens examined: MEXICO. Chiapas: Mirador for Chicoasen Dam along road to Tuxtla Gutierrez
to the Chicoasen, Municipio San Fernando, Breedlove 41494 (MEXU). Veracruz: Barranca de
Pachuquilla, a 2 km al SO de dicha poblacion, municipio de Puente Nacional, M. E. Medina A. y M
Ortiz D.702 (XAL),; Paso de la Milpa, municipio de Actopan, F. Ventura A. 18001 (ENCB, XAL); Cerro
del Brujo, 3 km al N de Jacomulco, municipio de Jalcomulco, G. Castillo Campos y P. Zamora C. 8757
(XAL); AINW de Jalcomulco, entre qu apan y Jal conmred municipio de Jalcomulco, G. Castillo-
Campos, et al. 8347 (XAL); El Hatito El Hatito-Cano Prieto, Bee de Paso
de Ovejas, J. E. Gonzdlez H. 184 (XAL); Mata de Cana, municipio de Puente Nacional, F Ventura A.
10706 (ENCB). PANAMA. Provincia Los Santos: Las Trancas-Guararé, Alcibiades Veldzques 24(MEXU ).
Corchorus trilocularis resembles C. orinocensis in its herbaceous habit and the
size of the leaves. However, C. trilocularis is distinguished by the presence of
small sepals and petals and the fimbriate stigma; by the ovate-lanceolate leaf
1306 BRIT.ORG/SIDA 20(3)
SAR RN
a
ti
1 fruits: B. Mature flower, seen laterally: C. Lateral view of
1 y
. pos a
the fruit; D. Seeds. (based on A. Veldzquez 24)
COLMENERO AND FERNANDEZ, NEW RECORDS OF CORCHORUS FOR MEXICO 1307
blade Clinear-lanceolate in C. orinocensis), and, most remarkably, by its three-
valved fruits with pubescence of simple and fasciculate or palmate hairs, the
latter extremely small and imperceptible at first look (Fig. 3),as opposed to the
two-valved fruits with only simple hairs found in C. orinocensis.
The presence of Corchorus trilocularis in America has not been established
previously with certainty, because specimens with two or three-valved fruits
have been identified as Corchorus orinocensis. In the earlier works of Macbride
(1956) for Peru and Robyns (1964) for Panama, the descriptions of Corchorus
orinocensis correspond to three-valved material.
In the revision of the Neotropical species of the genus Corchorus, Tiliaceae,
Martinez (1981) assigned Ecuadorian specimens to Corchorus trilocularis.
In the revision of the family Tiliacae for the Flora of the Guyanas, Jansen
and Meijer (1995) have concluded that only the two-valved specimens should
be assigned to C. orinocensis based on Kunth’s 1823 description.
Corchorus trilocularis is widely distributed in Africa, Asia, and Australia,
including in the xerophytic regions of Arabia and India (Bhandari 1978; Edmons
1990). At present there is no clear indication of the date of arrival in the Ameri-
cas. In Mexico it has been collected in the states of Chiapas and Veracruz, in
disturbed areas of deciduous and semi-deciduous tropical forest andin Panama
in tropical rain forest.
Although Corchorus trilocularis is of little economic importance, it is used
at the domestic level; for example, in Africa, the leaves are used in traditional
medicine to reduce swellings; an infusion of leaves and branchs is used to pro-
duce an alcoholic drink; the seeds are used for certain stomach disorders and to
induce vomiting; and it is used as an ornamental plant and as food for livestock
(Edmons 1990).
CONCLUSIONS
With these two new reports of Corchorus, the number of species known for
Mexico rises to six. Corchorus olitorius is presently known only from Sinaloa,
although its presence in the region of the Caribbean and South America as a
useful plant has long been known. Its origin in the Indo-Malay region and its
discovery in the Pacific coastal plain in Mexico leads to the supposition that
trade of agricultural products, mainly seeds with Oriental countries, has been
the route of introduction of the species into Mexico.
In Mexico Corchorus trilocularis behaves as weed adapted to the disturbed
conditions in southern Mexico, in dry and sub-humid warm vegetation types
and in cloud mountain forest. In the Old World it isa characteristic species of the
arid zones of Africa, Arabia, and India, in areas of cultivations, grasslands, savan-
nas, and xerophyllous shrubs, while it is also present in the warm-humid zones
of India and Australia. Its presence in the Ecuador and part of Central America
leaves unanswered questions about this apparently disjunct distribution.
1308 BRIT.ORG/SIDA 20(3)
In the specimens of C.trilocularis collected in Mexico and Central America,
we observe that the pubescence of the mature capsule is of simple and fascicu-
late trichomes similar to specimens collected in Australia (Halford 1995). Speci-
mens collected in Africa present mature capsules with simple and palmate tri-
chomes (Edmons 1990).
It seems likely that more intensive collection in Mexico, Mesoamerica, and
South America will lead to a more definite hypothesis as to the manner and
route of introduction of the species into the New World, including whether the
introduction is recent or ancient.
ACKNOWLEDGMENTS
The authors thank Pedro Leyva for the specimen drawing, and curators of the
herbaria cited here for access to their valuable collections. The first author
thanks COTEPABE, IPN (Project CGPI S99001) for financial support to visit the
Missouri Botanical Garden herbarium in August 2000.
REFERENCES
BHANpARI, M.N. 1978. Flora of the Indian Desert. Edit. Pawan Kumar Sharma.
Dimitri, MJ.y F.Riat A. 1952. Las plantas cultivadas en la Argentina. Tiliaceas. Ministerio de
Agricultura y Ganaderia, vol. 7. Fasc. 123.
Epmons, J.M. 1990. Herbarium survey of African Corchorus L. species. Systematic and
Ecogeographic Studies on Crop Genepools 4. International Board for Plant Genetic
Resources, Rome. Italy
Fryxet, PA. 2001. Tiliaceae. In: Flora Novo-Galiciana 3:68-109. The University of Michigan,
Ann Arbor, U.S.A.
Guat, M.D. 1998.La familia Tillaceae Juss.en el estado de Guerrero. México. Tésis de Maestria.
Facultad de Ciencias. Division de Estudios de Posgrado. Universidad Nacional Aut6énoma
de México. México, D.F
Hatrorb, D.A. 1995. Notes on Tiliaceae in Australia, 2. A revision of the simple-haired spe-
cies of the genus Corchorus L. Austrobaileya 4:297-320.
Howarb R.A. 1989. Tiliaceae. In: Flora of the Lesser Antilles, Leeward and Windward Islands.
5:184-199. Arnold Arboretum, Harvard University, Cambridge,U.S.A
Irvine, E.R. 1961.Woody plants of Ghana, with special reference to their uses. Oxford Uni-
versity Press.
JANSEN, M.J.J. and W. Meuer. 1995, Tiliaceae. In: Flora of the Guianas. Ser. A. Phanerogams.
Koeltz Scientific Books. Konigstein, Germany.
Linnaeus, C. 1753. Species plantarum. A facsimile of first edition. Printed for The Ray Soci-
ety.London, England (1957). Vol. |.
Macerioe, J.F. 1956. Tiliaceae. In: Flora of Peru. Field Mus. Nat. Hist., Ser. Bot. 13(3A, 2):
413-442
Martinez, M.D. 1981.The Neotropical species of the genus Corchorus, Tiliaceae. Thesis, Uni-
versity of Kentucky. Lexington, Kentucky, U.S.A.
COLMENERO AND FERNANDEZ, NEW RECORDS OF CORCHORUS FOR MEXICO 1309
Mai, R.M. 1995. Fibras vegetales en el mundo.Aspectos botanicos, calidad y utilidad. Edit.
Trillas, México, D.F
RicarbDo, N.N.E., E. Pouyu R.,and PP. Herrera. 1995. The synanthropic flora of Cuba. Fontqueria
42:367-429.
Ropyns, A. 1964. Tiliaceae. In: Flora of Panama. Annals Missouri Bot. Gard. 51:1—35.
Ropricuez, F.A. 2000. Tiliaceae. Fasciculo (3) 5.Flora de la Republica de Cuba. Serie A.Plantas
Vasculares. Koeltz Scientific Books. Konigstein, Germany.
SCHUMANN, K. 1886. Tiliaceae. In: Martius, V.PF., and Eichler, A.G., eds. Flora Brasiliensis 12(3):
118-200.
Simmons, N.W. 1979. Evolution of crop plants. Longman. New York, U.S.A.
Stanbtey, PC. 1923. Tiliaceae. In: Trees and shrubs of Mexico. Contr. U.S. Natl. Herb. 23:
734-746.
STANDLEY, P.-C. and J.A. Stevermark. 1949. Tiliaceae. In: Flora of Guatemala. Fieldiana Bot. 24:
1310 BRIT.ORG/SIDA 20(3)
BOOKS
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Robert F BARNES, C. Jerry NELSON, MICHAEL COoLLins, and KENNETH J. Moore (Editors).
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Physiology of Forage Plants; 5) Environmental Aspects of Forage Management; 6) Grasses for North-
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dix; Common and Botanical Names of Forages; Glossary; Index.
RicHArb J. Sorre (Editor). 2003. The Agricultural Notebook, 20th Edition. (ISBN
0-632-05829-3, pbk.). Blackwell Science, Osney Mead, Oxtord OX2 OEL, UK
(Orders: lowa State Press, 2121 State Avenue, Ames, [A 50014, U.S.A: 1-800-
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SIDA 20(3): 1310. 2003
ADDITION TO FLORA MESOAMERICANA: A NEW RECORD
OF THELYPTERIS (THELYPTERIDACEAE)
FOR CHIAPAS, MEXICO
Miguel A. Pérez Farrera Ramon Riba
Escuela de Biologia UAM-lztapalapa, A.P 55-535
Universidad de Cienci {as Artes del Estado de Chiapas México, D.F. 09340
Tuxtla Gutiérrez, Chiapas, 29,000, MEXICO MEXICO
Maria E. irene
lodoPR
cid de Biologia
Universidad de Ciencias y Artes del Estado de Chiapas
AP. 782, Tuxtla Gutiérrez, Chiapas, 29,000, MEXICO
ABSTRACT
The
ypteris rhachiflexuosa Riba, described from Veracruz is recorded from Chiapas, where it was
lin North Highlands. Additional data ak listribution, habitat | phology are discussed.
RESUMEN
Thelypteris nna caalt exuosa Ribs, date a Veracruz, es registrado para las Montanas del Norte de
Chiap as. Se discute inf ibuci habitat y mor fologi: a.
INTRODUCTION
The genus Thelypteris comprises about 900 species—distributed around the
world—with high diversity in the tropical and subtropical regions. One hun-
dred thirty three (133) species have been recorded for Mesoamerica and about
50% of these are found in Chiapas (Smith 1995).
The Northern Highlands of Chiapas is one of the less well known areas from
a botanical viewpoint (Breedlove 1981). Nevertheless, a few studies have been car-
ried out in this area (Pérez & Sarukhan 1970; Lopez-Mendoza 1980), and it has
been considered one of the most important regions for endemic taxa (Schutzman
et al. 1988; Went 1989, 1998). Unfortunately it is also one of the most altered places,
due mainly to anthropogenic activities (cattle raising and sun coffee plantation).
This has resulted in loss of both primary vegetation and floristic diversity.
During a floristic exploration of the tropical forests in the Northern High-
lands in 1999, we collected several samples of ferns, among them a Thelypteris
with a flexuose rachis and 2-4 pairs of alternate, undulate pinnae.
This specimen proved to be Thelypteris rhachiflexuosa Riba, oy
known only from the region of Los Tuxtlas, Veracruz. This species was collected
and described by Riba (1989) and recorded as endemic to Veracruz (Ibarra-
Manriquez et al. 1997; Riba & Pérez-Garcia 1997; Riba 1998).
joel
SIDA 20(3): 1311-1315. 2003
1312 BRIT.ORG/SIDA 20(3)
Voucher specimen: MEXICO. Chiapas: Mpio. Amatan, Mt. Palmar, 10 km E of Amatan, Northern High-
lands, 23 Me vy 1999, 800 m, Pérez-Farrera 1967; (Herbarium of Escuela de B Biologia a, U ee 1; CHIP)
(Fig. 1) .Thelypteris rhachiflexuosa is being grown in the Botanical Garden “Faus la” in the
085.
Instituto de Historia Natural under accession number
In Chiapas this species occurs in “bosque tropical perennifolio” (tropical rain
forest) according to Rzedowski (1978), or “selva alta perennifolia” (Miranda &
Hernandez-X. 1963). This vegetation is comprised of three strata. The first stra-
tum is dominated by Brosimum alicastrum Sw., Dialium guidnense (Aubl.)
Sandw., Manilkara zapota (L.) Royen, Guatteria anomala R.E. Fries,
Calophyllum brasiliense Camb., Pouteria sapota Jacq.) H.E. Moore & Stearn,
and Pterocarpus hayensii Hemsl. In the middle stratum are usually Bursera
simaruba (L.) Sarg., Pimenta dioica (L.) Merrill, Miconia trinervia (Sw.) D. Don.
ex Loud., Oreopanax xalapensis (Kunth) Decné & PL, Trophis racemosa (L.) Ur-
ban, Saurauia belizensis Lundell, and Dendropanax arboreus (L.) Planch. &
Decne. The third stratum or understory typically consists of Astrocaryum
mexicanum Liebm., Chamaedorea concolor Martius, Chamaedorea elegans
Martius, Chamaedorea tepejilote Liebm., Chamaedorea arenbergiana Wendl.
Desmoncus chinantlensis Liebm. Begonia nelumbiifolia Cham. & Schlecht.,
Anthurium pedatoradiatum Schott, A. schlechtendalii Kunth. In addition, some
epiyphytes and hemiepiyphytes are found in these habitats, e.g., Monstera
deliciosa Liebm., Anthurium pentaphyllum (Aublet) G. Don. var. bombacifolium
(Schott) Madison, and Polypodium sp. Thelypteris rhachiflexuosa is sometimes
found in secondary vegetation or coffee plantations.
The soils in this area are red clay derived from lutitas (Lopez-Mendoza 1980).
The topography is abrupt, with steep slopes (70%). The landscape is karstic and
the bedrock consists of a complex of Eocene and Oligocene marine limestone
(Lopez-Hernandez 1994; Pérez & Sarukhan 1970). This species occurs at eleva-
tions from 600-800 m (Fig. 2).
DISCUSSION
Thelypteris rhachiflexuosa is in the subgenus Goniopteris. In Mesoamerica about
41 species are known in this subgenus, with about 34 species in Mexico
(Yatskievych, pers.comim.),and approximately 17 in Chiapas (Smith 1995). There
are still taxonomic problems at the species level.
This record for Chiapas extends the distribution of this species into
Mesoamérica. This taxon has a distribution pattern similar to that of other
woody species restricted to the Caribbean slope, an area including Los Tuxtlas,
Veracruz and northern mountains of Chiapas (Wendt 1998). We expect that T.
rhachiflexuosa will also be found in the State of Tabasco.
Thelypteris rhachiflexuosa is similar to T. oroniensis L.D. Gomez, endemic
to Costa Rica, in its zig-zag rachis, but differs from that species in its meniscioid
secondary veins. The specimen of T. rhachiflexuosa collected in Chiapas dif-
PEREZ ET AL., THELYPTERIS RHACHIFLEXUOSA NEW FOR CHIPAS, MEXICO 1313
1314 BRIT.ORG/SIDA 20(3)
[ o 20 40 60 80 0 120
=
Kilometers
Limites estatales |
_
@ T. rhachiflexuosa
TABASCO
8 00 ie
\
=
VERACRUZ ‘
OCEANO
PACIFICO
Fic. 2. Distribution of The/
7F r
fers slightly from the voucher collected in Veracruz in having more setulose
sori. Thelypteris rhachiflexuosa sometimes grows with Thelypterisghiesbreghtii
(Hook.) C.V. Morton and differs from this species with regard to straight rachis,
costae, veins and leaf tissue moderately to densely hairs below.
ACKNOWLEDGMENTS
The authors thank Jesus de la Cruz Rodriguez, Emerit Meléndez Lopez, Tomas
Acero Acero, Oscar Farrera Sarmiento, and Francisco Hernandez Najarro for
their help in the field work and processing of plants. Special thanks to Blanca
Pérez Garcia and M. en C. Eleazar Carranza for their comments in the manu-
script, Alan Smith and George Yatskievch for helpful comments, and Biol. Laura
Noble Camargo for help with the map. The first author thanks the Escuela de
Biologia de la Universidad de Ciencias and Artes de Chiapas for financial sup-
port to explore the northern zone of Chiapas.
—
PEREZ ET AL., THELYPTERIS RHACHIFLEXUOSA NEW FOR CHIPAS, MEXICO 1315
Note.—The first and third authors would like dedicate this manuscript to
the memory of Dra. Ramon Riba y Nava Esparza, who passed away three hours
after checking the identity of this species.
REFERENCES
Breeolove, D. 1981. Introduction to flora of Chiapas. California Academy of Science Press.
California U.S.A.
IBARRA-Manriquez G., M. Martinez-Ramos, R. Dirzo, and J. NUNEZ-FARFAN. 1997. La vegetacion. In:
GONZALEZ S.,E.R. Dirzo and R. Voct, eds. Historia Natural de los Tuxtlas. Universidad Nacio-
nal Auténoma de México. México, D.F.
Lopez-Menpoza, R. 1980. Tipos de vegetacidn y su distribucién en el estado de Tabasco y
norte de Chiapas. Cuadernos Universitarios . Ser. Agronomia 1.Chapingo, México
Lopez-HernAnoez, E.S. 1994. La vegetacion y la flora de la sierra de Tabasco. Universidad
Juarez Autonoma de Tabasco, México, Villahermosa Tabasco.
MirANDA F.and E. HerNANoez X. 1963. Los tipos de vegetacién de México y su clasificacion .
Bol. Soc. Bot. México 28:20-179.
Perez J.A.and J. Sarukhan .1970.La vegetaciéon de la Regidn de Pichucalco, Chiapas, Inst.
Natl. Inv. For. Pub. Esp. 5: 13-48.
Riga, R. 1989. A new species of Thelypteris subg. Goniopteris from the State of Veracruz,
México. Amer. Fern. J. 79:122-124
Riga, R. and B. Peérez-Garcia. 1997. Pteridofitas. In: Gonzalez S., E.R. Dirzo and R. Vogt, eds.
Historia Natural de los Tuxtlas. Universidad Nacional Auténoma de México. México,
DE.
Riga, R. 1998. Pteridofitas mexicanas: distribucién y endemismo. In: Ramamoorthy TP, R.
Bye, A. Lot y J.Fa, eds. Diversidad bioldgica de México: orfgenes y distribucién. Univer-
sidad Nacional Autonoma de México, México, D.F.
Rze00wski, J. 1978.La vegetacion de México. Ed. Limusa, México, DF.
ScHUTZMAN, B., A.P. Vovioes, and B. DeHcan. 1988. Two new species of Zamia (Zamiaceae,
Cycadales) from Southern Mexico. Bot. Gaz. 149:347-360.
Smith, A.R. 1995. Thelypteridaceae. In: R.C. Moran and R. Riba, eds. Pteridofitas. Flora
Mesoamericana. Vol. |. Psilotaceae a Salvinaceae. Universidad Nacional Autonoma de
México. México D-F.
Wenot , 7. 1989. Las selvas de Uxpanapa, Veracruz-Oaxaca, México: evidencia de refugios
floristicos cenozoicos. Anales Inst. Biol. Univ. Nac. Autonoma México, Ser. Bot. 58:29-54.
Wenor,T. 1998. Composicion, afinidades floristicas y origenes de la flora arbdrea del dosel
de los bosques tropicales humedos de la vertiente mexicana del Atlantico. In:
Ramamoorthy T.P., R. Bye, A. Lot y J. Fa, eds. Diversidad bioldgica de México: origenes y
distribucion. Universidad Nacional Aut6noma de México, México, DF.
1316 BRIT.ORG/SIDA 20(3)
BOOKS
FROM UNIVERSITY OF ARIZONA PRESS
TuHeopore H. FLEMING and ALFONSO V ALIENTE-BANUET (Editors). 2002. Columnar Cacti
and Their Mutualists: Evolution, Ecology, and Conservation. (ISBN 0-8165-
2204-9, hbk.). The University of Arizona Press, 355 S. Euclid, Ste 103, Tuc-
son, AZ 85719, US.A. (Orders: 520-621-1441, fax 520-621-8899;
www.uapress.arizona.edu). Price not given, 371 pp., b/w photos, maps, 6"
x 91/4"
From the preface: This book is the result of a conference held in Tehuacan City, Puebla, Mexico (1998)
and focuses on the evolution, ecology (sensu lato), and conservation needs of columnar cacti and
their vertebrate mutualists. There are 17 chapters divided into three sections: 1) Geology and Evolu-
tion; 2) Anatomy and Physiology; 3) Population and Community Ecology and Conservation.
NatHan F Sayre. 2002. Ranching, Endangered Species, and Urbanization in the
Southwest: Species of Capital. (ISBN 0-8105-2158-1, hbk.). The University
of Arizona Press, 355 S. Euclid, Ste 103, Tucson, AZ 85719, U.S.A. (Orders:
520-621-1441, fax 520-621-8899; www.uapress.arizona edu). $48.00, 320 pp,
18 b/w photos, 4 maps, 6" x 9".
Contents: 1) The social production of the endangered masked bobwhite; 2) The cattle boom in the
Altar Valley; 3) The formation of ranching; 4) Producing nature; 5) The urbanization of ranching; 6)
Restoring grass and quail; 7) Producing a state of nature; 8) “Where wildlife comes natur: ally?” 9)
Counterleiting conservation. Notes, References, Index.
SIDA 20(3): 1316. 2003
POLYGONUM DOUGLASII SUBSP. JOHNSTONII
(POLYGONACEAE) NEW FOR CANADA
Mihai Costea! Francois J. Tardif
Department of Plant Agriculture Department of Plant Agriculture
University of Guelph U uelph
Guelph, Ontario, NIG 2W1, CANADA Guelph, Ontario, NIG 2W1, CANADA
coste_amihai@hotmail.com ftardif@uoguelph.ca
Polygonum douglasii E.L. Greene subsp. johnstonii (Munz) Hickman is native to
western North America. Its presence in southwestern Canada was to be expected
since it has been recorded from several adjacent American states—Washing-
ton, Idaho, Montana and North Dakota (Kartesz 1999)—that share common habi-
tats with the southern parts of the western Canadian Provinces. It has not been
recorded from Saskatchewan until now (Vernon Harms, pers. com.). Subspecies
johnstonii grows on sandy soils, open rocky places in dry grass communities
and Ponderosa forests. It flowers from June to August. Four point collections in
the grasslands from southern Saskatchewan are indicated on the map that ac-
companies the herbarium label of the specimen cited. Since its occurrence is
also probable in Alberta and British Columbia, further research is necessary to
determine its exact distribution in Canada. The closest related taxon is
Polygonum douglasii subsp. douglasii (Hickman 1984). Polygonum douglasii
subsp. johnstonii differs from subsp. douglasii mainly through the erect, almost
sessile or short-pedicellate achenes. Subspecies douglasii has spreading or re-
flexed, long-pedicellate achenes). Other distinguishing characters, an identifi-
cation key and line drawings of the two taxa can be found in Hickman (1993).
Voucher specimen: CANADA. Sasxatcuewan: Grasslands Park, 9 mi SE of Val Marie, 17 Jul 1989, E.R.
Hooper 11110 (USAS).
ACKNOWLEDGMENTS
We thank Vernon Harms for confirming the novelty of the taxon in
Saskatchewan; George Ledingham for the additional information about the col-
lection from USAS; and Luc Brouillet for reviewing the manuscript.
REFERENCES
Kartesz, J.T. 1999. A synonymized checklist and atlas with biological attributes for the vas-
cular flora of the United States, Canada and land. |In:Kartesz, J.T.and C.A.Meacham.
1999. Synthesis of the North American flora, version 1.0. North Carolina Botanical gar-
den, Chapel Hill, North Carolina.
Corresponding author
Lad bof
SIDA 20(3): 1317-1318. 2003
1318 BRIT.ORG/SIDA 20(3)
Hickman, J.C. 1984. Nomenclatural changes in Persicaria, Polygonum and Rumex
(Polygonaceae). Madrono 31:249-252
Hickman, J.C. 1993. Polygonaceae In: J.C.Hickman, ed. The Jepson manual: Higher plants of
California. Univ. of California Press, Berkeley. Pp: 854-895.
KOENIGIA ISLANDICA (POLYGONACEAE) NEW FOR UTAH
Mihai Costea! Francois J. Tardif
Department of Plant Agriculture Department of Plant Agriculture
University of Guelph University op Guel Iph
Guelph, Ontario, NIG 2W1, CANADA Guelph, Ontario, N1G 2W1, CANADA
=
coste_amihai@hotmail.com ftardif®uoguelph.ca
Koenigia islandica L. isa circumpolar species that extends south in North and
South America, in scattered alpine summits of the Rocky Mountains and the
Andes (Love & Sarkar 1957; Freeman, unpublished). It occurs in alpine com-
munities growing in permanently moist gravel or moss, especially around per-
sistent nival basins near small streams, ponds, and lakes at elevation up to 3800
m (Freeman, unpublished). Sub-national conservation status has been assessed
to Sl (critically imperiled) in Montana, Wyoming (U.S.A.), Alberta and Manitoba
(Canada), S2 (imperiled) in Colorado (U.S.A.) and $2S3 in British Columbia,
Labrador and Newfoundland (Canada). The species is possibly extirpated (SH)
from Ontario, Canada (NatureServe Explorer 2002).
ucher specimen: U.S.A. UTAH. Duchesne Co.: Ashley Forest, Uinta Monnaie Uints draine age,
Giller Bench E of Gilbert Peak, TSNR4W S24 N1/2 USM, elevation ca. 3600 m
growing in a seep, apparently rooted in moss, 9 Aug 2000, S. Goodrich, A. oe E Bartlett an
F ae 26308( 5) (BRY, NY).
Koenigia islandica can be mistaken with seedlings of Epilobium spp. or with
dwarf plants of Polygonum minimum Small. In the latter case, it can be easily
distinguished by its opposite or whorled leaves and the flowers with 3 tepals
and 3 stamens (P minimum has alternate leaves and flowers with 5 tepals and 8
stamens).
ACKNOWLEDGMENTS
We warmly thank Stanley Welsh for confirming our finding and Craig Free-
man for sharing with us his unpublished treatment of Koenigia for FNA.
REFERENCES
Freeman, C.C. (unpublished). Koenigia L. in Flora of North America, unpublished manu-
script prepared for Vol. 5.
Love, A.and P. Sarkar. 1957.Chromosomes and relationships of Koenigia islandica. Canad. J.
Bot.35:507-514
NatureServe Explorer: AN ONLINE ENCYCLOPEDIA OF LIFE [Web application]. 2002. Version 1.6. Ar-
lington, Virginia, USA: NatureServe. Available: http://www.natureserve.org/explorer.
(Accessed: March 17, 2003).
\Correspondina author
_OFresp
SIDA 20(3): 1319. 2003
1320 BRIT.ORG/SIDA 20(3)
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Gary PAUL NABHAN. Forewords by WeNDELL Berry and MiGueL ALTIERI. 2002. En-
during Seeds: Native American Agriculture and Wild Plant Conservation.
(ISBN 0-8165-2259-6, pbk.). The University of Arizona Press, 355 S. Euclid,
Ste 103, Tucson, AZ 85719, U.S.A. (Orders: 520-621-1441, fax 520-621-8899,
ww w.uapress.arizona.edu). $19.95, 250 pp., 6" x 9"
Contents: Part 1: A New World Perspective; 1) The flowering of diversity; 2) Diversity lost: The wet
and the dry tropics; 3) Fields infused with wildness; 4) Invisible Erosion: The rise and fall of native
farming; 5) A spirit earthly enough: Locally adapted crops and persistent cultures; 6) New and old
ways of saving: Botanical gardens, seed banks, heritage farms, and biosphere reserves, P
Local Parables; 7) Wild-rice: The endangered, the sacred, and the tamed; 8) The eal anc
anomaly: Wild America sunflowers; 9) Lost gourds and spent soils on the shores of Okeechobee; 10)
Drowning ina shallow gene pool: The factory turkey; 11) Harvest time: Northern plains agricultural
change; 12) Turning foxholes into compost heaps, shooting ranges into shelterbelts. Biological Es-
t Le
the ae
Oo
i
says, Literature Cited, Index.
RAYMOND M. TurNeR, Ropert H. Wess, JANICE E. Bowers, and JAMES RODNEY HastTiNGs.
2002. The Changing Mile Revisited: An Ecological Study of Vegetation Change
with Time in the Lowser Mile of an Arid and Semiarid Region. (ISBN 0-
8165-2306-1, hbk.). The University of Arizona Press, 355 S. Euclid, Ste 103,
Titeson, AZ. 05719, Suk. (Orders: 320-021-1441 Ta 20-02-6899:
ww w.uapress.arizona.edu). $75.00, 334 pp., 9" x 12"
Contents: 1) The desert habitat; 2) The influence of American Indians, Spaniards, Mexicans; 3) The
influence of Anglo-Americans; 4) The oak woodland; 5) The grassland; 6) The desert; 7) The pattern
of change; 8) Change and cause. Appendixes: 1) Climate stations in Arizona and Sonora; 2) Common
and Latin plant names; Details of photographic stations and photo credits. Notes, Literature Cited,
About the Authors, Inc
SIDA 20(3): ): 1320, 2003
NOTES ON PWO SPECIES OPAGALINIS
(SCROPHULARIACEAE)
INCLUDING AGALINIS HARPERI, NEW TO TEXAS
Eric L. Keith John Hays
Raven Environmental Services Florida Park Service
PO. Box 6482 13798 SE Federal HY
Huntsville TX 77342, U.S.A. Hobe Sound, FL 33455, U.S.A.
ABSTRACT
New distribution records of Agalinis species are reported including Agalinis harperi Pennell new to
Texas.
RESUMEN
Se citan nuevas | le Agalinis incluyendo Agalinis harperi Pennell nueva para
Texas.
Agalinis harperi Pennell grows in the southeastern United States from South
Carolina westward to Louisiana (Godfrey & Wooten 1981; Vincent 1982). It is
on the Louisiana Natural Heritage Program’s watch list for rare species (Louisi-
ana Natural Heritage Program 1999). Hays (1998) recently clarified the nomen-
clature for this species, previously named A. pinetorum.
A population of approximately 500 plants was discovered in northern
Newton County growing in an open rocky barren on Scrappin Valley, a hunt-
ing preserve and Habitat Management Area (HMA) for the Red-cockaded wood-
pecker owned by Temple-Inland Forest Products Corp. The species has not been
reported for Texas (Hatch et. al. 1990; Jones et. al. 1997), but it has been collected
in four Louisiana parishes bordering Texas (MacRoberts 1989; Thomas & Allen
1998: Vincent 1982). Agalinis harperi is similar to A. purpurea (L.) Pennell but
can be distinguished by the more narrowly-linear, usually ascending leaves (0.5-
1 mm) and by the longer distance between nodes, and thus by a more sparse
vegetative appearance.
Voucher specimen: TE Valley, open, rocky Catahoula barren in RCW clus-
ter 2368E, 31.144499° N an 93.798482° W, 15 Oct 1999, Keith s.n. (BRIT.)
Agalinis filicaulis (Benth.) Pennell is also found in the coastal plain from Geor-
gia westward to southwestern Louisiana (Godfrey @ Wooten 1981; Vincent
1982). It is listed as an Sl species by the Louisiana Natural Heritage Program
(Louisiana Natural Heritage Program 1999). Until recently, only one record of
this species was known west of the Mississippi River in Allen parish (Vincent
1982: Thomas & Allen 1998; Louisiana Natural Heritage Program 1999). One
SIDA 20(3): 1321-1322. 2003
1322 BRIT.ORG/SIDA 20(3)
new record in Beauregard Parish and a collection from Vernon Parish previ-
ously reported in Thomas and Allen (1998) extend the range of this species.
The Vernon Parish specimen was found on the edge of a nearly level pitcher
plant bog, while the Beauregard Parish specimen was found ina wet open pine
savannah. Both of these areas are burned frequently and have a predominately
herbaceous understory. This species should be looked for in similar habitats in
eastern Texas.
Voucher specimens: LOUISIANA. Beauregard Parish: in a wet pine savannah in RCW cluster 6230A
ee 4 mi E of Singer, 30.637805° N and 93.342741° W, 30 2001, Keith 26 (BRIT). Ve
non Parish: edge of pitcher plant bog, 2.5 mi ESE of Fort Polk in Fort Intensive Use Area, 18 5 Sep
1997, Keith sn. (NLU
we
ACKNOWLEDGMENTS
Many thanks to the employees of Temple-Inland Forest Products Corp. includ-
ing Don Dietz, Stan Cook, Bill Goodrum, Robert Wilson, and Jason Sebesta who
have always been helpful and accommodating. Michael MacRoberts of Bog Re-
search and an anonymous reviewer provided many helpful suggestions.
REFERENCES
Goprrey, R.and J.W. Wooten. 1981. Aquatic and wetland plants of the southeastern United
States. Volume II, Dicotyledons. University of Georgia Press.
Hatcn, S.L., K.N. GANDHI, and L.E. Brown. 1990. Checklist of the vascular plants of Texas, MP-
1655. Texas Agric. Exp. Sta., College Station.
Hays, J.F. 1998. Priority of the name Agalinis harperi (Scrophulariaceae) over the names
Agalinis delicatula and Agalinis pinetorum. Sida 18:369-370.
Jones, S.D.,J.K. Wiprr, and PM. Montcomery. 1997. Vascular plants of Texas. A comprehensive
checklist including synonomy, bibliography, and index. University of Texas Press. Austin.
LOUISIANA Natural Heritace Procram. 1999. Rare plant species of Louisiana. NatureServe
website. http:// \Np/us/la/laplants.html.
MacRoseets, D.T.1989.A decanted hes kist and atlas of the vascular flora of Louisiana.
Dicotyledoneae: Fagaceae to Zygophyllaceae. Louisiana State University in Shreve-
port, Shreveport, LA.
THomas, R.D. and C.M. Aiten. 1998. Atlas of the vascular flora of Louisiana, Vol. Ill: Dicotyle-
dons Fabaceae — Zygophyllaceae. Published by the Louisiana Department of Wildlife
& Fisheries Natural Heritage Program
VincENT, K.A. 1982. Scrophulariaceae of Louisiana.M.S. Thesis. Southwestern Louisiana Univ.,
Lafayette.
OX YPOLIS TERNATA (APIACEAE) DELETED FROM THE
TEXAS FLORA
Bruce A. Sorrie Michael H. MacRoberts
3076 Niagara-Carthage Road and Barbara R.MacRoberts
Whispering Pines, NC 28327, U.S.A. Bog Research, 740 Columbia
Shreveport, LA 71104, U.S.A.
dM f Life Sciences
Louisiana State University in Shreveport
Shreveport, LA 71115, U.S.A.
Suzanne Birmingham Walker
14671 State Highway 87 South
Shelbyville, TX 75973, U.S.A.
Rodgers (1950), Watson (1982), Johnston (1990), Texas Natural Heritage Program
(1995), Jones et al. (1997), and Kartesz and Meacham (1999) include Oxypolis
ternata (Nutt.) A. Heller in the Texas flora, apparently on the basis of a 1947
Lincoln Constance annotation of a 1945 V.L. Cory specimen (Cory 49902 [GH])
from Hardin County that was identified initially as Cynosciadium digitatum
DC. The Constance annotation reads: “?Oxypolis ternata.” This apparently led
Rodgers (1950) to include O. ternata in the Texas flora (Rodgers did not anno-
tate Cory 49902) and Watson later (see Johnston 1990:57) to include it in her Big
Thicket National Preserve plant list from which it has subsequently been cop-
ied by others. A search of Watson’s vouchers revealed no Oxypolis ternata
(Brown n.d.).
We examined Cory 49902 and a duplicate at BRIT and found them to be
narrow leaved O. rigidior (L.) Raf., which isa common morphological variant
of that species over much of its range. Both specimens have paired leaflets and
reticulate veins, characters of O. rigidior but not of O. ternata. Another trouble-
some Texas specimen has been Correll 34924 [TEX] collected in Angelina Co. in
1967 and annotated to O. ternata by A.O. Tucker in 1980. It was annotated to O.
rigidior by B.L. Turner in 2000 and reconfirmed to be O. rigidior by Tom Wendt
(pers.comm.) in 2003. We also examined the Mathias and Constance (1961) treat-
ment of Texas Apiaceae and found that they listed Cory 49902 as an example of
O. rigidior but did not mention O. ternata as occurring in Texas. A search of
ASTC, BRIT, DUKE, F FLAS, FSU, NCU, SBSC, SHST, TAMU (on line), TEX, and
VDB revealed no Texas specimens of O. ternata. We have collected specimens
of O. rigidior from Tyler County, adjacent to Hardin County, that are almost
identical in appearance to Cory 49902. These have been deposited at TEX.
As far as we can discern, O. ternata occurs no closer to Texas than the cen-
SIDA 20(3): 1323-1324, 2003
1324 BRIT.ORG/SIDA 20(3)
tral Florida panhandle (Wunderlin 1998; Florida Atlas of Vascular Plants
www plantatlas.usf.edu)).
=
ACKNOWLEDGMENTS
Bill Carr and Jason Singhurst, the Texas Parks and Wildlife Department, and
the curators and staff of the herbaria where we worked, all aided with the study.
Tom Wendt and Lindsay Woodruff examined a specimen at TEX.
——
REFERENCES
Brown L.E. n.d. Vouchered list of Big Thicket National Preserve plant collections at Rice
and Lamar universities. Unpublished report. Big Thicket National Preserve, Beaumont.
Jounston, M.C. 1990. The vascular plants of Texas: A list up-dating the “Manual of the vas-
cular plants of Texas.” Second edition. Privately printed.
Jones, S.D., J.K. Wiprr, and P.M. Montaomery. 1997.Vascular plants of Texas. University of Texas
Press, Austin.
Kartesz, J.T.and C.A. Meacham. 1999. Synthesis of North American flora. Version 1.0. Chapel
Hill,
Matias, M.E. and L. Constance. 1961. Umbelliferae. Flora of Texas. Vol. 3. Part 5. Texas Re-
search Foundation, Renner
Ropaers, C.L. 1950. The Umbelliferae of North Carolina and their distribution in the south-
east. J. Elisha Mitchell Sci. Soc. 66:195-266,
Texas Natural Heritact Procram. 1995. Special plant list. Unpublished report. Texas Parks
and Wildlife Department, Austin.
Watson, G.E. 1982. Vegetational survey of the Big Thicket National Preserve. Upublished
report. Big Thicket National Preserve, Beaumont.
WUuNDERLIN, R.P. 1998. Guide to the vascular plants of Florida. University Press of Florida,
Gainesville.
ANOTHER JUNIPER (CUPRESSACEAE) IN JEFF DAVIS
COUNTY TEXAS? WE'RE NOT CONVINCED
John P. Karges James C. Zech
The Nature Conservancy Department of Biology
PO. Box 2078 Sul Ross State iversity
Fort Davis, TX 79734, U.S.A. Alpine, TX 79832, U.S.A
jkarges@tnc.org, seas
Within volume twenty, number two, of this journal, Griffith and Bartel (2002)
answered our request (Karges & Zech 2001) for further study and documenta-
tion of Cupressus arizonica within the Davis Mountains of Jeff Davis County,
Texas. While the effort has been made, we found several problems with the at-
tempt, including the main points of concern that follow. Of the three additional
specimens sampled from Jeff Davis County there is no indication that any were
collected from the Davis Mountains, let alone the Davis Mountains Preserve
(DMP). We’re puzzled as to why fresh material of the Jeff Davis County C.
arizonica or Juniperus deppeana from the DMP were not included for chloro-
plast DNA analysis. Both would have been made available upon request. Griffith
and Bartel (2002) cite two specimens of the Jeff Davis County C. arizonica.
Karges and Hedges 2480 is the only existing specimen, a second specimen, Karges
s.n., was never collected. Griffith and Bartel’s (2002) reference to, and discus-
sion of, a mixed specimen is most puzzling, since neither Hedges (pers. comm.)
nor Karges collected reproductive material.
Perhaps the chloroplast DNA data does suggest that the Jeff Davis County
Arizona Cypress is Alligator Juniper. However, this study’s discre} ies, along
with the admittedly low 63% bootstrap, leave us unconvinced. We still recog-
nize the tree in question as C. arizonica, and most importantly, continue to en-
courage additional, careful, study of this unique tree.
REFERENCES
GrirFitH, M.P. and S.C. Bartet. 2002. A Cypress (Cupressus arizonica, Cupressaceae) in Jeff
Davis County, Texas? Sida 20:585-592.
Karces, J.P.and J.C. Zech. 2001. Cupressus arizonica (Cupressaceae) new to the Davis Moun-
tains of West Texas. Sida 19:719-721.
SIDA 20(3): 1325. 2003
1326 BRIT.ORG/SIDA 20(3)
BOOK NOTICES
Davib H. Gisson. Introduction by JOHN B. ROHRBACH. 2002. Casyouland Vision: The
Colorado Plateau of Southeast Utah. (ISBN 0-57441-1560-X, hbk.). Waterous
& Co. (Orders: Folio Gallery Inc., 4404-15" Street S.W. Calgary, A Alberta T2T
4B2, CANADA). Price not given, introduction, 44 b/w photos (six-by-sev
enteen-centimeter negative), 14" x 10".
The dust jacket notes, “Canyonland Vision: The Colorado Plateau of Southeast Utah captures the
sweeping majesty of the harsh but poetic canyons of the American Southwest. John Rohrbach, Asso-
ciate Curator of Photographs at the Amon Carter Museum in Fort Worth, Texas, provides an insight-
ful introduction to Gibson's work.”
The response | received after showing this book t I as, “wow is this Ansel Adams”?
No, this is David Gibson of Dallas, Texas and be loves nature ae ieee It’s easy to see that
David wie nature y onli at these magnificent photographs done in panoramic format. | can
just stare at t y. There are pics that include aspens, cottonwoods, Chamisa, ancient pines,
ete.
The breathtaking photographs will stir your soul and your sense of wonder. | can't help but
remember the quote by J.B.S. Haldane, “The world will not perish for want of wonders, but for want of
wonder.” The Colorado Plateau of southeast Utah is filled with beautiful and inspiring wonders. When
you look at David Gibson’s photographs in Canyonland Vision you will be filled with “wonder.” |
wonder how he did it! | wonder when I will get to see these poetic canyons myself. Soon I hope!—
Barney Lipscomb, Botanical Research Institute of lexas, 509 Pecan Street, Fort Worth, TX 76102-4060,
U.S.A, barney@britorg.
CHRISTOPHER T. Martine. Illustrated by Rachie A. FicLey. 2002. Shrubs and Vines
of New Jersey. (No ISBN, pbk. spiral bound). New Jersey Forest Service, Forest
Resource Education Center, 370 E. Veterans Highway, Route 527, Jackson,
NJ 08527, U.S.A. (Orders: 908-928-0987, njfsfrec@bellatlantic.net
wwwstate.nj.us/dep/forestry, check ae to NJ Forest Service). $10.00,
114 pp., line drawings, maps, 4 1/2" x
This is a nice pocket guide to the shrubs and vines of New Jersey. Each taxon comes with an illustra-
tion and includes information on common name, Latin name, family, other names [common], wet-
land indicator status, NJ status, habitat and range; descriptive information is given on eat leaves,
al
stems, flowers, and fruit. W —o species might be confused with other taxa, a g ris
provided on similar species
SIDA 20(3): 1326. 2003
A ROCKHOUSE MICROHABITAT IN THE WEST CROSS
TIMBERS OF NORTH CENTRAL TEXAS
George M. Diggs, Jr. Robert J.O’Kennon
a Biology Botanical Research Institute of Texas
College 509 Pecan Street
San see 75090, U.S.A. Ft. Worth, TX 76102, U.S.A.
and bokennon@brit.org
Botanical Research Institute of Texas
gdiggs@austincollege.edu
ABSTRACT
A rockhouse microhabitat located in the xeric savannah/ woodland community of the West Cross
Timbers of North Central Texas is discussed and an associated disjunct population of Arisaema
triphyllum is reported.
RESUMEN
Se discute un microhabitat rocoso en la comunidad sabana/arbolado de West Cross Timbers en el
centro-norte de Texas y se cita una poblacion disyunta asociada de Aris triphyllum.
“Rockhouses” or rockhouse cliff formations are deep recesses in bedrock cliffs
that can maintain microhabitats that are highly buffered from the extremes in
temperature and moisture typical of the surrounding macroenvironment
(Farrar 1998). As discussed by Walck et al. (1996), in comparison with the out-
side macroenvi ,rockhouse habitats tend to be more shaded, have higher
temperatures during the winter, lower temperatures during the summer, and
have lower rates of evaporation and higher relative humidities. Not only is there
less evaporation, but water often enters via groundwater seepage, in some cases
providing a relatively stable supply of moisture. In addition, condensation on
the rock walls further increases humidity locally. The result is a rockhouse ca
provide microclimatic conditions very different (more moderated) from those
typically found in a given geographic area.
There are numerous such rockhouse haoitats in various areas of the east-
ern United States, particularly in sandstone strata. Several recent papers (Walck
et al. 1996; Farrar 1998) have discussed these rockhouses and the interesting
plant taxa now known to occur in them. According to Walck et al. 1996), 11
plant taxa are endemic or nearly endemic to sandstone rockhouses in the east-
ern U.S. Particularly interesting is the occurrence in rockhouses of a number of
bryophytes and ferns with tropical affinities (Farrar 1998).
While relati vely common in the eastern U.S : such khouse environments
seem unlikely in the West Cross Timbers of North Central Texas—particularly
SIDA 20(3): 1327-1333. 2003
1328 BRIT.ORG/SIDA 20(3)
since hard surface rocks are unusual in most of the region. Such rocks, however,
can be found in the area known as the Palo Pinto Country in the northwestern
portion of the West Cross Timbers. This rather rugged region is underlain by
the oldest rocks exposed in North Central Texas, deposited during the Pennsyl-
vanian Period. These rocks vary, but are “largely made up of soft, impure shales
alternating with harder, coarse, brown sandstone and conglomerates” (Hill
L901). In one area of western Parker County, where Pennsylvanian sandstones/
conglomerates are exposed, deep recesses have formed that are worthy of the
term rockhouse. Striking rock walls approximately 40 feet (2 meters) tall form
narrow passageways and recesses. It has been known for a number of years that
one unusual plant distributional record is associated with this environment.
Cary ovata (Mill.) K. Koch, shag-bark hickory Quglandaceae), is disjunct to this
area by about 100 miles (275 kilometers) from its main region of occurrence in
Fast Texas (e.g., Lipscomb 2321, 1977—BRIT). The site is also the westernmost
locality in Texas for Asplenium platyneuron (L.) Britton, Sterns, @ Poggenb.,
ebony spleenwort (Aspleniaceae) (Turner et al. 2003).
The moderated microclimate of this sheltered rockhouse microhabitat
stands in rather sharp contrast to the macroclimate of the exposed upland West
Cross Timbers macrohabitat/community. While yearly average precipitation
in the area is approximately 29 inches (74 cm) (Griffiths @ Orton 1968; Hatch
et al. 1990), rainfall is unevenly distributed with summers typically being ex-
—
tremely hot and dry. Asa result, the upland vegetation has a distinctly dry area
aspect—a savannah/woodland with stunted Quercus stellata (post oak) and
Quercus marilandica (blackjack oak),an understory of grasses (Diggs et al. 1999),
and even such xerophytic, mainly western species as Ber beris trifoliolata Moric.
(agarito), Echinocereus reichenbachii (Terscheck ex Walp.) E Haage (lace cac-
tus), Juniperus ashei J. Buchholz (mountain-cedar), Mimosa borealis A. Gray
(catclaw), Opuntia leptocaulis DC. (pencil cholla), Zanthoxylum hirsutum
Buckley (prickly-ash), and Ziziphus obtusifolia (Hook. ex. Torr. & A. Gray.) A.
Gray (lotebush).
Several years ago, another disjunction was discovered by one of us
(O’Kennon). A large population of Arisaema triphyllum (L.) Schott, jack-in-the-
pulpit (Araceae), was found in the mesic microhabitat between rock walls (Figs.
land 2) in the same extensive rockhouse as Carya ovata (voucher: O’Kennon
16024—BRIT). Interestingly, even though we estimate the population at approxi-
mately 2,000 above ground stems (the number of genetically distinct individu-
als is unknown), it had not been reported previously in the literature, possibly
due to confusion with the related widespread species, Arisaema dracontium
(L.) Schott, (green-dragon). The locality information was provided to Turner et
al. (2003) for inclusion in the Atlas ofthe Vascular Plants of Texas. The large num-
bers of individual stems present and their occurrence through a rather exten-
sive area of the rockhouse argues against any type of recent human interven-
TEXAS 1329
=}
a
4
a]
h
=
~
a]
Fic. 1
(photo by R.J. O’Kennon).
1330 BRIT.ORG/SIDA 20(3)
tion (e.g., transplantation). Observation of the population over two growing sea-
sons revealed that relatively few plants produce inflorescences (roughly one in
LOO during the first week of April), even fewer produce fruits (during one sea-
son only three individuals were observed with mature fruits), and most repro-
duction is probably by vegetation means. At the site, digging indicated that rhi-
zomes/underground stolons could be traced between different stems signifying
vegetative expansion of the population. Additionally, several transplanted
plants produced offsets (traceable to the original plant) the following spring
approximately 40-50 cm from the parent plants. It thus seems clear that the
plants are reproducing vegetatively, and that the number of genetically differ-
ent individuals probably can be determined only by molecular analyses. T
apparent lack of extensive sexual reproduction is interesting in light of Kral’s
—
—
re
(1966) observation that other relict species in East Texas were also apparently
reproducing predominantly via vegetative means. The disjunction between the
western Parker County site and the nearest known East Texas location (Hend-
erson County) is approximately 140 miles (225 kilometers). The occurrence of
this species in the West Cross Timbers is quite unexpected given its preference
for low or moist woods. However, the special microclimate (particularly reduced
evapotranspiration and persistent substrate moisture) apparently has allowed
the long term survival of this mesic species in an otherwise rather inhospitable
macroenvironment.
The taxonomy of Arisaema triphyllum is controversial. Detailed study of
these extremely variable plants led Huttleston (1949, 1981, 1984) to conclude
that the variation seen in the species was best represented by recognizing a
single species with four subspecies; he indicated that “..plants of the four sub-
species are very distinctive and readily identified, at least in living condition.
Since many of the key characteristics..are obscured or lost during drying, it is
notalways possible to identify herbarium specimens to subspecies.” At the other
taxonomic extreme, because “much overlap occurs in expression of the charac-
teristics supposedly defining infraspecific taxa,” because numerous interme-
diate forms exist, and because the ranges of the subspecies overlap, Thompson
(2000) recently recognized a single variable species without infraspecific taxa.
Thus, depending upon one’s treatment of the variation present, the disjunct
Parker County population would be either Arisaema triphyllum (no subspe-
cies recognized) or A. triphyllum subsp. triphyllum. In a forthcoming flora of
East Texas, we are treating the taxon as A. triphyllum subsp. triphyllum.
As discussed by Kral (1966), MacRoberts and MacRoberts (1997), and oth-
ers, Texas has numerous plant occurrences which represent relicts of past cli-
matic conditions. Further, from the study of Texas bogs (e.g., Potzger @ Tharp
1954; Bryant 1977; Holloway & Bryant 1984; Bryant & Holloway 1985) and from
knowledge of modern day plant distributions (e.g., Turner et al. 2003), itis clear
TEXAS 1331
Fic. 2. Closeup of Arisaema triphyllum (photo by R.J.0’Kennon).
that Texas vegetation was profoundly influenced by the climatic change asso-
ciated with glaciation, even though actual glaciers were hundreds of miles to
the north of Texas. The Parker County occurrence of Arisaema triphyllum thus
probably represents an “Ice Age holdover” (MacRoberts & MacRoberts 1997)
froma colder and wetter period of the Pleistocene when vegetational areas were
shifted considerably to the south and west. It is known, for example, that at
15,000 years before present, the mean annual air temperature of Texas was 5° C
less than at present, conditions were moister, and there was a more widespread
forest mosaic over most of Texas (Bryant 1977; Stahle @ Cleaveland 1995). Other
modern day reflections of such past conditions can be seen in the many species
and genera typical of East Texas showing up in isolated disjunct pockets on the
Edwards Plateau (e.g. Hamamelis virginiana L., witch-hazel) as well as many
other interesting disjunct species and communities (Palmer 1920; Diggs 2002).
While rare in Texas, rockhouses do provide an interesting window into bio-
geographic history. As noted by Farrar (1998), the disjunct occurrences of plant
species in rockhouses “document, as clearly as do fossils, a record of changing
climate and vegetation.” When this phenomenon is taken together with the
study of bogs, and an understanding of other current day plant distributions,
the impact of glacial history on the vegetation of Texas becomes increasingly
understandable.
1332 BRIT.ORG/SIDA 20(3)
ACKNOWLEDGMENTS
Thanks to James Peck and Donald Farrar for reviewing the manuscript and
making helpful suggestions.
REFERENCES
Bryant, V.M.Jr. 1977.A 16,000 year pollen record of vegetational change in central Texas.
Palynology 1:143-156.
Bryant, V.M. Jr.and R.G. Hottoway. 1985. A late-Quaternary paleoenvironmental record of
Texas: An overview of the pollen evidence. In: V.M. Bryant, Jr. and R.G. Holloway, eds.
Pollen records of late-Quaternary North American sediments. Pp. 39-70. American
Assoc. of Stratigraphic Palynologists Foundation, Dallas, TX.
Dicas, G.M., Jr. 2002. East Texas as a unique habitat. In: Native Plant Society of Texas. Special
habitats. Pp. 53-66. 2002 Symposium Proceedings, Houston.
Dicas, G.M., Jr. B.L. Liescome, and R.J.O’Kennon. 1999, Shinners & Mahler's illustrated flora of
North Central Texas. Sida, Bot. Misc. 16.
Farrar, D.R. 1998. The tropical flora of rockhouse cliff formations in the eastern United
States. Bull. Torrey Bot. Soc. 125:91-108.
GrirFiTHs, J.F.and R. Orton. 1968. Agroclimatic atlas of Texas—Part |. Precipitation probabili-
ties. Texas Agric. Exp. Sta. Misc. Publ. No. 888.
Harcu, S.L., KN. GANDHI, AND L.E. Brown. 1990. Checklist of the vascular plants of Texas. Texas
Agric. Exp. Sta. Misc. Publ. 1655:1-158.
Hitt, R.T. 1901.Geography and geclogy of the Black and Grand prairies, Texas. 21st Annual
Report of the U.S. Geological Survey, Part Vil-Texas. U.S. Government Printing Office,
Washington, D.C
Ho.toway, R.G. and V.M. Bryant. 1984. Picea glauca pollen from late-glacial deposits in cen-
tral Texas. Palynology 8:21-32.
Hutteston, D.C. 1949. The three subspecies of Arisaema triphyllum. Bull. Torrey Bot. Club
76:407-413.
Huttieston, D.C. 1981. The four subspecies of Arisaema triphyllum. Bull. Torrey Bot. Club
108:479-481,
Huttieston, D.C. 1984. The North American species of Arisaema (Araceae)- “Jack-in-the-
Pulpit.” Aroideana 7:14-18.
Krat, R. 1966. Observations on the flora of the southeastern United States with special
reference to northern Louisiana. Sida 2:395—408.
MacRoeerts, B.R. and M.H. MacRoserts. 1997. Floristics of beech-hardwood forests in East
Texas. Phytologia 82:20-29.
Patmer, EJ. 1920. The canyon flora of the Edwards Plateau of Texas. J. Arnold Arbor. 1:233-
239.
Porzcer, J.E.and B.C. THarp. 1943. Pollen record of Canadian spruce and fir from a Texas bog.
Science 98:584-585.
Stante, D.W. and M.K. CLeavetanp. 1995. Texas paleoclimatic data from daily to millennial
time scales. In: J. Norwine, J.R. Giardino, G.R. North, and J.B. Valdés, eds. The changing
TEXAS 1333
climate of Texas: Predictability and implications for the future. Chapter 7. Pp. 49-69.
GeoBooks, Texas A&M Univ., College Station.
THompson, S.A. 2000. Araceae. In: Flora of North America Editorial Committee, eds. Fl. North
Amer. 22:128-142. Oxford Univ. Press, New York and Oxford.
Turner, B.L., H. NicHots, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas, Vol.
monocots. Sida, Bot. Misc. 24
1:Dicots, Vol. 2: Ferns, g |
Watck, J.L., JM. BASKIN, and C.C. Baskin. 1996. Sandstone rockhouses of the eastern United
States with particular reference to the ecology and evolution of the endemic plant
taxa. Bot. Rev. 62:311-362.
1334 BRIT.ORG/SIDA 20(3)
BOOKS
FROM UNIVERSITY OF NORTH TEXAS PRESS
CampBeLL and LYNN LOUGHMILLER (Compilers & Editors). Foreword by Francis E.
ABERNETHY. 2002. Big Thicket Legacy. (ISBN 0-57441-150-X, hbk.). Univer-
sity of North Texas Press, PO Box 311336, Denton, TX 76203-1336, U.S.A.
(Orders: 940-565-2142, fax 940-565-4590, rchrisman@unt.edu, www.untedu/
untpress). $14.95, 256 pp., 42 b/w photos, | map, 7" x 10"
This book was first published in 1977 and this paper back printing brings this classic on Big Thicket
history back into print. Key Words: Texas, Big Thicket (Texas), folklore, natural history.
GERALDINE ELLis WATSON. 2003. Reflections on the Neches: A Naturalist’s Odys-
sey Along the Big Thicket’s Snow River. (ISBN 1-57441-160-8, hbk.). Uni-
versity of North Texas Press, PO Box 311336, Denton, TX 76203-1336, US.A.
(Orders: 940-505-2142, fax 940-505-4590, rchrisman@untedu, www.untedu/
untpress). $14.95, 256 pp., Ll b/w photos, 29 drawings, 20 maps, 7" x 10".
If there was ever a dust jacket that made you want to oP n up a book it is the one on this book. It
its stunning buttress-like trunks, re-
)wit
pictures the glorious bald cypress (laxodium d
flecting on the Neches River. This book is a reflection of Geraldine Watson
“The book is a series of essays about a river and its
and her love of nature,
East Texas, the Neches River, and the Big Thicket.
environment, and the kind of people such environs produce, which includes myself.”
| conservationist, was one of several
Geraldine Watson, botanist, plant ecologist, naturalist, anc
dle instrumental in raising attention and serving as a national conscience to save at least some
peo}
thing of the Big Thicket in East Texas. Thank you Geraldine for your hard work and for giving us a
book full of history and memories! Thanks to the TLL. pee Foundation for a generous grant to
help publish the beak and to the Big Thicket Association for their assistance.—Barney Lipscomb, Bo-
tanical Research Institute of Texas, 509 Pecan Street, ae auth IX 76102-4060, U.S.A.,
barney@britorg.
SIDA 20(3): 1334. 2003
WILLIAM DEAN REESE
(1928-2002)
Ronald A. Pursell Paul L.Redfearn
Department of Biology Biology Department, Box 14
The Pennsylvania oe ed Southwest Missouri State University
University Park PA 16802-5301 “ eld MO 70504-0095
rap ek. Ir426g@smsu.edu
William Dean Reese, ‘Bill’ to family members and his many friends, died, age
73, on 4 February 2002 in Lafayette, Louisiana after a short battle with cancer.
Bill was born in Baltimore, Maryland on 10 September 1928. His formative
years were spent ona dairy farm, and asa result he always considered himself a
‘farm boy at heart. Perhaps asa result of his experiences on the farm Bill hada
practical approach to life and an unquenchable appreciation for nature. This
view never left him. Bill strived to pass this view on to his children and grand-
children. Realizing that his children and grandchildren would never experi-
ence the joys of farm life he decided to record memories of his years on the farm,
primarily for their benefit. These memories were privately published in 1997 as
Lone Hickory. Reminiscences from the Reese Farm at Gwynnbrook, Maryland,
1930s-1940s, with corrections made in 1998.
Bill enrolled in the University of Maryland in 1945, but his undergraduate
studies were interrupted by two tours in the United States Army. In 1946-48, as
a member of the Signal Corps, he was stationed in the Philippines; between
1950-51 he was a cook in Fort Hood, Texas. Bill’s great love of cooking (and eat-
ing!) good food was perhaps initiated by this experience. In 1953 Bill was gradu-
ated from the University of Maryland with a B.S. in Botany. It was during his
undergraduate studies that Bill’s interest in bryophytes was piqued. In one of
his biological science laboratories Bill observed the release of living
antherozoids from moss antheridia, an observation that led to his second
bryological publication, On observing bryophyte antherozoids (Bryologist
58:335-336. 1955). In the Fall of 1953 Bill began graduate studies in bryology at
Florida State University under Dr. Ruth S. Breen, graduating with an MS. in
Botany in 1955 and a Ph.D. in Botany in 1957.
Upon graduation, Bill was offered and accepted an appointment as Assis-
tant Professor of Biology at The University of Southwestern Louisiana (now The
University of Louisiana at Lafayette). He remained at this university through-
out his 30-year professional career, advancing to Associate Professor (1962) and
Professor (1966). Bill was an outstanding teacher and served as primary advisor
to 11 graduate students. In 1970 he was named University of Southwest Louist-
SIDA 20(3): 1335-1338. 2003
1336 BRIT.ORG/SIDA 20(3)
WILLIAM DEAN REESE
(1928-2002)
PURSELL AND REDFEARN, WILLIAM DEAN REESE 1337
ana Foundation Distinguished Professor. Upon his retirement in 1987 the Uni-
versity appointed him Edwin L. Stephens Honor Professor Emeritus, an appoint-
ment he held until his death. In addition to his teaching responsibilities, Bill
served the University and Biology Department as an able ad rator: Chair-
man of the Biology Department (1974-77) and Department Head (1981-86).
Bill was a towering figure in American bryology. His primary interest cen-
tered on the tropical/subtropical family Calymperaceae. His studies on this
family began with his Ph.D. dissertation and continued for the remainder o
his life. Never tiring of these mosses, he delved into their systematics and phy-
logeny, the evolution and functional morphology of their asexual reproduction,
and the evolution of their niche specificity. At his death he left an incomplete
manuscript, Fertility in Calymperaceae: Calymperes, Mitthyridium and
Syrrhopodon. Bill’s interest in bryology, however, was not limited to the
Calymeraceae. He was also a superb floristic bryologist with particular inter-
est in the mosses of the southern United States. His book Mosses of the Gulf Coast,
which treats the species from the Rio Grande (southern Texas) to the Apalachicola
River (western Florida), is the standard floristic work for that region.
Throughout his career, Bill had a total of 154 bryological publications.
Bill was a consummate botanist. Although his principal interests lay in
bryology, Bill was an acknowledged authority on the taxonomy of vascular
plants. He took great delight in showing visitors the many interesting native
flowering plants he had transplanted around his home. His professional inter-
est in vascular plants led to the publication of several papers as well as the book
Acadiana Flora. Native and Naturalized Woody Plants of South-Central Loui-
siana, completed with his long-time colleague at the university, Garrie Landry.
At the time of his death he was working on another book., Mamou: Acadian
Folklore, Natural History, and Botany of Erythrina herbacea L., with Charles
Allen, a former graduate student. Away from botany, he had considerable knowl-
edge of fungi, particularly the basidiomycetes, and was an avid Or nibHo/ORIst
delighting in pointing out to visitors the many spectacular bird th-
ern Louisiana.
Billanda colleague, John Thieret, were the first (and only) botanists to com-
plete an exhaustive survey of the vascular flora of the five ‘salt domes’ [Bell Is-
land, Cote Blanche Island (accessible only by boat), Weeks Island, Avery Island,
and Jefferson Island] located along the Louisiana coast. The results of this sur-
vey were published as Botanical Study of the Five Islands of Louisiana. Bill was
immensely proud of his part in this exploratory work and continued to revisit
the accessible areas long after the survey was completed.
Bill found great enjoyment in field work. Collecting specimens of bryo-
phytes and vascular plants was one of his great pleasures and greatly enhanced
his research. In addition to collecting extensively in North America he also par-
ticipated in expeditions to Mexico, the Amazon Basin in South America, the
mae)
1338 BRIT.ORG/SIDA 20(3)
West Indies, Australia, and China. His earliest recorded collection, a specimen
of Quercus falcata Michx., was made just outside Tallahassee, Flordia in August
1955, and his last identified collection (Reese 18,593), Pseudoleskeella tectorum
(Brid.) Kindb., was made in Otero County, New Mexico on 15 June 2000. These
collections (numbering about 35,000 specimens) are now in MO and are avail-
able for study. Eventually the collections will be intercalated in the Crosby Bryo-
phyte Herbarium (MO). Record books and journals are also on file at MO.
In addition to his duties as teacher, researcher and administrator, Bill was
an exceptional editor. Although he was exacting and had little patience with
foolishness, he balanced his criticisms with encouragement and insightful com-
ments, a reflection of his quiet and even nature. When a manuscript was re-
turned by Bill one could be certain that it was considerably improved, regard-
less of how many times it had been edited previously. Bill was twice Editor of
The Bryologist (1970-74; 1988-1995), and Associate Editor of The Bryologist
(1977-1980), and an editor for and an unwavering supporter of the Bryophyte
Floraof North America project, for which before his death he completed manu-
scripts in the Brachytheciaceae, Caly mperaceae, C1 y phaeaceae, H ypnaceae,
Leucodontaceae, Leucophanaceae, Meteoriaceae, Pterobryaceae, Ptychomitriaceae,
Racopilaceae, and Splachnobryaceae.
Bill served The American Bryological and Lichenological Society as Secre-
tary- Treasurer (1969-1970), Vice-President (1977-1979), and President (1979-
L981). In addition, Bill was a member of the American Association for the Ad-
vancement of Science, Association of Southeastern Biologists, British Bryological
Society, International Association of Bryologists, International Association for
Plant Taxonomy, Nordisk Bryologisk Forening, Sociedad Latinamericano de
Briologia, Society of the Sigma Xi, and a charter member of the Hedwigian So-
ciety (Missouri Botanical Garden). In 1988 he was appointed a Research Asso-
ciate by the Missouri Botanical Garden.
Bill is survived by his wife, Lorelle, two daughters, Jean Harvill and Chris-
tine Proctor, twin sons, David and William, and five grandsons.
Bryology has lost one of its pre-eminent students, his family a loving and
wonderful husband, father and grandfather, and those of us who knew him well,
a faithful and loyal friend.
The tribute to Bill published in Bryologist 105(4):625-634. 2002 [2003] in-
cludes a list of taxa associated with Bill, and a bibliography of his botanical
publications.
BOOK REVIEW
JouN FE Nunn. 2002. Ancient Egyptian Medicine. (ISBN 0-8061-3504-2, pbk.) (Or-
ders: University of Oklahoma Press, 4100 28th Avenue, Norman, OK 73069-
8218; www.oupress.com, 405-307-9048 fax). $29.95, 240 pp, 45 b/w illus-
trations, 32 tables, | map, 7" x 10".
When one considers the topic of Egyptology, the thoughts which arise are sure to concern them-
selves with the elaborate tombs and pyramids that can be found in any picture guide to the Ancient
Egyptians; although this is an important facet of study, it allows other, more subtle aspects of the
culture to be ignored. Within the 240 pages of his work, John Nunn presents a comprehensive, yet
detailed, account of one of the lesser studied topics within Ancient Egyptian culture: medicine. Nunn
carries the reader froma brief background of the aspects of Egyptian culture influencing their medi-
cal practice to studies of their healers, drug therapies and to their more specialized branches. Ap-
proaching the subject as a retired anesthesiologist as well as an Egyptologist, Nunn is in a unique
position toactasan authority on the merging of these two subjects.
The book begins with a brief geographical and historical placement of the Ancient Egyptians,
establishing the “unique environment in which their medical practice could be developed and re-
corded” by means of discussion of methods of food production, nutritional state, life expectancy,
writing, etc. The primary concern of this first chapter is to set the stage and the background for the
reader to be able to follow the remainder of the book. After establishing that the culture had the
leisure to engage in activities other than defense and food production, the discussion moves to the
primary source of the ee aims Nunn males, the medical papy
Through an ext ilation and investigation pe ious surviving medical texts writ-
tenon papy es as vay Ws). sean reaches his ee: ane the medical practice of the
At
ncient Egyptians he difficulties of translation. Other important sources
of information used in the investigation anil ale human remains and artistic renditions of the dis-
ease state. In particular, the embalming techniques and environmental conditions (a dry and hot
climate) of the region acted to preserve the human remains in excellent detail (causing them to des-
iccate faster than they would putrely). There is a conscious effort ae to avoid the pitfalls of vali-
dating the past in terms of modern concepts, while yet highlighting the fact that certain medical
procedures practices by the Egyptians are in fact up to modern standa
The remaining chapters carry the topics mentioned at the beginning to deeper levels; for ex-
ample, carrying the topic of the patterns of disease, to intricate discussions about such things as
parasitic, viral and bacterial diseases, deformities, and ageing processes. Furthermore the book is
littered with documented studies regarding each topic, thus also serving as a reference to further
reading on the particular topics. The readers will chuckle as they read the various treatments for
disease states, such as those found within the chapter on specialized medical branches. Regarding
sex determination during pregnancy: “Emmer (bedet) and barley (it), the lady should moisten with
her urine every day, like dates and like sand in two bags. If they all grow, she will bear a child. If the
barley grows, it means a male. If the emmer grows, it means a female. If they do not grow, she will not
bear a child.” The interesting aspect about this case is the unusual findings Nunn documents of stud-
ies done in 1963 by Ghaliounghui et al., testing the treatment.
The format of the book is quite easy to follow, it increases in complexity as the reader's back-
ground is established, and where a subject is touched upon that will later be clarified, the location is
specified. Although there are no color illustrations, those few well ania di lagreits clar ay the con-
cepts dealt with in the text. The many
}
g with
herbal remedies of certain, less certain and dneenens aw as well as 29 others. While it is aes
SIDA 20(3): 1339. 2003
1340 BRIT.ORG/SIDA 20(3)
that the majority of am will be able to follow the entire book with complete understanding, th
discussions are all s
>
iciently in-depth to maintain a specialist’s interest while also maintaining
that of a ae new tot ce field.
-
=
rile the information contained within this 2002 work is merely a reprint of the 1996 cloth
saat ee the improved presentation of the material makes it less liable to being overlooked. TI
book manages to bridge a gap in its partic
1€
ular field, ne and ee: upon other a
tions which have at best been incomplete and muddling. Nunn has presented a concise and well-
rounded approach to the study of medicine in Ancient Egyptian ¢ hire are will i both the
merely interested reader and the avid scholar, fully satistied.—Tiana Francesca Franklin, Botanical
Research Institute of lexas, 509 Pecan Street, Fort Worth, TX, 76102,4060, U.S.A, tianaff@hotmail.com.
NEW BOOKS
Davib PiMeNnteL (Editor). 2002. Biological Invasions: Economic and Environmen-
a ee of Alien Plant, Animal, and Microbe Species. (ISBN 0-8493-0830-
k.). CRC Press LLC, 2000 N.W. Corporate Blvd., PO Box 31225, Boca
a PL 33431, U.S.A. (Orders: 800-272-7737, fax 800-374-3401; Canada
& International 561-994-0555, fax 561-989-8732, www! cpl ess.com). Price
not given, 369 pp., graphs/Tigs., 7" x 10"
Forty-five contributors provide detailed information on oS of the invasive-species prob-
lem from six continents. The book is divided into nine sections: 1] tralia; WI Brazil:
IV British Isles; V aes New Zealand; VU one Africa; VILE United States; IX see Overview. The
book is informative and will serve “scientists, policymakers, and the
public on the seriousness of
non-native species invasions, which can cause extinctions and significant costs to the environment,
economy, and public health.”
Eric T. Jones, Resecca J. McLAIN, and JAMES WEIGAND (Editors). 2002. Nontimber
Forest Products in the United States. (ISBN 0-7006-1165-7, pbk.). Univer-
sity Press of Kansas, 2501 West 15" Street, Lawrence, KS 66049, U.S.A.(Or-
ders: 785-864-4154, fax 785-864-4586, www.kansaspress.ku.edu). $29.95,
445 pp., 2 figures, 6" x 9"
From press release: “Nontimber Forest Products in the United States provides the first comprehensive
examination of these abundant resources throughout the United States, illustrating nontimber for-
est products’ diverse importance, Sees mae people who harvest th
hem, and outlining the steps
hat are being taken to ensure access to them
DUANE A. SMITH. 2002. Mesa Verde National Park: Shadows of the Centuries, Revised
Edition. ISBN 0-87081-084-5, pbk.). University Press Colorado, 5589 Araphoe
Road, Suite 206C, Boulder, CO 80303, U.S.A. (Orders: 720-406-8849, fax 720-
406-3443). $14.95, 272 pp., 03 b/w photos, 2 maps, 6" x 9"
SIDA 20(3): 1340. 2003
ANNOUNCEMENTS 1341
THE RUPERT BARNEBY AWARD
The New York Botanical Garden is pleased to announce that Jason Alexander,
currently a graduate student in the Department of Botany & Plant Pathology,
Oregon State University, is the recipient of the Rupert Barneby Award for the
year 2003. Mr. Alexander will be studying the systematics of various species of
Astragalus in the western United States.
The New York Botanical Garden now invites applications for the Rupert
Barneby Award for the year 2004. The award of US$ 1,000.00 is to assist research-
ers to visit The New York Botanical Garden to study the rich collection of
Leguminosae. Anyone interested in applying for the award should submit their
curriculum vitae, a detailed letter describing the project for which the award is
sought, and the names of 2-3 referees. Travel to the NYBG should be planned for
sometime in the year 2004. The application should be addressed to Dr. James L.
Luteyn, Institute of Systematic Botany, The New York Botanical Garden,
200' Street and Kazimiroff Blvd., Bronx, NY 10458-5126 USA, and received no
later than December 1, 2003. Announcement of the recipient will be made by
December 15.
Anyone interested in making a contribution to THE RUPERT BARNEBY
FUND IN LEGUME SYSTEMATICS, which supports this award, may send their
check, payable to The New York Botanical Garden, to Dr. Luteyn.
THE DELZIE DEMAREE TRAVEL AWARD
Graduate students in plant systematics are eligible to apply for the Delzie
Demaree Travel Award, a $250.00 stipend to help defray expenses related to at-
tendance at the Missouri Botanical Garden 50‘ Annual Systematics Sympo-
sium (10-11 Oct 2003). The 2003 symposium: Species Reconsidered: Conse-
quences for Biodiversity and Evolution.
The application should include a letter from the applicant telling how sym-
posium attendance will benefit his/her graduate work and letter of recommen-
dation sent by the major professor. Please mail letters of application to: Dr. Donna
MLE. Ware, Herbarium, Biology Department, The College of William and Mary,
Williamsburg, VA 23185, U.S.A.
Anyone interested in making a contribution to Delzie Demaree Endowment
Fund, which supports the travel award, may send their check, payable to Bo-
tanical Research Institute of Texas, to Barney Lipscomb, 509 Pecan Street, Fort
Worth, TX 76102-4060, U.S.A. Thank you.
SIDA 20(3): 1341. 2003
Te lavishly illustrated monograph is the most
comprehensive botanical treatment of beans to
date. It begins with a brief history about the former
taxonomical treatments of the genus, and goes on
with the taxonomical criteria and a presentation
about discriminant characteristics. It presents a full
description of each section and species, its distribu-
tion and habitat, relationships with other species,
uses and potentially useful traits, and historical notes.
Color pictures, line drawings, and distribution maps
lead easily to the right identification of each species.
Available from
Botanical Research Institute of Texas Press
509 Pecan Street
Fort Worth, TX 76102-4060, USA
E-mail: sida@brit.org = Fax 1-817-332-4112
www.brit g [si bm23toc.htm
Sida, Bot. Misc. No. 23, 2002
issn 0833-1475, isbn 1-88878-11-1+7" « 10", xviii + 300 pp,
97 b/w figures and distribution maps, 5 color plates (60 figs.).
$40 + p&h = (USA $4; international $8)
Texas residents add $3.30 sales tax.
Taxonomy, Distribution,
and Ecology
of the Genus Phaseolus
(Leguminosae-
Papilionoideae)
in North America, Mexico
and Central America
By
George F. Freytag
Daniel G. Debouck
he Atlas of Texas covers about 6000 taxa. This is
Ee result of 54 years of herbarium and fieldwork
by B.L. Turner, beginning in 1948 at Sul Ross State
University, Alpine, Texas. In short, the senior author
has examined personally, touched, or “pored over”
an estimated several hundred thousand sheets in
the preparation of the forthcoming Atlas volumes.
Contents include an introduction, atlas of Texas
plants arranged alphabetically by family, by genus,
by species, and an index.
Available from
Botanical Research Institute of Texas Press
509 Pecan Street
Fort Worth, TX 76102-4060, USA
E-mail: sida@brit.org = Fax 1-817-332-4112
www.brit.org/sida/sbm/sbm24toc.htm
Sida, Bot. Misc. No. 24, 2003, issn 0833-1475,
isbn (vol. 1) 1-889878-08-1 = isbn (vol. 2) 1-889878-09-X = 7 1/2" < 10"
Vol. 1 approx. 630 pp.+ Vol. 2 approx. 275 pp.
Vol. 1 $50 + p&h* = Vol. 2 $40 + p&h* = Set $80 + p&h*
*USA: $10 (vol. 1); $9.50 (vol. 2); $12 (set)
*International: $12.50 (vol. 1) «$11.50 (vol. 2); $25 (set)
Texas residents add sales tax: $4.13 (vol. 1) = $3.30 (vol. 2) « $7.43 (set)
Atlas of the
Vascular Plants
Atlas of the
Vascular Plants of Texas
By
B.L. Turner,
Holly Nichols, Geoffrey
Denny, Oded Doron
6
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Barney L. Lipscoms, EDITOR
Botanical Research Institute of Texas
509 Pecan Street
Fort Worth, Texas 76102-4060, USA
817 332-444] / 817 332-4112 FAX
Electronic mail: sida@brit.org
Home page at the URL: http://www.brit.org/sida/
SIDA CONTRIBUTIONS TO
BOTANY WAS FOUNDED BY JOHN W. THIERET, ASSOCIATE EDITOR
Ltoyp H. SHINNERS (LEFT) Dept. of Biological Sciences
Northern Kentucky University
IN 1962. INHERITED BY
Highland Heights, KY 41099 USA
WILLIAM F. MAHLER
(RIGHT), DiREcToR EMERITUS FELIX LLAMAS, CONTRIBUTING SPANISH EDITOR
of BRIT in 1971, AND Dpto. de Botanica, Facultad de Biologia
Universidad de Leon
SINCE 1993, IT HAS BEEN ,
E-2471 Leon, SPAIN
PUBLISHED BY Brit Press.
Do
Guidelines for contributors are available upon request
and on our SIDA home page as well as the last pages
of each volume.
Subscriptions for year 2004:
$35. Individual
$60. USA Institutions
$70. Outside USA
numbers issued twice a year
{ )
; | | | |
| } | | )
« SIDA, ¢ ONTRIBUTIONS TO BOTANY
Votume 20, NUMBER 4, PAGES 1343-1836
22 December 2003
COPYRIGHT 2003
BOTANICAL RESEARCH INSTITUTE OF TEXAS
BRIT PravTeD I THE UNITED STATES OF AMER
PROS
ISSN 0036-1488
Szda, Contributions to Botany volume 20 (in 4 numbers)
ISSN 0036-1488
Copyright 2002, 2003
By
Botanical Research Institute of Texas
509 Pecan Street
Fort Worth, Texas 76102-4060, USA
http://www. brit.org/sida
DATES OF PUBLICATION
No. 1, pp. 1-460: 19 Jul 2002
No. 2, pp. 461-876: 23 Nov 2002
No. 3, pp. 877-1341: 29 Jul 2003
No. 4, pp. 1343-1836: 22 Dec 2003
Index, included with number 4 2003
For contents, see index or the unnumbered pages
forming the front cover of the separate issues.
SIDA
CONTRIBUTIONS
TO BOTANY
VOLUME 20
2002—2003
TABLE OF CONTENTS
SYSTEMATICS
]
Castilleja albobarbata Sp. nov. (Orobanchaceae) from Sierras
Manantlan and Cacoma, Jalisco, Mexico
Hucu H. Ittis, Guy L. Nesom, and J. MARK EGGER
Ipomoea tacambarensis (Convolvulaceae)
de Michoacan, México
ELFAZAR CARRANZA GONZALEZ
especie nueva del estado
Portulaca matthewsii (Portulacaceae), a new species from the
“Sierra Gorda” Biosphere Reserve, Querétaro, Mexico
GILBERTO OCAMPO ACOSTA
A new species of Chaptalia (Asteraceae: Mutiseae) from Mexico and
rediscovery of Chaptalia mexicana
LeticiA CABRERA R. and Guy L. Nesom
Taxonomy of the Polemoniaceae: the subfamilies and tribes
VERNE GRANT
A synopsis of Machaeranthera (Asteraceae: Astereae), with recog-
nition of segregate genera
Davip R. MorGan and RONALD L. HARTMAN
A new species of Arida (Machaeranthera sect. Arida— Asteraceae:
Astereae) from Trans-Pecos, Texas
B.L. Turner and Guy L. Nesom
Further additions to the genus Ardisia subgenus Graphardisia
(Myrsinaceae)
Jon M. RICKETSON and JOHN J. Pipoty HI
A revision of the Psidium salutare complex (Myrtaceae)
Lesuizt R. LANDRUM
Revision and typification of some species of Ixora (Rubiaceae) from
Central and Southern Brazil
Piero G. DELPRETE
A new section of Baccharis (Asteraceae: Astereae), and notes on
allied taxa
DanieEt A. GIULIANO and Guy L. Nesom
Two new combinations in Bolivian Gentianella (Gentianaceae)
JAMES S. PRINGLE
Botanical summary of a lowland ultrabasic flora in Papua New Guinea
W. TAKEUCHI
New combination in Eurybia (Asteraceae: Astereae) from North
America
Luc BROuILLET
1343
1351
1357
1363
1371
1387
1417
1423
1449
1471
1481
1485
1491
1561
A new species of Brazoria (Lamiaceae) from the central mineral
region of Texas
Matt W. TurNER
Two new species of Liatris series Graminifoliae (Asteraceae:
Eupatorieae) from the southeastern United States
Ropert Krat and Guy L. Nesom
New combinations in Xanthisma (Asteraceae: Astereae)
Guy L. Nesom
A new species of Solidago (Asteraceae: Astereae) from north
central Alabama
BRIAN R. KEENER and Ropert KRAL
Solidago faucibus (Asteraceae: Astereae), a new mesic forest
goldenrod from the Appalachian Mountains
Tuomas FE Wirsotpt and JOHN C. SEMPLE
New names and combinations in goldenrods, Solidago (Asteraceae:
Astereae)
JOHN C. SEMPLE
Pappus variation in Solidago (Asteraceae: Astereae)
JENNIFER L.A. Hoop and JOHN C. SEMPLE
Typilication, taxonomy, and a new subspecies of Polygonum
sawatchense (Polygonaceac) from North America
Minal Costea and FRANCOIS J. TARDIF
Typification and nomenclatural history of Trachelospermum
difforme (Apocynaceae)
ALEXANDER KRINGS
New names and new combinations in the flora of Hispaniola
HENRI ALAIN LIOGIER
Glandularia verecunda (Verbenaceae): species or fruit-form?
B.L. TURNER
New combination in Astereae (Asteraceae)
Guy L. Nesom and Haroi_p RopiNson
DEVELOPMENT AND STRUCTURE
Anatomia comparada de la epidermis foliar en el grupo de especies
Solanum sessile, de la secci6n Geminata (Solanaceae)
CARMEN BENITEZ DE RojAS and CHIARA BERLINGIFRI
DOCUMENTED CHROMOSOME NUMBERS
Documented chromosome numbers. 2003: 2. Chromosome num-
bers of the rare Texas species, Arida blepharophylla and A. mattturneri
.s
(Asteraceae: Astereae)
A. MICHAEL POWELL
1565
1573
1585
1589
1595
1605
1617
1631
1641
1645
1647
1649
1651
1663
FLORISTICS AND ECOLOGY
Aspectos ecoldgicos y estado de conservacion de Mammillaria
theresae (Cactaceae
IRMA LORENA LOPEZ-ENRIQUEZ, M. SOCORRO GONZALEZ-ELIZONDO,
MARTHA GONZALEZ-ELIZONDO, and JORGE ALBERTO TENA-FLORES
Type localities of vascular plants first described from Ohio
RoNALD L. Stuckey and JAMEs S. PRINGLE
The east-west transition of the flora in Texas: a biogeographical
analysis
MicHaet H. MacRoserts and Barsara R. MACROBERTS
Rediscovery of Tephrosia jamnagarensis (Fabaceae), an endangered
and narrow endemic plant species of Saurashtra, Gujarat, India
PS. NaGar, S.J. Sata, and S.J. PATHAK
Does Polygonum patulum (Polygonaceae) grow in North America?
Minat Coste and Francois J. TARDIF
Polygonum aviculare subsp. rurivagum (Polygonaceae) in North
America
Minat Costea and Francois J. TARDIF
Uvularia (Liliaceae) in Texas
Jason R. SINGHURST, WILLIAM FE Carr, WENDY J. LEDBETTER, and
WALTER C. HOLMES
Fagopyrum esculentum (Polygonaceae), new for Texas
Ropert J. O7KENNON, CAREN McLemore, and AMANDA K. NEILL
Triglochin concinna (Juncaginaceae), a new family, genus, and
species for Texas
B.L. TURNER and Matt W. TurNER
Andropogon glomeratus var. glaucopsis (Poaceae: Andropogoneae)
documented in Louisiana
Davin J. Rosen and JosEPH K. WIPFF
Carex conoidea (Cyperaceae) new to Arkansas and notes on its
occurrence in Arkansas and Missouri
Paut M. McKenzir, Tim Smit, and C. THEO WITSELL
Additions to the flora of Cleveland County, Arkansas: collections
from Moro Bottoms Natural Area, a state-protected old-growth
bottomland hardwood forest
DANNY SKOJAC, MARGARET S. DEVALL, and BERNARD R. PARRESOL
Arkansas flora: additions, reinstatements, exclusions, and re-
exclusions
James H. Peck
1665
1677
1693
1701
1707
1709
1713
1717
1721
1723
1727
1731
1737
Northward range extension in Florida of the invasive fern Lygodium
microphyllum (Lygodiaceae)
Rospert W. PEMBERTON
1759
Muhlenbergia dubia (Poaceae) in central Texas
GEOFFREY C. DENNY
1763
Rotala rotundifolia (Lythraceae) new to Florida
KATHLEEN C. Burks, VERNON V. VANDIVER, JR., DaviD W. HALL, and
COLETTE
COMMEMORATIVE
C. JACONO
1765
In memoriam: Chester Morrison Rowell, Jr. (1925-2003)
Jim BLASSINGAME
1771
Book reviews AND notices 1362, 1370, 1386, 1448, 1470, 1560, 1564, 1572, 1584, 1594,
1604, 1650, 1662, 1664, 1676, 1706, 1712, 1726, 1758, 1762, 1770, 1773
Sida, Contributions to Botany Update
Reviewers for Volume 20, 2002-2003
Index to volume 20, 2002-2003
Titles of Articles With Authors
Authors
Botanical Names
Subject
New Names and New Combinations
Sida Guidelines for Contributors
Index to new names and new combinations in Sida 20(4), 2003
Ardisia nigrescens subsp. donnellsmithii
ae z) Ricketson . Pipoly, comb.
stat. nov—143
Avdizis verdisepala eee S& Pipoly,
sp. nov—14
Arida = R. Morgan & ay
mb. et stat. nov—1
a (R.C. a. & RR.
oon D.R. Morgan & R.L.
ae comb. nov—1413
Arida blepharophyll (A. Gray) DR.
Mo R.L. Hartman, comb.
oe
Arida carnosa (A. Gray) D.R. Morgan &
R.L. Hartman, comb. nov—1413
Arida coulteri (A. Gray) D.R. Morgan &
R.L. Hartman, comb. nov—1414
Arida crispa (1T.S. Brandegee) DR.
Morgan & R.L Hartman, comb.
nov.—1414
Arida mattturneri -e L.
sp. nov.—14
oo
Turner & G. Nesom,
Arida parviflora (A. Gray) D.R. Morgan
&R.L. Hartman, comb. nov—1414
Arida i aoe (Kunth) D.R. Morgan @ R.L.
man, comb. nov—1414
ones see (M.L. Arnold & R.C.
kson) D.R. Morgan & R.L.
k fecetes comb. nov—1414
Asteraceae tribe Astereae subtribe
Bellidinae (Cass. e
Nesom & Ac Robinson, ‘comb. et
stat. nov.—
Baccharis sect. tae alae Giuliano &
G.L. Nesom, sect. nov—1481
Brazoria enquistii MW. Turner, sp. nov.
—1
a as Iltis @ Nesom, sp.
ov.— 1343
Glapialia hidalogoensis a Cabrera &
Nesom, sp. nov.—
Dieteria asteroides var. gandulos (B.L.
Turner) D.R. an oi RL.
Hartman, comb. nov—139
Dieteria asteroides var. lagunensis (D.D.
Keck) D.R. Morgan R
man, comb. nov—1393
Dieteria bigelovii (A. Gray) D.R. Morgan
& R.L. Hartman, comb. nov.
4
3
Dieteria bigelovii var. commixta (Greene)
D.R & R.L. Hartman,
4
ar
Dieteria canescens var. ambigua (B.L.
Turner) D.R. Morgan & RL.
Hartman, comb. nov.—1396
Dieteria canescens var. aristata
(Eastwood) D.R. Morgan & RL.
Hartman, comb. nov—1397
Dieteria canescens subsp. glabra a
ray) D.R. Morgan
Hartman, comb. nov—1396
Dieteria canescens var. glabra (A. Gray)
rgan & R.L. Hartman, comb.
nov.—1397
Dieteria canescens var. incana (Lindl.)
Morgan @ R.L. Hartman, comb.
nov.—1396
Dieteria canescens var leucanthemifolia
Greene) D.R. Morgan L
Hartman, comb. nov—
Dieteria canescens var. nebraskana (B.L.
Tu ner) D.R. Morgan @ RL.
artman, comb. nov—1397
Dieteria canescens var. ee iles (Nutt.)
DR n@&@RL.Hartman,comb
Dieteria canescens Val. shastensis (A.
Gray) D.R. Morgan 1
Hartman, comb. nov—1396
Dieteria canescens var. ziegleri (Munz)
.R. Morgan @ R.L. Hartman,
comb. nov—1396
Eurybia glauca var. pulchra (SF Blake)
Brouillet, comb. et stat. nov—1562
Gentianella boliviana oo JS. Pringle,
comb. nov—14
Gentianella nacauialy Secs ae ex
‘J e, comb. no
Gia verecunda ae ve
(Moldenke) B.L. Turner, comb.
Ipomoea tacambarensis EF. Carranza, sp.
nov.—1
Leandra urbaniana (Alain) Alain, comb.
nov.— 1645
Liatris patens Nesom & Kral, sp. nov.
—1579
Liatris savannensis Kral & Nesom, sp.
nov—1574
Matelea monticola Alain, nom. nov.
—1645
Maytenus urbanii Alain, nom. nov.
45
Polvoann tcl oblivium
sees & Tardif ces nov.
—1
7
Portulaca snail G. Ocampo, sp.
nov—1
Psidium eee var. decussatum (DC.)
Landrum, comb. nov—1463
Psidium saluta natum
(Cambess.) es comb. nov.
—1463
Psidium salutare var. pohlianum (O.
Berg) Landrum, comb. nov—1466
Psidium salutare var. sericeum (Cambess.)
Landrum, comb. nov—1467
Psychotria bulilimontis Takeuchi, sp. nov.
1514
i, Var. nov—
Sagraea rugosa Alain, nom. nov.—1645
Psychotria bulilimontis var. aestuarii
Takeuc 19
(Rydb.) 1C Semple, comb. et stat.
nov.—1606
Solidago arenicola B.R. Keener & Kral, sp.
nov.—1589
Solidago ser. Auriculatae Semple, ser. nov.
—1606
pardazee ser, ne Semple, ser.
no
Solidago faucibus Wieboldt, sp. nov.
pune ee )
oe
Solidago eae
mple, ae stat.
esis kralii eae sp.n
Solidago lepida subsp. fallax Fernald)
Semple, stat. nov—
Solidago lepida var. ae te peo] Se
emple, comb. nov—1
Solidago subsect. parade Juz.)
Semple, comb. nov—160
Solidago adona subsp. chapmanii (A.
ray) iG, Seranle comb. nov.
Salidago ser. Odorae (Mackenzie in
Small) Soni. con et Stat. nov.
—1606
aa ae ee strictula (Torr. &
y) J.C. Semple, stat. nov.
Solidago oe subsp. pulverulenta
C. Semple, an et stat.
op
Var. cronquistiana
ey var. Eaaeie
Suitcase paisa subsp. azorica
(Hochst. ex Seubert) Semple,
comb. et stat. nov—1615
Solidago sempervirens subsp. mexicana
..) Semple, comb. et stat. nov.
clean ru
-
Se)
—1615
Solidago speciosa subsp. pallida (Porter)
Semple, comb. et stat. nov. 5
Sohdago:: ie subsp. sencilla, (Torr.
ray) Semple, comb. et stat.
nov.—
Solidago velutina subsp. californica
Nutt.) Semple, comb. et stat.
¢ 1
Solidago velutina subsp. sparsiflora (A,
ae Semple, comb. et stat. nov.
—1616
Trichadenia sasae Takeuchi, sp. nov.
—1522
Xanthisma arenarium (Benth.) D.R.
Morgan = R.L. Hartman, comb.
nov.—
Xanthisma ple chiacsplillaee (A. Gray)
D.R. Morgan & R.L. Hartman,
comb. nov—1403
Xanthisma section Blepharodon (DC.)
. Morgan @ R.L. Hartman,
com b. nov—1401
Xanthisma coloradoense (A. Gray) D.R.
Morgan & R.L. Hartman, comb.
403
nov.—1
Xanthisma crutchfieldii (B.L. Turner) D.R.
Morgan & R.L. Hartman, comb.
nov.— 1403
Xanthisma glaberrimum (Rydb.) Nesom
)Kennon, comb. nov.—1586
Xanthisma gracile (Nutt.) D.R. Morgan &
R.L. Hartman, comb. nov—1405
Xanthisma grindelioides (Nutt.) D.R.
organ & R.L. Hartman, comb.
nov—1403
Xanthisma grindelioides var. depressum
(Maguire) D.R. Morgan @& R.L.
Hartman, ee nov.— 1404
Xanthisma gymnocephalum (DC.) D.R.
Morgan & RL Hartman, comb.
nov.— 1404
Xanthisma gypsophilum (B.L. Turner)
D.R. Morgan & R.L. Hartman,
comb. nov—1404
Xanthisma section Havardii (R.C. Jack-
son) D.R. Morgan & R.L. Hartman,
comb. nov—1410
= incisifolium (1 : eee
om, comb. nov—
eaniae eee SE Bla ke) DLR.
M one 7 R.L. Hartman, comb.
nov—
Xanthisma j a (Greene) D.R. Morgan
@ R.L. Hartman, comb. nov.
—1406
Xanthisma restiforme (B.L. Turner) D.R.
Morgan & R.L. Hartman, comb.
nov.— 5
Xanthisma ccna (M.C.
ee & RL. Ha
comb. nov—1405
Xanthisma section Sideranthus (Nutt. ex
Nees) D.R. Morgan @R.L. Hartman,
comb. nov.—1405
Xanthisma spinulosum var. chihuahuanum
(B.L. Turner & R.L. Hartman) DR.
Morgan . R.L. Hartman, comb.
nov.—
aay ee se var. glaberrimum
ydb.) D.R. Morgan & RL.
ve tman, comb. nov—1408
ear spinulosum var. gooddingii
A. Nelson) D.R. ee = RL.
ee tman com z nov—
Xanthisma s m vat. Garin
cv onaston): D.R. Morgan & RLL.
Hartman, comb. nov—1409
Xanthisma spinulosum var. par Sa xum
(B.L. Turner “ R. = Hartman) D.R.
Morgan — res oa
nov.—
Xanthisma a on var. scabrellum
(Greene) D.R. Morgan & RL.
Hartman, comb. nov—-1409
Xanthisma spinulosum (Pursh) D.R.
Morgan & R.L. Hartman, comb.
nov.— 1406
Xanthisma stenolobum (Greene) D.R.
Morgan : R.L. Hartman, comb.
nov.—
eae aac ere (Wooton &
tandley) D.R. Morgan & RL.
eee comb. nov v—-14
Xanthisma wigginsii (S.F Blake) D.R.
Morgan & R.L. Hartman, comb.
nov—1409
ohnston)
rtman,
a
CASTILLEJA ALBOBARBATA SP. NOV. (OROBANCHACEAE)
FROM SIERRAS MANANTLAN AND
CACOMA, JALISCO, MEXICO
Hugh H. IItis Guy L.Nesom
Department of Botany Botanical Research Institute of Texas
University of Wisconsin 09 Pecan Street
Madison, WI 53706, U.S.A. Fort Worth, TX 76102-4060, U.S.A.
J.Mark Egger
Un iversity of ‘ashington Herbaritym
Box 35532
Seattle, WA 98195-5325, U.S.A.
ABSTRACT
“1, ]f£ ] 1443 . e Jon 14
A new species, Castilleja albobarbata Iltis @ Nesom, is d Manantlan
and Sierra Cacoma of Jalisco, México. It differs ae C integrifolia primarily in features of vestiture
and is geographically disjunct from the nearest populations of C. integrifolia by about 450 kilometers.
Kry Worps: Ca stilleja Orobanchaceae. S pl lari Sierra de M tlan, Sierra Cacoma, Jalisco
México, biogeography, rare plants
RESUMEN
Se describe una a sobens nueva, Castilleja albobaxbata oa . Nesom, ce a sleita de eee y
a
Sierra C Meéxico.$
del indumento - esta cena pecan tiesmentena unos 5 450 niomedas de la ob acion mas proxima
de C. integrifolia.
oe albobarbata IItis @ Nesom, sp. nov. (Figs. 1-3) Tyr: MEXICO. JALISCO:
Mpio. Cuautitlan, [Sierra de Manantlan Biosphere Reserve], top of sharp crest of
the Sierra de Manantlan eee E of Cerro Las Capillas, along highest point
of gravel road from Cerro La Cumbre to Fos Jataines: 19 km due SSE of El Chante
(19° 33'19" N, 104° 09' W), cliffs of basalti , 2800-2860 m
elev. 10 Oct 1980, H.H. Iltis 3227 with R.Guzmdn-M. (HOL OTYPE: WIS; ISOTYPES: LEX,
ZEA).
Castillejae integr . iae L.f. similis sed vestimento trichomatibus brevissimis antrorse appressis et
rhata differr
corollarum galea
Herbaceous perennials, usually with numerous (up to 15) erect to ascending or
sprawling stems originating from the common base, the stems 2-4(-7) dm tall,
basally woe), simple or few-branched in the upper half; stems, leaves, and
ular, moderately to sparsely invested with short (ca. 0.05-0.1 mm
long) white, flattened, antrorsely ascending-appressed hairs. Leaves yellow-
green, spreading to slightly deflexed, entire, not clasping or auriculate, those of
the lower stems linear to narrowly lanceolate, 25-45 mm long, 2-4 mm wide,
SIDA 20(4): 1343-1350. 2003
1344 BRIT.ORG/SIDA 20(4)
Fic 2 Fl fF net iat Ihak jae ee llorti )
ILTIS ET AL., A NEW SPECIES OF CASTILLEJA FROM MEXICO 1345
® Castilleja albobarbata
© Castilleja integrifolia
Fic. 3. Distributi f Castill i i ifolia in Mexi iC. albobarbata. Modified after Nesom 1992.
becoming narrowly ovate-lanceolate near the inflorescence with a subclasping
base and up to6 mm wide, grading into the floral bracts. Floral bracts entire,
lanceolate, 15-35 mm long, markedly enlarged from the uppermost leaves, the
lower bracts completely green, the terminal persistent and conspicuous, mark-
edly differentiated from the lower, red and obovate to widely ovate. Flowers in
a racemose, somewhat secund inflorescence; pedicels 10-20 mm long, ascend-
ing; calyces 29-35 mm long, slightly gibbous abaxially, minutely hispid-stri-
gose, brick-red with a yellow-green apical margin or sometimes yellow at the
base, less commonly yellow-green throughout, the abaxial cleft 20-25 mm deep,
the adaxial cleft 2-5 mm deep, the primary lobes with 2 short teeth or acute
notches; corollas 40-45 mm long, the tube pale yellowish, becoming brick-red
near the lower lip; the galea yellowish with red flanges, 25-27 mm long, 5/8-5/
9 the length of the corolla, exserted (2-)7-15 mm from the calyx, producing a
long (ca. 12 mm), densely pilose, adaxial, toothbrush-like beard of multicellu-
lar, white trichomes 2-3.5 mm long, the corolla otherwise glabrous, the lower
lip composed of 3, small, thickened, greenish-yellow, incurved teeth 1-2 mm
1346 BRIT.ORG/SIDA 20(4)
long; stigma barely thickened, ca. 0.3 mm wide; anthers 2-2.5 mm long. Mature
fruits not observed.
Additional collections examined: MEXICO. Jalisco. Mpio. Cuautitlan: Sierra de Manantlan, area of
ne collection near Cerro Las Capillas, pine woods with Arbutus - actin Ae 2800 m, 31
g 1986, Cuevas 1617 (WIS) alone 1980, Iltis et al. 2507 (WIS - Sierra de Manantlan, ca. 15
mi of Autlan by way of Chante, rocky outcrops in cy press-pine — ft, 30 Jul 1949, Wilbur
Wilbur 1989 (MICH, NY); Mpio. Cuautitlan: Sierra de Manantlan, rocky summit of canyon 3 mi
‘| Guizar, 9300 ft [near type locality?], flowers red, white, and green, 23 Nov 1968, Boutin and
Meo t 2550 (MICH); Mpio. moe de Allende: [Sierra Cacomal, 4 km due SE of Cuale along dirt road
between El Tuito turnoff (5 km N of El Tuito and 32.2 mi by road E of Mex. Hwy 200 to Minas de
Zimapan) and Talpa de Allende, rare, one plant, with Begonia and Prochnyanthes mexicana atop large
rock outcrop in pine forest, 20° 22’ N, 105° 04' W, 1920 m (altimeter), 18 Aug 1993, Cochrane et al.
13136 (IBUG, TEX, WIS, ZEA)
5
The closest relatives of Castilleja bata are within a group of species iden-
tified as Castilleja subg. Castilleja by Holmgren (1976) and sect. Castilleja by
Nesom (1992). These species are characterized by (1) a racemose inflorescence,
(2) colored calyces but usually green floral bracts, G) calyces with a deeply cut
(anterior) abaxial cleft and a shallow (posterior) adaxial one, and (4) corollas
with the galea equalling or longer than the tube. A key to most of the Mexican
taxa with this character constellation (including “white-bearded Castilleja Itis
& Nesom ined.” = C. albobarbata), as well as detailed discussion and distribu-
tion maps for many of them, were provided in Nesom (1992).
Within sect. Castilleja, C.albobarbata is most similar to C. integrifolia. This
pair is distinguished among its relatives by long pedicels 10-20 mm long), a
dense beard on the upper side of the galea, and loose, racemose inflorescences
terminated by a cluster of colored bracts. The new species differs from C.
integrifolia in its short, ascending-appressed (antrorsely oriented) stem hairs,
compared to the cauline vestiture in C. integrifolia of slightly longer (mostly
0.2-0.4 mm long), deflexed (retrorsely oriented) and closely appressed hairs.
The orientation of stem hairs is consistent within both species, although it may
be variable in related species (e.g., Castilleja tenuiflora Benth.). The galea of C.
albobarbata produces a beard similar in density and trichome length to that of
C. integrifolia, but the beard of the new species is bright white and a striking
recognition feature.
Castilleja integrifolia is a wide-ranging and variable species, distributed
from the paramos of the Andes Mountains in Colombia and Venezuela, broadly
through the mountainous regions of most of Central America and southern
Mexico, and on northward in the Sierra Madre Oriental of northeastern Mexico
as far as northern Nuevo Leon and Coahuila. A map of its distribution in Mexico
is shown in Nesom (1992, Map 2). The type specimen was collected in Colom-
bia (Mutis 82, holotype: LINN, fiche!). Within its broad range, C. integrifolia is
fairly common in appropriate habitat and mostly consistent in its basic mor-
phological features. Regional variants exist, however, and these are incompletely
ILTIS ET AL., A NEW SPECIES OF CASTILLEJA FROM MEXICO 1347
understood. One taxon, described as C. chiapensis Brandegee, is based on plants
from the mountains of central Chiapas, having a densely villous vestiture of
golden-yellow hairs. A second form, apparently endemic to central Oaxaca and
described as C. longibracteata Mart. & Gal. is distinguished by its broadly obo-
vate, limbriate-tipped apical bracts. Type material for another variant, C.
integrifolia var. alpigena L. Williams from Guatemala, appears to be referable
to C. pectinata Mart. & Gal. The range of C. integrifolia in Mexico most closely
approaches that of C.albobarbata in central Guerrero and Edo. México, across a
gap of about +50 kilometers. The morphological distinction between the two
taxa is consistent but not great - recognition of C. albobarbata at specific rank
emphasizes its wide geographic disjunction from C. integrifolia.
This taxonomic disposition is further supported by the discovery of
Castilleja albobarbata at the far northwestern end of Sierra Cacoma, an exten-
sive, two-parted, southeast-to-north west oriented mountain range of the Sierra
Madre del Sur that begins where Sierra de Manantlan ends at Puerto Los Mazos.
This locality is about 33 kilometers southeast of Puerto Vallarta and extends
the range of C. albobarbata more than 110 kilometers northwest of the area of
the type locality at Cerro Las Capillas. It also suggests that other stations of
this rare species may yet be found at high elevations in between, in that vast
area of rough topography still undercollected despite many years of concerted
botanical activity.
The type locality of Castilleja albobar bata is most remarkable floristically
and ecologically (Vazquez-G. et al. 1995). Perched atop an enormous, sheer and
windswept, south-facing cliff, this crest of the central section of Sierra de
Manantlan forms the very edge of the Sierra Madre del Sur and reaches its high-
est elevation (2800-2860 meters) at Cerro Las Capillas. Below, one can see on
any clear evening, far beyond the parallel ridges of ever-lower hills, the shim-
mering Pacific Ocean at Manzanillo, some 60 kilometers to the south. Be it due
to the near continual roaring winds from off the Pacific coast or the high alti-
tude, be it the dry crumbly sandstone conglomerate or evidence of frequent
fires, the vegetation on the crest is one of an open, burnt-over pine-oak parkland
(Pinus hartwegii Lindl. sensu lato, see comments below), the scattered trees
understoried by ericaceous shrubs and tussock grass (“sacaton”), giving the
appearance of a Mexican subalpine “sacatonal.” The low, evergreen ericaceous
scrub consists of Pernettia prostrata (Cav.) DC., Vaccinium confertum Kunth,
Comarostaphylos discolor (Hook.) Diggs subsp. manantlanensis Diggs, and the
remarkable diminutive Arbutus occidentalis McVaugh & Rosatti, the latter two
here at their type locality. Amongst our scattered and rare Castilleja are small
patches of anas yet undescribed Alchemilla(A. manantlanensis Iltis, nom. prov)
seen nowhere else, and of the robust grass tussocks of a giant Agrostis known
apparently only from this one site. With fully 8 dm tall culms from a woody,
rhizome-like base, and with many, crowded, long leaves, A. novogaliciana
1348 BRIT.ORG/SIDA 20(4)
McVaugh is unusual in other ways as well. The author himself (McVaugh
1983:42) noted that “it is quite unlike any other Mexican species of Agrostis and
indeed may be out of place in that genus.”
On the gentler, cooler, moister, north-facing slopes, a few hundred meters
below the crest and less susceptible to fire, is a mixed conifer-deciduous cloud
forest of Abies religiosa (Kunth) Schltdl. @ Cham., Pinus hartwegii, Cupressus
lindleyi Endl, and Quercus spp. In its dense understory another Castilleja grows
in abundance, the yellow and red-flowered C. macvaughii N. Holmgren, ashrub
to 1.5 or even 2 meters tall, strictly endemic to the high elevation cloud forests a
few kilometers east and west of Cerro Las Capillas. Castilleja macvaughii shares
its cool habitat with the recently described, more widely distributed but al-
ways local Symplocos sousae F. Almeda and other endemics such as Ageratina
manantlanensis B.L. Turner. Other, more widespread Castilleja species, at
slightly lower elevations, include C. nervata Eastw. and C. pterocaulon N.
Holmgren.
Finally, even the pines at Cerro las Capillas present a taxonomic problem.
“Pinus hartwegii” sensu lato (as noted above) is listed in Vazquezet al. (1995:106)
as P. durangensis Martinez, according to the botanists R. Cuevas-G. and S. Car-
vajal-H. In the discussion, however, the local pine expert J. Pérez de la Rosa sug-
gested it to be P martinezii E. Larsen, while WB. Critchfield identified these
collections as an aberration of P. hartwegii. Considering the great local ende-
mism at this locality, it may yet prove to be an undescribed local variant, and
thus Critchfield correct.
The concentration of so many endemics on the slopes or at the very top of
Sierra de Manantlan, some 40 species in all (Vazquez et al. 1995; Hernandez
1995:72-8]), suggests that this mountain, like so many others in Mexico, acted
not only as a “survivium” (refugium) of a once more extensive montane and
subalpine flora during periods of glacial cooling and of its remnants in the
present post-glacial warming but, in addition, because of its isolation, as an
“Inselberg,” an island of autochthonous speciation, even if now only a few hect-
ares in extent.
Similarly, the Sierra Cacoma station for Castilleja albobarbata lies in the
center of a hotbed of endemism, which though only 30 to 40 kilometers south-
east to southwest of the vacation mecca of Puerto Vallarta, has until recently
remained relatively uncollected. Its exploration has been facilitated by the now
already deteriorating east-west gravel road that starts out (from Mexican High-
way 200) 5 kilometers north of El Tuito, goes east for 52 kilometers to the
Castilleja locality near the now abandoned Minas de Zimapan, and continues
beyond for 47 kilometers to Talpa de Allende. This “brecha” Tuito-Talpa opened
up since the 1960's vast, previously inaccessible, forested lands, from the sub-
tropical northern end of Sierra El Tuito (500-1880 m) south to the higher Si-
erra El Cuale (1000-2300 m) north of the road, and finally to the grand, rug-
ILTIS ET AL., ANEW SPECIES OF CASTILLEJA FROM MEXICO 1349
ged, isolated end of the Sierra Cacoma (with several peaks above 2600 m and
one to 2740 ml). Most of the collecting evidently occurred next to the road, as
with the single plant of Castilleja. Yet, with its rocky pine forest habitat con-
tinuing up the mountain slopes for another 1800 meters to equal the altitude of
the type collection, one can predict that it may yet reveal a much larger popula-
tion. And the cloud forests that probably crown the peaks also offer potentially
rich scientific rewards. In any case, all along the Minas de Zimapan road, these
pine to oak woods have yielded a cornucopia of rarities and recently described
local endemics: Quercus tuitensis L.M. Gonzalez and 9. cualensis L.M. Gonzalez
(Gonzalez-V. 2003a, b), Pinus jaliscana Perez de la Rosa, Muhlenbergia cualensis
Herrera & P. Peterson, and various novelties in the genera Aechmea, Agaristd,
Aristida, Triniochloa, Pedilanthus, Stevia, and Wedelia.
What the flora of the highest peaks may yet yield isa tantalizing question.
These are, after all, extensions of Sierra de Manantlan and equivalent in alti-
tude to that of Cerro Las Capillas (2740 vs. 2860 meters). Here is a real botani-
cal challenge to any young botanist, for soon it may be too late. Not only are
lumbering and cattle grazing expanding up the mountains to supply an ever
growing Mexican population, but global warming, world-wide, is a fact bota-
nists need to consider.
What indeed will be the fate of many of these high-altitude microcenters
of biodiversity, the “Inselbergs” of evolution, these biogeographic survivia, in
consequence of global warming? Surely many of these endemics and rarities
would soon be pushed off their high mountain perches by a warmer climate
and displaced by pine forests or even cloud forests advancing from below. But
as for the large number of rarities at Las Capillas, the type locality of Castilleja
albobarbata, their survival for the time being is assured by protection in the
Core Zone of the Sierra de Manantlan Biosphere Reserve (Guzman-M. and Iltis
1991; Halffter 1987; Vazquez et al. 1995).
py
ACKNOWLEDGMENTS
We are grateful to MICH for a loan of specimens, to Ted Cochrane for his excel-
lent label data, and Rich Spellenberg for his review of the manuscript. Field
work was supported by the Instituto Manantlan de Ecologia y Conservacion de
la Biodiversidad (IMECBIO), Universidad de Guadalajara, and the National His-
tory Museums Council, F.K. and O.N. Allen Herbarium Fund, and the Davis
Fund, all of the University of Wisconsin-Madison.
REFERENCES
Guzman-M.,R.and HH. litis. 1991. Biosphere reserve established in Mexico to protect rare
maize relative. Diversity 7:82-84.
GONZALEZ-VILLAREAL, LM. 2003a. Quercus tuitensis (Fagaceae, Quercus sect. Lobatae), a new
deciduous oak from western Jalisco, Mexico. Brittonia 55:42—48.
1350 BRIT.ORG/SIDA 20(4)
GONnzALEZ-VILLAREAL, L.M. 2003b. Two new species of oak (Fagaceae, Quercus sect. Lobatae)
from the Sierra Madre del Sur, Mexico. Brittonia 55:49-60.
Hacrrter, G. 1987.La reserva de la biosfera Sierra de Manantlan y la conservacion in situ de
los recursos bidticos. Revista Soc. Mex. Hist. Nat. 39:27-34.
Hernanoez-L.,L.1995.La flora endemica de al Sierra de Manantlan, Jalisco—Colima, Mexico:
Observaciones preliminares. In Vazques et al.,loc. cit., Flora de Manantlan pp. 72-81.
Hoimoren, N.H. 1976. Four new species of Mexican Castilleja (subgenus Castilleja,
Scrophulariaceae) and their relatives. Brittonia 28:195—208.
McVauch, R. 1983. Agrostis. Flora Novo-Galiciana (Gramineae) 14:40-43.
Nesom, G.L.1992.Taxonomy of the Castilleja tenuiflora group (Scrophulariaceae) in Mexico,
with an overview of sect. Castilleja. Phytologia 73:389-415.
VAzauez-G,., A., R. Cuevas R., 7.5. CocHrane, H.H. luis, FJ. SANTANA-M., and L. Guzman-H. 1995. Flora
de Manantlan: Plantas Vasculares de la Reserva de la Biosfera de Manantlan, Jalisco-
Colima, Mexico. Sida, Bot. Misc.. 13:1-315.
IPOMOEA TACAMBARENSIS (CONVOIVULACEAE),
ESPECIE NUEVA DEL ESTADO DE MICHOACAN, MEXICO
Eleazar Carranza Gonzalez
ea)
Instituto de oy A.C.
R
Cent Bajic
Pdtzcuaro, i h., MEXICO
carranza @inecolbajio.edu.mx
RESUMEN
Se describe como especie nueva y se ilustra a Ipomoea tacambarensis (Convolvulaceae), que se
encuentra en la depresion del Balsas en el estado de Michoacan, México. Esta planta, aunque de
OMOEA
—
apariencia arbustiva, es una hierba perenne que alcanza hasta 2.2 m de alto. Es similar a |
stans Cav., pero se distingue por su mayor tamano (de hasta 2.2 m de alto), las hojas mas grandes (de
hasta de 35 cm de largo) y las inflorescencias multifloras.
8
PALABRAS CLAVE: Ipomoea, Convolvulaceae, Michoacan, México
ABSTRACT
[pomoed tac cass nsis As onvolv nae en, eo ~ sees depression in the state of Michoacan,
a
Mexico, i it does not undergo consid-
erable secondary growth, and is thus re: ‘lies a oe Sena hen to22mt ale It is very similar to
[pomoea stans Cay, but it is distinguished by its greater s size (to 2.2 m tall vs. to lm sae renee? leaves
(
(to 35cm long vs. to 11 cm long) and many flowered inflorescence (vs. {1 y)
Key Worpbs: [pomoed, Convolvulaceae, Michoacan, México
En la vertiente hacia la Cuenca del Balsas, en el Eje Neovolcanico y dentro del
municipio de Tacambaro, Michoacan, se encontr6 recientemente una planta
herbacea vistosa y de porte erecto que pertenece al género Ipomoea
(Convolvulaceae). Esta especie crece en laderas y canadas con encinares, pinares
o con bosque tropical caducifolio. Comunmente forma parte de asociaciones
vegetales secundarias en pequenas areas dedicadas al pastoreo de ganado bovino,
oen lugares con cierto grado de alteracion y relativamente cercanos a caminos
y poblados.
Debido al tipo de habito tan conspicuo y por encontrarse relativamente cerca
de la carretera, es raro que esta planta haya pasado desapercibida por los botanicos.
Sin embargo, la zona ha sido poco explorada desde el punto de vista bioldgico,
lo que con seguridad ha limitado el conocimiento de varios hallazgos mas.
Ipomoea tacambarensis [:. Carranza, sp. nov. (Figs. 1 y 2). Tipo: MEXICO.
MICHOACAN. Municipio de Tacambaro: + 1 km al N de Paso de Morelos, canada
perturbada, con transicion de bosque tropical caducifolio y bosque de encino, 1550-
1600 m, 3 Sep 2002, FE. Carranza y V. Steinmann 6392 (HOLOTIPO: IEB; ISOTIPOS: para
distribuirse en MEXU, ENCB, XAL, EBUM).
SIDA 20(4): 1351-1356. 2003
1352 BRIT.ORG/SIDA 20(4)
Fic. 1. Ipomoea tacambarensis (basado en el ejemplar tipo: £. Carranza y V. Scena 6392 cane fs aspecto de una
a. B. Flor abierta, mostrando la disposicién de los estambres. C. Rama de ilo, con sépalos y
anes E. Semilla.
Differt a L stans. habitu ad 2.2 m alto: caule ad 3 cm diametro ific (30-)40 cm supra basin;
foliis (10-)16-30(-35) cm longis, (5-)12-24 cm latis, petiolis canaliculatis ad tt cm longis, ad : mm
diametro; inflorescentiis terminalibus vel axilla has ltiflori li go-ovatis, (13-)15-17
mm longis, 6-8 mm latis, aequalibus vel subaequalibus; stylo 3.5- 4c cm ee
Planta herbacea, erecta, de hasta 2.2 m de alto; tubérculo de 20 a 40 cm de largo
y de 10a 20 cm de diametro, tallos principales 3 a 5, algo angulosos, huecos, de
ARRAN A 1353
/
| /
pee = }
a
——
(ces
Fig. 2. Variacién en tamano de las hojas (A-B) y en la forma de los sépalos (a—b) de /} t is (A, a)e
Ipomoea stans (B, b).
hasta 3 cm de grosor cerca de la base, ramificados a (30-)40 a 50 cm arriba de la
base, ligeramente puberulentos; lamina foliar de contorno oblongo-ovado a
anchamente eliptico, de (10-)16 a 30(-35) cm de largo, de (5-)12. a 24cm de ancho,
apice redondeado a obtuso, base truncada a subcordada, margen profundo e
irregularmente eee odentado-lobulado, lobul dos, mucronulados
venacion conspicua en el envés, superficie puberulenta,a veces diminutamente
pilosa en el envés, primordialmente sobre las nervaduras, peciolo de (1.2-)2 a
4.5(-6) cm de largo, de 2 a 3(-5) mm de grueso, piloso, acanalado por el haz;
inflorescencias axilares y terminales, en cimas dicasiales y monocasiales, flores
(3-)5a 15, pedunculo primario de (6-)9 a 18(-20) cm de largo, de 2.a 3(-4) mm
de grueso, glabrescente o piloso, pedtinculos secundarios de 7 a 15(-25) mm de
largo, glabros, a veces pilosos, bracteas caedizas, linear-lanceoladas u oblongo-
1354 BRIT.ORG/SIDA 20(4)
ovadas, membranosas a escariosas, de 12 a 17 mm de largo, de hasta 6 mm de
ancho, margen erésulo, 0 a veces irregularmente ciliado, pedicelos de (8-)12 a
30 mm de largo, glabros; sépalos escariosos, iguales o inconspicuamente
subiguales, oblongo-ovados, de (13-)15 a 17 mm de largo, de 6a 8 mm de ancho,
los interiores ligeramente mas anchos, glabros, rara vez los exteriores apenas
pildsulos; corola infundibuliforme, de color rojo vino a azul-purpureo, de 7 a 9
cm de largo, de 6a 9cm de diametro en el limbo, el tubo mas palido que el limbo;
estambres desiguales, de 2 a 4 cm de largo, filamentos blancos, pubescentes en
la base, anteras de 5a7 mm de largo; estilo de 3.5a 4 cm de largo, glabro, blanco,
estigma globoso, algo bilobado; fruto una capsula subglobosa a anchamente
ovoide, bilocular, 4-valvada, de (8-)9 a 11 mm de largo, de (7-)8 a 9 mm de
diametro, glabra, de color pajizo a café oscuro; semillas 4, elipsoides, + 3-
anguladas, de color café, de (5-)6 a 8 mm de largo, de 4a 6mm de ancho,
diminutamente pubescentes.
Fjemplares examinados: MEXICO. Michoacan. Municipio de Tacambaro: + 8 km de Paso de Morelos,
camino a Villa Madero, 3 2003, E. Carranza y V. Steinnman 6397 (IEB); al N de Paso de Morelos, 3
Ago 2003, E. Carranza y P. Carrillo 6390
Debido al habito sui de esta sate las hojas sublaciniadas 0 irregularmente
dentado-lobuladas, fruto bilocular con semillas diminutamente pubescentes,
la especie debe incluirse en Ipomoea serie [yrianthinae (House) D. F Austin, al
lado de L. ancisa, I. sescossiand e I. stans (McDonald 2001).
El aspecto general de esta planta es similar al de Ipomoea stans Cav,
principalmente en lo que respecta al porte erecto y la forma de las hojas. Sin
embargo, I. tacambarensis tiene sépalos iguales o apenas subiguales, oblongo-
ovados, los interiores un poco mas anchos, hojas generalmente de 16 a 30 cm de
largo y 12 a 24 cm de ancho y cimas multifloras. En contraste I. stans posee
sépalos generalmente desiguales, los externos elipicos, los internos
angostamente elipticos, hojas de 2.5 a 7 cm de largo, de 0.6 a 3.cm de ancho y
flores solitarias. En el cuadro | se senalan con mayor detalle las diferencias en-
breestas especies.
Ecologia.—El area donde prospera I. tacambarensis tiene suelos de tipo rego-
sol y acrisol que se asientan sobre rocas de origen igneo, principalmente areniscas
y conglomerados, e1 es 1500 > 1950 m de altitud. Los climas subhtmedos tanto
calidos c enel sitio tipo. Las condiciones ambientales
de esta region se encuentran con diferentes grados de alteracion, ocasionados
basicamente por vias de comunicacion, agricultura y pastoreo. Por lo menos en
algunas zonas, estas plantas se ven favorecidas con dichas actividades.
Vegetacion.—La vegetacion donde crece esta especie varia desde la ecotonia
de los bosques tropical caducifolio y de encino, en las partes con menor altitud,
hasta el bosque de pino-encino en lo mas alto. En la primer zona destacan
especies de los géneros Quercus, Bursera, Colubrina, Annona, Cordia y Ficus.
Ademas se encuentran plantas propias de ambientes perturbados, por ejemplo
i 1355
Cuabro 1. Comparacion morfoldgica y distribucion geogréfica de /pomoea stans e Ipomoea
tacambarensis.
Ipomoea stans Ipomoea tacambarensis
Habito Hierba erecta, hasta 1 m de alto Hierba erecta, de mas de 2 m de alto
Tallo Hasta de 5 mm de grueso Hasta de 30 mm de grueso
Hoja Lamina de 2.5—7(-11) cm de largo, — Lamina de (10-)16-30(-35) cm de largo,
de 0.6—3(—5) cm de ancho, de (5—)12-24 cm de ancho, membranacea;
subcoriacea; peciolo de 2-4(-6) mm_ peciolo de (12-)20-45(-60) mm
de largo, de 0.5—-1 mm de grueso, de largo, de 2-3(-5) mm de grueso,
cilindrico acanalado
Inflorescencia Axilar; flores solitarias, rara vez en Terminal o axilar;cimas con (3-)5-15 flores:
pares; pedunculos de 0.8-5(-8.5) pedunculos de (6-)9- 0) cm de
cm de largo, de 1-2 mm de a largo, de 2—3(-4) mm de grt ; pedicelos
pedicelos de 5-12 mm de largo de (8-)12-30 mm de Rice
Sépalos Desiguales, a veces nee los Iguales 0 apenas subiguales, oblongo-
externos elipticos, de 10-23 mm ovados, de (13—)15-17 mm de largo, de
de largo, pubescentes 6-8 mm de ancho, los interiors ee
los internos angostamente elipticos mas anchos, glabros,rara vez los exteriores
pilosulos
Estilo 2.5—3 cm de largo 3.5-4 cm de largo
Distribucion Ampliamente distribuida en Restringida hasta donde se conoce, a una
regiones semidridas de México region semihumeda de Michoacan, en la
(Dgo., Zac,, S.L.P.,, Gto., Qro., Hgo., vertiente sur del Eje Neovolcanico
Jal., Mich., D.F., Mor., Pue., Tlax., Ver., hacia la Depresion del Balsas.
Oax.)
de los géneros Verbesina y Acacia, que manifiestan claramente la alteracion de
la vegetacion. La zona de mayor altitud, donde también destacan elementos
indicadores de disturbio, concentra como especies dominantes las que son
propias de estas comunidades sobresaliendo Pinus oocarpa y Quercus
magnon OH.
igual que otras especies de Ipomoea, a I. tacambarensis se le conoce con
el nombre de “quiebraplatos.” De acuerdo con los pobladores del lugar, tradi-
cionalmente, los tallos y hojas son hervidos y utilizados a manera de banos,
para contrarrestar los efectos de la picadura de animales venenosos (alacranes,
viboras y otros); acompanados frecuentemente por la ingestion de un té
preparado con el tubérculo de las mismas plantas.
El nombre de la planta alude al municipio de Tacambaro en el estado de
Michoacan, lugar donde crecen estas poblaciones.
Distribucién.—Hasta ahora, solo se conoce de la localidad tipo y en otro
sitio relativamente cerca de ésta. Se sabe ademas, por informacion de los
habitantes de la zona, que se encuentra en varias de las numerosas cafiadas y
i
1356 BRIT.ORG/SIDA 20(4)
laderas similares, aledanas al area de colecta. Es probable que se trate de una
planta endémica a esta region.
Fenologia.—Florece de agosto a septiembre y los frutos maduran algunos
meses después, encontrandose sus restos atin al principio de la floracion del
siguiente ano.
AGRADECIMIENTOS
Deseo expresar mi agradecimiento a J. Rzedowski por la revision critica del
manuscrito, en especial la diagnosis en latin, lo mismo que a A. McDonald y V.
Steinmann. Igualmente agradezco a este ultimo y a P. Carrillo por sucompania
durante las exploraciones de campo, asi como a R. Cardenas por la elaboracion
del dibujo. Finalmente, este trabajo ha sido posible gracias al apoyo econdmico
brindado por el Instituto de Ecologia, A.C. (cuenta 902-07).
REFERENCIA
McDonato, A. 2001. Revision of Ipomoea series Tyrianthinae (Convolvulaceae). Lundellia
PORTULACA MATTHEWSII (PORTULACACEAE),
A NEW SPECIES FROM THE “SIERRA GORDA”
BIOSPHERE RESERVE, QUERETARO, MEXICO!
Gilberto Ocampo Acosta
ituto de E Ecologia, A. C, Centro pia del Bajio
Pdatzcuaro, Michoacan, MEXICO
ee ee
ABSTRACT
Portulaca matthewsii is proposed and described as a new species. It belongs to subg. Portulaca sect.
Catoclasis subsect. Conocarpae, and has been collected in the northern pare va ne lia Borda”
Biosphere Reserve. in the Mexican state of Querétaro. The
Mexican species of this subsection by the fibrous ne by the srcllates tuberculate surface of the
seed.
Key worbs: Mexico, Portulaca, Portulacaceae, Querétaro, “Sierra Gorda” Biosphere Reserve
RESUMEN
gp]
aciencia. Est on se ubica
Se propone y describe a P
enel subgénero Portulaca, seccion Catoclasis, shea Conoc fears y ha sioaieeeine en a a porcion
norte de la Reserva de la Biosfera “Sierra Gorda”, en el estado mexicano de Querétaro. La entidad
nueva se diferencia de las especies de la subseccion conocidas para México por la presencia de raices
fibrosas, asi como por la superficie de la semilla que es estelulado-tuberculada.
PALABRAS CLAVE: México, Portulaca, Portulacaceae, Querétaro, Reserva de la Biosfera “Sierra Gorda”
During preparation of the treatment of Portulacaceae for the Flora del Bajio y
de Regiones Adyacentes, Mexico, some specimens of the genus Portulaca col-
lected in the “Sierra Gorda” Biosphere Reserve in the state of Querétaro were
detected to have a campanulate capsule operculum. This feature, in addition to
others, places the specimens in subsect. Conocarpae D. Legrand (subg. Portu-
laca sect. Catoclasis D. Legrand (Legrand 1962)). (Even though Legrand’s ap-
proach is focused mainly in the American species of Portulaca and does not
consider the variation presented by the genus ina global scale, his systematic
arrangement has been very useful in characterizing the New World Portulacas;
nevertheless, a modern revision for the whole genus is considered necessary ).
Beyond this placement, they could not be determined to species, so here they
are proposed as:
nald
Research stpported by the Instituto de Ecologia, A.C. (account number 902-07), by the Consej
Ciencia y Tecnologia a ee Mexico) and by the Comision Nacional para el Conocimiento y Uso de la
Biodiversidad (Conabio, Mexic
SIDA 20(4): 1357-1361. 2003
1358 BRIT.ORG/SIDA 20(4)
Portulaca matthewsii G. Ocampo, sp. nov. (Fig. 1). Type: MEXICO. QUERETARO. Mu-
nicipality of Landa: Puerto del Gato, 3 km al W de Acatitlan de Zaragoza, bosque
de encino, ladera de cerro, 1450 m, 29 Jul 1988, E. Gonzdlez 44 (HOLOTYPE: IEB;
ISOTYPES: MEXU, QMEX, XAL).
rassiuscula; radix fibrosa; caules usque ad 15 cm longi, prostrati vel decumbentes;
Herba ni
folia Aeaaa subteretia, lanceolata, seen lanceolata, oblanceolata vel elliptica, 3-10 mm longa,
0.6-2.5(-3) mm lata, apice acuta vel subacuta, aliquando obtusa; sepala deltata, rubra, (3-)3.5-4.5(-5)
mm longa, 2.5-4(-4.5) mm lata; petala ee purpurea, ee vel oblongo-obovata, (4~)5.5-
8 mm longa, (2.5-)3-4 mm lata; stamina (13-)16-28; stylus (3-)4-5 mm longus, lobis stigmatosis 3(-
4); capsula (2.5-)3-4(-4.5) mm longa, operculum campanulatum; semina reniformia, nigra vel
coracina, (0.5-)0.6(-0.7) mm longa, testa stellato-tuberculata.
Fleshy perennial herb with fibrous roots; stems several from the base, usually
suffrutescent in their lower portion, prostrate to decumbent, 4-15 cm long,
sparsely branched above, glabrous; axilar hairs scarce, 3-5(-6) mm long; leaves
alternate; petiole 1-1.5(-2) mm long; lamina lanceolate, oblong-lanceolate or
oblanceolate to elliptic, 3-10 mm long, 0.6-2.5(-3) mm wide, subterete in cross
section, apex acute to subacute, rarely obtuse; flowers clustered in a head in
groups of 4-7, subtended by an involucre of 6-8(-10) leaves, surrounded by hairs
3-4.5(-5) mm long: sepals reddish, deltate, clasping, (3-)3.5-4.5(-5) mm long,
2.5-4(-4.5) mm wide, apex acute, margin scarious; petals magenta to purple,
obovate to oblong-obovate, (4-)5.5-8 mm long, (2.5-)3-4 mm wide, apex ob-
tuse toemarginate, usually apiculate; stamens (13-)16-28; filaments connate at
the base and forming a tube 0.6-0.8 mm long, the filaments (2-)2.5-3.5 mm long;
anthers 0.4-0.6(-0.7) mm long; style (3-)4-5 mm long, stigma lobes 3(-4); fruit-
ing pedicel 0.5-0.7 mm long; capsule (2.5-)3-4(-4.5) mm long, 2-3 mm in di-
ameter, circumscissile in the lower third; operculum campanulate; seeds (0.5-)
0.6(-0.7) mm long, reniform, black to bluish-black, testa with tubercles stellate
at the base.
Additional specimens examined: MEXICO. Querétaro. Seana eae 3-4 km al N de L
Parada, 20 Jun 1991, B. Servin 1146 (EB), + 2 km de La Tinaja, camino Lindero, 15 Aug 1989, E.
al ss 1976 (EB). Municipality of Landa: + 0.5 km al SW de El Lobo, pe 2002, G. Ocampo & E.
2 (IEB); 1 km al SW de El Lobo, | Aug 1987, J. Rzedowski 44051 (ENCB, IEB), of a - al Ede
= ne por la ee a Xilitla, 25 Sep 2002, S. Zamudio & V. Steinmann 12125 (IEE g MEX
5 km (by road) E of the turnoff to La Vuelta, 25 Sep 2002, V. W. Steinmann & S. SZamaudio 2832
ea 12 kmal 7 7: El Lobo, sobre la carretera a Landa, Aug 1987, J. Rzedowski 43969 (ENCB, IEB).
Portulaca matthewsii is found in the municipalities of Jalpan and Landa,
Querétaro, in the northern portion of the “Sierra Gorda” Biosphere Reserve. This
region is part of the Sierra Madre Oriental mountains system, and it is highly
probable that the new entity also occurs in closely adjacent areas of the state of
San Luis Potosi. The new species grows in rocky open places, on limestone sub-
strate, in pine-oak forest, oak forest, oak-juniper forest and submontane scrub,
as well as in grassland, from elevations of 1300 to 1600 m (4300-5200 ft). It has
been collected with flowers and fruits from June to September. The local abun-
OCAMPO ACOSTA, A NEW SPECIES OF PORTULACA FROM MEXICO 1359
Ga
WNW,
he:
SZ X
We SRA wr ay Pe
Se) SEE, NY
\A VAN ae A awh AY S cay
y \ 14 Cy ™~ SF, 4 RV yt
S 4 ' , \ j)
mm
2mm
Fic. 1 Dravtist sph ii. A. Habit. B Capsules C Seed showing 4 close-up to a tubercle.
Taste 1. Comparison of Portulaca matthewsii with other species of subsection Conocarpae found in Mexico.
P.matthewsii P mexicana P rzedowskiana
Root Fibrous Fascicled, thickened, Thickened, suberous
not suberous
Stem 4-15 cm long to 10 cm long 4-26 cm long
Shape Subterete, lanceolate, oblong- Terete, linear to sublinear Flat to subterete, lanceolate, some
lanceolate or oblanceolate to times oblong-lanceolate to
lliptic oblanceolat
peauae ellipti anceolate
Size 3-10 mm long 4-10 mm long 4-20 mm lon
0.6-2.5(—3) mm wide 1.5-2 mm wide 1-3(-4.5) mm wide
Apex Acute to subacute Subacute to obtuse Acute
eanste (3-)3.5-4.5(-5) mm long 4.5-5 mm long (3-)4.5-6.5 mm long
ig 2.5-4(-4.5) mm wide 1.5-2 mm wide 4-5.5(-6.5) mm wide
Color Magenta to purple Yellow Pink to purple
Petals Size (4-)5.5-8 mm long 4.5-6 mm long (5.5-)6.5-11 mm long
(—2.5)3-4 mm wide (3-)3.5-6(-7) mm wide
No. of stamens (13-)16-28 14-21 22—44
Si Size (3-)4-5 mm long 1.5-3 mm long 4.5-6(-8) mm long
ee No.of stigma lobes 3(-4) 4-5 a4 o)
Capsule (2.5-)3-4(-4.5) mm long 3-4.5 mm long 3.5-6 mm long
Color Black to bluish-black Black Bluish-black
Seeds Size (0.5-)0.6(-0.7) mm long 0.5 mm long 0.6-0.7(-0.8) mm long
Surface Stellate-tuberculate Diminutively tuberculate Stellate
Vegetation Pine-oak forest, oak forest, oak- Desert scrub, grassland Disturbed subtropical scrub,
juniper forest, submontane rassland
Habitat wae 9
grassland
Altitude
1300-1600 m (4300-5200 ft)
1800-2550 m (5900-8400 ft)
1600-2500 m (5200-8200 ft)
Distribution
Querétaro; possibly also San Luis
Potosi
ntral Mexico, (Colombia?
ae ferina Poelln,)
Michoacan, Estado de México,
orelos
O9EL
(p)0z vaIs/D¥O'LINS
OCAMPO ACOSTA, A NEW SPECIES OF PORTULACA FROM MEXICO 1361
dance of this species is reported from scarce to abundant, so it can be consid-
ered that is not in danger of extinction at present.
The new taxon belongs to subg. Portulaca sect. Catoclasis D. Legrand
subsect. Conocarpae D. Legrand. This subsection is characterized by the size of
the capsules (2.5 to 5(-6) mm long), dehiscing in its lower third portion, by its
campanulate to tubular-campanulate (rarely hemispherical) operculum, as well
as by its linear to oblong-elliptic leaves that are sometimes more or less com-
pressed (D. Legrand 1962). Only two species from this subsection were previ-
ously reported to occur in Mexico: P mexicana P. Wilson (Torreya 28:29. 1928)
and P. rzedowskiana Ocampo (Sida 20:487. 2002). Portulaca matthewsti pri-
marily differs from these by having fibrous roots and by the stellate-tubercu-
late testa of the seed. In addition, P mexicana is distinguished from P matthewsii
by its yellow petals, and terete, linear to sublinear leaves. Portulaca
‘zedowskiana differs from the new entity by its larger dimensions of sepals,
petals and capsules; and higher number of anthers and stigma lobes (Table 1).
Etymology.—The new species is named in honor of James F Matthews,
whose research on Portulaca represents a very important contribution to the
knowledge of the genus in North America.
ACKNOWLEDGMENTS
lam grateful to Victor W. Steinmann, James Henrickson and John McNeill for
helpful comments and corrections to the manuscript; and to Rogelio Cardenas
for the illustration of the new species.
7
REFERENCE
LecRAND, D.1962.Las especies americanas de Portulaca. Anales Mus. Hist. Nat. Montevideo,
2¢ ser, 7:1-147.
1362 BRIT.ORG/SIDA 20(4)
BOOK REVIEW
ELizABETH A. (BERNDT) GeorGe. Watercolour paintings and illustrations by
MARGARET PIFRONI. 2002. Verticordia: The Turner of Hearts. (ISBN 1 876268
46 8, hbk.). University of Australia Press, Crawley, Western Australia 6009,
ww duau. (Orders: International Specialized Book Services,
5824 NE Hassalo St, Portland, OR 97213-3644, U.S.A. 503-280-8832 fax;
www.sbs.com; info@isbs.com). $84.95, 422 pp., 28 color photos, 39 figs. +4
maps, 149 color plates, 6 tables, glossary, 81/2" x 111/2"
This magnum opus isa product of ten years of collecting, mounting, and photographing 1400 speci-
mens of Verticordia (Myrtaceae) by the author and more than 250 volunteer helpers. Furthermore,
the artist, Margaret Pieroni, worked over a period of eighteen years to paint the 147 species, subspe-
cies, and varieties pictured in the book. She wanted to paint each plant in full flower from the actual
specimen and not only had to depend on chance to achieve that, but over the years the number of
ult of this collaboration is a very comprehensive study of a
lor
known species kept increasing. The res
genus unique to Western Australia and the Northern Territories. It is a splendid introduction
anyone who wants to become familiar with this less well known genus and a valuable resource
for
the specialist.
The enormous amount of material about Ve oe has b been well-organized by the author.
rigins to its cultivation today,
Her introduction discusses every aspect of the genus: fro
from chromosome studies to its use in art and craft.
The following section contains tables of species arranged by chronological order of naming, by
earomesonie number, by flowering characteristics, and by habit. There is a good a y,and a key
illustrates each subgenus and sectio
taxa and species A lined arawing
"The main part of the book is devoted to the illustrations and treatments of the species. Eac
the 102 color plates is faced by a page of textand a map show ing where the species may be found. The
m is to convey information, and these
h of
watercolors are in the genre of botanical painting where t
paintings show —— scr ane of oa specimen by the artist. i a me of the reproductive sys-
A It especially when the paint-
This is so Z
tem of the plant i
ing isona colored enn The decision to show the owering twig reched the possibility of
poriey ing the habit of this shrubby plant. The spectacular masses of color can be seen in the photo-
s.
icordia. It is a worthwhile project as the
The author’s aim is to share information about Vert
rium of about
genus may not be well-represented in herbaria outside of Australia. The BRIT Herba
00 specimens has only ten examples of this interesting genus.—Ruth Ginsburg, Botanical Re-
search Institute of Texas, Fort Worth, TX, 76102-4060, U.S.A.
SIDA 20(4): 1362. 2003
A NEW SPECIES OF CHAPTALIA (ASTERACEAE: MUTISEAE)
FROM MEXICO AND REDISCOVERY
OF CHAPTALIA MEXICANA
Leticia Cabrera R.! Guy L.Nesom
Department of Biological Sciences Botanical Research Institute of Texas
University of Texas at Brownsville 509 Pecan Street
brownsville, TX 78520, U.S.A. Fort Worth, TX ie 02-4060, U.S.A.
ABSTRACT
A new species, Chaptalia hidalgoensis L. Cabrera & Nesom, is described and illustrated. It is known
only from the type collection made in Mun. Zimapan, Hidalgo. The occurrence of Chaptalia mexicana
Burkart is documented here, with descriptio 1 illustration, apparently for the first time since the
type was collected by Ehrenberg in 1840. The recent collection was made in Mun. Pinal de Amoles,
oe aro. Both of these Mexican species appear to be members of sect. Chaptalia.
RESUMEN
Una nueva especie, Chaptalia hidalgoensis L. Cabrera & Nesom, se describe e ilustra; ésta se le conoce
anicamente de la coleccion tipo del Mun. Zimapan, Hidalgo. También documentamos la existencia
de Chaptalia mexicana Burkart, proporcionando una descripcion completa e ilustracion;
eearauiied esta especie no se habia vuelto a colectar desde la colecta tipo de Ehrenberg en 1840
Esta coleccion reciente es del Mun. Pinal de Amoles, Querétaro. Ambas especies mexicanas parecen
ser miembros de la seccion Chaptalia.
Research by the first author toward a treatment of Mutisieae of the Bajio region
of México has brought to light two interesting discoveries in Chaptalia Vent. A
irst known collection of C. mexicana Burkart
oH
new species is described, and the f
since its initial discovery in 1840 is documented.
Chaptalia hidalgoensis L. Cabrera @ Nesom, sp. nov. (Fig. 1). Type: MEXICO.
HIDALGO: Mun. Zimapan. Verdosas, bosque de pino, encino y Juniperus, 1750 m, 20
Nov L991, VM. Huerta 1352 (HOLOTYPE: IEB!).
tu
is flosculis centralibus fungenter staminatis—
ree lyratifoliae et alio sect. Chaptaliae simi
fert ab omnibus coniunctione rhizomatum, scaporum bracteatorum, foliorum coriaceorum
ee um, owes um pistillatorum interiorum numerosorum, et acheniorum non rostratorum;
differt a C. lyratifoliae scapis bracteatis, phyllariis marginibus glandulosis, flosculis pistillatis
exterior ee seen ac flosculis aisilianis interioribus multo paucioribus, et flosculis
bisexualibus corollis brevioribus.
Plants perennial, fibrous-rooted, rhizomatous. Leaves coriaceous, 3-5 cm long,
blades L.5-3.6 cm long, 1.1-2.1 cm wide, elliptic-oblong to ovate, lyrate, with an
ovate to ovate-cordate terminal portion and I(-2) pairs of basal lobes, margins
‘Current address: Leticia Cabrera R., Biological Sciences Department, Northwest Missouri State University, 800
Ur niversity Drive, Maryville, MO 64468, U.S.A.
SIDA 20(4): 1363-1369. 2003
1364 BRIT.ORG/SIDA 20(4)
nif
NA) fis
whi /
es Z
Fic. 1. Habit of Chaptalia hidalgoensis, from the type.
crenate to denticulate, revolute, apex acute to obtuse and mucronate, the base
cordate, lower surface white-gray tomentose, upper surface villous-arachnoid,
glabrescent. Scapes 1-3 per plant, 8-12 cm tall, villous to almost glabrous,
tomentulose close to the head, 6-9-bracteate, bracts linear-lanceolate, 3-5.2 mm
long, also with 3 bract-like phyllaries close to or subtending the involucre (ie.,
bracts transitional to phyllaries). Heads solitary, 1.1-1.7 cm tall, 14-18 cm wide;
involucres turbinate, 0.8-0.9 cm tall, phyllaries 3-seriate, green, herbaceous, all
acute, scarious at the margins, with the distal area usually dark-red, the dorsal
surface somewhat arachnoid, soon glabrous, the outermost linear-lanceolate,
ca. 2.8mm long, 0.5-0.7 mm wide, margins sparsely glandular, the central lan-
ceolate, +-5.2 mm long, 0.8-1 mm wide, margins sparsely glandular, the inner-
most ca.8 mm long, ca. 1mm wide, margins glabrous. Florets trimorphic—outer
CABRERA R. AND NESOM, A NEW SPECIES OF CHAPTALIA FROM MEXICO 1365
pistillate and ligulate, middle pistillate with reduced ligules, inner bisexual
and tubular-bilabiate: outer florets 15-21 in | series, ligules white-cream in color,
sometimes somewhat purple with age, with an abaxial purplish midstripe at
maturity, 10 mm long, ca. | mm wide, the tube 2.8 mm long, the inner lips 1-3,
filamentous, l-1.2 mm long, the style 5.2 mm long, including the branches 0.2-
0.3 mm long; middle florets ca. 7, ligules ca. 6 mm long, ca. 0.4 mm wide, the
tube almost as long as the ligule, inner lip absent; style 6 mm long including
the ca. 0.5 mm long branches; inner florets ca. 22, corollas bilabiate, creamy in
color, 7mm long, including the 5mm long tube and the recurved lobes 2-3 mm
long; stamens 4.5 mm long, appendage 1.5 mm long, tails ca. 1.2 mm long, t
ecae ca. L.8 mm long. Achenes: (outer pistillate florets) immature but apparently
fertile, ca. 2 mm long, papillose over whole surface, cylindric; pappus biseriate,
ca. 5mm long; (inner pistillate florets) fertile, ca. 2.5mm long, fusiform, appar-
ently not beaked, papillate; pappus uniseriate, 5-5.5 mm long; (bisexual flo-
rets) sterile, ca. 2 mm long, cylindric, with few hirsute hairs; pappus biseriate,
6.2 mm long. Known only from the type collection, where it was reported as
scarce.
Chaptalia hidalgoensis (as wellas C. mexicana, below) appears to bea mem-
ber of sect. Chaptalia, as outlined by Nesom (1995)—recognized by the produc-
tion of functionally staminate (pseudo-bisexual) central florets, ligules of the
outer pistillate florets witha purplish, abaxial midstripe at maturity, and heads
nodding in bud. Previously, only one of the six species of sect. Chaptalia was
known to produce consistently bracteate scapes—both of the species discussed
inthe present manuscript have bracteate scapes. The behaviour of the heads in
the two species treated here in detail is inferred from their putative relatives.
Within sect. Chaptalia, C. hidalgoensis mostly closely resembles C.
lyratifolia Burkart in its coriaceous and lyrate leaves. One of the plants of the
Huerta collection shows production of short rhizomes (the other plant collected
without any below-ground parts) and because of the similarities of Chaptalia
hidalgoensis to C. lyratifolia, which characteristically is colonial through pro-
duction of creeping or stoloniform rhizomes, it seems probable that C.
hidalgoensis also has a colonial habit. A summary of distinctive contrasts of
the new species with C. lyratifolia is given in the key below (following com-
ments on C. mexicana). The latter species is distributed from central Nuevo Leon
and adjacent Coahuila and Tamaulipas into central San Luis Potosi.
—
a
~~
Rediscovery of Chaptalia mexicana
Chaptalia mexicana Burkart was described and illustrated (Burkart 1944) from
a single collection, cited below. A more recent collection from Querétaro is the
second known and documents its continued existence, although the species,
which was inadvertently omitted in Nesom’s review of the North American
and Central American species (1995), apparently is rare elsewhere.
1366 BRIT.ORG/SIDA 20(4)
Chaptalia mexicana Burkart, Darwiniana 6:543. 1944 (Fig. 2). Typr: MEXICO. [No
other locality data], 1840, Ehrenberg 1042 (HOLOTYPE: B, fide Burkart 1944).
Recent collection: smite Querétaro: Mun. Pinal de Amoles, mas o menos 2 km al N de Puerto de
Tejamanil, b _ ladera de cerro, canada, 2400 m, 27 Mar 1989, FE. Carranza 157] (EB).
The sheet au three eae
Plants perennial, fibrous-rooted, producing rhizomes at least to 1.4 cm long.
Leaves 3.5-9.2 cm long, dissected into 5-7 pairs of somewhat retrorse lobules,
these increasing in size distally, the terminal lobe 1.6-3 cm long, 1.1-2.1 cm wide,
ovate to ovate lanceolate, with the margins retrorse-crenate, the lower surface
gray-white tomentose, the upper surface dark-green with red-tinges, lightly
villous to glabrous with age, the midvein very prominent, villous-tomentose,
ark red, becoming white-tomentose distally; petioles 0.8-1.5cm long, winged,
white-villous. Scapes |-2 per plant, 4-10.5cm tall, white-tomentose, (2-)4-6[8-
10}-bracteate, bracts linear-lanceolate to trullate, 4-5.5 mm long, 0.5-0.8 mm
wide. Heads solitary, 1.4-2 cm tall, 1.1-1.8 cm wide; involucres 0.9-1.3 cm tall,
turbinate-campanulate, phyllaries 3-seriate, acuminate, the outermost red-dark,
somewhat herbaceous, linear-lanceolate, 4-5.5 mm long, 0.5-0.8 mm wide, scari-
ous at the margins, arachnoid on the dorsal surface, the central red-dark to green,
somewhat scarious, lanceolate, 10-11 mm long, ca. | mm wide, scarious on the
margins, somewhat villous on the dorsal surface, the innermost more or less
scarious to somewhat greenish, lanceolate, 12-13 mm long, ca. 1 mm wide, gla-
brous dorsally. Florets trimorphic—outer pistillate and ligulate, middle pistil-
late with reduced ligules, inner bisexual and tubular-bilabiate: outer florets
15-17 in 1 series, ligules white-cream, with an abaxial purple midstripe when
mature, [7]12-14 mm long, 1.1-1.8 mm wide, the tube 3-3.5 mm long tube, the
inner 2 lips ca. [1]L.5-2.1 mm long, filamentous; style ca. 5.8-8 mm long, includ-
ing the [0.6]0.2 mm long branches; middle florets 7 mm long, 0.7 mm wide, the
tube 2.5-3 mm long, inner lip ca.0.5 mm long; style 8 mm long including the ca.
0.2 mm long branches; pappus 5 mm long; inner florets 17-20, 8.5 mm long, the
tube ca. 4.5 mm long, the semi-erect to recurved lobes ca. 4 mm long; stamens
ca.6 mm long, the filaments 2 mm long, the tails ca. 1.2 mm long, the append-
age ca. 1.5 mm long, thecae ca. 2.5 mm long, fertile. Achenes: (outer pistillate
florets) fertile, ca. 3-4 mm long, fusiform, papillose; pappus biseriate, 4-7 mm
long; (inner pistillate florets) fertile, 3.5 mm long, fusiform, apparently not
beaked, sparsely papillose, pappus 5 mm long; (bisexual florets) sterile, ca. 3
mm long, narrowly cylindric, glabrous; pappus uniseriate, 8 mm long. Square
brackets [] enclose data from Burkart description.
The scape of the Ehrenberg type of Chaptalia mexicana (as illustrated by
Burkart) appears to have 8-10 bracts, vs. (2-)4-6 in the recent Carranza collec-
tion, and several other measurements in Burkart’s description vary from the
recent collection, but the combination of bracteate scapes and the highly dis-
CABRERA R. AND NESOM, A NEW SPECIES OF CHAPTALIA FROM MEXICO 1367
Fic. 2. Habit of Chaptalia mexicana, from Carranza 1571.
—
sected leaves is so remarkable that it is difficult to conclude that these are not
the same species.
To arrive at an identification of Chaptalia mexicana using Burkart’s key
(1944, p. 524), which places the species in sect. Lieberkuhna, one must follow
the choice “Flores centrales hermatroditas” (vs. “Flores centrales del capitulo
masculinas, con ovario atrofiado.”). His protologue for the species described
“flores centrales hermatfroditas” (p. 544), but he explicitly noted further down
in the description that he did not see the achenes (probably meaning mature
achenes, since his illustration included essentially similar ovaries at the base
of each of the three kinds of corollas). It seems clear that Burkart would have
placed this species in his sect. Euchaptalia if he had observed central florets
—-
1368 BRIT.ORG/SIDA 20(4)
with sterile achenes. Our observation is that ovaries of the central (pseudo-bi-
sexual) florets of Carranza 1571 are glabrous and remain narrowly cylindric,
while those of the pistillate florets are papillose and swell with the production
of embryos.
Ehrenberg’s type collection apparently did not include rhizomes, and the
rhizomes on the Carranza collection are short, but because of the similarities
of C. mexicana to C. lyratifolia and C. estribensis (leaves coriaceous and lyrate to
cordate; style branches U-shaped, see comments below), it seems likely that the
former also is colonial through production of creeping or stoloniform rhizomes
KEY TO THE SPECIES OF SECT. CHAPTALIA
ey basally attenuate, not lyratiform; plants without rhizomes, occurring indi-
ually.
, peas t istillate florets in 1 series, prominently ligulate; fertile achenes
with a ma Sigh constricted neck.
talia pringlei Greene
2. Scapes ebracteate; pistillate florets in 2 series, those of the outer series ai
those of the inne
r series with ligules absent or greatly reduced; fertile achenes
oe a neaty mot oe
enatijlat
spatulate,
with a narrowly attenuate petiolar ca
tile achenes 6-9 mm long, including the slender neck 1/2-1/2 as long as
arsely papillate over the whole surface. Chaptalia ee Nesom
3. ee ee to elliptic-obovate, without a petiolar region; fertile achenes
3.5-5.2 mm long, including the slender neck 1/4-1/5 as long as the achene,
glabrous except for the sparsely papillate neck region.
Chaptalia tomentosa Vent.
Leaves aaa cordate and/or distinctly lyratiform; plants rhizomatous an
colo
4. Leaf a thin-herbaceous, sometimes reddish. Chaptalia hintonii Bullock
4. Leaf blades thick-coriaceous, not r
»:
Leaf blades cordate, without es Seasons with orange- ay vestitu
on the abaxial surface.
ptalia ep ak Nesom
5. Leaf blades cordate to truncate at base, distinctly lyrate, with oe vestiture
on the abaxial surface
6, Leaves dissected into 5-7 pairs of somewhat retrorse, roughly equal-size
lobules; scapes bract
haptalia mexicana Burkart
6. Leaves lyratiform, ay with one terminal primary lobe and 0-2(-4)
smaller basal lobules; scapes bracteate or ebracteate
7. Scapes bracteate; ee Bac sparsely alandular outer pistillate
florets 15-21, inner lip 1 m long; middle florets (pistillate) ca. 7,
fewer than outer; corollas a ead florets 7 mm lon
Chaptalia
hidalaoensis Cabrera & Nesom
7. . ebracteate; phyllary margins eglandular; outer pistillate floret
24-38, inner lip absent or less than 0.5 mm long; mee florets (ist
late) pa more than 90, usually 3 more numerou rollas
of bisexual florets 9-12 mm long. pereien lyratifolia Burkart
Generic placement
Katinas (1998) transferred Chaptalia hintonii to the genus Gerbera Gmel.,
which otherwise (except for one South American species) has been recognized
CABRERA R. AND NESOM, A NEW SPECIES OF CHAPTALIA FROM MEXICO 1369
only from Africa and Asia. Her rationale for the transfer emphasized the pro-
duction of staminodes on the outer pistillate florets of C. hintonii, a feature
characteristic of Gerbera but usually not of Chaptalia, and the “U-shaped” style
branches of C. hintonii. We acknowledge that generic limits among taxa of the
Gerbera complex are somewhat ambiguously drawn and may yet require taxo-
nomic adjustments, but it does seem clear that C. hintonii is a member of the
group of five species within sect. Chaptalia (as recognized here) characterized
by production of stolons and cordate to lyratiform leaves (C. estribensis, C.
hidalgoensis, C. hintonii, C. lyratifolia,and C. mexicana). All species of this group
also produce the U-shaped style branches. Style branches are more slender and
“V-shaped” in the three species of sect. Chaptalia with basally attenuate leaves
and without stolons (C. madrensis, C. pringlei, and the type of the genus, C.
tomentosa)—the style branch morphology is more like that of species of sect.
Leria (DC.) Burkart, which includes C. nutans (L.) Polak., C. texana Greene, and
others. The similarities of C. tomentosa and its two closest relatives to the five
others here placed in sect. Chaptalia, and the apparent geographic integrity of
the whole group, are taken here as evidence of close evolutionary relationship
(Nesom 1995). A more detailed discussion of this problem is presented ina pair
of papers by Katinas and Nesom (submitted).
ACKNOWLEDGMENTS
Loans of specimens to the first author from F GH, IEB, MEXU, MICH, and USCH
are greatly appreciated. We are grateful for helpful reviews from the editor and
Beryl B. Simpson.
REFERENCES
Burxart, A. 1944. Estudio del género de Compuestas Chaptalia con especial referencia a
las especies Argentinas. Darwiniana 6:505—594,
Nesom, G.L. 1995. Revision of the North and Central American species of Chaptalia
(Mutisieae: Compositae). Phytologia 78:153-188.
Katinas, L. 1998. The Mexican Chaptalia hintonii is a Gerbera (Asteraceae, Mutisieae). Novon
8:380-385.
1370 BRIT.ORG/SIDA 20(4)
BOOK REVIEW
Jeremy D. GARRETT. CORAL MCCALLISTER CLlustrator). 2003. Landscaping for Wild-
life: A Guide to the Southern Great Plains. (ISBN: 0806134895, pbk.). Uni-
versity of Oklahoma Press, 4100 28th Ave., Norman, OK 73069-8218, U.S.A.
(Orders: www.oupress.com, 800-627-7377, 800-735-0476 [orders only], 405-
364-5798 fax). $28.00, 209 pp., b/w and color illustrations, 81/2" x 11".
From Landscaping for Wildlife —People want to attract wildlife to their yards for a variety of rea-
sons, but probably foremost among these is that birds and other animals bring us pleasure. The exu-
berant songs of nese — snes cites the playful antics of young squirrels at feeders, and the
sight of colorful | g al wildflower patches provide us with both daily and last-
ing joy.”
Through careful landscaping, you can make your property a year-round waystation for a wide
variety of animal species, from birds, butterflies, and squirrels to bats, turtles, and opossums. Garrett
explains how specific plants draw animals to your property by providing food and shelter sources
To assist readers in designing their own “wildscape,” detailed diagrams and plant listings ac-
company photographs of several Oklahoma eee en of Wildlife Conservation-certified
wildscaped properties. The detailed diagrams will help you inven
LUL and VV ildscape YUU OWT beV’P
erty. A woodworking chapter is complete with patterns of nesting and feeding stations designed for
individual wildlife species. Each section includes tables that list the common name and the scien-
tific name of the plant or wildlife species. The tables discuss how the plants are used or what animal
uses them and the relationship between flora and fauna is definec
The detailed diagrams are especially useful in visualizing what a particular wildscaped plot
will look like. Each includes a color photograph, a bird's-eye or plan view and an interpretive sketch
on which the plants are identified. The bird’s-eye view could be used as a guide for planting. The
eleven case studies cover six kinds of settings: sunny, uel one moist waldeea pes. ileal
meadows, and butterfly and rock gardens. All of the multi-l
sion of ¢
“ ter sources rate their own chapter and cover several types. Three types of ponds, a re-circu-
lating stream, and a side yard marsh are illustrated though the use of case studies. Common aquatic
and wetland plants dis ssed, as well as wetland-related wildlife species
ap i orovide a ee checklist and a monthly accnnescslenaey Sources for seeds
ants are listed, and there is a bibliography of field guides and references. Internet sites and
related eaneaien and agencies 2 are ene! ed.
Landscaping for Wildlife | I d like with specific, detailed; yet easy-
to-follow instructions for meeting water vigetls feeding preieeaces and nesting sof the
wildlife found in the southern Great Plains. Although written specifically for the southern Great
Plains the concept is one that could be used anywhere with appropriate substitutions for locality.
Written by Jeremy D. Garrett in association with the Oklahoma Department of Wildlife Conserva-
tion and ae singe by Coral McCallister’s elegant and charming illustrations of wildlife as well
=
and p
as of s habitats and wildscapes.
The Oklahoma aDe par tment of Wildlife Conservation manages Ok lahoma'’s wildlife resources
and habitat to | ific, educational, aesthetic, economic { | benefits for present
and future generations of hunters, aon: and others who appreciate wildlife. Jeremy D. Garrett is
president of NaTour Communications, a natu
sulting firm. Coral McCallister is a biological
illustrator for the Department of teas at the. University of Oklahoma.—Gary Jennings, Botanical
Research Institute of Texas, Fort Worth, 1X, 76102-4060, U.S.A
SIDA 20(4): 1370. 2003
TAXONOMY OF THE POLEMONIACEAE:
THE SUBFAMILIES AND TRIBES
Verne Grant
Section of Integrative se and
Plant Resources Cent
Tee of Texas ee
Austin, TX 78712, U.S.A.
ABSTRACT
This paper ised f | classificati f the eee es of the Polemoniaceae
papel
based onasyninests of new and old charact d pI I The geners
il Itl bfamilies (Cobaeoideae id A I
The! five topical and subtropical genera, which are basal in the family, fall into two eau y groups:
subfam. Cobaeoideae, containing Cantua, Cobaea, Bonplandia, and Loeselia,; and subfam.
Acanthogilioideae, containing Acanthogilia. The eomnperate herbaceous genera, which are derived,
also fall into two primary groups. The subfam. Pol d d derived from
the Cobaeoideae, contains Polemonium, eolinane. Navarretia, Phlox, Linanthus, etc. The second main
be Gilieae, which is related to the tribe Acanthogilieae
group of
and incl bed in the same sublamily Acanthogiliideae with it. The Gilieae contains the genera Gilia,
is system differs in numerous respects from the recent system of Porter
Eriastrum, Ipomopsis, etc. This sy
and Johnson (Aliso, vol. 19(1), 2000), based on a molecular cladistic approach. Different goals and
methods produce different results. It is suggested that other plant groups, which have been revised in
recent times acca to molecular elagistic ev vadence ane criteria, should also be revised by taxo-
1 d for the useful features that they alone
nomic methods. U
provide.
RESUMEN
En este trabajo se presenta una clasificacion formal revisada de las subfamilias y tribus de las
] ¢ ee et 1: P ] ree
y vie)
plasice! Le os Sp Heres se pempan en nueve uanus y tres subfamilias (Cobaeoideae, Polemonioideae,
I g y subtropicales, que son basales en la familia, quedan
bf ssbaenideae con Canta, Conga, ik nein saci Bien
d mbién
Aemiopitodexé con ae Los genero
estan en dos grupos primarios. La subfam. Pole
Cobaeoideae, contiene a Polemonium, Collomia, Navarretia, Phlox, Linanthus, etc. El segundo grupo
Gilieae, que se relaciona con la tribu
iesisnadd y eidentenents ae de
principal de géneros herbaceos templados es la tribu
Acanthogilieae y esta incluida también en la misma subfamilia nes Gilieae contiene
los géneros Gilia, Eriastrum, Ipomopsis, etc. Este sistema difiere en numerosos aspectos del sistema
reciente de Porter y Johnson (Aliso, vol. 19(1), 2000), basado en un método cladistico molecular.
mre s A ieee _ Pd aT ] ] | ee [4 . | ]
J 6 1 fa am b
id : ] ] Pen ] ] ea <
lasificaciones taxonémicas actualizadas se necesitan por
}
q
también por métodos taxonémicos. Las
] Feri 4] 1] ]
L
Y
SIDA 20(4): 1371-1385. 2003
1372 BRIT.ORG/SIDA 20(4)
INTRODUCTION
Research on systematics of the Polemoniaceae is going on in several laboratories
at present and is producing new evidence concerning relationships. This paper
presents a revised classification of the primary subdivisions of the family—the
subfamilies and tribes—in the light of the currently available information.
An earlier system (Grant 1959) was constructed before there was much
pollen-morphological evidence or any DNA evidence. Both of these lines of
evidence are very important for classification (see Materials and Methods for
references). The two types of evidence were used ina recent system (Grant 1998;
see Table 1).
I did not appreciate the full potential of the pollen evidence in 1998. It was
used there to distinguish groups at low and middle taxonomic levels. In the re-
vised system presented here it is used as a key character separating subfamilies.
An important molecular survey of the family was carried out by Johnson
etal. (1996), using the chloroplast gene matK and a cladistic approach. This study
recognized informal groupings based on the DNA cladograms, many of which
differed from those in the Grant (1959) system. I took up some of these new
groupings but rejected others in a revised taxonomic classification (Grant 1998,
1999; Grant & Day 1999). The taxonomic system of 1998-1999 differs signifi-
cantly from the molecular cladistic groupings of Johnson et al. (1996).
Cladistic treatments of other organellar genes were made by various work-
ers in this same period. The studies most relevant to this paper are listed in
Materials and Methods. A complete list of molecular studies in the family up to
2000 is given by Porter and Johnson (2000).
Porter and Johnson (2000) then used the available molecular evidence as
the main basis for a formal system of the family. The primary clades or deepest
branchings in the DNA cladograms become subfamilies in their system, the
secondary clades become tribes, and the third-order clades become genera or
small sets of genera. Formal taxonomic names are assigned to the groups. Good
descriptions of phenetic characters are given for the groups. However, it is diffi-
cult to tell what role these phenetic characters play in defining the groups; sets
of diagnostic characters are lacking. The groups are basically clades. Porter and
Johnson (2000, p. 55) describe their system as “a phylogenetic classification of
the Polemoniaceae.” It is what Mayr and Bock (2002) call a cladification.
Again, major differences exist between the cladistic system (Porter &
Johnson 2000) and the taxonomic system (Grant 1998, 1999; Grant & Day 1999).
A detailed analysis and discussion of these differences is given in Grant (2001).
The incongruences persist in a comparison of the molecular cladistic system
with the revised taxonomic system presented in this paper. Some of the differ-
ences are shown in Table |
GRANT, TAXONOMY OF THE POLEMONIACEAE
Tasle 1. Comparison of recent classification systems of the family
1373
Grant 1959
Grant 1998, 1999; Grant and Day 1999
Tropical and subtropical taxa
Tribe 1. Cantueae.
Cantua, Huthia
Tribe 2. Cobaeeae.
baea
O
Tribe 3. Bonplandieae.
Bonplan Loeselia.
Temperate ta
Tribe 4. eee eae.
Polemonium, ie Allophyllum
Gymnosteris, Phlox, Microsteris.
Tribe 5. Gilieae
Gilia /pomopsls Eriastrum, Langloisia
Navarretia, Leptodactylon, | thus
subfam. 1. Cobaeoideae.
t
Tribe 2. Cobaeeae.
Cobaea
Tribe 3. Bonplandieae.
landia.
Bonpla
Biss 4. Loeselieae.
selia
os Acanthogiieae
Acanthog
Subfam. 2. Jaane
Tribe 6. Polemonieae.
Polemonium, an Allophyllum,
Navarretia, Phlox, Microsteris, Gymnosteris
Tribe 7. Gilieae.
Gilia, lpomopsis, Eriastrum, Langloisia
Tintinabulu
Tri
8. | on eee
Leptodactylon, Linanthus, Maculigilia
Porter and Johnson 2000
System proposed here (2003)
Subfam. 1. Acanthogilioideae.
Tribe 1. eae
Acanthogi
Subfam.2,Cobacoidese
Tribe 2. Can
Cantua el ae
Tribe 3. Cobaeeae.
Cobaea.
ice 4. Popoentiene
Bonplandia.
Subfam. 3. Polemonioideae.
Tribe 5. Polemonieae.
Polem
Tribe 6. ae sie
Loeselia pareseata Dayia, Bryantiel
la
Aliciella, lpomopsis, Microgilia, Eriastrum
Langloisia, Loeseliast
Tribe 7. Gilieae.
Collomia, pu, Navarretia, Gilia,
Saltugilia, Lathro
Tribe 8. Phloc
Phlox ee ee Linanthu
Leptosiphon.
rum
Subfam. 1. Cobaeoideae.
Tribe 1. Cantueae.
Cantua, (incl. Huthia).
Tribe 2. Cobaeeae.
Cobae
Tribe 3 Ponadee
Bonplandia
Tribe 4. Die
Loeselia.
Subfam. 2. Polemonioideae.
Tribe 5. Polem
Polemonium, elon Via, Allop
Navarretia.
Tribe 6. Phlocidea
Phlox, Microsteris, armor
Tribe 7. Leptodac
=~
ayllum
ede.
Leptodactylon [. Macul
igilia
Gilia, lpomopsis, Eriastrum, Langloisia
Tintinabulum.
1374 BRIT.ORG/SIDA 20(4)
Cladists, on finding differences between cladistic and taxonomic systems
of the same group, commonly attribute the differences to inadequacies in the
taxonomy. Johnson et al. (1996) expressed this view of the Grant (1959) system
very clearly. Indeed, the 1959 system was old and in need of repair in 1996. How-
ever, there are also other explanations to consider.
The basic consideration is that one system is a product of taxonomy and
the other a product of cladistics. The two approaches have different criteria,
goals, and methods. Taxonomy groups organisms according to similarity and
difference, cladistics groups them into ancestor-descendant lineages. Taxa are
not the same as clades. Taxonomy uses the traditional definition of monophyly
whereas cladistics uses a special definition of its own. In taxonomy a mono-
phyletic group is any group of organisms descended from a recent common
ancestor; in cladistics a group to qualify as monophyletic must contain all the
descendants of the common ancestor (see reviews of Mayr & Bock 2002, and
Grant 2003).
The differences in the definition of a systematic unit and of monophyly
automatically lead to incongruence between taxonomic and cladistic systems
of the same group. A group circumscribed in a given way can be monophyletic
in taxonomy but non-monophyletic in cladistics. And cladists frequently and
incorrectly declare a taxonomic group to be non-monophyletic when it is mono-
phyletic by the taxonomic definition. Many of the disagreements between the
taxonomic and cladistic systems of the Polemoniaceae (and other plant groups)
can be attributed to the differences in working concepts and definitions (see
Grant 2001, 2003).
Where incongruences between rival systems are due to the use of different
working concepts, one system cannot be said to be right and the other wrong;
both systems may be right by their own respective standards. The view held by
many cladists, including Johnson et al. (1996) in the Polemoniaceae, that a new
cladistic treatment should replace a preexisting taxonomic treatment, is not
justified insofar as it is based on the working concepts. In fact, one could argue
just the opposite. We should have both cladistic systems and up-to-date taxo-
nomic classifications, so that consumers of systematic biology can have achoice.
Other incongruences between taxonomic and cladistic systems of the
Polemoniaceae are due to the evidence used. Here we are comparing taxonomy
with molecular cladistics, phenetic characters with DNA sequence variation,
and broad databases with narrow ones. This aspect will be considered in the
Discussion section.
An up-to-date formal taxonomic classification of the Polemoniaceae is
needed to complement the valuable cladistic treatment of Porter and Johnson
(2000). The present paper deals with the subfamilies and tribes. An updated
taxonomic treatment of the genera and sections of the temperate herbaceous
Polemoniaceae is also needed. Work has begun on selected temperate genera.
GRANT, TAXONOMY OF THE POLEMONIACEAE 1375
MATERIALS AND METHODS
I compiled a list of conventional taxonomic characters from the older litera-
ture. I then sorted out the characters a aistinguish groups at higher levels in
the family, and set aside those that are di ically useful for genera and spe-
cies. Character expressions were ees in the University of Texas Herbarium
where desirable.
Family-wide surveys of particular microscopic and chemical characters
are very helpful. Such surveys are available for: chromosomes (Grant 1959, ch.
6); pollen (Stuchlik 1967a, 1967b; Taylor & Levin 1975); wood anatomy (Carlquist
et al. 1984); corolla venation (Day & Moran 1986); flavonoids (Smith et al. 1977,
1982); and organellar DNA (several authors listed below). The older studies of
chromosomes size (in Grant 1959) need to be repeated by modern methods of
determining DNA quantity.
Some of the characters vary at the upper taxonomic levels, and are used in
this paper, while others vary at the generic and infrageneric levels. This is illus-
trated by the flavonoids.
The flavonoids occurring in the family fall into three main groups: (A) the
common types, kaemplferol, quercetin, and myricetin; (B) 6-methoxyflavonols;
and (C) C-glycosylflavones (Smith et al. 1977). Variation in these is more useful
taxonomically at the genus level than at the tribal level, and only a part of the
flavonoid evidence is included in this paper. Smith et als (1977) findings are
recorded below for the four tropical genera. One aspect, the presence or absence
of types B and C is recorded below for the temperate tribes.
Numerous features of the pollen show variation within the family. Day and
Moran (1986) found that the spatial distribution of the pores is especially use-
ful diagnostically. Most members of the family fall into one or the other of two
contrasting character states: pantoporate (pores scattered over the grain) or
zonocolporate (pores equatorial).
The corolla tube has five sets of three parallel and separate veins, one set
for each corolla lobe. In most members of the family the veins anastomose to
form a network in the corolla throat or lobes. In the tribe Leptodactyloneae,
however, the veins are non-anastomosing. The character may not be well ex-
hibited in small-flowered species of other tribes (Day & Moran 1986).
Family-wide studies of DNA sequences have been carried out with chloroplasts
(matk, ndhF), mitochondria (nad1B), and ribosomes (ITS) (Steele & Vilgalys
1994; Johnson et al. 1996; Porter 1997; Porter & Johnson 1998; Prather et al. 2000).
The DNA relationships are clearly portrayed in the cladograms, and I have used
these in constructing the taxonomic classification. However, | have not founda
way to express the molecular evidence in verbal terms for inclusion in the de-
scriptions. The Porter and Johnson (2000) system is a good representation of
the molecular clades (see Table 1).
1376 BRIT.ORG/SIDA 20(4)
The molecular evidence is of ten but not always in agreement with the phe-
netic evidence. In cases of a conflict | reexamine the phenetic evidence with
the thought that it is best to have some phenetic characters to support the mo-
lecular characters. In the recent past (Grant 1998, 1999) and again for the present
study I have found some such correlated phenetic and molecular characters,
and have made the appropriate taxonomic changes. If, however, molecular evi-
dence alone points in one direction, while two or more reliable phenetic char-
acters point toa different taxonomic conclusion, | follow the phenetics. In other
words, | do not assume that the molecular evidence is always the right guide to
follow.
The descriptions in the formal system consist mainly of the character states
of diagnostic characters. These can be regarded as the evidence supporting the
taxonomic groupings. Good complete descriptions are given by Porter and
Johnson (2000).
The nomenclatural paragraphs of the subfamilies and tribes contain the
essential original names and recent names. Additional synonyms may be found
in Grant (1959).
SYSTEM OF CLASSIFICATION
Fam. Polemoniaceae Juss. Polemoniaceae (as Polemonia) Juss.; Gen. PI. 136.1789. TyPE:
Polemonium caeruleum L.
Various life forms from small trees to annual herbs. Leaves mostly alternate,
but opposite in Phlox, Leptodactylon, and Linanthus. Floral plan 5-5-5-3. Co-
rolla sympetalous with epipetalous stamens, usually radial, sometimes bilat-
eral. Corolla veins mostly anastomosing in the corolla throat or lobes, but non-
anastomosing in Leptodactylon and Linanthus. Ovary superior and 3-carpellary,
Fruit generally a 3-celled capsule, but 1- or 2-celled in some species of
Navarretia. Original basic number x = 9; polyploids and aneuploids common.
Mainly American hemisphere; several species in Eurasia. About 347 spe-
cies. These are grouped here into three subfamilies and nine tribes. The species
are listed for the tropical and subtropical tribes and genera. This is not feasible
for the species-rich temperate groups; for these see the species lists in Grant
(1959) and Porter and Johnson (2000).
ANALYTICAL KEY TO TRIBES
1. Seeds mostly winged (sometimes wingless); chro e Il; tropical and sub
tropical.
2. Spiny desert shrub; deciduous green leaves in axils of spines; pollen zono-
colporate; endemic in Baja California Acanthogilieae
2. Not spiny desert shrubs with leaves as above; pollen pantoporate; widespread
in American tropics and subtropics.
ae eee
py bracts: life form
| | Ty] ; { J
3. Calyx WIOHY 2 not subtended
varies.
GRANT, TAXONOMY OF THE POLEMONIACEAE 1377
4. Climbing vines; calyx sepals divided to base; leaves with terminal tendrils
Cobaeeae
4. Not vines; calyx synsepalous; leaves without tendrils.
5. Small trees and shrubs; Andes Cantuea
5. Perennial herbs; Mexico and cuatemals Bonplandieae
3. Calyx membranous with no or only II gi individual fl
ers subtended by bracts; nee and herbs Loeselieae
. Seeds without wings; chromosomes mostly medium-sized to large, but small in 1
tribe; temperate and boreal.
6. Pollen zonocolporate; seeds sandy or pale-colored; mostly in xeric habitats, com-
mon in deserts Gilieae
6. Pollen pantoporate; seed color varies; in various habitats.
7. Seeds sandy or pale-colored; leaves opposite; leaves if cleft or divided are
palmately so; chromosomes small eptodactyloneae
7. Seeds dark brown or black; leaves alternate or opposite; leaves if divided are
pinnately so; chromosomes large or medium-sized
8, Leaves alternate; leaves pinnately compound or lobed Polemonieae
8, Leaves opposite, or absent in 1 small genus; leaves simple and entire ____- Phlocideae
Subfam. 1. Cobaeoideae (D. Don) Arn., Cobaeaceae D. Don, Edinburgh Philos. J. 10:109-
111. 1824. Subfam. Cobaeoidea Arn., Encycl. Brittanica, ed. 7, 5:121. 1832. Subfam.
