Au AN ARRAINDSAD FLORIDA TOVWA ilar
“eonraBuTiOns i
oe AR INDIA . Oi (Eu NSLA
Barney L. Lipscomb, Editor
Botanical Research Institute of Texas
Fort Worth, Texas 76102-4060, USA 817 332-4441 / 817 332-4112 FAX
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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
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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. (ISBN 0-88192-502-0, 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). $29.95, 144 pp, 148 color photos, 4 color plans, 4 b/w plans, 81/2" = 1"
Creating a “sanctuary” is a popular mae turning ee ntly in the recent crop of gardening lit-
's how y t as
sina hec tic
=
erature. British garden writer Stephe st environment. The emphasis in Ur on Sanctuaries is on the traditional b idea: but Anderton's bee
concepts will apply to smaller and more exotic areas as well. Anderton’s approach is a down-to-
earth yet mee pieeniatie of alternatives ranging from cutting-edge ene of pi contem-
porary plots to family-oriented onan and container gardens that make maximum use of mini- mum space.
Anderton's concepts remain constant throughout his narrative. In Part One: The Body of the Garden he addresses the fundamental basics of garden design. What type, full or partial, and time of sun does the plot receive? What type of soil does your garden have? What plants or design elements are already there, and will you include them or not? What is your preferred planting style: tradi- tional, tropical, minimalist? Do you want to incorporate a water feature? The scale of plantings as
move-
—
well as water and soil supplies must be considered carefully. To make it seem larger, the pace o ment through the area should be slowed and there must be a place to sit, w hich seems ania oe during t1 le design stage the fact that eventually one will want to spe nd leis
often overlooked. The broad scope of his vision encompasses all categories of stylistic Abie lexarions with the aim of directing the reader to a design path best suited to one’s fundamental predilections. By looking at innovative designs for varied styles of urban gardens Stephen Anderton shows how
they can be adapted to create spaces that are not only imaginative but are also imbued with a mood of peace and serenity
Part Two: The Spirit of the Garden follows Anderton's musing on different garden themes and ideas of how to make those themes a reality. He uses case studies to provide design solutions com-
plete with diagrams for gardens with magic with mystery, the natural approach, devoted to water, minimalist style, and more. A selected directory, complete with US hardiness zones, of 72 plants that
meet his criteria of those that are “tough, easy to grow, and ... have a long season”. Plants selected range from trees to perennials. rominent personality in the world of English Ciena Stephen Anderton writes a Saturday
column for The Times (London) and regularly contributes to Horticulture, The Garden, and Gardens
Illustrated. This unique resource for the city dweller should appeal to gardeners of all levels —Gary Jennings, Botanical Research Institute of Texas, Fort Worth, TX 76102-4060, U.S.A
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
Atwoop, N.D. and S.L. We tsk (In press). Sphaeralcea gierischii N.D. Atwood & S.L.Welsh.
Cronauist, A. ET AL. 1972-1997. Intermountain flora: vascular plants of the Intermountain 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.
54 BRIT.ORG/SIDA 20(1)
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
a
bs Buoar pso7
ey 90S'Lzg= Baly ( vwseunjes judy
PmeSUN
> Buaqsnqueg es
Buaquaddei3 uayonisjawey,
oe, savin ae _ \ ee <a 2 } “4 “ s Pe Gms y \,- os Oss Ls —— om, WAY IES @ 7K ; : APs, ares] & id J} aa teat é GE ye ers oe > awoaese ! =D. v& — “ Se! 5 j tN ee i HSV : = s
* «
=
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=
L
L
« <= s = i= S S —, wn. i i la cS 4 w 4 s- 4 o- _" “
< a +
58 BRIT.ORG/SIDA 20
=
1)
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.
REFERENCES
A.ten, G.R.1991.Field guide to the freshwater fishes of New Guinea. Christensen Research Institute Publ. 9:1-268.