Cobaeoideae Brand, Pflzr. 4(250):19. 1907. Type: Cobaea scandens Cav.
Various life forms from small trees to herbs, in tropical and subtropical zones.
Seeds mostly winged, sometimes wingless. Pollen pantoporate. Chromosomes
small
Tribe 1. Cantueae Peter, Nat. Pflazfam. 4(3a):45. 1891. Type: Cantua buxifolia Juss.
Small trees and shrubs. Leaves simple with broad blade or pinnately divided
and narrow. Calyx wholly herbaceous and synsepalous. Corolla radial to bilat-
eral. Seeds flat with broad wings. All three groups of flavonoids (A, B, and C)
present (see Materials and Methods for explanation of these groups). 2n = 54.
Distribution and taxa.—Andes. One genus, Cantua, with twelve species: C.
bicolor, C. buxifolia, C. candelilla, C. coerulea, C. cuzcoensis, C. flexuosa, C.
longiflora, C. longifolia, C. ovata, C. pyrifolia, C. quercifolia, C. tomentosa. The
former small genus Huthia has been submerged in Cantua by Porter and
Johnson (2000) and Alan Prather (pers. comm.) who is currently studying the
group. I am following these authors.
Tribe 2. Cobaeeae (D. Don) Meisn., Cobaeaceae D. Don, Edinburgh Philos. J. 10:109-
111. 1824. Tribe Cobaeeae Meisn.,, Pl. Vasc. Gen. 180, 273. 1839-1840. TYPE: Cobaea
scandens Cav.
Climbing vines. Leaves pinnately compound with a terminal tendril. Flowers
large and solitary. Calyx wholly herbaceous, sepals divided to base. Corolla ra-
dial. Seeds flat with broad wings. Has flavonoids of type A only. 2n = 52.
Distribution and taxa.—Tropical forests from Mexico to Peru. One genus,
Cobaea, with four sections and eighteen species: C. aequatoriensis, C. ascher-
1378 BRIT.ORG/SIDA 20(4)
soniana, C. bidaurita, C. campanulata, C. flava, C. gracilis, C. lutea, C. minor, C.
pachysepala, C. paneroi, C. penduliflora, C. pringlei, C. rotundiflora, C. scandens,
C. skutchii, C. stipularis, C. trianae, C. triflora. See the recent monograph of
Prather (1999).
Tribe 3. Bonplandieae Baill, Hist. Pl. 10:342. 1890. Typr: Bonplandia geminiflora
Cav.
Subshrubs with woody base and herbaceous shoot. Leaves simple with a broad
blade and serrate margin or lobed. Calyx wholly herbaceous and synsepalous.
Corolla bilateral. Seeds plump with narrow wings or wingless. Has flavonoids
of type B only. 2n = 3
Distribution and taxa.—Mexico and Guatemala. One genus, Bonplandia,
with two species, B. geminiflora and B. linearis.
Tribe 4. Loeselieae J.M. Porter & L.A. Johnson, Aliso 17:84. 8 Jun 1998. Tribe
Loeselieae V.E.Grant; Amer J. Bot. 85:744. 17 Jun 1998. TYPE: Loeselia ciliata
L.
Subshrubs, perennial herbs, and annuals. Leaves simple with broad blade and
entire or serrate margin. Inflorescence bracts with prominent veins subtend
individual flowers; these are usually large and leaflike, but may be small. Ca-
lyx synsepalous and membranous with no or only small herbaceous regions.
Corolla radial or bilateral. Seeds with narrow wings or wingless. Flavonoid
groups A and C present, but type B absent. 2n = 18.
Distribution and taxa.—Mexico to southern Texas and Arizona and to
northern South America. Often in the temperate zone of tropical mountains.
One genus, Loeselia, with fifteen species: L.amplectens, L. caerulea, L. ciliata, L.
cordifolia, L. glandulosa, L.grandiflora, L.greggii, L. hintoniorum, L.involucrata,
L. mexicana, L. nepetifolia, L. pumila, L. purpusii, L. rupestris, L. rzedowshi. See
Turner (1994) for a recent generic treatment.
Subfam. 2. Polemonioideae Brand., Pflzr. 4(250):30. 1907. Type: Polemonium
caeruleum L.
Subshrubs, perennial herbs, and annuals, of temperate and boreal zones. Seeds
not winged. Pollen pantoporate. Chromosomes mostly large or medium-sized,
but small in one tribe.
Tribe 5. Polemonieae Arn., Encycl. Brittanica, ed. 7, 5:121. 1832. Tribe Polemonieae
Meisn., Pl. Vasc. Gen. 273. 1839. TYPE: Polemonium caeruleum L.
Perennial and annual herbs. Leaves alternate. Leaves pinnately compound (in
Polemonium) or simple and pinnately lobed, or simple and entire in reduced
lorms (in the other genera). Capsule locules commonly containing | to few seeds
each; sometimes many-seeded in Navarretia. Seeds dark brown or black, some-
times plump and rounded, sometimes small. Pollen generally pantoporate, but
GRANT, TAXONOMY OF THE POLEMONIACEAE 1379
zonocolporate in some species of Collomia. Flavonoids of type C absent; type B
absent in Polemonium and Allophyllum, but present in Collomia and
Navarretia. Chromosomes medium-sized or large, x = 9 and 8.
Distribution and taxa.—North America, extending to Eurasia. Genera: Po-
lemonium, Collomia, Allophyllum, Navarretia. About 78 species.
Polemonium is set apart from the other three genera by leaf form and some
molecular characters Johnson et al. 1996). It could well be separated in a tribe
of its own, as in the Porter and Johnson (2000) system. On the other hand, it is
allied to the other genera by seed and pollen characters and by other molecular
evidence (Prather et al. 2000). A conservative disposition is made here.
Tribe 6. Phlocideae Dumort., Anal. Fam. PL, 25. 1829. Gruppe Phloginae Rchb.,
Handb. Nat. Pflzsystems, ed. 1, 194. 1837. Tribe Phlogieae Rohb. ex J.M. Porter @
L.A. Johnson, Aliso 17:84. 1998. Tyre: Phlox glaberrima L.
Subshrubs, perennial herbs, and some annuals. Leaves or at least the lower leaves
opposite; true leaves absent in the small genus Gymnosteris. Leaves simple with
narrow blade and entire margin. Capsule locules mostly l-seeded. Seeds brown,
sometimes plump and rounded, sometimes small. Flavonoids of type B lacking,
but type C present. Chromosomes medium-sized or large, x = 7 and 6.
Distribution and taxa.—North America, extending to Asia. Genera: Phlox,
Microsteris, Gymnosteris. About 66 species.
Phlox (x = 7) could be derived from Collomia (x = 8) in the Polemonieae.
Microsteris (x = 7) is a reduced annual derived from a perennial Phlox.
Gymnosteris (x =6) is a reduced leafless annual formerly thought to be close to
Collomia; but phenetic and molecular evidence now indicate that its closest
relative is Phlox or Microsteris (Porter & Johnson 2000). See Ferguson and Jansen
(2002) on molecular relationships in Phlox.
Tribe 7. Leptodactyloneae V.E. Grant, Amer. J. Bot. 85:746. 1998. TYPE:
Leptodactylon californicum Hook & Arn.
Subshrubs, perennial herbs, and annuals. Leaves mostly opposite. Leaves pal-
mately divided, or simple and entire in reduced forms. Corolla veins non-anas-
tomosing, in contrast to other tribes (see explanation in Materials and Meth-
ods). Capsule locules containing several to many seeds each, rarely l-seeded.
Seeds sandy or pale-colored, or sometimes brown in Leptodactylon. Flavonoids
of types B and C present. Chromosomes small, x = 9.
Distribution and taxa.—Semiarid and arid habitats including deserts in
western North America. Genera: Leptodactylon, Linanthus, Maculigiia. About
50 species.
In the system of Porter and Johnson (2000), Linanthus s.l. is subdivided
into two genera, Linanthus ss.and Leptosiphon, on cladistic grounds. These two
taxa are indeed well differentiated, but they are also united by some common
characters, and they could be treated as either subgenera of one genus or two
1380 BRIT.ORG/SIDA 20(4)
separate genera. A conservative treatment as Linanthus s.. is preferred here
because it shows the relationships of the two branches. Maculigilia is a recent
segregate genus for the old distinctive desert species Gilia maculata (see Patter-
son 1989; Grant 1999). Maculigilia maculata is probably a derivative of some
xerophytic member of Linanthus.
Leptodactylon and Linanthus form a closely related and isolated genus pair.
This group was formerly placed in the tribe Gilieae on macroscopic characters
(Grant 1959), but the more recent evidence of flavonoids and cpDNA and nrDNA
point to a relationship with Phlox (Smith et al. 1977, 1982; Johnson et al. 1996:
Porter 1997). Leptodactylon/Linanthus is placed in the tribe Phlocideae in the
Porter and Johnson (2000) system.
However, in other characters, Leptodactylon and Linanthus are unlike the
locideae (or Polemonieae); namely, palmate leaves, non-anastomosing corolla
veins, sandy or pale-colored seeds, small chromosomes, and an affinity for semi-
arid and arid habitats. The present treatment attempts to resolve the similari-
ties and important differences by placing these genera in a separate tribe
Leptodactyloneae next to the Phlocideae. This makes for a better definition of
both tribes.
—
Dp}
ri
Subfam. 3. Acanthogilioideae (V.E. Grant) J.M. Porter & L.A. Johnson, Aliso 19:60.
2000. Tribe Acanthogilieae VE. Grant; Amer. J. Bot. 85:744. 1998. Type: Gilia
thogilia gloriosa A.G. Day & R. Moran.
~—
gloriosa ae Acant
Shrubs, subshrubs, and herbs of arid habitats, commonly in deserts. Seeds
winged or wingless. Pollen zonocol porate Chromosomes mostly medium-sized
or large, but small in one tribe.
Tribe 8. Acanthogilieae VE. Grant, Amer. J. Bot. 85:744. 1998. Typr: Gilia gloriosa
Brand., Acanthogilia gloriosa A.G. Day & R. Moran.
Spiny desert shrub. Leaves dimorphic, with primary leaves modified as persis-
tent spines, and small deciduous green leaves in the axils of the spines. Capsule
locules containing 1-6 seeds each. Seeds flat with broad wings. Chromosomes
small, x =9
Distribution and taxa.—Endemic in central Baja California. One genus and
species Acanthogilia gloriosa. Detailed information about this interesting taxon
is given by Day and Moran (1986).
Tribe 9. Gilieae (Rchb.) VE. Grant, Nat. Hist. Phlox Fam. 120.1959. Gruppe Gilieae
Rchb., Handb. Nat. Pflzsystems, ed. 1, 194, 1837. Tribe Gilieae J.M. Porter & L.A.
Tohneon: Aliso 19:63. 2000. TypF: Gilia laciniata Ruiz & Pav.
Subshrubs, perennial herbs, and annuals; not spiny. Leaves not dimorphic; leaves
pinnately divided or dissected with narrow segments, or small and linear in
reduced forms. Capsule locules usually many-seeded. Seeds sandy or pale-col-
ored, mostly angular, sometimes banana-shaped. Pollen generally
GRANT, TAXONOMY OF THE POLEMONIACEAE 1381
zonocolporate, but deviating toward pantoporate in some species of Eriastrum.
Flavonoids of type B present; type C uncommon (occurs in Gilia and Langloisia)
or absent (Gilia, Ipomopsis). Chromosomes mostly medium-sized, sometimes
large; x = 9, 8, and 7.
Distribution and taxa.—Widespread in arid habitats, common in deserts.
The genera as treated here are Gilia, Tintinabulum, Ipomopsis, Eriastrum,
Langloisia. About 105 species.
A broad concept of Gilia as a genus composed of six sections is adopted
here and elsewhere (Grant 1999, 2001). Gilias.l. with a basic chromosome num-
ber of x = 9 in all sections (and x = 8 in part of one section) is basal in the tribe.
Tintinabulum (x = 9) is a small specialized genus close to Gilia. Ipomopsis,
Eriastrum, and Langloisia with x = 7 appear to be derivatives of Gilia (Grant
—
Molecular cladists consider Gilias.l.to be polyphyletic, and split it up into
a series of small genera Johnson et al. 1996; Porter & Johnson 2000). | contend
that this viewpoint is based, first, on the cladistic definition of monophyly and,
second, on an overreliance on the molecular cladograms, and that Gilia sl. is
monophyletic by the traditional definition of monophyly (Grant 1999, 2001).
PHYLOGENY
The Polemoniaceae is evidently derived from an ericalean stock (Brown 1938:
Porter & Johnson 1998; Johnson et al. 1999). The basal living forms of the fam-
ily are the tropical and subtropical genera. There are five of these (Cantua,
Cobaea, Bonplandia, Loeselia, Acanthogilia), all very different from one another.
In this group Cantua comes closest to the ericalean stock, while the other gen-
era exhibit derived characters of one sort or another.
he subtropical desert shrub, Acanthogilia, differs markedly from Cantua,
but also shares some inconspicuous but significant characters with it (Day &
Moran 1986). These workers suggest (p. 125) that “Acanthogilia may be a spe-
cialized desert descendant of a diploid line also ancestral to Cantua.”
One of the significant characters is the spatial distribution of pores on the
pollen grains, whether pantoporate or zonocolporate. Another is ecological pref-
erences: mesic or xeric. The pollen pore character has proven to be a valuable
marker for revising the classification of species groups and genera (Day & Moran
1986; Grant & Day 1999), and can be extended to higher taxa. Cantua and three
related genera (Cobaea, Bonplandia, and Loeselia, comprising the subfamily
Cobaeoideae) are pantoporate and mesophytic. Acanthogilia is zonocol porate
and xerophytic. Day and Moran’s (1986) hypothesis can be rephrased as the sug-
gestion of an early split between the ancestors of the pantoporate Cobaeoideae
and the ancestor of the zonocolporate Acanthogilia. The molecular evidence of
Prather et al. (2000) is in agreement with this hypothesis.
This split persists in the derived, temperate, mainly herbaceous tribes. Day
1382 BRIT.ORG/SIDA 20(4)
and Moran (1986) point to characters which Gilia and its close relatives share
with Acanthogilia. The tribe Gilieae, as circumscribed here and in Grant (1998),
is zonocolporate and xerophytic. It could well have an unknown ancestor in
common with Acanthogilia.
The mid-Eocene plant Gilisenium from Utah is close to Gilia ina number
of characters (Lott et al. 1998). It is desirable to determine its pollen pore char-
acter, if possible.
The tribes Polemonieae and Phlocideae are pantoporate and generally
mesophytic. They may be derived from a Bonplandia-like ancestor in the
Cobaeoideae. The Polemonieae and Phlocideae extend into cold northern areas
and are the only representatives of the family in northeastern North America
and Eurasia. The Leptodactyloneae is also pantoporate and is related to the
Phlocideae but is xerophytic.
The two main temperate phylads often occur in the same area, especially
in western North America, where some species or groups have crossed over from
one type of habitat to another. In this situation the two phylads have given rise
to species of annuals which converge in their general morphological charac-
ters. For example, some small-flowered species of Allophyllum (Polemonieae)
closely resemble certain small-flowered species of Gilia (Gilieae), and were
mistakenly treated as Gilia until recently (Grant 1999; Grant & Day 1999).
caer
DISCUSSION
Several causal factors contribute to the numerous differences between the taxo-
nomic and molecular cladistic systems of the Polemoniaceae. Two of these are
theoretical: (1) the use of different systematic units, namely, taxa and clades:
and (2) the application of different definitions of monophyly. These two factors
were discussed in the introduction. The two approaches also use different evi-
dence and this of course leads to different results. We will consider here: (3)
differences in the types of characters used; and (4) differences in the range and
breadth of the database.
(3) Molecular systematists working in the Polemoniaceae (and other plant
groups) use DNA sites in chloroplasts and mitochondria, which are parts of the
cytoplasmic genome. Phenetic characters used in taxonomy are determined
mainly by the chromosomal genome (see Grant 1975, 2003, for review). Discor-
dance between plastid and mitochondrial evidence, on the one hand, and phe-
netic characters on the other, can be expected and is often found. Ribosomes,
also used in molecular systematics of Polemoniaceae and other families, are a
part of the chromosomal genome. They can vary independently of plastids and
mitochondria, and for that matter, independently of unlinked chromosomal
genes. Here again, incongruence between the various sources of evidence can
be expected (see Grant 2003, for further discussion).
(4) Molecular cladistic treatments of the Polemoniaceae (and those of other
_—
GRANT, TAXONOMY OF THE POLEMONIACEAE 1383
plant groups) are based on one or a few genes. The treatment of Johnson et al.
(1996) blocks out informal systematic groups on the basis of one chloroplast
gene. Taxonomic systems, by contrast, are based on numerous phenetic char-
acters determined by scores or hundreds of genes and gene systems (Grant 1975,
for review). These contrasts between approaches represent the extremes. Mod-
ern taxonomists pues the aN mace of molecular systematists. And many
molecular in reaching their conclusions.
Factors (1) and 2) bring about different results but do not affect the accu-
racy of the results, as noted in the introduction. The type and range of charac-
ters (factors 3 and 4) do have an effect on the robustness of the results. Molecu-
lar cladograms are essentially gene trees, or character phylogenies to use an
older term. Single characters often work well in one part of a group but break
down elsewhere in the same group. If the goal isa natural classification of whole
organisms, one must assemble a broad range of characters, some of which may
conflict with others, and one should then follow the weight of the evidence, as
is standard procedure in taxonomy.
Taxonomic classifications thus have certain methodological advantages.
Another advantage lies in the area of convenience and desirability. Named simi-
larity groups, and hierarchies of such groups, are generally useful for identifi-
cation, information retrieval, and cataloging. It would be difficult to write a
flora, using clades instead of taxa, and sucha flora would be difficult to use if it
were written (see Diggs & Lipscomb 2002).
For several reasons, therefore, it is desirable to make an updated taxonomic
classification of the Polemoniaceae available. But why stop with the
Polemoniaceae. Numerous other plant groups have been revised recently by
molecular cladistic methods, but are represented in the literature by old taxo-
nomic treatments. These groups should also be re-revised by taxonomists, us-
ing the new molecular evidence, but handling all the evidence by taxonomic
methods. Plant taxonomy has much work to do.
ACKNOWLEDGMENTS
I discussed the problem with Billie L. Turner, Tom Wendt, and Karen A. Grant. J.
Mark Porter and Carolyn J. Ferguson read the manuscript. The help of these
colleagues is greatly appreciated.
REFERENCES
Brown, W.1938.The bearing of nectaries on the phylogeny of flowering plants. Proc. Amer.
Philos. Soc. 79:549-595.
Cartouist, S.,V.M. EckHart, and D.C. MicHener. 1984. Wood anatomy of Polemoniaceae. Aliso
10:547-572
Day, A.and R. Moran. 1986. Acanthogilia,a new genus of Polemoniaceae from Baja Califor-
nia, Mexico. Proc. Calif. Acad. Sci.44:111-126.
1384 BRIT.ORG/SIDA 20(4)
Diccs, G.M. and B.L. Liescoms. 2002. What is the writer of a flora to do? evolutionary tax-
onomy or phylogenetic systematics? Sida 20:647-674.
Ferauson, C.J. and R.K. JANseN. 2002. A chloroplast DNA phylogeny of eastern Phlox
(Polemoniaceae): implications of congruence and incongruence with the ITS phylog-
eny. Amer. J. Bot. 89:1324-1335.
Grant, V. 1959. Natural history of the phlox family; systematic botany. Martinus Nijhoff, The
Hague.
Grant, V.1975. Genetics of flowering plants. Columbia Univ. Press, New York, NY.
Grant, V. 1998. Primary classification and phylogeny of the Polemoniaceae, with com-
ments on molecular cladistics. Amer. J. Bot. 85:741-752.
Grant, V. 1999. Classification of the genus Gilia (Polemoniaceae), Phytologia 84:69-86.
Grant, V.2001.A guide to understanding recent classifications of the family Polemoniaceae.
Lundellia 4:1 2-24.
Grant, V. 2003. Incongruence between cladistic and taxonomic systems. Amer. J. Bot.
90:1 263-1270.
Grant, V. and A. Day. 1999. Transfer of some species from Gilia to Allophyllum and
Tintinabulum, and the effects of the transfer on the generic definition of Gilia
(Polemoniaceae). Phytologia 84:368-382.
Joxnson, L.A., L. Schutz, D.E. Soutis, and PS. Soutis. 1996. Monophyly and generic relation-
ships of Polemoniace based on matK sequences. Amer. J. Bot. 83:1207-1224.
Jounson, L.A., D.E. Soutis,and P.S. Souris. 1999. Phylogenetic relationships of Polemoniaceae
inferred from 18S ribosomal DNA sequences. Plant Syst. Evol. 214:65-89.
Lor, L.A, S.R. MancHester, and D.L. DitcHer. 1998. A unique and complete polemoniaceous
plant from the middle Eocene of Utah, USA. Rev. Palaeobot Palynol. 104:39--49.
Mayr, E.and W. Bock. 2002. Classifications and other ordering systems. J. Zool. Syst. Evol. Res
40:169-194.
Patterson, R. 1989. Taxonomic relationships of Gilia maculata (Polemoniaceae). Madrono
36:15-27.
Porter, JM. 1997. Phylogeny of Polemoniaceae based on ribosomal internal transcribed
DNA sequences. Aliso 15:57-77.
Porter, JM. and L.A. JoHNson. 1998. Phylogenetic relationships of Polemoniaceae; infer-
ences from mitochondrial nad7b intron sequences. Aliso 17:157-188.
Porter, JM. and L.A. JoHNson. 2000. A phylogenetic classification of Polemoniaceae. Aliso
19:55-91.
Prater, L.A. 1999. Systematics of Cobaea (Polemoniaceae). Syst. Bot. Monogr. 57:1-81.
Pratuer, L.A., C.J. Fercuson, and R.K. JANseN. 2000. Polemoniaceae phylogeny and classifica-
tion; implications of sequence data from the chloroplast gene ndhF. Amer. J. Bot.
87:1300-1308.
Smit, D.M., C.W. Giennic, J.B. Harsorne, and C.A. Witiams. 1977. Flavonoid diversification in
the Polemoniaceae. Biochem. Syst. Ecol. 5:107—115.
SmitH, D.M., CW. Gtennie, and J.B. HArBorNe. 1982. Flavonoid patterns in Leptodactylon and
Linanthus. Biochem. Syst. Ecol. 10:37-42.
GRANT, TAXONOMY OF THE POLEMONIACEAE 1385
STeeLe, K.P. and R. Vicatys. 1994. Phylogenetic analyses of Polemoniaceae using nucleotide
sequences of the plastid gene matkK. Syst. Bot. 19:126-142.
STUCHLIK, L. 1967a. Pollen morphology in the Polemoniaceae. Grana Palynol. 7:146-240.
STUCHLIK, L. 1967b. Pollen morphology and taxonomy of the family Polemoniaceae. Rev.
Palaeobot. Palynol. 4:325-33.
Taytor, T.N. and D.A. Levin. 1975. Pollen morphology of Polemoniaceae in relation to sys-
tematics and polliniation systems; scanning electron microscopy. Grana 15:91-112.
Turner, B.L. 1994. Synopsis of the North American species of Loeselia (Polemoniaceae).
Phytologia 77:318-337.
1386 BRIT.ORG/SIDA 20(4)
BOOK NOTICES
Mito L Harpsteab, THOMAs J. SAUER, and WILLIAM F. BENNETT. 2001. Soil Science
Simplified, 4th edition. (ISBN 0-8138-2942-9, hbk.). lowa State Univ. Press,
2121 South State Ave., Ames, lowa 50014, U.S.A (Orders: 800-862-6657,
ww w.isupress.com). $42.95, 225 pp., numerous b&w drawings and half-
tones, 6" x 9",
This book is suitable as an undergraduate text or as a reference for biologists or other scientists not
trained in soil science. It systematically covers the aa biological and chemical properties of
ue fertility; lassification and sual and soil man-
}
soils: soil formation, soil water, heat oe a
ion f | il colloi soil classi-
agement, conserva
fication, and soil surveys the sest helpful in smicnorente soil n
types of clays and their characteristics are exp laa Also, the various word roots, prefixes anc
bclas «plained. It includes a glossary and index.
3
—
J
ristics. The different
1 suf-
fixes used in the names of soil classes and
The weakest aspect of the book is the quality of the lasetians In many figures, the lines are
excessively heavy, black, and run-together. Many of the drawings are not well-executed and fail to
show the described features. As a result, much space is wasted that could have been better used by
replacing with text or photographs.—Roger W. Sanders, Associate Collections a Botanical Re-
Texas, 509 Pecan St, Fort Worth, TX 76102-4060, U.S.A., g.
search Institute of
WILHELMINA FEEMSTER JASHEMSKI and FREDERICK G. Meyer (eds.). 2002. The Natural
History of Pompeii. ISBN 0-521-80054-4, hbk.). Cambridge University Press,
The Edinburgh Building, Cambridge CB2 2RU, UK; 100 Brook Hill Drive,
West Nyack, NY 10994-2133, U.S.A. (Orders: 845-353-7500, 845-353-4141
fax, http://www.cup.org/). $170.00, 502 pp., 387 figs. (b/w, color, drawings,
etc.), 9" x 12",
From the dustjacket.—This volume brings together the work of geologists, soil specialists,
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ologists, Aiea ie and archaeologists, affor
ncient sites. The detailed and rigor-
zoologists, ornithologists, mammologists, herpet
ough picture of the landscape, flora and fauna of the a
ously scientific catalogues, which are copiously illustrated, provide weneeilist of the flora and
upon which future generations of scholars can continue to build.”
The catalogue of plants by Wilhelmina F Jashemski, Frederick G. Meyer, anc
enumerates and discusses in detail 184 plants. The chapter is 101 pages. The evidence for the plant
list comes primarily from the urban sites Pompeii and Herculaneum, and a few villas, which give a
popular ornamental plants ose ac ae the many home gardens, parks, and
1 those used for medicine and
ing a thor
‘auna
| Massimo Ricciardi
good poe of the
other public places, as well as plants that |
other Pai especially food There is an extensive list of references. This is an impressive work
of audiences.—Barney Lipscomb, Botanical Research Institute of Texas,
vad
that will be of interest.a variet
509 Pecan St, ae Worth, TX 76102-4060, U.S.A., barne y@britorg.
SIDA 20(4): 1386. 2003
A SYNOPSIS OF MACHAERANTHERA (ASTERACEAE:
ASTEREAE), WITH RECOGNITION OF SEGREGATE GENERA
David R. Morgan Ronald L. Hartman
Department of Biology Department Tae
Western Washington University University of Wyom
Bellingham, WA 98225-9160, U.S.A. Laramie, WY 82070-3 es co SA.
morgan@biol.wwu.edu rhartman@uwyo.edu
ABSTRACT
ee bara aus oF Machaera nthera si, naCORDOratty 6 moleculis, morphological, cytological,
tion of the genus. To bring the classification of Machaeranthera more in line with current wader:
standing oF -dtpevelunionayy history, revis are proposed. Machaeranthera is cir-
aT -]
and other species that ha i included in Machaeranthera are
apportioned into three addidonel genera. Dieterid is revived to accommodate three of these species,
Xanthisma is expanded to accomodate 16 more species, and the new genus Arida is proposed for
those remaining. New ae oe: for the species in Dieteria, Xanthisma, and Arida are provided.
RESUMEN
In ns eee recientes en Machaeranthera s.l. que incluyen datos moleculares, morfoldgicos,
citologicos, y quimicos dan como resultado unas relaciones que no concuerdan con ninguna
pani reciente del género. Para poner la clasificacion de Machaeranthera mas en linea con
el conocimiento actual de su historia evolutiva, se proponen revisiones de su taxonomia.
Machaerant nerd se cinGHSCEDS para melee dos SPECS y otras especies que se habian incluido en
Jieterid se rescata para acomodar tres
de estas especies, Xanthisma se expande: ae a incluir 16 especies mas, y se propone el nuevo género
Arida para las restantes. Se realizan las nuevas combinaciones para las especies de Dieteria,
Xanthisma, y Arida
The genus Machaeranthera Nees (Asteraceae: Astereae) was established to ac-
commodate the single species M. tanadcetifolia Nees. Since its inception, many
different circumscriptions of Machaeranthera have been proposed, and the
genus has included various parts of several other genera, with little agreement
on its composition or its relationships with other taxa (see Morgan and Simp-
son 1992 for a more detailed taxonomic history).
The most recent published treatment of Machaeranthera was by Hartman
(1990), who included 36 species in the genus, excluding several taxa that are
now placed in the genera Hazardia Greene (Hartman 1990), Rayjacksonia RL.
Hartman & M.A. Lane (Lane & Hartman 1996), Tonestus A. Nelson (Nesom 1991),
and Xylorhiza Nutt. (Hartman 1990). The circumscription of Machaeranthera
adopted by Hartman (1990) included species with both cyanic (white, blue, pink,
purple) and yellow rays, and apportioned them into two subgenera and eight
sections. We will refer to this circumscription as Machaeranthera s.l.
SIDA 20(4): 1387-1416. 2003
1388 BRIT.ORG/SIDA 20(4
=
Investigations of Machaeranthera s.l. using molecular evidence have helped
to clarify its systematics. Restriction site data from chloroplast DNA (cpDNA)
indicated that Machaeranthera section Psilactis (A. Gray) Turner & Horne was
not a close relative of Machaeranthera s.l. (Morgan & Simpson 1992; Morgan
1993). This section was removed from Machaeranthera s.l. by Morgan (1993),
leaving 30 species in the genus. More recent investigations of Machaeranthera
(sensu Morgan 1993) employed DNA sequence data from nuclear ribosomal
DNA internal and external transcribed spacers (n9rDNA) (Morgan 1997; Mor-
gan 2003). Comparison of this evidence with cpDNA data indicated that the
evolution of Machaeranthera and its relatives has been complex, involving as
many as seven occurrences of reticulate evolution. The molecular evidence also
indicated that other genera (e.g., Odnopsis Nutt., Pyrrocoma Hook., and
Xanthisma DC.) are closely related to various parts of Machaeranthera (Fig. LD.
The relationships supported by a combination of molecular, morphologi-
cal, cytological, and chemical evidence do not support any recent circumscrip-
tion of Machaeranthera. Therefore, a new classification is needed to bring the
taxonomy of the genus more in line with current understanding of its relation-
ships. A case could be made for including Odnopsis and Pyrrocoma in an ex-
panded Machaeranthera,and cpDNA evidence provides some support for such
a treatment (Fig. |). However, nrDNA data do not support this circumscription
(Fig. L). Furthermore, we contend that enlarging Machaeranthera through ac-
cretion would obscure the phylogenetic patterns within the complex, and would
result in a large, heterogenous, and unwieldy genus that would be difficult to
characterize satisfactorily. We therefore propose that Machaeranthera (sensu
Morgan 1993) be divided into four genera, as described below.
KEY TO THE GENERA TREATED HERE
. Ray corollas yellow (heads fallete) Xanthisma
ne ollas white isk drying), pink, red-purple, or purple,
or ners era adi ate
2. Leaves deeply sinnaurd to bipinnatifid throughout, at least many of the teeth
and lobes sharply acute with bristle tips, bristles 0.2-1 mm long; plants annual;
ee of subulate bristles, dorsi-ventrally flattened near base, bases
overlap Machaeranthera
Vg
Leaves ene to toothed or lobed, if pinnatifid to bipinnatifid throughout then
lobes often rounded, with or without an apiculum but not bristle tipped; plants
annual to strongly perennial; pappus of filiform to subulate bristles, terete to
flattened near base, bases overlapping or not overlap
3. Plants stror ney Petras Wen a Pralic hed caudex aprooted sometimes bi-
nia al ir + IAA | ~ P| +h
Ik covered with
one 0.3-3 mm long, Shen: forming an alveolate retict tlum:; pappus bristles
subulate, flattened near base, bases strongly overlapping Xanthisma
3. Plants taprooted annuals or short lived perennials (forming vegetative ro
tes in Arida blepharophylla, a species also exceptional, as indicated, in the
following characters); receptacles usually naked (or scales rarely to 0.5 mm
n
@)
“Ajaanadsad ,,“Y,, Pue ,"W,, PayelAaaqqe ae pusiyjUDY
c
\CDOOL;
J
7
wou aie AuabojAyd YNQJU ay} UO sayrueg ay) Mojaq sanjeA pue ‘SISA
c
1
tha
ba
ainyepuaw
‘
¢ ‘DI4 Woy
raed L “lf
Ws
+
VNU
AVING
-
.
JLIPF
cpDNA sa
67
99
100
p
74 100-——
52
89
49 oe
88
88 56
lsocoma veneta
glandulifera
essingia filaginifolia
onlooor
Xylorh
M. tanacetoa
nopsis wardii
oar: engemenn
Dieteria bigelov
Dieteria canescens
Pyrrocoma crocea
Pyrrocoma lanceolata
Arida riparia
rida ene
Arida
Arida sephoh
cephal
fe)
blephariphyllum
crutchfieldii
x KKK MK KRM KK
aa of a
rad
spinulosum
nrDNA
17
28
28
30
31
46
86
92
Isocoma veneta
psis
Pyrrocoma crocea
Pyrrocoma lanceolata
Hazardia squarrosa
ardia cana
Lessingia glandulifera
Lessingia filaginifolia
Haplopappus
M. tanacetifolia
Oonopsis wardii
Oonopsis engelmannii
Xylorhiza
peta seectees
Dieter
And 2s lpheoo
spinulosum
racile
gymnocephalum
coloradoense
xxx xX XK KK XX
59398
oO
no}
pF
©
Be]
ay
<
c
3
viscidum
VUIHINVYIVHVW 40 SISGONAS ‘NVWLYVH ONY NVOYOW
68EL
1390 BRIT.ORG/SIDA 20(4)
long); pappus bristles filiform, not flattened near base (or slightly so), bases
not (or slightly) overlapping.
Ray florets with prominent pappus (often eradiate in the glandular-pu-
D. canescens var. shastensis); leaves entire to ipenise and plants
variously pubescent with glandular and/or ae
plains, and basins of W U.S.A; SW Canada; extreme N Mexico Dieteria
4. Ray florets with pappus absent (5 species; nue in the glabrous
carnosa) or present; If pappus pie a cae ee to bieianatind
throughout or (ifleaves entire pt for bristles
or apiculate or bristle tipped teeth on leaf margins; deserts of SW U.S.A,;
Mexico Arida
Machaeranthera Nees, Gen. & Sp. Asterearum 224. 1832. Type: Aster tanacetifolius
Kunth.
Se eel ced Pappochroma Nutt., J. Acad. Nat. Sci. Phila. 7:34. 1834. Dieteria subgenus
) Nutt., Trans. Amer. Philos. Soc. 7:302. 1840. apie cue Pappochroma
(Ni itt.) Wa lpers, Rep. Bot. Syst. 2:587. 1843. Type: Chr Nutt.
Aster section Machaeranthera (Nees) Benth. & Hook. on PL 2272 1873. Aster subgenus
Machaeranthera (Nees) A. Gray, Proc. Amer. Acad. Arts 16:99. 1880.
Machaeranthera series Verae Cronquist & D.D. Keck, Brittonia 9:238. 1957, nom. illeg. ICBN: 52.1).
Herbs taprooted annuals or biennials, 5-100 cm tall. Stems erect to ascending,
much branched when well developed, moderately to densely pubescent with
glandular and nonglandular hairs. Leaves alternate; pinnatifid or bipinnatifid,
deeply lobed or dissected, moderately to densely pubescent with glandular and
nonglandular hairs, apices, including lobes and teeth, mucronate to bristle
tipped, bristles if present 0.2-1 mm long. Heads radiate. Involucres broadly tur-
binate to hemispheric. Phyllaries in 3-6 imbricate series, graduated in length,
appressed, spreading, or reflexed, moderately to densely pubescent with glan-
dular and nonglandular hairs; bases indurate, apices herbaceous. Receptacles
alveolate, flat to convex, naked or with scales to 0.4 mm long. Ray florets pistil-
late, fertile; corollas blue, violet, or purple. Dise florets bisexual, fertile, corollas
yellow, narrowly funnelform; tubes and limbs glabrous or glabrate, lobes 0.3-1
mm long, glabrous to pubescent. Cypselae of ray and disc florets distinctly di-
morphic, narrowly to broadly obovate, sparsely to densely pubescent; moder-
ately thick walled, with 4-9 ribs per face, 2.4-3.8 mm long, those of ray florets
~3 sided, rounded abaxially, of disc florets somewhat compressed laterally; pa-
ppus white to tawny, of ray florets 1/2 to nearly equalling length of disc florets;
bristles coarsely barbellate, in 1-3 series, dorsi-ventrally flattened and dissimi-
lar in width near base, bases overlapping, mostly 4.5-6.5 mm long. n = 4.
The circumscription of Machaeranthera adopted here results in a genus
of two species. We apportion the remaining 28 species of Machaeranthera
(sensu Morgan 1993) to Dieteria, Xanthisma, and Arida (see below). The three
species placed in Dieteria could potentially also be included in Machaeranthera
because of a number of similarities. All members of the two genera produce
flavonols, and all have chromosome numbers of n = 4, obovate fruits, and nar-
—
MORGAN AND HARTMAN, SYNOPSIS OF MACHAERANTHERA 1391
row phyllaries that are often reflexed (Hartman 1976, 1990; Turner 1987). This
relationship was also supported by cpDNA evidence (Fig. 1). However, nrDNA
evidence conflicted with cpDNA (Fig. 1), and the substantial conflict between
them complicates the issue by suggesting reticulate evolution in the ancestors
of Machaeranthera, Dieteria, or both. In addition, cpDNA and nrDNA evidence
both supported a close relationship between Machaeranthera and Oénopsis, and
expansion of Machaeranthera to pide Dieteria would also have to include
Oonopsis, which differs from Machaeranthera in many ways (Morgan & Simp-
son 1992). Additional synonymy for Machaeranthera is included in Turner
(1987)
KEY TO THE SPECIES OF MACHAERANTHERA
1. Involucres hemispheric; herbaceous phyllary apices spreading to reflexed; disc
corolla lobes mostly 0.3-0.7 mm long, glabrous or glabrate 1. Machaeranthera
tanacetifolia
1. Involucres broadly turbinate; herbaceous phyllary apices appressed; disc corolla
lobes mostly 0.7—1 mm long, pubescent 2. Machaeranthera tagetina
1. Machaeranthera tanacetifolia (Kunth) Nees, Gen. & Sp. Asterearum 225. 1832.
ster tanacetifolius Kunth, Nov. Gen. & Sp., ed. folio, 4:74. 1820. Aster chry-
santhemoides Willd. ex Spreng., Syst. Veg. 3:538. 1826. Type: MEXICO: “Colitur in
horto Mexicano,” s.d., Humboldt s.n. (HOLOTYPE: P nv, IDC microfiche 6209. 97.1116;
possible ISOTYPE: B- W; PHOTOISOTYPE: TEX).
Chrysopsis coronopifolia Nutt., J. Acad. aa Sci. Phila. 7:34. 1834. Dieteria coronopifolia (Nutt.)
Nutt., Trans. res r. Philos. Soc., ser. 2, 7:302. 1840. cence hera coronopifolia (Nutt.) A
Nelson, Bot. Gaz. ee 1904. TYP :U S.A. NORTH DAK “Towards the sources of the Bliss
souri” (probably near Fort cre Jul-Aug 1811, cc. s.n. ISOTYPE: GH; probable ISOTYPE:
Y)
2 esrieahcet ss tagetina Greene, Pittonia 4:71. 1899. Aster tagetinus (Greene)
e, Contr. U.S. Natl. Herb. 25:263. 1925. Type: U.S.A. ARIZONA. Cochise Co::
near aoe Huachuca, 1891, Wilcox s.n. (HOLOTYPE: US).
Dieteria Nutt., Trans. Amer. Philos. Soc., ser. 2, 7:300. 1840. LECTOTYPE, here desig-
nated: Aster canescens Pursh.
asnie nen section Hesperastrum A, Gray, Proc. Amer. Acad. Arts 6:539. 1865. Aster subge-
Hesperastrum (A. Gray) A. Gray, Proc. Amer. Acad. Arts 16:97. 1880. Aster section
on (A. Gray) A. Gray, Syn. Fl. N. Amer. 1(2):174. 1884. TYPE: Machaeranthera
shastensis A. Gray.
Machaeranthera subgenus Dieteria (Nutt.) Greene, Pittonia 3:59. |
ee nera series Variabiles Cronquist & D.D. Keck, Brittoni: 20: 237, 1957, nom. illeg. (CBN:
ioe
Herbs ae annuals, biennials, or perennials, 10-100 cm tall. Stems erect
to ascending, much branched when well developed, glabrous to variously pu-
bescent with glandular and/or nonglandular hairs. Leaves alternate; entire to
irregularly serrate or dentate, moderately to densely pubescent with glandular
1392 BRIT.ORG/SIDA 20(4
and/or nonglandular hairs, apices, including lobes and teeth, spinescent due
to indurated apiculae, bristles absent. Heads radiate or eradiate. Involucres tur-
binate, campanulate, or hemispheric. Phyllaries in 3-12 imbricate to
subimbricate series, graduated in length, appressed, spreading, or reflexed, gla-
brous or variously pubescent with glandular and/or nonglandular hairs; bases
indurate, apices herbaceous, margins scarious. Receptacles alveolate, convex,
naked or scales to 0.3 mm long. Ray florets pistillate, fertile (sterile or absent in
D. canescens var. shastensis): corollas white, blue, violet, or purple. Disc florets
bisexual, fertile, corollas yellow, + funnelform, tubes and limbs glabrous or
glabrate, lobes 0.2-0.7 mm long, glabrous to minutely pubescent. Cypselae of
ray and disc florets somewhat dimorphic, linear to obovate, glabrous to moder-
ately pubescent; thin walled, smooth or with 4-6 ribs per face, 3-6 mm long,
those of ray florets generally flattened, rounded abaxially, of disc florets flat-
tened laterally; pappus white to brown or reddish brown, of ray florets 2/3 to
nearly equalling length of disc florets; bristles somewhat barbellate, in 1-3 se-
ries, at most somewhat dorsi-ventrally flattened and dissimilar in width near
base, bases not to slightly overlapping, mostly 4-8 mm long. n = 4.
Although similar to Machaeranthera in several ways (see above), the mem-
bers of Dieteria may be distinguished from it by foliage and fruit characters.
Dieteria has entire to serrate or dentate leaves with thin-walled fruits that are
mostly longer than 3 mm, while Machacranthera has leaves that are deeply
lobed or dissected and thick-walled fruits that are mostly 3 mm or less long.
There isa great deal of morphological variation in Dieteria, which has resulted
ina large number of species names. The treatment adopted here follows that of
Turner (1987) in recognizing three species containing 16 varieties, with 3 vari-
eties in D. asteroides, 3 in D. bigelovii, and 10 in D. canescens. Additional syn-
onymy for Dieteria is included in Turner (1987).
—
KEY TO THE SPECIES OF DIETERIA
Phyllaries and peduncles both with prominent glandular pubescence.
2. Leaves covered with stiff glandular hairs 1. Dieteria asteroides (var. glandulosa)
2. Leaves glabrous to variously pubescent, but not covered with stiff glandular hairs.
3. Midstem leaves linear-lanceolate to linear or linear-oblanceolate (ovate to
obovate in var. /eucanthemifolia), 1.5-5 mm wide; herbaceous eee apices
acute to acuminate (1-3 mm long) cs tra canescens
3. Differing from the above in at least one respect; midstem leaves lanceolate
oblanceolate and 5—15 mm wide, or herbaceous phyllary apices a acum
nate (2-6 mm long), or both ieteria bigelovii
. Phyllaries and peduncles variously pubescent with glandular = nace ular
hairs, but rarely with prominent glandular pubescence on bo
4, Phyllaries usually pubescent throughout, with pu ee on ott herbaceous
apices and indurate bases, apices acute to long-acuminate, 1-6 mm long; mid
ele leaves 6-20 mm wide (2-6 mm in var. i 1. Dieteria asteroides
4. Phyllaries only occasionally pubescent throughout, with pubescence mostly con
MORGAN AND HARTMAN, SYNOPSIS OF MACHAERANTHERA 1393
fined to herbaceous mee apices acute to acuminate, 1-3 mm long; mid-stem
leaves 1.5-6(-8) mm wide 3. Dieteria canescens
1. Dieteria asteroides Torr, in Emory Report 142.1848. Machaeranthera asteroides
(Torr) Greene, Pittonia 3:63. 1896. TYPE: U.S.A. NEW MEXICO: “elevated land between
the Del Norte and the waters of the Gila,” 16 Oct 1847, Emory s.n. (HOLOTYPE: NY).
Machaeranthera canescens var. latifolia A. Gray, Pl. Wright. 2:75. 1853. Aster canescens var. latifolia
(A. Gray) A. Gray, Syn. Fl. N. Amer. 1(2):206. 1884. TYPE: U.S.A. New Mexico: “Near the copper
mines,” Sep-Oct 1851, C. Wright 1152 (LECTOTYPE: GH [designated by Turner 1987
ISOLECTOTYPES: GH, MO, NY, PH, UC, US).
Aster canescens Pursh var. tephrodes A. Gray, Proc. Amer. Acad. Arts 16:99. 1880. Mach }
des (A. Gray) Greene, Pittonia 4:24. 1899. Aster tephrodes (A. Gray) S.F Blake, Contr. U.S.
Natl. Herb. 25:563. 1925. Typification unclear: In Bot. Calif. 1:322.1876, Gray mistakenly based
the name Aster incanus toad A. oe on Diplopapyis incanus pind for plants occurring
in southern California. H ster
canescens var. tephrodes for ieee plants, noting that they also ecemed in Arizona and New
Mexico. Although sens that A. c. var. tephrodes “..was named A. incanus in the Botany of
California, but...cannot be Lindley’s plant,” Gray apparently did not select a type specimen
for Aster canescens var. tephrodes. The only material mentioned by Gray in connection with
any of these names was collected by “Capt. E.k. Sm n “Guadalupe Canyon, Sonora,” but
we have not been able to locate the specimens. nae isey) placed Aster incanus, Aster
canescens var. tephrodes, Machaeranthera tephrodes, and Aster tephrodes in synonymy un-
der Machaeranthera canescens var. incana.
KEY TO THE VARIETIES OF .DIETERIA ASTEROIDES
1. Stems glabrous to glandular-pubescent; leaves covered with stiff glandular hairs
1 asteroides var. glandulosa
. Stems pubescent, puberulent, or canescent, glandular pubeccance sparse or lack-
ing; leaves puberulent or canescent, stiff glandular hairs sparse or lacking.
2. Involucres hemispheric; herbaceous phyllary apices long-acuminate, 3-6 mm
long; mid-stem leaves 6-20 mm wide, usually clearly serrate or serrulate
la teroides var. asteroides
2. Involucres broadly turbinate (rarely hemispheric); herbaceous phyllary apices
acute to acuminate, 1-3 mm long; mid-stem leaves 2-6 mm wide, entire to slightly
serrulate 1c. D. asteroides var. lagunensis
la. Dieteria asteroides var. asteroides
1b. Dieteria asteroides var. glandulosa (B.L. Turner) D.R. Morgan & R_L. Hart-
man, comb. nov. BASIONYM: Machaeranthera asteroides a senor var.
glandulosa B.L. Turner, ee 60:77. 1986. TYPE: U.S.A. ARIZONA. Marico :
U.S. highway 60, 2.6 mi E of Queen Creek tunnel, 4200 ft, 9 Sep 1975, de
Keil & Lehto L18904 Loon PE: LL! ISOTYPES: ASU, CSU, NY).
1c. Dieteria asteroides var. lagunensis (D.D. Keck) DR. Sooae & R. L, Hartman,
comb. nov. BASIONYM: Machaeranthera lagunensis D.D. Kec 8.1957.
Machaeranthera asteroides (Torr) Greene var. lagunensis aa Keck) B.L. Turner,
Phytologia 60:77. 1986. TyPE: U.S.A. CALIFORNIA. San Diego Co. 2 mi S of the main
recreation area, Laguna Mountains, 5200 ft, 20 Aug 1952, PA. Munz & EK. Balls
17948 (HOLOTYPE: NY; ISOTYPE: RSA).
1394 BRIT.ORG/SIDA 20(4)
2. Dieteria — (A. Gray) D.R. Morgan & R.L. Hartman, comb. nov. BASIONYM:
Aster bigelovii A. Gray, Pacific Railway Survey 4:97. 1857. Machaeranthera Ee
(A. Gray) Greene, Pittonia 3:63. 1896. TYPE: U.S.A. NEW MEXIco. Bernalillo C ’
royos in the Sandia Mountains,” LO Oct 1853,J.M. Bigelow s.n. (HOLOTYPE: GH; ISOTYPE:
NY)
KEY TO THE VARIETIES OF DIETERIA BIGELOVIL
1. Involucres hemispheric, 2-3 times broad as tall; phyllaries 90-100, mostly 0.5-1 mm
wide at midpoint, herbaceous apices long-acuminate, 2-5 mm long _____ 2. D. bigelovii var.
mucronata
1. Involucres broadly turbinate to hemispheric, 1-2(—3) times broad as tall, phyllaries
25-100, mostly 1-2 mm wide at midpoint, herbaceous apices acute to long-acumi-
nate, 2-6 mm long
2. Phyllaries 25-50, apices acute to acuminate, 2-4 mm long; ray florets 12-30
2b. D. bigelovii var.commixta
2. Phyllaries 50-100, apices long-acuminate, 3-6 mm long; ray florets 30-60
2a. D. bigelovii var. bigelovii
2a. Dieteria bigelovii var. bigelovii
2b. Dieteria bigelovii var. commixta (Greene) D.R. Morgan @ R.L. Hartman, comb
nov. BASIONYM: Machaeranthera commixta Greene, Pittonia 4:71. 1899.
Machaeranthera canescens (Pursh) A. Gray var. commixta (Greene) S.L. Welsh,
Great Basin Naturalist +3:316. 1983. Machaeranthera bigelovii (A. Gray) Greene
var. commixta (Greene) B.L. Turner, Phytologia 60:77. 1986. TyPE: U.S.A. UTAH.
Garfield Co.: Henry Mountains, Bromide Pass, 10,000 ft, 27 Jul 1894, M.E. Jones 5695y
(HOLOTYPE: US; ISOTYPE: POM).
a
2c. Dieteria bigelovii var. mucronata (Greene) D.R. Morgan & R.L. Hartman,
comb. nov. BASIONYM: Machaeranthera mucronata Greene, Pittonia 4:72. 1899.
Aster adenolepis S.F. Blake, J. Wash. Acad. Sci. 30:471. 1940, nom. nov, non Aster
mucronatus Sheldon, 1903. Machaeranthera bigelovii (A. Gray) Greene var.
mucronata (Greene) B.L. Turner, Phytologia 62:238. 1987. Type: U.S.A. Arizona.
Coconino Co.: Thompson Canyon, 8500 ft, 19 Sep 1894, M.E. Jones 6065bl (LECTO-
TyPE: US [designated by Turner 1987]).
—
3. Dieteria canescens (Pursh) Nutt., Trans. Amer. Philos. Soc., ser. 2, 7:300. 1840.
BASIONYM: Aster canescens Pursh, Fl. Amer. Sept. 2:547, 1813. Machaeranthera
canescens (Pursh) A. Gray, Pl. Wright. 1:89. 1852. Type: U.S.A. NORTH DAKOTA: “on
the denuded banks of the Missouri” (vicinity of Fort Mandon), 1811, Nuttall s.n.
(HOLOTYPE: BM; possible ISOTYPE: NDG).
Dieteria viscosa Nutt., Trans. Amer. Philos. Soc., ser, 2, 7:301. 1840. Aster canescens Pursh var. viscosus
(Nutt) A. Gray, Proc. Amer. Acad. Arts 16:99 1880. Machaeranthera viscosa (Nutt.) Greene,
Pittonia 4:22. 1899. Machaeranthera canescens (Pursh) A. Gray var. viscosa (Nutt.) Piper, Contr.
U.S. Natl. Herb. 11:576. 1905. Type: U.S.A. NEBRASKA. Scotts Bluff Co. “near Scott’s Bluff, on the
Platte,” 1834, Nuttall s.n. HOLOTYPE: BM).
Dieteria divaricata Nutt., Trans. Amer. Philos. Soc. ser. 2, 7:301. 1840. Machaeranthera divaricata
Nutt.) Greene, Pittonia 4:23. 1899. Type: U.S.A. “denudated plains of the Rocky Mountains
and Oregon, common,” 1834, Nuttall s.n. (HOLOTYPE: GH),
Dieteria pulverulenta Nutt., Trans. Amer. Philos. Soc, ser. 2, 7:300. 1840. Machaeranthera
MORGAN AND HARTMAN, SYNOPSIS OF MACHAERANTHERA 1395
tt.) Greene, Pittonia 4:23. 1899. Type: U.S.A. “arid plains towards the sources
of oe Platte? 1834, Nuttall s.n. (HOLOTYPE: PH).
,Pittonia 3:61. 1896. Aster leiodes Blake, Contr. U.S. Natl. Herb.
25:563. 1925, nom. nov, non Aster laetevirens Greene (1900). TyPE: U.S.A. NEVADA. Elko Co:
Sey uaa: ae 1894, E.L. Greene s.n. (HOLOTYPE: ae ISOTYPE: ees
{ ia A. Nelson, Proc. Biol. Soc. Wash. 20:3 :
(Pursh) A. hee var. latifolia (A. Nelson) S.L. Welsh, oo. eae Mem. 9:213. 1987.
se Se aie canescens (Pursh) A. Gray var. monticola Dorn, Vascular Pl. Wyo. 295. 1988.
Salt Lake Co. Big Cottonwood Canyon, 8950 ft, 9 Aug 1933, A. O. Garrett
me j rh ISOTYPES: GH, LL!, US)
KEY TO THE VARIETIES OF DIETERIA CANESCENS
1. Phyllaries spreading to reflexed, rarely appressed.
2. Involucres mostly (9-)10-16 mm tall.
3, Suffruticose perennials; involucres (10-
ls
12-16 mm tall; phyllary oa densely
glandular 3f.D escens var.ziegleri
3. Biennials or short-lived perennials;involucres (9-)10-15 clea:
ces densely appressed-pubescent, glands if present obscured by appressed
n j.D. canescens var. nebraskana
2. Involucres mostly 6-10(-12) mm tall.
4. Stems canescent; gianeuler aur naalie sparse or lacking.
Plants usually y erect stem; branches stiff and straight,
widely divaricate 3b. D. canescens var.incana
5. Plants with a single stem or with multiple ascending stems; branches
flexible and a loosely spreading or (usually) ascending but not
widely diva
6. Ray one ae or absent; phyllaries 3-5(—7) seriate 3e.D. canescens
var. shastensis
6. Ray florets pistillate and fertile; phyllaries 5-10 seriate 3a. D. canescens
var. canescens
4. Stems evidently glandular, frequently also puberulent or canescent, occa-
sionally glabrous.
7. Stems canescent, glands usually sessile or subsessile; most heads sessile
or with peduncles shorter than involucre height 3d.D. canescens
var. sessiliflora
7. Stems with prominent stipitate glands, als y puberulent or
canescent; most peduncles longer than hier neigh
8 Midstem and upper leaves ovate, oblong or obovate, 1.5-4 times
ee than wide 3c. D. canescens er aaa
8. Hea and upper leaves linear-lanceolate to linear, more than
es longer than wide 3h. D. canescens var. aristata
. Phyllaries pres or occasionally spreading, rarely aa
9. Ray florets sterile or absent e. D, canescens var. shastensis
9. Ray florets pistillate and fertile.
10. Phyllaries densely appressed-pubescent, etal pubescence lacking or
g. D.
sparse and obscured by appressed pubes canescens var.ambigua
10. Phyllaries glabrous to glandular- Te ay seed pubescent or
canescent.
1. Stems with | inent stipitate glands, occasionally als ent, rarely
glabrous 3h D. canescens var. aristata
1396 BRIT.ORG/SIDA 20(4)
11. Stems glabrous to puberulent or canescent, stipitate oie sparse or
lacking 3i D. canescens var. glabra
(i) Dieteria canescens subsp. canescens
3a. Dieteria canescens val. canescens
3b. Dieteria canescens var. incana (Lind|.) D.R. Morgan & R.L. Hartman, comb.
nov. BASIONYM: Diplopappus incanus LindL, Bot. Reg. 20:t. ae ee Dieteria
incana (Lindl) Torr. @ A. Gray, Fl.N. Amer. 2:100. 1841, Machaeranther
(Pursh) A. Gray var. incana (Lind) A. Gray, Bot. Wilkes Exp. Phan. 1702) 340. 1874.
Machaeranthera incana (Lindl.) Greene, Pittonia 3:62. 1896. TYPE: U.S.A. OREGON:
“Columbia River,” described from seeds grown in London, 1830, Douglas s.n. (HO-
LOTYPE: BM; ISOTYPE: GH; ISOTYPE, microfiche: G-DC).
3c. Dieteria canescens var. leucanthemifolia (Greene) D.R. Morgan & R.L. Hart-
man, comb. nov. BASIONYM: Aster leucanthemifolius Greene, Erythea 3:119, 1895,
Machaeranthera leucanthemifolia (Greene) Greene, Pittonia 3:61. 1896.
Machaeranthera canescens (Pursh) A. Gray var. leucanthemifolia (Greene) SL.
Welsh, Great Basin Naturalist 43:316. 1983. Type: U.S.A. NEVADA. Esmeralda Co:
near Candelaria, 6000 ft, Jun 1886, WH. Shockley 268 (HOLOTYPE: CAS; ISOTYPES:
DS, US)
3d. Dieteria canescens var. sessiliflora (Nutt.) D.R. Morgan & R.L. Hartman,
comb. nov. BASIONYM: Dieteria sessiliflora Nutt., Trans. Amer. Philos. Soc.,, ser. 2,
7:301. 1840. Machaeranthera sessiliflora (Nutt.) Greene, Pittonia 3:60. 1896.
Machaeranthera canescens (Pursh) A. Gray var. sessiliflora (Nutt.) B.L. Turner,
Phytologia 60:78. 1986. TYPE: U.S.A. IDAHO(?): “Denudated plains of the Rocky Moun-
tains and Oregon,” 1836, Nuttall s.n. (HOLOTYPE: B
3e. Dieteria canescens var. shastensis (A. Gray) D.R. en @RLL. popes
comb. nov. BASIONYM: Machaeranthera shastensis A. Gray, Proc. Amer. Acad. Art
6:539. 1866. Aster shastensis (A. Gray) A. Gray, Bot. Calif. 1:322. 1876,
Machaeranthera canescens (Pursh) A. var. shastensis (A. Gray) B.L. Turner,
Phytologia 60:79. 1986. TyPE: U.S.A. CALIFORNIA. Siskiyou Co.; Mt. Shasta, 9000 ft,
1860-62, WH. Brewer 1385 (HOLOTYPE: GH; ISOTYPE: US).
3f. Dieteria canescens var. ziegleri (Munz) D.R. Morgan & R.L. Hartman, comb.
nov. BASIONYM: Machaeranthera canescens (Pursh) A. Gray subsp. ziegleri Munz,
Aliso 7:65. 1969. Machaeranthera canescens (Pursh) A. Gray var. ziegleri (Munz)
B.L. Turner, Phytologia 60:79. 1986. TyPE: U.S.A. CALIFORNIA. Riverside Co.: N side
of Santa Rosa Mountain, 6500-7500 ft, 30 Sep 1968, L.B. Ziegler s.n. (HOLOTYPE: RSA;
ISOTYPE: CAS).
(ii) Dieteria canescens subsp. glabra (A. Gray) D.R. Morgan & R.L. Hartman,
comb. nov. Machaeranthera canescens (Pursh) A. Gray var. glabra A. Gray, PL.
Wright. 1:89. 1852. Machaeranthera canescens (Pursh) A. Gray subsp. glabra (A.
Gray) B.L. Turner, Phytologia 62:239. 1987
3g. Dieteria canescens var. ambigua (B.L. Turner) D.R. Morgan & R_L. Hartman,
comb. nov. BASIONYM: Machaeranthera canescens (Pursh) A. Gray var. ambigua
B.L. Turner, Phytologia 60:77. 1986. TPE: U.S.A. ARIZONA. Coconino Co: Flagstaff,
MORGAN AND HARTMAN, SYNOPSIS OF MACHAERANTHERA 1397
28 Aug 1922, H. Hanson A/ (HOLOTYPE: TEX: ISOTYPES: ARIZ, F MO, NEB, NY, OSU,
PHIL, RMI, TEX).
3h. Dieteria canescens var. aristata (Eastwood) D.R. Morgan & R.L. Hartman,
comb. nov. BASIONYM: Aster canescens s Pursh var. aristatus Eastwood, Proc. Calif.
Acad. Sci., ser. 2,6:296. 1896. Macl canescens (Pursh) A. Gray var. dristata
(Eastwood) B.L. Turner, Phytologia 60:78. 1986. Type: U.S.A. UTAH. San Juan Co.:
Willow Creek, 14 Jul 1895, A. Eastwood 45 (HOLOTYPE: CAS).
Machaeranthera rigida Greene, Pittonia 4:25. 1899. Type: U.S.A. ARIZONA. Navajo Co.: “Kearn’s
Canyon,” 20 Aug 1897, Zuck 41 (HOLOTYPE: NDG).
3i. Dieteria canescens var. glabra (A. Gray) D.R. Morgan @ R.L. Hartman, comb.
nov. BASIONYM: Machaeranthera canescens (Pursh) A. Gray var. glabra A. Gray, PL.
Wright. 1:89. 1852. Aster canescens Pursh var. viridis A. Gray, Proc. Amer. Acad. Arts
16:99. 1880. TyPE: U.S.A. NEW MEXICO. Dona Ana Co.: Rio Grande Valley at Presidio
San Elizario on sand-bars, 22 Sep 1849, C. Wright 262 tS no. 1258) (LECTOTYPE:
GH [designated by Turner 1987]; ISOLECTOTYPES: GH, M )
Machaeranthera linearis Greene, Bull. Torrey Bot. Club 24:51. 1897, non Machaeranthera linearis
Rydb. 1900. Aster linearis (Greene) Cory, Rhodora 38:407. 1936. TYPE: U.S.A. NEW MEXICO.
Dona Ana Co: Mesilla Valley, 3900 ft, 6 Sep 1897, E.O. Wooton 444 (LECTOTYPE: NDG [desig-
nated by Turner 1987} ISOLECTOTYPES: DS, GH, MO, NY, RM!, UC, US).
3j. Dieteria canescens var. nebraskana (B.L. Turner) D.R. Morgan & R_L. Hart-
man, comb. nov. BASIONYM: Machaeranthera canescens (Pursh) A. Gray var.
nebraskana BL. Turner, Phytologia 60:78. 1986. TyPE: U.S.A. NEBRASKA. Sheridan
Co.: 2 mi E Ellsworth, sandhill prairie on dry, loose sand, 27 Aug 1968, S. Stephens
28307 (HOLOTYPE: NY; ISOTYPES: ARIZ, DS, GH).
Xanthisma DC., Prodr. 5:94. 1836. Type: Xanthisma texanum DC.
Eriocarpum Nutt., Trans. Amer. Philos. Soc., ser. 2, 7:320. 1840. TyPE: Eriocarpum grindelioides
Nutt.
Sideranthus Nutt.ex C. Neesin M. Wied-Neuwied, Reise Nord-America 2:440. 1841. Type: Amellus
spinulosus Pursh.
Centauridium Torr. & A. Gray, Fl.N. Amer. 2:246, 1842. Type: Centauridium drummondii Torr. &
A. Gray.
Dieteria subgenus Sideranthus (Nutt. ex Nees) Nutt, Trans. Amer. Philos. Soc., ser. 2, 7:301L 1840.
Machaeranthera subgenus Sideranthus (Nutt. ex Nees) R.L. Hartman, Phytologia 68:456. 1990.
There has been disagreement on the dates of publication for Eriocarpum and
Sideranthus. For example, Farr, Leussink, and Stafleu (1979) give them as Oct-
Dec 1840 and 1840, respectively. According to Greene (1895), Nuttall’s treatment
(p.320) of Eriocarpum appeared in 1840 (for pp. 283-356). Subsequently,
Sideranthus was published by C. Nees in 1841 (Stafleu @ Cowan 1981, p. 707).
Herbs taprooted annuals, biennials, or perennials, if perennial usually with
a much-branched, woody caudex or rhizomatous (forming loose colonies in X.
rhizomatum), 3-100 cm tall. Stems erect or spreading or sprawling, often much
branched, glabrous or moderately to densely pubescent with glandular or
nonglandular hairs. Leaves alternate; entire to serrate, dentate, pinnatifid, or
1398 BRIT.ORG/SIDA 20(4
bipinnatifid, moderately to densely pubescent with glandular or nonglandular
hairs, apices, including lobes and teeth, apiculate to bristle tipped, bristles if
present I-4 mm long. Heads radiate or eradiate. Involucres turbinate, campanu-
late, or hemispheric. Phyllaries in 2-8 imbricate series, graduated in length,
appressed, spreading, or reflexed, glabrous or moderately to densely pubescent
with glandular or nonglandular hairs; bases indurate, apices herbaceous or
achlorophyllous, stiff, leathery, or margins scarious. Receptacles alveolate, flat
to convex, scales laciniate, 0.3-3 mm long, often forming a reticulum. Ray flo-
rets pistillate, fertile (absent in X. grindelioides, X. johnstonii, X. restiforme); co-
rollas white, pink, red-purple, purple, or yellow. Dise florets bisexual, fertile,
corollas yellow, + funnelform, tubes and limbs glabrous to somewhat pubes-
cent, lobes 0.4-0.9 mm long, glabrous to pubescent. Cypselae of ray and disc
florets distinctly dimorphic, elliptic to obovate, oblong, or obscurely cordate,
sparsely to densely pubescent; thin to thick walled, with 3-9 nerves or ribs per
face, (1-)1.5-3.5 mm long, those of ray florets, if present, + 3 sided, rounded
abaxially, of disc florets compressed laterally; pappus usually whitish to brown
or reddish brown, of ray florets 1/2 to nearly equaling length of disc florets:
bristles coarsely barbellate, in 2-4 series, dorsi-ventrally flattened and dissimi-
lar in width near base, bases overlapping, mostly 3.5-8 mm long. n = 2, 3, 4, 8.
As defined here, the genus Xanthisma consists of the former
Machaeranthera sections Blepharodon, Sideranthus, Havardii, and Stenoloba,
plus the unispecific genus Xanthisma, which because of the rule of priority
becomes the name of the genus. The grouping of Xanthismaand these four sec-
tions of Machaeranthera (sensu Morgan 1993) was well supported by nrDNA
evidence (Morgan 2003), and, except for Arida blepharophylla (see discussion
below), cpDNA evidence did not support any conflicting relationships (Fig. 1).
Although Xanthisma includes species with both cyanic (blue, purple, pink, or
white) and yellow rays, several other morphological characteristics are shared
by its members, including short, turbinate, thick-walled fruits that are densely
pubescent, receptacular scales, leaves with marginal spines, and chromosome
numbers based on x = 4. Many of these species have in fact been grouped to-
gether at the generic or sectional level by authors such as Greene (1894) and
Hall (1928).
The following key is based largely on data from Hartman (1976, 1990) and
Semple (1974).
KEY TO THE SPECIES OF XANTHISMA
ner phyllaries, at least, with a basal “stalk,” abruptly enlarged into an ovate to or-
ie or elliptic blade, mostly 2-5 mm wide and glabrous; apices ee acum
nate to obtuse, or cea rounded, not bristle-tipped — niin texanum
1. Inner phyllaries, as with a
not markedly | Jed abo a ert oblonc
or lanceolate, mostly 1-2 mm wide; apices narrowly obtuse to long-attenuate, usu-
ally bristle-tipped.
MORGAN AND HARTMAN, SYNOPSIS OF MACHAERANTHERA 1399
2. Ray florets absent.
2 Pillay coulne leaves aos | ace cm ong if shorter or absent then plants
shion of basal leaves; stems
lacking axillary rosettes; NW New Mexico and N Arizona to S Cana
___5.Xanthisma
grindelioides
cm long, basal leaves not persisting on mature
plants; stems covered by axillary rosettes of tiny leaves; Coahuila and Nuevo
Leon, Mexico
3. Primary cauline leaves 0.2-1
4. Main stem leaves on new growth oblong to narrowly lanceolate; stems
canescent to slightly tomentose, the pubescence increasing the stem di-
ameter but slightly; plants much branched and bushy with flexible stems
8. a mevonaetenl
4. Main stem leaves deltoid to narrowly triangular; stems densely wool
tomentose, the pubescence increasing the stem diameter 2 to 3 tim
plants divergently but sparingly branched with rigid, ee like s
as restiforme
2. Ray florets present.
5. Ray floret corollas white, pink, purple, or laven
6. Plants dwarf, arising from a cluster of basal an stems leafy, up to 1 ae cm
tall, few to several, each 1 headed; montane to alpine, bias
ming pore coloradoense
6. Plants aes robust, basal leaves usually not aa on eae mature
plant;st mostly 15 to 40 cm tall, often branched, each
with 1 to 10 or more heads; cantial NM, W Texas; N half of Mexico.
7. Plants biennial or short-lived perennial herbs from a normal taproot;
ray flowers pink, purple, or lavender (often fading on old specimens);
peduncles moderately to densely glandular-stipitate, often with long
flexuous hairs intermixed; phyllaries usually REIDaCoUs in upper half,
long-attenuate, reflexed or widely sf ds, central Mexico
to S edge of the Chihuahuan Desert 6. ans gymnocephalum
7. Plants perennial from a woody, branched caudex, often with a thick-
ened taproot; ray flowers white (rarely purple in X. gypsophilum), a
with pink or purple streaks on the underside; peduncles villous, hispid,
or glandular-stipitate; phyllaries not herbaceous above but with a green
or purple strip or patch in the upper half, the tips acute to broadly
acuminate, appressed to squarrose; Chihuahuan Desert and vicinity and
mountains of W Texas and S New Mexico.
8, Peduncles and leaves moderately to densely vill | ded
ot alway slea
a
n the lower half to two-thirds of the stem, not areatly reduced in
size above (except for 1 or 2 reduced ai be stems u
Rese unbranched, bearing only on 4.X cn ae crutchfieldii
Peduncles and leave eae oe and/or ae to
ap eh aoe ab distributed along the stem usually for well
over two-thirds its length, oft Aste ase gradually reduced
in size al f| at least sparingly branched with
+
1 to 10 or more eae
9. Peduncles stipitat | ft | tl
5 to 14 pairs of Geih: SE New Mexico, W Texas; N ve
VVIUTT
su eae aeons
9, Peduncles hispid or hispidulose; leaves usually finely or obscurely
1400
BRIT.ORG/SIDA 20(4)
serrate or serrulate, usually with 12 to 25 pairs of teeth; south-
ral New Mexico to W Texas 2.Xanthisma blephariphyllum
c
5. Ray floret corollas yellow
10. Plants growing in loose colonies with stems arising from 3 to 10 or more
elongate (2-15 cm or more long), branched rhizomes, 1.5-6 mm in di-
ameter, arising from a woody rootstock; leaves without glands
0. Xanthisma rhizomatum
. Plants from a taproot or woody caudex, not rhizomatous; leaves often
glandular
=
—_—
. Cyp selae of disc florets oblong t Nall wly | II lI
. Cypselae of disc florets broadly obovate to obscurely and narrowly
cordate, the thick, Pony walle with 10-12 pF Fomine tas ribs, densely pu-
bescent f subulate, markedly dor-
see) flattened bristles de oe mm long; plants annual; leaves —
pinnatifid or bipinnatifid; SW Chihuahua, Mexico 5.Xanthisma
stenolobum
» thin,
flexible, smooth or with 6-16 obscure ribs, glabrous to moderately
pubescent, margins not thickened; pappus of filiform, only slightly
flattened bristles 3.5-7 mm long; plants annual to perennial; leaves
entire to bipinnatifid; occurring elsew here
12. Plants taprooted, annual; phyllaries ae tase to broadly acute, not
bristle tipped; leaves serrate or dentate, the teeth blunt or termi-
nating in a stiff callosity, not bristle tipped; cypselae oblong or
narrowly elliptic with 12-14 barely discernible nerves, sparsely
pubescent; lowlands of SE New Mexico and W Texas 17. Xanthisma
12. Plants perennial with much-branched caudices, if taprooted an-
nuals then ryaiies with ies acute to acuminate and
prominently bristle tipped apices; leaves various, if serrate or den-
tate, the teeth bristle tipped; ee elliptic to narrowly obo-
vate with 6-16 weak ribs, glabrous to moderately pubescent;
Canada to S U.S.; Mexico
13. Plants annual with herbaceous stems and bases; involucres
eglandular
14. Leaves with entire margins, not bristle lined, or lower
cauline and basal ones tridentate or sometimes pinnati-
i aes a by bristles 0.1-0.3 mm long; NE Baja
16. Xanthisma wigginsii
14. pene eri spinulose toothed or lobed ie usu-
ally 1.5-3 mm long; phyllaries (except innermost series)
tipped by Hees usually 0.6-1.5 mm long; a a
wher
thisma gracile
13. Plants perennial, stems with woody b
15. Bracts of peduncles apie and grading inte phy
ies; stems nearly leafless above base 13. Xanthisma junceum
15. Bracts of peduncles few and not grading i into si alae
stems leafy or leaves reduced near inflorescence.
16. Phyllaries strongly squarrose in upper ha leaves
thickish, usually oblanceolate to spatulate with
MORGAN AND HARTMAN, SYNOPSIS OF MACHAERANTHERA 1401
bluntly dentate to subentire margins; S Baja
California Sur, Mexico 11. Xanthisma arenarium
16. Phyllaries with appressed to spreading apices; le
thin, usually oblong to lanceolate with prominent
teeth or lobes; SW Canada, W U.S.A.;N half of Mexico
14. Xanthisma spinulosum
Xanthisma DC. section Xanthisma
Herbs taprooted annuals, rarely biennials, 20-90 cm tall, taproot sometimes
thickened. Leaves entire on upper stem, serrate below, pinnatifid to bipinnatifid
near base, teeth or lobes usually with apiculate callosities, not bristle tipped.
Heads radiate. Involucres turbinate to hemispheric or campanulate. Phyllaries
in 3-4 graduated series, the owe portion stalk-like, stiff, stramineous, enlarged
above into an ovate to orl ord d elliptic blade witha glabrous, green
pia apices acuminate or obtuse and cuspidate or broadly rounded, not bristle
tipped. Receptacles alveolate, scales 0.2-2 mm long, mostly distinct. Ray floret
corollas yellow. Cypselae distinctly dimorphic, 1.5-2.2 mm long, narrowly obo-
void to oblong, walls thick _ 8-16, often obscure ribs, sparsely pubescent;
ray cypselae 3 sided; disc cy} d laterally, generally 4 sided. Pap-
pus similar in ray and disc oes tawny: bristles 3-7 mm long, linear, dorsiven-
trally flattened basally, in 2 series. n = 4.
1. asians texanum DC., Prodr. 5:95. 1836. Tyre: U.S.A. TEXAS. Bexar Co.: woods
Medina, May 1828. J.L. Berlandier 2039 (LECTOTYPE: IDC microfiche 800.
es G-DC! [designated by Semple 1974], ISOLECTOTYPES: G-DC, GH, NY, US).
Xanthisma texanum DC. var. berlandieri A. Gray, Pl. Wright. 1:98. 1852. Xanthisma berlandieri
(A. Gray) J.K. Small, Fl. S.E. U.S. 1184. 1903. Type: U.S.A. SOUTHERN TEXAS: J.L. Berlandier s.n.
on 7 oe A. Trécul s.n.(SYNTYPES: GH).
KEY TO THE INFRASPECIFIC TAXA OF XANTHISMA TEXANUM
. Blades of phyllaries somewhat expanded above the proximal stalk, mostly orbicu-
lar to depressed elliptic: dimensions above the widest part 1-2 mm long, 1.5-3.5
m wide 1a. X. fae subsp. texanum
2. Phyllary apices rounded with little or no flare 1 mm from the apex 1b. X. texanum
subsp. texanum var. texanum
2. ee apices obtusely cuspitate, flared cca the ovate portion 1. a 2m
ong, 2-3 mm wide above the widest portio c.X.texanum subsp.
exanum var. orientale
. Blades of phyllaries abrupt! ded above proximal stalk, mostly ovate: dimen-
sions apove the widest pat 3. os 8 mm long, 2.5-5 mm wide 1d.X.texanum subsp
rummondii
la. Xanthisma texanum DC. subsp. texanum
lb. Xanthisma texanum DC. subsp. texanum var. texanum
1c. Xanthisma texanum DC. subsp. texanum vat. orientale Semple, Rhodora 76:14.
1402 BRIT.ORG/SIDA 20(4)
1974. Type: U.S.A. TEXAS. Hidalgo Co. 13.5 mi N of Edinburg on US. hwy 183, 8 Jul
1970, W. an 7642 (HOLOTYPE: MO).
Id. Xanthisma texanum DC. subsp. drummondii (Torr. & A. Gray) Semp
See 76:16. 1974. BASIONYM: Centauridium drummondii Torr. & A. G1 ray, : 1
N. Amer. 2:246, 1842. Xanthisma texanum DC. var. drummondii (Torr. & A. Gray)
A. Gray, PL Wright. £98. 1852. Xanthisma drummondii (Torr. & A. Gray) Hooker f.
Curtis’ Bot. Mag. 33:t.6275. 1877. Type: U.S.A. TEXAS: exact locality unknown, Nov
1835, T. Drummond 227 (LECTOTYPE: GH [designated by Semple 1974};
ISOLECTOTYPES: GHI, NY).
Xanthisma texanum DC. subsp. texanum var. orientale Semple forma rubrum Semple, Rhodora
76:15, 1974, TYPE: U.S.A. TEXAS. San Patricio Co.: 0.5 mi E of Ingleside on Texas hwy 361, 13 Jun
L971, .C. Seniple 602 (HOLOTYPE: MO).
This polymorphic species has always been maintained distinct from other gen-
era in the Tribe; in part due to the unique involucre. The relationship to mem-
bers of Machaeranthers sensu lato was never seriously considered until data
from DNA became available. According to Semple (1974) the following combi-
nation of features characterized the species: “heterocarpic fruit with a pappus
of bristly scales only, fruit pubescence of long white ascending hairs, recep-
tacle slightly convex with a persistent reticulate network of subulate scales.”
Interestingly, all of these characters are found in at least some of the taxa of this
newly expanded genus.
Semple (1974) recognized two forms of variety orientale, the typical one
having whitish lateral margins on the bracts and forma rubrum in which the
lateral margins were red. Forma rubrum is said to be rare but is found mixed
with the typical form throughout the range of var. orientale.
Xanthisma section Blepharodon (DC.) D.R. Morgan & R.L. Hartman, comb. nov,
BASIONYM: Haplopappus section Hees DC.,, Prodr. 5:346. 1836, “Aplopappus,”
non Blepharodon Decne. in DC., 1844 [Ascl epiadaceael.
Machaeranthera section Blepharodon (DC.) R.L. Hartman, Phytologia 68:443. 1990, TyPE:
Haplopappus gymnocephalus DC. “Aplopappus” (ectotype designated by Hall 1928, from
among the four species included)
Eriocarpum Nutt, Trans. Amer. Philos. Soc., ser. 2, 7:320. 1840. Haplopappus section Eriocarpaea
Benth. & J.D. Hook., Gen. Pl. 2:254. 1873, based on Eriocarpum Nutt., Trans. Amer. Philos. Soc.,
ser. 2, 7: nt 1840. TYPE: Erioc arpuim grindelioides Nutt.
eronquist & D.D. Keck, Brittonia 9:237, 1957, nom. illeg. ICBN:
24.3).
Herbs taprooted annuals, biennials, or perennials, 3-50 cm tall, often with a
much branched caudex or in loose colonies with stems arising from 3 to 10 or
more elongate, branched rhizomes, 1.5-6 mm in diameter, from a woody root-
stock. Leaves serrate or dentate, entire or rarely pinnatifid in a few species, teeth
or lobes usually bristle tipped. Heads radiate or eradiate. Involucres depressed
hemispheric to campanulate. Phyllaries in 3-6 graduated series, linear to lan-
ceolate, the lower portion thickened, stramineous, the upper fourth to half her-
MORGAN AND HARTMAN, SYNOPSIS OF MACHAERANTHERA 1403
baceous or with a green or purple tinged strip or patch, stipitate glandular to
canescent, appressed to reflexed; apices obtuse to long attenuate and usually
bristly tipped. Receptacles alveolate, scales often present, 0.3-1.5 mm long, dis-
tinct or forming a reticulum. Ray floret corollas white, pink, purple, or lavender
when present (yellow in Xanthisma rhizomatum). Cypselae distinctly dimor-
phic, (1-)L5-3.5 mm long, narrowly obovoid to oblong, the wee thick with 6-
18, often prominent ribs, often densely pubescent; ray cypsel ly 3 sided;
disc cypselae compressed laterally. Pappus white to tawny: bristles 3.5-8 mm
long, dorsiventrally flattened basally, in 2-4 graduated series, those of the ray
florets often much reduced in length. n = 4, 8.
—
2. Xanthisma blephariphyllum (A. Gray) D.R. Morgan & R.L. Hartman, comb.
nov. BASIONYM: Haplopappus blephariphyllus A. Gray, Pl. Wright. 1:97. 1852,
“Aplopappus.” Eriocarpum blephariphyllum (A. Gray) Greene, Erythea 3:14. 1895.
Machaeranthera blephariphylla (A. Gray) Shinners, Field & Lab. 18:38. 1950. TYPE:
U.S.A. Probably TEXAS: plains at the eastern base of the Guadalupe Mountains, 18
Oct 1849, C. Wright 288 (HOLOTYPE: GH}; ISOTYPE: NY}).
Eriocarpum serratum Greene, Bull. Torrey Bot. Club 25:119, pl. 331, fig. 3, 4. 1898. Sideranthus
serratus (Greene) Standley, Contr. U.S. Natl. Herb. 13:222.1910. Type: U.S.A. NEW MEXICO. Lin-
coln Co.:: White Mountains, S fork of Tularosa Creek E of the Mescalero Agency, 6800 ft, 30 Jul
1897, E.O. Wooton 251 (HOLOTYPE: NDGI ISOTYPES: B, K, MO!, NY!, RM!, UC).
Machaeranthera correllii Shinners, Field & Lab. 17:55. 1949. TyPE: U.S.A. TEXAS. Sueiia Co.:
Guadalupe Mountains, Pine Springs Canyon, 5700 ft, 15 Aug 1946, L. Shinners 9059
TYPE: SMUD).
3. Xanthisma coloradoense (A. Gray) D.R. ak a & R.L. Hartman, comb. nov.
NyM: Aster coloradoensis A. Gray, Proc. Amer. Acad. Arts 11:76. 1876.
Eriocarpum coloradoense (A. Gray) Greene, Erythea 2:110. 1894, “coloradense.”
Xylorhiza coloradoensis (A. Gray) Rydb., Bull. Torrey Bot. Club 32:124. 1905,
“Xylorrhiza coloradensis.” Machaeranthera coloradoensis (A. Gray) Osterhout,
Torreya 27:64. 1927, “coloradensis,” Haplopappus coloradoensis (A. Gray) R.L. Hart-
man ex R. D. Dorn, Vasc. Pl. Wyoming 295. 1988. Type: U.S.A. COLORADO: South
Park near Hamilton, 4 Aug 1875, E.L. Greene 679 (LECTOTYPE, here designated from
among syntypes: GH).
Xylorhiza brandegei Rydb., Bull. Torrey Bot. Club 32:124. 1905, “Xylorrhiza.” Machaeranthera
coloradoensis (A. Gray) Osterhout var. brandegei (Rydb.) TJ. Watson ex R.L. Hartman
Phytologia 68:444. 1990. TYPE: U.S.A. COLORADO: San Juan Pass, 1875, I.S. Br a 1165 (Ho-
LOTYPE: GHh; IsoTYPES: NY!, UC).
—
4. Xanthisma crutchfieldii (B.L. Turner) D.R. Morgan & R.L. Hartman, comb.
nov. ee M: Machaeranthera crutchfieldii B.L. Turner, Phytologia 26:116. 1973.
TYPE: MEXICO. NUEVO LEON: bare gypsum rock outcrop on south slopes, 17 mi E
of San Sion ue then 2 miS, 24 Oct 1970, B.L. Turner & J. Crutchfield 6324
(HOLOTYPE: TEX! ISOTYPE: MEXU)).
5. Xanthisma grindelioides (Nutt.) D.R. Morgan @ R.L. Hartman, comb. nov.
BASIONYM: Eriocarpum grindelioides Nutt., Trans. Amer. Philos. Soc., ser. 2, 7:321.
1840. Haplopappus nuttallii Torr. & A. Gray, Fl. N. Amer. 2:242. 1842, “Aplopappus,
~—
1404 BRIT.ORG/SIDA 20(4)
nom. nov. non Haplopappus grindelioides DC, Prodr. 5:348. 1836, “Aplopappus.”
Aster nuttallii Torr @ A. Gray) O. Kuntze, Revis. Gen. Pl. 1:318. 1891, non Torr. & A.
Gray, 1842. Sderanteuependeie desu Britton in Rydb,, Bull. Torrey Bot. Club
27:620. 1900. Machaeranthera grindelioides (Nutt.) Shinners, Field & Lab. 18:40.
1950. TyPE. U.S.A. WYOMING): on shelving rocks in the Rocky Mountain range,
Oregon, s.d., 1. Nuttall s.n. (HOLOTYPE: PH! ISOTYPE: BM).
KEY TO THE VARIETIES OF XANTHISMA GRINDELIOIDES
. Plants tufted, mostly 10-35 cm tall; leaves evenly distributed along the length of
the stem, the margins often ciliate or spinulose-toothed but the bristles usually
less than 1.5 mm long; peduncles less than 2 cm long, often leafy to the base of the
capitulum; heads solitary or 2 to 5 per stem 5a. X. grindelioides var. grindelioides
1. Plants densely cespitose, mostly less than 8 cm tall; leaves mostly crowded at the
base of the plant, the margins lined with conspicuous, white bristles up to 4 mm
long; peduncles 1-4 cm long, naked or with an occasional greatly-reduced leaf;
heads mostly solitary 5b. X. grindelioides var. depressum
5a. Xanthisma grindelioides var. grindeloides
5b. Xanthisma grindelioides var. depressum (Maguire) D.R. Morgan & R_L. Hart-
man, comb. nov. BASIONYM: Haplopappus nuttallii Torr. & A. Gray var. depressus
Maguire, Amer. Midl. Naturalist 37:144. 1947. Machaeranthera grindeloides (Nut)
Shinners var. depressa (Maguire) Cronquist & D.D. Keck, Brittonia 9:237. 1957. TYPE:
U.S.A. UTAH. Millard Co.: Warm Point, in shallow gravelly and sandy soil, 55 mi SW
of Desert Range Experiment Station Headquarters, 18 Jun 1941, B. Maguire 20859
(HOLOTYPE: NYb ISOTYPES: GH!, UC).
6. Xanthisma gymnocephalum (DC.) D.R. Morgan & R.L. Hartman, comb. nov,
BASIONYM: Haplopappus gymnocephalus DC, Prodr. 5:346. 1836, “Aplopappus.” As-
ter gymnocephalus (DC.) A. Gray, Proc. Amer. Acad. Arts 15:32. 1879, non (F
Ruprecht) B. Fedtschenko, 1915. Eriocarpum gymnocephalum (DC.) use
Erythea 2:109. 1894. Machaeranthera sym nocephala (DC.) Shinners, Field & Lab.
18:40. 1950. TyPE: MEXICO: “Environs de Mexico,” 20 Jul 1827, J.L. Ber ens 588
(LECTOTYPE, here designated: IDC eae 800. 867.116, G-DC!).
Machaeranthera setigera Nees, Linnaea 19:722. 1847. TyPF: “In terris Mexicanis,” De Berghes s.n.
(location of type unknown).
ee p a sealers DC. subsp. primitivus H.M. Hall, Publ. Carnegie Inst. Wash. 389:56,
6. 1928. Type: MEXICO. MEXICO. Calcareous bluffs, Flor de Maria, 31 Jul 1890, C.G. Pringle
Aon HOI ae CI ISOTYPES: GH, NY!, PH!, SMU! UC).
7. Xanthisma gypsophilum (B.L. Turner) D.R. Morgan & R.L. Hartman, comb.
Vv. BASIONYM: Machaeranthera gypsophila B.L. Turner, Phytologia 26:117. 1973.
Type: MEXICO. COAHUILA: 14 mi S of Cuatro Ciénegas, then W 18 mi to actively
blowing dunes of gypsum, I Apr 1970, B.L. Turner 6052 (HOLOTYPE: TEX! ISOTYPES:
MEXUI, MICH).
8. Xanthisma johnstonii (S.F. Blake) D.R. Morgan & R.L. Hartman, comb. nov,
BASIONYM: Haplopappus johnstonii S.F Blake, Proc. Biol. Soc. Wash. 54:18. 1941,
eg ae Machaeranthera johnstonit (S.F Blake) B.L. Turner, Amer. J. Bot.
60:8 3. TYPE: MEXICO. COAHUILA: in dry heavy alkaline soil, 1 mi S of
cna 22-24 Aug 1938, IM. Johnston 7066 (HOLOTYPE: GH ISOTYPE: US|).
MORGAN AND HARTMAN, SYNOPSIS OF MACHAERANTHERA 1405
9, Xanthisma restiforme (B.L. Turner) D.R. Morgan & R.L. Hartman, comb. nov.
BASIONYM: Machaeranthera restiformis B.L. Turner, Amer. J. Bot. 60:836, figs. 1-5.
1973. TypE: MEXICO. Coahuila: in gypsum soil 2.4 mi SW of Cuatro Ciénegas, 11
Apr 1970, B.L. Turner 6063 (HOLOTYPE: TEX!; ISOTYPES: MEXU!, MICH)).
10. Xanthisma rhizomatum (M.C. Johnston) D.R. Morgan & R.L. Hartman, comb.
nov. BASIONYM: Haplopappus rhizomatus M.C. Johnston, Rhodora 63:177. 1961.
SL soaaale heterophylla R.L. Hartman, Phytologia 68:445. 1990, nom. nov.
n Machaeranthera rhizomata A. Nelson & J.-F Macbride, Bot. Gaz.
( wioraille 62:148. 1916. Type: MEXICO. NUEVO ne Saltillo-Matehuala
highway, 5 mi N of junction of the side road to Galeana, 6000 ft, 8 Oct 1959, J. Gra-
ham & M.C. Johnston 4203 (HOLOTYPE: TEX!; ISOTYPES: KANU!, MEXU)).
es section Sideranthus (Nutt. ex Nees) D.R. Morgan & R.L. Hartman,
mb. nov. BASIONYM: Sideranthus Nutt. ex C. Nees in M. Wied-Neuwied, Reise
cae ree 2:440. 1841. Dieteria section Sideranthus NEE ex Nees) Nutt. ex
Walpers Repert. Bot. Syst. 2:5 587 3M
ex Nees) R.L. Hartman, Phytologia 68:456. 1990. TYPE: Amellus ee Pursh.
>
Machaeranthera section Stenoloba R.L. Hartman, Phytologia 68:459. 1990. TyPE: Eriocarpum
ee
Herbst ted annuals, biennials, or perennials, 3-80 cm tall, sometimes with
amuch branched caudex. Leaves entire, serrate, dentate, or pinnatifid to deeply
bipinnatifid, teeth or lobes usually bristle tipped, often markedly so. Heads ra-
diate. Involucres broadly turbinate to depressed campanulate or hemispheric.
Phyllaries in 4-8 graduated series, linear to narrowly oblong, the lower portion
rigid, light green to stramineous, the upper fifth to half with a green patch or
strip (sometimes blackish near apex), usually pubescent, often with glandular
hairs, appressed to squarrose; apices obtuse to acuminate, usually tipped by a
stiff and often pronounced bristle. Receptacles usually alveolate, scales 0.4-3
mm long, distinct or forming a reticulum. Ray floret corollas yellow. Cypselae
often dimorphic, 1.5-3.2 mm long, elliptic to broadly obovate or obscurely and
narrowly cordate, walls thin with 6-16 nerves or prominent ribs, glabrous to
densely pubescent, ray cypselae usually rounded on dorsal edge, obscurely 3
sided, often asymmetrical, and slightly shorter; disc cypselae flattened later-
ally. Pappus white to tawny; bristles 3.5-6 mm long, slightly to moderately dor-
siventrally flattened basally, in 2-4 markedly graduated series, those of the ray
florets sometimes reduced in length. n = 2, 3, 4, 8.
11. Xanthisma arenarium (Benth.) DR. Morgan & R.L. Hartman, comb. no
ius Benth., Bot. Voy. Sulphur 24.1844, * ‘Aplopappus-
ner ee arenaria (Benth.) Shinners, Field & Lab. 18:40. 1950. TYPE:
MEXICO. BAJA CALIFORNIA SuR: Cabo San Lucas, 1841, R. B. Hinds s.n. (HOLOTYPE:
K, photo at DS!).
12. Xanthisma gracile (Nutt.) D.R. Morgan & R_L. Hartman, comb. nov. BASIONYM:
Dieteria gracilis Nutt. Proc. Acad. Nat. Sci. Philadelphia 4:22. 1848. Haplopappus
gracilis (Nutt.) A. Gray, Pl. Fendler. 76. 1849, “Aplopappus,” non TS. Brandegee, 1889.
1406
13.
14.
BRIT.ORG/SIDA 20(4)
Aster dieteria O. Kuntze, Revis. Gen. Pl. £315. 1891, based on Dieteria gracilis Nutt.
Eriocarpum gracile (Nutt.) Greene, Erythea 2:109. 1894. Sideranthus gracilis (Nutt.)
A. Nelson, Bot. Gaz. (Crawfordsville) 37:266. 1904. Machaeranthera gracilis (Nutt.)
Shinners, Field & Lab. 18:41. 1950. Type: U.S.A. NEW MEXICO: near Santa Fe, s.d., W,
Gam bel s.n. (HOLOTYPE: GHI; ISOTYPE: K).
—
Haplopappus ravenii R.C. Jackson, Amer. J. Bot. +9:123. 1962. Type: U.S.A. ARIZONA. Yavapai Co.
open oak grasslands near Johnson Wash, ca. 10 mi S of the Verde River, 8 Jun 1959, R.C. Jack-
son 2680 (HOLOTYPE: KANU)).
Xanthisma junceum (Greene) D.R. Morgan & R.L. Hartman, comb. nov.
BASIONYM: Haplopappus junceus Greene, Bull. Calif. Acad. Sci. 1:190. 1885,
“Aplopappus.” Eriocarpum junceum (Greene) Greene, Erythea 2:108. 1894.
Sideranthus junceus (Greene) A. Davidson & Moxley, FL. S. Calif. 377. 1923.
Machaeranthera juncea (Greene) Shinners, Field & Lab. 18:40. 1950. Type: U.S.A.
CALIFORNIA. San Diego Co.: Cleveland, s.d., M.K. Curran s.n. dectotype designated
from among two syntypes by Hall 1928 w ith no indication of location; it was not
found in the followi ing relevant herbaria: CAS, DS, NDG, UC, RSA).
Xanthisma spinulosum (Pursh) D.R. Morgan & R.L. Hartman, comb. nov.
BASIONYM: Amellus spinulosus Pursh, Fl. Amer. Sept. 2:564. 1813. alee
spinulosus (Pursh) DC, Prodr. 5:347. 1836, “Aplopappus,” non Phil, 1873, 1
Brandegee, 1889. Starkea pinnata Nutt., Gen. N. Am. 2:169. 1818, nom. ee:
Machaeranthera pinnata Shinners, Field & Lab 18:41. 1950, non Machaeranthera
spinulosa oe 1899. Dieteria spinulosa (Pursh) Nutt., Trans. Amer. Philos. Soc.,
ser. 2, 7301. 1841. Eriocarpum spinulosum (Pursh) Greene, Erythea 2:108. 1894.
Seen us spinulosus (Pursh) R. Sweet ex Rydb., Bull. Torrey Bot. Club 27:621.
1900. TyPE: U.S.A. [SOUTH DAKOTA. Lyman or Brule Co. Reveal et al. 1999], in open
prairies on the Missouri, 15 Sep 1804, M. Lewis s.n. (LECTOTYPE: PH! [designated by
Hall 1928} ISOLECTOTYPE: UC).
Diplopappus pinnatifidus Hook. f., Fl. Bor-Amer, 2:22. 1834. Aster pinnatifidus (Hook.f) O. Kuntze,
Revis. Gen. PI. 1:313. 1891. Machaeranthera pinnatifida (Hook. f.) Shinners, Sida 1:295. 1964,
nom. nov. non Machaeranthera spinulosa Greene, Pittonia 4:24. 1899. TypE: CANADA.
SASKATCHEWAN: Rocky Mountains near ot. or’s Lake, s.d., Drummond 5.n. (HOLOTYPE: K)
Hapopsppmspiadsi us DC. var. canescens A. Gray, P. Fendler 75. 1846 ase TYPE:
S.A. NEW MEXICO: between Santa Fe and the Rio del Norte. May 1847, A. Fendler 395 (HOLO-
TYPE: GH).
Eriocarpum wootonii Greene, Bull. Torrey Bot. Club 25:120, pl. 330, fig. 1, 2. 1898. Sideranthus
wootonti (Greene) Standley, Contr. U.S. Natl. Herb. 13:222. 1910. Type: U SAL NEW MEXICO: White
Mountains, 19 Aug 1897, £.0. Wooton 518 (HOLOTYPE: NDGE ISOTYPES: NY!, UC).
—
Sideranthus puberulus Rydb., Bull. Torrey Bot. Club 27:622. 1900. Type: U.S.A. COLORADO. Chaffee
Co.: Salida, s.d., CB. Clarke 174 (HOLOTYPE: NYI ISOTYPE: UC!)
Sideranthuscarh ‘lus Rydb., Bull. Torrey Bot. Club 27:622. 1900. Hap! | Pursh)
DC. llus(Rydb.) S.F Blake, Contr. Gray Herb. 52:23. 1917,' Aplopappus1 TYPE: USA,
IDAHO, Babtiadl Co. Pocatello, 25 Aug 1892, AT. Mulford s.n. (HOLOTYPE: NYE Isc UC).
Sideranthus machaeranthera J.K. Small, Fl. S.E. US. 1186. 1903. Typ: U.S.A. TEXAS: in io soil,
oe the Brazos River, 1852, A.C. Schott 508a (HOLOTYPE: NY! ISOTYPE: UC).
Cula J.K. Small, FL. S.E. US. 1186. 1903. le ae 2 sarees subsp.
cotula J.K. Small) H.M. Hall, Publ. Carnegie Inst. Wash. 389:77. 1928. TYPE: U.S.A. OKLAHOMA
(Indian Territory): Cherokee Town and vicinity, 1868, E. aie 442A (HOLOTYPE: NY).
MORGAN AND HARTMAN, SYNOPSIS OF MACHAERANTHERA 1407
Sideranthus laevis Wooton & Standley, Contr. U.S. Natl. Herb. 16:180. 1913. Haplopappus spinulosus
(Pursh) DC. subsp. laevis (Wooton & Standley) H.M. Hall, Publ. Carnegie Inst. Wash. 389:78
1928. Machaeranthera laevis (Wooton & Standley) Shinners, Field & Lab. 18:40. 1950. TyPE:
U.S.A. NEW MEXICO. Eddy Co.: gypsum hills near Lakewood, 6 Aug 1909, E.O. Wooton s.n. (HO-
LOTYPE: USI IsOTYPES: UCI, USI).
Haplopappus texensis R.C. Jackson, Rhodora 64:142. 1962. Machaeranthei rd fevensis (RC. Jack-
son) Shinners, Sida 1:378. 1964. TYPE: U.S.A. TEXAS. Brooks Co. 1 g railroad right-
of-way about 7.5 mi S of Falfurrias, 7 Aug 1959, eee 2938- ue ae KANUD.
The following two binomials are not valid as the genus Sideranthus wasa nomen
nudum when the combinations were published: Sideranthus pinnatifidus Nutt.,
Fraser’s Catalogue, no. 81, 1813, non Aplopappus pinnatifidus Nutt. 1840 and
Sideranthus spinulosus (Pursh) Sweet, Hort. Brit., 227. 1826. Furthermore,
Eriocarpum australe Greene (Erythea 2:108. 1894) although validly published
has never been lectotypfied. The relevant synonyms are: Sideranthus australis
(Greene) Rydb., Bull. Torrey Bot. Club 27:621. 1900; Haplopappus spinulosus
(Pursh) DC. subsp. australis (Greene) H.M. Hall, Publ. Carnegie Inst. Wash.
389:77. 1928: and Machaeranthera australis (Greene) Shinners, Field & Lab.
18:42. 1950.
Xanthisma spinulosum “is an exceedingly complex, variable taxon. It is
doubtful that any ‘absolute’ key can be constructed so as to recognize unequivo-
cally the infraspecific categories proposed here. Instead, we have composed a
key that attempts to recognize character trends, in combination, that serve to
distinguisha given taxon from another, but the occasional (if not frequent) speci-
men will be found that keys to a given regional variety but belongs to yet an-
other. However, if one accepts the exception and recognizes our account as an
attempt to portray quite variable, regional populations and not asa key to indi-
viduals, then little trouble would be experienced in pinning a varietal name on
this or that collection” (Turner & Hartman 1976). The following key is adapted
from the aforementioned source.
=
KEY TO THE VARIETIES OF XANTHISMA SPINULOSUM
ao
. Involucres of ites heads 15-25 mm wide, heads mostly solitary on elongate
peduncles 2-1 or more long (shorter in ae California, Mexico); herbs and
subshrubs inet ee )50- 100 cmt tall: foliac ge relati ly sparse, CrOW wded belo Ww, in-
ternodes at mid-stem 0.5-2 mm or more long (short in Baja California, Mexico); W
Colorado to S California and Arizona; Baja California, Mexico and vicinity.
2. Heads borne on elongate, leafy’ eae cles,"the upper leaves not much reduce
subspinose; subshrubs up to 100 cm tall 14d. X. spinulosum var. pgooddinai
2. Heads borne on relatively short, ety peduncles,” the uppel leaves not much
reduced no | bs, but often v oody below or at the crown,
mostly 30- aon tall,
3. Heads a anes lower leaves generally 2—3 cm long, pinnatisect;
W Colorado, adjacent Utah, New Mexico, and Arizona 14f. X. spinulosum
var. paradoxum
1408 BRIT.ORG/SIDA 20(4)
3. Heads broadly turbinate to campanulate; lower leaves 0.5-2 cm long, den-
tate to nearly incised leaves; Baja California, Mexi
4. Involucres of pressed heads 22-25 mm wide; leaves large, decided
cised; coastal islands
x.
O
ir
spinulosum var. incisifolium
4. Involucres of pressed heads 15-20 mm mae small and nearly ser
rate or dentate; widespread on mainland Ag. X. seen var. scabrellum
. Involucre of pressed heads 8-15 mm wide, heads mostly i to 10 or more pers
on relatively terminal and lateral Pecnniels> M 5-2 cm long; herbs mostly 20- cm
tall; foliaa
bundant and not muc | 1s nein at nie stem 0.2—
0.5 cm or ee lon
5. Stems stiffly en Lineslunbenched: foliage glabrate or nearly so; heads nearly
sessile; pappus bristles profuse, exceeding as disc corollas; south Dakota to N
Texas
nulosum var. saa
5. Stems spreading to sprawling, usually much-branched, ae ie pubes
cent, rarely glabrous; pappus less pronounced in the head, equaling the disc
corolla; widespread, W Canada to SW U.S.A.;central Mexico
6. Involucres of pressed heads 8-12 mm wide; stems equally leafy throughout;
plants only rarely stiffly erect or woody at the base, mostly 10-40 cm tall;
widespread spinulosum var. spinulosum
6. Involucres of pressed heads 12-16 mm wide; stems mostly leafy below, up-
per leaves much reduced giving the heads a “long-peduncled" appearance;
jee stiffly erect and often ee at the base, ed 30-50 cm tall; W
s, S New Mexico, S Arizona; N Mex
X. spinulosum var. chihuahuanum
(i) Xanthisma spinulosum subsp. ne
14a. Xanthisma spinulosum var. spinulosum
14b. Xanthisma spinulosum var. chihuahuanum (B.L. Turner & R.L. Hartman)
D.R. Morgan @ R.L. Hartman, comb. nov. BASIONYM: Machaeranthera
pinnatifida (Hook. f.) Shinners var. chihuahuana B.L. Turner & R.L. Hartman,
Wrightia 5:311. 1976. Haplopappus spinulosus (Pursh) DC. var. a uanus (BL,
Turner & R.L. Hartman) K.N. Gandhi, Phytologia 69:276. 1990. TypF: MEXICO.
CHIHUAHUA: Sierra del las Monillas, ca. 16 mi SE of El Morreon on a to lake on
Rio Conchos, 13 Jul 1972, A. M. Powell 2447 (HOLOTYPE: TEX).
14c. oe spinulosum var. glaberrimum (Rydb.) D.R. Morgan & R_L. Hart-
an, comb. nov. BASIONYM: Sideranthus glaberrimus Rydb., Bull. Torrey Bot. Club
7 621. 1900. Sideranthus spinulosus (Pursh) R. Sweet ex Rydb. var. glaberrimus
(Rydb.) A. Nelson, in J.M. Coulter & A. Nelson, Man. Bot. Rocky Mt. 499. 1909.
Haplopappus spinulosus(Pursh) DC. var. glaberrimus (Rydb.) S.F Blake, Contr. Gray
Herb. 52:24. 1917, “Aplopappus.” Haplopappus spinulosus (Pursh) DC. subsp.
glaberrimus (Rydb.) H.M. Hall, Publ. Carnegie Inst. Wash. 389:77. 1928.
Machaeranthera alle as (Hook. f.) Shinners forma glaberrima (Rydb.)
McGregor, Trans. Kansas Acad. Sci. 60:162. 1957. Machaeranthera pinnatifida
(Hook. f.) Shinners var. glaberrima (Rydb.) B.L. Turner & R.L. Hartman, Wrightia
5:311. 1976. TyPE: U.S.A. NEBRASKA. Thomas Co.: Middle Loup River near Thedford,
19 Aug 1893, PA. Rydberg 14034 (HOLOTYPE: NY! ISOTYPES: GHI, US!).
—
(ii) Xanthisma spinulosum subsp. gooddingii
14d. Xanthisma spinulosum var. gooddingii (A. Nelson) D.R. Morgan & RL.
MORGAN AND HARTMAN, SYNOPSIS OF MACHAERANTHERA 1409
Hartman comb. nov. BASIONYM: Sideranthus gooddingii A. Nelson, Bot. Gaz.
(Crawfordsville) 37:266. 1904. Haplopappus gooddingii (A. Nelson) Munz & LM.
Johnston, Bull. Torrey Bot. Club 49:44. 1922, “Aplopappus.” Haplopappus spinulosus
Pursh) DC. subsp. gooddingii (A. Nelson) S.F Blake, Contr. U.S. Natl. Herb. 25:543.
1925, “Aplopappus.” acdc pinnatifida (Hook. f.) Shinners subsp.
gooddingii (A. Nelson) B.L. Turner & R.L. Hartman, Wrightia 5:313. 1976.
Machaeranthera pinnatifida (Hook 1) Shinners var. gooddingii (A. Nelson) BL.
Turner & RL. Hartman, Wrightia 5:314. 1976. TYPE: U.S.A. NEVADA: “The Pockets,”
30 Apr 1902, L.N. Goodding 667 (HOLOTYPE: RMI, ISOTYPE: USI).
Sideranthus viridis J.N. Rose & Standley, Contr. U.S. Natl. Herb. 16:19, pl. 15.1912. TyPE: MEXICO.
SONORA: Pinacate Mountains, 21 Nov 1907, D.T. MacDougal s.n. (HOLOTYPE: US! ISOTYPE: UC).
14e. Xanthisma spinulosum var. incisifolium (1.M. Johnston) D.R. Morgan & R.L.
an, comb. nov. BASIONYM: Haplopappus arenarius Benth. var. incisifolius
I.M. Johnston, Proc. Calif. Acad. Sci. ser. 4, 12:1190. 1924, “Aplopappus.” Haplopappus
spinulosus (Pursh) DC. subsp. incisifolius (.M. Johnston) H.M. Hall, Publ. Carnegie
Inst. Wash. 389:75. 1928. Machaeranthera pinnatifida (Hook. f.) Shinners var.
incisifolia (LM. Johnston) B.L. Turner @ R.L. Hartman, Wrightia 5:315. 1976.
Machaeranthera incisifolia (ILM. Johnston) G.L. Nesom, Phytologia 69:112. 1990.
TYPE: MEXICO. BAJA CALIFORNIA: S San Lorenzo Island, 9 May 1921, I.M. Johnston
3529 (HOLOTYPE: CAS! ISOTYPES: NY!, UCD).
14f. Xanthisma spinulosum var. paradoxum (B.L. Turner & R.L. Hartman) DR.
organ & R.L. Hartman, comb. nov. BASIONYM: Machaeranthera pinnatifida
(Hook. f.) Shinners var. paradoxa B.L. Turner & R.L. Hartman, Wrightia 5:314. 1976.
Haplopappus spinulosus (Pursh) DC. var. pega (B.L. Turner & R.L. Hartman)
Cronquist, Intermountain FI. 5:199. 1994. Type: U.S.A. COLORADO. Montrose Co.:
Paradox, 21 Jun 1912, E. P Walker 147 (HOLOTYPE: NY} ISOTYPES: DS!, GH!, RMI, USD.
14g. Xanthisma spinulosum var. scabrellum (Greene) D.R. Morgan & R_L. Hart-
man, comb. nov. BASIONYM: Eriocarpum scabrellum Greene, Erythea 2:108. 1894.
Haplopappus spinulosus (Pursh) DC. var. scabrellus (Greene) S.F Blake, Contr. Gray
Herb. 52:24. 1917, “Aplopappus.” Haplopappus spinulosus (Pursh) DC. subsp.
scabrellus (Greene) H.M. Hall, Publ. Carnegie Inst. Wash. 389:74. 1928.
Machaeranthera scabrella (Greene) Shinners, Field & Lab. 18:42. 1950.
Machaeranthera pinnatifida var. scabrella (Greene) B.L. Turner & R.L. Hartman,
Wrightia 5:314. 1976. TYPE: MEXICO. BAJA CALIFORNIA: foothills Los Angeles Bay,
Dec 1887, E. Palmer 539 (HOLOTYPE: NDGI; ISOTYPE: NY!).
sae ie ae arenarius Benth. var. rossii 1M. Jo oliiston, Proc. Calif. Ace Sci., ser. 4, 12:1191. 1924,
appus.” TYPE: MEXICO. BAJA CALIFORNI San Marcos
nat 12 May 1921, LM. Johnston 3627 (HOLOTYPE: CASI ISOTYPE: US).
15. Xanthisma stenolobum (Greene) D.R. Morgan @ R.L. Hartman, comb. nov.
BASIONYM: Eriocarpum stenolobum Greene, Erythea 2:109. 1894. Haplopappus
stenolobus (Greene) H.M. Hall, Publ. Carnegie Inst. Wash. 389:65. 1928
Machaeranthera stenoloba (Greene) Shinners, Field & Lab. 18:40. 1950. TYPE:
MEXICO. CHIHUAHUA: Sierra Madre, Arroyo Ancho, 16 Oct 1887, C.G. Pringle 1303
(HOLOTYPE: NDGI; ISOTYPES: GH, NY!, PH).
16. Xanthisma wigginsii (S.F Blake) D.R. Morgan & RL. Hartman, comb. nov.
1410 BRIT.ORG/SIDA 20(4)
BASIONYM: Haplopappus wigginsti S.F. Blake, Proc. Biol. Soc. Wash. 48:169. 1935,
“Aplopappus.” Machaeranthera wigginsii (S.F Blake) R.L. Hartman, Phytologia
68:458. 1990. TyPE: MEXICO. BAJA CALIFORNIA: Sierra San Pedro Martir, 18 Sep 1930,
IL. Wiggins & D. Demaree 4914 (HOLOTYPE: USI).
Xanthisma section Havardii (R.C. Jackson) D.R. Morgan & R.L. Hartman, comb.
nov. BASIONYM: Haplopappus section Havardii R.C. Jackson, Univ. Kansas Sci. Bull.
46:479. 1966. Machaeranthera section Havardii (R.C. Jackson) R.L. Hartman,
Phytologia 68:458. 1990.
Herbs taprooted annuals, 15-45 cm tall. Leaves serrate or dentate, often coarsely
so, teeth blunt or terminating in stiff callosities, not bristle-tipped. Heads radi-
ate. Involucres broadly turbinate. Phyllaries in 5-7 graduated series, linear to
narrowly oblong, the lower portion rigid, stramineous, the upper fifth to half
witha green patch or strip, densely glandular, appressed; apices obtuse to acute.
Receptacles alveolate, scales 0.1-0.5 mm long, mostly distinct. Ray floret corol-
las yellow. Cypselae essentially similar, 2.5-3 mm long, oblong or narrowly el-
liptic, sometimes slightly asymmetrical, flattened laterally, walls thin with 12-
14 barely discernible nerves, sparsely pubescent. Pappus similar in ray and disc
florets, white; bristles 5-7 mm long, filiform, not dorsiventrally flattened ba-
sally, in 2-3 poorly defined series. n = 4.
17. Xanthisma viscidum (Wooton G Standley) D.R. Morgan & R.L. Hartman,
comb. nov. BASIONYM: Sideranthus viscidus Wooton @ Standley, Contr. US. Natl.
Herb. 16:180. 1913. Haplopappus viscidus (Wooton & Standley) S.F Blake, J. Wash.
Acad. Sci. 28:486. 1938. Machaeranthera viscida (Wooton & Standley) R.L. Hart-
man, Phytologia 68:459. 1990. Type: U.S.A. NEW Mexico. Eddy Co. near Hope, 3
Aug 1905, E.O. Wooton s.n. (HOLOTYPE: U 3).
Haplopappus havardii Waterfall, Rhodora 45:352. 1943. Machaeranthera havardii (Waterfall)
Shinners, Field & Lab. 18:40. 1950. Type: U.S.A. TEXAS. Culberson Co. 9 mi E of Van Horn, s.d.,
UT. Waterfall 4153 (HOLOTYPE: GHE Isorypes: ARIZ!, MOL NY),
Arida D.R. Morgan G R.L. Hartman, comb. et stat. nov. BASIONYM: Machaeranthera
section Arida R.L. Hartman, Phytologia 68:446. 1990. Typ: Machaeranthera arida
B.L. Turner & D.B. Ho
Herbs taprooted annuals or rarely short lived perennials (often much
branched, rhizomatous, forming vegetative rosettes in A. blepharophylla), 10-
60(-80) cm tall. Stems erect to ascending or sprawling, often much branched,
glabrous to variously pubescent with glandular and nonglandular hairs. Leaves
alternate; entire, dentate, lacerate to deeply pinnatifid or bipinnatifid, glabrous
to densely pubescent with glandular or nonglandular hairs, apices, including
lobes and teeth, apiculate to bristle tipped, bristles 0.1-1 mm long. Heads radi-
ate or eradiate. Involucres turbinate to depressed hemispheric. Phyllaries in 4-
8 imbricate series, graduated in length, appressed, spreading, or reflexed, gla-
brous or moderately to densely pubescent with glandular or nonglandular hairs:
bases indurate to herbaceous, apices herbaceous, margins sometimes scarious.
MORGAN AND HARTMAN, SYNOPSIS OF MACHAERANTHERA 1411
Receptacles indistinctly alveolate, convex, naked or nearly so, or (in A.
blepharophylla) with scales up to 0.5 mm long. Ray florets pistillate, fertile (ab-
sent in A. carnosa), corollas light to dark blue. Disc florets bisexual, fertile, co-
rollas yellow, + funnelform, tubes and limbs generally glabrous, lobes 0.3-0.5
mm long, glabrous to minutely pubescent. Cypselae of ray and disc florets some-
what dimorphic, narrowly oblong, moderately to densely pubescent; thin
walled, with 5-11 filiform nerves per face, 1-3.2(-4) mm long, those of ray flo-
rets, if present, obscurely 3 sided, rounded abaxially, of disc florets only slightly
compressed laterally; pappus white or whitish (tawny in A. blepharophylla), of
ray florets 1/3 to nearly equaling length of disc florets; bristles minutely
barbellate, in 2-3 poorly differentiated series, at most somewhat dorsi-ventrally
flattened and dissimilar in width near base, bases not to slightly overlapping,
mostly 2.5-4.5(-5) mm long. n = 5.
Four of the nine species that are here included in Arida (A. blepharophylla,
A. parviflora, A. riparia, and A. turneri) were included in molecular investiga-
tions of Machaeranthera s.l.(Morgan & Simpson 1992; Morgan 1997, 2003). The
conflicting relationships supported by cpDNA and nrDNA for these species
suggested substantial reticulate evolution among the four species and with spe-
cies of Dieteria, Pyrrocoma, and Xanthisma, complicating efforts to produce a
taxonomy that best reflects current understanding of the systematics of Arida.
In classifying the species of Arida, one issue to consider is the proper tax-
onomy for A. blepharophylla. Chloroplast DNA and nrDNA supported conf lict-
ing relationships for this species (Fig. 1), suggesting that it has experienced re-
ticulate evolution (Morgan 1997, 2003). Since the taxonomy of A. blepharophylla
should match its evolutionary history as closely as possible, it would be desir-
able to base its classification on what kind of reticulate evolution has occurred.
As described by Morgan (1997), there are two major possibilities. First, because
A. blepharophylla exhibits characteristics of both parental lineages, it could
have originated through hybrid speciation; the best treatment in this situation
might be toestablish A. blepharophylla asa monospecific genus descended from,
but separate from, both parental lineages.
The second evolutionary possibility is that A. blepharophylla isa close rela-
tive of A. riparia that has acquired chloroplast DNA from the ancestor of
Xanthisma gymnocephalum, the best treatment in this case might be to clas-
sify it with A. riparia. Here we choose the latter disposition. According tonrDNA
evidence, the nuclear relationships of A. blepharophylla are with A. riparia. In
addition, the majority of its characteristics, including chromosome number and
fruit morphology, are shared with A. riparia rather than with X.
gymnocephalum (Nesom et al. 1990). It is possible that Arida blepharophylla is
a hybrid species, but in our view additional genetic data would be necessary to
convincingly demonstrate the occurrence of hybrid speciation. In the absence
of such data we propose that A. blepharophylla be considered a close relative of
1412 BRIT.ORG/SIDA 20(4)
A. riparia that has experienced chloroplast introgression, and that its taxonomy
reflect this history.
With Arida blepharophylla classified with A. riparia, the next question is
whether the nine species of Arida should all be maintained in the same genus.
If A. blepharophylla is placed with A. riparia as described above, cpDNA evi-
dence would support keeping all the species together because it supported a
group composed of A. parviflora, A. turneri, and A. riparia (Fig. 1). However,
nrDNA evidence separated these four species into two lineages (Fig. 1). As de-
scribed in detail by Morgan (2003), there are two primary explanations for this
conflict, both involving reticulate evolution. The first explanation considers the
alliances supported by nrDNA to be closer to the true species relationships. The
corresponding classification would split Arida into two pene
The second explanation considers the alliances supported by cpDNA to as
closer to the species relationships for Arida. In supporting the g mposed
of A. parviflora, A. turneri, and A. riparia, cpDNA evidence euasesiéd that the
two lineages of Arida are closest relatives. In this case, the corresponding tax-
onomy would keep the two lineages together, retaining all species of Arida in
the same genus. This is the classification adopted here because, in addition to
cpDNA, several additional pieces of evidence support this treatment. All spe-
cies of Arida have n = 5, all that have been investigated produce flavones, and
hybridization experiments suggest much genetic similarity between the two
lineages (Turner & Horne 1964; Turner et al. 1975; Stucky 1978; Hartman 1976,
1990; Nesom et al. 1990). This array of characteristics unites the species of Arida
and distinguishes them from other taxa in the Machaeranthera complex; we
interpret the conflicting results from nrDNA data as resulting from nrDNA
introgression, as explained by Morgan (2003).
The following key is based in part on data in Hartman (1976, 1990), Nesom
(2003), Nesom et al. 1990), and Turner (1986). It includes the new species Arida
mattturneri (Turner & Nesom 2003).
KEY TO THE SPECIES OF ARIDA
ae cypselae sulieee args eradiate in A. carnosa).
2. Leaves entire to
3. Ray florets eG
3. Ray florets present.
4. Plants annual;leaf margins entire and glabrous or wit
each spinulose tipped; involucres hemispheric, 10-16 mm wide when fresh;
phyllaries linear lanceolate to oblong lanceolate, narrowly acute to
acuminate or attenuate
Plants perenni al rhi omatous, f
3. Arida carnosa
11-8 teeth per side,
8. Arida riparia
egetative rosettes; leaf margins es-
sentially entire with 8-20 cilia c or ee per side, cilia 0.4-0.8(-1.5) mm
long; involucres turbinate, 5-8 mm wide when fresh; phyllaries oblong
ovate to obovate, broadly acute to short acuminate __ 2. Arida blepharophylla
ao
MORGAN AND HARTMAN, SYNOPSIS OF MACHAERANTHERA 1413
2. Leaves (at least some) pinnatifid to bipinnatifid.
5. Receptacles 8-11 mm wide; phyllary apices spreading to reflexed___—_—s9. Arida turneri
5. Receptacles 2-7 mm wide; phyllary apices mostly appressed 7. Arida parviflora
ey cypselae epappose (except in occasional plants).
Mid-stems essentially glabrous or with few sessile glands.
7. Leaves deeply pinnatifid; an Colorado, N New Mexico, Utah, and E Arizona;
C
hihuahua and Coahuila, co 7. Arida parviflora
7. Leaves above basal rosette, a present enulte or toothed.
8 Stems 10-45 cm long, jing to procumbent; SW Sonora, Mexico
4. Arida coulteri
8 Stems 50-80 cm long, erect; W Texas 6. Arida mattturneri
6, Mid-stems densely glandular pubescent.
9. Involucres 3-6 mm tall, 5-10 mm wide; mid-stems d ular, also with
longer, crisp, Sabie hairs interspersed; S Nevada fence S Arizona;
Sonora, Mexi 1. Arida arizonica
9 Involucres ae mm tall, 10-15 mm wide; mid-stems densely apatite
nonglandular hairs absent;N Baja California Sur and adjacent W Sonora, Mexico
5. Arida crispa
1. Arida arizonica (R.C. Jackson & R.R. Johnson) D.R. Morgan & R.L. Hartman,
comb. nov. BASIONYM: Machaeranthera arizonica R.C. Jackson & R.R. Johnson,
Rhodora 69: 476, fig. 1.1967. Type: U.S.A. ARIZONA. Pima Co.: Oregan Pipe Cactus
National Monument, low, rocky hillsides and sandy soils around Quitobaquito
Springs, 31 Mar 1962, R.C. Jackson & R.R. Johnson 3043-1 (HOLOTYPE: KANU,
ISOTYPES: ARIZ!, KANU)).
sa eel arida B.L. Turner & D.B. Horne, Brittonia 16:324. 1964. Machaeranthera coulteri
A. Gray) B.L. Turner var. arida (B.L. Turner & D.B. Horne) B.L. Turner, Phytologia 61:144. 1986.
. PE: sh CALIFORNIA. San Bernardino Co:: ae bige L7 mi SW of Kingston, 2600 ft,
15 May 1941, CB. Wolf 10635 (HOLOTYPE: DS! IsOTYPES: NY!, TEX!, UC).
scans: ammophila Reveal, Bull. Torrey ene hehe 1970. TyPE: U.S.A. NEVADA. Nye
o.: Amargosa Desert, on the W edge of Ash Meadows, 0.3 mi W of Carson Slough, 21 Aug
ee JL. Reveal & N.H. Eee 1882 (HOLOTYPE: US! ISOTYPES: GH!, NY!, RMD.
2. Arida blepharophylla (A. Gray) D.R. Morgan & R.L. Hartman, comb. nov.
BASIONYM: Aster blepharophyllus A. Gray, Pl. Wright. 2:77. 1853. Machaeranthera
gypsitherma G.L. Nesom, Vorobik, and R.L. Hartman, Syst. Bot. 15:638. 1990, nom.
nov, non Machaeranthera blephariphylla (A. Gray) Shinners, Field & Lab. 18:38.
1950. TYPE: U.S.A. NEW MEXICO. Hidalgo Co. in subsaline soil, Las Playas Springs, 7
Oct 1851, C. Wright 1164 (LECTOTYPE: GH! [designated by Nesom et al.1990};
ISOLECTOTYPES: CGE, GH!, MO! NY! P. PH!)
3. Arida carnosa (A. Gray) D.R. Morgan & R.L. Hartman, comb. nov. BASIONYM:
Linosyris carnosa A. Gray, Pl. Wright. 2:80. 1853. Aster carnosus (A. Gray) A. Gray ex
; msley, Biol. Cent.-Amer., Bot. 2:120. 1881, non J.-B. Gilibert, 1781. Leucosyris
carnosa (A. Gray) EL. Greene, Fl. Francisc. 384. 1897. Machaeranthera carnosa (A.
Gray) G.L. Nesom, Phytologia 67:439. 1989. TYPE: MEXICO. SONORA: W of Chiricahui
Mountains, Sep 1851, C. Wright 1187 (HOLOTYPE: GHk ISOTYPES: PHI, US!).
Bigelowia intricatus A. Gray, Proc. Amer. Acad. Arts 17:208. 1882. Aster intricatus (A. Gray) 5.F
Blake, J. Wash. Acad. Sci. 27:378. 1937. Machaeranthea carnosa (A. Gray) G.L. Nesom var.
1414 BRIT.ORG/SIDA 20(4)
py
intricata (A. Gray) G.L. Nesom, Phytologia 82:106. 1997. TypE: U.S.A. CALIFORNIA: Mohave
Desert, at Lancaster Station, s.d., CC. Parry s.n. (HOLOTYPE: GH!)
4. Arida coulteri (A. Gray) D.R. Morgan & R.L. Hartman, comb. nov. BASIONYM:
Psilactis coulteri A. Gray, Pl. Fendler. 72. 1849. Machaeranthera coulteri (A. Gray)
B.L. Turner & D.B. Horne, Brittonia 16:322. 1964. TYPE: MEXICO. SONORA: probably
the vicinity of Guaymas, n.d., [. Coulter 295 (HOLOTYPE: GH!).
5. Arida crispa (TS. Brandegee) D.R. Morgan @ R.L. Hartman, comb. nov.
BASIONYM: Psilactis crispa TS. Brandegee, Proc. Calif. Acad. Sci, ser. 2, 2:169. 1889.
Machaeranthera crispa (TS. Brandegee) B.L. Turner & D.B. Horne, Brittonia 16:321.
1964. Type: MEXICO. BAJA CALIFORNIA SUR: alkaline soil, San Joaquine, San Igna-
cio, | Apr 1889, T.S. Brandegee s.n. (HOLOTYPE: UCI; IsoTYPES: GH!, US).
6. Arida mattturneri B.L. Turner & G.L. Nesom, Sida 20:1418. 2003. Typr: U.S.A.
TEXAS. Presidio Co.: ca. 2.2 mi NNW of Ruidosa in “Blumberg Canyon,” (ca. 30° 00’
45" N, 104° 44'00" W), 20 Jul 2003, M.W. Turner 100 (HOLOTYPE: TEX; ISOTYPES: NY],
SRSG)
7. Arida parviflora (A. Gray) D.R. Morgan & R.L. Hartman, comb. nov. BASIONYM:
Machaeranthera parviflora A. Gray, Pl. Wright. 1:90. 1852. Aster parviflorus (A.
Gray) A. Gray in W.H. Brewer & S. Watson, Bot. California 1:322. 1876, non C. Nees,
1818, nec W. Darlington, 1826. Aster parvulus S.F. Blake in Tidestrom, Contr. U.S.
Natl. Herb. 25:563. 1925. Type: U.S.A. TEXAS: “collected in expedition from W Texas
to El Paso, N. Mex.,” May-Oct 1849, C. Wright 271 (HOLOTYPE: GHI, IsoTYPES: NY1,
US).
Machaeranthera tanacetifolia (Kunth) C. Nees var. pygmaea A. Gray, Pl. Wright. 2:74. 1853. Aster
tanacetifolia Kunth var. pygmaeus (A. Gray) A. Gray, Syn. Fl. N. Amer. 1(2):206. 1884.
Machaeranthera pygmaea (A. Gray) Wooton & PC. Standley, Contr. U.S. Natl. Herb. 16:189.
1913. Type: U.S.A. NEW MEXIco: near El Paso, 1851, C. Wright 1395 (HOLOTYPE: GHk ISOTYPES:
NY!, USI).
8. Arida riparia (Kunth) D.R. Morgan & R.L. Hartman, comb. nov. BASIONYM: As-
ter riparius Kunth, Nov. Gen. Sp., folio ed., 4:72. 1818. Machaeranthera riparia
(Kunth) A.G. Jones, Syst. Bot. 8:85. 1983. Tyre: MEXICO: “In humidis juxta lacum
Cuiseo. .. alt. 900 hex.” n.d, EW.H.A. von Humboldt & A.A. Bonpland 4308 (HOLO-
TYPE: P; ISOTYPES: B-W, P
Aster sonorae A. Gray, Pl. Wright. 2:76. 1853. Machaeranthera sonorae (A. Gray) Stucky, Amer. J.
Bot. 65:132. 1978. Type: U.S.A. SOUTHERN ARIZONA: low valley W of the Chiricahui Mountains,
Sep 1851, C. Wright 1163 CHOLOTYPE: GH! ISOTYPES: CGE, PHI, USD.
9. salar turneri (M.L. Arnold @ R.C. Jackson) D.R. Morgan & - L. Hartman, comb.
Ov. BASIONYM: Machaeranthera turneri M.L. Arnold & R.C. Jackson, Syst. Bot.
3.209 1979. TYPE: MEXICO. CHIHUAHUA: 3.3 mi N of eee ui, along highway 45, |
Aug 1964, R.C_ Jackson 4005 (HOLOTYPE: TTC).
ACKNOWLEDGMENTS
We thank Guy Nesom for reviewing the manuscript, John F Pruski and Victo-
ria C. Hollowell for detailed suggestions on an earlier version of this work, and
MORGAN AND HARTMAN, SYNOPSIS OF MACHAERANTHERA 1415
James L. Reveal for advice on several nomenclatural questions.
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599-615.
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71:122-127.
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Mexico (in press).
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(Asteraceae: Astereae). Syst. Bot. 15:638-642.
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(Asteraceae): Proper usage of infraspecific categories. Rhodora 76:1-19.
StaFLeu, FA.and R.S. Cowan. 1981.Taxonomic literature, Lh—O, ed. 2, vol. 3. Bohn, Scheltema
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Stucky, J. 1978. Hybridization between Aster and Machaeranthera and its taxonomic sig-
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(Asteraceae- Astereae). Phytologia 61:143-146.
1416 BRIT.ORG/SIDA 20(4)
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A NEW SPECIES OF ARIDA (MACHAERANTHERA
SECT. ARIDA—ASTERACEAE: ASTEREAE)
PROM TRANS ECOS. TEXAS
B.L. Turner Guy L.Nesom
Plant Resources Center Botanical Research Institute of Texas
The University of Texas 09 Pecan Street
Austin, 1X 78713, U.S.A. Fort Worth, TX 76102-4060, U.S.A.
ABSTRACT
Arida mattturneri Turner & Nesom, sp. nov. is described from collections made near Ruidosa in
Presidio County, Texas. The plants are perennial, lavender-rayed, and occur in gypseous soils along
shallow perennial streams and seeps. It isa member of Machaeranthera sect. Arida, now segregated
as the genus Arida, and is similar to Arida turneri in significant respects, especially the bipinnately
parted leaves. The new species is illustrated by photographs.
RESUMEN
Se describe Arida mattturneri Turner @ Nesom, sp. nov. a partly ee colecciones hechas cerca de
Ruidosa, en Presidio County, Texas. Las pl. antas son perennes | d aparecenen sue los
y
yesosos a lo largo de < permanentes poco profundos y lugares rezumantes. Es un oe de
Arid
ee ae sect. Arida, ahora segregado como género Arida, y es semejat u
aspectos importantes, especialmente en las hojas bipinnado-partidas. La nueva especie se slustes a
mediante fotografias.
In the spring of 2001, Ms. Charmaine Delmatier of the U.S. Fish and Wildlife
Service, Austin, Texas, collected sterile material of plants occurring along the
gypseous, seepy walls of a canyon near Ruidosa, Texas (Presidio County), on
the property of James Blumberg. She asked the senior author to identify the
material concerned and, because of its peculiar habitat, there was little prob-
lem in tentatively identifying it as Machaeranthera gypsitherma Nesom,
Vorobik, and Hartman (1990), a rare species previously unknown to Texas—it
now apparently is extinct at its type locality in southern New Mexico and oth-
erwise known from only a single population in Chihuahua, Mexico.
Ina later visit to the Presidio County site (spring of 2003), to collect flow-
ering material of this remarkable discovery, the present authors encountered
essentially sterile material of yet another Machaeranthera, which is the sub-
ject of the present paper. The senior author, along with his son Matt and James
Blumberg, revisited the site in the ensuing summer, at which time abundant
flowering material of the plants concerned was collected. Examination of the
material clearly indicates its status as a previously undescribed species.
We follow recent morphological and molecular studies and taxonomy of
Ronald Hartman and David Morgan, as well as their forthcoming treatments
SIDA 20(4): 1417-1422. 2003
1418 BRIT.ORG/SIDA 20(4)
for the Flora of North America volumes of Asteraceae, in dividing the species
of Machaeranthera (sensu Hartman 1990) among four segregate genera, includ-
ing Machaeranthera sensu stricto but not including Psilactis (sensu Morgan
1993). Nomenclatural combinations to formally effect these taxonomic changes
are in this current issue of Sida (Morgan & Hartman 2003).
Arida oeemaaal B.L. Turner & G. Nesom, sp. nov. (Figs. 1, 2). Type: UNITE D
oc . TEXAS: Presidio Co.: ca, 2.2 mi NNW of Ruidosa in es 7 ‘anyon,
growing aie perennial sulphur streams and around tinajas (ca. 30° 00' 00" N,
LOo+ 44° ( Jo" W). 20 Jul 2003, Matt W. Turner 100 (HOLOTYPE: TEX; ISOTYPES: NY,
SRSC).
Aridae turneri (Arnold & Jackson) Morgan & Hartman similis vestimento glanduloso, foliis
bipinnatifidis lobis mucronatis, flosculis radii lavandulaceis, acheniis 1.0-1.5 mm longis nervis
tenuibus numerosis, et pappo brevi, sed differt plantis perennibus foliis basalibus persistentibus,
ut-
capitulis minoribus flosculis radii paucioribus, phyllariis oblongi-lanceolatis apicibus rotunc
acutis, et Flosculis disci pappo carenti.
Perennial herbs from a thick, lignescent taproot, caudex simple, stems, leaves,
and phyllaries sticky-viscid from resin globules exuded by sessile or minutely
stipitate glands, without other kinds of hairs; leaves and phyllaries with indu-
rate mucros but not bristle-tipped. Stems erect, 50-80 cm tall. Leaves basal and
cauline, basal in persistent rosettes, bipinnately parted, abruptly passing into
merely pinnately parted lower and midcauline, gradually reduced upwards into
entire, scale-like bracts; basal +-6 cm long, 1.5-3.5 cm wide, broadly obovate in
outline, ultimate segments ovate to linear-lanceolate or linear. Heads ca. 10-13,
terminal on elongate bracteate peduncles 5-10 cm long, collectively forming a
broad, open capitulescence; involucres campanulate, 5-6 mm high, 6-8 mm
wide, phyllaries in 5-6 graduate series, oblong-lanceolate, 1.0-1.5 mm wide, out-
ermost |-2 mm long, innermost 5-6 mm long, distal 1/3-1/2 slightly widened
then rounded-acute, green and slightly turgid-thickened; receptacles plane, ca.
5mm wide, glabrous. Ray florets 9-13, pistillate, fertile, tubes ca. 1.5 mm long,
ligules 9-10 mm long, lavender, markedly drooping in full bloom, tightly coil-
ing during fruit maturity. Dise florets 40-100, fertile, corollas narrowly cylin-
dric, yellow, 3.5-4.5 mm long, tubes 1.2-1.5 mm long, grading into the nearly
glabrous throats 2.0-2.8 mm long, lobes 5, narrowly deltoid, erect, 0.3-0.5 mm
long. Achenes |.0-1.5 mm long, obovate-oblong in outline, 8-12 nerved, com-
pressed but more or less 3(-4) sided, densely sericeous; pappus of disc achenes
of numerous whitish bristles mostly 3-4 mm long, in 2-3 series and variable in
length ona single achene, absent on ray achenes. Chromosome number, 2n = 10
(Powell 2003, this issue).
The species is named for the senior author's son, Matt Warnock Turner, who
participated in collection of type materials. Matt once confided to his father
that if ever a plant were to be named for him, he would like it to be very distinct,
relatively rare, and confined to Texas. With such a pitch, the senior Turner de-
om
TURNER AND NESOM, A NEW SPECIES OF ARIDA FROM TRANS-PECOS, TEXAS 1419
spaired of ever finding a plant with such stringent ‘cultural’ parameters. The
present novelty fulfills all such requirements, and with considerable filial pride,
he has proposed the name concerned. The name also is appropriate, since its
close relative, Arida turneri (Arnold & Jackson) Morgan @ Hartman, is named
for his father, thus both the plants and their eponymy bear a genetic connec-
tion. We also note that Matt isa first-rate systematist, evidenced by his account
of the southwestern U.S.A. species of Baileya (Turner 1993) and the more east-
ern Brazoria (Turner 1996).
Following Hartman’s conspectus (1990) of the genus Machaeranthera, the
features of Arida mattturneri place it as a member of sect. Arida R.L. Hartman
(now genus Arida (R.L. Hartman) Morgan & Hartman): leaves without bristle-
tipped lobes, ray corollas lavender, achenes 1.5-3 mm long with thin walls and
numerous, thin, superficial nerves, and pappus of disc achenes of relatively short
bristles, absent on ray achenes. The chromosome number of A. maitturneri is n
= 5 (from plants at the type locality; Powell 2003), which is constant for other
species of Arida. All other species of Machaeranthera (sensu Hartman 1990)
have chromosome numbers based on x = 4.
Among taxa of Arida, the only previously described species with
bipinnatifid leaves is A. turneri, which is here hypothesized to be a close rela-
tive of the new species. Both species also occur in close proximity to saline or
gypseous seeps and springs. The two are distinguished as follows:
—
. Plants perennial; basal leaves persistent; involucres 6-8 mm wide, receptacles ca.5
mm wide; phyllaries oblong-lanceolate, 1.0-1.5 mm wide, distal 1/3-1/2 slightly
widened, then with rounded-acute apices; ray florets 9-13, corollas 10-12 mm long,
a
chenes ;disc corollas 3.5-4.5 mm lon Arida mattturneri
1. Plants annual pas leaves deciduous before flowering; involucres 12- oe mm wide,
receptacles 8-11 mm wide; phyllaries linear-lanceolate, 0.8-1.0 mm wid ver with
narrowly acute to acuminate apices; ray florets 41-59, corollas 11- 1S mm eng,
achenes pappose; disc corollas 4.5-6.0 mm long Arida turneri
Arnold and Jackson (1978), as well as Hartman (1990), observed that the closest
relative of Arida turneri is A. (Machaeranthera) parviflora (A. Gray) Morgan
& Hartman. Asin A.turneri, plants of A. parviflora are annual with basal leaves
deciduous before flowering, but the latter differs primarily in its smaller and
less dissected leaves and smaller heads (shorter and narrower involucres and
fewer and shorter ray and disc florets). Ray achenes in some populations of A.
parviflora are epappose but pappose in others. Differences between the two
species appear to be mainly quantitative, but they have been collected in close
proximity apparently without intergrades. Even so, morphological evidence is
equivocal in suggesting relationships among the three species under consider-
ation, and it remains to be discovered whether the two annuals (A. turneri and
A. parviflora) or the two with bipinnatifid leaves (A. turneriand A. mattturneri)
are more closely related.
1420 BRIT.ORG/SIDA 20(4)
Fic. 1 Upper h f | f Arid [ t | li y, April 2003 l n 1 og
at same site, July 2003.
TURNER AND NESOM, A NEW SPECIES OF ARIDA FROM TRANS-PECOS, TEXAS 1421
Fic. 2. Flowering heads showing (left, at ant
CU Tay I
When originally described, Arida turneri (as Machaeranthera turneri;
Arnold and Jackson 1978) was primarily known by material from the vicinity
of Meoqui, Chihuahua, Mexico, but subsequent collections have been made in
Mpio. Julimes, Chihuahua, in the vicinity of Banos de San Diego (McDonald
2549, BRIT, TEX); Nesom & Vorobik 5479, TEX; and Nesom 6543b, TEX). The only
other known extant locality for Arida blepharophylla (A. Gray) Morgan & Hart-
man (= Machaeranthera gypsitherma) also is Banos de San Diego, thus all three
species in the discussion are rare and have similar ranges. Arida mattturneriis
now the only one of the three known from only a single locality.
ACKNOWLEDGMENTS
We are grateful for the assistance of Mr. James Blumberg; he not only gave us
free access to his property, he also accompanied the two Turners in their acqui-
sition of type material. Thanks also to Mike Powell for providing the chromo-
some count and to Ron Hartman for review comments.
REFERENCES
Arnot, M.L.and R.C. Jackson. 1978. Biochemical, cytogenetic and morphological relation-
ships of a new species of Machaeranthera sect. Arida (Compositae). Syst. Bot. 3:
208-217.
Hartman, R.L. 1990. A conspectus of Machaeranthera (Asteraceae: Astereae). Phytologia
68:439—-465.
1422 BRIT.ORG/SIDA 20(4)
Morcan, D.R. 1993. A molecular systematic study and taxonomic revision of Psilactis
(Asteraceae: Astereae). Syst. Bot. 18:290-308
Moroan, D.R. and R.L. Hartman. 2003. A synopsis of Machaeranthera (Asteraceae: Astereae).
Sida 20:1387-1416.
Nesom, G.L., L.A. Vorosik, and R.L. Hartman. 1990. The identity of Aster blepharophyllus A. Gray
(Astereae: Asteraceae). Syst. Bot. 15:638-642.
Powett, A.M. 2003. Documented chromosome numbers 2003:2. Chromosome numbers
of two rare species of Machaeranthera sect. Arida (Asteraceae: Astereae). Sida 20:
1663.
Turner, MW. 1993. Systematic study of the genus Baileya (Asteraceae: Helenieae). Sida
15:491—508.
Turner, M.W. 1996. Systematic study of the genus Brazoria (Lamiaceae), and Warnockia
(Lamiaceae), a new genus from Texas. Pl. Syst. Evol. 203:65-82.
FURTHER ADDITIONS TO THE GENUS ARDISIA
SUBGENUS GRAPHARDISIA (MYRSINACEAE)
Jon M.Ricketson John J. Pipoly Ill
Missouri Botanical Garden Fairchild Tropical Garden
PO. Box 299 11935 Old Cutler Road, Coral Gables (Miami)
St. Louis, MO 63166-0299, U.S.A. Florida 33156-4242, U.S.A.
jon.ricketson@mobot.org jpipoly@fairchildgarden.org
ABSTRACT
Revision of the group of species formerly placed in Oerstedianthus Lundell revealed that the group is
synonymous with Ardisia subgenus crear) Mez oa ies incorporates those taxa in the
bg
,inclu
ing an emended descrip-
subgenus, necessitating further revision of
tion of the subgenus, a key to the species and sabenecies ee wach updated pal ioe of the four
added taxa and new illustrations for each. Four additional taxa are recognized, including one new
D Ardisia verdisepala Ricketson & Pipoly, and one new combination: cs nigrescens subsp.
onnellsmithii (Mez) Ricketson & Pipoly. Seven binomials are relegated to synonymy within A.
Ly wn
tuerckheimii Donn. Sm. In addition, Ardisia nigrescens Oerst., A. nigrescens subsp. donnellsmithii
(Mez) Ricketson & Pipoly and A. tuerckheimii Donn. Sm. are lectotypified.
RESUMEN
Una revision del grupo de especies anteriormente clasificadas dentro del género Oerstedianthus
Lundell revelo que el erape se ubica mejor dentro del género Ardisia subgénero Graphardisia Mez, y
por lo tanto se rel a sinonimia. El propésito del presente trabajo es para incorporar ese
grupo dentro del subgenero y actualizar su circunscripcion, claves para identificar las especies y
les lla uevas ilustraciones para cada una. Se reconocen cuatro taxa
subespecies, d
adicionales, ae -inclaye una capecie nueva para la ciencia: Ardisia a verdisepala Ricketsou ou pe
] 1]
<I
siente binomiales a la sinonimia baie A. buerekheimnii Dean: Sm, Ademas, se lectotipiican Ardisia
nigrescens Oerst., A. nigrescens subsp. donnellsmithii (Mez) Ricketson & Pipoly y A. tuerckheimii
Donn. Sm.
INTRODUCTION
The pantropical genus Ardisia Sw. is the largest within the Myrsinaceae, con-
taining perhaps as many as 500 species (Chen & Pipoly 1996). Its circumscrip-
tion has been problematic owing to a lack of comprehensive treatment since
that of Mez (1902) in Engler’s Das Pflanzenreich over a century ago. In prepara-
tion of our treatment of the Myrsinaceae for Flora Mesoamericana, we have criti-
cally reexamined taxa segregated by Lundell from Ardisia, including Gentlea
(Ricketson & Pipoly 1997), Graphardisia (Pipoly & Ricketson 1998), Chontalesia
and species related to it, now known to belong to the genus Hymenandra (Pipoly
& Ricketson 1999a), Zunilia which is now known to belong to Ardisia subgen.
Graphardisia (Pipoly & Ricketson 1999b), Ardisia rarescens Standl. which was
SIDA 20(4): 1423-1447, 2003
1424 BRIT.ORG/SIDA 20(4)
placed in Amatlania by Lundell and is now known to belong to Ardisia subgen.
Acrardisia (Pipoly & Ricketson 2000), Auriculardisia, Amatlania and
Valerioanthus now all belonging to Ardisia subgen. Auriculardisia (Ricketson
& Pipoly 2003) and the segregate genera, Yunckeria Lundell (=Ctenardisia
Ducke) and Ibarrea Lundell (=Ardisia subgen. Ardisia). Since our study of the
subgenus Graphardisia (Pipoly & Ricketson 1998, 1999b), including Lundell’s
segregate genera Graphardisia and Zunilia, it has become evident that the taxa
comprising the segregate genus Oerstedianthus (Lundell 1981) are best placed
within Ardisia subgenus Graphardisia.
According to Lundell (981:139-140), “The genus [Graphardisia] is related
to Oerstedianthus Lundell.” He separated the two as follows:
“Filaments strictly glabrous; stems and inflorescence glabrous; punctations of all parts
dense and blackish; bracts and bractlets usually foliaceous and often persistent;
ae and see Bre accrescent, usually blackened, usually ribbed or with dense
elevated black glands : Graphardisia
4 . apes pe | Ia}
yee ti pl aweace with gla anad-t i ly Qlabr
usually puberulent, hirtellous or r dense hirsute-tomentose; punctations of all nate
conspicuous but not prominently raised, pellucid or rarely blackish; bracts and
sepals small, not accrescent; sepals not ribbed with glands Oerstedianthus’
However, our studies of Ardisia subgenus Graphardisia (Pipoly & Ricketson
1998, 1999b) have clearly shown the presence of raised or sessile glandular-papil-
late filaments in all taxa. In addition, because we are not sure of the polarity of
this character state, or if it can have multiple origins, its utility is dubious. Fo-
liar and other punctations are said to be inconspicuous in the taxa newly added
in this work, but they are normally prominent and occasionally conspicuous.
Finally, the sepals are generally small, but they are comparable to those found
in other members of subgenus Graphardisia, (ie. Ardisia bartlettii Lundell and
A. weberbaueri Mez). Lundell (98L:141) also notes: “Oerstedianthus is related
to Graphardisia (Mez) Lundell. Both have similar linear-lanceolate anthers
which dehisce by apical pores.” Therefore, we find it necessary to relegate the
genus Oerstedianthus Lundell to synonymy under Ardisia subgenus
Graphardisia.
TAXONOMIC TREATMENT
Ardisia subgenus Graphardisia Mez in Eng]., Pflanzenr. lV. 236 (Heft 9):78. 1902:
Lundell, Wrightia 3:192-198. 1966. Graphardisia (Mez) Lundell, Phytologia
48:139. 1981; Lundell, Phytologia 59:429-433. 1986. Type: Ardisia opegrapha Oerst.
(LECTOTYPE, designated by Lundell, Phytologia 48:139. 1981)
—
Oerstedianthus Lundell, Phytologia 48:14L 1981. Syn. Nov. Type: A ee nigrescens Oerst. 1861:130.
1862. Oerstedianthus nigrescens (Oerst. - undell, Phytologia 48:141. 1981.
saat - undell, eae 49:353. 1981. TYPE: Ardisia se oats ita L ne Wrightia 3:29. 1962.
x] Lundell) Lundell, cereepaee L.
Subshrubs to trees. Branchlets mostly terete, ow puberulent, hirsute or
1425
rarely, glandular-granulose. Leaves petiolate; blades membranous to subcoria-
ceous, densely and prominently black (rarely pellucid-) punctate and punc-
tate-lineate, the margins entire to crenulate or serrate. Inflorescence terminal,
rarely lateral, pinnate to tripinnately paniculate, pyramidal to obpyramidal
rarely globose, the ultimate branches corymbose, at times in high anthotactic
spirals and thus appearing umbellate, the rachis often densely and prominently
black punctate and punctate-lineate; inflorescence and floral bracts minute to
foliaceous, caducous to persistent, often resembling the vegetative leaves but
acropetally reduced in size. Flowers 5 or 6-merous, densely and prominently
black punctate and punctate-lineate; calyx with sepals free or nearly free, large,
accrescent and clasping fruits at maturity, corolla white, pink, lavender or
purple, rotate, the lobes imbricate in bud, basally short-connate and sparsely
to densely yellow glandular-granulose within at the base, densely and promi-
nently black punctate-lineate; stamens inserted at corolla tube base, the fila-
ments basally connate to form an inconspicuous tube, the tube free from the
corolla tube, the apically free portions of the filaments with scattered yellow
glandular-granulose or glandular-papillate, less than half the length of the
anthers, the anthers ovoid-lanceoloid, linear or lanceoloid, prominently apicu-
late, dehiscent by subapical pores; ovary globose, the style slender, equaling the
sepals, 2—3 times longer than the ovary, the placenta apiculate, the ovules
pluriseriate, biseriate, or apparently uniseriate (few in number and ina very
high anthotactic spiral). Fruit globose or oblongoid, densely punctate and punc-
tate-lineate, basally surrounded by persistent, clasping sepals.
Distribution.—A small distinctive subgenus of seven species with nine sub-
species found from Mexico to Bolivia and adjacent Brazil.
Ecology.—Members of the subgenus occur in diverse humid to pluvial veg-
etation types, lowland, premontane, montane, and cloud forests.
The subgenus is defined by: 1) black, or rarely, reddish-black, punctations
or punctate-lineations on all leaf and floral parts; 2) sepals often accrescent
and usually clasping the developing fruit; 3) linear-lanceolate, concolorous,
apiculate anthers with subapical, poricidal dehiscence; and 4) styles that are
2-3 times longer than the ovary. Among the species of Ardisia, it is notable that
the branchlets, petioles and inflorescence rachises among species of subg.
Graphardisia are never tomentose, but may be glabrous, puberulent, rarely hir-
sute or glandular-granulose. Therefore multicellular trichomes are not found
in the subgenus. Species of the subgenus are often used for home decoration
and for use in Christian churches for religious holidays (Pipoly, pers. obs.).
KEY TO TAXA OF ARDISIA SUBGENUS GRAPHARDISIA
1. Stoloniferous subshrubs mostly less than 1(—2) m tall; leaf blade margins sharply
and irregularly dentate; corolla tube and filaments yellow glandular-granulose;
Amazonian Ecuador to Bolivia and adjacent Brazil. A. weberbaueri
1426 BRIT.ORG/SIDA 20(4)
Shrubs to small trees (0.5—)2—6(-30) m tall, without stolons; leaf blade margins en-
tire, undulate or crenulate to serrate; corolla tube yellow glandular-granulose; fila-
ments glabrous or sessile to stalked glandular-
papillate; Mexico to the Choco Re-
gion of Colombi
a.
2. Sepals oblong, 4.2-8 mm long, apically broadly rounded to obtuse, the margins
entire
A. opegrapha
. Inflorescence obpyramidal; leaf blades oblanceolate or rarely obovate, 3.5—
7.5(-8) cm wide, 3 or more times longer than wide.
4. Floral bracts caducous; stamens 5.2-6.5 mm ie 2.5-3 mm long;
dals 5-8 mm lon
egrapha subsp. opegrapha
al bracts persistent; stamens 3.8-5 mm long; ne 1.5-2 mm long;
yals 4.2—5.2 mm long
=
pegrapha subsp wagneri
—)8-14.5
3. Inf ones globose; leaf blades elliptic to broadly elliptic 5
e, 2-2.5 times longer than wide opegrapha subsp. 5 oaaunens
2 seh ovate, 1.5-3.2(-4) mm long, apically acute to mua the margins
subentire to entire or erose.
5. Corolla lobes ovate, ellipt
Colombia
6. Sepals 1.
ic or lanceolate; filaments glabrous; Panama and
A. bartlettii
1.8mm long; petal lobes 6-6.5 mm long; stamens 3.5— 4
long; ee base tapering; tall wet forest A. bartlettii subsp. bartlettii
pals 2-2.5 mm long; petal lobes 7-8 mm long; stamens 4.5—5.7 mm long;
style base stylopodic; strand vegetation and beach forests
A. bartlettii
subsp. lilacina
5 G ~|| LAF | fi] | i] H
Tl tall feo Alaa
= PaVltac LU SLGIRNGU qgianaular papil
late: Oa aca, Mexico ‘6 Nicara
7. Branchlets and ieee nee rachis glabrous; anthers 3—3.2 mm long
A. verapazensis
8 Corolla lobes 6.5-6.7 mm long; free sala of filaments 3.3-3.4 mm
long; style 3-3.1 mm long; fruit 6.5-9 mm in diam.; fruiting style 7-8.6
mm fis susually only the basal portion eee nt A. verapazensis subsp.
erapazensis
8. Corolla lobes 5.7-5.9 mm long; free portion of filaments 2.6-2.8 mm
long; style 5.5-5.9 mm long; fruit 5—6.1 mm in diam, fruiting style 8.8
9.4mm long, the entire style usually persistent
A. verapazensis
subsp. cucullata
7. Branchlets and inflorescence rachis long-hirsute with reddish hairs or very
sparsely to densely puberulent of simple trichomes; anther
long.
2-5.) mm
9. Branchlets very sparsely to densely some .A.tuerckheimii
9. Branchlets scattered to densely lo g hirsute wi with reddish hairs.
10. Calyx lobes oblong, 3.4-3.8 mm; ste lobes 8-8.4 mm; filaments
as wide as anthers, the anthers basally hastate. _ 2.A.verdisepala
10. Calyx lobes ovate, 1.9-2.7 mm; corolla lobes 4.8-8 mm; filaments
abruptly constricted apically, the anthers basally aed wae __ 1.A. nigrescens
11. Leaf margins crenulate or serrate; calyx lobes 1.9-2.1 mm
1a. A. nigrescens subsp. nigrescens
ll. Leaf oa bisa, entire to slightly Siar or serrate; calyx
lobes 2.2-2.7 A. nigrescens subsp. donnellsmithii
1. Ardisia nigrescens Ocrst., Vidensk. Meddel. Dansk Naturhist. Foren
1427
Kjobenhavn 1861:130. 1862. Icacorea nigrescens (Oerst.) Standl., Contr. U.S. Natl.
Herb. 23:1109. 1924. Oerstedianthus nigrescens (Oerst.) Lundell, Phytologia 48:142.
1981. Type. MEXICO. VERACRUZ: Prope Colipa, without elev, Mar 1842 (fr), F
Liebmann 28B (LECTOTYPE: C, here designated, F neg. # 22952, in part).
Shrubs or small trees 1-12 m tall, to 7.5cm in diam. Branchlets |-5 mm in diam.,
densely long hirsute reddish hairs throughout. Leaves: with leaf blades mem-
branous to chartaceous, ovate or elliptic to oblong, 3.5-18.2 cm long, 1.5-8.2 cm
wide, with an acumen 0.8-2.1 cm long, basally obtuse, inconspicuously to con-
spicuously, often prominently punctate and punctate-lineate, sparsely hispid
above and below, often densely hispid along the midrib above, densely hirsute
along the midrib below, the secondary veins 15-24 pairs, the margins entire or
crenulate to serrate; petioles 0.5-L.6 cm long, densely hispid along above, densely
hirsute below. Inflorescences 1.5-6 cm long, 2-8 cm wide, shorter than the leaves,
the rachis with dense, long hirsute with reddish hairs, the branches 3-18-flow-
ered corymbs; peduncles nearly absent to 1.2 cm long, densely long hirsute with
reddish hairs; inflorescence branch bracts oblong to oblanceolate, 4-25 mm
long, 1-14 mm wide, scattered long hirsute with reddish hairs; floral bracts simi-
lar to the inflorescence bracts but, when present, to 0.3-0.8 mm long, 0.1-0.2
mm wide; pedicels 0.7-1.4 cm long, conspicuously and prominently punctate
and punctate-lineate, with densely long hirsute simple reddish hairs. Flowers
5-6-merous; calyx lobes ovate, 1.9-2.7 mm long, 1.1-1.5 mm wide, apically acute,
sparsely hirsute outside, glandular-granules within near the base, the margins
entire, densely hirsute-ciliolate; corolla 6.5-10 mm long, the tube 1.2-2.4 mm
long, the lobes +.8-8 mm long, 2.2-4.8 mm wide; stamens 4.6-6 mm long, the
filaments 1.5-2.8 mm long, the staminal tube 0.5-1 mm long, the apically free
portion abruptly constricted 1-2.1 mm long, the anthers 3.3-4.5 mm long, 0.8-
Ll mm wide; pistil 5-6.4 mm long, the ovary 0.8-1.2 mm long, the style 4-4.7
mm long, the ovules 5-7. Fruits 5.5-7.3 mm in diam.
In Oersted’s (1862: p. 130) original description he cites two localities, both
collected by Liebmann: “Crescit in Mexico, ubi cl. Liebmann Martio
fructificantem prope Colipa et Jecaltepec legit.” The Museum Botanicum
Hauniense (C) clearly houses the two collections used by Oersted, “Mexico, Prope
Jecaltepec, Mar. 1842 (fr), E Liebmann 28A” and “Mexico, Prope Colipa, Mar 1842
(fr), F Liebmann 28B.” Both collections are in fruit, but the 28A specimen has
narrower leaves with no attached fruits and the 28B specimen has wider leaves
with many attached fruits. Unfortunately, the photograph taken by J.E Macbride
during the Field Museum of Natural History’s project to photograph European
types (F neg. no. 22952), is a combination of both specimens, although prima-
rily of the 28B specimens. Thus, because the 28B specimen is more complete
and more widely seen via the F photograph, we elect to designate the Liebmann
28B (C) specimen as the lectotype. Although, CL. Lundell annotated the 28A
specimen in 1966 as the “TYPE!” he did not formally publish a lectotypification.
1428 BRIT.ORG/SIDA 20(4)
Within Ardisia subgen. Graphardisia, A. nigrescens is most closely related
to A. verdisepala, because of the long, hirsute, with reddish tricomes through-
out the vegetative parts of the plant. However, Ardisia nigrescens may be easily
distinguished by its shorter, ovate calyx lobes (1.9-2.7 mm long), and shorter
corolla lobes (4.8-8 mm long). Ardisia verdise pala has longer oblong calyx lobes
(3.4-3.8 mm long) and longer corolla lobes (8-8.4 mm long).
la. Ardisia nigrescens Oerst. subsp. nigrescens (Fig. 1).
Shrub or small tree |.5-5 m tall. Branchlet slender, terete, 1-3 mm in diam., with
dense, long hirsute simple reddish hairs throughout. Leaves: with leaf blades
membranous, elliptic, 3.5-11.2 cm long, 1.5-5.2 cm wide, apically abruptly
acuminate, the acumen 0.8-1.4 cm long, basally obtuse, midrib impressed above,
prominently raised below, the secondary veins 15-24 pairs, prominulous above,
prominently raised below, inconspicuously to conspicuously, often prominently
punctate and punctate-lineate, sparsely hispid above and below, often densely
hispid along the midrib above, densely hirsute along the midrib below, the mar-
gins crenate to serrate, flat; petic les slender, canaliculate, 0.5-1.lcm long, densely
hispid along above, densely hirsute below. Inflorescence terminal, rarely lateral,
erect, pinnate paniculate, 1.5-5 cm long, 2-5.5 cm wide, shorter than the leaves,
the rachis with dense, long hirsute simple reddish hairs, the branches 3-7 flow-
ered corymbs; peduncles nearly absent to | cm long, with dense, long hirsute
simple reddish hairs; inflorescence branch bracts caducous, membranous, ob-
long to oblanceolate, 2-2.5cm long, 0.1-0.4 cm wide, apically acute, basally acute
on sessile or short petioles, the veins prominent, conspicuously and prominently
punctate and punctate-lineate, with scattered, long hirsute, simple reddish hairs,
the margins entire; floral bracts similar to the inflorescence bracts but, 0.3-0.8
mm long, 0.1-0.2 mm wide; pedicels slender, terete, 0.7-1.4. cm long, conspicu-
ously and prominently punctate and punctate-lineate, with dense, long hir-
sute simple reddish hairs. Flowers 5-6-merous, white to pink; calyx lobes mem-
branous, ovate to narrow ovate, 1.9-2.1 mm long, L1-1.3mm wide, apically acute,
conspicuously and prominently punctate and punctate-lineate, sparsely hir-
sute outside, granular-glandules within near the base, the margins entire,
densely hirsute-ciliolate; corolla membranous, 6.5-10 mm long, the tube 1.2-
2.4mm long, the lobes oblong, 4+.8-8 mm long, 2.2-4.8 mm wide, apically acute
to rounded, conspicuously and prominently punctate and punctate-lineate,
glabrous except with scattered yellow papillae at the base within, the margins
entire, erose; stamens 5-6 mim long, the filaments 1.9-2.8 mm long, the stami-
nal tube 0.5-1 mm long, the apically free portions abruptly constricted, 1.3-2.]
mm long, epunctate, with scattered yellow papillae near the base, the anthers
ree, lanceolate, 3.5-4.4 mm long, 0.9-L.1 mm wide, apically apiculate, basally
subcordate, the connective inconspicuously punctate; pistil 5-5.5 mm long, gla-
brous, inconspicuously punctate, the ovary ovate, 0.8-1.1 mm long, the style 4-
—y
‘ # -
Es
e Pow a eae
a pall Pe Be wie
ee > E
7
fi
os
MF
#,
*
oa
inst,
pa 2.)
Seca
=
rete Loe
eno s aa
aah:
y
ot
7 area a
~
———
ae CO
Fic. 1. Ard.
subsp. nigrescens A. Fl fant h RB Detail of al
P.Hg
men, showing adaxial surface. E. Fruit.A—B, Ed from lectotype, F. Liet 28B (C).C&D drawn from H. Herndndez
&R. Torres 873 (MOQ).
ial leaf surf C. Flower. D. Sta-
1429
1430 BRIT.ORG/SIDA 20
=
4)
4.4 mm long, stylipodic, inconspicuously punctate and punctate-lineate, the
ovules 5-7. Fruit globose to oblong, 6.9-7.3 mm long, 6.2-6.6 mm in diam., in-
conspic uously punctate and punctate-lineate.
Distribution.—Ardisia nigrescens subsp. nigrescens is known from Mexico
(Puebla, San Luis Potosi, Oaxaca, Veracruz, Chiapas and Tabasco) and in Nica-
ragua Jinotega and Zelaya). It grows at 50-1000 m elevation.
Ecology and conservation status.—Subsp. nigrescens occurs in primary de-
ciduous and evergreen, lower montane rain forests. This subspecies does not
appear to be threatened.
Etymology.—The epithet is based on the latin ‘niger’ or ‘nigrescens’ mean-
ing ‘becoming black’ and refers to the black hispid trichomes.
Ardisia nigrescens subsp. nigrescens is most closely related to subsp.
donnellsmithti because they have similar reddish, hirsute trichomes and ovate
calyx lobes. However, subsp. nigrescens has crenulate or serrate leaf margins
and shorter calyx lobes (1.9-2.1 mm long), whereas subsp. donnellsmithii has
mostly entire to slightly crenulate or serrate leaf margins and longer calyx lobe
(2.2-2.7 mm long).
a
Specimens examined: MEXICO. Chiapas: Mpio. Ocosingo, Laguna Ocotal Grande, 3,300 [t [1,006 ml],
14 Apr 1967 (fr), D. Breedlove 15702 (F, LL); Mpio. Palenque, 6-12 km S of Palenque on the road to
Ocosingo, 300 m, 22 Feb 1972 (fr), D. Breedlove 24232 (F LL, MO, NY); Mpio. La Libertad, 15-20 km
towards Chancala on road to Se a rom the Palenque-Ocosingo road, 280 m, 4 Jan 1981 (ID, D.
Breedlove 49112(GH, MO, NY, TEX); Mpio. La Libertad, 10 . towards Chancala on mac to Bonompak
from the Palenque- Ora ite 280 m, 16 Jan 1982 (ED, D. Breedlove & F Almeda 57370 (DUKE, LL,
MO, NY, TEX): Mpio. Ocosingo, near Laguna Ocotal Gri aoe ca 25-30 kim SE of Monier erro) Libano,
ca. 950 m, 26 Jul 1954 (fD, R. Dressler 1507 (GH); Mpio. Palenque, a lo largo del Rio Nututtin, ca. | km
‘TG, LL, MEXU,
Mpio. San Cristobal de Las Casas, Santa Cruz en San Filipe, without elev, 15 Nov 1986 ([L), A
vat [& M. Concepcion Mtz. de Lopez 9499 (GH, MO, NY, TEX). Oaxaca: Ejido Roberto Colorado,
near Tuxtepec, without elev, 9 Dec 1943 (fl-bud), E. Hernandez X.102(LL); District Tuxtepec, Chiltepec
and vicinity, ca. 20 m, Jul 1940-Feb 1941 (ster), G. Martinez-Calderén 209 (LL); ca. 200 m, 20 Nov
1941 (fr), G. Martinez-Calderén 828 (GH). Puebla: as Ajenjelre, without elev, Aug 1951 (fr), D.
Ramirez C.s.n.(MEXU);, Mpio. Hueytamalco, Paxta 1, 4 Nov 1983 (i), f: Ventura A. 20767 (MEXU,
MO). San Luis Potosi: Mpio. San Antonio, Lejem, w ae elev, | May 1979 ([r), J. Alcorn 2937 (TEX-2
& A. Lundell 7130 (LL).
Tabasco: Mpio. Tacatalpa, Cerro del Madrigal, km 7 de la E Tacotalpa hacia Tapijulapa, without elev,
: on 1983 aad G ssa etal. 3935 (TEX-2 sheets), S side of Mex. Hwy. 186, 2.2 mi W of Tabasco
ni EF of Macuspana, without elev, 23 Dec 1973 (ED, A. Reznicek etal. 187 (MICH).
Vepacnite: 34 kin & aes uca, hacia Tuxpan, 280 m, 12 Nov 1970 (fD, F Chiang 301 (F GH); 20 km de
Tepetzintla, hacia Tantoyuca, without elev. 18 Mar 1971 (fr), F Chiang 377 (K GH, MEXU); 20 km al N
de Martinez de La Torre, without elev, 2 May 1963 (fr), A. Gomez P7885 (LL); Jecaltepec, without elev,
Mar L841 (fr), k Liebmann 28A (C “lectoparaty pe), Mpio, Puente Nacional, Barranca de Palmillas, 2
km al SE de dicha poblacion, 19° 12'N, 096° 45’ W, 450 m, 24 Jul 1985 (fr), M. Medina A. & F Vazquez
B. 379 (F), Mpio. Misantla, Cerro Fspaldilla, an old isolated volcanic plug ca. 5 km N of Miscantla, 19°
58' N, 096° 50' W, +00 m, 14 Jun 1986 (fr), G. Schatz etal. II95 (NY, US); Without locality, without elev,
without date (Ir), L. Schnée & P Maury s.n.(P-2 sheets); 6 mi N of Tuxpan on Hwy. 165, without elev,
2 Jan 1959 ({1), B. Thompson & B. Fields 379 (TEX), Mpio. Martinez de la Torre, El Mirador, 100 m, 16
rio sa la carretera, without elev, 14 Jan 1985 ({), H. Herndndez & R. Torres 813
Mc
sheets), Mpio. Tamazunchale, Tan ae 200 m, Jul 1937 (fr), C. Lundel
1431
Nov 1976 (fl, fr), E Ventura A. 13621 (MEXU, MO). Without further locality in Mexico: Without local-
ity, without elev., 1865-1866 (fr), M. Hahn 21 (kK); San Pablo, without elev, Jun 1841 (fr), F Liebmann
15341 (F); Without locality, without elev., 1841-1842 (fr), W. Karwinsky 450 (H); 1787-1804 (£D, M.
Sessé et al. 679 (F. F neg. no. 47119); 730 (F F neg, no.47121). NICARAGUA. Jinotega: Cerro San Pedro,
Comarca Kilambe, 13’ 36' N, 085° 38-39 W, 820 m, 21 Jul 1980 (f{1), J. Sandino 181 (MO). Zelaya: 20 kin
NW de Alamicamba, 13° 36' N, 084° 22' W,50 m, 16 Apr 1971 (fr), E. Little 25392 (MO, US).
1b. Ardisia nigrescens Oerst. subsp. donnellsmithii (Mez) Ricketson & Pipoly,
mb. et stat. nov. (Fig. 2). Ardisia donnellsmithii Mez, Bull. Herb. Boiss. ser. I,
3:235. 1903. Ardisia nigrescens Oerst. var. donnellsmithii (Mez) Lundell, Wrightia
3:99, 1964. Oerstedianthus donnellsmithii (Mez) Lundell, Phytologia 48:141. 1981.
Ardisia multilineata Mez, Repert. Spec. Nov. Regni. Veg. 16:311. 1920, nom. superfl.
Type. GUATEMALA. ALTA VERAPAZ: Cubilquitz, 350 m, Sep 1900 (fl), H. von
Ttirckheim 7766 (HOLOTYPE: B-destroyed; LECTOTYPE: here designated, US;
ISOLECTOTYPES: GH, LL, F neg. no. 55603).
Shrub or small tree 0.9-6-(12) m tall, 2.5-7.5 cm in diam. Branchlets slender,
terete, 2-5 mm in diam., with dense, long hirsute simple reddish hairs through-
out. Leaves: with leaf blades membranous to chartaceous, ovate or elliptic to
oblong, 5-18.2 cm long, 2.1-8.2 cm wide, apically abruptly acuminate, the acu-
men 1.2-2.1 cm long, basally obtuse, the midrib impressed above, prominently
raised below, the secondary veins 15-24 pairs, prominulous above, prominently
ly to conspicuously, often prominently punctate
raised below, in onspicuous p
and punctate-lineate, sparsely hispid above and below, often densely hispid
along the midrib above, densely hirsute along the midrib below, the margins
mostly entire or slightly crenate to slightly serrate, flat; petioles slender,
canaliculate, 0.6-1.6 cm long, densely hispid along above, densely hirsute be-
low. Inflorescences terminal, rarely lateral, erect, pinnately paniculate, 3-6 cm
long, 4-8 cm wide, shorter than the leaves, the rachis with dense, long hirsute,
simple reddish hairs, the branches 14-18 flowered corymbs; peduncles nearly
absent to 1.2 cm long, with dense, long hirsute, simple reddish hairs; inflores-
cence branch bracts caducous, membranous, oblong to oblanceolate, 0.4-2.5
cm long, 0.1-1.4 cm wide, apically acute, basally acute on sessile or short peti-
oles, the veins prominent, conspicuously and prominently punctate and punc-
tate-lineate, with scattered, long hirsute, simple reddish hairs, the margins en-
tire; floral bracts apparently absent, with no apparent scars; pedicels slender,
terete, 0.7-1.3cm long, conspicuously and prominently punctate and punctate-
lineate, with dense, long hirsute, simple reddish hairs. Flowers 5-merous, white
to pink; calyx lobes membranous, ovate, 2.2-2.7 mm long, 1.2-1.5 mm wide,
apically acute, conspicuously and prominently punctate and punctate-lineate,
sparsely hirsute outside, granular-glandules within near the base, the margins
entire, densely hirsute-ciliolate; corolla membranous, 7.8-8.7 mm long, the tube
1.6-2 mm long, the lobes oblong, 6.3-7.5 mm long, 2.8-3.3 mm wide, apically
acute to rounded, conspicuously and prominently punctate and punctate-
lineate, glabrous except with scattered yellow papillae at the base within, the
1432 BRIT.ORG/SIDA 20(4)
hth
wOSTSs ie ere
haat 1A, ae 2 SS:
(at wed:
iz sic om
en
7
pe TAS
a
—
CR
1G. 2. Ardisia ni 0 subsp al Ismithii (Mez) Ri wering branch. B. Detail of abaxial
E. Fruit. A—-D drawn from lectotype, H. von Tiirckheim 7766
Fy
leaf surface. C Flo ower. D. Sta h
(US). E drawn from G. Davidse & A. Brant 32362 (MO).
1433
margins entire, erose; stamens 4.6-5.5 mm long, the filaments 15-2 mm long,
the staminal tube 0.5-0.7 mm long, the apically free portions abruptly con-
stricted, 1-1.5mm long, epunctate, with scattered yellow papillae near the base,
the anthers free, lanceolate, 3.3-4.5 mm long, 0.8-11 mm wide, apically apicu-
late, basally subcordate, the connective inconspicuously punctate; pistil 5.4-
6.4 mm long, glabrous, inconspicuously punctate, the ovary ovate, 0.8-L2 mm
long, the style 4-4.7 mm long, inconspicuously punctate and punctate-lineate,
the ovules 12-15. Fruit globose, 5.5-7.2 mm long, 6.2-6.6 mm in diam., incon-
spicuously punctate and punctate-lineate.
Distribution.—Ardisia nigrescens subsp. donnellsmithii is known from
Mexico (Chiapas), Belize (Stann Creek and Toledo) and Guatemala (Alta Verapaz,
Huehuetenango and Petén). It grows from 50-950 m.
Ecology and conservation status. —Subsp. donnellsmithii occurs in lower
montane rain forest.
Etymology.-This taxon was dedicated to John Donnell Smith (1829-1928), who
collected extensively in Guatemala and Central America.
In Mez’s (1903) original description he lists two collections by H. von
Turckheim numbers 7766 and 7918, both in the Mez Herbarium (B). These origi-
nal collections were destroyed in 1943 during WWII, necessitating the need to
select a lectotype. Although, C.L. Lundell labeled the US sheet of H. von
Tiirckheim 7766 as the “Holotype,” he failed to actually publish a lectotypi-
fication of the collection. However, from the available material, the US sheet is
the best material, and we here select it as the lectotype for this taxon. The only
known sheet of H. von Tiirckheim 7918 is also at US but is missing the inflores-
cence and has only a few loose fruits. Unfortunately, Mez (1920) re-described
this same material as Ardisia multilineata using the same H. von Ttirckheim
collections, thus making this name superfluous.
Specimens examined. MEXICO. Chiapas: Mpio. ee ace 46 km N of Ocozocoautla on road to
Mal Paso, 700 m, 8 Oct 1974 (fr), D. Breedlove 38709 (D 2 sheets); Mpio. Palenque, 50 km SW of
Palenque on road to Ocosingo near Colonia Uieule Galvan, ae m, 9 Nov ee (fr), a pice alie
47348 (CAS): Mpio. La Libertad, 15 201
Ocosingo road, 280 m, 4 Jan 1981 (fr), D. Breedlove 49160 (LL, ae ue Oc osingo, on Bro near
lake at Naja, 300 m, 3 Mar 1981 (fr), D. Breedlove 49970 (LL, MO, NY); Mpio. Chilon, near and along
road to Pozo Cuevas above Agua Azul, 600 m, 14 Jan 1982 (fr), D. ae & F Almeda 57228 (LL,
MO); Mpio., La Independencia, ridges 45-50 km E of Lagos de Montebello National Park on road to
Ixcan from Santa Elena, 760 m, 22 Jan 1982 (f), D. Breedlove 57759 (CAS); Mpio. Ocosingo, S of Santo
Domingo on road to Bonampak and Echeverria from Chancala, +55 m, 25 Jan 1982 (fr), D. Breedlove
& F Almeda 57927 (MO); Mpio. Ocosingo, near Laguna Ocotal Grande, ca. 25-30 km SE of Monte
i: Libano, ca. 950 m, 10 Aug 1954 (1D, R. Dressler 1637 (GH, US); Rio Salinas, without elev. 8 Feb
4 (fD), C. Lundell 17823 (LL); Mpio. Ocosingo, Bonampak, 350 m, 12 Jun 1984 (fr), E. Martinez S.
ss (MO); 6537 (MO); Mpio., 4 km W of Crucero Corozal along Palenque road, Boca Lacantum, 180
m, 10 Aug 1984 (fr), E. Martinez S. 6901 (MEXU); 19 Sep 1984 (fl), E. Martinez S. 7622 (MO, TEX);
Mpio. Ocosingo, Sierra la Cojolite, 8 km W of Crucero Corozal 7 km E of Bonam pak, 540 m, 20 Sep
1984 (fl), E. Martinez S. 7658 (MEXU); Mpio. Ocosingo, 10 km SE of Crucero Corozal, road to Boca
1434 BRIT.ORG/SIDA 20(4)
Lacantum, 200 m, 18 Aug 1984 (ED, E. Martinez S. 7411 (FTG, MO); Mpio. Ocosingo, near Marques of
Comillas, 6 km SE of Ejido Benemérito de las Americas, toward Flor de Cacao, 160 m, 8 Oct 1984 (fr),
FE. Martinez S. 8059 (MO); Mpio. Ocosingo, 6 km S of Campamento COFOLASA, and 24 km SE of
Crucero Corozal, Palenque-Boca Lacantum road, 200 m, 17 Oct 1984 (f1), E. Martinez S. 8427 (MO):
Mpio. Ocosingo, 5 km S of Crucero Corozal, on road to Boca Lacantum, 250 m, 22 Oct 1984 (f1), E.
Martinez S.& G. Aguilar 8630 (MO); Mpio. Ocosingo, Boca Lacantum, 120 m, 28 Oct 1984 (fr), E.
Martinez S. & G. Aguilar 8790 (MO); Mpio. Ocosingo, ee eine COFOLASA 24 km SE of El
Crucero Corozal, on Palenque-Boca Lacantum road, 220 m,7 Dec 1984 (fl), E. Martinez S. 9318 (MO):
Mpio. Ocosingo, 10 km S of Ejido Ber rito de Las Americas, ro 7 o Flor de Cacao, Zona Marquez
de Comillas, 120 m, 9 Dec 1984 (FD, E. Mariiners 9361 (MO); Mpio. Ocosingo, Boca Lacantum, anu
Rio Lacantum, 120m, 10 Dec 1984 (fr), E. Martinez S. 9581(MEXU ), Mpio. Ocosingo, 2 km W of Cruc
Corozal, Palenque-Boca Lacantum road, 180 m, 13 Feb 1985 (fr), E. Martinez S. 10106 (MO, TEX): ieee
(MO); 22 Feb 1985 (fr), E. oe S. 11075 (MEXU); Mpio. Ocosingo, Crucero Corozal, 180 m, 5 Apr
1985 (fr), E. Martinez S. 11531 (MEXU); Mpio. Ocosingo, 19 km NW of Crucero Corozal, road to Palenque,
600 m, 8 Sep 1985 (fl), EF. Martinez S. 13454 (MO, TEX); Mpio. Ocosingo, a 3 km NW of Vertice del
Chixoy road a Boca Lacantum, 120 m, 11 Sep 1985 (fr), E. Martinez S. 13616 (MO); Mpio. Ocosingo, 15
km NW of Boca Lacantum, road to Palenque, 220 m, 13 Sep 1985 (fr), E. Martinez S. & G. Aguilar
13707 (MO), Mpio. Ocosingo, Neuvo Veracruz, 33 km W of Vertice del Rio Chixoy, road to Chajul,
ee Comillas, 130 m, 10 Jan 1986 (fr), E. Martinez S. 15982 (MEXU): Mpio. Ocosingo, 2 km N of
Naja, road to here 900 m, 24 May 1987 (fr), E. Martinez S. 21333 (MO); Mpio. Ocosingo, 2 km S of
Naja, Lo" 58'N 1° 36’ W, 940 m, 24 Sep 1988 (f1, fr), W. Stevens & E. Martinez S. 25849 (FTG, MO):
ee Oxesin near Chamisal, 760 m, 3 Dec 1976 (£1), P. Valdivia Q. 2407 (LL, MO). BELIZE. Stann
Cre Aiddlesex, without elev. 25 May 1939 ({r), P. Gentle 2798 (A, FE MO, NY, US); 200 ft {61 m], 24 Jul
ne W. Schipp 287 (A, F. GH, MO, NY); 23 Nov 1929 (f1), W. Schipp 478 (A, BM, E.G, GH, K, MO, NY,
TEX), 1928 (fr), N. Stevenson V (US). Toledo: Maya Mountains, canyon along Bladen Branch from Ri-
chardson Creek to Quebrada de Oro, 16° 31-33' N, 088° 46-49 W, 100-200 m, 12 Mar 1987 (fr), G.
Davidse & A. Brant 32362 (BRH, F FTG, LL, MO): Maya Mountains, Bladen Nature Reserve, Ek Xux
canyon, 1.8 airline km NW of the Ek Xux archeological site, 16° 30'57" N, 088" 55' 10" W, 320 m, 18 May
1996 (fr), G. Davidse 36051 (BRH, MO, SEL); Balo Camp, upper reach of Golden Stream, without elev,
12 Apr 1944 Ur), P Gentle 4522 (LL-2 sheets); Jacinto Creek, Rio Grande, without elev. 18 Oct 1944
(f), P Gentle 4895 (LL-2 sheets); Temash River, without elev, 27 Feb 1945 (fr), P Gentle 52. “ (LL-
sheets, PH, TEX); Ridge beyond Columbia, without elev, 25 Feb 1947 (fr), P Gentle 6177 (LI Deets
TEX): Sat Road, 14 Miles, San Antonio-Punta Gorda Road, without elev. 5 May 1949 (fn), p Gentle
6729 (LL-3 sheets); Cero, without elev, 15 Feb 1950 (fr), P Gentle 6980 (LL-2 an TEX); Between
Orange Point and Moho River, een elev, 7 Apr 1952 (fr), P Gentle 7639 (F, LL-2 sheets, US); Camp
3,6-8 km SE of Union ei trail from Camp 3 toward the Jimmy cut trail, e 3 14" N, 089° 04" 37"
W, 605-700 m, 16 Feb 1997 (fr), I’ Hawkins 1443A (MO): Toledo, without elev, 10 Nov 1906 (fl), M. Peck
9576(GH, K, NY), Forest Home, 200 ft [61 m], 2 Nov 1933 (fl), W. Schipp 482 (EF MO). GUATEMALA. Alta
Verapaz: Sebol, ca. 6 km from village near Rubelquiche, without elev, 13 Apr 1964 (fr), E. Contreras
4254 (LL-2 sheets, TEX); Sebol, on old Petén road, without elev, 14 Apr 1964 (fr), E. Contreras 4291
(GH, LL-2 sheets, TEX): Sebol, without elev, May 1964 (fr), E. Contreras 4667 (LL): Sebol and vicinity,
without elev. Jul 1964 (fD, E. Contreras 5414 (LL-3 sheets); Near Finca Sepacuite, near Cajabon, with-
out elev, 26 Apr 1902 (fr), O. Cook & R. Griggs 767 (US); Semacoch, without elev, 8 Mar 1905 (fr), G.
Goll 261 (US), Xalavé, 1,500 ft [457 m], 25 Jun 1920 ({D, H. Johnson 228 (A, F. US): Forest in valley,
“pantano,” 2.5 mi W of Cubilguitz, 250-300 m, 28 Feb 1942 (fr), J. Steyermark 44284 (F); 44322 (F):
1.5-2 mi Sof Cubilguitz, 300-350 m, | Mar 1942 (fr), }. Steyer else al US). S side of Cerro Chinaja,
between Sachaj and Sacacac, 150-180 m, 20 Mar 1942 (fr), J. S ‘mark 45161 (F); Cubilquitz, 35
Dec 1900 (fr), H. von Tuerckheim 7918 (US-isolectoty pe); Aug i904 (ED, H. von Tuerckheim e707 578
(US-3 sheets); Oct 1906 (fl), H. von Tuerckheim 11163 (BR-2 sheets, F G-3 sheets, LL, MO).
Huehuetenango: Between Ixcan & Rio Ixcan, Sierra de los ee econ 150-200 m, ee 1942
1435
(fr), J. pect mark 49253 (F). Petén: La Cumbre, without elev. 30 Sep 1966 (fr), E. Contreras 6290 (LL-2
sheets, TEX); La Cumbre, bordering Rio Chacte, E of km 135, without elev, 19 Jul 1976 (fr), FE. Contreras
EC. Lundell 20069 (LL-2 sheets); Sayaxche, oh airport, without elev., 9 Feb 1964 (fr), C. Lu . ell
17923 (LL); Sayaxche, on Arroyo Santa Cruz trail, without elev. Ll Feb 1964 D, C. Lundell 17978 (L
Sayaxche, in canada 2 km NE of village on ae Cruz trail, without elev. Feb 1964 (fr), C. Lu a
18039 (LL); (£1), C. Lundell 18040 (LL-2 sheets, Nas Laguna Petexbatun, forest bordering the la
without elev, 2 Apr 1964 (fr), C. Lundell 18133 (LL-2 sheets), Between Finca Yalpemech & Chinaja,
50-100 m, 28 Mar 1942 (fr), J. Steyermark 45427 (FU 1S) W & NW of Chinaja, between Rio Chinaja &
6 mi W of Rio San Raman, 50-70 m, 29 Mar 1942 (f1), J. Steyermark 45493 (F).
Ke,
2. Ardisia verdisepala Ricketson & Pipoly, sp. nov. (Fig. 3). Type. MEXICO. CHIAPAS:
Mpio. Ocosingo, 15 km SW of Ocosingo on road to San Cristobal Las Casas, 1,372
m, 1 Sep 1981 (£1), D. Breedlove 52591 (HOLOTYPE: MO: IsOTYPES: CAS, GH, LL, NY).
] f]
Propter longi-hirsutos A. nigrescens valde arcte affinis sed
ab ea lobis srivcuis oblongisque (non ovatisque) 3.4-3.8 (nec 1.9-2.7) mm longis praeclare distat.
Shrub to small tree |.2-6.]1 m tall. Branchlets slender, terete, 1-2.5 mm in diam.,
with dense, long hirsute simple reddish hairs throughout. Leaves: with leat
blades, membranous, elliptic to oblong, 3.3-10.2 cm long, 1.5-4.8 cm wide,
apically abruptly acuminate, the acumen 0.5-1.6 cm long, basally acute to ob-
tuse, the midrib impressed above, prominently raised below, the secondary veins
12-19 pairs, prominulous above, prominently raised below, conspicuously and
prominently punctate and punctate-lineate, sparsely hispid above and below,
often densely hispid along the midrib above, densely hirsute along the midrib
below, the margins serrate, flat; petioles slender, canaliculate, 3.5-8 mm long,
with dense, long hispid reddish hairs above and below. Inflorescences terminal,
rarely lateral, erect, pinnately paniculate, 3-6 cm long, 3-6.5 cm wide, shorter
than the leaves, the rachis with dense, long hirsute, simple reddish hairs, the
branches 5-7 flowered corymbs; peduncles nearly absent to 1.5 cm long, with
dense, long hirsute, simple reddish hairs; inflorescence branch bracts caducous,
membranous, oblong to obovate or oblanceolate, 7.5-15.5 mm long, 3.5-6 mm
wide, apically acute, basally acute on sessile or short petioles, the veins promi-
nent, conspicuously and prominently p and punctate-lineate, with scat-
tered to dense, long hirsute, simple reddish hairs, the margins entire, floral bracts
apparently absent above, with no apparent scars, below similar to the inflores-
cence branch bracts but, early caducous, usually linear, 3.8-6.6 mm long, 1.3-
1.5mm wide; pedicels slender, terete, 1.5-2.5 cm long, conspicuously and promi-
nently punctate and punctate-lineate, with dense, long hirsute, simple reddish
hairs. Flowers 5-merous, white to pink or reddish; calyx lobes membranous,
narrowly oblong, 3.4-3.8 mm long, 1.2-1.7 mm wide, apically acute to rounded,
conspicuously and prominently punctate and punctate-lineate, sparsely hir-
sute outside, glabrous within, the margins entire, scattered hirsute-ciliolate;
corolla membranous, 9.2-9.8 mm long, the tube 1.2-1.4 mm long, the lobes ob-
long, 8-8.4 mm long, 3.2-5.4 mm wide, apically acute to rounded, conspicu-
ously and prominently punctate and punctate-lineate, glabrous except with
1436 BRIT.ORG/SIDA 20(4)
man
<
ty
tea
a eee
TI wg eee
4 hh ical
S eX
A ‘
Cue A
L
Lip
& Pinoly A. Fl inab h. B. Detail of adaxial leaf f Detail of abaxial
eee
C
F. Fruit. A-D drawn from holotype, D. Breedlove 52591 (MO).
}
leaf surface. D. Flower. E. Stamen, showi laxial f.
F drawn from A. Méndez Ton 5667 (MO)
1437
scattered yellow papillae at the base within, the margins entire, erose, stamens
5-5.8 mm long, the filaments 1.7-2.2 mm long, the staminal tube 0.5-0.7 mm
long, the apically free portions as wide as the anthers, 1.4-1.7 mm long,
epunctate, with scattered yellow papillae near the base, the anthers free, lan-
ceolate, 3.8-4.3 mm long, 1-1.2 mm wide, apically apiculate, basally hastate,
the connective inconspicuously punctate; pistil 5.5-6.8 mm long, glabrous, in-
conspicuously punctate, the ovary ovate, 1-15 mm long, the style 4.5-5.6 mm
long, inconspicuously punctate and punctate-lineate, the ovules 14-17. Fruit
globose, 5.4-7 mm long, inconspicuously punctate and punctate-lineate.
Distribution —Ardisia verdise pala is endemic to the area around Ocosingo,
Chiapas, Mexico. It grows from 550-1,372 m.
Ecology and conservation status.—Ardisia verdisepala occurs in evergreen pine-
oak, lower montane and montane rain forests. Although, no specimens have
been collected since the 1980s, it seems to be locally common and not pres-
ently threatened.
Etymology—The epithat verdisepala comes from the greenish color of the se-
pals, which is evident because of the reduced number of punctations on the
calyx lobes.
Ardisia verdisepala has affinities to A. nigrescens because of the similar
vestiture, but is easily separated by the narrowly oblong calyx lobes verses the
ovate lobes of A. nigrescens. Ardisia verdisepala also has affinities to A.
tuerkheimii, because both have narrow calyx lobes; however, A. verdisepala is
immediately separated from the virtually glabrous A. tuerckheimii, by the well-
developed vestiture.
PARATYPES. MEXICO, Chiapas: Mpio. Ocosingo, 6-8 km N of Ocosingo along road to Bachajon, 900 m,
9 Nov 1971 (ED, D. Breedlove & A. Smith 22120 (MO), 22121(LL, MO, NY); 24 Sep 1972 (£1), D. Breedlove
7927 (LL, MO, NY); Mpio. Ocosingo, adjacent to Laguna Ocotal Grande, 800 m, 6 Feb 1973 (fr), D.
Breedlove 33006 (LL, MO); Mpio. Bachajon, 3 km N Bachajon-Ocosingo road on road to Palenque, 1,170
m, 13 Jan 1981 (fr), D. Breedlove 49445 (CAS, DUKE, LL, MO, NY. TEX); Mpio. Ocosingo, 70 km SW of
Palenque on road to Ocosingo along the Jol Uk'um, 550 m, 14 Jan 1981 Cf), D. Breedlove & B. Keller
49485 (DUKE, LL, MO, NY, TEX); Mpio. Bachajon, 3 kim No are area road at Temo on
road to Palenque, 1,170 m, 31 Aug 1981 (fD, D. Breedlove 52487 (CAS, DUKE, F, LL, MO, NY); Mpio.
Ocosingo, 80 km SW of Palenque on road to Ocosingo along the Jol Uk’um, 760 m, oe 1982 (f1, D.
Breedlove & E Almeda 57210 (CAS, GH, LL, MO, NY): Mpio. Yajalon, K’ak’ate’el, 1,100 m, 15 Jul 1982
(fr), A. Méndez G.4445(MEXU, MO): Mpio. Oxchuc, La Cascada de Mesbija, 1,300 m, 29 Nov 1982 (fD,
A. Méndez T. 5097 (MEXU, MO); Mpio. Yalalon, Rancho San Luis, 1,000 m, 10 Feb 1983 (fr), A. Méndez
T. 5469 (ASU, MEXU, MO, TEX); Mpio. Yajalon, Banco de grava, 1,000 m, 10 Mar 1983 (fr), A. Méndez T.
5640 (MEXU, MO, TEX); Mpio. Yajalon, Rancho San Luis, 900 m, 15 Mar 1983 (fr), A. Méndez T. 5667
(MEXU, MO, TEX); 15 Apr 1983 (fr), A. Méndez T. 5837 (MEXU, MO); 15 Apr 1983 (fr), A. Méndez T.
5842 (MEXU, MO); Mpio. Yajalon, Arroyo en la Graba, 800 m, 16 May 1983 (fr), A. Méndez T. 6052
(MEXU, MO, TEX); Mpio. Sabanilla, Finca Morelia, 650 m, 5 Jun 1983 (fr), A. Méndez T. 6112 (MEXU,
MO, TEX); Mpio. Yajal6n, El Azufre, 700 m, 20 Jun 1983 (fr), A. Méndez T6211 (MEXU, MO, TEX).
3. Ardisia tuerckheimii Donn. Sm., Bot. Gaz. 13:74. 1888. (Fig. 4). Oerstedianthus
tuerckheimii (Donn. Sm.) Lundell, Phytologia 48:142. 1981. Type. GUATEMALA.
1438 BRIT.ORG/SIDA 20(4)
ALTA VERAPAZ: Pansamala, mountain forests, 3,800 ft [1,158 m], Aug 1886 (fD, H.
von Tiirckheim 1035 (LECTOTYPE: US sheet no. 40099, here designated, US neg. no.
2390; ISOLECTOTYPES: F (F neg. no. 68257), G-4 sheets, GH-2 sheets, K, LL (F neg. no.
55673), M, MO (LL neg. no. 71-116), NY-2 sheets, PH, US sheet no. 1330942 (US neg.
no. 2391)).
Ardisia mitchellae ILM. Johnst., Contr. Gray Herb. 81:96. 1928. Oerstedianthus mitchellae (.M.
Johnst.) Lundell, Phytologia 48:142. 1981. Type. HONDURAS. ATLANTIDA: vicinity of Tela, near
a river, 50 ft. [15 ml], 4 Apr 1926 (1), E. Mite nee H; ISOTYPE: US).
Ardisia carlsonae Steyerm.,, Ceiba 4:301, 1955. Oerstedianthus carlsonae (Steyerm.) Lundell,
Phytologia 54:285. 1983. Type. MEXICO. Culapas: Around Los Arcos, Los Lagos, 3 mi NW of
Rancho San José and 34 mi SE of Comitan, 1600 m, 15—20 Apr 1949 (fl), M. Carlson 1744
(HOLOTYPE: F; ISOTYPE; EAP not seen
Ardisia erythrocarpus Lundell, Wrightia 2:59. 1960. Oerstedianthus eryth rocarpus (Lundell)
Lundell, Phytologia 4+8:141. 1981. Type. GUATEMALA. PETEN: San Luis, km 52 of road S of
village, without elev, 10 Jul 1959 (fr), C. Lundell 16267 (HOLOTYPE: LL-TEX (F neg. no. 55604);
ISOTYPES: LL-TEX, TEX, US).
Ardisia brevipes Lundell, Wrightia 3:97. 1964. Oerstedianthus brevipes (Lundell) Lundell,
Phytologia 48:141. 198L Type. MEXICO. VERACRUZ: Suchilapa, without elev, 10 Mar 1930 ([D),
C. Mell 576 (HOLOTYPE: US; ISOTYPE: F (F neg. no. 68136, LL neg. no. 1971-24), LL-TEX (F neg. no.
55610), NY).
Ardisia hirtella Lundell, Wrightia 3:98. 1964. Oerstedianthus hirtella (Lundell) Lundell,
Phytologia 48:141. 1981. Type. BELIZE. EL Cayo: Retiro, without elev. 30 Jun-Aug 1936 CD, C.
Lundell 6302 (HOLOTYPE: LL-TEX (F neg. no. 55615; ISOTYPES: F (F neg. no. 68223), GH-2 sheets,
LL, MEXU, NY-2 sheets, TEX, US).
Ardisia trinitariae Lundell, Wrightia 5:62. 1974. Oerstedianthus trinitariae (Lundell) Lundell,
Phytologia 48:142. 1981. Type. MEXICO. CHIAPAS: Municipio de La Trinitaria, slope at the Lago
de Monte Bello, ), 25 mi E of La Trinitaria, 5,100 ft [1,554 ml], 13 Apr 1965 (fl), D. Breedlove 9721
Races L-TEX, (F neg. no, 55631): IsOTYPES: CAS-DS, F (F neg. no. 68449).
Lundell, Wrightia 5:63. 1974. Oerstedianthus tuxtepecanus (Lundell) Lundell,
paorolegia 48142 1981. TYPE. MEXICO. OAXACA: Rancho “Los Caracoles,” ca. de Tuxtepec,
without elev, Jul 1963 (fr), J. Alcocer y C. Morales s.n. (HOLOTYPE: LL, (F neg. no. 55632).
Ww
A
Shrubs to small trees 0.5-15 m tall, 2.5-25 cm in diam. Branchlets slender, ter-
ete, 1-3 mm in diam.,, very sparsely to densely puberulent of simple trichomes.
Leaves: with leaf blades membranous, elliptic, 2.8-12.5cm long, 0.9-5.8 cm wide,
apically abruptly long acuminate, the acumen 0.5-2.2 cm long, basally obtuse,
the midribimpressed above, prominently raised below, the secondary veins 14-
30 pairs, prominulous above, prominently raised below, inconspicuously to con-
spicuously punctate and punctate-lineate, glabrous to scattered puberulent of
simple trichomes on the upper and lower surface, especially along the midvein,
the margins entire to crenate or serrate, flat; petiole slender, canaliculate, 0.5-
L.l cm long, glabrous. Inflorescences terminal, erect, pinnately paniculate, 1.5-
12 cm long, 1.5-7 cm wide, shorter to slightly longer than the leaves, the rachis
glabrous to densely puberulent of simple trichomes, the branches 4-9-flow-
ered corymbs; peduncles nearly absent to 1 cm long, glabrous; inflorescence
branch bracts caducous, membranous, oblong to obovate or oblanceolate, 3-9
mm long, 1-3 mm wide, apically acute, basally acute on sessile or short peti-
1439
f m.A. Flowering branch.B. D il of abaxial leaf surf C. Flower. D. Stamen, showing
eee E. Fruit. A-D drawn from lectotype, H. von Tiirckheim 1035 (US). E drawn from E. Contreras 6668 (LL).
oles, the veins prominent, conspicuously and prominently punctate and punc-
tate-lineate, glabrous, the margins entire; floral bracts apparently absent in the
upper portion of the inflorescence with no apparent scars; pedicels slender, ter-
ete, 11-18 cm long, conspicuously and prominently punctate and punctate-
lineate, glabrous to densely puberulent with simple trichomes. Flowers 5-
1440 BRIT.ORG/SIDA 20(4)
merous, white to pinkish; calyx lobes membranous, ovate to narrow oblong or
oblanceolate, 1.5-3 mm long, 0.6-2 mm wide, apically acute to rounded, con-
spicuously and prominently punctate and punctate-lineate, glabrous to scat-
tered papillae inside and out, the margins entire, scattered hirsute-ciliolate; co-
rolla membranous, +-10.2 mm long, the tube 0.7-1.9 mm long, the lobes oblong,
4.5-9.1 mm long, 1.6-5 mm wide, apically acute to rounded, conspicuously and
prominently punctate and punctate-lineate, glabrous except rarely witha few
yellow papillae at the base within, the margins entire, erose; stamens 4.7-6.2
mm long, the filaments .8-2.9 mm long, the staminal tube 0.2-0.9 mm long,
the apically free portion 1.7-2.5 mm long, epunctate, with scattered yellow pa-
pillae near the base, the anthers free, lanceolate, 3.2-5.1 mm long, 0.6-L6 mm
wide, apically apiculate, basally subcordate, the connective inconspicuously
punctate; pistil 5.2-8.3 mm long, glabrous, inconspicuously punctate, the ovary
ovate, 0.8-1.5 mm long, the style 4.2-6.7 mm long, inconspicuously punctate
and punctate-lineate, the ovules 8-15. Fruit globose, 5-6.4 mm long, inconspicu-
ously punctate and punctate-lineate.
Distribution.—Ardisia tuerckheimii is known from southern Mexico
(Oaxaca, Veracruz and Chiapas), southern Belize (Cayo and Toledo), northern
Guatemala (Petén, Huehuetenango, Baja Verapaz, Alta Verapaz and Izabal) and
northeastern Honduras (Cortés, Atlantida and Yoro). It grows from 800-1,500 m.
Ecology and conservation status.—Ardisia tuerckheimii occurs in primary
and secondary, pine-oak-Liquidambar, lower ane and montane forests. This
—
species appears to be quite common and not in any immediate danger.
Etymology.—This taxon was named in honor of the German-born collector
Hans von Turckheim (1853-1920), who collected extensively in Guatemala and
Santo Domingo.
In John Donnell Smith’s (1888) original description he clearly based this
taxon on the H. von Ttirckheim 1035 collection, and it is agreed that his types
are housed at the Smithsonian Institution (US). However, three specimens are
—
housed at US with no indication as to which sheet is the holotype, thus necessi-
tating the need to select a lectotype. One of these sheets US no. 1339941 was
photographed by the Field Museum (F neg. no. 55673) and was de-accessioned
and sent asa gilt toC.L. Lundell (LL-TEX). The remaining two US sheets are of
approximately equal value, one labeled as US no. 40099 and the other US as no.
1339942; the latter appears to have an original hand written label. Although
C.L. Lundell labeled the US no. 40099 sheet as the “Type,” he did not formally
lectotypify the collection. Because the US no. 40099 sheet is of slightly better
quality and has a lower accession number, making it more likely that John
Donnell Smith used this material, at least in part, for his original description,
we designate this sheet as the lectotype.
Ardisia tuerckheimii is an extremely variable species, which has led to its
over description. Many of the collections from Mexicoare glabrous or nearly so,
~—
1441
and gradually become more puberulent, as one moves south towards Hondu-
ras. However, because of this gradual gradation from glabrous to densely pu-
berulent no clear justification can be found to recognize any of these proposed
species and all are synonymized here. In fact the types of Ardisia mitchellae
and A. hirtella were originally compared to A. nigrescens (including subsp.
donnellsmithii), and not even compared to A. tuerckheimii, and separated based
mostly on the differences in the indument, which is clearly different. A.
erythrocarpa is unique only for its slightly larger fruits and slightly ciliated
larger sepals. Populations of A. brevipes are unique only for their apparent
subsessile fasciculate inflorescences with shorter peduncles, and with what are
described as “bulbous” anther bases, and involuted petals apically. Populations
of A.carlsonae are unique only for their slightly larger flowers, slightly smaller
leaves and slightly obtuse anther. The type of A. trinitariae has slightly longer
pedicels, slightly larger flowers, “bulbous” anthers bases and slightly oblong
leaves. Finally, A. tuxtepecana differs slightly by its slender habit and smaller
more densely puberulent leaves with slightly longer pedicels. Close examina-
tion of the total variation in this species shows that the slight differences, dis-
played by the synonymied taxa are not discontinous but fall within the range
of variation of A. tuerckheimii.
od
Specimens examined. MEXICO. Chiapas: Mpio. La Trinitaria, slope at the Lago of Monte Bello, 25 mi
E of La Trinitaria, 5,100 ft [1,554 ml], 13 Apr 1965 (fr), D. Breedlove 9733 (F LL); 5,200 ft [1,585 m], 17 Aug
1966 (f1, fr), D. Breedlove 14978 (F LL, US); Mpio. La Trinitaria, forest along the Comitan River at its
sumidero, Lagos de Montebello, 42 km NE of La Trinitaria, 1,300 m, 23 Oct 1971 ED, D. Breedlove & R.
Thorne 21166 (F LL, MO, NY); Mpio. La Trinitaria, ee of Laguna Tzikaw, Monte Bello National
Park, 1,300 m, 16 Nov 1972 (fD, D. Breedlove & R. Dressler 29601 (MO, TEX); Mpio. Las Margaritas, on
the W side of De Miramar E of San ne a m, ll Feb 1973 (fr), D. Breedlove 33302 (MO),
Mpio. Ocozocoautla de Espinosa, 32 km N of Ocozocoautla on road to Mal Paso, 800 m, 6 Oct 1974
(fr), D ee 38200 (DS, LL, MO); Mpio. La Trinitaria, + km E of Laguna Tsiskaw near Dos Lagos,
300 m, 19 Oct 1974 (fD, D. Breedlove 38764 (CAS, MO); Mpio. La Trinitaria, E of Laguna Tziscoa,
Monte Bello National Park, 1,380 m, 18 Dec 1980 (fr), D. Breedlove 48740 (LL, MO); Mpio. La Trinitaria,
Near Cinco L ae Lagos de Montebello National Park, 1.372 m, 5 Oct 1981 (£D, D. Breedlove 53339
(DUKE, FE GH, LL, MO, NY); 13 Oct 1981 (fr), D. Breedlove 53425 (LL); Mpio. La Trinitaria, Cinco Lagu-
nas, Lagos de Montebello National Park, 1,600 m, 24 May 1988 (fr), D. Breedlove 68667 (CAS), A Jatu
del Tigre, 30 km NW Ocozocuantla, without elev, 26 May sr eae it onsale eae Ledisi
Ocosingo, a los lados de la carretera, without elev, 7 Dec 1980 (1), 1356 (MEXU,
O
MO): Mpio. La Trinitaria, slope on the E side of L a) a aw in region of the Lagos ae nee
4.800 ft [1,463 ml], 5 Jul 1967 (fr), A. Shilom T. 2599 (F, LL); Mpio. La Trinitaria, near Parque Nacional
Lagunas de Monte Bello, 40 km E of La Trinitaria, | a m, 26 Oct 1989 ([1), J. Soto et al. 13479 (BM).
Oaxaca: Mpio. Santa Maria Chimale apa, Dto. Juchitan, Los Angeles, before Arroyo Seco, to Santa Maria
Chimalapa, 33 km E of Mezquite, 16° 52’ N, 94° 44’ W, 250 m, 24 Mar 1988 (fr), A. pas etal. Bie
(MEXU, MO); 1715 (MEXU, var 2 km before C Sy Ae along Uxpanapa Road, 12° 0
W, 120 m, 4 Nov 1978 (fr), G. Castillo C. 408 (LL, MEXU); Mpio. Juchitan, 24 km = . Lazaro
Cardenas,Santa Maria Chimalapa Road. o m, : Mar 1982 (fl), R. Cedillo T. & R. Torres 1144 (LL,
MEXU); Mpio. Santa Maria Chimalapa, near Santa Maria, 1655’ N, 094° 40! 30" W, 300 m, 15 Sep 1984
(fr), H. Hernandez G. 433 (LL, MO): Mpio. Santa Maria Chimalapa, Rio Milagro, ca. 3 km W of Santa
Maria, 16° 55’ N, 094° 42’ W, 190 m, 13 Feb 1985 (fr), H. Herndndez G. 842 (LL, MO); Mpio. Santa Maria
1442 BRIT.ORG/SIDA 20(4)
Chimalapa, Arroyo Palomares, ca. 2.5 km N of Santa Maria, 16" 55’ N, 094° 41' W, 250 m, 19 Feb 1985
(fl), H. Herndndez G.896 (LL, MO): Mpio. Santa Maria Chimala pa,ca. 1.5 km NE of Santa Maria through
Paso Tzajinmjeun del Rio del Cortes, at junction of Paso Lagarto, 16° 55' N, 094° 40' 30" W, 250 m, 27
Feb 1985 ({l), H. Herndndez G.935 (LL, MO): Mpio. Santa Maria Chimalapa Arroyo Sangre, ca. 2 kmal
E de Santa Maria, 16° 54 30" N, 094" 40 W, 250 m, 12 Mar 1985 (EL), H. Hernandez G. 985 (LL); Mpio.
Santa Maria Chimalapa, ca. 10 aa W of Santa Maria via Chicosaja, 16°53’ N, 094° 46' W, 250 m, 30 Mar
1985 (1D, H. Herndndez G. oe .. MO); Mpio. Santa Maria Chimalapa, Canada del Rio Negro, at
Arroyo Huapond’s mouth, ca. ve S de Santa Maria, 16° 56'N, 094° 39' 30" W, 300 m, 14 Oct 1985 (fr),
H. Herndndez G. & C. Gonz oer 1637 (FTG), Mpio. Santa Maria Chimalapa, Cerro Azul-Escol os
ca.8 km SW ol Santa Maria, S of road to Lazaro Cardenas, 16°51’ 30" N, 094° 43’ W, 400 m, 19 Oct |
(ir), H. Herndndez G.& C.Gonzdlez L. L681 (FTG), Mpio. Santa sae \ per oe mountain range at
V,50
Paso Jugcuypac del Rio Verde, ca. 8 km N of Santa Maria, 16° 57' 30" N, 09 00 m, 29 Oct 1985
(fr), H. Hernandez G.& C. Gonzalez L. 1769 (FTG), Mpio. Santa cn chal Aro Sangre, ca.
0.2-0.5 km from mouth of Rio Milagro, ca. 3 km SE of Santa Maria. 16 54'N, ( 130" W, 200 m, 28
Dec 1985 (fr), H. Herndndez G. 1994 (FTG, LL); Mpio. Valle Nacional, 2 km W of royo Seco, 17° 54° N,
096° 18' W, 200-300 m, 12 Oct 1992 (£D), J. Meave del Castillo et al. 1480 (MEXU, MO); Mpio Santa a aria
Jacatepec, Dtto. Tuxtepec, Predio El Aguila, W of San Agustin, entering by La Reforma, 28 kn e
Tuxtepec, road to Matias Romero, 17’ 50' N, 096° 06' W, 550 m, 19 Jan 1988 (fr), R. Torres C. en
MO), Mpio. Santa Maria Chimalapa, Dtto. Juchitan, Los Angeles before Arroyo Seco, 23.3 km E of
Lazaro Cardenas, toward Santa Maria Chimalapa, entering from F] Mezquite, 16° 50' N, 094° 45° W.
400 m, 21 Jan 1988 (fr), R. Torres C.& E. Martinez S. 11163 (MEXU, MO): Mpio. Santa Maria Jacatepec,
Dtto, Tuxtepec, Predio del Aguila, at San Agustin, 25 km W of la Reforma, Ayozintepec highway, 17°
50' N, 096: 06' W, without elev. 21 Feb 1988 (fr), R. Torres C.& L. Cortes A. 11500 (MEXU, MO); Mpio.
San Juan Guichicobi, 7 km E of Sarabia, without elev.,6 ~ 1974 (fr), M. Vazquez T.1395(FTG, MEXU,
MO); Mpio. Matias Romero, ca. 22 kim S of Esmeralda, ca 9 km S$ of La Floresta sawmill, hills $ of Rio
Verde, 17° OL N, 094° 49' W, 290 m, 26 Mar 1981 (£1), I. Wendt et al. 3081 (LL-2 sheets, MO): Mpio. Matias
Romero, 7.2 km W of Esmeralda, along La Laguna-Sarabia, then 2.6 km S, 17° 07'N, 094° 49’ W, 200 m,
8 Mar 1982 (£D, 1. Wendt et al. 3623 (LL). Veracruz: Mpio. Catemaco, Lado NE of Lago Catemaco E of
Coyame, ca. 450 m, 27 Oct 1971 (fr), |. Beaman 5186 (ELL): Mpio. San Andrés Tuxtla, Estacién Biologic a
de Los Tuxtlas, ca. 20 km N of Catemaco, “Eblotrolotu,” without elev, ca. 1970 (f1), J. Calzada 227 (F
LL, MEXU), 6 Apr 1972 (ED, J. Calzada 723 (F MEXU); Mpio. San Andrés Tuxtla, Estacion Biologica de
Los Tuxtlas, without elev, 26 Oct 1972 (fr), R. Cedillo T. & J. Calzada 48 CF, LL, MEXU); 500 m, 8 Apr
1972 £1), R. Cedillo T. 164(E LL, NY-2 sheets); Mpio. Catemaco, Estacion Biologica Tropical Los Tuxtlas,
150 m, 16 May 1981 (£D, R. Cedillo T. & D. Lorence 702 (FLL); Mpio. San Andrés Tuxtla, Estacién
Biologica de Los Tuxtlas, UNAM road to La Laguna Escondida, 18° 35' N, 095° 04 W, without elev, 29
Mar 1979 £1, M. Dillon et al. 1831 (ER MO, NY, TEX); Mpio. Hidalgotitlan, kms 0-2 del camino Plan de
Arroyos-Alvaro Obregon, 17° 15' N, 094° 40 W, 130-150 m, 14 Apr 1974 (fr), B. Dorantes 2802 (MEXU,
MO); Mpio. Hidalgotitlan, Cedillo-La Escuadra road, 150 m, 30 Sep 1974 (fr), B. Dorantes 3576 (MEXU,
MO); Mpio. Hidalgotitlan, road from Hermanos Cedillo to la Laguna, 17° 15 N, 094° 40' W, 140 m, 9
Oct 1974 (fr), B. Dorantes 3617 (MEXU, MO); Mpio. Jesus Carranza, km 10 Cedillo-Francisco Villa
road, 17° 20' N, 094" 40' W, 150 m, 18 Jan 1975 (fr), B. Dorantes 3941 (F LL, MEXU, MO): Hills between
Playa Escondida and Fstacion Biologica Los Tuxtlas, 50-100 m, 27 May 1981 (fr), A.Gentry et al. 32337
(MO); Coatzacoalcos-Las Choapas, 2 km from the exit, 50 m, 18 Nov 1968 (fr), A. Gomez P 3909 (MEXU):
Mpio. San Andrés Tuxtla, Lote 67, Estacion Biologica Tropical Los Tuxtlas, 350 m, 28 Mar 1983 ({L), G.
Ibarra M. & L. Cortes A. 523 (MEXU, MO); 18 34-36 N, 095° 04-09' W, 160 m, 3 Nov 1983 ([r), G. [barra
M.& C. Lundell 990 (MEXU, MO); 200 m, 18 Apr 1987 (fl), G. Ibarra M. & N. Pérez N.3092 (FTG, MEXU,
MO); 300 m, 13 Apr L991 (LD, G. [barra M.& S. Sinaca C. 3554 (MEXU, NY); Mpio. Mecayapan-Soteapan
border, pe of spur on N side of Volcan San Martin a in, ca. LOO m below the summit ridge, 6
Pajapan, 18° 18' 30" N, 094" 43' 30 W, 1,100 m, 15 Jul 1982 (ster), M. Nee et al. 25027 (F): Mpio.
junction with road to Sontecomapan
fa
km N
Crenies along road from Catemaco to Coyame, 2.5 km Eo
1443
and 0.5 km from N edge of pe ee oy aco, 6 km NE of Catemaco, 18° 27' N, 095° 03' 30" W, 350
m, 20 Apr 1983 (fl), M. Nee et al. 26675 (F, LL, MO, NY); Mpio. San Andrés Tuxtla, Estacion Bioldgica
de Los Tuxtlas, “Ebitrolotu, meee ; ne rs D.Pinero 15B(MEXU, MO); Mpio. San Andrés
Tuxtla, Estacion Bioldgica de Los Tuxtlas, without elev. 28 Oct 1981 (fr), I. Ramamoorthy 2617 (MEXU,
MO), 25 Mar 1982 (f1), T. Ramamoorthy 3457 (MEXU, MO); 17 Sep 1982 (fr), T. Ramamoorthy 4148
(MEXU, MO); 22 Apr 1970 (f{D, M. Rosas 1879 (MEXU); Mpio. San Andrés Tuxtla, Estacion Biologica de
Los Tuxtlas, Laguna Zacatal 2.5 km NW, 18’ 34-36 N, 095° 04-09' W, 200 m, 9 Apr 1985 (£1), S. Sinaca
& E. Aparicio 31 (MEXU, MO). Mpio. Hidalgotitlan, brecha Hnos. Cedillo-La Escuadra, 200 m,
ve 1974 (fl), M. Vazquez T. et al. 125 (BM, MEXU), Mpio. Hidalgotitlan, Rio Soloxuchil between
Hermanos Cedillo and La Escuadra, 17 17' N, 094° 38' W, 150 m, 3 Apr 1974 (fD, M. Vazquez T etal.
333 (F); Mpio. See entaes er anos Cedillo-La Escuadra por El Caminoa Pancho Villa, 17° 16'N,
094° 37' W, 152 m, 20 Oct 1974 (fr), M. Vazquez T. et al. 1241 (F), Mpio. Jess Carranza, 2 km N of
Poblado, 17° 16' cs 094" 40 . os m, | Nov 1983 (fr), M. Vazquez T. et al. 2654 (LL); 17 Apr 1983 (ED, A.
Villalobos C. et al. 28 (LL, MO); Mpio. San Andrés Tuxtla, Estacion Bioldgica de Los Tuxtlas, ca. 20 km
N of Catemaco, 140 m, 6 Apr 1973 (f1), A. Villegas H. 84 (F LL); Mpio. hills S of Minatitlan, ca. 27 km E
of La Laguna, 17’ 14'N, 094° 18' W, 250 m, 21 Feb 1981 (fr), T. Wendt et al. 2911 (LL); Mpio. Hidalgotitlan
Laguna-Sarabia, | km W of Rio Chachijalpa, 30 km al W de La Laguna, 17° 12'N, 094° 46’ W, 100 m, 26
Feb 1981 (f1), T. Wendt et al. 2935 (LL); Mpio. Hidalgotitlan, Cenotes 0.3 km W of La Raya, 6.5 km N of
La Laguna, along road to Poblado 7, 17° 19' N, 094° 30’ W, 130 m, 30 Mar 1981 (fl), I. Wendt et al. 3106
(LL); Mpio. Minatitlan, hills S of Poblado II, ca. 27 km E of La Laguna, 17° 14' N, 094° 17' W, 300 m, 3
Jun 1981 (fr), T. Wendt et al. 3407 (LL); Mpio. Minatitlan, 2 km N of Uxpanapa, Poblado 12, along road
to Poblado 13, 17° 14'N, 094° 13' W, 130 m, 17 May 1983 (fr), T. Wendt et al. 4107 (LL); Mpio. Minatitlan,
oe S of Poblado lL, ca. 27 km E of La Laguna, 17° 14' N, 094° 30' W, 250 m, 26 Apr 1985 ({r), T. Wendt
al. 4841 (LL, NY). BELIZE. Cayo: Mountain Pine Ridge, without elev, 24 Feb 1931 (fr), H. Bartlett
i (F); El Cayo, without elev., 13 Apr 1931 (fr), H. Bartlett 12926 (F NY); San Antonio, without elev,
6 May 1931 (fr), H. Bartlett 13076 (F); Vicinity of La Flor at Rio de la Flor, 6 mi S of Grand de Oro,
1,700-2,000 ft [518-610 ml], 3 Jun 1973 (fr), T. Croat 23750 (MO), Along road between Millionario and
Grand de Oro, 1,700 ft [518 m], 2-3 Jun 1973 (fr), I. Croat 23689 (LL, MO); El Cayo road, without elev, 16
Feb 1938 (fr), P Gentle 2181 (LL, NY); Vaca, without elev, 11 Apr 1938 (fr), P Gentle 2476 (A, F NY);
Humming Bird Highway, without elev, 24 Mar 1955 (fr), P Gentle 8637 (LL-2 sheets, TEX); Rio Frio
Caves road, Augustine, Mountain Pine Ridge Forest Reserve, 460 m, 26 May 1989 (fr), J. Meave & A
Howe 1404 (MO); Maria Camp, 800 m, 24 Mar 1954 (fr), A. seaales R. 103 (F), At Rio Frio Cave near
Augustine, ca. 300 m, 5 Jul 1970 (fr), D. Spellman 1583 (MO), Vicinity of Puenta natural, Chiquibul
Forest Reserve, 1,700-1,800 ft [518-549 m], 26 Apr 1969 (fr), G. sis 30122 (LL): Millionario, 2,000
ft[610 ml, 12 Dec 1968 (FL), G. Proctor 29864 (LL); San Pastor track, 450 m, 15 Apr 1995 (fr), C. Whitefoord
9014 (BR); Chiquibul Forest Reserve, Engineers’ Camp, 500 m, 27 Apr 1998 (fr), C. Whitefoord 10378
(MO). Toledo: Ca. 40 km SW of Punta Gorda Dolores Estate, 15° 59° N, 089° 13’ W, without elev. 30 Jan
1990 Ur), M. Balick et al. 2500(TEX): First second bridges near Columbia Forest Station, without
elev, 26 Jun 1972 (fr), J. Dwyer 9954 (LL, MO); Edwards Road beyond Columbia, without elev. 8 Jun
1947 (fr), P Gentle 6226 (GH, LL-2 sheets, - TEX): Near San Antonio, without elev., 22 Nov 1951 ({D)
P Gentle 7525 (GH, LL-2 sheets, PH, TEX); SW-ern Maya Mountains, Columbia River Forest Reserve,
Union Camp, 16° 23' N, 098° 09' W, 700-750 m, 5 Apr 1992 (fr), B. Holst 4015 (MO); Columbia River
Forest Reserve, Union Camp, ca. 3 km W of SW end of Little Quartz Ridge, 16° 23' 53" N, 089° 07.34" W,
00-750 m, 22 Feb 1997 (fr), B. Holst 5975(BRH, F FTG, MO, SEL); Temash River, Pueblo Birejo, 1,700 ft
[518 ml, 19 Feb 1934 (£1, fr), W. Schipp S-692 (A, BM, EK GH, K, MO, NY-2 sheets), Union Camp, 750 m, 15
May 1979 (fr), C. Whitefoord 1739 (MO); Blue Creek, without elev, 12 Jun 1981 (fr), C. Whitefoord 3208
(MO); 2 Oct 1992 (Fl), C. Whitefoord 8275(BM). GUATEMALA. Alta Verapaz: Tucuru, Finca de Remedios,
without elev, 20 ne 1979 (£1), J. Boeke & S. Utzschneider 2948 (NY); On Coban Road, km 285-286,
between Chiracte and Chapultepec farm, without elev, 24 May 1964 (fr), E. Contreras 4773 (LL-2
sheets); 4775 (LL Beane Chapultepec farm, ca.1 km N, without elev. 26 May 1964 (fr), FE. Contreras
aan
1444 BRIT.ORG/SIDA 20(4)
4809 (GH, LL-2 sheets, TEX); Chahal, bordering Rio Chiyt, 3.5 km NW W, 28 Sep 1968 (fD, E. Contreras
7742 (DUKE, FG, LL-3 sheets); Chahal, on Sepur Ranch, eaecen Chahal, 2 km S, without elev,
8 Oct 1968 ([D, E. Contreras 7860 (FE LL-3 sheets); Secoyoeti, near the Finca Sepacuite, without elev, 14
Apr 1902 Ur), O. Cook & R. Griggs 61 CUS); Tamahu, 2,500 ft [m], 17 os 1920 (ED), H. Johnson vere (F
US), Between Coban and Finca Chimoté, near Rubeltein, 800-1,500 m, 25 Feb 1942 (f1, fr), J. Steyermark
44205 (F). Baja Verapaz: Nino Perdido, bordering Rio San Jose, 8 km N, without elev., 26 May 1977 (f1-
bud), C. Lundell & E. Contreras 20986 (LL-2 sheets, PH); 28 May 1977 (fr), C. Lundell & E. Contreras
21012 (LL-2 sheets, PH); Paujal, 1000 m, Oct 1912 (f(D, H. von Turckheim 3922 (US). Huehuetenango:
Cerro Jolomtac, above Finca San Rafael, Sierra de los Cuchumatanes, 900-1,200 m, 22 Jul 1942 (fr), J.
Steyermark 49143 (PF); Between Cerro Chiblac and Finca San Rafael, between Ixcan and Finca San
Rafael, Sierra de los Cuchumatanes, 200-800 m, 24 Jul 1942 (fr), J. Steyermark 49475 (F): Cerro Victo-
ria, across river [rom Finca San Rafael, Sierra de los Cuchumatanes, 800 m, 27 Jul 1942 (1), |.
Steyermark 49644 (F). Izabal: El Estor, without elev. 5 Mar 1972 (fr), E. Contreras 11145 (DUKE, LL-2
sheets); 17 Mar 1972 (fr), F. Contreras 11357 (DUKE, LL-2 sheets); Jocolo, without elev, 25 Nov 1920
(ED, H. Johnson 1024 (F); Cadenas, ca 6 km S of the village, on old brecha to Rio Chocon Arriba, with-
out elev, 20 Feb 1975 (fr), C. Lundell & E. Contreras 19016 (LL-3 sheets); El Estor, bordering Rio Sarco,
without elev, 26 May 1975 (ir), C. Lundell & E. Contreras 19358 (LL-2 sheets); Carosal bordeando Rio
Sorstun, without elev, 21 Jan 1969 (fr), R. Tun O. 120 (FLL); Entre caserios Ceja y Cienaga, 15° 40 N,
O89 W, without elev. 9 Mar 1972 (fr), R. Tin O. 2300 (EF LL). Petén: Remate, on Tikal Road ca.6 km NE
of village, without elev, 16 Mar 1960 (fr), E. Contreras 678 (LL-2 sheets, TEX): Dolores, bordering Ar-
royo Ixcol, E of village, 800 m, 14 Apr 1961 (fr), E. Contreras 2071(LL-3 sheets, TEX); Dolores, between
kms 83-84 of Machaquila Road, without elev. 24 Apr 1961 (fr), E. Contreras 2171 (G, LL-2 sheets):
Dolores, bordering pineland ca 2 km south of village, without elev. 3 May 1961 (fr), E. Contreras 2243
(DUKE, LL-2 sheets, PH, TEX); Dolores, km 79 of Machaquila Road, without elev, 14 Jul 1961 (£1), E.
Contreras 2005 (DUKE, LL-2 sheets, TEX); Dolores, ca. 2 km S, without elev, 18 Jul 1961 (fD, E. Contreras
2625 (LL-2 sheets); Santo Toribio, bordering the village, without elev, 28 Jul 1961 (fD, E. Contreras
2703 (LL-2 sheets, TEX); Milpa Grande at Macanché, without elev, 5 Mar 1966 (fr), E. Contreras 5511
(ELL): Cadenas, W of ie 106-167 of Poptun ae without elev. 12 Aug 1966 (EL, fr), E Contreras 5956
(GH, LL-2 sheets, TEX); Chinchila, 3 km from San Luis on Sebol Road, without elev, 12 Oct 1966 (£1),
E. Contreras 6384 (G, LL-3 sheets); Poptun Road, km 82, without elev. 14 Nov 1966 we E. Contreras
6595(LL-3 sheets); La Cumbre, at km 142 of road, without elev. 2 Mar 1967 (fr), E. Contreras 6634 (LL
2 sheets); Seamay, without elev. 6 Mar 1967 (fr), E. Contreras 6668 (FG, LL-3 sheets); San Pedro on
Cadenas Road, without elev, 13 Aug 1967 (fr), E Contreras 7018 (FE, G-2 sheets, LL-3 sheets); San Pedro
on Cadenas Road, bordering river, without elev. 9 Jan 1970 (fr), EC s 9481 (DUKE, LL-3 sheets):
Ca 5miSot Tikal National Park, without elev. 19 Jun 1973 (fr), J. Dwyer 11262 (LL, MO); San Luis, km
52 of road S of village, without elev, 12 Jul 1959 (fr), C. Lundell 16382 (F LL-3 sheets, TEX); Cadenas,
bordering Rio teen a Dios, without elev. 7 Sep 1975 (fr), C. Lundell & E. ne reras 19805 (LL-2
sheets, NY, PH); La Cumbre, on Pusila Road, N ca. 4.5 km, without elev, 30 Aug 1976 ([D, C. Lundell &
E. Contreras een |L-2 sheets), La Cumbre, Sapurul, without elev, 18 Apr 1977 (fr), C. Lundell
& E. Contreras 20829 (LL-2 sheets); Santa Elena a Petén en orillando camino para Puerto Mendez,
ie km Lo4, without elev, 17 Mar 1970 Ur), R. Tun O.774a (ELI > 777 (BLL, MO, NY); Santa Elena,
behind the school at Colpeten, ca. 200 m, 5 Jun 1970 (fr), R. Tun O. 82 (F LL); San Luis, along road to
Poctun, km 119, without elev, 3 Dec 1970 ({D, R. Tun O. 1451 (F LL, MO, NY); 117, without elev, 6 Dec
1970 (fr), R. Tun O. 1479 (ELL); Along road to Arroyo Dolores, ca 300 mS of Dolores, without elev, 16
Feb 1971 (fr), R. Tun O. 1587 (FLL); Dolores, without elev, 14 Feb 1971 (fr), R. Tun O. 1563 (F LL); Santa
Elena, along road to La Libertad, km 9, SE side, without elev, 8 May 1971 (fr), R. Tun O. 1753 (E LI
MO); Poptun, along road to San Luis, km 117, ca 125m W of road, without elev, 7 Jan 1972 Ur), R. Tun
O, 2158 (FLL, MO, NY). HONDURAS. Atlantida: Mpio. Esparta, 19.2 km EF of Tela on the Tela-Ceiba
Hwy. ca. L5 km Sof the hwy., near San Francisco de Soco, 15 44' N, 087° 21’ W, 50-100 m, 15 Apr 1994
1445
(fl), A. Brant & R. Ztiniga 2842 (F MEXU, MO); Along road for municipal water supple of Tela,
Lancetilla Botanical Gardens, on road ca. 2 mi WSW of Tela and S of main hwy., 15 44'N, 087° 27' W,
70-90 m, 9 Feb 1987 (fr), I. Croat & D. aad Seema MO), Tela, Lancetilla Valley, va alley above
Experiment Station, along stream and IT
30" W, 30-60 m, 4 Nov 1988 (fr), J. MacDoneel etal. 3159 (MO); Vicinity of Tela, 500 ft [152 ml], 6 oe
1926 ({L), E. Mitchell 98 (GH); Lancetilla, 7 km SW de Tela, 150 m, 3 Feb 1986 (fr), E. Mondragon 128
(MO); Vicinity of San Alejo, at base of hills S of San Alejo near Rio San Alejo, 150-270 m, 22-27 Apt
1947 ({D, P Standley 7616 (F); Lancetilla Valley, near Tela, 20-600 m, 6 Dec 1927-20 Mar 1928 (fr), P.
Standley 52643 (A, FE US), 52983 (A, F US); 53120 (A, F US); 55598 (F); 56563 (A, FE US); 56803 (A, F US),
56882 (F GH, US). Cortés: Along Rio Lindo near Carrizal, 550 m, 12 Apr 1951 (ED, L. Williams & A.
Molina R. 17804 (FE GH, US). Yoro: Rio Texiguat watershed, 15° 30' 10" N, 087° 20' 32" W, 220 m, 18 Apr
1995 (fl), T. Hawkins & M. Merello 767 (F MEXU, MO, NY, TEX); Near Progreso, without elev. 26 Mat
1929 (fl), W. Hottle 26 (F). Unknown locality in Honduras: Puerto Sierra, forest along Highland Creek,
without elev, 18 Jan 1903 (fr), P Wilson 69 (NY, US).
ACKNOWLEDGMENTS
We thank the Missouri Botanical Garden and the Flora Mesoamericana Project,
as well as the Fairchild Tropical Garden, for their support, to bring Ricketson
and Pipoly together to work on the project. We gratefully acknowledge the loans
from many herbaria that made the study possible, along with our collaborators
who supplied us with much of the critical material. Richard G. Walter (formally
of CS) and Gerrit Davidse (MO) generously gave of their time to provide thor-
ough reviews of the entire manuscript. Special thanks are due to Roy Gereau for
his critical review of the Latin diagnoses published herein. The drawings were
prepared by Jon Ricketson. Finally, the assistance of Jon Ricketson’s volunteer,
Mary Bard, greatly facilitated the annotation and movement of specimens.
REFERENCES
Cuen, C. and J.J. Piroty. 1996. Myrsinaceae. In: Wu Zheng-yi & P. Raven, eds. Flora of China.
Science Press, Beijing, People’s Republic of China, and Missouri Botanical Garden, St.
bouls, MO-US A. 1521-38.
Lunbewt, C.L. 1981. Neotropical Myrsinaceae IV. Phytologia 48:137-142.
Mez, C. 1902. Myrsinaceae. In: A. Engler, ed. Das Pflanzenreich IV. 236 (Heft 9):1-437.
Mez, C. 1903. Additamenta Monographica 1903. Bull. Herb. Boiss. ser. II, 3:224—238.
Mez, C. 1920. Additamenta monographica 1919. Repert. Spec. Nov. Regni. Veg. 16:308-
Orrstep, A.S. 1862. Myrsineae Centroamericanae et Mexicanae. Vidensk. Meddel. Dansk
Naturhist. Foren Kjabenhavn 1861:11 7-142.
Pipoty, JJ.and J.M. Ricketson. 1998. A revision of the genus Ardisia subgenus Graphardisia
(Myrsinaceae). Sida 18:433-472.
Pipoty, J.J. and J.M. Ricketson. 1999a. Discovery of the Indio-Malesian genus
Hymenandra(Myrsinaceae) in the neotropics, and its boreotropical implications. Sida
18:701-
1446 BRIT.ORG/SIDA 20
—
>
)
Pipovy, J.J.and J.M. Ricketson. 1999b. Additions to the genus Ardisia subgenus Graphardisia
(Myrsinaceae). Sida 18:1145-1160.
Pipovy, JJ.and J.M. Ricketson. 2000. Discovery of Ardisia subgenus Acrardisia (Myrsinaceae)
in Mesoamerica: Another boreotropical element? Sida 19:275-283.
Ricketson, JM. and JJ. Pirovy. 1997. A synopsis of the genus Gentlea (Myrsinaceae) and a
key to the genera of Myrsinaceae in Mesoamerica. Sida 17:697—707.
RickeTson, J.M.and J.J.Pipovy Ill. 2003. Revision of Ardisia subgenus Auriculardisia (Myrsinacae).
Annals Missouri Bot. Gard. 90:1 79-317.
Smith, J.D. 1888. Undescribed plants from Guatemala. Ill. Bot. Gaz. 13:74—77.
NUMERICAL LIST OF TAXA
la. Ardisia nigrescens Oerst. pac oe
b. Ardisia nigrescens Oerst. subs] ithii (Mez) Ricketson & Pipoly
|
2. Ardisia ies es eons & ia
_Ardis ia tuerckheimii Don
Lisl OF ExXsiCCATAE
The figures in parentheses refer to the numbers from the numerical list of taxa.
Collection numbers in boldface type indicate type specimens.
J. Alcocer, J. & C. Morales s.n. (3); Alcorn, J.2937 (3); 9481 (3); 11145 (3); 11357 (3 fagtalen E.&
(la) C. Lundell 20069 (1b); Cook, O. & R. Griggs 611
3); 767 (1b); Cowan, C. et al. 3935 (1 jae. AR
23689 (3); 23750 (3); Croat, T.& D. Hannon 64596
Balick, M. et al. 2500 (3); Bartlett, H. 11742
(3);12926 (3);1 3076 (3); Beaman, J.5 186 (3); Boeke,
J.&S.Utzschneider 2948 (3);Brant, A.& R. Zuniga
2842 (3); Breedlove, D.9721 (3); 9733 (3); 14978 Davidse, G. 36051 (1b); Davidse, G. & A. Brant
(3); 15702 (1a); 24232 (1a); 27927 (2); 33006 (2); 32362 (1b); Dillon, M. et al. 1831 (3); Dorantes, B.
33302 (3); 38200 (3); 38709 (1b); 38764 (3); 47348 2802 (3); 3576 (3); 3617 (3); 3941 (3); Dressler, R.
(1b); 48740 (3); 49112 (1a); 49160 (1b); 49445 (2): 1507 (1a); 1637 (1b); Dwyer, J.9954 (3); 11262 (3).
49970 (1b); 52487 (2); 52591 (2); 53339 (3);53425 Gentle, P2181 (3); 2476 (3);2798 (1b);4522 (1b):
(3); 37739 1D); O8667 13); Breedlove; DBE _4g95 (1b) 5231 (1b) 6177 (1b); 6226 @)r6729
Almeda 57228 (1b),57210 (2),57370(1a):57927 (14). 6980 (1);7525 (3);7639 (1b);8637 (3);Gen-
(1b); Breedlove, D. & R. Dressler 29601 (3); try, A.et al, 32337 (3); Goll, G, 261 (1b);Gémez P,
Breedlove, D. & B. Keller 49485 (2): Breedlove, D. A. 3909 (3): 7885 (1a
& A.Smith 22120 (2):22121 (2);Breedlove,D.&R
Thorne 21166 (3). Hahn, M. 21 (1a); Hawkins, T. 1443A (1b); Hawk-
ins, T.& M. Merello 767 (3); Hernandez X., E. 102
1a);Hernandez G., H.433 (3);842 (3);896 (3);935
(3); 985 (3); 1056 (3); 1994 (3); Hernandez G., H. &
Gonzalez L. 1637 (3); 1681 (3); 1769 (3);
i)
—
—
Calzada, J. 227 (3); 723 (3);Campos, A.et al. 1702
(3); 1715 (3); Carlson, M. 1744 (3); Castillo C., G.
408 (3); Cedillo T., R. 164 (3); Cedillo T., R. & J.
Calzada 48 (3);Cedillo T, R.& D. Lorence 702 (3), Hernandez G,H.&R.Torres 813 (1a);Holst,B.4015
Cedillo T., R.& R. Torres 1 144 (3); Chiang, F. 301 (3): 5975 (3): Hottle, W. 26 (3).
ened? oo ee (3); os erat Ibarra M.,,G.& L.Cortes A.523 (3);IbarraM.,G.& C.
(3); 2243 (3); 2605 (3); 2625 (3); 2703 (3);4254 (1b); 1 ndell 990 (3); Ibarra M,, G. & N. Pérez N. 3092
4291 (1b); 4667 (1b); 4773 (3); 4775 (3); 4809 (3); 3): Ibarra M.,G.& S, Sinaca C, 3554 (3).
5414 (1b); 5511 (3); 5956 (3); 6290 (1b); 6384 (3);
6595 (3); 6634 (3); 6668 (3); 7018 (3); 7742 (3); 7860 Johnson, H. 228 (1b); 1007 (3); 1024 (3).
—a
—
1447
Karwinsky, W. 450 (1a).
Liebmann, F.28A (1a); 28B (1a); 15341 (1a):Little,
E.25392 (1a);Lundell, C.6302 (3); 16267 (3): 16382
(3); 17823 (1b); 17923 (1b); 17978 (1b); 18039 (1b);
18040 (1b); 18133 (1b); C. Lundell & E. Contreras
19016 (3); 19358 (3); 19805 (3); 20283 (3); 20829
(3); 20986 (3); 21012 (3); Lundell, C.& A. Lundell
7130 (Ja).
MacDougal, J.et al. 3159 (3); Martinez S.,E.6523
(1b); 6537 (1b); 6901 (1b); 7411 (1b); 7622 (1b
7658 (1b); 8059 (1b); 8427 (1b); 9318 (1b); 9361
(1b); 9581 (1b); 10106 (1b); 10132 (1b); 11075 (1b);
11531 (1b); 13454 (1b); 13616 (1b); 15982 (1b);
21333 (1b); Martinez S.,E.& G. Aguilar 8630 (1b);
8790 (1b); 13707 (1b); Martinez-Calderon,G. 209
(1a);828 (1a); Meave,J.& A. Howe 1404 (3); Meave
del Castillo, J. et al. 1480 (3); Medina A., M. & F.
Vazquez B.379 (1a);Mell,C.576 (3); Méndez G,, A.
4445 (2); Méndez T., A. 5097 (2); 5469 (2); 5640
(2); 5667 (2); 5837 (2); 5842 (2); 6052 (2); 6112 (2);
6211 (2); Mendez T.,A.& M. Concepcion Mtz. de
Lopez 9499 (1a); Miranda, F.6289 (3); Mitchell, E.
66 (3); 98 (3); Molina R., A. 103 (3); Mondragon, E.
128 (3).
Nee, M. et al. 25027 (3); 26675 (3).
Peck, M. 576 (1b); Pinero, D. 15B (3); Proctor, G.
29864 (3); 30122 (3).
Ramamoorthy, |. 2617 (3); 3457 (3): 4148 (3);
Ramamoorthy, T.et al. 1356 (3);Ramirez C.,,D.s.n.
(1a);Reznicek, A.et al. 187 (1a); Rosas, M. 1879 (3).
pat,
,
=
Sandino, J. 181 (1a); Sessé, M. et al.679 (1a); 730
(1a); Schatz, G.et al. 1195 (1a); Schipp,W. 287 (1b):
478 (1b);482 (1b);S-692 (3):Schnée, L.& P Maury
s.n. (1a); Shilom T., A. 2599 (3): Sinaca C., S. & E.
Aparicio 31 (3); Soto, J.et al. 13479 (3): Spellman,
D. 1583 (3); Standley, P. 7616 (3); 52643 (3);52983
3); 53120 (3); 55598 (3); 56563 (3); 56803 (3);
56882 (3); Stevens, W.& E. Martinez S.25849 (1b);
Stevenson V.,N. (1b); Steyermark, J. 44284 (1b);
44383 (1b); 45161 (1b); 49253 (1b); 45427 (1b);
45493 (1b); 44205 (3); 49143 (3); 49475 (3); 49644
(3).
Thompson, B. & B. Fields 379 (1a); Torres C.,, R.
11039 (3); Torres C., R. & L. Cortes A. 11500 (3);
Torres C.,R.& E. Martinez S. 11163 (3);Tun O,,R.
20 (3); 774a (3); 777 (3); 1182 (3); 1451 (3); 1479
(3); 1587 (3); 1563 (3); 1753 (3); 2158 (3); 2300 (3):
Turckheim, H. von 1035 (3); 3922 (3); 7766 (1b);
7918 (1b); 8707 [11978] (1b); 11163 (1b).
Valdivia O.,P. 2407 (1b); Vazquez T., M. 1395 (3);
Vazquez T.,M.et al. 125 (3);333 (3); 1241 (3); 2654
(3); Ventura A.,F. 13621 (1a); 20767 (1a);Villalobos
C,,A.et al. 28 (3); Villegas H.,, A. 84 (3).
Wendt, T. et al. 2911 (3); 2935 (3); 3081 (3); 3106
(3); 3407 (3); 3623 (3); 4107 (3); 4841 (3);
Whitefoord, C. 1739 (3); 3208 (3); 8275 (3);9014
(3); 10378 (3);Williams,L.& A. Molina R. 17804 (3);
Wilson, P. 69 (3
—
5:
pas
1448 BRIT.ORG/SIDA 20(4)
BOOK NOTICES
R.P.(Rop) RANDALL. 2002. A Global Compendium of Weeds. (ISBN 0-9587439-8-
3, hbk.). R.G.and FJ. Richardson, PO. Box 42, Meredith, Victoria 3333, Australia.
(Orders: www.weedinfo.com.au, richardson@weedinfo.com.au, +61-3-5286-
1533 fax, VISA, MasterCard, BankCard, AE, checks sas dollars only]
payable to R.G. and FJ. Richardson). $A165.00 plus $AL0 postage within
Australia, overseas postage: please ask for a quote, 944 pp., almost 21,000
entries, 6 3/4 x 93/4".
From the backcover— Predicting which plants will become weeds is difficult, with the single most
eae nt indicator of a species’ weed potential being a documented weedy history.
Jntil now, the most a acca coverage of the world’s ers i a was produced in 1979
im Randall has compil list of almost 21,000 en-
=)
a
and | ~ 6400 species
tries, as well asa soramieuenaive dex containing more than 15,000 alternate scientific names and
27,000 common names in numerous languages.”
“A Global Compendium of Weeds is specilically designed to give anyone interested in the weed
potential of a planta cae on its status with, most importantly, further avenues of finding more
information throug] sive reference listing.”
laving receiv ede a recent request for information on dragon grass or Chinese alpine rush (Poa
sphondy ‘lodes), |checked the list < und to my surprise found it listed. The ntry came Wi ith eight 1 ‘eter-
ences that presumably lead you to additional information. A couple of the references also include
websites that | checked..two worked and one ” id already on inged addresses. Not a surprise. The
index isextensive and includ ). Table Lis the source codes
(references with the taxonomic and sect entries) with feferenices and includes 300 references.
Table 2 is the s ich region represented in the book. This an impressive compilation.—
Barney Lipscomb, es search Institute of lexas, 509 Pecan St, Fort Worth, 1X 76102-4060, U.S.A.,
barney@britorg.
CARMINE STAHL and RitA McELvANEY. 2003. Trees of Texas: An Easy Guide to Leaf
Identification. (ISBN 1-58544-242-9, hbk.). Texas AGM University Press,
John H. Lindsey Bldg., Lewis St. 4354 TAMU, College Station, TX 77843-
4354, U.S.A. (Orders: wlawrence@t edu, http://www.tamuedu/upress,
800-826-8911). $29.95 hbk., 338 pp., 18 color, 270 b/w photos, | map, Bib.,
Index, 7 appendices, 8 1/2" x 11",
This book contains numerous life-sized leaf, flower, and fruit imi oe With more than 200 native
and naturalized species organized by leaf shape, it of species. The genus Crataegus
(hawthorn) has 25-30 species in Texas and ie ae and McE lvaney cover nine species. Three species of
Prunus are cover and yet there are a dozen or more in the state. Nothing is wrong here but one needs
to keep in mind that Trees of lexas: An Easy Guide to Leaf Identification is a guide to the more com
mon trees of Texas; a very nice one at that with photographs that are sharp and crisp with detail.
“Texds, 509 Pecan St, Fort Worth, TX 76102-4060,
—Barney Lipscomb, Botanical Research Institute o
U.S.A, barney@britorg.
SIDA 20(4): 1448. 2003
A REVISION OF THE PSIDIUM SALUTARE
COMPLEX (MYRTACEAE)
Leslie R. Landrum
ELSE,
Sc hool Or cig Sciences
Arizona State University
Tempe, AZ 85287-4501, U.S.A.
les. landrum@asu.edu
ABSTRACT
1]
lex is revised with keys, descriptions, maps, and illustrations and is con-
The Psidium salutare
sidered to consist of P. Lares inum (with no additional daa udee taxa) and P salutare a a
oats
total of five varieties). This complex ranges from eastern temperate South America to Mex
Central America. The following taxa are lectotypified: Calycolpus 7 orus Sagot, ie acutata
O. Berg, Myrtus blanchetiana O. Berg, eens O. Berg, Myrtus cuspidata var. tetramera O.
Berg, Myrtus incana O. Berg, Myrtus mucronata Cambess., Myrtus nivea O. Berg, Myrtus pubescens O.
Berg, Myrtus rigida O. Berg, Sea iapnton O. Berg, Myrtus sericea var. fruticosa O. Berg, Myrtus
suffruticosa O. Berg, Myrtus suffruticosa var. latifolia O. Berg, Psidium guayabita A. Rich., Psidium
pohlianum O. Berg, Psidium salutare var. laxum O. Berg and Psidium salutare var. subalternum O.
erg. The following new combinations are made: Psidium salutare var. decussatum (DC.) Landrum,
Psidium salutare var. mucronatum (Cambess.) Landrum, Psidium salutare var. pohlianum (O. Berg)
Landrum, and Psidium salutare var. sericeum (Cambess.) Landrum
RESUMEN
Sel ision del pl i le Psidi salutar >| descripciones mapas e ilustraciones
E] leininel p] +t fixe | sf: long inci Dead :
plej | y un total de
cinco variedades). E lej lesde el este de Sur América templada hasta México y
Centro América. Se seleciniad Con para los siguientes taxa: Calycolpus parviflorus Sagot,
Myrtus acutata O. Berg, Myrtus blanchetiana O. Berg, Myrtus mere O. Berg, Myrtus cuspidata
var. tetramera O. Berg, Myrtus incana O. Berg, Myrtus mucronata Cambess., Myrtus nivea O. Berg,
Myrtus pubescens O. Berg, Myrtus rigida O. Berg, Myrtus sellowiana O. Berg, Myrtus sericea var.
fruticosa O. Berg, Myrtus suffruticosa O. Berg, Myrtus suffruticosa var. ee O. Berg, Psidium
ee A.Rich, Psidium aati Berg, Psidium salutare var. laxum O. Berg y Psidium salutare
r. subalternum O. Berg. Se hacen las siguientes combinaciones nuevas: Psidium salutare var.
a ussatum (DC.) Landrum, Psidium salutare var. mucronatum (Cambess.) Landrum, Psidium salutare
var. pohlianum (O. Berg) Landrum, y Psidium salutare var. sericeum (Cambess.) Landrum.
Psidium (Myrtaceae) is a genus of at least 50 and perhaps as many as 100 species
(McVaugh 1968) with a natural range from Mexico and the Caribbean to Uru-
guay and northern Argentina on the American continents and extending to
some east Pacific islands (e.g. Galapagos). A few weedy and/or cultivated spe-
cies have been spread by humans around the World to tropical and subtropical
climates (e.g, Pguajava L., P.cattlei Sabine). The great variance in estimates
of numbers of species is due to complexes of entities that might themselves be
considered single variable species or species groups. One such group is the
SIDA 20(4): 1449-1469. 2003
1450 BRIT.ORG/SIDA 20(4)
Psidium salutare is complex, which ranges from Mexico to Uruguay and in-
cludes such commonly accepted species as P. laruotteanum, P. salutare, P.
luridum, Pincanum, and P. pohlianum. The purpose of this paper is to treat the
P salutare complex, as it may be several years before a monograph of the whole
genus will be completed.
Psidium is distinguished from other genera by acombination of floral and
seed characters discussed in Landrum and Sharp (1989), namely, flowers 5-
merous with multiovulate locules; placenta often peltate; seed coat rough or
dull, not lustrous, covered with a pulpy layer when wet; hard portion of seed
—
coat (5-)8-30 cells thick at narrowest point, the cells thick-walled, elongate,
overlapping. The P. salutare complex is differentiated from other species of the
genus by 1) a usual habit of subshrub or shrub (but reaching tree size in some
populations); 2)a habitat mainly of grasslands or low shrubby growth (cerrado)
that is frequently burned; 3) an ability to sprout back from underground stems
even after fires; +) leaves with brochidodromous venation with a well marked
—
marginal vein that closely follows the margin and tertiary veins that form a
reticulate-dendritic pattern; 5) relatively small flower buds (4-9 mm) with the
calyx open; and 6)a peltate placenta with one or two rows of ovules on the edge
of the each lamina.
Another group of similar range (northern Argentina to Venezuela), habit,
and habitat is the Psidium cinereum complex (P. cinerum DC,, P grandifolium
DC., Paustrale Cambess., P missionum D. Legrand, and P.suffruticosum O. Berg).
That complex differs in having leaves with brochidodromous venation (some-
times blending into acrodromous venation proximally) with a marginal vein
that broadly arches between the laterals and thus does not closely follow the
margin, relatively large flower buds (often over lcm long) with the calyx open
or closed, and a non-peltate, only slightly protruding placenta with numerous
irregular rows of ovules.
A third group that is sometimes confused with the P. salutare complex is
the P oligosperma DC. complex, which includes P sartorianum (O. Berg) Nied.
It differs in having the calyx closed or nearly closed and being shrubs to rela-
tively large trees of woodland or forest habitats.
The great variability in the Psidium salutare complex is reflected in the
numerous names that have been proposed, including at least 44 basionyms at
the specific level and several more subspecific names. After several years of in-
termittent study of herbarium specimens and fieldwork | have concluded that
the P. salutare complex is best considered two species: P. salutare and P.
laruotteanum. | recognize P salutare as containing five varieties that correspond
closely with geographic regions that sometimes overlap. All the varieties rec-
ognized here have been recognized previously as species by other taxonomists,
e.g., Berg 1857-1859), de Candolle (1828), Legrand and Klein (1977), and Rotman
(1976). Because Lo
ten find specimens that are intermediate between the enti-
LANDRUM, REVISION OF THE PSIDIUM SALUTARE COMPLEX 1451
ties I recognize as varieties of P salutare, | suspect that they freely interbreed
when growing sympatrically. I strongly recommend accepting them at the va-
rietal level but realize others may disagree. Thus, in the discussion of the non-
typical varieties I indicate the name that the entity would have if recognized at
the specific level.
For illustrations I have used portions of scanned herbarium specimens. These
images can be viewed in their entirety in color in the Image Library on the ASU
Herbarium website http://lifesciences.asu.edu/herbarium/. A list of exsiccatae
will also be made available at the same website once this paper is published.
Ecology.—The species of Psidium salutare complex are similar in ecologi-
cal preferences and often grow side by side (e.g., near Brasilia, Brazil). Both grow
in fire-induced savannas or scrub vegetation (cerrado). Both sprout back from
underground stems after fire or other disturbance. The climates in which they
grow often have distinct dry and wet seasons, with freezing temperatures be-
ing rare or non-existent. Commonly associated with them are other genera of
Myrtaceae, e.g., Campomanesia, Eugenia, and Myrcia and the vegetation is of -
ten dominated by Poaceae, Fabaceae, and Asteraceae.
KEY TO THE SPECIES AND VARIETIES OF THE P. SALUTARE COMPLEX
1. Young leaves densely covered with hairs beneath, the lower surface of the leaf
often hidden
2. ee sively lanate, sey less that 1.5 cm wide; apex usually sharply acute to
uptly acuminate; of leaves of two lengths, short and tangled and long
a nearly straight; cna. Uruguay, Bolivia, Paraguay, Rio Grande do Sul
razil). P. salutare var. sericeum
2. Leaves yellowish to grayish short tomentose, often over 1.5 cm wide; apex acute
to rounded;hairs of leaves uniform in length, all tangled; Paraguay, Bolivia, Parana
(Brazil), ieee to Venezuela and Costa Rica P.laruotteanum
1. Young leaves glabrous to moderately sigeecnt beneath.
3. Venation clearly evident above and below, the marginal vein within 1 mm of the
margin; leaves often narrowly elliptic or lanceolate, often 3 or more times as
long as wide, often lustrous; leaf apex often apiculate; calyx-lobes longer than
calyx tube plus se tube; Argentina, Uruguay, Paraguay, Rio Grande do
Sul, Santa Catarina, Par . salutare var. mucronatum
3. Venation clearly deen or raarene marginal vein nee more than 1 mm from
argin; leaves mostly less than 3 times as long as wide, lustrous or not; leaf
usually without an apiculum; calyx-labes longer or shorter than calyx tube
ane hypanthial tu
4. Leaves narrowly ee to elliptic, mostly less than 1.5 cm wide, the apex usu-
ally rounded; peduncle 0.5—1.5 cm long; Parana to Goias. Psidium salutare var.
ecussatum
4. Leaves various, often lanceolate, usually over 2. cm wide, = often acute;
peduncle 0.4-3 cm long; Reteg Ue to Mexico and Caribbea
5. Leaves 4-9 cm long, 2-5 cm wide, elliptic to obovate; ces shrubs to
small trees; plants glabrous; central Brazil, Bolivia to Venezuela. Psidium salutare
var. pohlianum
1452 BRIT.ORG/SIDA 20(4)
5. Leaves 3.3~7 cm long, 1-3.3 cm wide, usually lanceolate, ovate, or ate
normally subshrubs; plants glabrous or pubescent; Paraguay to Mexico al
Caribbean. Psidium ice var. salutare
Psidium laruotteanum Cambess., in Saint-Hilaire Bras. merid. 2:282. 1833. (Figs.
, 3). TYPE: BRAZIL: “Prope Alto da Varginha (prov. Minas Geraes),” Saint-Hilaire
s.n. (HOLOTYPE: Pl, = F-36412!, = ASU photo!).
Psidium aerugineum O. Berg, in Mart. Fl. bras. 14(1):391. 1857. Type: BRAZIL: “in campis prov. Rio
Grande do Sul, oe n. (HOLC OTYPE: B, lost; ISOTYPE: LE, = ASU photol).
! sids Ir ticosa O. Berg, in Mart. FI. Bras. 14(1):448. 1857. Type: BRAZIL: “in prov. Ceara,”
- Gare 1611 & eee 1610 in “hb. Vindob. et Mart.” LECTOTYPE: Gardner 1611 W!
ae letnype by Landrum, 1986), ISOLECTOTYPES, BRI, K!, G, = F-23367! SYNTYPE: Gard-
ner LOO WI, ISOSYNTYPES KI, Sl, US!).
basant a O. Berg, in Mart. FL. bras. 14(1):601. 1859. Type: BRAZIL: “prope Paracatu et
Psidium
Jugi prov. S. Pauli,” Riedel s.n. (HOLOTYPE: LE, = ASU photo!).
Psidium oaieeste ns O. Berg, in Mart. FL. bras. 14(1):600. 1859. Typr: BRAZIL: “Serra da Chapada
prov. Minarum,” Riedel s.n. (HOLOTYPE: LE= ASU photo! lsoTyPE: Pl, = F-36410).
Psidium aerugineum var. angustifolium O. Berg, in Mart., Fl. bras. 14(1):60L 1859. Type: BRAZIL:
(HOLOTYPE: LE; probable HOLOTYPE = ISOTYPE of Psidium aerugineum).
ea a — in Engh & Prantl, Naturl. Pflanzenfam. 3, Abt. 7:66. 1893. New name for
B
ticosa O. Berg.
a warmingianum ee Enum. Myrt. bras. 28. 1893. New name for Campomanesia
suffruticosa ©. Ber
Psidiur rH Val. ve
Santa,” L nil & Warming s.n. (HOLOTYPE: C
Psidium savannarum Donn. Sm,, Bot. Gaz.244. 1897. Type: COSTA RICA: “Savana at Buenos Ayres,
Comarca de Puntarenas,” Tonduz CR-4033 (HOLOTYPE: CR; ISOTYPE: BR).
Psidium allel — ) Burret, Notizbl. Bot. Gart. Berlin-Dahlem 15:485. 1941.
Rodr, Myrt. Paraguay 16. 1903 Type: PARAGUAY: “Ipe hu...Sierra Maracayu,”
rticillata Kiaersk., Enum. My rt. bras. 28.1893. TYPE: BRAZIL: “Lagoa
Myrtus for
Hassler 5079 (HOLOTYPE: Gl, = ASU photol).
sae tat yense Barb. Rodr.ex Chodat & Hassl., Bull. Herb. Boissier 7:797. 1907 TyPE: PARA-
pr. Vaqueria Capibary,” Hassler 4387 (HOLOTYPE: G!, = ASU photol).
ree quinguedentatun Amshoff, Recueil Trav. Bot. Neer. 39:164. 1942. TypE: SURINAM: “Up-
Sipaliwini R. near Brazilian frontier,” Rombouts, H. E. 329 (HOLOTYPE: U; IsoTYPES, MOI,
NY).
Shrub up to ca. 1.5m high, often less than 0.5 m high, with new shoots arising
from a woody subterranean base or rhizome, densely tomentose to sparsely
pubescent over most surfaces; hairs toca. 1.5 mm long, grayish, yellowish-white,
to rusty, usually curled and tangled; young twigs usually densely tomentose,
remaining so for more than I year, the bark of older twigs becoming rough and
scaly. Leaves normally opposite (rarely in whorls of 3, or spirally arranged)
obovate, oblanceolate, elliptic, or oblong, 3-9 (-11) cm long, 1.6-4.5(-6) cm wide,
1.6-3.2 times as long as wide; ies rounded, acute, or acuminate; base obtuse to
cuneate; petiole 0-2 mm long, ca. 2 mm wide, tomentose; midvein flat or slightly
raised above, prominent below, usually more densely hairy above than surround-
ing blade, the venation brochidodromous with 7-13 pairs of lateral veins, these
weak to prominent, slightly raised above in mature leaves, ascending at an angle
LANDRUM, REVISION OF THE PSIDIUM SALUTARE COMPLEX 1453
XN
\\
wt
\N
wt oe
Ne c
Fic. 1. Psidium Jaruotteanum. A, Ribas & Pereira 1812 (ASU), young branch of 0.7 m shrub; note large leaves. B & C,
ea 53638 (ASU). B, middle portion showing full stature of plant with new growth arising from ground level
stem; C, close view of flowers.
of ca. 45 degrees, nearly straight, joining an equally prominent, shallowly arc-
ing marginal vein near (1-2 mm) the margin, the tertiary veins weaker, alter-
nating with the laterals, branching, arising from the marginal vein; blades stiffly
coriaceaous at maturity, drying gray-green to reddish-brown (under hairs), the
lower surface usually densely tomentose, the upper surface usually sparsely to
moderately pubescent. Flower bud ovoid to pyriform, +-9 mm long; peduncles
solitary in the axils of leaves or bracts, 0.2-3 cm long, 1-1.5 mm wide, normally
uniflorous (rarely triflorous), densely pubescent; bracteoles linear to narrowly
elliptic, 5-8 mm long, 1-1.5 mm wide, densely pubescent, caducous at about
1454 BRIT.ORG/SIDA 20(4)
anthesis; calyx open, prolonged ca. 1 mm beyond the ovary summit, the lobes
about triangular, |-4 mm long, 1-3 mm wide, densely pubescent within and
without, sometimes less pubescent than the hypanthium; petals suborbicular
to obovate, 8-10 mm long, subglabrous; hypanthium funnel-shaped, 2-5 mm
long, densely tomentose; disk ca. 4 mm across; stamens 70-200, 5-9 mm long;
anthers suborbicular, 0.5-1 mm long; style 6-8 mm long; ovary (2-)3(-4)-locu-
ar; ovules 7-19 per locule, reflexed, usually uniseriate on the margin of a peltate
placenta. Fruit subglobose, 1.2-2 cm long; seeds 5-20, more or less rounded, 4-7
mim long.
Representative specimens. BOLIVIA. Santa Cruz: Velasco Prov. PN. Noel Kempff M., C ae
-as Gamas (14°48'52"S, 60°24'08"W), 900 m, 1 Nov 1995 (fr), Killeen & Grinwood 7819 (A
BRAZIL. Distrito Federal: ca. 20 km NE de Brasilia na estrada Brasilia/Fortaleza, ‘ a 9 1967
cap) 69 (HB): Plano Piloto, Brasilia D.F, 20 Sep 1962 ({1), Heringer 9007 (HB, NY); Horto do
Guara Brasilia D.F, 5 Jan 1962 (fr), Heringer 881021 (HB): Jardim Botanico de Brasilia (15'52'S, 47°50'W),
1070 m, 31 Aug 1995 (st), Poenca etal. 1444 (ASU, UB); Chapada da Contagem, road NE edge of P.N. de
Brasilia, Rua 4, 1160 m, 10 Sep 1995 (ED), Proenca et . 1456 CASU, UB); entre Sobradinho/Planaltina,
KM 20, 3 Sep 1995 (st), Proenca et al. 1473 (ASU, U i Parque Boca da Mata, 3 Sep 1995 (fr),
Rezende 186 (ASU); S side of Campus, 16 Nov 1977 oS Universidade de Brasilia, laxonomy Class 552
(US). Goias: Rod. GO-118, 5-8 kim a W de Alto Paraiso, 1200 m, 16 Oct 1990 ({), Hatschbach 54613
(ASU); Sao Joao da Alianea, 30 Oct 1979 (fr), Heringer 2650 (NY); Cidade Ecletica (50 km de Brasilia),
LO Nov 1973 1D), Heringer 12962 (NY, UB); Serra dos Pireneus, ca. 20 km E of Pirenopolis, 1000 m, 16
Jan 1972 (fr), Irwin et al. 34281 (NY); Mun. de Mineiros (17 28'S, 52°31'W), 800 m, 12 Dec 1983 (fr),
Magnago 334 CARB); Cristalina, ca. 2 km N of turnoff to Cristalina on highway to Brasilia, 1150 m, 9
Sep 1995 (st), Proenca etal. 1447 (ASU, UB); Padre Bernardo, 10 km S$ de Brazlandia (15'43'S, 48°12'W),
13 Nov 1990 (fr), Viera et al. 610 (ASU). Mato Grosso: Chapada dos Guimaraes, just above Cachoeira
Furada, Reserva Buriti, 720 m, 12 Oct 1973 ({D), Prance et al. 18827 (NY); Alto do Araguaia, 15 km NO
de Alto do Araguaia na estrada para es eee 7" pe 53 oe Ww ) 18 Sep 1996 ae Proenca et al. 1526
(ASU, UB). Mato Grosso do Sul: Brasilandia XI rio Pardo (21'40'30"S, 52°40'W), 23 Oct
1981 (CD, Dambros 215 (ARB); 6 kim N pov. ie lara margen esq. estr. que ie se? Uniflora, Gemae
outras (20°24'S, 52°55'W), 12 Nov 1981 (fr), Guimaraes 1339 (HBR, RB); a 6 km do Posto Sao Sebastiao -
Tres Lagoas (20°05, 5US1'W), 26 Jan 1982 (fr), Guimaraes 1367 Ct ne RBC ees. Capao Redondo,
LL Nov 1973 (f(D, Hatsch bach 33063 (MBM); sais Reserva do Exércit oe 30'S, 54°42'W), 19 Sep 1996
(st), Proenga etal. 1550(ASU, UB). Mi lov. Bri aie Belol te, Paracatu, 3 Jun 1960
(LD, Heringer 7566 (UB); Serra do Espinhaco, ca. 48 km W of Montes Claros, 950 m, 25 Feb 1969 (fr),
Irwin etal. 23866 (MO, NY); Morro das Pedras, ca. 37 km NE of Patrocinio, 1000 m, 29 Jan 1970 (fr),
Irwinetal. 25604 (NY); Serra do Cipo, between Veuda Noiva and Alto do Palacio (ca. 1915'S, 43°40'W),
1000-1400 m, 31 Jan 1982 (fr), Landrum 4242 (MBM, NY); entre Cruzilha e Mindurim, 27 Nov 1967
(fr), Mattos 15225 (SP): prox. ao Horto Florestal, 27 Nov 1940 C1). Occhionis.n.(RB): Diamantina. rd.to
Sao Joao da Chapada, 17 km NW of jet. with rd. from Curvelo-Diamantina, 1180 m, 23 Nov 1985 (fr),
Thomas et al. 4868 (ASU). Parana: Parque Vila Velha, Mun. Ponta Grossa, 850 m, 4 Oct 1963 (FL),
Hatschbach 10234 (MBM); Chapadao S. Antonio (Mun. Arapoti), LI Oct 1968 (fD), Hatsch bach 19977
(MBM); Rio SAna, Mun. Cerro Azul, 6 Oct 1977 (1D, Hatschbach 40353 (ARB), Mun. Senges, Rod. PR-
ll, prox, km 252, 18 Nov 1989 ({D, Hatschbach 53638 (ASU). Sao Paulo: Botucatu, 14 km EF of Sao
Manuel (22°45'S, 48°25'W), 550 m, 2 Nov 1973 ({1), Gottsberger L-21173 (ASU); Mun. Moji-Guac¢u: Faz.
Campininha, 9-11 km NNW de Padua Sales, 28 Sep 1960 (fr), Mattos 8339 (SP); 3 km ao sul de Pilar do
Sul, estado de Sao Paulo, 21 Oct 1966 (y{L), Mattos 14088 (SP); Itararé, campos de Sao Pedro na Serra de
Bom Sucesso, Faz. Ventania, OOO m, 21 Oct 1966 (Ir), Mattos 14119 (SP); estrada Itararé - Itapeva, a
ponte do Rio Verde (24°05'00'S, 49'12'06"W), 14 Nov 1994 (fr), Souza et al. 7244 (ASU).
LANDRUM, REVISION OF THE PSIDIUM SALUTARE COMPLEX 1455
COSTA RICA. Puntarenas: on road to Salitre, ca. 2 km from Buenos Aires, ca. 500 m, 11 Sep 1989
(st), Landrum 6560 (ASU); Buenos Aires de Osa, Barrio el Carmen, 22 Jul 1987 (f1), Sanchez & Poveda
1260 (MO).
GU
YANA. Pakaraima Mis; near base of Malakwalai-Tipu (4°48'N, 60°12'W), 700 m, 9 Jul 1994
(fr), sea & hes 5522 (ASU
ARAGUAY. Canindeyu: Teac. Reserva Natural del Bosque Mbaracayu, Nandurocai (ca.
24 ae 55 oe a 19 Nov 1995 (st), Landrum 8860 Sone FCQ).
VENEZUELA. Bolivar: Santa Elena de Uairén, 2-4 Mar 1972 (£D, Bunting & Holmquist 4764 (NY):
Roscio, ca. 2 km NW de San Ignacio de Yuruani (5 O1'N, 61°8'W), 900 m, 20 Jun 1983 ( (fr), tiicbed &
Alarcon 7512 (MO, NY, VEN). Zulia: Sierra de Perija, above Pishikakao, 400-1850 m, 1-3 Apr 1972 (ED.
Steyermark et al. 105766 (MICH).
Psidium rufum DC. (= P. widgrenianum O. Berg), which ranges from Parana to
Goids and Bahia, is sometimes confused with P. laruotteanum. They are con-
trasted in the key below.
. Shrub usually no more than 1.5 m high; hairs grayish, yellowish, or reddish brown,
usually curled and tangled, A ati the lower leaf surface; midvein and lateral
veins flat or es raised above; petiole 0—2(—3) mm long; calyx-lobes sence
or subtriangular, 1-4 mm lo P.laruotteanum
1. Shrub or tree to : m high; ae mainly reddish brown, mostly erect, mee not
Seas lower leaf su are ;midvein and often lateral veins impres
petiole (2-)3 lobes mainly broadly rounded,0.5-2 mm ne
triangular and up to 4mm long) P.rufum
Psidium laruotteanum is variable as to le - size and shape (Fig. 1). Smaller plants,
especially new sprouts, tend to have small elliptic leaves and larger plants tend
to have large oblanceolate or obovate leaves. Since a similar pattern is seen
throughout the range, I do not suspect a genetic basis for this difference.
Berg cites the type of Psidium acrugineum var. angustifolium as being at
LE but cites no collector or locality. A specimen at LE, collected by Sellow, an-
notated by Berg as Psidium aerugineum, | considered to be an isotype of that
taxon. I suspect that this same specimen is the holotype of Psidiumaerugineum
var. angustifolium, but it was never annotated by Berg as such. It has relatively
narrow young leaves, which Berg mentions in the protologue of var.
angustifolium. I have found two other cases where Berg described a taxon at LE
but seems not to have annotated a specimen (Psidium riedelianum O. Berg, in
Mart., Fl. bras. 14(1):602. 1859) and P. pohlianum var. brevipes discussed below
under P. salutare var. pohlianum.
Psidium salutare (H.B.K.) O. Berg, Linnaea oo 356. 1856. ies eae iS ELBA.
Nov. gen. sp. 6:132. 1823. TyPE: “Carichanam, ad ripam Ori ” Humboldt &
Bonpland s.n. (HOLOTYPE: P, = F-36905! ISOTYPE: eB. =B1263/11 ae at MICH)).
Subshrub or shrub up to ca. 1.5 m high (often less than 0.5 m high), with new
shoots arising from a woody subterranean base or rhizome, with shoots often
short lived, or in one variety sometimes reaching tree size (up to 10 m high),
glabrous, glabrous except for disk and calyx-lobes within, or sparsely to mod-
1456 BRIT.ORG/SIDA 20(4)
erately pubescent on young growth, or silvery lanate in one variety; hairs when
present whitish, 0.3-1 mm long; young twigs glabrous to densely pubescent,
reddish-brown, becoming grayish, the older bark gray to reddish-brown, be-
coming flaky. Leaves opposite or alternate on some shoots (rarely ternate), ovate,
lanceolate, elliptic, narrowly elliptic, obovate, oblanceolate, 1-)2-9 em long, 0.6-
5.5 cm wide, 14-5 times as long as wide, the margin entire to somewhat revo-
lute; apex obtuse, acute to acuminate, abruptly acuminate, sometimes apicu-
late; base cuneate, obtuse, or rounded; petiole 0-2 mm long, |-1.5(-2) mm wide;
midvein normally flat or slightly raised above, prominent below, the venation
brochidodromous, with 5-12 pairs of lateral veins, these prominent to scarcely
visible, leaving the midvein at an angle of ca. 45 degrees or less, nearly straight,
joining a clear shallowly arcing, equally prominent marginal vein near (0.2-2
mm) the margin, the tertiary veins forming a dendritic pattern between the
laterals; blades stiffly coriaceous at maturity, drying reddish-brown to gray-
green, dull or lustrous above, the cellular pattern sometimes visible with a dis-
secting scope. Flower buds pyriform, +-7 mm long; peduncles, axillary,
uniflorous or triflorous, 0.4-3.5 cm long, 0.5-0.8 mm wide; bracteoles linear to
lanceolate, deciduous or persisting, 2-9 mm long, 0.5-2 mm wide; calyx open,
cup-like, tearing ca. 1 mm between the lobes at anthesis, the lobes broadly
rounded to ovate-triangular, 0.5-5(-6) mm long, 2-3 mm wide, glabrous to lanate
without, usually puberulent (less often glabrous) within; petals obovate to sub-
orbicular, 5-11 mm long; hypanthium obconic to campanulate, 2-4 mm long,
extended beyond the ovary summit ca. | mm; disk 3-4 mm across, puberulent
or glabrous; stamens 100-200, 5-12 mm long; anthers subglobose to oblong, 0.3-
0.8 mm long; style 5-8 mm long; ovary 2-3-locular; ovules 9-48 per locule,
uniseriate or biseriate along edge of the placenta, this strongly to scarcely peltate.
Fruit globose to subglobose, 8-10 mm in diam.; seeds 4-20, 4-8 mm long,
subovoide.
Psidium salutare is a widespread and variable species. There are a few
recognizable forms that correspond to geographic regions that I recognize as
varieties, but the limits between these varieties are often unclear. Legrand and
Klein (1977, p. 723) also noted difficulty in distinguishing these entities, but
chose to retain them at the specific level. One might speculate that these mor-
phological types represent adaptive syndromes of characters for somewhat dif-
ferent niches. In-depth studies of the entities, including population samples,
field studies, estimates of genetic distinctness and exchange, and transplant
experiments should prove interesting in the future, especially for sympatric
varieties, ¢.g., var. sericeum and var. mucronatum, or var. decussatum and var.
pohlianum. As recognized here, there is still considerable variation in var.
salutare to which specimens unassignable to any of the other varieties are ten-
fou
tatively placed.
LANDRUM, REVISION OF THE PSIDIUM SALUTARE COMPLEX 1457
Psidium salutare var. salutare
Psidium salutare (HBK) O. Berg, Linnaea 27:356, as to type. 1856. (Figs. 2E, 4, 5).
Myrtus arayan HBK, Nov. gen. sp. 6:133. 1823. Type: PERU [ECUADOR]: “prope Gonzanamam
eruvianorum, ad ripam fluminis Catamayo,” Humboldt & Bonpland s.n. (HOLOTYPE: P=
36874).
Psidium ciliatum Benth., in Hook. J. bot. 2:318. 1840. Type: BRITISH GUIANA: “dry savannahs,”
Schomburgk s.n. (HOLOTYPE: K; ISOTYPES [Schomburgk 365], P!, = ASU photo!, W, = F-
ty eee Benth., Plantas Hartwegianas, 174. 1845. TYPE: MEXICO: “Popayan,” Hartweg
977
Qu
a
ay
(HOLOTYPE: K, = ASU photo!).
Radium A. Rich., Ess. Fl. Cub. 581.1846. TYPE: CUBA: “Vuelta de Abajo,” J.M. Valenzuela
s.n., (2 sheets at Pl, “TYPE” LECTOTYPE, here designated], = ASU photo!).
Psidium oe var. subalternum O. Berg, Linnaea 27:357. 1856. TYPE: BRITISH GUIANA: “ad
flu acutu,” Rich. Schomburek 498 & 1252; Rob. S. 365 (SYNTYPES B, lost, ISOSYNTYPES
soimabieh 365], PI[LECTOTYPE, here designated], W, = F-31434! = ao photo).
Psidium oerstedeanum O. Berg, Linnaea 27:360. 1856. TyPE: COSTA RICA & saci {AL _
“Provincia Guanacaste (Oersted) & Rincon in Guatemala dich: eds.)
Friedrichsthal 1226(SYNTYPES W and “hb. Oersted. nol6”; SYNTYPE: C[Oerst ne sa
Psidium salutare var. laxum O. Berg, Linnaea 27:357. 1856. TYPE: TENEZUE LA: One ad Up
Otto 987 (HOLOTYPE: B, lost; ISOTYPE: LE [LECTOTYPE, here designated], = ASU photo!; sot
ISOTYPE: W = F-31435).
Psidium salutare var. stricta O. Berg, Linnaea 27:356. 1856. Inadmissible name to be replaced by P.
sdalutare var. salutare.
Myrtus rigida O. Berg, in Mart., Fl. bras. 14(1):417 1857. TyPE: BRAZIL: “ad Paranapitanga distr.
— prov. S. Pauli,” Sellow s.n. (HOLOTYPE: B, lost; ISOTYPE: P![LECTOTYPE, here designated],
31, ASU photo ol)
Moras Mancheton O. Berg, in Mart. FI. bras. 14(1):418. 1857. TyPE: BRAZIL: “Bahia,” Blanchet
B, lost; ISOTYPES, W, G = F-23481), P![LECTOTYPE, here designated], = ASU photo!).
Myrtus sagraea O. Berg, Linnaea 30:710. 1860. TYPE: CUBA: without locality, De la Sagra s.n. (HO-
LOTYPE: P!, =ASU photo!)
asa lanceolatum O. fae Linnaea 30:704. 1861. TyPE: BRAZIL: without locality, (HOLOTYPE:
ASU photo!).
mes: ear var. oblongata Griseb., Cat. pl. Cub. 91. 1866. TYPE: ee without locality.
hi oe 2436 (HOLOTYPE: GOET, IsoTYPES, MICH!, MO!, NY!, = ASU photo!
Psi abita var. angustifolia ee Cat. pl. Cub. 91. 1866. TYPE: Ae Bat locality,
Wr ight 2436a (HOLOTYPE: GOET
Calycolpus parviflorus Sagot, Ann. Sci. Nat. (Paris) VI. 20:181.1885. TYPE: FRENCH GUIANA: with-
out locality, Leprieur s.n. (HOLOTYPE: Pl, 2 specimens, [*TYPE” LECTOTYPE, here designated], =
ASU photo!).
Psidium deltosepalum Barb. Rodr. ex Chodat & Hassl., Bull. Herb. Boissier 7:799. 1907. TYPE: PARA-
GUAY: “pr. a Capibary,” Hassler 4400 (HOLOTYPE: G!, 3 specimens, =ASU photos!
ISOTYPES, P)).
Psidium val l Barb. Rodr. ex Chodat & Hassl., Bull. Herb. Boissier 7:798. 1907. TYPE: PARA-
cae og Valenzuela,” Hassler 6947 (HOLOTYPE: Gl, 2 specimens; ISOTYPE: NY).
wrayan (HBK) Burret, Notizbl. Bot. Gart. Berlin-Dahlem 15:484. 1941.
aaa Lundell, Amer. Mid. Nat. 29:483. 1943. Type: BELIZE: TOLEDO Dist: “Monkey River,
Jenkins Creek,” 1 Aug 1942 (fr), Gentle 4062 (HOLOTYPE: MICH: IsoTYPES MO], NY).
Myrcianthes reptans D. Legrand, Bol. Univ. Parana Fac. Farm. 27:1- 3 1971. Type: BRAZIL: Parana
“Mun. Palmeira, Faz. S. Amelia,” Hatschbach 17697 (HOLOTYPE: MVM; ISOTYPE: HB!, =ASU photo!).
1458 BRIT.ORG/SIDA 20(4)
oA
“ .
Fic. 2. Psidium salutare young fruits. A, Irwinet al. 10699 (NY), var. decussatum. B,
var. mucronatum.C, Irwin et al. 9101 (NY), var. pohlianum. D, Venturi 7582 (F), var. sericeum.E, Burch
6180 (NY), var. salutare:
occ iNV\
Usually a subshrub less than 0.5 m high; leaves elliptic, lanceolate, oblanceolate,
ovate, or obovate, 3-7 cm long, |-3.3.cm wide, 1.6-3.2 times as long as wide, gla-
brous to moderately pubescent; venation obscure to moderately pronounced,
the marginal vein usually about | mm from margin; apex usually without an
apiculum; peduncle 1-5 cm long, uniflorous or triflorous; calyx-lobes shorter
or longer than the hypanthial tube plus calyx tube, acute to rounded.
LANDRUM, REVISION OF THE PSIDIUM SALUTARE COMPLEX
1459
1000%
9 200 400 600 800 m
+ T T
(eames T 1
QO 100 200 300 400 600 600 miles
eu 20
40. —
Fic. 3. Distribution of Psidium laruotteanum.
Representative specimens. BELIZE. El Cayo: Mountain Pine Ridge, San Agustin, Jul -Aug 1936 (fr),
Lundell 6560 (MICH, MO, NY); upper 200 m of Baldy Beacon, ca. 1000 m, 10 Jul 1970 (fr), Spellman
1647 (MO).
BOLIVIA. Santa Cruz: PN. Noel Kemplf Mercado, Los Fierros (14°3620"S, 60°51'30"W), 200 m,
26 Oct 1993 (fr), Killeen et al. 5943 (ASU): PN. Noel Kempff Mercado, 6 km NE del campamento Las
Gamas (13°53'41'S, 60°48'46"W), 850 m, 28 Oct 1995 ([]), Rodriguez & Surubi 549 (ASU).
BRAZIL. Ceara: Chapada do Araripe. ca. 15 km SW of Crato on BR-122 (7°30'S, 39°35'W), 860 m,
9 Jan 1983 (fl), Plowman & Tavares Cacula 12704 (F). Distrito Federal: margem do lago Paranoa, 9 Nov
1978 (fr), Heringer 701 (NY). Goias: Corubaiba, Rio Corumba, Foz do Corrego (17°48'S, 48°21'W), 21
Sep 1993 (FL), Da Silva et al. 1880 (SP); Serra do Caiapo, 35 km S of Caiaponia on road to Jatai 7'12'S,
51°47'W), 800-1000 m, 29 Oct 1964 (I), Irwin et al. 7552 (CAS); Serra dos Cristais, ca. 1O km W of
Cristalina (17'S, 48'W), 1200 m, 4 Mar 1966 (fr), Irwin et al. 13449 (MICH, MO, NY). Mato Grosso:
Barra do Gareas, 210 km along new road NNE of village of Xavantina, Corrego do Gato, ca. 450 m, +
Oct 1968 (fl), Eiten & Eiten 9058 (US). General Carneiro, Meruri [Meruré], Sep 1963 (f(D, Hartmann
369 (SP); | km NE of Garapt (13° 12'S, 52° 34’'W), 300-400 m, 1 Oct 1964 (FD), Irwin & Soderstom 6506
(MICH, MO, NY, RB); Mirassol D'Oeste, entre Caceres e Porto Esperidia, 23 Oct 1983 (fD, Saddi 3477
(RB). Minas Gerais: Sa |
Sac Preto, P E. do Rio Preto (ca. 18°6'S, 43°20'W), 20 Feb 2002 (fr),
U): Beltim, Serra da Caveira, 1600 m, 1] Mar 1945 (fr), Williams & Assis 6227 (MO).
Para: Martins Pinheiro, Campina do Mangaba, 28 Feb 1975 (yfr), Coradin 141 (MICH); Maracana, ca.
73 airline km NE of Castanhal, Martins Pinherio (0°52'S, 47°35'W), 50 m, 6 Apr 1980 (fr), Davidse
L ] 13 A5SQ22(AC
OTTLE/QEAL
1460 BRIT.ORG/SIDA 20(4)
110 a =
an y _—
s Z
ae oe :
aa
°
30
°
, -
ark, ‘ ee
s woeoeeeree “-
) _ eee fee
7 : ee :
a, °
eaters e : a
ae = . ° = a
en _ o o? . ;
— ee eee, ,
—_ sail ; 7
20|-— ‘ .
Le. Pp
/
ae o@ Seis
oa
© 200 400 800 800 1000km ..
i 0 0
|
0 100 200 300 400 500 800 miles ; :
°
——a ee ° :
— “ \ ol
a ~ :
_ —— | /
Fig. 4. Distribution of Psidium salutare var. salutare in northern part of range.
17923 (NY). Parana: Lapa, Rio Passa Dois, 900 m, 5 Oct 1958 (fl), Hatsch bach 5048 (MBM); Palmeira
Rod. Do Caté, Rio Tibagi, 780 m, 18 Nov 1962 (1), Hatsch bach 9636 (MBM): Campo Moura, 14 Oct 1965
({D, Hatsch bach 12997 (MBM); Balsa Nova, Barra do Rio dos Papagaios, 14 Mar 1968 (fr), Hatschbach
13749 (MBM), Ponta Grossa, Vila Velha,7 Oct 1969 (f{D, Hatsch bach 22326 (MBM); Guarapuava, Entre
Rios, 21 Oct 1969 (£D, Hatschbach 22551 (MBM); Jaguariaiva, Lageado 5 Reis, 18 Sep 1975 (f1),
Hatschbach 37098 (MBM), Alm. Tamandaré Rodovia dos Minerios, Rio Barigui, 9 Feb 1982 ([1),
Hatschbach 44564 (MBM): Palmeira, Faz. Sant; — ca.65 km W of Curitiba (ca. 25'25'S, +9°50'W), 2
Dec L981 (ED, Landrum 3966 (MBM, NY): Rio Br l.along road to Cerro Azul (ca. 25'S, 40'20'W),
7 Jan 1982 (fr), Landrum 4112 (CAS, MBM, MICH, MO, NY). Rio Grande do Sul: Fazenda Faxinal.
Arroio dos Ratos, 13 Apr 1982 (f{D), Hagelund 13890 (CTES). Santa Catarina: Campo Eré, 24 km W ol
Campo Eré Q6° 22'S, 53° L'W), 900-1000 m, 7 Dec 1964 (f1), Smith & Klein 13826 (MICH, NY).
Tocantins: Arraias, km 15-20 a W na rod. para Parana, 10 Nov 1991 (f1), Hatsch bach 56024 (MBM).
COLOMBIA. Caqueta: San Vicente del Caguan, laguna “El Retorno,” (74'17.0'W, 1'5.43'N), 265
m, 26 Jan 1990 ({D), Betancur & Porras 1517 (MO). Los Llanos: Boyaca, La Comarca, ca. 15 km E of
Barranca de Upia, 300 m, 12 Feb 1939 (£1), Haught 2602 (MICH). Sta. Marta: Onaca, 2500 ft, 28 Aug
1898 (1D, Smith 2509 (NY). Vichada: Parque Natural “El Tuparro,” ca. 11 km NE of Fl Tapon on rd. to
Centro Administrativo (5'12'N, 69°4'W), 18 Mar 1985 (fD, Zarucchi & Barbosa 3749 (ASU),
COSTA RICA. Guanacaste: entre La Cruz et Liberia, 30 Jan 1909 (11), Brenes s.n. (NY): Lomas
Barbudal, Bagaces, 100 m, 10 May 1984 (fr), Gomez & Herrera 23021 (ASU); 7 km NW of Bagaces,
Comelco Ranch, 29 May 1971 (fD, Heithaus 128 (MO); Santa Rosa National a across road from
entrance (10° 53'N, 85° 35'W), 250-300 m, 28 Jun 1977 ({l), Liesner 2700 (MO): al norte de la ciudad de
Liberia, camino al Parque Nac. Rincon de la Vieja, 2 Oct 1985 (fr), Gomez 23687 (MO).
JBA. Isla de la Juventud (Isla de Pinos): La Canada, 16 Feb 1916 ({D, Britton et al. 14401 (NY):
Sierra de Los Caballos, 2 Mar 1916 ({1), Britton 15160 (NY,US): near Nueva Gerona, 1904 (f1, fr), Curtiss
350 (MO, NY, US). Pitar del Rio: vicinity of Herradura, 26 Aug 1910 (fr), Britton et al.6450 (NY): Pinar
del Rio to Vinales, 200 m, 12 Sep 1910 (fr), Britton 7309 (NY); near el Guama, 9 Mar 1900 (f1), Palmer &
Riley 212 (US); Laguna Jovero to Laguna Herradura, 12 Dec L911 (st), Shafer 10922 (NY): Arroyo del
Sumidero, 7-9 Aug 1912 ({D), Shafer & Br Leén 13680 (NY); San Juan y Martinez, 7 May 1988 (fD,
Urquiola et al. 4522 (ASU): Mantua, 8 Nov 1990 (fr), Urquiola et al. 6678 (ASU).
=
LANDRUM, REVISION OF THE PSIDIUM SALUTARE COMPLEX 1461
Fic. 5. Distributi f Psidi lut. lutare in South A ica ( ircles) iP. sal sericeum (dots)
DOMINCAN REPUBLIC. Sanchez Ramirez: Cotui, 6 Sep 1952 (fr), Jimenez 2433 (MICH, US).
ECUADOR. Loja: Zambi, camino a Tablaso (3°55'S, 79°30'30"W), 1500 m, 18 Mar 1995 (st), Eynden
& Cueva 294 (ASU, QCA).
EL SALVADOR. Ahuachapan: 1923, Padilla 633 (US).
UATEMALA. Chiquimula: Volcan Ipala, near Amatillo, 900-1510 m, 25 Oct 1939 ED, Steyermark
30553 s
>
GUYANA. Rupununi Northern Savana, Annai Hillside, stand 37, ca. 350 ft, 7 Oct 1963 (fD,
Goodland a3 (NY); Potaro-Siparuni, Pakaraima Mts., Tay-klay-o Creek, upper Ireng River (4°50'N,
59°58'W ), 650-825 m, 19 Jan 1993 (yfr), Henkel 880 (ASU); 7 km SW of Aishalton, Boidkorodai Mt, S
Rupununi Savana (2°25'N, 59°20'W), 220 m, 18 Nov 1993 (fr), Henkel 3479 (ASU); Kanuku Mts.,
Rupununi R., Bush Mouth near Witaru Falls (3" 4'N, 59°28'W), 90-120 m, 9 Feb te eee Hae
et al.87 (ASU, MO); NE from Karasabai, to Yourora Creek (4°03'N, 59°30'W), ca. 100 m, 7 Mar
McDowell 2154 (ASU).
1462 BRIT.ORG/SIDA 20(4)
HONDURAS. Comayagua: Siguatepeque, Escuela Nac ronal de Ciencias Forestales, 26 May 1972
(f1), sath 6180 (EAP, MO, NY). El Paraiso: Guinope, 2.5 mi S of the intersection of El Zamora ano-
Morolica and Guinope rd (13°51'50"N, 86°58'43"W), 1370 m, enn 1904 (I1), Davidse et al. 35004 (A
between Manzaragua and San Lucas, 1350 m, 22 Sep 1968 (fr), Molina 22698 (EAP. NY); 10 mi S sl
Yuscaran, ca. 1700 ft, Jul 1962 (yfr), Webster 12011 (MICH): 5 km E of Ojo de Agua, 760 m, 4 Sep 1946
(fr), Williams & Molina 10484 (EAP. MICH, MO). Gracias a Dios: Caserio de Rus-Rus, vaguada de Rio
Rus-Rus, Jul 1977 (fr), Nelson & Romero 4126 (EAP, MO). Franciseo Morazan: road from Comayagua to
Tegucigalpa, 6.7 km SW of Parque Aurora, near KM 27, ca. 1300 m, 27 Aug 1989 (fr), Landrum 6480
(ASU), ca. 25 km NW of Tegucigalpa on road to Comayagua, ca. 1000 m, 27 Aug 1989 (fr), Landrum
6520 (ASU).
MEXICO. Chiapas: Concordia, 543 m, 16 Jun 1945 (fl), Matuda 5905 (MO): Rocky plains,
Monserrate, Jun 1925 (FI), Purpus 10306 (US). Guerrero: Agua del Obispo, 34 km W ol Chilpanginge
on road to Acapulco, ca. 900 m, 23 Jul 1989 (fr), Landrum & Landrum 6338 (ASU, MEXU). Nayarit:
Mina Esperanza Rosa Morada, ({D, Ortega 6658 (US); foothills between Acaponeta y Pedro Pablo, 2
Aug 1897, Rose 1950 (US). Veracruz: Ejido Fl Nigromante, 6 Oct 1971 (st), Chavelas et al. 4230 (MEXU).
ICARAGUA. Chontales: Hda. Veracruz (12° 11-12'N, 85 21-22'W), 120-475 m, 4-6 Aug 1983
(£D, Stevens 22419 (MO). Nueva Segovia: Monte Rico, 5 kmal NE de El Jicaro (13° 44'N, 86° 5'W), ca 705
m, 3 Sep 1984 (fr), Moreno 24611 (MO)
PANAMA. Toboga Island, Gulf of Panama, 0-250m, 20 May 1911 (fL), Pittier 3581 (US). Canal
Zone: Ancon Hill, Nov-Dec 1923 (ED, Standley 26340 (MO, US). Chiriqui: Boquete, 4000 ft, 29 Jun
1938 (1D, Davidson 835 (MO). Panama: hills NE of Hacienda La Joya, 50-300 m, 9 Dec 1934 (1), Dodge
etal. 16899 (MO); between Rio Pacora and Chepo, 3 Apr 1969 (fl), Dwyer 5101 (MICH)
RAGUAY: Caaguazu: Ea. La Esmeralda, 11 Dec 1982 ({r), Schinini 22902 (CTES, MICH).
VENEZUELA: Apure: Pedro Camejo, near mouth of Cano San Miguel (67°17'W, 6'33'N), 38 m, 29
Apr 1977 (1), Davidse & Gonzalez 12373 (MICH, MO); 11 km directly E of Paso de San Pablo, along
banks of Rio Capanaparo (67 39'W, 7°2'N), 45 m, 8-9 May 1977 (Ir), Davidse & Gonzalez 12953 (MICH,
MO, VEN). Bolivar: Cedeno, Foraneo La Urbana, 30 m, 2 Feb 1989 ({1), Cuello 743 (MO): Caroni, Puerto
Ordaz (817'N, 62°55'W), 30 m, 20 Nov 1997 (I), Diaz & Marin 3384 (ASU); Mun. Asc. Farreras, Maripa-
Aripao (7 29'N, 65:20'W), 80 m, Feb 1990 (fr), Elcoro 686 (MO); Piar, Cerro Tomasote (7°48'N, 62°02'W),
540 m, May 1986 (fD, Fernandez 2742 (MO); Represa Guri, 1-5 km S of dam (7'45'N, 63°0'W), 20-240
m, | Apr 1l98L (ID, Liesner & Gonzalez 11064 (ASU, MO, VEN): km 11 on Puerto Ordaz-Cerro Bolivar
Railroad, 300-350 m, 26 Oct Chiguao (6°32'N, 63'8'W), May Jun 1987 (£1), Stergios 11113 (MO): E ol
Miamo, Altiplanicie de Nuria, between Hato de Nuria and camp, +00 m, 23 Jan 1961 (fr), Steyermark
88711 (NY, VEN); entre San Felix y Puerto Ordaz, opposite the Fabrica de Alfareria, 26 Jun 1964 (fr),
Steyermark 9429] (MICH). Guarico: Est. on de los Llanos, ca. 12 km SE de Calabozo (8'56'N,
67 25'W), 75m, 15 May 1987 (Ir), Ramirez 2182 (MO). Mérida: | Beguilla and Mucuchachi, 1065-
2430 m, + May 1944 (fl, fr), Steyermark 56312 (NY, VEN). Portuguesa: Llanos de Araure, 9 Apr 1925
({1), Pittier 11748 (NY). Sucre: Cumbre de Montana de Mochima, 18 km SE de Cumana (10°20'N, 64°
20'W), 350-400 m, 16 Sep 1973 Ur), Steyermark et al. 108604 (MO, VEN). Zulia: Miranda, entre la via
El Consejo-El Pensado y el Limite con el Edo. Falcon, 21 May 1980 (f1), Bunting & Stoddart 9238 (NY).
pms
Psidium salutare var. salutare (P. salutare in the traditional sense) is sometimes
contused with P. oligospermum (=P. sartorianum). Their ranges are similar, both
extending to central Mexico and the Caribbean and as far south as southeast-
ern Brazil and Paraguay. They are compared in the key below.
1. Calyx open in the flower bud (closed corolla clearly visible); calyx lobes either evi
dent or the margin of the calyx merely sinuate; subshrubs of open grasslands or
scrub vegetation; petals 5-11 mm long; hairs when present ca. 0.5 mm long.
P. salutare var. salutare
LANDRUM, REVISION OF THE PSIDIUM SALUTARE COMPLEX 1463
eae.
1. Calyx closed in the flower bud (closed corolla hidden); calyx calyptrate or with ter-
minal lobes (appearing puckered); shrubs to large trees, usually of forested habi-
tats; petals 3-6 mm long; hairs when present up to ca.0.2 mm long. . oligospermum
Psidium salutare var. salutare as understood here ranges from Paraguay to the
Caribbean and Mexico and is quite variable. There are two local forms that might
merit recognition as varieties: |) Populations in Cuba have the calyx-lobes con-
nate about 2-3 mm beyond the staminal ring; these have long gone under the
name P guayabita. 2) In Parana to Rio Grande do Sul, Brazil and Paraguay there
are populations with rather large elliptic to obovate, thinly pubescent leaves
and include the types of Myrcianthes reptansand Psidium valenzuelense. Simi-
lar specimens occur in Central America, so it appears that P salutare var. salutare
is a morphologically flexible entity throughout its range.
Psidium salutare var. decussatum (DC.) Landrum, comb. nov. (Figs. 2A, 6). Psidium
decussatum DC., Prodr. 3:235. 1828. TYPE: BRAZIL: “prov. Minarum,” Martius s.n.
(HOLOTYPE: M, = F-19710}, = ASU photo!).
Usually a subshrub less than 0.5 m high; leaves elliptic to narrowly elliptic, ()
2-5.3 cm long, 0.7-1.7 cm wide, (1.5-)2.5-4.5 times as long as wide, glabrous
(rarely sparsely pubescent); venation obscure, the marginal vein about 0.5mm
from margin; apex without an apiculum; peduncle 0.5-L5 mm long, uniflorous
(rarely triflorus); calyx-lobes shorter or about as long as hypanthial tube plus
calyx tube, acute or rounded.
Representative sce out Distrito Federal: 12 kin W of Taguatinga, 1250 m, 26 Nov 1905 (1), Irwin
et al. 10699 (NY); 2 km E of Lago Paranoa, DF-6, 1000 m, 26 Feb 1970 (fr), Irwin et al. 26671 (NY);
pada da Contagem, rd. NE edge Parque Nac. de Brasilia, Rua 4, 1160 m, 10 Sep 1995 (st), Proenca et
al. ae (ASU, UB). ao lado da Reserva Biologica das Aguas Emendadas, 3 Sep 1995 (st), Proenca et a
1496 (ASU, UB). Goias: 10 km W of Cristalina (17'S, 48°W), 1200 m, 5 Mar 1966 (fr), Irwin et al. 13548
(NY). Minas Gerais: Morro das Pedras, ca. 37 km NE ol Patrocinio, 1000 m, 29 Jan 1970 (fr), Irwin eta
25600 (NY). Parana: Colombo, Capivari, 4 Nov 1971 (fr), Hatsch bach 27711 (MBM), Rio Branco do sul,
along road to Cerro Azul (ca. 25'S, 49°20'W), 7 Jan 1982 (fr), Landrum 4110 (MICH, NY); Bocaitva do
Sul, Serra da Bocaina, 16 Jan 2001 (fr), Ribas & Barbosa 3166 (ASU).
Oo
—
If this entity is recognized at the specific level the name Psidium decussatum
DC. should be used. It seems to freely inter;rade with var. pohlianum near
Brasilia.
Psidium salutare var. mucronatum (Cambess.) Landrum, comb. nov. (Figs. 2B,
6). Myrtus mucronata Cambess., in Saint-Hilaire, FI. Bras. merid. 2:295. 1833. TyPE:
Saint-Hilaire s.n. “ad ripas fluminum Rio de la Plata et Uruguay in provincia
Cisplatina,” Martius s.n. (three specimens, P!, all marked as ISOTYPE, one= F neg.
364 361, [LECTOTYPE, here designated]).
Myrtus lurida Spreng., Syst. Veg. 2:480. 1825. Type: URUGUAY: “Monte Video,” Sello s.n. (HOLO-
TYPE: B?, probably lost).
pee poe oe in Saint-Hilaire, Fl. Bras. merid. 2:296. 1833. TyPE: “Encapamento do
parte occidentali provinciae Cisplatinae,” Martius s.n. (HOLOTYPE: PI, =
F-36439), = ASU photo!).
1464 BRIT.ORG/SIDA 20(4)
500 £00 MILES
peppy
| ° 200 400 oo 800 KILOMETERS
| |
/
Fic. 6. Distributi f Psidi I hli ( ircles), P salutare var. d ( ), and P salutare
var. mucronatum (dots).
Myrtus cuspidata O. Berg, in Mart, FI. bras. 14(1):415. 1857. Type: URUGUAY: “in Montevideo,”
Sellow s.n. (SYNTYPES|ty pes of the varieties] B, lost: ISOTYPE of Myrtus cuspidata var. pentamera
at SG oly ECTOTYPE, here designated], = ASU photo).
i r pentamerd O. Berg, in Mart. Fl. bras. 14(1):415. 1857. Type: URUGUAY: “in
" Mourevideo® Selle sn. (HOLOTYPE: B, lost; ISOTYPE: SGO!, = ASU photo!). Name to be replaced
with Myrtus cuspidata var. cuspidata.
LANDRUM, REVISION OF THE PSIDIUM SALUTARE COMPLEX 1465
Mvyrt ida
var. tetramera O. Berg, in Mart., Fl. bras. 14(1):415. 1857. Type: URUGUAY: “in
” Montevideo? Sellow s.n. (HOLOTYPE: B, lost; ISOTYPE: SGO! [LECTOTYPE, here designated], = ASU
photo!).
Myrtus mucronata var. perforata O. Berg, in Mart., Fl. bras. 14(1):416. 1857. Illegitimate name to be
fo}
replaced by M. mucronata var. mucronata.
Myrtus mucronata var. opaca O. Berg, in Mart., Fl. bras. 14(1):416. 1857. Type: URUGUAY: “in
Montevideo,” Sellow s.n. (HOLOTYPE: B).
Myrtus ovalis O. Berg, in Mart, Fl. bras. 14(1):417 1857. TyPE: URUGUAY: “in Montevideo,” Sellow
$.n. eee OTYPE: B).
{ ffrut O. Berg, in Mart., Fl. bras. 14(1):418 1857. pa URUGU aS “in Montevideo,”
oe n. es aa the v B, ae ISOTYPE of Myrtus suffruticosa var. latifolia
at P![lectotype, here designated], = ASU p
Myrtus ay ae var. latifolia O. Berg, in FL. bras. 14(1):418. 1857. Type: URUGUAY: “in
Montevideo,” Sellow s.n. (HOLOTYPE: B, lost; ISOTYPE: P! [LECTOTYPE, here designated], = ASU
ase
iticosa var. angustifolia O. Berg, in Matt. Fl. bras. 14(1):419. 1857. TyPE: URUGUAY:
nm Monrevileo? Sellow s.n. (HOLOTYPE: B,
an tusacutata O. Berg, in Mart., Fl. bras. ser 5. 1857. TYPE: URUGUAY: “in Montevideo,” Sellow
s.n. (HOLOTYPE: B, lost; IsoTYPEs, P [two specimens] , one = F-30424! [LECTOTYPE, here desig-
nated).
Myrtus sellowiana O. Berg, in Mart., Fl. bras. 14(1):413. 1857. TYPE: BRAZIL: “ad Tapanhoacanga
prov. Minarum,” Sellow s.n. (HOLOTYPE: B, lost; ISOTYPE: P!, = F-36445! [LECTOTYPE, here desig-
nated], = ASU photo!).
Saran, thea Griseb., Pl. lorentz. 91. 1874, and in Goett. Abh. 19:139. 1874. TYPE: ARGENTINA:
Cordoba, in monte Cerro negro pro. San Bartolo., Tucuman,” Hieronymus? (HOLOTYPE: prob-
ably GOET, possible isorype: P = F-30422).
Myrtus mucronata var. thea (Griseb.) Griseb. Abh. Konig]. Ges. Wiss. Gottingen 24:127. 1879.
Psidium luridum (Spreng.) Burret, Notizbl. Bot. Gart. Berlin-Dahlem 15:484. 1941
Psidium pubifolium Burret, Notizbl. Bot. Gart. Berlin-Dahlem 15:484. 1941, New name for Myrtus
ovalis O. Berg.
Psidium luridum var. cinereum Mattos, Loefgrenia 64:2. 1975. Type: BRAZIL: Santa Catarina
“Campo Eré, 17 km W of Campo Eré, ca. 26’ 22'S, 53° 08' W, 900-1000 m,” Smith & Klein 13807
(HOLOTYPE: HBR, ISOTYPE: R!, = ASU photo).
estan pubifolium f. nanum Rotman, Darwiniana ieee 1976. TYPE: BRAZIL. Santa Catarina,
Campo Eré, 37 km W of Campo Eré, ca. 26° 22'S, 53" 08'W, 900-1000m,” Smith & Klein 13807
(HOLOTYPE: MVM, ISOTYPE: RI, = ASU photol).
Usually a subshrub less than 0.5 m high; leaves mostly narrowly elliptic to lan-
ceolate, (1.5-)2-6 cm long, 0.7-2.3 cm wide, (1.4-)1.5-5 times as long as wide,
glabrous to subglabrous, often lustrous; venation pronounced, raised on both
surfaces, the marginal vein usually within | mm of the margin; apex apiculate;
peduncle often over 2 cm long, uniflorous; calyx-lobes usually longer than
hypanthial tube plus’calyx tube, usually acute.
eae specimens. ARGENTINA. Corrientes: 2.5 kim de la ciudad de Monte Caseros, 28 Dec
1968 (fr), Carnevali 1317 (CTES); Paso de Los Libres, Bonpland, costa rio Uruguay, 19 Jan? 1945 (fr),
Ibarrola 2135 (NY); Santo Tomé, 33 km N de Santo Tomé, 27 Jan 1976 ({D, Krapovickas & Cristobal
28934 (CTES, NY); Ituzaingo, Rincon Ombti Chico, 3-5 Jul 1974 (£D, Krapovickas et al. 25477 (CTES,
MICH): Mercedes a Ita Cora, Ayo. Pay-Ubre, dep. Mercedes, 2 Feb 1974 (fr), Quarin & Gonzalez 2069
1466 BRIT.ORG/SIDA 20(4)
(CTES), Beron de Astrada, 15 km W of Ita Ibaté, Ayo. Santa Isabel, 16 Jan 1977 (1D), Schinini 14105
(CTES), Estacion Experimental INTA, Dep. Empedrado, 7 Dec 1978 (f1), Schinini 16225 (CTES); Co.
Nazareno (Co. de Susini), 15 Feb 1979 (fr), Schinini et al 17192 (CTES). Entre Rios: Santa Ana, dep.
Federacion, 15 Oct 1968 (f{D, Gomez Sosa 99 (CTES). Misiones: Posadas, Bonpland, 1] Jan 1908 (f1),
Ekman 2048 (MICH, NY): Candelaria Bonpland arroyo Martires Chico, 16 Jan 1976 ({D), Krapovickas
& Cristobal 28785 (CTES); Cainguas, Monte Carlo, 205 m, 2 Feb 1955 (fr), Montes 14806 (NY); Apostoles,
29 pe 1948 (fr), Schulz 6889 (CTES). Tucuman: Villa Nongues, Jan 1918 (fr), Lillo 1351 (MO)
ZIL. Parana: Palmas, Rio Chopim, 7 km abaixo da nascente, 20 Nov 1990 ({L), Hatschbach
een MBM). Rio Grande do Sul: Sao Francisco de Paula, ({r), Rambo 30793 (MICH); Bom Jesus,
Fazenda B. Velho, 4 Jan 1947 (fr), Rambo 35177 (MO, NY).
PARAGUAY. Itapua: Arroyo Guazu Acati, 15 Sep 1983 ({)), Basualdo s.n.(FCQ); Capitan Miranda,
4.2 km N of entrance to Hotel Tirol (ca. 2712'S, 55°45'W), ca. 210 m, 13 Aug 1995 (st), Landrum 8665
(ASU). Paraguari: National Park Ybycui, NE corner of the park on Arroyo Corrientes (2603'S, 56'50'W),
21 Dec 1988 (fr), Zardini et al. 9015 (PY).
URUGUAY. Artigas: ruta 30,7 km S de Artigas, 10 Dec 1995 (fr), Solis Neffa et al. 242 (CTES);
Cerro Largo: S of Melo, Vie 9 Jan 1944 (fr), Bartlett 21279 (MICH). Maldonado: Sierra de Animas,
ca.65 km E of Montevideo (ca. 34°45'S, 55°30'W), 22 Nov 1981 ({), Landrum 3856 (NY). Montevideo:
Punta ta Fpl, Legrand 2710 (MICH); a eel orillas del rio Uruguay, Sta. Sofia, 15 Nov 1942
CD), Rosengurtt et al. B-4183 (MO, NY): Rocha: Santa Teresa, (f1), | egrand eg 1064 (MICH). San José:
Rincon Gallinas, 5m, Dec 1931 (LD, Herter 88052 (NY, oA le los Cuervos, 17
1944 (fr), Legrand 3338 (NY).
a
an
There are many intermediates between typical var. mucronatum (which is gla-
brous) and typical var. sericeum (which is densely covered with silvery hairs).
The fact that these varieties also have similar distributions, leads me to suspect
that there is a simple genetic difference between them. Intermediates have
mainly been identified as var. sericeum.
Psidium salutare var. mucronatum frequently grows with P missionumand
can easily be confused with that species. The two species are contrasted in the
key below.
. Leaves 2-4.5 cm long, 0.7-2.3 cm wide, 1.5—5 times as long as wide; marginal vein
distinct, Se following the margin; placenta protruding, peltate; style 5—6 mm
long, glabrov P, maretate? var. mucronatum
. Leaves 2.5- ate long, 1.1-4 cm wide, 1.8-3.5 times as long as wide; marginal vein
evident only in distal shes n of leaf, arching broadly between laterals; placenta
protruding only slightly, not peltate; style 7-9 mm long, usually with a few scat-
tered hairs. P. missionum
—
If the entity here called Psidium salutare var. mucronatum is recognized at the
specific level, the name P.luridum (Spreng.) Burret should be used. There may
be no type specimen of Myrtus lurida in existence, having been at B and now
destroyed. Burret (1941), who studied the type shortly before it was destroyed,
listed Myrtus cuspidata asa synonym. That coupled with Sprengel's protologue
leave little doubt as to its identity. The type of Myrtus ovalis at B was also de-
stroyed, but the protologue is sufficient to consider ita synonym of this variety.
Psidium salutare var. pohlianum (O. Berg) Landrum, comb. nov. (Figs. 2C, 6).
Psidium pohlianum O. Berg, in Mart. Fl. bras. 14(1):390. 1857. Type: BRAZIL: “ad S.
LANDRUM, REVISION OF THE PSIDIUM SALUTARE COMPLEX 1467
Luzia in prov. Goyazensi,” Poh] 913, Sellow s.n. (SYNTYPES B, lost; SYNTYPE [Poh] 913],
W, = F-31431! ISOSYNTYPE [Sellows.n_], Pl, [LECTOTYPE, here designated], = ASU photo!).
Psidium pohlianum var. brevipes O. Berg, in Mart., Fl. bras. 14(1):601. 1859. TYPE: BRAZIL: “prope S.
Carlos prov. S. Pauli,” Riedel s.n. (apparent HOLOTYPE 2 sheets at LE, =ASU photos!).
—
Shrub or tree to 10 m high; leaves mostly elliptic, to obovate, or oblanceolate, 4-
9cm long, 2-5.5cm wide, 1.4-2.7(-3.5) times as long as wide, glabrous; venation
pronounced, raised on both surfaces, the marginal vein usually about | mm from
the margin; apex usually without an apiculum; peduncle 0.4-2 cm long, often
triflorous; calyx-lobes shorter or about as long as hypanthial tube plus calyx
tube, rounded to obtuse.
—
Representative specimens. BOLIVIA. Santa Cruz: P. N. Noel Kempff Mercado, Pista Las Gamas,
4481'S, eer 815 m, 9 Nov 1993 (fr), Guillén & Centurién 1023 (ASU); PN. Noel Kempff
Mercado, 6 SW del rere Las Gamas (14°49'36"S, 60°23'10"W), 850 m, 30 Oct 1995 (fr),
Rodriguez & ee 592 (A N. Noel ae _ Huanchaca I, (13'53'55'S, 60°48'46"W),
850 m, 3 Nov 1995 (fr), Rodriguez & Surubi 630 (A
BRAZIL. Bahia: near Junco, ca. 15 km WNW 7 town of Rio de Contas (41°55'W, 13°32'S), 22 Jan
1974 (fr), Harley 15596 (ASU, MICH). Ceara: Crato, Barreiro Grande- Agreste, 2 Apr 1995 ({r), Silveira
22103 (RB). Distrito Federal: 10 km E of Planaltina, 1150 m, 10 Oct 1965 (yfr), Irwin et al. 9101 (NY);
Chapada da re ee rd. NE edge Parque Nac. de Brasilia, Rua 1, 1160 m, 1160 m, 10 Sep 1995 (fD),
Proe ax et al. 1455 (ASU, UB). Goias: Serra dos Cristais, ca. 12 km E of Cristalina (17°S, 48°W), 1200 m,
9 Mar 1966 (fr), Irwin et al. 13226 (CAS, MICH, NY): Serra dos Cristais, ca. 3 km W of Cristalina (17’S,
48'W), oe m, 3 Mar 1966 (fr), Irwin et al. 13367 (CAS, NY). Minas Gerais: Palacio, Serra do Cipo, 150
km N of Belo Horizonte, 1450 m, 20 Feb 1968 (fD, Irwin et al. 20561 (F); 4 km N of Patrocinio, 1000 m,
31 Jan 1970 (fr), Irwin et al. 25733 (NY). Sao Paulo: Itirapina, Estrada de Gratina, 2 Feb 1993 (fr), Fabio
de Barros 2515 (ASU)
VENEZUELA. Amazonas: Atures, ca. 30 km al N de Puerto Ayacucho y a unos 5 km al NE de
Galipero (5°48'N, 67'20'W), 80 m, 9 May 1980 (yf), Huber 5210 (MICH, MO, NY, VEN); Atures, Rincones
de Chacorro, 30 km N de Puerto Ayacucho, 5 km NE de Galipero (5'48'N, 67°20'W), ca. 80 m, 27 Feb
1982 ({D, Huber 6289 voce MO, NY, VEN): Atures, road NE from Puerto Ayacucho, 27.5 km towards
El Burro, 28 Apr 1984 (fr), Plow & Guanchez 13756 (CAS, FE MO, NY, RB). Bolivar: Rio Parguaza, 13
Apr 1946 (fD, Velez 2374 (US, VEN)
jer
If the entity here called Psidium salutare var. pohlianum is recognized at the
specific level , the name Psidium pohlianum O. Berg should be used. As men-
tioned above, this variety intergrades with var. decussatum in the vicinity of
Brasilia.
Two specimensat LE fit the protologue of Psidium pohlianum var. brevipes,
but are annotated by Berg as to species only. lassume these specimens to repre-
sent the holotype of var. brevipes. One specimen is mixed with a fragment of P.
guineense.
Psidium salutare var. sericeum (Cambess.) Landrum, comb. nov. (Figs. 2D, 5).
Myrtus sericea Cambess., in Saint-Hilaire, Fl. Bras. merid. 2:295. 1833. TYPE: BRA-
ZIL: “Capilha de Mercedes... provinciae Cisplatinae, necnon... Rincao de Saneloés
ad ripam amnis [bicuy in provincia Missionum,” Saint-Hildire s.n. (HOLOTYPE: PI, =
F-36446! ISOTYPE: P!, = ASU photo!).
1468 BRIT.ORG/SIDA 20(4)
Myrtus nived O. Berg, in Mart., Fl. bras. l4+(:414. 1857. Type: URUGUAY: “ad PI... dos Inforcados in
Montevideo,” Sellow s.n. (HOLOTYPE: B, lost; ISOTYPES P! [LECTOTYPE, here designated], =ASU
photol, W, = F-31405).
Myrtus sericea var. fruticosd O. Berg, in Mart. Fl bras. 14(1):414. 1857. Type: URUGUAY: “ad Cerro
in Montivedeo,” Sellow s.n. (IOLOTYPE: B; ISOTYPE: PHLECTOTYPE, here a ae ASU photo).
Myrtus sericea var. suffruticosa O. Berg, in Mart., PL bras. 14(1):414. 1857. Tyee: BRAZIL: illegiti-
mate name to be replaced by M. sericed var. seri¢
Myrtusincana O, Berg, in Mart., Fl. bras. 14(1):416. — TyPE: BRAZIL: “ad Cassapava in Rio Grande
do Sul,” Sellow s.n. (HOLOTYPE: B; ISOTYPE: P! [LECTOTYPE, here designated], = ASU photo!).
Myrtus pubescens O. Berg, in Mart, FI. bras. 14(1):415. 1857. Type: BRAZIL: “ad Andre’ Ferrina,”
Sellow s.n. (HOLOTYPE: B; ISOTYPE: P![LECTOTYPE, here designated], = F-36441!, = ASU photo!).
Psidium incanum (O. Berg) Burret, Notizbl. Bot. Gart. Berlin-Dahlem 15:485. 1941.
Psidium niveum (O. Berg) Herter, Rev. Sudamer. Bot. 7:221. 1943.
Psidium tomentellum Burret, Notizbl. Bot. Gart. Berlin-Dahlem 15:485. 1941. New name for Myrtus
sericea Cambess.
Usually a subshrub less than 0.5 m high; leaves elliptic, ovate, obovate, nar-
rowly elliptic, oblanceolate to lanceolate, 2-7 cm long, 0.6-2.8 cm wide, 2-3.7
times as long as wide, densely covered with silvery gray hairs when young; ve-
nation pronounced under hair cover, the marginal vein usually within | mm of
the margin; apex apiculate; peduncle often over 2 cm long, uniflorous; calyx-
lobes usually longer than hypanthial tube plus calyx tube, usually acute.
Representative specimet SES ENEINE, Corrientes: Santo Tome, ruta 41, 5-6 km al N de Galarza (ca.
28°4'S, 56°39"W), 17 Nov 1904 cab) ‘(al6402(CTES); Estancia Ana Cua, l7 Dec 1970 (f)), Cammevan
2243 (CTES),; Estancia Garruchos, cachuera, Ayo. ree 6 Feb 1972 (fr), Krapovickas et al 211
(CTES); General Paz, Arroyo Sta. Isabel at ruta 12 E of Ita Ibaté (ca. 57°30'W, 27°20'S), 9 Dec 1987 (fr),
Landrum 5701 (ASU, CTES); Estancia El Recreo, 21 km E Bonpland, costa rio Uruguay, Dep. Paso de
los Libres, 18 Nov 1973 (fl, Lourteig et al 2737 (CTES); Monte Caseros, Dep. Monte Caseros, 19 Feb
L975 (fr), Prause O(CTES); Empedrado, Estacion Experimental INTA, 7 Dec 1978 (£1), Schinini 16188
(CTES); LO sos S de Bella Vista, 8 Nov 1978 ({1), Schinini G& Ahumada 15894 (CTES, MO); 1 km S de
Mercedes, ant Curuzu Cuatia, Co. eee camara Iressens et al 2420 (CTES).
Entre Rios: ( ener. Dec 1946 (ED, Meyer 11007 (LIL). s, Lareto, Casa de Drewes, 26
Jan 1908 (fr), Ekman 2056 (MICH, NY): San José, Feb 1961 (st), Martinez C rovettoSD-I(CTES). Tucuman:
oe ua, Cerro aie Campo, 1000 m, Nov 1978 (1), Venturi 7582
BOL a Cruz: Prov. Florida, + km N of center of Samaieata (1808'S, 63°52'W), 2000-
2100 m, iis ie 1992 ae Nee & Vargas 43465 (ASU)
B
RAZIL. Rio Grande do Sul: Continuacao da estrada Alegrete-Cerro do Tigre, apos o Cerro do
Tigre, direc sonieiteni 11 Feb 1990 (fr), Falke Sene Sobral 15235 (MBM); Porto Alegre, Montserrat,
13 Nov 1941 (1D, Emrich 8380 (LIL); Fazenda Faxinal, Arroio dos Ratos, 5 Nov 1980 (fD, Hagelund
13420 (CTES, NY); Morro das Abertas, 9 Jan 1949 ({1), Rambo 39674 (LIL): Sao Vicente do Sul, estrada
a Cacequi, rio Ibicui, Dec 1985 (st), Sobral & Marchiori 4544 (UB).
PARAGUAY. Central: Ita, Granja lsapy, orilla arroyo Lazarillo, 30 Jan 1966 ({r), Krapovickas et al
12231 (CTES). Cordillera: Ypacaray, 6 Dec 1950 (fr), Sparre & Vervoost 814 (LIL). Misiones: San Juan
Bautista, ca. 8.5 km along road to Pilar, ca. 170m, 8 Novy 1995 (Ir), Landrum 8790 (ASU); Ea. La Soledad,
3km Sde Santiago (56 46'W, 2710'S), 3-4 Feb 1988 (fr), Schinini & Vanni 26108 (ASU, CTES). Paraguari:
Rt. 1, between Quindy and Caapucu, near km 246 (26'S, 57°15'W), ca. 250 m, 7 Nov 1995 (st), Landrum
3764 (ASU).
URUGUAY. Cerro Largo: Arroyo Zapallar, 22 Dec 1961 (fr), Praderi 740 (LIL). Durazno: Est. Las
Palmas, Mar 1922 (st), Osten 16563 (NY). Montevideo: La Colorada, 17 Nov 1947 (fD, Legrand 2711(NY).
LANDRUM, REVISION OF THE PSIDIUM SALUTARE COMPLEX 1469
Paysandu: Chapicuy, orillas del rio Uruguay, Sta. Sofia, 15 Nov 1942 CD, Rosengurtt B- 3250 (MO, NY).
Rivera: Tranqueras, ({l), Legrand 4145 (MICH, NY). San José: Rincon Gallinas, Dec 1931 (f), Herter
8051 (MO). Tacuarembo: Cerro Dos Hermanos, Mar 1922 (fr), Osten 16651 (NY).
This entity has long gone by the name Psidium incanum (O. Berg) Burret, which
is the correct name if one recognizes it at the specific level. It most closely re-
sembles var. mucronatum and intergrades with it.
ACKNOWLEGMENTS
lam ereterl to the curators of the following herbaria who kindly allowed me
to study their on loan, as images, or asa visitor (ALCB, ASE, BR, CAS,
(CEPEC GES, FAP. FoPCO 4G, GH. AB HBR TRB. TUBS. kK CE, Lil, MBM,
MEXU MICH, MO, NY, MY/OXF PPY,OCA,OCNE,R, RB, SGO, SP TERA, UB,
UC, US, USM, USP, VEN, WIS). Alexander N. Sennikov has been especially help-
ful in providing images of types at LE. My Latin American colleagues C. Nel-
son, R. Bye, I. Basualdo, E. Zardini, C. Proenga, M. L. Kawasaki, A. Krapovickas, G.
Hatschbach, and M. Sobral have helped in many ways. Herbarium visits and
fieldwork was supported by a Fulbright American Republics grant and by a
National Geographic Society grant. N. Snow and an anonymous reviewer of-
fered many helpful suggestions.
REFERENCES
Bera, O. 1857-1859. Myrtaceae. In: C.F.P. von Martius, Flora brasiliensis 14(1):1-655.
Burret, M. 1941. Myrtaceen — Studien. Notizbl. Bot. Gart. Berlin-Dahlem 15:479-500.
CANDOLLE, A.P. DE. 1828. Myrtaceae. In: Prodr. Syst. Nat. reg. Veg. 3:207—296.
Lanorum, L.R.1986.Campomanesia, Pimenta, Blepharocalyx, Legrandia,Acca, Myrrhinium, and
Luma (Myrtaceae). Flora Neotropica. Monogr. 45:-178.
Lanorum, L.R. and W.P. SHare. 1989. Seed coat characters of some American Myrtinae
(Myrtaceae): Psidium and related genera. Syst. Bot. 14:370-376.
Lecranp, C.D. and R.M. Kein. 1977. Psidium. Flora llustr. Catarin. [MIRT]:684—724.
McVauch, R. 1968. The genera of American Myrtaceae — An interim report. Taxon 17:354—
Rotman, A. 1976. Revision del género Psidium en la Argentina (Myrtaceae). Darwiniana
18-444
1470 BRIT.ORG/SIDA 20(4)
BOOK NOTICES
Cc t ae IMA 1 IE] LY cee? |
fod r
GeyATA Ajitvscl 2002. Wildflowers of Texas. Revised Edition. (ISBN 0-940672-
73-1, pbk, laminated). Shearer Publishing, +06 Post Oak Road, Fredericksburg,
TX 78624, US.A. (Orders: 800-458-3808, ww wshearerpub.com). $19.95, 524
pp., numerous color photographs, 4 1/2 x 7 1/2".
From the inside cover
ae
lap.—*IN this revised edition of her popular Wildflowers of Texas, native-plant
>xpert Geyata Ajilvsgi gives lay readers the most comprehensive field guide currently available on
the state's abundant wildflowers. With the addition of 105 species, the book now contains, informa-
tion on 482 of the most common wildflower species found in the state's major vegetation zones—
from the mountains in the west to the swamplands in the east, from desert to piney woods to coastal
plain, from the Panhandle to the sa ae Rio Grande Valley. Each entry reflects the most cur-
rent scientific knowledge about the plant, such as recent changes in taxonomic classification.”
The ae is nice a color showsereohe on the left side of the page and text on the right.
Every plant list family, bloom period, description, habitat, anda general
note about the cme It is fun to just read the vbout infdlivicluel species. The field
guide includes a glossary, an illustrated glossary (floral structures, vegetative structures), selected
bibliography that includes many up-to-date references, and an index of Latin names anda wealth of
common names.—Barney Lipscomb, Botanical Research Institute of Texas, 509 Pecan St, Fort Worth,
|X 76102-4060, U.S.A., barney@britoreg
Ropert H. MoHLeNBROCK. 2002. The [lustrated Flora of Illinois. Grasses: Bromus
to Paspalum. Second Edition. (ISBN 0-8093-2 359-1, hbk.) Southern Illinois
University Press. RO. Box 3697, Carbondale, IL 62902-3697, U.S.A. (Orders:
618-459-6633, 618-453-1221 fax, wwwsiuedu/‘siupress). $60.00, 416 Pp,
284 line drawings, 6" « 9".
Ropert H. MOHLENBROCK. 2002. Vascular Flora of Illinois. (ISBN 0-8093-2421-0,
pbk.) Southern Illinois University Press. PO. Box 3697, Carbondale, IL 62902-
3697, US.A. (Orders: 618-459-6633, 618-453-1221 fax, wwwsiuedu/‘siupress).
$50.00, 736 pp, | map (county map of Illinois), 6" x 9"
—
The Illustrated Flora of Illinois. Grasses: Bromus to Paspalum. Second Edition The second edition fea-
tures 22 newly discovered taxa since the first edition in 1972. “For each new species, a full-page illus-
tration is provided. This second edition updates the status of Illinois grasses. The book features 263
figures from the first edition plus 21 new figures for this edition by Paul W. Nelson.” The drawings are
excellent. Keys and distribution maps are included.
Vascu
—
ie second
and enlarged edition in 1986. The latest edition covers over 3200 taxa, an increase of almost 300
ar Flora of Illinois is the sequel to the first edition published in 1975 and to t
since 1986. Besides keys to genera and species, common names, {lowering times, habitats, and geo-
graphical distribution, the taxa below the rank of species are recognized and may be identified by
keys.—Barney Lipscomb, Botanical Research Institute of Texas, 509 Pecan St, Fort Worth, 1X 76102-4060,
U.S.A, barney@britore.
SIDA 20(4): 1470. 2003
REVISION AND TYPIFICATION OF SOME SPECIES OF IXORA
(RUBIACEAE) FROM CENTRAL AND SOUTHERN BRAZIL
Piero G. Delprete'
Institute sles C ele
The Nev k Botanical Ga
Bronx, WW? 10458-5126, USA.
pdelprete@nybg.org
ABSTRACT
In connection with the treatment of Ixora for the Flora Fanerogdmica do Estado de Sdo Paulo, Flora
é
Ilustrada Catarinense and Flora de Goids e Tocantins, it was necessary to nnd hake a taxonomic
study of the taxa occurring in Central and Southern Brazil. F S] ecur in the state of
Sao Paulo, and 12 taxa originally described ey Muller argv are here treated as synonymous
with the recognized species. Only one native the state of Santa Catarina. Full typi-
fication, synonymies, brief 3 iptions, Sacre cited anda . to the Ixora of Sao Paulo are here
presented.
Key worps: Lxora, lxoroideae, Rubiaceae, typification, Brazil.
RESUMEN
E nexion con el tratado de Ixora por la Flora Fanerogdmica do Estado de Sdo Paulo, la Flora
Ilustrada Catarinense y la Flora de Goids e Tocantins fue necesario realizar un estudio taxonomico
de los taxa presentes en Brasil central y meridional. Cinco especies nativas ocurren en el estado de
Sao ae y doce taxa originariamente descritos por Muller Argoviensis son trat eid aqui como
idas. Solamente “atarina
Se presentan aati tipificaciones y si S pletas, descripciones, especimenes citados y una
clave para las especies de Ixora de Sao P: ul
Ixora is a pantropical genus of about 350 species, with most of them present in
tropical Asia and the islands of the South Pacific, about 37 species in continental
Africa, about 35 species in Madagascar, and about 45 species in the Neotropics.
Although the African species have been recently studied by De Block (1998), re-
visions for the Neotropical or the Asiatic species have never been attempted.
During the preparation of the treatment of Ixora for the Flora Fanerogdmica
do Estado de Sdo Paulo (Delprete, in press) and Flora Ilustrada Catarinense
(Delprete et al., submitted), and Flora de Goids e Tocantins, it was necessary to
examine material from many Brazilian and international herbaria, and to study
the type specimens of the species described by Bentham (1850) and Miller
Argoviensis (1875, 1881).
Johannes Miller (1828-1896) was renamed by CEP. von Martius (coordi-
'Current address: National Herbarium of the Netherlands, Utrecht University branch, Plant Systematics,
Heidelberglaan 2, 3584 CS Utrecht, The Netherlands, E-Mail: pdelprete@hotmail.com
SIDA 20(4): 1471-1480. 2003
1472 BRIT.ORG/SIDA 20(4)
nator of Flora Brasiliensis) as Muller Argoviensis (because he was native of a
small town in the canton of Aargau, Switzerland), to distinguish him from other
contemporary botanists with the same last name. The personal herbarium of
Muller Argoviensis was subsequently included in the Candolle Herbarium (G-
DC). Because of their historical value, the specimens of this herbarium are not
sent out on loan. In order to examine the type specimens of the Ixora species
described by Muller Argoviensis, a set of about 80 black and white negatives
were made by the staff of the G-DC Herbarium and sent to NY. The negatives
were used to make large photographs, which were mounted on NY herbarium
sheets. The present work would have not been possible without the close ex-
aminations of these photographs.
The most complete treatment of Brazilian species of lxora still remains
that of Muller Argoviensis (1881), where he expanded the original descriptions
of his new species published a few years before (Miller Argoviensis 1875), and
cited additional specimens of contemporary collectors. In his treatments, he
distinguished the species of Ixora based on leaf shape, presence of colleters in-
side the calyx, and presence and shape of leaf-like bracts subtending the sec-
ondary branches of the inflorescences. All these characters are here consid-
ered of little taxonomic significance, and most of Muller Argoviensis new taxa
jemi
i
here studied are reduced to synonymy. In addition, the fruits of the species exam-
ined are quite uniform, in being spherical to subspherical and 5-7 mm in diam-
eter, usually reddish to deep red, and consequently without any useful taxo-
nomic characters. Most of the characters considered of taxonomic significance
in the definition of the species presented below were found in the stipule shape,
length of the stipular arista, leaf venation, inflorescence size and architecture,
number of inflorescence secondary branches, flower bud shape (ovoid or ob-
long-ellipsoid at apex), corolla tube length, corolla lobes length and shape, and
corolla tube/lobe ratio. A key to the species of the State of Sao Paulo is pre-
sented below.
Full typification and citation of specimens examined is here presented
because this information can not be included in the concise format of the Flora
Fanerogamica do Estado de Sdo Paulo. It is my hope that this work will provide
an initial contribution towards the clarification of Ixora from central and south-
ern Brazil; however, considerable work remains to be done before a revision of
the Brazilian species of this difficult genus can be completed.
Ixora brevifolia Benth., Linnaea 23:448. 1850. Type: BRAZIL. MINAS GERAIS: without
locality, 1845-1846 (1D, Widgren 1119 (LECTOTYPE: UPS! V-134607, here selected:
ISOLECTOTYPE: UPS n.v. V-102640; not at GOET, K, S).
Ixora thyrsoidea Muell. Arg., syn. noy., Flora 58:455, 459. 1875. TYPE: BRAZIL. BAHIA: without lo-
cality, 1857, Blanchet s.n. (HOLOTYPE: G-DC n.v., photos at NY}).
Ixora warmingii Muell. Arg,, syn. nov., Flora 58:455, 459.1875. TyPE: BRAZIL. MINAS GERAIS: Lagoa
Santa, s.d. [1863-1865
. Warming s.n. (LECTOTYPE: G-DC n.v,, here selected, photos at Cl, NY).
DELPRETE, TAAUNUNLT UF 1473
Ixora nse serene syn. nov., in Mart., FI. Bras. 6(5):460. 1881. Type: BRAZIL. R10 DE JAN-
Without ee ei [1861-1895], Glaziou 10932 (HOLOTYPE: G-DC n.v, photos at NY},
ISOTYPE: Bt, photo at NY).
Ixora membranacea Muell. Arg., syn. nov., in Mart. Fl. Bras. 6(5):458. 1881. TyPE: BRAZIL. RIO DE
JANEIRO: without locality, s.d. [1861-1895], Glaziou 10947 (HOLOTYPE: G-DC n.v,, photos at NY};
ISOTYPE: Bt, photo at NY).
Tree up to 20 m tall, to 35cm at dbh; bark smooth; branchlets glabrous. Stipules
4-8 x 3-4 mm, base deltoid, arista 4-6 mm long, glabrous outside, densely seri-
ceous-pubescent and intermixed with colleters inside. Leaf blades narrowly el-
liptic, oblong elliptic to obovate, (4-)8-20 = 15-6(-8) cm, apex acute or acumi-
nate, base acute to round, subcoriaceous to coriaceous, glabrous; secondary veins
15-40 each side, tertiary venation subparallel-reticulate; petioles 4-10 mm long,
glabrous. Inflorescences sessile or pedunculate, paniculate, multiflorous, 5-12
x 3-8 cm, with 2 or 3 lateral branches, these sparsely branches or corymbose;
peduncles 2.5-4 cm long, or absent; bracts subtending the secondary branches
narrowly triangular, 1-4 mm long; bracteoles triangular, 0.4-0.7 mm long.
Flower buds oblong-ellipsoid at apex. Pedicels absent or to 2 mm long. Flowers
sessile or short-pedicellate, fragrant. Calyx 0.7-1 mm long; lobes triangular or
ovate, glabrous. Corolla 6-9 mm long, cream-white; tube 2.6-3.7 x 1-15 mm,
glabrous outside, pubescent inside; lobes oblong, 3-5 x 0.8-2 mm, apex round,
glabrous. Stamens 3.5-4.5 mm long: anthers narrowly oblong, ca. 2.5 mm long,
apex acute. Style ca. 4-6 mm long; style branches narrowly oblong, ca. 2 mm
long. Fruit globose, 5-6.5 mm diam., glabrous, red.
a
Selected specimens examined. BRAZIL. Bahia: Rd to Abaira, ca. 8 km N of town of Rio da Contas, ca.
13°33), 41°47'W, 1000 m, 18 Jan 1972 ({1), Harley et al. 15251 (NY); Serra da Agua de Rega, ca.24 km N of
Seabra, rd to Agua de Rega, 1000 m, 25 Feb 1971 (fr), Irwin et al. 31058 (NY); Serra da Agua de Rega, ca.
28 km N of Seabra, rd to Agua de Rega, 1000 m, 27 Feb 197] (fr), Irwin et al. 31184 (NY); rd. BA, 12 km
from Mucugé, L entrance of Fda. Paraguacu, 16 Dec 1984 (f1), Lewiset al. s.n. (SPF 36789) (NY). Distrito
Federal: Bacia do Rio Sao Bartolomeu, 18 Dec 1979 (f1), Heringer et al. 2950 (NY), 14 Jan 1981 (fr), 5985
(NY); Corrego da Papuda, near barra of Rio Sao Bartolomeu, 22 Jan 1981 (fr), Heringer et al. 6026 (NY),
Granja do Tamandua, 6 Oct 1965 ({D, Heringer et al. 10613 (NY); Rio Corumba, 120 km from Brasilia,
18 Jan 1968 (fL, Heringer 11618 (NY). Minas Gerais: Serra do Cantagalo, Sao Tomé das Letras, 3 Nov
1984 (fr), Mello Silva et al. s.n. (SPF 35862) (NY). Goias: Formosa, 20 Oct 1961 (f1]), Heringer 10740
(NY): Serra dos Pireneus, ca. 9 km S$ of Corumba de Goias, 1000 m, 1 Dec 1965 (£1), Irwin et al. 10889
(NY); Serra do Caiapé, ca. 50 km S of Caiaponia, rd to Jatai, 900 m, 27 Jun 1966 (fr), Irwin et al. 17885
(NY); Serra do Caiap6, 66 km of Jatai, 22 Oct 1964 (FD, Prance & Silva 59565 (NY), Mun. Goiania, rd.
GOM-9, 2 km from Escola de Agron. e Veterinaria, 23 May 1968 (fr), Rizzo & Barbosa 1050 (UFG);
Mun. Goiania, rd. Goiania-Leopoldo de Bulhoes, 18 km from Goiania, 2 Jul 1968 (f1), Rizzo & Barbosa
1570 (UFG): Mun. Goiania, rd GOM-6, 16 km from Goiania, 2 Nov 1968 ({D, Rizzo & Barbosa 2622
(UFG[2)); Mun. Goiania, Morro dos Lobos, 3 Dec 1968 ({1), Rizzo & Barbosa 291] (UFG), Mun. Goiania,
rd Goiania-Senador Canédo (GOM-7), km 12, 5 Dec 1968 (£1), Rizzo a ol ae 2989 (UFGI2]), 27 Dec
ule fl), 3159 (UFG); Mun. Goiania, rd. Goiania-Senador Canédo (G ), km 12, 4 Mar 1969 (£1),
220 & Bar bosa 3905 (UFG); Mun. Jatai, towards Perolandia, 20 ae rom ee Claro, 9 Nov ae ot
Seren (UFG); Rio Paranaiba, 20 km from Itumbiara, 23 Nov 1972 ({D, Rizzo & Barl G),
21 Dec 1972 (£1), 8697 (UFG); Mun. Goiania, Universidade Federal de Goias, Campus II, Bosque rene
de Saint-Hilaire, 17 Nov 1978 (f1), Rizzo et al. 10045 (UFG), 18 Nov 1997 (fl), 12258 (UFG). Sao Paulo:
1474 BRIT.ORG/SIDA 20(4)
Mun. Tatui, rd Sorocabana, | Feb 1946 (£D, Amaral 36 ( ee : Mun. Moji Mirim, 27 Oct 1993 ({D, Arbocz
l4 TAC), 14 Nov 1993 ED, 61 UAC); Mun. Santo A 3 e, Fda. Palmital, 3 Dec 1993 (fl), Arbocz
88 IAC); Mun. Anhembi, Fda. Barreiro Rico, 15 Dec 1981 (f ae César s.n. (HRCB 3048), 30 Sep 1982 (Ir),
sn. (HRCB 3333) (ARCB); Mun. Agudos, Fda. Cia. Cervejaria Brahma, 23 Oct 1997 (f1), Cristianini &
Assis Camargo 651 (SP), 20 Nov 1997 (fD, 718 (SP); Mun. Sao Paulo, Parque Estadual da Serra do Mar,
Nucleo Curucutu, 2359'S, 46'44'W, 19 Dec 1996 (f1), Garcia et al. 967 (NY): Mun. Moji Guagu, Fda.
Campininha, 17 Jan 1977 (f{D, Gibbs & Leitdo Filho 4275 (NY, UEC); Mun. Itararé, Ibiti, Estacao
perimental Monte Alegre, | Aug nese Kuhlmann s.n (IAC 28616) (IAC, SP); Mun. Ilha do C: arose)
1-14 Dec 1979 (tr), Leitdo Filho et al.s.n.(UEC 10823) (UEC); Mun. Pereira Barreto, noes Lagoinha IL,
Nov 1985 (f1), Martins etal. 95 (HRCB, SP. ao Mun. Campinas, Bosque dos Jequitibas, s.d. (1) ie 5
7757 UEC), 7758 (U ie Me SC ce Fda. Sta. Carlota, 15 Dec 1999 ({1), Nicolawesd? 2091 (NY), 2092
(SP); Mun. Améric Sep 1992 (st), Rocha 17 (ESA); Mun. Moji Guacu, Reserva Biologica
e Estacao Esperimental, 21 Jun 1988 (st), Rossi et al. 1195 (SP); Mun. Porto Ferreira, Reserva Florestal,
22 Nov 1975 (£1), Silva et al. sn. (UEC 13842) (UEC); Mun. Aguas de Lindoia, rd. to Barao de Ataliba, 9
May 1995 (fr), Tamashiro et al. 1039 (SP UEC)
7
~
fi
A species occurring in the states of Minas Gerais, Goias, Distrito Federal, Mato
Grosso, Bahia, Rio de Janeiro, Espirito Santo and Sao Paulo, usually found in the
moist forests of the coastal states, and in the gallery forests of the cerrado biome
of central Brazil. Many specimens have been traditionally annotated as Ixora
warmingii, which is a later synonym of this species.
Because Ixora brevifolia Benth. was originally described by Bentham (1850),
the type material was expected to be housed at K. After two distinct searches
by the K herbarium staff, no type material (Pohl s.n.and Widgren 1119) pertain-
ing to this species was found at this institution. Therefore, a search of duplicate
specimens of Widgren 1119 was initiated in the main herbaria where Widgren's
collections were distributed (C, GOET, S, UPS; Vegter, 1988). Two duplicates of
Widgren 1119 were found at UPS, one of wihighs is here selected as the lectotype
of Lxora brevifolia.
am)
—
Ixora gardneriana Benth., Linnaea 23:448. 1850. Type: BRAZIL. RIO DE JANEIRO: woods
of Tijuca, Dec 1840, Gardner 5496 (HOLOTYPE: K!, one at NY} ISOTYPE: K!).
Ixora schottiana Muell. Arg., syn. nov., Flora 58:455, 459. 1875. TYPE: BRAZIL. RIO DE JANEIRO: Serra
Tingua, s.d. [1817-1820], Schott 889 (LECTOTYPE: G-DC n.v.. here selected, photos at NY ISOTYPE:
K!, photo at NYD.
Ixora riedeliand Muell. Arg., syn. nov., Flora 58:455, 459. 1875. TYPE: BRAZIL [Southern Brazill:
without locality, s.d. [1821-1836], Riedel s.n. [*478"] (HOLOTYPE: G-DC n.v, photos at NY! dubi-
ous ISOTYPE: NY!).
Ixora obscura Muell. Arg., syn. nov., Flora 58:455, +59. 1875. Type: BRAZIL [Southern Brazil with-
out locality, s.d. [1815-1817], Sello s.n. (HOLOTYPE: G-DC nv, photos at NYE ISOTYPE: Bt [Sello
6018], photo at NY).
Ixora truncata Muell. Arg. syn. nov., in Mart. Fl. Bras. 6(5):459. 881. TYPE: BRAZIL. RIO DE JAN-
EIRO: Rio de Janeiro, s.d. [1861-1895], Glaziou 10933 (LECTOTYPE: G-DC nv, here selected. pho-
tos at NY! ISOLECTOTYPE: Bt, photo at NY)).
Tree; bark smooth: branchlets glabrous. Stipules 2.5-3.5 « 1.5-2 mm, base del-
toid, arista 15-2 mm long, glabrous outside, densely sericeous-pubescent and
intermixed with colleters inside. Leaf blades elliptic or ovate, 5-13 x 2.5-5.5cm,
DELPRETE, TAAUINNUNTT UE 1475
apex acute, base acute, subc oriaceous glabrous; secondal y veins 6-14 each side,
tertiary venation reticulate; petioles 5-12 mm long, glabrous. Inflorescences
pedunculate, paniculate, pauci- or multi-florous, 5-11 x 3.5-6 cm; peduncles
0.5-2.5 cm long: bracts subtending the secondary branches often foliose, nar-
rowly triangular to obovate, 5-10 mm long; bracteoles | or 2 per flower, trian-
gular, 2-3 mm long, or absent. Flower buds ovoid at apex. Pedicels 2-7 mm long.
Flowers pedicellate (rarely sessile), fragrant. Calyx 0.8-1 mm long; lobes del-
toid, glabrous. Corolla 8-9 mm long, cream-white; tube 4.5-5.5 x 1-1.2 mm, gla-
brous outside, pubescent inside; lobes ovate, 3.5-4 x 2-2.5mm, apex round, gla-
brous. Stamens 3.5-4 mm long; anthers narrowly oblong, 3.2-3.7 mm long, apex
apiculate. Style 10.5-11 mm long: style branches narrowly oblong, 7-2 mm long,
Fruit unknown.
Illustration.—Mart., Fl. Bras. 6(5), Tab. 9. 1881 (as Ixora schottiana).
Additional specimens examined. BRAZIL. Sao Paulo: Sao Paulo, Fda. Itatuba, 23 Dec 1931 (fD), Ama-
ral sn. (SP 28628) (NY, SP); Sao Paulo, Jardim Botanico, tree n. 179, 9 Dec 1931 ({)), Hoehne s.n. (SP
28588) (NY[2], SP); Mun. Campinas, Bosque de Sao José, 13 Sep 1994 ({D, Santin & Bertani s.n. (UEC
33566) (NY, UEC).
A very rare species, and certainly in danger of extinction, that occurs in the
Atlantic forests of the states of Rio de Janeiro (Serra do Tingua e Serra da Estrella)
and Sao Paulo (Fda. Itatuba and Bosque de Sao José).
The holotype specimen has the stamp “Herbarium Benthamianum’” anda
label with the handwriting “5496, Ixora Gardneriana Benth, Linnaea 13:448"
and the print “Brasilia, Gardner, 1841.” Whereas the isotype specimen has the
stamp “Herbarium Hookerianum’ and the handwritten label “5496, Faramea
sp.,a large shrub - woods of Tijuca, Dec./[18]40.” Because the label of the isotype
was handwritten by Gardner, the locality and date reported on this specimen
are considered more accurate.
Ixora heterodoxa Muell. Arg., Flora 58:454, 459. 1875. Type: BRAZIL. RIO DE JAN~
FIRO: near Rio de Janeiro, s.d. [1861-1895], Glaziou 6154 (HOLOTYPE: G-DC nv, pho-
tos at NY ISOTYPE: Bt, photo at NY).
Ixora burchelliana Muell. Arg., syn. nov., Flora 58:454, 458. 1875. TYPE: BRAZIL. SAO PAULO: “Prov
Sao Paulo,” s.d. [1825-1830], Burchell 3208 (HOLOTYPE: fragments G-DC n.v., photos at NY!:
ISOTYPE: BR).
Ixora di phophyll Muell. Arg., syn. nov., Flora 58:454, 458.1875. TYPE: BRAZIL. Rio de Janeiro:
Without locality, Glaziou 705 (HOLOTYPE: C!, photo at NY}; IsSOTYPES: BRI, K!; photo-K at NY).
Ixora heterophylla Muell. Arg., syn. nov., in Mart. Fl. Bras. 6(5):68. 1881. TyPE: BRAZIL. RIO DE
JANEIRO: Serra Estrella, in fruticetis umbrosis, s.d. [1821-1836], Riedel 342 (HOLOTYPE: G-DC
n.v,, photos at NY! lsoTYPE: BR).
Shrub 2-3(-5) m tall, rarely trees 5-8 m tall, 10-15 cm at dbh; bark smooth;
branchlets glabrous. Stipules 5-6 x 2.5-3.5 mm, base deltoid, arista 2-3.5 mm
long, glabrous outside, densely sericeous-pubescent and intermixed with
colleters inside. Leaf blades ovate, elliptic to obovate, 5-18 = 2.5-7 cm, apex acute
1476 BRIT.ORG/SIDA 20(4)
or acuminate, base acute, membranaceous to chartaceous, glabrous; secondary
veins 6-16 each side, tertiary venation reticulate; petioles 5-20 mm long, gla-
brous. Inflorescences pedunculate, paniculate, multiflorous, 6-9 x 4.5-9 em,
rachis usually filiform, red; peduncles 1.5-4 cm long, glabrous; leaves subtend-
ing the inflorescence smaller than those of lower nodes: bracts subtending the
secondary branches narrowly triangular to lanceolate, 4-7 x 1-2 mm long:
bracteoles | or 2 (rarely 3) each flower, triangular, 2-3 mm long, or rarely ab-
sent. Flower buds ovoid at apex. Pedicels absent or 2-4 mm long. Flowers sessile
or short-pedicellate, fragrant. Calyx 0.7-1 mm long; lobes triangular, glabrous.
Corolla 9-12 mm long, cream-white; tube 6-8.5 x 0.7-0.9 mm, glabrous outside,
sparsely pubescent inside; lobes oblong, 2-35 x 1.7-2.5(-3.5) mm, apex round
(rarely microscopically apiculate), glabrous. Stamens 1.6-1.9 mm long; anthers
narrowly oblong, 1.5-L8 mm long, apex apiculate. Style 8.5-13 mm long; style
branches ovate, ca. | mm long. Fruit subglobose, didymous, 5-7 x 4-6 mm, gla-
brous, red to blackish.
—
Selected specimens examined: BRAZIL. Parana: Mun. Guaraquecaba, Rio Murato, 1] Dec 1970 (1), G.
Hatsch bach 25807 (UB). Rio de Janeiro: Taquara da Tijuca, | Nov 1972 (fl), D. Sucre 9722 (NY, UB). $a0
Paulo: Mun. Iguape, Estagao Ecologica Juréia-Itatins, 7 Mar 1993 (fr), Anunciacdo et al. 190 (NY), 29
Oct 1994 ED, Anunciacdo & Rossi 392 (NY); Mun. Ubatuba, Picinguaba, ll Jan 1993 ({]), Assis 65 (NY):
Mun. Ubatuba, 19 Apr 1995 (fr), Assis & Carneiro 529 (NY); Mun. Cananéia, Ilha do Cardoso, 10 Feb
1981 (ED, Baitello 46 (SPSF 7879), 25 Oct 1989 (f1), Cordeiro et al. 569 (LAC): between Ubatuba and
Caraguatatuba, LO m, 22 Aug 1976 (ED, Davis et al. 59883 (UEC); Mun. Ubatuba, 1] Mar 1989 (fr), Furlan
et al.677 (HRCB, SPSF), 12 Mar 1989 (fr), 780 (SPSF. NY), 788(NY), 8 Dec 1989 ([1), 1014 (NY); Mun. Sete
Barras, Fda. Intervales, 12 Mar 1994 (fr), Galetti et al. 119 (SP); Mun. Cananéia, I]ha do Cardoso, ll Fe
1981 ED), Jung-Mendacolli 479 DAC), 31 Oct 1985 (f), Kirizawa 1529 IAC): Sao Paulo. Jardim Botanico,
4 Dec 1951 (fD, Kuhlmann 2806 (SP, UEC); Mun. Cubatao, Vale do Rio Piloes, 4 Oct 1988 (fl), Leitdo
Filho & Pagano s.n. UEC 20804) (UEC); Mun. Cubatao, 23 Sep 1896 (f1), Loefgren s.n. (SP 11557) (SP):
Mun. Peruibe, Estagao eee ee de Juréia, 11 Nov 1982 ({D, Rodrigues & Figueiredo s.n. (UEC 15667)
(UEC), 17 Nov 1982 (f1), sn. (UEC 15679) (NY, UB, UEC): Mun. Peruibe, Juréia, 6 Nov 1990 (£1), Simdo-
Bianchini 252(NY); Mun. | wee Estagao Ecologica Juréia-ltatins, Oct 1991 (f1), Sobral & Attili 7320
(HRCB); Mun. Peruibe, Foz do Rio Guarau, 9 Oct 1995 (f1), Souza et al. 9267 (NY),
A common species occurring in the states of Rio de Janeiro, Sao Paulo and Parana,
distinguishable by its leaf blades membranaceous to chartaceous, with tertiary
(subsecondary) veins parallel to the secondary veins, inflorescences
pauciflorous and with delicate-thin rachis (often filiform), and flower buds
narrowly-tubular at base and ovoid at apex.
The names Ixora heterodoxa, 1. dimorphophylla and 1. burchelliana are
synonymous and have equal priority because they were published in the same
article by Mtiller Argoviensis (1875). The name I. heterodoxa is chosen over the
others because its holotype is a complete specimen with flowers in bud and in
anthesis, while the holotype of I. burchelliana (G-DC) is only represented by a
leaf fragment. No type speci sof | dimorphophyllaare present at G-DC, and
its holotype is at C, as it was indicated by Muller Argoviensis (1875).
_
DELPRETE, TRAVINYINET VE 1477
Ixora syringiflora (Schltdl.) Muell. Arg., Flora 58:455. 1875. Psychotria syringaeflora
Schltdl. Linnaea 28: 516. 1856. TYPE: BRAZIL [Bahia?|: locality unknown, s.d.[1828-
1856], Blanchet 3348 (HOLOTYPE: Bt, photo at NY).
Ixora benthamiana Muell. Arg., syn. nov., Flora 58:455, 459. 1875. Type: BRAZIL. RIO DE JANEIRO:
Rio Sao Joao, s.d. [1817-1820], Pohl 2209 (LECTOTYPE, G-DC n.v., here selected, photo at NY!).
Tree to 10 m tall; branchlets glabrous. Stipules 4-6 = 2.5-3.5 mm, base deltoid,
arista 3-4 mm long, glabrous outside, densely sericeous-pubescent and inter-
mixed with colleters inside. Leaf blades elliptic, oblong-ovate, to narrowly obo-
vate, 5.5-21 x 3-6.5 cm, apex acute, commonly deltoid-acuminate, base acute,
membranaceous to chartaceous, glabrous; secondary veins 16-34 each side, ter-
tiary venation reticulate; petioles 5-20 mm long, glabrous. Inflorescences pe-
dunculate, paniculate, multiflorous, LO-14 x 10-14 cm, with 4-5 lateral branches
laxely paniculate; peduncles L.5-4 cm long, glabrous; leaves subtending the in-
florescence smaller than those of lower nodes; bracts subtending the second-
ary branches narrowly triangular to narrowly elliptic, 6-15 x 1-4 mm long;
bracteoles | or 2 each flower, ovate to oblong, 0.7-1 mm long, or rarely absent.
Flower buds oblong-ellipsoid at apex. Pedicels absent or 2-5 mm long. Flowers
sessile or short-pedicellate, fragrant. Calyx 0.7-1 mm long; lobes triangular or
ovate, glabrous. Corolla 6.5-7.5 mm long, cream-white; tube 2-2.9 x 0.8-1 mm,
glabrous outside, sparsely pubescent at distal portion inside; lobes oblong, 3.8-
4.6 x 15-18 mm, apex round, glabrous. Stamens ca. 4.5 mm long; anthers nar-
rowly oblong, ca. 3 mm long, apex apiculate. Style 6.5-8.5 mm long; style
branches narrowly oblong, 1.5-2 mm long. Fruit unknown.