Davies, H.L.and D.S. Hurcuison, 1980a. Papua New Guinea 1:250,000 geological series map. Ambunti, sheet SB 54-4. Geological Survey of Papua New Guinea, Dept. Minerals and Energy
Davies, H.L. and D.S. HurcHison. 1980b. Explanatory notes to accompany the Ambunti 1:250,000 geological map. Report 80/1.Geological Survey of Papua New Guinea, Dept. Minerals and Energy. Pp 1-25.
Dinc Hou. 1960. Thymelaeaceae. Flora Malesiana ser. 1, 6 (1):1-48.
Gentry, A. 1988. Changes in plant community diversity and floristic composition on envi- ronmental and geographical gradients. Ann. Miss. Bot. Gard. 75: 1-34.
GIDEON, O.G. 1998. Systematics and evolution of the genus Tapeinochilos Mig. (Costaceae- Zingiberales). Ph.D. thesis. James Cook University of North Queensland, Australia.
Hammermaster, E.T. and J.C. Saunpers. 1995a. Forest resources and vegetation mapping of Papua New Guinea. PNGRIS Publ. 4, Canberra, CSIRO and AIDAB.
HamMmermaster, E.T. and J.C. Saunbers. 1995b. Forest resources and vegetation mapping of Papua New Guinea. 1:250,000 vegetation map overlays separately issued as working copies to PNGRIS Publ. 4, Canberra, CSIRO and AIDAB
JoHNs, R.J. 1986. The instability of the tropical ecosystem in New Guinea. Blumea 31:341- SF i
Louman, B.and S. Nichoits. 1995. Forestry in Papua New Guinea. In:N.Sekhran and S. Miller, eds. Papua New Guinea country study on biological diversity. Colorcraft Ltd, Hong Kong. Pp 155-167.
NATIONAL StaristiCAL Orrice. 2001. Papua New Guinea 2000 census preliminary figures. In: Weekender Post Courier. August 3, 2001: 10-11.
PAUMANS, K,, (ed.) 1976. New Guinea vegetation. CSIRO and Australian National University Press, Canberra.
Papua New Guinea Forest AuTHority. 1998. Forest resource acquisition general information. PNG Forest Authority.
Pickam, C.J. and H.L. Davies. 1987. Terranes and the accretion history of the New Guinea orogen. J. Austr. Geol. Geoph. 10:193-211,
70 BRIT.ORG/SIDA 20(1)
SoHMER, S.H., R. Kiapranis, A. ALLISON, and W. TakeucH!. 1991 (unpublished). Report on the Hunstein River expedition — 1989. Pp 1-70.
STEENIS-KauUsEMAN, MJ. vAN. 1950. Malaysian plant collectors and collections. Being a cyclo- pedia of botanical exploration in Malaysia and a guide to the concerned literature up to the year 1950. FI. Male. ser. |, 1:1-639.
Takeuchi, W.and M. Gouman. 2002. The identity of eaglewood (Gyrinops, Thymelaeaceae), a new economic resource for Papua New Guinea. Sida 20:261-267.
TakeucHi, W.and S.S. Renner. 2002. A generic record for Faika (Monimiaceae) in Papua New Guinea. Fl. Male. Bull. 13:54-55.
Townsenp, G.W.L. 1968. District officer, from untamed New Guinea to Lake Success, 1921- 46. Pacific Publications, Sydney, 1-271.
VetoKame, J.F,,W. Vink, and D.G. Fropin. 1988. XI. Ledermann’s and some other German locali- ties in Papua New Guinea. FI. Male. Bull. 10: 32-38.
Warner, JN.and C.O. Grassi. 1958.The 1957 sugar cane expedition to Melanesia. Hawaiian Planters’ Record 55:209-236.
Wewzen, PC. van. 1997. Increased speciation in New Guinea: tectonic causes? In:J.Dransfield, M.J.E. Coode, and D.A. Simpson, eds. Plant diversity in Malesia Ill. Proceedings of the Third International Flora Malesiana Symposium 1995. Pp 363-387.
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. REFERENCES Avexanper, E.J. 1955. Thelesperma, North American flora, Ser. II, Pt. 2:65-69. Avise, J.C. 1994. Molecular markers, natural history, and evolution. Chapman and Hill,
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