Illustration.—Mart., Fl. Bras. 6(5), Tab. 10. 1881 (as Lxora benthamiana)
Additional specimens examined. BRAZIL. $40 Paulo: Mun. Sao Paulo, Fda. Itatuba, 23 Dec 1931 (1D,
Amaral s.n.(SP 28628) (K, MO, NY, SP): Mun. Porto Ferreira, Reserva Estadual de Porto Ferreira, 2 Dec
1981 (fl), Bertoni s.n. (UEC 16894) (NY, UEC); Mun. Campinas, Reserva Municipal de Santa Genebra, 7
Novy 1988 ({L), Leitdo Filho & Morellato s.n. (UEC 22901) (UEC).
This species easily recognized by its laxely paniculate, many-branched,
multiflorous inflorescences. It isa very rare, or probably extinct, species known
only from four historical collections from the states of Rio de Janeiro and Sao
Paulo (and probably Bahia, Blanchet 3348, the type specimen).
Ixora venulosa Benth., Linnaea 23:446. 1850. Type: BRAZIL. MINAS GERAIS: Caldas,
s.d. [1841-1874], Regnell 1275b (LECTOTYPE, K!, here selected, photo at NY}).
Shrub to 5 m tall or rarely tree to 9 m tall; bark rugose; branchlets glabrous.
Stipules 4-8 x 3-4 mm, base narrowly triangular to oblong, arista 1-3 mm long,
glabrous outside, densely sericeous-pubescent and intermixed with colleters
inside. Leaf blades narrowly elliptic, oblong-elliptic, narrowly obovate to obo-
vate, (1.5-)4-14 = 1-6 cm, apex acute, often acuminate, base acute or decurrent,
chartaceous to subcoriaceous, glabrous; secondary veins fine and dense, 30-60
each side, tertiary venation subparallel to the secondary (sometimes somewhat
—
1478 BRIT.ORG/SIDA 20(4)
reticulate); petioles 2-15 mm long, glabrous. Inflorescences pedunculate, pan-
iculate, pauciflorous, 2-8 x 1.5-6 cm, with 1 or 2 lateral branches, these sparsely
branched; peduncles 1-6 cm long, glabrous; bracts subtending the secondary
branches narrowly triangular, 7-10 x ca.1 mm long, to narrowly elliptic, 10-13
« 3-5 mm; bracteoles | or 2 each flower, narrowly oblong-triangular, or absent.
Flower buds ovoid at apex. Pedicels absent or to 2 mm long. Flowers sessile or
short-pedicellate, fragrant. Calyx 0.3-1.2 mm long; lobes triangular or ovate,
glabrous. Corolla 8.5-10.5 mm long, cream-white; tube 4.5-6.5 x 0.7-0.9 mm,
glabrous outside, sparsely sericeous-pubescent inside; lobes oblong-ovate, 3.5-
3.8 = L5-2.2 mm, apex round, glabrous, sometimes with undulate margin. Sta-
mens 3.5~-4 mm long; anthers narrowly oblong, 2.5-3 mm long, apex apiculate.
Style 6.5-8.5 mm long; style branches oblong, 1.3-1.7 mm long. Fruit globose, 5-
6.5mm in diam.,, ee red.
Selected specimens examined: BRAZIL. Minas Gerais: Monte Belo, Fda. Monte Alegre, 21 Feb 1982
(fr), cae 300 (NY); Monte Bela *da. Lagoa, Mato Olaria, tree n. 498, 2 Nov 1987 (f1), Vieira 1334
(NY). Parana: Mun. Antonina, Reserva Biologica de Sapitanduva, 2525'S, 48°43'W, 28 Nov 1984 (f1),
Cervi etal. s.n.(UPCB 22718) (NY); Mun. Para nagua, Serra da Prata, 50 m, 2 Nov 1962 (fl), Hatsch bach
9458 (HBR, NY); Mun. Morretes, Prainhas, 50 m, 29 Nov 1966 (f1), Hatsch bach 15290 (HBR, NY): Mun.
Sao José da Boa Vista, Rio Jaguariaiva, Corredeira Paulista, 19 Nov 1970 ({D, Hatschbach 25569 (NY):
Mun. Guaraquecaba, trail to Paraquara, 28 Oct 1971 ({)), Hatschbach 27697 (US), Mun. Cascavel,
Autodromo de Cascavel, 25 Oct 1975, Hatsch bach 37352 (US); Mun. Cerro Azul, Rio Turvo, 5 Oct 1977
(fD, Hatschbach 40215 (NY); Mun. Marumbi, Serra do Mar, Nov 1971, Kuniyoshi 3121 (US); Mun. Cerro
Azul, Mato Preto, Rio Ribeira, 10 Feb 2001 (fr), Ribas & Silva 3214 (US); Mun. Sao José dos Pinhais,
Castelhanos, Rio Arraial, 30 Oct 1996 (f1), Silva & Saldanha 1742 (NY, US); Mun. Guaratuba, Rio
Arraial, Fda. Bamerindus, | Nov 1996 (f1), Silva & Saldanha 1772 (US). Rio de Janeiro: Rio de Janeiro,
s.d., Glazious.n. (NY). Santa Catarina: Mun. Garuva, Trés Barras, 21 Jan 1958 ({r), Reitz & Klein 6223
(FLOR, HBR, NY, US): sen Joinville, s.d., Schenk 1209 (K. Schum. in Mart., Fl. Bras. 6(6):399. 1889)
Sao Paulo: Mun. Teodoro Sampaio, Parque Estadual do Morro do Diabo, 22 Jun 1994 (fr), Baitello 681
(SP, SPSF); Mun. Corumbatai, rd. Washington Luiz, 22°15'S, +7°42'W, 27 Oct 1993 (fl), Barreto et al. 1512
(ESA); Mun. Cabretva, 2316'S, +7°02'W, 16 Mar 1994 (fr), Barreto et al. 2175 (ESA. NY); Mun. Campinas,
Fda. Sao Vicente, Ll Oct 1989 (fl), Bernacci s.n. (ESA 13210, VEC 25044) (ESA, UEC), 29 Aug 1990 (fr),
H7(NY), Mun. Tieté, Instituto Agronémico, 20 Jul 1994 (fr), Bernacci et al. 528 (NY, UEC): Dto. Souza,
Condominio Colinas do Atibaia, 12 Sep 1995 ({D, — et al. 186 (ESA, NY, UEC); Mun. Ipetina,
11 Oct 1985 (1D, Catharini & Mantovani 471(ESA, NY); Mun. Anhembi, Fda. Barreiro Rico, 9 Oct 1979
(ED, César sn. CHRCB 2404) (HRCB), 16 Oct L981 ({),s.n = ), 15 Dec 1981 (fr), s.n. CHRCB 3049) (HRCB,
NY), Mun. Itirapina, 21 Dec 1983 (£1), Cesdrs.n.C HRCB 3770) (HRCB, NY); Mun. Assis, Estacao Ecologica,
9 Apr 1992 (fr), Durigan s.n. (SPSF eae F); Mun. Eldorado, Caverna do Diabo, 2438'S, 48°23'S, 9
Feb 1995 (fr), Fernandes et al. 32695 (ESA, NY, UEC); Mun. Botucatu, Fda. Sao Joao, 19 Oct 1988 (fL),
Gabriel s.n. (HRCB 10569) (HRCB): Mun. Guarulhos, Aeroporto Internacional, 22 Nov 1984 (1),
Gandolfi et al. s.n. (ESA 5605) (ESA, NY); Mun. Sao Pedro, Cachoeira da Peroba, 22°32'S, 47°56'W. 22
Feb 199] (fr), Gandolfi et al. s.n. (ESA 32657) (ESA, NY); Mun. Sao Paulo, Butantan, | Dec 1921 (f1),
Gehrt s.n.(SP7538)(NY, SP); Mun. Valinhos, area de reforma agraria, 16 Aug 1994 (fr), June-Mendacolli
et al. 609 (NY); Mun. Jundiai, Estagao eu eh do IAC, 28 Sep 1994 (f1), Jung-Mendagolli et al.
624 (IAC, NY, UEC), 5 Apr 1995 (fr), 1412 (NY, UEC); Mun. Ipanema, 20 Oct 1887 (f1), Loefgren s.n. (NY,
SP): gu Sao Paulo, Instituto de i ade Universitaria, 740 m, 19 Aug 1972 (fr), Klein 10210
(HBR, NY, US), 23 Nov 1973, 10980 (HBR, NY, US); Mun. Amparo, Monte Alegre, Fda. Sta. Isabel, 07
May er (ft), onan: & Lemos 1179 (NY, SP), Mun. Itararé, Ibiti, 900 m, 26 Jul 1946, Kuhlmann &
=
aan
DELPRETE, TAAUNUMT UF
1479
Kuhn 1386 (AC, NY), 29 Jul 1946 (st), Kuh] mann 1393 (SP); Mun. Sao Jodo da Boa Vista, 12 Nov 1947
(fl), Kuhlmann 1484 (IAC, NY); Mun. Jundiai, Serra do Japi, 7 Nov 1981 (f{1), Leitdo Filho et al.s.n.(UEC
13102) (NY, UEC), si: Ae 13146) (UEC); ie Santa Rita do Passo Quatro, Parque Estadual de
Vassununga, 26 Oct 1978 (fl), Martins s.n. (UEC 9999) (UEC), s.n. (UEC 10000) (UEC); Mun. Paulinia,
Fda. Sao Francisco, 20 ae ), Martinse ve sn feces Mun Aguas da P 1, Reserva
Estadual, 47°20'W, 21°52'S, 1120 m, 2] Mar 1994 (fr), Martins et al. 31413 (NY, SP); Mun. hee Fda.
Grota Funda, 16 Nov 1987 ({1), Meira Neto et al. s.n.(UEC 21316) (UEC); Mun. Avai, Rio Batalha, 9 Apr
1997 (fr), Miranda & Miranda 322 (SP), 16 Oct 1997 (£1), 376 (NY); Mun. Guarei, Sarandi, 23°20'S,
48°14'W, 9 Nov 1980 ({]), Neves & Cerantola IPH-USP 37 (UEC); Mun. Cajuru, Fda. Santana, 3 Oct 1999
(f1), sane et al. 1792 (SP); Mun. Jat, Fda. Santo Antonio, 14 Oct 1988 ({1), Nicolini s.n. HRCB 11930)
(HRCB); Mun. Américo Brasiliense, Fda. Serrinha, 25 May 1944 (fr), Pickel s.n. (SPSF 1108) (SPSF);
Mun. aes Bairro sais sitio a Francisco, 4 — 1946 (fl), Pickel s.n. (SPSF 2836) (SPSF):
Mun. Ipetina, R Cinco, LO May 1989 ({r), Rodrigues & Zandoval s.n.
(ESA 6510) oe NY); Mun. Itu, 25 Dec 1897 ({1), Russel 198 (SP); Mun. Campinas, Reserva Municipal
de Santa G 3 Jul 1991 (st), Zickel sn. (UEC 30350) (UEC); Moji Guacu, Reserva Biologica, 2 Jun
1992 (f1), Zifirine é Romaniuc Neto 16 (NY, SP).
PARAGUAY: Alto Parana: Rese em biolegiea, near Rio Itabd, embalse Itaipti of Rio Parana, 2505'S,
54°05'W, 16 Oct 1996 ({1), Schinini et al. 31471 (N
A species readily distinguishable for the leaf blades with fine, dense venation
(30-60 secondary veins each side), common in the moist Atlantic forests of the
Brazilian states of Rio de Janeiro, Sao Paulo, Parana, and Santa Catarina, and in
the gallery forests of the cerrado biome of Minas Gerais, and occasionally found
in Paraguay.
KEY TO THE NATIVE SPECIES OF IXORA OF THE STATE OF SAO PAULO, BRAZIL
. Leaf blade with dense, equal ee aa venation (30-60 each side); tertiary veins
dense and parallel to the secondary ve
I. venulosa
1. Leaf blade with lee unequal secondary venation (6-40 each side); tertiary
veins subreticulat
2. Corolla lobes ae than the tube; flower buds ovoid at apex
3. Corolla tube 6-8.5 mm ah lobes 2—3.5 mm long; stipules 5— 4 x 2.5-3.5mm,
with an arista 2-3.5 mm lo I. heterodoxa
3. Corolla tube 4.5-5.5 mm ong lobes 3.5-4 mm long; stipules 2.5-3.5 x 1.5-2
with an arista 1.5 mm lon
2. éeiais lobes longer than the tube; fee pues oblong- ellipsoid at apex.
4. Inflorescence 10-14 cmin diam
branched and laxely Eo aati |. syringiflora
4. Inflorescence 3-8(-10) cm in diam., with 2-3 secondary branches, these
sparsely branched and corymbiform
I.gardneriana
—5 secondary branches, these densely
|. brevifolia
ACKNOWLEDGMENTS
| am extremely grateful to Sigrid Jung-Mendacolli (IAC), coordinator of the
Rubiaceae treatment for the Flora Fanerogamica do Estado de Sdo Paulo, for
inviting me to contribute Ixora for the flora, for her productive collaboration
and for arranging the loans of Ixora from the herbaria of the state of Sao Paulo
to the NY herbarium. I am also much indebted to the Conselho Nacional de
Pesquisa e Desenvolvimento Cientifico e Tecnologico (CNPq), Brazilian Govern-
1480 BRIT.ORG/SIDA 20(4)
ment, for granting permission to perform field and herbarium work in the states
of Goids and Santa Catarina, during 1998-2000. | also wish to thank the direc-
tors and curators of the following herbaria for sending material on loan and for
facilitating the access to their collections: BR, C, ESA, FLOR, HBR, HRCB, IAC,
K, MO, NY, S, SPSPSE UB; UFG, UEC, UPCR, UPS and US. My sincere eratitude
goes to the staff of the G-DC herbarium for producing and sending the 80 black
and white negatives of Ixora types, and to The New York Botanical Garden for
printing the photos used in this study. Lalso wish to thank the staff of the Kew
herbarium (Kk), Jochen Heinrichs (GOET), and Arne Anderberg (S) for search-
ing lor the type specimens of Ixora brevifolia in their institutions. lam particu-
larly grateful to Roland Moberg, Curator of the UPS herbarium, for sending the
type specimen of Ixora brevifolia on loan to NY, and for additional informa-
tion. Special thanks are also due to Claes Persson (GB) and Petra de Block (BR)
for the constructive comments on the manuscript.
REFERENCES
BenTHAM, G. 1850. /xora. In: Plantae Regnellianae. Linnaea 23:446-448.
De Brock, P, 1998. The African species of /xora (Rubiaceae—Pavetteae). Opera Bot. Belg. 9:
l=28,
Devprete, PG. (In press) dxora.In:M.G.L.Wanpertey, G.J. SHEPHERD & A.M. Giuuiett! (Coordinators),
Flora Fanerogamica do Estado de Sao Paulo. Editora Hucitec, Sao Paulo.
Devprete, PG.,L.B. Smith, and RJ.Downs. (submitted) Rubiaceae. |n:A. Reis (Editor), Flora llustrada
Catarinense. Herbdario Barbosa pc areis Itajal, Santa Catarina.
MULLER Arcoviensis, J. 1875. Rubiaceae brasilienses novae. Flora 58:449-459.
Mutter Arcoviensis, J. 1881. /xora (Rubiaceae. Tribes |-VI). In: C.F.P. Martius, A.G. EicHter, and |.
Urean, Flora Brasiliensis, Vol. 6(5):57—75. F. Fleisher, Lipsia.
VecTer, 1H. 1988. Index Herbariorum, Part Il (7), Collectors Tt/mZ. Regnum Veg. 1
987-1213.
=.
ds
A NEW SECTION OF BACCHARIS (ASTERACEAE:
ASTEREAE), AND NOTES ON ALLIED TAXA
Daniel A. Giuliano Guy L.Nesom
Division Plantas Vasculares Botanical Research Institute of Texas
nse us La Plata 509 Pecan Street
Fort Worth, TX 76102-4060, U.S.A.
1900 La Plata, ARGENTI NA gnesom@brit.org
giuliano@museo.fcnym.unlp.edu.ar
ABSTRACT
; new section of Baccharis is described: sect. Punctatae, includi i ies f ly placed in sect.
identes(B bigelovii B.crassicuneata, B. mexicana, B.sulcata "B the ciples and B. zamoranensis).
—
Baccharis palmeri is excluded from sect. Aristidentes while the South American B. hirta is added.
Six species (B. auriculigera, B. huairacajensis, B klattii, B. maxima, B. volubilis, and B. zumbadorensis)
are added to sect. Tubulatae. A key is provided to differentiate these allied sections, as well as the
related sect. Bogotenses.
RESUMEN
Sed ib ion de Baccharis: sect. Punctatae, la cual incl
ubicadas en la seccion Aristidentes (B bige lovii. B. crassicuneata. B. mexicana, B ve ata 'B thesioides
y B. zamoranensis). Baccharis palmeri es excluida de! aseccion Aristidentes, mientras que la especie
sudamericana B. hirta es agregada a ella. Seis 5 (B.auriculigera, B. huairacajensis, B. klattii, B.
maxima, B. volubilis y B. zumbadorensis) son neregad as a la seccion Tubulatae. Se provee una clave
para diferenciar estas secciones afines, incluyendo asimismo la seccion Bogotenses.
Ina revision of the infrageneric taxonomy of North and Central American spe-
cies of Baccharis, Nesom (1990) proposed the section Aristidentes to include 12
species. Later, the same author (Nesom 1998) recognized two informal groups
within this section: the “Baccharis multiflora complex”, including 6 species (B.
brevipappa (McVaugh) G.L. Nesom, B. mexicana Cuatrec., B. multiflora Kunth -
the type species of the section-, B. serrifolia DC., B. sordescens DC, and B.
zamoranensis Rzed.), and the “Baccharis thesioides complex”, comprising 3 spe-
cies (B. bigelovii A. Gray, B. sulcata DC., and B. thesioides Kunth). With further
observation, the differences between these two groups have proved to be more
significant, especially when two closely related South American taxa, sect.
Bogotenses Cuatrec. and sect. Tubulatae Cuatrec., are also considered. Effectively,
the “B. multiflora group” shares more features with these South American taxa
than with the “B. thesioides group” (see key below); therefore, this taxon is here
given formal recognition at sectional rank, as follows.
Baccharis sect. Punctatae Giuliano & GL. Nesom, sect. nov. Type: Baccharis bigelovii
Gray
SIDA 20(4): 1481-1484. 2003
1482 BRIT.ORG/SIDA 20(4)
Sectioni Aristidenti similis, sed plantis glutinosis nidi pilosis vestitis, foliis uni- vel obscure trinervatis
et ak minoribus differt.
Erect shrubs or subshrubs, glutinous, covered by minute tufts of (microscopic)
biseriate glandular hairs; on both leaf surfaces, each tuft is placed in a depres-
sion of circular contour, altogether forming a characteristic dotted pattern.
Leaves pseudopetiolate, uni- or trinervate, margins aristate-serrate (rarely en-
tire). Capitula pedicellate, arranged in simple or compound corymbs, altogether
forming a false panicle. Involucres campanulate. Receptacles flat or slightly
convex, alveolate, epaleate. Ratio of carpellate: staminate florets ca. L:1. Carpel
late corollas filiform, apically truncate and crowned by long and short biseriate
hairs. Achenes terete, 1.2-2.2 mm long, 5-8-ribbed, glabrous or with few
uniseriate hairs with apiculate terminal cell; pappus bristles in 1-2 series, per-
sistent, slightly accrescent. Staminate corollas campanulate, 5-lobed at apex:
style branches lanceolate, free; pappus bristles apically broadened.
Includes 6 species, distributed from the southwestern United States to
southern Mexico.
LB. bigelovii A. Gray, Rep. U.S. Mex. Bound., Bot. 84. 1859.
crassicuneata G.L. Nesom, Phytologia 84:43. 1998.
mexicana Cuatrec., Brittonia 12:195. 1960.
sulcata DC., Prodr. 5:419. 1836.
. thesioides Kunth, in Humboldt, Bonpland & Kunth, Nov. gen. sp. 4
[quarto]: 61. 1820.
6. B. zamoranensis Rzed., Brittonia 24:400. 1972.
VIB WN
mo
Formerly, B. mexicana and B. zamoranensis were included in the “B. multiflora
complex” of sect. Aristidentes (Nesom 1998) but now are classified within
Punctatae, as they are glutinous plants dotted with tufted hairs.
Section Punctatae is closely related to sect. Aristidentes, sect. Bogotensesand
sect. Tubulatae; the four taxa form a complex characterized by the capitulescence
architecture (a false panicle of compound corymbs), achenes with 5 8 pale ribs
glabrous or with few uniseriated hairs, and pappus with persistent accrescent
bristles in 2-3 series. These sections can be differentiated by the following features:
1,
Plants glutinous, dotted by minute (microscopic) tufts of glandular hairs, without
other vestiture, Leaves uni- or obscurely trinervate. Achenes 1.2-2.2 ong.
Sect. Punctatae
. Plants non-glutinous,non dotted by minute hair tufts but densely or loosely clothed
by conspicuous (non-microscopic), non-glandular uniseriate hairs with acuminate
terminal cell, often incurved. Leaves usually reticulate-nervate, trinervate in some
species. Achenes 2.5-4 mm lo
2. Scan de nt shru Ibs, densel pube scent
with long uniseriate conic hairs accompa-
nying the uniseriate hairs with acuminate terminal cell. Capitula often grouped
in few-he nflorescences; carpellate involucres 7-9 mm long. Staminate
florets ue aie elongate limb and short straight lobes Sect. Tubulatae
scandent or erect shrubs, without conic hairs but densely or loosely clothed by
GIULIANO AND NESOM, A NEW SECTION OF BACCHARIS 1483
uniseriate hairs with acuminate terminal cell. Capitula usually grouped in many-
headed inflorescences; carpellate involucres 3-6 mm long. Staminate florets with
short limb and developed recurved lobes.
3. Scandent shrubs, subglabrous, with few isolated hairs. Sect. Bogotenses
3. Erect shrubs, densely pubescent. Sect. Aristidentes
The following couplet provides a summary of the contrasts between sect.
Punctatae and the taxon from which it was segregated, sect. Aristidentes:
_ Plants glutinous, dotted by minute tufts of (microscopic) glandular hairs, without
other vestiture. Leaves uni- or obscurely trinervate. Achenes 1.2-2.2 mm long.
Sect. Punctatae
1. Plants non-glutinous, non dotted by minute hair tufts but densely clothed by con-
spicuous (non-microscopic), non-glandular uniseriate hairs with acuminate termi-
nal cell, often incurved. Leaves usually reticulate-nervate, trinervate in some spe-
cies. Achenes usually 2.5-3 mm long. Sect. Aristidentes
Further notes on sect. Aristidentes
Several of the species originally included in sect. Aristidentes are here relocated
to the new section Punctatae. Baccharis palmeri Greenm. is also here excluded
from sect. Aristidentes since it lacks the pubescence characteristic of this taxon,
and it is placed in sect. Angustifoliae Baker emend. Cuatrec., considering that it
produces solitary capitula at the apex of the branches, which often are grouped
to form false ¢ apit ulesc ELLCES
On the other hand, one species needs to be added to sect. Art stidentes, since
it presents its typical features: B. hirta DC, from southern Brazil and Uruguay.
This species was previously classified in the formerly heterogeneous sections
Trinervatae DC. (Candolle 1836), Cuneifoliae DC. (Hooker & Arnott 1840,
Oblongifoliae DC. (Baker 1882) and Paniculatae Heering (Heering 1904). With
the 4 South American species added to sect. Aristidentes by Giuliano (2001), the
number of species for the section becomes 11: B. brevipappa (Mc Vaugh) G.L. Nesom,
B. hirta DC.,B. lilloi Heering, B. malibuensis R.M. Beauch.& Henrickson, B. mul-
tiflora Kunth, B. plummerae A. Gray, B. polygama Ariza, B. pulchella Sch. Bip. ex
Griseb., B. racemosa (Ruiz & Pav.) DC., B. serrifolia DC, and B. sordescens DC.
ame
Addition of species into sect. Tubulatae
Section Tubulatae was founded by Cuatrecasas (1967), who included 3 Andean
species (B. fraterna Cuatrec., B. granadina Cuatrec., and B. grandiflora Kunth).
Here it is considered necessary to add 5 species more, as they accord with the
diagnostic features of this taxon: the Andean B.auriculigera Hieron. (previously
placed into sect. Trinervatae, Hieronymus 1905), B. huairacajensis Hieron,
(placed in sect. Oblongifoliae when originally described; Hieronymus 1901), B.
klattii Benoist (never classified before), B. volubilis Kunth (formerly in sect.
Oblongifoliae; Candolle 1836), and B. maxima Baker, endemic of the state of Rio
de Janeiro in Brazil (placed in sect. Oblongifoliae when described; Baker 1882).
Another species, the Venezuelan endemic B. zumbadorensis V.M. Badillo, is
1484 BRIT.ORG/SIDA 20(4)
also placed here although staminate specimens are still unknown, since it has
scandent habit, long conic hairs, reticulate-nerved leaves, large few-headed ca-
pitula, and large achenes, which fit properly in Tubulatae. The nine species of
this section are mainly Andean in distribution.
ACKNOWLEDGMENTS
We are grateful to Scott Sundberg for his insightful and researched review of
the manuscript. This study was carried out partly (D.A.G.) by means of a fel-
lowship (Beca de Formacion de Postgrado”) conceded by the Consejo Nacional
de Investigaciones Cientificas y Técnicas de la Republica Argentina (CONICET).
REFERENCES
Baker, J.G. 1882. Compositae: Baccharis.In:C.F. Martius (ed,), Fl. bras.6 (3):37-100. Munchen,
Wien, Leipzig.
Canpoite, AP. be. 1836, Compositae: Baccharis. In: Prodr.5:398-429, Paris (Treuttel et Wirtz).
Cuarrecasas, J. 1967. Revision de las especies colombianas del género Baccharis. Revista
Acad. Colomb. Ci. Exact. 13(49):5-102.
Giuliano, D.A. 2001. Clasificacién infragenérica de las especies argentinas de Baccharis
(Asteraceae, Astereae). Darwiniana 39:131-154.
HeeRING, W.C.A. 1904. Die Baccharis-Arten des Hamburger Herbars. Jahrb. Hamburg. Wiss.
Anst. 21 (Beih. 3):1-46.
Hieronymus, G.H.E.W. 1901. Baccharis.In: Plantae Lehmannianae praesertim in Columbia et
Ecuador collectae, additis quibusdam ab aliis collectoribus ex iisdem regionibus allatis
determinatae et descriptae. Compositae II. Bot. Jahrb. Syst. 28:588-594,
ligRONYMUS, G.HL.E.W. 1905. Baccharis. In: Plantae peruvianae a claro Constantino de Jelski
collectae. Compositae. Bot. Jahrb. Syst. 36:477—483.
Hooker, W.J.and G.A.W. Arnott. 1841.Compositae: Baccharis.In:Flora of South America and
the islands of the Pacific. J. Bot. (Hooker) 3:21—44,
Nesom, G.L. 1990. Infrageneric taxonomy of North and Central American Baccharis
Asteraceae: Astereae). Phytologia 68:40—-46.
Nesom, G.L. 1998. Two newly recognized species of Baccharis (Asteraceae: Astereae) from
Mexico. Phytologia 84:43-49.
—
TWO NEW COMBINATIONS IN BOLIVIAN
GENTIANELLA (GENTIANACEAE)!
James S. Pringle
Royal Botanical Gardens
PO. Box 399, Hamilton, Ontario
CANADA L8N 3H8
jpringle@rbg.ca
ABSTRACT
Gentianella boliviana and G. inaequicalyx cognized as species and the nomenclatural combi-
nations are validated.
RESUMEN
Se reconocen como especies Gentianella boliviana y G. inaequicalyx, y se validan las combinaciones
nomenclaturales.
Two new combinations are required for identifications and for use in the Cata-
logue of the Vascular Plants of Bolivia, in progress at the time of this writing,
edited by Peter Moller Jorgensen.
GENTIANELLA BOLIVIANA
Gentiana boliviana Pax, G. hicronymi Gilg, G. lobelioides Gilg, and G. peruviana
(Griseb.) Gilg, which had been recognized as four species by Gilg (1916), were
treated by Ho and Liu (1993) as a single species, which they called Gentianella
peruviana (Griseb.) Fabris. Nomenclaturally, this treatment is incorrect. The
name Gentianella peruviana (Griseb.) Fabris was based on Gentiana
limoselloides[var] B peruviana Griseb. The name Gentiana peruviana (Griseb.)
Griseb. was based on this varietal name as to the basionym but not as to the
plants described. When it was published in 1879, it was an illegitimate hom-
onym of Gentiana peruviana Lam. 1786, which is a taxonomic synonym of
Centaurium cachanlahuen (Molina) B.L. Rob. The name Gentiana peruviana
(Griseb.) Gilg 1896 was based on G. limoselloides [var] B peruviana Griseb. both
as to basionym and as to plants described, but by typification was a superflu-
ous combination. It likewise is illegitimate. There was no obstacle to the use of
the epithet peruviana in Gentianella by Fabris (1958), but, as he recognized,
the priority of the epithet peruviana at the rank of species, as applied to any of
the Gentianella taxa discussed here, dates only from its use in this legitimate
combination in 1958. The epithets hieronymi, boliviana, and lobelioides have
priority at the rank of species in this context dating from their publication in
1 ntribution No. 117 from the Royal Botanical Gardens, Hamilton, Ontario, Canac
SIDA 20(4): 1485-1490. 2003
d.
1486 BRIT.ORG/SIDA 20(4
=
—
legitimate combinations in 1896, 1909, and 1916, respectively. Therefore, if the
four taxa recognized by Gilg in 1916 are treated as conspecific, the correct name
for the species thus circumscribed is Gentianella hieronymi (Gilg) Fabris. The
name Gentianella peruviana (Griseb.) Fabris is correct only when the species
to which it is applied is more narrowly defined, as it was by Fabris.
Within G. peruviana, Ho and Liu (1993) recognized two varieties: var.
peruviana, in which they included G. hieronymi of Gilg 1916); and var. boliviana
(Pax) T.N. Ho, which was equivalent to Gentiana boliviana plus G. lobelioides
of Gilg 1916). Having studied nomenclaturally significant material of both taxa,
| have chosen to recognize G. bolividna as a species rather than to consider it
conspecilic with Gentianella hieronymi.
Gentianella boliviana (Pax) J.S. Pringle, comb. nov. BASIONYM: Gentiana boliviana
Pax, Repert. Spec. Nov. Regni Veg. 7: 243. 1909. Gentianella peruviana (Griseb.)
Fabris var. boliviana (Pax) T.N. Ho, Bull. Nat. Hist. Mus. London, Bot. 23:64. 1993.
Type: BOLIVIA. LA PAz: Chacaltaya, 30 km from La Paz, 4800 m, Buchtien 1482
(HOLOTYPE: B, destroyed; ISOTYPE: US, microfiche MO}).
—
—
Gentianella boliviana was further represented in my studies by Beck 7477 and
8022 (HAM, [LPB]), Menhofer X-2312 (HAM, [LPB], Solomon 5066 (MO), Solomon
& Moraes R. 13462 (MO), and Ruthsatz 1057 and 1105 (both HAM). (Brackets
indicate replicates not seen by me.) All of these specimens were collected near
the type locality in Depto. La Paz.
Gentianella hieronymi is native to Provs. Jujuy and Salta, Argentina. The
type collection is Hieronymus & Lorentz 15 (uo.otye: B, destroyed, photo Fi:
isoryPrs: CORD, K!). It and G. boliviana are well separated geographically.
Gentianella boliviana and G. hieronymi differ as indicated in the follow-
ing descriptions:
Gentianella boliviana (Pax) J.S. Pringle. Herbaceous perennial, |-4 cm (includ-
ing peduncles and flowers). Flowering stems usually only | or 2, occasionally
to 6, decumbent, unbranched. Leaves all basal or cauline leaves | pair, all leaves
narrowly obovate to linear-oblanceolate, +-10(-16) x 1-4 mm, rounded or ob-
tuse. Flowers solitary; peduncles 1-3 cm. Calyx 5-8 mm, lobed 0.45-0.55x its
length, lobes narrowly obovate to oblong, rounded to obtuse or occasionally
subacute. Corolla light blue with darker veins, widely campanulate, 10-16 mm,
lobed 0.54-0.65x its length, lobes oblong-obovate, rounded at apex.
Gentianella hieronymi (Gilg) Fabris. Herbaceous perennial (?), 2-7 cm (including
peduncles and flowers). Flowering stems usually 5-12, decumbent, unbranched.
Basal leaves narrowly obovate to oblanceolate, 8-12 x 2-4 mm, obtuse; cauline
leaves usually none, occasionally or | or 2 pairs, elliptic to linear, 6-10 X 1-2 mm,
obtuse. Flowers solitary; peduncles |-4 cm. Calyx 8-14 mm, lobed 0.45-0.65 its
length, lobes oblong, rounded at apex. Corolla pale lilac, campanulate, 18-25 mm,
lobed 0.52-0.60 its length, lobes elliptic-obovate, subacute.
PRINGLE, 1487
Among the several low-growing, solitary-flowered species of Gentianella
native to Depto. La Paz, G. boliviana is especially distinctive. The plants of this
species are generally smaller than those of any other Gentianella species in La
Paz. Its distinctive features include calyx lobes about as long as the tube, usu-
ally rounded at the apex, and corollas 10-16 mm long, with widely spreading
lobes slightly longer than the tube, widely rounded at the apex “obtusissimus”
in the original description). The corollas of both of the specimens at MO were
described as “pale blue” and those of Beck 8022 as “verde-azul palida.” In her-
barium specimens, this color, when well preserved, is reminiscent of that of
Amsonia tabernaemontana Walter of North America, and is an instant guide
to the recognition of G. boliviana. In the other solitary-flowered Gentianella
species in Depto. La Paz the corollas are predominantly white, in most cases
marked with dark blue-violet.
Gentianella hieronymi, which is not native to Bolivia, differs most conspicu-
ously from G. boliviana in its larger, more floriferous plants, more narrowly
campanulate calyces; larger, pale lilac, less widely opening corollas, and pro-
portionately narrower corolla lobes that are subacute at the apex.
Many low-growing species of Gentianella with solitary- or few-flowered
stems are present in Argentina, Bolivia and Peru, and not all of the taxonomic
questions pertaining to these species can be resolved within a single paper.
Gentiana lobelioides, now called Gentianella lobelioides (Gilg) Zarucchi, is
known only from the type specimen, Weberbauer 955 (holotype B, destroyed,
photo F!), from Depto. Puno, Peru, directly across the border from Depto. La Paz,
Bolivia. From my study of published descriptions and a photograph of the type
its inclusion within G. boliviana seems reasonable. Notably, the corollas were
described as “pallide-coerulei” (Gilg 1916), and those of the type specimen ap-
pear to have been ca. 11 mm long. However, since the type was in poor condition
when photographed and I have encountered no specimens from Peru referable
to G. lobelioides, its disposition is not within the scope of this paper. With G.
boliviana (and probably G. lobelioides) excluded, populations referable, respec-
tively, to G. hieronymis.str.in Argentina and G. peruvianas. str. in Depto. Cuzco,
Peru, are widely separated. These species were considered distinct by Fabris
(1958), who emphasized differences in the proportionate lengths of the calyx
lobes and tube as well as in corolla size. From descriptions (Gilg 1916; Fabris
1953, 1958) and photographs of the type specimens their recognition as sepa-
rate species seems tenable, but since | have not seen specimens referable to G.
peruviana s. str, further discussion of that species is not included here.
GENTIANELLA INAEQUICALYX
Ho and Liu (1993) included five species recognized by Gilg (1916) within their
circumscription of Gentianella silenoides (Gilg) Fabris. They treated this spe-
cies as comprising three varieties: var. silenoides, equivalent only to Gentiana
1488 BRIT.ORG/SIDA 20(4)
silenoides of Gilg (1916); var. striaticalyx (Gilg) T.N. Ho, within which they in-
cluded Gentiana striaticalyx Gilg, G. anthosphaera Gilg, and G. herzogii Gilg;
and var. indequicalyx (Gilg) T.N. Ho, based on Gentiana inaequicalyx Gilg. The
epithets silenoides and inaequicalyx both have priority at species rank dating
from 1896, the other epithets from 1906 or 1916. Ho and Liu indicated that they
had seen no specimens relevant to the typification of the epithets silenoides,
striaticalyx, and herzogii, and cited no specimens that would exemplify their
concepts of these taxa. They had seen isotypes of the names Gentiana
antho pus and G. indequicalyx.
| have examined photographs of the type specimens of all five of these epi-
thets, as well as isotypes, paratypes, and more recent specimens from sites near
the respective type localities that corresponded closely to the respective type
specimens. My studies have in part supported Ho and Liu’s revision, but have
eas
also indicated that some modifications would be appropriate. Specifically,
Gentianella silenoides s. str. and the taxon treated here as G. indequicalyx are
distinctly dissimilar and conform well to standards for the recognition of spe-
cies in South American Gentianella.
Gentianella inaequicalyx (Gilg) Fabris ex J.S. Pringle, comb. nov. BASIONYM:
Gentidana inaequicalyx Gilg, Bot. Jahrb. Syst. 22:324. 1896. Gentianella silenoides
(Gilg) Fabris var. inaequicalyx (Gilg) T.N. Ho, Bull. Nat. Hist. Mus. London, Bot.
23.04.1993. TYPE: BOLIVIA. LA PAZ: Vicinis Sorata, 2650-3000 m, eae 365 (LEC
TOTYPE, here designated, G, photos Fl, NY!: lsoTYPES: GH], K!, NY!,S, US). PARATYPES:
BOI IVIA. COCHABAMBA: Vic. Cochabamba, Bang 1143 (GH, KI, an Coch abamba:
Tunari, 3000-4000 m, Kuntze s.n. (probably B, destroyed).
Gentidnella inaequicalyx was further represented in my studies by Luteyn &
Dorr 13799 (HAM, [LPB, NY), from Depto. La Paz, Bolivia, and Vargas C.& Cruz
G. 4980 (HAM, INY, USZ)), from Depto. Santa Cruz, Bolivia. Existing data indi-
cate that the range of G. inaequicalyx is more northern than that of any of the
other entities included in G. silenoides s. lat. by Ho and Liu.
The type of the name G. silenoides is Lorentz & Hieronymus 877 (holotype
B, destroyed, photo Fl), from Depto. Tarija, Bolivia, This species was further rep-
resented in my studies by Bastidn 1183 and 1312 (1AM, |[LPB)), Ehrich 199 (HAM,
[LPB]), and Solomon 10197 and 10262 (HAM, |[MO}), all from Depto. Tarija, Bo-
livia. It has also been reported from the adjacent Prov. Salta, Argentina (Fabris
1953).
Gentianella inaequicalyx and G. silenoides differ as indicated in the fol-
lowing descriptions:
Gentianella inaequicalyx (Gilg) Fabris ex J.S. Pringle. Herbaceous annual, 6-25
cm. Primary stem usually solitary, erect or curved only at base; branches all +
strongly ascending. Basal and proximal cauline leaves mostly oblanceolate to
elliptic, 8-20 x 1-4 mm, obtuse, often absent at flowering time; distal
—
Caves
PRINGLE, 1489
gradually narrower, lanceolate to linear, 5-30 x 1-5 mm, acute. Inflorescence a
dense cyme; flowers sessile or pedicels up to 1 cm. Calyx 6-15 mm, lobed 0.50-
0.75x its length, lobes often distinctly unequal, linear, subacute. Corolla white
with blue-violet tinge or blue-violet throughout, tubular-campanulate, open-
ing narrowly, 11-22 mm, lobed 0.35-0.45x its length, lobes oblong-ovate, ob-
tuse or abruptly short-acuminate.
Gentianella silenoides (Gilg) Fabris. Herbaceous annual, 8-30 cm. Primary stem
usually solitary, erect or + decumbent; branches widely spreading to ascend-
ing. Basal and proximal cauline leaves mostly elliptic, 8-20 x 1-4 mm, obtuse
to acute, sometimes absent at flowering time; distal leaves gradually narrower,
lanceolate to linear, 7-30 X 1-4 mm, acute. Inflorescence a diffuse cyme; pedicels
1-10 cm. Calyx 9-15 mm, lobed 0.55-0.95x its length, lobes usually subequal,
narrowly linear, acute to acuminate. Corolla white to pale blue-violet, narrowly
campanulate, opening + widely, 15-26 mm, lobed 0.45-0.55x its length, lobes
elliptic-ovate, acute to acuminate.
Plants of G. inaequicalyx are stiffly erect, with dense, many-flowered in-
florescences in which the flowers are sessile or on pedicels less than | cm long.
Its aspect, consequently, is similar to that of the North American G. quinquefolia
(L.) Small. Its corollas are more narrowly tubular than those of G. silenoides,
and are proportionately less deeply lobed. Gentianella silenoides differs in its
diffuse inflorescence with widely separated flowers on pedicels 1-10 cm long,
conforming to Gilg’s (1916, in the key) description, “Flores in cymas laxas
dispositi.” Its aspect is consequently similar to that of another Bolivian species,
G. lythroides (Gilg) Fabris ex T.N. Ho & S.W. Liu, which differs in its more deeply
lobed corollas.
The other reductions to synonymy or varietal status within G. silenoides
by Ho and Liu have been supported by my studies, in particular the reduction
of G. herzogii. Solomon 10262, which, having corollas 28-36 mm long, would be
identifiable as G. herzogii, and Solomon 10197, with corollas 15-18 mm long, in
the size range given by Gilg (1916) for G. silenoides, were collected at the same
locality in Depto. Tarija. They exhibit only modest differences in the size of the
plants and in the size of the leaves, peduncles, and flowers, rather than any quali-
tative differences. The distinction between var. silenoides and var. striaticalyx
(including plants recognized by Gilg both as Gentiana striaticalyx and as G.
anthosphaera), is so slight that even varietal status might be questioned.
Further studies of taxa in this complex would be desirable. Gentianella
pallidolilacina (Gilg) TN. Ho & S.W. Liu was accepted as a distinct species by
Ho and Liu (1993), but studies of additional material might indicate that it
should be included within G. inaequicalyx. If these taxa were to be united (ex-
clusive of G. silenoides) the epithet inaequicalyx would have priority.
1490 BRIT.ORG/SIDA 20(4)
REFERENCES
Fapris, H.A. 1953. Sinopsis preliminar de las Gencianaceas argentinas. Bol. Soc. Argent. Bot.
232-259.
Fasris, H.A. 1958. Notas sobre Gentianella del Peru. Bol. Soc. Argent. Bot. 7:86-93.
Giic, E. 1916. Gentianaceae Andinae. Bot. Jahrb. Syst. 54(Beibl. 118):4-122.
Ho, T.N.,and S.W. Liu. 1993.New combinations, names and taxonomic notes on Gentianella
(Gentianaceae) from South America and New Zealand. Bull. Nat. Hist. Mus.London, Bot
BOTANICAL SUMMARY OF A LOWLAND ULTRABASIC
FLORA IN PAPUA NEW GUINEA
W. Takeuchi
| ; dA ld Arhoretil4m
Ly Ve ha ny
PIG VGIG UTTHVEEL
\
Research Associ Ite, Village D
+
velopr FICHE
Lae, Morobe P. Province 41 1, PAPUA NEW GUI he
ABSTRACT
The ultrabasic flora of the Kamiali Wildlife Man agement Area is described from a recent series of
ne surveys. The general account and checklist also incorporates information from earlier col-
ns. Two new species are presented: Psychotria bulilimontis (Rubiaceae), and Trichadenia sasae
ctio
et icaceae). Other taxa of special interest are briefly discussed.
Key Worps: botanical survey, serpentine, ultrabasics
JAPANESE ABSTRACT
DAT V BAA HK OREM OVC, Be pbc HOM Die
Bo gs ena ANA aegaeie b fel heckuva 1 1 AGH BD HSRC Stik >
Hate LE HEITR AM Ith ote. ZOOM. tria bulilimontis (7 BAB) & Hla ae sasae
(AA SVRIZTOWTHRTSZEL es EIT PURE V Mi DZ) EIT OUT & fi HAZ
%—O— BS ARAB. RECA. HEEL Be
INTRODUCTION
After more thana century of scientific publication on the Papuasian flora, the
ultrabasic (i.e. ultramafic or serpentine) vegetation of Papua New Guinea (PNG)
is still largely unknown in spite of its unusual qualities. Multidisciplinary sur-
veys were initiated in 1997 to address the need for information from such areas,
using the Kamiali Wildlife Management Area (KWMA) as a focal site for
Diogcr assessment (Figs. 1-2). The selected location offered a number of
PNG environments, as it included a com-
plere elevational sequence of natural growth habitats from sealevel to cloudy
sumunits, and its exploration could be conducted with cost-effective itinerar-
ies. In the following narrative, botanical results from the recent investigations
are presented and integrated with earlier work in the Papuan Ultrabasic Belt.
HISTORICAL BACKGROUND
The first collections from southern Morobe were made in the Waria drainage
(Fig. 3) by R. Schlechter (in 1908; see Schlechter 1911-14) after which there was
a general neglect of the district until the NGF/LAE collections of the former
Department of Forests. Most of Schlechter’s specimens were later lost during
the WWII destruction of the Berlin Herbarium. In 1990, Clements rediscovered
SIDA 20(4): 1491-1559. 2003
2
3 ss ce
oe!
on
2
ee
. - Indonesian Papua
e J
gy 400 km :
LO?
oa
Fic. 1. Island of New Gui Shaded lowland i laf the P TT eee
76rL
(p)02 VOIS/ONO'LINS
TAKEUCHI, ULTRABASIC FLORA IN PAPUA NEW GUINEA 1493
\ pu BE
wd} hake Drase ‘ Mu CN
qo Sa
AE tare ere ae
Fic. 2. Kamiali Wildlife M A (KWMA) Light | | KWMA it 1 y Darker-colored
perimeter: customary lands ‘daimed by the Kamiali people Shaded elliptical principal y sites. Contours:
250; 500; 1000 then subsequently in increments of 10 pted Village Develoy Trust fil i
several of the lost German taxa and described some new ones (Clements &
Ziesing 1990). Exploration of nearby Natter Bay (e.g. by Croft in 1976) also
yielded interesting new material (Fig. 4). Since 1908 there has otherwise been
little collections activity between the Waria River and Paiewa (Paiawa).
Particularly during the 1970s, the area around Buso was used as a training
site by the Bulolo Forestry College (now Bulolo University) and many speci-
mens were taken during periodic visits by expatriate botanists of that general
period (inter alia B. Conn, D. Foreman, A. Gillison, E. Henty, M. Jacobs, A. Millar,
C. Ridsdale, and H. Streimann). Although the Buso locality is thus relatively
well documented in comparison to adjacent areas, it was later despoiled by log-
ging subsidiaries of Rimbunan Hijau. Confronted by mounting environmental
concerns from landowners and advocacy groups, the last logging company
(Timber Products Marketing Corporation) was forced to leave in 1995, but by
1494 BRIT.ORG/SIDA 20(4)
Fic. 3. A typical non the Waria River.
+
a]
>
a
Fic. 4. Natter Bay. C
polyandra
TAKEUCHI, ULTRABASIC FLORA IN PAPUA NEW GUINEA 1495
then an extensive territory between Buso and Morobe Bay had been harvested
by commercial operators (Fig. 5).! Athough remnant Anisoptera-dominant
stands are still scattered along the S Morobe coast (e.g. at Siboma or ‘Simpoma’),
most of the low elevation forests are currently in early successional recovery.
The Kamiali Wildlife Management Area (KWMA) is now the only significant
primary-growth habitat in the Huon electorate southeast of Lae.
In 1992-1993, the villages of SE Morobe (i.e. Kamiali, Buso, Kui, Siboma,
and Paiewa) developed a cooperative proposal for environment-sensitive alter-
natives to the concessional logging which was then occurring (Zibe-Kokino
1993). Known as the Lasanga Island-Lake Trist Conservation Project, the initia-
tive was based on the principles of Integrated Conservation and Development
(ICAD), an economic model which promotes smallscale enterprise as a substi-
tute for ecologically-disruptive activities.
Since its early beginnings at Crater Mt, the ICAD methodology has devel-
oped into the principal present-day mechanism for environmental conserva-
tion in PNG, as evidenced by the growing body of literature produced by its
proponents (e.g. James 1996; Johnson 1997; McCallum & Sekhran 1997; Pearl
1994; Saulei & Ellis 1998; Wagner 200la, in press). Several major programs are
currently being implemented under the ICAD format, most notably by testbed
projects in the Adelbert Mts, Crater Mt, Hunstein, Kamiali, Kikori, Milne Bay,
and Tonda. However, some conspicuous failures have occurred (McCallum &
Sekhran 1997).
Many issues and problems encountered at other ICAD localities were also
experienced by the Lasanga-Lake Trist initiative, in particular the inability of
stakeholders to achieve a consensus as each clan group fought to maximize its
own benefits. After the original design for a 250,000 ha easement was compro-
mised by funding difficulties, the proposal was recast as a 47,000 ha conserva-
tion area including only the Kamiali territory (Anon. Sept. 19, 1996). This re-
duction in scope enabled the project to proceed by focusing participation on a
more homogeneous combination of landowners, but at a probable loss in pro-
tection of biodiversity values.
The present KWMA isa linked land-sea conservation zone, of which only
29,285 ha is terrestrial environment (ibid.). While there is little consensus on
the minimum size requirement for protected areas, it is unlikely that the exist-
ing KWMaA is large enough to serve as a stand- alone « conservation unit. The
self-sustainability of biotic ities arguably ly larger
territories, especially in view of the destructive effects of probable disturbance
‘Additional timber ion bloc} I d by the National Forest Plan (Anon. 1996) for
Salamaua, Lake Trist. and Waria Valley. Placed in eae ie a country-wide forestry moratorium, ue poe
tions against new logging projects were recently lifted. However as a result of
ous years, it will be difficult for industrial operators to reenter the general area, in part Bees shine environ-
mental ad now conducted by organizations such as VDT.
1496 BRIT.ORG/SIDA 20(4)
4}
Salamaua
Lokanu-Tambu Bay
%
?
"ROVINcg
NORTHERN PROVINCE
Fic. 5.F producti i h Morobe (striped blocks 3—7). 3. Kui Buso TP (Timber Purchase); 4. Kui Buso exten-
sion TP; 5. Maiama-A TP; 6. Maiama-B TP; 7. Morobe TP. Other localities f I Iso indi
I AMarth D * Adantaod £
t
The Waria River
\
TAKEUCHI, ULTRABASIC FLORA IN PAPUA NEW GUINEA 1497
mechanisms such as cyclonic storms and El Nino events. In depauperate for-
ests with monodominant canopies (e.g. Hawaii),a complete suite of species and
forest successional stages can be encompassed by tracts as small as 100-200 ha
(Mueller-Dombois 1980). But New Guinea floristic environments are among
the richest in the paleotropics, and the minimum-area requirements are sev-
eral orders of magnitude higher Jeffries (1996) for example, gives 50,000 haasa
lower limit for biodiversity protection, while Beehler and Alcorn (1993) go as
high as 800,000 ha. Even within that wide interval, the existing Kamiali ease-
ment falls substantially under target. Against this background, the impending
expansion of the KWMA toa southern boundary near Paiewa will help to ad-
dress some of the biological requirements implicit in ICAD design. Sanctioned
by local communities in 2002, the new development will bring ca. 300,000 ha
of terrestrial environment under conservation management (Kisokau & Siga,
pers. comm. Feb. 2003)
Although traditional cultures in PNG are often depicted as being based on
consultation and consensus, achieving a determination of common purpose or
directed action is never easy. Centrifugal forces are powerful in local politics, as
clan, subclan, and — self-interests are more important than any percep-
tion of the common good (Martin 1998). Expansion of ICAD operations from
Kamiali into adjacent territories is certain to aggravate the difficulties inher-
ent in management of customary lands. Those same problems previously led
to the polarization of the original Lasanga-Lake Trist conservation plan. The
successful resolution of conflicts arising from parochial motivations will de-
termine the effectiveness of the enlarged VDT conservation area at Kui-Siboma
and of possible future expansion along the coastal corridor to Oro (Northern)
Province. Whether this can actually be achieved within the context of the ICAD
philosophy is very much an ongoing issue of considerable future significance
to conservation in PNG
The Physical Environment
Climate.-The KWMA lies mostly within Morobe’s lowland and premontane
climatic zones, where the mean annual rainfall is 3,000-4,000 mim. The wet-
test months generally occur during January-April, when prevailing winds are
northwesterly, and the driest in May-August when the winds are replaced by
southeasterly trades (McAlpine et al. 1983). Even during the dry season, mean
monthly rainfalls are generally around 200 mm, and the vegetation is rarely
subjected to water shortages under normal circumstances (ibid: 140). Most cli-
matic classifications would describe the project sites as tropical everwet or
perhumid, reflecting the overall absence of moisture deficits.
In the western Pacific, El Nino oscillations are typically accompanied by
severe droughts in rainforest environments like the Bowutu Range. During the
1997-98 disturbance, coastal tracts adjacent to the KWMA were subjected to
1498 BRIT.ORG/SIDA 20(4)
devastating fires which destroyed substantial areas of forest (Fig. 6). Formerly
covered with primary growth, Lasanga burned out of control during the 1997-
98 drought and is now a weedy island. In marked contrast to the El Nino epi-
sodes, the periods of peak rainfall are often accompanied by severe flooding in
the lowlands and numerous landslides on higher slopes. In 1999 the La Nina
rains resulted in torrential surges along the main rivers emptying into Nasau
and Sachsen Bays, completely destroying the subsistence gardens on the Bitoi
Delta. As the KWMA rivers overflowed their banks, the flood waters flattened
wide channels through the surrounding alluvial forest. The botanical team sub-
sequently enjoyed unprecedented access and easy collecting along the corri-
dors of felled canopy, which years later, are now becoming clogged with im-
penetrable thickets of Calamus.
Geology.—Much of New Guinea's diversity has been shaped by a complex
and dynamic geological past. Although the southern part of the island is an
extension of the Australian craton, the northern districts are a melange of former
island arcs which rafted onto the mainland during the northward migration
of the Australian landmass (Dow 1977. Jaques & Robinson 1975; Pigram &
Davies 1987). The Papuan Peninsula, consisting of the SE-trending ‘tail’ of New
Guinea, represents the remnants of an accreted arc (Hamilton 1979).
The Bowutu Range forms a major portion of the Papuan Peninsula and is
also PNG’s most extensive elevational series of ultramafic landscapes (Dow &
Davies 1964). This general region is part of a terrane complex paleohistorically
distinct from the rest of northern New Guinea, having docked with the main-
land ca. 10 m.y. after the Sepik terranes (Pigram & Davies 1987). The area’s un-
usual geology has led to its recognition as a separate physiographic province,
extending from the Kamiali coastline to the craggy summits of the Owen Stan-
ley Range (ie. the Papuan Ultrabasic Belt, cf. Bain 1973; Loffler 1977; Thompson
S Fisher 1965). Between Lokanu-Tambu Bay and Kui village, the ultrabasic belt
forms a continuous series of lowland environments. The offshore islands such
as Lasanga and Musik are part of a different (nonserpentine) geological series
reaching southwards to the Waria River (Dow & Davies 1964) and are excluded
from consideration in this paper
Soils—Ultrabasic environments are characterized by substrates low in
silica, but with high magnesium and iron contents. In addition to deleterious
magnesium/calcium ratios, ultrabasic soils are frequently associated with phy-
totoxic rations of nickel, chromium, and cobalt (Brooks 1987; Whitmore
1975). The influence of heavy metals is often cited as being responsible for the
stunted vegetation of serpentine habitats, though it should be noted that low-
statured canopies are not an invariable feature of ultramafic forests (see Fox &
Tan 1971; Proctor et al. 1988).
Bowutu populations from at least one species [Rinorea bengalensis (Wall.)
TAKEUCHI, ULTRABASIC FLORA IN PAPUA NEW GUINEA 1499
Fic. 6. Bulili Ridge. Forest burn from the 1997-98 El Nifio drought.
OK], are known to accumulate nickel (Brooks 1987). There are no reports of
heavy metal accumulation in other native plants. A brief discussion of serpen-
tine floras in New Guinea is provided by Brooks (ibid.), Paijmans (1976), and
van Royen (1963), but there is a conspicuous absence of floristic data in such
accounts. A summary of ultrabasic vegetation is otherwise available for the
tropical far east (Proctor 1992). While it itis generally becogmices that ultrabasics
are associated withd ily discerned in aerial photographs
(Paijmans 1976), the species composition of the communities remains poorly
known for Papuasia. Comparative studies from Western Malesia show wide
variation in the structure and endemism of such floras (Proctor et al. 2000).
Ultrabasic terrain in PNG is characteristically composed of massive ridges
with uniform slopes and unstable soils (Loffler 1977; Paijmans 1976). The topo-
graphic instability of the study area is reflected in the presence of old landslip
scars on many mountainsides (Figs. 7-8). Earthquakes with shallow epicenters
are common in northern New Guinea (Hamilton 1979), and can aggravate lo-
cal tendencies for landsliding. Quake-triggered landslides have been respon-
sible for many fatalities in Morobe Province.
he KWMA hill habitat is unsuitable for agriculture (Bleeker 1975a, 1975b)
because of erosion hazards and the general infertility associated with serpen-
1500 BRIT.ORG/SIDA 20(4)
tine substrates. In contrast, the flood plain of the major streams (Bitoi, Tabali,
Arawiri, and Saia),* have deep deposits of organic alluvium (Fig. 9). The Bitoi
Delta between the Areta (N) and Daunawa (S) branches isa particularly fertile
tract in which nearly all the Kamiali food gardens are concentrated. Unlike
many Highlands areas where intensified cropping has led to lowered
productivities (see Levett & Bala 1995), the KWMA agricultural system has sur-
plus capacity capable of supporting twice the present population (Bein et al. in
press, in submission). The high productivities are due to the presence of rich
delta soils and their renewal by seasonal floods (ibid.).? These factors have en-
abled the adoption of a shortened 7-year garden cycle (vs.an estimated 20 years
in normal slash and burn rotations). At current rates of population increase,
the existing subsistence system can probably satisfy local needs for 35 more
years without any expansion in area (Wagner 2001b).
The Social Environment
The total population of Kamiali was 520 in August 1997. There are two major
On contemporary maps the Arawiri (Ariwiri) is shown as the Alealer River, and the Saia as the Sela River (Roya
Australian Survey Corps Series T601, Nasau sheet 8383, Edition 1-AAS, 1:100,000)
Martin (1998) specifies a Bitoi crop rotation of 3-4 years and suggests the standard rotation period at nearby
Kui village is 10-15 years. Levett and Bala (1995) give 7—20 years as the standard swidden cycle for PNG
TAKEUCHI, ULTRABASIC FLORA IN PAPUA NEW GUINEA 1501
nee eet
ae
Fic.8.As for figure 7, showing tt ional ity. TI | includ inly Di is [i is. Machaerina
ic
]
glomerata, M rubiginosa Myrtella beccarii
1502 BRIT.ORG/SIDA 20(4)
Fic. 9. Saia River. S | fist ies t t di din the cl flowi ftho KWMA
clans, the Gara and Tabali, and at least 12 subclans (Martin 1998, 1999). Kela is
the indigenous tongue, one of approximately 200 comprising the Austronesian
language group of northeastern New Guinea (McElhanon 1984, Wurm 1985).
Nearly all villagers also speak the lingua franca of PNG (ie. Tok Pisin). With
the recent establishment of a community school, most children have acquired
at least some understanding of English.*
Seemingly unstructured to western observers, local village life is actually
governed by an extensive network of customary practices and expectations.
Inheritance and property rights are matrilineal by tradition, but considerable
change is occurring asa result of exposure to modern influences. Martin (1998,
1999, 2001) and Wagner (2001b) provide an informative discussion of social
developments in their specific connection to conservation.
Despite their retention o
customary lifestyles, Kamiali inhabitants are
surprisingly sophisticated and informed, a direct result of the quality-of-life
improvements introduced by the Village Development Trust within the past 10
years. Several community leaders for example, are computer literate and main-
tain international email contacts from VDT’s offices in Lae. As another sign of
‘There are 6 principal Kela-speaking villages on the Huon coast, divided into two apparent dialect groups. Kamiali
has its closest linguistic and kinship affinities to the hamlets of Buso and Kui immediately to the southeast, the
latter centers having been established by migration of Kamiali settlers during a former time of epidemic (Martin
1998)
TAKEUCHI, ULTRABASIC FLORA IN PAPUA NEW GUINEA 1503
cultural change, researchers are required to establish a Memorandum of Un-
derstanding with village-appointed representatives, disclosing the purpose of
an intended study and affirming its obligations to the Kamiali people.
The KWMA villagers are dependent on marine resources for their liveli-
hood (mainly reef fishing) supplemented by a limited range of garden crops.
Cash incomes are small and sporadic, on a household basis typically falling
between PGK 500-4,000 per year (at current rates | PGK=0.25 USD), mostly
from the sale of fish, sea shells, and betel nuts (Martin 1998; Wagner 2001b).
Although the Kamiali territory consists predominantly of primary forest rich
in animal life, only three individuals from the village population (Enok Nasa,
Tani Jena, Tom Ziena) are known to actively hunt wild game such as cuscus,
pigs, cassowaries, and wildfowl. For a people with copious forest resources, re-
markably little use is made of the terrestrial environment. Even among the small
hunting fraternity, forest forays are apparently undertaken mainly for recre-
ation, and overnight trips are rarely attempted. To a certain extent, the appar-
ent avoidence of inland habitats may be due toa belief in animalism and spirits
(masalai), even though all villagers profess devotion to the Christian (Lutheran)
faith. For whatever reason, there is essentially no human presence in the inte-
rior areas above 500 m. Since subsistence farming is confined to the flood plain
on the Bitoi Delta (a tract of ca. 460 hectares), human impacts on the environ-
ment are practically nonexistent (Martin 1998; Wagner 2001b).
The Logistical Environment
In geographic regions with the environmental qualities of New Guinea, inves-
tigators have little difficulty identifying biologically suitable localities for bo-
tanical research. During the selection of potential study sites, the logistical limi-
tations imposed by poor civil infrastructures often take precedence over the
biological concerns. When viewed in this context, the KWMA offers a particu-
larly attractive combination of site attributes.
ition to exceptional floristic quality, the coastal location of the
KWMA and its proximity to Lae (the second largest city in the country) pro-
vide researchers with unusually cost-effective logistics. There is no other con-
servation locality in PNG offering equivalent accessibility and scientific value
in one package. While many natural areas are of similar interest, their remote-
ness and associated travel costs discourage longterm study. Although the
KWM4A has no roads or airstrips, the site can be conveniently reached by out-
board-equipped dinghies (‘speedboats’).’
On the shores of Nasau Bay the Village Development Trust maintains a
permanent training center, guest houses, on-site staff, wireless communications,
and sea transport. Primary forest is immediately adjacent to the accommodations,
‘Travel time is 2.5 hours from Lae by 40 hp outboard and 1.5 hours by 70 hp. Charter rates for 40 and 70 hp
speedboats are PGK 600 and PGI p ively (rOur \dtrip Lae-Kamiali).
1504 BRIT.ORG/SIDA 20(4)
allowing easy access to serpentine vegetation. With electric generators lor the
facilities available on continuous standby, researchers are able to live and work
under conditions conducive to high morale anc
on
productivity. Of particular
importance to scientists using customary lands, access and intellectual prop-
erty issues have been successfully negotiated via VDF brokered agreements with
local landowners. In recognition of these infrastructural advantages, the KWMA
was selected as the principal PNG site for elevation-sequenced studies in the
Pacific-Asia Biodiversity Transect network (PABITRA; Takeuchi 2003a, digital
images on http://wwwsenckenberg.uni-frankfurt.de/odes/).
In conformity with the ICAD concept, ecotourism is actively encouraged
as an environment-friendly enterprise within the conservation zone. Kamiali’s
scenic shoreline of white-sand beaches is currently a featured scientific attrac-
tion, with nearly 5 km of waterfront serving as active nesting sites for the en-
dangered leatherback turtle. Substantial numbers of science-oriented visitors
arrive during the haul-out season in November-March to observe the turtle tag-
ging operations (Kisokau & Dutton 2002; Lindgren 1999a). In marked contrast
to the spartan conditions at most research venues, the K WMA's ambience and
infrastructure are comparable to a vacation resort.
—
a"
Kamiali’s checkered history forms a colorful backdrop to these present-day
assets. During WWII Nasau Bay was the site of a major battle between Allied
and Japanese forces (see Lindgren 1999b, 2000, for a popular account),and many
war artelacts, including aircraft and ordnance, are still present in the area (Figs.
LO-11). Local guide services are available at modest cost to tourists and other
Visitors.
The Biotic Environment
Although Paijmans (1976) believed that no native plants are restricted to ultra-
basics, there is mounting evidence that such substrates are in fact associated
with substantial endemism. In addition to the presence of localized endemics,
serpentine environments are also noted for distributional anomalies, particu-
larly by taxa occurring at elevations lar below their usual centers. This pattern
isespecially pronounced on the Bowutu ultrabasics, where the Massenerhebung
effect of coastal mountains (see Grubb 1971; Grubb & Stevens 1985; Whitmore
1975) combines synergistically with substrate effects. Many low-elevational
records can be expected from such situations, since the KWMA is PNG’ only
conservation locality where serpentine communities are disposed ina continu-
ous sequence from sealevel to cloudy summits.
Nearly all information from Bowutu environments are from easily-studied
coastal localities. The montane and inland areas have been neglected despite
their presumed biotic significance. In recognition of these deficiencies, the
Bowutu Range is included among PNG’s 16 terrestrial unknown areas and as
one of the five most critical watersheds requiring conservation action (Sekhran
TAKEUCHI, ULTRABASIC FLORA IN PAPUA NEW GUINEA 1505
inl I 4 JIC cnn ik CD
Fic. 10. p I
At n fehd tee soe ra
used In neat y 9,4} t t L | 7
L ee oe 1 + n t . | (Nn
Curh TT g DNG d | 1 ] he KWMA" g ( Sept 19, 1996), but
Pa hihte: . | £ 4 DI uv TWienl
Eo ee ee |
& Miller 1995). Johns (1993) had listed the district with 42 other localities of
greatest floristic importance for PNG. The estimate of 4,000+ plant species for
the area is among the highest of any existing conservation site (ibid.).
In 1997 a multidisciplinary survey was concluded within the KWMA, lo-
cusing primarily on marine and land-fauna documentation (Bein et al. 1998:
Bein 1999). The botanical inventory started in 1998 immediately after the ani-
mal surveys had ended. Taken collectively, the biotic assessments have exam-
ined coastal and alluvial swamps, lowland rainforest, premontane foothill for-
est, and mossy cloud forest. The collections density (CD: 890 plant specimens
per 100 sq km) is very high by Papuasian standards, but this figure is mislead-
ing because much of the collecting has been confined to the lowlands.° The
montane communities are still poorly known, even though most of the floris-
tic diversity is probably concentrated at the higher elevations. Approximately
20 new plant species have been discovered during the ongoing investigations
(Huynh 2001, 2002; Pipoly & Takeuchi in submission: Stevens in submission;
The overall CD for Papuasia is only 30 collections per 100 sq km (Frodin 1990).The atypical nature of the Kamiali
sampling coverage is evidenced by the near-absence of collections from above 500 m elevation.In other PNG
environments, the montane areas are usually much better known than the lowland habitats.
1506 BRIT.ORG/SIDA 20(4)
fa Pty Pe lek bolic
Fic. 11. Rusting 81 mm mortar shells from the KWMA. In 1943 Nasau
32 Bde (Burns 1991). Photo Karol Kisokau.
Takeuchi 2001, 2003b). The findings obtained to date provide evidence of a
continuing potential for discovery in one of PNG’s most distinctive floristic
Wehh VIEVJLILIIC LIS.
MATERIALS AND METHODS
In contrast to Rapid Biological Assessments, the botanical survey was based on
a plan involving numerous visits to the KWMA. Fieldtrips of 1-2 weeks dura-
tion were conducted at ca. 4 month intervals during the 4-year survey. Wit
a
1a
schedule of repeated visits, fertile specimens were acquired throughout the year,
and even infrequently-flowering taxa could be collected. Because of the KWMA's
logistical advantages, studies employing such itineraries can achieve a more
complete and cost-effective coverage than efforts based on single visits. The
benefits of conducting small-area inventories over extended periods of time
have been discussed by earlier commentators (e.g, Prance 1977).
Forest communities were examined by walk-through collecting and visual
assessment, operating either from campsites established in the bush, or from
the KWMA science facilities at Lababia Ridge and Kulindi (Pigs. [2=|4). The
explorations were generally confined to the lowland zone below 500 m. Plant
collections were usually made in sets of 3-5 duplicates, but if a specimen was
believed to have special significance, this was increased to ca. 10 duplicates. At
TAKEUCHI, ULTRABASIC FLORA IN PAPUA NEW GUINEA 1507
the Kulindi Science Center, electric generators enabled gatherings to be dried
by artificial heat immediately after collecting in adjacent forests. Otherwise
when operating from bushcamps, the — were field-pressed in news-
print and plastic bags, then soaked with 70% ethanol for eae: pe ess-
ing in Lae (Fig. 15). Silica-dried samples for DNA sequencing were als
if specialists had placed earlier requests lor assistance.
In Appendix 1, the survey specimens have been tabulated together with
earlier numbers from the ultrabasics at Buso and Kui.’ The ultrabasic boundary
extends southeast from Kui village, passing immediately inland from Braunshweig
Harbor at Siboma (Simpoma). Specimens from Kui are appended with a (K) on
the checklist, to indicate when the material is from the ultrabasic boundary
and may have been obtained from non-serpentine terrain. Natter Bay and Paiewa
represent Tertiary substrates of different age and origin from the Bowutu ultra-
basics (Dow & Davies 1964), and are thus omitted from the compilations.
The Paiewa drainage is intersected by an extensive network of logging
tracks along which a few collections were made under the NGF series (e.g.
Gillison NGF 22489-22500, 25011-25049). Some of the upstream areas at Paiewa
are inside the ultrabasic belt, but the locality data are too ambiguous to deter-
mine the substrate status of such collections. Jacobs’s numbers from ‘opposite
“Author citations for binomials are provided in the text for names which are not in the appendix.
)
BRIT.ORG/SIDA 20(4
1508
Fic. 13. Bulili Ridge. Survey team in ultrabasic forest.
TAKEUCHI, ULTRABASIC FLORA IN PAPUA NEW GUINEA 1509
Fic. 14 TLAY ‘lise, cr fa. tite! Vilind: hak we fh Dp
y. Natural-growth ultrabasic forest
J
eral area.
Lasanga Island’, are similarly difficult to interpret with respect to their sub-
strate association.
Much of the methodology and rationale for the Kamiali surveys have been
discussed elsewhere (Takeuchi 2003a). As with many recent efforts, the itiner-
ary was implemented by village teams involving institutional parataxonomists
D. Ama, Nathan, and/or A. Towati, or as a larger workforce supervised by col-
league B. Siga and the writer.
GENERAL DESCRIPTION OF THE VEGETATION
The physiognomy of the KWMA lowland flora is similar to a montane forest,
with a preponderance of small-leaved species and a general absence of vining,
cauliflorous, or compound-leaved plants (Fig. 16). Plank-buttressed trees are rare.
To some extent, these characteristics are due to the anomalous presence of
higher-elevation species in lowland habitats. Many sealevel distributional
records were documented by the recent surveys (e.g. see discussion on
Astronidium morobiense).
The KWMaA regrowth phase (between sealevel to at least ca. 500 m elev.) is
dominated by Commersonia bartramia, Decaspermum bracteatum, Dicranopteris
1510 BRIT.ORG/SIDA 20(4)
Fic. 15. Processing of plant specimens in the field. Gatherings are individually wrapped in newsprint to protect the
td
TAKEUCHI, ULTRABASIC FLORA IN PAPUA NEW GUINEA 1511
*]
>
=]
Fic. 16 Tha DIK . ig . y
linearis, Myrtella beccarii, Schuurmansia henningsii, and Timonius paiawensis.
Many of the seral or pioneer taxa normally found in Mamose environments are
absent or uncommon.® Cyathea, Cypholophus, Dendrocnide, enlarge
Glochidion, Gouania, Leea, Macaranga except for M. bifoveata), Mal lotus, Musa,
Mussaenda, Omalanthus, Pipturus, and Trichospet ly miss-
ing or are insignificant in the floristic succession on ultrabasics (Fig. 17). Even
the aggressive weed Piper aduncum rarely manages to establish in seemingly
opportune situations. Similar patterns have been noted in the Raja Ampat ser-
pentine district of Irian Jaya (Takeuchi 2003b).
Within the lowland ultrabasic flora as a whole, a number of ordinarily
speciose families are clearly under-represented. The impoverished groups in-
clude Araceae, Begoniaceae, Combretaceae, Dilleniaceae, Gesneriaceae,
Malvaceae (sens. str.), Marantaceae, Moraceae, Urticaceae, Verbenaceae, and
Vitaceae. The representation by ferns and their allies is also poor in compari-
son to regular substrates at comparable levels of rainfa
8Mamose is the administrati ion in northern PNG istin Madang, Morobe, and the Sepik provinces.
The name is an acronym formed » the first two letters aie ees and is also rendered as'Momase’ by the
same f Although ‘Mamose' is universally understood within PNG, its use is virtually unknown outsid
the country. There is a certain sly in its usage, because the eal refers » coincidence to a cae of PNG
defined by tectonic and biotic similarities, and thus represe y environmen
1512 BRIT.ORG/SIDA 20(4)
TAKEUCHI, ULTRABASIC FLORA IN PAPUA NEW GUINEA 1513
In addition to the impoverished groups, floristic depauperation is further
evidenced by the absence of several families which should be present in low-
land environments of northern PNG. The families which were not seen included
Commelinaceae (notably Amischotolype and Pollia), Cycadaceae,
Dichapetalaceae, Hernandiaceae, Opiliaceae, Ruscaceae (Dracaena angustifolia
Roxb.), and Tetramelaceae (Octomeles sumatrana Miq.). In contrast,
Euphorbiaceae, Myrtaceae, and Rubiaceae (except for Ophiorrhiza and
Mussaenda) are exceptionally common. Other families represented by large
numbers of individuals are Burseraceae, Clusiaceae, Cyperaceae, Meliaceae,
Myristicaceae, Myrsinaceae, and Pandanaceae. However there isan obvious gen-
eral reduction in floristic diversity, as evidenced by the low species counts rela-
tive to other New Guinea environments. The PABITRA transect investigations
are currently providing quantitative data for defining these patterns (Sengo in
prep.).
One direct consequence of low alpha diversity in KWMA habitats is the
disproportionate presence of a small group of plants. Certain taxa are nearly
everywhere in the lowland ultrabasic forest. Among understory shrubs and sub-
shrubs, these ‘weedy’ elements include Casearia aff. erythrocarpa (sp. 2nov.),
Conandrium polyanthum, Fittingia tubiflora, Geniostoma rupestre, Ixora sp. (sp.
A onchecklist), and Syzygium trivene sens. str. The understory fern Lindsaea
obtusa is the most ubiquitous pter idophyte. Anisoptera thurifera ssp. po lyandra,
Hopeaglabrifolia, Myristica chrysophylla, Syzygium furfuraceum, and Tristant-
opsis macrosperma are similarly abundant in tall-growth canopy. Gymnostoma
papuana (exposed ridges and riverbanks) and Stenocarpus moore i (streambanks
and landslips) often form locally monodominant stands in repetitively dis-
turbed habitats. The subcanopy and middle layers in hill forest below 500 m
are dominated particularly by Brackenridgea forbesii, Canarium spp. Garcinia
spp., Gordonia papuana, Gymnacranthera farquhariana var. zippeliana,
Haplolobus floribundus, Polyosma cf. forbesit, and Syzygium effusum sens. lat.
The KWMA lowland vegetation is thus characterized by a limited number
of very common species. The disharmonic nature of the flora is similar to insu-
lar environments from more isolated stations within the Pacific, and the pre-
vailing patterns are reversed to some extent only in the coastal and alluvial
flats bordering the major waterways. In such locations, some of the missing
taxonomic elements reappear, presumably due to a reduction in soil toxicity.
Alluvial substrates in the KWMA have acidity values to pH 4, unlike the ultra-
basic substrates on ridges (Bein, pers. comm. March 2000). There are thus two
general floristic associations within the lowland KWMaA: 1) a macrophyllous,
liane-rich community characteristic of riverine/swampy environments, and
2) a highly disharmonic ultrabasic flora of contrasting generic composition
and with some of the attributes of a montane vegetation.
Although Alyxia acuminata sens. lat., Freycinetia spp., Hugonia jenkinsii,
1514 BRIT.ORG/SIDA 20(4)
and Psychotria olivacea are moderately common in ultrabasic forest, clim bing
taxa are generally notable by their absence or rarity. Apocynaceae
(Asclepiadoideae), Calamus, Convolvulaceae (with the exception of Erycibe),
Dichapetalum, Flagellaria, Gouania, Lomariopsidaceae, Mucuna, Stenochlaena,
Tecomanthe, Uncaria, and all Vitaceae, are often absent or rare even from open
streamcourses and other edge environments where such taxa are ordinarily
prevalent. Understories are also remarkably clear, and physical passage th rough
the forest is decidedly easier than with most lowland communities.
On the current forest mapping system for PNG (Hammermaster & Saun-
ders 1995a, b) the Bowutu ultrabasic vegetation is primarily classified asa small-
crowned lowland hill forest (code Hs, ibid.), reflecting the overall stunting of
the serpentine canopies. KWMaA ultrabasic stands are easily detected in aerial
photos because of their depauperate appearance, changing only at the transi-
tion to regular substrates, where the canopy develops larger, medium-sized
crowns (code Hm, e.g. at the Kui ecotone).
In contrast to ultrabasic communities, the alluvial habitats are primarily
seral environments, often Gymnostoma-dominant and subject to seasonal {lood-
ing. These riverine successional areas are generally restricted to the coastal
floodplains where the KWMA%S clear-flowing streams emerge onto the lowlands
(see Hammermaster & Saunders 1995b: Salamaua SB 55-15 overlay). In other
low-lying sections near the coast, like the beachfront opposite Lababia Island,
the presence of poorly drained flatland is associated with Metroxylon-
monodominant swamps (Wsw; ibid.). The coastline vegetation also includes
scattered seagrass shallows (Enhalus acoroides), Bruguiera-R hizophora man-
groves, and tidal estuarine forest (Fig. 18), but the areas encompassed by such
communities are below the resolution of existing vegetation maps.
DESCRIPTIONS OF NEW SPECIES
RUBIACEAE
Psychotria bulilimontis Takeuchi, sp. nov. (Figs. 19-21). Type: PAPUA NEW
GUINEA, MOROBE PROVINCE: Kamiali Wildlife Management Area, lower slopes of
Bulili Mt, multistoried ultrabasic forest, 07° 18.5’ S, 147° 07.5 E, 20 m, 6 Oct 2002 (1,
fr), W. Takeuchi & D. Ama 16428 (HOLOTYPE: LAE! isoTyPES: A. BISH. BO, BRIT,
CANB, k, L, MO, NSW. US).
4 Se eae] | et f “1 Lal
5 1.
} }
Species 5 2g pay rhachidi
inflorescentiarum teretibus statim distinguitur.
Understory shrub 3-4 m in height, erect, often polelike, sparingly or moder-
ately branched, outer bark thin, dark gray, smooth. Branchlets terete. slightly
compressed at the top, subapical diam. 2-3(-4) mm, spreading, weak, surfaces
green in the leaf-bearing intervals, furfuraceous at the stipular scars, otherwise
glabrous, internodes (1-)2-6 cm long. Leaves diverging in one plane, glabrous;
blades fleshy or subcoriaceous, without domatia or cystoliths, adaxially dark
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ULTRABASIC FLORA IN PAPUA NEW GUINEA
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TAKEUCHI
1516 BRIT.ORG/SIDA 20(4)
Fig. 19. Psychotria bulilimontis var. bulilimontis in the ultrabasic forest on Bulili Ridge. White arrow: stem base of the
type individual.
TAKEUCHI, ULTRABASIC UA NEW GUINEA 1517
Fic. 20. Psychotria bulili ti' bulilimontis. The ty thering (W. Takeuchi & D. Ama 16428).
1518 BRIT.ORG/SIDA 20(4)
a)
Fic. 21. Psychotria bulilimontis var. bulilimontis. Closer view of tk
(W. Takeuchi & D. Ama 16428).
TAKEUCHI, ULTRABASIC FLORA IN PAPUA NEW GUINEA 1519
green, abaxially mid-green, bifacially olivaceous after drying, linear-elliptic,
(7-)12-21 by (1.5-)3.5-6.5 cm, apex gradually acuminate, margins entire, base
cuneate, equal; venation pinnate, secondaries 7-12 pairs, (7-)10-22 mm apart,
at the lamina center diverging 55-70° from the midrib, arcuate,
brochidodromous, tertiary nervation areolate, all venation plane or weakly
raised on upper surfaces, prominulous beneath; petiole 8-20 by LO-2.0 mm,
adaxially channelled or flattened, rounded beneath; stipules valvate, linear-
acuminate, to 15 by 4 mm, brownish-green, caducous, adaxially furfuraceous
at the base, otherwise glabrous, apex bifid, the arms 3-4 mm long, filiform. In-
florescence axillary from the subapical and lower nodes, glabrous, lax,
paniculiform, ultimately cymose, the primary branches opposed or 3-verticil-
late, all axes spreading, light green, delicate (0.5-1.0 mm wide), peduncle (LO-
)50-85 mm long, primary branches to 25 mm long; peduncular bracts cadu-
branching points, acuminate, 1-3 mm long;
—
cous; rachis bracts persisting at the
floral bracts triangular, minute. Flowers (measurements from spirit-preserved
material) tetramerous, glabrous on exterior surfaces, 5.0 mm long, sessile; ca-
lyx infundibular, 2.7 by 1.8 mm, limb dentate, teeth ca. 0.2 mm long, erect at
anthesis; corolla sympetalous, valvate, white, tube 2mm long, pilose ina |mm
wide band at the mouth, lobes ovate, 2.0-2.5 by 1.8 mm, apex acute; stamens
alternipetalous, erect, glabrous, anthers exserted, oblongoid, ca. 0.8 mm long,
basifixed, introrse, filaments very short, attached at the sinus; gynoecium gla-
brous, ovary globular, recessed at the summit, style included, ca. 0.6-0.7 mm
long, stigma lobes ca.0.5 mm long, coherent. Fruits shiny orange-red when ripe,
juicy, subglobose, 5 mm diam. in vivo, crowned by the calycine residue; pyrenes
2, pale yellow-brown, 2-3-ridged on the back, inside farinaceous, no ruminations.
Distribution and ecology.—Psychotria bulilimontis is known only from low-
land environments within the Kamiali Wildlife Management Area. The nomi-
nate variety is restricted to the ultrabasics.
Etymology.—The epithet commemorates the type locality.
PARATYPE: PAPUA NEW GUINEA. Morobe Province: Kamiali Wildlife Management Area, lower slopes
of Bulili Mt, multistoried ultrabasic forest, 07’ 18.5'S, 147° 07.5'E, 20 m, 25 Aug 2002 (f1, fr), W. Takeuchi,
D. Ama, B. Siga, & Nathan 16416 (A, L, LAED.
Axilliflorous Psychotria are rare in New Guinea. Only P axilliflora Merr. &
Perry and P dipteropoda Laut. & K. Sch. are known to flower from subapical
nodes. The new species is easily distinguished from its axilliflorous congeners
by the globose fruits (fusiform and compressed in P.axilliflora) and by the deli-
cate, terete axes of the inflorescence (conspicuously flattened and broad in P.
dipteropoda).
Psychotria bulilimontis var. aestuarii Takeuchi, var. nov. (Figs. 22-23). Tyre: PAPUA
GUINEA. MoroBE PROVINCE: Kamiali Wildlife Management Area, banks of
the Saia River near Hessen Bay, alluvial-estuarine forest, 07° 21.6'S, 147° 07.3' E,
1520 BRIT.ORG/SIDA 20(4)
Fic. 22. Psychotria bulili j tuarii.A ted dupli f | theri g. The delicate, umbelliform
_— F
t istic (W. Takeuchi. B. Siga, & A. Towati 14993).
abd:
a a
pa
SRA
Fic. 23. Hessen Bay. Estuarine f habi f tt lation for Psvchotria bulili, ; aestuari
y i
TAKEUCHI, ULTRABASIC FLORA IN PAPUA NEW GUINEA 1521
5-10 m, 15 Jan 2001 (fl), W. Takeuchi, B. Siga, & A. Towati 14993 (HOLOTYPE: LAE!
ISOTYPES: A, BISH, BO, CANB, K, L, MO, US).
a var. bulilimonti laminis ano sine ad cea rotund: atis vel cordatis, indumento denso denique
} vise.
| isti
Understory shrub |-2(-4) m in ae erect, oan or moderately branched.
Branchlets obliquely ascending, weakly compressed near the summit, terete
below, subapical diam. 2-4 mm, hirtellous, hairs persisting, internodes (2-)4-
7(-11) cm long. Leaves distichous, spreading; lamina usually fleshy, adaxially
dark dull green, abaxially pale green or brownish-green, discolorous after dry-
ing, turning black or fuscous above and rufous-brown beneath, upper surfaces
glabrous, marked by linear cystoliths or not, lower surfaces hirtellous, hairs or-
ange- or reddish-brown, densest on midribs and veins, domatia absent; blades
variable, narrowly to broadly elliptic, ovate, or obovate, (9-)13-23 by (3-)6-11
cm, apex acuminate, margins entire, base subcordate, rounded, (or cuneate):
venation pinnate, secondaries 10-12 pairs, (7-)15-20(-28) mm apart, at the
lamina center diverging (35-)50-70' from the midrib, arcuate, more or less
brochidodromous but reticulating freely beyond the commissural loops, ter-
tiary nervation conspicuously and coarsely areolate, all venation plane or im-
pressed on upper surfaces, prominulous beneath; petiole 5-20 by 15-2.0 mm,
adaxially channelled or flattened, rounded beneath, indument like the
branchlets; stipules valvate, caducous, lanceolate-ovate, to 15 by 5 mm, apex
bifurcately divided into 3-4 mm long filiform caudae, exterior surfaces coarsely
hairy, inside furfuraceous-barbate along the base, otherwise glabrous. Inflores-
cence axillary from the subapical and lower nodes, umbelliform or paniculiform,
the ramifications verticillately branched for 1-3 orders, ultimately cymose, all
axes spreading, lax, hirtellous, delicate (0.5-1 mm wide), dull light green; pe-
duncle (10-)35-95 mm long, primary rays to 31 mm long, secondary rays to 16
mim long; peduncular bracts caducous, linear-acuminate, to 3.0 by 0.5 mm, ra-
chis bracts subpersisting at the branching points, acuminate, ca. 1 mm long;
bracteoles triangular, ca. 0.5 mm. Flowers (measurements from spirit-preserved
material) tetramerous, 5.0 mm long, sessile; calyx tubular, 2.0 by 15 mm, pu-
berulent, limb dentate, teeth reflexed at anthesis; corolla sympetalous, val vate,
white, outer surfaces glabrous, tube 1-2 mm long, mouth pilose, lobes oblong,
2.6-27 by 1.2-1.3 mm, apex obtuse or acute; stamens alternipetalous, erect, gla-
brous, anthers oblongoid, ca. 0.5 mm long, basifixed, introrse, filaments very
short, attached at the sinus; gynoecium glabrous, ovary recessed at the summit,
style exserted, ca. 4.5 mm long, stigma conspicuously bilobed. Fruits imma-
ture, globose.
Distribution and ecology.—Known only
Wildlife Management Area.
Etymology.—The varietal name reflects the estuarine forest habitat of the
——
‘rom coastal forests in the Kamiali
type population.
1522 BRIT.ORG/SIDA 20(4)
PARATYPES: PAPUA NEW GUINEA. Morobe Province: Kamiali cee Reesor Area, banks ol
the Saia in near Hessen Bay, alluvial-estuarine forest, 07° 21.6' 07.3’ E, 5-10 m, 15 Jan 2001
(HD, W. Takeuchi, B. Siga, & A. Towati 14990 (A, L. | Re me WwW. se ei Siga, & A. Towati 14995
(A, BO, K, L, LAE!, MO); ditto, W. Takeuchi, B. Siga, & A. Towati 15002 (A, BISH, BO, CANB, K, L, LAF!,
MO, US).
Variety destuarii is clearly referable to Psychotria bulilimontis. The lax and
delicate inflorescence from lowermost axils is very distinctive and character-
istic of the species. However unlike the nominate variety, the alluvial plants are
very hairy shrubs with broad leaves and a pronounced tendency for
umbelliform branching on the inflorescence. In contrast, the ultrabasic popu-
lations (var. bulilimontis) are essentially glabrous, with narrow leaves gradu-
ally tapered at both ends, and a more paniculiform inflorescence. Both variet-
ies have heterostylous flowers.
SALICACEAE
Trichadenia sasae Takeuchi, sp. nov. (Figs. 24-27). Typr: PAPUA NEW GUINEA.
MOROBE PROVINCE: Kamiali Wildlife Management Area, base of Bulili Ridge at
Kulindi, ultrabasic forest, 07° 18'S, 147° 08' E, 0-5 m, 4 Feb 2003 (f 1), W. Takeuchi,
D. Ama & B. Siga 16561 (HOLOTYPE: LAE}; isotypes: A, BO, CANB, K, L).
aliic congenerihie lami
Species haec a
tis usque ad 31.5 cm longisque 27.0 cm latis, dense
velutinis fructibus magnis 4 cm diametris eel Ve ‘ler recognoscitur
Canopy tree 25 m tall, unbuttressed, highly branched, outer bark gray, thick,
rough-textured, slash orange-brown, without exudate, sapwood pale yellow.
Branchlets terete, 12-15 mm in diam. below the leaf spray; indument
subpersisting, orange-brown velutinous, usually with a monolayer of erect sep-
tate hairs, sometimes also with an underlayer of crispate hairs; periderm dark
gray, longitudinally fissured; abscission scars conspicuous, patelliform, 4-10 by
7-12 mm. Leaves spirally congested in apical tufts, spreading; blades coriaceous,
bichromatic in vivo, adaxially dark green, abaxially yellowish-green, olivaceous
insicco, the mature laminae manifestly cordate, 22.5-3L.5 by 20.5-27.0 cm, apex
obtuse, (emarginate, or mucronate), margins entire or irregularly repand, depth
of basal sinus (2.0-)2.5-4.0(-5.5) cm; upper surfaces pilosulous on veins, gla-
brescent between, lower surfaces velutinous; venation plinerval-palmatiform
at the petiole, pinnate above the base, craspedromous, secondaries 5-8, (15-)30-
70(-85) mm apart, at the lamina center diverging (45-)50-55(-60)’ from the
midrib, straight (or arcuate), sometimes sparingly anastomosing before the
margin, tertiary nerves scalarifom, reticulum conspicuously and bifacially ar-
eolate, adaxially impressed, all venation prominent below; petioles strictly cy-
lindrical, 5-10 cm by 2.5-4.0 mm on apical leaves, progressively longer on lower
leaves (to 18 cm by 5.0 mm), swollen at both ends, geniculate, indument like the
branchlets; stipules caducous, subulate, 7-12 mm long, densely hairy. Flowers
unknown. Infructescence solitary from axils of attached leaves, densely
velutinous; peduncle ca. 10 mm long, rachis terete, 10-20 by 3-7 mm, unbranched;
TAKEUCHI, ULTRABASIC FLORA IN PAPUA NEW GUINEA 1523
Fic. 24. Trichadenia sasae. C f f thet individual (W. Takeuchi, D. Ama & 8. Siga 16561).
bracts linear-acuminate, 1.5-2.0 mm long, caducous; pedicels 1.5-3.0 by 5-6 mm,
articulated at the rachis. Fruits indehiscent, pendulous, globose, 40-43 by 38-
43 mm, bilobed or obscurely trigonous, (38-47 by 46-55 mm), epicarp pale gray-
ish-brown through all stages of maturation, lenticellate, laxly hirtellous (gla-
brous to naked eye), surface scrape bright green, no exudate, pericarp thin (ca. 1
mim), crustaceous, entire in vivo, usually collapsing and coarsely rugose after
drying; mesocarp at first straw-pale brown, later yellow and baccate, ripe pulp
fleshy, no odor; seeds 1-3, endosperm copious, white, turning pink after sec-
tioning.
Distribution and ecology.—Trichadenia sasae is known with certainty only
from the closed-canopy forest bordering Nasau Bay. All confirmed sightings
have occurred on the ultrabasics.
Etymology.—The new species is named after colleague Sasa Zibe-Kokino, a
professional forester and prominent conservation advocate, currently serving
as the Member of Parliament for Huon electorate.
PARATYPE: PAPUA NEW GUINEA. Morobe Province: Kamiali aioe een Area, base of Bulili
Ridge at Kulindi Science Center, ultrabasic forest, 07’ 18'S O8' E m, 25 Nov 2001 (fr), W.
Takeuchi, D. Ama & B. Siga 15589 (A, BO, CANB, L, LAE, MO, an
The genus Trichadenia was previously represented in the Malesian region only
by the widely distributed T. philippinensis Merr. A second species (T. zeylanica
BRIT.ORG/SIDA 20(4)
Fic. 25. As for figure 24, looking into the crown from below.
TAKEUCHI, ULTRABASIC FLORA IN PAPUA NEW GUINEA 1525
Nn L 11 LI £ tA thari
Fic. 26. Trichadenia sasae
D 4h A KA a detetaa k LI . an ee |
Fic. 27. Trichadenia sasae
1526 BRIT.ORG/SIDA 20(4)
Thwait.) is endemic to Ceylon (Sleumer 1954). The new Trichadenia is easily
recognized by its large cordate leaves, dense indument, and large fruits.
The field appearance of T. sasae is similar to the Sterculia ampla-
macrophylla complex and is thus almost impossible to identify froma distance.
The type tree was for several years misidentifiec
eu
as a Sterculia due to the gross
similarities in habit. There are apparently several individuals of this species
along the Bulili coast but few have been seen in fertile condition. The species is
not common within the KWMA.
NOTES ON OTHER TAXA
CONVOLVULACEAE
Erycibe spp.—Erycibe is generally an uncommon genus in PNG. When last
treated by Hoogland (1953a, b) and Oostroom (1953, 1955), most taxa were known
from only asmall number of specimens. The current species concepts are ill-
defined, in part due to the former scarcity of materials for study and by what
appears to have been an excessive splitting of the variation. As is generally true
of canopy lianes with brightly colored fruits and inconspicuous flowers, most
of the modern gatherings consist of fruiting material which are less useful than
[lowering ones. The genus is greatly in need of revision.
In contrast to most PNG localities, Erycibe is well represented within KWMA
alluvial habitats, but all existing specimens are in fruit and cannot be keyed out.
While many Morobe collections are apparently conspecific with the KWMA
species, none of the sheets has been reliably identified and the assigned names
seem to have been taken up by successive collectors in uncritical fashion. The
KWMA Erycibe are neither rare or new, but the difficulties involved in their iden-
tification are symptomatic of a greater problem with the genus as a whole.
CYPERACEAE
Cladium mariscus (L.) Pohl sens. lat—A cosmopolitan species, but rare in New
Guinea and not previously recorded for PNG (Hay 1984). LAE has no recent
accessions from New Guinea other than the survey voucher.
FABACEAE
Maniltoa sp. (Verdcourt sp. E)-The only Maniltoa recorded in the ultrabasic
lowlands is apparently an undescribed taxon closely related to Maniltoa
schefferi K. Sch. & Hollrung. On Verdcourt’s (1979) alphabet list of novelties,
the KWMA plants are referable to species E, distinguished by sessile ovaries
and fruits. Maniltoa E is very common within the hill forest and consistently
maintains its separate character. There are no transitional forms to M. schefferi.
The KWMA Maniltoa is apparently restricted to the serpentine zone and is ar-
guably best regarded as a subspecies. In other characters, species E is so similar
to M. schefferi that it would be inappropriate to establish a new species on just
one differentiating trait.
TAKEUCHI, ULTRABASIC FLORA IN PAPUA NEW GUINEA : 1527
LECYTHIDACEAE
Barringtonia spp.—At least five species of Barringtonia have been documented
on the ultrabasics (B. apiculata, B. asiatica, B. lumina, B. pinifolia, and B.
racemosa; see Jebb 1991). KWMA Barringtonia are partitioned by habitat into
two groups, with B. asiatica and B. racemosa occurring mainly along the coast,
and the remaining species ranging into the upper drainages and interior hill
forest. Barringtonia pinifolia is particularly common within alluvial commu-
nities near the base of the main range. In comparison, the distinctive B. lumina
was seen only a few times along the Saia River and is apparently rare.
Barringtonia is widely employed as a fish poison in lowland New Guinea
(ibid.; Payens 1967; Peekel 1984) but Kamiali villagers app tly use only Derris
trifoliata Crop dynamit’) for fishing. This situation is certainly not due toa lack
of availability, since Barringtonia spp. are present in large, easily-accessed popu-
lations around the main village and fishing camps. Some respondents claim that
Derris is more effective on the reefs than Barringtonia, but this assessment is not
universal. Although the use of plant poisons for fishing isa well-established prac-
tice in Kamiali society, it has been traditionally discouraged except in special
circumstances (Martin 1998, 1999). The growing frequency of reef bombing and
Derris ‘dynamiting’, are manifestations of the changes occurring within Kela
society, and of the declining influence of customary observances.
MALVACEAE
Sterculia sp. nov. aff. ampla-macrophylla group
Sterculia sp. ?nov, aff. shillinglawii Fv.M.
The KWMA has two possible novelties in Sterculia.
Tantra (1976: 75-6) regarded Jacobs 9639 from the Buso-Kui foothills as a
new species in the ampla-macrophylla-morobeensis group, but formal descrip-
tion was deferred pending acquisition of more complete material. No new col-
lections have been made since the initial gathering in 1973. A series of numbers
from the current survey also cannot be identified with Tantra’s conspectus
(ibid.), and may represent another undescribed species.
MELASTOMATACEAE
Astronidium morobiense Maxw. (Figs. 28-29).—The species was previously
known only from mossy fagaceous forest between 1350-1800 m near Wagau
(Maxwell & Veldkamp 1990). However recent determinations (ex S.S. Renner,
pers. comm. March 2003) indicate that the species is distributed southwards
along the Morobe coast as far as Natter Bay, with occurrences in both ultrabasic
and non-serpentine habitats.
In marked contrast to the environment specified in its protologue, the
KWMaA populations of A. morobiense are found primarily at low elevations. On
Bulili Ridge, the species is even found at sealevel, immediately behind the
highwater line and in forest occasionally subject to saltwater damage. Between
BRIT.ORG/SIDA 20(4)
1528
1i, KWMA.
wilt
RULING
TAKEUCHI, ULTRABASIC FLORA IN PAPUA NEW GUINEA 1529
Fic. 29. As for figure 28,
=]
(W. Takeuchi & D. Ama 16223).
1530 BRIT.ORG/SIDA 20(4)
Lokanu-Tambu Bay and Kui village (Fig. 5), the only remaining hill forest is on
the headlands bordering Sachsen and Hessen Bays (e.g. Cape Dinga and Cape
Roon). The communities in or near these areas often have taxa at their lowest
elevations of record (e.g. Astronidium morobiense, Bhesa archboldiana, Gnetum
costatum, Hunga papuana, Lophopetalum torricellense, Mackinlaya schlechteri,
Nastus schlechteri, Ormosia calavensis, Paphia, Psychotria archboldii, and
Syzygium richardsonianum),.
MYRSINACEAE
Fittingia urceolata Mez and EF. tubiflora Mez (Figs. 30-31).—These obviously
related species are sympatric and vegetatively indistinguishable. Fittingia
urceolata is noticeably more frequent on the ultrabasic substrates than its con-
gener, but the two often occur side by side outside the serpentine (e.g. the foot-
hills at Paiewa). The only distinction between the species is in the color of the
drupes (red for F tubiflora and white for F urceolata; Sleumer 1988).
Sleumer (ibid.) suggested the apparent difference in fruit color might not
be an adequate basis for discrimination between the species, although he ac-
cepted the separation in his revision. Field observations from the recent sur-
veys show that the color contrasts are indeed consistent and effective discrimi-
nators. In F tubiflora the red color is evident even in the ovary of the anthetic
flower (label data for 14429) and is preserved as the fruit develops through a
marble-like phase and eventually into the spongy-textured drupe from which
the generic name is derived. Whether or not a species pair should be upheld on
the basis of a single contrasting character is a matter of subjective judgment,
but the color distinction is certainly not some transient feature related to fruit
maturation—ie. white fruits do not ripen into red fruits or vice versa. It may be
more appropriate to relegate the taxa to varietal rank, but in any event, there is
a readily detectable difference in living material?
tu
MYRTACEAE
Syzygium trivene (Ridley) Merr & Perry and S. lorentzianum |aut—Syzygium
lorentzianum was known to Hartley and Perry (1973) only from the type. The
species is supposedly distinguished from S. trivene by slight differences in leaf
form and by the longer igs buds (ibid.: 177).
Within the KWMA, S. trivene sens. str. is one of the most common
subarborescent taxa of riverine understories and interior forest. From exami-
nation of the survey’s many numbers, it is evident that the characters separating
this species from S. lorentzianum are part of an intergrading series of variation.
In general, S. lorentzianum is the name of a robust form growing in exposed
areas (e.g. along streambanks) while S. trivene is the spare variant of closed
“Most of the survey duplicates were distributed indicating a synonymy between the names (i.e. as F tubiflora =
F. urceolata.
TAKEUCHI, ULTRABASIC FLORA IN PAPUA NEW GUINEA 1531
Fic. 30. Fittingi lata. Tt i ; paquely pening
(W. Takeuchi, A. Towati, B. Siga, & M. Kavua 16172).
BRIT.ORG/SIDA 20(4)
|
og,
ne eae,
aes, gl igs
£
|
‘
Fic, 31. Fittingia tubiflora. \n contrast to the congener, fruits are red from the start of their development (there is no
green phase). The initially hard pericarp becomes spongy-textured and somewhat juicy when ripe (W. Takeuchi, B. Siga,
& A. Towati 14997).
TAKEUCHI, ULTRABASIC FLORA IN PAPUA NEW GUINEA 1533
1]
forest. The distincti determined and thus undeserv-
ing of formal recognition. Syzygium trivene should be regarded as a synonym
of S. lorentzianum, since the latter has the older epithet.
POACEAE
Nastus schlechteri (Pilger) Holtt., or aff -The condensed panicles and aristate
basal glumes are distinctive. Apparently known only from the type (Schlechter
19720), which was not seen by Holttum (1967) and presumably lost at Berlin.
The KWMA collections are from atypically low elevations for the genus,
having been obtained from ca. 100 m, but the vouchers otherwise conform to
Holttum’s species description and the key Gbid.).
RUBIACEAE
Psychotria archboldii Sohmer, P. mayana Takeuchi, P. melanocarpa Merr. &
Perry—Indument and fruit color are the principal means for separating these
taxa. Psychotria archboldii is entirely glabrous on all parts, while P mayana
and P. melanocarpa have stelliform hairs. Although P. archboldii has red fruits,
the drupes are white in P mayana and black in P melanocarpa. The three spe-
cies are apparently geographically separated. Only P.archboldii var. archboldii
has an appreciable distribution, occurring from Morobe to Milne Bay and off-
shore islands (var. multinervia is known by a single collection from Gulf Prov-
ince), while P mayana and P. melanocarpa have been found only in Madang and
Western Provinces, respectively. Apart from these distinctions, the plants are
obviously related and can be plausibly regarded either as sibling species, or as
geographically separate subspecies. Herbarium specimens from the archboldii-
mayana-melanocarpa complex are visually striking, with bright orange-brown
or rufous leaves. The large fruits are invariably jet black on exsiccatae, irrespec-
tive of the coloration in vivo.
In P archboldii asa whole, there is considerable doubt about the fruit color,
a character widely employed by Sohmer (1988) as a basis for species separa-
tions among the Papuasian congeners. Although variety archboldii is recorded
as having red fruits, the KWMA populations of this variety were collected with
white drupes. The fruits of P archboldii var. multinervia are also supposedly
white, according to the label for the only existing specimen (though the variety
was formally described as having red fruits)
The discrepancies could be explained if P. archboldii has distinct color
phases with the fruit maturing from white to red, or less plausibly from red to
white. However such color changes have never been substantiated for any
Papuasian Psychotria. If fruit color has no significance in this species complex,
the present separations should be dismantled and recognition given only to
geographic subspecies or races.
~
Psychotria croftiana Sohmer—Ps ychotria croftiana isa large-leaved species pre-
1534 BRIT.ORG/SIDA 20(4)
viously known from three fruiting collections on the Huon coast (Sohmer 1988).
It isa common and conspicuous shrub in the ultrabasic forest but also occurs
on the nonserpentine substrates southeast of Buso. The distinctively papillate-
hairy flowers were unknown until the recent surveys. A supplementary descrip-
tion of the flowering plant can be provided from the new material:
Inflorescence paniculate, pyramidal, exceeding the leaves at anthesis, to 37
by 29 cm, all axes glabrous; peduncle 7-13 cm long, first order branches 3-4-
verticillate, to 17 cm long; rachis bracts acuminate, 2-6 mm long, abruptly subu-
late at the apex, subpersisting, abaxially glabrous, adaxially furfuraceous at the
base; floral bracts inconspicuous, linear or linear-acuminate, ca. 0.5 mm long;
pedicels I-2 mm long; cymes lax. Flowers (measurements from rehydrated
material) 5(-6)-merous, 2.5 by 3.5 mm at anthesis; calyx infundibular (shallowly
cupular when dry), parted about halfway to the base, lobes triangular, equal,
ca. 0.5 mm long; corolla white, obovoid in bud, tube 1.7 by 1.8 mm, glabrous on
all exterior surfaces, mouth not clearly barbate, lobes adaxially papillate-hairy,
acute, 1.0 by 1.2mm, reflexed at maturity; stamens alternipetalous, anthers erect,
oblongoid, 0.6-0.7 mm long, filaments short, inserted about half way down the
tube; gynoecium glabrous, ovary dome-shaped, recessed at the top, style 0.7 mm
long, stigma 0.3 mm long, slightly expanded, columnar, 5-6-sulcate, not or only
weakly exserted.
Specimens examined: PAPUA NEW GUINEA. Morobe Province: Kamiali Wildlife Management Area,
lower slopes of Bulili Mt, ultrabasic forest, 07° 18.5'S, 147° 07.5' FE, 40 m, 6 Oct 2002 (f(D, W. Takeuchi &
D. Ama 16430B (A, BO, CANB, K, L, LAE!, MO, US): base of Lababia Ridge, ultrabasic forest, 07° 16'S,
147° 00° E,75 m, Oct 2002 (1D, W. Takeuchi & D. Ama 16578(A, LAF!).
Psychotria waiuensis Sohmer—Psychotria kamialii Takeuchi, Edinburgh J. Bot.
58:159-161. 2001. Type: PAPUA NEW GUINEA. MoroBE PROVINCE: Kamiali Wild-
lite Management Area, 1.5 km W of Lababia Village, 0717'S, 147° 06'E, 5-10 m
Nov 1999, W. Takeuchi & J. Sengo 14310 (HOLOTYPE: LAE; ISOTYPE: K).
Current fieldwork indicates that the recently described Psychotria kamialii
must unfortunately be regarded as a synonym of P. waiuensis. As the most ob-
vious point of distinction from its closest congeners, the monocaulous habit of
P. kamialit was the principal character used to establish that species. Because
of anemphasis on architectural form, the equivalence to P. waiuensis was over-
looked. There are several points which can be made in relation to the synonymy,
First, itis very unlikely that P waiuensisisa 10 feet (3 m) tall shrub as indi-
cated in the diagnosis. Psychotria waiuensis is not rare, and every plant seen
during the recent surveys were monocaulous dwarfs occurring as helophytes
in shallow mud or standing water In this respect the plant is similar to the higher
elevation forms of P ramadecumbens Sohmer. The type gathering for P kamialii
lor example, was a colony sample from sago swamp, taken from separate 0.5m
individuals. The KWMA populations have never been seen growing on firm
ground as medium-sized branching shrubs. Although P. waiuensis is described
—
TAKEUCHI, ULTRABASIC FLORA IN PAPUA NEW GUINEA 1535
as ‘about 3 m’ (Sohmer 1988) this figure is probably a label error. It is more plau-
sible that the first collections at Waiu Bay (Braunschweig Harbor on most maps)
were also monocaulous subshrubs, as suggested by the fact that the earlier sheets
show no indication of branching. Most of the bayfront at Waiu is a coastal
swamp similar to the type locality for P kamialii,soa general stunting of woody
plants would also be expected at the earlier collection site. Because the actual
habit of this swamp species was not appreciated, Sohmer's (1988) key will go
astray at fork 107, where there is a couplet separation between monocaulous
and branching shrubs. Psychotria waiuensis will actually key out to P
inconspicua Merr. & Perry, as happened during the initial evaluation of P.
kamialii.
As presently known, P. waiuensis occurs only between Kamiali and Waiu,
an area consisting mainly of ultrabasics. However future collectors will prob-
ably find this species further south along the coastline, since the swampy lor-
ests which the plant inhabits are not restricted to serpentine. Judging from its
current habitats, P. waiuensis may be present in the coastal wetlands near the
Morobe-Waria Rivers, where extensive areas of comparable environment can
be found.
Coastal swamps are such disagreeable places that it is easy to understand
why only one collection existed of P. waiuensis prior to the recent surveys, even
though the species is moderately common. The presence of saltwater croco-
diles (Crocodilus porosus) and swarms of mosquitos are disincentives to loiter-
ing in these shoreline areas. The most promising strategy for collectors is to work
the ecotone along the margins of the swamps, rather than their interior parts.
If the putative differences between P. waiuensisand P kamialiiare derived from
substrate conditions, different growth forms should be found at the contact
between swamp and dry land.
SALICACEAE
Homalium d’entrecasteauxense Craven—Formerly known only by the type col-
lection from Normanby Island (Craven 1979). In the survey vouchers, stamens
are consistently arranged in 3-membered fascicles. Branchlets are puberulent.
Steenis (1982) expressed doubt over the number of new species recognized
by Craven (1979), and of their distinction from the variable H. foetidum (Roxb.)
Benth. The taxonomic issues are still unresolved because of the limited num-
ber of specimens available for study (7 out of the 10 Papuasian species are known
only from the types). Multiple collections showing the variation within indi-
vidual populations are much needed.
The new gatherings from the KWMA suggest that staminal number is an
effective criterion for splitting H. d’entrecasteauxense from the foetidum com-
plex. This separation is corroborated in the field by differences in stature and
habit. The Kamiali specimens of H. d’entrecasteauxense were taken from frail,
1536 BRIT.ORG/SIDA 20(4)
4m tall understory shrubs, but H. foetidum is ordinarily a robust timber-sized
tree. The survey specimens also confirm the differences in indument between
H. dentrecasteauxense and H. maneauense Craven (see modified key in Steenis
1982: 564).
THELYPTERIDACEAE
Plesioneuron croftii Holtt—Formerly known in the literature only by the type
collection from Natter Bay, but now also represented by newer material from
the Waria basin and Buso.
Plesioneuron croftii is similar to P. dryas Holtt,, differing primarily in the
color of the aerophores (dark in P.dryas and pale in P croftii; see Holttum 1981).
The recent collections are closest to P croftii but have dark aerophores, and seem
to combine characteristics of that species and its congener. Although the posi-
tion of the sori near costules is of some value in the identification of P croftii
(ibid: 399), the distinction is small and inconstant, and probably unworthy of
specific rank.
=
DISCUSSION
—
Whether or not floristic patterns discerned in the KWMA are of general appli-
cation to other floras is unknown, especially as the ultrabasic patterns are dif-
ficult to separate from those applying to the Papuan Peninsula as a whole.
Although the distribution of ultrabasic substrates can be extracted from the
geological literature, information on the associated vegetation is sparse and
unsubstantiated. There are no checklists or forest descriptions of representa-
tive formations anywhere in PNG. The patterns found at Kamiali may be unique
to the Bowutu communities, but at least within the K WMA, the lowland ser-
pentine has been sampled to very high intensities, and is presently one of the
better known lowland floras in New Guinea. Approximately 1,915 specimens
have been compiled from a lowland area of 215 sq km, for a collections density
(CD) of 890 collections per 100 sq. km!° The current checklist includes 130
families, 412 genera, and ca. 710 morphospecies. Most of the lowland taxa from
the Kamiali area have probably been recorded. In view of the general depau-
peration of serpentine floras, the species counts primarily reflect the sampling
saturation achieved by the surveys, rather than any special richness of the ser-
pentine plant life. AtJosephstaal for example, an inventory of lowland rainforest
below 400 m recordeda total of 139 families, 445 genera, and 730 morphospecies
(Takeuchi 2000), an outcome very similar to the Bowutu study. However the
—
1989) notes that the current benchmark of 50-100 collections per 100 sq km does not necessarily
Stevens
represent a well-collected locality. The Bowutu inventory demonstrates that even at much higher sampling
c ct 1 f C
|
hlichon
cf
{ toh 1
1equ
intensities, unknown taxa will escape detection.Once ana or a specific
Adit
| if
flora, further progress will be dependent on h are distributed through space and time,
rather than the mere acquisition of more material.
TAKEUCHI, ULTRABASIC FLORA IN PAPUA NEW GUINEA 1537
Josephstaal assessment involved only one month of survey work, while the ul-
trabasic itinerary was much more intensive, consisting of repeated fieldtrips
over a period of four years.
The results at Kamiali have been constrained by the surveys’ strict and de-
liberate focus on the lowland zone below 500 m. On Mt Kinabalu, the speciose
character of the vegetation has been attributed primarily to the presence of
ultrabasic outcrops in the montane elevations (Beaman & Beaman 1990). Spe-
cies counts in the KWMA are likely to increase considerably when investiga-
tions are extended into the higher habitats where most of the diversity in New
Guinea environments are probably concentrated. A substantial number of
Bowutu taxa have already been typified from NGF collections made at Lake
Trist (elev. 1800 m), mainly by Henty in 1966 (NGF 29004-29138). Many of these
montane serpentine records are still known only from the original collection, a
situation similar to the Kinabalu ultrabasics (ibid.). Existing evidence suggests
that a fair number of species remain undiscovered. Unlike Mt Kinabalu how-
ever, the KWMA lowland ultrabasics have relatively few species, which for the
most part are already well-represented in collections from Buso, Kamiali, and
Kui. The recent high-intensity surveys have started to reveal the presence of
rarer endemics such as Discocalyx kaoyae, Paphia megaphylla, and Trichadenia
sasae. Mammea papyracea (NGF 39414), Pouteria gillisonii Vink (NGF 25627),
and P pullenii Vink (NGF 25625), known only from their types, are also prob-
ably rare plants from the ultrabasics (cf. Stevens 1995; Vink 2002).
Despite the extensive work done by earlier botanists at Buso and Kui, the
lowland ultrabasics are still a fruitful venue for new material. Even supposedly
well-collected localities can be a source of floristic discovery when explora-
tion routes are slightly altered from the previously-used tracks (Stevens 1989).
The recent discoveries are a case in point. Papuasian collectors often retrace the
field itineraries of previous investigators due to the convenience of following
well-used trails. Yet because plant distributions are typically very patchy across
seemingly homogeneous or comparable habitats (Oatham & Beehler 1998),
slight changes in exploration schedules can be immensely productive. In the
KWMaA, adjacent ridges often have very different compositions and species fre-
quencies even across the same geological substrates (Sengo in prep.).
Some idea of the conspicuous contrasts in plant distributions and frequen-
cies can be seen when comparing the collections from Buso and Kamiali. For
example, Kairothamnus phyllanthoides and Plesioneuron croftii are common
at the former locality, but have not yet been recorded from immediately adja-
cent Kamiali habitats. Conversely, although Psychotria bulilimontis and
Timonius sp. ?nov. are plentiful in the KWMA hill forest, they have yet to be
collected at Buso. Distributional peculiarities are further exemplified by plants
that are essentially restricted to the ultrabasics, having high frequencies inside
1538 BRIT.ORG/SIDA 20(4)
the KWMA, but which are also known from one or two collections on norma
substrates elsewhere on the Papuan Peninsula (e.g. Astronidium morobeense,
Guioa grandifoliola, Hunga papuana, and Zanthoxylum novoguineense).
Patchiness of the plant distributions on ultrabasics occurs over a range of
spatial scales. At one end, are the localized differences reported by Oatham and
Beehler (1998) at Lakekamu. These may be due to the dynamic fragmentation
—
of the forest caused by disturbances such as storms and changes in streamcourse.
Ona larger scale, plant distributions can vary markedly along different ridges
or rivers, even though there are no obvious habitat factors separating the occur-
rences (Frodin 1990; Grubb & Stevens 1985; Kalkman &@ Vink 1970). The differ-
ent levels of patchiness have obvious implications for floristic exploration. Per-
functory surveys will probably miss many of the most significant plants,
particularly those with quirky distributions. These difficulties are compounded
by unpredictable phenologies and the resulting element of chance in the re-
sults achieved by surveyors.
—_
Within the Bowutu tract, taxa endemic to the ultrabasic belt include
Calophyllum streimannii,!! Discocalyx kaoyae, Freycinetia curvata, E kamialiensis
F mediana, F rubripedata, Kairothamnus phyllanthoides, Myristica filipes,
Paphia megaphylla, Psychotria bulilimontis var. bulilimontis, and Solanum
symonianum. The percentage of strict endemics in the local flora is small com-
pared to more extreme environments (e.g. the Waigeo ultrabasics in Irian Jaya,
where rainfalls are much lower than in the KWMA,; cf. Takeuchi 2003b). Hu-
mid conditions appear to reduce the intensity of effects from serpentine sub-
strates. An inverse relationship between rainfall and the severity of the serpen-
tine syndrome has been noted for other areas (e.g. Kruckeberg 1985).
An overwhelming majority of the KWMA species occur on both sides of a
serpentine contact, although there are apparent differences in relative frequen-
cies when passing across such boundaries. The floristic contrasts are primarily
ones of degree rather than quality. With so many lowland areas being altered
by anthropogenic development, the natural distributions of many native spe-
cies will become difficult to determine in the future. This will complicate com-
parative studies on PNG’s ultrabasic ecosystems . Most of the coastal habitats
south of Kamiali are now fragmented by logging damage, and the real range of
plants comprising the Papuan Peninsular flora is increasingly obscured by for-
est destruction.
en
en)
In view of the general connection between serpentine effects and rainfall,
the growing incidence of El Nino-associated droughts is likely to result in future
floristic change which will amplify existing patterns. As rainfalls are reduced,
ultrabasic effects should increase, and floristic distinctions across the substrate
contact at Kui will become more acute. This is likely to be accompanied by a
"But C. streimannii may be present in Western Province (see Stevens 1995)
} \ L
TAKEUCHI, ULTRABASIC FLORA IN PAPUA NEW GUINEA 1539
sharper physiognomic discontinuity at the contact (i.e. a more obvious reduc-
tion in tree statures and crown densities across the boundary). Floristic attenu-
ation can be expected to involve mainly the generalist species that lack serpen-
tine-coping mechanisms. Since many of these pane (e.g. Canarium) are used
by Kamiali villagers, a reduction in forest ethnobotanical values is probable.
Future changes in beta-diversity patterns can also be anticipated in the mon-
tane parts of the ultrabasic belt. The KWMAs mossy forests above 500 m were
severely affected by the recent El Nino episodes of 1997-98 and 2002. Repeated
droughts could result in a lifting of the ecotone and an associated degradation
of watershed services.
APPENDIX 1. LOWLAND PLANTS RECORDED FROM KAMIALI
Voucher source for occurrence record: LAE = collections from the Lae Herbarium
institutional (new) series; NGF = collections from the New Guinea Force series;
nv = non visus, specimen not found at LAE but cited in the literature and/or
entered in LAE logbooks; sn = sin numéro (without number); SR = sight record;
numbers without prefix = W. Takeuchi et al. (usually with D. Ama, B. Siga,and/
or A. Towati). Other collectors indicated by name.
The checklist is a compilation of collections (below ca. 500 m) from t
ultrabasic zone between Kamiali and Kui. Specimens from the present survey
have been determined by WT unless otherwise noted. Determinations to ear-
lier collections are based on annotations from published revisions or from spe-
cialist tickets on herbarium sheets.
—
Me
FERNS AND FERN ALLIES
ADIANTACEAE Diplazium esculentum (Retz.) Sw.; 15251
Acrostichum speciosum Willd.; 14896 BLECHNACEAE
Adiantum hollandiae v.AN.R. 15228 eset entdele er
ore acalomelanos (L.) Link, 15062 Blechnum vittatum Brack; 14815, 214973, 15059,
norammya hornecncjc
g is (Hook.) J. Smith; 14468 15102
Syngramma grandis (Copel) Co Cini; RAE 52308 Stenochlaena milnei Underw.; SR, Tabali
. riverbanks
Taenitis blechnoides (Willd.) Sw.; 14642, 14659;
Kog 30; NGF 44261; Palis 27 CYATHEACEAE
Cyathea macgillivrayi (Bak.) sale 15036, 15356
ASPLENIACEAE ae taes Cyathea werneri Ros.; 1488
Asplenium laserpitiifolium Lam.; 14722, 14840, Cyathea sp, subsection ene 15205
14915, 14917
Asplenium cf. musifolium Mett.; 14705 DAVALLIACEAE
Asplenium nidus L. var. nidus; yeas Unkau 96 Davallia heterophylla Sm.; 14615B, 15390; NGF
Asplenium phyllitidus Don ssp. malesicum 24493
Holttum; NGF 45123, 45124; Palis 30 Davallia parvula Hook. & Grev.; 14391
Davallia pectinata Sm.; 14411, 14538
ATHYRIACEAE Davallia repens (L.f.) Kuhn; 15215
os id ca, ee Davallia solida (G. Forst.) Sw. var. solida; 14358,
mS irse hee 14539, 14686, 15225
Dipl
1540
DENNSTAEDTIACEAE
Hypolepis tenuifolia (Forst. f.) Bernh. ex Presl;
15067
idi iquilinum (L.) Kuhn; 15060
DRYOPTERIDACEAE
Dryopolysti ae UM eee (Cesati) Copel;
ea 14685, 14878; Manseima 19;
Palis |
GLEICHENIACEAE
Dicranopteris linearis (Burm. f.) Underw. var.
subferruginea (Hieron.) Nakai; 14609, 15052
GRAMMITIDACEAE
Ctenopteris blechnoides (Grev.) Wagner &
Grether; 14776, 14794, 15141, 15172, 15214;
Conn et al. 208; NGF 45189; Palis 31
HYMENOPHYLLACEAE
Hymenophyllum sp. (Mecodium); 15231
fea (Cephalomanes) atrovirens Kunze;
5) kia 17;Gawi 22; Ron 2
eee vanes (Selenodesmium) obscurum Bl;
14674, 15185; Damas 1; Palis 29; Rau 288
Trichomanes sp., Nesopteris-Selenodesmium
group; 16619
LINDSAEA GROUP
Lindsaea ensifolia Sw. ssp. agatii (Brack.) Kramer
8, 15336
Lindsaea ensifolia Sw. ssp. ensifolia; 15140
Lindsaea ensifolia Sw.x ?L. obtusa J. Smith; 14880
srs te eal (Gaud.) Desv.; 14326, 14668,
4789, 14842, 15352, 16433; Gawi 24
inna / Copel,; 15226; NGF 39415
Lindsaea obtusa J. Smith; 14319, 14390, 14407,
ne 14669, 14690, 151748, 15180, 15440;
Gawi 23; NGF 4512
Lindsaea repens (Bory) Thwaites var. sessilis
(Copel.) Kramer; 15186; NGF 394 ae
Sphenomeris retusa (Cav.) Maxon; 14
peed dium longipinnulum (Ces.) C. at 5196;
27;NGF 45126,45127
LYCOPODIACEAE
Huperzia cf. carinata (Poiret) Trevisan; 14716,
14932
,
Huperzia phlegmaria (L.) Rothm.;
521]
Huperzia squarrosa (Forst.f.) Trevisan; 14812
Palhinhaea cernua (L.) Vasc. & Franco; 14643,
15051
14710, 14843,
BRIT.ORG/SIDA 20(4)
MARATTIACEAE
Marattia sp., aff. ?megaptera Copel; 15155, 15158,
OLEANDRACEAE
Nephrolepis falcata (Cav.) C. Chr; 14621
Nephrolepis hirsutula (Forst.) Presl; 15232
OPHIOGLOSSAC
Ophioglossum ana um : f. pendulum; 14914;
51690; NGF 4522
PARKERIACEAE
Ceratopteris thalictroides (L.) Brongn.; 15253
POLYPODIACEAE
Drynaria rigidula Bedd,; 14536
Drynaria sparsisora ee Moore; 14657, 14950
Lecanopteris deparioides (Ces.) Baker; 15223,
6635; NGF 45219
sca sinuosa (Wall. ex Hook.) Copel.
14546, 15079; Bellamy B12; NGF 44206
Microsorum ne idria (Burm.f) een
Microsorum sibomense Copel.; NGF 45182
Pyrrosia ee (Alston) Morton var. em
1526
14360
PSILOTACEAE
Psilotum nudum (L.) Beauv.; 14637
PTERIDACEAE
Pteris blumeana Agardh; 14684, 14717
Pteris tripartita Sw.; 15061, 15073, 15077
SCHIZAEACEAE
lygodium microphyllum (Cav,) R. Br 1
god um pat Baker; 14406, oS ve
str.as L.dimorphum)
lygodium versteegii ene 14323, 15103, 15182;
Gawi 25; Kairo 752; NGF 44201
Schizaea dichotoma (L.) Sm.;1
Kog 27; Kwangut 33; NGF 4
itata (L.) Sw.; 1 fe 5006; NGF
ie 2 a 9545;
Schizaea dig
39417
Schizaea wagneri Selling; NGF 45170
SELAGINELLACEAE
Selaginella cf. schlechteri Hieron.; 14409
Selaginella cf. suffruticosa V.AN.R.; Palis 32
Selaginella sp., aff. ’melanisica Kuhn; 14809
Selaginella sp. oe latifolia, puberulipes, or
suffruticosa); 1
Selaginella sp.; Ss
TAKEUCHI, ULTRABASIC FLORA IN PAPUA NEW GUINEA
TECTARIA GROUP
Tectaria bamleriana (Rosenst.) C. Chr; SR, Bullili;
NGF 45220
Tectaria durvillei (Bory) Holttum; NGF 22884 (K,
nv)
Tectaria sp.,aff. ?cristovalensis (C.Chr.) Alston; NGF
22884 (K)
THELYPTERIDACEAE
Plesioneuron croftii Holttum; Conn et al. 177
Manseima 26 (723, two nos. given on label)
Pneumatopteris sogerensis (Gepp) Holttum
67
Sphaerostephanos multiauriculatus (Copel.)
Ho cei
ahaa SS D4
(Brause) Ros.; on 14961,
VITTARIACEAE
Antrophyum sp., callifolium Bl. facies; SR, Lababia
Monogramma sens. lat., closest to Vaginularia
trichoidea Fee; Kwangut 37
Vittaria angustifolia Bl; 15391; Unkau 98
Vittaria elongata Sw. var.angustifolia (Bory) Thw.;
15216
Vittaria elongata Sw. var.elongata; Manseima 20
Vittaria cf. scolopendrina (Bory) Thw.; 14775
aff. Sea
15010, 1
GYMNOSPERMS
GNETACEAE
Gnetum costatum K. Sch.; 14320, 14765, 14965,
15415, 15420, 15577, 16579, 16586; Conn et
al. 296; NGF 25677
Gnetum gnemon L. var. gnemon; 14739, 14741,
14764, 14783, 16562; LAE 68531
Gnetum latifolium Bl. var. latifolium; 14952
PODOCARPACEAE
Podocarpus neriifolius D. Don; 14330, 15086,
5110; NGF 28082A (K)
MONOCOTS
AMARYLLIDACEAE
Crinum asiaticum L.; SR, Tabali riverbanks
ARACERE
Colocasia esculenta (L.) Schott; SR, cult. Lababia
NGF
Cyrtosperma macrotum Becc. ex Engl.;
45172
Holochlamys beccarii Engl.; NGF 25681
Scindapsus sp.; 14638, 15080
1541
Spathiphyllum schlechteri (Engl. & Krause)
Nicolson; 15157, 16595; Conn et al. 205, 217,
237;LAE 51676;NGF 45 149:Vi Kairo 311,
312,314,317
Xanthosoma sagittifolia
Lababia
(L.) Schott; SR, cult.
ARECACEAE
Areca envied [eee SR, cult. Lababia
Cal
rungi cs SR, Sai
a River
Calamus sp., aff. brevifolius Becc.; 14956; NGF
24479 (nv)
Calamus spp.; 14426; Jacobs 9561, 9698; NGF
24469
Calyptrocalyx sp., aff. ?stenochista Burret; 14398
Calyptrocalyx sp.; Kjaer AB511
Caryota rumphiana Mart.; SR, Kulindi
Cocos nucifera L.; SR, cult. Lababia
Cyrtostachys glauca H.E. Moore; NGF 24460
Cyrtostachys sp.; Kjaer AB 512
Heterospathe cf. muelleriana (Becc.) Becc.; Kjaer
513
Hydriastele microspadix Becc.; 14547, 15043,
5142; LAE 52052; NGF 24427, 24477
Livistona sp.nov.;Kjaer AB 514; NGF 24466, 25682
Metroxylon sagu Rottb,; SR, Tabali River
Nypa fruticans Wurmb; SR, Tabali R
Orania lauterbachiana Becc.; NGF ee 24480
Orania sp.; Kjaer AB 515
BROMELIACEAE
Ananas comosus (L.) Merr.; SR, cult. Lababia
CYPERACEAE
Cladium mariscus (L.) Pohl sens. lat.; 14630
Cyperus pedunculatus (R. Br.) Kern; LAE 52301
Cyperus rotundus L. ssp. retzii (Nees) Ktk., 14974
Eleocharis geniculata (L.) R. & S.; 14533B; LAE
52324; NGF 44152, 44203
Fimbristylis cymosa R.Br; 14531, 14533A
Gahnia aspera (R. Br.) Spreng.; 14578, 15063
Hypoletrum nemorum (Vahl) aa ;LAE 52340
oe ina glomerata (Gaud.) Koyama; 14443,
0, 14805, 15337, 16569; " 72468; NGF
ise 8 (K), 39432
Machaerina mariscoides (Gaud.) Kern; 14850,
16571;LA )
Machaerina rubiginosa (Spreng.) Koyama; 14629
Mapania baccifera C.B. Clarke ssp. baccifera;
14573, 14658
Mapania macrocephala (Gaud.) K. Sch.; 14396,
14818
1542
Mapania macrocephala (Gaud.) K. Sch. ssp.
nacrocephala; 14791; Conn et al. 262; LAE
0
5229
Paramapania parvibractea (Clarke) Uittien;
640, 14656, 15083, 15563, 16612
Schoenus falcatus R. Br; 14624, 14762, 14849
Schoenus laevinux (KUk.) Ohwi; 14907
Scleria ciliaris Nees sens. lat. 15122, 1544]
Scleria polycarpa Boeck.; 14862
DIOSCOREACEAE
Dioscorea alata L.; SR, cult. Lababia
Dioscorea esculenta (Lour) ae SR, cult. Lababia
Dioscorea nummularia Lamk.; 14346, 15070
ery
FLAGELLARIACEAE
Flagellaria indica L.; 14331
HYDROCHARITACEAE
Enhalus acoroides (L.f.) Royle; Jacobs 9693; NGF
45190
LAXMANNIACEA
oe pet (L.) A. Chev.; 14457, 14732,
4826, 15044, 16439; NGF 24453
MARANTACEAE
Phacelophrynium sp.; 14645 (distr. as PArynium
0611
, 14564, 14930
MUSACEAE
Musa cultivars; SR, cult. Lababia
ORCHIDACEAE (Vogel et al. colls. det. by Vogel
& Schuiteman; other colls. det. by Howcroft)
Acri aie javanica ae ex BL. var. javanica;
412, 15262;NG 94
na liliifolia (J. a Ormerod; Vogel et al.
20030827
eal spp.; 14601; Vogel et al.
roses ieee 14415, 14748, ined LAE
77:NVogel et al. 20030793, 200
licula reflexa Bl; Vogel et al. se
Aopendicul a spp.; 15229; Vogel et al. 20030789,
2003079
Ascoglossum calopterum (Rchb.f) Schltr; 16443
Bromheadia finlaysoniana (Lindl.) Miq.,; 16568;
Vogel et al. 20030831
Bulbophyllum blumei (Lindl) JJ.Sm.; 15201
ee um La imum ale Rolfe; Vogel et
0030837, 20030838
ce, um ie mea sn Lindl; 14594
BRIT.ORG/SIDA 20(4)
Bulbophyllum sessile (Koen.) J.J. Sm.; Vogel et al.
20030818
ane sp., sect. Aphanobulbum; 15389
sae cent sp., sect. Sestochilus; Vogel et al.
0807, 20030814, 20030817
Perrin spp.; 15392;LAE 52323;NGF 31624;
Vogel et al. 20030808, 20030809, 20030810
20030811, 20030812, 20030813, 20030815,
20030816, 20030817, 20030829, 20030836,
20030855
Cadetia mnOMN ek Schltr; se 45195 (det.
Sch Vogel et al. 20030800
badeta ee pp. err NGF fe (no flow-
ers)
Ceratostylis sp; 14641
Claderia cf. papuana Schltr; Vogel et al.20030788
Claderia sp.; Vogel et al. 20030856
ee spp.; Vogel et al. 20030850,
2003085
Coelogyne i Lindl; 15154,15219
Coelogyne cf. fragrans Schltr; Vogel et al.
030796
=
Dendrobium austrocaledonicum Schltr; 15071
Dendrobium bracteosum Reichb.; SR, Saia River
Dendrobium coeloglossum Schltr; Vogel et al.
0804
Dendrobium hosei Ridl.; 15227; LAE 52310 (600
elev.)
Dendrobium lawesii F.v.M.; LAE 52307, 52308,
5230 )0 m elev.)
Dendrobium ?lineale Rolfe; SR, Saia River
Dendrobium macrophyllum A.Rich; 14597, 14599;
t al. 20030843, 20030845, 20030846
Dendrobium spectabile (Bl.) Miq.; 14603; LAE
2330: NGF 45191; Vogel et ne 20030822,
20030842
Dendrobium viridiflorum F.M. Bailey; Vogel et al.
20030840
Dendrobium sp., sect. Amblyanthus; NGF 45162
(ny)
Dendrobium sp., sect. Brevisaccata; Vogel et al.
2 0805, 20030857
Dendrobium sp., sect. Ceratobium; 15075
Dendrobium sp., sect. ?Distichophyllae; Vogel et
al. 20030806
Dendrobium sp., sect. Latouria; Vogel et al.
30847
Dendrobium sp., sect. Oxystophyllum;Vogel et al.
TAKEUCHI, ULTRABASIC FLORA IN PAPUA NEW GUINEA
Dendrobium sp., sect. Spatulata; Vogel et al.
2003082
Dendrobium spp.; 14647; Vogel et al. 20030802,
Dipodium spp.; a et al. ene ee a
Eria spp.; NGF 45111;Vogel et os
cats bcd aie ) Lindl;
leuca Schltr; . 25685
16603; NGF 24478 (nv)
henavrin
Lep idoaunena minor sch
Malaxis sp.; NGF 4
Neuwiedia veratri an o 14431, 14644, 15096,
6584; NGF 24478; Vogel et al.
20030833
Paphiopedilum violascens Schlitr.; 16618; NGF
45147 (nv)
Phreatia sp.; Vogel et al. 20030852
Plocoglottis sp., maculata facies; 15055, 15087
15159, 16565, 16594; LAE 52037; NGF ;
5192,477
Plocoglottis sp.,sakiensis facies; 14353;NGF 25688
Plocoglottis spp.; 14665 (inflorescence terminal);
Vogel et al. 20030794
Podochilus sp., scalpelliformis facies; 14824
Pseuderia cf. similis Schltr; 14489, 15035, 15187,
15217
Pseuderia spp.; Vogel et al. 20030834, 20030848
Rhynchophreatia sp.;
?Sarcanthopsis sp.; 15075, 16443
Spathoglottis plicata Bl.; Vogel et al. 20030820
Spathog ottis plicata BI. ssp. humilis Howcroft
ed.;1 yes 14452, 14524, 14631,15163;NGF
45095, 47766
SP neat portusfinschii Krzl.; Vogel et al.
20030819
Tainia sp.; Vogel et al. 2003079
Tropidia disticha Schltr.; a : o Rau 286
Tropidia sp.; Vogel et al. 2
Vanilla sp.;Vogel et al. une ene
Vrydagzynea novaguineensis J.J.Sm.; NGF 45120
genus indets.; 2, 14897, 15069 (bottled);
Vogel 20030857
PANDANACEAE (Freycinetia dets. by K.L. Huynh)
Freycineti t 47
—
+
CIC
Fone nes (Sav.ex Lam.) Merr.; 14587,
14589, 14596, 16539; Jacobs 9668; NGF
Freycinetia glaucescens Huynh; 14428
1543
jE ; tint realy.
Is Huynh; 16474
Freycinetia longiramulosa Huynh; 15259
dies macrostachya Mart; 14593
Freycinetia mediana Huynh; 15233
ie inetia neoforbesii Huynh; NGF 45155
Freycinetia oraria Huynh; 14414, 14528, 14731
Freycinetia pluvisilvatica Hiavehe 14694
Freycinetia rubripedata Huynh; 16440
Freycinetia sachsenensis Huynh; 14384
Freycinetia takeuchii Huynh; 14351, 14552,16475
Freycinetia tenuis Solms.; NGF 31628 (kK)
Freycinetia sp., ?marginata Bl.; NGF 24488
(staminate)
Freycinetia spp.; 14694; LAE 52341 (550 m elev.)
Pandanus cernuifolius Merr.& Perry; 14421, 14442
Pandanus lustrorum Stone, vel aff; 15559
Pandanus tectorius Parkinson; 14379
Pandanus sp.; SR, Kulindi, 30 m canopy trees
POACEAE
Bambusa forbesii (Ridl.) ee 15360
Bambusa vulgaris Schrad. i SR, Bitoi
et eee ONG BS soy Tals —
is (Retz.) Trin.; 14438
ee cf. brownii (Kunth) Nees; 14532
— tenella (L.) P Beauv.ex Roem. & Schult;
4542
Fri aie triseta Nees ex Steud.; 14530
Bailey) Merr.; 14816
Ichnanthus vicinus (F.M .
haemum mutica L.; 14439
Nastus schlechteri (Pilger) Holttum, vel aff; 14583,
14755, 15235, 15395, 16604, 16642
aspalum conjugatum Berg.; 14562
Paspal biculare G. Forst.; 14543
Saccharum edule Hassk.; SR, cult. Lababia
Saccharum officinarum L.; SR, cult. Lababia
Sacciolepis indica (L.) Chase; NGF 45193
rochloa mutica (Forssk.) T-Q. Nguyen; SR, Kulindi
Zea mays L.; SR, cult. Lababia
SMILACACEAE
Smilax cf. calophylla Wall. ex DC; 14655, 15016
1556]
Smilax cf.ovatolanceolata Koyama; 14742, 15023
Smilax sp.,‘australis-zeylanica facies’; 14567
TRIURIDACEAE
Sciaphila tenella Bl.; NGF
Sciaphila sp.; Jacobs ae re 45144
ZINGIBERACEAE
Pleuranthodium sp., Psychanthus facies; 14459,
14957, 14963
1544
Riedelia corallina Val., NGF 2445
Riedelia cf. hollandiae Val.; es 16602, 16617
Riedelia sp. A; ?15049, Hohe
Ri ra B; 14383, 1 14754, 14960, 16623
tae sp.C; 14403, va 15353, 16610 (14403,
648 distr. as oe tephro-
Shia NGF 24499, 477
DICOTS
ACANTHACEAE
Acanthus ilicifolius L., sens. lat. (spineless form);
Graptophyllum sp.; LAE 52757 (nv)
Ruellia sp. ee ium); 15252
Strobilanthes sp.; 15269
ACTINIDIACEAE
Saurauia sp., aff. schumanniana Diels; 14356,
14417, 14557, 14579, 14623, 14787, 14918,
15032, 15045, 15443;NGF 25670,47765;Rau
565
ANACARDIACEAE
Buchanania macrocarpa Laut.; 15221, 16589;
NGF 24484
ae brevipetiolata Volk.; SR, Tabali
Bive
Cas eects montanum Laut.; 15066; NGF
39416, 45110
Euroschinus papuanus Merr.& Perry; NGF 25666
Semecarpus aruensis Engl; NGF 45151
Semecarpus australiensis Engl;Jacobs 9646; NGF
Semecarpus cf. brachystachys Merr. & Perry;
14486, 14614, 15005, 15358; NGF 28085 (K)
Semecarpus bracteatus Laut., vel aff.; 14821,
14948, 15046
Semecarpus cassuvium Roxb.; Jacobs 9559; LAE
6
Semecarpus forstenii Bl; Jacobs 9594
Semecarpus sp.,aff. *schlechteri Laut. NGF 39436
ANN
Cyathocalyx cf. polycarpum C.T.White (papuanus-
polycarpum group); 14819, 14910, 15402
NGF 24489 (nv), 28075 (K, nv), 31634 (K, nv),
1642 (K, nv), 45141 (nv
Goniothalamus aruensis Scheff, NGF 25663,
5160
Goniothalamus sp.; Jacobs 9539
BRIT.ORG/SIDA 20(4)
Haplostichanthus longirostris (Scheff.) Heusden;
NGF 25002 (K)
Mitrella sp.(Fissistigma s.lat.); 14693, 14912, 15388
?Petalo See megalopus K. Sch., vel aff; 14652,
15167, 15177 (all nos. without flowers, pos-
sibly an rid
Polyalthia cf. oblongifolia Burck.; NGF 25006 (K),
31638 (K)
Polyalthia spp.; 14865, 16631; Jacobs 9666, 9688;
NGF 24456
i. Hae aii Sincl.; Conn et al.
47,NGF 45137;Rau 613
Pe ride sp., aff. *filipes a & K.Sch,; 14822
sp., cordata-rosenbergiana facies; 14859,
ae 14919
Xylopia sp., aff. sual Diels; 14941
genus indet.; LAE 52034
—
, 15580
APOCYNACEAE
Alyxia acuminata K.Sch., sens. lat.; 14698, 14766,
14977, 14985, 15025, 15145, 16563; LAE
he ann noe 28072 (K)
Hemsl.; 15433
Cerbera floribunda K. Sch.; 14516,15344
Cerbera manghas L.; 15134
Hoya lauterbachii K.Sch.; NGF 22888 (nv)
Hoya sp. A; 1 es 14933, 15206, 15240
nye sp. B; 1
Ichne Care US VWOTIGTIG (Schitr,) Middleton; Jacobs
Melodinus cf. novoguineensis (Wernh.) Pichon;
14679
Ochrosia coccinea (Teijsm. & Binn.) Miq.; 14422,
15040
Parsonsia Eaten oe Mabberley;
4839, 15133, 15
Parsonsia ee (Teijsm. ; a ) Boer|.; 15137
Parsonsia curvisepala K.Sch,; 14718, 14853, 14971;
SF 2567
NGF 2 ,45128
Sarcolobus retusus K. Sch.; 14354, 14606, 15138,
16566; Bellamy 5; Conn 174; Jacobs 9680;
Tabernaemontana aurantiaca Gaud.; 14395
Tabernaemontana pandacaqui Lam.; 14858,
15224
genus indet.; 15212
CEAE
aya celebica (Harms) Philipson; SR, near
Cape Roon
ARALIA
Mackin
ie
TAKEUCHI, ULTRABASIC FLORA IN PAPUA NEW GUINEA
Mackinlaya 7 echteri (Harms) pa 14671,
1509 ate 15572; NGF 4516
gel ( Piso 15264
Bleranata stahliana Warb.; NGF 6205
Schefflera sp.; 14797, 14804, 14833
ARISTOLOCHIACEAE
Aristolochia momandul K. Sch.; 14394
ASTERACEAE
Adenostemma lavenia (L.) O. Ktze.; 15248
Bidens pilosa L. var. minor (Bl) Sherff; 14436
Erigeron sumatrensis Retz.; 15204
en biflora (L.) DC; SR, near Tabali
BEGONIACEAE
Begonia sp., brachybotrys-pseudolateralis facies;
15424
BIGNONIACEAE
Ph hee cana (Lf) K.Sch.; 15268
a (Andr.) Steen.; 14852
a (Bl) ane 24467,
ae dendrophila
39434
BIXACEAE
Bixa orellana L.; SR, cult. Lababia
BURSERACEAE
eae um seal Leenh.; NGF 28060 (K)
Burtt; NGF 28092
ak um vitiense A Gray; Jacobs 9672
Canarium sp.,close to vitiense A.Gray; NGF 31648
(K)
Haplolobus floribundus (K. Sch.) HJ.Lam; Jacobs
9673; LAE 52055; NGF 24468
CANNABACEAE
Gironniera celtidifolia Gaud.; 14404, 14708, 14779,
14921, 15378, 16640; LAE 51667; NGF 25680,
14483, 15064, 15173,
, 187; NGF 31633 (K);
47768
Trema cannabina Lour.;
15346; Conn et al. 185
CARICACEAE
Carica papaya .; SR, cult. Lababia
CASUARINACEAE
Gymnostoma papuanum (S. Moore) L.A.S.
Johnson; 14444, 14778, 14780; NGF 25665
CELASTRACEAE
Bhesa archboldiana (Merr. & Perry) Ding Hou;
42, 15056, 15074 (distr. as B. robusta),
15571
1545
Loeseneriella macrantha (Korth.) A.C. Smith;
obs 9619
Lophopetalum torricellense Loes.; 16600; LAE
Salacia chinensis L.; 16425
Salaci arpa K. Sch; LAE 52316
Salacia papuana (Loes.) Ding Hou; LAE 52320
Salacia sp.; NGF 45109 (carpological coll. miss-
ing, sheet sterile)
CHRYSOBALANACEAE
Hunga papuana (Baker f.) Prance; 14922, 14929,
14940, 15584
Maranthes corymbosa Bl; 14556, 14712, 14920,
14937, 14942, 14968; NGF 31644 (K)
Parastemon versteeghii Merr.& Perry; 15021; NGF
45157
— ree CT. White ssp. whitei Prance;
14 15015; Jacobs 9583; NGF 28063,
ao
ada train re taal nee pe P. Stevens)
a llum goniocarpum 14981; NGF
ee
ATA
Ca ie inophyllum L;14535,15111;Swaine
66
Calophyllum morobensis Stevens; Johns sn (LAE
sheet 129181, nv); NGF 24490 (nv)
aa pi oman Laut.; 14892, 14905,
5347, 16564; LAE 52312; NGF 24487
eh um streimannii Stevens;LAE 52755;NGF
24285, 24491, 28084 (K)
Calophyllum sp.; 14535
Garcinia cf.assugu Laut.; 16632
Garcinia celebica L.; 14416, 14419, 16413
Garcinia dulcis (Roxb.) Kurz; 15406; NGF 28066
(K)
inia h ji Laut.; 15050; NGF 24458
ia latissima Miq.; NGF 24447, 28073
Gana ledermanii Laut.; 14333, 14348, 14617,
14734, 14867, 15404
Garcinia maluensis Laut.;
31646 (K)
15412, 16574; NGF
Garcinia eee Stevens; 14886, 14888, 15169,
151921 16583; NGF 24449, 45165
Garcinia sp., ass ie u-maluensis facies; 14651,
4689, 14713, 14729, 14830, 14863, 14890,
14955, 14987, 15001, 15199, 16436; LAE
5276]
Garcinia sp.,aff. ?sabangensis Laut.; 14433, 14769,
14808, 16641
1546
Garcinia sp. ?nov.;ledermanii facies but not Cam
bogia; 15379, 15409; NGF 24455
Garcinia spp.; 14325, 14799
Mammea papyracea Stevens; NGF 39414
TACEAE
Lumnitzera littorea (Jack) Voight; 14378, 14902:
LAE 52059; hes aot 44153
lerminalia canalice Exell; 15256
pa LeSR, Lababia
Terminalia rubiginosa K.Sch.; LAE 52043
Terminalia sepicana Diels; NGF 39412
CONVOLVULACEAE
Erycibe cf. hellwigii Prain; 14585, 14682;LAE 51691;
NGF 2447), 24498, 45100 (keys between F.
and E. puberula)
pee batatas (L.) Lamk; SR, cu a ‘a
[pomoea pes-caprae (L.) R.Br; 14
Ipomoea pes-caprae (L.) R. Br. ssp ae (L.)
Ooststroom; NGF 22879
Merremia peltata (L.) Merr.; SR, Tabali River
CUCURBITACEAE
Cucumis sativus L.;
CUNONIACEAE
Ceratopetalum succirubrum C.T. White; sn
(sterile)
Weinmannia fraxinea (D.Don) Miq,; 15053, 15054,
15150
DILLENIACEAE
Tetracera nordtiana Fv.M. var. moluccana
Hoogl.; 15004
DIPTEROCARPACEA
Anisoptera thurifera a Bl. ssp. polyandra
(BI.) Ashton; SR, throughout KWMA; LAE
51965, 52039
Hopea cf. glabrifolia C.T.White; 14620
EBENACEAE
Diospyros cf. elliptica (J.R. & G. Forst.) Green; NGF
28070 (K), 31631 (K), 31632 (K), 31647 (K),
45168, 45169 ors inflorescences cymose
ost simple)
ers ferrea Canes Bakh., sens. lat.; 14366,
14555, 14559, 14574, 14715, 14847, 14943,
14975, 15033, 15439 (preceding nos. distr. as
D. elliptica but inflorescence not cymose),
15387; NGF 39096
Diospyros novoguineensis Bakh.; 15371, 16417;
LAE 51679; NGF 24457, 28078 (K), 45148
Terminal).
-SR, cult. Lababia
—
Mart.)
5
sec
—
BRIT.ORG/SIDA 20(4)
Diospyros papuana Val. ex Bakh.; NGF 45135
Diospyros cf. sogeriensis Bakh.; 15029, 15098,
15381; NGF 24450, 24500
Diospyros sp., ?ferrea group’; NGF 25010 (K)
ee
itium parvifolium Schltr; NGF 31627 (kK)
aeocarpus dol een Schltr. ssp.
dolichostylus; NGF 4
aeocarpus jean — LAE 51670,
52758; NGF 44209,
aeocarpus sphaericus ae K. Sch.; 14418,
14548
Elaeocarpus sp.,‘sepikanus group’; LAE 52048
Elaeocarpus sp. ‘nov. 5, aff. miegei fide Coode;LAE
52295; NGF 25668, 28089
Sloanea pulchra (Schitr.) A.C. Smith ssp.
morobensis Coode; NGF 25005 (K), 28067 (kK),
E
aul
E
Sloanea sogerensis Bak. f., sens. lat.; 14695; Conn
et al. 197; NGF 28065 (K), 31649 (K), 45117
(most colls. represent the ‘schumannii facies’
cf. Coode 1981)
ERICACEAE
Paphia megaphylia Stevens ined.; 15383B, 16644
ERYTHROXYLACEAE
sca ecarinatum pd 14763, 14768,
72, 14786, 14796, 1
EUPHORBIACEAE sens. lat. (including
Phyllanthac
Actephila 9 lindleyi (Steud.) Airy Shaw; 14386,
14456, 14639, 14677, 15030, 15354, 16592;
Jacobs 9538; LAE 52031; NGF 45136
Antidesma moluccanum Airy Shaw var.
moluccanum; 15417
Antidesma polyanthum K. Sch. & Laut, 14856,
15014
Antidesma cf. sarcocarpum Airy Shaw; 14972;
NGF 28061 (K)
Antidesma sphaerocarpum Muell. Arg.;
14493, 15038, 15414, 15568; LAE 52051
4516/7
14474,
> NGF
Aporosa brassii Mansf; NG
Aporosa praegrandifolia (S. Hoo Schot; 15168;
AE 52049; NGF 45134, 4517
Breynia cernua (Poir,) a a
4834, 14967; NGF 47
Claoxylon_ aff. Leite Airy Shaw var.
ledermannil, 14738, 14751 (nos. distr. as
14423, 14580,
TAKEUCHI, ULTRABASIC FLORA IN PAPUA NEW GUINEA
group ‘Purpurescentia’), 15386, 15416, 16587;
LAE 51669
Cleistanthus pedicellatus Hook.f; 14749 (distr. as
Kairothamnus phyllanthoides)
Cleistanthus sp., cf. myrianthus (Hassk.) Kurz
16621; NGF 44608
Cleistanthus sp., aff. pedicellatus Hook. f; 14473
(leaves obtuse, capsule lobed)
Codiaeum variegatum (L.) Bl. var. moluccanum
(Decne) Muell. Arg.; Jacobs 9596; NGF 45143
Croton - choristadenius K. Sch.; 14747, 15151,
GF 45101, 45102
aes cf. labios Schodde; 15255
Euphorbia hirta L.; 14435
Glochidion granulare Airy Shaw; 16412
Glochidion sp.; 14571
Kairothamnus phyllanthoides (A. Shaw) A. Shaw;
LAE 52046, 73372; NGF 24462, 44211, 44602,
45108
Macaranga angustifolia Laut. & K. Sch. 15375;
Macaranga bifoveata J.J. Smith; 14525, 14582,
14586, 14807, 15359, 16431
Macaranga polyadenia Pax & Hoffm.; SR, passim
Macaranga tanarius (L.) Muell. Arg.; NGF 44609
(nv)
Manihot esculenta Crantz; SR, cult. Lababia
Phyllanthus cf. ee : oo 14321, 14387
Phyllanthus sp.; NGF
Pimelodendron ae inicum Hassk.; 14788,
15405; NGF 31640 (K), 31643 (K)
Suregada glomerulata (Bl.) Baill; 14367, 15013,
15034
FABACEAE
Abrus pulchellus Thwaites ssp. pulchellus; NGF
62046
achis hypogaea L.; SR, oe ee
A hid, 7 I, yi Fv.
Archidendron mo lle (K. ae ) Wit, vel aff; 16524;
21; LAE 52760; NGF 28090 (K)
eat ptenopum Verdc.; 16616; NGF
cen Sp.;
aesalpinia crista . ce 22896
ie ia retusa 37
Dahibergi cance ee (Dennst.) Prain; 14882
Derris indica (Lam.) J.J. Bennett; 14619, 15123;
25686 (nv)
Derris trifoliata Lour.; SR, Tabali
1547
Desmodium umbellatum (L.) DC; 14361, 14450,
15117
Inocarpus cf. papuanus Kostermans; 14/719
Intsia oe (Colebr.) O. Kuntze; SR, Bulili shore-
lin
Maopsehants lauterbachii Harms; SR, Saia
Pe ce sp., aff. schefferi K.Sch.& des 14318
15582; sp. E in Verdcourt (1
Mucuna schlechteri Harms; | pe 4966, 15377;
Jacobs oe NGF 25673, 45096
rmosia calay Azaola ex Blanco; 14958
Gonben tomentosa L. ssp. tomentosa; Martin
2520; NGF Bane
Vigna unguiculata (L.) Walp.; SR, cult. Lababia
7,14517 (nos.
K),
FAGACEAE
Lithocarpus vinkii Soepadmo; 1431
distr. as L. celebicus); NGF 24492, 31622
44213
GENTIANACEAE
Fagraea amabilis S.Moore; 14462 (610 m)
Fagraea berteroana A.Gray ex Wall.; Jacobs 9542;
—
pais
Fagraea ceilanica Thunb.; 14369, 14598, 15020
Fagraea racemosa Jack ex Wall; 14760, 14908,
14979, 15340; Jacobs 9622;LAE 51665,52041,
52335, 68510; NGF 22885 (K)
pal a ae
eschynanthus cf. kermesinus Schltr; 14504
ascot cf. leptocladus C.B. Clarke; 1446]
Boea lawesii H.O. Fo eee re
Boea mollis Schltr; NGF 477
es oe schumanniana ie vel aff; LAE
516
oe sp., sect. Centrosi ace: NGF 45173
Cyrtandra sp
14458, see
GOCDENIACERE
et L Oxo} hyllum P haeotrichium;
7R. Br; SR, Kulindi
Seaevola sericea Va hl; 14363
ICACINACEAE
Platea latifolia Bl.; 154
Rhyticaryum longifolium K. Sch. & Laut.; 14653,
14664, 14877; Jacobs 9671
LAMIACEAE
Callicarpa longifolia Lamk.; NGF 22883 (K)
Gmelina ledermanni H.J. Lam; 14365, 14487,
1548
14924, 15026, 15114, 15343; Jacobs 9536;
NGF 28077 (K); Rau 609
Leucas flaccida R. Br; NGF 25684
Plectranthus sp., aff. ?parviflorus Willd.; 14437
(does not key)
P, 1 serratifolia L; 14441, 14893; NGF 47784
LAURACEAE
Actinodaphne nitida Teschn.; NGF 31637 (K)
Cassytha filiformis L.; 14527, 14540
Cinnamomum eugenoliferum Kosterm.; NGF
45130
Cryptocarya cf. densiflora Bl; LAE 52315
Cryptocarya laevigata Bl.; 14709, 14947, 14953,
5153, 16588, 16608
Cryptocarya multipaniculata Teschn.; 15241
Cryptocarya novoguineensis Teschn.; 16633; NGF
28086 (K)
Cryptocarya pie Teschn., vel aff; 15578
ndra ae Teschn.; NGF 28068 (K)
erate forbesii Gamble; NGF 39098, 45187
Litsea sp., collina facies; 15348
ee guppyi facies; 14703, 14978, 15244; NGF
24472
Persea americana Mill. SR, cult. Lababia
LECYTHIDACEAE
Barringtonia apicu
F 28081 (K)
Barringtonia asiatica (L.) Kurz; SR, Lababia, also
| colls. Lasanga
Barringtonia lumina Jebb ined; SR, Saia River (un
——
ata Laut; 15113; LAE 52053;
C : ‘
a _ ia Jebb ined.; 14560, 14576,
14681, 14704 (distr. as B. calyptrocal bs : val.
bori a 515), 16438, 16585; Jacobs 955
Barringtonia racemosa (L.) Spreng.; 1 oe
arringtonia sp., aff. ?acutangula (L.) Gaertn.;
153604
Barringtonia sp. A; NGF 25676, listed as uniden
tified ‘group 2’ in Jebb (1991)
Barringtonia sp.B; 14866 (distr.as B.cal ta but
not that species)
LINACEAE
Hugonia jenkinsii Fv.M.; 14632, 15112, 15160; Ja-
bs 9603; NGF 24470
LOGANIACEAE
Geniostoma rupestre J.R.& G.Forst.; 16597; Conn
et al. 175, 178, 186, 207; Jacobs 9597, 9613;
BRIT.ORG/SIDA 20(4)
NGF 24486,47763-Rau 283,590, 606,608,614,
617
Geniostoma rupestre J.R. & G. Forst. var. rupestre;
14448, 14480, 14714, 14727, 14752, 14832,
14838, 14855, 14931, 15072, 15442
Neuburgia corynocarpa (A. Gray) Leenh.;
15258, 15363
15245,
Strychnos minor Dennst.; 14316, 15024, 15027
NTHACEAE
Amyema scandens ( Tiegh.) Danser ssp. scandens;
Decaisnina aie (K. Sch.) Barlow; 14551,
ae F 45196
Dendropt urvata (Bl) Miq.; 14430, 14529
genus dee NGF 47761
MAESACEAE
Maesa haplobotys F.v.M.; 14923, 15238, 15384,
16620
MALVACEAE
Rrovwsnlowsi7 ar
g Kurz; 15275
Commersonia bartramia (L te 14410, 14460,
823, 15047, 15342; Swaine 10
Heritiera littoralis Ait.; 15249; LAE 52333
Hibiscus tiliaceus L.; SR, Tabali River and Lababia
beachfront
Kleinhovia hospita L.; SR, Lababia
Microcos sp.; NGF 28069 (K)
Sida cordifolia \.; 14434
Sterculia cf. lepidostellata Milbr.; 14820
Sterculia morobeensis Tantra; Jacobs 9663; NGF
28093 (K)
Sterculia ae hee Mildbr; 14721;
nn et al. 2 2045; NGF 28076 (kK)
fea Sp. oe al sot Fv.M.; 14591,
14743, 14803, 1
Sterculia A i co (1976); Jacobs 9639
Thespesia populnea (L.) Sol.ex Correa; SR, Lababia
TOMATACEAE
Astronidium morobiense Maxw.; 15351, 15560,
16223 (dets.S.S.Renner); NGF 25658 (nv) and
45180 (nv) could be this sp
Medinilla sp., aff. a Bak. f;, 14795
Medinilla spp.; 15365, 1
Melastoma Seneca Lis SSp.
malabathricum; 14372,15121 (distr.as M.af-
fine D.Don
Poikilogyne sp.; NGF 45232
TAKEUCHI, ULTRABASIC FLORA IN PAPUA NEW GUINEA
MELIACEAE
Aglaia brownii C.M. Pannell; 14911; NGF 24446
Aglaia cf. rimosa (Blanco) Merr.; 14666, 15076
Aglaia sapindina (Fv.M.) Harms; 14427, 15230,
266, 15366, 15574, 16222; Jacobs 9576,
9607; NGF 45140; Warete-Namorong /
Aglaia tomentosa Teijsm. & Binn.; 15407, 15422,
16643
Aglaia sp.,aff.? / Merr.& Perry; 14332,
14482, 15179 (distr. as A. silvestris, vel aff.),
15419, 15576; NGF 45183
ey sp., ait Peete (F.v.M.) Harms;
5175, 1 15350; LAE 52054
soho areas (Wall.) R.N. Parker;
5434
14393,
Chi haces lasiocarpus (Miq.) Val.; 15257, 15429
Chisocheton lasiocarpus (Miq.) Val. entity
‘schlechteri’. 15041, 15104, 15161, 15176,
16599; LAE 52343
Chisocheton lasiocarpus (Miq.) Val. entity
‘weinlandii’, LAE 62059
Chisocheton sapindinus Stevens; 14336, 15411,
15574, 16414; NGF 45140
Dysoxylum arborescens (Bl.) Miq.; LAE 51686
Dysoxylum papuanum (Merr.& Perry) Mabb.;LAE
52319
\ Pal
Benth.; NGF 24482
Xylocarpus granatum Koen.; 14634, 14899
MEMECYLACEAE
Memecylon sp.,sepicana-schraderbergense facies;
14670; NGF 25003
MENISPERMACEAE
Hypserpa polyandra Becc. var. polyandra; 14553,
14744, 15078; Jacobs 9653; LAE 51673; NGF
473
Macrococculus pomiferus Becc.; LAE 62057
ana
Philioson; 154
a obl ongata Philipson, vel = 14334, 15012
Levieria nitens Perkins; NGF 24439 (nv)
ee irsuta (Warb.) Perkins; 14575, 14706,
GF 25687
AE
Artocarpus communis J.R. & G. Forst.; SR, Tabali
River
us adenosperma Miq.; 14359
j
f
Ficus ampelas Burm. f. 14565, 14581, 14592,
14726; NGF 4511
1549
Ficus arfakensis King; 15263
Ficus benjamina L.;LAE ae a NGF 44154
Ficus calodictya Surnmerh.; 14962
Ficus au Laut. & K.Sch.; ee 15222,15272
Ficus Warb.; 14687, 14964
Ficus irritans ene 14691, 14970
Ficus itoana Diels; 14
Ficus odoardi King; i ave
nee He n Warb.; 1, 14569, 14584,
14696, 15210, or 15569B (some
nos. aie with F. subtrinervia Laut. & K.
Sch.
Ficus rhizophoriphylla King; NGF 28087 (K)
Ficus trachypison K. Sch. var. pallida Corner; LAE
2759
Ficus wassa Roxb.; NGF 31645 (K)
Parartocarpus venenosus (Zoll.& Mor.) Becc. ssp.
papuanus (Becc.) Jarrett; NGF 25001 (K)
Prainea papuana Becc.; NGF 45150
MYRISTICACEAE
Gymnacranthera farquhariana (Hook. f. & Th.)
Warb. var. zippeliana (Miq.) R. Schouten;
14337, 14680, 14872, 14945, 14954A, 15410,
6626, 16627, 16628
Horsfieldia hellwigii (Warb.) Warb.; 15435
Horstieldia laevigata (BI.) Warb. var. laevigata; LAE
5] 30
Horsfieldia pilifera Markgrf,; Jacobs 9609, 9609A;
56; NGF 28080 (nv)
lorsfieldia sylvestris (Houtt.) Warb,; 15271
Horsfieldia tubercular (K. Sch.) Warb. var.
tuberculata; |
Spe csp Sincl. ssp. chrysophylla;
14735,14951, a Jacobs 9574;LAE 52032,
aie Sin Ch
entrecasteauxensis ee ne ie ie
14602, 15018, 15082, 15130; LAE 52029
Myristica cornutiflora Sincl.ssp.elegans de Wilde;
14590, 14
Myristica we de Wilde; 14566, 14700, 1493
182, 281 (nv); LAE 51668, Bie
ee vee 6
Myristica globosa Warb.; Jacobs 9572
Myristica markgraviana A.C. Smith; LAE 52027
Myristica sulcata Warb.; Jacobs 9685; LAE 52028
Myristica tubiflora Bl.; NGF 2 42
Myristica umbrosa Sincl.; 14600, 15188; LAE
52033, 52047, 52287, es NGF 25659
Myristica
1550
Myristica sp. ?nov.; 14340, 14661, 15011, 15109,
15397, 16424, 16573
Myristica sp.; 16591
ideas
ilatum (L.) Blanco; 15274
conancium en thum (Laut. & K. Sch.) Mez;
14338, 14447, 14570, 14761, 14774, 14825:
LAE 52040, 52050
Discocalyx kaoyae Pipoly & Takeuchi ined; 16441
Discocalyx orthoneura K. Sch.; NGF 24451
Discocalyx sp.; Jacobs 9584 ea
se oe Mez; 14425, 14429, 14445,
4572, , 14997, nes 16580, 16582,
oe nuk 52302
Fittingia urceolata Mez; Conn et al. 284; Jacobs
9644; NGF 45139,45163; Rau 291
Myrsine leucantha (kK. Sch.) Pipoly; NGF 24429,
45235 (nv)
oe ne rawacens
44603
1s A. DC; 14604, 15562; NGF
Myrsine sp., (Rapanea ?lamii Sleumer); 14469
MYRTACEAE
Decaspermum bracteatum (Roxb,) AJ. Scott var.
bracteatum; 14611, 14770, 14782, 14792,
15007, 16567; LAE 52322
Eucalyptopsis papuana C.T. White; LAE 52296,
2331, 52334; NGF 28097 (K)
Lophostemon suaveolens (Sol. ex Gaertn.
Wilson & J.T.Waterh,; 15116
Myrtella beccarli Fv.M.; 14349, 14380, 14
15583; Bellamy 6; Jacobs 9650; LAE oe
72469; NGF 44151; Paijmans 1564; Swaine 8;
PG.
aed
Is insignis Diels; NGF 25018
Psidium guajava L.; SR, cult. Lababia
Rhodamnia blairiana Fv.M. vat. propingua (C.T.
Scott; 14405, 14841; NGF 44215
ene pinnatinervis C.T.White; LAE 52304
m)
Syzygium buettnerianum
: E
na
NO
K. Sch.) Niedenzu, vel
; 297
Syzygium claviflorum (Roxb.) Cowan & Cowan;
14477
Syzygium effusum (A. Gray) C. Muell., sens. lat.;
14315; NGF 24483, 45159
Syzygium cf. flavescens (Ridley) Merr. & Perry;
14341
Syzygium furfuraceum Merr. & Perry; 14740,
14944; NGF 31635 (K), 31650 (K)
BRIT.ORG/SIDA 20(4)
/ 15194; NGF 39095,
Cyazvaiiim lentonelsrsim ‘ .
fe Diels
47770; Swaine 9
longipes Merr, & Perry; 14388
Svzvaqfum
Syzygium nutans (K. Sch.) Merr. & Perry; Jacobs
G
Syzygium richardsonianum Merr. & Perry, vel aff;
14733, 14954B, 15084, 15144, 15399; LAE
51663; NGF 24476, 39431
Syzygium pace (aa os & Perry; LAE
52289 7,;NGF 4
Syzygium ae ies : — NGF 24428
Syzygium trivene (Ridley) Merr. & Perry; 14381,
14402, 14413, 14484, 14518, 14588, 14618,
14688, 14737, 14851, 14969, 14994, 14998,
15396, 15437, 15569A, 16434, 16581; Jacobs
9617, 9645; LAE 51664; NGF 24497, 25657,
39419, 44155
Syzygium cf. trivene (Ridley) Merr. & Perry; 14773,
14811, 14835, 14845; NGF 25669 (noncon-
formist nos. with persisting bracts but cf.
Hartley & Perry 1973: 213)
Syzygium viburnoides Diels; 14479
Syzygium xylopiaceum (Diels) Merr. & Perry; Ja
obs 96
Syzygium sp.,=Acmena acuminatissima (Bl.) Merr.
& Perry; NGF 28095 (K)
Syzygium sp., aff. malaccense
—
L.) Merr. & Perry;
Syzygium sp., aff. 2rostratum (BI.) DC; 15127
Syzygium sp. ?nov.; aff. pyriforme Merr. & Perry;
15382, 16607
Syzygium sp. ?nov.,aff.couplet 57a or 88a; 14723,
Syzygium sp.; 15095
Syzygium sp.; 15428
Tristaniopsis macrosperma (F.v.M.
J.T.Waterh.; 14343, 15058, 15131,
24464, 44607-Vinas & Kairo 295
Tristaniopsis oreophila (Diels) P.G. Wilson & J.T.
Waterh.; NGF 42638
Xanthostemon petiolatus (Val.) P.G.Wilson; 15147,
PG. Wilson &
15579;NGF
nS
NEPENTHACEAE
i neoguineensis Macf.; 14453, 14697;
837 (distr. as N. mirabilis), 15573, 16435;
nies 9651, 9658; LAE 52313; NGF 24461,
24463;Rau 5/1;Watt 31
NYCTAGINACEAE
Pisonia longirostris Teijsm. & Binn.; LAE 62049
TAKEUCHI, ULTRABASIC FLORA IN PAPUA NEW GUINEA
NYSSACEAE
Mastixia kaniensis Melch. ssp. kaniensis; LAE
52
OCHNACEAE
Brackenridgea forbesii Tieg.; 16629; NGF 25004 (kK),
28091 (K), 44205
<<
Schuurmansia henningsii K. Sch.; 15220; NGF
31620 (K), 39413, 39430
OLACACEAE
Anacolosa papuana Schellenb.; 16624
OLEACEAE
peer ee Roxb.; LAE 62048
Is (Lingels oe Kiew; 14385,
14607, 14 672, ae 15149 (distr. as C.
ramiflorus); Bellamy 9;LAE ae NGF 24465;
Chionanthus sessiliflorum (Hemsl.) Kiew; LAE
Jasminum schumannii Lingelsh.; LAE 62050
OXALIDACEA
toe L.; 14408, 14817
DACEAE
_ aie ca Koord. var. celebica; 14660,
927, 15037, 15209, 15400, 15403; LAE
mie
PASSIFLORACEAE
Adenia heterophylla (BI.) Koord.ssp. heterophylla;
NGF 62062
Passiflora foetida L.; SR, Lababia
PENTAPHYLACACEAE
Eurya sp./?spp.; 14463; NGF 24431 (nv),45234 (nv)
PIPERACEAE
Piper aduncum L.; 15273
Piper celtidi aie ae 15243
Piper lessertianum (Miq.) C. DC.; 14432, 14455,
14828
Piper macropiper Pennant; 14561
Piper novoguineense Warb.; 14345
Piper pseudoamboinense C. 25678
Piper versteegii C.DC.;LAE 5
Piper sp.; 14314
POLYGALACEAE
Eriandra fragrans Royen & Steen.;
15239, ]
P lyvaala nani.
b
ata L.; 14375
14376, 14683,
14777, 14871, 14873, 14874, 14875, 14946,
5380
1551
POLYOSMACEAE
Polyosma sp.,aff. forbesii Val; 14329, 14466, 14884,
14894, 15090, 15413; NGF 52337
PROTEACEAE
Bleasdalea papuana (Diels) Domin; LAE 52344
50m
Finschia chloroxantha Diels; 14364, 15146; LAE
52060, 52345; NGF 45119, 47783
Grevillea papuana Diels; 14449, 14521
Helicia obtusata Sleum.;LAE 51685, 52038, 52288,
2
52298
Stenocarpus moorei Fv.M.; 14526, 14534, 14984,
14989, 15394, 16426; LAE 52044, 52294,
52326; NGF 39091
RHAMNACEAE
Alphitonia excelsa (Fenzl) Reiss. ex Endl. (sensu
Schirarend); 15335, 15374; Jacobs 9667
Alphitonia macrocarpa Mansf, 15408; NGF 25664
RHIZOPHORACEAE
Bruguiera gymnorrhiza (L.) Lamk; 14377
Ceriops tagal (Perr.) C.B.Rob.; 15148B; Bellamy 16
LAE 68539
>
eee axillaris Bl.; 15091, 15570; Jacobs
9
NGF 24494, 31641 (K)
Rhizophora apiculata Bl; LAE 68537, 68543
Rhizophora stylosa Griff; 15143
ROSACEAE
Rubus fraxinifolius Poiret; LAE 62052
RUBIACEAE
Anthorrhiza echinella Huxley & Jebb; 14470 (distr.
5M dia tuberosa); NGF 39094, 45103
pe Aas pasa ate (F.v.M.) Puttock; Jacobs
95
Atractocarpus sp., =Randia sphaerocarpa K. Sch.
ut.; NGF 2445
Atractocarpus sp.; 14342, 14370, 14472, 14676,
14707, 14711, 14928, 15081, a 15370;
NGF 25009 (K), 39093, 45152, 4
Canthium sp., sens. lat.; 14654, sd oo 15136,
15237;Heads a NGF 39429, pee 47760,
47772; Rau 287
Gardenia lamingtonii F.M. Bailey; NGF 45185
Gardenia sp. nov.; 14864, 15101, 15558, 15575;
NGF 25675, 4515
Gynochthodes aff. YaMNeNatG (Val.) Hosokawa;
14692, distr.as ‘R
Hedyotis corymbosa fi é — 14541
Hedyotis pubescens Val.; NGF 45176
1552
Hydnophytum hellwigii Warb.; 15207; NGF 25662,
45105
Hydnophytum cf. radicans Becc.; LAE 62058
Hydnophytum s
Hydnophytum sp. - ee Jacobs 9669
lxora He : 14322 , 14392, 14625, 14628,
14 14887, ne 14936, 14938, 14976,
ee 5003, 15119, 15341; LAE 51675,
52042, 62061; NGF 24495, 31630 (K), 45179
Ixora sp. B; 14481, 15009, 15105, 15385, 15581,
16420, 1660
Lasianthus sp., papuanus-chlorocarpus facies;
15178, 15197
Morinda citrifolia L.; 14424, 15368; Conn 320; Ja-
702; LAE 52293; NGF 47769
Morinda umbellata L.; 14478, 14635
ferruginea K. Sch; 14757
Meee tuberosa oP 14730
lis (L.) L.; 15254
ot se NGF 22886 (K
Perry; NGF 22489
?Porterandia, =Randia pale he Wernh. Jacobs
9689; NGF 51688, nee
Psychotria amplithyrsa Val; 15166; NGF 51680
Be a dii So et var. archboldii;
15028, 16415, 16422
ae ai mints var. aestuarii Takeuchi;
993, 14995, 15002
Psychotria bulilimontis Takeuchi var. bulilimontis;
6416, 16428
ples croftiana Sohmer; 14490, 15048,
15566, 16430B, 16578; Jacobs 9636
ree diplococca Laut. & K. Sch. var.
diplococca; 15432
Psychotria cf. lefopal oea Merr. & Perry; 15234,
15236, 16430A
Psychotria leonardii Merr. & Perry; 15057
Psychotria leptothyrsa Miq. var. leptothyrsa; 14492:
Jacobs 9567
Mirssqenda
aus
—
<a
ns ed tenuiflora
Psychotria membranifolia Bartl, ex DC; 14488,
14831, 15184, 15203, 16575, 16577,
6606
Pct Arlee ale (Laut. & K. Sch.) V
506
8, 15339, 15426, 15427; NGF 31651
Pe cice micrococca (Laut. & K. Sch.) Val; Conn
t al. 278 (nv)
beets sri Ain 14771, 14827, 14898,
0, 15108, 15189, 15357, 16437; Jacobs
pes NGF e083 (K)
valuden sis Sohmer;
14310;NGF 25626
f sychotria v
BRIT.ORG/SIDA 20(4)
as 74A, 16593;Vinas
& Kairo
~—
Psychotria sp. ae LAE 62053
Ps sp., =Canthium ae igerum (Val.) B.L.
Burtt; 14382, 14784, 14810; NGF 24475, 25667
le oe hydrophyllacea Gaertn.; 14636; NGF
ewe
me jus paiawensis S. Darwin; 14362, 14440,
ioe 14522, 15065, 15338; peu et al. 301,
313; LAE 52328; NGF 39097, 4776
Timonius timon (Spreng.) Mert. var. es Isis S,
Darwin; 14362B; Jacobs 9626; NGF 4420
Peat fimo (Spreng.) Merr. var. timon; 15367
limonius sp., aff. rufescens (Miq.) Boerl; 14798,
14870
Timonius sp. ?nov.; 14328, 14663, 14781, 15106,
1556/7
Uncaria lanosa Wall. var. appendiculata (Benth.)
Ridsd.; 14/67
Urophyllum glaucescens Val., vel aff; 14401, 14891,
15089; soe se 9692;LAE 52056,52299,
52338; NG 448, 28094 (K), 31621 (K),
31626 Noe 607,612
Xanthophytum papuanum Wernh,; 14454, 16418;
NGF 44210, 4518
UTACEAE
Acronychia trifoliolata Zoll.& Mor; 14901, 15118,
15148, 16419
Flindersia laevicarpa White & Francis var.
heterophylla (Merr. & Perry) Hartley; 14519
15124, 15132; NGF 31636
alfordia drupifera F.v.M.; 14344
aa et ll (Seem,) Hartley; 14420, 14523,
14736, 1503
Melicope ae oides (F.v.M.) Hartley; 15373
Zanthoxylum novoguineense Hartley; 14699,
14759, 14785, 14801, 14813 (distr.as Euodia
aff. ‘alata-mollis group’) 16645; LAE 52311,
52329; NGF 25690
genus indet.,; 15564
SABIACEAE
Sabia pauciflora Bl.; 15431
SALICACEAE
Casearia erythrocarpa Sleumer, vel aff. 14327,
14339, 14368, 14451, 14701, 14806, 14846,
14861, 14909, 14988, 15355; NGF 25671,
28082 (K)
Casearia cf.macrantha Gilg; 14848, 14916, 14939,
15162: NGF 52045
TAKEUCHI, ULTRABASIC FLORA IN PAPUA NEW GUINEA
Casearia sp., aff. flexicaulis K. Sch.; te 14545,
14605, 14616, 14829, 14959, 15115 (14357,
14616, 14829 distr. as Glochi a sp.); LAE
81; NGF 25661
Casearia sp., closest to ?7novoguineensis Val.;
15418, 1
Homalium d’entrecasteauxense Craven; 14650,
14876
Scolopia novoguineensis Warb.; 14612, 14627
Trichadenia sasae Takeuchi; 15589, 16561
SANTALACEAE
arene nies R.Brown; ies 15
6423; LAE 51666; NGF 394
Sc eum qurantiacum eee : K.Sch.) Pilger;
5198; NGF 31629 (kK), 45131,45174
SAPINDACEAE
Alectryon ferrugineus (Bl.) Radlk; 14728, 14802,
15247
120, 15372,
Arytera cf. litoralis Bl; 14925
Cnesmocarpon discoloroides Adema; 15092; NGF
25007
Dictyoneura obtusa Bl. i race’ oblique
leaflets); 15362, 1642
Euphorianthus euneurus an ) Leenh,; LAE 52318
Guioa grandifoliola aly 14980; Conn et al.
195, 319 (nv); NGF 4515
Guioa rigidiuscula Radlk., or Padme com-
plex’; 15208, 16614
Harpullia longipetala Leenh.,; 15165; NGF 256/79
Harpullia ramiflora Radlk.; LAE 62055 (ny)
Mischocarpus pyriformis (F.v.M.) Radlk. ssp.
papuanus (Radlk.) Ham; NGF 47775
Pometia pinnata J.R.& Ss Rots NGF 31639
Hk; NGF 44208,
45099
?Synima aff. cordierorum (F.v.M.) Radlk.; 14550,
14577, 14633
genus indet.; NGF 45216 (nv;
Elattostachys in LAE logbook)
SAPOTACEAE
Magodendron mennyae Vink; 16570; Conn et al.
194; Jacobs 9537; NGF 24436, 45146
Palaquium cf.warburgianum Schltr; 15085
Pichonia sp., ?lauterbachiana (HJ. Lam
Pennington; 14558, 14568, 14793, 14982,
14986 (nos. distr.as Pouteria sarcospermoides
listed as
pane
1553
H.J. Lam); LAE 51674; NGF 39435, 44605,
45107
Pouteria ledermannii (K. Krause) Baehni; 14471,
ets. W. Vink)
Pouteria luzoniensis (Merr.) Baehni; 14355
Pouteria maclayana (F.v.M.) Baehni; Jacobs 9573
Pouteria obovoidea H.J.Lam;NGF 24496
genus indet.; NGF 45188
SIMAROUBACEAE
Quassia indica (Gaertn.) Nooteboom; 14389
SOLANACEAE
Capsicum anuum L.; SR, cult. Lababia
Solanum anfractum Symon; Jacobs 9558; NGF
2
olanum melongena L.; SR, cult. Lababia
Solanum symonianum Takeuchi; 12027
STEMONURACEAE
Gomphandra ?pseudoprasina Sleum.; LAE 62056
(nv)
ee ammui (Kaneh.) Sleum.;
4756, 14857, 14983; NGF 24481 (nv)
SYMPLOCACEAE
Symplocos cochinchinensis (Lour.) S. Moore;
14464, 15008, 1515
THEACEAE
Gordonia papuana Kobuski; 15093, 16576; NGF
14400,
THY MELAEACEAE
ices coccinea (Gaud.) F.v.M.; 14745, 15017,
15218, 15565; NGF 28062 (K)
Pee macrocarpa (Scheff.) Boer|.; 16601
(Hemsl.) Chew; 15260
Poikilc SPEFITIUTT! SP. 14934
RTICACEAE
Dendrocnide longifolia
ERBENACEAE
Stachytarpheta cayennensis (Rich.) M.Vahl; 14537
VITACEAE
Cissus sp., javana-discolor group’; Jacobs 9592
a 5p.; NGF 28074 (K; Leiden det. ae
T, pedunculare)
WINTERACEAE
Zygogynum sp.; 14465
FAMILY INDET.; 14373, 14374, 14675
1554 BRIT.ORG/SIDA 20(4)
ACKNOWLEDGMENTS
The botanical surveys of the KWMA were supported by the Botanical Research
Institute of Texas, Harvard University, and the John D. and Catherine T.
MacArthur Foundation. | thank my associates at the Village Development Trust
lor their assistance with the Kamiali program, including the Hon. Sasa Zibe-
Kokino (VDT co-founder and Minister of the PNG Dept. of Environment and
Conservation), Aung Kumal (Executive Director), Karol Kisokau (Director of
Research and Conservation), John Sengo (Manager of Integrated Conservation
and Development), Bing Siga (Field Coordinator) and Madlene Kavua (Science
Officer). Hitofumi Abe (Ecosystem Research Group, University of Western
Australia) wrote the Japanese translation. John Pipoly (FTG) provided the Latin
diagnoses. Kipiro Damas (LAE) brought the new Trichadenia to my attention.
Parataxonomists Demas Ama and Ali Towati were my principal associates in
the field. Parataxonomist Nathan (no surname), also assisted with the collect-
ing until his recent and untimely death from acute cerebral malaria.
The following specialists provided determinations and/or shared informa-
tion on their respective groups: L. Craven (noncapsular Myrtaceae), H. Fortune
(Cunoniaceae), N.H.S. Howcroft (Orchidaceae), K.-L. I luynh (Pandanaceae), M.
Jebb (Aristolochiaceae, Lecythidaceae), J. Kessler (Annonaceae), A. Kjaer
(Arecaceae), D. Middleton (Apocynaceae), J. Pipoly (Myrsinaceae), S.S. Renner
(Melastomataceae), A. Schuiteman (Orchidaceae), PE Stevens (Clusiaceae,
Ericaceae, passim), W. Vink (Sapotaceae), E. de Vogel (Orchidaceae), and P. Wil-
son (capsular Myrtaceae).
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assessment report, vol. 1. PNG Dept. of Environment and Conservation, Boroko, Papua
New Guinea. Pp 121-26.
1560 BRIT.ORG/SIDA 20(4)
BOOK NOTICES
S.S. KHARKEVICH (chief ed.) and N.N. Tzvetev (vol. ed.). 2003. Vascular Plants of
the Russian Far East. Volume 1: Lycopodiophyta, Juncacaceae, Poaceae
(Gramineae). (ISBN 0-8093-2421-0, pbk.) Science Publishers, Inc. c/o Enfield
Distribution Co., 234 May Street, P.O. Box 699, Enfield, NH 03748, U.S.A.
(Orders: 603-632-7377, 603-632-5611 fax, http:/ WW w.scipub.net,
sales@scipub.net, complete catalog at info@scipub.net/catalog). $165.00,
506 pp, 26 plates of b/w is 162 maps, 6" x 91/2".
ist tskogo Dal Vostoka, 1985. There
This volume is actually a translation of Sosudistye r ¢
escr noe include name ad nomenclatural citation, Rus-
—
—_—
are keys to genera and oo ee d
5 chromosome number, and distribution by region foll owed by
|
sian name, morphol : p
‘vation. Running heads with fam-
o
ecology, s encene flowering time, economic uses,
ily/genus would have been nice; the index does work.
DOUGLAS J. FutuYMA, H. BRADLEY SHAFFER, and DANIEL SIMBERLOFF (eds). 2002. Annual
Review of Ecology and Systematics: Volume 33, 2002. (ISBN 0-8243-1433-6,
hbk; ISSN 0066-4162). Annual Reviews Inc., 4139 El Camino Way, P.O. Box
10139, Palo Alto, CA 94303-0139, U.S.A. (Orders: www.AnnualReviews.org,
800-523-8635, 650-493-4400, 650-424-0910 fax). $145.00 (USA), $150.00
(Int'l), 828 pp., 6" x 9".
Volume 33 of Annual Reviewof Ecology and Systematics contains a mix 25 articles: Saproxylic Insect
Ecology and the Sustainable Management of Forests; Conus Venom Peptides: Reflections from the
Biology of Clades and Species; Troubleshooting Molecular Phylogenetic Analyses; The Early Radia-
tions of Cetacea (Mammalia): Evolutionary Pattern and Developmental Correlations; The Mesozoic
Radiation of Birds; Plant Ecological Strategies: Some Leading Dimensions of Variation Between Spe-
cies, Reproductive Protein Evolution; The Causes and Consequences of Ant Invasions; Gulf of Mexico
Hypoxia, a.k.a. “the Dead Zone”, The (Super) Tree of Life: Procedures, Problems, and Prospects; Ho-
aissance of Community-Based Marine Resource Man-
mogenization of Freshwater Faunas; The Ren
Uv-B Radia-
agement in Oceania; Nutrient Cycling by Animals in Freshwater Ecosystems; Effects of
tion on Terrestrial and Aquatic meee Producers; The Evolution and Maintenance of Androdioecy;
Mast Seeding in Perennial Plants: Why, How, Where?; Disturbance to Marine Benthic Habitats by
Trawling and Hees iplieabions for Marine Biodiversity; Phylogenies and Community Ecology;
s in Plant Ecology; The Quality of the Fossil Record: oe ante . aeneaael
malaie idy in ae lants; Herbivore Offense; The Pacific Salm s: Wha
e of Recovering Species, Estimating namin Times a
able Isoto
Analyses; Neo
ence Brings to the Challen
ogenetic and pope Genetic Timescales; The Fate of Clades in a World of Recurrent
a
Data on Phy
Climatic eet Milankovitch Oscillations and Evolution.
volume has a Subject Index, Cumulative Index of Contributing Authors (vols. 29-33) and
"ac
a Index of Chapter Titles (vols. 29-33)
SIDA 20(4): 1560. 2003
NEW COMBINATION IN EURYBIA
(ASTERACEAE: ASTEREAE) FROM NORTH AMERICA
Luc Brouillet
Herbier Marie-Victorin
Institut a recherche en biologi ie végétale
Montréal
01 Sherbrooke a Montreal eae CANADA, H1X 2B2
email: luc. brouillet@umontreal.ca
ABSTRACT
An bination i l here f ifi in Eurybia from North America: Eurybia
lineal (Nutt) GL. Reson var. pulchra (S.F. Blake) Brouillet, comb. nov.
RESUMEN
1 c aps
Eurybia de Norte América: Eurybia
glauca (Nutt.) G.L. Nesom var. piilchiva (S.E Blake) Brouilles comb. nov.
INTRODUCTION
The North American genus Eurybia, segregated from Aster by Nesom (1994),
has now been shown convincingly to be distinct from the Eurasian Aster s.str.
(Noyes & Rieseberg 1999; Brouillet et al. 200la, b; Bastien & Brouillet 2002;
Semple et al. 2002). Nesom (1994, 1997) provided most of the combinations
needed in this genus. Nevertheless, during preparation of the treatments of this
genus for the Flora of North America project, differences in taxonomic con-
cepts from those suggested by Nesom (1994) have led me to propose a new com-
bination.
Eurybia glauca (Nutt.) G.L. Nesom
In his taxonomy of Eurybia, Nesom (1994) recognized Eurybia pulchra (SF
Blake) G.L. Nesom asa species distinct from Eurybia glauca (Nutt.) G.L. Nesom,
justifying it by the smaller leaves, more acute phyllaries and glandularity. Yet,
except for glandularity, which does not appear to vary gradually from one taxon
to the other, the other features all fit within the range encountered in E. glauca,
apart from a slight tendency toward larger reproductive features in E. pulchra;
leaf size ranges overlap considerably when a large sample of EF. glauca is con-
sidered. A morphometric study would be needed to determine the exact ampli-
tude and significance of size differences between the two taxa. The ranges of
the two taxa appear to be parapatric in southern Utah-northern Arizona, with
E. pulchra restricted to the vicinity of the Grand Canyon while E. glauca ranges
widely to the north and east into the Southern Rocky Mountains; it must be
SIDA 20(4): 1561-1563. 2003
1562 BRIT.ORG/SIDA 20(4)
noted, however, that populations of the latter are found in Arizona south and
east of the Grand Canyon. Nonetheless, no mixed population of the two taxa
has been reported. The restricted range and distinct glandularity justify that E.
pulchra be recognized as a taxon. Nevertheless, glandularity alone does not seem
to justify recognizing EF. pulchra asa distinct species. lagree with Nesom (1994)
that the inclusion by Cronquist (1994) of E. pulchra within FE. wasatchensis is
misguided as it is based only on the shared possession of glands. Several char-
acters distinguish E. pulchra and E.wasatchensis, including leaf glaucousness
and phyllary habit (appressed vs squarrose), which the former shares with E.
glauca. Therefore, | prefer to recognize the former asa variety of the latter, as i
done by Welsh et al. 1987), even though it was originally described as a a:
~—
species.
Eurybia glauca (Nutt.) G.L. Nesom var. glauca
Eurybia glauca (Nutt.) G.L. Nesom var. pulchra (S.F Blake) Brouillet, comb. et
stat. nov. BASIONYM: Aster glaucodes S.F. Blake subsp. pulcher S.F Blake, Proc. Biol.
Soc. Washington 35:174. 1922. Eurybia pulchra(S.F Blake) GL. Nesom, Phytologia
77:261. 1994.
The combination Aster glaucodes Blake var. pulcher (Blake) Kearney & Peebles,
reported in Welsh et al. (1987), is not valid because Kearney and Peebles (1960)
did not ma
—
<e a formal status change with proper reference to the basionym.
The name was thus never validly published at the varietal rank. These author
perhaps unwittingly referred to the infraspecific taxon asa variety instead of a
subspecies and might not have intended to make a new combination.
The origin of glandularity in E. glauca var. pulchra and E. wasatchensis, as
compared to the eglandular E. glauca var. glauca, could be explained by consid-
ering the other members of section Herrickia, to which these taxa clearly be-
long: Eurybia horrida (Wooton & Standl.) G.L. Nesom (formerly Herrickia
horrida) and Aster kingii (Brouillet et al. 2001a, b). All members of section
Herrickia are glandular except E. glauca var. glauca. It is more parsimonious to
explain the lack of glands in the latter by a loss of glands after speciation. This
hypothesis would imply that the presence of glands in var. pulchra is ancestral
within the species, ee that var. glauca may represent a relatively more recent
expansion of an eglandular phenotype to the range of the species as known
today, from a more restricted original area perhaps close to the San Juan river
basin where var. pulchra is restricted today.
—
lane |
ACKNOWLEDGMENTS
| would like to thank Guy L. Nesom (Botanical Research Institute of Texas) and
Kanchi Gandhi (Harvard University Herbaria) for their useful comments on
the manuscript. This project was supported in part by a research grant from
NSERC (Canada).
BROUILLET, NEW COMBINATION IN EURYBIA 1563
REFERENCES
Bastien, J.and L. Brouittet. 2002.Phylogeny of North American and Eurasian asters (Astereae:
Asteraceae) using coDNa and combined ITS-coDNA sequence data. Botany 2002 Meet-
ing, Madison, WI, Aug. 2-7, 2002. Abstract 636, p. 162.
Brouittet, L.,G.A.ALLEN, J.C. Semete, and M.l10.2001a.ITS phylogeny of North American asters
(Asteraceae: Astereae): basal grade to North American lineages and distinct from Eur-
asian ones. CBA/ABC Meeting,. Ketowna, B.C., June 23-27, 2001. p.31 (abstract).
Brouitter, L., G.A. Auten, J.C. Semete, and M. Ito. 2001b.ITS phylogeny of North American as-
ters (Asteraceae: Astereae). Botany 2001 Meeting,. Albuquerque, NM, Aug. 12-16, 2001.
Cronauist, A. 1994. Asterales. Vol. 5. ln: A. Cronquist, AH. Holmgren, N.H. Holmgren, J.L. Re-
veal, and PK. Holmgren, eds., Intermountain flora. New York Botanical Garden, Bronx.
Kearney, |.H. and R.H. Peestes. 1960. Arizona flora. Univ. California Press, Berkeley and Los
Angeles.
Nesom, G.L. 1994. Review of the taxonomy of Aster sensu lato (Asteraceae: Astereae), em-
phasizing the New World species. Phytologia 77:141-297.
Nesom, G.L. 1997. Taxonomic adjustments in North American Aster sensu latissimo
(Asteraceae: Astereae). Phytologia 82:281-288.
Noyes, R.and L.H. Riesesera. 1999.ITS sequence data support a single origin for North Ameri-
can Astereae (Asteraceae) and reflect deep geographic division in Aster s.|.Amer.J.Bot.
86:398-41 2.
Semete, J.C., $.B. Hearo, and L. Brouittet. 2002. Cultivated and native Asters of Ontario
(Compositae: Astereae). Univ. Waterloo Biol. Series 41.
WetsH, S.L., N.D. ATtwoop, L.C. Hicains, and S.Gooprich. 1987.A Utah flora. Great Basin Natural-
ist Memoir no. 9.
1564 BRIT.ORG/SIDA 20(4)
BOOK NOTICES
JAMes B.Pripps with Rosert J. OKENNON and Ron W. Lance. 2003. Hawthorns and
Medlars. (ISBN 0-88192-591-8, hbk.). Royal Horticultural Society Plant
Collector Guide. Timber Press, Inc. 133 S.W. Second Ave, Suite ee Portland,
OR 97204-3527, US.A.(Orders: wwwtimberpress.com, mail @ti press.com,
DUS22 (261.0, 1-000-32 79000, 505-227 - 3070 fax). $24.95, 180 pp., 75 color
photos, + b/w photos, 2 tables, 11 drawings, 3 maps, 7 x 9"
Hawthorns (Crataegus) and medlars (Mespilus) are in the rose family (Rosaceae). This 180 page book
is beautifully illustrated and adorned with excellent color photos from a variety of sources. The au-
thor brings together in one place a broad mix of information on these two groups of closely related
aie ae re are ie rhaps 140 species af Camara in the world”; c ae seven the book covers
| ilabl iapters include
ifr
am |
CALC
11 lawthorns and Medl: urs in 2 olklore; 2) Natural History ‘an Consaation: Structure and Repro-
ductive Biology; 3) Practical Value; 4) Cultivation; 5) Breeding and Selection; 6) Classification and
Identification; 7) Hawthors; and 8) Medlars and Intergeneric Hybrids. There are four appendices: 1)
Preparation of Herbarium specimens; 2) Checklist of Crataegus and Mespilus Species; 3) Tabular
Key to Hawthorns and Medlars; and 4) Conversion Charts. A Glossary, Bibliography, and Index com-
plete the book.—Barney Lipscomb, Botanical Research Institute of Texas, 509 Pecan St, Fort Worth, TX
76102-4060, U.S.A, barney@britorg
rs
PAUL L. COMEAU, YASMIN S. COMEAU, and WINSTON JOHNSON. 2003. The Palm Book of
Trinidad and Tobago including the Lesser Antilles. (ISBN 0-9740870-0-9,
pbk.). The International Palm Society, 6729 Peacock Road, Sarasota, FL 34242,
U.S.A. (Orders: The Allen Press, PO. Box 1897, Lawrence, KS 66044-8897,
US.A,, 785-843-1274 [credit card orders only| online order wwwpalms.org).
$39.95, 108 pp., numerous color photos, distribution maps, 7 1/2 x 10".
— nts.—Foreword, Preface, Ano eceme ae: 1) ae 2) Learning to Identify Native Palms
ntification Key lor the Palms), 3) Pal liar Distributions, 4) Palms of the Lesser Antilles
—
G . Nomenclatural Notes on the Palin Making Sense of Common Names, Palm Species aad
Their Synonyms, Known Introduced Palms, Photograph Data, References, and hi
e local name(s), the geogre ses al disibation visa eenoity
e) abhor
“Each species account incluc
(distribution map shows each species ean al rang
tinguishing features, reese with other similar ac habitat prelerence ollinarars anc adic:
tributors, economic uses and conservation statues.” Twenty-two species of palms are recognized, dis-
cussed in detail and photogre phe in their natural habitat.
SIDA 20(4): 1564. 2003
A NEW SPECIES OF BRAZORIA (LAMIACEAE)
FROM THE CENTRAL MINERAL REGION OF TEXAS
Matt W. Turner
Univ. of Texas at Austin
McCombs School of Business
1 Uni i nig/Seaton
Austin, TX 78712, U.S.A.
ABSTRACT
Brazoria enquistii M.W. Turner sp. nov. from the central mineral region of Texas (Mason, Llano and
Burnet counties) is described. It is similar to B. truncata (Benth.) Engelm. & A. Gray from which it
differs in having longer floral bracts with more pronounced ciliation, shorter internodes, and dis-
tinctions in the lobes of the calyx. Its habitat and edaphic preferences, unique to the genus, are also
discussed. Descriptions, photographs, dot-maps, and keys to species are Eee noe
RESUMEN
Se describe Brazoria enquistii M.W. Turner sp. nov, de la region mineral central de Texas (condados
de Mason, Llano y Burnet). Se asemeja a B. truncata (Benth.) Engelm. & A. Gray de la que se distingue
por tener las bracteas florales mas largas con ciliacion mas destacada, los entrenudos mas cortos, y
unas distinciones en los naan ce Ci an Su habitat y Piclerenclas enanae unicos en el sGerisEe)
también se discuten. Se of ;, fotos y de es}
Is I
As noted previously in my overview of the genus (Turner 1996), the only three
collections of Brazoria that were known to me from the central mineral region
of Texas prior to 1996 (from collections housed at BRIT, GH, K, LL, OKL, TAES,
TEX) possibly constituted an undescribed taxon, but lack of extant popula-
tions precluded their recognition. Subsequently, sightings of populations of the
taxon by Marshall Enquist in this region prompted three years of additional
fieldwork by myself. These studies have convinced me that the plant is
undescribed and is deserving of specific recognition.
TAXONOMY
Brazoria enquistii M.W. Turner, sp. nov. (Figs. 1, 2 & 3). Type: U.S.A. TEXAS: Mason
o.:ca. Ll airline mi ESE of Mason, off Lower Willow Creek Rd at crossing of Wil-
low Creek, 4 mi E of US Hwy 87, ll May 1996, M.W. Turner 64 (HOLOTYPE: TEX;
ISOTYPES: BRIT, GH, MO, TAES, US).
Brazoriae truncatae (Benth.) Engelm. & A. Gray var. truncatde similis sed racemis compactissimis
cum internodis 2.5-4.0 m; bracteis floralibus 8- oan, cum ciliis 2-3 mm; lobis inferioribus calycum
leniter apiculatis, plerumque habentibus unum “dentem” per lobum. Plantae indescriptae
conservandae sunt.
peel
Erect annual herbs mostly 15-40 cm high, simple, or branched at base; if latter,
branches immediately ascending from base. Stems square, pubescent below
SIDA 20(4): 1565-1571. 2003
1566 BRIT.ORG/SIDA 20(4)
inflorescence on all sides. Leaves opposite; lower leaves, if persistent, usually
oblanceolate (occasionally spatulate), 5-9 cm long, 1-2 cm wide, often tapering
toa long-winged petiole; margins undulate to denticulate; upper leaves oblong
to elliptic, denticulate, usually sessile and partly clasping. Mature spikes
racemoid, densely flowered, with lower internodes mostly 2.5-4.0 mm long;
{lowers decussate. Bracts at base of pedicels mostly ovate-orbicular, 8-12 mm
long; margins ciliate, with hairs mostly 2-3 mm in length. Calyces broadly cam-
panulate, 5-8(-10) mm long, bilabiate; the two lower lobes mostly truncate with
one “tooth,” woolly-tomentose at base, with many eglandular hairs reaching 2
mm in length; upper middle lobe usually truncate or shallowly mucronate;
upper lateral lobes mucronate to apiculate; capitate-glandular hairs scattered
over both lower and upper lobes. Corollas 15-25 mm long, bilabiate, pale laven-
der and usually distinctly veined, sparsely puberulent along the upper surface;
upper lip bifid; lower lip trifid, lobes usually emarginate; throats with annulus
usually perpendicular to axis of corolla 2-3 mm above ovary, maculate, with
distinct hump on lower surface beneath stamens. Stamens 4, fertile, the ante-
rior pair longer than the posterior. Filaments ca. | mm wide at base of free por-
tion, conspicuously winged, villous, arched inward near apices; upper pair free
from adnation to corolla for 3-4 mm; lower, innermost pair free for L.0-1.5 mm,
with mutually opposing “bulges” or “shoulders” that, together with overlapping
hairs, hold style in epipetalous position. Thecae apiculate laterally at base, dark
purple and ringed with white; upper pair slightly smaller than lower (ca. 1.2
mm vs 2.0 mm); ventral (rarely dorsal) surface of connective tissue between
thecae usually with several to many white hairs. Styles 10-15 mm long, bi-
lobed, glabrous; branches linear-lanceolate. Nutlets mostly 1.5-2.5(-3.5) mm
long, strongly trigonal, with incurved hispidulous hairs, rarely glabrous.
Distribution.—The most geographically isolated species of the genus,
Brazoria enquistii is known only from three counties—Burnet, Llano, and Ma-
son (Fig. 1)—and I have only been able to locate extant populations in the latter
two of these. Many attempts to ascertain distribution extensions (beyond what
is reported here) have been made in the surrounding vicinity over the past five
years to no avail. Although only ca. 130 km separates the taxon (Llano/Burnet
Co. border area) from the nearest populations of B. truncata (Bastrop Co.) (Fig.
2), the intervening distance presents the sand-loving genus with a barrier of
limestone, the central mineral region being renowned as a granitic outcrop
(which weathers to sand) surrounded by calcareous soils. This geographic iso-
lation, together with its infrequent occurrence and short blooming period, have
kept the taxon in relative obscurity.
Habitat.—Brazorid enquistii, unlike the eastern species of the genus which
tend to prefer the loose white sands of xeric slopes, grows almost exclusively
along granitic streambeds of the Llano River watershed. Only four of the 16
populations discovered by the author (sites of Turner 71, 72,95 and 96 ) are not
TURNER, A NEW SPECIES OF BRAZORIA FROM TEXAS 1567
Colorado River
McCulloch Co.
San Saba Co. Burnet Co.
Lats
~~
Scale
1 om = ca3 mi (ord B lum)
0 | 6 9 2 1s
lections known to date and cited herein.
along such streambeds (but even two of these are within 100 m of one). More
specifically, the species is rarely found near the water’s edge, but rather occurs
on the raised, sparsely-vegetated sand embankments within 10-20 m of adja-
cent streams. The species seems to prefer embankments along permanent wa-
ter, and is only rarely found in dry washes and gullies. One of the largest popu-
lations (estimated at 10,000+ individuals at the type locality) was discovered
ona large, flat, sand island in the middle of Willow Creek as it approaches the
Llano River. Indeed, the species seems to flourish in creek beds as they widen
downstream near their mouths (presumably because the waterways accumu-
late more sand), and becomes less abundant upstream as the creeks become
narrower. Only three of the 22 sites known to date (both historical and extant)
lie south of the Llano River.
Phenology.—All known collections are from the month of May.
Remarks.—Brazoria enquistii most closely resembles B. truncata var.
truncata. The most obvious distinguishing character of the taxon is its long
and well-developed bracts (see Fig. 3 and key following), which almost conceal
the developing buds, in stark contrast to the shorter bracts of B. truncata whose
buds, left exposed to view, display themselves in the compact rows that have
earned that species the common name of “rattlesnake flower.” Another obvi-
ous character is the internode length which is the smallest of the genus (2-4
1568 BRIT.ORG/SIDA 20(4)
KILOMETERS
150
| | |
Fic. 2. Distributi cp : + / eas n ).B.a ia (cl aes pe dp tee ta ( Pr ):B
enquistii (closed circles).
mm vs. 5-6 mm for B. truncata var. truncata, 8-13 mm for B.t. var. pulcherrima
(Lundell) M.W. Turner, and 8-20(-30) for B. arenaria Lundell) and provides for
an extremely compact spi
Less obvious characters distinguishing the taxon from B. truncata include
a longer bract ciliation (2-3 mm vs. 1-2 mm), a distinct shape to the lower lobes
of the calyx (truncate or with only | “tooth” vs. with 3-5 “teeth”), and a subtle
difference in the shape of the upper middle lobe of the calyx (truncate to shal-
lowly mucronate vs. mucronate to apiculate).
So far as known, the first recorded collection of the taxon was made in 1910
in Kingsland (Llano Co.), Hastings s.n.(LL). Some 20 years later, the second re-
corded collection is from an area six miles northwest of Valley Spring in Llano
Co., Bauman’s [sic] 6286 (TAES). Some 30 years elapsed before the third collec-
tion was made (in 1962 in Inks Lake State Park, Burnet Co., along Clear Creek,
ar
Ce
TURNER, A NEW SPECIES OF BRAZORIA FROM TEXAS 1569
Fic.3.Tt i finfl level i ies of B ja. Top row, AC: Brazoria enquisti.
hl ra] Ih fal L L L iwi rat ca.12?mm):
J
J | 4 + h * 4
i: kK c . .
sides: Bottom row, D- F: Brazoria eupeata var. truncata. a stare fra brat hidden (cf. “attlesnake flower,” the com-
mon name for this species); E, bracts do not (or y) subt ; g y f
1570 BRIT.ORG/SIDA 20(4)
Correll & Ogden 25300 (LL)). An extant population has been found at a loca-
tion approximating the second collection (Turner 69); however, the author's re-
peated attempts at locating living populations in the Kingsland area and in
western Burnet Co. have been unsuccessful. The Kingsland area may have be-
come overdeveloped and/or the damming of the Colorado River for Lake LB]
(in 1951 with the Alvin Wirtz Dam) may have influenced the habitat. Repeated
visits to Inks Lake State Park, with special attention to Clear Creek and Spring
Creek, have likewise revealed nothing. A 1996 sighting by Marshall Enquist from
the Llano/Burnet Co. line area (near Buchanan Dam), prompted this investiga-
tion, but local roadwork, immediately thereafter, completely altered the site and
no further sightings have been made in the area to date. At the present, the west-
ern edge of Burnet Co. represents the eastern limit for the species.
Brazoria enquistii is the most geographically restricted taxon of the ge-
nus, distributed primarily within just two counties. Like B. truncata var.
pulcherrima, which is restricted to about five counties in northeastern Texas
(Turner 1996), B. enquistii can be locally abundant, which would likely pre-
clude it from endangered species status. On the other hand, it is more infre-
quent in occurrence, tending to appear in isolated patches (due to its special-
ized habitat), rather than being loosely scattered along the roadside. The number
of individuals at a given location can vary sharply from year to year: the type
locality mentioned above contained over 10,000 plants in 1996, but only about
% of that number in 1999.
—
—
Choice of rank.—The taxon is here treated at specific rank for the following
reasons: it exhibits several distinguishing morphological characters (outlined
above); it has a habitat and edaphic preference distinct from the other taxa of
the genus (raised embankments of streambeds in purely granitic sand vs. xeric
slopes of loose, white, non-granitic sands in open woodlands and stabilized
dunes); it is geographically and edaphically isolated by the surrounding cal-
careous soil, which erects a barrier to gene exchange; and no populations inter-
mediate to its related taxa have been discovered.
Etymology.—The species is named for Marshall Enquist, an attorney in the
Austin area, who called to my attention extant populations in the central min-
eral region, without which this paper would not have been possible. Mr. Enquist’s
excellent floristic guide (Enquist 1987), as well as his active research on native
Texas flora Cincluding Anemone, Crataegus, Desmodium,and Prunus), have been
an inspiration to botanists and naturalists alike.
Specit |: U.S.A. Texas. Burnet Co.: Inks Lake State Park along Clear Creek, 18 May 1962,
Correll & Ogden 25300 (LL). Llano Co.: Kingsland, 12 May 1910, Hastings s.n. (LL); 6 mi NW of Valley
Spring, 8 May 1933, Bauman’s|sic] 6286 (TAES), ca.6 mi WSW of Llano on FM 152 at Six Mile Creek. 1]
May 1996, Turner 60(TEX); ca. 18 mi due W of Llanoon TX 29 in Weide Creek. 14 May 1996, Turner 61
(TEX); Castell, 19 May 1991, Enquist 1934 (TEX); In Castell at Llano River crossing on FM 2768, 14
May 1996, lurner62 (TEX); ca. 6 airline mi due W of Valley Spring, on CR 405, 2.1 mi Sof TX 7Lat San
TURNER, A NEW SPECIES OF BRAZORIA FROM TEXAS 1571
Fernando Creek crossing, 18 May 1997, Turner 69 (TEX), ca. 2.5 mi E of Llano,on FM 2241, 15 mi NE
of jct TX 29, along Wrights Creek, 16 May 1999, Turner & Robbins 97 (TEX). Mason Co.: ca. 0.5 mi W
of Castell on TX 152 in creek bed, 14 May 1996, Turner 63 (TEX), ca. LI airline mi W of Llano, off CR
404 at San Fernando Creek crossing, 19 May 1996, Turner 65 (TEX), ca. 7 airline mi ESE of Mason, off
Art Hedwigs Hill Rd, 3 mi N of US Hwy 87 & L6 mi S of TX 29 along Willow Creek, 18 May 1997,
Turner 68 (TEX): 2.7 mi S of Pontotoc (jet TX 7D on Hickory Grove Rd, along San Fernando Creek, 18
May 1997, Turner 70 (TEX); 2.2 mi Sof Pontotoc (jet TX 71) on Hickory Grove Rd, 18 May 1997, Turner
71 (TEX); Hillis Fly Gap Ranch, 2.5 mi W of Pontotoc on TX 71, then 3.1 mi Son Fly Gap Rd, then 0.2
mi W toentrance, 12 May 2002, Hillis 286 (TEX); 2.0 mi Sof TX 7Lon Fly Gap Rd at creek crossing, 18
May 2002, Hillis 293 (TEX); ca. 14 airline mi ENE of Mason on Hickory Grove Rd, 2.5 mi N of jet with
Union Rd, 18 May 1997, Turner 72 (TEX); ca. 12 mi due F of Mason on Union Rd, 0.6 mi N of TX 29 at
crossing of Martin Creek, 10 May 1998, Turner 79 (TEX), ca. 1] airline mi ENE of Mason, on Eaton
Cemetery Rd, 1 mi E of FM 1900 at crossing of Martin Creek, 10 May 1998, Turner 80 (TEX); ca. 8 mi
ENE Mason, on N Art Rd, 2.4 mi N of jct TX 29,75 m N of creek (branch of E Willow Creek), 16 May
1999. Turner & Robbins 95 (TEX): ca. 8 mi ENE Mason, on N Art Rd, 2.8 mi N of jet TX 29, 16 May 1999,
Turner & Robbins 96 (TEX).
A REVISED KEY TO THE SPECIES OF BRAZORIA
1. Floral bracts ciliolate, with hairs 0.1-0.2 mm in length; mature plants often basally
branched with branches recumbant at base; basal leaves usually persistent, oblan-
ceolate to spatulate; southern Texas
- Floral bracts ciliate, with hairs 1-3 mm in length; mature plants often not basally
ranched, or, if so, with branches immediately ascending from base; basal leaves
rarely persistent, usually oblong to oblong-oblanceolate; central, and southcentral
to northeastern Texas.
Floral bracts at base of pedicels mostly 8-12 mm in length; mature spikes very
densely flowered, with lower internodes mostly 2.5-4.0 mm long; lower lobes of
calyces shallowly apiculate with mostly one “tooth per lobe; central mineral re-
gion (Burnet, Llano, Mason Co.) B. enquistii
Floral bracts at base of pedicels mostly 4-8(-9) mm in length; mature spikes less
densely flowered, with lower internodes mostly 5-13 mm long; lower lobes of
calyces dentate with mostly 3-5 “teeth” per lobe; not in central mineral region
B. arenaria
—
N
N
ex
SS B. truncata
ACKNOWLEDGMENTS
Lam grateful to the Directors and Curators of the following institutions for the
loan of approximately 450 herbarium specimens: BRIT, GH, K, LL, OKL, TAES,
TEX. | would also like to thank Gayle Turner for assistance with the Latin diag-
nosis, David Hillis for bringing to my attention additional locations of B.
enquistii near his ranch, and Tom Wendt and Lindsay Woodruff for the use of
the Plant Resources Center (TEX-LL) and for the steadfast care and kindness
that they have extended to me in answering my many questions.
REFERENCES
Enouist, M. 1987.Wildflowers of the Texas hill country. Lone Star Botanical, Austin, TX.
Turner, M.W. 1996. Systematic study of the genus Brazoria (Lamiaceae), and Warnockia
(Lamiaceae), a new genus from Texas. PI. Syst. Evol. 203:65-82.
1572 BRIT.ORG/SIDA 20(4)
BOOK NOTICES
tA | LT ee |
CHRISTOPHER NEWsHOLME. 1992 (Paper back reprint 2003). Willows: The Genus Salix.
(ISBN 0-88192-565-9, pbk.). Timber Press, Inc. 133 S.W. Second Ave, Suite
450, Portland, OR 97204-3527, U.S.A. (Orders: www.timber press.com,
mail@timberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax).
$19.95, 256 pp., 65 color photos, numerous line drawings, 6 x 9",
Contents.—Color Hlustrations, Acknowledgments, Preface, Introduction, General Characteristics ol
Willows, Management and Cultivation, Ornamental Trees and Shrubs for | -arge Gardens, Parks and
Estates, Ornamental Trees and Shrubs for Small Gardens, Dwarf and Ground- hugging Alpine Spe-
cies for Rock Gardens, Glossary, Bibliography, General Index, and Botanical Index.
IAN HALL, STEVEN STEPHENSON, PETER BUCHANAN, WANG Yun,and ANTHONY COLE. 2003.
Edible and Poisonous Mushrooms of the World. (ISBN 0-88192-586-1, hbk.).
Timber Press, Inc. 133 S.W. Second Ave, Suite 450, Portland, OR 97204-3527,
US.A.(Orders: www.im berpress.com, mail@timberpress.com, 503-227-2878,
1-800-327-5680, 503-227-3070 fax). $39.95, 372 pp, 253 color photos, 6 color
illustrations, 3 tables, 6 x 9",
This valuable book includes detailed descri iptions, keys to help identify many common and impor
tant mushrooms, edible and not so edible. The authors offer suggestions and instructions on critical
identification, mushroom toxins and guidelines to follow when mushroom hunting. The excellent
r - harry - 1 eolorfiil
SANDRA KNAPP. 2003. Solanum Section Geminata (Solanaceae). (ISBN 0-89327-
441-0, pbk.). Flora Neotropica Vol.84. The New York Botanical Garden Press.
200" Street & Kazimiroff Boulevard, Bronx, NY 10458-5126, US.A.(Orders:
718-817-8721, 718-817-5126 fax, http://wwwanybgorg, ee
$60.00, 404 pp., 204 figs. (b/w photos, line draw ings, maps), 7 x LO"
Contents.—Abstract/Resumen, Introduction, History of Classification of Solanum, Nomenclatural
and Taxonomic History of Section Geminata, Mor phology, Ecology and Natural History, Species
Groups and Species Concepts, Taxonomic Treatment, Doubtful Names, Excluded Names. Acknowl-
edgments, Literature Cited, Numerical List of Taxa. F xsiccatae, Index to Scientific Names, and
Appendices.
he 126 species plus four insufficiently known taxa are discussed in context with their puta-
ryms, description, chromo-
tive relationships.” Each taxonomic entry includes nomenclature, syl
some number Gf known), illustration (line drawing/b/w photo), acai bution (map included), se
d, local names (if applicable) and general comments on rel: atonchinara to
lect ed spec ime ns ex amin >,
other taxa, morphology and natural history, This isa m ajor accomplishment for the Flora Neotropica
series.
SIDA 20(4): 1572. 2003
TWO NEW SPECIES OF LIATRIS SERIES GRAMINIFOLIAE
(ASTERACEAE: EUPATORIEAE) FROM THE
SOUTHEASTERN UNITED STATES
Robert Kral and Guy L.Nesom
Botanical Research Institute of Texas
09 Pecan Street
Fort Worth, TX 76102-4060, U.S.A.
ABSTRACT
Two species of Liatris series Graminifoliae are described for the first time and illustrations and dis-
tribution maps are provided for both. Liatris savannensis Kral & Nesom, sp. nov, is similar to L.
gra bee but differs in its larger heads with more numerous florets and broader, erose-lacerate
ry margins. It occurs on the western coast of peninsular Florida, south of the known ae of
L. graminifolia. Liatris patens Nesom & Kral, sp. nov, is similar to L. gracilis in its consistently pe-
dunculate heads and hirtellous-strigose stems and peduncles but differs in longer peduncles and
larger heads with more numerous florets; it is similar to L. graminifolia in head size and number of
florets but differs in its hairy stems and peduncles and longer peduncles. Liatris patens occurs in
southeastern Georgia and adjacent South Carolina, where it apparently is allopatric in range with its
close relatives L.graminifolia and L. gracilis.
RESUMEN
Se describen por primera vez dos especies de Liatris serie Graminifoliae ¥ se aportan ilustraciones y
mapas de distribucién para ambas. Liatris savannensis Kral @ Nesom, sp. nov., es similar a L.
eceunyong pete se diferencia pe sus cabezuelas mas grandes con flosculos mas numerosos y
y rados. Aparece en la costa oeste de la peninsula de Florida,
Oo
| del 4 ida de L.graminifolia. Liatri ‘ & Kral, sp. nov, es similar a L. graci-
lis por sus cabezuelas peduncul adas, y tallos y aie ulos hirto-estrigosos pero se diferencia =
tener pedtinculos mas largos y cabezuelas mas grandes con muchos mas flésculos; es similar a |
nae por el tamano de las cabezuelas y numero de fldsculos pero se diferencia por sus tallies
el y pedtnculos pelosos y mas largos. Liatris patens aparece en el sureste de Georgia y parte
adyacente de Carolina del Sur, donde aparentemente es alopatrica con sus parientes cercanos L.
graminifolia y L. gracilis
The species of Liatris series Graminifoliae (Gaiser 1946) are characterized by a
pappus of barbellate bristles, corolla tubes pilose within, phyllaries nearly flat
to rounded, obtuse, or acute at the apex, and sessile or pedunculate heads with
3-17(-20) florets. Gaiser recognized five species in series Graminifoliae: L.
graminifolia Willd., L. helleri Porter, L. gracilis Pursh, L. regimontis (Small) K.
Schum., and L. turgida Gaiser. Stucky (1991, 1992) and Stucky and Pyne (1990)
recently clarified concepts related to several related taxa, reviving use of the
name L. virgata Nutt. (to include L. regimontis and L. graminifolia var. smallii
(Britt.) Fern. & Grisc.), and adding L. cokeri Pyne & Stucky to the group. Liatris
gholsonii L. Anderson, an endemic of the Apalachicola River bluff system in
SIDA 20(4): 1573-1583. 2003
1574 BRIT.ORG/SIDA 20(4)
the Florida panhandle and close relative of L. gracilis, was recently described
(Anderson 2002). The present study adds two more species to this group—the
first endemic to a small region on the western coast of peninsular Florida, the
second known from a broader area of southeastern Georgia.
In the fall of 1964, during a field study of slash pine-sabal palmetto sa-
vanna adjacent to brackish marshland in the vicinity of Punta Gorda, Florida,
the first author collected an unusual Liatris. Additional collections were made
ne next 15 years,
es,
from other sites in two adjacent counties over the course of t
but a final determination of its identity was not made and other work inter-
vened. After +0 years, its description is presented here
Liatris savannensis Kral @ Nesom, sp. nov. (Fig. 1, 3). Type: UNITED STATES.
FLORIDA. Charlotte Co: wet pine/cabbage palm flats, ca. 3 mi S of Punta Gorda, 7
Oct 1979, R. Kral 64559 (HOLOTYPE: US; ISOTYPES: FSU, NY, VDB, VSC).
1a . ot {] 5 5 i |
nw}
shyllariis marginibus
se ae similis, s
lati litt eT, spicatde etms sed differt caulibus gli indulosis, foliis manifeste
punctatis mar bus ciliatis, et tubis corollarum interne pilosis.
i fo)
Plants perennials, corms ranging from globose to irregularly cylindric, narrowly
ovoid, or fusiform and caudex-like. Stems stiffly erect, (80-)90-130 cm tall,
simple or (rarely) ascending-branched, terete, irregularly multicostate and shal-
lowly grooved, glabrous but (including inflorescence axis) often sparsely but
evidently sessile-glandular. Leaves in high spiral, the oldest approximate, mostly
broken away by anthesis, but, if present elongate (-30 cm), essentially narrowl y
linear-lorate, those immediately above on stout internodes to 5 cm or more,
pandurate, gladiate-lorate, (17-)20-40 cm long, with broad, clasping-based,
intermittently pilose-ciliate petiole bases, gradually narrowed, then dilating to
linear-elliptic, linear-spatulate or linear oblanceolate blades 3-10 mm wide, the
petiolar nerves convergent to few laterals (ultimately no laterals), the median
nerve broad, sulcate or compound, apex variously acute, callused, margin en-
tire, shghtly thickened, surfaces pale green with darker glandular punctation,
glabrous except for some intermittent pilosity along abaxial midrib; upstem
leaves gradually shortening, narrowing in tighter spiral, sessile, mostly ascend-
ing, proximally pilose-ciliate, transitional to inflorescence bracts, the lower-
most exceeding, the uppermost exceeded by, heads. Inflorescence mostly de-
terminate, spiciform, rarely branched, its axis typically at least as au as the
leafy part of the plant, the heads sessile or on peduncles mostly 1-2 mm long.
Heads spreading- ascending, turbinate-campanulate; involucre just past anthe-
sis (7-)8-10 mm high from base to involucral apex, (8- )9-11(-12) mm wide
(pressed); phyllaries ciliate, in (3-)4(-5) series, the lowermost shortest, |.5-3 mm,
ovate to oblong, fleshiest, greenish, glandular-punctate, 3-nerved, progressively
longer inward, 4-6 mm, becoming obovate, then spatulate 7-9(-10) mm,
apically broadly rounded, witha hyaline, pinkish-purple, erose to lacerate bor-
KRAL AND NESOM, NEW SPECIES OF LIATRIS FROM SOUTHEASTERN US 1575
tt i. ome ie ral rs \ : I { 5 a | 1 fh } 1 ae | by ae eh ae RE
Fic.1 ype).a. Habit, (rig
view. b. Midstem.c. Infl (left), subtending bract, at ght).d. Head, with subtendi g bract.e. Phyl-
laries of ies f t (left) toi i ht). f. B f midcauline leaf, abaxial side. g. Adaxial
£ P| Ti h ee 12 i £] llafte\. £1 +h ce futinhe)
view of sp I;
1576 BRIT.ORG/SIDA 20(4)
der (0.2-)0.4-1 mm wide, intermittently ciliolate, the dorsal areas of longer
bracts greenish to reddish-green or purple, distinctly glandular-punctate, with
a sharp, low, rounded buttress delimiting the hyaline border. Florets 9-16(-17):
corolla tubes at anthesis yellowish or pale with pinkish-purple lobes, sessile-
glandular, tubular portion 5 mm, lobes 5, spreading, triangular, 2 mm long; in-
ternally pilose in the region of filament insertion witha broad annulus of whit-
ish, crisped hairs. Achenes cylindric, base slightly narrowed, apex with a short
neck at juncture with pappus rim, ca. 4 mm long, blackish, with ca. 10, paler,
raised ribs, densely hirtellous on the ribs, sparsely so on the intervals; pappus
mostly 5 mm long, bristles 30-40, pale purple-brown distally, strongly
barbellate.
Etymology and habitat.—The epithet “savannensis” alludes to the savanna
habitat of the species. The habitats are characterized as open sandy flats, usu-
ally moist or wet sites, dominated by slash pine, sabal palmetto, or a mixture of
the two. Collections of flowering plants have been made mostly from mid-Sep-
tember through mid-October. Flowering less commonly may begin in August
and extend into November.
Additional collecti ined. UNITED STATES. FLORIDA. Charlotte Co.: S side of Punta Gorda
on sandy peat of ms a flatwoods bordering brackish marsh, 21 Sep 1964, Kral 22919 (VDB, 2
luplicates distributed as L. spicata); sandy palmland by brackish marsh, E side of Punta Gorda, 21
Sep 1973, Kral 51978 (I ve GH, MO, TEX, VDB); wet grassy clearings in pine-palmetto flats by US
Hwy 41, L mis of Punta Gorda, 7 Oct 1979, Kral 64561 (FSU, GH, MICH, MO, NY, TEX, VDB): abun-
dant in marshy lands, each side of RR tracks, within city limits, $ side of Punta Gorda, 29 Sep 1961,
Lakela 24684 (BRIT, FLAS, USF). Hillsborough Co Sera River State Park, 20 mi NE of Tampa
on US 301, 18 Oct 1977, Arcuri 587 (USF), along Fla. H y 581, ca. 5 mi N of Tampa, roadside, 6 Oct
1977, Funk 48 (USF); along SR 581, ca. 5 mi N of ae 6 Oct L977, Gregory 18 (USF); margin of flat,
open Saas with scattered palmetto, N of Tampa Air Port, Kennedy Road, 13 Sep 1961, Lakela
24637 (USF): flat conta palmetto, and along boulevards NW of International Airport, Tampa,
24 Sep i Lakela 24670 (BRIT), margin of pineland and palmetto, along US 301, ca. | mi So
Hillsborough River State eo NE of Tampa, | Oct 1963, Lakela 26508 (FLAS, USF); frequent along
SR-581, ca 5 mi N of Tampa, 6 Oct 1977, L HOS 30 (U SF); Rocky Creek, disturbed site near Rocky
Creek and Smear esee Hwy, 7 Oct 1977, Mo 1 Giesy 169 (USB); disturbed area at edge of oak
woodland along SR-581, ca. 5 mi N of Tampa, 6 Oct 107, Willett 32 (USF ), flat ground edge of pine
flatwoods, 5 mi N Fletcher Avenue on 30" Street, Tampa, [Oct 1977], Winkler 36 (USF); Flatwoods
Park, 2.5 mi NW of Morris Bridge, ca. 7 mi E of Lutz, pine flatwoods, 22 Oct 1994, Wunderlin 10618
USF). Lee Co.: Business Rte. 41, N of Ft. ere in front of ae library, ine oe 7 Nov 1985,
Brown s.n. (USF); [no specific locality], | Aug 1933, Buswell s.n. (USF). Manate 2B Bradenton, dry
sand hills, Palma Sola, 19 Aug 1945, Gaiser, Butts,and Arnold s.n.(F 4 AS). Sarasota Co.: T. Mabry Carlton,
|
Jr. Memorial Reserve, pine fl 5 between wet prairies, grassy area Siiounded ey saw palmetto,
13 Oct 1997, preane 97-34 (USF): T. Mabry Carlton, Jr. Memorial Reserve, disturbed a parallel
to So. FPL grade, 997, Edmondson 97-35 (USF): sandy peat o open pine: pi aus sammock atN
side of Sort ie 1979, Kral 64571(+4 duplicates,
ginal to flat pinelands, 10 mi S of Venice, US 41, with Monarda, Coreopsis, and Rhuscopallina 20 Sep
1961, Lakela 24648 (FLAS, USF).
Two species of series Graminifoliae grow in the general area of Liatris
savannensis. Liatris gracilis occurs throughout Florida but grows in distinctly
KRAL AND NESOM, NEW SPECIES OF LIATRIS FROM SOUTHEASTERN US 1577
drier, sandier, more upland ecological systems; it differs from L. savannensis in
its hairy stems and peduncles, smaller, consistently short-pedunculate heads,
fewer florets per head (3-6(-7)), and shorter achenes. Liatris graminifolia is more
similar to L. savannensis in vestiture and head size, but the geographic range of
L. graminifolia apparently extends southward only to the vicinity of Levy
County, apparently separated by roughly 150 kilometers from the nearest
known population of L. savannensis in Hillsborough County.
Liatris savannensis closely resembles L. spicata (L.) Willd., which occurs
throughout Florida, and has commonly been identified as such. The two are
similar in their tall stems, narrow leaves, essentially glabrous vestiture, simi-
lar-sized heads in a spicate inflorescence, and phyllaries often with a broad,
scarious, pinkish rim. The first author identified his initial collections as L.
spicata, but in 1968, going back to the material and (finally!) opening a corolla,
noted pilosity within it,an annulus of crisped hairs at the level of the filament
bases. We follow Gaiser (1946) in weighting the presence of this pilosity as an
indicator of relationship (Nesom in prep.), with the implication that L.
savannensis isa member of series Graminifoliae, and observe that similarities
between L. savannensis and L. graminifolia also are strong.
The following pair of couplets outline significant morphological differences
first between Liatris savannensisand L. spicata and then between L. savannensis
and L. graminifolia.
couplet L:
. Stems minutely sessile-glandular; leaves prominently punctate, proximal margins
sparsely but distinctly ciliate with hairs 1-2.5 mm long; corolla tubes pilose within
at the region of filament insertion; florets 9-16(-17) per head. Liatris savannensis
. Stems egletigulet! leaves not ae naa or if so, then inconspicuously, margins eciliate
with a few scattered hairs 0.3-0.8 mm long; corolla tubes glabrous within; florets
(5-)6-10(-14) per head. Liatris spicata
ae et 2:
. stems minutely sessile-glandular;involucres (7—)8-10 mm high, (8-)9-11(-12) mm
de; inner phyllaries spatulate, 3.5-4 mm wide, hyaline, distinctly erose to lacerate
ie (0.2-)0.4-1 mm wide; florets 9-17 per head; achenes 4—4.5 mm long. Liatris
savannensis
. Stems eglandular; involucres 6-8(—9) mm high, 6—7.5 mm wide; inner phyllaries
oblong to obovate or slightly spatulate, 1-2.2 mm wide, with hyaline, relatively
smooth-edged borders 0.1-0.3 mm wide; florets 6-10 per head; achenes 3-4 mm
long. Liatris graminifolia
—
—
Other differences exist between Liatris savannensis and L. graminifolia where
comparisons are less diagnostic. Stems of L. savannensis range taller and leaves
are longer. Corms of L. savannensis range from globose to elongate; we have ob-
served only globose corms in L.graminifolia. The basal and lower cauline leaves
of L. savannensis are essentially strap-shaped Clorate), usually with little in-
crease in width distally, while similar leaves of L. graminifolia almost always
1578 BRIT.ORG/SIDA 20(4)
Sh ee
SS eae
Wy)
y
%
ale
Y
;
Fic 7 ae . (£. Vv, I1cocordc\ a Habit 1 Ji L iva Fa | + h Wl 1 £, re ££ ot eee |
} / U P rr vt
rescence base. c. Middl f infl d. Basal leaf, abaxial view (left), adaxial view (right). e. Base of
ack lane whacked (below), adaxial ( \f£B f medi line leaf, abaxial side (left), a lower bracteal
leaf (right). g. Head in anthesis. i. A recently fertilized floret. j. Anther tube with 1 style branches. k. A spread
mature corolla, inside view.
KRAL AND NESOM, NEW SPECIES OF LIATRIS FROM SOUTHEASTERN US 1579
show a distinct petiolar portion in the lower 1/4-1/2 of the total leaf length,
distinctly widening into a ‘blade’ portion. Heads of L. savannensis are consis-
tently sessile or on shorter peduncles (peduncles 0-2 mm long) compared to
slightly longer ones, on average, in L. graminifolia (peduncles 0-3(-7) mm long).
In addition to differences in overall phyllary size and margin, the medial, green,
glandular-punctate portion of L. savannensis phyllaries is consistently raised
above the hyaline border by a low, rounded ledge similar to that commonly
seen in the large heads of L. scariosa (L.) Willd., which is perhaps closely related
to ser. Graminifoliae.
Liatris graminifolia occurs over a wide area and in various habitats (espe-
cially northward), but meaningful infraspecific taxa are difficult to recognize,
and comparisons to it in the present study refer to the species as a whole. South-
ern elements of L. graminifolia have phyllaries that are distinctly obovate to
subspatulate, similar to those of L. savannensis, in contrast to oblong phyllar-
ies, which are characteristic of plants from the northern part of the range of
the species (roughly South Carolina and northward, see Fig. 3). The southern
plants have been recognized as L. graminifolia var. elegantula (Greene) K. Schum.
(the type from Alabama), but intergradation in phyllary morphology occurs
over a wide area and oblong phyllaries are found even in some southern popu-
lations. Some descriptions of L. graminifolia have attributed more variability
in vestiture and floret number to the species than we do, but it seems likely
that these descriptions included the two species recognized and segregated for
the first time in the present study.
A second previously undescribed species of Liatris has been recognized
primarily on the basis of herbarium studies connected with preparation of a
taxonomic treatment of the genus for the developing Flora of North America
(FNA) volumes.
Liatris patens Nesom & Kral, sp. nov. (Fig. 2, 3). Type: UNITED STATES. GEORGIA.
Laurens Co. drier margins of wet sandy meadow 9 mi E of Dublin, 300 ft elev., 19
Oct noe A. ee 4876 (HOLOTYPE: BRIT).
Liatrig Cc] 1
iaet
is, differt pedunculis
longioribus « et -caulibus ac pedunculis hirteli “strigosis Liatri he ans capitaus consistente
liffer
ones flosculis coc :
Plants perennial, arising from a globose to depressed-globose corm 7-14 mm in
diameter. Stems stiffly erect, 35-95 cm tall; stems and peduncles moderately to
densely hirtellous-strigose with antrorsely curved, ascending-appressed, mul-
ticellular white hairs 0.2-0.4 mm long. Leaves punctate-glandular, glabrous
except sparsely ciliate along 1-5 mm of the proximal margins; basal and lower
cauline narrowly oblanceolate to linear-oblanceolate, 9-18 cm long, narrowed
toa petiolar portion 1/4-1/2 the leaf length, the broader (blade) portion 2-4(-
1580 BRIT.ORG/SIDA 20(4)
: Ly Pd “S
mL
ny
@ Liatris savannensis ATs
a fe Sp
A Liatris patens we | ;
“et T CT
eo,
A
Fic 3.6 eee I . fyi . * ty;
set
7)mm wide, gradually or abruptly becoming much shorter and linear, upper
cauline l-2 cm long, ca. 0.5 mm wide, grading into inflorescence bracts. Inflo-
rescence racemose, determinate, forming a loose column (10, 20-)25-40 cm long,
3-6 cm wide. Heads erect or nearly so, on divergent-ascending peduncles |-2.5(-
3) cm long, peduncles initially diverging from the inflorescence axis at angles
of 45-90 degrees, subtended by a linear bract 5-12 mm long, minutely subu-
late-bracteate, rarely (especially above damage and mostly on lower peduncles)
1-2-branched; involucres turbinate-campanulate, (5-)6-7.5 mm high, 5-7 mm
wide (pressed); phyllaries in (2-)3-4(-5) series, strongly graduate in length, erect,
KRAL AND NESOM, NEW SPECIES OF LIATRIS FROM SOUTHEASTERN US 1581
nearly flat, green with an evident midvein, broadly obovate to broadly oblong-
obovate, apex rounded to nearly flat, innermost 1.5-2.5 mm wide, (5-)6-7.5 mm
long, outermost mostly ovate, ca. 1/5-1/3 as long as the inner, all evenly punc-
tate-glandular, outer sparsely strigose-villous with white hairs like those on
the peduncles, inner glabrous, margins hyaline and pinkish-purple (like the
corollas), 0.1-0.3 mm wide, even to slightly erose, usually densely ciliate. Flo-
rets 7-12; corollas pinkish-purple, sessile-glandular, tubular portion 4-5(-6) mm
long, lobes 5, spreading, narrowly triangular, 2-2.5 mm long; internally pilose
in the region of filament insertion with a broad annulus of whitish, crisped
hairs. Achenes obtriangular in outline, slightly flattened, with ca. 10 nerves,
2.5-2.8 mm long, densely but weakly hirsute on nerves and intervals; pappus
5-6 mm long, usually equal or longer than the corolla tubes, bristles strongly
barbellate, distally purplish.
Etymology and habitat.—The epithet “patens’ alludes to the spreading, rela-
tively long peduncles characteristic of the species. Collections have been made
in sandy pinelands of southeastern Georgia and adjacent South Carolina, ap-
parently mostly commonly in moist sites. Known collections have been made
in late September through November.
Additional collections examined. UNITED STATES. GEORGIA. Bryan Co.: Fort Stewart Military ne
ervation, Training Area C18, N side of Fort Stewart Road 58, ca. 1.8 mi E of ict with oe Rte. 6
occasional on disturbed sandhill, 5 m elev, with Vaccinium myrsinites, Quercus geminata, 25 —
1992, Zebryk 672 (VDB). Bulloch Co.: 4.5 mi NW of Portal in open wooded area, 12 Oct 1962, Boole
1311 (BRIT, TEX, US). Candler Co.: sandy add See — above bog by hiway, ca. 3 mi NW of Aline,
9 Oct 1975, Kral 56936 (VDB). Charlton Co.: plentiful in open disturbed area [along] Ga 177, 13 mi NE
of Edith, 8 Oct 1966, D’A o 1223 (VDB). McIntosh Co.: sandy flatwoods along US 17, near Middleton
Church at Eulonia, 20 Oct 1987, Angerman s.n. USCH); clearcut woods on north side of Eulonia, 20
Oct 1987, Angermans.n. (USCH). Pierce Co.: sandy rise in pine flatwoods, 10 mi N of Bristol on Ga. 1
11 Oct 1991, Kral 79978 (VDB): i plan ae by US 301, S side of Jesup by Landmark Motel, 9 ae
1975, Kral 59916 (VDB). Screven ni a. 2 mi from Blue Springs near a “Carolina Bay,” 10 Oct 1959,
Boole 997 (BRIT). Tattnall Co.: ca. 5 mi oa er eee County line on Route 280 from Claxton,
17 Oct 1959, Boole 1018 (BRIT). Wayne Co.: sandy er . pine-palmetto flats, by US 341, 2 mi NW of
Everett, | Oct 1976, Kral 59205 (VDB). SOUTH CAROLINA. Beaufort Co.: N of Victoria Bluffs Heri-
a STEEN, near Waco Bluffs development, N . Rte 218, N of Ebaons ae alone roadside
through I | tt spodosol soils, + Nov 2001,
Mesil n 16055(CLEMS) Jasper Son yi 408600 E, 358900 N, Pinus palustris/Pteridium/Quercus
umil f both mesic and dry areas, 10 Aug 1995, McMillan
1101 b(USCH).
=
Collections of Liatris patens have previously been identified as L. graminifolia
or L. gracilis, both of which appear to be sympatric with the new species (Fig.
3). The apparent geographical coherence of L. patens, recorded here for ten coun-
ties in Georgia and two in South Carolina, is matched by its morphological con-
sistency. Liatris patens is similar to L. gracilis in its cauline and peduncular
vestiture and its pedunculate heads, but the new species has larger heads with
more numerous florets and the peduncles are longer and arch upward to bear
1582 BRIT.ORG/SIDA 20(4)
erect or nearly erect heads. Liatris patens is similar to L.graminifolia in its rela-
tively large heads and number of florets but different in its hirtellous-strigose
stems and peduncles and long-pedunculate heads in a broadly racemoid inflo-
rescence. Peduncles of L.graminifolia often are slightly elongate (O-3(-7) mm),
sometimes more in plants with damaged stems, but the peduncles are almost
always straight and divergent at ca. 45 degrees.
Distinctions among Liatris patens and close relatives are summarized in
the following contrasts.
. Stems and peduncles glabrous or nearly so; head lly sessile or on short (O a
7) mm) peduncles, usually in a densely <atettenm arrangement; involucres 7-12 m
high; florets 6-14 per head. Liatris Saran
. Stems and peduncles distinctly hirtellous-strigose with white hairs; heads on di
vergent, arcuate-ascending peduncles and loosely racemoid or nearly sessile but
sods: involucres (4—-)5—7.5 mm high; florets 3-6(-7) or 7-12 per head.
2. Peduncles (O-)2—10(—12) mm long, straight or slightly sioner sce involucres cy-
lindric-campanulate, (4-)4.5-6(-7) mm high, 3-4(-5) mm wide;
per head.
o
Liatris gracilis
2. Peduncles 10-25(-—30) mm long, arching-ascending; involucres turbinate-cam-
panulate, (5-)6-7.5 mm high, 5-7 mm wide; florets 7-12 per head. Liatris patens
ACKNOWLEDGMENTS
We greatly appreciate the help of Bruce Hansen and Richard Wunderlin, who
sent us photocopies of the collections of Liatris savannensis housed at USF John
Nelson, who sent photocopies of the USCH collections of L. patens from MclIn-
tosh Co,, Ga.,and Patrick McMillan, who sent a photocopy of the CLEMS collec-
tion from Beaufort Co.,, S.C. The collection citations from those institutions are
based on the photocopies, which are filed in the BRIT herbarium. A loan of
specimens was received from FLAS. The distinction of Liatris patens also has
been recognized by Melanie DeVore, who began a study of the genus in connec-
tion with an FNA treatment but subsequently moved to other responsibilities.
Bruce Hansen and the editor provided useful comments on the manuscript. We
note that although neither species described here appears to be rare, at least
during the periods when most of the collections were made, both are restricted
in range and their conservation status needs to be evaluated by local and re-
gional field study.
REFERENCES
Avexanper, E.J. 1933. Laciniaria.|n:J.&. Small, Manual of the southeastern flora. Univ.of North
Carolina Press, mee Hill. Pp. 1331-1335
ANDERSON, L.C. 2002.1 sgho! i (Asterac Eupat ),a new blazing star from the
Apalachicola River bluffs and ravines in Florida. Side 20:97-103.
Cronauist, A. 1980. Vascular flora of the southeastern United States, Vol. |. Asteraceae. Univ.
of North Carolina Press, Chapel Hill.
KRAL AND NESOM, NEW SPECIES OF LIATRIS FROM SOUTHEASTERN US 1583
Gaiser, L.O. 1946. The genus Liatris. Rhodora 48:165-183, 216-263, 273-326, 331-382,
Stucky, J.M. 1991. Affinity of Liatris cokeri Pyne & Stucky (Asteraceae), a sandhills endemic
of the Carolinas, and its widely distributed relative, L. graminifolia Willd. Amer. Mid.
Naturalist 125:323-330.
Stucky, J.M. 1992. Liatris virgata (Asteraceae) in the southeastern United States. Sida 15:
177-183.
Stucky, JM. and M. Pyne. 1990. A new species of Liatris (Asteraceae) from the Carolina
sandhills. Sida 14:189-208.
1584 BRIT.ORG/SIDA 20(4)
BOOK NOTICES
Horticulture/Gardening
ROGER PHittirsand Martyn Rix. 2002. The Botanical Garden I: Trees and Shrubs.
(ISBN 0-55297-591-6, hbk.). Firefly Books Lt., 3680 Victoria Park Avenue,
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499-8412, 416-499-1142 fax). $75.00 (US), $95.00 (CANADA), 492 pp. 2000+
color photos, 81/2 x 11"
—
ROGER PHitiies and Martyn Rix, 2002. The Botanical Garden II: Perennials and
Annuals. (ISBN 1-55297-592-4, hbk.). Firefly Books Lt., 3680 Victoria Park
Avenue, Toronto, Ontario M2H 3K] CANADA. (Orders: wwwlireflybooks.com,
416-499-8412, 416-499-1142 fax). $75.00 (US), $95.00 (CANADA), 540 pp.,
2000+ color photos, 81/2 x 11".
Phe Botanical Garden Land ILis an impressive two volume set of modern day botany. It is billed as “a
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useful in a future lecture. Enjoy!
Each taxonomic entry contains Name (genus, species and common names, date of discovery,
and range), Description (detailed and concise in the scientific style), Key Recognition Features,
Evolution and Relationships, Ecology and Geography, and Comments.—Barney Lipscomb, Botanical
Research Institute of Texas, 509 Pecan St, Fort Worth, TX 76102-4060, U.S.A, barney@britorg.
DALLAS Baxter (ed). 2003. Big Bend Gardener’s Guide. (No ISBN, pbk.). Native
Plant Society of Texas, Big Ben Chapter, PO Box 1319, Alpine, TX 79831 (Orders:
PO Box 1319, Alpine, TX 79831, bigbendwoman@brooksdata.net). $10.00,
32 pp., numerous line drawings, | map, 81/2 x 11"
Contents. ~Welcome, Why Use Native Plants?; Planting Zone Map; Weather Data by Community;
What Zone Are We In?, Tree Pruning Do's and Don'ts; Xeriscape: Landscaping for an Arid Climate:
On NOT Following the Rules of Landscape Design; Plant List for the Trans-Pecos Region; Big Bend
Places to See; The Deer Problem; Planting for Wildlife Chart; The Joys of Solarizing: Going Buggy:
Integrated Pest Management; Catchwater Systems; Collecting/Harvesting Plants from the Wild: How
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Some Thoughts on Soil; and Recommended Reading.
anta Tree, What is Making Me Sneeze? A Few Words on Plant Allergies; Cacti: Food for Thought:
SIDA 20(4): 1584. 2003
NEW COMBINATIONS IN XANTHISMA
(ASTERACEAE: ASTEREAE)
Guy L.Nesom
Botanical Research Institute of Texas
509 Pecan Street
Fort Worth, TX 76102-4060, U.S.A,
ABSTRACT
1 +] ‘ sofia
comb M al -incisifolia
Xanthisma incisifolium (I.M. Jol N
lal . {PD
= Machaeranthera incisifolia), and Xanthisma 1b.) Nesom QO Kennon, comb. nov.
(Machaeranthera pinnatifida var. glaberrima), are transferred at specific rank to the genus
Xanthisma, newly expanded by Morgan and Hartman to include species segregated from
Machaeranthera sensu stricto.
RESUMEN
Xanthisma incisifolium (1M. Jo! N comb Mac} } tifida var. incisifolia
= Machaeranthera incisifolia), y > anti an glaberrimum (Rydb.) } Nesom & O’Kennon, comb. nov.
(Machaeranthera pinnatifida var. glaberrima), se transfieren con rango especifico al género
Xant
Machaeranthera sensu stricto
hisma, ampliado de nuevo por Morgan y Hartman para incluir especies segregadas de
Molecular evidence (Morgan 2003) indicates Machaeranthera Nutt. sensu lato
(Hartman 1990) is not monophyletic if the genera Oonopsis Nutt., Pyrrocoma
Hook., and Xanthisma DC. (sensu Semple 1974) are excluded from it. In conse-
quence, Morgan and Hartman (2003) have restricted Machaeranthera to two
species and assigned the remaining species to three other genera, including
Xanthisma. The latter is enlarged by Morgan and Hartman from a monospe-
cific taxon to one including 17 species, and two additional taxa at specific rank
are formally recognized here. Each of those added here is a member of the
Xanthisma spinulosum (Pursh) Morgan @ Hartman complex (=
Machaeranthera pinnatifida sensu lato) and has been treated at varietal rank
by Turner and Hartman (1976), Hartman (1990), and Morgan and Hartman
(2003)
Xanthisma incisifolium (1.M. Johnston) Nesom, comb. nov. BASIONYM: ee
arenarius Bentham var. incisifolius IM. Johnston, Proc. Calif. Aca
12:1190, 1924. Machaeranthera pinnatifida (Hook. f.) Shinners var. eee vi |
Johnston) Turner & Hartman, Wrightia 5:315. 1976. Machaeranthera incisifolia
(LM. Johnston) Nesom, Phytologia 69:112. 1990. Xanthisma spinulosum (Pursh)
Morgan & Hartman var. incisifolium (1M. Johnston) Morgan & Hartman, Sida
20:1409. 2003.
SIDA 20(4): 1585-1588. 2003
1586 BRIT.ORG/SIDA 20(4)
This Mexican taxon was earlier treated at specific rank (Nesom 1990), where it
was noted that it is morphologically non-intergrading with other ‘phases’ of
the Machaeranthera pinnatifida complex. It is restricted to San Lorenzo Island
of Baja California Norte and the islands of San Esteban and Tiburon of Sonora.
Xanthisma glaberrimum (Rydb.) Nesom & O’Kennon, comb. nov. BASIONY™M:
Sideranthus glaberrimus Rydb., Bull. Torrey Bot. Club 27:621. 1900.
Machaeranthera pinnatifida (Hook. f.) Shinners var. glaberrima (Rydb.) Turner
& Hartman, Wrightia 5311. 1976, Xanthisma spinulosum (Pursh) Morgan & Hart-
verrimum (Rydb.) Morgan & Hartman, Sida 20:1408. 2003.
—
man var. gla
—
Xanthisma glaberrimum is an entity of the Great Plains, occurring from the
Texas panhandle region northward into southeastern South Dakota and ex-
treme west-central Minnesota. Xanthisma spinulosum (var. spinulosum) bor-
ders the distribution of X. glaberrimum on the south, west, and north. Turner
and Hartman (1976, p. 311) observed that “Populations of |X. glaberrimumlseem
fairly well-marked throughout most of Kansas and Nebraska and southern
South Dakota, but to the western periphery of these regions they intergrade
over a broad area into |X. spinulosum], especially in Oklahoma and the pan-
handle region of Texas.” The somewhat contrasting observation is made here
that reproductive isolation in this area of sympatry is at least as significant as
intergradation, and in view of its morphological distinction and apparent iso-
lation, a balance of evidence indicates that recognition at specific rank is ap-
propriate for Xanthisma glaberrimum.
In recent floristic studies centered in Hutchinson, Moore, and Potter coun-
ties of the Texas panhandle (Nesom and O’Kennon in prep.), we have observed
many sites where populations of Xanthisma glaberrimum and X. spinulosum
grow in close proximity without intermediacy. A large array of herbarium col-
lections also indicates that two morphological ‘nodes’ exist in this region (see
contrasts in key below) and confirms the field experience. Evidence of intro-
gression is seen, but even in such instances, X. spinulosum that shows charac-
teristics of X.glaberrimum remains distinct from the latter at a population level,
indicating the existence of reproductive isolation. For example, from Hutchin-
son County, Nesom & O’Kennon 164 and 165(BRIT) represent populations of X.
glaberrimum (in typical form) and X. spinulosum (with strict, few-branched
stems, similar to X. glaberrimum in habit but glandular and densely tomen-
tose) that were growing at the same site—the two entities were distinct, with no
intermediates observed in the field. Without the apparent introgression, the
distinction is even more obvious. The direction of introgression appears to be
primarily toward X. spinulosum—this species s| greater inter-populational
variability, especially when it occurs in close proximity to X. glaberrimum,
while the latter maintains a more stable morphology. In our field work, we have
referred to these as the “gray thang” (= X. spinulosum) and “green thang” (= X.
NESOM, NEW COMBINATIONS IN XANTHISMA 1587
glaberrimum) and rarely find it necessary to get closer than 5 meters to make
an identification.
Collections at BRIT indicate that both taxa coexist in at least 10 of the 20
counties in the upper Texas panhandle region—this is considerably more of a
mix than shown by the map in Turner et al. (2003). Herbarium material at BRIT
indicates that a broad area of sympatry also occurs in adjacent Oklahoma. The
area of sympatry was not indicated by the map in Turner & Hartman (1976,
Fig. 42), and the Atlas of the Flora of the Great Plains (Great Plains Flora Asso-
ciation 1976) made no distinction between Machaeranthera pinnatifida var.
pinnatifida and var. glaberrima.
Turner and Hartman (1976, p. 311) noted that “In eastern Wyoming, east-
ern Colorado, eastern New Mexico and the panhandle of Texas, the more typi-
cal, mostly pubescent, populations [of Machaeranthera pinnatifida var.
pinnatifida] pass into the var. glaberrima, the two varieties often occurring near
one another, but it is believed that this has resulted primarily from migration
along road shoulders in relatively recent time, of var. pinnatifida into the re-
gions of var. glaberrima, and perhaps vice versa.” In contrast, however, Nesom
and O’Kennon observe that the two taxa grow both with and without the other
at many sites that appear to be relatively undisturbed and that are relatively
distant from highway sides. The kind of morphological intergradation seem-
ingly implied by the phrase “pass into” would be the impression from study of
herbarium collections and reflect the recognition of gene flow (as also postu-
lated above). Only the repeated field experience led us to interpret this intergra-
dation with further perspective, and it is significant that Turner and Hartman's
observation that the two occur “near one another” agrees with our own.
_ Stems usually unbranched until the upper third, the heads usually distinctly clus-
tered; leaves strictly ascending, narrowly oblong in outline, 1-pinnatifid, midportion
mm wide, lobes oblong-lanceolate to lanceolate or triangular, sometimes
shallowly toothed, glabrous or less commonly lightly tomentose, eglandular or less
commonly glandular; involucres cupulate. Xanthisma glaberrimum
_ Stems branched from midstem or below to the upper third, the heads diffusely
arranged; leaves loosely ascending to spreading, oblong to obovate in outline, es-
pecially the basal and lower cauline, 1-2-pinnatifid, midportion 0.5—-1(-1.5) mm
wide, lobes linear to lanceolate, usually shallowly toothed, usually ee to
varying degrees, slightly to densely tomentose, rarely without eglandular hairs;
volucres shallowly hemispheric. Xanthisma spinulosum var. ; oinuioails
=
Plants of Xanthisma glaberrimum with light tomentum were annotated in 1971
by R.C. Jackson as “var. glaberrima > subsp. wootonii (Greene) Jackson” (the com-
bination by Jackson never published), but the type of Eriocarpum wootonii
Greene was collected in the White Mountains of Lincoln Co., New Mexico, and
is part of X. spinulosum. Its similarity in vestiture to some plants of X.
glaberrimum is interpreted here as parallel. It seems likely that relatively un-
1588 BRIT.ORG/SIDA 20(4)
common plants of X. glaberrimum showing glandularity reflect gene flow from
X. spinulosum, although their close relationship also suggests that parallelism
is a possibility. Differences in pappus morphology noted by Turner and Hart-
man (1976) between X. glaberrimum and X. spinulosum have not been con-
firmed in the present study.
—
ACKNOWLEDGMENTS
Review comments by Ron Hartman, Bob O’Kennon, and the editor are
appreciated.
REFERENCES
Great PLAINS FLoRA Association (R.L. McGregor, coord.; T.M. Barkley, ed.). 1976. Atlas of the
Flora of the Great Plains. lowa State University Press, Ames.
Hartman, R.L. 1990. A conspectus of Machaeranthera (Asteraceae: Astereae). Phytologia
68:439-465.
Morcan, D.R. 2003.nrDNA external transcribed spacer (ETS) sequence data, reticulate evo-
lution, and the systematics of Machaeranthera (Asteraceae). Syst. Bot. 28:179-190
Morcan, D.R.and R.L. Hartman. 2003.A synopsis of Machaeranthera (Asteraceae: Astereae).
Sida 20:1387-1416.
Nesom, G.L. 1990. Notes on variation in the Machaeranthera pinnatifida complex
(Asteraceae) in Mexico with a new combination. Phytologia 69:108—114.
Sempte, J.C. 1974, The phytogeography and systematics of Xanthisma texanum DC.
(Asteraceae): Proper usage of infraspecific categories. Rhodora 76:1-19.
Turner, B.L.and R.L. Hartman. 1976. Infraspecific categories of Machaeranthera pinnatifida
(Compositae).Wrightia 5:308-315.
Turner, B.L., NicHots, H., Denny, G., and O. Doron. 2003. Atlas of the vascular plants of Texas.
Vol. 1. Sida, Bot. Misc. No. 24
my
A NEW SPECIES OF SOLIDAGO (ASTERACEAE: ASTEREAE)
FROM NORTH CENTRAL ALABAMA
Brian R. Keener! Robert Kral
The Unive ity of Alabama Vanderbilt University } ferbarium
Biodiversi ity gee Systematics Botanical Research Institute of Texas
Tuscaloosa, AL 35487-0345 U.S.A. 509 Pecan Street
Fort Worth, TX 76102-4060, U.S.A.
ABSTRACT
Anew go ldenrod. Solid arenicol _ has k ndi SCOVE 1 from Blount C ounty, Alabama. It is hereby
€
described, figured, and contra asted hel her Solid S plumosa Small, S s. simplex be
and S. erecta Pursh, all of which have similar mor pholosies and are from the southeastern U.
RESUMEN
Una nueva vara de oro, Solidago ar _ha sido descubierta en el condado de Blount de Al: abama.
Aqui se describe, se ae y se contrasta con otras tres especies de Solidago, S. plumosa Small, S
simplex Kunth y S. erecta Pursh, que son del suroeste de los Estados Unidos y tienen nieriblosies
similares.
The description below is based on several specimens, all from the same local-
ity, which is the only locality from which S. arenicola has been collected to
date. The collections were made along the Locust Fork River in Blount County,
Alabama. The habitat at this location is shady, acidic woods in deep sandy allu-
vium on the river flood plain. These woods are commonly inundated by swiftly
flowing water when the river overflows following heavy rains during the win-
ter and early spring months. The drier summer and fall months aided by the
porous sandy alluvium bring about a relatively dry substrate as the river re-
cedes toa very small sluggish flow, well within the main channel.
Solidago arenicola B.R. Keener & Kral, sp. nov. (Fig. 1). Type: U.S.A. ALABAMA, Blount
Co. along E side of Locust Fork Branch of Black Warrior River, 0-1/4 miS of Swann
Bridge and ca. 1 mi W of Cleveland by Swann Bridge Road, 7 Sep 2002, R. Kral
93190 & B.R. Keener (HOLOTYPE: US; ISOTYPES: F GA, MO, NCU, NY, TENN, UNA,
USCH, VDB, WAT)
Planta inter S. eae Small et S. simplex Kunth et vars et S. erecta Pursh, species America borealis
orientalis; differt a S. plumosa caulibus brevioribus, foliis basalibus brevioribus in ambitubus (ad
apiceny anOBIeS in eee: anes voneuS achenils dongiompus: ener a = are vere ae iis
paioEhS involucris altioribus; floribus eeieanibes ceegheresionions on Siecitseraious
longioribus
'The Gaivenicy of West Alabama, Department of Biological and Environmental Sciences, Livingston, Alabama
35470, U.S.A.
SIDA 20(4): 1589-1593. 2003
1590 BRIT.ORG/SIDA 20(4)
Wg )
elit ilE|
ih
iN fa
Fic 1 Solid Penta RR K & Kral (f } ype).A. Habit. B. |
g C. Lower cauline
leaf, adaxial view. D. Mid f abaxial E.Upy f aday FE { upper porti
peduncle. G. | hyllary t H. Median phyllary.1. Inner phyllary. J. Ray floret. K. Disc floret post anthesis (left);
expended anther.
KEENER AND KRAL, A NEW SPECIES OF SOLIDAGO FROM ALABAMA 1591
Perennial (35-)40-80 cm tall from a short, horizontal to ascending rhizome,
this and its branches apically caudex-like, to5 mm thick, mostly imbedded in
imbricated chaffy bases of older leaves. Stems erect, single or 2-3 approximate
from rhizomal branches, terete, 2-4 mm thick at base, shallowly multicostate,
glabrous save in inflorescence and its branches, there sparely to moderately
hirtellous with short (0.2 mm or less), narrowly triangular, spreading or
antrorsely curved multicellular hairs. Leaves alternate in high spiral, ascend-
ing toerect, margins coarsely scabro-ciliate, with ciliae antrorsely incurved and
claw-like, the basal and lowermost leaves mostly spathulate to oblanceolate,
10-15cm x 1.5-4cm, blade below middle entire, gradually attenuate, toa slightly
dilated clasping base, distal margins supramedially, coarsely ascending-serrate
with long, shallowly convex or level-backed shallow teeth, apices acute to nar-
rowly rounded, often to short, brownish, callused tips; distal cauline leaves
gradually narrowing and shortening, ranging progressively to narrowly oblan-
ceolate or narrowly elliptic, with apices more narrowly acute, margins entire,
the uppermost 3.5-6cm x 6-10 mm, there grading to widely ascending to spread-
ing or even ref lexed bracteal leaves; leaf surfaces glabrous, adaxially deep green
with only the impressed midvein evident, abaxially pale with midvein raised
and with sharply contrasting level network of deep green vein reticulum. Heads
arranged into racemose or narrowly paniculate inflorescences, mostly with
primary branches widely to narrowly ascending, the lowest usually longest
with longer (often reflexed) bracts, from nodes 2-4 cm distant, and most flo-
riferous, progressively shorter, becoming simple or with few-headed cymules,
or primary axis simply a raceme of heads. Ultimate penduncles stiffly, widely
ascending, angulate, slightly compressed, ca. 0.5 mm thick, stubby-bracteolate,
scattered-hirtellous. Heads radiate, essentially turbinate, mature involucres 7-
9 mm high, 0.7-1 cm wide at distal most point, phyllaries loosely spirally im-
bricate, 1-1.3mm wide with pale, scarious-edged borders, and abaxially sunken
midrib, this bordered by a narrow green band dilating apically toa narrow dia-
mond, around a pale, elliptic depression, all with pilose-ciliate apices, lower
ones 3-3.5 mm long, transitional from peduncular bracts, more green and
shorter than upper, gradually grading to 5-6 mm, narrower, more acute from
narrowly elliptic or oblong to linear-oblanceolate with broader, thinner pale
borders. Pappus bristles of mature achenes of more or less equal lengths, 4-5
mm long, white, fine and finely barbellate, without evident dilated tips. Ray
florets 6-10, carpellate, corolla tube ca. 3 mm long, ca. as long as pappus, blade
mostly narrowly elliptic, 3-4 mm long, yellow, stigma branches to ca. 1 mm
long, dull brown, mature achenes cylindric to somewhat elliptic 2-3.5 mm long.
Disc florets 11-16, perfect, corolla at anthesis ca. 6 mm, dull yellow, tube ca. 3
mm, throat with 5 divergent, narrowly triangular lobes ca. 2 mm; staminal ring
ca. 2.5mm long; stylar cylinder at anthesis 4-4.5 mm, stigma branches barely
divaricate, ca. 1.5 mm. Mature achenes cylindric to somewhat compressed, 3-4
1592 BRIT.ORG/SIDA 20(4)
mm long, typically 5-ribbed, glabrous, abruptly narrowed toa short “neck” sur-
mounted by narrow, brownish pappus disc.
DISCUSSION
Following Nesom’s (1993) taxonomic alignment of Solidago L., we find that S.
darenicola is part of section Solidago; however, it is unclear as to which subsec-
tion the new species should be placed, it having affinities for species in both
Solidago and Albigula (Raf.) Nesom. Within the subsection Solidago, a species
complex, loosely referred to as the “Simplex Group,” seems to share similar
morphologies with S. arenicola (Semple per. com.). The Simplex group is cur-
rently thought to be composed of three species which include S. plumosa Small,
a distinct rarity known only from along the Yadkin River in North Carolina, S.
simplex Kunth, a more complex and widespread set of morphologies, trans-
a)
continental and essentially northern but with one subspecies from the rocky
banks of the Potomac River in eastern Virginia [ssp. randii (Porter) Ringius var.
racemosa (Greene) Ringius] and S. spathulata DC, which is of dune areas re-
stricted to California and Oregon (Cronquist 1980; Cronquist 1994; Hitchcock
& Cronquist 1973). The hypothesized relationship with the Simplex Group and
S. arenicola is based on a suite of characters that S. arenicola shares with these
three species, including caudiciform rhizomes, basally disposed, spathulate or
oblanceolate, glabrous leaves, racemose or thrysoid-paniculate inflorescences
that lack recurved-secund branching, heads turbinate/campanulate with 3-4
seriate phyllaries, ray florets 6-10 per head, and achenes up to 3 mm or more
long. Of the subsection Albigula, which includes several species found in the
southeastern USS., only S. erecta Purs
—
1 could be considered close enough to S.
arenicola morphologically to cause confusion about their identity. The wide-
f
ranging S. erecta is known to occur from coastal Massachusetts and New Jersey,
west to Indiana and south to northeastern Mississippi, central Alabama, and
Georgia. In addition to the characters mentioned above associated with the Sim-
plex Group, which apply here as well, S. erecta and S. arenicola also sometimes
share almost indistinguishable foliage (stem leaves average larger in length and
width in S. erecta), as well as very similar achene shape. (Cronquist 1980; Small
1933).
Differences that help distinguish the new species from similar species of
Solidago that are known to occur in the southeastern U.S. are detailed in the
below:
—~
. Achene surface sees Short-hairy S. simplex
mg surface glabro
. Inflorescence axis al branches glabrous: ac
venes 2—3 mm long S. plumosa
2. Inflorescence axis and branches pubescent; achenes 3-4 mm long.
3. Number of heads per stem usually more than 40, involucres 4—6.5 mm high,
disc flowers usually 6-10 per head, disc corollas ca.4 mm long __ S. erecta
KEENER AND KRAL, A NEW SPECIES OF SOLIDAGO FROM ALABAMA 1593
3. Number of heads per stem usually less than 30, involucres 7-9.5 mm high,
disc flowers usually 12-16 per head, disc corollas ca.6 mm long S.arenicola
ACKNOWLEDGMENTS
We are indebted to John Semple for his continued communication and valu-
able advice over the course of this project. We are also extremely appreciative
of Laurel Hodges who aided in the translation of the English abstract to Span-
ish. We would like to thank Robert Haynes and Steve Ginzbarg who were very
supportive and helpful during this project. We would also like to thank Guy
Nesom and another anonymous reviewer whose suggestions proved to be ex-
tremely helpful in improving the quality of this paper.
REFERENCES
Cronouist, A. 1980. Asteraceae. Vol. 1., Vascular flora of the southeastern United States.
University of North Carolina Press, Chapel Hill. [Solidago, pp. 116-133]
Cronauist, A. 1994. Intermountain flora vascular plants of the Intermountain West, USA.
Volume Five Asterales. The New York Botanical Garden, Bronx. [Solidgo, pp. 242-250]
HitcHcock C.L.and A.Cronauist. 1973. Flora of the Pacific Northwest. University of Washing-
ton Press, Seattle. [Solidago, pp. 549-551]
Nesom, G.L. 1993. Taxonomic infrastructure of Solidago and Oligoneuron (Asteraceae:
Astereae) and observations on their phylogenetic position. Phytologia 75:1-44.
Smatt, J.K. 1933. Manual of the southeastern flora. Hafner Publishing Company, New York.
(Solidago, pp. 1344-1360]
rae
1594 BRIT.ORG/SIDA 20(4)
BOOK NOTICES
Horticulture/Gardening
A.istair Scott (Line drawings by Hrarier INsi). 2002. A Pleasure in Scottish Trees.
CISBN 1-84018-568-6, hbk.). Mainstream Publishing, Edinburgh and Lon-
don. (Orders: Ti alalgat Square, No. Pomfret, VT, U.S.A., 05053, 800-423-4525,
wwwtralalgarsquarebooks.com). $24.95, 202 pp., color photos, drawings,
x 9"
—
This book is described by the publisher as containing a combination of elements of forestry, identifi-
cation, natural woodlanc
s with a celebration of the wonderlul wealth of trees flourishing in Scot-
land. The table of contents include: Native Woodlands, Native Trees, The Restoration of Native Wood-
land, Traditional Fstates, Some Classic Sites of Plantation Forestry, Community Woodlands, Particular
Conifers (29 taxa), Particular Broadleaves (31 taxa), followed by a Bibliography. There is NO index.
TIMBER PRESS
Horticulture/Gardening
KiRSTEN ALBRECHT LLAMAS. 2003. Tropical Flowering Plants: A Guide to Identifi-
cation and Cultivation. (ISBN 0-88192-585-8, hbk.). Timber Press, Inc. 133
S.W. Second Ave, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders:
www.timberpress.com, mail@timberpress.com, 503-227-2878, |-800-327-
5680, 503-227-3070 fax). $24.95, 180 pp., 75 color photos, 4+ b/w photos, 2
tables, 11 drawings, 3 maps, 8 1/2 x 11"
Key Worbs: 1) Tropical plants; 2) Identification; 3) Pictorial Works. From the dustjacket—“Tropical
Flowering Plants: A Guide to Identification and Cultivation bridges a long-standing gap between
obscure references in tropical botany and the gardener’s need for an accurate, practical guide with
clear photographs. Incorporating the latest advances in plant taxonomy from the definitive text of
Dr. Walter Judd, the book isa rare work of scrupulous research—a hat
will be as useful to the gardener as it is to the botanist.” From the Acanthaceae (Acanthus) to the
Zygophyllaceae (Guaiacum), this book is an encyclopedia of beautiful photographs and concise and
specific descriptions of tropical plants. The book ends with three appendices: 1) Invasive anc
Poten-
tially Invasive Species; 2) Rare, Endangered, and Threatened Species; 3) Plants for Coastal Landscap-
ing; and 4) Xerophytic Plants. There is a Glossary, Bibliography, List of Web Sites and an Index of
Scientific and Common Names.
SIDA 20(4): 1594. 2003
SOLIDAGO FAUCIBUS (ASTERACEAE: ASTEREAE),
A NEW MESIC FOREST GOLDENROD
FROM THE APPALACHIAN MOUNTAINS
Thomas F.Wieboldt John C.Semple
mee) Herbariu WAT Herbarium
rtment, Virginia Tech Fedak ee ology
"Bineeburg, VA 24061, U.S.A. rsity of Waterloo
wieboldt@vt.edu ites ans N2L 3G1 CANADA
jcsemple@sciborg.uwaterloo.ca
ABSTRACT
Solidago faucibus, sp. nov. is described from two disjunct areas of the Appalachian Mountains: South
Carolina and a contiguous area of Kentucky, Tennessee, Virginia, and West Virginia. The species is a
decaploid (2n = 90) and has its affinities with the Solidago arguta group. It is distinctive in its ex-
tremely large leaves, especially on basal off-shoots, generally shorter inflorescence branches, large
capitulae, uniformly pubescent achenes, and in forming large clonal (few-flowering) patches. Popu-
lation variation, distribution, and habitat are discussed.
RESUMEN
Solidago faucibus, sp. nov, se describe de dos areas disyuntas de los Montes Appalaches en Carolina
del Sur y un area contigua de Kentucky, Tennessee, Virginia, y West Virginia. La espe
decaploide (2n = 90) y tiene afinidades con el grupo de Solidago arguta. Se diferencia por sus ae
extremadamente grandes, especialmente en los renuevos basales, generalmente ramas de la
inflorescencia mas cortas, capitulos grandes, aquenios uniformemente pubescentes, y en formar
grandes céspedes clonales (que producen pocas flores). Se discute la variacién poblacional,
distribucion, y habitat.
INTRODUCTION
The existence of an anomalous goldenrod became known to the first author
while doing fieldwork in The Breaks of Russell Fork in Buchanan and Dicken-
son Counties, Virginia, in late summer of 1982. The plant was notable for its
large size, especially its basal off-shoot (rosette) leaves. Initially there was un-
certainty regarding its proper status, as there was some thought of its possibly
being a hybrid (A. Cronquist, pers. comm.). Over the next 10 years, additional
populations of the plant were found in eastern Kentucky, southern West Vir-
ginia, other locations in southwestern Virginia, and most recently in Tennes-
see. Notable in this regard, the populations showed a definite affinity for spe-
cific habitat parameters. That the plant recurs in the landscape over a fairly
large geographic area and occupies a specific habitat type argues for its recog-
nition as a species. It is herewith described as new.
SIDA 20(4): 1595-1603. 2003
1596 BRIT.ORG/SIDA 20(4)
money faucibus Wieboldt, sp. nov. (Fig. 1). Typr: UNITED STATES. VIRGINIA. Lee
dry hardwoods around top of N-facing oe ai on Powell River, just
ee Wilder Bend at river mile 135.6, SW ol Jonesville, 24 Sep 1992, TF Wieboldt
8429 (HOLOTYPE: VPI; IsOTYPES: NY, US, VDB, WAT).
Ex alfinitate Solidago arguta Aiton, sed foliis basalibus laminis truncatis basaliter ad S. harrisii Steele
similis, differt ramis floriferis brevibus, capitulis secundis inconspicuo, phyllaria nervis utroque
costae latere, marginibus parallelis, apicibus rotundatis sparse ciliatis. Habitat in sylvis humidas
{ }
APPTTTITe FAUCTDUS,
Herbaceous perennial from a branching caudex, generally producing |-2 (sev-
eral) lateral off-shoots terminated by a cluster of large rosette leaves. Stems erect
solitary, terete, 3-6 mm wide at base, 7.5-15 dim tall, glabrous below the inflores-
cence (and usually into the lower portion of the inflorescence). Leaves dull green
above, pale below, the veinlets forming distinct areolae. Basal and lower cauline
leaves with a pair of primary lateral veins arising 2-3 cm above the base of the
blade and running most of its length. Basal offshoot leaves very large, petiolate,
abruptly tapering toa winged petiole 10-20 cm long; blade ovate 12-22 cm long,
7-15 cm wide, broadly tapering to truncate or slightly cordate (rarely) base,
apices acute, margin singly or doubly serrate with teeth |-3(-6) mm. Lower stem
leaves largest, slightly smaller than rosette leaves, petiolate, abruptly tapering
toa winged petiole, ovate, progressively reduced upwards and becoming sessi
glabrous abaxially, scabrous-hispidulous adaxially at least along margins, ser-
rate with teeth 1-3 mm long, 4-7 per cm. Inflorescence variable, mostly narrow
and elongate, consisting of a slightly arching main axis 12-60 cm long with
axillary and terminal racemiform branches 2.5-7(-13) cm, averaging about 4
cm, or, in sunny situations, wider and compounded, consisting of multiple in-
[lorescence branches with short secondary racemes, leafy bracted below,
branches and peduncles pubescent with short, stiff, owardlly curved, multi-
cellular hairs. Heads large, few to many, inconspicuously secund on longer
branches, appearing fasciculate on shorter branches, peduncles 2-5 mm (longer
on South Carolina plants, +-LO(-13) mm). Involucres 4.5-7 mm. Phyllaries 10-
18, in 4-6 series; median phyllaries averaging 3-4.5 mm long by 1-1.4 mm wide,
lanceolate, broadly acute to obtuse, glabrous except for fimbriate-ciliate mar-
gins towards tip, ascending; midvein heavy and dark, slightly resinous and
raised in distal portion, a pair of lateral veins commonly present, sometimes
evident as interrupted lines, or occasionally only asa short resinous line in the
distal half. Ray florets +-6; strap 2.2-3.5 mm, 0.8-L7 mm wide. Dise florets 5-
7(-11), corollas 4.0-5.6 mm, lobes 1.6-2.5 mm, stigmatic lobes 1-1.3 mm, anthers
~2.9 mm. Achenes 2.5-3.2 mm long, short pubescent at least on upper half,
somewhat variable within populations; pappus bristles 3-4.7 mm.C
number 2n =
Cou
ay)
ge)
AroOMosome
Etymology.—The species epithet faucibus is taken from the phrase “in
faucibus crescens” meaning “growing in the gorges” (Stearn 1983). The common
name “Gorge Goldenrod” is suggested in reference to its preferred habitat.
1597
WIEBOLDT AND SEMPLE, A NEV OF CIES VE
yt
) median phyllary
I y
}
Fa
t, inflorescence
Fic. 1. Solid
1598 BRIT.ORG/SIDA 20(4)
Additional speci ined: KENTUCKY: Breathitt Co.: Robinson Forest, 9 Oct 1982, W. Meijer
sn. (KY). Clay Co.: base of E-facing slope with much Laportea canadensis, Redbird River, 0.1 mi N of
Kates Creek, Daniel Boone National Forest, Creek ville Quad., 37°2'12"N, 83°32'25' W, 30 Jul 1992, Allen
C. Risk 6330 (MDKY). Leslie Co.: Red Bird River, mesic hardwoods at toe of SW-facing slope at mouth
of Bowen Creek, 1.5 km SE of Creekville, 24 Sep 1992, LE Wieboldt 8422, J. Campbell, and D. Taylor
(VPD; abandoned picnic area, hemlock woods, US-421 S of Stinnel, just S of where highway crosses
the river, 14 Sep 1991, J.C. Semple 9619 and B.A. Suripto WAT); moist soil, shaded lower slope in south
llowing ravine, 1.5 mi SE of FS. Rd. 1530 from junction KY 66 and FS. Rd. 1530 in Bowen Creek, 17 Sep
1993, James D. Kiser 371 (MDKY); low steep banks near gullies, along Kentucky River opposite Fron-
tier Nursing Services, 9 May 1990, J. Campbell s.n. (KY); same locality, Jul 1991 & 6 Sep 1991, J. Camp-
bells.n. (KY). Letcher Co.: wooded ravine and hillsides near big Everidge entrance, gravelly road bank,
full sun, Lilly Cornett Woods State Preserve, + Oct 1974, J. Stuart Lassetter 2361 (EKY). Owsley Co.:
oe Brushy Creek, Sturgeon, 22 May 1990, J. Campbell s.n. XY). SOUTH CAROLINA: Pickens
E of US-178, 17 Oct 1981, Semple and C amine wski 6170 WAT); in rich deciduous wood-
lands with circ eamnenta or ultramafic soils on slopes above Eastatoe Creek, 0.8 km E of bridge on
St. Rt. 143 (Roy F Jones Hwy), 8 Sep 2001, PD. McMillan 5866 (CLEM); dry, medium-aged succes-
woodland S$ of = : Jones Hwy, LL km W of jet. Hwy 11, 2 Oct 2002, TLE Wieboldt 11083 (VPD.
acing slope above
Qu
siona
TENNESSEE: Campbell Co.: mesic hardwoods dominated by sugar maple, toe of N-
Hwy 25W where Clear Fork cuts through Pine Mountain, 4 mi W of Jellico, 18 Jun 2003, LE Wieboldt
11206 (VPD; Hancock Co.: 02-302, Short Mt. Rd. 0.1-0.2 mi above church. 7 Oct 2002, H.R. DeSelm s.n.
(TENN). VIRGINIA: Buchanan Co.: Breaks Interstate Park, rich hardwoods along Grassy Creek, Oct
1982, F Levys.n.(VPD; base of mesic hardwood slope, Knox Creek 4.6 km NW of Hurley, 14 Sep 199]
CF Wieboldt 7913 and D.W. Ogle (VPD. Dickenson Co.: Breaks Interstate Park, mesic, rich hardwoods
along Russell Fork just above Garden Hole parking, 25 Aug 1982, LF Wieboldt 4483, A.B. Davenport, F
Levy, M. Medley, and M. Palmer (VPI); same locality, 22 Sep 1992, TF Wieboldt 8417 (VPD. Giles Co.:
en Lyn, lo Aug 1973, LJ. Uttal 10217 (VPI. Lee Co.:
ledges, summit, and adj. woods around high, N-facing limestone cliff, Powell River just above Wilder
Bend, 6.8 km SW of Jonesville, 24 Sep 1985, LF Wieboldt 5893 (VPD, (type station). Scott Co.: seepage
area near base of high, E-facing cliff along Guest River 0.15 km above confluence with Clinch River,
8.5 km SE of Coeburn, 9 Aug 1985, TE Wieboldt 5825 and A.B. Wieboldt (VPI). Tazewell Co.: oak-
dominated mixed hardwoods on narrow, rocky (limestone) ridgeline between Clinch River and In-
dian Creek at Cedar Bluff, 6 Aug 1997, LF Wieboldt 9769 (VPD. Wise Co.: Guest River gorge, 14 Sep
1984, D.W. Ogle s.n. VPD. WEST VIRGINIA: McDowell Co.: very dry hillside, Route 52, Keystone, 9
Sep 1962, F. Rusinke s.n. WVU). Mercer Co.: mesic stream bottom woods from toe of slope to rocky
riverbank, 0.75 mi below Eads Mill along Bluestone River, 2 Oct 1984, LE Wieboldt 5352 and C. Douglas
(VPD; near base of NE-facing outcrops, mesic hardwoods just above road along New River 0.25 mi W
of Virginia line, 30 Aug 1985, TF Wieboldt 5945 (VPD. Duplicate specimens are to be distributed.
limestone bottomland, deep woods, Rt. 649 N of G
The range of Solidago faucibus coincides closely with that of the Appalachian
coalfield region, which may account, in part, for its having gone unrecognized
for so long. This rugged terrain of narrow winding valleys cut into the Appala-
chian and Cumberland Plateaus is, with few exceptions, poorly botanized due
to difficulties of travel and remoteness from population centers.
Solidago faucibus appears to have affinities to Solidago arguta Aiton. The
Solidago arguta group has been variously defined by different authors (see Morton
1973, and references therein). Shared characters with the core members of the
group include having the basal leaves largest and progressively reduced up-
wards, secund flowering branches, glabrous round stems (below inflorescence),
WIEBOLDT AND SEMPLE, A NEW SPECIES OF 1599
hirtellous pedicels and inflorescence branches, and glabrous abaxial leaf sur-
faces (Small 1933). The confusion and difficulties in the group are amply de-
scribed by Morton (1973). Asa direct result of these concerns, considerable hesi-
tancy has been exercised in naming this species. The species bears a close
resemblance to Solidago arguta var. harrisii (Steele) Cronq. In fact, the earliest
known collection of the species is that of E.L. Braun who reported it as Solidago
harrisii (Braun 1943; Campbell 1993). Solidago harrisii was described by Ed-
ward S. Steele (1911) from the shale barrens of the central Appalachians. It was
subsequently considered part of the endemic flora of these unique habitats
(Keener 1983, but see also Braunsweig et al. 1999). Whether treated as a species
or a variety of S. arguta, the taxon has subsequently been found to occur with
high fidelity in association with limestone cliffs over a considerably broader
region. While Braun’s plant proves to be S. faucibus, true S. arguta var. harrisii
has since been found in remote areas of eastern Kentucky J. Campbell, pers.
comm.).
Cytology.—Chromosome counts of 2n = 90 have been determined from three
locations. First, Semple et al. (1993) reported a chromosome count of 2n = 90 for
S. aff. flexicaulis L. from Leslie County, KY. Solidago flexicaulis was otherwise
known to be tetraploid or hexaploid (Semple et al. 1984). Leslie County, KY, is
one of the areas where multiple populations of S. faucibus are known, the
voucher (Semple & Suripto 9619 WAT) is S. faucibus. Second, Semple et al. (1984)
reported 2n =ca. 96 for S. cf. glomerata Michaux from Pickens Co., South Caro-
lina. The voucher (Semple & Chmielewski 6170 WAT) was reexamined and also
found to be a plant of S. faucibus. Review of notes indicate the count was re-
done later and confirmed to be 2n = 90, but not yet corrected in the literature.
Third, when plants of an arguta type were found in South Carolina by Patrick
McMillan, photos were sent to the first author to see if they might be the as yet
unnamed taxon. After seeing the plants in the field in the fall of 2002, the first
author confirmed that they were the same despite a few minor differences. Live
plants were sent to the second author who determined that they also were 2n =
90. In Solidago, chromosome numbers of 2n = 90 are otherwise known only for
S. lancifolia Torr. & A. Gray, from which S. faucibus is readily distinguished (S.
subsect. Glomeruliflorae, Rachel Cook and J.C. Semple, pers. comm.).
DISCUSSION
The most striking feature of Solidago faucibus is its large rosette leaves which
often form large patches (clones?) with few flowering stems. In addition, sev-
eral diagnostic characters serve to distinguish S. faucibus from members of the
S. arguta group. Inflorescence shape is of critical importance in goldenrod tax-
onomy. The disposition of heads along the upper side of inflorescence branches
is made conspicuous in S. arguta by the widely spreading branches which are
1600 BRIT.ORG/SIDA 20(4)
often naked or few-flowered proximally. In S. faucibus, the branches of typical
woodland plants are short, floriferous to the leaf axil, and inconspicuously se-
ye secund nature of the
—~
cund, often suggesting an elongated axillary thyrse. T
inflorescence may be obscured even more in pressed specimens. Often, only
the longest (basal-most) branches display the secund condition. Given an abun-
dance of sunlight in disturbed areas and along woods roads, primary branches
may be more elongate with secondary branching being similarly thyrsoid to
short racemose. It is these open-grown plants with their more open architec-
ture that somewhat disguises their identity since their primary branches may
be clearly secund. Nevertheless, the basal off-shoots and involucral bracts
readily identify them.
The involucral bracts of Solidago faucibus provide additional key charac-
ters which are especially helpful in distinguishing it from S. arguta var. harrisii.
Median involucral bracts clearly show the presence of a lateral vein on either
side of the midvein. It is sometimes discontinuous and may show more clearly
in the distal half. The median phyllaries are parallel-sided with a rounded,
sparsely ciliate tip. These two features clearly differentiate it from S. arguta var.
harrisii, which has only a single midvein and phyllaries that taper most of their
length with barely discernible marginal hairs at the tip. Although leaf shape of
S. argutad var. harrisii is nearly identical, that variety’s widely spreading inflo-
rescence normally identifies it without having to resort to involucral characters.
The multinerved condition of the phyllaries does not fit Nesom’s charac-
terization of Solidago subsect. and ser. Argutae (Mackenzie in Small) Nesom
(Nesom 1993). However, multinerved phyllaries occur in other species in other
subsections as well: S. roanensis Porter in subsect. Albigulae (Raf.) Nesom; S.
glomerata and S. lancifolia in subsect. Glomeruliflorae Torr. & A. Gray) Nesom:
the well known oligoneurate species of sect. Corymbosae Torr. & A. Gray (the
genus Oligoneuron in Nesom 1993, 2000).
Due to its abruptly narrowed leaf bases and more constricted inflorescence
compared with typical S. arguta, the new species can be confused with S.
flexicaulis. The large size is an important consideration because herbarium
—
specimens may lack the lower stem leaves which are crucial in aligning it with
S.argutd rather than Solidago subsect. Glomeruliflorae. Solidago flexicaulis, as
well as other members of its subsection, has its largest leaves near mid-stem or
even closer to the inflorescence. Lower stem leaves are generally absent at flow-
ering time.
Typical Solidago arguta is found commonly throughout the region in drier
sites. On rare occasions, it is found within a few feet of Solidago faucibus, in
which case the two are readily distinguishable with no intermediates being
found. Any intermediates would be cytologically very distinct being either 2n
= 54 (= 9+ 45) or 2n = 63 (=18 + 45).
WIEBOLDT AND SEMPLE, A NCW OF CVICS VUE 1601
Population variation.—Apart from the unifying characters discussed above,
Solidago faucibus displays considerable population variation in several mor-
phological characters. These include leaf vestiture, leaf toothing, inflorescence
branch length, involucre length, peduncle length, and disposition of capitulae
(secund or not). A geographical trend is noted in South Carolina plants although
not all plants display it to the same degree.
Leaves are sparsely scabrous-hirsutulous above. The zone of hairs is typi-
cally marginal but varies in width up to about 1 cm wide, especially toward the
leaf tip. Scabrous hairs sometime occur more widely on veins. South Carolina
plants are scabrous-hirsutulous over most or all of their upper surfaces. Mar-
ginal teeth of basal and rosette leaves vary from single, low (1 mm) serrations
to doubly serrate with jagged teeth (3 mm). Individual populations are quite
uniform, however. Inflorescence branch length is quite variable and appears to
be partly environmentally controlled. The longest branches occur on South
Carolina plants, which display a general tendency toward elongation of parts.
Peduncles are longer (4-10 mm vs. 2-4 mm), involucres average longer, although
the ranges are only slightly skewed toward the long end of the range, and stig-
mas and anthers are longer. Plants from The Breaks (Buchanan and Dickenson
Counties, Va.) with uniformly tightinflorescence branches, when transplanted
to full sun, produced elongate primary branches with short, secund secondary
branching. These populations also have the least coarsely toothed leaves and
leaf blades with noticeably longer length:width ratios. As noted above, heads
on the short inflorescence branches are inconspicuously secund in most popu-
lations. This condition is barely noticeable in the Bluestone River population
(Mercer Co., W. Va.), which otherwise fits well into the species concept. South
Carolina plants are similarly thyrse-like with the slight secund condition be-
ing further obscured by the longer peduncles. South Carolina populations seem
to flower very infrequently, however, so relatively few flowering stems have been
observed. The rosette leaves of South Carolina plants are more truncate to
slightly cordate at the basal angle, and basal rosettes are produced on longer
off-shoots than the more northern populations.
The high polyploid condition may be a factor in population variation.
Multiple copies of the genome could provide an inherent variability with dif-
ferent phenotypic expressions being favored in different populations. It is not
known whether the 10x chromosome number is the result of sequential autop-
loid events with divergence from a single species of Argutae, the result of al-
lopolyploid events involving several species of Argutae, or the result of a com-
bination of autopolyploidy and allopolyploidy. If allopoly ploidy were involved,
then S. faucibus would likely include greater genetic variation.
Habitat.—Solidago faucibus occupies the most mesic habitats of any spe-
cies in the S. arguta group. It is especially tolerant of deep shade, not uncom-
1602 BRIT.ORG/SIDA 20(4)
monly prowing under Tsuga canadensis (L.) Carr, eastern hemlock. Neverthe-
less, it is also tolerant of a wide range of moisture conditions and bedrock types.
The species most often occurs along rivers and major streams where they are
entrenched in the surrounding landscape. In the Cumberland and Appalachian
Plateaus and the Ridge and Valley Province, gorge-like conditions frequently
occur where streams cut through more resistant rocks. It is near the base of
these steep, often rocky, slopes that S. faucibus is most often found, being con-
spicuously absent from the gentler terrain of intervening areas. In this rugged
landscape, forests are frequently intact and typically support a very mesic,
mixed hardwood association. In limestone districts, the species may occupy
considerably more open cliff sites.
KEY TO SPECIES OF SOLIDAGO SER. ARGUTAE
[oR
1. Leaves minutely scabrous, puberulent or short villous, blades attenuate at base.
2. Leaves scabrous on upper surface, the hairs very short and with swollen bases;
bogs, swamps, and seepage in S. patula
2. Leaves soft villose or puberule
3. Rays usually 7-12; leaves modest to densely short villose; spring bloom
ing; coastal plain of N.C,‘ S.verna
3. Rays none, rarely 1-2:1 25d and finely puberulent;fall blooming;open
woods, coastal plain and size ORY e C. to Ala. (possibly Miss. 5. brachyphylla
1. Leaves een Ol ee so, or if scabrous above, the blades abruptly contracted
or truncate at
4. Phyllaries striate, ae ns enlarged, involucres 4.5—7 mm:Va.,W.Va., Ky., Tenn
aes
S. faucibus
SiC.
4. Boylan § Not striate, only main
. Plants without slender stoloniferous rhinomes .arguta (including S. harrisil)
: Plants with slender stoloniferous rhizomes in addition to main more deeply
situated rhizome or cau
». Lower leaves elliptic to <n (ovate), acute, with obtuse to attenuate
base; upper stem leaves quickly reduced and ascending to appressed
S.ludoviciana
6. Lower leaves mostly ovate, acute to acuminate, with truncate to obtuse
base; upper stem leaves gradually reduced, rarely appressed S.tarda
y rarely more striate in S. argut ta).
_s
ACKNOW
-EDGMENTS
The first author would like to thank his wife, Ali Wieboldt, for her artistic tal-
ents in creating Figure 1. Thanks also to many botanists, Foster Levy, Julian
Campbell, Max Medley, in particular, who have known the plant, encouraged
the work, and extended unlimited grace in waiting for it to be named. Thanks
to Patrick McMillan for sharing his discovery on the plants in South Carolina,
and sharing a day in the field. We thank curators at CLEM, EKU, KY, MDKY,
MUHW, NCU, and WVU for making specimens available to the first author.
Appreciation is extended to Guy Nesom and Alan Weakley for their critique
WIEBOLDT AND SEMPLE, A NEW SPECIES OF 1603
and helpful suggestions concerning the draft manuscript. Duncan Porter and
Andrew Becker helped resolve questions about the proper form of the species
epithet.
REFERENCES
Braun, E.L. 1943. An annotated catalogue of the spermatophytes of Kentucky. John S. Swift
and Company, Cincinnati, OH.
Braunswei, S.H., E.T. Nitsen, and T.F. Wiesotor. 1999. The mid-Appalachian shale barrens. In:
R.C. Anderson, J.S.Fralish, and J.M. Baskin, eds. Savannas, barrens, and rock outcrop com-
munities of North America. Cambridge University Press, Cambridge, MA. Pp. 83-98.
Campeett, J.J.N. 1993. Appendix A: Notes on three globally rare plant species known in the
Rugged Eastern Area of the Appalachian Plateau. In: Cooperative inventory of endan-
gered, threatened, sensitive and rare species, Daniel Boone National Forest, Redbird
Ranger District. Pp. 103-128.
Keener, C.S. 1983. Distributional biohistory of the endemic flora of mid-Appalachian shale
barrens. Bot. Rev. 49:65-115.
Morton, G.F. 1973.The taxonomy of the Solidago arguta-bootii complex. PhD dissertation,
University of Tennessee, Knoxville
Nesom, G.L. 1993. Taxonomic infrastructure of Solidago and Oligoneuron
(Asteraceae:Astereae) and observations on their phylogenetic position. Phytologia
Nesom, G.L.2000. Generic conspectus of the tribe Astereae (Asteraceae) in North America,
Central America, the Antilles and Hawaii. Sida, Bot. Misc. No. 20. Botanical Research
Institute of Texas, Ft. Worth.
Semete, J.C., G.S. Rinalus, C. Leeper, and G. Morton, 1984. Chromosome numbers of golden-
rods, Euthamia and Solidago (Compositae: Astereae). Il. Additional counts with com-
ments on cytology. Brittonia 36:280-292.
Sempte, J.C., J. ZHANG, and C. Xianc. 1993. Chromosome number determinations in Fam.
Compositae, Tribe Astereae. V. Eastern North American taxa. Rhodora 95:234-253.
Smatt,J.K. 1933. Manual of the southeastern flora. University of North Carolina Press, Chapel
Hill.
Stearn, W.T. 1983. Botanical Latin, 3" ed. David and Charles, North Pomfret, VT.
Steeve, E.S. 1911. New and noteworthy plants from the eastern United States. Contr. U.S.
Natl. Herb. 13:359-374.
1604 BRIT.ORG/SIDA 20(4)
BOOK NOTICES
TIMBER PRESS
Horticulture/Gardening
JOHN C. MANNING, PETER GOLDBLATT, and DEE SNUMAN. 2002. The Color doa ae
of Cape Bulbs. (ISBN 0-88192-547-0, hbk.). Timber Press, Inc. 133 $
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5680, 503-227-3070 fax). $59.95, 486 pp., 611 color photos, 2 color maps
tables: 8 1/2: 11".
The title just about says it all. The book is filled with excellent color photographs, it is encyclopedic
(nearly 1200 species), and it is all about bulbous and beautiful plants that grow in the Cape Floral
Region at the southern tip of Africa. Included at the end is a Conversion Table, Specialist Suppliers,
Glossary, References, Index of Synonyms, and an Index of Common Names. Well done thou good
and faithful servants! See notice (Sida 19(2), 2000
and Manning) Wi
ona previous book by two of the authors (Goldblatt
—
dflowers of the Fairest Cape.
Davib More and Joun Wiitr. 2002. The Ilustrated Encyclopedia of Trees. (ISBN
0-88192-520-9, hbk.). Timber Press, Inc. 133 S.W. Second Ave, Suite 450,
Portland, OR 97204-3527, U.S.A. (Orders: www.timberpress.com,
mail@timberpress.com, 503-227-2878, |-800-327-5680, 503-227-3070 fax).
$79.95, 800 pp., +500 color plates (watercolors), 71/4 x 10 3/8".
Even though the trees selected and illustrated ar |
mmonl y cultivated in England and continental
Europe, many are found here in American ee and many are native to North America. This isa
beautiful work that took 12 years compile. Deciduous trees are illustrated with both a summer paint-
ing in full leaf and a winter silhouette. Both deciduous and evergreen trees are illustrated as well
ae a close- aap artwork detailing leaves or needles, bark, blossoms, fruit, nuts, or cones. More than
18 ies and cultivars are detailed in this beautif fully done work.
ALLAN M. Armitace and Jupy M. LAUsHMAN. 2003. Specialty Cut Flowers: The Pro-
duction of Annuals, Perennials, Bulbs, and Woody Plants for Fresh and
Dried Cut Flowers. Second Edition, Revised and Enlarged. (ISBN 0-88192-
579-9, hbk.). Timber Press, Inc. 133 S.W. Second Ave, Suite 450, Portland, OR
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photos, 84 line eee | map, 6 x 9".
Contents.—Pretace, Acl ledg
. Introduction, Postharvest Care, Drying and Preserving, Cut
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Suitable for Cut Flower Production; Appendix III: Useful Conversions:
JS De paainent of Agricul-
ture Hardiness Zone Map, Index of Botanical Names, and Index of
Common Names.
SIDA 20(4): 1604. 2003
NEW NAMES AND COMBINATIONS IN GOLDENRODS,
SOLIDAGO (ASTERACEAE: ASTEREAE)
John C.Semple
Department of Biology
University of Waterloo
Waterloo, Ontario, CANADA N2L 3G1
jcsemple@sciborg.uwaterloo.ca
ABSTRACT
{ bi 1: Solid. i t. Multiradiatae
The following new Solidago are |
Solidago subsect. Humiles, Solidago ser. Auriculatae, Solidage ser. Odorae, Solidage ser. Drummondiani,
S. altissima subsp. gilvocanescens, S. kralii, 8. lepida subsp. fallax, S. lepida var. salebrosa, S. odora subsp.
chapmanii, S. patula ie strictula, S. aaa ula subsp. pulverulenta, S. rugosa var. cronquistiana, S.
sempervirens subsp. az sp. mexicana, S. speciosa subsp. pallida, S. stricta subsp.
rica, S.
gracillima, S. velutina cen californicl aie S. velutina subsp. sparsiflora.
RESUMEN
—
os siguientes nombres y combinaciones nuevos en Solidago: Solidago subsect.
Se proponen \
Multiradiatae, Solidago subsect. Humiles, Solidago sev. Auriculatae, Solidago ser. Odorae, Solidago scr.
Drummondiani, S. altissima subsp. gilvocanescens, S. kralii, S. lepida subsp. fallax, S. lepida var. salebrosa,
S. odora subsp. chapmentt: S. aad subsp. strictula, S. eee ula subsp. pulverulenta, S. rugosa vat.
cronquistiana, S. azorica, S bsp. mexicana, S. speciosa subsp. pallida,
S. stricta subsp. aellene: S. selueean subsp. caliente y S. velutina subsp. sparsiflora
The following new names and combinations were determined to be needed dur-
ing work to prepare the treatment of Solidago for Flora North America (Semple
& Cook, submitted). Ten of the new combinations treat taxa as subspecies that
usually have been recognized as varieties. The definitions of the ranks follow
Semple (1974). In each case, the subspecies is morphologically distinct and has
a distribution that is nearly allopatric from the other subspecies within the re-
spective species. Intermediates between the subspecies occur in the areas of
sympatry. Ranges of varieties overlap considerably, with pure and intermedi-
ate populations occurring intermixed within the range of the variety. Hamil-
ton and Reichard (1992) noted inconsistencies in application of the ranks to
taxon, and some of the combinations proposed here correct such inconsisten-
cies within Solidago in North America. Subsection and series names are pro-
posed to fit my opinions on how species should be grouped within the genus; a
full infrageneric nomenclature is included in Semple and Cook (submitted).
Solidago subsect. dieeuaare ey Quz.) Semple, comb. nov. BASIONYM: Soli one
Multiradiatae Juz., Fl. U.R.SS. 25: 47.1959. Type: Solidago multiradiata L
SIDA 20(4): 1605-1616. 2003
1606 BRIT.ORG/SIDA 20(4)
Solidago subsect. Humiles (Rydb.) Semple, comb. et stat. nov. EASTON: Solidago
(sp.-grouq p) Humiles Rydb, Fl. Rocky Mts. 868. 1917. Type: Solidago humilis A. Gray
(1884), non Pursh (1814). [= S S. simple x Kunth|
Solidago ser. Auriculatae Semple, ser. nov. TyPr: Sol
Blake
idago auriculata Shuttlew. ex S.F
Solidagini subsect. Argutae (Mackenzie) Nesom accedens sed foliis auriculatissimis differt.
The leaves are distinctly auriculate clasping. The basal leaves are largest and
petiolate like other Argutae.
Solidago ser. Odorae (Mackenzie in Small) Semple, comb. et stat. nov, BASIONYM:
Solidago (sp.-group) Odorae Mackenzie in Small, Man. SE. Fl. 1345, 1346, 1933. Sol-
idago subsect. Odorae (Mackenzie in Small) Nesom, Phytologia 75:10. 1993, Type:
Solidago odora Aiton.
Solidago ser. Drummondiani Semple, ser. nov. Tyr: Solidago drummondii Torr. & A.
G
Tay.
Solidagini ser. Venosae G. Don accedens sed foliis caulis ovatis triplinervis differt.
Lower mid stem leaves ovate, serrate and with two, large, lower lateral veins
(triple-nerved) as well as additional less enlarged upper lateral veins
Solidago altissima |. subsp. gilvocanescens (Rydb.) J.C. Semple, comb. et stat.
nov. BASIONYM: Solidago canadensis L. var. gilvocanescens Rydb., Contr. US. Natl.
Herb. 3:162. 1895. Solidago gilvocanescens (Rydb.) Smyth, Trans. Kansas Acad. Sci.
1 1S 1899. Doi id igusoeaing Sellen. db. ) Lunell, Amer. Mid]. Naturalist 5:43. 1917.
is L.ssp.gilvocan s(Rydb.) Love & Léve, Taxon 31:358. 1982.
Se aleissirna {. var. gilvocanest scens (Rydb.) Semple Phytologia 58:430. 1985.
Type: U.S.A. NEBRASKA. Hooker Co.: Cody’s | oo the head of the Dismal R, sandy
soil, lO Aug 1893, Rydberg 1 peolmionnat S, photol; IsoTYPES: GH!, NY2, photos!)
Subspecies gilvocanescens includes diploids and tetraploids of S.altissima found
across the Great Plains from southern Canada to Texas. Subspecies altissima
includes hexaploids throughout its range in eastern North America from the
edge of the Great Plains eastward from Nova Scotia to northern Ontario and
eastern Manitoba south to northern Florida and eastern Texas; a few tetraploids
are known from Arkansas and Tennessee. The subspecies differ in head and
Horet size and are difficult to distinguish where the two ranges overlay
Solidago kralii Semple, sp. nov. (Figs. 1-11). Typr: U.S.A. GeorGIA. Pulaski Co. ca. 1-
1.5 mi S of jet. US-341 and GA-230 at Hartford, longleaf pine scrub hills, 10 cv
1975, R. Kral 56345 (HOLOTYPE: VDBI; ISOTYPE: MOI).
Solidagini plumosa Small accedens sed capitulescentiis viscidis resinosisimis differt.
Herbaceous perennials from creeping rhizomes. Stems 6.5-12 dm tall, glabrous
to sparsely strigulose in capitulescence, of ten copiously viscid resinous in capitu-
lescence. Basal leaves
s petiolate, blade gradually tapering to winged petiole,
oblanceolate, (2.5-)1
0-20 cm long, (4-)15-28 mm wide, main vein prominent,
SEMPLE, NEW NAMES AND COMBINATIONS IN SOLIDAGO 1607
wn Sere WAAL 150271
10
Hiyttnyt
=o
=o
=
LANTS OF Georgia
Pulaski Co,: 10 Aug 1975 R,Kral 56345
Solidago speciosa -Nutt, 4
Longleaf pine scrub sandhills ca, 1l-1.5 m | i
Be Jet. US 341 and Ga. 230 at Hartford.
St OR Kral
Fic. 1. Holotype of Solidago kralii, R. Kral 56345 (VDB).
BRIT.ORG/SIDA 20(4)
1608
5 ie,
hattratint
iar
£ Calid
6. Capitulum. 7. Involucre.8. Mature cypsela
|
vt
—2 Net
Fias. 2
mid vein. 4. Stem. 5. P
Imm
SEMPLE, NEW NAMES AND COMBINATIONS IN SOLIDAGO 1609
Fig | £ Cals. kralii 9. R 4 veg ey Pe rere Bs Jui R 'Hite c "¢¢ [Oc Gxr alia
Ly
s.9-11.E a J , J P
11216).10. Habit of 1 ll robust individual; Rict 1 Co., Georgia (Semple & Semple 11217).11.Large bee visitor on
yal 4 | 1, Daal | ol rad - fod I oc My 11208).
t f J \ P I
membranous, glabrous, viscid, margins shallowly serrate apically, teeth less
than | mm long, finely ciliate; rosettes present at flowering, forming at the ends
of elongated rhizomes, first leaves produced the smallest. Lower to mid stem
leaves similar to basal to sessile and linear elliptic, quickly reduced, mid stem
leaves 4-8 cm long, 6-9 mm wide, reduced upward, viscid, entire. Upper stem
leaves sessile, linear elliptic to linear, 10-35 mm long, 1-3 mm wide, reduced
into capitulescence, glabrous, viscid. Capitulescence narrowly thyrsiform
1610 BRIT.ORG/SIDA 20(4)
paniculiform, 9-30(-40) cm long, 4-7(-12) cm wide, longer branches ascend-
ing, 2-4(-15) cm long; heads 1-9(-15) per branch. Peduncles sparsely strigulose,
somewhat to copiously resinous, naked below to bracteolate near heads,
bracteoles usually 1-3. Involucres campanulate, 5-7 mm high. Phyllaries in 3-4
strongly graduated series, the outer ovate 1.5-2 mm long, middle ones 3-4 mm
long, narrowly ovate, to 1.5 mm wide, inner linear lanceolate, apex obtuse,
rounded to slightly cuspidate, ciliate, surfaces often copiously resinous, very
sparsely finely strigose and obscured by exudate. Ray florets 3-5(-11), strap 2.5-
3.5mm long, 1-15 mm wide. Disc florets 10-16(-20), corolla 5-6 mm long, lobes
1-L5C-1.8) mm long. Cypsellae fusiform to narrowly obconic, 3-5 mm long, 5-8
golden brown thin ribs darker than intercostal portions, glabrous; pappus
bristles 4-5 mm long, sometimes strongly clavate. 2n = 18.
Flowering August-September. Turkey oak and pine scrub sandhills; Ga., S.C.
Kral’s Goldenrod is closely related to S. simplex Kunth, S. plumosa Small
and S.arenicola Kral & Keener, the latter two and S. kralii possibly being diver-
gent and isolated relicts of acommon ancestor more widely distributed across
the southeastern United States during late glacial times. Solidago kralii is of-
ten much more copiously viscid resinous then these other three species. The
involucres of S.arenicola are much taller than those of S. kralii and S. plumosa.
Recently made field collections of S. kralii tended to stick somewhat to the
newsprint in which specimens were dried. Solidago kraliiis nearly always found
in full sun on sandhills (Fig. 9); the sites can be highly disturbed. Population
sizes varied from one to several dozen or more individuals, the exact number
not being determined during recent field observations. Individuals were ob-
served in the field with more than three dozen robust shoots up to | m tall (Fig.
10). In general, plants of S. arenicola observed in the field in September 2003
were much shorter, had fewer shoots, and were only found in partially shaded
sandy soils of river banks in northern Alabama. Solidago plumosa can be as tall
as S. kralii, but it is known is only from the type location in Stanly Co., North
Carolina on a very limited section of the rocky margins of the Yadkin River, a
habitat observed in September 2003 that was, prior to dam construction, prob-
ably similar to the habitat of tetraploid S. simplex var. racemosa (E.L. Greene)
Ringius along the Potomac River near the Great Falls of the Potomac in Mary-
land observed in 1984 (Semple & Ringius 7663 WAT, Ringius and Semple 1987).
Chromosome counts for S. kralii determined from two locations were both
were diploid: 2n = 18, U.S.A. Georgia. Pulaski Co.: S of Hartford, GA-230 1 km
SE of US-341/GA-27, 23 October 2001, R. Cook et al. 701 (WAT, shoots well past
blooming); 2n = 9}, U.S.A. Georgia. Richmond Co.: US-1, NE of Blythe, S of Ellis
Pond, steep roadside embankment, sand hill, 7 Sep 2003, J. Semple & B. Semple
11217 (WAT).
The species is named in honor of Dr. Robert Kral (VDB) who collected the
specimens that first brought the species to my attention. He is well known for
SEMPLE, NEW NAMES AND COMBINATIONS IN SOLIDAGO 1611
his work on the flora of the southeastern United States over many years and for
his numerous collections deposited in VDB and duplicates in many other her-
baria in eastern North America.
—
Additional collections: U.S.A. GEORGIA. Ben Hill Co.: US-129 at S edge of Bowens Mill town limits,
just N of GA-182, 7 Sep 2003, J. Semple & B. Semple 11212 (WAT); S of Bowens Mill, US-129 0.4 km S of
GA-182,7 Sep 2003, J. Semple & B. Semple 11216 (WAT); 8.2 mi N of Fitzgerald, sandhills by US-129, 25
Sep 1975, Kral 56788 (MO, VDB). Pulaski Co.: W side of Hartford, US-341 0.9 km E of GA-26, just E of
radio station, 7 Sep 2003, J. Semple & B. Semple 11208 (WAT). Richmond Co.: Ft. Gordon Military Res.,
turkey oak sandhills, S.B. Jones 15123 (USF). SOUTH CAROLINA. Aiken Co.: N of Graniteville, Inter-
state-20, embankment, road right-of-way, 8 Sep 2003, J. Semple & B. Semple 11218 (WAT).
Solidago lepida DC. ae fallax (Fernald) Semple, stat. nov. BASIONYM: ee
lepida DC. var. fallax Fernald, Rhodora 17:9-10. 1915. FL.S. Brit. Columbia 30
Solidago elongata Nutt. var. fallax (Fernald) G.N. Jones, Univ. Wash. Publ. Biol. : 7
1936. Solidago canadensis L. var. fallax (Fernald) Beaudry, Naturaliste Canad. 95:37.
1968. TYPE: CANADA. NEWFLOUNDLAND: Harry’s River, gravelly thicket, 18 Aug
1910, Fernald & Wiegand 4108 (HOLOTYPE: GH}; IsoTYPE: NY photo!).
A discussion of all species in Solidago subsect. Triplinervae is in preparation to
explain the taxonomic treatment of the subsection in Semple and Cook (sub-
mitted). In the treatment, S. lepida is the very sparsely to moderately glandular
member of the subsection. Subspecies lepida is primarily a western taxon found
from Alaska through the mountains to northern Californa and New Mexico
and across northern Canada to Ontario and rarely to New Brunswick and the
Gaspé, Québec. Subspecies fallax occurs in Newfoundland south to New Brun-
swick and rarely to northern Ontario. It has upper stems leaves with more nu-
merous and larger serrations than occur on leaves of either var. lepida or var.
salebrosa of subsp. lepida. Subspecies fallax is only very sparsely glandular with
minute stipitate glands on the phyllary margins or tips.
Solidago lepida DC. var. salebrosa (Piper) J.C. Semple, comb. nov. BASIONY™: Sol-
idago serotina Ait. var. salebrosa Piper in Piper & Beattie, Fl. Palouse Region, 185.
1901. Solidago canadensis L. var. salebrosa (Piper) M.E. Jones, Bull. Univ. Montana,
Biol. ser. 15:49. 1910. Solidago salebrosa (Piper) Rydb., Fl. Rocky Mts. 870, 1067. 1917.
Solidago gigantea salebrosa (Piper) Friesner, Butler Univ. Bot. Stud. 4:196. 1940. Sol-
idago gigantea Ait. var. salebrosa (Piper) Friesner, Butler Univ. Bot. Stud. 5:113. 1941.
Solidago canadensis L. ssp. salebrosa (Piper) Keck, Aliso, 4:104. 1958. TYPE: U.S.A.
WASHINGTON: Pullman, Piper 1580 (HOLOTYPE: WS! ISOTYPE: GH!)
Minute stipitate glands occurs on capitulescence structures (upper most stem
leaves, bracts, peduncles and/or phyllaries) in S. lepida var. salebrosa, which
differs from var. lepida in having arching, elongated lower branches in the
Peres |
CaplIlulesce;nce.
Solidago odora Aiton subsp. aume (A. Gray) J.C. Semple, comb. nov.
ASIONY™: Solidago chapmanii A. Gray, Proc. Amer. Acad. 16:80. 1880. Solidago odora
Ait. var. chapmanii (A. Gray) Cronq., Dalene ZU DITA NO EE: SYN YPESHW. SA.
FLORIDA: “pine barrens, Chapman s.n.(GHI, NY photo!). Levy Co.: Nov 1877, Dr. Gar-
1612 BRIT.ORG/SIDA 20(4)
lesi S1g nated). The
ection at
ber s.n.(GHI, LECTOTYPE, oe GH, ISOLECTOTYPE, here c
Garber collections at GH are marked ‘nsp. A. Gray’; the Chapman col
GH is not. The Lectotype was selected because the label includes in Gray’s hand
“odorless form” and has heads in anthesis: its capitulescence has elongated lateral
amage to the apex of the primary axis. The other Garber collec-
ean
branches due toc
tion has two shoots, both with primarily fruiting heads.
eels patula Muhl. ex Willd. subsp. strictula (Torr. & A. Gray) J.C. Semple,
NOV. BASIONYM: pee ery Muhl. var. strictula Torr. & A. Gray, Fl. N. Amer.
321 3. 1841. Type citation: “North ae to Florida! and Louisiana!” SYNTYPES:
U.S.A. LOUISIANA: Drummond s.n. (GH!, LECTOTYPE designated here because it is a
more complete specimen; narrow- pclae secund capitulescence and low. ver
NA label annotated by Gray as
A
portion of stem with petiolate leaves, T. & G.
patula [beta] strictula”). Hale s.n. (GH! narrow-paniculate capitulescence and por-
ame sheet with the Lectotype). Another collection by
1) “Jacksonville, Louisiana. T. Drummond.
Hooker misit January 1835”, 2) “1870 Herb. From Herb J. Gay purchased by Dr.
Hooker”; 3) “S. patula var. strictula” in Gray’s hand on his SYN. FL. N. AMER. label;
and 4+) an annotation by Dr.G. Morton 1970 with the identification “Solidago patula
ile thi cimen was seen by Gray at some point, there smonaaicatien
it was seen betore 184 and, therefore, is rejected as a possible syntyp
tion of upper stem; on the s
Drummond at GH has several labels:
Solidago puberula Nutt. subsp. pulverulenta (Nutt.) J.C. Semple, comb. et stat.
nov. BASIONYM: Solidago pulverulenta Nutt.,Gen. Pl. 16L. 1818. non Bush (1918). Sol-
idago puberula var. pulverulenta (Nutt.) Chapm., Fl. South. U.S. 210. 1860. TYPE:
U.S.A. GEORGIA: s.d., Nuttall s.n. (HOLOTYPE: not seen; possible soTyPE: DH-122552)).
Solidago rugosa Mill. var. cronquistiana Semple, var. nov. Figs. 12-18). Type: U.S.A.
Nort CAROLINA. Mitchell Co. top of Roan Mt.,6250 [t.el., Forest Rd.-30 near park-
ing lot, 15 Sep L991, Semple & Suripto 9666 (HOLOTYPE: WAT, ISOTYPES to be distrib-
uted: MO, NCU, NY).
Solidagini rugosi var. asperiaccedens sed capitulescentia ramis brevibus foliis subtentibus nunc vix
iffert.
longioribus nune brevioribus et caulibus sparsim hispidis dil
a var. aspera anit var. rugosd cm villlose morph). The leaves are lly
strongly rugose like those of var. dspera, but not as densely hairy. The branches
of the elongated capitulescence are usually short and more like those seen in
morphs of var. rugosa traditionally assigned to the var. villosa, but lacking the
dense pubescence of the latter. The lowest capitulescence branches of very ro-
bust plants can be elongated and repeat the pattern of the upper portion of the
capitulescence (e.g. Semple & Suripto 9829), a similar phenomenon is seen in
the virgate capitulescences of robust plants of other species of goldenrods such
as S. hispida and S. bicolor. Based on field observations and knowledge of type
material in S. rugosa, my lirst impression of the type collection made in 1991
was that it did not fit well into any previously described race of the S. rugosa
complex. Additional collections were made in the Appalachian Mts. of North
Carolina and extreme northern Georgia. The taxon likely also occurs at higher
SEMPLE, NEW NAMES AND COMBINATIONS IN SOLIDAGO 1613
WaT
HARRIES
DEPART OF BOUOSY
\ uNReriner? OF TERED
{TREO ORD
Lamp me cama ty te
Multivariate Studies In Solidago
S. rugosa Complex ee eeecren men No: SP
J.C. Semple & E Ransom
1995 University of Waterloo
AT HERBARIUM
University of Waterloo, Ontano
Solidago rugosa Mt
var. cronguistiana J.C, Semple, var now
oe" Compasitac
J.C, Semple & eee Agus Sunpto
sun 1S Sept 1991
“ 9666
ec
U.S.A, North Carolina, Mitchell Co.: top of Roan
Mb., 6250 2. eb, For Rd,-30 (Roan High BlutT Rd).
an
Large clone by entrance to parking lot
sre WAT: Isotypes: MO, NY, UNOC
HOLOTYPE
Fig 1? Holot £ Cnfid cronauistiana
. vP J J - J
1614 BRIT.ORG/SIDA 20(4)
r)
Lay
+ Siti ele
- ——
Calf ict, (S, le & Suri 9 ).13. Capitulescence
t
ics. 13-18. Details of tk pI | JJ 3 y 4 b v
of wild plant with wasp visitor. 14. Mid stem leaf, abaxial surface. 15. Detail of abaxial surface of upper mid stem leaf.
just bel itul 18. Enlarged f lateral vein st in 15. Scale bars in 16-
Ce
16. Upper mid stem. 17
17 equal 1 mm.
SEMPLE, NEW NAMES AND COMBINATIONS IN SOLIDAGO 1615
elevations in adjacent eastern Tennessee and western Virginia. Allchromosome
determinations made for the new variety were tetraploid. Overall, var.
cronquistiana appears closer to var. aspera than to var. rugosa and is thus as-
signed to subsp. aspera.
The new variety is named in honor of the late Dr. Arthur Cronquist, whose
work on composites is well known and whose occasional phone calls (usually
at night or on weekends) on matters Astereae were both enlightening and en-
tertaining.
aoe collections of var. cr -GEORGIA. Rabun Co.: GA-15 2 km N of Tallulah Falls,
21 Sep 1991, Semple & Suripto 982 28 ( WAT). NORTH CAROLINA. Avery Co.: Beech Mountain, Beech
Me Parkway just W of Red Oak Rd., 4 Sep 2002, Semple 11131 (WAT); E of Linville, Grandfather Mt,
ca. 5000 ft el., 15 Sep 1991, full sun, Semple & Suripto 9674 (WAT), partial shade, Semple & Suripto
9675 (WAT); US-221 S of Linville, 17 Sep 1991, Semple & Su ae 9698 (WAT), between Linville and
Roseborough, Roseborough Rd., F of Blue Ridge Parkway, 3 Sep 2002, Se mple & B. Semple 11120 WAT).
Mitchell Co.: top of Roan Mt.,ca. 6260 ft el., Roan High Bluff Rd. (For Rd.-130), 15 Sep 1991, a &
Suripto 9667 (WAT): : ou 1999, Semple 10808 (WAT). Watauga Co.: Deep Gap, US-221 just N of US
431, 5 Sep 2002, Semple & B. Semple 11135 (WAT).
Collections of aff. var. Sl sal U.S.A. GEORGIA. Rabun Co.: GA-15 2 ae N of Tallulah
Falls, 21 Sep 1991, Semple & Suripto 9829 (WAT, sample includes robust plants wi gated lower
capitulescence branches). NORTH CAROLINA. Wilkes Co.: US-42] 11.8 km E of Watauga Co. line, 17
1991, Se mple & Sur ipto 9707 (WAT, the ca pitulesce nee is di amaged). Duplicates of Se mp rle collec-
tions to be distributed.
Solidago sempervirens L. subsp. azorica (Hochst. ex Seubert) Semple, comb. et
stat. nov. BASIONYM: Solidago azorica Hochstetten ex Seubert, Fl. Azor. 31. t. 10.
1844. Solidago sempervirens L. var. azorica (Hochst. ex Seubert) St. John, Rhodora
17:27. 1915. Type: AZORES, Hochstetten 107 (not seen), St. Michael, s.d., Hccnt? Esq.
s.n., d.Schp-Bip. (GH). The label is handwritten and difficult to read.
Solidago sempervirens L. subsp. mexicana (L.) Semple, comb. et stat. nov. BASIONYM:
Solidago mexicana L., oe Pl. 879. 1753. non Kunth (1818), nec Berlandier ex DC.
(1836). Solidago sempervirens L. var. mexicana (L.) Fern., Rhodora 37:447. 1935.
SYNTYPES: Hort. Clifford 409, Solidago | (LECTOTYPE designated here if not done so
previously: BM). Herb. Linn. 998.13 [?] (LINN). Both specimens are immature [rag-
ments and thus not ideal as type material for a taxon to be distinguished on the
basis of mature head characteristics. The Clifford specimen is larger and more
complete.
Solidago speciosa Nutt. subsp. pallida (Porter) Semple, comb. et stat. nov,
BASIONYM: Solidago speciosa Nutt. var. pallida Porter, Bull. Torrey Bot. Club 19:130.
1892. Solidago pallida (Porter) Rydb., Bull. Torrey Bot. Club 33:153. 1906. SYNTYPEs:
U.S.A. COLORADO. [Jefferson Co.:] Bergen Park, Aug 1877, E.L. Greene (LECTOTYPE
eae as NY ex Columbia!). Canada: Mt. Mackay, Sep 1889, Britton (not seen;
S not occur in Canada).
eoudeee stricta Aiton subsp. gracillima (Torr. & A. Gray) Semple, comb. et stat.
v. BASIONYM: Solidago gracillima Torr. & Gray, Fl. N. Amer. 2(2):215. 1842.
. NTYPES: U.S.A. FLORIDA: “middle,” Dr Chapman s.n. (LECTOTYPE, here designated:
NY; specimen with the capitulescence with three long secund branches;
1616 BRIT.ORG/SIDA 20(4)
ISOLECTOTYPES or just oi if not duplicates of the lectotype collection:
KEW)! NY). Chapman ‘tion labels often lack sufficient information to de-
termine which a eer from rie same location are duplicates of a single collec-
tion or unicates of separate collections.
Solidago velufing; DG subsp. californica (Nutt.) Semple, comb. et stat. nov.
BASIONYM: S ornica Nutt., Trans. Amer. Phil. Soc. ns. 7:328. TYPE: U.S.A.
CALIFORNIA: Sons. Baaban Nuttall s.n. HOLOTYPE: BMI),
oS
Solidago velutina DC. subsp. sparsiflora (A. Gray) Semple, comb. et stat. nov.
BASIONYM: Solidago sparsiflora A. Gray, Proc. Amer. Acad. 12:58. 1877. TypE: U.S.A.
ARIZONA: near Camp Lowell, 1874, Rothrock 706 (HOLOTYPE: GH)).
ACKNOWLEDGMENTS
This research was supported by a Natural Sciences and Engineering Council of
Canada Discovery Grant. Guy Nesom is thanked for his useful review comments
and assistance with the Latin diagnoses. Brenda Semple is thanked for her as-
sistance in the field.
REFERENCES
Hamitton, C.W. and S.H. ReicHaro. 1992. Current practice in the use of subspecies, variety,
and forma in the classification of wild plants. Taxon 41:485—498
Rinaius, G.S.,and J.C. Semete. 1987. Cytogeography of the Solidago spathulata - S. glutinosa
complex (Compositae: Astereae). Canad. J. Bot.65:2458-2462.[S. glutinosa = S. simplex]
Sempte, J.C. 1974. The phytogeography and systematics of Xanthisma texanum DC.
(Asteraceae): proper usage of infraspecific categories. Rhodora 76:1-19
Sempe, J.C. and R.E. Cook (submitted) Solidago. |n: Flora North America Editorial Commit-
tee, eds. Flora of North America. Vol. 20. Asteraceae. Oxford University Press.
—
PAPPUS VARIATION IN SOLIDAGO
(ASTERACEAE: ASTEREAE)
Jennifer L.A. Hood John C.Semple'
Department of Biology
iversity of Waterloo
Waterloo, Ontario, CANADA N2L 3G1
jcsemple@sciborg.uwaterloo.ca
ABSTRACT
The pappus in the goldenrod genus Solidago (Asteraceae: Astereae) was examined in 75 species and
compared with variation in 14 species of 12 related genera in subtribe Solidagininae in the narrow
sense following Nesom (2000) and on ao e asters on two een basal to pine Norte American
Clade of the tribe. Solidago has f one whorl of
barbellate bristles with ae exception of a few ee in sev ae sections, sometimes treated as dis-
tinct genera. Species of Solidago sect. Corymbosae ( pe eae have long been recognized as hav-
ing a biseriate pappus with clavate bristles similar to that of other genera of the subtribe. However,
nearly all species of Solidago displayed some evidence aoe the pappus bristles occur in two
t less heteromorphic whorls of bristles, here designated as rhe primary outer wien and the
rd
primary inner whorl. A few species also occasionally hada thi y outer whorl
of a few bristles. In the most heteromorphic species, the primary outer whorl of bristles was slightly
shorter and had tips that gradually tapered. ther primary inner whorl of slightly longer bristles had
distinctly clavate tips up to several times as broad as the bristle below the tip. As well, the bases of
the primary outer whorl were clearly external to those of the primary inner clavate whorl in some
species. In contrast, most species in several subsections of the genus exhibited little evidence of a
biseriate pappus or clavate bristle tips. The pappus of each species was scored on several traits: 1)
non-clavate to distinctly clavate bristle tips, 2) evidence of alternating non-clavate and clavate bristles
or shorter and longer penis ane 3) ev ene of ov ee Denar outer and inner whorls of
bristles. One species of S y short p bristles of S. sphacelata were less
than half the ner of “the cypse a body. TI he biseriate pappus sl Brintonia discoidea was usually
tl y species of Solidago; the species has been included in
tinted
Solidago by some authors.
Fe
RESUMEN
Se examino el vilano en 75 Resa ae pene ro ee cas SIE?) y se comparo. con Lie
variacion en 14 especies de
oo Neon (2000) e : . ies teraceas de dos géneros bacales al clado Norte ciericans
de latribu it lor de un vilano simple que consiste en un verticilo de
7
oe barbel wane con la excepcion de unas pocas especies en varias secciones, a veces tratadas como
géneros diferentes. Las especies de Solidago sect. Corymbosae (Oligoneuron) han sido reconocidas
por presentar un vilano biselaado con acu clavadas Snatates alas oi otros BeuetOS de la subtribu.
sde
embargo, casi todas las es p g g | | jue | as sedas
J] ] en | ea | Ye i ra | “_ |
UMOL VWELLILITIL
i t y verticilo primario interno. Unas pocas aes tan bio tienen ocasionalmente
un tercer verticilo, el verticilo secundario externo, mucho mas corto y de unas pocas sedas. En la
‘Author of correspondence.
SIDA 20(4): 1617-1630. 2003
1618 BRIT.ORG/SIDA 20(4)
especie mas heteromorlica, el verticilo pri io ext le sedas era ligeramente mas corto y tenia
extremos gradualmente alilados. El verticilo PUSTEMOURECEG, de seca npvaninnie mo) tiene
extremos claramente clavados de hasta varias
Asi mismo, las bases del verticilo primario externo eran claramente externas a 1s del verticilo
pu trio interno clavado en algunas especies. Por contra, la mayoria oe las especies de varias
straban pocas pruebas de un s de las sedas
claysdios EI ila ae bg lnaiasespecie ie escruti nen varios eapecits Wiactenns de las sedas
de no-clavadas a distintamente clavadas 1 edas no-clavadas y clavadas
o sedas mas cortas y mas largas, y 3) ev le ncia de sO Aa amiento de i ver Heiss de sedas primario
cie de Solidag Wide
externoe interno. Una espec sphacelata
eran menos de la mitad de la loneined del cuerpo de lavetoselt 1. El vilano biseriado de de Brintonia
]
normalmente tenido con plgmentos antoclanicos, Cosi aque ho ocurre en le as espectes
de Solidago; la especie ha sido incluida en Solidago por algunos autores.
INTRODUCTION
Previously the genus Solidago L. was considered different from closely related
genera in regards to its pappus characteristics. While related taxa such as
Sericocarpus Nees, Tonestus A. Nels., Petradoria E.L. Greene, Chrysothamnus
Nutt. and Ericameria Nutt. were considered to have a pappus in 2-3 series (1-2
for Tonestus), the genus Solidago was stated to have a pappus in a single series
(e.g., Semple et al. 1999; Nesom 2000). It has also been described as simple
(Fernald 1950; Correll et al. 1970). Some treatments do not mention anything
about the seriate nature of the pappus in the genus description (Radford et al.
1968; Cronquist, 1980; Gleason and Cronquist 1991), although variations are in-
dicated in species descriptions. Some treatments state that the pappus of Sol-
idago exhibits some variation. Gleason and Cronquist (1991) recorded the pap-
pus as being “of numerous equal or sometimes unequal capillary bristles” for
species from the northeast U.S.A. and adjacent Canada, while Cronquist (1994)
noted the pappus as being “equal or sometimes somewhat unequal” in Great
Basin species. Some of the differences in description are due to differences in
which taxa were included in the genus Solidago, particularly the generally rec-
ognized clavate tipped members of Solidago sect. Corymbosae Torrey & A. Gray.
For example, Cronquist (1980) included these flat-topped goldenrods, but ex-
cluded Brintonia discoidea (Elliott) E.L. Greene, the Mock Goldenrod. In con-
trast, Nesom (2000) excluded the former species placing them in the genus
Oligoneuron Small, but included Brintonia discoidea as S. discoidea (Elliott)
Torrey GW A. Gray. Nesom (1991) discussed morphological reasons for treating B.
discoidea within Solidago. Semple (submittted) treated Brintonia as a separate
genus based on morphology (phyllary traits, disc corolla traits, pappus traits),
field observations and DNA studies (Beck and colleagues, pers.comm.). Solidago
sphacelata Raf. has been treated as Brachychaeta sphacelata (Raf.) Britt. due to
its distinctive short pappus, but the general consensus recently is to retain it
within Solidago (Cronquist 1980; Nesom 1991, 1993; Semple and Cook, submit-
ted). Other authors in the past have included Euthamia (Nutt.) Nutt.,
HOOD AND SEMPLE, PAPPUS VARIATION IN SOLIDAGO 1619
Oreochrysum Rydberg, or Petradoria in Solidago, there by increasing the range
of variation in pappus traits within the latter genus (e.g., Fernald 1950; Kearney
& Peebles 1951; Harrington 1964; Semple et al. 1999).
The details of pappus bristle shape have generally been overlooked and
descriptions have included various terms. Most have described the pappus of
Solidago as “capillary” or “hair-like” (Fernald 1950; Radford et al. 1968; Correll
& Johnston 1970; Cronquist 1980, 1994; Gleason & Cronquist 1991). Some treat-
ments mention the pappus of Solidago as being “barbellate” or “clavate” in some
species (Cronquist 1980, S. ptarmicoides (Torrey & A. Gray) Boivin; Semple et al.
1999, S. houghtonii Torrey & A. Gray, S. ptar micoides (as S. asteroides Sem ple) 5.
sempervirens L.). Pappus bristles of Solidago and closely genera have been de-
scribed as being “barbellate” (Nesom 2000). Nesom (1993) in his overview of
Solidago and Oligoneuron used pappus bristle apex features as taxonomic char-
acters: “strongly clavate,” “attenuate,” and “slightly but distinctly dilated toward
the apex.” He listed these in descriptions of sections and subsections, and noted
the character state of pappus bristles for some species in particular.
The occurrence of a biseriate to multiseriate pappus has long been reported
in other genera in the Tribe Astereae in both the North American Clade, which
includes Solidago (see Semple et al. 2002), and the Basal Grade of South Ameri-
can and Old World Genera. The more basal genera in the North American Clade
are biseriate or triseriate with bristles of equal or different lengths. These in-
clude most of the genera included traditionally in the Old World genus Aster
L.: Doellingeria Nees, Eucephalus Nutt., lonactis E.L. Greene, Oclemena ELL.
Greene, Sericocarpus. Most species in subtribe Chrysopsidinae Nesom are
biseriate, having a short outer whorl and a much longer inner whorl. In con-
trast, the pappus of some genera is typically uniseriate, e.g. Symphyotrichum
Nees. In some cases the usual barbellate bristles are absent or very reduced, e.g.
ray floret cypselae of Heterotheca sect. Heterotheca, Aphanostephus DC. The
number of bristles varies greatly between species within and between genera
as well. Thus, within the North American Clade of genera of Astereae there
exists a range in the number of whorls, their relative lengths and their shapes
(see Nesom 2000 for general descriptions of genera). No detailed systematic
examination of the pappus of most or all species of Solidago has been reported.
Our study was undertaken to fill in this gap in knowledge.
MATERIALS AND METHODS
A preliminary survey was undertaken to examine under the dissecting micro-
scope the pappus bristles of one or two specimens of representative species of
the sections and subsections of Solidago. Subsequently, a more rigorous survey
was conducted involving 75 species of Solidago and 17 species of 14 other gen-
era listed in Table 1. Two methods of observation were employed, and a system
of ranking the degree of the clavateness of bristle tips was developed. Most ob-
1620
Taste 1.Pappus characteristics of Solidago taxa and related genera.
BRIT.ORG/SIDA 20(4
)
Taxon Clavate! Alternating Overlapping Comments
SOLIDAGO L. sect. SOLIDAGO
subsect. Solidago
virgaurea 3 Yes (random)
aibeece Multiradiatae ateeen: Semple
S. cutleri a) Yes Yes (slight)
5. multiradiata 2 Yes Yes (slight)
S. spithamea 2 Yes
subsect. Humiles (Rydberg) Semple
S.arenicola 4 Yes Yes (random)
ralti 4 Yes Yes (slight) short obscure secondary
outer whorl
S. plumosa | Yes Yes (random)
S. simplex 5) Yes Yes (random)
S. spathulata | Yes Yes (random)
subsect. oo (A ae Nesom
S. buckle Yes Yes thickened length
Si hincanen mm ; Yes Yes much overlapping of bases
S. oriental! 3 Yes Yes much overlapping of bases
S. petiolaris 3 Yes Yes (slight)
S. wrightii y Yes Yes (random)
subsect. Albigula (Raf.) Nesom
Yes Yes Overlapping very distinct in
places
5. erecta 4 Yes Yes possible secondary outer
whorl, very thickened tips
S. hispida 3 Yes Yes (slight)
S. puberula 3 Yes Yes
S. roanensis 2 Yes Yes (random)
S. sclaphila 3 Yes Yes
S, speciosa 4 Yes Yes
S. squarrosa 3 Yes Yes (random) — bristles highly crowded
subsect. Glomeruliflorae (Jorrey & A.Gray) Nesom
3
S.albopilosa Yes No
S.caesia Yes Yes (slight)
S.curtisil 3 Yes No
S. flexicaulis 3 Yes Yes
S.glomerata 3 Yes Yes (slight)
S.lancifolia 3 Yes Yes (slight)
S.macrophylla 2 Yes No
subsect. ees (Mackenzie) Nesom
ser. Argutae (Mackenzie in Small) Nesom
sf i 4 Ye Yes (random)
S. fauciba 2 Yes Yes (slight)
S. ludoviciana 2 Yes Yes (slight)
5. patula 2 Yes Yes
at base
very large cypselae
seemingly short pappus
HOOD AND SEMPLE, PAPPUS VARIATION IN SOLIDAGO
1621
Taxon Clavate’ Alternating Overlapping Comments
ser. Auriculatae Semple
5. auriculata 3 Yes
er. Brachychaeta (Torrey & A. Gray) Nesom
S. sphacelata 1 No very reduced; alternating
very short and short bristles
subsect. Junceae (Rydberg) Nesom
S.confinis Yes Yes thick bristles, long-short
alternation, pronounced
overlap
S.gattingerl 0 Yes No
S.guiradonis 2 Yes Yes
S.juncea | Yes No
S. missouriensis 0 Yes No ng-short alternation, some
oe bristles are thicker
S. pinetorum | Yes No
S. spectabilis 1-2 Yes No thickened tips, crowding
at base
subsect. Maritimae (Torrey & A. Gray) A. Gray
S. sempervirens 2 Yes reduced secondary outer
whorl (subsp. sempervirens
and —
S. stricta 2 Yes Yes (subsp. stricta and gracillima)
S. pulchra 2 Yes Yes (slight) ee length, very
reduced secondary outer
whorl
S.uliginosa ] Yes Yes (slight)
subsect. Venosae (G.Don in Loudon) Nesom
ser. Venosae
S.ulmifolia 0 Yes Yes overlapping very distinct at
times; alternating short and
long bristles
S.delicatula 1 Yes Yes (random)
S.latissimifolia ] Yes Yes (slight) some prominent overlap
at base of whorls
S. rugosa ] Yes No
S. fistulosa 1 Yes No crowding at base
ser. Odorae (Mackenzie) Semple
S.odora 2 Yes Yes long-short alternation,
strong overlap, possible
secondary outer whorl
subsect. ee is & A. Gray) A. Gray
S.altiplan Yes Yes (slight)
Ss oa 0) No possibly alternating, but
only very weakly (bot
subspecies)
S. canadensis 0 No No possibly alternating, but
a
only very wea
1622
Taste 1. cont.
BRIT.ORG/SIDA 20(4)
Taxon Clavate' Alternating Overlapping Comments
S. elongata 2 Yes Yes (random)
S. gigantea 0 No No
S. juliae | Yes Yes (random)
S. leavenworthii l Yes Yes (random)
5. lepida
subsp. fallax | Yes No
subsp. lepida | Yes No (var. /epida and var.
salebrosa)
S.muelleri 1 Yes No
S. shortil ) Yes Yes (slight)
S. tortifolia 0 Yes
subsect. Nemorales (Mackenzie) Nesom
S.nana 3 Yes
S.nemoralis ] Yes No (subsp. nemoralis and
decemflora)
S. mollis 2 Yes No
S.radula 2-3 Yes No
S.velutina
subsp. californica 2 Yes No
subsp. sparsiflora 1-2 Yes Yes (slight)
subsp. velutina 2 Yes Yes (slight)
SOLIDAGO L. sect. ae iets Torrey & A. Gray
S. houghtor 2 Yes Yes
S. nitida 3 Yes Yes
S.ohioensis 3 Yes Yes
S. ptarmicoides 4 Yes Yes
S. riddellii 2 Yes Yes
S. rigida 2 Yes Yes strong overlapping (subsp.
rigida, humilis, and glabrata)
placement uncertain
S. ericamerioides 2 Yes Yes (slight)
Soins Other Genera of the Solidagininae sensu Nesom (2000)
Chrysoma paucif
osculosa
Yes (slight)
Yes
bristles tinted with antho-
cyanins; biseriate pappus
strong overlapping;
bristles; primary outer whorl
slightly shorter bristles
tapering to weakly clavate;
primary inner whorl of
longer somewhat clavate
bristles
HOOD AND SEMPLE, PAPPUS VARIATION IN SOLIDAGO 1623
Taxon Clavate’ Alternating Overlapping Comments
Chrysothamnus viscidiflorus 1 Yes Yes obvious biseriate pappus
ss icameria laricifolia 3 Yes Yes obvious biseriate pappus
ricameria discoideum ] Yes Yes biseriate pappus
Euthamia graminifolia 0 Yes No variation in bristle length
obscures weak pattern of
short-long alternation of
ristles
dlachia corymbosa 0-2 Yes No weakly biseriate with the
two whorls alternating;
primary outer slightly
shorter, tapering; primary
er longer, weakly clavate
utierrezi th 0 No No crown eee some-
what erose scales of
varying lengths and widths
Hesperadoria scopulorum ] Yes Yes multiiseriate pappus;
primary whorls of many
Longer bristles, one shorter
outer secondary whorl o
fewer bristles
Oreochrysum parry! 1 Yes Yes (slight)
Petradoria pumila ] Yes Yes obvious biseriate pappus
Serico
S. linifolia 4 Yes Yes obvious biseriate pappus;
outer whorl slightly shorter
han |
S. oregonensis 4 Yes No possible slight overlapping
at base of whorls
Tonestus pygmaeus 4 Yes Yes
Genera Basal to the North American Clade
Doellingeria umbellata 3 Yes Yes triseriate; short outer third
whorl
Eucephalus
E. breweri 2 Yes Yes triseriate pappus
E. elegans 4 Yes Yes biseriate, possible weakly
triseriate
' arbitrary ranking of 0 (not clavate) to 4 (strongly clavate)
2 treated as Solidago discoidea by Nesom (1993)
1624 BRIT.ORG/SIDA 20(4)
servations were made using a dissecting scope or a compound light microscope.
Some observations were made using dried and gold coated specimens on a
Hitachi S-570 Scanning Electron Microscope (SEM). At least five different fruits
[rom each species were observed under the dissecting microscope at a maxi-
mum of 70x; most observations were made at 30. In some cases multiple doz-
ens of specimens of a species were examined. For the most part, observations
were made on specimens in the WAT Herbarium, but additional material on
loan from BRIT, F GH, JEPS, NY, TEX and UC (Holmgren et al. 1990) was also
examined.
Observations on the compound microscope were made from slides pre-
pared as follows. For each species, two ray floret and two disc floret cypselae
with corollas were mounted in DepeX mounting medium under a cover slip.
Observations at 00x on the compound microscope were made similarly to the
observations under the dissecting scope at 30x-70«. Observations made on the
two kinds of scopes were compared and any discrepancies were resolved by re-
examining specimens.
Each taxon was scored in the following manner:
1. Cypselae were assigned a number [rom 0-4 depending on the thickness
of the tip of the bristle in relation to the thickness along the length: 0 = no ob-
servable increase in thickness and the bristles tapered to a point (Fig. 1); 1 =a
ne tip, or the bristles not tapering gradu-
—
very slight increase in thickness near t
ally; 2 = a slight to moderate increase in thickness near the tip (Fig. 2); 3 =a
strong and obvious presence of clavate tips; and 4 = very noticeable increase in
thickness, very obvious clavate nature of the bristles (Fig, 3). A set of reference
light-microscopy photomicrographs were used to maintain consistency in as-
signing values.
2. Cypselae were scored as showing no evidence of alternation of primary
"
inner and outer whorls of bristles or as showing evidence of alternating wit
the notes on the nature of the alternation (No or Yes in Table 1, respectively).
3. Cypselae were scored for evidence of overlapping at the bases of the outer
and inner whorls of bristles. The taxa examined were graded in three catego-
apping of the bristles at the base: Yes = complete overlap
—
ries regarding the over
of bristles; Yes (slight) = slightly overlapping with outer whorl bristles consis-
tently appearing external to inner whorl bristles; and Yes (random) = overlap-
ping of some outer bristles external to inner bristles but not consistently so for
the two whorls.
4. Notes were made when a secondary outer whorl (a third whorl) of very
short barbellate bristles was observed. Such bristles were not easily observed
and occurred in low numbers, thus the presence or absence of such bristles was
not tabulated.
Digital photomicrographs were taken using a Nikon CoolPix 990 camera
manually held against the ocular lens of either the dissecting or compound
HOOD AND SEMPLE, PAPPUS VARIATION IN SOLIDAGO 1625
Fics. 1-3. Pappus bristle tip variation in Solidag ing elect icrographs, scale bar = 0.3 mm. 1. Non-clavate
bristle tip, (0 value in Table 1). 2. Weakly clavate bristle tip, 5. odora (2 value in Table 1). 3. Strongly clavate
bristle tip, S. erecta (4 value in Table 1).
microscope. Pictures were taken of specimens under the compound light mi-
croscope with either below stage or above stage lighting. SEM photomicrographs
were made using Ilform Pan F film commercially developed and subsequently
digitized by scanning the negatives. Final digital illustrations were made using
CorelDraw 10® from digital images edited with Corel PhotoPaintl0® (Corel Corp.).
RESULTS AND DISCUSSION
The lack of a standard set of terms for pappus bristle whorls in Solidago and
other genera lead to some confusion in discussing the results between the au-
thors and for the reviewer. The following labels are presented as potential fu-
ture “standard” terms for discussing pappus whorls in the Astereae. Because a
biseriate pappus is found in many genera of the tribe, we recommend that these
two whorls of bristles be referred to as the “primary outer whorl” and the “pri-
mary inner whorl.” The primary inner whorl of ten is longer and has slightly to
strongly clavate bristle tips. The third, often much shorter, outer whorl present
in some genera can be referred to as the “secondary outer whorl.” These labels
for bristle whorls are used consistently throughout this paper. The term “sec-
ondary” with appropriate modifiers can be applied to any additions whorls, e.g.
a secondary innermost whorl of non-clavate long bristles.
The pappus of the genus Solidago is usually biseriate, although obscurely
so in some species. In most species the pappus exhibited either alternation of
1626 BRIT.ORG/SIDA 20(4)
shorter and longer bristles, alternation of clavate and non-clavate bristles, or
some degree of basal overlapping of outer and inner bristles of the primary
inner and outer whorls. Table | lists observations on the degree of clavateness
of bristle tips, presence or absence of evidence of alternation of whorls of
bristles, degree of basal overlapping of outer and inner long whorls, and spe-
cific observations on 75 species of Solidago and 17 species of 14 related genera.
Species of Solidago are grouped within Table | by section, subsection and series
lollowing the treatment of the genus by Semple (Semple et al. 1999: Semple 2003;
Semple, submitted; Cook and Semple, submitted, treatment of Solidago for Flora
North America). There is a large range in size of the bodies and length
ol the pappus bristles among species of Solidago (Figs. 4-9). Bristles can be much
bristle length ranged
from
—
shorter than to four times longer than the fruit body, and
from 0.25-7.5 mm. Cypsela bodies varied from 0.5-5 mm in length anc
glabrous to densely short strigose. The number of bristles per cypsela ranged
from about 25 (Fig. 9, S. canadensis) to more than 40 (Fig. 6, S. auriculata). On
average, pappus bristles were slightly shorter than the disc corollas (averaging
about 9/10 as long comparing the mature pappus to the disc corolla length at
anthesis based on data in Semple and Cook, submitted).
Some of the pappus bristles in the majority of species of Solidago were ob-
served to be more or less clavate, and these were the primary inner whorl bristles
in taxa where position of the whorls could be determined. About 25% of the
species examined had, relative to other species, broad clavate bristle tips on the
primary inner whorl of bristles, e.g, 5. auriculata (Fig. 6), S. erecta and S. argute
Almostall species exhibited some indication of alternating clavate br ae ae
non-clavate bristles (Fig. 6), and some exhibited alternating between short
bristles and long bristles, e.g. S. ulmifolia, (Fig. 13). Alternation of short and long
bristles indicated a biseriate pappus even if little difference was observed in tip
traits of the two whorls. Variation in the pappus of each species was observed
but was of limited range; that is, bristles of a whorl could vary from scores 0
0-2 or 3-4 for degree of clavateness, but not to the degree that some fruits had
only non-clavate bristles while others of the same species had only strongly
SL
——
clavate bristles.
Some species had very clear overlapping of the bases of the primary whorls
of bristles, e.g. S. rigida (Fig. 10) and S. odora (Fig. 12), while others (e.g, S. bi-
color and S. ulmifolia) clearly had overlapping of some bases of the pappus
bristles while other bristles on the same fruit did not appear to be overlapping,
Many other species exhibited only slight overlapping of the bristle bases (eg,
S.glomerata and S. multiradiata) or random overlapping of some bristle bases
but notin a clear primary inner whorl and primary outer whorl pattern. (eg, S.
leavenworthii and S. simplex). Some species appeared to have no overlapping
—
bases, e.g. S. altissima (Fig, Ly)
Some species of Solidago had a third, much reduced, secondary outer whorl
HOOD AND SEMPLE, PAPPUS VARIATION IN SOLIDAGO 1627
Fics.4—9. Length and I iation pappus bristles in Solidago; whi leb 1mm.4-5. Solidago sphacelata,
' A r ll resnactively 6 rey eee .. lat, +) H 1 ith di I] +e hod scale
tA r P é J J er
bar = 1 mm. 7-8. Solidag phyla, i t d mat ypselae, respectively. 9. 5 lidag densis, mature
cypsela.
of a few short barbellate bristles: e.g, S. erecta and S. kralii, S. rigida (Fig. 10)
and S.sempervirens. This condition was difficult to observe and required search-
ing at 70-100x. As noted in the Materials and Methods, detection of the pres-
ence of a third whorl was difficult because t
a
ire few small bristles making up
1628 BRIT.ORG/SIDA 20(4)
Fics, 10—13. Variation in pappus bristle whorls in Solidago ( ing el icrographs). 10. Biseriate senses lg
5. ae scale bar = 0.43 mm; indi I pappus bristle. 11. Uniseriate | §. altissima, scale
100 :m Bi papy ith clearly lapping i 1 out whorls ofbristles, 5; forasenle se mm.
13. Biseri ith st horl ( )and| i whorl of bristles, 5. u/mifolia, scale bar = 1mm.
the whorl could easily be missed. Thus, other species of Solidago than those
noted in Table | may also occasionally produce a secondary outer whorl. Within
the North American Clade of the Astereae a short outer whorl of bristles is
<nown in other species, but it is not always clear whether the short bristles are
the primary or the secondary outer whorl, e.g. all species of Doellingeria, some
species of Eucephalus, almost all species of Erigeron, most species of the
goldenaster subtribe Chrysopsidinae (Semple 1996; Nesom 2000; Semple et al.
HOOD AND SEMPLE, PAPPUS VARIATION IN SOLIDAGO 1629
2002). In Doellingeria, the short outer bristles clearly make up the secondary
outer whorl. In other genera, the outer short bristles may be the secondary outer
whorl or the primary outer whorl. Additional research is needed on these genera.
Within each section and subsection variation was seen in each of the fea-
tures examined, but patterns were still present. The strongest evidence for a
biseriate nature of the pappus of Solidago was observed in members of S. sect.
Corymbosae and in S. subsect. Albigulae. Members of subsect. Junceae, subsect
Maritimae, subsect. Venosae, subsect. Triplinervae nearly always were observed
to have non-clavate to weakly clavate bristle tips. In three species in the latter
subsection, no evidence of a biseriate pappus was detected: S. altissima, S.
canadensis, and S. gigantea. These species had no observable alternation, no clav-
ate qualities to the bristles and no overlapping at the bases.
SYSTEMATICS IMPLICATIONS
The determination that Solidago species generally have a biseriate pappus with
a primary inner whorl of clavate bristles means the genus is not aberrant within
the Solidagininae sensu Nesom (2000). Of the taxa examined, the most distinct
appus was that of Gutierrezia sarothrae which had erose scales rather than
barbellate and sometimes clavate bristles. The results mean that pappus traits
are not useful in separating Oligoneuron (= S. sect. Corymbosae here) from Sol-
idago. If authors wish to do so, then the argument will need to rely on other
differences. Within the genus Solidago, there is sufficient variation in pappus
traits among members of subsections and series that pappus traits can not be
used as diagnostic features of any infrageneric group with one exception. The
very short pappus of S. sphacelata is unique and is diagnostic for ser.
Brachychaeta, if the series is treated as monotypic, as done here. Nesom (1993)
defined the series by its cordate to truncate basal leaves with winged petioles,
strongly reduced cauline leaves, 0-3 ray florets, and markedly short pappus
bristles; he included S. sphacelata, S.auriculataand S. brachyphylla. The bristles
of the latter two species are about 2/3 the length of the corollas, while those of
S. sphacelata are about 1/4 to 1/5 the length of the corollas.
The most obvious conclusion to be drawn is that older descriptions of pappus
traits of members of the Astereae may not be accurate or sufficiently detailed.
Future descriptions should be based on very careful observations at high di-
ssecting scope magnifications or lower powered compound microscope mag-
nifications. Use of the labels “ Dima inner and outer whorls” and “secondary
whorl(s)” should facilitat dditional research on pappus whorl
—
variation in the Astereae.
ACKNOWLEDGMENTS
This research was supported by a Natural Sciences and Engineering Council of
Canada Discovery Grant to JCS and a University of Waterloo Undergraduate
1630 BRIT.ORG/SIDA 20(4)
Research Internship to JLAH. The technical assistance of Dale Weber on the
scanning electron microscope is gratefully acknowledged. Guy Nesom is
thanked for his useful review comments.
REFERENCES
CorreLt, D. S., M.C. JoHNsTON. 1970. Manual of the vascular plants of Texas. 1970. Texas Re-
search Foundation, Renner
Cronouist, A. 1980. Vascular flora of the southeastern United States — Vol, 1 Asteraceae. The
University of North Carolina Press, Chapel Hill.
Cronauist, A. 1994. Intermountain flora vascular plants of the Intermountain West, U.S.A:
Volume 5 Asterales. The New York Botanical Garden, Bronx.
Fernato, M.L., 1950. Gray’s manual of botany 8" Ed.D. Van Nostrand Company, New York.
Gieason, H.A.and A.Cronouist.1991,Manual of vascular plants of northeastern United States
and adjacent Canada, second edition. The New York Botanical Garden, Bronx.
Harrincton, H.D. 1964, Manual of the plants of Colorado. 2nd ed. Swallow Press, Chicago.
Houmaren, P.K., NH. HouMGren and L.C. Barnett. 1990. Index herbariorum. Part |. The herbaria
of the world, ed. 8. Regnum Veg. 120:1-693.
Kearney, T.H.and R.H. Prestes. Arizona flora. University of California Press, Berkeley.
Nesom, G.L.1991.Morphological definition of the Gutierrezia Group (Asteraceae: Astereae).
Phytologia 71:252-262.
Nesom, G.L. 1993. Taxonomic infrastructure of Solidago and Oligoneuron (Asteraceae:
Astereae) and observations on the phylogenetic position. Phytologia 75:1-44.
Nesom, G. 2000. Generic conspectus of the tribe Astereae (Asteraceae) in North America,
Central America, the Antilles and Hawaii. University of North Carolina, Chapel Hill.
Raprorb, A.E., H.E. AHLes, CR. Bett. 1968. Manual of the vascular flora of the Carolinas. The
University of North Carolina Press, Chapel Hill.
Sempte, J.C. 1996. A revision of Heterotheca sect. Phyllotheca (Nutt.) Harms (Compositae:
Astereae): the prairie and montane goldenasters of North America. Univ.Waterloo Biol.
Ser. 37:1-164.
Sempte, J.C. 2003. New names and combinations in Solidago (Asteraceae: Astereae). Sida
20:1605-1616.
SemPe, J.C. (submitted). Brintonia E.L.Greene.|n: Flora North America Editorial Committee,
eds. Flora of North America. Vol. 20. Asteraceae. Oxford University Press.
Sempce, J.C. and R.E. Cook. (submitted). Solidago. |In: Flora North America Editorial Commit-
tee, eds. Flora of North America. Vol. 20. Asteraceae. Oxford University Press.
Sempte, J.C. GS. Rinaius and J.J. ZHanc. 1999. The goldenrods of Ontario: Solidago L. and
Euthamia Nutt. 3 Edition. Univ. Waterloo Biol. Ser. 36:1—90.
Sempce, J.C., S.B. Heard and L. Brouitter. 2002. Cultivated and native asters of Ontario
(Compositae: Astereae):Aster L. (including Asteromoea Blume, Diplactis Raf.and Kalimeris
(Cass.) Cass.), Callistephus Cass., Galatella Cass., Doellingeria Nees, Oclemena E.L.Greene,
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Virgulus Raf). Univ. Waterloo Biol. Ser. 41:1-134.
TYPIFICATION, TAXONOMY, AND A NEW SUBSPECIES OF
POLYGONUM SAWATCHENSE (POLYGONACEAE) FROM
NORTH AMERICA
Mihai Costea'! Francois J. Tardif
Department of Plant Agriculture Department of Plant Agriculture
University of Guelph University of Guelph
Guelph, Ontario, NIG 2W1, CANADA Guelph, Ontario, NIG 2W1, CANADA
coste_amihai@hotmail.com ftardif@uoguelph.ca
ABSTRACT
A new subspecies of Polygonum sawatchense Small is described and illustrated for the forthcoming
treatments of the genus Polygonum (Polygonaceae) in Flora of North America and Jepson Manual,
Flora of California: Polygonum sawatchens
e subsp. oblivium. Polygonum sawatchense is typified and
| Je }
its taxonomy and distribution in North America are discussed.
RESUMEN
Se describe y se ilustra una nueva subespecie de Polygonum sawatchense Small para los futuros
tratamientos del género Polygonum (Polygonaceae) en la Flora de Norteamérica y el Jepson Manual,
Flora de California: Pol itcl | |
lye subs] Polygonum sawatchense se ha tipificado
y se discute su taxonomia y distribucion en Norteamérica.
INTRODUCTION
Small (1893) described Polygonum sawatchense from some specimens collected
by Katherine Brandegee in Sawatch Range, Colorado, U.S.A., and deposited in
CAS. When Hickman (1984) proposed to treat this species at subspecific rank
within P douglasii Greene, he observed that these collections were presumably
destroyed during the earthquake and fire of 1906. He did not propose a lecto-
type or neotype and instead adopted the epithet of the heterotypic P douglasii
var. johnstonii Munz, calling the subspecies P. douglasii subsp. johnstonii (Munz)
J.C. Hickman (Hickman 1984). He thus cited the holotype of P. douglasii var.
johnstonii when he made the new combination P. douglasii subsp. johnstonii
(Hickman 1984). However, based on his annotations from various herbaria (e.g.
CAS, DS, UC, RSA) later Hickman considered plants that agree morphologi-
cally with the type of P douglasii subsp. johnstonii to be “intermediates between
P. douglasii subsp. douglasii and subsp. johnstonii,” whereas the type of
Polygonum douglasii var. johnstonii published by Munz (1935) is actually within
the range of variation of typical P sawatchense Small. In fact, Hickman (1993)
shifted the meaning of P douglasii subsp. johnstonii to an undescribed taxon.
During the preparation of the accounts of Polygonum s.str. for Flora of North
‘Corresponding author
= Sve J
SIDA 20(4): 1631-1640. 2003
1632 BRIT.ORG/SIDA 20(4)
America and the forthcoming revision of the Jepson Manual, Flora of Califor-
nia, it was concluded that P.sawatchense should be accepted at the specific rank.
Plants that were referred to by Hickman (1993) as “P douglasii subsp. johnstonii”
are described here as a new subspecies, P. sawatchense subsp. oblivium.
MATERIAL AND METHODS
Descriptions and details of distribution of taxa are based on collections from:
A, ALTA, CAS, DAO, DC, DS, GH, FE JEPS, LL, MT, MTMG, NY, OAC, POM, QFA,
RSA, SASK, TEX, UBC, UC, USAS and US. Stem papillae are easily observed on
young stems or toward apices of mature stems and branches, at magnifications
higher than 50. Measurements of leaves, ocreae and petioles were made at the
middle of the main stem. The state characters, “flowers closed” and “flowers
wide-open” were observed on herbarium specimens. However, it should be noted
that in fruit, [lowers are always closed. Description of perianth refers to the fruit-
ing perianth, which was measured from the joint with the pedicel. Pictures were
taken with a Scanning Electron Microscope Hitachi $-570 at 15 KV. Samples
were collected from herbarium specimens within the holdings of GH, NY and
RSA; these collections were indicated with a “*” in the lists of representative
specimens examined. Samples were coated with 30 nm gold using an Emitech
K 550 Sputter Coater.
Typification of Polygonum sawatchense
A specimen collected by Brandegee from the Sawatch Range, Colorado is de-
posited in NY (00323810), but it was not cited by Small (1893) in the protologue
and, furthermore, it is damaged. A neotype is designated below from one of the
specimens cited one year later by Small (1894) as “nearly like the origina
every respect.”
—_—
in
Taxonom
Hickman (198+) proposed a new taxonomic concept for P ius ala by includ-
ing within it five other species: P. sawatchense, P. spergulariaeforme Meisn. ex
Small, P majus Meisn,, P nuttallii Small and P austiniae Greene. Hickman (1984)
justified these new combinations by the intermediate patterns of variation he
observed between some of the taxa: “intermediates sometimes outnumber ty pi-
cal forms.” For example, he reported such frequent intermediates between P
douglasii and P. sawatchense (Hickman 1984). However, even in this case Hick-
man admitted later that they “both may grow together without local intergra-
dation” (Hickman 1993). We found that Hickman’s intermediates (in herb.) are
in most of the cases within the range of variation of each taxon, which has
been poorly understood. Polygonum sawatchense is more variable than previously
known and two subspecies, one of them formally undescribed, are recognized
here. The two subspecies may partially intergrade in the areas of sympatry.
COSTEA AND TARDIF, TAXONOMY OF POLYGONUM SAWATCHENSE 1633
Polygonum sawatchense Small, Bull. Torrey Bot. Club 20: 213, Le pe. 1893. TyPE:
U.S.A. Yellowstone Park, 1885, T. Tweedy s.n. (NEOTYPE, d: USD.
a. P. sawatchense subsp. sawatchense (Fig. la)
- Polygonum douglasii Greene var. johnstonii Munz, Man. So. Calif. Bot. 131:597. 1935; P. douglasii
subsp. sohastori (Munz) J.C. Hickman, Madrono 31:250. 1984.
Polygonum exile Eastw., Proc. Calif. Acad. Sci. Ser. 3, Bot. 2:286. 1902.
Polygonum triandrous Coolidge, Madrono 20:266. 1970.
Stems green to brownish, usually branched from base, 4-50 cm long, glabrous
(Fig, 2a), with smooth faces and sometimes narrow winged angles. Leaves; ocreae
glabrous; blades linear, narrow-oblong to oblanceolate, 15-45 x 2-8(-12) mm,
margins usually revolute, glabrous (Fig. 2c); upper leaves abruptly reduced to
scale-like bracts, or much longer than the flowers they subtend. Inflorescences
axillary or of long, open spike-like racemes, 5-15 cm long; cymes (1-)2-4-flow-
ered; bracts equalling, somewhat longer, or much longer than the flowers. Flow-
ers closed; pedicels erect, 1-4 mm, ae n (2.5-)3-3.5 mm, the tube 25-40% of
the perianth length, the tepals naviculate (Fig. 2e), greenish or reddish with
white or pink borders, with midveins unbranched or with a few branches at
base. Stamens 3-8; anthers pink to purple, 130-160 pm long. Achenes black,
trigonous, 2.5-3 mm, ovate or elliptic, smooth.
Distribution and ecology—CANADA: Alberta, Saskatchewan. U.S.A.. Ari-
zona, California, Colorado, Idaho, Montana, Nevada, New Mexico, North Da-
kota, South Dakota, Utah, Washington, Wyoming. MEXICO: Baja California. It
grows in dry meadows, pastures, sagebrush and forests on sandy, gravelly or
rocky substrates. Flowering Jun-Aug.
It is important to emphasize that narrow-leaved plants regarded
man as intermediates between P. douglasii subsp. douglasii and his “subsp.
johnstonii,” belong in fact to P. sawatchense subsp. sawatchense. Even with this
narrow circumscription, P sawatchense subsp. sawatchense is variable. “Typi-
cal” P. sawatchense plants described by Small from the Rocky Mountains, Colo-
rado are relatively dwarf, 6-15 cm tall, dull green-brownish with persistent
leaves that gradually transform into leaf-like bracts, 3-5 times longer than the
flowers they subtend. Plants described as P. douglasii var. johnstonii by Munz
from San Bernardino Mountains, California are taller, to 50 cm, with caducous
green-glaucous, basal leaves, sharply differentiated from the bracts, which are
equal to or up to 3 times as long as the flowers they subtend. However, we found
a continuous transition between these two extremes, and a further subdivision
of P. sawatchense subsp. sawatchense is apparently not possible. Collections of
Patterson (1885) from the Rocky Mountains, near Golden, Colorado illustrate
very well this transition. For example, Patterson 131 (UC, NY) collected from
elevations ranging from 3000 to 3300 m is “typical.” Small himself observed
that another collection (Patterson 128; NY, UC, US) from an elevation of 1800 m
—
by Hick-
1634 BRIT.ORG/SIDA 20(4)
Fic. 1. Pol { f general habit; a. P } I p. sawatchense, based on PA. Munz & M. Johnston
8506 (RSA), b. P tch bsp. oblivium, based on FW. 0 260 (RSA)
J
COSTEA AND TARDIF, TAXONOMY OF POLYGONUM SAWATCHENSE
IG. a pt 1 gy wer s p lyg L | f | L
subsp. oblivium (scale bar = 0.3 mm), c—d. Margin of leaves:
0.5 mm), b. P sawatchense
p. sawatchense, d. P. sawatchense
subsp. oblivium (scale bar = 0.3 mm), e-f. Fruiting perianth: e. P. sawatchense subsp. sawatchense, f. P. sawatchense
subsp. oblivium (scale bar = 1 mm).
1635
1636 BRIT.ORG/SIDA 20(4)
“is more elongated and rather straggling” and that “this difference is most likely
due to their unlike habitats and altitudes” (Small 1894).
Representative specimens examined. CANADA. ALBERTA. Cypress Hills Region: Chappice Lake, 23
Jun 1990, ].H. Hudson 4921(SASK). SASKATCHEWAN. Moose Jaw Region: Grasslands Park, 9 mi SE of
Val Marie, 17 Jul 1989, E.R. Hooper ILO (USAS). Swift Current Region: Altawan, 49E14’30°N LOSES W.
8 Jul 1999, ].H. Hudson 5407 (SASK). U.S.A. ARIZONA. Apache Co.: 3.5 mi E of McNary, roadside,
ponderosa pine forest, 13 Aug 1974, DJ. Pinkava 12343 et al. (US, NY). Coconino Co.: Buch Springs
Ranger Station, 7500 {t, Sep. 1936, R.E. Collom 599 (US); Flagstaff, 2 Aug 1884, M.E. Jones 3957 (RSA*).
Cochise Co.: Chiricahua Mountains, Coronado National Forest, base of Barfoot Peak, 8200 ft, 4 Sep
1983, D. Ward & D. Peterson s.n. (RSA*); road to Rustler Park, 7600 ft, 14 Oct 1943, TH. Kearney & R.H
‘ebles 15104 (UC). Gila Co.: Workman Creek Canyon, near end of car road, Sierra Ancha Viouncains,
5500 ft, 27 Aug 1946, FW. Gould 3734 (GH). Greenlee Co.: Apache National Forest, 13 mi Sof Strayhorse
Campground, just N of Blue Lookout, ponderosa pine, 24 Jul 1973, D.J. Pinkava 112804 et al. (US, NY).
CALIFORNIA. Fresno Co.: Hutchinson Area, Piute Canyon, 10 mi E Florence, 12100-12500 ft, 18 Aug
1955, CH. Quibell 5847 (RSA). Inyo Co.: Head of Wyman Canyon, White Mountains, 10000 ft, 19 Jul
1955, PA. Mung 21043 (RSA). Lassen Co.: Harvey Valley, 9 Jul 1934, ].T Howell 12528(GH, RSA*). Modoc
Co.: one side et Benet Mountain, 2.8 mi N of 8 mi creek on the road from Lake Annie, 30 Jun 1984,
All an ); Duncan Horse Camp, | Jul 1934, ].T. Howell 12317 (GH, NY). Mono Co.: Sweetwater
Mountains, site ol Star City, 8500 ft, 12 Jul 1975, M. De Decker 3958 (RSA): Sierra Nevada, Little Lake,
3300 m, 20 Jul 1969, M DeDecher 2163854): Nevadai¢ Co.: se Valley, 5500 ft, 30 Jun 1913, A.A. Heller
10897 JEPS, NY, US). Riverside Co.: Santa Rosa Peak, Virgin Springs, 7500 ft, 14 Aug 1938, PA. Munz
15359 (RSA*). San Bernardino Co.: San Bernardino ee bellow Dollar Lake, 8700 ft, 22 Aug
1922, PA. Mung 6190 (RSA*, UC), Tulare Co.: Head of Monache Meadows on § Fork of Kern River,
8050 It, 13 Jul 1950, PA. Munz 14994 (RSA), Mineral King, 8000 ft, 19 Jul 1951, L.S. Rose s.n. (RSA, UC).
Ventura Co.: Pinos Mountain, 8400 ft, 30 Jul 1922, FW. Peirson 3223 (RSA). COLORADO. Conejos
Co.: 7 mi N of Conejos Campground, NW of Antonito, 19 Jul 1952, W.A. Weber 7862 (RSA*). Grant
Co.: ne ky Mountain National Park, 2.25 mi WSW of Mount Bryant, granite soil origin, 8500 ft, 4
Aug 1936, Sawyer & Rutter 8b (UC). Gunnison Co.: 3 mi N of Almont, on road to Crested Butte,
8500 {t, 2 Aug 1955, W.A. Weber 9376 (GH, RSA, UC). Larimer Co.: Moraine Park, 11 Aug 1925, G.E.
Osterhout 6464 (RSA*). Montezuma Co.: Mesa Verde National Park, near Cedar Tree Tower ruins, ca.
7100 ft, 26 Jul 1949, H. Bailey & V. Bailey 3106 (UC). Rio Grande Co.: West Fork San Francisco Creek,
Del Norte Ranger District, Rio Grande National Forest, 7 Aug 1984, B.H.M. Movers 437 (NY). IDAHO.
Blaine Co.: Ketchum, 5877 ft, 20 Jul 191, A. Nelson & J. Macbride 1246 (RSA). NEVADA. Clark Co.:
Charleston Mountains, Charleston Park, 2270 m, 29 Jul 1938, LW. Clokey 7901 (GH, NY, RSA*, UC);
Pipeline Canyon to Rainbow Falls, FE of Cathedral Rock, Charleston Mountains, 8200 ft, 30 Jul 1938,
P Train 243 (GH, NY, UC). Elke Co.: Lamoille Canyon, Ruby Mountains, 20 Jul 1938, EW. Pennell &
RL. Shaffer 23307 (GH). Humboldt Co.: Pine Forest Range, 1] mi N of Leonard Creek Ranch, 6400 ft,
27 Jun 1964, N.H. Holmgren & J.L. Reveal 1120 (GH, NY, RSA*, UC). Mineral Co.: Cottonwood Creek
Canyon, + mi below Laphan Meadows Mountain, Grant Rd., 7800 ft, 22 Jun 1940, P Train 4109 (NY,
UC). Nye Co.: Pine Creek Canyon, frequent, gravelly sagebrush slopes, 20 Jul 1940, B. Maguire & A.H.
Holmgren 25861 (GH, NY, UC, US). Washoe Co.: 20 mi NE of Gerlach, 5500 ft, 20 Jun 1942, C. Mc-
Millan & K.H. McKnight 147 (NY). NEW MEXICO. Catron Co.: between Luna and the Arizona border
on Hwy. 180, 14 Aug 1973, R. Spellenberg & M. Spellenberg s.n.(NY). Chaves Co.: Roswell, 21 May 1931,
Nelson 11587 (UC). Colfax Co.: vicinity of Ute Park, 2200-2900 m, 26 Aug 1916, PC. Standley 1 3847
ra) Grant Co.: Pinos Altos Mountains, 24 Sep 1931, M_E. Jones 28608 (RSA). Lincoln Co.: White Moun-
tains, 7400 ft, 25 Aug 1907, E.O. Wooton & PC. Standley s.n. (US). McKinley Co.: Zuni Mountains, 8 mi
E of Hwy. 32 on road to ren inside Cibola National Forest, 16 Aug 1973, R. Spellenberg & M.
Spellenberg s.n. (NY). San Miquel Co.: TM. O'Connor Ranch, South Rim Pasture, 6900 ft, 1.5 mi E of
Las Vegas, 28 Jul 1984, S.A. Hill 14631 (NY, RSA*). Taos Co.: Apache Springs, Apache Peak, Carson
—
—=
COSTEA AND TARDIF, TAXONOMY OF POLYGONUM SAWATCHENSE 1637
Forest, 3000 m, 22 Aug 1923, W.W. Eggleston 19162 (US). Valencia Co.: Mount Taylor, San Mateo Moun-
tains, College experimental area, 20 mi E of Grants, 8300 [t, 4 Sep 1935, KF. Parker eeeae UTAH.
Grand Co.: Uintah and Ouray Indian Reservation, 59 mi S of Ouray, along Weaver Reservoir Rd.,
8500 ft, 30 Jul 1965, N.H. Holmgren etal. 2309(GH, NY).Juab Co.: Ridge above junction of Nebo Loop
Rd. and Salt Creek Rd., 6400 Ft, | Jul 1983, kK. Thorne & J. Thorne 2523 (NY). Salt Lake Co.: Big Cotton-
wood Canyon, 8000 ft, 17 Aug 1941, J.F Brenckle et al. s.n. (GH, NY, US). San Juan Co.: S side of Abajo
Mountains, 2000-2500 m, 28-29 Jul 1911, PA. Rydberg & A.O. Garrett 9270 (NY, US). Washington Co.:
Pine Valley Mountains, 6760 ft, 2 Aug 1967, J.L. Gentry Jr 2256 (NY); Zion National Park, East Rim
trail, Echo Canyon, Ll Jul 1949, H. Bailey & V. Bailey 3537 (UC). WYOMING. Crook Co.: Bear Lodge
Range, Reuter Canyon, 5 mi N of Sundance, 5800 ft, 12 Jul 1960, CL. Porter & M.W. Porter 8374 (RSA*,
UC). Fremont Co.: Togwotee Pass, 26 Jul 1939, J. Brenckle 39-139 & S.D. Mellete (RSA, NY, UC, US).
Natrona Co.: Southern Powder River Basin, 10 mi SW of Waltman along Powder River, 0400 [t, 6 Jul
1993, R.L. Hartman 40486 (UC). Weston Co.: Black Hills, Elk Mountain Area, Clifton Canyon, above
Last Canyon, 4700 ft, 8 Aug 1984, H. Marriott 8809 (NY). MEXICO. BAJA CALIFORNIA: Sierra, San
Pedro Martir, Dead Horse Meadow, SW end of Venado Blanco, see Jul 1988, S. Boyd 2628 et al. (RSA*);
Vallecitos Meadow, 2400 m, 26 Aug 1973, R. Moran 21127(R
Cy
b. ga a sawatchense subsp. oblivium, Costea & Tardif, subsp. nov. (Fig.
E: U.S.A. CALIFORNIA. Siskiyou Co. Salmon Mountains, High Lake Basins
n the vicinity of English Peak, Marble Mountain Wilderness Area, gully near
neon SE of Tom Taylor Cabin, elevation 6700 ft, 13 Aug 1967, FW. Oettinger 260
(HOLOTYPE: RSA).
Polygonum douglasii subsp. johnstonii auct
DtICLs
acai
Polygono sawatchensi subsp. sawatchense alfinis sed caulis scabri-papillosis et foliis oblongi-elhi
vel ellipticis, marginibus papillosi-denticulatis differt.
Stems green, simple or with a few branches from base, 5-15(-25) cm long, pap-
illose scaberulous on and between ribs; papillae white, patent, dense, conic-
cylindrical, 90-150 um long (Fig. 2b). Leaves; ocreae 4-10 mm, scaberulous, with
the free part lacerate or disintegrating into a few fibers; blade elliptic to oblong-
elliptic, gradually decreasing in size upwards and persistent at flowering-fruit-
ing stage, 8-20(-25) x 5-10 mm, acute at both ends; margins plane, papillose-
denticulate; papillae similar to those present on the stems (Fig. 2d).
Inflorescences mostly axillary or if a terminal spiciform raceme present, then
less than 5cm long; cymes-1-3(-4)-flowered. At least some flowers wide-open;
pedicels 1-3 mm. Perianth 3-4 mm, the tube 20-30% of the perianth length,
tepals oblong-elliptic, cucullate, naviculate and overlapping (Fig. 2{), greenish-
white or greenish-yellow (sometimes flushed with purple) with white borders
the mid-veins greenish or brown, usually branched at base. Stamens 8; anthers
pink to purple, rarely whitish, 130-200 um. Achenes black, trigonous, 2.6-3.3
min, elliptic to elliptic-ovate, smooth.
Distribution and ecology.—CANADA: British Columbia. U.S.A.: California,
Idaho, Nevada, Oregon and Washington. It has the same ecology as subsp.
sawatchense, but unlike the latter may also grow in shaded or moist places. Flow-
ering Jun-Au
This taxon has been usually referred toas P. douglasii subsp. johnstonii (e.g.
Le
1638 BRIT.ORG/SIDA 20(4)
by Hickman 1993). The protologue of P. douglasii var. johnstonii indicates only
that it has erect pedicels, | mm long (Munz 1935). Hickman (1984) studied the
holotype of P. douglasii var. johnstonii from RSA (Munz & Johnston 8506) when
he included P sawatchense within his new subspecies of P. douglasii. He noted:
“the type shows several features of subsp. douglasii but it is closer to the ex-
treme of subsp. johnstonii so the latter epithet can be retained at subspecific
rank” (Hickman 1984). However, only almost 10 years later, in the Jepson Manual
(1993) did it become apparent what Hickman understood by “subsp. johnstonii.”
He described this taxon as having “persistent leaves, all elliptic” (Hickman 1993).
This intrigued us because Munz’s holotype has caducous, narrow, oblong-lan-
ceolate leaves. Even Munz specified later (Munz & Keck 1968) that, except for
the erect and short pedicels, P. douglasii var. johnstonii is like P douglasii (var.
douglasii), namely with narrow, oblong-lanceolate leaves. Apparently, Hickman
changed the original meaning of this name, from the plants of Munz, which
are within the range of variation of typical P sawatchense, to a taxon that has
not been formally described. These plants are called here P. sawatchense subsp.
oblivium. This contusion induced Hickman to consider plants of P. douglasii
with broad leaves (var. latifolium Greene), as well as typical plants of P.
sawatchense, to be intermediates “between subsp. douglasii and subsp.
johnstonii” (Hickman, in herb.).
The stems of Polygonum sawatchense subsp. oblivium are scaberulous-pap-
illose (Fig. Ib, 2b), the leaves are oblong-elliptic to elliptic (Fig. 1b), papillose-
denticulate on the margins (Fig. 2 d) and the flowers, at least some of them, are
open. Polygonum sawatchense subsp. sawatchense is entirely glabrous, the leaves
are linear to oblanceolate (Fig. la, Fig. 2a, c), and flowers are normally closed.
The tepal surface pattern of the two taxa is also different (Costea & Tardif, un-
published). Both subspecies occur in California, Idaho, Nevada and Washing-
ton. However, P. sawatchense subsp. sawatchense is more widely distributed,
extending from Mexico, Arizona and New Mexico to Utah, Wyoming, Montana,
South Dakota, North Dakota and Canada - Alberta and Saskatchewan. Although
the two taxa are usually distinct, intermediate variation patterns may occur in
the areas of sympatry, and this is w
ry we treated them as subspecies. For ex-
ample, glabrous plants with more or less broad and gradually decreasing leaves,
as well as papillose-scaberulous plants with relatively narrow leaves, may be
occasionally encountered.
Munz, who described P. douglasii var. johnstonii (1935), observed that this
was diflerent from P sawatchense subsp. oblivium, but he usually identified the
latter plants as P.austiniae (e.g. Munz 18361, RSA). Polygonum sawatchense subsp.
oblivium dilfers from P. austiniae through its erect pedicels and much larger
achenes. Polygonum sawatchense subsp. oblivium has also been
brea - Cement en Or
requently con-
fused with P minimum. They differ in respect to the morphology of leaves and
ocreae, stems, size of fruiting perianth and achenes, as well as the pattern of
COSTEA AND TARDIF, TAXONOMY OF POLYGONUM SAWATCHENSE 1639
epicuticular wax sculpture of tepals (Costea & Tardif, unpublished). In par-
ticular, P minimum has leaves that are hardly reduced distally, funnelform
ocreae, 1-4 mm long, papillae on stems and ocreae are conical, short, with a
swollen base, and flowers and achenes are usually much smaller.
Representative specimens examined. CANADA. BRITISH COUUMBIA: Road up to Windy Joe, Man-
ning Park, roadside, +800 [t, + Aug 1957, K. Beamish 7856 * UBC). U.S.A. CALIFORNIA. EI Do-
rad neadow near Fallen Leaf Lake, 21 Jun 1920, A.M. ae 822 (NY). Humboldt Co.: Grouse
Mountain, nal moist flat, 5000 ft, 30 Aug 1933, J.P. Tracy 13023 (UC); 14 Jul 1935, |.P. Tracy 14042
(UC). Modoc Co.: common in dry forest openings; with Abies concolor; Northern Juniper Woodland,
7 mi as of Fort Bidwell, 6150 ft, 7 Jul 1955, RJ. Weatherby 1608 (RSA). Nevada Co.: Castle Peak, 31
Jul 1903, A.A. oe SM. GH*, NY, RSA). Plumas Co.: Scattered annual growing in volcanic grave
with Purshi lentata; Red fir forest, Bear Ridge near ea ake; 7200 ft, 2 Jul 1955, RJ. Weatherby
1570 (RSA); ca. 0.5 mi “ ow Red Rock lookout, ca. 8 mi S of Susanville, bare reddish gravel and out-
crop between road and cliff, ca. 7000 ft, 3 Jul 1989, B. eae 8642 (UC). Siskiyou Co.: rocky summit,
ridge road to Deadwood Lookout, mountains west of Yreka, 25 Jun 1952, M. Ownbey & SJ. Preece, Jr.
3345 (NY*); Salmon Mountains, open site northeast side of creek bed, Tom Taylor cabin, 6700 ft, 11
Aug 1968, I: W. Oettinger 471 (RSA): Paynes Springs, 2 mi SE from Medicine Lake, sandy shallow-soil
basins on recent lava flows situated in Abies magnifica lorest, 6550 ft, 26 ieee ae D.W. Taylor 9075
(UC); Grass Lake, 18 Jul 1948, H.L. Mason 13623 (RSA*, UC). Truckee Co.: 1/2 mi W of Lake Van Norden,
7000 [t, 22 Jun 1934, N. French 478 (RSA). IDAHO. Elmore Co.: W side Lava oo at headwaters
of Russell Gulch, Boise National Forest N of Prairie, ead soil on granite, dry shade of forest,
7500 ft; 15 Jul 1981, B. Ertter & |. W. Grimes 4531 ( Valley Co.: Dry slope 3 mi N and W
Payette Lake, 6000 ft, 25 Jun 1946, CL. Hitchcock & C. V Muhlick 13971(NY). NEVADA. Elko Co.: among
rock outcrops on hillsides above Maggie Summit, Bull Run mountains; with Prunusand Amelanchier,
6700 ft; 20 Jun 1980, J.W. Grimes et al. 1646 (NY*). Humboldt Co.: Sonoma Range, Water Canyon,
sandy steep E facing slope with sagebrush; frequent, 6800 [t, 10 Jul 1964, N.H. Holmgren & |.L. Reveal
1374 (NY*); Jackson Range, ca. 50 mi WNW of Winnemucca; halfway up road to mines at head of
aie Creek, dry NW facing slope between streamlet oS rimrock outcrops; 2100 m, 14 Jun 1987, B.
rtter 7146 et al. (RSA, UC). OREGON: Woods. Achoco Forest, 22 Jun 1932, M.E. Peck 17068 (NY*).
Grant Co.: Malheur National Forest; Straw ee ae iin Wilderness, on Strawberry Mountain,
woods of Pseudotsuga, Abies grandis, Larix and Pinus, 6500 ft, locally abundant on dry slopes, 12
July 1973, M.F. Denton 3188 (US); Aldrich deers slopes between Fields Creek summit and Fields
Peak, 5500 ft, 20 Jul 1962, A.R. Kruckeberg 5465 (UC). Josephine Co.: common in a neglected field,
summit of the Siskiyou shee Da pine cet 4500 tt,.21 jum 1922, AvAy Heller sade):
Oregon Caves, 6000 ft, 24 Jun 19 ©. Parks 4892 (UC). Lake Co.: dry flat under aspens and pines
near meadow. Willow Creek eae camp, Warner ee ains; 3800 [t, 24 Jun 1973, PA. Munz 18361
RS ew Co.: Juniper Springs, 1350 m, 15 Jun 1896, J.B. Leiberg 2275 (UC). WASHINGTON.
ou
open Se pes es along ridge leading N of Big Butte on the breaks of the Grand Ronde River,
acl ct, oy 939, Meyer 1622 (GH*). Okanogan Co.: lower wooded slopes of Billy Goat Moun-
tain, 4000 ft, 10 Jul oe W. Thompson 10891(NY). Wallowa Co.: Ice Lake Trail, Wallowa Mountains,
26 Jun 1936, A. Eastwood & J.T. Howell (NY).
ACKNOWLEDGMENTS
We are much indebted to John McNeill and Guy Nesom for their helpful criti-
cism and suggestions on earlier drafts of this paper. We warmly thank all cura-
tors of the mentioned herbaria for the prompt preparation and expedition of
the loans. Giles Waines and Margriet Wetherwax helped us with some old ref -
1640 BRIT.ORG/SIDA 20(4)
erences from California, and Laura Echarte checked our Spanish translation of
the abstract.
REFERENCES
Hickman, J.C. 1984. Nomenclatural changes in Persicaria, Polygonum and Rumex
Polygonaceae). Madrono 31:249-252.
Hickman, J.C. 1993. Polygonaceae. In: J.C. Hickman, ed. The Jepson manual: higher plants of
California. University of California Press, Berkeley. Pp: 854-895,
Munz, PA. 1935.A manual of southern California botany. Claremont Colleges, Claremont,
CA.
Munz, PA. and A.D. Keck. 1968. A California flora. University of California Press, Berkeley.
Smait, J.K. 1893. Further notes on American species of Polygonum. Bull. Torrey Bot. Club
20:213-217.
Smatt, JK. 1894. New and interesting species of Polygonum. Bull. Torrey Bot. Club 21:
168-173.
fled:
TY PIFICATION AND NOMENCLATURAL HISTORY OF
TRACHELOSPERMUM DIFFORME (APOCYNACEAE)
Alexander Krings
Herbarium, Department of Botany
North Carolina State University
Raleigh, NC 27695-7612, U.S.
Alexander_Krings@ncsu.edu
ABSTRACT
This study d f 1 with ing vine Trachelospermum
difforme. Although not ated: in the literature, a holotype of Echites difformis i Waltek was discovered
in the Thomas Walter herbarium of the Natural History Museum, London. Types and nomenclatural
history are discussed for other synonyms and combinations.
RESUMEN
Sed | iad la enredadera Trachel liffor Aunque
no citado en la | Tea fue descubier to un aE DO}On DS en a Hevbarie: de Thom as Wa ler en vel Mitse0
de Historia Natural, Londres. Sed
i rE
The twining vine Trachelospermum difforme (Walter) A. Gray (Apocynaceae)
is native to the eastern United States and was first described as Echites difformis
Walter in 1788. However, Walter did not indicate specimens he had studied. His
herbarium is housed at the Natural History Museum (BM), London, and photo-
graphs thereof may be found in the collections of the Gray Herbarium (GH) at
Harvard University. Walter’s herbarium contains 122 sheets, a great many of
which are plants of his Flora Caroliniana (Walter 1788; Stafleu & Cowan 1988).
Although numerous authors proposed new combinations based on Walter's
Echites difformis, none cited an appropriate type specimen. As typification is
an ICBN requisite (Greuter et al. 2000), this study seeks to document types for
names associated with the species Trachelospermum difforme.
In his treatment of Forsteronia, Alphonse de Candolle (1844) cites two
specimens for the entity Trachelospermum difforme: “in Virginia infer. (Pursh)
Carolina! et Mississippi (Torr! in h. Boiss.).” However, John Torrey was not born
until 1796 and Frederick Pursh visited North America for the first time in 1799—
well after Walter’s Flora had been published (Stafleu @ Cowan 1983, 1986). For
his new combination based on Echites difformis, John Miers (1878) cited a
Florida specimen (“vs. in herb. Mus. Brit. Florida (Chapman)”). However, this
specimen also cannot be considered as Walter's type as it was collected by Alvan
Chapman (1809-1899) in 1845, many years after Walter’s death in 1789. Lack-
ing further citation in the literature of potential types seen by Walter, a search
of the Walter herbarium at BM was undertaken by the author. Fortunately, the
SIDA 20(4): 1641-1644. 2003
1642 BRIT.ORG/SIDA 20(4)
search revealed a specimen of the taxon (Walter 215)—which should thus be
considered the holotype of Echites difformis Walter.
In 1803, André Michaux proposed the name Echites puberula in his Flora
Boreali Americana for the taxon in question. Three specimens that can be con-
sidered types for this name reside at the Muséum National d'Histoire Naturelle
(P) in Paris. One specimen exhibits flowers in bud and a handwritten note
“Foresteronia difformis A. DC.” Another specimen exhibits fully opened flow-
ers (but no labels) and the last exhibits fruiting material. From Michaux’s
protologue it is clear that both [lowering and fruiting materials were available
—:
to him at the time of description.
In 1817, Poiret published the name Iabernaemontana populifolia and indi-
rated Seeing a specimen in “herb. Desfont.” Poiret’s name has long been cited in
synonymy with Trachelospermum difforme. However, Poiret’s type at Fl-W does
not belong to Irachelospermum (stems stout; lamina decurrent onto petioles).
Recent annotations suggest it belongs to Logania populifolia (Lam.)
Leeuwenberg (see also Leeuwenberg 1991). A small sachet (Scm x 5cm)of likely
the same material resides at P, but bears no indication of locality (Aymonin, per.
comin.). As the type belongs to another taxon, Tabernaemontana populifolia
Poir. is here excluded from synonymy with Trachelospermum difforme (Walt.)
A. Gray.
Miers, in proposing a new combination based on
Thyrsanthus populifolius (Poir) Miers., Apocyn. S. Amer. 99. 1878.), indicated
seeing a specimen collected by “Marshal [sic!’ from Georgia in “herb. Mus. Brit.”
This specimen is indeed Trachelospermum difforme (BM! and not Logania
populifolia, however Miers’ combination cannot be applied to the former spe-
cies as ICBN Art. 7.4 holds that new combinations, in all circumstances, are
typified by the type of the basionym (Greuter et al. 2000).
In 1836, Rafinesque proposed the name Echites salicifolia. However, this
name isa nomen illegitum since Echites salicifolia Willd. ex Roem. & Schult.
(=Allamanda cathartica L., Apocynaceae) was validly published in 1819
(Systema Vegetabilium 4:796). No type specimen could be located for
Ralinesque’s name. Unfortunately, as Stuckey (1971) notes, Rafinesque’s her-
y destroyed by rats after his death and virtually all the re-
rites salicifolia Raf. is not
Poiret’s name (1.e.,
barium was partial]
maining material was discarded by Elias Durand. Ec]
listed among Rafinesque’s North American vascular plants at PH (Stuckey 1971),
nor are potential type specimens found in the collections of GH and NY. In-
quires with P and DWC also provided no material.
TAXONOMIC SUMMARY
Trachelospermum difforme (Walter) A. Gray, Syn. Fl. N. Amer. 2:85. 1878. ae
difformis Walter, Fl. Carol. 98. 1788. Forsteronia ¢ lifformis (Walter) A.DC., Prodr
37.1844. Secondatia difformis (Walter) Benth. @ Hook.f., Gen. Pl. 2:710, 1376
pay
KRINGS, NOMENCLATURE OF TRACHELOSPERMUM DIFFORME 1643
nom. invalid. Thyrsanthus difformis (Walter) Miers, Apocyn. S. Amer. 99:1878.
Thyrsanthella difformis (Walter) Pichon, Bull. Mus. Natl. Hist. Nat. ser. 2, 20:192.
48. Type: Walter 215 (HT:BM)).
Echites puberula Michx., Fl. Bor. Amer. 1:120. 1803. Type: Michaux s.n. [protologue: “in sylvis
Carolinae inferioris”| (ST: P; IDC Fiche! [three specimens: one with flowers in bud, one with
flowers, and one with fruits]).
Echites salicifolia Raf., New FL N. Amer. 4:59. 1838 (1836’), nom. illeg.; non Willd. ex Roem. &
Schult. 1819. TyPE: unknown.
ACKNOWLEDGMENTS
I thank the curators and staff of the following herbaria for their kind assistance:
BM (particularly V. Noble), DWC, FI (particularly E. Luccioli for digitizing im-
ages of Poiret’s type), GH, NY, P (particularly G.G. Aymonin), and PH; the staff
of the NC State University, Duke University, and University of North Carolina
Libraries; Paul Fantz; and Wendy Worley. K. Gandhi and J. Bauml provided
thoughtful reviews of a previous version of this manuscript.
REFERENCES
DeCANDOLLE, A.P. 1844. Apocynaceae. Prodromus systematis naturalis regni vegetabilis
8:317-489
Gray, A. 1878. Synoptical flora of North America. Ivison, Blakeman, Taylor, and Co., New
York.
GreUTER, W., J. MCNEILL, FR. Barrie, H-M. Burbert, V. DEMOULIN, T.S. Fitcueiras, D.H. NICOLson, PC. Sitva,
J.E. Skoa, P. TReHANE, NJ. TurtAno, and D.L. Hawkswortu. 2000. International Code of Botani-
cal Nomenclature. Regnum Veg. 131. Koeltz Scientific Books, KOnigstein
HARRIMAN, N.A. 1981.A generic index to the IDC microfiche (#6211) of the Michaux her-
barium arranged according to the alphabetical generic index of Michaux's Flora Boreali-
Americana, fascimile of the 1803 edition, with many added generic names. Published
by the author, Oshkosh.
LeeuwenserG, A.J.M.1991.A revision of Tabernaemontana. Royal Botanic Gardens, Kew.
Michaux, A. 1803. Flora Boreali-Americana, vol. 1. Caroli Crapelet, Paris.
Miers, J. 1878.On the Apocynaceae of South America. Williams & Norgate, London.
PicHon, M. 1948. Classification des Apocynées: XV, genres Trachelospermum, Baissea et
Oncinotis. Bull. Mus. Natl. Hist. Nat. 2, 20:190-197.
Poiret, J.L.M. 1811. Tabernemontana. Encyclopédie Méthodique, Botanique, suppl. 5:276.
RaFiNesaue, C.S. 1836. New flora and botany of North America. Murray Printing Co. (Photo-
lithographers), Cambridge.
Stareu, F.A.and R.S.Cowan. 1976. Taxonomic literature: A selective quide to botanical pub-
lications and collections with dates, commentaries and types, Vol. |: A-G. Bohn,
Scheltema, and Holkema, Utrecht.
StaFveu, F.A.and R.S. Cowan. 1981.Taxonomic literature: A selective guide to botanical pub-
lications and collections with dates, commentaries and types, Vol. |: A-G. Bohn,
Scheltema, and Holkema, Utrecht.
1644 BRIT.ORG/SIDA 20(4)
Starceu, F.A.and R.S. Cowan. 1983. Taxonomic literature: A selective guide to botanical pub-
lications and collections with dates, commentaries and types, Vol. IV: P-Sak. Bohn,
Scheltema, and Holkema, Utrecht.
Starteu, E.A.and R.S. Cowan. 1986. Taxonomic literature: A selective guide to botanical pub-
lications and collections with dates, commentaries and types, Vol. VI: Sti-Vuy. Bohn,
Scheltema, and Holkema, Utrecht.
Starveu, FA.and R.S. Cowan. 1988. Taxonomic literature: A selective guide to botanical pub-
lications and collections with dates, commentaries and types, Vol. Vil: W-Z. Bohn,
Scheltema, and Holkema, Utrecht.
Stuckey, R.L. 1971. C.S. Rafinesque’s North American vascular plants at the Academy of
Natural Sciences of Philadelphia. Brittonia 23:191-208.
Water, |. 1788. Flora Caroliniana. Murray Printing Co. (Photolithographers), Cambridge.
NEW NAMES AND NEW COMBINATIONS
IN THE FLORA OF HISPANIOLA
Henri Alain Liogier
Botanical Research Institute of Texas
e
Fort Worth, TX 76102-4060, U.S.A.
In revising the published Flora of Hispaniola (1981, 1989, 1994, 2000), I have
found a need to make revisions in nomenclature of some taxa. Some names were
antedated by previous publications, and this came to my attention when I was
verifying publication dates through the W3TROPICOS (2003) database on the
web page of the Missouri Botanical Garden.
The following changes are made to correct inadvertent errors | have made
for lack of complete bibliographies in the past. | am very grateful to
W3TROPICOS for enabling me to catch these errors. | also want to thank the
Botanical Research Institute of Texas, for the use of its library and the facilities
given to me for my research. The National Botanic Garden in Santo Domingo
has a very good plant collecting program underway, along with a number of
detailed florulas.
ASCLEPIADACEAE
Matelea monticola Alain, nom. nov. Matelea sylvicola Alain, Phytologia 22:168. 1971,
non Matelea sylvicola L.O. Williams, Fieldiana, Bot. 32:57. 1968
CELASTRACEAE
Maytenus urbanii Alain, nom. nov. Maytenus impressa Urb. & Ekm., Ark. Bot. 22
A(8):67. 1928, non Maytenus impressa Reissek, Fl. Bras. 11(1):29. 1861.
MELASTOMATACEAE
Leandra urbaniana (Alain) Alain, comb. nov. BASIONYM: Ossaea urbaniana Alain,
Brittonia 20:158. 1968. Ossaea polychaeta Urb. & Ekm., Ark. Bot. 23A()):27. 1931.
non Ossaea polychaete Urban & Ekm., Ark. Bot. 22A(17):60. 1929. Leandra polychaeta (Urb. &
Ekm.) Alain, Sida 18:1026. 1999, non Leandra polychaeta Cogn. in A. DC. & C. DC, Monogr.
Phan. 7:1186. 1891
Sagraea rugosa Alain, nom. nov. Melastoma scabrosum L., Syst. Nat. ed. LO2):1022.
1759. Ossaea scabrosa (L.) DC., Prodr. 3:169. 1828. Clidemia scabrosa(L.) Griseb., Mem. Amer.
Acad. ns. 8:184. 1861. $ ] (L.) Alain, Moscosoa 8:7. 1994, non Sagraea scabrosa
oO
Naud., Ann. Sci. Nat. Bot., ser. 3, 18:97. 1852, nec Seem., Bot. Voy. Herald 124. 1854.
SIDA 20(4): 1645-1646. 2003
1646 BRIT.ORG/SIDA 20(4)
REFERENCES
Liocier, AH. 1981- La Flora de la Espanola, Vol. 1. Phytologia Memoirs Ill.
Liocier, AH. 1989. La Flora de la Espanola, Vol. 5.San Pedro de Macoris, Dom. Republic.
Liocier, AH. 1994. La Flora de la Espanola, Vol. VI, San Pedro de Macoris, Dominican
Republic.
Liocier, A.H. 2000. La Flora de la Espanola, Vol. 9. Jardin Botanico Santo Domingo.
Tropicos. 2003. Click on w3TROPICOS (www.mobot.org) to access the taxonomic
database.
GLANDULARIA VERECUNDA (VERBENACEAE):
SPECIES OR FRUITFORM?
B.L. Turner
University of Texas at Austin
Plant Resources Center
I University Station FO404
Austin, TX 78712-0471, U.S.A.
In the most recent issue of Sida, Henrickson (2003) treated Glandularia
verecunda Umber as a distinct species, this in sharp contrast to the present
author's treatment of the taxon as but a fruit-form of G. quadrangulata (Heller)
Umber (Turner 1999). My reason for this disposition followed several years of
field work in which I could find no differences between the two hypothetical
species except that of a terminal appendage on the apex of G. verecunda, this
nicely illustrated by Henrickson’s Fig. 1.
Glandularia verecunda was first proposed by Umber (1979), the species
name said to be derived from the Latin word verecundus or “bashful,” presum-
ably in reference to the taxon’s proclivity to go unnoticed, both in the herbarium
and in the field. Indeed, Umber’s description of the taxon nearly matches that
of G. quadrangulata, except for the terminal appendage on the fruit. Turner
(1999) clearly noted that Umber himself remarked that G. verecunda is “indis-
tinguishable from G. guadrangulata except for the seeds,” meaning the pres-
ence of an appendage on the nutlets.
Glandularia verecunda and G. quadrangulata are partially if not largely
sympatric over most of their distributions (not “largely allopatric” as stated by
Henrickson, his use of the term sympatric apparently restricted to
intrapopulational occurrences of the two taxa).
fter a discussion “Of the three nomenclatural possibilities: 1) being in-
consequential forms of each other; (2) being geographical subunits of a species,
ie. subspecies or varieties of one species; or (3) being separate species that merely
co-occur in some areas,” Henrickson surmises that the two taxa are distinct spe-
cies “that are very similar vegetatively but can co-occur without interbreeding
as no intermediates have been found.” Immediately thereafter, he concluded
his discussion with a terse paragraph, as follows:
“If there was evidence of interbreeding, they [sic] the two taxa could be recognized at the vs
cific level. But this does not exist. Therefore the two taxa will here be recognized as distinct species.
Henrickson comments further that “No plants have ever been discovered in
j herr ae,
nature with nutlets intermediate in structure between those of G. quadrangulata
and G. verecunda ... Furthermore, no artificial hybridization has been done to
SIDA 20(4): 1647-1648. 2003
1648 BRIT.ORG/SIDA 20(4)
determine the condition that would be expressed in a hybrid.” Needless to say,
this same comment might be applied to most any albino form occurring in natu-
ral populations; nor does one spend much effort seeking out an intermediate
plant to prove the point!
Fruit forms (or fruit polymorphism) in natural populations are relatively
rare, but are relatively common in some groups such as, for example, the genus
Valerianella (Valerianaceae). The interested reader is referred to the work of
Ware (1983), who detailed numerous fruit forms in Valerianella,several of which
had long been recognized as species. For those interested in the genetics of such
fruit types, the work of Xena de Enrech and Mathez (1998) might be consulted.
Fruit forms also occur in other Texas plants, for example in the genus
Thelesperma (Asteraceae). The species T. curvicarpum Metchert, first described
in 1963 from Burnett and Coleman counties, Texas, was subsequently shown to
be but a fruit-form (Greer 1997). And it is likely that my description of
Cynoglossum hintoniorum Turner (Boraginaceae) amounts to no more than a
ruit-form of the widely distributed C. amabile Stapf & Drumm. (Turner 1995).
Finally, it should be reemphasized that | have called to the fore in the above-
mentioned publication several populations of G. quadrangularis in which
appendaged and nonappendaged fruits occur intermixed. The two forms can
be separated only by the appendages on their nutlets, but | have no trepidation
in recognizing this fruit-form as a named taxon, as follows:
ane)
Glandularia quadrangulata forma albida (Moldenke) B.L. Turner, comb. nov
BASIONYM: Verbena pumila forma albida Moldenke, Phytologia 9:188. 1963
= Glandularia verecunda Umber
Umber (1979 treated the forma as a synonym of his concept of G. verecunda. It is
unfortunate to have to take up the name “albida’ for this fruit-form, but the Code of
Botanical Nomenclature mandates use of the earliest name in rank as having priority.
REFERENCES
Greer, LF. 1997. Thelesperma curvicarpum (Asteraceae), an achene form in populations of 7.
simplicifolium var. simplicifolium and T.filifolium var. filifolium. Southw.Naturalist 42:242-244,
Henrickson, JH. 2003. Studies in Glandularia (Verbencaeae): the Glandulaia quadrangulata
complex. Sida 20:953-967
Turner, B.L. 1995. A new species of Cynoglossum (Boraginaceae) from Oaxaca, Mexico.
Phytologia 79:306-308.
Turner, B.L. 1998. Texas species of Glandularia (Verbenaceae). Lundellia 1:3-16.
Umeer, R.E.1979. The genus Glandularia (Verbenaceae) in North America. Syst. Bot.4:72-102.
Ware, D.M. 1983. Genetic fruit polymorphism in North American Valerianella
(Valerianaceae) and its taxonomic implications. Syst. Bot. 8:33-44.
XENA DE Enrich, N.and J. MatHez. 1998. Genetic control of fruit polymorphism in the genus
Fedia (Valerianaceae) in the light of dimorphic and trimorphic populations of F
pallescens. Pl. Syst. Evol. 210:199-210.
NEW COMBINATION IN ASTEREAE (ASTERACEAE)
Guy L.Nesom Harold Robinson
Botanical Research Institute of Texas NMVNH - Botany Section
509 Pecan Stre t Smith nian Inctitttion
Fort Worth, TX 76102-4060, U.S.A. Box 37012
Washington, DC 20013-7012, U.S.A.
The following combination is necessary to complete a set of subtribal names
for the developing treatment of Asteraceae tribe Astereae (Asteraceae) by Guy
Nesom and Harold Robinson for “The Families and Genera of Vascular Plants”
(Vols. I-V, 1990-2003: K. Kubitzki, General Editor; J.W. Kadereit and C. Jeffrey,
Volume Editors for Asterales).
Asteraceae tribe Astereae subtribe Bellidinae (Cass. ex D. Don) G. Nesom & H.
son, comb. et stat. nov. BASIONY™: Asteraceae tribe Bellideae Cass. ex D.
Don in R. Sweet, fae FL. Gard. 1: ad t. 38. 1830. TYPE: Bellis L.
REFERENCE
Kusirzki, K.(ed.). 1990+. The families and genera of vascular plants. 5+ vols. Springer-Verlag,
Berlin.
SIDA 20(4): 1649, 2003
1650 BRIT.ORG/SIDA 20(4)
TIMBER PRESS
Horticulture/Gardening
Ropert Noip. 2003. Columbines: Aquilegia, Paraquilegia, and Semiaquilegia. (ISBN
0-88192-588-8, hbk.). Timber Press, Inc. 133 S.W. Second Ave, Suite 450,
Portland, OR 97204-3527, U.S.A. (Orders: www.timberpress.com,
mail@timberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax).
$24.95, 193 pp., +5 color photos, 8 watercolors, 4 line drawings, 6 x 9".
Contents.—Pretace, Acknowledgments, Cultivation, Pests and Diseases, Propagation, Morphology and
Distribution, Doves and Eagles, Taxonomic Tangles. Some Columbine Cousins, The Species, Hybrids
and Curiosities, Sources of Seed, Glossary, Bibliography, and Index. Excellent photography and the
watercolors are beautiful.
Lewis J. Marrurws. 2002. The Protea Book: A Guide to Cultivated Proteaceae.
(ISBN 0-88192-553-5, pbk.). Timber Press, Inc. 133 S.W. Second Ave, Suite
450, Portland, OR 97204-3527, U.S.A. (Orders: www.timberpress.com,
mail@timbet press.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax).
$34.95, 184 pp., 286 color photos, 6 x 9",
Contents. —Acknowledgments, Introduction, Cultivating proteas, Proteas in the garden, Commercial
cultivation, Identification, Glossary, References, and Index. This beautiful 184-page guidebook cov-
ers most of the cultivated members of Proteaceae family not just the genus Protea.
Ethnol ical/E ic/Medicinal/Useful Plants/Toxic
JEAN H. LANGENHEIM. 2003. Plant Resins: neue Evolution, Ecology, and Eth-
nobotany. (ISBN 0-88192-574-8, pbk.). Timber Press, Inc. 133 $.W. Second
Ave, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders:
www.timberpress.com, mail@timberpress.com, 503-227-2878, 1-800-327-
5680, 503-227-3070 fax). $49.95, 612 pp., 47 color photos, 30 b/w photos,
70 line drawings, 4 maps, 4 tables, 7 3/8" x 10 3/8".
Contents.—Preface, ancien 1) What Plant Resins Are and Are Not, 2) Resin-Producing
Plants, 3) How Plants Secrete and Store Resin, 4) Amer: Resins Through Geologic Time, 5) Ecol ogical
Roles of Resins, 6) The eiinet botany of Resins, 7) Oleoresins, 8) Fragrant and Medicinal Balsams, 9)
Varnish and Lacquer Resins, 10) Miscellaneous Resins, and 11) Future Use of Resin. Appendix 1:
Xesin-Producing Conifers, Appendix 2: Resin-Producing Angiosperms, Appendix 3: Skeletons of
Characteristic Components of Fossil Resins, Appendix 4: Age, Location, and Plant Source of Amber
Deposits, Appendix 5: Common Names, Plant Sources, and Uses of Resins. Glossary, References, Plant
Index, and Subject a
Ic recked to see if Paul t (S ppl 1 Dragon Tree) wa li | Sl producing
angiosperm and it was not. The leaves emcee a very sticky resinous substance aie tis hard to re-
move from the hands.—Barney Lipscomb, Botanical Research Institute of Texas, 509 Pecan St, Fort
Worth, TX 76102-4060, U.S.A., barney@britorg.
SIDA 20(4): 1650. 2003
ANATOMIA COMPARADA DE LA EPIDERMIS FOLIAR
EN EL. GRUPO DE ESPECIES SOLANUM SESSILE,
DE LA SECCION GEMINATA (SOLANACEAE)
Carmen Benitez de Rojas y Chiara Berlingieri
Universidad oli a ibaa
LCariiltad de Agr nomin | tanica Aartcola
eh
Prolonga ion Avenida 19 de Abril,
Area Universitaria, Apartado Postal 4579
Maracay. Edo. Araqua, VENEZUELA
cbenitez@telcel.net.ve berles@cantv.net
RESUMEN
= mean y deserinel = ca Raclesistieas de la epidermis foliar en amas SUpentlcles de oe taxones
Bitter, S. pene Benitez, S. appositifolium Ruiz & Fav, S. sessile Ruiz & Pav, S. triste Jacq y S.
turgidum S. Knapp. Se utili 1 cm?de la porcién media de hojas
secas (especimenes de herbariolie o frescas y eee en FAA al 70%. Fueron aplicadas técnicas
anatomicas eonvencionales per macerado y tinciOn. Se evaluan las siguientes variables: presencia,
tipo dimensiones y ul
tipo, longitud, ubicacion my densidad de tricomas;
forma y grosor de la pared de las células epidermicas normales e indice estomatico. Los resultados
muestran diferencias en algunas de las car as anatomicas consideradas, especialmente las
referidas a los tipos tricomaticos y a la ubicacion de estomas. Dichos resultados se comparan y han
permitido la separacion de las especies mediante una clave.
ABSTRACT
pnee character istics of leaf blades were studied and described from taxa belonging to the spe-
g Solanum sessile represented in Venezuela by the following species: Solanum
prendre Bitter, S. obovalifolium Benitez, S. oppositifolium Ruiz & Pav, S. sessile Ruiz & Pav, S.
triste Jacq. and S. turgidum S. Knapp. Middle portions of leaf blades were sampled from herbarium
specimens and from fresh sie material fixed in FAA 70%. Conventional anatomical procedures for
macerations and staining were used. The following criteria were evaluated: presence, type, dimen-
n and location of stomata; presence, type, length, location and density of trichomes; shape and
thickness of the wal Mor ae epidermal aes and stomatal index. Results show there are differ-
lly with regards to trichome ty
and the location of stomata. These characteristics ¢: can be Rea to eet collections of Solanum to
the species level. A key to the species of Solanum section Geminata using these characters is pre-
sented.
La Secci6n Geminata del género Solanum con 122 especies acomodadas en 16
grupos (Knapp 2002), es una de las secciones dentro del género con mayor
numero de especies, localizadas principalmente en regiones tropicales de
América del Sur y con unas pocas especies representadas en el Viejo Mundo; en
Venezuela estan presentes 34 especies constituyendo el 23% del total mundial
SIDA 20(4): 1651-1661. 2003
1652 BRIT.ORG/SIDA 20(4)
y el 32% del total de especies del género Solanum en Venezuela (105). Dichas
especies estan ampliamente distribuidas, principalmente en selvas humedas
primarias de los pisos meso y microtémicos y en selvas semideciduas y
secundarias de los pisos macrotérmicos (Knapp 2002).
Las especies que conforman esta seccion exhiben caracteres morfol6gicos,
que en términos generales pueden ser utilizados para el reconocimiento de las
mismas como miembros de la seccién:; tales son su condicién de arbustos o
arboles, inflorescencias caracteristicas del tipo cimas helicoidales naciendo
Opuestas o alternas a una hoja que puede ser geminada o no. La importancia
taxonomica de caracteres anatomicos foliares ha sido ampliamente reconocida
por muchos investigadores (Solereder 1908; Metcalfe y Chalk 1950; Carlquist y
Holt 1961).
Entre dichos caracteres, los estomas que varian en forma, numero,
orientacion y arreglo de las células subsidiarias que los rodean, han
desempenado un papel importante en los procesos evolutivos de las especies
vegetales, desde el punto de vista anatomico y taxondmico, lo cual ha justificado
su estudio por lineas diferentes y en diversas familias, como lo indicado por
Ahmad (1964) quien realizando estudios epidérmicos en 28 especies
pertenecientes al género Solanum, encontr6 mayor frecuencia estomatica en
especies de dicho género comparado con otros géneros, reportando anomalias
estomaticas en algunas especies tales como en: S. macranthum Dunal, Solanum
nigrum L., y Solanum decorticans Sendtn.
En 1979 Bessis y Guyot estudiaron estomas en miembros de algunas tribus
de las Solanaceae, concluyendo que la distribucion de tipos de estomas puede
ser utilizada para establecer divisiones sistematicas y sugerir relaciones
filogenéticas entre los miembros de dicha familia. Rashib y Beg (1989)
elaboraron una clave foliar, como herramienta taxonémica, para algunas
especies de la familia Solanaceae, en la que incluyeron caracteristicas de estomas
y tricomas. También los tipos tricomaticos asi como su densidad y frecuencia,
han sido de alto valor diagnéstico para la definicién de las especies, caracter
que ha sustentado la clasificacion infragenérica dentro de Solanumestablecida
por Seithe (1962, 1979). Dada la utilidad de los caracteres anatomicos foliares
como rasgos de valor diagnéstico, en el presente trabajo se estudian y describen
las caracteristicas de la epidermis foliar en ambas superticies, de los taxones
del Grupo de especies Solanum sessile representados en Venezuela a saber
Solanum chlamydogynum Bitter, S. obovalifolium Benitez, S. oppositifolium Ruiz
S& Pav, S. sessile Ruiz & Pav, S. triste Jacq. y S.turgidum S. Knapp. Los caracteres
evaluados y los resultados obtenidos se comparan intentando extraer
informacion anatomica destacable entre las especies.
MATERIALES Y METODO
rs
El material estudiado procede de colecciones de herbario y de aquel fresco fijado
BENITEZ Y BERLINGIERI (A : a
en FAA, segtin como se enumera en el Anexo 1. Se utilizaron segmentos de
aproximadamente1em2 de la porcion media de hojas pertenecientes a exsiccatas
del Herbario Victor Manuel Badillo, Universidad Central de Venezuela (MY) 0
fijadas directamente en FAA al 70% (Formalina, acido acético glacial, etanol
70%). Enel primer caso, previo al estudio anatomico, las secciones de hoja fueron
hidratadas sumergiéndolas en agua hirviente por 5-10 min. Para estudiar las
caracteristicas de la epidermis foliar, se realizaron macerados de la porci6n
media de la lamina, colocando segmentos de aproximadamente 1 cm (que
incluyeran nervaduras) en una soluci6n de hipoclorito de sodio 5,25% (cloro
comercial) 0 en acido nitrico 5%-acido crémico 5% (1:1), en una estufa a 56-58"
C. Luego se realizo un lavado profuso de la epidermis con agua destilada y se
tind con azul de toluidina acuosa 0.5%, para finalmente montarla en agua-
glicerina (1:1).Las laminas se observaron y fotografiaron con un microscopio
binocular marca Leitz. Ademas, se realizaron algunas mediciones con la ayuda
de un ocular micrometrado. La caracterizacion cualitativa y cuantitativa se ha
obtenido mediante el analisis de las siguientes variables: Presencia, tipo,
dimensiones y ubicacién de estomas; presencia, tipo, longitud, ubicacion y
densidad de tricomas; forma y grosor de la pared de las células epidérmicas
normales, etc. Ademas se determin6 el indice estomatico (LE) mediante la
siguiente formula establecida por Salisbury (1927)
n’ de estomas (S)
LE= x LOO
n’ de estomas (S) + n° de células epidérmicas (E)
RESULTADOS
Solanum chlamydogynum Bitter
Las células de la epidermis adaxial tienen contornos poligonales (Fig. 1) y
paredes celulares ligeramente engrosadas, mientras que las de la epidermis
abaxial poseen contornos poligonales a ligeramente ondulados y paredes
delgadas (Fig. 2). Hoja anfiestomatica; estomas de la epidermis adaxial
principalmente del tipo anisociticos (largo 25,03 um y ancho 14,67 wm),
ubicados proximos y sobre las nervaduras (Fig. 1), aunque pueden aparecer
algunos aislados en la lamina. En la epidermis abaxial los estomas son con mayor
frecuencia anisociticos (LE.= 19,78-22,51); largo 19,27-23,2 um y ancho 14,83-
17,2 um). Presencia de tricomas protectores y glandulares en ambas epidermis.
En la epidermis adaxial, los tricomas protectores aparecen simples y ramificados
sobre la lamina y nervaduras en muy bad densidad (0,22/mm2); los protectores
simples son multicelulares uniseriados con apice agudo y con longitud y numero
variable de células. Los protectores ramificados (multicelulares) en su mayoria
tienen dos ramas que emergen al mismo nivel (una mas desarrollada que la otra),
aunque también se observan con mayor cantidad de ramas. En la epidermis
1654 BRIT.ORG/SIDA 20(4)
Anexo 1.Relacion del material garante utilizado para el estudio anatomico.
Colector N°coleccibny — Especie Localizacion geografica Condicion del
herbario de Material
dsito
Ruiz Zapata, 4568 MY S.chlamydogynum — Edo. Aragua- Municipio De herbario
Thirza. et al. Tovar. Selva nublada
Benitez de Rojas = 6193 MY S.chlamydogynum — Edo. Araqgua- Municipio FAA 70%
C.E.et al. Tovar. Selva nublada
Badillo, VictorM. 3848 MY S.obovalifolium Edo. Aragua- Municipio De herbario
M.B. lragorry. Selva
nublada
Benitez de Rojas 4908 MY S.obovalifolium Edo. Aragua- Municipio De herbario
CE. etal. M.B.lragorry. Selva
nublada
Morillo, G.y M. 5132 MY S. oppositifollum Edo. Amazonas- De herbario
Hasegawa MunicipioAtures. Selva
humeda macrotérmica
Benitez de Rojas — 2401 MY 5. sessile Edo. Mérida- Municipio ~~ De herbario
CE. etal. Arzobispo Chacon. Selva
nublada
Benitez de Rojas 805 MY S. triste Edo. Sucre.-Distrito De herbario
CE. et al. Montes. Selva nublada
Knapp, S.et al. 6/763 MY S. turgidum Edo. Sucre-Peninsula de De herbario
Paria. Selva nublada
abaxial, los tricomas protectores son todos multicelulares ramificados (Fig. 2)
y se encuentran en una alta densidad sobre lamina y nervaduras (8,84-15,94/
mm~); los mismos son de mayor tamano que los de la epidermis superior y
generalmente tienen dos ramas principales que emergen a un mismo nivel, las
que luego se ramifican en un numero variable de ramas de orden mayor (Fig. 2).
-os tricomas glandulares en las dos epidermis son multicelulares simples (D
adaxial= 2,39/ m2; D abaxial= 1,47-2,78/ mm2), no distinguiéndose claramente
el numero de células que los conforman.
—
Solanum obovalifolium Benitez
La epidermis adaxial presenta células con contornos poligonales, de paredes
ligeramente engrosadas (Fig. 3), mientras que la abaxial esta formada por células
de paredes ligeramente onduladas y mas delgadas. Hoja hipostomatica (LE.=
7,73-12,5); estomas con mayor frecuencia del tipo anisociticos, aunque también
se observan diaciticos (largo 19,5-20,] jum y ancho 16,4-16,5 um). En la epider-
mis adaxial se observan tricomas glandulares. En la epidermis abaxial estan
presentes tricomas protectores y glandulares, con una densidad de 3,27 y 4,75-
7,2/mm- respectivamente. Los primeros son simples, cortos y unicelulares (Fig.
+) y los segundos son simples, cortos, multicelulares uniseriados; sin embargo,
no fue posible distinguir el numero de células que lo conforman.
1655
Fics. 1-4. Solanum chlamydogynum Figs. 1-2. 1. Epidermis adaxial 100.0 I ;
f de las células epidérmi | i f tor ramificado. 2. Epidermis abaxial 100. Véase tricomas
P los y f las epideé les. Sol bovalifolium Figs. 3-4. 3. Epidermis
adaxial 400 «.4. Epdermis abaxial 400 =. N6 i p imple unicelular. B d la: Figs. 1-2 = 100
Lim; Figs. 3-4 = 10 um
th do |
1656 BRIT.ORG/SIDA 20(4)
Ha sido indicado (Metcalfe y Chalk 1979), que el tricoma glandular es de
naturaleza delicada, por lo que éstos son mejor detallados en material vivo oen
especimenes tratados mediante secado por punto critico.
Solanum oppositifolium Kuiz & Pay.
La epidermis adaxial presenta células epidérmicas normales con contornos
poligonales a ligeramente ondulados y paredes celulares ligeramente
engrosadas, mientras que la epidermis abaxial esta formada por células con
contornos ondulados y paredes relativamente delgadas. Hoja anfiestomatica (LE.
superior 4,01; LE. inferior 13,69); estomas del tipo anisociticos en ambas epider-
mis (Fig. 5), siendo de largo 25,37 tm y ancho 15 um en epidermis adaxial y de
largo 24,37 um y ancho 18,5 um en la epidermis abaxial. Presencia de tricomas
en ambas epidermis. En la superior todos se encuentran desprendidos,
observandose tnicamente las células basales (D= 0,83/ mm7), por lo que se hace
necesario repetir el procedimiento con una maceracion mas suave. Sobre las
nervaduras se observan tricomas protectores simples, cortos y unicelulares. En
la epidermis inferior se encuentran en mayor frecuencia tricomas glandulares
alargados, de forma columnar, ubicados en toda la lamina (D=2,43/ mm?) y
sobre nervaduras; la parte inferior de este tricoma se tine intensamente con el
azul de toluidina (Figs. 6,7). Ubicados principalmente sobre las nervaduras 0
proximos a ellas, se encuentran tricomas protectores cortos unicelulares, aunque
en ocasiones pueden observarse sobre la lamina.
See sessile Ruiz W Pay.
La epidermis adaxial esta constituida por células de contornos poligonales y
paredes celulares relativamente gruesas (Fig. 8), mientras que la abaxial tiene
células epidérmicas normales con contornos poligonales y paredes ligeramente
engrosadas, pero mas delgadas que la epidermis superior (Fig. 9). Ambas epi-
dermis presentan ornamentaciones a manera de papilas en la cuticula, siendo
estas con mejor definicion en la epidermis superior (Fig. 8). Hoja anfiestomatica:
estomas de la epidermis adaxial unicamente presentes en muy baja frecuencia
sobre las nervaduras principales o muy cerca de éstas (largo 29,67 um y ancho
11,78 jum). Estomas de la epidermis abaxial oie alas del tipo anisocitico
(Fig. 9) (LE: 13,37; largo 23,1 um y ancho 14,6 pm). Presencia de tricomas
glandulares simples, multicelulares uniseriados, ees ubicados sobre la
lamina y nervaduras de ambas epidermis (Fig. 9) (D adaxial =0,82/ mm2 y D
abaxial =4,4/ mm2).
Solanum triste Jacq.
Células epidérmicas normales, de la epidermis adaxial, con contornos ligera-
mente ondulados y paredes relativamente delgadas. En la epidermis abaxial,
las células tienen contornos ondulados y paredes delgadas (Fig. 11). Hoja
antiestomatica; epidermis superior con estomas eee ey anisociticos
Ct 4
TOFU e
TONES
- Et i
afnetises
Senshi v7 radthey
e
om
Figs. 5—9. Solanum oppositifolium Figs. 5—7.5.Epidermis adaxial 100. Notese la | ia de est la lamina. 6
Epidermis abaxial 100. Vista general. 7. Detall i I imple unicelular 400 x. Solanum sessile Figs. 8-
9. 8. Epidermis adaxial 100. 9. Epidermis abaxial 100. Notese el tricoma glandular simple (indicado con flecha).
Barra de escala: Figs.5—6, 8—9= 100 Lim; Fig. 7=10 lum
1658 BRIT.ORG/SIDA 20(4)
proximosa las nervaduras (Fig. 10) argo 22,77 um y ancho 15,07 um) y epider-
mis inferior con estomas en su mayoria anisociticos (Fig. 1) Cargo 24,67 pm y
ancho 18,43 um) ubicados en la lamina (IL. E. = 14,73). Tricomas protectores y
glandulares presentes en ambas epidermis sobre la lamina y nervaduras. Los
primeros son de dos tipos, simples cortos, multicelulares uniseriados (Fig. 10 y
12) y ramificados cortos (multicelulares, con ramas cortas cercanas a la base),
siendo este ultimo tipo de tricoma mas ramificado en la epidermis inferior (Fig.
12). La densidad de los tricomas protectores fue de 6,08/mm? en la superficie
adaxial y 13/mm¢en lasuperficie abaxial. Los tricomas glandulares son simples
cortos, multicelulares uniseriados (D=1,26/ mm? en ambas epidermis).
Solanum turgidum S. Knapp
Células epidérmicas normales de la epidermis adaxial con contornos
poligonalesa ligeramente ondulados y paredes relativamente delgadas (Fig. 13),
mientras que las de la epidermis inferior tienen contornos medianamente
ondulados y paredes delgadas (Fig. 14). Hoja anfiestomatica; estomas
principalmente del tipo anisocitico, los de la epidermis adaxial ubicados
proximos o sobre las nervaduras (Fig. 13) Cargo 20,53 pm y ancho 13,3 nm),
muy aisladamente se encuentran estomas dispersos sobre la lamina (Fig. 13).
Los estomas de la epidermis abaxial se encuentran en mediana densidad sobre
toda la lamina (Fig. 14) (1-E. 11,37; largo 19,57 um y ancho 14,8 4m). Presencia de
tricomas glandulares en ambas epidermis. En la superior, los mismos se
encuentran colapsados, dispersos en toda la lamina (D=0,33/ mm2). En la cara
abaxial se observan constituidos por pie unicelular y cabeza formada por varias
células (D=1,1/mm~).En esta misma epidermis, sobre las nervaduras principales
se observan tricomas protectores simples, cortos unicelulares. El pie unicelular
del tricoma glandular se tine mas intensamente con el colorante, lo que dificulta
su observacion.
CLAVE DE LAS ESPECIES
1. Tric omas unicamente glar \dule ire Solanum sessile
1. Tricomas protectores y glandulares
2. Tricomas protectores siempre simples y cortos en la epidermis abaxial.
3. Hoja hipoestomatica Solanum obovalifolium
3. Hoja anfiestomatica.
4, Estomas en la epidermis adaxial ubicados en toda la lamina
Solanum
oppositifolium
4, Estomas de la epidermis adaxial sdlo proximos a/o sobre las nervaduras
Solanum turgidum
2. Tricomas protectores ramificados, excepcionalmente simples en la epidermis
abaxial.
5. Tricomas con ramas cortas. Paredes celulares onduladas en la epidermis
abaxial Solanum triste
. Tricomas con ramas largas. Paredes celulares poligonales a ligeramente
onduladas en la epidermis abaxia __ Solanum chlamydogynum
BENITEZ Y BERLINGIERI AIA PAR 1659
a
5
5, eam
ey
me
ED
Since, RB BB:
ses aa. onda eae. oo
= — nD. wig
oe ea ae a Se ee eS
ter 2
SB OY IB Niles sn ik ay se
Fics. 10-14, Solanum triste Figs, 10-12.10. Epidermis adaxial 400. Ndtese estomas cercanos ala nervadura y tricoma
protector simple. 11. Epidermis abaxial 400 .Obsé paredes celul J 1 lle de tri protect
ramificado corto 400 x. Solanum turgidum Figs. 13-14.13.Epidermi t
estomas en la nervadura. 14. Epidermis abaxial 400. Vista general. Barra de escala: Figs. 10-12, 14= 10 tm; Fig. 13=
100 Lim.
110 Vict f | J 4 n
Ox«
1660 BRIT.ORG/SIDA 20(4)
CONCLUSIONES
Rasgos del sistema dérmico representan caracteres tiles para la caracterizacion
de las hojas en especies de Solanum seccion Geminata, pudiendo ser de ayuda
en la identificacion de especies, lo cual ha sido indicado para otros géneros de
Solanaceae (De Pasquale et al. 1991). El tipo de hoja segtan la ubicacion de los
estomas en ambas epidermis ademas de su distribucion bien sea en la lamina o
cerca de las nervaduras, representan caracteres de valor diagnostico para las
especies estudiadas. Respectoa la ubicacion de los estomas, este rasgo permitid
diferenciar las especies, no asi el tipo de estoma ya que se evidencié similitud en
todas las especies estudiadas. Los tricomas hasta ahora observados, basicamente
protectores, son de valor para la diagnosis de las especies, caracter que ha
sustentado la clasificacion infragenérica dentro de Solanum (Seithe 1962, 1979).
Aunque la mayoria de los tricomas glandulares se colapsan con la metodologia
—
utilizada en este estudio, no se consideré realizar un examen mas detallado de
ellos, ya que vistos al microscopio Optico son muy similares en todas las especies.
Caracteres Cuantitativos como densidad y longitud de tricomas reflejan
variacion condicionada por el ambiente y edad de la planta y raramente pueden
ser usados para diferenciaciones taxonomicas ( Metcalfe y Chalk 1950); por lo
que dichos caracteres se han considerado como rasgos complementarios a la
caracterizacion de
ea
as especies. El reconocimiento anatomico de la epidermis
foliar como una fuente de evidencias taxonomicas ha permitido separar las
especies mediante una clave.
AGRADECIMIENTO
Al Fondo Nacional de Investigaciones Cientificas y Tecnologicas por e
financiamiento de esta investigacion con base en el proyecto FONACIT SI-
98002242.
REFERENCIAS
Aumab, KJ. 1964, Epidermal studies en Solanum. Lloydia 27:243-250.
Bessis, J. and M. Guyor. 1979. An attempt to use stomatal characters in systematic and
phylogenetic studies of Solanaceae. In: J.G. Hawkes, R.N..Lester and A.D. Skelding, eds.
The biology and taxonomy of the Solanaceae. Academic Press. Pp. 321-328.
Cartouist, S.R. and W. Hott. 1961. Comparative Plant Anatomy. Claremont, California.
tures of the genus Datura (Solanaceae), Section Dutra. Plantes médicinales et
phytothérapie. 25(1): 23-31.
Knapp, S. 2002. Solanum section Geminata (Solanaceae). Flora Neotropica Monograph 84:
1-404.
BENITEZ Y BERLINGIERI { 1661
Mertcatee, C.R.and L.CHatk. 1950. Anatomy of the dicotyledons Vol. |, Clarendon Press, Ox-
ford. Pp. 965-978.
Metcatre, C.R.and L.CHatk. 1979. Anatomy of the dicotyledons Vol. |, Clarendon Press, Ox-
ford. Second edition. Pp.63-75.
RasHig, A.and A.R. Bec. 1989. A foliar key to the family Solanaceae. Pakistan J. Forest. 39:
107-111.
Sauissury, EJ. 1927.On the causes and ecological significance of stomatal frequency, with
special reference to the woodland flora. Phil. Trans. Roy. Soc. London 216:1-65.
Sette, A. (NEE VON Horr). 1962. Die haararten der gattung So/anum L.und ihre taxonomische
Verwertung. Bot. Jahr. Syst. 81:261-336.
SertHe, A. (NEE VON HorF). 1979. Hair types as taxonomic characters in Solanum.|n:J.G. Hawkes,
R.N. Lester and A.D. Skelding, eds. The biology and taxonomy of the Solanaceae. Aca-
demic Press. Pp. 307-319.
Sotereder, H. 1908. Systematic anatomy of the dicotyledons Vol. 1 y 2. Clarendon Press,
Oxford. Pp. 1183.
1662 BRIT.ORG/SIDA 20(4)
BOOK NOTICES
cel } t bf Vo - IRA d ee
l/Useful Plants/Toxic
Joun and MARGARET CANNON. 2003. Dye Plants and Dying. (ISBN 0-88192-572-1,
pbk.). Timber Press, Inc. 133 S.W. Second Ave, Suite 450, Portland, OR 97204-
3527, U.S.A. (Orders: www.timberpress.com, mail@timberpress.com, 503-
227-2878, 1-800-327-5680, 503-227-3070 fax). $19.95, 129 pp., 48 color
paintings, 7 3/16 x 9 3/4",
This is a lovely book deserving of a readership beyond those mainly concerned with the fiber arts.
For the historian and ethno-botanist the use of plants for color arose in the far distant past. For the
botanical novice, the e ite of the plants and the ay and checklist of scientific
names offer a good intr . There is material for the conservationist and the bio-
chemist. For the art ge the oe the book a delight
This combination of art and science makes this book important. It offers exact instructions for
als
craft dyers and accurate botanica rmation about the plant sources of the coloring material. T
illustrations are accurate and beautiful. The color one ae mordant symbols are incorporated
into the eect the ple ate soas to make a pleasing whole. This is cellent book.
Itcoversa wide range of natural plant dyes and provides accurate botanical int I
each plant.—Ruth Ginsburg, Botanical Research Institute of Texas, Fort Worth, TX, 76102-4060, U.S.A
MARIANNA Appel KUNow. 2003. Maya Medicine: Traditional Healing in Yucatan.
(ISBN 0-8263-2864-4, hbk.). University of New Mexico Press, 1720 Lomas
Blvd. NE, | University of New Mexico, Albuquerque, NM 87131-0001, US.A.
(Orders: 800-249-7737, 505-277-9270 fax, unmpress.com). $29.95, 160 pp.,
36 line drawings, 6 x 0",
Review forthcoming in Sida 21(1), 2004.
Joun Bors. 2001. Natural Compounds in Cancer Therapy: Promising Nontoxic
Antitumor Agents from Plants & Other Natural Sources. (ISBN 0-9648280-
I-4, pbk.). Oregon Medical Press, 315 10" Avenue N, Princeton, MN 55371,
U.S.A. (Orders: 800-610-0768, 612-395-5239 fax, sales@ompress.com,
www.ompress.com). $32.00, 520 pp., 81/2 x 11".
Review forthcoming in Sida 21), 2004.
SIDA 20(4): 1662. 2003
DOCUMENTED CHROMOSOME NUMBERS. 2003: 2.
CHROMOSOME NUMBERS OF THE RARE TEXAS SPECIES,
ARIDA BLEPHAROPHYLLA AND A. MATT TURNERI
(ASTERACEAE: ASTEREAE)
A. Michael Powell
Be
Biology De f
Sul Ross State University
Alpine, TX 79832, U.S.A.
Meiotic chromosome counts for the exceedingly rare Texas species, Arida
blepharophylla (A. Gray) Morgan & Hartman and A. mattturneri B.L. Turner &
Nesom (cf. description in this issue, p. 1418) are reported, both being diploid
with 2n =5 pairs. The two taxa were found growing together or near each other
in ‘Blumberg Canyon’, ca. 2 mi NNW of Ruidosa, Presidio Co., Texas.
Vouchers for the collecti Ari
B.L. Turner 23-91(SRSC, TEX).
| i, Matt Turner LOO(NY, SRSC, TEX); Arida blepharophylla,
Arida blepharophylla (= Machaeranthera gypsitherma Nesom, Vorobik & Hart-
man) was also previously counted as diploid (2n =5 pairs) from a population in
the state of Chihuahua, Mexico (Nesom et al. 1990). Free hand drawings of the
chromosomes, from buds collected on 28 Aug 2003 by B.L Turner, are illustrated
in Fig. Ll.
Fic 1. Meiotic chromosomes of Arida mattturneri (left side, 5 bivalents at metaphase 1); and A. blepharophylia (5
bivalents and 3 centric fragments, at diakinesis)
REFERENCE
Nesom, G.L., L.A. Vorosik, and R.L. Hartman. 1990. The identity of Aster blepharophylla
(Asteraceae: Astereae). Syst. Bot. 15:638-642.
SIDA 20(4): 1663. 2003
1664 BRIT.ORG/SIDA 20(4)
BOOK NOTICES
Crk } a VE . ./KA Jj;
/Useful Plants/Toxic
KeivIN CHAN and Henry Ler (ed). 2002. The Way Forward for Chinese Medicine.
(ISBN 0-415-27720-5, hbk.). Taylor & Francis, 1] New ae Lane, London
EC4P 4EE, UK and 29 West 35" Street, New York, NY 10001, U.S.A. (Or-
ders: info@tandf.co.uk, www.tandf.co.uk, 44 (0) 207 = 9855, fax 44 (0)
207-842-2298, standing orders call +4 (0)1264 343071. US and Canada
Customers: By mail: Taylor @ Francis Customer Service, 10650 Toebben
Drive, Independence, KY 41051, U.S.A., Toll Free Tel: 1-800-634-7004, Toll
Free Fax: 1-800-248-4724, email: cserve@routledge-ny.com). $96.00, 191 pp.,
T1060".
Fhe Way Forward for Chinese Medicine is basically an introduction to Chinese Traditional Medicine.
It takes a look at acupuncture, qigong and related practice, as well as chemical treatment using Chi-
nese medicinal materials for the prevention and cure of illness. The book als he modern
use of Chinese Traditional Medicine in cancer research, issues of toxicity, minced or aiae
tion of Chinese herbal materials using DNA ee inting, and how modernization of Chinese medi-
cine is taking place in China
Contents.— Section One: Some Key Issues in Chinese Medicine; 1) The Historical Evolution ol
Chinese Medicine and Orthodox Medicine in China; 2) Philosophical Aspects of Chinese Medicine
from a Chinese Medicine Academician; 3) The Theory and Application of Functional Food in Chi
nese Medicine; 4) Understanding the Toxicity of Chinese Herbal Medicinal Products: Section Two:
Special Aspects of Clinical and Scientific Practice 1n Chinese Medicine; 3) The Practice of Chinese
Medicine in TCM-KLINIC KOTZTING in Germany; 6) The Practice of Chinese Medicine in the Na
tional Health, Service Environment in the United Kingdom; 7) Treatment of Irritable Bowel Syn-
drome Using Chinese Medicine; 8) The Progress of Using Chinese Herbal Mec
icines in Cancer Re-
search; 9) DNA Techniques for the Authentication of Chinese Medicinal Materials; Section Three:
Progress of Chinese Medicine and Related Practice in Some Countries and Regions; 10) The Progress
of Chinese Medicine in Australia; 11) The Progress of Chinese Medicine in Mainland China; 12) The
Progress of Chinese Medicine in Some Countries in Europe: Belgium, Germany and Holland; 13) The
Progress of Chinese Medicine in Hong Kong SAR, China; 14) The Progress of Kampo Mec
Japan; 15) The Progress of Chinese Medicine in Taiwan; 16) The Progress of Chinese Mec
Unitec
Vers
icine in
icine in the
Kingdom; 17) The Progress of Chinese Medicine in the United States of America; Section Four:
Concluding Issues; 18) A Curriculum Model for Teaching and Training Chinese Medicine in English;
19) An Orthodox Practitioner’s View of Chinese Medicine; 20) The Way Forward for Chinese Medi
cine; and Index.
SIDA 20(4): 1664. 2003
ASPECTOS ECOLOGICOS Y ESTADO DE CONSERVACION DE
MAMMILLARIA THERESAE (CACTACEAE)
Irma Lorena Lopez-Enriquez
| Jorge Alberto Tena-Flores
Instituto Politécnico Nacional
CHDIR Unidad Durango
Apartado postal 738
34000, Durango, Dgo XICO
RESUMEN
tadod vacion de Mammillaria
Se analizan algunos aspectos de la ec Dore cistr ibucion y
le la Si te Coneto, localizada
theresae Cutak (Cactaceae)
al posi ede a Sierra Madre Occidental y en el limite
bi
en el centro del estado de Durango (Méxi
ercidenral del Desierto cnt ahuense Por s isl la Sierra de Coneto es de
s ecologico y fi r -es la Unica loeancad conocida para M. ETE SIEY para un 1 genero
secienbernents deserito(Megacora Onagraceae). Mammillaria the 1a por
conoce oe alments con los nombres de peyote y nevoulle ysu
blacion esta siend la debidoa | le un camino. Se evaluo el eae actual de la
siete
poblacion de uy theresae mediante muestreos es vegetacion y monitoreo de 126
una superficie de 1.7 ha en manchones distribuidos en un area de 2.5 km’,
sitios. La
oe sobre s sustrato rocoso. Debido a su ecm RESETS Ges tamano de pep placion y
M.t]
| c .
disturbi habitat
OY
La informacion recente a ser aplicada en el diseno de un plan nat la prorecciGn de la especie
PALABRAS CLAVE: Mammillaria, Cactaceae, conservacion, endemismo, extincién, Durango
ABSTRACT
We analyzed some aspects of the ecology, distribution, and conservation status of Mammillaria
inenesae Cutak (Cactaceae), a narrow ende mic to the Sierra de Coneto in central Durango, Mexico.
land phytogeographical importance. It is located at the
this sierra |
east of the Sierra Madre Occidental ae on mn the western moMncaes are the Chibiahuan Desert, and it
is the only known locality for M.t g ¢ vgraceae )
i | hich is locally | t nes
Mammillaria theresae isa highly
Its population is being severely aflee ted aiataly due to t
present situation of the species as well as the environmental factors with which it is related. We
studied the population size from seven plots sampled across its distribution area. Density, cover and
were measured, and 126 individual plants were
The species grows over L7 ha distrib-
truction of a food, We evaluated the
frequency of M. theresae and its associated species
monitored to record its fenology and dispersal mechanisms.
uted over approximately 2.5 square km Ai an elev. ational range beeen 2180 and ee mas.l.
Because its restricted distribution, size of the popul
tion of its habitat, it is considered as an endangt species. This inf
plan to protect the species from further decli
andi
orm nation can be used to ae a
Key worps: Mammillaria, Cactaceae, conservation, endemism, extinction, Durango, Mexico.
SIDA 20(4): 1665-1675. 2003
1666 BRIT.ORG/SIDA 20(4)
Las cactaceas representan un recurso unico por su importancia ecoldgica y
economica. Son apreciadas como plantas decorativas, por sus formas raras y
caprichosas y por sus flores vistosas y exoticas, lo que las hace codiciadas por
aficionados y coleccionistas (Sanchez-Mejorada 1979, 1982, 1982a; Becerra 2000:
Benitez y Davila 2002). Ademas de suimportancia como ornamentales, también
se utilizan como alimento, como medicinales y como materia prima en la in-
dustria rural (Rzedowski y Rzedowski 1985). De México se conocen alrededor
de 822 especies, de las cuales mas de 350 pertenecen al género Mammillaria, la
mayoria endémicas del pais. Algunas cactaceas estan entre las plantas mas
amenazadas del reino vegetal, principalmente porque durante varias décadas
han sido objeto de un intenso saqueo de plantas y semillas con el fin de satisfacer
el mercado internacional (Benitez y Davila 2002), y muchas estan sujetas a
presiones de colecta y a la destruccion de su habitat. Regularmente tienen tasas
de crecimiento muy bajas y ciclos de vida muy largos, y sureproduccién es muy
lenta sobre todo después de un evento de perturbacion (Hernandez y Godinez
1994; Arias 1998). En la Norma Oficial Mexicana (NOM-059-ECOL-2001) se
reportan 285 especies de cactaceas en alguna categoria de riesgo.
Mammillaria theresae Cutak (Cactaceae) es una especie de distribucion
restringida a una pequena area en la Sierra de Coneto, en el estado de Durango
(Glass y Foster 1985; Bravo 1991; Gonzalez et al. 1997). Forma parte de la serie
Longiflorae (Hunt 1981, 1987), particularmente buscada por la belleza de sus
flores, y es altamente apreciada por coleccionistas, particularmente europeos,
por su originalidad y rareza (Fig. 1 y 2). La especie aparece por lo menos en 135
sitios de Internet, es cultivada en jardines botanicos de todo el mundo y aparece
en catalogos de muchas companias que la propagan artificialmente para venta.
Hasta hace 15 anos, era tanta su demanda que la poblacién rural del area la
conoce con los nombres de “peyotillo” y “peyote” al considerar erroneamente
{.
que su demanda se debia a que la planta posee propiedades alucin6genas.
Esta incluida en la Norma Oficial Mexicana como especie amenazada;
Vovides (1981) la cita entre las especies en peligro, y en el libro rojo de datos de
la IUCN (www.redlist.org) esta considerada en la categoria de especies
vulnerables. Actualmente su poblacion esta siendo afectada por la ampliacion
del camino entre Coneto y Nuevo Ideal, que sera un tramo de la carretera entre
Torreon, Coahuila y Culiacan, Sinaloa. El disturbio ocasionado por laampliacion
del camino y las actividades de extraccion esta afectando en forma severa tanto
a las poblaciones de M. theresae como de otras especies.
El objetivo de este trabajo es contribuir al conocimiento de M. theresae,
especialmente en lo que se refiere al tamano y densidad de su poblacién, su
distribucion, la especificidad de su habitat y el estado actual en que se encuentra
la poblacion.
pr
hig gy . a
hi ne Pee dol tnd d | i 14 qd
Fig. 3
io ( ta topografica 1:250 000 de
INEGI).
Fic. 4. Habitat de M illaria th Nolina ji primer plano, con Pinus cembroides.
1668 BRIT.ORG/SIDA 20(4)
METODOLOGIA
Tomando como base la localidad tipo de M. theresae, se efectuaron recorridos y
observaciones a lo largo de la sierra de Coneto, apoyados en mapas de Instituto
Nacional de Estadistica, Geografia e Informatica (INEGD. Se llev6 a cabo una
busqueda intensiva en diferentes épocas del ano, particularmente en areas
ecologicamente alines a la zona de donde se tenia reportada la especie.
En la unica localidad donde se encontro la especie se detectaron siete
poblaciones, cada una de las cuales se recorrié para determinar su superficie.
En cada poblacion se trazaron tres lineas Canfield de 20 m (Canfield 1941; citado
en Mueller-Dombois y Ellenberg 1974) para cuantificar la cobertura y la
frecuencia de M. theresae yYsu flora acot npan inte, ya lo largo de la linea media
en cada lugar se coloc6 un bastidor de | m*a través de 10 m para determinar su
densidad. Adicionalmente, en cada poblacion se establecieron puntos fijos de
muestreo de un metro cuadrado para monitorear la [enologia, variacion en
tamano y establecimiento de plantulas de esta especie, estudiandose un total de
126 individuos. Se Hevaron a cabo colectas botanicas de la flora acompanante:
el material se proceso, identificd y deposito en el Herbario CHDIR y sus
duplicados seran posteriormente distribuidos a otros herbarios. Para determinar
iu
el estado de conservacion de M. theresae se consideraron los criterios
especificados en la Norma Oficial.
Descripcion del area de estudio
La Sierra de Coneto se localiza en los municipios de Nuevo Ideal y Coneto de
Comontort, en el centro del estado de Durango. Fs una serrania aislada, ubicada
al oriente de la Sierra Madre Occidental y en el limite occidental del Desierto
Chihuahuense, entre los 24° 54 y 24°59 N y los 104° 45’ y 104° 49 W; su altitud
varia entre los 2100 y los 2700 m s.n.m. (Fig. 3 y 4). El clima del area es templado
semiseco (BS; kw) de acuerdo al sistema de Koeppen, modificado por FE. Garcia
(1973). La vegetacion dominante es un matorral esclerofilo de Quercus
depressipes y Arctostaphylos pungens con Pinus cembroides, Quercus y Yucca.
En el estrato herbaceo destacan Selaginella schaffneri, S. rupincola, S. pilifera,
Sedum pringlei y diversas gramineas y compuestas. Este tipo de ecosistemas
a
contribuyen a la génesis y al mantenimiento de endemismos y por lo tantoa la
riqueza de la flora (Rzedowski 1978). De la zona se conoce también Megacorax
graciclanus, género y especie descubiertos durante el desarrollo de este proyecto
y reclentemente descritos (Gonzalez et al. 2002).
RESULTADOS Y DISCUSION
Distribucion y abundancia
Mammillaria theresae tiene una distribucion muy restringida. Se localizo
tnicamente en la zona conocida como Puerto de Coneto y sus alrededores, entre
los 2180 y los 2320 m, en una superficie de 1.7 ha distribuidas sobre un area de 2.5
1669
km2. Se distribuye en siete manchones (poblaciones) de tamano y densidad muy
diversos, en pendientes de 0’ a 45° desde la parte media de las laderas hasta el pie
de monte, donde es mas abundante. Manifiesta peeenee por sitios con fuerte
t
afl y suelomuy también
se localiza debajo de piedras. Crece solitaria 0 en colonias; la densidad promedio
de los individuos en la zona en donde mejor se desarrolla es de 1.2/m?, aunque
llega aalcanzar (incluyendo plantulas y juveniles) hasta 78 plantas/m? en el sitio
de mayor densidad. En las poblaciones examinadas, M. theresae cubre el 0.059%
de la superficie y presenta una frecuencia relativa de 1.11% (Cuadro D).
Aldesarrollarse en fisuras de la roca madre, esta especie afronta condiciones
de escasa o nula humedad la mayor parte del atlo, quedando completamente
expuesta al sol y al viento. Como resultado de la erosion que esta sufriendo el
suelo las plantas en ocasiones son arrastradas por las lluvias y la raiz queda al
descubierto.
Flora acompanante
Las especies herbaceas que sobresalen tanto en cobertura como en frecuencia
relativa son Selaginella schaffneri y S. rupincola, asi como varias gramineas
(Cuadro 1). Las herbaceas con mayor frecuencia en el area pertenecen a las
familias Selaginellaceae, Crassulaceae, Portulacaceae, Gramineae, Compositae
y Euphorbiaceae. Mammillaria theresae se encuentra asociada principalmente
a Sedum pringlei, Selaginella schaffneri, S. rupincola y S. pilifera, asi como a
gramineas, otras herbaceas y musgos.
Los elementos lenosos dominantes en la zona son: Pinus cembroides,
Quercus depressipes y Arctostaphylos pungens, mientras que Pinus chihuahuana,
Quercus eduardii, Q. emoryi, Garrya wrightii, Perymenium sp., Yucca sp., Nolina
durangensis, N. juncea y Agave parryi son escasos. En canadas y sitios con mayor
humedad ambiental se encuentran Q. oblongifolia, Ceanothus greggii, Purshia
mexicana y, de manera muy aislada, Arbutus arizonica, Quercus rugosa, Q.
sideroxyla y Comarostaphylis polifolia; en sitios mas secos prevalecen Quercus
chihuahuensis, Mimosa aculeaticarpa var. biuncifera, M. dysocarpa, Dalea bi-
color y Verbesina chihuahuensis. La cobertura aérea y frecuencia de las
principales especies lenosas se presenta en el Cuadro 2
o menos dos elementos se conocen como endémicos a la Sierra de
Coneto: M. theresae y Megacorax gracielanus, género y especie descubiertos
durante el desarrollo de este trabajo. Este descubrimiento confirma la necesidad
de mayor exploracion botdanica, y no se descarta la posibilidad de encontrar
mas elementos de interés en el area.
Fenologia
Durante la época de lluvias (mayo-julio) la planta aumenta de volumen,
alcanzando su mayor tamano (hasta 2.6 cm de diametro y 4.2 cm de altura)
entre junio y agosto, periodo en el que se registran incrementos desde 0.3 hasta
1670 BRIT.ORG/SIDA 20(4)
Cuapro 1. Cobertura y frecuencia de Mammillaria theresae y su flora acompanante, roca, suelo y
grava. Datos en orden de importancia de acuerdo a la cobertura.
Componente Longitud de Cobertura Frecuencia Frecuencia
intercepcin (m) Relativa (%) Relativa (%)
Roca 158.86 37.824 346 20.13
Grava 80.844 19.249 320 18.62
Selaginella schaffneri 67.319 16.028 261 15.18
y S. rupincola
Suelo 42.643 10.153 283 16.46
Gramineas 38.697 9.214 309 17.98
Piedra 9.86 2.348 AQ 2.33
Musgos 3.906 0.930 33 1.92
Herbaceas 3.233 770 31 1.80
ipes 3.19 0.760 7 0.41
Aoaie cas 3.185 0.758 12 0.70
Arctostaphylos pungens 2.39 0.569 4 0.23
plas pilifera 1.78 0.424 17 0.99
Vim 1 ] 1.05 0.250 7 0.41
y M. on arpa
Otros arbustos 0.95 0.226 4 0.23
Polypodiaceae 0.635 0.151 10 0.58
Purshia mexicana 0.41 0.098 | 0.06
Sedum pringlei 0,336 0.080 / 0.41
Pinus cembroides 0,29 0.069 2 0.12
Mammillaria theresae 0.247 0.059 19 111
aes rax — cielanus 0.105 0.025 | 0.06
Opt 0.05 0.012 2 0.12
es ee sp. 0.02 0.005 3 0.17
otal 420.00 100.00 1719 100.00
15cm del diametro original. Durante 2002, en julio se present una tendenciaa
la reduccion del volumen. En la Figura 5 se muestra el cambio en diametros de
la mayoria de los individuos muestreados (promedio), y en la Figura 6 se presenta
el cambio ocurrido en el individuo de mayor diametro y en los de menor
diametro registrados; la linea truncada de la Figura 6 indica la desaparicion,
debido a la ampliacion del camino, del individuo mas grande. El diametro de
los individuos adultos varia entre 0.5 y 2.6 cm. Cuando la planta alcanza su
mayor tamano, los tubérculos del tallo se alargan, Ilegandoa medir 0.2a0.5cm,
y se separan entre si hasta 0.4 cm, y la planta toma un color morado-verdoso.
En la estacion seca (enero-abril), las plantas pierden volumen, contrayéndose
hasta mas de un 50% de su tamano y permaneciendo enterradas en el suelo y
fisuras de las rocas hasta el nuevo ciclo de Iluvias, lo que dificulta su localizacion.
En el area se encuentran pequenas rocas en forma redondeada de aspecto muy
similar al de M. theresae.
1671
Cuaoro 2.Cobertura aérea y frecuencia de los principales el lef de la vegetacion donde
ocurre Mammill 1 theresae.
Especies Longitud de Cobertura Frecuencia Frecuencia
intercepcion (m) Relativa (%) relativa (%)
Pinus cembroides 14.69 47.76 5 15.63
Quercus depressipes 5.69 18.50 9 28.13
ey os pungens 1.86 6.05 4 12.50
Mimosa acanthocarpa 47 15.28 5 15.63
var. biuncifera y M. dysocarpa
Dalea bicolor 0.31 1.01 1 3.13
Perymenium sp. 0.46 1.50 3 9.38
Garryd Sp. 0.36 1.17 ] 3.13
Selene gregil 0.2 0.65 | 3.13
2.31 7.51 2 6.25
ibe na sp. 0.18 0.59 1 3.13
Total 30.76 100.00 32 100.00
Durante 2002, M. theresae presento dos periodos de floracién, uno a
mediados de mayo y otro a principios de julio. Con base en estudios en vivero,
Kleine & Kluger (2002) reportan que la planta florece de manera intermitente
desde las primeras semanas de mayo hasta mediados de julio, presentando dos
o tres periodos de floracion. La flor es de color violeta palido a intenso, de 2.5a
4.7 cm de largo (incluyendo el tubo de 0.8a 2.5cm) y hasta de 4.5 cm de diametro,
con 18 a 32 pétalos; las anteras son de color amarillo intenso. En un periodo de
floracion se desarrollan una a tres, raramente cuatro, flores por individuo. La
flor dura de dos a tres dias, abriendo por la manana y cerrando por la tarde. Se
observaron flores ramoneadas y separadas de la planta. El fruto es rojo, diminuto,
de aproximadamente 2 mm de ancho. Las semillas permanecen retenidas en la
planta por varios meses, protegidas entre los lobulos del tallo y particularmente
en la base de éste, en un engrosamiento formado por el tallo que se compacta en
la época seca con la parte superior de la raiz.
Dispersion
La dispersion se lleva a cabo principalmente por arrastre, ya que el fruto y las
semillas son movidos por el agua 0 el viento. Las semillas se establecen en las
fisuras de las rocas o sobre el suelo sro. Adicionalmente, las plantas muertas
o desenterradas por efecto de la erosion, son arrastradas por el agua o el viento
y dispersan las semillas que aun permanecian adheridas entre los lobulos del
tallo y en el engrosamiento en la base de éste, dando origen a nuevas colonias.
Las plantulas aparecen en el mes de agosto, tanto aisladas como alrededor de la
planta madre. No se observaron indicios de dispersion por animales, aunque
cabe la posibilidad de que ocurra.
1672 BRIT.ORG/SIDA 20(4)
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Estado de conservacion
Mammillaria theresae esta incluida en la Norma Oficial Mexicana (NOM-059-
ECOL-2001) como especie amenazada. Hernandez y Godinez (1994) y Hunt
(1987), la reportan como especie amenazada, conocida tnicamente de la
localidad tipo, mientras que Vovides (1981), la cita como especie en peligro de
extincion. Los criterios para definir “rareza” en grupos taxonomicos particulares
son variables (Ceballos 2001). Se consideran especies raras aquellas que tienen
distribuciones restringidas, poblaciones poco abundantes 0 combinaciones de
ambos lactores. Las especies de distribucion restringida son mas vulnerables a
1673
la extincién como resultado de actividades antropogénicas que destruyen o
modifican el ambiente, caso en el que se encuentra M. theresae. La Union
Mundial para la Naturaleza (IUCN) considera especies raras a las que tienen
areas de distribucién menores de 50 000 km? y clasifican a M. theresae en la
categoria de especies vulnerables. La especie no se incluye en el Apéndice I de la
ITES
Se han realizado estudios para conservar especies de cactaceas que se
encuentran en alguna categoria de riesgo. Hernandez y Sanchez (2000), reportan
384 especies que se encuentran en 19 colecciones de jardines botanicos en la
Republica Mexicana, en los que se Hlevan a cabo acciones para conservar y
reproducir especies en riesgo. No se tienen reportes de que M. theresae se
encuentre jardines botanicos de México, a excepcion probablemente del jardin
botanico del Grupo Cante, en Guanajuato. La especie se ha reproducido con €xito
en Europa en diversos jardines botanicos y en viveros para venta. Aunque en el
Instituto Tecnolégico Agropecuario No | de Durango se estan reproduciendo
las semillas con éxito, no existe un programa de dispersion de la especie en
México.
Aplicando los criterios del Libro Rojo de Datos IUCN 1980), M. theresae
califica en la ee acne en peligro de extincion. El mismo resultado se obtiene
al aplicar el méto acion del riesgo de extincion de especies que marca
el Anexo ca Ide la NOM-059-ECOL-2001. Por su distribucion muy
restringida (menos de 2 km2), su poblacién pequena y el disturbio y la
fragmentacion de su habitat debido a la ampliacion del camino, M. theresae es
evidentemente una especie en peligro. Por tanto, se requieren acciones
inmediatas de proteccion que eviten su desaparicion.
CONCLUSIONES
Mammillaria theresae presenta una distribucion muy restringida, y hasta ahora
se conoce Gnicamente del area conocida como Puerto de Coneto y sus
alrededores. La bisqueda de M. theresae ha permitido el descubrimiento de un
género y especie nuevos, lo que parece indicar que el area representa una zona
de interés floristico que debe ser estudiada con mas detenimiento.
De los diversos reportes en que se menciona a esta especie, la mayoria
coinciden en que se encuentra en la categoria de amenazada. Sin embargo, el
analisis de sus condiciones actuales (area de distribuci6n muy pequena,
crecimiento lento y alteracién y fragmentacion en su habitat) senala que se
encuentra en peligro de extincion. Para que el desarrollo social y econdmico
vayana la par con la conservacion de los ecosistemas, es necesario que las nuevas
obras estén avaladas por un estudio de impacto ambiental previo. En el caso de
la ampliacion del camino que afecta a la poblacion de M. theresae, de existir un
estudio de impacto ambiental, no se consider6 que el area representaba la unica
1674 BRIT.ORG/SIDA 20(4)
localidad conocida de una especie, y no se aplicaron acciones que permitan
mitigar el impacto en esa poblacion. Es urgente el establecimiento de un
programa para la proteccion de esta especie y de su habitat.
AGRADECIMIENTOS
Los autores agradecen a la COFAA e Instituto Politécnico Nacional por el aks
brindado para el desarrollo de este trabajo (CGPI-20010347); a M. Pinedo y $
Acevedo el apoyo brindado en camp ;a Abel Garcia Arévalo por su participacion
durante muestreos previos, y Barney Lipscomb y un revisor anénimo por
sugerencias que permitieron mejorar el manuscrito.
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1675
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Wiley & Sons. N.Y. Toronto.
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BRIT.ORG/SIDA 20
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1676 )
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species of Irigonella, their pharmacological
describing t
Materials, dealing with the chemi ts of the s}
properties and the marketing ol aaa seed are also discussed.”
Contents.—1) ee we Botany; 3) Physiology: a Cc oe ation; 5) Breeding; 6) Nutrition
ean ises; 8) Weeds; 9) Cl | constit 10) Pharmacological
7) i)
and use of fertilize
properties; 11) ee (ane and Index.
Yono-su Zien (ed). 2002. Tea: Bioactivity and Therapeutic Potential. (ISBN 0-
415-27345-5, hbk.). Medicinal and Aromatic Plants—Industrial Profiles. Volume
17. Series Editor: Roland Hardman. Taylor & Francis, || New Fetter Lane,
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Contents.~1) Tea and Health - An Overview: 2) Botanical Classification of Tea Plants: 3) Green Tea.
Black Tea and Semi-fermented Tea; 4) The Chemistry of Tea Non-volatiles; 5) The C hemistry of Tea
Volatiles; 6) Biochemical and Cellular Bases for Tea Activity; 7) Pharmacological Fffect of Caffeine
and Related Purine Alkaloids: 8) The Effects of Tea on the Cardiovascular System; 9) Antimicrobial
Activity of Tea Products; 10) Anticarcinogenic Activi ity of Tea; 11) Amina Activity of Tea Prod-
icine; 13) Agronomy niet commercial
ucts; 12) Therapeutic Uses of Tea in Traditional Chinese Medi
Production of Tea; and Index.
SIDA 20(4): 1664. 2003
TYPE LOCALITIES OF VASCULAR PLANTS
FIRST DESCRIBED FROM OHIO
Ronald L. Stuckey James S. Pringle
Herbarium Royal Botanical Gardens
The Ohio State University . Box 399
Museum of Biological Diversity Hamilton, Onteno CANADA L8N 3H8
1315 Kinnear Road jpringle@rbg.ca
Columbus, OH 43212-1192, U.S.A.
Type localities are those places where biological specimens were obtained that
subsequently were used as the basis for the scientific name and description of
any organism when it was first described as new to science. Throughout the
history of classical and descriptive biology, countless numbers of organisms,
or taxa, have been named and described. Along with each new taxon, usually
but not always, is recorded its original source location, or type locality. These
type localities are deserving of permanent record, because they are the special
places to which a biologist must return if he is to obtain living material from
descendents of the original populations. A biologist may need to verify the iden-
tity and description of the taxon in the area where the original population oc-
curred, obtain material for genetic and developmental studies, and/or acquire
populational and life-history information. With the expansion of taxonomic
studies into the broader realms of biosystematics and population analysis that
has come into prominence within the past half century, much more impor-
tance is now accorded the type localities than in earlier times. These classic
sites are significant biogeographical reference points at the international, na-
tional, and state levels. They should be spared from permanent destruction, such
as the flooding of a river valley by construction of a dam on a stream or river,
the building of a housing or industrial complex, the construction of a highway,
or the tilling of the land for agricultural purposes. Type localities therefore may
serve as criteria for evaluating areas or sites as potential nature sanctuaries, as
discussed by Stuckey (1994).
Information on the type localities of plants first described from the United
States is inadequately documented as a subject by itself. Although the informa-
tion is recorded and scattered throughout the descriptive historical and botani-
cal literature, it has not been brought together into accessible and usable data
banks. Taxonomic monographs regularly include information on the type lo-
calities for names applied to the taxa discussed in those papers, but they deal
only with small groups of species, and the organization is taxonomic rather
than according to the geographic origin of the specimens. A project to record
SIDA.20(4): 1677- 1692. 2003
1678 BRIT.ORG/SIDA 20(4)
type localities for species within a genus was presented in a catalogue of type
specimens for names in the large genus Carex (Cyperaceae) (Shetler et al. 1973).
It included an index to the states, provinces, and countries from which the speci-
mens were obtained. Lists of the type specimens deposited therein have been
compiled for many herbaria, but these references list specimens according to
the repositories rather than according to their geographic origin. “Botanical
gazetteers” have been compiled fora few states, but do not deal specifically with
type localities. At the state level, we are aware of published lists of type locali-
ties for only four states, viz. Illinois Jones 1952), lowa (Lammers 1985), New
Mexico (Standley 1910), and West Virginia (Core 1936). A few smaller lists cover
either a region or a specific county in the United States. Below are listed over 60
type localities for Ohio, from which a total of 197 vascular plants were first de-
scribed from the state. Stuckey and Wentz (1974) have previously catalogued
the type specimens from these localities that were deposited in the herbarium
of The Ohio State University.
That the precise location of the original sources of many of these new taxa
described from Ohio is not given is not surprising. Many of the taxa were named
in the nineteenth century, when precision as to location was of little concern,
since no International Code of Botanical Nomenclature existed and little em-
phasis was placed on the source location of new biological entities. In those
days it was considered sufficient to record the location of a newly described
plant from such broadly defined areas as the State of Ohio, the shore of Lake
Erie, the Miami River valley, or the Ohio River. The last locality, of course, means
from along the banks of the Ohio River or from its vicinity. The most notable
example of this situation is the 112 or more taxa described between 1808 and
1840 from Ohio by Constantine Samuel Rafinesque, who listed the source lo-
calities as “Ohio,” “Lake Erie,” or “Ohio River.” Rafinesque’s 17 vascular plants
named from the shore of Lake Erie! and nine noted as being from the Ohio River
are included in the list below, although some of Rafinesque’s plants may have
been obtained on the Kentucky side of the river. With the exception of one name
in current usage and two upon which currently used names are based, the many
taxa described by Rafinesque merely as from “Ohio” are not listed here, but they
can be determined from Index Rafinesquianum (Merrill 1949). André Michaux’s
taxa described from the Ohio River are also listed, but the exact locations along
the river where these plants were obtained are not known. Michaux is known to
have conducted some botanical exploration near the Ohio River in the vicinity
| pat | 1 £ ae
'Some or perhaps all of th shore of Lake Frie may have been observed at
Sandusky, Erie County, where he waited three days in late May 1826 for a steamboat from Detroit that took him
to Buffalo. However, during the route to Buffalo he did stop briefly at such places as Cleveland and Fairport
(Rafinesque 1826, pp.6—7; 1836a, PP 80-81). He also noted later that the shore of Lake Erie near Sandusky repre-
sented a locality of ' ‘great t botanical interest” that aff € ecies of plants aaa le 1836b, pp. 28-
5p || |
29),
1679
of Louisville, Kentucky, and in what is now southern Illinois, as well as near the
present site of Portsmouth, Ohio. Some spe recorded by Michaux as hav-
ing been obtained from the “Ohio River” or its banks are, therefore, likely from
localities outside present-day Ohio.
In other situations only the nearest town was mentioned, with no specific
sites or habitats given for the plant. Some sites are sufficiently well described so
that their locations can be determined today. Among the best-known type lo-
calities in Ohio from which vascular plants have been described are Beaver Pond
in Adams County, Cedar Bog (Cedar Swamp) in Champaign County, Cedar Point
and other locations near Sandusky in Erie County, the limestone cliffs along
the Scioto River north of Columbus and other sites in Franklin County,
Hoffman’s Prairie (Wright Brothers Prairie) east of Dayton in Greene County,
Rocky Fork Creek in Highland County, Ofer Hollow in Jackson County, and
Cranberry Island in Buckeye Lake in Licking County. Significant type locali-
ties that have disappeared through agricultural or industrial development in-
clude the Oxford Prairie in Erie County, the Darby Plains in Madison County,
and Van Cleve’s Prairie near Dayton in Montgomery County.
The general locations of the type localities listed below are shown on a
map of Ohio (Fig. 1), and two additional maps show portions of the state where
many type localities are concentrated in small areas (Figs. 2, 3). The localities
are concentrated mostly in the Miami valley in southwestern Ohio, in Frank-
lin and adjacent counties in central Ohio, and near Lake Erie in northern Ohio.
These locations reflect the pioneer botanical exploration in Ohio, first in the
Miami River valley in the early 1800s, in central Ohio mostly in the mid 1800s,
and in northern Ohio primarily about 1900 and sas The es unusual habi-
tat types represented are fens, wet prairies, and li s. These habitats
are not well represented in the states to the east of his whew most of the de-
scriptive botany had been written before the early 1800s. It is understandable,
therefore, that plants characteristic of these habitats that had not previously
been named and described were first described from places in Ohio.
Unlike the other state lists mentioned above, which were arranged alpha-
betically by taxon, this list for Ohio is organized by locality, beginning with the
largest unit, Ohio, followed by sections of the state including the shore of Lake
Erie, the Miami River valley, and the Ohio River valley, and then alphabetically
by county. Those taxa with more specific locality data are listed under each
county. If two or more localities in Ohio were cited when a taxon was first de-
scribed, as for example with Valeriana ciliata Torr & A.Gray, all such localities
are listed. All of the names based on specimens from the same type locality are
listed under that locality, in alphabetical order. An index to names of taxa, by
genus only, begins on page 1691.
Double asterisks denote plant names that are currently accepted as the
correct names for the respective taxa. Single asterisks denote names that are no
BRIT.ORG/SIDA 20(4
1680
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longer in use, or at least are not accepted in the standard references consulted
in the present study, but which have been superseded by currently accepted
names based directly upon them. Single asterisks are also used for the names ol
taxa originally described as species that are now considered to be interspecific
hybrids. In these cases, the present-day name generally includes the original
epithet, meaning the adjectival component of the name in a nomenclatural
combination that was published later. The type localities remain the same.
When the name based ona specimen or specimens obtained in Ohio is no longer
in use for a taxon, the currently accepted name is given in brackets when pos-
sible. Currently accepted names and taxonomic synonymy have been deter-
1681
FRANKLIN Boundaries
25 Mar 1851-1990
Ww \ ‘:
{
Rs
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NG
Flinta ge Westervill aa Sess
28\936 / SOL
ink Hube New
orthingto Ridee Aue _~ Ale
a ree ( Jersey .
n
Shadevill
ér] y
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op hd
—] Ly
ateriea
Z Lockbourne =< iachullle
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—— Ty Lithopolis
14
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(¢ ~
Point
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Fr 7 7 ot Ly
that county in Ohio.
mined primarily from Kartesz (1994), Kartesz and Meacham (1999), and those
volumes of the Flora of North America North of Mexico that had been published
at the time of this writing. For some old names not listed even in synonymy in
these works, the modern equivalents have been determined from older floras,
such as those by Britton and Brown (1913), Fernald (1950), and Gleason (1952),
and from monographs, notably those by Boivin (1944) on Thalictrum, Davis et
al. (1967-1970) on Rubus, Fernald (1905) on Eriophorum, Heiser et al. 1969) on
Helianthus, Mackenzie (1931-1935) on Carex, and Palmer (1956, 1961) on
Crataegus. When no current status or taxonomic equivalent is given for aname
published at the rank of variety, it may be inferred that varieties are now gener-
1682 BRIT.ORG/SIDA 20(4)
WESTERN LAKE ERIE
LUCAS CO
|
WOOD CO |
Miles |
0 10 SANDUSKY CO ; :
| i |
R.L. Stuckey |
Fic 3 AA 44h Nhi af | g tl L 1 P4 fVAl + 1 L t L q by } ape } yt 1 1 ££
vascular pl 1 I 1f I y f Oh
ally not recognized within that species. In these cases, when a synonym is given
lor the species, it should not be assumed that the minor variant described asa
variety is equivalent to the entire species. Current status is not indicated for
names published at the rank of form (Latin forma, abbreviation “f.”), because
names at this rank are not usually included in modern floras. In most cases, the
original epithet applied to the form, transferred if necessary to the currently
accepted name for the species, which is indicated here, would be correct for
anyone wishing to use names for these individual variations.
In some cases it has not been possible to equate an older name with any
currently accepted taxonomic equivalent. When the original description is de-
ficient in detail and the original specimens no longer exist, it may not be cer-
tain to which currently recognized taxon an old name, based on plants occur-
ring in Ohio, was applied, although taxonomic research continues to resolve
some of these long-persistent questions. This situation is an especially frequent
problem with plants named by Ralinesque. The best single source for their in-
terpretation is Merrill’s 1949) Index Rafinesquianum. Other publications that
are especially significant with regard to plants originally described from Ohio
specimens include the first author's studies of the plants named and described
by Daniel Drake (Stuckey 1969) and Thomas Nuttall (Stuckey 1966, 1967), and
his studies of the plants described by CS. Kunth and E.G. von Steudel from speci-
mens obtained in the Miami River valley by Joseph Frank (Stuckey 1974).
1683
THE PLANTS AND THEIR TYPE LOCALITIES
OHIO
“Acalypha rhomboidea Raf., New FI. 1:45. 1836
[sometimes included in Acalypha virginica L.,
r. rhomboidea (Raf) Cooperr].
ssc eens Riddell, WJ. Med. Phys. Sci.
1835. Reprint p. 34. 1835 [Aesculus
vie Willd].
Cakile americana Nutt.,Gen.N.Amer.PI.2:62.1818
[Cakile edentula (Bigelow) Hook. subsp.
edentula
Carex laxi flora Lam. var. latifolia Boott, Ill. Carex p.
38, pl. 93. 1858 [Carex albursina E
Carex alata Torr. var. ferruginea ce ee
Amer. Acad. Arts 37:477, pl. 2. 1902 [Carex
suberecta (Olney) Britton].
**Carex comosa Boott, Trans. Linn. Soc. London
20:11
Carex sos sii ae & Torr. f. glabrescens Kuk.
Engl., Pflanzenr. 4(20):588. 1909.
oe foenea Willd. var. B Boott, Ill. Carex p. 118,
pl.376.1862 Pin ieee pis oe
Carex foenea Willd.v anual,
ed. 5, p. 580. 1867 [Carex Aubereee a
7
Britton].
Carex ise! Lam. var. blanda (Dewey) Boott
racillima Boott, Ill. Carex p. 38, pl.91,
. 2. ee on gracilescens Steud].
*Carex monile Tuck., Enum. Meth. Caric. p.20. 1843
[Carex vesicaria L.var.monile (Tuck.) Fernald].
*Carex tenera eee! var. suberecta Olney, Carices
eo r. 3:1871 [Carex suberecta (Olney)
Brit a
pene ridgwayi Sarg.,J.Arnold Arbor.6:2.1925
[Crataegus mollis (Torr.& A.Gray) Scheele].
*Cyanotris scilloides et Amer. Monthly Mag. &
Crit. Rev. 3:356. 1818 [Camassia sciloides (Raf,)
Cory].
**Fleocharis erythropoda Steud., Syn. Pl.
Glumac.2:76.1854 ee known as Eleocharis
calva Torr., an invalid na
Eriophorum gracile Henan ex Roth [var] B
paucinervium Engelm., Amer. J. Sci. Arts
46:103. 1844 [E iophor Ulli (Or rellum Nutt.].
“Eriophorum latifolium be [var] 8B
a Engelm., Amer. J. Sci. Arts
1844, “viridi-carinatum" in
ie (Engelm.) Fernald].
**Fuphorbia commutata Engelm. ex A.Gray,
Manual, ed. 2., 9. 389.1856
Fimbristylis frankii Steud, Syn. Pl Glumac. 2:11
1855 [Fimbristylis autumnalis (L.) Roem. !
Schult].
*Uodysarlim pau iflorum Nutt _ Gen. N. Amer. Pl.
2:109. 181 8 [Desmodium pauciflorum (Nutt.)
DC].
**Helianthus Um Torr. & A.Gray, FI. N.
soph fis ‘a Riddell, WJ. Med. Phys.
516. 1835. Reprint p. 84. 1835
[Hudronhbvill im macron
iP
[Hy y hyllum Nutt.]).
** Juncus brachycarpus Engelm., Trans. Acad. Sci.
St. Louis 2:467. 1868
“Juncus canadensis J.Gay var. brachycephalus
Engelm., Trans. Acad. Sci. St.Louis 2:474. 1868
Juncus brachycephalus — (Engelm.)
Buchenaul].
Lysimachia revoluta Nutt. Gen.N. Amer. Pl. 1:122
8 [Lysimachia quadriflora Sims].
**Dog sy Ivestris A Gra\ /Mant ial ed 1,p.596. 1848.
Scutellaria ambigua Nutt., Gen. N. Amer. Pl. 2:37.
1818 [Scut lla ia nervosa Pursh].
Scutellaria versicolor Nutt. Gen. N. Amer. PI. 2:38.
1818 [Scutellaria ovata Hill subsp. eee
**Stachys cordata Riddell, WJ. Med. Ph
579. 1836.Reprint p. 15.1 oes eee ate
; We Ch tel .
literature, but no valid reason exists to re-
me S.cordata Riddell].
ea oe Greene, Pittonia 3:368.
898 [now usually treated as Hymenoxys
herbacea (Greene) Cusick, but sometimes
retained in Tetraneuris].
**Tradescantia ohioensis Raf., Précis Découv.
miol. p.45. 1814; see also New FI.N. Amer.
2:84, 1836 [1837].
Vernonia altissima Nutt., Gen. N. Amer. Pl. 2:134.
1818 [Vernonia gigantea (Walter) Trel.subsp.
gigantea]
LAKE ERIE, SHORE OF (Probably at Sandusky,
Erie County)
Aesculus muricata Raf., Alsogr. Amer. p. 68. 1838
esculus glabra Willd].
Asclepias dasypus Raf., Atlantic J. 1:
clepias purpurascens L].
152.1832 [As-
1684
Asclepias rotundifolia Raf, Atlantic J. |
not Mill.
JESm J.
Caprifolium dentatum Raf., Atlantic J.1
[Lonicera dioica L.].
Cornus punctata Raf., Alsogr. Amer. p. 62. 1838
Cornus alternifolia LJ.
Cornus suffruticosa aa Atlantic J. 1
lsogr. Amer. p.61.
Fragaria e latior Raf., nae J. “
Ehrh. 1792 nor Wight & Arn.
Fragaria serotina Raf., Atlantic J. 1 2.1832.
Lathyrus incurvus Raf. Atlantic J. | 7 1832, not
Roth 1787 nor Willd. 1802 nor Rchb. 1832.
Lobelia falcata Raf., New FI. 2:18. 1836 [1837] [Lo
delia kalmii LL].
Lonicera eriensis Raf., New FI.3:18-1
nei era hirsuta Eaton].
Lys 1 sessilifolia Raf., Atlantic J. 1
ie ena ia suede Sims].
Mentha traxigona Raf., Autik. Bot. p. 114. 1840.
Samolus petiolatus Raf., Herb. Raf. p. 41. 1833
1:152. 1832,
1768 [Asclepias amplexicaulis
151.1832
1:151. 1832;
52. 1832, not
foe
9.1836 [1838]
=
1:151. 1832
[Samolus valerandi L. subsp. paviflorus (Raf.)
Hultén].
he brevipetalum Raf., Med. FI. 2:100. 1830;
Autik. Bot. p. 134. 1840 [Trillium flexipes Raf].
lium rotundifolium Raf., Med. FI.2:97. 1830 [Tril-
lium erectum L].
Viola eriocarpa Raf., Atlantic J. 1:152. 1832
[nomenclaturally distinct from Viola
eriocarpa Schwein. (1822), although prob-
i || H | +]
ably taxonomically equivalent].
MIAMI VALLEY; MIAMI COUNTRY
ri
=
bass cea is een ia Michx. var. vitacea
Kne 70, 1893 [Parthenocissus
vi see, : ae
a ee us sel Riddell, WJ. Med. Phys.
Sci.8:5 oe Reprint p.70.1835,not Roem.
819 nor Willd.ex Spreng. 1824 [/po-
moea ae LJ.
ae as Steud, Syn. Pl. Glumac. 2:199.
5 (Carex rosea Schkuhr ex Willd].
a gracilescens Steud, Syn. Pl.Glumac. 2:226,
1855. 1849.
Carex steudelii Kunth, Enum. PI. 2:480. 1837 [Carex
jamesii Schwei
Eragrostis cognata syn. PI. Glumac. 1:273.
1854 [Eragrostis a nacea (Michx.) Nees ex
>
Steud ].
BRIT.ORG/SIDA 20(4)
**Fragrostis frankil
Glumac. 1:273.1
a ene unionis St eud.6 n. PL Glumac. 1:273.
4 [Eragrostis pectinacea (Michx.) Nees ex
Set
Valerianella triquetra Hochst. & Steud. ex
Shuttlew., Flora 20:211. 1837 [Valerianella
chenopodiifolia (Pursh) DC].
OHIO RIVER
Asplenium angustifolium Michx., Fl. Bor.-Amer.
2:265. 1803 [Diplazium pycnocarpon
(Spreng.) M.Broun].
Discovi es le Raf. J.Phys. Chim. Hist. Nat. Arts
89:9 9 [?Lesquerella globosa (Desv.)
C.A.Mey. ex Steud., Syn. PI.
—
em
Eup qtoritm longipes Raf,, Atla ntic J.1:1 : : 3):
a ium pectinatum Raf, pel J.1:17.1832,
Wall. 1831 nor Small 1
ae thymifolia Michx., - ae 22) 2.
1803, not L. 1753 [Chamaesyce maculata (L.)
Small; sometimes retained in Euphorbia, as
E. maculata L.]).
Hibiscus hastatus Michx., Fl.Bor.-Amer. 2:45. 1803,
not Lf. 1781 nor Cav. 1787 [Hibiscus laevis
All.].
llysanthes riparia Raf., Ann. Nat. 1:13
[Lindernia dubia (L.) Pennell var. dubia].
Lithospermum angustifolium Michx., Fl. Bo
Amer. 1:130.1803, not Forssk. 1775 oe
& Mog. 1893 [Lithospermum incisum Lehm.].
Planera gmelinii Michx., Fl. Bor.-Amer. 2:248. 1803,
“qmelini’ (Planera aquatica (Walter) J.-.Gmel.]
Podalyria coerulea Michx., Fl. Bor.-Amer. 1:264.
1803 [Baptisia australis (L.) R.Br.ex W.T.Aiton
var. australis}.
*“Polygonum amphibium L. [var] B emersum
Michx., Fl. Bor.-Amer. 1:240. 1803
Potamageton 0 natans L. [var] B Michx., Fl. Bor-
Amer. 1803 [Potamogeton epihydrus
Raf].
**Potentilla paradoxa Nutt.in Torr. & A.Gray, FIN,
er. 1:437.1840, not Schur ex Nyman 1878.
Scutellaria radicata ae Atlantic J. 1:16. 1832.
**Spermacoce glabra Michx., Fl. Bor.-Amer. ie
3, not Roxb. 1820 nor Sessé & Mog. 1
Tilia fulva Raf., Alsogr. Amer. 45. 1838 “ie
ericana L.].
Vitis odoratissima Donn var. atropurpurea Raf,
Med. FI. 2:132. 1830.
1820
co
pa
oe
=
1685
“Vitis riparia Michx., Fl. Bor.-Amer. 2:231. 1803.
ADAMS COUNTY
01. Beaver Pond: **Asplenium xinexpectatus
E.L.Braun ex C.V.Morton, Amer. Fern J.46:152.
"1956" [1957]. [= Asplenium cryptolepis
Fernald x A. rhizophyllum (L.) Link; earlier bi-
nomial xAsplenosorus inexpectatus E.L. 7
ex Friesner, Butler Univ. Bot. Stud.4:154. 1
was not validly Sublishedl
. Prairie, north part of Jefferson Township,
southeast of Scrub Ridge, 2 '/2 miles north
of Ohio Route 348: **Silphium terebinthina-
ceum Jacq. var. lucy-brauniae Steyerm
Rhodora 53:133-135.1951.
ae COUNTY
Old prairie near Hamilton: Amaranthus
vs imus Riddell, WJ. Med. Phys. Sci. 8:367.
. Reprint p. 41. 1835 [Amaranthus
ae (Moq,) J.D.Sau
04. Small prairie halfa mile an + ede
Euphorbia herronii Riddell, WJ.Med. Phys. Sci.
8:58. 1835. Reprint p. 32. 1835 [Euphorbia
dentata Michx,].
CHAMPAIGN COUNTY
05. Cedar Swamp [now Cedar Bog], Urbana:
*Valeriana ciliata Torr. & A.Gray, Fl. N. Amer.
2:49. 1841 [Valeriana edulis Nutt. ex Torr. &
A.Gray var. ciliata (Torr. & A.Gray) Cronquist].
CLARK COUNTY
06. Near Springfield: *Valeriana ciliata Torr. &
A.Gray, FI.N. Amer. 2:49. 1841 [Valeriana edulis
Nutt. ex Torr. & A.Gray var. ciliata Cronquist]
COSHOCTON COUNTY
07. North Appalachian Experimental ie a
near Fresno, Chili,and Canal Lewisville:
Aralia spinosa os : subinermis eae Casta-
nea 9:54. ]
j=)
NI
Daucus carota - : fischeri Moldenke, Castanea
Daucus i L. f. goodmanii eMMalelene: Casta
944,“goodmani
Lobelia ee L. f. albiflora Moldenke: Castanea
9:65. 1944.
Lycopodium flabelliforme (Fernald) Blanch. f.cluter
Moldenke, Castanea 9:32. 1944
[Diphasiastrum digitatum (A.Braun) Holub
Morus alba L.f.nigrobacca Moldenke, Castanea
9:51.1944
=
Plantago rugelii Decne. f. fasciculata Moldenke,
Castanea 9:65. 1944.
Tovara virginiana (L.) Raf. f. rubra Moldenke,
Boissiera 7:4. 1943; see also Castanea 9:40.
1944 [Polygonum virginiana L.}. Although
one might infer otherwise from Moldenke's
paper in Castanea, this and the following
two names were validly published in
Boissiera.
Tracaulon sagittatum (L.) Small f. subalbidum
Moldenke, Castanea 9:41. 1944 [Polygonum
sagittatum L,].
1 €iu KAAIA | DEERE
74-5. 1943; see also Castanea 9:49. 1944,
Vernonia altissima Nutt. f. alba Moldenke,
Boissiera 7:5. 1943; see also Castanea 9:64.
1944 [Vernonia gigantea (Walter) Trel.subsp.
gigantea].
08. Roscoe: Stachys glabra Riddell, W.J.Med.Phys.
Sci. 9:580. 1836. Reprint p. 16. 1836 [Stachys
tenuifolia Willd).
CUYAHOGA COUNTY
09. Cuyahoga: Hieracium watsonianum Gand.,
Bull. Soc. Bot. France 65:49. 191
10. Parma: Hieracium ohioense Gnd. Bull. Soc.
Bot. France 65:48. 1918.
11. Rocky River, deep gorge about one- ow
mile from Lake Erie: Cerastil
webbii Jennings, Ohio Naturalist 9-441. 305
see also Ohio Naturalist 10:136. 1910
[Cerastium arvense L. subsp. velutinum (Raf.)
Ugbor. var. velutinum,]
DELAWARE COUNTY
12. Delaware: Crataegus ellipticifolia Sarg., J. Ar-
nold Arbor. 3:194. 1 71990 [Craimeque commacin
Sarg.].
ERIE COUNTY
13. hee Island, bottom of deep northwest
quarry by glacial grooves, N portio
nie Island, Lake Erie: **Juncus x
M.Reinking, Brittonia 33:175. 1976 [=
alpinoarticulatus Chaix x J. torreyi Coville].
14. Cedar Point: *Helianthus luxurians E.Watson,
Pap. Michigan Acad. Sci. 9:464. 1929
[Helianthus x/uxurians E.Watson; pro sp.;= H.
giganteus L. x H.grosseserratus M.Martens].
15. Huron River at mouth: eee esculen-
Gen. MN Amer. Pl. 1:219. 1818
s (Raf.) Cory].
tum Nutt.,
—
[Camassia scillo
1686
16. Milan: Crataegus propinqua Ashe, J. Elisha
Mitchell Sci. Soc. 20:53-54. 1904 [Crataegus
iracunda Beadle}.
17. Oxford Prairie: Solidago moseleyi Fernald,
Rhodora 10:93. 1908 [Futhamia gymno-
Lae ides Greene; often included in Solida
5 S.gymnospermoides (Greene) fl
Thalictrum ce reene, Amer. Midl. Natu
ist 2:294. 19 2 {Thalictrum dasycarpum i
cher & Aveé-Lall.].
18. ae een marcida Ashe, J. Elisha
Mitchell Sci. Soc. 20:53. 1904 [Crataegus
iracunda Beadle].
Crataegus tenera Ashe, J. Elisha Mitchell Sci. Soc.
20:52-53. 1904 [Crataegus macrosperma
he].
19. a ek Bay: Vicia douglassii Torr. Amer. J.
6.1822 [?Vicia cracca
FAIRFIELD COUNTY
20. Sugar Grov
J. Elisha Mitchell Sci. Soc. 16:79.1899;see also
E.J.Palmer, J. Arnold Arbor. 6:57. 1925
e]
[C, rat 1egus intricataq Lane
Crataegus pol\ bracteata Ashe,
21. Clearecr: Tigh R.19W,sec.10,1.5 miles east
of Clearport on the south side of County
Highway 69, east of the covered bridge over
Clear Creek: **Cystopteris xwagneri R
Moran, Castanea 48:224. 1983 [= Cystopteris
tennesseensis Shaver x C. tenuis (Michx.)
Desv.].
alae eounly
Reyley: T)
SC haffneri Camp, Rhodora 42: 55. 1940.
*Asclepias sullivantii Engelm. ex
A.Gray, Manual, ed. 1, 366. 1848.
Aster laxifolius Nees [var.] B /aetiflorus Torr. &
A.Gray, FI.N. Amer. 2:138. 1841 [Aster borealis
eek Prov.ssometimes segregated
Aster as el boreale Torr.
Gray) A.Love & D.Love].
ae conjuncta Boott, I. Carex 3:122, pl. 392.
1862.
*Carex sullivantii Boott in A.Gray, Amer. J. SCi.
42:29. 1842 [Carex xsullivantii Boott, pro sp.;
= Carex hirtifolia Mack. x C. gracillima
Schwein].
Crataegus ohioensis Sarg., J. Arnold Arbor. 3:1
1922 [Cra sec arborea Beadle].
Fedia patellaria Sull.ex A.Gray, Manual, ed. 1, 183.
23. Columbus: *
BRIT.ORG/SIDA 20(4)
1848 [Valerianella umbilicata (Sull.) Alph.
Wood}.
*“Fedia umbilicata Sull., Amer. J. Sci. 42:50. 1842
Neat a umbilicata eae ) Alph.Wood].
He ji Britton, Man. FI. N. States
p. 994. 1901, “kellermani"; see also WA.
Kellerman, Ohio Naturalist 2:179-181, 1902
(Helianthus xkellermanii Britton, pro sp.; =
Helianthus salicifolius A.Dietr. x H.
grosseserratus M.Martens]
“Rudbeckia sullivantii ete & Beadle,
Biltmore Bot. Stud. 1: 1901 [Rudbeckia
fulgida Aiton var. sullivantii (C.L.Boynton &
Beadle) Cronquist].
24. Columbus, cat-tail swamp, three miles west:
“Coreopsis discoidea Torr.& A.Gray, FI.N. Amer.
2:339.1842 [Bidens ee (Torr. & A.Gray)
Britton]
25. Columbus, rocky limestone banks of the
Scioto River: **Arabis patens Sull, Amer. J.Sci.
42:49. 1842, not Royle ex Hook.f. & Thomson
1861.
Annual Rep. Missouri Bot. Gard. 14:181. 1903
(Lonicera dioica L. var. dasygyna (Rehder)
Gleason].
Lonicera aa var. Pa FILN. Amer.
2:6. 1841 [ Raf].
iss
rene ie ee bal [var] B? [sic] sullivantii
Iph.Wood, Class-book Bot., ed. 2, p. 298.
1847, sent on the Ohio component of
Lonicera parviflora var. B et sensu Torr. &
A.Gray, FIN. ee ei , but nomen-
claturally distinct; type ie determined
from Torrey and Gray [Lonicera reticulata
Raf].
26. Columbus, two miles south: **Solidago
ohioensis Riddell, W.J. Med. Phys. Sci. 8:497.
1835. Reprint p.57.1835 [sometimes segre
from Solidago as Oligoneuron
ohioense (Riddell) G.NJones].
27. Dublin, calcareous ravine and steep decliv-
ity on the Scioto River: Prenanthes proteo-
phylla fiche W.J.Med. Phys. Sci.8:490. 1835.
Reprint p 1835 [Prenanthes alba L
“rium (i ee
a
Between Flint and Glenmary: Crataegus
anki ea J.Amold Arbor.4:100.1923
(Crataegus dissona Sarg].
29. Scott's Plains, 12 miles east from Worthing-
on: **Solidago riddellii Frank in Riddell, WJ.
Med. Phys. Sci.8:497.1835.Reprint p.57.1835
[sometimes segregated from Solidago as
Oligoneuron riddellii (Frank) Rydb.]. Riddell
probably should have written west instead
of east, because of the prairie habitat in that
area west of Worthington, rather than in the
forest east of Worthington (R.LS.).
30. Worthington: Aster carneus Nees var.
ambiguus Torr. & A.Gray, Fl. N. Amer. 2:133.
1841 [Aster lanceolatus Willd. var. interior
(Wiegand) Semple & Chmiel.; sometimes
segregated from Aster as Symphyotrichum
lanceolatum subsp. lanceolatum var. interior
Wiegand) G.L.Nesorm].
eerie 's Riddell, WJ.Med.Phys. Sci.
8:4 . Reprint p. 55. 1835 sometimes
segregated from Aster as es ichum
oolentangiense (Riddell) G.L.Nesom].
Prenanthes parviflora Riddell, W.J. ie Phys. Sci.
90. 1835. Reprint p. 50 [Prenanthes
altissima L].
aes labo Riddell, WJ. Med. Phys. Sci. 9: 580.
Reprint p. 16. 1836 [Stachys tenuifolia
a
GALLIA CO
31. ioe er horseyi E.J.Palmer, Ohio
J. Sci. 56:211-212. 1956 [Crataegus intricata
Lange].
32. Gallipolis, banks of the Ohio River: **Collin-
verna Nutt., J. Acad. Nat. Sci. Philadelphia
1:190. 1817.
Discovium ohioense Raf., Autik. Bot. 17. 1840
(?Lesquerella globosa (Desv.) S.\Watson].
GREENE COUNTY
33. Rien Elie eight miles east from Day-
Riddell, WJ.Med
Ales Sci. 8:367. 1835. Reprint p. 41. 1835
[Amaranthus tuberculatus (Mogq.) J.D.Sauer].
"saldogo rs i Frank in Riddell, WJ.Med.Phys.
1835. Reprint p. 57. 1835. This lo-
cee is now referred to as the Wright Broth-
ers Prairie or Wright Patterson Natural Area,
which is located within the Wright Patter-
son Air Force Base east of Dayton, Ohio
Ptogoeica
/ . i}
var. onigAE remid Roden 30: 43. 1928
1687
[Asplenium ruta-muraria L., usually not rec-
ognized at the varietal level; when accepted,
the correct epithet is in doubt as of this writ-
ing, but evidently would not be ohionis].
HAMILTON COUNTY
35. Cincinnati: Aesculus maxima D.Drake, Natu-
ral and Statistical View, or Picture of Cincin-
nati and the Miami Country ... p. 78. 1815
(1 81 6] TAescuits flava Aiton].
Enslen Nutt.,Gen.N.Amer. Pl. 1
lomaneun laeve (Michx.) Pers.].
a eee albidum Nutt., Gen. N. Amer. PI.
3.1818
1:164.1818
pre aranaicia Nutt.,Gen.N. Amer. PI. 2:29.
1818 [Synandra hispidula (Michx.) Baill].
36. Three si north of Cincinnati: *Quercus
leana Nutt, Sylva 1:13.1849 [Quercus x/leana
Nutt. pro sp.;= Quercus imbricaria Michx. x
QO. velutina Lam
37. Near Terrace Park: **Viola xbrauniae Grover
ex Cooperr,, Michigan Bot. 25:108. 1986 [=
Viola rostrata Pursh x V. striata Aiton].
HARDIN COUNTY
38. Mt. Victory: Crataegus meiophylla Sarg., J.
Arnold Arbor. 3:198. 1922 [Crataegus
margaretta Ashe].
HIGHLAND COUNTY
39. Rocky Fork Creek, 3/4 mile above junction
with Paint Creek: apes sullivantii Torr. &
A.Gray, Fl. N. Amer. 1:575. 1840; Sullivantia
ohionis Torr. & A ae en 42:22.1842,
me for same taxon [Sullivantia
ae (Torr.& A.Gray) Britton. This species
is the type of the genus name Sullivantia,
which was published in the same paper].
HOCK'NG COUNTY
40. Benton Township: Above Keifel Rd., 0.3 mi
NE, jct of Big Pine Creek rd., Sect 7, Benton
Township: *Lycopodium xbartleyii Cusick,
Amer.Fern J.77:100. 1987 [Huperzia xbartleyi
(Cusick) Kartesz & Gandhi; identified in origi-
nal description by Cusick as Lycopodium
lucidulum Michx. x L. porophilum F.E.Lloyd &
U = Huperzia lucidula (Michx.) Trevis.
x H. ies a (FE.Lloyd & Underw.} Holub].
JACKSON COUNTY
41. Liberty Township, Ofer Hollow: *Calama-
grostis insperata Swallen, J. Wash. Acad. Sci.
1688
25:413. 1935 [Calamagrostis porteri A.Gray
subsp. insperata (Swallen) C.W. Greene].
LAKE COUNTY
42. Painesville: “Lonicera glaucescens Rydb. var.
dasygyna Rehder, Annual Rep. Missouri Bot.
sard. 14:181. 1903 [Lonicera dioica L. var.
ecu tee Gleason].
Pani rneri Scribn.ex Britton & A.Brown, Ill.
Fl. N. U. S: @ 501 1898. [Dichanthelium
inearifolium (Scribn. ex Britton & A.Brown
ould; often retained in Panicum, as P
pane jum Scribn. ex Britton & A.Brown].
Ribes cynosbati L.var.glat F
—
=
= —
—
Id, Rhodora
156. 1905.
43. Richmond: *Helianthus brevifolius E.Watson,
ichigan Acad. Sci. 9:448. 1929
(Helianthus x brevifolius E.Watson; pro sp.; =
Helianthus grosseserratus M.Martens x H.
mollis Lam.} .
LICKING COUNTY
44, Buckeye Lake, Cranbert y Island: Acer En
Li oe ) Detmers,“rubro-carpum,”
Ohio J. Sci, 19:236,1919
Acer rubrum L. vat. eee Detmers, Ohio J. Sci.
19:235,1919,
45. Granville: Scutellaria ovata Hill subsp. pseudo-
venosa Epling, Univ. California Publ. Bot. 20:56.
1942 [Scutellaria ovata Hill subsp. ovata].
LORAIN COUNTY
46. Oberlin: Ribes cynosbati L. var. glabratum
Fernald, Rhodora 7:156. 1905; see also O.E.
Jennings, Ohio Nat. 6:492—495, 1906.
47. ee Silas River: Aloitis foliosa Gre
Leafl. Bot. Obs. & Crit. 1:94. 1904 oer
quin ae (L.) Small subsp. occidentalis
(A.Gray) J.M.Gillett].
LUCAS COUNTY
. Toledo: Staphylea brighamii J.F.Macbr.,,
Rhodora 20:129. 1918 [ [Staphylea trifolia LJ.
MADISON COUNTY
49. Darby Plains, 15 miles west of Columbus:
oo Sull., Amer. J. Sci. Arts
42:49. 1842.
=
Helianthus cinereus Torr. & A.Gray var. sulli
orn. & A.Gray, FILN. Amer. 2:234. 1842 7
Aus xcinereus Torr. & A-Gray; pre
Helianthus mollis L. x H.occidentali eis
ivantii
elian
BRIT.ORG/SIDA 20(4)
MEIGS COUNTY
50. North of Dexter; sandstone exposures on
mesic slop above Leading Creek, Co. Rt 10,
0.25 mi (0.02 km) SW of Twp Rt 27, Sec. 6,
Salem Twp:Polypodium xincognitum Cusick,
Amer. Fern J. 92:241. 2002 [= Polypodium
appalachianum Hauffler & Windham x P
virginianum L.].
MONTGOMERY COUNTY
51. Dayton: Prenanthes miamiensis Riddell, WJ.
Med. a Sci. ae 1835.Reprint p.50.1835
[Pr 6 Michx,].
— de Dayt on, O." Prenanthes ovata
dell, WJ. Med. Phys, Sci. 8:490. Reprint p.
835 on anthes alba LJ].
52. os Cleve’s Prairie, Dayton: “50 ONaagO
ohioensis Riddell, W.J. Med. Phys. Sci. 8:497.
1835. Reprint p.57. 1835 eee segre
aa from Solidago as Oligoneuron
=
ohioense (Riddell) G.NJones].
OTTAWA COUNTY
53. Catawba Island: Persicaria laurina Greene,
Leafl. Bot. Observ. Crit. 1:35. 1904;see also OLE.
Jennings, Ohio meal 6:492-495, 1906.
54. South Bass Island: **Polygo
cum L.var.eglandulosum J, C: Miyars, Castanea
7:74, 1942
—_ corel L.H.Bailey in Core, Franz Theodore
Lab. Contrib. 9:70. 1948 [Rubus alum-
a Bailey],
ee eriensis |.H.Bailey in Core, Franz Theodore
Stone Lab. Contrib. 9:70, 1948 [Rubus fron
num pens) VONI-
dosus Bigelow].
Rubus gordonii L.H.Bailey in Core, Franz Theodore
Stone Lab, Contrib. 9:70. 1948 [Rubus deamii
L.H.Bailey].
PORTAGE COUNTY
55. Garrettsville: ele ees Ashe,J.Elisha
Mitchell Sci. Soc. 20:53. 1904 [Crataegus
racunda Bea
56. Windham: pion angustum (Willd.) C.Pres|
rFern J.9:86,1919
[Athyrium filix-femina L.var.angustum (Willd.
G.Lawson; oo as A.angustum
(Willd.) C-Presl].
57. Wood miles Glen: Cystopteris fragilis (L.)
Bernh.var.cristata Hopkins, Ohio Nat. 10:181.
1910, not EJ.Lowe 1869 [Cystopteris tenuis
(Michx.) Desv.].
a
ar.cristatum | lopkins, Amer
—
RICHLAND COUNTY
58. Mansfield: Crataegus decens Ashe, J. Elisha
Mitchell Sci. Soc. 19:19. 190 [Crataegus
lucorum Sarg.].
Crataegus habilis Ashe, Bot. Gaz. 35:435. 1903,
**Crataegus indicens Ashe, J. Elisha Mitchell Sci.
Soc. 19:27.1903
Crataegus mansfieldensis Sarg., J. Arnold Arbor.
4:103.] [Crataegus irrasa Sarg].
Crataegus onusta Ashe, J. Elisha Mitchell Sci. Soc
19:22. 1903 [Crataegus pruinosa (H.L.Wendl.)
K.Koch].
a se Ashe, J. Elisha Mitchell Sci. Soc.
903 she].
‘Crat qedarnemacroasnerma
Crataegus a nsonii Ashe, Bot. ee 35:435.1903,
wilkinsoni biessans crus-galli L.].
Thalictrum amabile G e, Amer, Midl. Natural-
ist 2:294.1912 = erate revolutum DC].
ROSS COU
59. Chillcothe, pees miles southwest:
Scutellaria saxatilis Riddell, WJ. Med. Phys.
Sci. 9:578. 1836. Reprint p. 14.1835.
SCIOTO COUNTY
60. cic banks of the Scioto River: re
ans , Gen. N. Amer. 7 1:164. 1818
haa laeve (Michx.) Pe
Cie ees Nutt.,Gen.N. os Pl.2:38.
S utel| wiaincangq Bie hler].
61. Junction of the Ohio River and Scioto River:
Eupatorium falcatum Michx., Fl. Bor. Amer.
toriadelphus, as E. purpureum (L.) R.M.King &
H.Rob].
**Kyllingia pumila Michx.,Fl.Bor.-Amer. 1:28. 1803
[ etimes placed in Cyperus, as C.
tenuifolius (Steud.) Dandy].
TRUMBULL COUNTY
62. Leavittsburg: Crataegus marcida Ashe, J.
1689
Elisha Mitchell Sci. Soc. 20:53.
[Crataegus iracunda Beadle].
TUSCARAWAS COUNTY
63. Dennison: Crataegus kellermanii Sarg., Trees
& ee 2:239. 1913 [Crataegus subor-
biculata
Dover: en retusa eee WJ. Med.
1836. Reprint p.17.1836, not
1904)
Phys. Sci.9:581.
Rodrigues 1882.
** Helianthus occidentalis Riddell, WJ. Med. Phys.
Sci.9:577. 1836. Reprint p. 13.1836
**linum sulcatum Riddell, ae ae Phys. Sci.
9:574. 1836. Reprint p. 1
65. Near Muskingum oe a ie River,
probably near Gnadenhutten]: **Carex
muskingumensis Schwein. Ann.Lyceum Nat.
Hist. New York 1:66. 1824; 1:312. 1825.
Leen COUNTY
6. [South] Lebanon: otk macrophylla
Nutt. Gen.N. Amer. PI. 2:49. 1818 [Dasistoma
macrophylla (Nutt.) Raf.
WASHINGTON COUNTY
67. Marietta: Crataegus mariettensis Sarg.,J.Arnold
Arbor. 3:194. 1922 [Crataegus dissona Sarg.].
aero putnamiana Sarq., J. Arnold Arbor.
102. 1923 [Crataegus chrysocarpa Ashe].
Sophonn petiolatum Nutt., Gen. N. Amer. PI.
818 [Stylophorum diphyllum (Michx.)
ies
- a
68. Bank of the Muskingum River at Marietta:
epee seb ee Michx., Fl. Bor.-Amer.
. 1803; see also Proc. Amer. Philos. Soc.
oo 1889 - isticia americana (L.) VahlJ.
WAYNE COUNTY
69. Rocky wooded hillsides: Adiantum pedatum
L. var. laciniatum Hopkins, Ohio Naturalist
10:180. 1910.
ACKNOWLEDGMENTS
The first author (R.L.S.) extends his thanks to his former students, Marvin L.
Roberts, J. Perry Edwards, Leslie L. May, and Karen Fries, who have given assis-
tance in this research.
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Stann ey, PC. 1910.The type localities of plants first described from New Mexico.Contr.U.S.
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Stuckey, R.L. 1966. Thomas Nuttall’s 1816 Ohio valley plant collections described in his
“Genera’ of 1818. Castanea 31:187-198
Stuckey, R.L.1967.The “lost” plants of Thomas Nuttall’s 1810 expedition into the Old North-
west. Michigan Bot. 6:81-94.
Stuckey, R.L. 1969. An overlooked plant name (Aesculus maxima) of Daniel Drake and his
lost herbarium. Castanea 34:185-192.
Stuckey, R.L. 1974. Dr. Joseph C. Frank's botanical work in Ohio, with a list of his 1835 type
specimens of vascular plants in American herbaria. Castanea 39:263-272.
Stuckey, R.L. 1994. Type localities as criteria for natural areas. Newsletter, Ohio Department
of Natural Resources, Division of Natural Areas and Preserves 16(4):5.
Stuckey, R.L., and W.A. Wentz. 1974. Vascular-plant type specimens in The Ohio State Uni-
versity Herbarium, Ohio J. Sci. 74:20-35.
INDEX TO GENERIC NAMES
Eriophorum 1683
Erythronium 1687
Eupatoriadelphus 1689
Eupatorium 1684, 1689
Euphorbia 1683, 1684, 1685
Acer 1688 Cerastium 1685
Acer 1688
fees 1687
Aloitis Cornus 1684 Euthamia 1686
ls ee 1685, 1687 Crataegus 1683, 1685, 1686 Fedia 1686
Ampelopsis 1684 1687, 1688, 1689 Fimbristylis 1683
Arabis 1686 Cyanotris 1683 Fragaria 1684
Aralia 1685 Cynanchum 1689 Gentianella 1688
Asclepias 1683, 1684, 1686 Cyperus 1689 Habenaria 1689
Asplenium 1684, 1685, 1687 Cystopteris 1686, 1688 Hedysarum 1683
Asplenosorus 1685 Dasistoma 1689 Helianthus 1683, 1685, 1686,
Aster 1686, 1687 Daucus 1685 1688, 1689
Hibiscus 1684
Hieracium 1685
Desmodium 1683
Dichanthelium 1688
Calamagrostis 1687, 1688
Camassia 1683, 1685
Carex 1683, 1684, 1686, 1689
Caprifolium 1684
Diphasiastrum 1685
Diplazium 1684
Discovium 1684,1687
Eleocharis 1683, 1688
Enslenia 1687, 1689
Eragrostis 1684
Huperzia 1687
Hydrophyllum 1683
noxys 5
Juncus 1683, 1685
1692
Justicia 1689
Lesquerella 1687
Lindernia 1684
Linum 1689
Lithospermum 1684
Lobelia 1684, 1685
Lonicera 1684, 1686, 1688
Lycopodium 1685, 1687
Lysimachia 1683, 1684
entha 1684
Morus 1685
Oligoneuron 1686, 1687, 1688
Panicum 168
Parthenocissus 1684
Plantago 1685
Scutellaria 1 1683, 1684, 1688,
1689
Seymeria 1689
Silphium 1685
Solidago 1686, 1687, 1688
Spermacoce 1684
Stachys 1683, 1685, 1687
BRIT.ORG/SIDA 20(4
Staphylea 1688
Stylophorum 1689
Sullivantia 1687
eee 68/
Synandra
a ae
Thalictrum 1686, 1689
Tilia 1684
Tracaulon 1685
Valerianella 1684, 1686
Vernonia 1683, 1685
Vicia 1686
Viola 1684, 1687
Vitis 1684, 1685
)
THE EASPWES! TRANSITION OF THE PLORA IN TEXAS:
A BIOGEOGRAPHICAL ANALYSIS
Michael H.MacRoberts and Barbara R. MacRoberts
Bog Research, 740 Columbia
Shreveport, LA 71104, U.S.A.
an
Herbarium, Museum of Life Sciences
Louisiana State University-Shreveport
Shreveport, LA 71115, U.S.A.
ABSTRACT
Quantitative methods were used to determine _ transition zone between the eastern and western
flora in Texas. The transition between the s f region and the
western plains/brushy-savanna is a 300 km wide area extending from around 95 Geprees bi to 99 de-
grees west longitude. Across this area, eastern an
the range edge for the two floras. This finding agrees meee with ee ee biogeographic
analyses
Key worps: West Gulf Coastal Plain, flora, biogeography, east-west floristic transition, Texas
RESUMEN
Se usaron métodos cuantitativos para determinar la zona de transicion entre la flora oriental y
eras y la
=
occidental en Texas. La transicion entre la region sur oriental de bosques deciduos/coni
occidental de llanuras/sabana arbustiva es un area de 300 vu de anche que se exUaice desde unos
95 grados a 99 verades de longitud oeste. ae |
separan | g Este ies concuerda
re re ]
oO oO
en gran medida con andlisis previamente.
INTRODUCTION
Despite the fact that the North American coastal plain is one of the continent's
major floristic provinces (Dice 1943; Takhtajan 1986), quantitative papers on its
phytogeography are poorly represented in the botanical literature (Sorrie &
Weakley 2001). Even more poorly represented are such papers on the West Gulf
Coastal Plain (MacRoberts et al. 2002; MacRoberts & MacRoberts 2003). While
several quantitative studies are now available on endemism within the coastal
plain (Sorrie &@ Weakley 2001; MacRoberts et al. 2002), studies on diversity, spe-
cies richness, dispersion barriers, and floristic boundaries are lacking (Currie
1991; MacRoberts & MacRoberts 2003).
It has been long known that the eastern part of the West Gulf Coastal Plain
(east Texas, south Arkansas, west Louisiana, southeast Oklahoma) is part of the
eastern and notably the southeastern United States (Thorne 1993; Christensen
2000; Delcourt & Delcourt 2000), but it is unclear where the floristic east-west
SIDA 20(4): 1693-1700. 2003
1694 BRIT.ORG/SIDA 20(4
transition occurs. An examination of the ecoregional literature indicates a rather
confused picture of this transitional area (MacRoberts & MacRoberts 2003).
The purpose of this paper is to determine where floristic east meets west
inthe West Gulf Coastal Plain.
METHODS
1) In order to obtain a characteristic sample of the southeastern flora at the
same latitude as Texas, we used Kartesz and Meacham (1999) to determine the
distribution of all North American vascular plants whose distribution includes
Georgia, Florida, Alabama, and Mississippi (although not necessarily restricted
to that area) and that cross the Mississippi River into Louisiana and/or Texas
(although not necessarily restricted to that area). Thus, species restricted to only
the above listed states are included (e.g., Xyrisdrummondii Malme) as well as
those found in all of the contiguous 48 states and Canada (e.g., Potamogeton
pusillus L.). Exotic species and hybrids are excluded. Subspecies and varieties
are lumped with species. Species whose taxonomic status or distribution was
questionable are excluded. Our initial data set contained 1320 species.
Since there is no thorough county-by-county floristic inventory of the West
—
Gulf Coastal Plain, we used a single source of data, which consists of samples
from those counties. We plotted the distribution of species that met our crite-
rion by Texas counties using the Texas Consortium database (TAMU-BWG Her-
barium Specimen Browser, a consortium of seven Texas herbaria with 130,000
records online). Of the 1320 species in the initial data set, 1138 were in the TAMU
database by county. In the preliminary analysis all data were converted to per-
centages because the counties have been unevenly collected and reported. Thus,
Bowie County had 489 species in the TAMU database, of which 302 were from
our data set (302/489 = 0.62) and Madison Co. had L016 species, of which 452
were from our data set (452/1016 = 0.44).
2. In order to do the same thing but from a west-to-east perspective, we
sampled the western flora by roughly the same methods as described above.
Because both New Mexico and Mexico border Texas on the west, we selected
three counties in west Texas (Taylor, Tom Green, Edwards), listed the species
found in them from TAMU data base, and randomly selected 313 from the list
on the basis of their western affinity ound in west Texas, New Mexico and/or
Mexico [TAMU; Kartesz & Meacham 1999]). We used a much smaller sample
from the west because we realized that the eastern sample was larger than nec-
essary. Because the sample is smaller, to make numbers comparable with the
east we multiplied the percentages by 2.
We then converted all of the county data in | and 2 above to the Owen and
Schmidly (1986) quadrat system for Texas to standardize area and to even out
irregular county boundaries and percentages. This also allowed us to use Owen
and Schmidly’s data on environmental variables of biological importance. The
ACROBERTS AND MACROBERTS, ANALYSIS OF THE FLORA OF TEXAS 1695
quadrats are 63.9 km ona side, overlaid on a Lambert’s conical projection map
of Texas. We converted the county data to quadrat data by overlaying the quad-
rat map onto our county map and then averaging the percent of species of the
counties that occurred in each quadrat.
3) Having found a clear east-west transition zone from the results of num-
bers l and 2 (see below), we wished to document the floristic affinities of Texas
along an east-west gradient more thoroughly. To do this, we selected four Texas
counties (Tyler, Madison, Bell, Irion) running across Texas at approximately 31
degrees North latitude and sampled them to determine their floristic relation-
ship to the surrounding states. Using the TAMU database, we selected a ran-
dom list of between 200 and 250 native species from each county. We then plot-
ted the United States/Canada distribution of all species in the four lists using
Kartesz and Meacham (1999). As before, the results are expressed as percentage
of the sample (e.g., Louisiana had 97 percent of the species in Tyler County).
RESULTS
Figure | shows the percentage of eastern and western species in samples ac-
cording to longitude across Texas. Figure 2 shows these data summarized for
Owen and Schmidly (1986) quadrats by ten percent intervals. Figure 3 shows
the distribution across North America of the flora of four Texas counties.
DISCUSSION
Anexamination of the shape of the curves in Figure | and the gradient expressed
in Figure 2 shows that along Texas’ 1250 km east-west axis, the area of most
significant change in eastern and western species is the approximately 300 km
wide zone from Houston and Tyler on the east (approximately 95 degree W. lon-
gitude) to Wichita Falls and San Antonio on the west (approximately 99 de-
grees W. longitude). On either side of this area, the change is gradual. Accord-
ing to our analysis, Fort Worth, Austin, and Corpus Christi appear to be the
point at which east and west are balanced (97 degrees West longitude),
The relationship between county affiliation in the broader context of the
entire North American continent north of Mexico shows the same pattern. Tyler
County in east Texas is floristically eastern. Madison County farther west is
mainly eastern, Bell County in the center of the transition zone is intermediate,
and Irion County in the west is western (Fig. 3).
While ours is not a study of species richness, ecoregion boundaries, or di-
versity, nor is it concerned with cause of the species transition, it has implica-
tions for such studies. For example, Currie (1991) in a study of species richness
has shown that the east-to-west loss of tree species across the gulf coastal plain
correlates most strongly with actual evapotranspiration, which is correlated
with primary production and is, therefore, a measure of available environmen-
tal energy.
1696 BRIT.ORG/SIDA 20(4)
Western Eastern
% Species
w
—
|
|
102° 100° 98° 96° 94°
Texas
Fic. 1 Dp + ft +t ld +
oo
Our study also indicates that similar variables are important in determin-
ing species distribution in the West Gulf Coastal Plain. Using Owen and
Schmidly’s (1986) data, we calculated the correlation coefficient of “productiv-
ity” for eastern and western species and found a strong correlation between pro-
ductivity across Texas and the number of eastern and western species. Eastern
species correlated positively (Pearson = 0.91, Spearman = 0.92) while western
species correlated negatively (Pearson = - 0.82, Spearman = - 0.77). The transi-
tion zone between east and west is after all the precipitation transition zone as
well.
Central Texas has long been known to be an east-west transition zone, at
least for vertebrates (Blair 1950; Webb 1950; Gehlbach 1991; Ward et al. 1994;
Schmidly 2002). Dice (1943) referred to this area as the “Texan Biotic Province”
between the eastern forest region ( ‘Austroriparian”) and the semi-arid brushy /
savanna-grasslands (“Comanchian”) of the western part of Texas. Blair (1950)
recognized that the “Texan” was a “transitional region’—even using the word
“ecotone”—but could find no alternative but to keep it a distinct province.
Gehlbach (1991) solved the problem simply by pointing out that central Texas
is a transition zone, not a biotic province, between the eastern deciduous forest
region (Austroriparian Biotic Province) and western semi-arid brushy-savanna
region (Comanchian Biotic Province). Characteristic plants and animals of both
the Comanchian and Austroriparian occur locally in the Texan. More recently,
Diggs et al. (1999:4) have characterized north central Texas floristically as “a
ANALYSIS OF THE FLORA OF TEXAS 1697
20-29
10-19
p ‘ ¢ 1 soe | 1: Ps Ieoh tan. (1002) ' ia -
Fic.2
y\
ten percent intervals.
1698 BRIT.ORG/SIDA 20(4)
1 Lipa bat ae lsc T
ler, Madison, Bell, and
Fic. 3. P t f states. provinces
f 7 ZF r J f r é
JF
Irion counties, Texas.
broad ecotone between eastern deciduous forest and western grassland.” We
concur with these characterizations. From a floristic perspective, there is an
east-west transition in the area roughly designated as Texan Biotic Province.
Paleoenvironmental studies indicate that the current floristic character of
Texas may be fairly recent. Pleistocene and Holocene climatic changes appear
to have been significant with the invasion of the West Gulf Coastal Plain by
southeastern and Mexican species since the last glacial retreat (Bousman 1998:
Delcourt & Delcourt 2000).
We do not consider our overall findings particularly novel. Years ago, zo-
ologists came to basically the same conclusions based on vertebrate distribu-
tion. West Gulf Coastal Plain botanists, however, have been slow to address bio-
geographic questions quantitatively. As a result, botanists and botanically
inclined ecologists have created a plethora of qualitatively defined “ecoregions”
across Texas (Webb 1950: MacRoberts & MacRoberts 2003). We believe that zo-
ologists currently have a better understanding of the biogeographic regions of
Texas than do botanists, but improved plant distributional data (e.g, Turner et
al. 2003) and the replacement of qualitative by quantitative methods of inves-
tigation should soon provide botanists with the opportunity to rectify this situ-
ation. With better data, and thus the ability to make finer comparisons, it will
EOGRAPHICAL ANALYSIS OF THE FLORA OF TEXAS 1699
be interesting to see just where the transition is most pronounced and how west
and south Texas separate from the remainder of Texas and the southeast.
ACKNOWLEDGMENTS
Laurence M. Hardy aided with the figures. Bruce Sorrie commented on an ear-
lier version of the paper. Thanks also to Guy Nesom and Fred Gehlbach for their
reviews of the manuscript.
REFERENCES
Biair, W.F. 1950. The biotic provinces of Texas. Texas J. Sci. 2:93-117.
Bousman, C.B. 1998. Paleoenvironmental change in central Texas: the palynological evi-
dence. Plains Anthropologist 43:201-219
CurisTeNseN, N.L. 2000. Vegetation of the southeastern coastal plain. In: M.G. Barbour and
W.D. Billings, eds. North American terrestrial vegetation. Cambridge Univ. Press, New
York. Pp. 397-448.
Currie, D.J. 1991. Energy and large-scale patterns of animal- and plant-species richness.
Amer. Naturalist 137:27-49.
Detcourt, H.R. and PA. Decourt. 2000. Eastern deciduous forest. In: M.G. Barbour and W.D.
Billings, eds. North American terrestrial vegetation. Cambridge Univ. Press, New York.
Pp. 357-395.
Dice, L.R. 1943. The biotic provinces of North America. Univ. Michigan Press, Ann Arbor.
Diccs, G.M., B.L. Lirscome, and RJ. O’Kennon. 1999. Shinners & Mahler's illustrated flora of
north central Texas. Sida, Bot. Misc. 16:1-1626.
GEHLBACH, F.R. 1991.The east-west transition zone of terrestrial vertebrates in central Texas:
a biogeographical analysis. Texas J. Sci.43:415-427.
Kartesz J.T.and C.A. Meacham. 1999. Synthesis of North American flora.Version 1.0. North
Carolina Botanical Garden, Chapel Hill.
MacRoserts, M.H., B.R. MacRoserts, B.A. Sorric, and R.E. Evans. 2002. Endemism in the West
Gulf Coastal Plain:importance of xeric habitats. Sida 20:767-780.
MacRoserts, M.H.and B.R. MacRoseats. 2003.West Gulf Coastal Plain ecoregions. Sida 20:1 247-
1276.
Owen, J.G.and D.J.ScHMIDLY. 1986. Environmental variables of biological importance in Texas.
Texas J. Sci. 28:99-117.
ScHmioLy, D.J. 2002. Texas natural history:a century of change. Texas Tech Univ. Press, Lub-
bock.
Sorrle, B.A.and A.S. WeAKLEY. 2001. Coastal plain vascular endemics: phytogeographic pat-
terns. Castanea 66:50-82.
TAKHTAJAN, A. 1986. Floristic regions of the world. Univ. California Press, Berkeley.
TAMU (www.csdl.tamu.edu/FLORA/tracy2/main1 htm).
THorne, RF. 1993. Phytogeography. In: Flora of North American Editorial Committee, ed.
Flora of North America. Vol. 1.Oxford Univ. Press, New York. Pp. 132-153.
1700 BRIT.ORG/SIDA 20(4
=
Turner, B.L., H. NicHots, G.C. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas.
Sida, Bot. Misc. 24:1-888.
Warp, R., E.G. Zimmerman, and T.L. Kinc. 1994. Environmental correlations to terrestrial reptil-
ian distribution in Texas. Texas J. Sci.4:21-26,
Wess, W.L. 1950. Biogeographical regions of Texas and Oklahoma. Ecology 31:426-433.
REDISCOVERY OF TEPHROSIA JAMNAGARENSIS
(FABACEAE), AN ENDANGERED AND NARROW ENDEMIC
PLANT SPECIES OF SAURASHTRA, GUJARAT, INDIA
PS. Nagar SJ. Sata
Department of Biosciences Department of Biosciences
Saurashtra University Saurashtra University
Rajkot-360 005, INDIA Rajkot-360 005, INDIA
S.J.Pathak
Department of Biosciences
Saurashtra University
Rajkot-360 005, INDIA
ABSTRACT
During recent botanical explorations of Saurashtra, Gujarat, India, the narrowly endemic plant spe-
cies Tephrosia jamnagarensis was rediscovered after a lapse of 3 decades. A full description of the
species is provided along with photographs: a key to closely related species is included.
RESUMEN
Taeanas S | Gujarat, India, | i lémica Tephrosia
Sanecanee fue redescubierta despurs de un intervalo eae tres Gerad se omeeel una ceScupCIOn
completa de la especie junto con fotografias y se incluye par |
proximas as.
INTRODUCTION
During a floristic survey of Jamnagar District (Nagar 2002), Tephrosia
jamnagarensis Santapau—an endangered and endemic species of Saurashtra
(Kothari & Hajra 1983; Nayar & Sastry 1988; Santapau 1958; Shah 1983)—was
collected from the lower slopes of the Khad Khambhaliya Vidi (Grassland),
Lalpur Taluka, Jamnagar District, Gujarat, India. Fieldwork in 2001 verified the
occurrence from Jamnagar District, though not from the same locality.
Tephrosia jamnagarensis was not recognized as a distinct species in earlier
floras of the state (Cooke 1901-1903; Thaker 1910). The specific epithet
jamnagarensis refers to the type locality.
The species was collected for the first time by Santapau (Santapau 7522)
near Rozi, Jamnagar District, Gujarat, India in 1945 (Santapau 1958) and for the
second time some LI years later (24 Aug 1954) from Victoria Bridge, Jamnagar
District (Santapau 1962). A few plants were collected in 1972 from wastelands
near the cultivated fields of Bhadbhut, close to Bharuch (South Gujarat), a dif-
ferent sub-humid geographical region (Shah 1978; Vyas 1973). A recent survey
SIDA 20(4): 1701-1705. 2003
1702 BRIT.ORG/SIDA 20(4)
for T. jamnagarensis in these same locations did not locate it. The areas have
been mostly converted to agricultural land (Nagar 2000; Rao 2002). The recent
collections of T. jamnagarensis represent not only the rediscovery of this very
interesting taxon—alter a gap of almost 3 decades—but are also the first avail-
able collections that allow for a full description of the flower
With this recent collection in flower and fruit, a detailed description of the
species with floral characteristics, phenology, and habit and habitat distribu-
tion is given; photographs are provided.
DESCRIPTION OF TEPHROSIA JAMNAGARENSIS AND |
COMPARISON WITH TEPHROSIA STRIGOSA
Tephrosia jamnagarensis Santapau in Proc. Natl. Inst. Sci. India 24B:133,t. 1.1958.
- a _ Type: INDIA. GUJARAT. Jamnagar District: near Rozi, 16 Oct 1945, Santapau
2 (HOLOTYPE: BLAT).
Erect or suberect, annual herb; stems simple or sparsely branched, covered with
whitish appressed hairs. Leaves simple, 3-5.2 x 5-9 mm, linear, glabrous
adaxially, densely hairy with silvery appressed hairs abaxially, subobtuse and
clearly apiculate at apex, acute at base; lateral nerves 25-30 pairs, parallel; mar-
gin entire, with a nerve running from near the base to the apex very near the
margin; the nerves covered with hairs, distinct on the adaxial surface, equally
distinct on the abaxial surface. Petioles 2-3 mm, hairy; stipules subulate, 3-4
mm, hairy. Flowers single or in pairs, axillary; pedicels 2-3 mm, about as long
as the petioles, densely hairy. Calyx, 1-3 mm, hairy, the teeth subulate, filiform,
subequal. Corolla mauve, 2.5-3 mm (standard 3 mm, wings 2-2.5 mm, keel 2.5
mm). Legumes compressed, ca. 20 x 5 mm, densely and patently hairy with
grayish tinge, oblique at both ends, apiculate; seeds 5-6, reniform, dull or matt,
brownish.
Tephrosia jamnagarensis is similar to T. strigosa (Dalz.) Sant. & Mahesh, dif-
lering mainly by its broader (twice as broad as in T. strigosa), densely pubes-
cent pods and by its larger leaves with densely appressed hairs on the abaxial
surface, which is clearly silvery or densely argenteo-canescent, making it very
distinct from TF. strigosa. The pedicels are shorter and stouter than in T.strigosa.
The seeds are not separated by any internal partition (partitioned in T. strigosa).
—
Voucher specimens: INDIA. Bhs eerie District: Lalpur Taluka: Khad khmbaliya Vidi (Grass-
land), 17 Oct 2001, PS. Nagar 1221, 1222, 1234, 1235 (sauuni= Saurashtra University).
Additional specimens examined: I - Gujarat. Jamnagar District: near Rozi, 16 Oct 1945,
Santapau 7522 (HoLotype: BLAT): ae 12 Oct 1972, KJ. Vyas 3095 (spu); Bharuch, 18 Sep 1972,
KJ. Vyas eee. 21 Sep KJ. Vyas 2946 (spud. Spec 5 other than the holotype are depos:
ited in the herbarium of Department of Biosciences, oe Patel University (spu), Vallabhvidyanagar,
Gujarat, India.
Phenology and Associated Species —The plants were in flower from September
to October, in fruit, from October to November. They grew in grassland with
NAGAR ET AL., REDISCOVERY OF TEPHROSIA JAMNAGARENSIS 1703
; a rae “¢ A Hahit | eat:R. Adayial surface: C. Abaxial surf. D. Flowering twig: E. Calyx:
g twig J
Fic.1 h
F. Ovary; G. Pod; H. Seed ornamentation; I. Seeds.
1704 BRIT.ORG/SIDA 20(4)
Chrysopogon fulvus (Spr) Choiv, Sehima nervosum (Rottl.) Stapf, Hetropogon
contortous (L.) P. Beauv. ex R. & S,, Aristida sp. Borreria stricta (L.£.) K. Schum,
Zornia diphylla auct., Indigofera cordifolia Heyne ex Roth, Indigofera tinctoria
L.,and Alysicarpus vaginalis (L.) DC.
Habitat and Population Status.—The plants were growing on calcareous
gravelly soil. The species is very rare. A quantitative analysis of the population
indicated an estimated density of 0.24 plants/m? and frequency of occurrence
of 6% (India: Gujarat. Jamnagar District. Khadkhambaliya Region, Lalpur Taluka
(Nagar 2002). The rarity of the plant is probably due to overgrazing, habitat
destruction, lack of protection, and probably other biotic factors. The plant has
been listed as rare under threat category (Botanical Survey of India 1993; WCMC
1994).
Distribution.—The geographical range of T jamnagarensis Sant. is restricted
to two separate regions: the western group of populations in northwestern
Jamnagar (Rozi and Khadkhambaliya) and the southern population in south-
west Bharuch.
Potential Value.—Several species of Tephrosiale.g., T. purpurea (L.) Pers.Jare
sometimes grown as green manure and as cover crops (Santapau 1962); T.
purpurea is useful medicinally [the whole plant is used internally as purgative,
laxative, and tonic; externally it is applied on skin boils (Thaker 1910)} Itis likely
that T. jamnagarensis may also have similar properties.
ACKNOWLEDGMENTS
We thank the Department of Biosciences, Saurashtra University, Rajkot, for fi-
nancial support; S.M. Almeida for providing access to the Blatter Herbarium
(BLAT), Mumbai, India and for confirming our identification of the species; and
A.S. Reddy for giving access to the Departmental Herbarium, Department of
Biosciences, Sardar Patel University, Vallabh Vidyanagar, India.
HL
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\tra
NAGAR ET AL., REDISCOVERY OF TEPHROSIA JAMNAGARENSIS 1705
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(BOTANY-A complete and comprehensive account of the flora of Barda Mountain
(Kathiawad), Gujarati Printing Press, Bombay.P.717.(2nd revised ed.;ed.B.G. Shah (1952),
Sastu Sahitya Vardhak Karyalaya, Anmedabad. P. 733).
Vyas, KJ. 1973. Contribution to the floristics and phytosociology of the river Narmada in
Gujarat State. Ph.D. thesis. S.P. University, Vallabh Vidyanagar, Gujarat.
WCMC 1994. Threatened plants of the World. http://www.wemc.org.uk/species/plants
redlists.Atm.
1706 BRIT.ORG/SIDA 20(4)
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Editor: Roland Hardman. Taylor & Francis, 11 New Fetter Lane, London
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SIDA 20(4): 1706. 2003
DOES POLYGONUM PATULUM (POLYGONACEAE)
GROW IN NORTH AMERICA?
Mihai Costea' Francois J. Tardif
Department of Plant Agriculture Department of Plant Agriculture
University of Guelph University of Guelph
Guelph, Ontario, NIG 2W1, CANADA Guelph, Ontario, NIG 2W1, CANADA
coste_amihai@hotmail.com ftardif@uoguelph.ca
ABSTRACT
Plants previously considered Polygonum ea Bieb in California are within the range of varia-
tion of Polygonum ramosissimum Michx. complex. The real Polygonum patulum is recorded now for
the first time from Alabama, Illinois, on a Washington, where it has occurred asa rare ruderal.
RESUMEN
Las plantas consideradas previamente Polygonum patulum en California se encuentran dentro del
rango de variacion del complejo de Polygonum ramosissimum. El verdadero Polygonum patulum es
citado ahora por primera vez en Alabama, Illinois, Utah y Washington, donde ha aparecido como un
ruderal raro.
The Eurasian and North African Polygonum patulum M. Bieb. has been listed
inall standard Californian floras (e.g., Munz @ Keck 1968; Hickman 1993). How-
ever, Hickman (1993) noted that the “applicability of this name to Californian
plants remains uncertain.” After comparing these plants with the European
specimens of P patulum, we can confirm that they do not belong to this spe-
cies. The Californian plants resemble P patulum in having a terminal spiciform
inflorescence (with the leaves in the upper part of inflorescence reduced to
bracts which are shorter than the flowers they subtend) and small flowers and
fruits of 2.5-3 mm, and 2.2-3 mm, respectively. However, the achenes of P
patulum are striate-tubercled, and the inner and outer tepals have about the
same length. Plants from California have smooth to roughened achenes, and
the inner tepals are evidently shorter than the outer tepals. Many of these plants
are identical to some of the specimens quoted by Small when he described P.
exsertum (Small 1894): United States of America. Missouri: St. Louis, Sep 1845,
G. Engelmann s.n. (NY). NEBRASKA: Valentine, 1891, J.M. Bates s.n. (NY). Unfor-
tunately, the protologue and the subsequent descriptions of P. exsertum have
placed emphasis only on the much elongated late-season achenes, which are
taxonomically irrelevant (e.g. Fernald 1950). The morphology of the perianth
and achenes during early summer has been not explored. Consequently, P.
exsertum has been considered by all the authors to be asynonym of Polygonum
ramosissimum (e.g. Gleason 1952; Mertens & Raven 1965; Wolf & McNeill 1986).
\Correspondina author.
I
SIDA 20(4): 1707-1708. 2003
1708 BRIT.ORG/SIDA 20(4)
Polygonum ramosissimum is a complex species similar in that regards to P
aviculare (Costea & Tardif 2003). Polygonum exsertum and possibly other spe-
cies previously synonymized with P ramosissimum, may deserve taxonomic
recognition as infraspecific taxa of P. ramosissimum. Nevertheless, further re-
search is needed to understand the patterns of variation within P ramosissimum.
For the time being, it is safe to conclude that what has been considered P
patulum in California is within the range of P. ramosissimum variation.
The real P patulum isa rare adventive in North America. During the prepa-
ration of the genus Polygonum for Flora of North America we have found only a
few collections in the United States of America that can be ascribed to this taxon,
and these are new floristical records. Except for P. ramosissimum, P. patulum
may be mistaken only with P.argyrocoleon Steud. ex Kunze. The latter has smooth
achenes and the terminal spiciform inflorescence is congested, dense, with
cymes having 4-5 flowers. Polygonum patulum has striate-tubercled achenes,
and the terminal inflorescence is open, with cymes having 1-4 flowers.
Typical speci |: U.S.A. ALABAMA. Mobile Co.: Mobile, adventive with ballast, Oct 1893,
C. Mohr s.n. (NY). ILLINOIS. Union Co.: 4.3 mi S of Jonesboro, disturbed clay, rather common road-
side weed, 23 Sep 1999, S.R. Hill 32090 (NY). UTAH. Uintah Co.: NW corner of Pelican Lake, 1250 m,
in moist soil of edge of lake, 8 Aug 1983, E. Neese 14645 (NY). WASHINGTON. Grays Harbor Co.:
Montesano, in fields, Sept 1917, J.M. Grant (NY).
ACKNOWLEDGMENTS
We thank James Reveal and Cecil Slaughter, Jr. for providing critical edits to
earlier drafts of the manuscript. We also warmly thank the curators from ACAD,
ALTA, CAS, DAO, DC, DS, GH, F JEPS, LL, MT, MTMG, NFLD, NSPM, NY, PO,
QFA, QUE, RSA, S, SASK, TEX, UBC, UC and US for preparing the (usually) vo-
luminous Polygonum loans.
REFERENCES
Costea, M. and FJ. Taroir. 2003. Nomenclatural changes in the genus Polygonum, section
Polygonum (Polygonaceae). Sida 20:987-997.
FERNALD, M.L. 1950. Gray’s manual of botany. 8th ed., American Book Co., Boston, MA.
Gteason, H.A. 1952. The new Britton and Brown illustrated flora of the northeastern United
States and Canada. Lancaster Press. Lancaster, PA.
Hickman, J.C. 1993. Polygonum L. In: Hickman, J.C. Ed., The Jepson manual: higher plants of
California, University of California Press, Berkeley. Pp. 866-892.
Mertens, T.R. and PH. Raven. 1965. Taxonomy of Polygonum section Polygonum (Avicularia)
in North America. Madrono. 18:85—92,
Munz,P.A.and D.D. Keck. 1968. A California flora. University of California Press, Berkeley, CA.
SMALL, J.K. 1894. New and interesting species of Polygonum. Bull. Torrey Bot. Soc. 21:168-
es
Wotr, S.J.and J. McNett. 1986. Synopsis and achene morphology of Polygonum section
Polygonum (Polygonaceae) in Canada. Rhodora 88:457-479.
POLYGONUM AVICULARE SUBSP. RURIVAGUM
(POLYGONACEAE) IN NORTH AMERICA
Mihai Costea' Francois J. Tardif
Pearman! of Pl ie It Department of Plant Agriculture
versit University of Guelph
Guelph hanes Nt 1G WI, CANADA Guelph, Ontario, NIG 2W1, CANADA
coste_amihai@hotmail.com ftardif@uoguelph.ca
ABSTRACT
The occurrence of Polygonum aviculare subsp. rurivagum in North America is confirmed. The taxon
is a rare adventive from Europe growing in sunny, dry and trampled ruderal habitats.
RESUMEN
La octane de oot aviculare subsp. rurivagum en Norteamérica es confirmada. Fs un taxon
ropa que crece en lugares ruderales, soleados, secos y pisoteados.
The occurrence of the European Polygonum aviculare L. subsp. rurivagum Qord.
ex Boreau) Berher in North America has been uncertain. Love and Léve (1956)
reported it from eastern North America but they did not indicate any location.
Furthermore, their results have been generally disregarded because of some
doubtful chromosome counts, and because no specimens or vouchers have been
quoted or found (e.g., Mertens & Raven 1965; Wolf & McNeill 1986). During the
preparation of the genus Polygonum for Flora of North America we have en-
countered this taxon in scattered locations in North America and we compared
it with European material. Some relatively recent European floras and treat-
ments of Polygonum have preferred to treat this taxon asa species (P rurivagum
Jord. ex Boreau, e.g., Akeroyd 1993; Do Amaral Franco & Da Luz Rocha Afonso
1995; Stace 1997) following the study of Styles (1962). Nevertheless, its classifi-
cation under P aviculare is apparently the best solution (Karlsson 2000; Costea
& Tardif 2003). Itisa rare but homogenous taxon. Its extreme morphology, with
pauciflorous cymes (1-3 flowers), small flowers, 2.3-3.2 mm, and narrow-ellip-
tic to linear-lanceolate leaves, 1.5-5 mm wide, brings it close to P. aviculare subsp.
neglectum (Besser) Arcang,, with which it has been frequently misidentified.
The subsp. rurivagum differs by the long ochreas, 8-12.5 mm, with strong veins
and the free part disintegrating into persistent fibers, and by the leaves with
lateral veins strongly raised on the abaxial part of the leaves. The subspecies
neglectum has ochreas of 4-8 mm, with inconspicuous veins and free part leav-
ing almost no fibrous remains and leaves without strongly raised lateral veins.
For differences between subsp. rurivagum and the other subspecies of P.
—
‘Corre pondina author.
Vv ~]
SIDA 20(4): 1709-1711. 2003
1710 BRIT.ORG/SIDA 20(4)
aviculare from North America see Costea and Tardif (2003). For iconography
see Raffaelli (1982, Fig. 12), Karlsson (2000, Fig. 96). The plant grows as a rud-
eral in dry, sunny and often trampled habitats, in pavement cracks, gravel and
sandy a Flowering occurs from June to October
Ts a |
y] | *ANADA. ALBERTA: S of Cypress Hills, elias eae ee
lo Aug 1951, EH. Moss 9632 (ALTA); 2.25 mi S of Fort Saskatchewan, 8 Aug 1944, EH. Moss & G.H.
lurner 6642 (ALTA, DAO); Ma-Me-O Beach, edge of paved highway, 33 Jul 7 G.H. Turner 9535
(ALTA, DAO). MANITOBA: Otterburne, en bordure de la voie ferrée, au nord de la localité, 13 Aug
1956, J.-P Bernard 5586a (QFA). NEW BRUNSWICK. Cloucester Co.: near Caraquet, on riverbank, 22
Jul 1945, W.G. Dore & E. Gorham 45753 (MT). NEW FOUNDLAND. Hermitage Distr.: Harbor Breton,
brackish pond behind beach, muddy gri — shore, 19 Jul 1989, A. Bouchard et al, 89250 (MT). QUE-
BEC. Comté de Bagot: St-Pie, 40°30'N, 72°54W, sous le pont traversant la riviére Yamaska; terrain
vague, prés des ae du pont du cote est de la riviére, avec Polygonum ramosissimum, 12 Sep 1992,
M. Blondeau 92154 (QFA). Comté de Rimouski: Le Bic, pa airie saumatre avec: Salicornia, Spartina
patens, Limonium, etc., 16 Aug 1962, Rolland-Ger 8202 (MT). SASKATCHEWAN. Regina Distr:
Reg ae eae empty lot N of Luther Care Home, 21 Aug 2000, G.E Ledingham 13707 (USAS).
FORNIA. Kern Co.: Breckenridge seven Road, at the $ fork of Lucas Creek, 1833
m, sea pine rest 27 Jul 1905, E.C. Twisselmann [182 S). Sonoma Co.: near Kenwood, 18 Sep
1932, J. 1. Howell 10780 (CAS). INDIANA. Howard Co.: F of ea in dry cindery soil, 22 Jul 1941,
“Chas. M. Ek.” sn. (NY). IOWA. Marion Co.: Knoxville Twp, wet clay adjacent to water in strip mine,
23 Jul 1957, LV. Bruggen sn. (DAO). MAINE. Androscoggin Co.: Valley of Androscoggin River, S of
Polland, 1895, K. Furbish s.n. (GH). Aroostook Co.: Valley of Meduxnekeag River, roadside, 12 Aug
1909, MLL. Fernald s.n. (GH). Cumberland Co.: Brunswick, Sep 1891, K. Furbish s.n. (GH).
MASSACHUSETS. Hampshire Co.: Ryan Road, North: eee waste ground, gravel pit, 11 Aug 1978,
H.E. Ahles 86179 (QFA). Middlesex Co.: N of Wobum, 1951, G.H. Hilton s.n. (GH). Worcester Co.: S of
Royalston, 1907, J.A. Bates s.n. (GH). NEW YORK: New York ae near Van Cortland Park, in sandy
soil, 17 Jul 1891, EP Bicknell s.n. (NY). Rhode Island. Providence Co.: Elmwood, 30 Aug 1893, J. Col-
lins s.n. (GH). (SOUTH?) DAKOTA: (Augustana?) College, 1889, J.C. Zuffeys.n.(US). WEST VIRGINIA.
Hardy Co.: immediately E of Trout Run in cleared area leading to automatic Beacon Tower, 9 Aug
L941, H.A. Allard 9389 (US).
—
ACKNOWLEDGMENTS
We warmly thank James Reveal and Cecil Slaughter, Jr. for providing critical
edits to earlier drafts of the manuscript. We also thank the curators from ACAD,
ALTA, CAS, DAO, DC, DS, GH, F JEPS, LL, MT, MTMG, NFLD, NSPM, NY, PO,
QFA, QUE, RSA, S, SASK, TEX, UBC, UC and US for preparing the (usually) vo-
luminous Polygonum loans. Laura Echarte checked the Spanish translation of
the abstract.
REFERENCES
Axeroyb, J.R. 1993. Polygonum L.In:T.G. Tutin, N.A. Burges, A.O. Chater, J.R. Edmondson, V.H.
Heywood, D.M. Moore, D.H. Valentine, S.M. Walters and D.A. Webb, eds. Flora Europaea
(2nd ed.), Vol. 1, University Press, Cambridge, U.K. Pp. 91-97
Costea, M. and FJ. Tarbir. 2003. Nomenclatural changes in ca genus Polygonum, section
Polygonum (Polygonaceae). Sida 20:987-997.
COSTEA , 1711
Do Amaral Franco, J. and M. Da Luz RocHa Afonso. 1995. Espécies da sect. Polygonum
existentes em Portugal. Stud. Bot. 14:153-175.
Love, A. and D. Love. 1956. Chromosomes and taxonomy of eastern North American
Polygonum. Canad. J. Bot. 34:501-521.
KARLSSON, T.2000. The genus Polygonum L.|n:Jonsell, B.ed., Flora Nordica vol. 1,Stockholm.
Pp. 255-273.
Mertens, T.R.and PH. Raven. 1965. Taxonomy of Polygonum section Polygonum (Avicularia)
in North America. Madrono. 18:85-92.
Rarractu, M. 1982. Contributi alla conoscenza del genere Polygonum L.4.Le specie italiane
della sectia Polygonum.Webbia 35:361-406.
Stace, C.A. 1997. New flora of British Isles, 2nd ed., Cambridge University Press, Cambridge,
Srytes, B.T. 1962. The taxonomy of Polygonum aviculare and its allies in Britain. Watsonia.
5177-214,
Wotr, $.J.and J. McNett. 1986. Synopsis and achene morphology of Polygonum, section
Polygonum (Polygonaceae) in Canada. Rhodora 88:457-479.
1712 BRIT.ORG/SIDA 20(4)
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look at I cultivation, chemical and/or |
current and/or future produces: etc.
pharmacological aspects, toxicology,
Massimo Marrei (ed.). 2002. Vetiveria: The Genus Vetiveria. (ISBN 0-415-27586-5
hbk.). Medicinal and Aromatic Plants—Industrial Profiles. Volume 20. Se-
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From Taylor & Francis adie —‘The genus Vetiveria is one of the most versatile genera in plant
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many other applications.”
Contents.—1) Introduction to the Genus Vetiveria; 2) Anatomy, Biochemistry and Physiology;
3) Collection, Harvesting, Processing, Alternative Uses and Production of Essential Oil; 4) Chemical
Constituents and Essential Oil Biogenesis in Veliveria zizanioides; 3) Ethno pnarmacevey and Phar-
sete Properties of Vetiveria z
Propet
_
zizanioides—Including Pharmacologic and Pharmacokinetic
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and ee Needs, and Environmental Import e, 9) Beyond the Vetiver Hedge—Organizing
Vetiver’s Next Steps to Global Acceptance, and index.
SIDA 20(4): 1712. 2003
UVULARIA (LILIACEAE) IN TEXAS
Jason R. Singhurst William F. Carr and Wendy J.Ledbetter
Wildlife Diversity Program The Nature Conservancy of Texas
Texas Parks and Wildlife Department San Antonio, TX 78295-1440, U.S.A.
Austin, TX 78704, U.S.A
Walter C. Holmes
Department of Biology
Baylor University
Waco, TX 76798-7388, U.S.A.
ABSTRACT
This report d h | distribution of the Uvularia perfoliata and U. sessilifolia
in the state of Texas. A key to species, notes, distribution (including a map), and a list of exsiccatae
are presented.
RESUMEN
Secita y sed tala presencia distribucién de Uvularia perfoliata y U. sessilifolia en el estado
y P yy
de oe Se presenta una clave de especies, notas, distribucion (incluyendo un mapa), y una lista de
exsicatas.
Uvularia is a small genus of five species native to eastern North America from
Nova Scotia and southern Quebec south to Florida and westward to Texas and
North Dakota (Wilbur 1963; Utech & Kawano 2002). The plants are rhizoma-
tous perennial herbs with leafy, forked aerial stems. Leaves ey be sessile or
perfoliate. Plants normally have one or occasionally t hor yellow flow-
ers that are campanulate in shape.
Our objectives are to document knowledge of the expanded distribution
of Uvularia sessilifolia and Uvularia perfoliata in eastern Texas, summarize
the known history of both species within Texas, and provide a means of identi-
fying the species in the state. The paper is based upon recent field studies con-
ducted in the eastern portion of the state and examination of herbarium speci-
mens from selected herbaria (ASTC, BAYLU, & TEX).
oO
KEY TO THE SPECIES OF UVULARIA IN TEXAS
1. Leaves rounded at the base, perfoliate 1. U. perfoliata
1. Leaves narrowed at base, sessile, not perfoliate _ 2. U. sessilifolia
1. Uvularia perfoliata L
Uvularia perfoliata is distributed from New England and Ontario south and
westward to Florida, Texas and Arkansas (Wilbur 1963, Utech &@ Kawano 2002).
Within Texas, the species is known from Sabine, Nacogdoches, Shelby, and San
SIDA 20(4): 1713-1716. 2003
114 BRIT.ORG/SIDA 20(4)
Augustine Counties (Fig. 1),a distribution that is basically contiguous with the
distribution in Louisiana presented by Thomas & Allen (1993). The species oc-
curs in mixed beech-maple series hardwood forests.
Brown (1972) first mentioned the occurrence of Uvularia pefoliata in Texas
but provided neither specific location nor other documentation. Brown made
trips to rich woodlands in eastern Texas and was generally consistent about
making vouchers for his photos, but he either failed to do so in the case of U.
perfoliata or the collection was lost or destroyed (Mark Mayfield, pers. comm.).
The first documented record of this species in the state was Nixon and Damuth’s
(1987) report for Sabine County. In 1989, Bridges and Orzell presented an ac-
count of the species in Texas, citing seven collections, all from Sabine County.
Turner et al. (2003), in the Atlas of Vascular Plants of Texas, illustrate the distri-
bution of the species as being Sabine, Shelby, and St. Augustine counties. The
— is reported as occurring in Nacogdoches County for the first time.
In addition to distinctive perfoliate leaves, the species is characterized by
short | cm long rhizomes, scarious smooth-margined leaves, and terete hollow
aerial stems. In the following list of exsiccatae, only one specimen per county is
cited
Specimens examined: Nacogdoches Co.: 6 mi ENE of Cushing, upper spring creek drainage that drains
into Barnes Lake, Apr 1994, Singhurst 4912a (BAYLU). Sabine Co.: 4.1 mi E of Geneva, 22 Mar 1985,
Nixon 14347(ASTC). San Augustine Co.: 0.7 air mi NW of Black Ankle, Sabine National Forest, Tenaha
Ranger District, Compartment 65, 12 Apr 1989, Orzell 9209 (TEX). Shelby Co.: 0.3 air mi ESE of FM
2261 and FS Rd 106 jet in FE Hamilton, Sabine National Forest, Tenaha Ranger District,Com
50, 12 Apr 1989, Orzell 9261 (TEX).
artment
|
2. Uvularia sessilifolia L.
The distribution of Uvularia sessilifolia extends from southern Canada (Ontario
and Nova Scotia), south to Florida and west to Texas, Oklahoma, and North
Dakota (Wilbur 1963, Utech & Kawano 2002). Uvularia sessilifolia has been
documented in three locations in Newton County and one location in Jasper
County (Fig. 1), approximately 280 km from the Cass County location cited in
this paper,and 210 km from its nearest Louisiana record in Lincoln Parish (Tho-
mas & Allen, 1993). The species occurs in moist hardwood and bottomland for-
ests, but in more xeric sites than U. perfoliata. The distribution map of this spe-
cies in Flora of North America (Utech & Kawano 2002) shows it occurring in
extreme northeast Texas, perhaps Bowie or possibly Lamar and Red River coun-
ties (the small scale of the map prevents accurate determination of county),
which all border Oklahoma and the Red River. Examination of specimens from
these counties determined to be U. sessilifolia showed them to be immature and
sterile specimens of Polygonatum biflorum (Walt. Elliott. (Liliaceae), a species
that is uncommon and sporadic where it occurs in east Texas. The Cass County
specimen cited is the only report of the species outside of the longleaf pine re-
gion of southeast Texas (Newton and Jasper counties).
SINGHURST ET AL., UVULARIA IN TEXAS
1715
Size
=
tame
sertieetescesars
Srsssersecense ys
SBN sasmtencerest
KO A EEC
UL ASRS
yaanecst
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aH
Au
Newtor
font “> Jasper
Fic. 1 i, er ee ee fyi t z. p ype bg { 4 gl \ Ail Pes 1 \ my
In addition to sessile non-perfoliate leaves, Uvularia sessilifolia can be dis-
tinguished from U. perfoliata by its longer rhizome of 10 cm or more in length,
scarious margined leaves that are very finely serrulate, and aerial stems that
are angled and solid. In the following citations of specimens, both the Cass
county and Jasper County records are reported for the first time. Because of the
1716 BRIT.ORG/SIDA 20(4)
aes of the species in the state, all collections are cited.
|: Cass Co.: State R ional Area, base of Dogwood Trail at jct. of wood
nave 9 Mar 2002, Singhurst a aia U). Jasper Co.: Temple Inland Distinctive Site, Beef Creek,
Sof Rayburn Country, jet. of R255 and private forest road between Rayburn Country and Lake Sam
Rayburn Dam, | mi S of forest na 6 Mar 2002, Singhurst 11213 (BAYLU). Newton Co.: Little Quick-
sand Creek on FM 1414 ca. 2.8 mi E of jct of Tex. Hwy 87 N of Newton, 3 Apr 1997, Carr 15983 (TEX):
Hickman Creek ca. 0.4 mi W of jet. with FM 645, 3 Apr 1998, Singhurst 6591 & Watson (BAYLU):
Mack Creek, ca. 4 mi FE of Newton on FM 1414, 4 Mar 2000, Holmes 10795, Singhurst, & Caldwell
(BAYLU).
ACKNOWLEDGMENTS
We are grateful to Tom Wendt (TEX) for his assistance and Mark Mayfield (for-
merly of LSU, now of KSC) for information on the collections of Claire Brown
We also acknowledge the cooperation of the United States Army Corps of En-
gineers for granting access to public lands near Lake Wright Patman in Bowie
and Cass counties. Richard Carter (VSC) and Heather White (BAYLU) reviewed
preliminary versions of the manuscript.
REFERENCES
Bribes, E.L.and S.L.Orzett. 1989, Additions and noteworthy vascular plant collections from
Texas and Louisiana, with historical, ecological, and geographical notes. Phytologia
66:12-69.
Brown, C.A.1972.Wildflowers of Louisiana and adjoining states. Louisiana State University
Press, Baton Rouge, LA.
Nixon, E.S.and S.C . DamutH. 1987. Five additions to the Texas flora. Sida 12:421—422.
THomas, R.D. and CA. Atten. 1993. Atlas of the vascular flora of Louisiana. Vol. |: Ferns and
fern allies, conifers, & monocotyledons. Louisiana Dept.Wildlife & Fisheries, Baton Rouge.
Turner, B.L., H. NicHots, G, Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas, Vol.
2, Sida Bot. Misc. 24, Botanical Research Institute of Texas, Ft. Worth
Utech, FH. and S. Kawano. 2002. Uvularia In: Flora of North American editorial committee
eds.), Flora of North America. vol. 26. Magnoliophyta: Liliidae: Liliales and Orchidales.
Pp. 147-150.
Witsur, R. L. 1963. A revision of the North American genus Uvularia (Liliaceae). Rhodora
65:158-188
—
FAGOPYRUM ESCULENTUM (POLYGONACEAE),
NEW FOR TEXAS
Robert J.O’Kennon Caren McLemore
Botanical Research Institute of lexas Botanical Research Institute of Texas and
University of North Texas
Fort Worth, TX 76102, U.S.A. Denton, TX 76203, U.S.A
okennon@brit.org clm0093@unt.edu
Amanda K. Neill
Botanical Research Institute of Texas
09 Pecan Street
Fort Worth, [X 76102-4060, U.S.A.
aneill@brit.org
Collections of Fagopyrum esculentum Moench (Polygonum fagopyrum L.), buck-
wheat, were made in May and June of 2003 in the Lyndon B. Johnson National
Grasslands near Decatur, Texas (Wise County). This native of Central Asia is
reported for Oklahoma, New Mexico, and Louisiana by Kartesz (1999). Despite
this proximity, it has not previously been reported for Texas (Correll & Johnston
1970; Stanford 1976; Hatch et al. 1990; Diggs et al. 1999). Since other populations
of F esculentum have not been found in North Central Texas, it is unknown if
the species is more widely established.
The generic name Fagopyrum is derived from Latin “fagus,” meaning beech,
and “pyros,” meaning wheat, because of the resemblance of the plant's seed to
the beech-nut. The English name is derived from the German “buchweizen”
(Fernald 1950) or acorruption of “boek-weit,” the Dutch form of the word “buck-
wheat” (Grieve 1931).
Fagopyrum esculentum is one of a continuing procession of introduced
species (ie. those originating outside the United States) finding its way into the
Texas flora (O’Kennon et al. 1999). Taxa such as F esculentum are variously re-
ferred toas “alien,” “exotic,” or “foreign.” Ina recent floristic study of North Cen-
tral Texas (Diggs et al. 1999), it was determined that 17.7% of the flora 394 spe-
cies) consisted of introduced species.
Lyndon B. Johnson National Grasslands is comprised of numbered “units”
scattered throughout much of north central Wise County. Fagopyrum
esculentum was found in two widely separated units. At the collection site in
Unit 71, two populations, approximately 300 meters apart with about twenty
individuals at each site, were observed growing along a moist unpaved lime-
stone road, in the roadbed and in the ditch along the road. Plants were young
and about 12-18 cm tall. In Unit 3,18 km west of Unit 71, about 100 plants were
SIDA 20(4): 1717-1720. 2003
1718 BRIT.ORG/SIDA 20(4)
observed in a sandy open woodland. Individuals in this population were as tall
as Zc.
Fagopyrum esculentum (Fig. 1) can be recognized by the following descrip-
tion (modified from Barkley 1986): annual erect herb, from a slender taproot, to
50 cm tall. Stem glabrous to pubescent at the nodes. Leaves long-petiolate be-
O’KENNON ET AL., FAGOPYRUM ESCULENTUM NEW FOR TEXAS 1719
low, becoming sessile above, acuminate, the base broadly sagittate, cordate or
truncate; stipules glabrous to puberulent, acute, sheathing but open on one side.
cemes axillary, single or in corybiform to paniculate groups, these 1 per leaf
axil, long-pedunculate and usually exceeding the subtending leaf; bracts gla-
brous, ovate, acute; pedicels not jointed. Perianth white to creamy, streaked tan,
5-lobed, connate below, 3-4 mm long; stamens 8; styles 3, free. Achene strongly
exserted, 3-angular, shiny, dark brown. (n = 8, 2n = 16) May-Sep(-Oct). Culti-
vated fields, disturbed places, waste ground; escaped in many parts of US. &
the world; Asia. Introduced.
Curiously, the first record of Fagopyrum in Texas was found ina plant scrap-
book created in 1925. This bound collection of pressed plants, notes, and illus-
trations was donated to BRIT at some time in the past, but the source is un-
known. The plants were collected in Fort Worth, Texas, by MJ. Joyslin. Though
a pressed specimen of F esculentum and a fine watercolor of the subject were
both preserved on one of the pages, they had been misidentified. As a result of
awareness of the new Wise County collections, the scrapbook page was
serendipitously recognized as the first, albeit unofficial, collection known from
the state.
Voucher seine TEXAS. Wise Co.: along limestone roadway through Grand Prairie in Unit 71 of
the Lyn B. Johnson National Grasslands, N33° 18' 35", W097" 36' 35", 30 May 2003, O’Kennon,
McLemore none Neill 18432 ae wie sae ae open forest of Western Cross Timbers in Unit 3
of the Lyndon B. Johnson Nati N33° 22' 30", W097" 46'19", 5 Jun 2003, O’Kennon and
McLemore 18453 (BRIT).
ACKNOWLEDGMENT
Thanks to Guy Nesom of BRIT for assistance in identifying this plant.
REPERENGES
Barkcey, T.M. (ed.). 1986. Flora of the Great Plains. Great Plains Flora Association. Univ. Press
of Kansas, Lawrence.
Corrett, D.S. and M.C. JoHnston. 1970. Manual of the vascular plants of Texas. Texas Re-
search Foundation, Renner.
Dices, G.M. Jr, B.L. Lirscome, and R.J. O’Kennon. 1999. Shinners & Mahler's illustrated flora of
North Central Texas. Sida, Bot. Misc. 16.
FerNaco, M.L. 1950. Gray’s manual of botany. Dioscoroides Press, Portland, Oregon
Grieve, M. 1931.Buckwheat.|n: C.F. Leyel,ed. A modern herbal. Harcourt, Brace & Company.
Electronic version first published in 1995 by Dover Publications, Inc. Accessed in Au-
gust 2003 at http://botanical.com/botanical/mgmh/b/buckwh81.html.
Hatch, S.L., K.N. GANDHI, and L.E. Brown. 1990. Checklist of the vascular plants of Texas. Texas
Agric. Exp. Sta. Misc. Publ. No. 1655.
Jaume Saint-Hitaire, MJ.H. 1828-1832.La flore et la pomone frangaises: ou, Histoire et figure
en couleur, des fleurs et des fruits de France ou naturalisés sur le sol frangais. Vol 5.
Paris, Chez l‘auteur.
1720 BRIT.ORG/SIDA 20(4)
Kartesz,J.1.1999.A synonymized checklist and atlas with biological attributes for the vas-
cular flora of the United States, Canada, and Greenland. First edition. In: Kartesz, J.T.,and
CA. Meacham. Synthesis of the North American flora, version 1.0. North Carolina Bo-
tanical Garden, Chapel Hill.
O'KENNON, RJ. T.M. Barkley, G.M. Diaas, Jr.,and B. Liescome. 1999. Lapsana communis (Asteraceae)
new for Texas and notes on invasive exotics. Sida 18:1277-1283.
STANFORD J.W. 1976. Keys to the vascular plants of the Texas Edwards Plateau and adjacent
areas. Howard Payne University, Brownwood, TX.
TRIGLOCHIN CONCINNA JUNCAGINACEAE),
A NEW FAMILY, GENUS, AND SPECIES FOR TEXAS
B.L. Turner Matt W. Turner
The University of Texas at Austin The University of Texas at Austin
is nt Re ecourses ey McCombs School of Bu
MBA Progra i Ofee:
en 1X 7871 12- 0471, ae 1 University Station B6004
Austin, TX 78712-0205, U.S.A.
Triglochin isa genus of about 20 species, most of these confined to aquatic habi-
tats in the more temperate regions of the world (Thieret 1988). It has not previ-
ously been reported for Texas (cf. Haynes & Hellquist 2000; Turner et al. 2003),
although at least two species of the genus have been collected in New Mexico
(Correll & Correll 1972).
On July 26, 2003 a survey for plants along a saline aquatic habitat near
Ruidosa (Presidio County), Texas revealed Triglochin concinna var. debilis (M_E.
Jones) J.T. Howell (Fig. l) growing along the embankment of a perennial sulfur
stream. We have followed the taxonomy of Thorne (1993) in recording this taxon,
although we are aware that Haynes and Hellquist (2000) position the latter in
their broad concept of T. maritima L.
Voucher specimen: U.S.A. TEXAS. Presidio Co.: ca 2.2 mi NNW of Ruidosa growing along perennial
sulphur streams and cant tinajas. M.W. Turner Sones TEX).
The species was noted to be relatively uncommon at the site, although at least
four or five plants were noted along a path of about 40 meters. It was growing
with a number of other rarely encountered semiaquatic species of the Trans-
Pecos such as Anemopsis californica, Samolus sp., and Eustoma exaltatum.
The sampled site sits upon the property of Mr. James Blumberg of Ruidosa,
Texas. We are grateful to him for permitting us entry to his property; indeed, he
accompanied us on our impromptu visit, providing helpful comments on this
or that collecting site.
REFERENCES
Correll, D.S. and H.B. Corrett. 1972. Aquatic and wetland plants of southwestern United
States. EPA, Washington D.C.
Haynes, R.A. and C.B. Hettquist. 2000. Triglochin, in Fl.N. Amer. 22:44-46.
THieret, J.W. 1988. The Juncaginaceae in the southeastern United States. J. Arnold Arbor.
69:1-23.
SIDA 20(4): 1721-1722. 2003
1722 BRIT.ORG/SIDA 20(4
Fics. 1-2. 1. Triglochi 1 t hof t i f | y (Presidio C y, Texas) of Mr James
J t
er t
Rlamhern: 2. WWluctrati f R th f19C0\ U dh Lb anftho Rritich Cl fid OLA-i)
be J J
T. maritimum
THorne, R.F. 1993. Triglochin. The Jepson manual: higher plants of California. Berkeley: Univ.
of California Press.
Turner, B.L., H. NicHots, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas.
Sida, Bot. Misc. 2:649-888.
ANDROPOGON GLOMERATUS VAR. GLAUCOPSIS
(POACEAE: ANDROPOGONEAE) DOCUMENTED
IN LOUISIANA
David J.Rosen Joseph K.Wipff
U.S. Fish and Wildlife Service Herbarium, Pure Seed Testing, Inc.
17629 El Camino Real, Suite 211 PO. Box 449
Houston, TX 77058-3051, U.S.A. Hubbard, OR 97032, U.S.A.
ABSTRACT
>
Andropogon glomeratus (Walt.) B.S.P. var. glaucopsis (Elliott) Mohr is reported for Louisiana. A key to
separate the Louisiana taxa in the A. virginicus complex is provided.
RESUMEN
Se cita Andropogon glomeratus (Walt. B.S.P. var. glaucopsis (Elliott) Mohr de Louisiana. Se aporta una
clave para separar los taxa del complejo A. virginicus en Louisiana.
The Andropogon virginicus complex comprises nine closely interrelated spe-
cies (Campbell 1983). In Louisiana, the complex is represented by four of the
nine species: A. glomeratus (Walt.) B.S.P, A. gyrans Ashe, A. liebmannii Hack,
and A. virginicus L. The taxonomic difficulty of this complex is due to the strong
similarity of its members, and the paucity and subtlety of the characters used
to separate the taxa (Campbell 1983). A review of the literature reveals a differ-
ence of opinion in the recognition of the infraspecific taxa of the A. virginicus
complex in Louisiana.
Campbell (1983) reported the following taxa as occurring in Louisiana:
Andropogon liebmannii Hack. var. pungensis (Ashe) CS. Campb., A. glomeratus
(Walt.) B.S.P. var. pumilus (Vasey) Vasey ex L.H. Dewey, A.gyrans Ashe var.gyrans
and var. stenophyllus (Hack.) CS. Campb., and A. virginicus L. var. virginicus
and var. glaucus Hack. Thomas and Allen (1993) reported A. glomeratus var.
glomeratus, A.gyrans var.gyrans, A. liebmannii var. pungensis, and A. virginicus
in Louisiana, and listed A. glomeratus (Walt.) BSP. var. glaucopsis (Elliott) Mohr
as an undocumented sight record from St. Tammany Parish. Allen (1992) rec-
ognized A. liebmannii, A. glomeratus, A. gyrans, and A. virginicus as occurring
in Louisiana, but did not recognize any infraspecific taxa.
Fieldwork in southeastern Louisiana has resulted in the collection of
Andropogon glomeratus (Walt.) B.S.P. var. glaucopsis (Elliott) Mohr from St.
Tammy Parish.
4
Andropogon glomeratus (Walt.) B.S.P. var. glaucopsis (Elliott) Mohr, Bull. Torrey
SIDA 20(4): 1723-1725. 2003
1724 BRIT.ORG/SIDA 20(4)
—
Bot. Club 24:21. 1897. [= A. glaucus Muhl., A. capillipes Nash, Cympogon glaucua
Schult.]| LOUISIANA. ST. TAMMANY PARISH: N of Bayou Paquet Road, 0.3 mi E of its
intersection with Transmitter Road, along fire-line that follows refuge boundary
W of dirt road, 26 Oct 2001, Rosen & Jones 1664 (BRCH-Oregon, NO)
Keys to separate the taxa modified from Campbell (1983) and Allen (1992) fol-
low. See Campbell (1983) for a list of representative specimens from Louisiana.
The classification of taxa in the Andropogon virginicus complex provided here-
with follows that of Campbell (2003).
KEY TO THE LOUISIANA TAXA IN THE ANDROPOGON VIRGINICUS COMPLEX
1. Racemes (2—)4—6(-13) terminating each peduncle; sheaths villous A. liebmannii
var. pungensis
ees 2(—3) terminating each peduncle; shedths glabrous to villous.
. Plants usually less than 1 m tall; leaves usually less than 3 mm wide; raceme
sheaths inflated, usually more than 4.6 cm lonc A. gyrans
2. Plants larger, usually 1 mormore tall; teat more than 3 mm wide;racem
eee usually less than 4.6 cm lo
Blades usually more than 37 cm on stem sheaths often scabrous; ligules
usually more than 1 mm long (if less than 1 mm long the keels of the first
glume often scabrous below the middle) A. glomeratus
3. Blades usually less than 31 cm long; stem sheaths smooth (rarely somewhat
scabrous); ligules less than 1 mm long; keels of the first glume scabrous only
above the middle A. virginicus
KEY TO THE VARIETIES OF ANDROPOGON GYRANS IN LOUISIANA
1. Ligules usually less than 0.8 mm long; plants of well-drained sites A. gyrans var. gyrans
. Ligules (0.8-)1.1(-1.5) mm long; plants of wet sites .gyrans var. stenophyllus
KEY TO THE VARIETIES OF ANDROPOGON GLOMERATUS IN LOUISIANA
. Leaves and culms oe glaucous A. glomeratus var. glaucopsis
. Leaves and culms gr
2. Keels of first penis essiann below the middle; plants ee m tall
glomeratus var. pumilus
2. Keels of first glume scabrous above the middle only; plants (0.6- ‘ OC .6) m tall
Ralemicitiay var. glomeratus ee d in Louisiana)
KEY TO THE VARIETIES OF ANDROPOGON VIRGINICUS IN LOUISIANA
1. Leaves and culms conspicuously glaucous __ A. virginicus var. glaucus
1. Leaves and culms green A. virginicus var. virginicus
ACKNOWLEDGMENTS
We are grateful to Stanley D. Jones (BRCH), Fred Werner (USFWS), and two
reviewers for their helpful comments.
REFERENCES
Aten, C.M. 1992. Grasses of Louisiana, 2nd ed. Cajun Prairie Habitat Preservation Society,
Eunice, LA.
ROSEN AND WIPFE 1725
Campsett, C.S.2003. Andropogon. |n: Flora of North America Editorial Committee, eds. Flora
of North America North of Mexico, Vol.25(2).Oxford Univ. Press, New York. Pp.649-664.
Campselt, C.S. 1983. Systematics of the Andropogon virginicus complex (Gramineae). J. Ar-
nold Arbor. 64:171-254.
THomas, R.D.and C.M. Auten. 1993. Atlas of the vascular flora of Louisiana, |: Fern allies, coni-
fers,and monocotyledons. The Natural Heritage Program, Wildlife Div., Louisiana Dept.
of Wildlife & Fisheries, Baton Rouge.
1726 BRIT.ORG/SIDA 20(4)
BOOK NOTICE
TAYLOR & FRANCIS
Medicinal and Aromatic Plants—Industrial Profiles
Each volume in ~ series disct usses the ania and botany of each genus, and provides an in-c
vation, chemical and/or pharmacological aspects, toxicology,
er
epth
look at areas s ¢
current and/or fut ure eee etc.
WING MING KEUNG (ed.). 2002. Pueraria: The Genus Pueraria. (ISBN 0-415-28492-
9, hbk.). Medicinal and Aromatic Plants—Industrial Profiles. Volume 23. Series
Editor: Roland Hardman. Taylor & Francis, 11 New Fetter Lane, London EC4P
4EE, UK and 29 West 35" Street, New York, NY 10001, U.S.A. (Orders:
info@tandf.co.uk, www.andf. yee 44 (0) 207 583 9855, fax 44 (0) 207-
842-2298, standing orders call 44 (0)12604 343071. US and Canada Cus-
tomers: By mail: Taylor & Francis Customer Service, 10650 Toebben Drive,
Independence, K Y 41051, U.S.A,, Toll Free Tel: 1-800-634-7064, Toll Free Fax:
1-800-248-4724, email:cserve@routledge-ny.com). $95.00, 290 pp, numerous
b/w photos drawings and tables, 7" « 10".
From Taylor & Francis website.—“Pueraria has been woven in Asian societies for more than two mil-
lennia, having been used in cooking, weaving, decorating and treating human ailments. This is the
first book of its aoe to _ me blished on this medicinal and industrial plant genus. It has been as-
sembled with a br se readership in mind, including students, educators and oe
and treatment pes as well as research scientists from a broad range of disciplines.”
Contents.~1) Pueraria: botanical characteristics; 2) Ethnobotany of Pueraria species: 3)
Pueraria (Ge) in traditional Chinese herbal medicine; 4) Puerarta tuberose DC: contraceptive elfi-
hemical ts of Puerarid plants: identification and meth-
cacy and toxicological profile; 5) C
ods of analysis; 6) Chemical components and irhaenaclogs ol the ecibvenating plant abies
1 Rg) pI}
2,
—
mirificd; 7) Biosynthesis and ni peace enaias !
system; 9) Preclinical suidienel kudzu ( (Pueraria lo
rata)
of Pueraria isotlavon
as a treatment for sicehiel as 10) Human studies of kudzu as a treatment for alcohol abuse: 11)
effec sof | strogen-dependent diseases: ren ean cancer ol
C ee
the breast, aca ia ee ene 12) ¢ C ‘hema s and hepatoprotective effect of Pueraria sa-
ponins, 13) Mammalian metabolism of Pueraria isoflavonoids; 14) Synthesis of n aes occurring
isol eae their analogs; 15) Research and development of Pueraria(Ge)-based medicinal fg
nd chen:
ucts in China; 16) Kudzu, a valuable potential commercial resource: food, paper, textiles
cals; 17) Baand or foe? Changing cultural definitions of kudzu; and Index.
SIDA 20(4): 1726-. 2003
CAREX CONOIDEA (CYPERACEAE) NEW TO ARKANSAS
AND NOTES ON ITS OCCURRENCE
IN ARKANSAS AND MISSOURI
Paul M.McKenzie Tim Smith
U.S. Fish and Wildlife Service Missouri Department of Conservation
608 E. Cherry St.; Room 200 O. Box 180
Columbia, MO 65201-7712, U.S.A. Jefferson City, MO 65102-0180, U.S.A.
C.Theo Witsell
Arkansas Natural Heritage Commission
1500 Tower Building
ter St.
Little Rock, AR 72201, U.S.A.
The range of Carex conoidea Schkuhr ex Willd. extends from Newfoundland
and Nova Scotia west to Manitoba and south to Missouri, Illinois, Indiana, Ohio,
and North Carolina; disjunct and possibly introduced in Arizona (Radford et al.
1964: Mohlenbrock 1999; Yatskievych 1999; Naczi & Bryson 2002). Habitat listed
for the species includes “bogs,” “low ground, and moist grassy places,” “wet
meadows, wet prairies,” and moist depressions of upland prairies (Radford et
al. 1964; Voss 1972: Mohlenbrock 1999; Yatskievych 1999; Naczi & Bryson 2002).
The species has not been previously documented from Arkansas (Hyatt 1998;
Smith 1988; Arkansas Vascular Flora Committee 2002; Naczi & Bryson 2002).
On 8 May 2003, while conducting a plant survey on Searles Prairie Natural
Area in Benton County, Arkansas, we discovered 32 clumps of Carex conoidea
scattered throughout the western half of the preserve. The species is readily
recognized by the combination of light green leaves, scabrous peduncles and
culms below the inflorescence, bracts that subtend the uppermost pistillate
spikes equaling or exceeding the staminate spike, and its plump, lustrous green
perigynia with ca. 20 impressed nerves. Carex conoidea can be readily distin-
guished from other members of the Carex section Griseae (L.H. Bailey)
Kukenthalin Arkansas by the combination of its scabrous peduncles and culms
below the inflorescence, the fewer number of impressed nerves (others in the
Section usually have 40 or more nerves), and the position of the staminate spike,
well-exserted above the uppermost pistillate spike.
Various authors (e.g., Steyermark 1963; Mohlenbrock 1999; Yatskievych
1999) depict the perigynia of C. conoidea as being tapered at both ends. Illustra-
tors undoubtedly only had access to herbarium specimens or over-mature
SIDA 20(4): 1727-1730. 2003
1728 BRIT.ORG/SIDA 20(4)
perigynia, as fresh, mature perigynia of C. conoidea are plump and inflated, and
watermelon shaped. We have noticed the perigynia of C. conoidea shrink and
become tapered at both ends upon drying in a plant press
Clumps of Carex conoidea at the Searles Prairie site extend from swales in
the prairie to the drier margins of areas just upslope of the depressions. Despite
the field characters listed above, identification of C.conoidea in the field can be
difficult because many of the tufts may have few flowering culms and the culms
often lean on or are hidden by adjacent, taller vegetation.
Searles Prairie represents the last intact remnant of the Osage Prairies that
once surrounded the town of Rogers, Arkansas in Benton County. It was used
as a hay meadow and cut annually until it was donated to the Arkansas Natu-
ral Heritage Commission in 1988. Prescribed fire was introduced to the prairie
in 1996, with additional prescribed burns in December 2001 and February 2003.
Searles Prairie is a low loamy prairie that probably exists because of shallow,
impervious clay subsoil and fluctuations in water depth and persistence. This
provides graminoids and other ground-covering forbs a competitive advantage
over trees (Arkansas Natural Heritage Commission 2002). The Jay and Chero-
kee silt loam soils are characterized by layers of moderately well-drained silt
loam, to somewhat poorly drained silt loam, silty clay loam, and clay, This prai-
rie remains wet for long periods of time after rains due to the clay (United States
Department of Agriculture 1977) and poor drainage.
Plants associated with C. conoidea at Searles Prairie include Agrostis
hyemalis, Andropogon gerardii, Carex brevior, C. bushii, C. buxbaumii, C.
festucacea, Cephalanthus occidentalis, Cicuta maculata, Eleocharis verrucosa,
E. wolfii, Eryngium yuccifolium, Eupatorium perfoliatum, Galium obtusum,
Juncus spp. Orbexilum pedunculatum, Panicum acuminatum, P scoparium,
Physostegia angustifolia, Potentilla simplex, Pycnanthemum tenuifolium, Ra-
nunculus laxicaulis, Rubus spp, and Sisyrinchium angustifolium.
—
Voucher Specimens: U.S.A. ARKANSAS. Benton Co.: Searles Prairie Natural Area: inter. of US. Rt. 62
and Dixieland Rd. in Rogers, AR; TZONR30WS35SE1/4SE1/4SW1/4; Bentonville South 7.5’ Quad. 8
May 2003, McKenzie 2081 (DOV, MO), Witsell 03-0232 (anhe-Arkansas Natural Heritage Commis-
sion, LRU); & LL Jun 2003 Witsell 03-0474 (anhe-Arkansas Natural Heritage Commission, APCR,
LRU, peh-pers. herb. of Philip E. Hyatt, STAR, UARK).
Carex conoided is apparently a good indicator of high quality, well-managed
prairies in Missouri as well as Searles Prairie in Arkansas (Yatskievych 1999;
pers. obs.). The lack of records in some prairies may reflect the lack of survey
efforts by individuals familiar with the species’ field characters or the diff iculty
in seeing the diagnostic inflorescences among other prairie vegetation. Searches
for the species by the senior author in five prairies in Benton and Pettis coun-
ties, Missouri in early June 2002 failed to yield additional populations. Searches
in two additional Arkansas prairies by the authors on 9 May 2003 and five
MCKENZIE ET AL., CAREX CONOIDEA NEW TO ARKANSAS 1729
additional sites by Witsell in Arkansas in May and June 2003 failed to docu-
ment the species at additional localities. Subsequently, however, McKenzie and
Smith discovered the species at Hite Prairie in Morgan County and Taberville
Prairie in St. Clair County, Missouri on 6 and 7June, 2003, respectively [McKen-
zie 2085 (DOV, MO), 2090 (DOV, MO)]. These coll represent only the third
and fourth extant records of this sedge in Missouri. The plants were difficult to
see at the two Missouri sites, however, because the 20 and 30 clumps with few
flowering culms had inflorescences leaning on or hidden among adjacent, taller
vegetation. Despite the limitations mentioned above, future searches for C.
conoidea should be conducted in other high quality prairies throughout Ar-
kansas and Missouri from about | May to | June depending on the latitude of
the locality to be surveyed.
Botanical nomenclature listed herein follows Yatskievych (1999) or
Yatskievych and Turner (1990).
ACKNOWLEDGMENTS
We are grateful to Rob Naczi of Claude E. Phillips Herbarium, Delaware State
University, and Philip E. Hyatt, U.S. Forest Service, Atlanta, Georgia for their
assistance with this report. Critical reviews by Charles T. Bryson, USDA-ARS
Southern Weed Science Research Unit, Stoneville, Mississippi, and an anony-
mous reviewer improved an earlier draft of this manuscript.
REFERENCES
ArKANSAS Natural HeriTaGe Commission. 2002. Searles prairie natural area management plan.
Arkansas Natural Heritage Commission. Little Rock
ARKANSAS VASCULAR FLORA Commitee, eds. 2002. Draft checklist of the vascular plants of Ar-
kansas. Internal Document 7.0. Little Rock.
Hyatt, PE. 1998. Arkansas Carex (Cyperaceae): a briefly annotated list. Sida 18:535-554.
MouHtensrock, R.H. 1999. The illustrated flora of Illinois. Sedges: Carex. Southern Illinois Uni-
versity Press. Carbondale and Edwardsville.
Naczi, R.F.C. and C.T. Bryson. 2002. Carex section Griseae. In: RW. Ball, K. Gandhi, R.W. Kiger, D
Murray, J.L. Zarucchi, A.A. Reznicek, J.L. Strother, eds. Flora of North America North of
Mexico, vol. 23 Magnoliophyta: Commelinidae: Cyperaceae. Oxford Univ. Press, New
York. Pp. 448-461.
Raprorb, A.E., H.E. AHies, and C.R. Bett. 1964. Manual of the vascular flora of the Carolinas.
University of North Carolina Press, Chapel Hill.
Smith, E. B. 1988. An atlas and annotated list of the vascular plants of Arkansas. Second
Edition. Published by the author. Fayetteville, Arkansas.
STEYERMARK, J.A. 1963. Flora of Missouri.lowa State University Press, Ames, lowa [errata added
at second printing, 1968].
Unitep States DEPARTMENT OF AGricutture. 1977. Soil Survey of Benton County, Arkansas.
1730 BRIT.ORG/SIDA 20(4)
Voss, E.G. 1972. Michigan flora. Part 1.Gymnosperms and monocots. Cranbrook Institute
of Science and University of Michigan Herbarium, Ann Arbor. Kingsport Press.
Yarskievycn, G. 1999. Steyermark’s flora of Missouri- Volume 1- Revised Ed. Missouri Depart-
ment of Conservation, Jefferson City.
Yarskievycn, G., and J. Turner. 1990. Catalogue of the flora of Missouri. Missouri Botanical
Garden Monograph in Systematic Botany No. 37. Braun-Brumfield, Inc., Ann Arbor,
Michigan.
ADDITIONS TO THE FLORA OF CLEVELAND COUNTY,
ARKANSAS: COLLECTIONS FROM MORO BOTTOMS
NATURAL AREA, A STATE-PROTECTED OLD-GROWTH
BOTTOMLAND HARDWOOD FOREST
Danny Skojac' and Margaret S. Devall
U.S. Forest Service, Southern Research Station
Southern Warcdia oods ! ahorator,
PO. Box 227 Stoneville, MS 38776, U.S.A.
Bernard R. Parresol
U.S. Forest Service | Southern Research Station
PO. Box 2680 Asheville, NC 28802, U.S.A.
ABSTRACT
An annotated list of 38 additions to the vascular flora of Cleveland County, Arkansas is presented.
The additions presented were elec rom Moro Bottoms Natura al Area, astate pl otec sted old growl th
bottomland hardwood forest located in the northwest region of the county.
RESUMEN
Se presenta una lista comentada de 38 adiciones a la flora vascular de Cleveland County, Arkansas.
Las adiciones presentadas se recolectaron en Moro Bottoms Natural Area, un bosque pantanoso viejo
protegido por el estado, ubicado en la parte noroeste del municipio.
This paper reports 38 additions to the vascular flora of Cleveland County, Ar-
kansas. Collections were taken from windthrow gaps? areas within Moro Bot-
toms Natural Area, a 40 ha old growth bottomland hardwood forest located in
the northwestern portion of Cleveland County. The area rests on the floodplain
of Moro Creek, a tributary of the Ouachita River, approximately 48 km/30 mi
upstream from the mouth of Moro Creek (Peacock 1983). Moro Bottoms sup-
ports a diverse stream-bottom community dominated by numerous bottom-
land oaks (Quercus spp.), sweetgum (Liquidambar styraciflua L.), bald cypress
(Jaxodium distichum (L.) Richard), and several hickories (Carya spp.). The study
area and most of Cleveland County reside in the west gulf coastal plain physi-
ographic region of Arkansas. Moro Bottoms is an excellent example of a mature
stream-bottom forest becoming old growth through gap regeneration (Devall
et al. 2001). The stand contains many old growth features, including little to no
ineresponcing errs Dann Skojac, email, dsko}j ac@fs.fed.us.
2A severe thunderstorm swept across the Moro Bottoms tract during the summer of 1989, downing trees and
A+}
creating large openings in the forest car lopy COMMOT! ily referred to as windthrow gaps.
SIDA 20(4): 1731-1736. 2003
1732 BRIT.ORG/SIDA 20(4)
evidence of human-induced disturbance and numerous large diameter trees.
Devall et al. 2OO1) reported cherrybark oak (Quercus pagoda Raf.) and
sweetgum to have diameters at breast height up to 149 cm/59 in. Additionally,
asample of larger red oaks and white oaks were determined to be approximately
150 years and 250 years of age, respectively (Guldin et al. 1995). Presently Moro
Bottoms and adjacent areas are owned and protected by the Arkansas Natural
Heritage Commission and the Arkansas Nature Conservancy. The land was
purchased from Georgia Pacific Corporation in 1986.
Collecting of specimens was initiated in the spring of 1990 and continued
seasonally through the fall of 1992. A more detailed description of sampling
methods is discussed in Devall et al. (200). Collecting efforts and identifica-
tions were performed by the authors; some material presented includes collec-
tions made by E. Sundell. Specimens are housed in the herbarium collection at
the U.S. Forest Service, Souther n Hardwoods Laboratory [SHL] (432 Stoneville
Road, Stoneville, MS 38776), with some duplicates and voucher material de-
posited in the University of Arkansas-Monticello Herbarium [UAMI. Smith
(1988) and The PLANTS Database (USDA Natural Resources Conservation Ser-
vice 2002) were consulted to verify the undocumented occurrence of species
within Cleveland County we present.
The following annotated list is arranged alphabetically by family,and then
by genus and species. Generalized species and habitat descriptions, supplemen-
tal to the description of the study area discussed above, cite Radford et al. 1968)
and Godfrey and Wooten (1979 and 1981); range data within the state of Arkan-
sas are taken from Smith’s (1988) Atlas. Collector, number and herbaria of de-
posit are listed last. Accession numbers are given for those specimens housed in
the SHL herbarium.
—
ARISTOLOCHIACEAE
Aristolochia serpentaria L.—Small herb of bot-
tomland forests and woodland margins;
Smith (1988) reports this species for 24 Ar-
kansas counties, mostly in the western half
o the oe it is neweNet listed for —
—
w counties in so
Dddl dana
1B y
eastern Arkansas; Sundell 9136 (UAM).
ASCLEPIADACEAE
Asclepias perennis Walter—A common herb of
alluvial forest, usually in low lying areas on
heavy clays; reported in Smith (1988) for
most eastern counties bordering the Missis-
sippi River and inland into the coastal plain
region of the state; present in neighboring
Bradley, Dallas, zs and Lincoln counties;
Devall s.n. (SHL: 710).
ASTERACEAE
Brickellia eupatorioides (L.) Shinners—Disturbed
woodland sites and thickets; a suprising find
for Cleveland County, listed by Smith (1988)
in western Arkansas counties of quite dif-
ferent physiography than the coastal plain
soils of Cleveland County; Devall s.n. (SHL:
=e
).
Eupatorium hyssopifolium L.—A weedy plant of
woodland borders and openings, probably
seeded in from adjacent upland mixed pine-
hardwood stands where it would be more
common; only reported by Smith (1988) in
our Arkansas counties: Jefferson, Lincoln,
Bradley, and Calhoun; Deval s.n. (SHL: 738).
Mikania scandens (L.) Willd—A perennial climb-
ing vine usually associated with disturbed
areas and margins of stream-bottom forests,
one
SKOJAC ET Al
AREA, ARKANSAS 1733
wet woods; Widespread across southern
Arkansas and up the Mississippi River bor-
counties (Smith 1988); Deval! s.n. (SHL:
18), Sundell 9129 (UAM)
CALLITRICHACEAE
Callitriche heterophylla Pursh—Aquatic herb of
ditches ene — and standing
water; read, occurring in several Ar-
ioe ees iriee neighboring Bra-
dley and Drew counties; Devalls.n. (SHL:933).
CORNACEAE
Cornus stricta Lam.—Small understory tree of
bottomland hardwood forests, usually oc-
curring on heavier textured soils than other
dogwoods found in the bottoms; listed in
Smith (1988) as C. foemina, this species is
found in most Arkansas counties, excluding
those in the Ozarks, also reported for all
counties contiguous to Cleveland County;
Devall s.n. (SHL: 704, 745).
CRASSULA
Penthorum ae L—Rhizomatous herb com-
mon to wet depressional areas, ditches and
slough margins, heavy clay soils; widespread,
found in several Arkansas counties, includ-
ing neighboring Drew, Bradley, Calhoun,
Dallas, Grant, and Jefferson counties (Smith
1988): Devalls.n. (SHL: 703,711), Sundell 9142
(UAM).
EUPHORBIACEAE
Acalypha virginica L—Weedy annual of dis
turbed sites i openings, common; found
in alm ery county in the state (Smith
1988); eee (SHL: 767).
FABACEAE
Desmodium paniculatum (L.) DC. var. dillenit
(Darl.) Istey—Woodland margins and open-
ings; Smith (1988) reports this species for
several Arkansas counties north and west of
Cleveland County, but is reported for neigh
boring Bradley and Drew counties; Sundell
34 (UAM).
Dioclea multiflora (Torrey & A.Gray) Mohr—Her-
baceous vine common in disturbed wood-
land openings, alluvial forests; reported in 15
Arkansas counties by Smith (1988), mostly
in the eastern and southern regions of the
state; Devall s.n. (SHL: 748), Sundel! 9140
(UAM
HYDROPHYLLACEAE
Hydrolea uniflora Raf—Somewhat decumbent
herb of wet ditches, swampy areas, and wet
woods; Smith (1988) reports this species for
several southern and central Arkansas coun-
ties, including neighboring Bradley, Drew,
Calhoun, and Jefferson counties; Devall s.n.
(SHL: 746,747).
HYPERICACE
Hypericum walteri J.G.Gmel.—Perennial herb of
wet floodplain forests, often associated with
cypress swamps and bogs; this species is
reported in 15 Arkansas counties, mostly in
the southern portion of the state; reported
for neighboring Bradley, Dallas, and Jeffer-
son counties (Smith 1988); Sundel! 9144
(UAM)
LAMIACEAE
Lycopus virginicus L.—Stoloniferous, running
herb of bottomland forests, shaded ditch-
banks and wet areas; emit (1988) reports
this i ross
nee state, as well as neighboring Jefferson,
Lincoln, Drew, and Calhoun counties; Deval
5.n. (SHL: 765).
ONAGRA
Ludwigia ek (L.) Ell —Mat forming peren-
nial common to swampy forests, pond Mar-
gins, and exposed shallow depressions,
somewhat invasive; scattered across the
nN numerous counties; reported for
neighboring Bradley and Drew counties
(Smith 1988): Devall s.n. (SHL:709).
POLYGONACEAE
Brunnichia cirrhosa Banks ex Gaertn.—Invasive
woody vine of woodland openings, margins,
and disturbed sites; widespread across the
ties of the Ozarks; Smith (1988) reports this
species for all counties pte a to Cleve-
land County; Devall s.n. (SHL: 754).
RUBIACEAE
Oldenlandia boscii (DC.) Chapm.—Nearly pros-
trate perennial with linear, opposite leaves;
alluvial forests, shallow depressions; very lim-
1734
ited range in Arkansas, only reported for 13
counties in Smith's (1988) Atlas, including
neighboring Bradley, Drew, and Dallas coun-
ties; Devall s.n. (SHL:698).
SAXIFRAGACEAE
tea virginica L.—Deciduous shrub of bottom
land forests, particularly on minor bottom
sites away from the Mississippi floodplain
forests in the eastern part of the state; widely
distributed across the state except in the
upland counties of the northwest and those
counties bordering the Mississippi River
(Smith 1988); Devall s.n.(SHL: 755).
SCROPHULARIACEAE
Lindernia dubia (L.) Pennell—Annual herb found
along ditch edges and muddy flats border
ing waterways, bottomland forests and
Openings; occurring in almost every county
in the state, including neighboring Drew,
Bradley, Calhoun, Dallas, and Jefferson coun
ties (Smith 1988); Devalls.n. (SHL:735), Sundell
9131,9143 (UAM)
SOLANACEAE
a“ sera L.—Weedy annual found in
gs and disturbed sites; Smith's (1988)
ee ee a sporadic distribution across
the state, but comments on its probable
tirren ] IH] Ard 7 |
—
=
for neighboring Bradley county to the south;
Sundell 9138 (UAM)
URTICACEAE
Boehmeria cylindrica (L.) Swartz—Very common
woodland herb of bottomland hardwood
net shaded ditches and woodland-edge
otones; found in nearly every Arkansas
cunt neighboring Jefferson, Lin-
coln, Drew, Bradley, and Dallas counties
6).
(Smith 1 ee eid (SHL: 705, 706
VITACEAE
Vitis cinerea Bat ex, MMet ae os
WOOdY VII f distur and
openings; lees nee pubescent, some
what aggressive in habit, restricted mostly
to floodplain forest settings; Smith (1988)
reports this species in several Arkansas
counties across all physiographic regions of
the state, including neighboring Jefferson,
BRIT.ORG/SIDA 20(4)
Lincoln, Drew, Bradley, and Dallas counties;
Devall s.n. (SHL: 757
Vitis palmata Vahl—Climbing woody vine with
glabrous leaves, common to bottomland
forests and disturbed woodland openings;
distribution in Arkansas appears limited, re
ported by Smith (1988) in 14 counties
eral other counties withou
vouchered material, mostly in ine oe
River floodplain, the tal _and Arkan-
sas River valley counties; Devalls. n. (SHL: 756).
<a
CYPERACEAE
Carex ed Schwein.—Cespitose peren-
nial of floodplain forests and wet meadows
and openings; perigynia somewhat ee
and round, tightly arranged in a compact
=pInE; os ay 3; anes jesticted in
with unvouchered reports for several other
eastern counties (Smith 1988); Devall s.n.
(SHL: 751,752
Carex debilis Michx.—Cespitose sedge of low-
land woods, seepage areas and wet clear-
ings; pistillate spikes loosely flowered (3-4
perigynia lanceolate tapering gradually toa
beak; Smith (1988) reports this species in 13
counties mostly in the coastal plain coun-
ties of southern Arkansas including neigh-
boring Grant, Jefferson, Lincoln, Drew, and
Bradley counties; Devall s.n. (SHL:
729,730,731).
Carex festucacea Schkuhr ex Willd.—Tufted pe-
nnial of bottomland hardwood forests and
wet openings; little documentation of this
species in Arkansas, Smith’s (1988) Atlas re-
ports only 2 counties where vouchered
material has been presented; our record is
among the first reports for this species pres-
ence in the southeastern region of the state;
Devall s.n. (SHL: 732).
Carex intumescens Rudge—Tufted perennial of
alluvial forests, similar to the rhizomatus C.
louistanica; pistillate spikes somewhat g
bose, loosely flowered, perigynia inflated
gradually tapering to a beak; this species is
well distributed across the southern half of
the state, Smith (1988) reports this species
in neighboring Lincoln, Drew, Bradley, and
Dallas counties; Devall s.n. (SHL: 713).
er
=
fev)
>
SKOJAC ET Al AREA, ARKANSAS 1735
Carex reniformis (Bailey) Small—Cespitose pe- kansas, this species is found in 16 counties
rennial mon to alluvial forests and wet but with no discernable pattern depicting
openings; pistillate spikes many flowered, its range in the state. Smith (1988) reports
ovoid, often tightly arranged terminally, this species for Bradley County to the south;
perigynia orbicular, relatively flat, conspicu- Sundell 9145 (UAM).
ously ke reported in Smith's (1988) At- — Erianthus strictus Baldwin—Robust grass asso-
las for 10 Arkansas counties mostly in the ciated with swamp margins and ditches;
nea half of the state including neigh- pikelet pi ly apy !;reported
boring Drew, Bradley, and Calhoun counties; in Smith's (1988) Atlas in several southern
Devalls.n. (SHL: 766). Arkansas counties including Drew, Bradley,
Carex tribuloides Wahlenb.—Abundantly com- and Calhoun counties to the south; Devall
sedge of bottomland hardwood forests s.n. (SHL: 742
and low lying wet areas; pistillate spikes —_ Leersia lenticularis Michx.—An often decumbent
ovoid to obovoid, tightly arranged termi- grass with distinctive lenticular florets ar-
nally, perigynia flat, narrowly ovate, winged ranged loosely in an open panicle; this spe-
apical reported by Smith (1988) in 18 Ar- cies is associated with bottomland hard-
nsas counties mostly in the southern half wood forests and other shaded swampy
of the state including neighboring Jefferson, settings; appears to be undocumented for
Bradley, and Dallas counties. No doubt this several Arkansas counties where it probably
species is present in all coastal plain and al- exists. Smith (1988) reports this plant in 12
luvial regions of the state; Devalls.n. (SHL:739, counties mostly in the southern Mississippi
: River floodplain and coastal plain provinces
Carex typhina Michx.—Cespitose perennial of the state including neighboring Drew an
found in alluvial woodlands; pistillate spikes Bradley counties; Devalls.n. (SHL: 749), Sundell
erect, cylindrical, perigynia ovoid abruptly 9125 (UAM).
tapering to a short slender beak; this spe- _Leersia virginica Willd—Slender, freely branch-
cies is not well documented in Arkansas, re- ing perennial grass common to alluvial
ported by Smith (1988) in only 8 counties woodlands and swamps; this species is re-
widely dispersed across the state; Devall s.n. ported by Smith (1988) in several Arkansas
(SHL: 762, 763). counties, primarily in the northern half of the
state, but is also documented in a few south-
ern counties as well, including neighboring
Drew and Bradley counties; Sundell 9124
(UAM).
Panicum dichotomum L.—Tufted perennial
grass common to bottomland forests and
wet clearings; Smith (1988) iene this sae
essentially throughout the
mented in neighboring een cs
Drew, Bradley, Dallas, and Grant counties; no
JUNCACEAE
Juncus coriaceus Mack.—Clumping, cespitose
rush with inflorescence oo lateral,
mon to swampy woods and depres-
sions; this rush has a ae limited ra ce in
Arkansas, reported by Smith (1988) in
counties all situated within the Ou ae
mountains and coastal plain physiographic
provinces of the state; Devall s.n. (SHL: 708).
POACEAE doubt this species is present in Cleveland
Agrostis perennans (Walter) Tuck— Tufted peren- County; Deva Ws. n. 1 (SHL: 71 5).
nial grass of alluvial forests and wet clear- — p \bundantly
ings; Smith (1988) reports this species in sev- grass of neodelen forests snd wet depies:
eral counties dispersed across the state in- sions; this species has a wide distribution,
cluding neighboring Jefferson and Bradley occurring in all physiographic regions of the
counties; Sundell 9126 (UAM). state, reported in Smith's (1988) Atlas for
Cinna arundinacea L.—Rhizomatous perennial neighboring Jefferson, Lincoln, Drew, Brad-
grass found in floodplain forests and on wet ley, and Grant counties; Devall s.n. (SHL: 770,
ditch banks; sporadic distribution across Ar- 771)
1736 BRIT.ORG/SIDA 20(4)
ACKNOWLEDGMENTS
The authors wish to thank the Arkansas Natural Heritage Commission and the
Arkansas Field Office of the Nature Conservancy for allowing us to collect speci-
mens and conduct research on the preserve. We express our appreciation to those
who helped with plant identification, including Eric Sundell, Seth Barnes,
Michael Lelong, Allen Moore, and Charles Bryson.
—"
REFERENCES
Devatt, M.S., BR. Parresot, and W.P. SmitH. 2001. The effect of herbivory by white-tailed deer
and additionally swamp rabbits in an old-growth bottomland hardwood forest. In:
Hamel, P.B.and L. Thomas, tech. eds. Bottomland hardwoods of the Mississippi Alluvial
Valley: characteristics and management of natural function, structure, and composi-
tion. 1995 October 28. Fayetteville, AR. Gen. Tech. Rep. SRS-42. USDA Forest Service,
Southern Research Station, Asheville, North Carolina. Pp. 49—64.
Goprrey, R.K. and J.W. Wooten. 1979. Aquatic and wetland plants of southeastern United
States: M otyledons. The University of Georgia Press, Athens.
Goprrey, R.K. and J.W. Wooten. 1981. Aquatic and wetland plants of southeastern United
States: Dicotyledons. The University of Georgia Press, Athens.
Gutoin, J.M., B.R. LockHart, and L. Peacock. 1995. Ecological character of a small-scale linear
wind event in an old-growth bottomland hardwood stand in south-central Arkansas.
http://ces.lisc.ernet.in/hpg/envis/proceed/lockhart.txt.htm (Accessed 5/20/02).
Peacock, L. 1983. Site preserve summary-Moro Bottoms. Arkansas Nature Conservancy,
unpublished report. Arkansas Field Office of The Nature Conservancy, Little Rock.
Rabrord, A.E., H. AHLES, and C.R. Bet. 1968. Manual of the vascular flora of the Carolinas. The
University of North Carolina Press, Chapel Hill, North Carolina.
Smith, E.B. 1988. An atlas and annotated list of the vascular plants of Arkansas, 2" ed.
Student Union Bookstore, University of Arkansas, Fayetteville.
United States DEPARTMENT OF AGRICULTURE, NATURAL RESOURCES CONSERVATION Service. 2002. The PLANTS
database, version 3.5.(http://plants.usda.gov). National Plant Data Center, Baton Rouge,
Louisiana. (Accessed 8/1/02).
ARKANSAS FLORA: ADDITIONS, REINSTATEMENTS,
EXCLUSIONS, AND RE-EXCLUSIONS
James H. Peck
Herbarium LRU
Depar CFTICHIL of Biology
University of Arkansas at Little Rock
Little Rock, AR 72204, U.S.A.
jhpeck@ualr.edu
ABSTRACT
Nine species are reported as new to the flora of Arkansas: Agalinis nuttallii Shinners
(Scrophulariaceae), Azolla caroliniana Willd. (Azollaceae), Crataegus thermopegaea Palmer (Ro-
), Cyrtomium falcatum (L.1.) K. Presl (Dryopteridaceae), Nerium oleander L. (Apocynaceae),
Polygala alba Nutt. (Polygalaceae), Punica granatum L. (Punicaceae), Ricinus communis L. (Euphor-
biaceae), and Schinus terebinthifolius Raddi (Anacardiaceae). They are a mixture of extinct,
overlooked, or urban exotic-escaped species, some of which are of special concern as they pose new
problems for urban poison control workers. Literature and herbarium work uncovered that 12
Arkansas species previously excluded actually have kn nown and valid vouchers, and they are re-in-
stated: eee Ces (L.) Willd. Mimosaceae), A 1(L.) Bernh. (Brassicaceae), Canavalia
Fabaceae), Cladium jamaicense Crantz eee Eragrostis trichodes (Nutt.)
ensiformis (L.)
Wood eres ee diptera Ellis (Styracaceae), Lygodesmia juncea (Pursh) D. Don (Asteraceae),
Prosopis glandulosa Torr. var. glandulosa (Mimosaceae), Rhynchospora colorata (L.) Pfeiffer
peraceae), Rosa canina L. (Rosaceae), Sagina procumbens L. (Caryophyllaceae), and Trifolium
stoloniferum Muhl. (Fabaceae). Another 13 species previously excluded and/or re-excluded from
ie were recently and i aaeaee! re-instated, requiring re-exclusion. Another 76 species are
here excluded, being rer sly as being from Arkansas, based either on literature reports
ona variety of clerical errors. These 21 additions and 89
that lack s voucher specimens or
subtractions to ne Ar Kansas [ as point to problems in achieving a uniform basis of ae iding which
1 ae |
VALE vouchers.
RESUMEN
A rR : Pees pi eas cad es ] ]
Se citan nueve especies nuevas para la flora de Arkansas:
ceae), Azollacaroliniana Willd. (Azollaceae), Crataegus thermopegaea Paloven Rosaceae) Cyaan
falcatum (Lf) K. Presl (Dryopteridaceae), Nerium oleander L. (Apocynaceae), Polygala alba Nutt.
uphorbiaceae), y Schinus
(Polygalaceae), Punica granatum L. (Punicaceae), Ricinus communis L.
terebinthifolius Raddi (Anacardiaceae). Son una mezcla de extintas, oe por alto, o especies
as exdticas escapaCas, alee os las c uales’s son de paar interés ya ls causan nuevos
Conel d
urban
1
TCDA
problemasa | J : : }
lI esneciescde Arl 1] laid ] ]
ji
stauradas: Acaci harm >siana (L.) Willd. (Mimosaceae), Arabis gl aba Cs nnn
L:
y validos S,Y son re
aBrassicaceae)) Canavalia ensiformis (L.) DC. (Fabaceae), Cladium jamaicense Crantz (Cyperaceae),
Eragrostis t les (Nutt.) Wood (Poaceae), Halesia d ipterd Ellis (Styracaceae), a ee
(Pursh) D. Don ee Prosopis glandulosa Torr. var. gla a sMimpsaCeae) ee ae
colorata(L.) Pfeiffer (Cyperaceae), Rosa canina L. (Rosaceae), Sagina } cE
y Trifolium an . (Fabaceae). Otras 13 especi luidas previ y/o re-excluidas
SIDA 20(4): 1737-1757. 2003
1738 BRIT.ORG/SIDA 20(4)
de Arkansas fueron reciente y erroneamente re-instauradas, y necesitan una re-exclusion. Otras 76
oe nae se exciyen aqui, por hi aberse clade erroneamente de Arkansas, basadas tanto en citas
g | plieg g varios errores administrativos, Estas 2] adiciones y
89 sul la flora de Arl los problemas de poner a punto una base uniforme
lf
oe a decidir que planté is incluir en una flora, y que constituye suficiente documentacion para los
validos.
plies 8
The preparation of a state flora remains the grandest state scientific enterprise
in most states. More than 1200 articles by nearly 100 botanists documented and
explained the diversity, abundance, and distribution of its flora and vegetation
(Peck & Peck 1988; Peck et al. 2000). Vascular plant floristics in Arkansas com-
menced with the 1819 exploratory travels, collections, and published list by
Thomas Nuttall 1835). Earlier expeditions commented on plants, but appeared
to either not have vouchers or none have survived from Arkansas locations
(Palmer 1927, Morton 1967, MacRoberts et al. 1997). Subsequent flora lists were
prepared by Lesquereux (1860) of 1320 species and by Branner and Coville (1891)
of 1534 species, but were developed from limited field work and were poorly
supported with extant vouchers. Buchholz and Palmer (1926) attempted to
verily, locate vouchers, or re-collect many old Arkansas records, and in the pro-
cess, added 534 species for a total of 2068. Demaree (1943) compiled all past
lists, personally added 694 species, and presented a total list of 2723 species. As
stated by Demaree (1943), many species on his list were not his records, but
were compiled at face value from prior lists. Demaree (1943) knew some were
of dubious identity, probably lacked vouchers, and contained records from
Oklahoma when it was part of the “Arkansas Territory” from 1819-1836. But
there was a war going on, no resources could be spared to validate the list, and
the fate of species-level biology in Arkansas was in doubt.
Edwin B. Smith inherited the Arkansas state flora enterprise. Through near
reroic efforts, being the last to attempt a “one man band” approach to a state
flora, he produced two annotated checklists with county maps. Smith (1978)
provided an annotated checklist with county dot maps for 2338 species, a list
of 200 possible additions, a list of 365 excluded species (many old reports ap-
peared to lack vouchers of any kind), and state maps with 40,941 county dots.
Smith (1988) presented a revised list and maps for 2469 species, 324 possible
additions, 375 excluded species, and maps with 49,510 county dots.
In 1998, a consortium of 16 botanists formed the Arkansas Vascular Flora
Committee to prepare a checklist, database, atlas, and manual. To date, the Ar-
kansas Vascular Flora Committee has located and verified vouchers for over
3350 taxa. Based upon the immensely helpful national electronic lists (Kartesz
1998, Kartesz & Meacham 1999; NRCS 1999), near monographic state floras from
adjacent Missouri and Texas, the atlas and manual of the Flora of the Great
Plains, checklists and atlases of Louisiana, Tennessee, and Texas, and the serial
a
—
—
PECK, ADDITIONS AND NOTES ON THE ARKANSAS FLORA 1739
volumes of the Flora North America project (Flora North American Editorial
Committee 1993+), another 80 taxa might yet be added.
Through herbarium search and review of literature and national electronic
floras, 110 noteworthy Arkansas species were identified that required addition
(9), reinstatement (12), re-exclusion (13), or exclusion (76). The nine additions
included overlooked but long-standing elements of the Arkansas flora and re-
cent escapes with properties of ecological concern (invasive) or human con-
cern (toxic). Twelve re-instated species included previously excluded, extinct,
or extirpated species that required re-instatement as their vouchers were finally
located, were replaced, or were erroneously excluded [criteria and judgment
differs with differing workers]. Thirteen species were re-excluded that were er-
roneously reported and re-reported as present in Arkansas. Another 76 species
were excluded as they lack a valid or known voucher to suppor their inclu-
sion. With this update, the listed flora of Arkansas is treated m
with that of floras of other states in the region and with national lists.
The purpose of this current report, beyond communication of state and
county noteworthy records, was to demonstrate the need for continued judg-
ment and interpretation of floristic information, whether in the older hard-
copy literature or in the newer data-based, on-line records. Records of each state
need to be reviewed by experts in taxonomic groups as well as local experts to
insure a correct name and proper documentation exists. Criteria for inclusion
need to be carefully followed regarding invasive, escaped, waif, and natural-
ized plants. Equally important is to exclude species erroneously included in
past lists. There may well be more exclusions than new additions (Ertter 2000).
Documentation of known and valid vouchers as to herbarium of deposit, iden-
tity, collector, and number should be the basis for inclusion, not just a passage
in the text, or a tone-map or dot-map. Literature errors may need to be removed
more than once from the database. Increased efforts at quality control and as-
surance through time and effort will result in a continually improved and more
powerful tool at both the state and national levels (Kartesz 2000).
ADDITIONS TO ARKANSAS FLORA
Agalinis nuttallii Shinners (Scrophulariaceae)—This is the “first report” for Ar-
kansas of this historical plant, now considered extinct. First collected and named
by Nuttall based on his work in Arkansas (Nuttall 1835, Palmer 1927) as Gerar-
dia longifolia Nutt. Although similar to G. holamantha (Pennell) Pennell, Pennell
(1935) asserted that it was distinct. The lack of conservation of the name Ger-
ardia required a new combination that was made by Shinners (1962) in honor
of its first collector. The species was listed for Arkansas by Lesquereux (1860).
Surprisingly, the species was not listed, discussed, or excluded by Smith (1978,
1988), but was listed as an Arkansas endemic by Kartesz and Meacham (1999).
1740 BRIT.ORG/SIDA 20(4)
Voucher: ARKANSAS: along the bank of the Arkansas, locality not specified, Nuttall s.n. (BM, K, PH).
Azolla caroliniana Willd. (Azollaceae)—The first collection of this species from
Arkansas was from a swamp at Spruell’s eee Area, 2 mi north of
Doddridge along US 71 on 19 May 1977 by B.L. Lipscomb. All other material of
this genus from Arkansas was referable to A. mexicana Schlecht. & Cahm. ex K.
Presl.
Voucher: ARKANSAS: Miller Co.: Lipscomb 2064 (LRU).
Crataegus thermopegaea Palmer (Rosaceae )—This species is reported based on
a collection by EJ. Palmer at Hot Springs, Arkansas in Apr and Oct 1925. It was
described asa new species by Palmer (1926). Palmer described 67 species as new
for Cratageus; this is the only one still recognized on national lists. The species
was not recognized by Tucker (1976) or Smith (1978, 1988), but was recognized
as an Arkansas endemic by Kartesz and Meacham (1999).
Vouchers: ARKANSAS: Garland Co.: Palmer 26843 (A), Palmer 26843 (G).
Cyrtomium falcatum (L.f.) K. Pres] (Dryopteridaceae)—This is an escaped ex-
otic species that can persist several years in Arkansas when its rhizome is well
protected; plants remain wintergreen in southern to central Arkansas. The spe-
cies is cultivated state-wide, escaping rarely, but persisting in urban waste ar-
eas, at least in Little Rock. A related species Cyrtomium fortuneiiJ.Sm.(Fortune’s
Net-vein Holly Fern) was collected in 1988 by Wilson Baker as part of a con-
tract with The Nature Conservancy to study plants in the batture lands of Desha
Co., Arkansas. Unfortunately, the voucher material was taken to Tall Timbers
Research Station, Georgia, and remains unmounted and unreported. Therefore,
the later collection from Little Rock was the first report of the genus from
Arkansas.
Voucher: ARKANSAS: Pulaski Co.: Peck 02744 (LRU).
Nerium oleander L. (Apocynaceae)—This is a very toxic horticultural shrub,
escaping from landscapes to urban waste places in Boyle Park, Little Rock, evi-
dently at least “root-hardy” and persisting in Arkansas.
Voucher: ARKANSAS: Pulaski Co.: Peck 02-845 (LRU)
Polygala alba Nutt. (Polygalaceae)—This species was first collected in Arkansas
on 21 May 1977 from a wooded hillside 12 mi north of Murfreesboro in Pike Co.
by B.L. Lipscomb,
Voucher: ARKANSAS: Pike Co.: Lipscomb 2177 (LRU).
Punica granatum L. (Punicaceae)—This horticultural shrub has persisted at old
homesteads in Arkansas and has escaped to urban waste places and to Boyle
Park, Little Rock, where it has persisted for decades.
PECK, ADDITIONS AND NOTES ON THE ARKANSAS FLORA 1741
Vouchers: ARKANSAS: Little River Co.: Tucker 16121 (APCR), Pulaski Co.: Luyet 8311 (LRU).
Ricinus communis L. (Euphorbiaceae)—A very toxic horticultural, suffrutescent
plant or semi-perennial herb, escaping from cultivation and establishing small
colonies in urban and rural waste or disturbed areas, now state-wide in Arkansas.
Vouchers: ARKANSAS: Bradley Co.: Leslie 1515 (UAM). aaa Co.: ee ai CL — Stone Co.: Le
onard 124 (LRU), and by the author and c wie at LRU in 22 cow 02369, Baxter
98117 co burne Co.: 96185, Faulkner Co.: 97321, Garland Co.: 97268, Grant Co.: obiaa Hot Spring
Co.: 072321 pend Co.: 96344, Izard Co.: 96127, Jackson Co.: 99176, Jefferson Co.: 02376, Lonoke
Co.: 02276, Marion Co.: 96432, Monroe Co.: 96518, Phillips Co.: 02333, Prairie Co.: 02312, Saline Co.:
02350, Searcy Co.: 98253, Van Buren Co.: 98278, White Co.: 99561, Woodruff Co.: 97566, and Yell Co.:
02467.
Schinus terebinthifolius Raddi (Anacardiaceae)—A toxic, invasive woody shrub-
tree, was introduced intoa backyard in Camden in southern Arkansas asa hor-
ticultural novelty. The tree with plentiful red berries was a Christmas tree
“present” from Florida. It was planted and had survived 3 winters, with root
suckers and seedlings produced around the original stock. The plants were eradi-
cated by the homeowner upon advice of the author before it became further
established or irreparably naturalized.
Voucher: ARKANSAS: Ouachita Co.: Peck 90117 (LRU).
REINSTATEMENTS TO ARKANSAS FLORA
Acacia farnesiana (L.) Willd. (Mimosaceae)—This species was first reported for
Arkansas by Demaree (1943). Tucker (1976) included this species, but consid-
ered it to be a “rare adventive.” Smith (1978, 1988) excluded it from the state
flora, as the specimen was sterile material [not unusual for woody plant collec-
tions], and he questioned the correctness of its identity. The voucher specimens
are of this species, collected from a 3-4 m tall tree adjacent to the Arkansas
River near Little Rock, but no longer present.
Voucher: ARKANSAS: Pulaski Co.: Palmer 29600 (A, UARK).
Arabis glabra (1.) Bernh. (Brassicaceae )—This species was first reported for Ar-
kansas by Branner and Coville (1891), compiled by Demaree (1943), and reported
by Fernald (1950). Smith (1978) suggested that this record might be a
misidentified Arabis laevigata (Muhl. ex Willd.) Poir, but he did not consult
the voucher. Al-Shehbaz (1988) noted that M. Hopkins (1937) reported this spe-
cies from Pulaski Co., AR. Smith (1988) excluded it again, but noted that “as it
has not been recollected in the last 50 years [1895], I assume it is no longer ex-
tant in the state.”
Voucher: ARKANSAS: Pulaski Co.: Hasse s.n. (NY).
Canavalia ensiformis (L.) DC. (Fabaceae)—Sauer (1964) reported this prehistoric
American Indian domesticate for Arkansas based on a collection by Harvey in
1742 BRIT.ORG/SIDA 20(4)
1885 and by Jennings in 1938. All other specimens, particularly at UARK, are
based on cultivated materials dating from 1930 tol959. The species was listed
for Arkansas by Kartesz and Meacham (1999), citing Sauer (1964).
Vouchers: ARKANSAS: locality not specified, Harvey s.n. (GH), Jennings s.n. (F).
Cladium jamaicense Crantz (Cyperaceae)—The species was reported first by
Branner and Coville(891). Demaree (1943) reported this species based on that
report and on his 1939 collection in Bradley Co., AR (Demaree 19433 BRIT, MO).
Without examining the vouchers, Smith (1978) listed it as “excluded” from Ar-
kansas, based on its reported range in the floras of Texas and the Carolinas.
Orzell and Bridges (1987) examined and verified the Demaree collections, but
considered it to be probably extirpated from the state. Smith (1988) listed it as
“excluded” in that it was only a waif and now extirpated from Arkansas. The
species was recently collected in four Arkansas counties.
—
Vouchers: ARKANSAS: Bradley Co.: Peck 98247 (LRU). Calhoun Co.: Peck 20220 (LRU). Ouachita
Co.: Peck 20240(LRU). Union Co.: Peck 20234 (LRU).
Eragrostis trichodes (Nutt.) Wood (Poaceae)—The species was reported from
Arkansas by Moore (1961) from two counties (Logan and Pulaski), listed by
Gould (1975) from southwestern Arkansas, but excluded from the flora by Smith
(1988), as one oT (Moore 470609 UAM) was a misidentified sheet of E.
hirsuta. The 2" specimen cited by Moore (1961) from Pulaski Co. is not at UARK.
Kartesz and Meacham (1999) listed this species as present in Arkansas citing a
Nuttall voucher at PH.
—_—
Vaiict - ARPWANCAGC
without I
locality data, Nuttall s.n.(PH): Pulaski Co.: Merrill 14 (UARK?).
Halesia diptera Ellis (Styracaceae)—The species was first reported for Arkan-
sas by Branner and Coville (1891), compiled by Demaree (1943), and reported
by Tucker (1976). Reveal and Seldin (1976) reviewed the taxonomy of this spe-
cies and its varieties. Smith (1978) included this species in his first checklist,
but he excluded it from the second edition (Smith, 1988), noting that it was col-
lected only once in 1887 by Letterman, that “it has not been re-collected in the
last century, land] it is probably no longer extant in the state.” R. G. Erickson
(1989) discovered a second population in Lafayette Co., Arkansas. The speci-
men was sent to APCR, but was apparently lost before being mounted. More
recently, a third population was located in Union Co. by R. Dale Thomas.
Vouchers: ARKANSAS: Nevada Co.: Letterman s.n. ILL), Union Co.: Thomas 109,060 (NLU).
Lygodesmia juncea (Pursh) D. Don (Asteraceae)—This species was first reported
for Arkansas by Buchholz and Palmer (1926). The first vouchered collection was
by Demaree (1943). Vuilleumier (1973) reported this species for Arkansas based
on Demaree’s 1937 collections. Tomb (1980) re-reported the species from Arkansas,
PECK, ADDITIONS AND NOTES ON THE ARKANSAS FLORA 1743
noting the locality as Magnet Cove, AR. Without examining vouchers or seeing
the plant in the field, Smith (1978, 1988) excluded this species as being “only a
waif,” and therefore not part of the “normal Arkansas flora.” Orzell and Bridges
(1987) reported the status of this species in Arkansas as being “probably extir-
pated.” Kartesz and Meacham (1999) reported this species as present in Arkan-
sas, but cited Smith (1988).
Vouchers: ARKANSAS: Hot Spring Co.: Demaree 16264 (BRIT, NY).
Prosopis glandulosa Torr. var. glandulosa (Mimosaceae)—Tucker (1976) reported
this species based on collections by D. M. Moore in 1954 and 1955 in Pulaski Co.,
noting that it was a “rare adventive” of potentially long duration that was col-
lected along the railroad tracks on the southern edge of Little Rock, apparently
brought in with livestock. Smith (1978, 1988) excluded ee species, consider-
ing it to be only a waif, and not “part of the normal flora.’
Vouchers: ARKANSAS: Pulaski Co.: Moore 54343, 55517 (UARK).
Rhynchospora colorata (L.) Pfeiffer (Cyperaceae)—First reported for Arkansas
by Demaree (1943) based on his collections at Warren Prairie, Bradley Co., Ar-
kansas. Smith (1978) listed this species for Bradley Co. and noted a potential
report from Pope Co., Arkansas. Orzell and Bridges (1987) reported “a stand of
thousands of plants from I _ittle River County,” but opined that the Bradley Co.
population was extirpated. Smith (1988) reported this species for Bradley Co.
and Little River Co., but noted that the Pope Co. record was erroneous, based on
diseased material of a Cyperus sedge. Similarly diseased sedge plants occur in
Bradley Co. eee the species was collected in another 10 counties.
Vouchers: ARKANSAS: Ashley Co.: ee 76 by RU). cpap! Co.: Demaree 15044 (F MO, BRIT, NY);
Demaree 19264 (GH, MO, gen BRIT, UARK); BRIT); Demaree 19269 (MICH, NLU, NY,
TEX, UARK); Logan 101 (UCAC). Galliois un Co.: Mar ian 22(L RU). Columbia Co.: Smith 32 (LRU). Drew
Co.: Smith 32 (LRU). cael Co.: Sontag 108 (LRU). Lafayette Co.: Smith 76 (LRU). Little River
Co.: Orzell 2726 (APCR, MO, NLU, BRIT, TEX, UARK). Miller Co.: Sontag 63 (LRU). Ouachita Co.: Be |]
99 (LRU). Union Co.: Peck 20235 i
Rosa canina L. (Rosaceae )—Smith (1978, 1988) excluded this species, based ona
Franklin Co. collection, as it “probably represents a local escape or was persis-
tent at an old home site; it should not be considered as a normal part of our
flora.” Yet, many species have entered the Arkansas flora by this method of in-
troduction. Kartesz and Meacham (1999) listed it as “present,” based on Smith
Voucher: ARKANSAS: Franklin Co.: Davis 449 (UARK)
Sagina procumbens L. (Caryophyllaceae)—First reported by Branner and Coville
(1891), and then listed by Demaree (1943). Without inspecting vouchers, Smith
1744 BRIT.ORG/SIDA 20(4)
(1978) excluded this species, suggesting that it wasa misidentified S.decumbens
(EL) Torr. and A. Gray. Crow (1978) noted that it was correctly named and was
collected in Pulaski Co. in May 1886. Smith (1988) re-excluded this species, not-
ing that it was “not re-collected in the intervening century.”
Voucher: ARKANSAS: Pulaski Co.: Hasse s.n.(MONTU).
Trifolium stoloniferum Muhl. (Fabaceae)—Smith (1978) did not list this species,
but noted it asa “possible addition” to the flora of Arkansas. Brooks (1983a, 1983b;
Batgie, 1985) reported that it had been collected from Arkansas in 1896. Smith
(1988) excluded it from the flora of Arkansas as it was “no longer extant—and |
am therefore excluding it.” This species has declined across its entire range since
settlement.
Voucher: ARKANSAS: Independence Co.: [:ggert s.n. (MO).
RE-EXCLUDED FROM ARKANSAS FLORA
Asclepias sullivantii Engelm. ex A. Gray (Asclepiadaceae)—First reported in
Arkansas by Buchholz and Palmer (1926). This report was based ona collection
from the southwestern corner of the University of Arkansas campus at Fay-
etteville, Washington Co. According to the card file by D. Moore kept at UARK,
EJ. Palmer observed it on 12 Jun 1923, and it was collected by Buchholz on 13
Jun 1923 (Buchholz 20145-6). The voucher is not at UARK or MO, but it might
be at the Arnold Arboretum. A second report was made erroneously by Smith
(1978, 1988) from Jackson Co., Arkansas, based on a misidentified collection by
D. Moore (Moore 440164 UARK). Another specimen of this species was collected
by EJ. Palmer from Jackson Co., Missouri, deposited at MO, and reported by
Woodson (1954). Woodson (1954) did not map this species in Arkansas. The
Palmer specimen label was typed on a “Flora of Arkansas” label, with the cor-
rected state name of Missouri hand written above the crossed-out “Arkansas.”
Asplenium ruta-muraria L. (Aspleniaceae)—Reported by Nuttall (1835) and
uncritically re-reported a dozen times from Arkansas for more than 165 years,
ut a voucher specimen has never been located at BM, K, P, or in any Arkansas
herbarium. This species was excluded many times from the flora of Arkansas
(Smith 1978, Taylor 1984, Smith 1988, Peck and Taylor 1995). Kartesz and
Meacham (1999) erroneously listed the species as present in Arkansas citing
Smith (1988).
Burmannia biflora |. (Burmanniaceae)—Reported by Demaree (1943, p. 21) as
his record, but without any comment. The species was not listed nor excluded
by Smith (1978, 1988). A specimen from Cass Co., Texas, adjacent to Texarkana,
a
Miller Co, Arkansas, exists at BRIT (per. comm., G. Diggs). The species occurs
nearby in Louisiana as well.
Dryopteris xneowherryi W.H. Wagner [D. goldiand » marginalis] (Dryopterida-
PECK, ADDITIONS AND NOTES ON THE ARKANSAS FLORA 1745
ceae)—Montgomery (1981) erroneously reported this hybrid from Arkansas in
a summary of Dryopteris hybrid morphology, systematics, and phytogeogra-
phy. Montgomery confused this hybrid with D. xleedsii which does occur in
Arkansas. Peck and Taylor (1995) and Peck (2000) specifically excluded this hy-
brid from Arkansas. Kartesz and Meacham (1999) erroneously listed this spe-
cies as present in Arkansas citing Montgomery (1981).
Escobaria missouriense (Sweet) D.R. Hunt var. similis (Englem.) N.P. Taylor
(Cactaceae)—Benson (1982) reported Coryphantha missouriense (Sweet) Britton
and Rose var. caespitosa (Engelmann) Benson from Miller Co., AR, based on a
specimen with only an indeterminent location of “Arkansas” [Dyck s.n., 1857
(MO)|. Zimmerman (1985) referred to this taxa as C. missouriensis (Sweet) Britton
and Rose var. similis (Englemann) A. Zimmerman and mapped it in Texas and
Oklahoma, but not in Arkansas. Orzell and Bridges (1987) excluded this find
by Benson (1982), considering it to be a Oklahoma record based on confusion
over the concept of Arkansas Territory. The “Arkansas Territory” (1819 - 1836)
also included what is modern Oklahoma. Smith(1988) agreed and cited Orzell
and Bridges (1987). However, to accept this interpretation, one must assume that
Arkansas had been a state for 21 years at the time the specimen was collected
and that the collector did not know this. Kartesz and Meacham (1999) errone-
ously listed the species as present in Arkansas, but cited Smith (1988) who ex-
cluded it.
Huperzia xbartleyi (Cusick) Kartesz & Gandhi [H. lucidula = porophila]
(Lycopodiaceae)—Smith (1988) reported this as a “possible addition” to the flora
of Arkansas, meaning that it might be found in the state in the indefinite fu-
ture, but that it had not as of yet been so documented. Peck and Taylor (1995)
specifically excluded this hybrid from the flora of Arkansas, noting that only
H. lucida Michx. was known in Arkansas. Kartesz and Meacham (1999) errone-
ously listed this hybrid for Arkansas citing Smith (988).
Krameria lanceolata Torr. (Krameriaceae)—This species was reported from Ar-
kansas by Nuttall (1835), listed by Lesquereux (1860), and again by Branner and
Coville (891). Demaree (1943) compiled the name into his list, but a voucher
has not been located. Smith (1978, 1988) excluded this species. Kartesz and
Meacham (1999) erroneously listed the species as present in Arkansas, citing
Demaree (1943). The origin of this record may be a Nuttall specimen from Okla-
homa. Alternatively, there is a specimen collected in 1836[C.W. Short s.n. MOJat
the “Arkansas [River] at Leavenworth,” a location within the Arkansas Terri-
tory, but now in present-day Oklahoma.
Lesquerella angustifolia (Torr. & A. Gray) Wats. (Brassicaceae)—This species was
collected by Nuttall (1835) from “prairies near the Red River, Arkansas.” Re-re-
ported by Branner and Coville (1891), Small (1913), and listed by Demaree (1943).
The collection location was actually in McCurtain Co., Oklahoma, then part of
Arkansas Territory (Goodman, 1966). Rollins and Shaw (1973) excluded it. Smith
1746 BRIT.ORG/SIDA 20(4
(1978) listed this species as a “possible addition” to the state flora. Smith (1988)
excluded this species as it was never documented from present day Arkansas.
The species was erroneously re-listed for Arkansas on some national e-lists.
Lesquerella gracilis (Hook.) Wats. var. repanda (Nutt.) Pays. (Brassicaceae)—This
species was collected by Nuttall (1835) from “prairies near the Red River, Ar-
kansas.” Re-reported by Branner and Coville (1891), Small (1913), and listed by
Demaree (1943). The location was actually in McCurtain Co., Oklahoma, then
partof Arkansas Territory (Goodman, 1966). Rollins and Shaw (1973) and Smith
(1978, 1988) excluded this species from the flora of Arkansas. This species was
erroneously re-listed for Arkansas on some national e-lists.
Lonicera albiflora Torr. & A. Gray (Caprifoliaceae)—This species was reported
from Arkansas by Small (1913). Ferguson (1966a) noted that the type locality
was “Fort Towson, Arkansas.” Fort Towson is in SE Oklahoma, another confu-
sion regarding Arkansas Territory which included present day Oklahoma. This
species was excluded by Smith (1978, 1988). Kartesz and Meacham (1999) erro-
neously re-listed the species as present in Arkansas, citing Smith (1988).
Rhynchosia minima (L.) DC. (Fabaceae)—This species was reported for Arkan-
sas by Greer (1978) based on a specimen collected in the “Arkansas Refuge [White
River?],” ostensibly in Phillips or Arkansas counties, but with a dot that was
placed in Desha Co. The specimen (Cory 45847 FSU) was actually from Aransas
Co., Texas, Aransas National Wildlife Refuge. Smith (1988) excluded this spe-
cies from Arkansas. Kartesz and Meacham (1999) erroneously re-listed this spe-
cies as present in Arkansas, citing Smith (1988).
Rivina humilus L. (Phytolaccaceae)—This species was collected by Nuttall at
“Red River, Arkansas” (voucher at NY), in present day Oklahoma, then Arkan-
sas Territory. The species was reported by Branner and Coville (1891). Demaree
(1943), who had never seen it in Arkansas, commented that this species must
be “probably rare in the state.” Smith (1978, 1988) did not list this species, and
he only considered it as a “possible addition.” Kartesz and Meacham (1999) er-
roneously re-listed the species as present in Arkansas, citing Smith (1988). Some
confusion may have resulted from uncorrected specimen labels on duplicates,
such as an EJ. Palmer specimen at MO (E,]. Palmer #30520), collected on 7 June
1926 from Brewster Co.,, Texas, typed on a “Flora of Arkansas” label blank with
the correct state typed above the strike-over of “Arkansas.”
Smilax auriculata Walt.(Smilacaceae)—This species was reported for Arkansas
by Duncan (1967). Tucker (1976) noted it was an enigmatic, 500 mi range ex-
tension reported by Duncan (1967), but a voucher was not located. Smith (1978,
1988) mapped it in his Arkansas atlas without seeing a voucher, referring to
Duncan (1967). Smith (1994) listed the species as a “possible addition” denoted
by an asterisk. Kartesz and Meacham (1999) erroneously listed the species as
present in Arkansas, citing Smith (1988). No Arkansas specimen vouchered this
record; it was mismapped by Duncan (1967). Moore (1941) reported that S.
—
—
—_—
—
PECK, ADDITIONS AND NOTES ON THE ARKANSAS FLORA 1747
walteri was present in Arkansas near Hot Springs, Garland Co. It was also known
from Hempstead Co. at this time (Buchholz 395 UARK). Coker (1944) re-reported
the presence of S. walteri in “swamps,” in Garland Co., Arkansas. The dot for
Garland Co. that belonged on the S. walteri map, was absent in Duncan (1967).
A clerical error was made: the dot for S. walteri was incorrectly mapped onto
the S.auriculata map and placed incorrectly in the adjacent Montgomery Co.
EXCLUSIONS FROM ARKANSAS FLORA
Agrostis capillaris L. (Poaceae)
species as a“possible addition ee that all
arlier records were based on cultiv -
terial (turf), not on escaped or eine!
plants Beyer euileivaciony Kartesz and
Meacham (19 neously listed the spe-
cies as “present,’ ‘citing Smith foe
ee gigantea (Walt.) Muhl. subsp.
a (Walt.) McClure (Poaceae)—Branner
oh COvIIE (1891 ) listed this variety for Ar-
t cit her. Efforts since
see ihee and paline’ ( 926) have failed to
locate a known and valid voucher. Kartesz
and Meacham (1999) listed this variety for
Arkansas, citing the compiled list of
Demaree (1943).
Asclepias rubra L. (Asclepidaceae)—This spe-
cies was listed by Kartesz and Meacham
(1999) for Arkansas, based faseries
of unpublished manuscripts (A. Radford et
al.,eds.Vascular Flora of Southeastern United
States). The species occurs to the south in
Louisiana and eastward along the Gulf
Coastal Plain (Woodson 1954), but a voucher
from Arkansas has ae been lOeatee
Boltonia diffusa Ell. var.d )
ranner and Coville (1891) listed this variety
for Arkansas without citing a voucher. Efforts
since Buchholz and Palmer (1926) have
failed to locate a known and valid voucher.
Kartesz and Meacham (1999) listed this va-
riety for Arkansas, citing the compiled list of
Demaree (1943).
Buddleyja lindleyana Fortune ex Lindl.
(Buddlejaceae)—Kartesz and Meacham
(1999) listed this species as present in Ar-
kansas and cited a reference to one of a se-
ries of unpublished manuscripts (A.Radford
et al.,eds. Vascular Flora of the Southeastern
United States). Although known as a horti-
—tThe species was listed by Moore (1961); Smith (1988) noted the
cultural shrub planted across the state, it has
ot been reported to escape in Arkansas.
Carex debilis Mich. var. rudgei Bailey
(Cyperaceae)—This variety was listed for
Arkansas by Buchholz and Palmer (1926),
compiled by Demaree (1943), tracked as C.
debilis Michx. var. pubens Gray by Arkansas
Natural Heritage Commission, reduced to
synonymy by Smith (1988), and reinstated
as present by Kartesz and Meacham (1999)
citing ANHC. In Vol. 23 of Flora North
America, the two varieties are considered to
be asynonym of var.pubens, already known
from Arkansas.
Carex microrhyncha Mack. (Cyperaceae)—
Kartesz and Meacham (1999) listed this spe-
cies for Arkansas based on a personal com-
munication with S. D. Jones; Vol. 23 of ee
North America reduced it to a synonym
C. umbellata Schkuhr ex Willd. see
known from Arkansas
Carex tenera Dewey Cena —The spe-
cies was listed by Smith (1968) as not
present in Arkansas, but he considered it to
be a “possible addition’; listed erroneously
s “rare” for Arkansas by Kartesz and
Meacham (1999) citing Smith (19 88).
Carex verrucosa Muhl.(Cyperaceae)—
cies was listed by Demaree (1943). It was
reported by Smith (1968) as a “possible ad-
dition,” as it lacked a known voucher. It was
reinstated erroneously by Kartesz and
Meacham (1999) as" ‘present, but now extinct.”
voucher. Efforts since Buchholz and Palmer
(1926) have failed to locate a known and
1748 BRIT.ORG/SIDA 20
4)
valid voucher. Kartesz and Meacham (1999) Crataegus succulenta Schrad. ex Link (Ro-
listed this variety for Arkansas, citing the saceae )—This exotic and horticultural spe-
compiled list of Demaree (1943). cies was listed by Tucker (1976), based on
Chloris cucullata Bisch. (Poaceae)—This spe the expert authority and memory of EJ.
cies was listed by Branner and Coville (1891), Palmer. Palmer visited UARK in 1955 to an-
compiled by Demaree (1943), excluded by notate specimens and left a hand-written list
Smith (1988), and erroneously re-listed as of Crataegus from Arkansas based in part on
"?present” in Arkansas by Kartesz and As MeIOry: Smith (1978, 1988) listed it only
Meacham (1999) citing Smith (1988). as “possible” for Arkansas, citing Tucker
Clinopodium vulgare L. (Lamiaceae)—This (1976).Kartesz and Meacham (1999) errone-
species was listed for Arkansas as Satureja ously listed it for Arkansas and re Smith
vulgaris (L.) Fritsch. by Branner and Coville 1988). The list was not extant a
(1891), compiled by Demaree (1943), and Crinum americanum L. ie spe-
erroneously re-listed by Kartesz and cies was listed by Branner and Coville (1891),
Meacham (1999). It is likely that this record compiled by Demaree (1943), excluded by
was based upon mis-identified Satureja Smith (1988), and reinstated as “present” by
arkansanum Ne ) House. rather than upon Kartesz and Meacham (1999) by citing Smith
a garden escape, but a voucher is lacking to (1988). [The species was now excluded, as
confirm Ai ‘ the record was not backed with a known
Collinsonia canadensis L. (Lamiaceae)—This voucher
species was reported by Demaree (1943) Cucurbita maxima Duchesne (Cucurbi-
and excluded by Smith (1968) as the record taceae)—This species was excluded by
lacked known voucher. The species was er- Smith (1988) referring to cultivated material
roneously re-instated by Kartesz and from Newton Co.collected by Ralph Thamp-
Meacham (1999), but they cited Smith son in 1977; reinstated erroneously by
(1988). Kartesz and Meacham (1999) based on
Corispermum americanum (Nutt.) Nutt. var. Smith (1988). The species is not known in
americanum (Chenopodiaceae)—This va- Arkansas beyond cultivation.
riety was not listed by Smith (1988). Kartesz_ Cyperus thyrsiflorus jumehen (Cyperace-
and Meacham (1999) listed this variety for ae)—Kartesz and Meacham (1999) listed this
Arkansas and cited Smith (1988). Kartesz and species for Arkansas, based on a text pas
Meacham (1999) did not list as present in sage in a monograph on Central and South
Arkansas the related C. americanum (Nutt) American species (Tucker 1994). There are
Nutt. var. rydbergii Mosyakin, which Smith neither dots, maps, nor vouchers support-
(1988) documented in Arkansas under C. ing this record. In Vol. 23 of Flora North
hyssopifolium L.Perhaps the compiler added America, on p. 185, the species was not
the wrong variety and deleted the correct mapped for Arkansas but occurs in the Gulf
variety of this species for Arkansas. states to the south
Crataegus monogyna Jacq. eee —This Desmodium canadense (\..) DC. (Fabaceae) —
exotic and horticultural species was listed This species was reported by Demaree
without a voucher by Tucker (1976), based (1943), listed as not yet known from the state
on the expert authority and memory of EJ. but“possible”by Smith (1978, 1988). The spe-
Palmer. Palmer visited UARK in 1955 to an- cies does not in Arkansas according to Isely
notate specimens and left a hand-written list (1998), but is found to the north in Missouri
of Crataegus from Arkansas based in part on and northeastern North America. Kartesz
his memory. Smith (1978, 1988) listed it only and Meacham (1999) considered it
as “possible” for Arkansas, citing Tucker “?present’in Arkansas, citing “Gleason (1963)"
(1976). Kartesz and Meacham (1999) errone- [1952? or Gleason pals Snare 1963?)
ously re-listed it for Arkansas and cited Smith Without a known and valid voucher, the spe-
(1988). The list was not extant at : cies remains excluded from the flora.
PECK, ADDITIONS AND NOTES ON THE ARKANSAS FLORA
Droseta intermedia Hayne (Droseraceae)—
species was reported from Conway Co.
by ae (1951); a es excluded this
species as it was bas lentification
of D. brevifolia Pursh. The species was rein-
tated ly by! {Meacha
(1999) by citing Smith (1968).
Eragrostis elliottii S. Wat. (Poaceae)—Kartesz
and m (1999) listed this species for
Aas cna the revised Flora of Missouri
by Yatskievych (1999), who cited Texas
manuals by Gould (1975) and Correll and
Johnson (1970).A known voucher is lacking.
ea Reece vere at eer ane Pe er
(Asteraceae)—Branner and Coville (1891)
listed this variety for Arkansas without cit-
ing a voucher. Efforts since Buchholz and
Palmer (1926) have failed to locate a known
and valid voucher. Kartesz and Meacham
(1999) listed this variety for Arkansas, citing
only the compiled list of Demaree (1943
Erigeron pulchellus Michx. var. pulchellus
(Asteraceae)—Branner and Coville (1891)
listed this variety for Arkansas without cit-
ng a voucher. Efforts since Buchholz and
Palmer (1 ee have failed to locate a known
and valid voucher. Kartesz and Meacham
ie58 ee ve variety for Arkansas, citing
only the compiled list by Demaree (1943).
Eupatorium album L. var. album (Asterace-
—Branner and Coville (1891) listed this
variety for Arkansas without citing a voucher.
Efforts since Buchholz and Palmer (1926)
have failed to locate a known and valid
voucher. Kartesz and Meacham (1999) listed
this variety for Arkansas, citing only the com-
piled list by Demaree (1943).
Eustoma exaltatum (L.) Salib.ex G.Don subsp.
russellianum (Hook.) Kartesz (Gentianace-
ae)—This variety was reported for Arkansas
by Moore (1951). Shinners (1957) corrected
the varietal name to E. exa/tatum (L.) Salisb.
subsp exaltatum. The chat ge WaSI ioted by
Smith (1988) and the Arkansas Natural Heri-
tage Commission. The incorrect variety was
re-instated by Kartesz and Meacham (1999)
as “present” and by citing the ANHC. Kartesz
and Meacham (1999) also listed the correct
varietal name for the species in Arkansas
Glycyrrhiza lepidota Pursh (Fabaceae)— This
je¥)
wp
1
1749
species was reported by Branner and Coville
(1891), compiled by Demaree (1943), and
noted by Steyermark (1963). It was only
listed as “possible” in Arkansas by Smith
(1978,1988) as a voucher was not found. It
was not mapped in Arkansas by Isely (1998).
The species was erroneously listed by
Kartesz and Meacham (1999) by citing only
the compiled list by Demaree (1943).
Hieracium marianum Willd. (Asteraceae)—
Kartesz and Meacham (1999) listed this spe-
cies for Arkansas based on personal com-
munication with Guy Nesom. Nesom does
not recollect this communication, but thinks
that this record is unlikely (pers. comm. Guy
Nesom).Lacking a known voucher, the spe-
cies was excluded.
Humulus japonicus Sieb. & Zucc. (Cannabace-
—This exotic and weedy species was
listed as “present but extinct” in Arkansas by
Kartesz and Meacham (1999), who cited
Flora of North America (Vol. 3). Lacking a
known voucher, the species was excluded.
Hydrophyllum macrophyllum Nutt. (Hydro-
phyllaceae)—All prior reports, such as in
Smith (1988) are misidentified material of
recently named Hydrophyllum brownei Kral
Bates. The species was erroneously re-in-
stated for Arkansas by Kartesz and Meacham
(1999) along with the correct species, H.
brownel.
Lindera benzoin (L
je)
©
.) Blume var. benzoin
(Lauraceae)—T| iS Val iety was first reported
by Branner and Coville (1891), compiled by
Demaree (1943), but not mapped by Tucker
(1976). Smith (1978, 1988) who assigned all
Arkansas material to L. benzoin var.pubescens
Willd., but the incorrect variety was rein-
stated by Kartesz and Meacham (1999) who
cited Demaree (1943)
Linum lewisii Pursh var. lewisii (Linaceae)—The
variety was listed by Branner and Coville
(1891), compiled by Demaree (1943), ex-
cluded by Smith (1988) noting that Rogers
(1963) mapped the species no closer than
north-central Mississippi. Kartesz and
Meacham (1999) reinstated this species as
“present” based on an “unattributed per-
sonal communication.”
Lobelia puberula Michx. var. puberula
os
1750
(Campanulaceae)—Branner and Coville
(1891) listed this variety for Arkansas with-
out citing a voucher. Efforts since Buchholz
and Palmer (1926) have failed to locate a
known and valid voucher. Kartesz and
Meacham (1999) listed this variety for Arkan-
sas, citing only the compiled list by Demaree
(1943).
Lobelia siphilitica |. var. ludoviciana A. DC.
(Campanulaceae)—This variety was re-
d by Demaree (1943) based on his own
=
@
je8) T
ey
=%
m (1999) citing Demaree (1943). All
herbarium material in Arkansas and at BRIT
with this name on Demaree collections was
annotated as var. siphilitica by R. L. Wilbur,
Duke University, in 1
Lonicera reticulata Raf. (Caprifoliaceae)—This
ecies was reported by Demaree (1943) as
L. prolifera (Kirchn.) Booth ex Rehd., but
voucher has not been located,; Kartesz and
Meacham (1999) listed it as present in Ar-
kansas citing Demaree (1943).
Lycopus xsherardii Steele [(L. uniflorus x
virginicus] (Lamiaceae)—Henderson (1962)
mapped this hybrid for Arkansas in the form
of a shaded overlap of both parent species,
but without dots evident or vouchers cited.
Listed for Arkansas by Kartesz and Meacham
(1999), based on Henderson (1962).
Lysimachia tonsa (Wood) Wood ex Pax & R.
Kunth (Primulaceae)—This species was
listed by Demaree (1943) as Steironema
tonsum (Wood.) Bicknell, but was exclude
by Smith (1988) as a voucher was lacking.
The species was erroneously reinstated as
present by Kartesz and Meacham (1999).
Magnolia pyramidata Bartr. ex Pursh
(Magnoliaceae)—This species was reported
for Arkansas by Hardin (1972), without
voucher citations. Tucker (1976) and Smith
(1977, 1988) excluded this species. Kartesz
and Meacham (1999) erroneously re-listed
this species for Arkansas based on Hardin
(1972). No nivale: eer are Raw
Speci
that fit herbarium sheets rather than normal
sized leaves of M. tripetala (L.) L., leading to
misidentifications. Any plant from Arkansas
BRIT.ORG/SIDA 20(4)
would seem to be extralimital, based on the
maps of both species in Little (1976).
Matelea carolinensis (Jacq.) Woods.
(Asclepidaceae)—This species was listed by
Branner and Coville (1891), compiled by
Demaree (1943), and treated as an additional
representative of M. decipiens (Alexander)
Woods. by Smith (1988). The species was re-
instated by Kartesz and Meacham (1999) cit-
ing only the compiled list by Demaree
(1943)
—
sata a rhees L.(Liliaceae)— This species
was listed by Branner and Coville (1891),
ae by Demaree (1943), and listed as
a “possible addition” but not present by
Smith a as vouchers were not found.
The species was erroneously reinstated as
ey Kartesz ne Meacham (1999) by
eng ule (1988).
Mi Pursh) Heimer! (Nyctaginace
ae)—This apace was es as aan
igustifolia (Pursh) B.L. Robbins . Branner
and Coville (1891), compiled by Demaree
(1943), and excluded by Smith ae asa
voucher was lacking. The species was erro-
neously re-listed as"?present” by Kartesz and
Meacham (1999) a Smith (1988
Nymphaea odorata subsp. tuberosa
seein) Wietsina e eis PIPE ace:
ae) |
variety for Arkansas, cs a text passage in
the Muenscher (1944). Later treatments by
Fassett (1957) and Crow and Hellquist (2000)
place this variety to the north of Arkansas.
Oenothera nutans Atkinson & Bartlett
(Onagraceae)—Kartesz and Meacham
(1999) listed this species for Arkansas, citing
Dietrich et al. (1997). The species was not
mapped in Arkansas, nor do voucher cita-
tions list any Arkansas specimens in that ref-
erenc
Opuntia soediines (Raf.) Raf. var. austrina
(Small) Dress (Cactaceae)—Kartesz and
Meacham (1999) listed this variety for Arkan-
sas, citing only the compiled list by Demaree
(1943). Great Plains Flora Association (1977)
mapped this variety west of Arkansas, again
a possible confusion based on Arkansas Ter-
ritory that included Oklahoma.
es
PECK, ADDITIONS AND NOTES ON THE ARKANSAS FLORA
Oxydendrum arboreum (.) DC. (Ericaceae)—
This species was first listed by Lesquereux
(1860), re-listed by Branner and Coville
(1891), listed again by Moore (1941), and
compiled by Demaree (1943). It was ex-
cluded by Tucker (1976) as a voucher was
lacking. It was re-excluded by Smith (1978,
1988). It was erroneously reinstated as
“present” by Kartesz and Meacham (1999).
The species is known as a street tree in Ar-
kansas, but it has not escaped.
Paronychia fastigiata (Raf.) Fern. var.paleacea
Fern. (Caryophyllaceae)—Demaree (1943)
reported this variety for Arkansas based on
his own collections; Kartesz and Meacham
(1999) listed this variety for Arkansas. All
Demaree specimens with this name in Ar-
kansas and at BRIT were annotated in 1985
by J.W. Thieret to be var. fastigiata, already
known from Arkan
Pentodon adn: (K. Schum.) Vatke
(Rubiaceae)—Smith (1988) noted its pee
tcounties in |
on MacRoberts (1988) and suggested that
it might be searched for in Arkansas; Kartesz
and Meacham (1999) erroneously listed it as
“present” in Arkansas, citing Smith (1988).
Physocarpus opulifolius (L.) Maxim. var.
opulifolius (Rosaceae)—First reported by
Buchholz and Palmer (1926) and compiled
by Demaree (1943). All Arkansas material
was assigned to P opulifolius (L.) Maxim. var.
intermedius (Rydb.) Robinson by Tucker
(1976). Smith (1978, 1988) agreed. Kartesz
and Meacham (1999) erroneously reinstated
the incorrect variety by citing Demaree
(1943).
Physostegia digitalis Small (Lamiaceae)—
Kartesz and Meacham (1999) listed this spe-
cies as “?present” in Arkansas, referring to
Cantino (1982). A review of this source
shows no text passage mentioning this spe-
cies in Arkansas, nor map dots in Arkansas,
or vouchers
Plantago major L. (Plantaginaceae)— This spe-
cies was listed by Branner and Coville (1891),
and compiled by Demaree (1943). It was
listed as a“possible addition” but not present
by Smith (1988) who noted the absence of
1751
a voucher. It was erroneously reinstated by
Kartesz and Meacham (1999) citing only the
compiled list by Demaree (1943
Polygala verticillata L. var. verticillata
(Polygalaceae)—Branner and Coville (1891)
listed this variety for Arkansas without cit-
ing a voucher. Efforts since Buchholz and
Palmer (1926) have failed to locate a known
d valid voucher. Kartesz and Meacham
(1999) listed this variety for Arkansas, citing
only the compiled list by Demaree (1943).
Populus xcanadensis Moench. [P deltoides x
nigra] (Salicaceae)—This cultivated hybrid
was listed for Arkansas by Kartesz and
Meacham (1999). All specimens of this hy-
brid in Arkansas are cultivated materials
from Washington Co. at
Populus xcanescens Small [P alba x tremula]
(Salicaceae)—This cultivated hybrid was
listed for Arkansas by Kartesz and Meacham
(1999). All specimens of this hybrid in Arkan-
sas are cultivated materials from Newton Co.
and so se a aie
Populus jackii iSa
ie cultivated hybrid is
planted in southern Arkansas, but although
persisting and expanding as clones, it is not
known from any wild population: noris it es-
caping from tree farms. The hybrid was lis
by Tucker (1976) as sedan in serdsen nt
plantings. Smith (1978, 1988) excluded it
from the flora as it had not spread or natu-
ralized. Kartesz and Mecham (1999) errone-
ously listed it for Arkansas by citing Smith
(1988).
Potamogeton epihydrus Raf. (Potamogeton-
aceae)—Kartesz and Meacham (1999) list
this ‘or Arkansas, citing a pers. Communica-
tion with Robert Haynes. Haynes has no such
recollection nor any records documenting
a voucher for this record.
ee shud te engeum: Rosaceae)
Robertson (1974) remarked that this species
aii Cy in Nanas ee on this passage,
Kartesz and Meacham (1999) erroneously
listed this species for Arkansas.
Pyracantha coccinea M. Roemer (Rosaceae) —
mith (1988) excluded this species as all
known vouchers were based on cultivated
aor
pus
daltajidecl
1752 BRIT.ORG/SIDA 20(4)
material, such as that at UARK collected by Meacham ee peeee ee ete
Ralph Thompson from Newton Co. The spe- sas, citing only th aree
cies was listed erroneously as “present” by (1943)
Kartesz and ee (1999) by citing Smith = Rhynchospora nivea Boecki. (Cyperaceae)—
(1988). This species was first reported for Arkansas
Quercus rubra L. var. ambigua (A. Gray) Fern. in Correll and Johnson (1970). It was prob-
(Fagaceae)—The variety was listed by ably based on misidentified Arkansas mate-
Kartesz and aualal, (1999) for Arkansas, rial of 8. colorata (L.) Pfeiffer. All white-bracted
based only o iled list by Demaree Rhynchospora in Arkansas are specimens of
(1943). Tuickere 1976) and Smith (1978, 1988) R.colorata, a larger ae coarser pie typi-
excluded this variety as they considered it cal of southeastern United Sta rthan
to be a trivial name not reflective of qeo- the Oklahoma - Texas ere is ite
graphic variation. Vol. 3 of Flora North and finer in forn
America does not use this varietal name. Rudbeckia bicolor ey cere re
Quereus epumndicols Palmer [Q. bicolor x and Coville (1891) listed this species for Ar-
This hybrid was listed by kansas without citing a voucher. Efforts since
panties (1943). It was excluded by Tucker Buchholz and Palmer (1926) have failed to
(1976) as one of the parents (Q. bicolor) is not locate a known and valid voucher. Kartesz
known from Arkansas; Smith (1978, 1988) and Meacham (1999) listed this variety for
concurred. The hybrid was re-listed by Arkansas, citing only the compiled list by
Kartesz and Meacham (1999) by citing Demaree (1943).
Demaree (1943). Perhaps some of the con- — Salix amygdaloides Anderss. (Salicaceae)—
fusion can be traced to a specimen cited by This species was reported by Smith (1978,
Palmer (1948) from Pulaski Co,, Illinois, [not 1988) for Crittenden Co, Arkansas, but it was
Arkansas. excluded by Argus (1986) as it was based on
Quercus xincomita Palmer [Q. falcata x a mis-identified specimen of 5. eriocephala.
marilandica] (Fagaceae)—This hybrid was Great Plains Flora Association (1977)
listed as present by Demaree (1943). Mate- mapped it in Franklin Co., Arkansas, but it
rials assignable to this hybrid for Arkansas was excluded by Argus (1986) as the ee
were misidentified specimens of Q. x bushii men was collected in Franklin Co., Kan
Sarg. [Q. marilandica x velutina],according to‘ Scutellaria incana Biehler var.punctata ichap.
Palmer (1948, p. 19) and with his annotations an) C. Mohr (Lamiaceae)—Kartesz and
made in 1955 at UARK. The hybrid was erro- acur (1999) listed this variety for Arkan-
neously re-listed for Arkansas by Kartesz and sas, citing Fernald (1950). Epling (1942) does
Meacham (1999). not map this variety in Arkansas, but does
Ranunculus cymbalaria Pursh (Ranuncula- map S. incana Biehler var. incana. Lacking a
ceae)—Kartesz and Meacham (1999) nee known and valid voucher, this variety was
i species for Arkansas, citing “Sida 1976.” excluded from the state flora.
In that source, the species’ distribution was. Scutellaria nervosa Pursh (Lamiaceae)—
summarized in a text passage (Keener, 1976) Kartesz and Meacham (1999) listed this spe-
as including Texas, Oklahoma, Missouri, IIli- cies for Arkansas, citing only the compiled
nois,and New Jersey, but not Arkansas; there list by Demaree (1943). Epling (1942) does
are no dots or vouchers suggestive of an not map this species in Arkansas. Lacking a
Arkansas occurrence. known and valid voucher, this species was
Ranunculus pusillus Poir. var. pusillus excluded from the state flora
(Ranunculaceae)—Branner and Coville Sesuvium verrucosum Raf. ere ae This
(1891) listed this variety for Arkansas with- species was reported for Arkansas by Smith
out citing a voucher. Efforts since Buchholz (1978, 1988), but without a voucher, referring
and Palmer (1926) have failed to locate a to text passages in Correll and Johnso
known and valid voucher. Kartesz and (1970). As a known and valid voucher has
PECK, ADDITIONS AND NOTES ON THE ARKANSAS FLORA
not been found in the last 30 years, this spe-
cies was excluded from the flora of Arkansas.
Smilax illinoensis Mangaly (Smilacaceae)—
Kartesz and Meacham (1999) reported this
species for Arkansas based on Mangaly
(1968). Mangaly (1968) did not report S
illinoensis in Arkansas. A clerical error was
made in reading Map 22 on p.75,wherein a
dot for S. pulverulenta was incorrectly as-
signed to S. illinoensis.
Sporobolus airoides (Torr.) Torr. (Poaceae)—
This species was reported by Buchholz and
Palmer (1926), compiled by Demaree (1943),
reported with a voucher by Smith (1977). It
was then reported as misidentified aaa
of Sporobolus cryptandrus (Torr.) A. Gray by
Smith (1988). It was tracked by ee
Natural Heritage Commission under the
species historic name. It was reinstated er-
roneously by Kartesz and Meacham (1999)
as “present” by citing only the compiled list
by Demaree (1943) and ANHC.
Tridens muticus (Torr.) Nash var. muticus
(Poaceae)—This variety was listed by
Demaree (1943) as Triodia mutica (Torr.)
Scribn.,and compiled as present 2 Kartesz
and Meacham (1999) citing oe (1943).
The variety was excluded, as the see lacks
a known and valid buat
Llter | 1 ta NAi-h +1 |
Kartesz and Meacham (1999) listed this spe-
cies for Arkansas, citing Taylor (1989). Text
passages state that this species was in Ar-
kansas, but other passages state that its dis-
1753
tribution is in the southern Gulf Coastal Plain
(Florida to Texas), not Arkansas. As a known
and valid voucher is lacking, the species was
excluded.
Utricularia striata LeConte ex Torr. (Lentibular-
iaceae)—Kartesz and Meacham (1999) listed
this species as present for Arkansas, citing
text passages in Correll and Johnson (1970).
Taylor (1989) did not list it for Arkansas. Lack-
ing a known and valid voucher, the species
was excluded.
Verbena xdeamii Moldenke (Verbenaceae)—
This hybrid was listed by Kartesz and
Meacham (1999) for Arkansas based only on
the compiled list by Demaree (1943). The
hybrid occurs to the north of Arkansas
Moldenke, 1980). A voucher has not been
located in Arkansas or at BR
Viola arvensis Murr. (Violaceae)—This species
as listed for Arkansas by Branner and
Coville (1891), compiled by Demaree (1943),
but excluded from Arkansas in Russell (1965),
and re-excluded in Smith (1978, 1988), who
agi the apenas Ne voucher. t was rein-
| ham
(1999) by on oa the compiled list by
Demaree (1
Zanthoxylum cai Buckl. (Rutaceae)—
This species was reportedly in Arkansas ac-
cording to Correll and Johnson (1970).It was
relisted by Kartesz and Meacham (1999). All
shrubby material in Arkansas is assignable
to Z. americanum P. Mill; larger plants are Z.
clava-herculis L. (Tucker 1976).
—
ACKNOWLEDGMENTS
Thanks are expressed to John Kartesz and Chris Meacham for their efforts and
assistance in extracting the Arkansas data from their helpful and powerful na-
tional database. Thanks are expressed to BRIT, MO, and UARK herbaria direc-
torsand staff for their assistance and dedication to floristic efforts, to the ANHC
director and staff for their assistance and excellent record keeping, and to my
fellow members and representatives of the Arkansas Vascular Flora Project. The
author welcomes being informed of vouchers that would clarify the status of
these plants in Arkansas.
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Ertter, B. 2000. Our undiscovered heritage: Past and future prospects for species-level
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Fassett, N.C. 1957. Manual to aquatic plants. (Revised by E. Ogden) Univ. Wisconsin Press,
Madison.
Feanaco, M.L. 1950. Gray's manual. 8" ed. American Book Co., NY.
FerGuson, |.K. 1966. The genera of the Caprifoliaceae in the southeastern United States. J.
Armold Arbor. 47:33-59.
Fiora NortH America Epiroria Committee. 1993+ . Flora of North America North of Mexico.
Vols. 1,2, 3,4, 22, 23, 26. Oxford University Press, NY.
FRIEDLAND, S. 1941. The American species of Hemicarpha. Amer. J. Bot. 28:855-861.
GLeASON, H.L. 1952. The new Britton and Brown illustrated flora of the northeastern United
States and adjacent Canada. 3 vols. New York.
=
PECK, ADDITIONS AND NOTES ON THE ARKANSAS FLORA 1755
Gteason, H.L.and A.Cronauist. 1963.Manual of vascular plants of northeastern United States
and adjacent Canada. Princeton.
Goopman, G.J. 1966. Notes on Oklahoma plants. Proc. Okla. Acad. Sci. 46:3-4.
Goutp, FW. 1975. The grasses of Texas. Texas A&M Univ. Press, College Station.
Great PLAINS Flora Association. 1977. Atlas of the flora of the Great Plains. lowa State Univ.
Press, Ames.
Great PLAINS Flora ASSOCIATION. 1986. Flora of the Great Plains. University Press of Kansas,
Lawrence.
Greer, J.W. 1978 A revision of the New World species of Rhynchosia (Lleguminosae-
Faboideae). Mem. N.Y. Bot. Gard. 31(1):1-168.
Haron, J.W. 1972. Studies of the southeastern United States flora: Ill. Magnoliaceae and
liciaceae. J. Elisha Mitchell Sci. Soc. 88:30-32.
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Kartesz, J.T. 1998. Biota of North America [BONAP]:A synonymized checklist of the vascu-
lar flora of the United States, Puerto Rico, and the Virgin Islands. [Http://
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6:22-283.
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PECK, ADDITIONS AND NOTES ON THE ARKANSAS FLORA 1757
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SPIRIDON E. Kintzios (ed.). 2003. Oregano: The Genera Origanum and Lippia. (ISBN
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” eneiniety of Ori Se 4) The Turkish Origanum species; 5) The chemistry of the genus Lippia
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PART 5: Phar ances 8) The biological/ pharmacological activity ¢ e Orig g PART 6:
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foodstulfs and in food preparation. PART 7: Biotechnology; 10) The biotechnology of Oregano (Orig
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ants genera Origanum and Lippiain the
=)
anum Sp. and L apt sp.).
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medical bibliog databases g Ip
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SIDA 20(4): 1758. 2003
NORTHWARD RANGE EXTENSION IN FLORIDA
OF THE INVASIVE FERN
LYGODIUM MICROPHYLLUM (LYGODIACEAE)
Robert W. Pemberton
nvasive Plant Research Laboratory
SDA aes tural Research Service
College Ave
Pe ae Fl 33314, U.S.A.
a
ABSTRACT
] T BY x ] 1] } } z. oat
A newly recognized population of
in Orange County in central Florida. T his population of by rece is ae to have persisted within
a cypress swamp for almost 15 years, during which tim area experienced single or multi-night
freezes of -2.5 to -5.5°C. a finding indicates that the ae could es in other USDA Cold Hardiness
Q
Zone 9 areas such as southern Texas, which the fern may be able to reach via its wind-borne spores.
et
RESUMEN
S Condado de Orange l centro de la Florida blacié id
um. Se ae que esta poblacion de maleza ha
J
del helecho invasivo y peligroso, aes m mic ae ae
durante tal periodo de tiempo
persistido durante 15 anos
el area experimento heladas de -2.5 hast a-5.5° C ae ante una o mas noches. Esta observacion indica que
la maleza podria vivir en otras areas de la Zona de Resistencia al Frio (USDA) Numero 9, tal como el sur
de Tej as, area que el helecho puede alcanzar por las esporas llevadas por el viento.
Lygodium microphyllum (Cav) R.Br. (Lygodiaceae), known as Old World climb-
ing fern, is considered to be the most serious weed of swamps and other moist
habitats in southern Florida (Pemberton & Ferriter 1998). The weed is a severe
problem because it dominates and displaces native vegetation, is spreading rap-
idly, and lacks adequate or affordable control measures. The fern is native to the
Old World subtropics and tropics (Pemberton 1998), and was first found to be
naturalized in Martin County in 1965 (Beckner 1968). Since then, the plant has
spread to infest numerous susceptible habitats in all the Florida counties south
of the northern shore of Lake Okeechobee. The weed covered an estimated
40,500 ha in Florida as of 1999 (A. Ferriter pers. comm.), and has expanded con-
siderably since then. For instance, the infestation in Everglades National Park
was estimated to occupy 810 ha in December 2002, but ca. 4,050 ha in August,
2003, despite aggressive aerial spraying with herbicides (T. Pernas and J. Taylor
pers. comm.). The fern occurs much more sporadically north of Lake
Okeechobee but is becoming more abundant. Prior to this report, the known
northern limits of L. microphyllum were near Nalcrest in southeast Polk County
and at a site in southern Brevard County (Pemberton & Ferriter 1998), both at
SIDA 20(4): 1759-1761. 2003
1760 BRIT.ORG/SIDA 20(4)
ca. 27°50 latitude. The Brevard County population is probably recent, first de-
tected in 1997 (specimen sent to and confirmed by the author by M.A. Poole),
but the Polk County population was present at least since 1979, when a her-
barium sample (Willson 353, FLAS!) was collected.
During January 2003, an unrecognized population of L. microphyllum was
examined in a cypress swamp east of Orlando. This site, at 28°30'42°N, repre-
sents a 70 km northern range extension of the plant.
Voucher specimens: U.S.A. Florida. Orange Co.: Eastern Service Area Wetland “ tment : acuity on
the W side of Alafa, Trail, 3 km S of Hwy. 408, ca. 8 km E of Orlando, along the
on bald cypress and other plants within the swamp, 27 Jan 2003, Robert Pemberton s.n.(U SF).
Patches of the fern are present across an area of ca. 2 ha within the site, where
the fronds twine up cypress tree trunks, Taxodium distichum (L.) Rich. forming
large skirts consisting of living intertwined fronds layered over dead but per-
sistent strands of rachis (“stems”). The large size of these skirts, like those in the
long-established populations of the weed in southern Florida, indicates that
this Orange County population is not new. Lygodium microphyllum was found
during a vegetation analysis of this wetland during 1989, 1990, and 1991, but
misidentified as Lygodium japonicum (Thunb.) Sw. (L. Swartz , Camp Dresser
and Mckee Inc., here forth CDM Inc., unpublished reports). Lygodium japonicum
is the other invasive climbing fern that has naturalized in Florida, and the two
species are often confused. Lygodium japonicum rarely occurs in deeper wet-
lands such as swamps, but it can’t survive constant standing water like L.
microphyllum. Lygodium japonicum has sterile pinnules (leaflets”) that are
lobed or dissected, whereas L. microphyllum only occurs at wet sites in Florida
and has sterile pinnules that are usually unlobed. Furthermore, L. japonicum is
common in northern and central Florida (and occurs widely in the Southern
states), but L. microphyllum occurs, with a few exceptions such as this one, in
the southern third of Florida.
The occurrence of this newly recognized population of L. microphyllum in
the northern part of Orange County, and its persistence for almost 15 years, dem-
onstrates the plant's ability to tolerate significant single-night freezes. This
population, which was first found by CDM Inc. staff during July1989, survived
the 1989 Christmas freeze when temperatures dropped to -2.5-5.5°C during the
three day period (Orlando International Airport Weather Station #6628
records). On all three dates (Dec. 23, 24, and 25), the daytime temperatures
climbed above Ireezing. The airport is ca. 13 km to the southwest of the popula-
tion. This more northerly occurrence indicates that the plant’s ability to endure
lower temperatures of short duration is greater than previously recognized. The
observations suggest that L. microphyllum may be able to colonize additional
areas in central Florida, and probably northward, especially along the relatively
mild east coast of the state. The fact that L. microphyllum has not come to domi-
nate the Orange County cypress swamp, despite being there for ca. 15 or more
——t
PEMBERTON 1761
years, suggests that the weed is not as aggressive in central Florida as it is in
southern Florida. The ability of this fern to colonize and persist in central
Florida, in USDA-Cold Hardiness Zone 9 (Cathey 1990), as well that wetlands
in southern Texas and Louisiana are climatically suitable for the weed. The plant
produces large numbers of spores and occurs on many oceanic islands within
its native range both of which suggest that it may be able to reach these areas
via its wind-borne spores (Pemberton & Ferriter 1998). Because this fern’s ga-
metophytes are self fertile (Lott et al. 2003), only one spore is needed to start a
new infestation. Biological control may have the potential to control this weed
(Pemberton, 1998); a suite of specialist insect and mite herbivores is currently
being evaluated (Pemberton et al. 2002; Goolsby et al. 2003).
ACKNOWLEDGMENTS
I wish to thank J. B. Lee and L. Swartz, CDM Inc., Maitland, Florida, for provid-
ing access to the infested swamp and for information from CDM reports on the
vegetation monitoring at the site. Amy Ferriter, South Florida Water Manage-
ment District and Tony Pernas and Jonathan Taylor, Everglades National Park,
kindly shared estimates of L. microphyllum abundance. Paul Pratt, USDA-ARS,
Ft. Lauderdale, read and improved an earlier draft of this note, and Forest W.
Howard, Univ. Florida, Davie, wrote the Spanish language abstract. Kathy Burks,
Florida Department of Environmental Protection, Tallahassee, and Colette
Jacono, US. Geological Survey, Gainesville, FL provided knowledgeable reviews
of the manuscript for Sida, which helped me produce better final version.
REFERENCES
Beckner, J. 1968. Lygodium microphyllum, another escaped fern in Florida. Amer. Fern J.
CatHey, H.M. 1990. USDA plant hardiness zone map. USDA Agric. Res. Ser. Misc. Publ. 1475.
U.S.Government Printing Office, Washington, DC.
Gootsey, J.A., 7. WricHt and R.W Pemeerton. 2003. Exploratory surveys in Australia and Asia
for natural enemies of Lygodium microphyllum. Biol. Control 28:33-46.
Lort, M.S., J.C. Voun, R.W. Pemserton and D.F. Austin. 2003. The reproductive biology of
Lygodium microphyllum and L. japonicum (Schizaeceae): implications for invasive po-
tential. Amer. J. Bot. 90:1144-152.
PEMBERTON, R.W. and A. Ferriter. 1998. Old World climbing fern (Lygodium microphyllum), a
dangerous invasive weed in Florida. Amer. Fern J.84:165-75.
PEMBERTON, R.W.1998.The potential of biological control for the management of Old World
climbing fern (Lygodium microphyllum) in Florida. Amer. Fern J.84:1 76-82.
Pemberton, R.W., J. Gootssy, and T. WricHt. 2002. Old World climbing fern (Lygodium
microphyllum (Cav.) R.Br). In: R. Van Driesche, B. Blossey and M. Hoddle, S. Lyon, and R.
Reardon, eds. Biological control of invasive plants in the eastern United States. US For-
est Service Forest Health Technology Enterprise Team-2002-04, Morgantown, West
Virginia. Pp.139-47.
1762 BRIT.ORG/SIDA 20(4)
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Medicinal and Aromatic Plants—Industrial Profiles
Each volume in the series discusses the history and botany of each genus, and provides an in-depth
look at areas such as commercial cultivation, chemical and/or pharmacological aspects, toxicology,
current and/or future products, ete.
Maria Lis-Batcuin (ed.). 2002. Geranium and Pelargonium: ba cnaig Gerannm
and Pelargonium. (ISBN 0-415-28487-2, hbk.). Medici
Industrial Profiles. Volume 27. Series Editor: Roland Hardman. ae &
Francis, ll New Fetter Lane, London EC4P 4EE, UK and 29 West 35" Street,
New York, NY LOOO1, U.S.A. (Orders: info@tandf.co.uk, www.tandf.co.uk,
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color plates 7" 10".
—
From laylor & Francis website.—This book dispels the myths and incorrect information surround-
ing this [these] particular genera and will be of interest to graduate students, scientists, industry, and
those using Geranium and Pelargonium as an alternative medicine
Contents.—1) General yee chor PART I; Geranium: 2) rican of nomenclature, usage and
cultivation of Geranium and Pelar 3) The taxonomy of Geranium species and culti-
vars, their origins and growth in the wild: 4) P ve toc scheniiste y of the genus Geranium; 5) Cultivation
and harvesting of Geranium macrorrhizum and Geranium sanguineum for medicinal use in Bul-
garia; 6) Essential oil of Geranium macrorrhizum L eon extraction, See lige and use; 7)
argonium; 8) The taxonomy o
Use of Geranium species extr as herbal medicines. PART 2; Pe
Pelargonium species and aes us, eckde origins and ee in an wild: 9) C i, and sales ol
P era es ints for ornamental use in the UK and worldwide; 10) Growing pelargoniums in the
garden, conservatory and for shows; 11) Phytochemiistry of the genus Pelargonium; 12) Pharmacol-
oBy of i irgonium essential ac a extracts in vitro and in vivo; 13) Antimicrobial properties of
Pelargonium extracts contrasted with that of Geranium; 14) Essential oils from different Pe largo-
nium | species and Soe their chemical composition (using GC, GC/MS) and appearance of
), 15) eae of Pelargonium based on alkaloids ane essential oils;
16) P Saher re cee re within the genus Pelargonium based on the RAPD-PCR method of
DNA analysis correlated with the essential oil composition; 17) Geranium essential oil:
its detection using GC, enantiomeric columns, toxicity and
Sarena ISO; adulteration anc
bioactivity; 18) ‘Rose-scented geranium’ a Pelarg d r the perfume industry; 19) Culti-
vation and AMS of Geranium oil from Pe aieaun species in India; 20) Micropropagation
and biotechnological appeeeas sto iene culture of ie aan nas sand production of essen-
tial oils of scenteds; 21 nd cosmetic prod-
ucts utilising Geranium oil; 23) New research: possible uses ae various pete lec f oils and
extracts as food preservatives; 24) Pelargonium reniforme and Pelargonium sidoides: their botany,
chemistry and medicinal use; 25) Interactions between arthropod pests and i aerate ue Cor-
relation of the chemical profiles of the essential oil of Pelargonium (Geranium
rately and in mixes, with their relaxant or stimulant properties in man and smooth muscle cena a-
tions in vitro; and Index
SIDA 20(4): 1762. 2003
MUHLENBERGIA DUBIA (POACEAE) IN CENTRAL TEXAS
Geoffrey C. Denny!
Plant Resources Center
University of Texas
Austin, TX 78713, U.S.A.
Muhlenbergia dubia Four. ex Hemsl. isa wide ranging species known from Texas
and New Mexico, south through northeastern and north central Mexico to
Michoacan, the state of México, Puebla, Oaxaca, and Veracruz (Correll &
Johnston 1970; Gould 1975; Powell 1994; Herrera-Arrieta 2001). In Texas, it has
been considered to be confined the Trans-Pecos region (Correll & Johnston 1970;
Gould 1975; Powell 1994). It is a species primarily found on rocky hilltops and
canyons in open oak-juniper/pine woodlands.
In 1912, M_E. Jones described a new species, Muhlenbergia metcalfei, from
Grant County, New Mexico. Hitchcock (1951), Gould (1975), Powell (1994), and
Jones et al. (1997) recognized M. metcalfei as distinct from M. dubia Four. ex
Hemsl. and M. rigida (Kunth) Kunth. These two species are often difficult to
distinguish from each other and both are reported as occurring in trans-Pecos
Texas. However, Correll and Johnston (1970) and Martin and Hutchins (1980)
recognized M. metcalfei as synonymous with M. dubia. Powell (1994) only ten-
tatively recognized these taxa as distinct ‘pending further studies.’ He com-
mented that it is ‘difficult to defend specific status for such close entities with
sympatric distribution, Peterson et al.(2001) reduced M. metcalfei to synonymy
with M. rigida (Kunth) Kunth in their synopsis of Poaceae, subfamily Chlori-
doideae, however, the reasons for this taxanomic placement are not presented.
There is no published record of Muhlenbergia dubia in central Texas. How-
ever, during field work fora flora of the Nature Conservancy of Texas’ Love Creek
Nature Preserve in Bandera County, Texas in the southern Edwards Plateau, M.
dubia was collected by the author. Surveys condu ted revealed numerous, large
populations primarily in the uplands of the preserve, a habitat very similar to
habitats in which the species occurs throughout its range. The original speci-
mens were collected from a population on a sloping, open area with various
grasses, scattered Sophora secundiflora (Ortega) Lag. ex DC, Juniperus ashei J.
Buchholz, and Diospyros texana Scheele, and a few individuals of Quercus
fusiformis Small and cacti. The substrate wasa very thin, rocky, limestone soil.
Subsequent collections were made from five other populations on the preserve,
and one ona roadside that is very similar in habitat in Kerr County, Texas.
A review of the TEX, LL and TAES herbaria revealed that Muhlenbergia
‘Current address: Dept. of Horticultural Sciences, Texas A&M University, College Station, TX 77843-21 33, U.S.A.
gcdenny@tamu.edu.
SIDA 20(4): 1763-1764. 2003
1764 BRIT.ORG/SIDA 20(4)
dubia actually was collected on the Edwards Plateau as early as 1967. A speci-
menof M. dubia, identified as M. metcalfei, from Edwards County, Texas, on the
western edge of the Edwards Plateau, was collected in 1967 by Albert Jenkins.
A specimen of M. dubia was collected by M.C. Johnston in Travis County, Texas
in 1977. It was, however, originally misidentified as M. rigens, a similar species
also from the trans-Pecos area of Texas. The existence of M. dubia on the Ed-
wards Plateau of Texas is also known to the native plant nursery industry in
central Texas (Dan Hosage, pers. comm.). Additional specimens should be ex-
pected from other counties throughout the Edwards Plateau. The species is rela-
tively easy to distinguish from other central Texas species of Muhlenbergia by
its contracted panicles, densely caespitose habit, and rounded, erect culms.
peed
Specimens cited: TEXAS. Bandera Co.: Love Creek Nature Preserve. ca. 10 mi W of Medina, uplands
of preserve, | Nov 2000, Denny 16 (TEX); 26 Nov 2000, Denny 60 (TEX); 26 Nov 2000, De nny 6l(TEX):
15 Oct 2001, Denny 359 (TEX); 15 Oct 2001, Denny 361 (TEX); 17 Oct 2001, Carr 20366 (TEX); stream
terrace near hunter's cabin; 2 Sep 2001, Denny 327 (TEX). Edwards Co.: (without Seen locality) 14
Oct 1967, Jenkins 49 (TAES). Kerr Co.: roadside of Highway 16, 9.3 mi N of Medin E side of road,
growing in and thin rocky soil, 29° 54.761’ N, 99° 15.204' W. 15 Oct 2001, es nny 358 (TEX).
Travis Co.: above the pour-olf in ae aa Hollow, the farthest up-river corner of Lake Austin, 26 Oct
1977, M.C. sme ies
REFERENCES
Corrett, D.S. and M.C. JoHnston. 1970. Manual of the vascular plants of Texas. Texas Re-
search Foundation, Renner.
Goulb, FW. 1975. The grasses of Texas. A&M University Press, College Station.
Herrera-Arrieta, ¥. 2001. Las gramineas de Durango, 1° ed. Durango: IPN, CIIDIR Unidad
Durango: CONABIO, Comision Nacional para el Conocimiento y Uso de la Biodiversidad
HircHcock, A.S. 1951.Manual of the grasses of the United States, 2" ed, Revised by A.Chase.
Reprinted 1971. Dover, New York.
Jones, S.D.,J.K. Wirrr,and PM. Montcomery. 1997.Vascular plants of Texas: A compreshensive
checklist including bi hy,and index. Austin: University of Texas Press.
Maatin, W.C. and C.R. Hutcuins. 1980. A Gane hacer Mexico, vol. 1.Braunschweig, J.Cramer,
West Germany.
Peterson, P.M.,R.J. Sorena, G. Daviose, T.S. Fitcueiras, FO. Zutoaca, and E.J. Juoziewicz. 2001. Cata-
logue of New World grasses (Poaceae):II. Subfamily Chloridoideae. Contr. U.S. Natl. Herb.
41:1-255.
Powett, A.M. 1994. Grasses of the Trans-Pecos and adjacent areas. University of Texas Press,
Austin.
ROTALA ROTUNDIFOLIA (LYTHRACEAE)
NEW TO FLORIDA
Kathleen C. Burks Vernon V. Vandiver, Jr.
Florida Department of Enviror mer ital Protection Department of Agronomy, | IFAS
3800 Commonwealth Blvd, MS 705 cer = lorida
Tallahassee, FL 32399, U.S.A. 7922 NW 71" Street
kathy.burks@dep.state.fl.us Gainesville FL ie USA.
David W. Hall Colette C. Jacono
3666 NW 13" Place United States Geological Survey
Gainesville FL 32605, U.S.A. Florida Caribbean Science Center
7920 NW 71° Street
Gainesville FL 32653, U.S.A.
ABSTRACT
Naturalized populations of the Asian amphibious species Rotala rotundifolia are documented for
three peninsular Florida counties. Digdneue nine characters and a comment on invasive potential
are also provided.
RESUMEN
Se documentan las poblaciones naturalizadas de la especie Asiatica anfibia Rotala rotundifolia para
tres condados de Florida peninsular. Se presentan también caracteres distintivos y un comentario
acerca del potencial de invasion.
Rotala rotundifolia (Buch.-Ham. ex Roxb.) Koehne is a creeping amphibious
perennial found mostly in mountainous areas across a broad native range ex-
tending from India to Japan and China (Cook 1979; TROPICOS 2003). It has
been widely available in the international aquarium trade for more than three
decades (e.g., Roe 1967; Stodola 1967) and more recently promoted in the water
garden trade (e.g., Schuck 2000). “Dwarf Rotala,” as it is often called in the trade,
was first collected in Florida as an established escape in 1996 by the second
author while consulting with water managers in the City of Coral Springs,
Broward County in southeast Florida. A robust stand was spreading along the
edge of a flood-control canal coursing through a residential area. Samples of
naturalized populations have since been collected from a second canal loca-
tion in Coral Springs, a flood-control canal in Palm Beach County to the north,
and flood-control canals in the community of Lehigh Acres, Lee County in
southwest Florida (Fig. 1).
Representative specimens: FLORIDA. Broward Co.: Coral Springs, one block N of Sample Road, in
and along edge of canal about 46 m E of the intersection of Cardinal Road (NW 40" Street) and
Woodside Drive, N26.2793 W80.2285, 8 Mar 1996, Vandiver s.n. (FLAS); Coral Springs, in and along
SIDA 20(4): 1765-1769. 2003
BRIT.ORG/SIDA 20(4)
1766
OKEECHOBEE
A
ST. LUCIE
L. Okeechobee
‘
. ‘ West Palm Beach
\ hee
Seat, ties
Fort Myers X
Pompano Beach
Fort Lauderdale
0 50
Kilometers
Fic. is florida ulseribution of fatale planaiohi ek Each aah sks) two or more hie Adega within a particular
i cal non-governmental).
W bank of canal on E side of Cypress Park, FE side of Coral — Drive, 0.5 km S of cara
Drive, N26.2476 W80.2662, 4 Apr 2000, Vandiver s.n. (FLAS, NY, SWSL, USF). Lee Co.: Lehigh Acres,
S bank and shallows of Able Canal, N26.6118 W81.6412, 12 ve 2002, Jacono 418 (FI es 10 Jun 2002
Burks 1215 (FLAS, FSU, FTG, USF). Palm Beach Co.: Indian Trail Improvement District, N ol
Lion Country Safari, Development Unit 5, two blocks S of North Lake Boulevard
Loxahatchee and |
(CR-809A), in floating mats by culvert at the intersection of Hamlin Boulevard pore Districts” alee
EZ).
Canal (lormerly 170" Avenue), N26.8089 W80.3114, 12 Jun 2001, Vandiver s.n. (FLAS, U
Additional small patches or mats were noted in connecting canals of the In-
dian Trail system, Palm Beach County. A June 2002 survey of the Lehigh Acres
canal system, Lee County, revealed additional patches at the junction of Charlie
Diversion and Hickey Creek Canals (N26.6435 W81.6626) and on the east side
of Harns Marsh, a filtration impoundment (N26.6520 W81.6872); none was
BURKS ET AL., ROTALA ROTUNDIFOLIA NEW TO FLORIDA 1767
found at the outfall to the Orange River, a natural waterway that is part of the
Caloosahatchee River basin. In North America, the species is known from just
one other point of introduction—Tuscaloosa, Alabama, where it is well estab-
lished around a campus pond (Haynes 2002, 2003 pers. comm.).
Only two congeners of this species are known to occur in the continental
United States and Canada: the widespread native “toothcup,” Rotala ramosior
(L.) Koehne, and another introduced Asian species, Rotala indica (Willd.)
Koehne, which is known from rice field edges in Butte County, California (Ma-
son 1957; Oswald 1994) and in five parishes of Louisiana (Thieret 1972 [Thieret
33544, 1972, FLAS! FSUI], R. D. Thomas [NLU] 2000 pers. comm.). All three spe-
cies are low growing, glabrous herbs found in or near water; the leaves are de-
cussate or rarely in whorls of three. Rotala rotundifolia (Fig. 2) is easily distin-
guished by a usually perennial habit; emersed leaves obovate to orbicular, to 2
cm long and 2.5cm wide; a conspicuous rose-pink inflorescence of small spike-
like racemes that elongate and become lax in post-anthesis; and 4-valved cap-
sules (Cook 1979, 1990; Graham 1975; Reed 1977).
In the water-plant trade, R. rotundifolia has sometimes been confused with
R. indica (e.g., Schuck 2000) and with an endemic of southwest India, Rotala
macrandra Koehne (see discussion in Ben] 1972). The annual R. indica has leaves
with distinct cartilaginous margins, solitary flowers or short lateral spikes in
upper leaf axils, calyx tubes longer than broad, and 2-valved capsules (Cook
1979). The Indian plant R. macrandra is much like R. rotundifolia in form and
habit, but its submersed leaves are usually longer, to 2.5 cm, with their bases
remaining more rounded (Rataj & Horeman 1977). Also, R. macrandra’s spike-
like racemes usually remain dense in fruit, not elongated, and its stamens and
styles are long exserted from the calyx tube (Cook 1979).
Presently, the naturalized populations of Rotala rotundifolia on Florida's
east coast are much reduced in size—though not entirely eradicated—having
responded well to routine tools of chemical control for aquatic plants that may
impede water flow in these canal systems. Aquatic-label glyphosate and the
aquatic-granular form of 2,4-dichlorophenoxyacetic acid have proven to be the
most successful herbicides in stopping the spread of plants (2003 pers. comm.,
T. Narrow, N. Sculy). Multiple tools are used in the canals, including sterile trip-
loid grass carp, or white amur (Ctenopharyngodon idella (Valenciennes)), as a
biocontrol agent. However, these herbivorous fish apparently are not inclined
to eat this species; they have been observed nibbling at submersed foliage of R.
rotundifolia and immediately spitting it out (2003 pers. comm.,, T. Narrow).
On the west coast in the Lehigh Acres system, the current situation is less
encouraging, with significant spread of shoreline patches and floating mats in
the canals (2003 pers. comm., K. Waugh). However, managers are trying differ-
ent tools and continuing to check for plants at the outfall water gates.
Florida’s widely separated populations no doubt represent multiple intro-
1768 BRIT.ORG/SIDA 20(4)
Fic. 2. Rotal FY oa Pee let . *° L "
Nehiriulorl ry 1
1 g , lallahassee, Florida. a p
like racemes. b, Submersed leaves, light green, decussate. Growing in a concrete vat with native marsh plants, this
[ ies t f { ll | i 1| f | 15 years i i | [ 1 d Pg led g |
between vats (by fallen stem fragments). Cultivation vouchered by Hall 1842, 1988 (FLAS) and Anderson 154317, 1995
(FSU).
BURKS ET AL., ROTALA ROTUNDIFOLIA NEW TO FLORIDA 1769
ductions, most likely by dumping of unwanted aquarium or water-garden
material. Within each canal system, the plants have probably spread by float-
ing stem fragments, which root adventitiously at lower nodes. Seeds have been
observed in collected plant samples, but their levels of viability and germina-
tion under Florida conditions are not known. Cook (1979) considered this spe-
cies self-incompatible and insect-pollinated based on his cultivation studies.
The proximity of natural waterways and wetland conservation lands to
the canal systems (Fig. 1) calls for close monitoring to prevent inadvertent trans-
port of viable stem fragments, and possibly seeds, to natural habitats. More edu-
cational effort is also needed to revise the age-old human habit of inappropri-
ate disposal of unwanted goods.
ACKNOWLEDGMENTS
Our thanks go to local resource managers who assisted in the field and shared
their experience, Christine Kooienga, Terry Narrow, Nick Sculy, and Ken Waugh;
to Rob Kipker for generating the GIS-based map; and to C.D.K. Cook for con-
firming identification of a duplicate specimen.
REFERENCES
Bent, A.V.G. 1972. Rotala macrandra Koehne (Lythraceae). Die Aquarien- und Terrarien-
Zeitschrift (Stuttgart) 25:198-201.
Cook, C.D.K. 1979. A revision of the genus Rotala (Lythraceae). Boissiera 29:1—156.
Cook, C.D.K. 1990. Aquatic plant book. SPB Academic Publishing, The Hague.
GraHam, S.A. 1975. Taxonomy of the Lythraceae in the southeastern United States. Sida
0-103
Haynes, R.R. 2002. Noteworthy collections: Alabama, Rotala rotundifolia. Castanea 67:216.
Mason, H.L.1957.A flora of the marshes of California. Univ. of California Press, Los Angeles
1969 Reprint).
Oswato, V. 1994. Manual of the vascular plants of Butte County, California. California Native
Plant Society, Sacramento.
Ratas, K. and TJ. Horeman. 1977. Aquarium plants. T.F.H. Publications, Neptune City, New
Jersey.
Reep, C.F. 1977. Economically important foreign weeds. Agric. Handbook 498. Agric. Re-
search Service, U.S. Dept. of Agric., Washington, D.C
Roe, C. D. 1967. A manual of aquarium plants. Shirley Aquatics Ltd., Shirley, Solihull,
England.
Scuuck, S. 2000. Accent on aquatics: Rotala rotundifolia. Pondkeeper 6:40-41.
Stopota, J. 1967.Encyclopedia of water plants. T.F.H. ide io City, New Jersey.
Tuieret, J.W. 1972. Rotala indica (Lythraceae) in Louisiana. Sida 5
TROPICOS. 2003. VAScular TROPICOS occurrence database. pane Rev. 1.5. Mis-
souri Botanical Garden. http://mobot.mobot.org/W3T/search/vast.htm! (accessed July
12,2003)
—
1770 BRIT.ORG/SIDA 20(4)
BOOK NOTICE
TAYLOR & FRANCIS
Medicinal and Aromatic Plants—Industrial Profiles
Each volume in the series discusses the history and botany of each genus, and provides an in-depth
look at areas such as commercial cultivation, chemical and/or pharmacological aspects, toxicology,
current and/or future products, ete.
Satyajit D. SARKER and Yuji MARuyAMA (eds.). 2002. Magnolia: The Genus Magno-
lia. ISBN 0-415-28494-5, hbk.). Medicinal and Aromatic Plants—Industrial
Profiles. Volume 28. Series Editor: Roland Hardman. Taylor & Francis, l1
New Fetter Lane, London EC4P 4EE, UK and 29 West 35" Street, New York,
NY LOOOI, U.S.A. (Orders: info@tandf.co.uk, www.tandf.co.uk, 44 (0) 207
583 9855, lax +4 (O) 207-842-2298, standing orders call 44 (0)1264 343071.
US and Canada Customers: By mail: Taylor & Francis Customer Service,
10050 Toebben Drive, Independence, KY 41051, US.A., Toll Free Tel: 1-800-
634-7064, Toll Free Fax: 1-800-248-4724, email: cserve@routledge-ny.com).
$120.00, 187 pp., numerous b/w photos drawings and tables, 7" = 10".
From laylor & Francis website —“The genus Magnolia consists of several medicinally important spe-
: & )
}
cies most of which come from Far East Asia. Many species of this genus traditionally been used
in China and Japan to treat various illnesses, from simple head: aches to complic: ated cancer, and be-
cause of their versatility have more recently been incorporated int Ily successful medi-
cine preparations. ove with the revival of interest in herbal and oriental traditional medicines, many
of these Magnolia-containing preparations have captured a significant proportion of the drug mar-
ket in the Western countries. In recent years several studies have been performed with Magnolia
species, resulting in the isolation of a number of bioactive compounds and discovery of new biologi-
cal and pharmacological activities.”
Contents.-1) ae aus ae Genus Magnolia; 2) Traditional Medicine and Magnolia; 3)
Phytochemistry of the Genus ;4) Bioactivity and Pharmacological Aspects of Magnolia; 5)
yaa Control of eae Bark 6) Distribution and Commercial Cultivation of Magnolia; and
ndex
BOOK RECEIVED
RONALD L. Sruckry (compiler). 2003. Linking Ohio Geology and Botany. Papers
by Jane L. Forsyth. (SBN 0-9668034-7-7, hbk.). Ronald L. Stuckey, RLS
Creations, Columbus, OH, U.S.A. (Orders: Ronald L. Stuckey, RLS Creations,
PO. Box 12455, Columbus, OH 43212-0455, U.S.A., 919-962-0522). $30.00,
338 pp., b/w photos, color photos, maps, drawings, graphs, tables, 8 1/2" x
|B
SIDA 20(4): 1770. 2003
IN MEMORIAM
CHESTER MOREISON ROWELL IR.
1925-2003
Jim Blassingame
2020 Mustang
Levelland, TX 79336-6704, U.S.A.
The botanical community of Texas expresses its sorrow upon the death of Ches-
ter M. Rowell of Marfa, Texas, June 5,
Chester was born December 2, 1925 in Burnet County, Texas the eldest son
of Chester Morrison Rowell and Irene Gahagan Rowell. He received his B.S. from
the University of Texas in 1947, his M.S. from Texas AGM University in 1949
and his Ph.D. from Oklahoma State University in 1967.
Chester taught at Texas A & M University from 1949 to 1957, Texas Tech
University from 1957 to 1970 and Angelo State University from 1970 to 1987. He
was the curator of the herbaria at Texas Tech University and Angelo State Uni-
versity. He served as chairman of the Department of Biology at Angelo State
University from 1973 to 1982. After his retirement he was an adjunct Professor
of Biology at Sul Ross State University.
Professor Rowell did research on the flora of Southwestern U.S.and Mexico,
authored several scientific papers, presented papers at scientific meetings and
was author and co-author of two books. Chester was the scientific editor of the
Native Plant Society of Texas. Dr. Rowell was a driving force in building the
herbaria at Texas Tech University and Angelo State University in to excellent
regional collections. He worked on toxic plants and endangered species in Texas.
He was an advisor to the Texas Department of Parks and Wildlife for floristic
and habitat studies.
Chester was the former President of the Southwestern Association of Natu-
ralists. Dr. Rowell was the recipient of many awards and honors. He was active
in professional societies and other organizations.
Dr. Rowell was a respected researcher, valued teacher, dedicated mentor
and generous friend to people all around the country. He had a lifelong love
affair with photography. His enthusiasm for teaching, learning and scholar-
ship, his quick wit and cooperative spirit, were a role model for all who knew
him.
Chester Rowell was a teacher and a friend to all of those interested in the
botany of any area. Early in his career he decided that his students should be
exposed to life in the field and not just in the laboratory. After this decision
1772 BRIT.ORG/SIDA 20(4)
Chester M. Rowell (1925-2003)
Chester's life became one continuous field trip or collecting trip. All of his
students and friends were given the opportunity many times to partake in such
pane
an experience. Chester was very proud to be a professional plant taxonomist.
However, he was most proud to be a teacher as a source of stimulation and ex-
citing guide for the intellectual development of young people and friends. He
insisted that events of each life should be recorded on film. If you didn’t have a
camera, he would help you remedy the oversight with encouragement ora gift,
usually the latter. Chester enriched the lives of most botanists, photographers,
students and friends that he came in contact with.
Chester died June 5, 2003 in Midland, Texas at the age of 77. His many
friends and associates will miss him greatly.
BOOK NOTICES 1773
BOOK NOTICES
TAYLOR & FRANCIS
Medicinal and Aromatic Plants—Industrial Profiles
Each volume in the series discusses the history and botany of each genus, and provides an in-depth
look at areas such as commercial cultivation, chemical and/or pharmacological aspects, toxicology,
current and/or future products, etc
Maria Lis-BaLcuin (ed.). 2002. Lavender: The Genus Lavandula. (ISBN 0-415-28486-
4 hbk.). Medicinal and Aromatic Plants—Industrial Profiles. Volume 29.
Series Editor: Roland Hardman. Taylor & Francis, 1l New Fetter Lane, London
EC4P 4EE, UK and 29 West 35" Street, New York, NY LOOO1, U.S.A. (Or-
ders: info@tandf.co.uk, www.tandf.co.uk, 44 (0) 207 583 9855, fax 44 (0)
207-842-2298, standing orders call 44 (0)1264 343071. US and Canada
Customers: By mail: Taylor & Francis Customer Service, 10650 Toebben
Drive, Independence, KY 41051, U.S.A,, Toll Free Tel: 1-800-634-7064, Toll
Free Fax: 1-800-248-4724, email: cserve@routledge-ny.com). $128.00, 268
pp., numerous b/w photos drawings and tables, 17 color plates 7" 10"
From I Taylor & Francis website.—With contributions from lavender resea rchers Idwid
will be of interest both to graduate students and scientists in ee and industry who are in-
volved with lavender, to professionals in the lavender growing and retail industry, the perfumery,
ood and cosmetics industries, and to those interested in lavender for alternative and conventional
this book
_—
medical use.”
Contents.—1) General introduction to the genus Lavandula; 2) The taxonomy of the genus
Lavandula L; 3) History of usage of Lavandula species; 4) History of nomenclature of Lavandula
species, hybrids and cultivars; 5) Lavender growing in England for essential oil production; 6) The
retail lavender nursery; 7) Lavender growing in Australia; 8) Naming and misnaming of lavender
he lavender type oils: theory
as)
cultivars; 9) Phytochemistry of the genus Lavandula; 10) Distillation of t
oil: Henrie ISO; adulteration and its detection using
ntiomeric columns and bioactivity; 12) Lavender oil and its therapeutic properties; 13)
he psychological
GC, e
ee of Lavandula essential oils os extracts in vitro and in vivo; 14) 1
effects of lavender, 15) Antimicrobial properties of lavender volatile oil; 16) Lavender oil and its use
in aromatherapy; 17) Perfumery uses of lavender and lavaridin oils; 18) Miscellaneous uses of laven-
der and lavender oil: use in hair products, food flavouring, tissanes, herbal pillows and medicinal
products; 19) New research into Lavandula species, hybrids and cultivars, 20) Further research into
hegnail a species: cell cultures of L. vera and rosmarinic acid ee ee 21) Lavandula x
slex: 22) C omparative study of aes oil pcos
and lees 11) Lavender esse ial
ac a and L. x allardii:a puzzling com
of organically grown lavender and lavandin; 23) Chemical pro
+ ial nfromt n ~yltiv ars
essential oils see different
iles of lv any oils and pharmacology; 24) Chemical composition o
species, hybrids and cultivars of Lavandula
PN. RaviINpRAN and KJ. MADHUSOODANAN aNieas) 2003. Cardamon: The Genus Elettaria.
(ISBN 0-415-28493-7, hbk.) land A Plants—Industrial Profiles.
Volume 30. Series Editor: Roland Hardman. ae & Francis, 11 New Fet-
ter Lane, London EC4P 4EE, UK and 29 West 35" Street, New York, NY 10001,
SIDA 20(4): 1773. 2003
1774 BRIT.ORG/SIDA 20(4)
U.S.A. (Orders: info@tandf.co.uk, wwwtand!.co.uk, 44 (0) 207 583 9855, fax
44 (O) 207-842-2298, standing orders call 44 (0)1264 343071. US and Canada
Customers: By mail: Taylor @ Francis Customer Service, 10650 Toebben
Drive, Independence, KY 41051, U.S.A., Toll Free Tel: 1-800-634-7064, Toll
Free Fax: 1-800-248-4724, email: cserve@routledge-ny.com). $128.00, 374
pp. figs. (b/w photos, line drawings, graphs), tables, 7" x 10"
From laylor & Francis website.—“A textbook and reference work for scientists and students of horti-
culture, botany and related fields, this book will be the main reference volume on cardamom for
many years to come.
Contents.~1) Introduction, 2) Botany, crop improvement and biotechnology of cardamom; 3)
Chemistry of cardamom; 4) Agronomy and management of cardamom; 5) Nutrition of cardamom:
6) Viral diseases of cardamom; 7) Diseases of cardamom (fungal, bacterial and nematode diseases):
8) Insect pests of cardamom: 9) | ae processing of ci ard umom; ne Industrial processing
and products of cardamom; 11) Cardam : 12) Properties 1-uses of cardamom, 13)
Cardamom - future vision; 14) Yield gaps and production constraints in cardamom; 15) Large carda-
mom (Amomum subuldtum Roxb.); 16) False cardamoms, Appendix 1: Sten: for cards umom;
Appendix 2: Selected recipes; and Index.
STEPHEN FREER. 2003. Linnaeus’ Philosophia Botanica. (ISBN 0-19-850122-6, hbk.).
Translated from the Latin in 1751 by Stephen Freer. Oxford University Press.
(Orders: Oxford University Press, 2001 Evans Road, Cary, NC 27513, U.S.A,
800-451-7556, 919-677-1303 fax, le $277.50, 402 pp., 10 plates
with numerous figs. 8 1/2" x I"
Whata fascinating book this is, the translation of Carol Linnaeus’ Philosophia Bot (The Science
of Botany). The work was originally published in Latin in 1751 in Stockhol | Amsterdam. Stephen
Freer’s translation is a blessing to botanists, taxonomists, historians, and other interested parties; it
gives a new lease on and insight into the work of the father of modern systematics, Carl Linnaeus.—
Barney Lipscomb, Botanical Research Institute of Texas, 509 Pecan St, Fort Worth, TX 76102-4060, U.S.A.,
oarney@britorg.
EpZaRrD Ernst (ed.). 2003. Hypericum: The Genus Hypericum. (ISBN 0-415-36954-
|, hbk.). Medicinal and Aromatic Plants—Industrial Profiles. Volume 31. Series
Editor: Roland Hardman. Taylor & Francis, 11 New Fetter Lane, London
EC4P 4EE, UK and 29 West 35'" Street, New York, NY 10001, U.S.A. (Or-
ders: info@tandf.co.uk, www.tandl.co.uk, 44 (0) 207 583 9855, fax 44 (0)
207-842-2298, standing orders call 44 (0)1264 343071. US and Canada
Customers: By mail: Taylor & Francis Customer Service, 10650 Toebben
Drive, Independence, KY 41051, U.S.A., Toll Free Tel: 1-800-634-7064, Toll
Free Fax: 1-800-248-4724, email: cserve@routledge-ny.com). $120.00, 241
pp. figs. (color and b/w photos, line drawings, graphs), tables, 7" x 10".
ohn’s wort (Hypericum perforatum) is one of the best-selling
rom Taylor S Francis website. ==" 5b
herbal medicines worldwide. This landmark book provides a summary of our current knowledge on
SIDA 20(4): 1774. 2003
BOOK NOTICES 1775
a wide range of issues. It covers botany and includes plant infections, cultivation, manufacturing,
standardisation, cpauly SOnEEG!: biochemistry, pharmacology and calea? scene uae this
wide range of topics this 5,51 s: Which
are the pl | ul i 1s? Is St iche s wort effective ops severe pene What
are its s mechanisins ol action? W nat are othe! one m effect L risks? Should St John’s wort prod-
? How does it compare to synthetic drugs for the
ve
same indication?”
Cont tents. mY By pen cam pean, 2) Coulclay sale u a cause of St John’s wort
(t breeding for a tolerant variety; 3) A virus caus-
ohn’s
—
jie vein ellewme and necrotic leaf spots of St ee wort (Hypericum perforatum L.); 4) St.
wort herb extracts: manufacturing, standardisation and eee errs 5) Culture and biotech-
nology of Hypericum; 6) Chemical constituents of Hypericum ssp.; 7) Determination of hypericins
jane]
and hyperforin in herbal medicinal ae 8) Secondary metabolites content of Hypericumsp. in
different stages and plant parts; 9) Herbal medicinal eeuns of St John’s wort: manufacturing and
quality control; 10) The potential of in vitro cultures of Hypericum perforatum ang of ae um
aneirosde um to be rgauce interesting phar maceuticl compounds; 11) 1s clinical pharmacol f
2) Hy] g he work of Professor oy unig
oe Rec eeoup a Seoul); 13) N hopl logical studi Indian Hy; per
Linn; 14) Hypericumin dhetreatmentol Cer and Index.
Amit KrisHNA De (ed.). 2003. Capsicum: The Genus Capsicum. (ISBN 0-415-29991-
8, hbk.). Medicinal and Aromatic Plants—Industrial Profiles. Volume 33. Series
Editor: Roland Hardman. Taylor & Francis, 11 New Fetter Lane, London
EC4P 4EE, UK and 29 West 35" Street, New York, NY 10001, U.S.A. (Or-
ders: info@tandf.co.uk, www.tandf.co.uk, 44 (0) 207 583 9855, fax 44 (0)
207-842-2298, standing orders call 44 (0)1264 343071. US and Canada
Customers: By mail: Taylor & Francis Customer Service, 10650 Toebben
Drive, Independence, KY 41051, US.A., Toll Free Tel: 1-800-634-7064, Toll
Free Fax: 1-800-248-4724, email: cserve@routledge-ny.com). $125.00, 275
pp. figs. (b/w photos, graphs), tables, 7" = 10"
From Taylor & Francis website —‘This book, written by experts, includes material that deals with a
variety of ieee Lue ethnobotany, chemistry, pharmacology, toxicology, cultivation, analy-
sis, t rcial aspects of Capsicum.
Contents.—1) Capsicum: historical and botanical perspectives; 2) A glimpse of the world trade
in Capsicum; 3) Chemistry and quality control of Capsicums and Capsicum products; 4) Pungency
ogy: 5) ey nthesis of capsaicinoids
rinciples in Capsicum - analytical determinations and toxico
I y
in Capsicum; 6) Biotechnological studies on Capsicum for metabolite } 1 plant improve-
ment; 7) Irrigational aspects of Capsicum; 8) The cultivation and processing of Capsicum in India; 9)
The preservation and production of Capsicum in Hungary, 10) Post-harvest handling and process-
ing of Capsicums; 11) Advances in post-harvest processing technologies of Capsicum; 12) The stor-
age of Capsicum; 13) Current requirements on paprika powder for food industry, 14) Adulterants,
contaminants and pollutants in Capsicum products; 15) Colour differences in peppers and paprikas;
16) Future perspectives of capsaicin research; and Index.
SIDA 20(4): 1775. 2003
1776 BRIT.ORG/SIDA 20(4)
Auster D. Muir and New D. Westcort (eds.). 2003. Flax: The Genus Linum. (ISBN
0-415-30807-0, hbk.). Medicinal and Aromatic Plants—Industrial Profiles.
Volume 34. Series Editor: Roland Hardman. Taylor & Francis, 11 New Fet-
ter Lane, London EC4P 4EE, UK and 29 West 35th Street, New York, NY LOOO1,
U.S.A. (Orders: info@tandf.co.uk, www.tandf.co.uk, 44 (0) 207 583 9855, fax
44 (0) 207-842-2298, standing orders call 44 (0)1264 343071.US and Canada
Customers: By mail: Taylor @ Francis Customer Service, 10650 Toebben
Drive, Independence, KY 41051, US.A., Toll Free Tel: 1-800-634-7064, Toll
Free Fax: 1-800-248-4724, email: cset nak a ee $135.00, 307
pp. figs. (b/w photos, line drawings, graphs), tables, 7" >
From Taylor & Francis website—*“This volume is a comprehensive review of the genus covering all
une of “e taxonomy, chemistry, cultivation, pharmacology and commercial uses of flax, and
the disease prevention potential of these plant
Conte a 1) ie nega history of the cultivation and uses flaxseed; 2) Cultivated flax
and the genus Linum L.: Taxonomy and germplasm conservation; 3) Chemical studies on the con-
stituents of Linum spp. 4) Cultivation of flax; 5) Principal diseases of flax; 6) Principal insect pests
of flax; 7) The contribution of a-linolenic acid in flaxseed to human health; 8) The role of flaxseed
lignans in hormone-dependent and independent cancer, 9) Flaxseed in the prevention of cardiovas-
cular diseases; 10) Flaxseed and flaxseed lignans: Effects on the progression and severity of renal
‘laxseed constituents and human health;
failure; 11) Mammalian metabolism of flax lignans; 12)
13) Traditional food and medicinal uses of flaxseed; 14) Use of flaxseed in animal
sumer products with aa fatty acid profiles; 15) Current oS trends and economic impor-
tance of oilseed flax; 16) Current regulatory status of fk: ‘cial products; and Index.
Guise M. KavaL Alt (eds.). 2003. Urtica: Therapeutic and Nutritional Aspects of
Stinging Nettles. (SBN 0-415-30833-X, hbk.). Medicinal and Aromatic Plants—
Industrial Profiles. Volume 37. Series Editor: Roland Hardman. Taylor &
Francis, ll New Fetter Lane, London EC4P 4EE, UK and 29 West 35" Street,
New York, NY 1OOOI, U.S.A. (Orders: info@tandf.co.uk, www.tandf.co.uk,
44 (0) 207 583 9855, fax 44 (0) 207-842-2298, standing orders call 44 (0)1264
343071. US and Canada Customers: By mail: Taylor & Francis Customer
Service, 10650 Toebben Drive, Independence, KY 41051, U.S.A., Toll Free Tel:
1-800-634-7064, Toll Free Fax: 1-800-248-4724. email: cserve@routledge-
ny.com). $90.00, 83 pp., | b/w photo, tables, 7" « 10"
From Taylor & Francis website —“This volume explores the historical and modern uses of the Urtica
genus. An account of the botanical aspects of the genus is followed by a comprehensive review of the
recent research into its biologically active constituents. With further coverage of the therapeutic
uses and its nutritional value, this book presents an absorbing and all-encompassing account of the
medicinal potential of Urtica, which will be of considerable interest to herbalists, health scientists
and botanists as well as researchers in the pharmaceutical industry,
Contents.-1) An introduction to Urtica (botanical aspects); 2) Historical and modern uses ol
Urtica;3) The chemical and pharmacologic abespeceson! Irtica: ak sisphinasliea ane utic ey Urtica;
5) The therapeutic uses ol Ur rica in benign y | : t { Urtica
7 ft
species; 7) Urtica products; and Index.
SIDA 20(4): 1776. 2003
BOOK NOTICES 1777
Traditional Herbal Medicines for Modern Times
YUKIO OGIHARA and Masaki ABuraDa~ (eds.). 2003. Sho-Saiko-To: Scientific Evalu-
ation and Clinical Applications. (ISBN 0-415-30837-2, hbk.). Traditional
Herbal Medicines for Modern Times. Volume 4. Series Editor: Roland Hardman.
Taylor & Francis, 1] New Fetter Lane, London EC4P 4EE, UK and 29 West
35th Street, New York, NY 10001, U.S.A. (Orders: info@tandf.co.uk,
www.andf.co.uk, 44 (0) 207 583 9855, fax 44 (0) 207-842-2298, standing
orders call 44 (0)1264 343071. US and Canada Customers: By mail: Taylor
& Francis Customer Service, 10650 Toebben Drive, Independence, KY 41051,
US.A., Toll Free Tel: 1-800-634-7064, Toll Free Fax: 1-800-248-4724, email:
cserve@routledge-ny.com). $120.00, 223 pp., 81 figs. (b/w photos, graphs),
tables, 7" x 10".
ae Peel eiaaS : Lees > iy f, | ] wo
From ar & Francis website.—“Sho-Saiko-To set practitioners
who | li Sho-Saiko-To and di er how this and other Kampo
age ae be effectively integrated into Western medicine.
ents.—1) Introduction; 2) Traditional usage and therapeutic seats 3) Crude drugs I:
a items, collection and cultivation, production etc.; Crude drugs H: phytochemical stud-
ies of ingredients and analytical evaluation; Crude drugs HI: structural eat vente of ingredi-
ents; Crude drugs IV: poenmacolaic al and On hemical studies of medicinal aloe of Sho-saiko-to;
4) Formulation L: quality assessment gy and quality control of Sho-saiko-to
extract formulation; Formulation 11: phi aimaedancnes toxicology and Se eee aes of Sho-saiko-
to; 5) Clinical studies; 6) Recent topics on Sho-saiko-to and other Kampo medicines; and Index.
Biographical/Bibliographical
UNIVERSITY OF TENNESSEE PRESS
Cuaries Boewe (ed.). 2003. Profiles of Rafinesque. (ISBN 1-57233-225-5, hbk.).
University of Tennessee Press, Suite 110, Conference Center Building, Knox-
ville, TN 865-974-3321, U.S.A. (Orders: fax 865-974-3724, tpost@utk.edu,
www.utpress.org). $45.00, 411 pp., +5 figs., | color plate, 5" x 9"
Asa graduate student, Charles Boewe, editor of Profiles of ieee a oped an extracurricular
cur oy about CS. eae » After reading this book one can easily understand the fascination
for this multi-faceted, ayaleyhn esas mave a cee a number of ae s by various writ-
] 14 | 1fyR
biography of Constantin ee Rafinesque
and at the same time sketched a picture of the state of 19* century natural science
In his introduction, the editor discusses the difficulties of finding decisive piouraitedl mate-
rial and depends on an address given by Francis W. Pennell in 1940 to supply the framework for an
authentic account Eee he: life. Pips Eee Sg — ce an overview of Rafinesque’s work in an
alphabe IRITIS ¢ ‘ccanuiacntined Another writer deals
with - oe as a field naturalist, taxonomist, ad bisiosical theoretician in terms of modern
knowledge.
Rafinesque regarded himself primarily as a traveler and nothing he encountered was beyond
his purview including the study of man. He was intrigued by the American Indians he met on his
journeys, studied their languages and customs and endeavored to explain the mysterious Indian
SIDA 20(4): 1777. 2003
1778 BRIT.ORG/SIDA 20(4)
mounds he saw. His contributions to ethnology and philology and the hoax of the Walum Olum are
the subjects of a fascinating section.
Another part of the book is concerned with Rafinesque as a writer. There is a dissection of his
two-volume Medical Flora. Another essayist s tribute to him as a popularizer of botany—a trend
in the 19" century—with his book Sc ie of Flor
The final section contains an account by ae James Audubon of Rafinesque’s three week visit
with him in Florida where Rafinesque collected “multitudes of plants, shells, bats, and fishes.” And
at the hae Boewe clears up the mystery of where Ralinesque is buried, but not quite.
ich of the essays is self-contained. Together they offer a splendid biography of an intriguing
de The copious notes and 45 figures add to the value of this excellent work.—Ruth Ginsburg,
Botanical Research Institute of Texas, Fort Worth, TX, 76102-4060, U.S.A.
Mary Coker Josuin with drawings by SANDRA BRooKs-MATHERS. 2003. Essays on
William Chambers Coker, Passionate Botanist. (ISBN 0-97 21600-0-0, hbk. ).
University of North Carolina at Chapel Hill Library and the Botanical Garden
Foundation, Inc. (Orders: BGF Book Sales, c/o North Carolina Botanical
Garden, Totten Center CB#3375, University of North Carolina, Chapel Hill,
NC 27599-3375, US.A., 919-962-0522). $30.00, 197 pp., b/w photos, color
photos, drawings, 6" x 91/4"
Howard GARRETT. 2003. Dear Dirt Doctor: Questions Answered the Natural Way.
(ISBN 0-292-72847-6, pbk.). University of Texas Press, PO. Box 7819, Aus-
tin, TX 78713-7819, U.S.A. (Orders 800-252-3206 512-471-4032, 800-687-
6046 fax). $16.95, 240 pp., 51/2" x 81/2".
Howard Garret, Dallas/Fort Worth’s champion of gardening without toxic chemicals, has revised
and updated his 1995 Dirt Doctor’s Guide to Organic Gardening to a fast-reading, fact-filled interac-
tion between the Dirt Doctor and his personal bounty of garden knowle dge and the plant woes anc
wisdom of his readers and talk show listeners. From the opening chapter on the common sense and
economics of gardening organically to the ¢ g asing oft
armadillos and mice with cayenne
pepper powder (it worked on one smart uae mouse in my house!), Mr. Garrett covers soil anc
bed preparation, compost, synthetic vs. natural fertilizers, pest control and beneficial insects, land-
scaping ideas and maintenance, trees, herbs, food crops, wildflowers, turf, and indoor house ee
The coverage is attention-grab
an
sing, not refried generalities. For instance, on pepe pea a reader t
relates a New Delhi research report on using peppermint oil to repel the pest and kill its ver :
sprinkled my rainwater barrels with the oil immediately. It worked. On the ee ol nies the
opening paragraph t il, garlic, clovers, or |
feven
the most di phan strains of E.coli. | have used this herbal sien ae sq uad for eight years to keep
me outola st ic . They deserve opening line promine enc
A grand salute is in order to Howard Garrett for a most impressive compilation of informa-
tion—with but one caveat! There are rumblings of discontent in his cornmeal paradise (see
page 71).
Humalfa Corporation’s website, www.hu-more.com a
idvises against the use of cornmeal with its
“Green Bag” product after disturbing results form the field. There is more research to be done on its
el fecti iveness.—Karen M. Foley, Fort Worth, Texds.
SIDA 20(4): 1778. 2003
BOOK REVIEWS/NOTICES/RECEIVED 1779
BOOK REVIEWS/NOTICES/BOOKS RECEIVED
Environmental/Ecology/Conservation/Landscape Ecology
Grec Gorpon. 2003. Landscape of Desire. (ISBN 0-87421-560-9, ppk.). Utah State
University Press, 7800 Old Main Hill, Logan, UT 84322-7800, U.S.A. (Or-
ders: 435-797-1362, Fax 435-797-0313). $19.95, 213 pp. 1 map, 6" x 9".
The old professor (AKA the old philosopher) takes a group of students from the Sierra Institute fora
six weeks exploration of Utah’s Canyon Country. He wants each aa to experience the place “ona
th * To help reinvent pend
Lae level” and develop a personal awareness 0
s ‘they metamorphose from adolescence to adulthood,” everyone takes a trail name. If the reader
ee t ee accompanying Sage, Yucca, Seeker, Bobofet, Seaweed, Metta, Mud, and ae leberry, he
can learn a great deal from this oe Gordon is extremely knowledgeable about the natural history
and folklore of this magnificent country and describes it ina ively manner as the group descends
snaoueH the 16 geologic formations s of me San aaa regio
Another Dee the triy tthe abuse of the land by motorbikes and all terrain
vehicles. This lead ination of the ih : question of the conflict between development
and niecesauen It’s no contest. Gordon is a disciple of eee and Abbe
I missed maps and pictures.—Ruth Ginsburg, Botanical Research eieteek Texas, Fort Worth,
TX, 76102-4060, U.S.A.
VACLAV SMIL. 2002. The Earth’s Biosphere: Evolution, Dynamics, and Change. (ISBN
0-262-19472-4, hbk.). The MIT Press, 5 Cambridge Center, Cambridge, MA
02142-1493, U.S.A. (Orders: 800-405-1619, mitpress-orders@mit.edu, http:/
/mitpress.mit edu). $32.95, 346 pp., 138 illustrations, 8" x 9".
Contents.—1) Evolution of the Idea: From Vernadsky toa Science of the Global Environment; 2) Life
in the Universe: Attributes, Constraints, and Probabilities; 3) Life’s Diversity and Resilience: Metabo-
lisms, Species, Catastrophes; 4) Energizing the Biosphere: Solar Radiation and the Earth’s Heat; 5)
Water ae Material Flows: Biospheric Cycles; 6) The Biosphere’s Extent: The Moveable Boundaries;
7) The Biosphere’s Mass and Productiv ity: Quantifying Life’s Presence and Performance; 8) The
Biosphere’s Dynamics and Organization: Fundamental Rules and Grand Patterns; 9) Civilization and
the Biosphere: The Earth Transformed by Human Action; 10) Epilogue; Appendixes; References; Sci-
entific Name Index; Name Index; and Subject Index.
BriAN ALEXANDER. 2003. Rapture: How Biotech Became the New Religion. (ISBN
0-7382-0761-6, hbk.). Basic Books, A Member of the Perseus Books Group,
387 Park Avenue South, New York, NY 10016, U.S.A. (Orders: 212-340-8100,
www.basicbooks.com). $25.95, 256 pp., 6" x 91/4"
Contents.-1) Waiting for the Rapture; 2) The Prophet; 3) The Endless Frontier, 4) Arise, Lazarus
Long! 5) The Immortal Mr. Steinberg; 6) Way Out West; 7) Bring On the Inquisition Water into
Wine; 9) Pop! Goes the Rapture; 10) The Rapture Rides in a Limo; Acknowledgments; Notes, and
Index.
This book basically sheds light on the modern science of cloning, stem cells, miracle drugs,
and genetic engineering.
SIDA 20(4): 1779. 2003
1780 BRIT.ORG/SIDA 20(4)
ArtHUR MoLFLLA and Joyce Bept (eds.). 2003. Inventing for the Environment. (ISBN
0-262-13427-6, hbk.). The MIT Press, 5 Cambridge Center, Cambridge, MA
02142-1493, U.S.A. (Orders: 800-405-1619, Sete rae http:/
/mitpress.mit.edu). $29.95, 398 pp., 49 illustrations, 7" x 91/4"
ribes tl any ways In which invention alfects the
From the dustj Jac ket.—“This ambitious
environment (here defined broadly to include all for mol interaction een humans and nature).
- e built environment and the
The book starts with nature itself and then leads
specific technologies in areas such as public health and aneieae
Contents.—On Nature and Meee. What Role Does Innovation Play in Urban Lanc
How do Innovations in City Planning Shape the Environment? How Do | ions in Architecture
t? How can Innovations in Alternative Energy Sources Affec
Sc ape s?
Affect the Environmen ‘t the Environ-
ment, How are the Principles of Industrial Ecology Applied to Benefit the Environment; Conclusion:
The New Environmentalism, and Index.
PAULOS. OLIVEIRA AND Ropert J. MARQUIS (eds.). 2002. The Cerrados of Brazil: Ecology
and Natural History of a Neotropical Savanna. (ISBN 0-231-12043-5, hbk.).
Columbia University Press, 136 South Broadway, Irvington, NY 10533-2599,
U.S.A. (Orders: 800-944-1844 fax, http:/ www.columbia.edu cu/cup/).
$29.95, 398 pp., +9 illustrations, 7" x 91/4"
Contents. 1) Introduction: Development of ota in the oe aos piso Historical Framework
and the Abiotic Environment: 2) Relation af S he Brazilian Cerrado;
3) Late Quaternary History and Evolution of the Cerrados as Rees led by Palynological Records; 4)
The Fire Factor; 5) Past and Current Human Occupation and Land-Use. PART I. The Plant Commu-
l
Flora of
nity: Composition, Dynamics, and Life-History. 6) Vegetation Physiognomies and Woc
the Cerrado Biome; 7) Herbaceous Plant Communities; 8) Patterns and Dynamics of P pales
tions, 9) The Role of Fire in Population Dynamics of Woody Plants; 10) Ecophysiology of Woody
oO
Plants. PART HL. The Animal Community: Diversity and Biogeography. 11) Lepidoptera in the Cerrac
Landscape and the Conservation of Vegetation, Soil, and Topographical Mosaics; 12) The Character
and Dynamics of t lo Herpetol. 13) The Avifauna: Ecology, a ogri ee and Behavior;
14) The Cerrado rn Diversity, eae and Natural History. Vv. Insect-Plant Interac-
tions. 15) Ant Foraging on Plant Foliage: Contrasting Fffects on the one ean
Herbivores; 16) Interactions Among Cerrado plants and Their Herbivores: Unique or typical? Robert
o Plant Communities. PART V. The
Ecology of Insect
J. Marquis; 17) Pollination and Reproductive Biology in Cerrad
Conservation of the Cerrados. 18) Biodiversity and Conservation Priorities in the Cerrado Region;
Index.
SARA OLDFIELD. 2002. Rainforest. (ISBN 0-262-15106-5, hbk.). The MIT Press, 5
Cambridge Center, Cambridge, MA 02142-1493, U.S.A. (Orders: 800-405-
1619, mitpress-orders@mit.edu, http://mitpress.mit.edu). $29.95, 160 pp.
200 color illustrations, 91/2" x 12 1/2".
us book is a great reminder of the incredible beauty and diversity of rainforests around the world.
The stunning photography is by Bruce Coleman Collection. Review forthcoming in Sida 21(1), 2004.
SIDA 20(4): 1780. 2003
BOOK REVIEWS/NOTICES/RECEIVED 1781
Loren M. Smitu. 2003. Playas of the Great Plains. (ISBN 0-292-70177-2, pbk; O-
292-70534-4, hbk.). University of Texas Press, PO. Box 7819, Austin, TX 78713-
7819, US.A., (Orders: 800-252-3206, 800-687-6046 fax). $24.95 (pbk), $60.00
(hbk), 264 pp., 34 b/w ane 6 sia Oe:
the Playas of the High Plains. There are eight
This book is about wetland ecology a
shapters broken down under three main sections; ares and their Environment: L) What is a Playa?
2) ce and Development. Ecosystem Aspects: 3) Flora; 4) Fauna; 5) Structure, Function, and Di-
versity. Conservation Aspects: 6) Historical, Cultural, and Current Societal ae of Playas; 7) Threats
to Proper Function of Playas; 8) Conservation Past, Present, and Future; Appendix; References; Index
James A. YouNGand CHARLIE D. CLEMENTS. 2002. Purshia: The Wild and Bitter Roses.
(ISBN 0-87417-491-0, hbk.). University of Nevada Press, Mail Stop 166, Reno,
NV 89557-0076, U.S.A. (Orders: 775-784-6573 x232, 775-784-6200 fax,
www.nvbooks.nevada.edu). $39.95, 280 pp., 35 b/w photos, 4 illustrations,
3 maps, 6" x 91/4".
hia is in the family Rosaceae. The book is an indepth look at the er and manage-
ment of habitat for these importz especies! plants. chatlomn a vital 1 the western and
] } eae 7 ]
The genus Purs
{ the wild
intermountain vegetation. T
and bitter roses to bitterbrush Slane commninices to insects one pant ee to management is-
sues related to Purshia species. The text is supplemented with line d gs and photographs
Plant Physiology/Plant Anatomy/Pathology/Agriculture
RicHarb C. Keatinc. 2003. Anatomy of the Monocotyledons. IX Acoraceae and
Araceae. (ISBN 0-19-850122-6, hbk.). M. Grecory and D.F Cutter (eds.). Oxford
University Press. (Orders: Oxford University Press, 2001 Evans Road, Cary,
NC 27513, US.A., 800-451-7556, 919-677-1303 fax, www.oup.com). $125.00,
Berg PP. 855 photomi« rog aphs Tig RS) plates, fale ee ace eae
This is volume nine in the continuation of the series Anatomy of the Monocotyledons. Based upon
anatomical, morphological, and molecular data, a new classification system of the Araceae is pre-
sented.
SIDA 20(4): 1781. 2003
1782 BRIT.ORG/SIDA 20(4)
BOOK NOTICES
IOWA STATE PRESS
(A BLACKWELL PUBLISHING COMPANY)
KRISTINA BOONE, TERRY MEISENBACH, and Mark Tucker. 2003. Agricultural Commu-
nications: Changes and Challenges. (ISBN 0-8138-2167-3, pbk.). Hardback
(9 edition 2000. lowa State Press, 2121 State Avenue, Ames, [A 50014, U.S.A.
(Orders: 1-800-862-6657, 1-515-292-3348 fax; www.iowastatepress.com).
$29.99, 144 pp., b/w photos and other illustrations, 6" x 9"
—
PETER F Frotiiott, Luis A.Bojorquez-T aria, and MARIANO HERNANDEZ-NARVAEZ. 2003.
Natural Resources Management Practices: A Primer. (ISBN 0-8138-0913-
4, pbk.). Hardback (1s) edition 2001. Iowa State Press, 2121 State Avenue,
Ames, IA 50014, U.S.A. (Orders: 1-800-862-6657, 1-515-292-3348 fax:
www.owastatepress.com). $34.99, 256 pp., illustrations, 6" x 9".
Cuartes B. Moss and ANpREw Scumitz. 2003. Government Policy and Farmland
Markets: The Maintenance of Farmer Wealth. (ISBN 0-8138-2329-9, hbk.).
lowa State Press, 2121 State Avenue, Ames, [A 50014, U.S.A. (Orders: 1-800-
862-6657, 1-515-292-3348 fax; www.iowastatepress.com). $59.99, 432 pp.,
tables and graphs, 6" x 9".
The book has 21 chapters that addresses farmland values and is divided into six sections: Section I:
Historical Perspectives on Farmland Values; Section H: Government Policies and Farmland Values;
Section HL: Capital Markets and Farmland Values; Section IV: Transaction Costs and Farmland Val-
ues; Section V: Urbanization, Environmental Quality, and Farmland Values; Section VI: Regional and
International Dimensions.
COLIN G. ScANes and Joun A. MirnANowsk1. 2004. Perspectives in World Food and
Agriculture: 2004. (ISBN 0-8138-2021-9, hbk.). lowa State Press, 2121 State
Avenue, Ames, IA 50014, U.S.A. (Orders: 1-800-862-6657, 1-515-292-3348
lax; wwwiowastatepress.com). $69.99, 485 pp., tables, graphs, drawings, 6"
—o
Perspectives in World Food and Agriculture: 2004, written by a team of international researchers,
scientists, and academics, is about global agriculture. It brings together cutting-edge agricultural
research in 25 chapters, in six broad sections: Section I: Frontiers in World Food; Section I: Frontiers
in Food; Seetion ILE Frontiers in Animal Agriculture; Section IV: Frontiers in Water; Section V: Fron-
tiers in Policy and Ethics; Section VE: Statistics and Trends in World Agriculture.
SIDA 20(4): 1782. 2003
BOOK REVIEWS 1783
BOOK REVIEWS
Dorotuy HiNnsHAW Patent. Photographs by WILLIAM MuNoz. 2003. Plants on the
Trail with Lewis and Clark. (ISBN 0-618-06776-0, hbk.). Clarion Books, a
Houghton Mifflin Company imprint, 215 a Avenue South, New York,
NY 10003, U.S.A. (Orders: http://www! ff linbooks.com /cata-
eee cfm?titleNumber=111018). $18, 00, 112 pp., color, b/w pho-
tos, 8" x
This companion to Animals on the Trail with Lewis and Clark (Clarion 2002) by Patent and Munoz
traces Lewis’ and Clark’s epic journey citing useful plants along the way and many of the plants they
introduced to science. Thongs speciically written for rchilenen in grades 4 - 8, the interesting text
1 {.
> a
and beautiful y
ALLYORI’ wanting tO know TM1OTe about
the feaponeanice of } tl f the expedition:
e book spans witha a beautiful two-page map that highlights the route of the explorers from
St. Louis in the midwest to Clatsop on the Pacific Ocean, showing camps and landmarks along
the way. Lewis and Clark ne a 2000-mile route with a crew of about 40 men with orders from
President Thomas Jefferson to find a navigable river passage across North America, describe, map
Pe | ] } ] - ee tase J a a | Si = -] J
4 LT
WIth
5 Ie
iF |
the soils, plant d animals tl don the journey. Patent and Munoz do an excellent job
I J y :
of helping the reader understand how plants were used by members of the Expedition and Lewis,
and Clark’s role in documenting these plants, many of which were new to science. The book dis-
cusses the importance of trees to the explorers’ success from providing wood for the keelboat and
pirogues to take the Expedition upriver, to supplying fuel for their many campfires. Patent also de-
scribes the plants used as food and medicine along the way, often citing the Indian tribe that intro-
euced ing acai de mem be rs to various new sources =k vitamins or ieedics exces from Lewis’
dc “reactions to
P
seeing enema iar plants ae landscapes for the first ti plants as foods. Munoz’s
striking color photographs on nearly every page spotlight Seomples of Pndigdual plants important
to the expedition and scenic views along the trail, and they give the reader glimpses of the beautiful
vistas the members of the Corps of Discovery must have witnessed on their incredible journe
A final section discusses the fate of the specimens collected by Meriwether Lewis with color
photographs of several select specimens. Patent explains Frederick Pursh’s role in describing, illus-
trating and naming the plants collected on the expedition. Appendices include a recommended list
of other books and websites regarding the Lewis and Clark Expedition, and a list of the surviving
plants in the Lewis and Clark Herbarium at the oa of Natural Sciences in Philadelphia with
information on each specimen’s date and location of collection.
book piowicee me with a great deal of helpful ees that I use in our school pro-
gram and teacher workshop “The Lewis and Clark Expedition Through the Eyes of Art and Science.”
The focus of this program is on the botanical Ss of Lewis and Clark, and I have sought
many sources for background informatio th ‘i s collected by Lewis and Clark and the
importance of plants on the expedition. ie book alone provided more helpful information than
many sources combined.—Amber Keller, Coordinator of School Services, Botanical Research Institute
of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A, akeller@britorg.
SIDA 20(4): 1783. 2003
1784 BRIT.ORG/SIDA 20(4)
PAUL MARTIN Brown. 2002. Wild Orchids of Florida, with References to the At-
lantic and Gulf Coastal Plains. (ISBN 0-8130-2439-0, pbk.). University Press
of Florida, 15 Northwest 15" St, Gainesville, FL 32611-2079, U.S.A. (Orders:
www.upf.com, 800-226-3822). $24.95, 428 pp., 447 color photos, 135 b/w
illus. 6" x 9"
PAUL MARTIN Brown. 2003. The Wild Orchids of North America, North of Mexico.
(ISBN 0-8130-2571-0, pbk.). University Press of Florida, 15 Northwest 15"
St, Gainesville, FL 32611-2079, U.S.A. (Orders: www.upf.com, 800-226-3822).
$27.95, 256 pp., 295 color photos, 247 b/w illus., 6" x 9".
The appeal ol orchids, and books about orchids, hardly requires explanation to anyone who has ever
admired or desired these plants. Even common orchids manage to maintain an air of elegant and
exotic (or erotic) mystery. The name Paul Martin Brown should be well known to American orchid
fanatics. He is the founder and editor of the North American Native Orchid Journal, and has pub-
lished several previous field guides on orchids. Most recently, he was a major contributor to the
Orchidaceae treatment in Volume 26 of Flora North America. Brown has produced two exemplary
orchid field guides, for North America and for Florida, with profuse color photographs and excellent
ustrations by Stan Folsom.
Wild Orchids of Florida covers L118 species and varieties, 26 growt rms, and 6 hybrids.
Approximately half the orchid species in North America are ‘found in Florida. 1 - book begins with
some information not typically found in field guides, including a history of orchid work in Florida, a
shapter wel orchids in general (morphology, habit, habitat), and a section on how to use a dichoto
mous key. The last is something that should definitely be included in any similar technical works
with suc i aw Hee and general appeal. Following this introduction, a key to the genera of orchids in
5. Briel
Florida is provided and supplemented with illustrations of some important ¢
explanations are then given concerning the logic behind scientific names; the nee ies ol common
3
=
4
es; variations in size, color, and flowering time; and range map use. A map of Florida is provided,
with all counties labelec
The orchids eee are divided into two sections: native and naturalized species, followed
by introduced, escaped, or wail species. Within these sections, ¢ - in alphabetical
order. Fach genus is introduced by a general description and statement of 1 ae event Florida, and
a key is given to the species, if more than one is found in the state. Each species is then treated with
nearly a [ull page of text, listing the Latin name, common name, world range, range within Florida
(hequently mrciicine status, if threatened or endangered), a description of the plant (habit, leaves,
ling size ranges, habitat, and flowering period. This is f followed a an a
cael cane ancl interest, Which may include more inf rarity
ol the species, life history, nomenclatural history, or a discussion of cee eee variation within
that species over a geographic range. Also on this page is a pen-at ink pestabon of the (usually)
entire plant, and a Florida range map, with one dot in each county led. The
illustrations generally depict the entire habit of the ees including oe ale and are detailed
se dr gs al The facing page is full of
enough so that most casas coul d be identified by
a
various color pl plants Or populations of each species, ee ups ol intlores-
cences and floweis or fruits. rs lew poor-quality photographs do not detract from the beauty or util-
ity of this book.
Afi | t ritl
1 “Resources and References” provides a great deal of useful information,
including a simple checklist of all taxa, recent literature references (listed by species), synonyms
SIDA 20(4): 1784. 2003
BOOK REVIEWS 1785
and misapplied names, and statistics on endangered and endemic orchid species in Florida. An in
teresting chapter called “Orch " Hunting in Florida” divides the state into regions, and then for each
region provides a discussion of habitats and a list of species. A special chapter that can be used out-
a of Blonde: but still within the Gulf Coastal Plains of the southeastern USA, lists all extra-Flo-
ridian orchid species and their ranges. The very brief glossary will probably require supplementa-
tion froma botany text for many users of the keys. Other than this, Wild Orchids of Florida isa perfect
field guide, with keys that will allow the identification of an orchid in-hand. Inversely, this book has
enough supplementary information that a reader can probably select a (relatively common) orchid
species from the book, and then go and successfully find it in the wild. Beyond these uses, the notes
on each species make for entertaining armchair reading, and the photos are so enjoyable that any
Floridian sand , other orchid fanciers) would be glad to own this boo
TheV
Cay ae as 3
3 species, 24 sieeacs and varieties, 103 growth/
color forms, and 24 hy bride The checklist is dered alphabetically by genus and species. Hybrids
are listed at the end of each genus. Generally, there are two tae per page. Each species entry gives
the Latin name and authority, synonyms and misapplied names, common name, nee sane list
of subspecific forms if any, references, and a brief comment ne t the plant's ey, rarity, or
preferred habitat. Non-native species are indicated with an asterisk next to the -atin name. Each
species is illustrated with a pen-and-ink drawing and a color photograph. a ne poole are excellent
] 1] | l UW 1
ALI USUALLY ott Upe of the flowe1 Or int lores« ClICe e photos not the draw-
ings have earenan of scale, and nothing about the size of the plants or pues is given in the
species descriptions.
My main complaint about this section of the book is the layout of the text. In an effort to take
up a minimum amount of page per entry, not enough space has been allowed between each element
(synonyms, rane relerences, comment) and some lines are indented in a confusing fashion. Final
comments for I form, rather than in sentences, and begin without aca ia
talized first letter, causing these to visually run into the previous element in a distracting way. F
many species, no references at all are given; for ober csibse or more are listed. The traduction
] f,,]] }
encouragement to orchid a ee toseek just for f
what blunted by the lack of any habitat information ‘A all for aon of the species Sonuavell Follow-
ing the main list of species, a shorter sec color photos of some of the forms or hybrids
already listed in the front section of the book. The next eae contains technical keys to all the
orchid genera and species in North America. The appendices include a list of excluded species, and
errata for Carlyle Leur’s classic works The Native Orchids of Florida (1972) and The Native Orchids of
the United States (1975). The glossary is handy, but is missing some terms used in the keys, such as
flabellate, pedicel, and rostellum. The term peduncle is misdefined as * the stalk of a flower or leaf,”
when it isin fact the stalk of anin
‘loresc ence. FON Onan the bibliography, a personal checklist sec-
| I e list.” The North American orchid book will be handy
for someone who has found an vorehia and ue like to neo it but does not desire much more
information beyond the correct name of the species. It is al g book to browse for the
sake of the photographs, and could be used to f ane oneself with the genera of North American
orchids.
Both books are durably “flexibound,’ and the fold-out endpages have useful metric and English
rulers on their margins.—Amanda - eas Botanical Research Institute of Texas, 509 Pecan St, Fort
Worth, TX 76102-4060, U.S.A., ane
tion is provided for those
rg iS
SIDA 20(4): 1785. 2003
1786 BRIT.ORG/SIDA 20(4)
Jens E. Mapsen, Ropert L. Mix, and Henrick BALSsLEV. 2001. Flora of Puna Island:
Resources on a Tropical Island. (ISBN 87 7288 854 7, pbk.). Aarhus Uni-
versity Press, Langelandsgade 177, DK-8200 Aarhus N, Denmark. (Orders:
www.unipress.dk, +45/89-42-53-80 fax). $39.95, 289 pp., 35 color photos,
10 b/w illus., 6 3/4" x 91/2".
Puna Island is situated in the Pacific Ocean near the equator, in the gulf of Guayaquil in Ecuador.
Though only 3 km from the coast, the small (855 sq. km) island has a flora rich in near-endemic
species. Puna Island also has an interesting cultural history of early human occupation (5000 BC),
>
es = island’s natural vegetation has remained fairly unaltered ces as to the main cate
1] a] (s ld f,
Well preserved remnant or tropical ary lorest—an extt ly
)
with less than 1% remaining in the neotropi
he text begins with an introduction to > Pund Island: the geography, climate, sociology, and
history of the region, including details on early exploration of the island, botanical and otherwise.
The second chapter ac dresses a ee of me a we is pany matorral, or thorny
deciduous forest. Seven pri salt marshes,
I
savannas, thorn scrubs, and ce sees with principal plants named for uk community Phe thorn
ices the most erates ny on as ae is sie described with quantitative structural
l-hectare plot. Trees greater
than 5 cm DBH Gismieres at breast- ee were identified and measured. The plot was found to
contain half the tree species on the entire island, with the families Bombacaceae and Bignoniaceae
being most abundant. A six-year study of forest dynamics and phenology is discussed.
e third ees summarizes the flora of Puna Island, which shows floristic similarities to
=
the re
ter of plant endemism—the area west of ae us between southern Colombia and northern ons
es and to the Galapagos Islands. This region encompasses what is known as the Guayas ce?
The native and naturalized fora of the is rises 428 species of vascular plants; of this, 15 %
of the plant species are endemic to the Guayas center. As in many floras throughout the world, the
most specious families are the Fabacese, Poaceae, and Asteraceae. No species are endemic to the is-
x
land itself, though rare and unusual plants are found there. nee 29% of the species on Puna
Island are shared with the Galapagos Islands, nearly 800 km to the west. At least 14% of the species
on Puna Island were introduced directly or indirectly by man and his domesticated animals. The
slants used by the people on Puna Island are discussed in the fourth chapter. Archaeological evi-
1
dence is ae for early agricultural crops (maize, cassava, pumpkin, sweet potatoes, peppers, and
cotton) < wellas early
I)
period ene introductions a O
is given concerning the use of many species ie lumber in shipbuilding over the past few centuries.
p, guava) in use in Pre-Columbian times. The early colonial
d World 5 li melon, and sugar cane. Much detail
—
An ethnobotanical and linguistic study of modern plant use and plant names on Puna Island fills
the rest ue bas ee with lists and discussions of edible plants, poisonous plants, useful woods,
al has dyes and soaps
T he bulk af this text is filled with keys and descriptions of all the species in the flora. This is
organized by family, genus, and species. Descriptions are brief and include human uses, vernacular
names, and information on nativity and distribution, as well as voucher specimen citations. Plants
are not euleaoise oe is no PeneGis alist: as such, but there is an index in which the alone atin
well spaced and browseable
rT
and historic people follows the plant index. A central section of the book groups the color photo
interest. The endpapers of the
plates of various habitats and a few plant species of ethnobotanica
book display colored maps of Puna Island, one combining physical, political, and natural features,
—
and one defining major plant communities. This book successfully builds upon the standard flora
SIDA 20(4): 1786. 2003
BOOK REVIEWS 1787
oem ee. oe historical and ethnogeographical data. This book will be of interest to any who
Columbian human occupation of coastal Ecuador, neotropical ethnobotany,
and early South oan agricultural practices. BlOweve er, if the reader is comloxtable SIE Cunt
]
q £ bea!
cal — without iustialions, T nel ion of Pund Isl Island will prima rily be a ve ry
can be used to identily pl ants anywhere in the coastal region of Ecuador—A manda K. Neill, Botani-
cal sa Institute of Texas, 509 Pecan St, Fort Worth, TX 76102-4060, U.S.A, aneill@brit.org.
Bertil NORDENSTAM, GAMAL EL-GHAZALY, and MonHaAMeD Kassas (eds). 2000. Plant
Systematics for the 21st Century: Proceedings from a symposium held at
the Wenner-Gren Centre, Stockholm, in September 1998. (ISBN 1-85578-
135-2, hbk.). Wenner-Green International Series, Vol. 77. Portland Press Ltd.,
59 Portland Place, London WI1B 1QW, U.K. (Orders: +44-020-7580-5530,
editorial@portlandpress.com, North American orders to: Princeton Univ.
Press, 41 William St. Princeton, NJ 08540, U.S.A.). $120.00, 376 pp., numer-
ous b&w drawings, charts, and halftones, 6 3/4" x 91/2"
The august title, * Plant systematics londas 21° ony may capture many purchasers for this book.
If so, the readers wil Be omy eisappoinicg with their econ. oe is not to eay there are not
£3} -| ] 11
But g 1 papers do not
fulfill the promise of ‘the title.
In aria this is a festschrift for two prominent and influential Swedish botanists, Vivi Lau-
rent-Té Edrtman, wae would have been 100 years old at the time of the sympo-
= Notice tl found in the mnupeluction:
nchoosing the themes of th posi I g view of the fields of these
ct ] ] 1 T Pe GS Parad re :
two sla namely plant
to)
berin ibutions of Tackholm and aedwnaie we also wished tos i t knowl-
gth
edge in ee fields. We also aimed to baelude'n new inicemaHion on some related and expanding re-
search areas of great sof anatomy and floral odours.”
Five Bapene ate ¢ biographical As commnemor tions e the eon nLpUEORS of Erdtman ane
Tackholm
ies in the flora of Egypt. Sie) more papers deal sane current ae in plant systematics includin
limits to computer analysis and nomenclatural challenges. Four papers examine new techniques
that have been applied to problems in particular families or orders. Finally the last paper records the
panel discussion used to wrap up the symposium
of the biological papers are case studies of the application of new methods to plant sys-
tematics, even the overview papers. A good example is Prance’s paper on the changing focus of re-
search on the Chrysobalanaceae over the last 40 years. Even Kallersjé and Farris’ paper serves pri-
marily to highlight the contribution of Farris’ computer algorithm on parsimony jackknifing.
Probably the best overviews of research advances are El-Ghazaly’s paper on new methods in pa-
lynology, Ferguson's paper on application of pollen morphology in systematics, and Chanda’s com-
ments on ie ae tion of ees ology to ee related fields.
W ll the biological papers are excellent summaries of new methods and application of
those pee ee over 6 nee decades of the oF century, only two are mul ae Pong
Rubieeits eoueabusion reviews ethes! state op |
oe : | ] : qd
and is concerned
s a Se ee Sa J
] i | | 1] : yO f
for the future of ple
taxa as global conservation becomes an increasin 19 important goal for biologists.
g gly g
SIDA 20(4): 1787. 2003
1788 BRIT.ORG/SIDA 20(4)
The panel discussion disappoints one looking for the future prospects of plant systematics. It
comes as no surprise that nearly all the speakers agreed that the top three priorities should be 1)
documentation of the world’s biodiversity before it is irreversibly lost, 2) conservation of that
biodiversity, and 3) continued research to better lerstand evolutionary mechanisms. Some of the
speakers mentioned increasing and improving herbarium and biological databases and retrieval.
However, no room was given to developing models on how the greatly expanded databasing shoulc
be achieved. Indeed the only other reference to this important issue was a couple of paragraphs in
Kubitzki’s paper.
Surprisingly, | found very little on the next phase tor molecular biology. That is, moving froma
simplistic comparisons of nucleotide sequences in a few analyzable genes to understanding the un-
derlying genetics of integrated character complexes of expressed - shenoty aes me in euan
assessment I found just the opposite, “To this end [of | |
should ee be based lecul aa
=)
or other traits can be mapped onto cladograms, whicl
While | understand his concern of avoiding circular reasoning by using alternate data sets, | see a
problematic attitude that is developing in systematics, even among the traditional morphologically
oriented researchers. Are we (as a biosystematics community) to form our classifications on one ora
few genes and force the interpretation of morphological characters, many of which derive from highly
correlated multi-gene complexes, onto those simple gene trees? Are we falling headlong again into
the trap of “one character taxonomy?” Perhaps we should be more prudent, realizing that these highly
sophisticated, but philosophically simplistic methods are not showin us phylogenetic history any
more an do the confusingly Basa morphological methods
<=
==
e this book falls short of its purported goal, nonetheless it is a valuable collection of ar-
ticles. It ‘ worth owning, ae for accessing the literature on Be a Egyptian botany, no-
menclature, and various modern systematic techniques.—Roger W. Sanders, Associate Collections
Manager, Botanical Research Institute of Texas, 509 Pecan St, Fort Worth, I “X 76102-4060, U.S.A.,
rnd @bhritor
&
Mark Bonta. 2003. Seven Names for the Bellbird: Conservation Geography in
Honduras. (ISBN 1-58544-249-6, hbk.). Texas ASM University Press, John
H. Lindsey Bldg., Lewis St. 4354 TAMU, College Station, TX 77843-4354,
U.S.A. (Orders: 1800-826-8911, w-lawrence@tamu.edu, www.tamuedu/
upress). $35.00, 250 pp., 35 b&w photos, 4 maps, 5 3/4" x 91/4",
Within the text, Mark Bonta takes an etl hi ypre oach to understan ling the relationship be
tween the local people of Honduras and a birds of their region, thus serv ing to create a bridge
between academic ene aphy and what he calls “on-the-ground conservation.” The recounting of
this approach and its results takes the form of individual and family histories and stories relayed
throughout the text. The particular intent of the text is to dissuade the neaaee thal dene: are en-
tirely destructive with a conscious purpose, and that in particular the rele ndu-
ran locals and their avifauna is not inherently detrimental to the ae Pour , Bonta see not
parade his findinese: as assurance that the birds have a secure place in the Sey landscape.
The text is divided into nine separate chapters, preceded by an introduction to conservation
geography, as it will be dealt with for the remainder of the text, and succeeded by an appendix list-
ing the birds recorded for the central Olancho department of Honduras as well asa glossary of Span-
ish terms. The first chapter delves into the subject of “ornithophilia” (a natural love for birds) through
the depiction of a young girl, Lucita, and her interaction with the birds in her area. Asa standard for
SIDA 20(4): 1788. 2003
BOOK REVIEWS 1789
chapters dealing with personal histories, Bonta further divides each into sections, the first few serving
as een examples of what in the latter sections he takes apart and reflects on in his own voice.
papier two estan tones the geogepince! and historical background of ane region and its bird
t a I varying
populz ation g the most in
| | that Honduras is a male-dominated society
relationships es birds CI papte
and the f relationships with birds are not the only ones; women and children
are oe voices to share their perceptions of the issue. Chapter four lays bare the situation within
Juticalpa, an urban center in Olancho, using case studies of two birds, the zopilote (Black Vulture)
the lla (cl ay colored Robin). Chapter five brings a the capacity in which large landowners
s, contrary to general expectation. Chapter six presents the other side of chapter
ae pignienting the Eure landscape of small domestic and fae ual villages and farmers, where
} | |
biodiversity is hig be most easily conserved. se aa seven exposes} mOnaneS as acoun-
try where the maou) of soil cover is not rainforest, and where many
her aed Le are also
~
of conser make their homesi f Ot
dealt with, such as those that are nocturnal. Guns eight is a case study centered about the
tane rainforest of Montana de Babilonia. In chapter nine Bonta presents several proposals sing
the conservation geography of Honduran avitauna, with the intent for broader applications
a message which Bonta intends be taken home is that the destruction in the
— is not accomplished with clear ee purposef ful intent, but rather asa side effect of certain
socio-economic factors. Therefore, to lessen its impact, we must take a “geographical approach to
conservation that cuts across artificial boundaries separating what is ‘natural’ from what is ‘cul-
tural’ in the landscape” (p.3). While the theme of conservation geography is implied throughout the
book, it is only in the last chapter where it is dealt with directly, and where it ceases to merely be an
“ethnography of birds.” The book does not resort to scientific and technical terms and therefore is
easily understood; it is for a general audience interested in avifauna, conservation, and the idea of a
“cultural landscape.’—Tiana Franklin, Botanical Research Institute of Texas, 509 Pecan St, Fort Worth,
TX 76102-4060, U.S.A, tfranklin@britore.
Paut E. Minnis (Ed.). 2003. People and Plants in Ancient Eastern North America.
(ISBN 1-58834-133-X, pbk.). Smithsonian Institution Press, 750 Ninth Street
NW, Suite 4300, Washington, DC 20560-0950, U.S.A. (Orders: 202-275-2300,
http://wwwsipress.siedu/). $34.95, 423 pp., 25 figs. 45 tables, 6" x 0"
This text is presented as the first in a two-volume set, consisting of eight chapters by contributing
North American anthropologists. Minnis chooses to extend the time frame of ethnobotany to the
past, and therefore deal with methods of paleoethnobotany. Using these methods, the contributing
authors delve into three subject areas: prehistoric use of plants, crop history and applications, and
human impacts on local environments. This first volume is loosely contained by geographical bound-
aries into, as the title indicates, an overview of Eastern North America; its fellow, as yet unpublished,
will cover the Serie Western North American region.
age introduction, Minnis provides an historical overview of the field of
PS eaoci ie quite easily explained for those of us that may have little use for such long-strung
terms. Following this are chapters that concentrate on the Central and Southern Woodlands (about
which much work has been done in the past), native plant use, crop domestication and food produc-
tion, anthropogenic eae and regional concentrations in the Northeast, the Great Plains, and in
the Caribbean Islands
SIDA 20(4): 1789. 2003
1790 BRIT.ORG/SIDA 20(4)
While it may be implied that the only data set for the conclusions drawn in the text consists of
micro- and macro-botanical remains, this is not entirely the case. In the foreword, Minnis clearly
establishes that along with botanical remains, such things as ancient field distribution and plant
images in art can be used to infer ancient ethnobotanical relationships. The contributing authors
present their portions particularly well although, having worked with botanists but not belonging
to the discipline themselves, tend to generalize and lump various evidences unto one name, whereas
in botanical nomenclature, ‘fruit’ would be inadequate and demand further explanation.
This compilation reads surprisingly well and holds interesting and significant information for
the concerned environmentally-minded reader, bringing attention to issues such as the fact that the
yuman food supply is derived from a play of only 20 crops, whereas paleoethnobotanical
—
majority of
evidence uncovers a suite of tens of thousands of crops cultivated in the past.—Tiana Franklin, Bo-
tanical Research Institute of Texas, 509 Pecan St, Fort Worth, TX 76102-4060, U.S.A, tfranklin@britorg.
SIDA 20(4): 1790. 2003
BIENNIAL REPORT AND UPDATE 2002-2003
SIDA, CONTRIBUTIONS TO BOTANY
4 ] : ] | ] 7} f ]
L 1
Source of ] world.
... bringing out the best in botanical science for plant conservation and eda cations
To all authors, we thank you for choosing Sida, Contributions to Botany as your
forum to disseminate information and knowledge gained by scientific inquiry.
Volume 20 represents the largest volume of Sipa ever published. Siva is pub-
lished twice a year with each issue seemingly increasing in size and in diver-
sity of quality papers. Volume 20 has 1836 pages, 202 authors, 165 published
contributions, and includes 229 new names and new combinations. Published
papers are available online in Adobe Acrobat format (PDF files); the PDF files
are true representations of the hard copy of Sipa. <http://www.brit.org/sida/
SCBCurlssue.htm>.
A word of thanks to the 228 individuals acknowledged on pages 1793-1794,
who generously supported Sipa through their time and expertise in reviewing
approximately 188 manuscripts submitted for volume 20. All manuscript sub-
missions are peer-reviewed by distinguished reviewers; your support is deeply
appreciated. If by chance you reviewed a manuscript and your name was left
out, the error rests solely on the shoulders of this editor and I truly apologize.
Please bring any omissions to my attention.
Below are the dates of publication for each of the four issues of volume 20,
subscriptions for calendar year 2004, and distribution of Sipa. We also bring to
your attention (see page 1792) the Mary M. Hennen Scientific Publications En-
dowment, established to provide support to the future of BRIT’s scientific publi-
cation program (Sida, Contributions to Botany and Sida, Botanical Miscellany).
The index to volume 20 (2002-2003) follows and the issue concludes with a
printed copy of the current guidelines to contributors. The online version is at
http://www.brit.org/sida/SubmitPaperhtm.
We thank all authors, reviewers, subscribers (individuals, institutions, or-
ganizations), and readers for your continued interest and support. It is our plan
to continue bringing you the best sources of current research in classical and
modern systematics with your continued support. Wishing you the best in 2004
and beyond.—Barney Lipscomb (BRIT), Editor; John W. Thieret (NKU), Associ-
ate Editor; and Félix Llamas (LEB), Contributing Spanish Editor.
Dates of publication
20(1), pp. 1-460: 19 July 2002
20(2), pp. 461-876: 23 November 2002 (40-year anniversary issue)
20(3), pp. 877-1341: 29 July 2003
20(4), pp. 1343-1836: 22 December 2003
SIDA 20(4): 1791. 2003
1792 BRIT.ORG/SIDA 20(4)
Subscriptions and distribution. —SipA is distributed in over 90 countries. There
are 900 subscriptions, 377 domestic and 523 foreign. Brazil has the most sub-
scribers (39) outside of the U.S.A. Sipa is available either through paid subscrip-
tion or scholarly exchange. Online information is available at http://
www.britorg/sida/subscription.htm.
Annual subscription rates for Sida, Contributions to Botany for 2004.
Domestic (USA only)
L)Subscribing institutions (universities, libraries, and other institutes
including agents)$60.00
2)Individual personal subscriptions$35.00
Outside the USA
subscribing institutions (universities, libraries, and other institutes
including agents)$70.00
ual personal subscriptions$35.00
2)Indivic
Mary M. Hennen Scientific Publications Endowment Established.—In honor of
his wile, Mary, BRIT research associate, professor emeritus of botany at Purdue
University, the world’s pre-eminent authority on the systematics and biogeog-
raphy of the rust fungi, Joe Hennen has established the Mary M. Hennen Scien-
tific Publications Endowment to provide support to the future of BRIT’s scien-
tific publication program (Sida, Contributions to Botany and Sida, Botanical
Miscellany). Joe and Mary (life-science librarian at Purdue University) have long
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mation about the BRIT Press or anyone interested in making a contribution to
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—
SIDA 20(4): 1792. 2003
INDEX
REVIEWERS: VOLUME 20 (2002-2003)
228 reviewers: several individuals reviewed more than one manuscript
Thank you for supporting Sida, Contributions to Botany.
Allen, Charles
Anderson, John L.
Anderson, Loran C.
rias, Salvador
Austin, Daniel F
mard C., Gerardo
Bain, John
Barkley, Theodore M.
Barnes, Richar
Barrie, Fred ‘i
Bauml, James A.
Berendsohn, Walter G.
Borchsenius, Finn
Bradford, Jason
Braun, Uwe
Brenner, David
Brouillet, Luc
Bryson, Charles T.
Burks, Kathleen
Canne-Hilliker, Judith M.
Cantino, Philip D.
Christensen, Knud Ib
Clary, Karen H.
Clayton, W.D.
Coile, Nancy C
Coons, oe) Patricia
Craven, Lyn A.
Croat, ae B.
Cusick, Allison W.
Davenport, LJ.
Davidse, Gerrit
Davila, Patricia D.
Petra
Delprete, Piero G.
e Vore, Melanie
Diamond, Alvin
Diggs, Jr, George M.
SIDA 20(4): 1793. 2003
Dorr, Laurence J.
Douglas, Andrew
Doyle, Robert
Drapalik, Donald J.
gh,
Endress, Mary E.
Engle, David M.
Faden, Robert B.
Farrar, Donald R.
Felger, Richard S
Ferguson, oe aay
Freeman, Craig C.
Fryxell, Paul A.
Gandhi, Kanchi N.
Gardner, Donald E.
Gehlbach, Fred
Gentry, Johnnie L.
George, Robert
Gereau, Roy
Gonzalez Elizondo
Socorro
Graham, Shirley A
Grant, Verne
Hammel, Barry E.
Hanlin, Richard T.
Daniel K.
Harris, Elizabeth M.
Hartman, Ronald L.
Hatch, Stephan L.
Haynes, Robes R.
Hill, Steven R
Hoagland, Bruce
Hodgson, Wendy C.
Eckenwalder, James E.
Egegli, Urs
Holmes, Walter C.
Horton, Diana
Howard, Ric oid A.
Howcrolt, N.HLS.
Hyatt, Philip E.
Jacono, Colette C.
Janni, Kevi
Jenkins, Fhilip D.
Johnson, Dale E.
Johnson, George P.
ones, Thomas H
Judd, Walter S.
Kaul, Robert B.
Keith, Eric L.
oe Joseph H.
Kral, R
Krings ie
Kron, Kathleen A
Lamont, Eric
Landry, Garrie
Lasseigne, Alex A.
ickey, Edgar B.
Liogier, Alain
Little, Damon P.
Llamas, Félix
MacDonald, Idu
MacDonald, John R.
Macklin, James A.
MacRoberts, Michael H
Middleton, David J.
Miller, James S.
1793
Miller, R
ci. William
Morales, J. Francisco
Nare- dora Amy
Nee
Neill, yee
Neill, David A.
Nelson, Allan D.
Nelson, John B.
Olesen, Bob
Omerod, Paul
Orzell, Steve L
Peck, James H.
Pedersen, Henrik
A
Peirson, Jess A.
Perkins, Kent D.
Persson, Claes
Peterson, Paul M.
Pipoly, II, John J.
Porter, J. Mark
Poston, Muriel
Prather, Alan
Rabeler, Richard
Ratter, James
Raz, Lauren
Reed, Monique Dubrule
Regalado, Jr, Jacinto C.
Rettig, Je
Reveal, James L
Rios, ona
Robart, Bruce
Rogers, oe 5)
Ryburn, Adam K.
Sanders, Roger
Sasidharan, N
Saunders, Richard
1794
Sc snare Andre
chultz, Joanna L.
Semple, John C.
Siedo, Stephen J.
Simpson, Beryl B.
Singhurst, Jason
Skean, Jr, James D.
Slaughter, Jr, Cecil R.
Smith, Alan R
Smith, Latimore
Snow, Neil
Solomon, James C.
Soltis, — E.
Sorensen, Pau
Spellenberg, oe
Stevens, Peter F.
SIDA 20(4): 1794. 2003
Stoynoff, Nick
Strother, John L.
Struwe, Lena
Stues
sen Scott
Sundell,
a
cae David M.
Sutton, David L.
Telfair Il, Ray C.
Thiede, Joachim
Thieret, John W.
Thomas, R. Dale
Thomas, William Wayt
nny Ralph L.
|
hrelkeld, Steven J.
Trock, Debra
Turner, B.L.
Ulloa U., M. Carmen
Umber, Ray
Utech, Fred
van Kley, James E.
Verhoek, Susan E.
Vincent, Michael
Walter, Richard
Weakley, Alan S.
Webster, Robert D.
Weec Ronald R.
Welsh, Stan
Wie
i Ronald
BRIT.ORG/SIDA 20(4)
Wiersema, John H.
Wilbur, Robert L.
Woodr a Lindsay
Wooc ichael
aren Richard D,
Wunderlin, Richard P.
Yatskievych, George A.
Zarucchi, James L.
Zomlefer, Wendy B.
Zona, Scott A.
INDEX
1795
INDEX TO 165 TITLES WITH 202 AUTHORS:
VoLumE 20 (2002-2003)
We know you have a choice. Thank you for choosing Sida, Contributions to
Botany.
Acommunity-level floristic reconnaissance
of the Raja Ampat Islands in New Guinea
by W. TAKEUCHI—20(3):1093
A cypress (Cupressus
Cupressaceae) in Jeff Davis County,
Texas? by M. Patrick GRIFFITH and STEPHANIE
C. BarteL—20(2):585
A floristic survey of Big Branch Marsh Na-
tional Wildlife Refuge, St. Tammany Par-
ish, Louisiana by Davio J. Rosen and Stan-
LEY D. Jones—20(3):1189
A new section of Baccharis (Asteraceae:
Astereae), and notes on allied taxa by
DanieL A. GiuLIANO and Guy L. Nesom—
20(4):1481
new species of Arida (Machaeranthera
sect. Arida—Asteraceae: Astereae) from
Trans-Pecos, Texas by B.L. Turner and Guy
L. Nesom—20(4):1417
new species of Brazoria (Lamiaceae)
from the central mineral region of Texas
by Mart W. TurNer—20(4):1565
A new species of Chaptalia (Asteraceae:
Mutiseae) from Mexico and rediscovery
of Chaptalia mexicana by Leticia CagrerA
R. and Guy L. Nesom—20(4):1363
new species of Dioscorea (Dioscore-
aceae) from the Andes of Venezuela by
L.J. Dorr and Basit Stercios—20(3):1007
A new species of Eriogonum (Polygona-
ceae) from south Texas by Guy L.Nesom—
20(1):31
new species of Gonolobus (Apocyna-
ceae: Asclepiadeae, Gonolobinae) from
southern Costa Rica by ALEXANDER KRINGS
—20(1):105
arizonica,
>
>
>
>
SIDA 20(4): 1795. 2003
A new species of Lithospermum (Bora-
ginaceae) from Nuevo Ledn, México by
B.L. TURNER—20(2):501
Anew species of Medinilla (Melastomata-
ceae) from Anamalai Hills, South India by
N. SASIDHARAN and P. SUJANAPAL—20(1):109
new species of Packera (Asteraceae:
Senecioneae) from the Edwards Plateau
of Texas by Desra K. Trock and Rosert J.
O'KENNON—20(3):945
A new species of Schistocarpha (Astera-
ceae: Heliantheae) from northwestern
Oaxaca and closely adjacent Guerrero,
México by B.L. TurNeR—20(2):505
A new species of Senecio (Asteraceae)
from Durango, Mexico by B.L. TurNeR—
20(1):175
A new species of Solidago (Asteraceae:
Astereae) from north central Alabama by
Brian R. Keener and Rosert KrRat—
20(4):1589
Anew species of Stylogyne (Myrsinaceae)
from Darién, Panama by Jon M. RickeTson
and JoHNn J. Piroty III—20(3):919
A new variety of Ipomoea costellata
(Convolvulaceae) from the Edwards Pla-
teau region of Texas by Rosert J.O’KENNON
and Guy L. Nesom—20(1):39
A revision of the Psidium salutare com-
plex (Myrtaceae) by Leste R. LawoRUM—
20(4):1449
rockhouse microhabitat in the West
Cross Timbers of North Central Texas by
Georcet M. Dicas, Jr.and Rosert J.O’KENNON
—20(3):1327
>
>
1796
A significant new population of the rare
semaphore pricklypear cactus, Opuntia
corallicola (Cactaceae) by KritH A. Brap-
Ley and Steven W.Wooomanste—20(2):809
A synopsis and nomenclator of the genus
Parathesis (Myrsinaceae) by Jon M.
RickETSON and JOHN J. Piroty III-——20(3):913
A synopsis of Machaeranthera (Astera-
ceae: Astereae), with recognition of seg-
regate genera by Davin R. Morcan and
RONALD L. HARTMAN—20(4):1 387
Addition to Flora Mesoamericana: a new
record of Thelypteris (Thelypterida-
ceae) for Chiapas, Mexico by Micuet A.
Perez FARRERA, RAMON Riga, and Maria E,
Lopez-Motna—20(3):1311
Additions to the Cyperaceae and Junca-
ceae flora of the Rio Mayo region, Sonora,
Mexico by Eric H. Roatson, THomas R. VAN
Devenper, and Ana LitiA REINA G.—
20(1):339
Additions to the flora of Cleveland County,
Arkansas: collections from Moro Bot-
toms Natural Area, a state-protected old-
growth bottomland hardwood forest by
Danny Skouac, Marcaret S. Devatt, and Ber-
NARD R. PARRESOL—20(4):1 731
Additions to the flora of Nags Head Woods
(Dare County, North Carolina) and the
outer banks of North Carolina by Atex-
ANDER KRINGS—20(2):839
Agave ovatifolia (Agavaceae), una nueva
especie de maguey del noreste de
México by Gree Starr and José A. VILLARREAL
Q.—20(2):495
Agrostis avenacea (Poaceae: Pooideae):
first record for the Mexican flora by Arey
Nava-Rojo, MariceLA GOmez-SANCHEZ, and
Manuel GONZALEZ-LEDESMA—20(1):423
An update to the vascular flora of Calcasieu
SIDA 20(4): 1796. 2003
BRIT.ORG/SIDA 20(4)
Parish, Louisiana by Ray Neytano—
20(1):431
Anatomia comparada de la epidermis fo-
liar en el grupo de especies Solanum
sessile, de la seccidn Geminata (Solan-
aceae) by Carmen Benitez pe Rosas and
CHIARA BERLINGIERI—20(4):165 1
Andropogon glomeratus var.glaucopsis
(Poaceae: Andropogoneae) docu-
mented in Louisiana by Davio J. Rosen and
Joseph K. Wiper —20(4):1723
Annotated checklist of the vascular flora of
the Hickory Creek Unit of the Big Thicket
National Preserve, Tyler County, Texas by
BarBARA R. MacRoserts, MicHaet H.
MacRoserts, and Larry
20(2):781
Another juniper (Cupressaceae) in Jeff
Davis County Texas? We're not con-
vinced by JoHN P. Karces and James C.
ZECH—20(3):1325
Arkansas flora: additions, reinstatements,
exclusions, and re-exclusions by James H.
Peck—20(4):1 737
Aspectos ecoldgicos y estado de conserva-
cidn de Mammillaria theresae (Cacta-
ceae) by IRMA Lorena Lopez-Enriquez, M.
MARTHA
GONZALEZ-ELIZONDO, aNd JorGE ALBERTO TENA-
FiLoreS—20(4):1665
Biological status of Hedyotis nigricans var.
gypsophila (Rubiaceae) in Texas by B.L.
TurRNegR—20(1):215
Biosystematic analysis of the Thelesperma
subnudum complex (Asteraceae) by
Curtis J. HANSEN, LOREEN ALLPHIN, and MicHael
D.WinDbHAM—20(1):71
Boerhavia coulteri var. palmeri, a new
varietal combination for Boerhavia
(Nyctaginaceae) of southwestern North
E. BrowNn—
Socorro GONZALEZ-ELIZONDO,
INDEX
America by RIcHARD SPELLENBERG—
20(1):151
Botanical summary of a lowland ultrabasic
flora in Papua New Guinea by W. Takeuchi
—20(4):1491
Carex conoidea (Cyperaceae) new to
Arkansas and notes on its occurrence
in Arkansas and Missouri by Paut M.
McKenzie, Tim Smith, and C. THeo WitsetL—
20(4):1727
Carlo Bertero (1789-1831) in the New
World by Piero G. DELprETE, GIULIANA FORNERIS,
and ANNALAURA PISTARINO—20(2):62 1
Castilleja albobarbata sp.
(Orobanchaceae) from Sierras Manant-
lan and Cacoma, Jalisco, Mexico by HucH
H. |uvis, Guy L. Nesom, and J. Mark Eacer—
20(4):1343
Celebration honors pioneer field botanist
André Michaux by Wittiam R. Burk—
20(2):861
Cirsium nuttallii (Asteraceae: Cynareae)
new to North Carolina and an illustrated
key to southeastern congeners by ALex-
ANDER Krincs, RANDY WeSTBROOKS, aNd JANINE
Lioyo—20(2):845
Commelina benghalensis (Commelina-
ceae) new to North Carolina and an up-
dated key to Carolina congeners by
ALEXANDER Krincs, MicHAEL G. BuRTON, and
ALAN C. York —20(1):419
Comparison of the morphology, flowering
NOV.
phenology, and life cycle type in plants
of Grindelia lanceolata (Asteraceae)
from cedar glades in middle Tennessee
and northern Alabama: a common gar-
den study by CuristopHer A. ADAMS, JERRY
M. Baskin, and Carol C. BASKIN—20(3):1059
Cryptocoryne beckettii complex
SIDA 20(4): 1797. 2003
1797
(Araceae) introduced at a Florida spring
by Cotette C. Jacono—20(2):819
Cynosurus echinatus (Poaceae) new to
Texas by R. Date THomas—20(2):837
Dahlia neglecta (Asteraceae: Coreopsi-
deae), a new species from Sierra Madre
Oriental, México by Dayte E. Saar—
20(2):593
Del Weniger (1923-1999), remarkable
teacher and naturalist by Marioric FOHN
EpLer—20(2):867
Documented chromosome numbers 2002:
1.Chromosome number of Stenanthium
(Liliales: Melanthiaceae) and its signifi-
cance in the taxonomy of tribe Melan-
thieae by Wenpy B. ZomLerer and Gerato L.
SmitH—20(1):221
Documented chromosome numbers 2002:
2. Chromosome numbers in North
American species of Cerastium (Caryo-
phyllaceae) by J.K. MortoN—20(1):227
Documented chromosome numbers 2002:
3. Chromosome counts for Packera
paupercula var. appalachiana (Astera-
ceae: Senecioneae) by Atison McKenzie
ManHoney, ViRGINIA W. Kone, and Roser R.
KowAl—20(2):605
Documented chromosome numbers 2003:
1. Chromosome number of Toxi-
coscordion nuttallii (Liliales: Melanthia-
ceae) and clarification of the genus by
Wenpy B. ZOMLEFER—20(3):1085
Documented chromosome numbers. 2003:
2. Chromosome numbers of the rare
Texas species, Arida blepharophylla
and A. mattturneri (Asteraceae: Astere-
ae) by A. MicHatt PowetL—20(4):1663
Does Polygonum patulum (Polygona-
ceae) grow in North America? by Minai
Costes and Francois J. TARDIF—20(4):1 707
1798
Effects of fire on two pitcher plant bogs
with comments on several rare and in-
teresting plants by Eric L.KeitH and N. Ross
Carrit—20(1):387
Endemism in the West Gulf Coastal Plain:
importance of xeric habitats by MicHAeL
H. MacRoserrs, BARBARA R. MacRoserts, BRUCE
A. Sorric, and Rosert E. Evans—20(2):767
Erratum—20(1):445
Euchiton (Asteraceae: Gnaphalieae) in
North America and Hawaii by Guy L.
NesomM—20(2):515
Fanonvril Mm +t (Dp
YYPY LUTTE
uler olygonaceae),
new for Texas by Rosert J. O'KENNON,
Caren McLemore, and Amanpa K. Newt—
20(4):1717
Festuca dentiflora (Poaceae: Loliinae:sect.
Glabricarpae),a new species from Peru
and taxonomic status of F. presliana by
DaNiet STANCIK and Pau M. Peterson—
20(3):1015
Flora and vegetation of the Mohawk
Dunes, Arizona by RicHarb STEPHEN FELGER,
Date Scott Turner, and MicHact F.Witson—
20(3):1155
Floristics of xeric sandylands in the post oak
savanna region of East Texas by Barsara
R. MacRoserts, MicHaeL H. MacRoeerts, and
James C. Cathey —20(1):373
Further additions to the genus Ardisia sub-
genus Graphardisia (Myrsinaceae) by
Jon M. Ricketson and JOHN J. Pipoty Il]—
20(4):1423
Glandularia verecunda (Verbenaceae):
species or fruit-form? by B.L. Tuaner—
20(4):1647
In memoriam: Chester Morrison Rowell, Jr,
(1925-2003) by Jim Biassincame—
20(4):1771
Ipomoea tacambarensis (Convolvula-
SIDA 20(4): 1798. 2003
BRIT.ORG/SIDA 20(4)
ceae), especie nueva del estado de
Michoacan, México by ELeazar CarrANza
GONZALEZ —20(4):1351
Key to the species of Ericaceae of Bolivia,
including two new species by James L.
Luteyn—20(1):1
Koenigia islandica (Polygonaceae) new
for Utah by Minai Costea and Francois J.
TARDIF—20(3):1319
Lectotypification of Mentzelia texana and
M. lindheimeri (Loasaceae) with an as-
sessment of their biological status by
Bituie L. TuRNeR—20(1):157
Lectotypification of Pseudognaphalium
biolettii (Gnaphalieae: Asteraceae) by
Guy L. Nesom—20(1):149
Lectotypification of Schlechtendal
Myrsinaceae from Schiede and Deppe
collections by Jon M. Ricketson and JOHN
J. Pirpoty III—20(3):909
Liatris gholsonii (Asteraceae: Eupa-
torieae), a new blazing star from the
Apalachicola River bluffs and ravines in
Florida by Loran C. ANDERSON—20(1):97
Listado preliminar de las Cactdceas y
Agavaceas de la Sierra“El Viejo,” Sonora,
region prioritaria para la conservacion
en México by Carlos G.VeLazco Macias and
GLariRO J. ALANIS FLORES—20(1):349
Muhlenbergia dubia (Poaceae) in central
Texas by Georrrey C. DENNY —20(4):1 763
Native flowering plant species new or oth-
erwise significant in Kentucky by Rosert
F.C.Naczi, RONALD L. Jones, F.JosePH METZMEIER,
Mark A. Gorton, and Timotuy J. WeckmAN—
20(1):397
Nephrolepis exaltata (Dryopteridaceae)
new to Arkansas by James H. Peck—
20(2):833
New and noteworthy orchids of the Bis-
INDEX
marck Archipelago, Papua New Guinea
by N.H.S. Howcrort and W. TakeucHi—
20(2):461
New combination in Astereae (Asteraceae
by Guy L. Nesom and Harotp Rosinson—
20(4):1649
New combination in Eurybia (Asteraceae:
Astereae) from North America by Luc
BrouiLLleT—20(4):1561
New combination in Salix (Salicaceae) by
Guy L. Nesom—20(2):523
New combination in Xylorhiza (Astera-
ceae: Astereae) by Guy L. Nesom—
20(1):145
New names and combinations in golden-
rods, Solidago (Asteraceae: Astereae) by
JOHN C. Semple —20(4):1605
New names and new combinations in the
flora of Hispaniola by Henri ALAIN Liocierk—
20(4):1645
New records in Asteraceae for Alabama
and Arkansas by Guy L. Nesom—
20(1):403
New records of Corchorus (Tiliaceae) for
Mexico by José Aurelio COLMENERO ROBLES
and Raract FERNANDEZ Nava—20(3):1299
New taxa of Crataegus (Rosaceae) from
the northern Okanagan-southwestern
Shuswap diversity center by J.B. PHipps
and R.J. O'KeNnNoN—20(1):115
New vascular plant records for Texas by Guy
L. Nesom and Rosert J. O'KENNON—
20(1):435
New vascular plant records for the grass-
land biome of central North America by
Craic C. FREEMAN, Cates A.Morse, and RONALD
L. McGrecor—20(3):1 289
Nomenclatural adjustments and com-
ments in Abronia and Acleisanthes
WS
SIDA 20(4): 1799. 2003
1799
(Nyctaginaceae) by RICHARD SPELLENBERG
and Jackie M. PooLte—20(3):885
Nomenclatural changes and innovations in
Panicum and Dichanthelium (Poaceae:
Paniceae) by Rosert W. FreckMANN and
MicHet G. LELONG-—20(1):161
Nomenclatural changes in the genus
Polygonum section Polygonum
(Polygonaceae) by Mikal Costea and
Francois J. TARDIF—20(3):987
Nomenclatural combinations in North
American Panicum subgenus Dichan-
thelium and subgenus Agrostoides
(Poaceae: Paniceae) by JoserH K. Wiprr
20(3):1043
Nomenclature of the heart-leaved hedge-
nettle, Stachys cordata (Lamiaceae) by
James S. PRINGLE—20(2):583
Northward range extension in Florida of
the invasive fern Lygodium micro-
phyllum (Lygodiaceae) by Rosert W.
PEMBERTON—20(4):1 759
Norwegian and Sami ethnobotany of
Veratrum album (Melanthiaceae) by
TorBJ@RN ALM—20(2):61 1
Notes on Carphephorus odoratissimus
(Asteraceae) in peninsular Florida, U.S.A.
by Steve L. Orzett and Eowin L. Brioces—
20(2):559
Notes on the flora of Texas with additions
and other significant records. Il by Larry
E. Brown and I. SANDRA Etsik—20(1):437
Notes on two species of Agalinis
(Scrophulariaceae) including Agalinis
harperi, new to Texas by Eric L. KeitH and
JoHN Hays—20(3):1321
Noteworthy vascular plants from Arkansas.
Il by Eric Sunoett, R. DALE THOMAS, CARL
Amason, and Curis DorFitr—20(1):409
1800
On the derivation of the generic name
lliamna (Malvaceae) by Wittiam A.Weser
and Paut A. FayxeiL—20(2):645
Oxypolis ternata (Apiaceae) deleted from
the Texas flora by Bruce A. Sorrig, MICHAEL
H. MacRoserts, Barsara R. MacRoserts, and
SUZANNE BIRMINGHAM WALKER—20(3):1323
Pappus variation in Solidago (Asteraceae:
Astereae) by Jennifer L.A. Hoop and JoHN
C. Sempte—20(4):1617
Perforation plate diversity in Mlicium
floridanum (Illiciaceae) with respect to
organs, provenance, and microtechnical
methods by Epwarp L. ScHNEIDER and
SHERWIN CARLQUIST—20(3):1047
Plants new to the United States and Florida
by RicHarp PW _ Bruce F. HANSEN, and
Loran C, ANDERSON—20(2):813
Polygonum aviculare subsp. rurivagum
(Polygonaceae) in North America by Mi-
HAI Costea and Francois J. TARDIF—
20(4):1 709
Polygonum douglasii subsp. johnstonii
(Polygonaceae) new for Canada by
Minal Costea and Francois J. TARDIF—
20(3):1317
Portulaca matthewsii (Portulacaceae), a
new species from the “Sierra Gorda” Bio-
sphere Reserve, Querétaro, Mexico by
GILBERTO Ocampo AcostA—20(4):1357
Pteroceras monsooniae (Orchidaceae), a
new species from India by N. SASIDHARAN
and P. SUJANAPAL —20(3):923
Rediscovery of Tephrosia jamnagarensis
(Fabaceae), an endangered and narrow
endemic plant species of Saurashtra,
Gujarat, India by PS. Naaar, S.J. Sata, and
S.J. PATHAK—20(4):1 701
Reinstatement of Viburnum ozarkense
lemic taxon of the
(Caprifoliaceae):an
SIDA 20(4): 1800. 2003
BRIT.ORG/SIDA 20(4)
Interior Highlands of Arkansas, Missouri
and Oklahoma by TimotHy J. Weckman—
20(2):849
Resource allocation patterns and pheno-
typic variation in the endangered Texas
wildrice (Zizania texana, Poaceae) by
PauLA Power—20(2):571
Revision and typification of some species
of Ixora (Rubiaceae) from Central and
Southern Brazil by Piero G. Detprere—
20(4):1471
Rotala rotundifolia (Lythraceae) new to
Florida by KatHieen C. Burks, VERNON V.
VANDIVER, Jr., Davio W. HALL, and Cotette C.
Jacono—20(4):1765
Rupert Barneby Award—20(1):446
Rust fungi (Uredinales) of Northwest Ar-
gentina by José R. HERNANDEZ and Joe F.
HenneN—20(1):313
Seed and fruit characters in selected
Spermacoceae and comparison with
Hedyotideae (Rubiaceae) by Enwarp E.
TERRELL and RicHArD P. WuUNDERLIN-
20(2):549
Senecio spribillei (Asteraceae: Seneci-
oneae), a new species from Montana,
U.S.A. by Witiiam A.Weser—20(2):51 1
Sinopsis del género Meliosma (Sabiaceae)
en Costa Rica y Panama, con tres nuevas
especies by J. Francisco Morates—
20(3):93 1
Six non-native species newly discovered in
the lowa vascular flora by ALLIson W.
Cusick —20(1):405
Soil-dependent fire frequency: a new hy-
pothesis for the distribution of prairies
and oak woodlands/savannas in North
Central and East Texas by Grorat M. Diccs,
Jrcand Perer C. ScHuLze—20(3):1139
Solidago faucibus (Asteraceae: Astereae),
INDEX
anew mesic forest goldenrod from the
Appalachian Mountains by THomas F.
Wiesotot and JOHN C. SempLe—20(4):1595
+
I p var. williamsiae,
var.nov. (Malvaceae) by Noet H. Houmaren
—20(1):47
Studies in Glandularia (Verbenaceae): the
Glandularia quadrangulata complex
by James HenrickKSoN—20(3):953
Symplocos kothayarensis (Symploca-
ceae),a new species from peninsular In-
dia by V. SUNDARESAN, S. RAJKUMAR, G.J. JOTHI,
and V.S. MANicKAM—20(3):927
Taxodiomeria (Taxodiaceae), an interge-
neric hybrid between Taxodium and
Cryptomeria from Shanghai, People’s
Republic of China by ZHANG JiAn-JUN, PAN
SHi-Hua, ZHU Wel-Jig, Niu Hui-Juan, Ye ZHENG-
Ji, ZHU Jian-Hua, and Hsu Pinc-SHenc—
20(3):999
Taxonomy of the most common weedy
European Echinochloa species (Poa-
ceae: Panicoideae) with special empha-
sis on characters of the lemma and cary-
opsis by Minai CosteA and Francois J.
TarbDiF—20(2):525
Taxonomy of the Polemoniaceae: the sub-
families and tribes by Verne Grant—
20(4):1371
The bracteoles in Amaranthus (Amaran-
thaceae): their morphology, structure,
function,and taxonomic significance by
Minat CosteA and Francois J. Tarbir—
20(3):969
The east-west transition of the flora in
Texas: a biogeographical analysis by
MicHaeL H. MacRoserts and Barsara R.
MacRoserts—20(4):1693
The flora of Oktibbeha County, Mississippi
SIDA 20(4): 1801. 2003
1801
by Anoreas Leiotr, SioNeyY McDaniet,and Tim
Nuttte—20(2):69 |
The genus Campomanesia (Myrtaceae) in
Atlantic rainforest fragments in Sergipe,
northeast region of Brazil by Myrna F.
Lanoim and Leste R. LANDRUM—20(1):205
The genus Packera (Asteraceae: Seneci-
oneae) in Colorado, U.S.A. by Desra K.
Trock—20(3):1023
The identity of eaglewood (Gyrinops,
Thymelaeaceae), a new economic re-
source for Papua New Guinea by W.
TakeucH! and M. Gotman—20(1):261
The John D. Freeman Herbarium (AUA): a
history and catalogue of vascular plant
type specimens by Curtis J. HANSEN—
20(3):1277
The name of the amaranth: histories of
meaning by Muai Costea and Francois J.
TArDIF—20(3):1073
The present status of Ledermann’s April
River localities in Papua New Guinea by
W. TakeucH| and M. GotmMaN—20(1):55
The resurrection and lectotypification of
eae
Miconieae): a hummingbird-pollinated
treelet endemic to Jamaica by Darin S.
Penneys and Watter S. Juoo—20(3):877
The taxonomy and morphology of
Macranthera flammea (Orobancha-
ceae) by Jit D. Atroro and Loran C.
ANDERSON—20(1):189
The varieties of Liatris elegans (Astera-
ceae) by Mark H. MayritLD—20(2):597
The vascular flora of the Cerrado in Emas
National Park (Goias, central Brazil) by
Marco ANTONIO BATALHA and FerNANDO Ros-
ERTO Martins—20(1):295
The vascular flora of the Laguna de Alegria,
1802
a national monument in the crater of
Tecapa Volcano, El Salvador by Denise
Wittiams, Rupys W. Herrera, and Jose L.
LINARES—20(2):675
The vascular flora of the Ogallala ecotone
on the Dempsey Divide, Roger Mills
County, Oklahoma by Craic C. FREEMAN
and Cates A. Morse —20(3):1217
Triglochin concinna (Juncaginaceae), a
new family, genus, and species for Texas
by B.L. Turner and Matt W. Turner—
20(4):1 721
Two new combinations in Bolivian
Gentianella (Gentianaceae) by James S.
~20(4):1485
Two new species of Festuca from South
America (Poaceae: Loliinae: sect.
Subulatae) by Daniel STANCIK AND PauL M.
PETERSON—20(1):21
as new PEGE? of Liatris series
+
PRINGLE
Eupatorieae)
from the ee United States by
Ropert Krav and Guy L.Nesom—20(4):15 73
Two new species of Parathesis (Myrsina-
ceae) from Mesoamerica and neotypifi-
cation of Parathesis rothschuhiana by
Jon M. RickeTtson and JoHn J. Pipoty |Il—
20(3):899
Two new species of Weinmannia
(Cunoniaceae: Cunonieae) from south-
ern Ecuador by Zachary S. Rocers—
20(1):179
Type localities of vascular plants first de-
scribed from Ohio by Ronatp L. Stuckey
and James S. PrincLte-—20(4):1677
Typification and nomenclatural history of
Trachelospermum difforme (Apo-
cynaceae)
20(4):1641
by ALEXANDER KrinGsS—
SIDA 20(4): 1802. 2003
BRIT.ORG/SIDA 20(4)
Typification, taxonomy, and a new subspe-
cies of Polygonum sawatchense
(Polygonaceae) from North America by
Mindat Costea and Francois J. Taroip-—
20(4):1631
Una especie nueva de Portulaca (Portula-
caceae) del norte de Michoacan (México)
by GiLBerTO Ocameo Acosta—20(2):487
Usos de Melastomataceae en el Ecuador
by ALINA Freire Fierro, DIANA FERNANDEZ, and
CATALINA QUINTANA—20(1):233
Uvularia (Liliaceae) in Texas by Jason R.
SINGHURST, WILLIAM F. Carr, Wenpy J. LEDBETTER,
and Watter C. Hoimes—20(4):1713
Vascular flora of Big Lake Bottom Wildlife
Management Area, Anderson County,
Texas by Kay M. FLemina, JASON R. SINGHURST,
and Watter C. Houmes—20(1):355
Vascular plants of Konza Prairie Biological
Station: an annotated checklist of spe-
cies in a Kansas tallgrass prairie by E. Gent
Towne—20(1):269
Vascular plants of the Mapimi Biosphere
Reserve, México: a checklist by Apet
Garcia-AREVALO—20(2):797
West Gulf Coastal Plain ecoregions by
MicHatL H. MacRoserts and Bareara R.
MacRoserts—20(3):1247
What is the writer of a flora to do? Evolu-
tionary taxonomy or phylogenetic sys-
tematics? by Georce M. Dicas, Jr. and
Barney L. Lipscomp—20(2):647
William Dean Reese (1928-2002) by RONALD
A. PursetL and Paut
20(3):1335
Yucca cernua (Agavaceae: series Rupi-
colae),a new species from Newton and
Jasper counties in eastern Texas by Eri
L. KeitH—20(3):891
L. REDOFEARN—
INDEX OF AUTHORS
1803
INDEX OF 202 AUTHORS: VOLUME 20 (2002-2003)
Thank you for choosing Sida, Contributions to Botany.
Adams, Christopher A—20(3):1059
Alanis Flores, Glafiro J—20(1):349
Alford, Jill D—20(1):189
Alm, Torbjarn—20(2):611
Allphin, Loreen—20(1):71
Amason, Carl—20(1):409
Anderson, Loran C.—20(1):97; 20(1):189;
20(2):813
Bartel, Stephanie C.—20(2):585
Baskin, Carol C —20(3):1059
Baskin, Jerry M—20(3):1059
Batalha, Marco Antonio—20(1):295
Benitez de Rojas, Carmen—20(4):1651
Berlingieri, Chiara—20(4):1651
Blassingame, Jim—20(4):1771
Bradley, Keith A—20(2):809
Bridges, Edwin L.—20(2):559
Brouillet, Luc—20(4):1561
Brown, Larry E—20(1):437; 20(2):781
Burk, William R—20(2):861
Burks, Kathleen C._—20(4):1765
Burton, Michael G—20(1):419
Cabrera R., Leticia—20(4):1363
Carlquist, Sherwin—20(3):1047
Carr, William F—20(4):1713
Carranza Gonzalez, Eleazar—20(4):1351
Carrie, N. Ross—20(1):387
Cathey, James C—20(1):373
Colmenero Robles, José Aurelio—
20(3):1 299
Costea, Mihai —20(2):525; 20(3):969;
20(3):987; 20(3):1073; 20(3):131 7;
20(3):1319; 20(4):1631; 20(4):1 707;
20(4):1 709
Cusick, Allison W.—20(1):405
Delprete, Piero G.—20(2):621; 20(4):1471
Denny, Geoffrey C—20(4):1 763
Devall, Margaret S—20(4):1 731
SIDA 20(4): 1803. 2003
Diggs, Jr, George M—20(2):647;20(3):1 139;
20(3):1327
Doffitt, Chris—20(1):409
Dorr, L.J.—20(3):1007
Egger, J. Mark—20(4):1343
Elsik, |. Sandra—20(1):437
Epler, Marjorie Fohn—20(2):86 7
Evans, Robert E—20(2):767
Felger, Richard Stephen—20(3):1 155
Fernandez, Diana—20(1):233
Fernandez Nava, Rafael—20(3):1299
Fleming, Kay M—20(1):355
Forneris, Giuliana—20(2):621
Freckmann, Robert W.—20(1):161
Freeman, Craig C—20(3):121 7; 20(3):1289
Freire-Fierro, Alina—20(1):233
Fryxell, Paul A—20(2):645
Garcia-Arévalo, Abel—20(2):797
Giuliano, Daniel A.—20(4):1481
Golman, M.—20(1):55; 20(1):261
Gdmez-Sanchez, Maricela—20(1):423
Gonzalez-Elizondo, M. Socorro—20(4):1665
Gonzalez-Elizondo, Martha—20(4):1665
Gonzalez-Ledesma, Manue|—20(1):423
Gorton, Mark A.—20(1):397
Grant, Verne—20(4):1371
Griffith, M. Patrick—20(2):585
Hall, David W.—20(4):1765
Hansen, Bruce F.—20(2):813
Hansen, Curtis J—20(1):71;20(3):1277
Hartman, Ronald L.—20(4):1387
Hays, Jonn—20(3):1321
Hennen, Joe F—20(1):313
Henrickson, James—20(3):953
Hernandez, José R.—20(1):313
Herrera, Rudys W.—20(2):675
Holmes, Walter C.—20(1):355; 20(4):1713
Holmgren, Noel H.—20(1):47
1804
Hood, Jennifer L.A.—20(4):1617
Howcroft, N.H.S.—20(2):461
Hsu, Ping-Sheng—20(3):999
IItis, Hugh H—20(4):1343
Jacono, Colette C.—20(2):8 19; 20(4):1765
Jones, Ronald L—20(1):397
Jones, Stanley D—20(3):1189
Jothi, G.J.—20(3):927
Judd, Walter S —20(3):877
Karges, John P—20(3):1325
Keener, Brian R—20(4):1589
Keith, Eric L——20(1):387;20(3):891:
20(3):1321
Kong, Virginia W.—20(2):605
Kowal, Robert R.—20(2):605
Kral, Robert—20(4):1573; 20(4):1589
Krings, Alexander—20(1):105;20(1):419;
20(2):839; 20(2):845; 20(4):1641
Landim, Myrna F—20(1):205
Landrum, Leslie R—20(1):205:20(4):1449
Ledbetter, Wendy J—20(4):1713
Leidolf, Andreas —20(2):69 |
Lelong, Michel G—20(1):161
Linares, José L.—20(2):675
Liogier, Henri Alain—20(4):1645
Lipscomb, Barney L.—20(2):647
Lopez-Enriquez, Irma Lorena—20(4):1665
Lopez-Molina, Maria E.—20(3):1311
Luteyn, James L.—20(1):1
Lloyd, Janine —20(2):845
MacRoberts, Barbara R—20(1):373:
20(2):/67; 20(2):781; 20(3):1 247:
20(3):1 323; 20(4):1693
MacRoberts, Michael H.—20(1):373;
20(2):/67; 20(2):781; 20(3):1 247:
20(3):1 323; 20(4):1693
Mahoney, Alison McKenzie—20(2):605
Manickam, V.S.—20(3):927
Martins, Fernando Roberto—20(1):295
Mayfield, Mark H—20(2):597
SIDA 20(4): 1804. 2003
BRIT.ORG/SIDA 20(4)
McDaniel, Sidney—20(2):691
McGregor, Ronald L.—20(3):1289
McKenzie, Paul M.—20(4):1 727
McLemore, Caren—20(4):1 717
Metzmeier, F. Joseph—20(1):397
Morales, J. Francisco—20(3):931
Morgan, David R.—20(4):1387
Morse, Caleb A.—20(3):1217;20(3):1289
Morton, J.K.—20(1):227
Naczi, Robert F.C —20(1):397
Nagar, PS.—20(4):1 701
Nava-Rojo, Arely—20(1):423
Neill, Amanda K.—20(4):1 717
Nesom, Guy L.—20(1):31;20(1):39:
20(1):145; 20(1):149; 20(1):403;:
20(1):435; 20(2):515; 20(2):523;
20(4):1 343; 20(4):1 363; 20(4):1417;
20(4):1 481; 20(4):1573; 20(4):1649
Neyland, Ray—20(1):43 1
Niu, Hui-Juan—20(3):999
Nuttle, Tim—20(2):691
O'Kennon, Robert J—20(1):39;20(1):1 15;
20(1):435: 20(3):945; 20(3):1 327;
20(4):1717
Ocampo Acosta, Gilberto—20(2
20(4):1357
Orzell, Steve L—20(2):559
Pan, Shi- Hua—20(3):999
Parresol, Bernard R.—20(4):1 731
Pathak, S.J—20(4):1 701
Peck, James H.—20(2):833; 20(4):1737
Pemberton, Robert W.—20(4):1 759
Penneys, Darin S.—20(3):877
Pérez Farrera, Miguel A—20(3):1311
Peterson, Paul M.—20(1):21;20(3):1015
Phipps, J.B —20(1):115
Pipoly Ill, John J—20(3):899; 20(3):909;
20(3):91 3; 20(3):919; 20(4):1423
Pistarino, Annalaura—20(2):62 |
Poole, Jackie M.—20(3):885
~—
487;
INDEX OF AUTHORS
Powell, A. Michael—20(4):1663
Power, Paula—20(2):571
Pringle, James S—20(2):583; 20(4):1485;
20(4):1677
Pursell, Ronald A—20(3):1335
Quintana, Catalina—20(1):233
Rajkumar, S.—20(3):927
Redfearn, Paul L.—20(3):1335
Reina G., Ana Lilia—20(1):339
Riba, Ramon—20(3):1311
Ricketson, Jon M.—20(3):909; 20(3):913;
20(3):919; 20(3):899; 20(4):1423
Roalson, Eric H—20(1):339
Robinson, Harold—20(4):1649
Rogers, Zachary S.—20(1):1 79
Rosen, David J —20(3):1 189; 20(4):1 723
Saar, Dayle E.—20(2):593
Sasidharan, N.—20(1):109; 20(3):923
Sata, S.J—20(4):1 701
Schneider, Edward L.—20(3):1047
Schulze, Peter C—20(3):1139
Semple, John C.—20(4):1595; 20(4):1605;
20(4):1617
Singhurst, Jason R.—20(1):355;
20(4):1713
Skojac, Danny—20(4):1 731
Smith, Gerald L—20(1):221
Smith, Tim—20(4):1727
Sorrie, Bruce A.—20(2):767; 20(3):1323
Spellenberg, Richard—20(1):151;20(3):885
Stancik, Daniel —20(1):21; 20(3):1015
Starr, Greg—20(2):495
Stergios, Basil—20(3):1007
Stuckey, Ronald L—20(4):1677
Sujanapal, P—20(1):109; 20(3):923
Sundaresan, V.—20(3):927
Sundell, Eric—20(1):409
Takeuchi, W.—20(1):55;20(1):261;
20(2):461; 20(3):1093; 20(4):1491
SIDA 20(4): 1805. 2003
1805
Tardif, Francois J—20(2):525; 20(3):969;
20(3):987; 20(3):1073; 20(3):131 7;
20(3):1319; 20(4):1631; 20(4):1 707;
20(4):1709
Tena-Flores, Jorge Alberto—20(4):1665
Terrell, Edward E.—20(2):549
Thomas, R. Dale—20(1):409; 20(2):837
Towne, E.Gene—20(1):269
Trock, Debra K—20(3):945; 20(3):1023
Turner, B.L—20(1):157;20(1):1 75;
20(1):215; 20(2):501; 20(2):505;
20(4):141 7; 20(4):1647; 20(4):1 721
Turner, Dale Scott—20(3):1155
Turner, Matt W.—20(4):1565; 20(4):1 721
Van Devender, Thomas R.—20(1):339
Vandiver, Jr., Vernon V.—20(4):1765
Velazco Macias, Carlos G.—20(1):349
Villarreal Q., José A—20(2):495
Walker, Suzanne Birmingham—20(3):1323
Weber, William A.—20(2):511;20(2):645
Weckman, Timothy J—20(1):397;
20(2):849
Westbrooks, Randy —20(2):845
Wieboldt, Thomas F—20(4):1595
Wilson, Michael F—20(3):1155
Williams, Denise —20(2):675
Windham, Michael D—20(1):71
Wipff, Joseph K.—20(3):1043; 20(4):1 723
Witsell, C. Theo—20(4):1 727
Woodmansee, Steven W.—20(2):809
Wunderlin, Richard P—20(2):549;:
20(2):813
Ye, Zheng-Ji—20(3):999
York, Alan C._—20(1):419
Zech, James C.—20(3):1325
Zhang, Jian-Jun—20(3):999
Zhu, Jian-Hua—20(3):999
Zhu, Wei-Jie—20(3):999
Zomlefer, Wendy B.—20(1):221;20(3):1085
1806
INDEX OF BOTANICAL NAMES
BRIT.ORG/SIDA 20(4)
: VOLUME 20 (2002-2003)
New names (229) in bold face
Abronia—20(3):877
umbellata var. breviflora—20(3):888
Aciotis—20(1):234
Acleisanthes—20(3):885
lanceolata var. megaphylla—
20(3):886
Agalinis
filicaulis—20(3):1321
harperi (Texas record) —20(3):1321
nuttallii (Arkansas record) —
(4):1739
Agarista boliviensis—20(1):4
Agavaceae—20(1):349; 20(2):495
havardiana—20(2):498
ovatifolia—20(2):495, 496 (illus),
497 (color photos)
parrasana—20(2):498
series Rupicolae—20(3):891
Agrostis avenacea—20(1):423, 424,425
(illus), 430 (map, color photo,
habitat)
Allium stellatum—20(1):441
Alstonia scholaris (USA/FL record) —
Alternanthera
caracasana (Kansas record) —
paronychioides—20(1):409
Amaranthaceae—20(3):969
Amaranth (history)—20(3):1073
Amaranthus—20(3):969; 1073 (history)
albus—20(3):976 (photo, anatomy)
blitoides—20(3):976 (photo,anatomy)
blitum
ssp. blitum—20(3):976 (photo,
anatomy)
ssp. emarginatus—20(3):976
(photo, anatomy)
SIDA 20(4): 1806. 20
ssp. oleraceus—20(3):976 (photo,
anatomy)
cruentus—20(3):975 (photo, anatomy)
hybridus—20(3):975 (photo,
anatomy)
hypochondriacus—20(3):975
(photo, anatomy)
powellii
ssp. bouchonii—20(3):975 (photo,
anatomy)
ssp. powelliim20(3):975 (photo,
anatomy)
retroflexus—20(3):975 (photo,
anatomy)
spinosus—20(3):977 (photo,
anatomy)
Anaphalis margaritacea—20(1):435
Andropogon
glomeratus var. glaucopsis—
20(4):1 723
ischaemum var. songaricus—
20(1):415
virginicus—20(4):1724
Anethum graveolens—20(1):405
Anodendron oblongifolium—20(1):59
(plant poison)
Apocynaceae—20(1):105
Aquilaria filaria—20(1):266
Araceae—20(2):819
Arachnis beccarii var. imthurnii—
20(2):466, 463, (illus), 468 (illus)
Ardisia
escallonioides—20(3):909
subgenus Graphardisia—
20(4):1423, 1424
nigrescens—20(4):1426
subsp. donnellsmithii—
20(4):1431, 1432 (illus)
INDEX OF BOTANICAL NAMES
subsp. nigrescens—20(4):1428,
1429 (illus)
tuerckheimii—20(4):1437, 1439 (illus)
verdisepala—20(4):1435, 1436
(illus)
Arida—20(4):1390, 1410
arizonica—20(4):1413
blepharophylla—20(4):1413, 1663
(photo, chromosomes)
carnosa—20(4):1413
coulteri—20(4):1414
crispa—20(4):1414
mattturneri—20(4):1414, 1418,
1420-1421 (color photos), 1663
(photo, chromosomes)
parviflora—20(4):1414
riparia—20(4):1414
turneri—20(4):1414, 1419
Arisaema triphyllum (Texas disjunct
record) —20(3):1327, 1329-1331]
(color photos)
Arthrostemma—20(1):235
Aster laevis var. purpuratus—20(1):437
Asteraceae—20(1):145, 149, 176,71,97;
20(2):505, 511,515,559, 597, 845
Astereae—20(4):1387, 1481, 1585,
1589, 1595, 1617
subtribe Bellidinae—20(4):1649
Coreopsideae—20(2):593
Eupatorieae—20(4):1573
Senecioneae—20(2):605;
20(3):1023
Astronidium morobiense—20(4):1527,
1528-1529 (photo)
Axinaea—20(1):235
Azolla caroliniana (Arkansas record) —
Baccharis—20(4):1481
sect. Aristidentes—20(4):1483
sect. Bogotenses—20(4):1483
SIDA 20(4): 1807. 2003
1807
sect. Punctatae—20(4):1481
sect. Tubulatae—20(4):1482
Barringtonia—20(4):1527
Bejaria aestuans—20(1):4
Bellucia—20(1):235
Berberis bealei—20(2):815 (Florida record)
Bergia texana—20(1):412
Bidens alba var. radiata—20(1):438
Blakea—20(1):235
Boerhavia
coulteri
var. coulteri—20(1):154
var. palmeri—20(1):151, 153
spicata—20(1):154
torreyana—20(1):154
wrightii—20(1):154
Boraginaceae—20(2):495
Bowlesia incana—20(1):410
Brachyotum—20(1):236
Brazoria
arenaria—20(4):1571
enquistii—20(4):1565, 1567-1568
(map), 1569 (color photos)
truncata—20(4):1571
Bromelia penguin—20(2):814 (USA/FL
record)
Bromus nottowayanus—20(1):397
Bulbophyllum pachyglossum—
20(2):474, 476 (illus)
Bulbostylis
barbata—20(1):340
pubescens—20(1):340
Cactaceae—20(1):349; 20(2):809;
20(3):1665
Cadetia echinocarpa—20(2):474, 477
(illus)
Calanthe triplicata—20(2):474, 478 (illus)
Calibrachoa parviflora (Colorado
record)—20(3):1290
1808
Calycanthus floridus var. glaucus
20(1):439
Campomanesia—20(1):205
aromatica—20(1):208
dichotoma—20(1):207, 210
guaviroba—20(1):208 (illus), 209
(photo), 210
viatoris—20(1):207, 208 (illus), 211
Canna glauca (Florida record) —20(2):815
Caperonia palustris—20(1):412
Caprifoliaceae—20(2):849
Carex
albicans var. australis (Kansas
record)—20(3):1290
buxbaumii—20(1):398
conoidea—20(4):1727 (Arkansas
record)
corrugata—20(1):398
oklahomensis—20(1):398
ouachitana—20(1):398
oxylepis var. pubescens—20(1):399
planostachys—20(1):340
roanensis—20(1):399
seorsa—20(1):390
swanii (Kansas record) —20(3):1 290
thurberi—20(1):341
Carphephorus odoratissimus—20(2):559,
561 (illus), 565 (map)
Carum carvi (Kansas record)—20(3):1291
Caryophyllaceae—20(1):227
Castilleja
albobarbata—20(4):1343, 1344
(color photos), 1345 (map)
integrifolia—20(4):1345 (map)
Catharanthus roseus (Kansas record) —
20(3):1291
Cavendishia
bracteata—20(1):6
Martii—20(1):6
pubescens—20(1):6
SIDA 20(4): 1808. 2003
BRIT.ORG/SIDA 20(4)
Cerastium—20(1):227-231 (chromosome
numbers)
Chamaesyce prostrate (Colorado record)
—20(3):1291
Chaptalia
estribensis—20(4):1368
hidalogoensis—20(4):1363, 1364
(illus), 1368
hintonii—20(4):1368
lyratifolia—20(4):1 368
madrensis—20(4):1368
mexicana—20(4):1 365, 1367 (illus),
1368
pringlei—20(4):1368
tomentosa—20(4):1368
Cirsium nuttalli—20(2):845 (North Caro-
lina record), 847 (b/w photos)
Citrullus lanatus var. lanatus—20(1):412
Cladium mariscus—20(4):1526
Clidemia—20(1):236
Commelina benghalensis (North Carolina
record) —20(1):419, 420 (illus)
Conostegia—20(1):237
Convolvulaceae—20(1):39; 20(3):1351
Corchorus—20(3):1299
olitorius—20(3):1301, 1302 (map),
1303 (illus)
trilocularis—20(3):1 302 (map), 1305,
1306 (illus)
Corybas epiphyticus—20(2):469, 470
(illus)
Costus pulverulentus (USA/FL record) —
20(2):814
Crataegus—20(1):115
atrovirens—20(1):141, 142 (illus)
castlegarensis—20(1):121, 122
(illus), 124 (map)
douglasii—20(1):1 20
enderbyensis—20(1):136, 137
(illus)
INDEX OF BOTANICAL NAMES
okanaganensis—20(1):131
var. wellsii—20(1):132, 133
(illus), 134 (map)
okennonii—20(1):1 21
orbicularis—20(1):138, 140 (illus)
phippsii—20(1):143
shuswapensis—20(1):128, 129
(illus)
suksdorfii—20(1):119
thermopegaea (Arkansas record
a
20(4):1740
williamsii—20(1):131
Cryptocoryne
beckettii—20(2):819, 826, 827 (color
photos)
undulata—20(2):822 (color photo)
wendtii—20(2):823, 827 (color
photos)
Cryptomeria—20(3):999
Cunoniaceae—20(1):179
Cuphea viscosissima—20(1):442
Cupressaceae—20(2):585
i@ IDressus arizonica—20(2):585,
20(3):1325
Cynoglossum zeylanicum—20(1):41 1
Cynosurus echinatus (Texas record) —
20(2):837
Cyperaceae—20(1):339; 20(3):1 727
Cyperus
arsenei—20(1):341
cephalanthus—20(1):440
diandrus (Kansas record) —20(3):1 291
manimae—20(1):341
var. asperrimus—20(1):341
var. divergens—20(1):341
var. manimae—20(1):342
sphaerolepis—20(1):342
Cyrtomium falcatum (Arkansas record) —
20(4):1 740
Dahlia neglecta—20(2):593, 595 (illus)
SIDA 20(4): 1809. 2003
1809
Demosthenesia
mandonii—20(1):8
pearcei—20(1):8
spectabilis—20(1):8
Dendrobium
antennatum—20(2):467 (illus)
archipelagense—20(2):464, 465
(illus), 467 (illus)
cuthbertsonii—20(2):479, 480 (illus)
gnomus—20(2):469, 471 (illus)
masarangense—20(2):479, 481
(illus)
rhodostictum—20(2):479, 482 (illus)
strepsiceros—20(2):467 (illus)
vexillarius var. hansmeyerense—
20(2):462, 463 (illus)
Deschampsia danthonioides (lowa
record) —20(3):1291
Dichanthelium—20(1):161
aciculare
subsp. angustifolium—
20(1):167
subsp. fusiforme—20(1):167
subsp. neuranthum—20(1):167
acuminatum
subsp. columbianum—
20(1):167
subsp. fasciculatum—20(1):167
subsp. implicatum—20(1):167
subsp. leucothrix—20(1):167
subsp. lindheimeri—20(1):168
subsp. longiligulatum—
20(1):168
subsp. sericeum—20(1):168
subsp. spretum—20(1):168
subsp. thermale—20(1):168
sect. Angustifolia—20(1):166
chamaelonche—20(1):168
subsp. breve—20(1):168
sect. Clandestina—20(1):164
1810
commutatum
subsp. ashei—20(1):169
subsp. equilaterale—20(1):169
subsp. joori—20(1):169
dichotomum
subsp. dichotomum (Kansas
record) —20(3):1292
subsp. lucidum—20(1):169
subsp. mattamuskeetense—
20(1):169
subsp. microcarpon—20(1):169
subsp. nitidum—20(1):169
subsp. roanokense—20(1):1 70
subsp. yadkinense—20(1):1 70
sect. Ensifolia—20(1):166
ensifolium subsp. curtifolium—
1):170
sect. Lancearia—20(1):166
sect. Lanuginosa—20(1):166
sect. Linearifolia—20(1):164
sect. Macrocarpa—20(1):165
sect. Nudicaulia—20(1):165
sect. Oligosantha—20(1):166
oliaqosanthes subsp
Y
scribnerianum—20(1):1 70
ovale
subsp. praecocius—20(1):1 70
subsp. pseudopubescens—
20(1):170
subsp. villosissimum—
20(1):170
sect. Pedicellata— 20(1):165
polyanthes (Kansas record)-
20(3):1292
portoricense subsp. patulum—
20(1):170
sect. Sphaerocarpa—20(1):166
sect. Strigosa—20(1):1 66
strigosum
subsp. glabrescens—20(1):1 71
SIDA 20(4): 1810. 2003
Didiplis diandra (Florida record
BRIT.ORG/SIDA 20(4)
subsp. leucoblepharis—
20(1):171
tenue—20(1):171
eons
Dieteria—20(4):1390, 1391
Dieteria
asteroides—20(4):1 393
var. asteroides—20(4):1393
var. glandulosa—20(4):] 393
var. lagunensis— 20(4):1393
bigelovii—20(4):1394
var. bigelovii—20(4):1394
var. commixta— 20(4):1394
var. mucronata—20(4):1 394
canescens—20(4):1 394
var.ambigua—20(4):1 396
var. aristata—20(4):1397
subsp. canescens—20(4):1 394
var. canescens—20(4):1 394
subsp. glabra—20(4):1 396
var. glabra—20(4):1 397
var. incana—20(4):1396
var. leucanthemifolia—
20(4):1396
var. nebraskana—20(4):1 397
var. sessiliflora—20(4):1 396
var. shastensis—20(4): 1396
var. ziegleri—20(4):1396
Diodia
dasycephala—20(1):415
teres—20(2):552 (photo, seeds)
virginiana—20(2):552 (photo, seeds)
Diogenesia
boliviana—20(1):8
racemosa—20(1):8
Dioscorea
coriacea—20(3):1012
lisae—20(3):1007, 1008-1009
(illus)
INDEX OF BOTANICAL NAMES
Diospyros papuana—20(1):266
Diplarpea—20(1):237
Disterigma
alaternoides—20(1):6
empetrifolium—20(1):6
ovatum—20(1):6
pallidum—20(1):6
pernettyoides—20(1):6
Dryopteridaceae—20(2):833
Echinochloa—20(2):525
colona—20(2):535
crus-galli—20(2):530 (photo,
anatomy)
hispidula—20(2):529 (photo,
anatomy)
oryzicola—20(2):529, 530 (photo,
anatomy)
oryzoides—20(2):529, 530 (photo,
anatomy)
Eleocharis
macrostachya—20(1):342
montevidensis—20(1):41 2
svensoniana—20(1):343
tuberculosa—20(1):399
yecorensis—20(1):343
Eleusine tristachya—20(1):415
xElyhordeum montanense (Kansas
record)—20(3):1292
Enydra fluctuans—20(2):814 (USA/FL
record)
Epilobium leptophyllum (Oklahoma
record) —20(3):1292
Ericaceae—20(1):1
Erigeron canus (Kansas record) —
20(3):1293
Eriogonum—20(1):31
multiflorum—20(1):36
riograndis—20(1):32, 33 (illus), 34
(map)
Erycibe—20(4):1526
SIDA 20(4): 1811. 2003
1811
Eucalyptus torelliana—20(2):814 (USA/
Florida record)
Euchiton—20(2):515
gymnoc¢ ephalus—20(2):516
involucratus—20(2):518
sphaericus—20(2):519
Euphorbia hirta—20(1):412
Eurybia
glauca—20(4):1561
var. glauca—20(4):1562
var. pulchra—20(4):1562
pee
Fagopyrum esculentum—20(1):415;
20(4):1717, 1718 (color illus)
Faika villosa—20(1):67, 68 (b/w photos)
Fatoua villosa—20(1):405
Festuca—20(1):21
cuzcoensis—20(1):21, 23 (illus)
dentiflora—20(3):1016, 1017 (illus)
tovarensis—20(1):24, 25 (illus)
Fimbristylis
complanata—20(1):343
decipiens—20(1):343
pentastachya—20(1):344
Fittingia
tubiflora—20(4):1530
urceolata—20(4):1530, 1531-1532
(photo)
Gaultheria
bracteata—20(1):5
buxifolia var. secunda—20(1):4
erecta—20(1):5
eriophylla var. mucronata—20(1):4
glomerata—20(1):5
hapalotricha—20(1):5
reticulata—20(1):5
vacinioides—20(1):4
Gaylussacia cardenasii—20(1):7
Gentiana andrewsii—20(1):399
Gentianaceae—20(4):1485
1812
Gentianella
boliviana—20(4):1485
yieronymi—20(4):1485
inaequicalyx—20(4):1487, 1488
silenoides—20(4):1489
Geranium
dissectum (Kansas record) —
20(3):1 293
molle (Kansas record)—20(3):1293
Glandularia
quandrangulata—20(3):953, 955
(illus), 958 (map), 963
racemosa—20(3):957 (illus), 958
(map), 965
verecunda—20(3):955 (illus), 958
(map), 964, 20(4):1647
forma albida—20(4):1648
Gleditsia triacanthos (Colorado record)—
20(3):1 293
Gonolobus
albomarginatus—20(1):107
barbatus—20(1):107
chiriquensis—20(1):107
denticulatus—20(1):108
edulis—20(1):107
heterophyllus—20(1):107
tenuisepalus—20(1):105, 106
(figs), 107
Goodyera rubicunda—20(2):479, 483
(illus)
Graffenrieda—20(1):237
Grindelia lanceolata—20(3):1059
Gypsophila elegans—20(1):435
Gyrinops—20(1):261
ledermannii—20(1):58, 262 (map)
Hedyotideae (Rubiaceae)—20(2):549
Hedyotis
nigricans
var. gypsophyla—20(1):215, 216
(map), 217, 218 (photo)
a
SIDA 20(4): 1812. 2003
BRIT.ORG/SIDA 20(4)
var. nigricans—20(1):215, 216
(map), 217 (photo)
Henriettella—20(1):237
Hibiscus syriacus (Kansas record) —
20(3):1293
Homalium d’entrecasteauxense—
20(4):1535
Hygrophila difformis (USA/FL record) —
20(2):814
Hypericum nudiflorum—20(1):414
Hypochaeris glabra——20(1):410, 438
Hypoxis Curtissiim20(1):414
Hyptis mutabilis—20(1):441
Ilex cornuta (Mississippi record) —
20(2):691
lliamna—20(2):645
liciaceae—20(3):1047
llicium floridanum—20(3):1047, 1050-
1053 (photos, anatomy)
Ipomoea
cordatotriloba—20(1):412
costellata
var. costellata—20(1):42, 43
(map)
var.edwardsensis—20(1):39, 40
(illus, color), 43 (map)
xleucantha—20(1):411
stans—20(4):1353 (illus)
tacambarensis—20(4):1351,
1352-1353 (illus)
Ixora—20(4):1471
bervifolia—20(4):1472
gardneriana—20(4):1474
heterodoxa—20(4):1475
syringiflora—20(4):1477
venulosa—20(4):1477
Juncaceae—20(1):339
Juncaginaceae—20(4):1721
Juncus
dichotomus—20(1):345
INDEX OF BOTANICAL NAMES
liebmanii var. polycephalus—
20(1):345
Khaya senegalensis (USA/FL record) —
20(2):814
Koenigia islandica (Utah record) —
20(3):1319
Lamiaceae—20(2):583
Lathyrus aphaca—20(1):413
Leandra—20(1):237
urbaniana—20(4):1645
Leptochloa dubia—20(1):443
Liatris—20(4):1573
elegans—20(2):597
var. bridgesii— 20(2):598, 599
(color photo)
var. carrizana—20(2):599 (color
photo)
var. elegans—20(2):599 (color
photo)
forma flabellata—20(2):603
var. ir nail (color
photo), 6
soln 201 97,98, 99 (fig)
gracilis—20(1):10
ae
patens—20(4):1578 (illus), 1579,
1580 (map)
savannensis—20(4):1574, 1575
(illus), 1580 (map)
Lilium longiflorum—20(1):441
Lindernia crustacean—20(1):416
Lipocarpha maculata—20(1):344
Lithospermum hintoniorum—20(2):501,
402 (color photo)
Loasaceae—20(1):157
Loreya—20(1):237
Lycoris radiata—20(1):414
Lygodium microphyllum—20(4):1759
Lyonia lucida—20(1):440
SIDA 20(4): 1813. 2003
1813
Lythraceae—20(4):1765
Lythrum lineare—20(1):442
Machaeranthera—20(4):1 387, 1388,
1390
tagetina—20(4):1391
tanacetifolia—20(4):1391
Macranthera flammea—20(1):189, 194,
195 (illus), 196 (map)
Mahonia bealei (Mississippi record)—
20(2):691
Maieta—20(1):238
Malvaceae —20(1):47; 20(2):645
Mammillaria theresae—20(4):1665, 1667
(color photos)
Maniltoa—20(4):1526
Marshallia caespitosa—20(1):410
Matelea monticola—20(4):1645
Maytenus urbanii—20(4):1645
Medicago minima—20(1):413
Medinilla—20(1):109
anamalaiana—20(1):110, 111
(illus), 112
beddomei—20(1):112
fuchsioides—20(1):112
malabarica—20(1):112
Melanthiaceae—20(1):221;20(2):611;
(3):1083
Melastomataceae—20(1):109, 233;
0(3):877
—
Meliosma—20(3):931
allenii—20(3):933
brenesii—20(3):933
chiriquensis—20(3):934, 935 (illus)
clandestina—20(3):934, 936 (illus)
cordata—20(3):937
depressiva—20(3):937, 938 (illus)
donnellsmithii—20(3):939
frondosa—20(3):940
glabrata—20(3):940
—
ee
1814
grandiflora—20(3):940
hartshornii—20(3):941
idiopoda—20(3):941
linearifolia—20(3):94 1
occidentalis—20(3):941
schlimii—20(3):942
subcordata—20(3):942
vernicosa—20(3):943
Melissa officinalis—20(1):414
Mentzelia
lindheimeri—20(1):157
texana—20(1):157
Meriania—20(1):238
Miconia—20(1):238
Mirabilis jalapa—20(1):414
Mitracarpus hirtus—20(2):554 (photo,
seeds)
Monochaetum—20(1):239
Monolena—20(1):239
Mouriri—20(1):240
Muhlenbergia dubia (Texas disjunct)—
20(4):1 763
Murdannia keisak—20(1):41 1
Myrsinaceae—20(3):899, 913,919;
20(4):1423
Myrsine coriacea—20(3):910
Myrtaceae—20(1):205; 20(4):1449
Myrtus communis—20(1):442
Najas minor—20(1):414
Nandina domestica (Mississippi record) —
20(1):411;20(2):691
Nastus schlechteri —20(4):1533
Nephrolepis exaltata (Arkansas record) —
20(2):833
Nerium oleander (Arkansas record) —
20(4):1 740
Nyctaginaceae—20(1):151;20(3):885
Oenothera
elata subsp. hirsutissima
SIDA 20(4): 1814. 2003
20(1):442
BRIT.ORG/SIDA 20(4
engelmannii (Colorado record) —
20(3):1 294
laciniata (Colorado record) —
20(3):1 294
Opuntia corallicola—20(2):809
Orobanchaceae—20(1):189; 20(4):1343
Orthaea
boliviensis—20(1):6
constans—20(1):5
ferreyrae—20(1):6
ignea—20(1):5
pinnatinervia—20(1):5
rusbyi—20(1):5
weberbaueri—20(1):5
Ossaea—20(1):240
Oxalis
corniculata—20(1):406
triangularis subsp. papilionacea—
20(2):814 (USA/FL record)
Oxypolis ternata—20(3):1323
Packera—20(3):1023
paupercula var. appalachiana—
20(2):605, 607 (map), 609 (photo
chromosomes)
plattensis—20(2):607 (map);
20(3):950 (map)
tampicana—20(3):950 (map)
texensis—20(3):945, 946 (illus), 950
(map)
tridenticulata—20(3):950 (map)
Panicum—20(1):161
aciculare
var. neuranthum—20(3):1044
var. ramosum—20(3):1044
acuminatum
var. ferventicola—20(3):1044
var. thermale—20(3):1044
amarum subsp. amarulum—
20(1):171
INDEX OF BOTANICAL NAMES
anceps subsp. rhizomatum—
20(1):171
sect. Antidotalia—20(1):163
capillare subsp. hillmanii—
20(1):171
commutatum var. equilaterale—
20(3):1044
dichotomiflorum
subsp. bartowense—20(1):171
subsp. puritanorum—20(1):1 71
hallii subsp. filipes—20(1):1 72
sect. Hemitoma—20(1):163
hirticaule
subsp. sonorum—20(1):1 72
subsp. stramineum— 20(1):1 72
ovale var. praecocius—20(3):1044
philadelphicum
subsp. gattingeri—20(1):172
subsp. lithophilum—20(1):1 72
rigidulum
var. abscissum—20(3):1045
subsp. abscissum—20(1):172
subsp. combsii—20(1):1 72
subsp. elongatum—20(1):1 72
subsp. pubescens—20(1):1 73
Parathesis—20(3):899, 913
subg. Laterales—20(3):915
sect. Calophyllae—20(3):915
sect. Laterales—20(3):915
melanostricta—20(3):910
subg. Parathesis
sect. Cubanae—20(3):916
sect. Versatiles—20(3):91 7
pseudocalophylla—20(3):899, 900
(illus)
pseudocrassiramea— 20(3):902,
903 (illus)
rothschuhiana—20(3):906 (illus),
SIDA 20(4): 1815. 2003
1815
Pedilochilus longipes—20(2):469, 472
(illus)
Pernettya prostrata—20(1):4
Phaius amboinensis—20(2):469, 473
(illus)
Phaleria coccinea—20(1):266
Phreatia loriae—20(2):474
Phyllanthus
tenellus—20(1):413
urinaria subsp. urinaria (Kansas
record)—20(3):1294
Plesioneuron croftii—20(4):1536
Pluchea yucatanensis—20(1):403
Poaceae—20(1):161, 21,423: 20(2):571,
Panicoideae—20(2):525
Loliinae—20(3):1015
Polemoniaceae—20(4):1371
Polyclita turbinata—20(1):7
Polygala alba (Arkansas record
20(4):1740
Polygonaceae—20(1):31;20(3):987;
20(4):1631;20(4):1717
Polygonum—20(3):987
aviculare
subsp. buxiforme—20(3):988
subsp. rurivagum—20(4):1 709
douglasii subsp. johnstonii (Cana-
dian record)—20(3):1317
fowleri subsp. hudsonianum—
20(3):994
hudsonianum—20(3):994
humifusum—20(3):995
subsp. caurianum—20(3):995
patulum—20(4):1 707
prolificum—20(3):995
ramossisimum subsp. prolificum—
20(3):995
sawatchense—20(4):1631, 1633
subsp. oblivium—20(4):1634
(illus), 1635 (photo), 1637
ao
1816
subsp. sawatchense—
20(4):1633, 1634 (illus), 1635
(photo)
setaceum—20(1):415
Striatulum var. texense—20(3):996
texense—20(3):995
Portulaca
matthewsii—20(4):1357, 1358,
1359 (illus)
rzedowskiana— 20(2):487, 489
(illus)
Portulacaceae—20(2):487; 20(4):1357
Pristiglottis montana—20(2):474, 475
(illus)
Psammisia
guianensis—20(1):6
pauciflora—20(1):6
Pseudognaphalium
biolettii—20(1):149
luteoalbum—20(1):403
Psidium
laruotteanum—20(4):1452, 1453
(photo), 1459 (map)
rufum—20(4):1455
salutare—20(4):1449, 1455, 1457
var.decussatum—20(4):1463,
1464 (map)
var.mucronatum—20(4):1463,
1464 (map)
var. pohlianum—20(4):1466,
1464 (map)
var. salutare—20(4):1457, 1458
(photo), 1460 (map)
var. sericeum—20(4):1467,
1461 (map)
Psychotria
archboldii—20(4):1533
bulilimontis—20(4):1514, 1516—
1518 (photos)
SIDA 20(4): 1816. 2003
BRIT.ORG/SIDA 20(4)
var.aestuarii—20(4):1519, 1520
(photos)
croftiana—20(4):1533
mayana—20(4):1533
melanocarpa—20(4):1533
waiuensis—20(4):1534
Pteroceras mMonsooniae— 20(3):923,
925 (illus)
Punica granatum (Arkansas record) —
20(4):1 740
Pyrus calleryana (Kansas record) —
20(3):1 294
Quercus arkansana—20(1):441
Raphanus raphanistrum—20(1):439
Rhynchosia tomentosa—20(1):440
Richardia
brasiliensis—20(2):554 (photo,
seeds)
humistrata—20(2):556 (photo,
seeds)
scabra—20(2):556 (photo, seeds)
tricocca—20(2):556 (photo, seeds)
Ricinus communis (Arkansas record) —
20(4):1 741
Rosaceae—20(1):115
Rotala rotundifolia—20(4):1 765, 1766
(map), 1768 (color photos)
Rubiaceae—20(1):215;20(2):549;
20(3):1471
Rudbeckia auriculata (Florida record)-
20(2):815
Rusbya taxifolia—20(1):7
Sagina
apetala (Kansas record) —
20(3):1295
procumbens—20(1):406
Sagraea rugosa—20(4):1645
Salicaceae—20(2):523
Salix exigua var. sericans—20(2):523
INDEX OF BOTANICAL NAMES
Sanicula smalliim20(1):410
Satyria
boliviana—20(1):5
neglecta—20(1):5
Schinus terebinthifolius (Arkansas
record) —20(4):1 741
Schistocarpha
bicolor—20(2):507 (map)
calzadana—20(2):505, 506 (photo),
507 (map)
eupatorioides—20(2):507 (map)
liebmannii—20(2):509 (map)
matudae—20(2):509 (map)
pedicellata—20(2):510 (map)
platyphylla—20(2):510 (map)
Scleria interrupta—20(1):344
Scoparia dulcis—20(1):416
Sedum sarmentosum (Kansas record)—
20(3):1295
Senecio
neowebsteri—20(2):5 13
sandersiana—20(1):175, 1 76 (illus),
177 (map)
spribillei—20(2):511, 512 (photo)
Silphium radula (Kansas record)—
20(3):1295
Siphonandra
boliviana—20(1):6, 13, 15 (illus)
magnifica—20(1):6
Solanaceae—20(4):1651
Solanum
chlamydogynum—20(4):1653, 1655
(photo, epidermis)
obovalifolium—20(4):1654, 1655
(photo, epidermis)
oppositifolium—20(4):1656, 1657
(photo, epidermis
sessile —20(4):1651, 1656, 1657
(photo, epidermis)
ir
SIDA 20(4): 1817. 2003
1817
troste—20(4):1656, 1659 (photo,
epidermis)
turgidum—20(4):1658, 1659 (photo,
epidermis)
Solidago—20(4):1589, 1617
altissima—20(4):1628 (photo)
subsp. gilvocanescens—
20(4):1606
arenicola—20(4):1589, 1590 (illus),
arguta—20(4): 1602
auriculata—20(4):1627 (photo)
ser. Auriculatae—20(4):1606
brachyphylla—20(4): 1602
ser. Drummondiani—20(4):1606
erecta—20(4):1592
faucibus—20(4):1595, 1596, 1597
(illus)
subsect. Humiles—20(4):1 606
kralii—20(4):1606, 1607-1609
subsp. fallax—20(4):1611
var. salebrosa—20(4):161 1
ludoviciana—20(4): 1602
macrophylla—20(4):1627 (photo)
subsect. Multiradiatae—
20(4):1605
odora—20(4):1628 (photo)
subsp. chapmanii—20(4):161 1
ser. Odorae—20(4):1606
patula—20(4):1602
subsp. strictula—20(4):1612
pinetorum—20(4):1625 (photo)
plumosa—20(4):1592
puberula subsp. pulverulenta—
20(4):1612
rigida—20(4):1628 (photo)
rugosa var. cronquistiana—
20(4):1612, 1613-1614 (photos)
1818
sempervirens
subsp. azorica—20(4):1615
subsp. mexicana—20(4):1615
simplex—20(4):1592
speciosa subsp. pallida—
20(4):1615
sphacelata—20(4):1627 (photo)
stricta subsp. gracillima—
20(4):1615
tarda—20(4): 1602
ulmifolia—20(4):1628 (photo)
velutina
subsp. californica—20(4):1616
subsp. sparsiflora—20(4):1616
verma—20(4): 1602
Spathoglottis plicata—20(2):479, 485
(illus)
Spermacoce
assurgens—20(2):553 (photo,
seeds)
prostrata—20(2):552 (photo, seeds)
tetraquetra—20(2):553 (photo,
seeds)
verticillata—20(2):553 (photo,
seeds)
Spermacoceae (Rubiaceae) —20(2):549
Sphaeralcea—20(1):47
ambigua—20(1):48
caespitosa—20(1):48
var. caespitosa—20(1):49, 50
(illus), 51 (map)
var. Williamsiae—20(1):48, 50
(illus), 51 (map)
Sphyrospermum
buxifolium—20(1):8
cordifolium—20(1):8
sessiliflorum—20(1):8
Stachys cordata—20(2):583
SIDA 20(4): 1818. 2003
BRIT.ORG/SIDA 20(4)
Stenanthium
densum—20(1):221, 224 (photo
chromosomes)
gramineum—20(1):224 (photo
chromosomes)
leimanthoides—20(1):223
Sterculia—20(4):1527
Stylogyne—20(3):919
darienensis—20(3):92 1
hayesii—20(3):921
pucuroensis—20(3):919, 920 (illus),
921
turbacensis—20(3):922
subsp. laevis—20(3):922
subsp. turbacensis—20(3):922
Symphyotrichum puniceum var.
scabricaule—20(1):403
Symplocaceae—20(3):928
Symplocos kothayarensis—20(3):928,
929 (illus)
Syzygium
lorentzianum—20(4):1530
trivene—20(4):1530
Tamarix parviflora—20(1):406
Taxodiaceae—20(3):999
x Taxodiomeria—20(3):999 1001
peizhongii—20(3):1001, 1002-
1005 (color photos)
Taxodium mucronatum—20(3):999,
2003 (color photo)
Tephrosia jamnagarensis—20(4):1 701,
1702, 1703 (photos)
Tessmanianthus—20(1):240
Tetrazygia fadyenii—20(3):877, 878, 882
(b/w photos), 883 (map)
Thelesperma
caespitosum—20(1):73 (map), 82
(photo chromosomes)
marginatum—20(1):73 (map), 82
(photo chromosomes)
INDEX OF BOTANICAL NAMES
pubescens—20(1):73 (map), 82
(photo chromosomes)
var. caespitosum—20(1):91
subnudum—20(1):71, 73 (map), 82
(photo chromosomes)
windhamii—20(1):91
Thelypteris rhachiflexuosa—20(3):1311,
1313 (photo), 1314 (map)
Themistoclesia
peruviana—20(1):7
unduavensis—20(1):7, 15, 16 (illus)
Thibaudia
axillaris —20(1):7
crenulata—20(1):7
densiflora—20(1):7
macrocalyx—20(1):7
regularis—20(1):7
Thymelaeaceae—20(1):261
Tibouchina—20(1):240
Tiliaceae—20(3):1299
Tococa—20(1):241
Toxicoscordion
brevibracteatus—20(3):1089
(photo, chromosomes)
fontanum—20(3):1089 (photo,
chromosomes)
fremontii—20(3):1089 (photo, chro-
mosomes)
micranthum—20(3):1089 (photo,
chromosomes)
nuttalliim20(3):1085, 1987 (illus,
map), 1089 (photo, chromosomes)
paniculatum—20(3):1089 (photo,
chromosomes)
venenosum—20(3):1089 (photo,
chromosomes)
Trachelospermum difforme—20(4):1641,
1642
Tragopogon pratensis—20(1):410
SIDA 20(4): 1819. 2003
1819
Trianthema portulacastrum (Colorado
record) —20(3):1 295
Trichadenia sasae—20(4):1522, 1523-
1525 (photo)
Tridax procumbens—20(1):438
Tridens flavus var. chapmanii (Kansas
record) —20(3):1295
Trifolium
lappaceum—20(1):413
nigrescens—20(1):413
Triglochin concinna—20(4):1721, 1722
(color, illus)
Trigonospermum annum (record for Ec-
uador)—20(2):675
Triolena—20(1):241
Uvularia
perfoliata—20(4):1713, 1715 (map)
sessilifolia—20(4):1713,1714,1715
(map)
Vaccinium
dependens—20(1):7
erythrocarpum—20(1):400
floribundum—20(1):7
Veratrum album—20(2):61 1
Verbena montevidensis—20(1):416
Verbenaceae—20(3):953
Veronica
hederifolia—20(1):416
persica (Colorado record)—
20(3):1296
polita—20(1):416
Viburnum
lantanoides—20(1):400
molle—20(2):856 (map)
ozarkense—20(2):849, 856 (map)
Viola arvensis—20(1):406
Wahlenbergia marginata—20(1):411
Websteria confervoides—20(1):41 2
1820 BRIT.ORG/SIDA 20(4)
Weinmannia—20(1):179 var. gooddingii—20(4):1408
bradfordiana—20(1):179, 181 var. incisifolium—20(4):1409
(illus) var. paradoxum—20(4):1409
cochensis—20(1):186 var. scabrellum—20(4):1409
condorensis—20(1):183, 184 (illus) subsp. spinulosum—20(4):1408
Xanthisma—20(4):1390, 1397, 1585 var. spinulosum— 20(4):1408
arenarium—20(4):1405 stenolobum—20(4):1409
blephariphyllum—20(4):1403 rexel)
section Blepharodon—20(4):1401 subsp. drummondii—20(4):1401
coloradoense—20(4):1403 SUS: texanum
crutchfieldii—20(4):1 403 var. orientale—20(4):1401
glaberrimum—20(4):1586 var. texanum—20(4):1401
gracile —20(4):1405 subsp. texanum—20(4):1401
grindelioides—20(4):1403 wiscidum —20(4)1410
var. depressum—20(4):1404 WiGaInS 2 OVN eae
var. grindelioides—20(4):1404 section Xanthisma—20(4):1401
gymnocephalum—20(4):1404 Xylorhiza |
gypsophilum—20(4):1404 glabriuscula—20(1):146 (map), 147
section Havardii—20(4):1 410 linearifolia—20(1 ):145, 146 (map)
incisifolium—20(4):1585 venusta—20(1):146 (map), 147
johnstonii—20(4):1404 Yucca
junceum—20(4):1406 cernua—20(3):892, 893 (illus), 894
restiforme—20(4):1405 (color photos)
rhizomatum—20(4):1405 pallida—20(3):897
section Sideranthus—20(4):1405 reverchonii—20(3):897
spinulosum—20(4):1406 rostrata—20(3):895
var. chihuahuanum— rupicola—20(3):897
20(4):1408 thompsoniana—20(3):897
var. glaberrimum—20(4):1408
Zannichellia palustris—20(1):417
subsp. gooddingii—20(4):1408
Zizania texana—20(2):57 1]
SIDA 20(4): 1820. 2003
SUBJECT INDEX
1821
SUBJECT INDEX: VOLUME 20 (2002-2003)
Alabama—20(1):403 (records);
20(3):1059
North Central—20(4):1589
Amaranth histories of meaning—
20(3):1073
Anamalai Hills (South India)—20(1):109
licium floridanum—20(3): 1047
Grindelia lanceolata—20(3):1059
Anderson County (Texas) —20(1):355
Andes (Venezuela) —20(3):1007
André Michaux—20(2):861
Apalachicola River (Florida) —20(1):97
Appalachian Mountains (Mesic forest
goldenrod)—20(4):1595
Arizona (Mohawk Dunes)—20(3):1155
Arkansas
Carex conoidea (record) —
20(4):1 727
Cleveland County—20(4):1731
state records—20(1):403, 409;
20(2):833, 849; 20(4):1 737
Atlantic Rainforest—20(1):205
Bertero, Carlo (1789-1831) in the New
World—20(2):621
Big Branch Marsh National Wildlife Ref-
uge (St. Tammany Parish, Louisi-
ana)—20(3):1189
Big Lake Bottom Wildlife Management
Area (Anderson County, Texas) —
20(1):355
Biosphere Reserve (Querétaro, Mexico) —
257
Bismarck Archipelago (Papua New
Guinea) —20(2):461
Blazing star—20(1):97
Bolivia—20(1):1; 20(4):1485
SIDA 20(4): 1821. 2003
Brazil—20(1):205, 295
Central and Southern—20(4):1471
Cacoma (Ualisco, Mexico) —20(4):1 343
Cactaceae (Mammillaria theresae)—
20(4):1665
Calcasieu Parish (Louisiana records)—
20(1):431
Canada (record) —20(3):1317
Carlo Bertero (1789-1831) in the New
World—20(2):621
Central Brazil—20(4):1471
Central mineral region (Texas) —
20(4):1565
Cerrado in Emas National Park (Goias,
central Brazil) —20(1):295
Chester Morrison Rowell, Jr. (1925-
2003)—20(4):1771
Chiapas (Mexico record)—20(3):1311
China (Shanghai) —20(3):999
Cleveland County (Arkansas)—
20(4):1 731
Conservacion (Mammillaria theresae)—
Coastal Plain—20(1):189
Colorado (U.S.A.)—20(3):1023
Common garden study (Grindelia
lanceolata)—20(3):1059
Costa Rica—20(3):931
southern—20(1):105
Dahlia—20(2):593
Darién (Panama)—20(3):919
Del Weniger (1923-1999)—20(2):867
Documented chromosome numbers
Arida blepharophylla and
mattturneri—20(4):1663
Cerastium—20(1):227
Stenathium—20(1):221
1822
Packera paupercula var.
appalachiana—20(2):605
Toxicoscordion nuttallii—
20(3):1085
Durango (Mexico)—20(1):175
Eaglewood (Gyrinops, [hymelaeaceae)—
20(1):26]
East Texas (Post oak savanna region) —
East-west transition (Texas flora)—
20(4):1693
Ecuador—20(1):233
Southern—20(1):1 79
Edwards Plateau (Texas) —20(1):39;
20(3):945
El Salvador (Laguna de Alegria) —20(2):675
Emas National Park (Goias, central Brazil)
—20(1):295
Endangered and narrow endemic
(Tephrosia jamnagarensis)
Ethnobotany—20(2):61 |
Ecuador—20(1):233
Evolutionary taxonomy or phylogenetic
systematics? —20(2):647
Florida—20(1):97; 20(2):559
introduced species —20(2):819
invasive fern (Lygodium
microphyllum)—20(4):1 759
Rotala rotundifolia (record) —
20(4):1765
State records—20(2):813
Freeman, John D., Herbarium (AUA)—
20(3):1277
Goldenrod—20(4):1605
Appalachian Mountains —20(4):1595
Grassland biome (records for central
North America)—20(3):1 289
Guerrero (México) —20(2):505
Gujarat (Saurashtra, India) —20(4):1 701
SIDA 20(4): 1822. 2003
BRIT.ORG/SIDA 20(4)
Hawaii—20(2):515
Heart-leaved hedge-nettle (Stachys
cordata)—20(2):583
Hickory Creek Unit (Big Thicket National
Preserve, Tyler County, Texas, check-
list) —20(2):781
Hispaniola—20(4):1645
Hummingbird-pollinated treelet
Tetrazygia fadyenii)—20(3):87 7
Hunstein Range (Papua New Guinea)—
20(1):61 (color photo)
India—20(3):923, 927
Saurashtra, Gujarat—20(4):1 701
Interior Highlands (Arkansas, Missouri and
Oklahoma)—20(2):849
Invasive fern (Lygodium microphyllum,
Florida) —20(4):1 759
lowa (records)—20(1):405
Jalisco (Mexico) —20(4):1343
Jamaica—20(3):877
Jasper county (eastern Texas) —20(3):891
Jeff Davis County (Texas) —20(2):585;
John D. Freeman Herbarium (AUA)—
20(3):1277
Kansas tallgrass prairie —20(1):269
Kentucky (records)—20(1):397
Konza Prairie Biological Station—
Laguna de Alegria (national monument
in the crater of Tecapa Volcano, El
Salvador) —20(2):675
Ledermann’s April River (Papua New
Guinea)—20(1):55
Lemma and caryopsis (Echinochloa)—
20(2):525
Louisiana
Andropogon glomeratus var.
glaucopsis (record) —20(4):1723
SUBJECT INDEX
Big Branch Marsh National Wildlife
Refuge (St. Tammany Parish, Louisi-
ana)—20(3):1189
Calcasieu Parish records—20(1):431
Lowland ultrabasic flora (Papua New
Guinea) —20(4):1491
Maguey—20(2):495
Mapim/ Biosphere Reserve (México,
checklist) —20(2):797
Mesic forest goldenrod (Appalachian
Mountains) —20(4):1595
Mesoamerica—20(3):899
Thelypteris record—20(3):1311
México—20(1):423 (record); 20(2):495;
20(3):1299 (records); 20(4):1 363,
1449
Biosphere Reserve (Querétaro)—
20(4):1357
Durango—20(1):175
Guerrero—20(2):505
Jalisco—20(4):1343
Michoacan—20(2):487; 20(4):1 351
Northwestern Oaxaca—20(2):505
Nuevo Leén—20(2):501
Sierra Madre Oriental—20(2):593
Sonora—20(1):339, 349
Michaux, André—20(2):861
Michoacan (México) —20(2):487;
20(4):1351
Mississippi (Oktibbeha County)—20(2):691
Missouri—20(2):849 (record)
Carex conoidea (record) —20(4):1727
Mohawk Dunes (Arizona) —20(3):1155
Montana (U.S.A.)—20(2):511
Moro Bottoms Natural Area (state-pro-
tected area, Cleveland Co., Arkansas)
—20(4):1731
Morphology (Grindelia lanceolata)—
SIDA 20(4): 1823. 2003
1823
Nags Head Woods (Dare County, North
Carolina)—20(2):839
Nevada—20(1):47
New Guinea (Raja Ampat Islands) —
20(3):1093
Newton and Jasper counties (eastern
Texas) —20(3):891
North America—20(2):515;20(4):1561,
1631, 1707, 1709
grassland biome records for central
North America—20(3):1 289
southwestern—20(1):151
North Carolina (record) —20(1):419
(record); 20(2):845 (record)
Nags Head Woods (Dare County)—
20(2):839
North central Alabama—20(4):1589
Northwest Argentina (rust fungi)—
20(1):313
Northwestern Oaxaca (México) —
Norwegian and Sami ethnobotany (Ver-
atrum album)—20(2):61 1
Nuevo Ledén (México) —20(2):501
Ogallala ecotone on the Dempsey Divide
(Roger Mills County, Oklahoma)—
20(3):1217
Ohio—20(4):1677
Okanagan (Canada) —20(1):115
Oklahoma—20(2):849 (record)
Ogallala ecotone on the Dempsey
Divide, Roger Mills County, Okla-
homa)—20(3):1217
Oktibbeha County (Mississippi)—
20(2):691
Panama—20(3):931
Darién—20(3):919
Pappus variation (Solidago) —20(4):1617
Papua New Guinea—20(1):261, 55
1824
Bismarck Archipelago—20(2):461
lowland ultrabasic flora—20(4):1491
Peninsular
Florida (U.S.A.)—20(2):559
India—20(3):927
People’s Republic of China (Shanghai)—
20(3):999
Peru—20(3):1015
Phylogenetic systematics (Evolutionary
taxonomy)—20(2):647
Pitcher plant bogs (Texas) —20(1):387
Post oak savanna region (East Texas) —
20(1):373
Prairies and oak woodlands/savannas
(North Central and East Texas) —
20(3):1139
Querétaro (Biosphere Reserve, Mexico) —
20(4):1357
Raja Ampat Islands (New Guinea) —
20(3):1093
Reese, William Dean (1928-2002)—
20(3):1335
Rio Mayo region (Sonora, Mexico)
Rockhouse microhabitat (West Cross
Timbers of North Central Texas) —
20(3):1327
Rowell, Jr, Chester Morrison (1925-2003)
—20(4):1 771
Rust fungi (Uredinales)—20(1):313
Sami ethnobotany (Veratrum album)—
20(2):611
Saurashtra (Gujarat, India) —20(4):1 701
Schiede and Deppe collections—
20(3):909
Schlechtendal Myrsinaceae—20(3):909
Seed and fruit characters
(Spermacoceae)—20(2):549
Semaphore pricklypear cactus (Opuntia
corallicola) —20(2):809
SIDA 20(4): 1824. 2003
BRIT.ORG/SIDA 20
—
4)
Sergipe (northeast region, Brazil) —
20(1):205
Shanghai (People’s Republic of China) —
3):999
Shuswap diversity center (Canada) —
20(1):115
Sierra
“El Viejo," (Sonora, México) —20(1):349
Gorda Biosphere Reserve
(Querétaro, Mexico) —20(4):1357
Madre Oriental (México) —20(2):593
Sierras Manantlan Valisco, Mexico) —
20(4):1343
Sonora (Mexico) —20(1):339, 349
South America—20(1):21;20(4):1449
South India—20(1):109
Southeastern United States—20(1):189;
20(4):1573
Southern Brazil—20(4):1471
Southwestern Shuswap diversity center
(Canada)—20(1):115
Tecapa Volcano (El Salvador) —20(2):675
Tennessee—20(3):1059
Texas—20(1):39, 215, 435,437 (records;
20(2):781 (checklist),837 (records);
20(4):1663, 1713
Anderson County—20(1):355
central—20(4):1 763
central mineral region—20(4):1565
East (Post oak savanna region) —
20(1):373
east-west transition of flora—
20(4):1693
Edwards Plateau—20(3):945
Jeff Davis County—20(2):585
Newton and Jasper counties —
20(3):891
Pitcher plant bogs—20(1):387
Prairies and oak woodlands/savan-
SUBJECT INDEX
nas (North Central and East
Texas) —20(3):1139
Rockhouse microhabitat (West
Cross Timbers, North Central
Texas) —20(3):1327
South—20(1):31
Species new to science
Arida mattturneri, sp. nov.—20(4):1418
Brazoria enquistil, SD. NOV.—
20(4):1565
Eriogonum riograndis, sp. nov.—
20(1):32
Ipomoea costellata var.edwardensis,
var. nov.—20(1):39
Liatris elegans var. bridgesii, var.
nov.—20(2):598
Liatris elegans var. kralii, var. nov.—
20(2):601
Packera texensis, so. nov.—20(3):945
Yucca cernua, sp. nov.—20(3):892
State Records
Agalinis harperi (state record) —
20(3):1321
Anaphalis margaritacea (state
record)—20(1):435
Aster laevis var. purpuratus (state
record) —20(1):437
Bidens alba var. radiate (state
record)—20(1):438
Calycanthus floridus var. glaucus
(state record) —20(1):439
Cynosurus echinatus (state
record) —20(2):837
Fagopyrum esculentum (state
record) —20(4):1717
Gypsophila elegans (state record) —
20(1):435
SIDA 20(4): 1825. 2003
1825
Hyptis mutabilis (state record) —
20(1):441
Lilium longiflorum (state record) —
20(1):441
Lyonia lucida (state record) —
20(1):440
Lythrum lineare (state record) —
20(1):442
Myrtus communis (state record) —
20(1):442
Oxypolis ternata (deleted from Texas
flora) —20(3):1323
Raphanus raphanistrum (state
record)—20(1):439
Tridax procumbens (state record) —
20(1):438
Triglochin concinna (a new family,
genus, and species) —20(4):1 721
Trans-Pecos—20(4):1417
Tyler County (Texas, checklist)—
20(2):781
wildrice—20(2):571
Tyler County (Texas, checklist) —20(2):781
Type localities (Ohio) —20(4):1677
Ukhop pok (Anodendron
oblongifolium)—20(1):59 (plant
poison)
United States (records) —20(2):813
Uredinales—20(1):313
Utah—20(1):47
Koenigia islandica (state record) —
20(3):1319
Venezuela (Andes) —20(3):1007
West Gulf Coastal Plain—20(2):767;
20(3):1247
William Dean Reese (1928-2002)—
20(3):1335
1826
BRIT.ORG/SIDA 20(4)
229 NEW NAMES AND NEw CoMmBINATIONS:
VOLUME 20 (2002-2003)
Abronia umbellata var. breviflora
(Standley) L.A. Galloway, comb. et stat.
nov.—20(3):888
Acleisanthes lanceolata var.megaphylla
(Fowler & B.L. Turner) Spellenb. & J.Poole,
comb. nov.—20(3):886
Agave ovatifolia Starr & Villarreal, sp.
nov.—20(2):495
Pests
| J it} ed ee
(Mez) Ricketson & Pipoly, comb. et stat.
nov.—20(4):1431
Ardisia verdisepala Ricketson & Pipoly, sp.
nov.—20(4):1435
Arida D.R. Morgan & R.L. Hartman, comb.
et stat. nov.—20(4):1410
Arida arizonica (R.C. Jackson & RR.
Johnson) D.R. Morgan & R.L. Hartman,
comb. nov.—20(4):1413
Arida blepharophylla (A. Gray) DR.
Morgan & R.L. Hartman, comb. nov.—
20(4):1413
Arida carnosa (A. Gray) D.R. Morgan & R.L.
Hartman, comb. nov.—20(4):1413
Arida coulteri (A. Gray) D.R. Morgan & R.L.
Hartman, comb. nov.—20(4):1414
Arida crispa (1.S. Brandegee) D.R. Morgan
& R.L. Hartman, comb.nov.—20(4):1414
Arida mattturneri 8.L. Furner & G.Nesom,
sp. nov.—20(4):1418
Arida parviflora (A. Gray) D.R. Morgan &
R.L. Hartman, comb. nov.—20(4):1414
Arida riparia (Kunth) D.R. Morgan & RL.
Hartman, comb. nov.—20(4):1414
Arida turneri (M.L. Arnold & R.C. Jackson)
D.R. Morgan & R.L. Hartman, comb.
nov.—20(4):1414
Asteraceae tribe Astereae subtribe
Bellidinae (Cass. ex D.Don) G.Nesom &
SIDA 20(4): 1826. 2003
H. Robinson, comb. et stat. nov.—
20(4):1649
Baccharis sect. Punctatae Giuliano & GL.
Nesom, sect. nov.—20(4):1481
Boerhavia coulteri var. palmeri (S. Wats.)
Spellenb., comb. nov.—20(1):153
Brazoria enquistii V\.W. Turner, sp. nov.—
20(4):1565
Castilleja albobarbata I|tis & Nesom, sp.
nov.—20(4):1343
Chaptalia hidalogoensis |. Cabrera &
Nesom, sp. nov.—20(4):1363
Crataegus atrovirens ee &
O’Kennon, sp. nov.—20(1):14
Crataegus castlegarensis J.B ce &
O’Kennon, sp. nov.—20(1):121
Crataegus enderbyensis J.8. Phipps &
O'Kennon, sp. nov.—20(1):136
Crataegus okanaganensis var. wellsii J.B.
Phipps & O’Kennon, var.nov.—20(1):1 32
Crataegus orbicularis J.B. ee &
O'Kennon, sp. nov.—20(1):1
Crataegus shuswapensis J. : — &
O’Kennon, sp. nov.—20(1):128
Dahlia neglecta D.E. Saar, sp. nov.—
20(2):593
Dendrobium archipelagense Howcroft &
Takeuchi, sp. nov.—20(2):464
Dendrobium vexillarius var. hans-
meyerense Howcroft & Takeuchi, var.
nov.—20(2):462
Dichanthelium aciculare subsp. angus-
tifolium (Elliott) Freckmann & Lelong,
comb. & stat. nov.—20(1):167
Dichanthelium aciculare subsp. fusi-
forme (Hitchc.) Freckmann & Lelong,
comb. & stat. Nov.—20(1):167
NEW NAMES AND NEW COMBINATIONS
Dichanthel | bso. neuran-
thum (Griseb.) Freckmann & Lelong,
comb. & stat. nov.—20(1):167
Dichanthelium acuminatum subsp.
columbianum (Scribn.) Freckmann &
Lelong, comb. & stat. nov.—20(1):167
Dichanthelium acuminatum subsp.
fasciculatum (Torr.) Freckmann &
Lelong, comb & stat.. nov.— 20(1):167
Dichanthelium acuminatum subsp.
implicatum (Scribn. ex Nash
Freckmann & Lelong, comb. & stat.
nov.—20(1):167
Dichanthelium acuminatum subsp.
leucothrix (Nash) Freckmann & Lelong,
comb. & stat. nov.—20(1):167
Dichanthelium acuminatum subsp.
lindheimeri (Nash) Freckmann &
Lelong, comb. & stat. nov.—20(1):168
Dichanthelium acuminatum subsp.
longiligulatum (Nash) Freckmann &
Lelong, comb. & stat. nov.—20(1):168
Dichanthelium acuminatum subsp.
sericeum (Schmoll) Freckmann &
Lelong, comb. & stat. nov.—20(1):168
Dichanthelium acuminatum subsp.
spretum (Schult.) Freckmann & Lelong,
comb. & stat. nov.—20(1):168
Dichanthelium acuminatum subsp.
thermale (Bol.) Freckmann & Lelong,
comb. & stat. nov.—20(1):168
t tifall
~
in B.L. Rob. & Fernald) Freckmann &
Lelong, stat. nov.—20(1):166
Dichanthelium chamaelonche (Trin.
Freckmann & Lelong, comb. nov.—
20(1):168
Dichanthelium chamaelonche subsp.
breve (Hitchc. & Chase) Freckmann &
Lelong, comb. & stat. nov.—20(1):168
eee)
SIDA 20(4): 1827. 2003
1827
Dichanthelium sect. Clandestina
Freckmann & Lelong, sect. nov.—
20(1):164
Dichanthelium commutatum subsp.
ashei (G. Pearson ex Ashe) Freckmann &
Lelong, comb. & stat. nov.—20(1):169
Dichanthelium commutatum subsp.
equilaterale (Scribn.) Freckmann &
Lelong, comb. & stat. nov.—20(1):169
Dichanthelium commutatum subsp.
joori (Vasey) Freckmann & Lelong,comb.
& stat. nov.—20(1):169
Dichanthelium dichotomum subsp.
lucidum (Ashe) Freckmann & Lelong,
comb. & stat. nov.—20(1):169
Dichanthelium dichotomum subsp.
mattamuskeetense (Ashe) Freckmann
& Lelong, comb. & stat. nov.—20(1):169
Dichanthelium dichotomum subsp.
microcarpon (Muhl. ex Elliott)
Freckmann & Lelong, comb. & stat.
nov.—20(1):169
Dichanthelium dichotomum subsp.
nitidum (Lam.) Freckmann & Lelong,
comb. & stat. nov.—20(1):169
Dichanthelium dichotomum subsp.
roanokense (Ashe) Freckmann &
Lelong, comb. & stat. nov.—20(1):1 70
Dichanthelium dichotomum subsp.
yadkinense (Ashe) Freckmann & Lelong,
comb. & stat. nov.—20(1):1 70
Dichanthelium sect. Ensifolia (Hitchc.in
B.L.Rob.& Fernald) Freckmann & Lelong,
stat. nov.—20(1):166
Dichanthelium ensifolium subsp.
curtifolium (Nash) Freckmann & Lelong,
comb. & stat. nov.—20(1):1 70
Dichanthelium sect. Lancearia (Hitchc.in
B.L.Rob.& Fernald) Freckmann & Lelong,
stat. nov.—20(1):166
1828
Dichanthelium sect. Lanuginosa (Hitchc.
in B.L.Rob. & Fernald) Freckmann &
Lelong, stat. nov.—20(1):166
Dichanthelium sect. Linearifolia
Freckmann & Lelong, sect. nov.—
20(1):164
Dichanthelium sect. Macrocarpa
Freckmann & Lelong, sect. nov.—
20(1):165
Dichanthelium sect. Nudicaulia Hitch-
cock & Chase ex Freckmann & Lelong,
sect. Nov.—20(1):165
Dichanthelium sect. Oligosantha (Hitchc.
in B.L.Rob. & Fernald) Freckmann &
Lelong, stat. nov.—20(1):166
Dichanthelium oligosanthes subsp.
scribnerianum (Nash) Freckmann &
Lelong, comb. & stat. nov.—20(1):1 70
Dichanthelium ovale subsp. praecocius
(Hitchc. & Chase) Freckmann & Lelong,
comb. & stat. nov.—20(1):170
Dichanthelium ovale subsp. pseudo-
pubescens (Nash) Freckmann & Lelong,
comb. & stat. nov.—20(1):170
Dichanthelium ovale subsp. villo-
sissimum (Nash) Freckmann & Lelong,
comb. & stat. nov.—20(1):170
Dichanthelium sect. Pedicellata Hitchc.&
Chase ex Freckmann & Lelong, sect.
nov.—20(1):165
Dichanthelium portoricense subsp.
patulum (Scribn. & Merr.) Freckmann &
Lelong, comb. nov.—20(1):170
Dichanthelium sect. Sphaerocarpa
(Hitchc.in B.L.Rob.& Fernald) Freckmann
& Lelong, stat. nov.—20(1):166
Dichanthelium sect. Strigosa Freckmann
& Lelong, sect. nov.—20(1):166
Dichanthelium strigosum
subsp.
SIDA 20(4): 1828. 2003
BRIT.ORG/SIDA 20(4)
glabrescens (Griseb.) Freckmann &
Lelong, comb. & stat. nov.—20(1):171
Dichanthelium strigosum subsp.
leucoblepharis (Trin.) Freckmann &
Lelong, comb. & stat. nov.—20(1):171
Dichanthelium tenue (Muhl.) Freckmann
& Lelong, comb. nov.—20(1):171
Dieteria asteroides var. glandulosa (B.L.
Turner) D.R. Morgan & R.L. Hartman,
comb. nov.—20(4):1393
Dieteria asteroides var. lagunensis (D.D.
Keck) D.R.Morgan & R.L. Hartman, comb.
nov.—20(4):1393
Dieteria bigelovii (A. Gray) D.R. Morgan &
R.L. Hartman, comb. nov.—20(4):1394
Dieteria bigelovii var. commixta (Greene)
D.R.Morgan & R.L. Hartman, comb nov.—
20(4):1394
Dieteria bigelovii var. mucronata
(Greene) D.R. Morgan & R.L. Hartman,
comb. nov.—20(4):1394
Dieteria canescens subsp. glabra (A. Gray)
D.R. Morgan & R.L. Hartman, comb. nov.
—20(4):1396
Dieteria canescens var. ambigua (B.L.
Turner) D.R. Morgan & R.L. Hartman,
comb. nov.—20(4):1396
Dieteria canescens var. aristata
(Eastwood) D.R. Morgan & R.L. Hartman,
comb. nov.—20(4):1397
Dieteria canescens var. glabra (A. Gray)
D.R. Morgan & R.L. Hartman, comb. nov.
—20(4):1397
Dieteria canescens var.incana (Lindl.) DR.
Morgan & R.L. Hartman, comb. nov.—
20(4):1396
Dieteria canescens var.leucanthemifolia
(Greene) D.R. Morgan & R.L. Hartman,
comb. nov—20(4):1396
NEW NAMES AND NEW COMBINATIONS
Dieteria canescens var.nebraskana (BL.
Turner) D.R. Morgan & R.L. Hartman,
comb. nov.—20(4):1397
Dieteria canescens var. iliflora (Nutt.)
D.R. Morgan & R.L. Hartman, comb. nov.
—20(4):1396
Dieteria canescens var. shastensis (A.
Gray) D.R.Morgan & R.L.Hartman, comb.
nov.—20(4):1396
Dieteria canescens var. ziegleri (Munz)
D.R. Morgan & R.L. Hartman, comb.
nov.—20(4):1396
Dioscorea lisae Dorr & Stergios, sp. nov.—
20(3):1007
Eriogonum riograndis Nesom, sp. nov.—
20(1):32
Eurybia glauca var. pulchra (S.F. Blake)
Brouillet, comb. et stat.nov—20(4):1562
Festuca is STANCIK & PM.Peterson,
sp. nov.—20(1):22
Festuca dentiflora E.B. Alexeev ex STANCIk
& PM. Peterson, sp. nov.—20(3):1016
Festuca is STANCIK & PM. Peterson,
sp. nov.—20(1):24
Gentianella boliviana (Pax) J.S.Pringle,
comb. nov.—20(4):1486
Gentianella inaequicalyx (Gilg) Fabris ex
J.S.Pringle, comb. nov.—20(4):1488
Glandularia verecunda forma albida
(Moldenke) B.L. Turner, comb. nov.—
20(4):1648
Gonolobus tenuisepalus Krings, sp.
nov.—20(1):105
Ipomoea costellata var. edwardsensis
O’Kennon & Nesom, var. nov.—20(1):39
Ipomoea tacambarensis E. Carranza, sp.
nov.—20(4):1351
SIDA 20(4): 1829. 2003
1829
Leandra urbaniana (Alain) Alain, comb.
nov.—20(4):1645
Liatris elegans var. kralii Mayfield, var.
nov.—20(2):601
Liatris elegans var. bridgesii Mayfield, var.
nov.—20(2):598
Liatris elegans var. elegans forma
flabellata (Small) Mayfield, comb.nov.—
20(2):603
Liatris gholsonii L.C. Anderson, sp.nov.—
20(1):98
Liatris patens Nesom & Kral, sp. nov.—
20(4):1579
Liatris savannensis Kral & Nesom, sp.
nov.—20(4):1574
Lithospermum hintoniorum B.L. Turner,
sp. nov.—20(2):501
Matelea monticola Alain, nom. nov.—
20(4):1645
Maytenus urbanii Alain, nom. nov.—
20(4):1645
Medinilla anamalaiana Sasidharan &
Sujanapal, sp. Nov.—20(1):1 10
Meliosma chiriquensis J.F. Morales, sp.
nov.—20(3):934
Meliosma clandestina J.F. Morales, sp.
nov.—20(3):934
Meliosma depressiva J.F. Morales, sp.
nov.—20(3):937
Packera texensis RJ. O’Kennon & D.K.
Trock, sp. nov.— 20(3):945
Panicum aciculare var. neuranthum
(Griseb.) Wipff, comb et stat. nov.—
20(3):1044
Panicum aciculare var.ramosum (Griseb.)
Wipff, comb nov.—20(3):1044
Panicum acuminatum var. ferventicola
(Schmoll) Wipff, comb. et stat. nov.—
20(3):1044
1830
Panicum acuminatum var. thermale
(Bol.) Wipff, comb. nov.—20(3):1044
Panicum amarum subsp. amarulum
(Hitchc. & Chase) Freckmann & Lelong,
comb. & stat. nov.—20(1):171
Panicum anceps subsp. rhizomatum
(Hitchc. & Chase) Freckmann & Lelong,
comb. & stat. nov.—20(1):171
Panicum sect. Antidotalia Freckmann &
Lelong, sect. nov.—20(1):163
Panicum capillare subsp. hillmanii
(Chase) Freckmann & Lelong, comb. &
Stat. Nov.—20(1):171
Panicum commutatum var. equilaterale
(Scribner) Wipff, comb. et stat. nov.—
20(3):1044
Panicum dichotomiflorum subsp.
bartowense (Scribn.& Merr.) Freckmann
& Lelong, comb. & stat. nov.—20(1):171
Panicum dichotomiflorum subsp.
puritanorum (Svenson) Freckmann &
Lelong, comb. & stat. nov.—20(1):171
Panicum hallii subsp. filipes (Scribn.
Freckmann & Lelong, comb. & stat.
nov.—20(1):172
Panicum sect. Hemitoma Hitchc. & Chase
ex Freckmann & Lelong, sect. nov.—
20(1):163
Panicum hirticaule subsp. sonorum
Vasey) Freckmann & Lelong, comb. &
stat. nov.—20(1):172
Panicum hirticaule subsp. stramineum
(Hitchc. & Chase) Freckmann & Lelong,
comb. & stat. nov.—20(1):172
Panicum ovale var. praecocius (Hitchcock
& Chase) Wipff, comb. nov.—20(3):1044
Panicum philadelphicum = subsp.
gattingeri (Nash) Freckmann & Lelong,
comb. & stat. nov.—20(1):1 72
ae
—
SIDA 20(4): 1830. 2003
BRIT.ORG/SIDA 20(4)
Panicum philadelphicum subsp. litho-
philum (Swallen) Freckmann & Lelong,
comb. & stat. nov.—20(1):172
Panicum rigidulum subsp. abscissum
(Swallen) Freckmann & Lelong, comb. &
stat. nov.—20(1):1 72
Panicum rigidulum subsp. combsii
(Scribn.& CR. Ball) Freckmann & Lelong,
comb. & stat. nov.—20(1):1 72
Panicum rigidulum subsp. elongatum
(Pursh) Freckmann & Lelong, comb. &
Stat. NOV.—20(1):1 72
Panicum rigidulum subsp. pubescens
(Vasey) Freckmann & Lelong, comb. &
stat. nov.—20(1):173
Panicum rigidulum var. abscissum
(Swallen) Wipff, stat. nov.—20(3):1045
Parathesis pseudocalophylla Ricketson
& Pipoly, sp. nov.—20(3):899
Parathesis pseudocrassiramea Ricket-
son & Pipoly, sp. nov.—20(3):902
Parathesis subg. Laterales Ricketson &
Pipoly, subg. nov.—20(3):915
Parathesis subg. Laterales sect. Calo-
phyllae Ricketson & Pipoly, sect. nov.—
20(3):915
Parathesis subg. Parathesis sect. Cuba-
nae Ricketson & Pipoly, sect. nov.—
20(3):916
Parathesis subg. Laterales sect.Laterales
Ricketson & Pipoly, sect.nov.—20(3):915
Parathesis subg. Parathesis sect. Versa-
tiles Ricketson & Pipoly, sect. nov.—
20(3):917
Polygonum aviculare subsp. buxiforme
(Small) Costea & Tardif, comb. et stat.
nov.—20(3):988
ie}
(Wolf & McNeill) Costea & Tardif, comb.
nov.—20(3):994
NEW NAMES AND NEW COMBINATIONS
Polygonum humifusum subsp. cauria-
num (Robinson) Costea & Tardif, comb.
et stat. nov.—20(3):995
Polygonum ramossisimum subsp. proli-
ficum (Small) Costea & Tardif, comb. et
stat. nov.—20(3):995
Polygonum sawatchense subsp. obli-
vium Costea & Tardif, subsp. nov.—
20(4):1637
Polygonum striatulum var.texense (M.C.
Johnston) Costea & Tardif, comb. et stat.
nov.—20(3):996
Portulaca matthewsii G.Ocampo, sp. nov.
—20(4):1358
Portulaca rzedowskiana G. Ocampo, sp.
nov.—20(2):487
Pseudognaphalium biolettii A. Anderb.
nom. nov.]—20(1):149
Psidium salutare var. decussatum (DC,)
Landrum, comb. nov.—20(4):1463
Psidium salutare var. mucronatum
(Cambess.) Landrum, comb. nov.—
20(4):1463
Psidium salutare var.pohlianum (O. Berg
Landrum, comb. nov.—20(4):1466
Psidium salutare sericeum
(Cambess.) Landrum, comb. nov.—
20(4):1467
Psychotria bulilimontis Takeuchi, sp.
nov.—20(4):1514
Psychotria bulilimontis var. aestuarii
Takeuchi, var. nov.—20(4):1519
Pteroceras monsooniae Sasicdharan &
Sujanapal, sp. nov.— 20(3):923
_
—
Var.
Sagraea rugosa Alain, nom. nov.—
20(4):1645
Salix exigua var. sericans (Nees) Nesom,
comb. nov.—20(2):523
Schistocarpha calzadana B.L. Turner, sp.
nov.—20(2):505
SIDA 20(4): 1831. 2003
1831
Senecio sandersiana B.L. [urner,sp.nov.—
20(1):175
Senecio spribillei W.A. Weber, sp. nov.—
20(2):511
Siphonandra boliviana Luteyn,sp.nov.—
20(1):13
Solidago altissima subsp. gilvo-
canescens (Rydb.) J.C.Semple, comb.et
stat. nov.—20(4):1606
Solidago arenicola B.R. Keener & Kral, sp.
nov.—20(4):1589
Solidago ser. Auriculatae Semple, ser.
nov.—20(4):1606
Solidago ser. Drummondiani Semple, ser.
nov.—20(4):1606
Solidago faucibus Wieboldt, sp. nov.—
20(4):1596
Solidago subsect. Humiles (Rydb.)
Semple, comb.et stat. nov.—20(4):1606
Solidago kralii Semple, sp. nov.—
20(4):1606
Solidago lepida subsp. fallax (Fernald)
Semple, stat. nov.—20(4):1611
Solidago lepida var. salebrosa (Piper) J.C.
Semple, comb. nov.—20(4):1611
Solidago subsect. Multiradiatae (Juz.)
Semple, comb. nov.—20(4):1605
Solidago odora subsp. chapmanii (A.
Gray) J.C. Semple, comb. nov.—
20(4):1611
Solidago ser.Odorae (Mackenzie in Small)
Semple, comb.et stat. nov.— 20(4):1606
Solidago patula subsp. strictula (Torr. &
A. Gray) J.C. Semple, stat. nov.—
20(4):1612
Solidago puberula subsp. pulverulenta
(Nutt.) J.C. Semple, comb. et stat. nov.
20(4):1612
Solidago rugosa var. cronquistiana
Semple, var. nov.—20(4):1612
1832
Solidago sempervirens subsp. azorica
(Hochst. ex Seubert) Semple, comb. et
stat. nov.—20(4):1615
Solidago sempervirens subsp.mexicana
(L.) Semple, comb. et stat. nov.—
20(4):1615
Solidago speciosa subsp. pallida (Porter)
Semple, comb.et stat. nov.—20(4):1615
Solidago stricta subsp. gracillima (Torr.&
A, Gray) Semple, comb. et stat. nov.—
20(4):1615
Solidago velutina subsp. californica
(Nutt.) Semple, comb. et stat. nov.—
20(4):1616
Solidago velutina subsp. sparsiflora (A.
Gray) Semple, comb. et stat. nov.—
20(4):1616
Sphaeralcea caespitosa var. williamsiae
N.H. Holmgren, var. nov.—20(1):49
Stylogyne pucuroensis Ricketson &
Pipoly, sp. nov.—20(3):919
Symplocos kothayarensis Sundaresan,
Jothi, Rajkumar & Manickam, sp. nov.—
20(3):928
xTaxodiomeria peizhongii Z.J. Ye, JJ.
Zhang et S.H. Pan, gen. et sp. nov.—
20(3):1001
Thelesperma pubescens
caespitosum (Dorn) CJ. Hansen, stat.
nov.—20(1):91
Thelesperma windhamii C.J. Hansen,
nom. et stat. nov.—20(1):91
Themistoclesia unduavensis Luteyn, sp.
nov.—20(1):15
Trichadenia sasae Takeuchi, sp. nov.—
20(4):1522
Var.
Weinmannia bradfordiana Z. Rogers, sp.
nov.—20(1):180
SIDA 20(4): 1832. 2003
BRIT.ORG/SIDA 20(4)
Weinmannia condorensis Z. Rogers, sp.
nov.—20(1):183
Xanthisma arenarium (Benth.) D.R. Mor-
gan & R.L. Hartman, comb. nov.—
20(4):1405
Xanthisma blephariphyllum (A. Gray) D.R.
Morgan & R.L. Hartman, comb. nov.—
20(4):1403
Xanthisma section Blepharodon (DC.)
D.R. Morgan & R.L. Hartman, comb.
nov.—20(4):1401
Xanthisma coloradoense (A. Gray) DR.
Morgan & R.L. Hartman, comb. nov.—
20(4):1403
Xanthisma crutchfieldii (B.L. Turner) DR.
Morgan & R.L. Hartman, comb. nov.—
20(4):1403
Xanthisma glaberrimum (Rydb.) Nesom
& O'Kennon, comb. nov.—20(4):1586
Xanthisma gracile (Nutt.) D.R. Morgan &
R.L. Hartman, comb. nov.—20(4):1405
Xanthisma grindelioides (Nutt.) D.R. Mor-
gan & RL. Hartman, comb. nov.—
20(4):1403
Xanthisma grindelioides var.depressum
(Maguire) D.R. Morgan & R.L. Hartman,
comb. nov.—20(4):1404
Xanthisma gymnocephalum (DC.) DR.
Morgan & R.L. Hartman, comb. nov.—
20(4):1404
Xanthisma gypsophilum (B.L. Turner) D.R.
Morgan & R.L. Hartman, comb. nov.—
20(4):1404
Xanthisma incisifolium (!.M. Johnston)
Nesom, comb. nov.—20(4):1585
Xanthisma section Havardii (R.C. Jackson)
D.R. Morgan & R.L. Hartman, comb.
nov.—20(4):1410
Xanthisma johnstonii (S.F. Blake) D.R. Mor-
NEW NAMES AND NEW COMBINATIONS
gan & R.L. Hartman, comb. nov.—
20(4):1404
Xanthisma junceum (Greene) D.R. Mor-
gan & R.L. Hartman, comb. nov.—
20(4):1406
Xanthisma restiforme (B.L. Turner) DR.
Morgan & R.L. Hartman, comb. nov.—
20(4):1405
Xanthisma rhizomatum (M.C. Johnston)
D.R. Morgan & R.L. Hartman, comb.
nov.—20(4):1405
Xanthisma section Sideranthus (Nutt. ex
Nees) D.R.Morgan & R.L.Hartman,comb.
nov.—20(4):1405
Xanthisma spinulosum (Pursh) D.R. Mor-
gan & R.L. Hartman, comb. nov.—
20(4):1406
Xanthisma spinulosum var. chihuahua-
num (B.L. Turner & R.L. Hartman) D.R.
Morgan & R.L. Hartman, comb. nov.—
20(4):1408
Xanthisma spinulosum var. glaberri-
mum (Rydb.) D.R.Morgan & R.L. Hartman,
comb. nov.—20(4):1408
SIDA 20(4): 1833. 2003
1833
Xanthisma spinulosum var. gooddingii
(A. Nelson) D.R. Morgan & R.L. Hartman
comb. nov.—20(4):1408
Xanthisma spinulosum var. incisifolium
(ILM. Johnston) D.R. Morgan & R.L. Hart-
man, comb. nov.—20(4):1409
Xanthisma spinulosum var. paradoxum
(B.L. Turner & R.L. Hartman) D.R. Morgan
& R.L. Hartman, comb.nov.—20(4):1409
Xanthisma spinulosum var. scabrellum
(Greene) D.R. Morgan & R.L. Hartman,
comb. nov.—20(4):1409
Xanthisma stenolobum (Greene) D.R.
Morgan & R.L. Hartman, comb. nov.—
20(4):1409
Xanthisma viscidum (Wooton &
Standley) D.R. Morgan & R.L. Hartman,
comb. nov.—20(4):1410
Xanthisma wigginsii (S.F. Blake) D.R. Mor-
gan & R.L. Hartman, comb. nov.—
20(4):1409
Xylorhiza linearifolia (7.J. Wats.) Nesom,
comb. nov.—20(1):145
Yucca cernua Keith, sp. nov.—20(3):892
1834 BRIT.ORG/SIDA 20(4)
SIDA GUIDELINES FOR CONTRIBUTORS
Sida home page: http://www.brit.org/sida/
Guidelines: http://www.brit.org/Sida/SubmitPaperhtm
Sida, Contributions to Botany is an international journal of systematic botany
containing primary research papers sensu lato, including anatomy, biogeogra-
phy, chemotaxonomy, cladistics, ecology, floristics, genetics & evolution, nu-
merical taxonomy, paleobotany, and palynology. Sida is open to all authors any-
where; coverage is not restricted to any geographical area.
All manuscripts submitted to Sida are considered by at least two review-
ers. Manuscripts may be submitted in English or Spanish. Page costs are $35 per
page but may be waived or reduced under certain circumstances. Please con-
tact the editor and make arrangements before publication. Manuscripts are not
rejected due to lack of financial support. Submit manuscripts and editorial ques-
tions to Barney Lipscomb, Botanical Research Institute of Texas, 509 Pecan
Street, Fort Worth, TEXAS 76102-4060, U.S.A. Phones: 1-817-332-4441 voice; l-
817-332-4112 fax. Our electronic mail address is sida@brit.org. Access our Sida
home page at the URL http://www.britorg/sida/
Please submit manuscripts in electronic format as an email attachment. If
you wish submissions are also accepted by USPS mail with a single hard copy
and on computer disk. Figures/illustrations may be submitted in low-resolu-
tion format Gpgs) as email attachments for review purposes only. Upon accep-
tance for publication, the manuscript may be submitted as an email attach-
ment or returned in the mail with a single hard copy and an electronic copy. If
you have figures/illustrations in electronic format they should be submitted in
tifformat, scanned at the absolute best quality possible. Digital figures may have
to be submitted on a CD-ROM disk if they are too large for email. Typically b/w
line drawings should be scanned at 600 dpi or higher and b/w half tones and
color photos at 300 dpi. If you need your figures/illustrations digitized, the edi-
torial staff at Sida will be happy handle any electronic digitization needs you
have; all original material can be returned if requested. Color digitization works
best from color transparencies (slides).
Consult the latest issue of Sida for format of articles and notes regarding
title, author, and address. Ranges of numbers are separated by double hyphens
(6 use an en dash. Footnotes, figure legends, appendices, and tables
should be on separate pages and inserted at end of manuscript following refer-
ences. Authors are encouraged to use the APG families, nearly all of which are
on /APweb/ which can be accessed through the Missouri Botanical Garden re-
search site (Angiosperm phylogeny).
SIDA 20(4): 1834. 2003
GUIDELINES FOR CONTRIBUTORS 1835
ABSTRACT
Every paper should include both an English and Spanish, or another major lan-
guage abstract.
SPECIMEN CITATIONS
Use the collector’s last name and number or (s.n) and underline. Abbreviate the
months of the year by the first three letters of the month without a period. Ex-
ample: Texas. Jeff Davis Co.:1 mi W of Fort Davis, 3 Jan 1972, Smith 118 (BRIT).
AUTHOR CITATIONS
Use the author abbreviations as given in the Author Abbreviations compiled at
the Herbarium, Royal Botanic Gardens, Kew (1992). Use et or the ampersand
(&) between two authors; for more than two authors, restrict to the first one
followed by et al. Do not italicize or underline in the manuscript the terms et, et
al., ex, or in.
LATIN DESCRIPTIONS OR DIAGNOSES; ENGLISH DESCRIPTIONS
Please provide a Latin diagnosis for each new taxon. After the Latin diagnosis,
an English or Spanish version is recommended highlighting the diagnostic char-
acters. Write plant descriptions as incomplete sentences with phrases separated
by semicolons.
FIGURES
Please present ALL figures (maps included) as a unified, single sequence of lig-
ures. A scale should be in or on the illustration or photograph; reduction will
automatically reduce the scale and subject proportionately. In halftones, sharp
glossy photographs with good contrast are necessary for good reproduction.
Please mount illustrative material (halftones, line drawings, etc.) on flexible
paper so that may be mounted and on a high-speed drum scanner if need. You
may also submit graphic materials in electronic format. Typically b/w line
drawings should be scanned at 600 dpi or hig er and b/w half tones and color
photos at 300 dpi. Please consult the editor for additional information if need.
Color figures can be used and an additional cost maybe charged. Color digitiza-
tion works best from color transparencies (slides). Please submit color prints
for reviewing purposes.
REFERENCES
This includes all of the literature cited in the text and may include other article
citations the author may deem desirable. Normal text references should be cited
SIDA 20(4): 1835. 2003
1836 BRIT.ORG/SIDA 20(4)
as follows: ‘Ricketson and Pipoly (1997) stated...” or ‘the latest revision (Ricketson
S Pipoly 1997) when reference is used as authority fora statement. When there
are three or more authors use only the name of the first author followed by et
al: ‘Barrie et al. 1992a) stated’. References at the end of the article are arranged
alphabetically and chronologically making use of a,b, etc. if an author had more
than one publication in a given year. Author’s names are printed in Cap/sMALL
caps, NOT ALL CAPS; please type author names in Cap/sMALL Cap or in Cap/
lower case except for initial capitals. Only the first letter of the initial word,
proper nouns, and proper adjectives of titles should be capitalized. Please note
there is no space between initials of authors. For journal abbreviations, use
Botanico-Periodicum-Huntianum. Reference examples:
Barrie, FR. CE. Jarvis, and J.L. Reveal. 1992a. The need to change Article 8.3 of the Code.
Taxon 41:508—512.
Ricketson, J. and J. Pipoly. 1997. A synopsis of the genus Gentlea (Myrsinaceae) and a key
to the genera of Myrsinaceae in Mesoamerica. Sida 17:697—707. [author's names in
cap/lower case]
Crawrord, D.J.1983.Phylogenetic and systematic inferences from electrophoretic studies
In: S.D. Tanksley and TJ. Orton, eds. Isozymes in plant genetics and breeding, Part A.
Elsevier, Amsterdam. Pp. 257—287. [author's names in cap/small cap]
Sivinski, R.C. and K. LicHTroor (eds.). 1995. Inventory of rare and endangered plants of New
Mexico. New Mexico Forestry and Resources Conservation Division. Energy, Minerals
and Natural Resources Department. Misc. Pub. No. 4.
ABBREVIATIONS
When the following abbreviations are used the period is omitted. Distance: mm,
dm, cm, m, km, ft, mi; directions (in caps): 5, N, E, W, months: first 3 letters only,
Jan, Feb, etc. Example: Potter Co.: 2 km W of Dot, 5 Jun 1971, Smith 118 (SMU).
DOCUMENTED PLANT CHROMOSOME NUMBERS
Refer to Sida 20(4):1663. 2003.
SIDA 20(4): 1836. 2003
T lavishly illustrated monograph is the most
comprehensive botanical treatment of beans to
date. It begins with a brief history about the former
taxonomical treatments of the genus, and goes on
with the taxonomical criteria and a presentation
about discriminant characteristics. It presents a full
description of each section and species, its distribu-
tion and habitat, relationships with other species,
uses and potentially useful traits, and historical notes.
Color pictures, line drawings, and distribution maps
lead easily to the right identification of each species.
Available from
Botanical Research Institute of Texas Press
509 Pecan Street
Fort Worth, TX 76102-4060, USA
E-mail: sida@brit. org" "Fax 1-817-332-4112
{cid } lsbhm? 3toc htm
Q)
www.brit
Sida, Bot. Misc. No. 23, 2002
issn 0833-1475,isbn 1-88878-11-1 +7" = 10", xviii + 300 pp,
97 b/w figures and distribution ba a 5 color plates (60 figs.).
h»=(USA 1 $8)
Texas residents sda $3.30 sales tax.
Taxonomy, Distribution,
and Ecology
of the Genus Phaseolus
(Leguminosae-
Papilionoideae)
in North America, Mexico
and Central America
By
George F. Freytag
Daniel G. Debouck
he Atlas of Texas covers about 6000 taxa. This is
Tin result of 54 years of herbarium and fieldwork
by B.L. Turner, beginning in 1948 at Sul Ross State
University, Alpine, Texas. In short, the senior author
has examined personally, touched, or “pored over”
an estimated several hundred thousand sheets in
the preparation of the forthcoming Atlas volumes.
Contents include an introduction, atlas of Texas
plants arranged alphabetically by family, by genus,
by species, and an index.
Available from
Botanical Research Institute of Texas Press
509 Pecan Street
Fort Worth, TX 76102-4060, USA
E-mail: sida@brit.org = Fax 1-817-332-4112
www.brit.org/sida/sbm/sbm24toc.htm
a, Bot. Misc. No. 24, 2003, issn 0833-1475,
isbn (vol. 1) 1-889878-08-1 «isbn (vol. 2) 1-889878-09-X = 7 1/2" * 10"
Vol. 1 approx. 630 pp. Vol.2 approx. 275’pp.
Vol. 1 $50 + p&h* «Vol. 2 $40 + p&h* « Set $80 + p&h*
*USA:$10 (vol. 1); $9.50 (vol. 2); $12 (set)
*International: $12.50 (vol. 1) «$11.50 (vol. 2); $25 (set)
Texas residents add sales tax: $4.13 (vol. 1) « $3.30 (vol. 2) « $7.43 (set)
Atlas of the
Vascular Plants of Texas
By
B.L. Turner,
Holly Nichols, Geoffrey
Denny, Oded Doron
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